The Fishes of the Amazon: Distribution and Biogeographical Patterns, with a Comprehensive List of Species

Fernando C.P. Dagosta; Mário De Pinna

We provide a general compilation of the diversity and geographical distribution of Amazonian fishes, updated to the end of 2018. Our database includes documented distributions of 4214 species (both Amazonian and from surrounding basins), compiled from published information plus original data from ichthyological collections. Our results show that the Amazon basin comprises the most diverse regional assemblage of freshwater fishes in the world, with 2716 valid species (1696 of which are endemic) representing 529 genera, 60 families, and 18 orders. These data permit a view of the diversity and distribution of Amazonian fishes on a basinwide scale, which in turn allows the identification of congruent biogeographical patterns, here defined as the overlapping distributions of two or more lineages (species or monophyletic groups). We recognize 20 distinct distributional patterns of Amazonian fishes, which are herein individually delimited, named, and diagnosed. Not all these patterns are associated with identifiable geographical barriers, and some may result from ecological constraints. All the major Amazonian subdrainages fit into more than one biogeographical pattern. This fact reveals the complex history of hydrographical basins and shows that modern basin-defined units contribute relatively little as explanatory factors for the present distributions of Amazonian fishes. An understanding of geomorphological processes and associated paleographic landscape changes provides a far better background for interpreting observed patterns. Our results are expected to provide a framework for future studies on the diversification and historical biogeography of the Amazonian aquatic biota.

THE F ISHES OF THE A MA ZON: D ISTRIBUTION AND B IOGEOGR APHICAL PATTERNS, WITH A C OMPREHENSIVE L IST OF S PECIES F E R N A N D O C . P. D A G O STA A N D M Á R I O D E P I N N A BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY THE FISHES OF THE AMAZON: DISTRIBUTION AND BIOGEOGRAPHICAL PATTERNS, WITH A COMPREHENSIVE LIST OF SPECIES FERNANDO C.P. DAGOSTA Universidade Federal da Grande Dourados Faculdade de Ciências Biológicas e Ambientais Dourados, Mato Grosso do Sul, Brazil MÁRIO DE PINNA Museu de Zoologia da Universidade de São Paulo São Paulo, Brazil BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 431, 163 pp., 22 figures Issued June 13, 2019 Copyright © American Museum of Natural History 2019 ISSN 0003-0090 CONTENTS Abstract. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5 Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6 Results and Discussion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8 Patterns of Distribution of Amazonian Fishes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12 Broadly Distributed Lineages. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12 Amazon Core . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 South American Lowlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 Amazon and Orinoco Lowlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16 Amazon and Paraguay Lowlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18 Amazon-only Lowland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20 Amazonas-Paraguay-Orinoco Lowland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20 Amazonas-Guiana-Orinoco Lowland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21 Other Cases of Amazonian Lowland Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22 Fossils of Lowland Amazonian Lineages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22 Eastern Lowland Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22 Marine Derived Lineages. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22 Deep Channel Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 Guiana Mangrove Province . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 Eastern Amazon (East of the Purus Arch) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 Amazon-core Uplands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28 Amazonian Uplands. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29 Guiana Shield (Atlantic and Amazonian Versants). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .30 Longitudinal Shield Correspondence among Amazonian Shield Versants. . . . . . . . . . . . . .31 Brazilian Shield . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .31 Extreme Shield: Chapada dos Parecis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33 Exclusive Faunal Sharing between Neighboring Basins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .34 Tapajós and Paraguay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .35 Tapajós and Xingu. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .35 Tapajós and Madeira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .35 Xingu and Paraguay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .37 Xingu and Tocantins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .37 Tocantins and Paraguay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .37 Tocantins and São Francisco . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .37 Tocantins and Upper Paraná . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38 Madeira and Paraguay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38 Branco and Essequibo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38 Negro and Orinoco . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38 Negro and Branco . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39 Negro to Trombetas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39 Poorly Sampled Neighboring Basins in the Guiana Shield. . . . . . . . . . . . . . . . . . . . . . . . .40 Cis-Andean Foothills. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 Central Blackwater Amazon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .41 Allopatric Branco-Tocantins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .44 2 Absence Patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .45 Endemism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .46 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .47 Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .47 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .48 Appendix 1. Taxonomic list of Amazonian fish species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .64 Appendix 2. Statistical test for species with distribution maps . . . . . . . . . . . . . . . . . . . . . . . . . . . 160 3 ABSTRACT We provide a general compilation of the diversity and geographical distribution of Amazonian fishes, updated to the end of 2018. Our database includes documented distributions of 4214 species (both Amazonian and from surrounding basins), compiled from published information plus original data from ichthyological collections. Our results show that the Amazon basin comprises the most diverse regional assemblage of freshwater fishes in the world, with 2716 valid species (1696 of which are endemic) representing 529 genera, 60 families, and 18 orders. These data permit a view of the diversity and distribution of Amazonian fishes on a basinwide scale, which in turn allows the identification of congruent biogeographical patterns, here defined as the overlapping distributions of two or more lineages (species or monophyletic groups). We recognize 20 distinct distributional patterns of Amazonian fishes, which are herein individually delimited, named, and diagnosed. Not all these patterns are associated with identifiable geographical barriers, and some may result from ecological constraints. All the major Amazonian subdrainages fit into more than one biogeographical pattern. This fact reveals the complex history of hydrographical basins and shows that modern basin-defined units contribute relatively little as explanatory factors for the present distributions of Amazonian fishes. An understanding of geomorphological processes and associated paleographic landscape changes provides a far better background for interpreting observed patterns. Our results are expected to provide a framework for future studies on the diversification and historical biogeography of the Amazonian aquatic biota. INTRODUCTION its fish species and available estimates suggest a number between 1300 and 3500 species (e.g., Géry, 1969; Lundberg et al., 2000, 2010; Junk et al., 2007; Albert et al., 2011; van der Sleen and Albert, 2017). Some studies have tried to correlate Amazonian fish distributions with underlying causal factors that might form the basis for a historical biogeography of these fishes: Eigenmann (1909), Géry (1969), Vari (1988), Jégu (1992a, 1992b), Hubert and Renno (2006), Ribeiro (2006), Albert et al. (2011), Albert and Carvalho (2011), Lima and Ribeiro (2011), Ribeiro et al. (2013), and Dagosta and Pinna (2017). Some other papers focused on only parts of the Amazon are still relevant for the recognition of distribution patterns, including: Kullander (1986), Jégu and Keith (1999), Pearson (1937), Crampton (2011), López-Fernández and Albert (2011), and Lujan and Armbruster (2011). The paradigm of vicariant biogeography (Croizat et al., 1974; Rosen, 1978; Nelson and Platnick, 1981) postulates that general distribution patterns demand general explanations, usually by associating lineage splitting with the origin of wide-ranging geographical barriers. Despite such conceptual clarity, in practice, the The Amazon basin is the largest hydrographic drainage on earth, covering ~6 × 106 km2 (larger still if estuarine coastal areas are included) (Sioli, 1984; Milliman and Farnsworth, 2011), or about one-third of South America. Its discharge is also the largest in the world, with about one-fifth of the entire freshwater volume on the surface of the planet (Callede et al., 2004). Its vast size is matched by an equally vast fauna and flora (Webb, 1995; Patton et al., 2000; Hoorn and Wesselingh, 2010; Cardoso et al., 2017), comprising the richest ecosystem on earth. Fishes are one of the faunal elements whose Amazonian biodiversity reaches superlative numbers. Despite such megadiversity and the attention it attracts, knowledge about the diversity and geographical distribution of Amazonian fishes is still not synthesized into a general framework that allows broad generalizations. Most data are scattered within thousands of species descriptions, and attempts to synthesize that information are inherently limited by the incomplete nature of their underlying databases. The Amazon basin still lacks a comprehensive list of 5 6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY association of biogeographical patterns with geographical barriers that contributed to taxonomic diversification is difficult because of factors such as dispersal across existing barriers, area coalescence (obliteration of preexisting barriers and resulting biotic dispersal), extinction, and others that make the detection and delimitation of historically cohesive areas (biogeographical units) a complex process (Harold and Mooi, 1994; Szumik et al., 2002). One way to approach the issue is to search for geographical homologies (i.e., biogeographical congruence) in the form of distributional congruence among unrelated taxa (Patterson, 1981; Nelson, 1994; Morrone, 2009) and the application of the notion of primary homology (de Pinna, 1991) as adapted to biogeography (sensu Morrone, 2001, 2009). Despite such caveats, there is ample consensus that the formulation of robust historical biogeographical hypotheses is critically important for understanding biotic diversification and in deciding what conservation policies to adopt (Cracraft, 1994; Crisci, 2000; Whittaker et al., 2005; Guedes et al., 2014). Analytical considerations aside, the identification of repeated patterns of geographical distribution is the first step toward a formulation of general biogeographical hypotheses. Such a task in itself can be daunting when dealing with large and complex taxonomic groups such as the Amazonian fishes. Basic knowledge about their taxonomy, phylogeny, and distribution has long been so irregularly scattered as to impede proper synthesis. On the other hand, knowledge about the systematics and distribution of Amazonian fish taxa has now accumulated to a degree that synthetic efforts are more enticing than ever before, both at specific and supraspecific levels. The purpose of this paper is to identify the taxonomic patterns of distribution of Amazonian fishes based on all data currently available in the literature and in some of the largest ichthyological collections with significant Amazonian holdings. The list compiled for this report is an expansion and refinement of the database published in Dagosta and Pinna (2017), the largest previously NO. 431 done on the distribution of Amazonian fishes and provides the first comprehensive list of Amazonian fishes. It permits the identification and delimitation of all repeated patterns of distribution. We also offer a discussion on the possible underlying causes for each of the patterns and on their potential as indicators of a general biogeographical history of the Amazon basin. We expect our work will provide a general framework for the categorization of forthcoming distributional data (new records, new species, and new clades) and facilitate future progress on the biogeography of Amazonian freshwaters. MATERIALS AND METHODS Species distributions were compiled from all the information available in the literature, in a total of over 1500 references (see Dagosta and Pinna, 2017: appendices 1–6; and appendix 1, herein, for distributional data on fish species in the Amazon and surrounding basins), including taxonomic revisions, species descriptions, inventories and faunistic lists. Additionally, primary data were obtained from the most relevant (in size and Amazonian coverage) ichthyological collections, namely Instituto Nacional de Pesquisas da Amazônia, Manaus (INPA), Museu Nacional do Rio de Janeiro, Rio de Janeiro (MNRJ ), Museu Paraense Emílio Goeldi, Belém (MPEG), Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP), and National Museum of Natural History, Washington, DC (USNM). Two smaller collections, LBP (Laboratório de Biologia e Genética de Peixes, Botucatu), and LIRP (Laboratório de Icitiologia de Ribeirão Preto, Ribeirão Preto), were also surveyed because of their unique holdings of material from critical portions of the Brazilian Shield. Examined material is listed in Dagosta and de Pinna, 2017: Supplementary Material 2, and includes all information on source of data (citation, date of publication, and catalog number, when based on collections), institutional catalog, and sample numbers along with published sources for each species. All published information utilized was 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES qualified as to accuracy regarding species identification and locality. Identifications in collections were verified by direct examination of specimens. Doubtful information was discarded. Compilation of the composition and geographical distribution of Amazonian fishes is updated to the end of 2018. Maps presented are intended to represent general patterns of distribution and individual species plots can vary slightly within those limits. Delimitation of the Purus Arch follows Sacek (2014). Water type of Amazonian Rivers follows Venticique et al. (2016) (SNAPP Western Amazon Group – Amazon Aquatic Ecosystem Spatial Framework Knowledge Network for Biocomplexity. Usage is granted according to a Creative Commons “CC BY 4.0”). Hydrographic shape used in figures 5 through 22 is from Lehner et al. (2008), courtesy from HydroSHEDS (hydrological data and maps based on shuttle elevation derivatives at multiple scales: http://www.worldwildlife.org/hydrosheds). The shaded relief of South America used in figures 5 through 22 is courtesy of NASA/JPL-Caltech. The delimitation of the Amazonian regions follows Dagosta and Pinna (2017). The taxonomic arrangement in appendix 1 follows Nelson (2006), except for the inclusion of recently described family Tarumaniidae (de Pinna et al., 2017), for considering Arapaimidae and Serrasalmidae as valid families, for including the genus Chalceus in Alestidae, and for adopting Cynolebiidae instead of Rivulidae. Within each family, genera are organized in alphabetical order. Species-level nomenclature follows Fricke et al. (2019), except for the validity of Astyanacinus, which is maintained herein since its synonymization within Astyanax is considered unjustified. In figure 1, estimates of the number of fish species in the Congo basin is from Snoeks et al. (2011), in the Mekong from Poulsen et al. (2004), in the Nile from Witte et al. (2009), in the Mississippi from Robinson and Buchanan (1988) and in the Ganges from Sarkar et al. (2012). Figures for other basins (Atrato, Capim, Cauca-Magdalena- 7 Sinú, Coppename-Suriname, Corentyne-Demerara, Essequibo, Maracaibo, Maroni-Approuague, Oiapoque, Paraná-Paraguay, and Parnaíba) are compiled from the present work. Estimates of fish diversity in continents follow Lévêque et al. (2008), except for Europe, which is based on Kottelat and Freyhof (2007); South America, based on Reis et al. (2016); and Central America, based on Matamoros et al. (2015). A distributional pattern is herein identified as the overlap (or major overlap) of geographical distributions of at least two species or monophyletic groups. Our definition of biogeographical pattern is deliberately broad and based solely on instantaneous geographical distributions of taxa. Thus, the recognition of a pattern herein is agnostic as to its causal reasons. It also does not preclude the possibility that some of the patterns are hierarchically arranged, i.e., that some patterns are actually subpatterns of larger ones. The selection and the delimitation of the distributional patterns are based on a visual analysis of map distributions of all Amazonian fishes, with consideration for previously published proposals. Repeated geographical distributions may be the result of ecological conditions, historical factors or both, and we consider that the identification and characterization of distribution patterns are relevant regardless of their underlying explanation and even without confirmation by rigorous statistical modeling procedures (which would be impossible at this point because of the heterogeneity of data sources). Actual geographical distributions are a result of past events and processes in combination with present-day constraints not necessarily related to the vicariant processes that generated that diversity. Many factors are brought into consideration when untangling the nature and meaning of a distribution pattern, and those are discussed separately in the relevant sections. Our notion of taxa assumes that they exist as empirically backed biological entities, corresponding to either species or monophyletic groups. Taxonomic entities that are demonstrably not monophyletic are not considered as valid evidence, 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 0 500 1000 1500 NO. 431 2000 2500 3000 Amazonas 2716 Congo 1250* Mekong 1200* Orinoco 1001 Nile 800* Paraná-Paraguay Essequibo Yangtze 362 299 2716 287 Mississippi 260 Coppename-Suriname 253 Cauca-Magdalena-Sinu Atrato 2954 293 Maroni-Approuague Ganges 3500 1411 514 Corentyne-Demerara Capim 546 740 233 176 143 118 Maracaibo 109 Parnaíba 108 Oiapoque 103 2434 260 SPECIES 3500 3000 2500 2000 1500 1000 500 0 Asia Africa Amazonas Rest of S. N. America Europe America Central America Oceania FIG. 1. Number of fish species in the Amazon basin and comparisons with other basins and continents. Left graph: species numbers in the Amazon and other large world basins. Bottom graph and map: species numbers in the Amazon basin compared to those in other continents. An asterisk indicates estimated numbers. For sources of data, see Material and Methods. even if formally named in classification, because they are not empirically supported as a basis for biogeographical inference. A chi-square test was used to verify the association of species’ distributions with biogeographical patterns proposed (i.e., whether or not the species has a random distribution) and to test whether water type influences the distributions of specific species (appendix 2). The test was applied only to species shown in maps. RESULTS AND DISCUSSION Our inventory shows that the Amazonian ichthyofauna is composed of 2716 valid species, included in 529 genera, 60 families, and 18 orders. Such figures make the Amazon drainage, by a wide margin, the basin with the richest fish fauna in the world (fig. 1), with a diversity equivalent to that of some entire continents (fig. 1). As a comparison, the estimated (i.e., not necessarily described) number of species in the second most diverse basin in the world (Congo) is less than half that of the Amazon. As in other Neotropical, African, and Asian drainages, the majority of Amazonian fishes belongs to the Otophysi (fig. 2), a group representing 80% (2193 spp.) of all Amazonian species. As in other Neotropical basins, the most species-rich orders are Characiformes and Siluriformes. The third largest order in Amazonian waters is the Perciformes, largely due to species of the family Cichlidae. Of the least diverse orders, 10 are from typically marine lineages that secondarily invaded Amazonian waters. The familial composition also follows that a pattern typical of the majority of continental waters in the neotropics, vastly dominated by small body-size species. Five families (Characidae, Loricariidae, Cichlidae, Cynolebiidae, and Callichthyidae) concentrate most of the diversity (1528 spp. or 56% of Amazonian species), with Characidae alone comprising nearly a quarter of all Amazonian fish species. 2019 9 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES A Characiformes B 1063 Siluriformes Characidae Perciformes Gymnotiformes Myliobatiformes 27 Clupeifomes 18 361 Cichlidae 286 Cyprinodontiformes 635 Loricariidae 956 166 Cynolebiidae 164 Callichthyidae 258 141 133 Anostomidae 91 Heptapteridae 87 Pleuronectiformes 9 Auchenipteridae 79 Beloniformes 9 Trichomycteridae 75 Osteoglossiformes 5 Curimatidae 74 Synbranchiformes 3 Doradidae 73 Pristiformes 2 Serrasalmidae 72 Batrachoidiformes 2 Apteronotidae 62 Tetraodontiformes 2 Pimelodidae 60 Anguiliformes 1 Lebiasinidae 49 Atheriniformes 1 Crenuchuidae 47 Carcharhiniformes 1 Sternopygidae 29 Ceratodontiformes 1 Potamotrygonidae 27 Species 0 200 400 600 800 1000 C Corydoras 0 1200 200 300 400 500 600 700 111 Hyphessobrycon 82 Moenkhausia 74 Apistogramma 70 Leporinus 55 Hemigrammus 49 Crenicichla 49 Creagrutus 43 Anablepsoides 36 Astyanax 34 Ancistrus 34 Hypostomus 30 Pimelodella 28 Melanorivulus 27 Cyphocharax 26 Gymnotus 25 Sternarchorhynchus 25 Characidium 24 Species 0 100 20 40 60 80 100 120 FIG. 2. Richest lineages of Amazonian fishes in number of species: A, All orders; B, families; C, genera. The most species-rich genera are those composed of small body-size species. Among the 10 richest genera, half are Characidae (fig. 2C), demonstrating that the diversity of that family in the Amazon is concentrated in few genera. The same happens with Corydoras, the genus with the most Amazonian species, comprising 111 of the 133 species of Callichthyidae in the basin. Among the most diverse Amazonian genera, three were the object of relatively recent revisions (Creagrutus, Cyphocharax, and Ster narchorhynchus). Such revisionary works increased significantly the number of Amazonian species in the respective lineages: 42% of the Amazonian species of Creagrutus were described in Vari and Harold (2001); 28% of 10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 0 A Scoloplacidae (6/0) Lepidosirenidae (1/0) Tarumaniidae (1/0) Gasteropelecidae (8/1) Chilodontidae (7/1) Hemiodontidae (26/5) Rhamphichthyidae (23/6) Cynodontidae* (10/3) Doradidae (73/24) Serrasalmidae (72/26) Potamotrygonidae (27/10) Acestrorhynchidae (10/4) Ctenoluciidae (5/2) Hypopomidae (24/10) Lebiasinidae (49/26) Apteronotidae (62/35) Auchenipteridae (79/44) Curimatidae (74/42) Anostomidae (91/57) Cetopsidae (26/17) Callichthyidae (133/87) Arapaimidae (3/2) Polycentridae (3/2) Gymnotidae (26/10) Sternopygidae (29/22) Pimelodidae (60/55) Aspredinidae (23/22) Characidae (635/657) Crenuchuidae (47/48) Pristidae (2/3) 20 40 60 80 100 100 89 11 88 13 84 16 79 21 77 23 75 25 73 27 73 27 71 29 71 29 71 29 65 35 64 36 64 36 64 36 61 39 60 40 60 40 60 40 60 40 57 43 57 52 43 48 51 49 49 51 49 40 * Roestes included in Cynodontidae 0 100 100 51 60 20 40 Heptapteridae (87/132) 38 Prochilodontidae (8/13) 37 Loricariidae (361/622) 33 Osteoglossidae (2/4) 33 Pseudopimelodidae (16/33) 32 Parodontidae (10/21) 31 Cynolebiidae (141/315) 28 Erythrinidae (5/13) 26 Achiridae (9/26) 24 Trichomycteridae (75/232) 21 Astroblepidae (17/64) 21 Belonidae (8/31) 15 Cichlidae (258/1459) 13 Pristigasteridae (5/33) 12 Synbranchidae (3/22) Engraulidae (12/144) 8 Cyprinodontidae (7/96) 7 Poeciliidae (17/257) 6 Anablepidae (1/17) 6 Sciaenidae (15/278) 5 Eleotridae (9/163) 5 Alestidae (5/111) 4 Batrachoididae (2/82) 2 Carcharhinidae (1/58) 2 Hemiramphidae (1/60) 2 Tetraodontidae (2/189) 1 Atherinopsidae (1/109) 1 Clupeidae (1/203) <1 Ophichthidae (1/336) <1 Gobiidae (1/1887) <1 % of species ocurring in Amazon NO. 431 40 60 80 100 60 62 63 67 67 68 69 72 74 76 79 79 85 87 88 92 93 94 94 95 95 96 98 98 98 99 99 100 100 100 % of species not ocurring in Amazon FIG. 3. Proportion of species occurring in the Amazon versus not occurring in the Amazon, by higher taxon. A. (above) By family. In green, percentage of species in the family occurring in the Amazon; in pink, percentage of species not occurring in the Amazon. B. (opposite page) Per genera (only 50 most species-rich genera shown). In green, percentage of species occurring in the Amazon; in blue, percentage of species not occurring in the Amazon. Fractions in parentheses represent actual numbers of species occurring (numerator) and not occurring (denominator) in the Amazon. Number of species in each family and genus follows Fricke et al. (2019). Cyphocharax in Vari (1992a), and 72% of Ster narchorhynchus in de Santana and Vari (2010). Thus, the position of those genera among the most diverse in the Amazon, especially in the case of Cyphocharax and Sternarchorhynchus, may be an artifact resulting from the lack of taxonomic revisions in other potentially more diverse yet poorly studied genera, for example, Chaetostoma, Knodus, and Rineloricaria. Interesting facts appear when the diversity of the richest Amazonian lineages is compared with equivalent data from outside the basin. The Amazonian fish community is formed both by typically Amazonian lineages (i.e., most or all of their diversity is in the Amazon basin) and by lineages that have greater diversity in other South American basins or in the ocean (fig. 3A). At the family level, it is clear that the Amazon dominates, by a wide margin, the alpha-diversity in the vast majority of primary-division lineages sensu Myers (1938), with relatively little diversity in the secondary and peripheral divisions. The only exclusively Amazonian family is the recently described Tarumaniidae (Scoloplacidae and Lepidosirenidae occur also in the Paraguay basin). Other interesting patterns are revealed by an examination of intrageneric diversity. The first one is that the most diverse genera in the Ama- 2019 11 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 0 B 20 40 Corydoras (111/64) Hyphessobrycon (81/75) 52 Hemigrammus (49/10) Crenicichla (49/46) 64 49 34 66 45 55 Cyphocharax (26/19) 58 Gymnotus (25/18) 58 Sternarchorhynchus (25/7) Characidium (24/44) 85 19 71 36 29 31 % of species ocurring in Amazon zon have the majority of their species in the basin (fig. 3B). Also, some megadiverse Neotropical genera are relatively poorly represented in the Amazon, such as Astyanax, Trichomycterus, Bryconamericus, Characidium, and Hemibrycon. This is probably related to the fact that the species of those genera in the Amazon are concentrated along its outer rims, which indicate that their presence in the region is relatively recent, caused by secondary geological events involving adjacent basins. Two cases that stand out are Astyanax and Trichomycterus, which have their Amazonian diversity concentrated respectively in the upper Tocantins and upper Ucayali, right at the divide with other drainages. The richest Amazonian tributary is the Rio Madeira (fig. 4), a fact that is attributed not only to its large size, but also to its hybrid nature (Dagosta and Pinna, 2017). The Rio Madeira drains one-third of the Amazonian lowlands and 25 58 65 35 70 30 93 Hypoptopoma (14/1) 93 7 30 46 54 67 33 55 Charax (12/5) 45 71 29 57 Aspidoras (11/13) 43 46 54 25 Curimata (11/2) Hemibrycon (11/40) 7 70 Centromochlus (12/6) Geophagus (11/20) 64 69 75 Cichla (14/1) Brycon (11/33) 6 45 42 Peckoltia (12/9) 27 81 Brachyhypopomus (20/8) Knodus (20/9) 15 54 73 Serrasalmus (22/5) 94 55 Cetopsis (12/10) 65 46 Potamotrygon (22/8) Bryconamericus (20/35) 22 6 91 Ituglanis (13/15) 42 78 Jupiaba (23/4) Chaetostoma (22/26) 9 Otocinclus (14/6) 42 35 79 Bryconops (14/6) 80 14 94 Steindachnerina (15/8) 51 20 21 Pimelodus (15/21) 80 Ancistrus (34/36) Astroblepus (17/64) Nannostomus (15/5) 36 20 100 100 Farlowella (16/13) 42 80 86 Pyrrhulina (18/0) Trichomycterus (17/162) 60 71 Aequidens (16/1) 48 58 40 Bujurquina (17/1) 17 52 Anablepsoides (36/20) Melanorivulus (27/33) 25 32 83 Creagrutus (43/31) 20 29 Hemiodus (18/3) 19 68 0 Rineloricaria (19/47) 48 75 Leporinus (55/26) Pimelodella (28/54) 100 81 Apistogramma (70/23) Hypostomus (30/121) 80 37 Moenkhausia (74/16) Astyanax (34/132) 60 63 75 85 15 35 22 65 78 % of species not ocurring in Amazon at the same time has tributaries associated with the Andean range and the Brazilian Shield, both areas particularly rich in fish endemics. The Rio Negro is also one of the most diverse Amazonian tributaries (fig. 4), despite its nutrient-poor and extremely acidic waters, which represent ecological barriers to numerous fish species (Goulding et al., 1988; Lima and Ribeiro, 2011). Despite that, the Rio Negro also harbors a large number of endemics (79 spp). High levels of endemism are also seen in the Marañon and Ucayali (25% and 16%, respectively; fig. 4), a likely result of the Andean range in the diversification of the Amazonian biota. The largest absolute number of endemic taxa (87 spp.) is found in the upper Rio Tocantins followed by the Marañon (81 spp.). Other basins draining Amazonian Shield regions, such as Juruena, Teles Pires, Jari, and Madeira Shield tributaries, also show high levels of endemism, 12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Amazonas main channel Middle-lower Madeira Negro Branco Mamoré Beni-Madre de Dios Guaporé Arabuaia Ucayali Trombetas Japurá Lower Tocantins Purus Putumayo Lower Xingu Upper Tocantins Upper Xingu Tapajós Marañon-Nanay Madeira Shield tributaries Teles Pires Napo-Ambyiacu Juruá Urubu-Uatumã Amazonas estuary Jari Iriri Juruena Jamanxim Tefé Xingu total Tapajós total Tocantins total Negro total Madeira total 71/8% 32/2% 79/12% NO. 431 922 708 682 23/4% 601 20/4% 563 34/7% 509 22/4% 506 68/16% 424 66/16% 416 18/4% 35% 414 14/3% 410 19/5% 403 10/3% 379 5/1% 378 13/4% 367 87/24% 358 15/5% 333 18/5% 332 80/25% 326 28/8% 314 63% 2% 22/7% 298 22/9% 257 4/2% 251 8/4% 211 5/3% 182 Species endemic to Amazon basin (1696 spp.) 15/8% 174 Amazonian species also occurring in neighboring basins (959 spp.) 19/11% 169 23/15% Poorly known species (61 spp.) 151 2/2% 109 1/1% 104 502 529 705 922 1062 0 200 400 600 800 1000 FIG. 4. Amazonian endemic species. Species richness (green) and endemic species (orange) for each of the main Amazonian tributaries or parts thereof. Pie chart indicates the proportion of species occurring exclusively in the Amazon and those also present in other basins. Minor endemic regions not included in graph are: Anapu (2 spp.), Coari-Urucu (1 sp.), Curuá-Paru do Oeste (2 spp.), Paru (2 spp.) and Javari (1 sp.). although relatively low total numbers of species. Such figures corroborate the hypothesis that highlands have proportionally fewer taxa with broad distributions and fewer cases of sympatry when compared with lowland rivers (Dagosta and Pinna, 2017). The majority of Amazonian fish diversity is exclusive, with 63% (1696 spp.) of its species found nowhere else (fig. 4). That number alone would place the Amazon as the richest basin in the world in fish species. listed is described as to its geographical (physical) boundaries and subsequently discussed as to its historical significance, possible causes (if any is identified) and any other relevant points. The taxa supporting each pattern is provided in the text. Patterns are arranged according to logical sections and subsections, with each of them given a name (when a pattern was already recognized in the literature by a widely used name, that name was maintained here) and a number reference, so as to provide an easy reference in future publications. PATTERNS OF DISTRIBUTION OF AMAZONIAN FISHES Broadly Distributed Lineages Below we present the general patterns of distribution herein recognized on the basis of information compiled in appendix 1. Each category This pattern includes lineages which are broadly distributed in major South American drainages such as Amazon, Coastal Atlantic, 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 13 FIG. 5. Broadly distributed lineages. A. Yellow area delimits the distribution pattern. B. Hoplias malabaricus (red dots; records from MZUSP), Erythrinus erythrinus (green diamonds; records from MZUSP), Hopleryth rinus unitaeniatus (yellow triangles; records from MZUSP). C. Synbranchus marmoratus, records from MZUSP. D. Callichthys callichthys (data from Lehmann and Reis (2004) with additional records from MZUSP). Parana-Paraguay, São Francisco, Orinoco, and Guianan basins (fig. 5A). This is the most common pattern for the majority of families of fishes in the Amazon and also includes a number of genera such as Acestrorhynchus (see González, 2015), Brycon (see Lima, 2017), Characidium, Crenicichla (see Ploeg, 1991), Corydoras, Eigen mannia, Geophagus, Gymnotus (see Albert et al., 2005), Hoplosternum (see Reis, 1997), Hyposto mus, Leporinus, Megaleporinus (see Ramirez et al., 2017), Pimelodella (see Slobodian, 2017), Pimelodus, Prochilodus, Serrasalmus, Schizodon, and Steindachnerina (see Vari, 1991). As to species-level taxa with such wide distribution, three categories exist: (1) well-studied ones that have been extensively revised and are demonstrably a single taxonomic entity, such as Acestrorhynchus falcatus (see González, 2015) and Callichthys callichthys (fig. 5D; see Lehmann and Reis, 2004); (2) species that have never been the object of detailed revisions but that nonetheless lack published evidence of comprising a complex of species, such as Corydoras aeneus, Leporellus vittatus, Leporinus fasciatus, Pinirampus piri nampu; and (3) taxa recognized or suggested as species complexes but that have not yet been disentangled, such as Eigenmannia virescens (L. Peixoto, personal commun.), Erythrinus erythrinus (Martins et al., 2012), Hoplerythrinus unitaeniatus (Giuliano-Caetano et al., 2001), Hoplias malabari cus (fig. 5B; see Dergam et al., 1998), Leporinus friderici (Silva-Santos et al., 2018), Sternopygus macrurus (Silva et al., 2008), and Synbranchus marmoratus (fig. 5C; see Torres et al., 2005). 14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Therefore, it is likely that this “pattern” may not actually be a congruent biogeographical pattern among species, but simply the result of a taxonomic artifact. More refined biogeographical patterns may be hidden under incomplete or deficient taxonomic definitions. It is interesting that none of the migratory fishes with notably vast living ranges, such as Brachyplatystoma spp., Prochilodontidae spp., and Curimatidae spp. include species with such broad ranges as those listed above (see Barthem and Goulding, 2007; Barthem et al., 2017). This is evidence that even species with extreme dispersal abilities meet with effective ecological/geographical barriers that keep them from achieving such broad ranges as defined in this pattern. Amazon Core The name given to this pattern has no relation to the origin of its components, but simply alludes to the region of greatest fish diversity in South America. The Amazon Core pattern is formed by fish lineages typical of the Amazonian biota that may be found both in lowlands and highlands of the Amazon basin and in adjacent basins such as Guiana drainages, Parnaíba, Capim, Orinoco, and Essequibo (fig. 6A) and that are absent in coastal drainages of southeastern Brazil, São Francisco, and Paraná-Paraguay. It is interesting to note that there is no species or lineage of fishes that is widely distributed throughout the high and low Amazonian lands not also present in adjacent basins of the Amazon such as Orinoco, Essequibo, or Guiana drainages. This is evidence that the Amazon basin is not an area of endemism, but instead is a historically composite area. Examples of the Amazon Core pattern are numerous and include supraspecific taxa such as Chilodontidae, Anostominae, genera Argonectes (see Langeani, 1996), Boulengerella (fig. 6B; see Vari, 1995), Cichla (see Kullander and Ferreira, 2006), Jupiaba (see Zanata, 1997; presence of the genus in Paraguay basin is hypothesized to be secondary according to Ribeiro et al., 2013), Mastigla NO. 431 nis, Panaque (see Lujan et al., 2010), and Semaprochilodus (see Castro and Vari, 2004). This pattern of distribution is also reflected in some individual species such as Moenkhausia collettii (fig. 6C), Moenkhausia oligolepis (fig. 6D) and Potamo trygon orbignyi (see Da Silva and De Carvalho, 2015). South American Lowlands As the name suggests, taxa following this pattern occupy only the lower altitudes of cisAndean South America, not occurring in Guiana and Brazilian Shield regions except in lowermost sectors of some large rivers such as Tapajós, Trombetas, and Xingu. Almost invariably, species with this pattern of distributions occur in the main channel of the Amazon and the Rio Madeira but exact limits vary according to taxon. Eigenmann was the first author to recognize that the ichthyofaunistic composition was different between South American high- and lowlands and named part of the latter as the “Amazon Province”: “East of the Cordilleras, and therefore east of the Magdalena basin, is found the most extensive and intricate fresh water system in the world. It forms a network of rivers practically uninterrupted, extending from the mouth of the Orinoco through the Casiquiare, Rio Branco, Rio Negro, Rio Madeira, Rio Guapore, Rio Paraguay, Parana and La Plata to Buenos Aires” (Eigenmann, 1909: 317). Eigenmann (1909) correctly pointed out that the pattern extrapolates the hydrographic limits of the Amazon basin, and it is possible to list countless other examples of species that occur in the lowlands of the Amazon that are also present in other lowland drainages such as Orinoco, Paraná-Paraguay, Essequibo, and drainages east of the mouth of the Amazonas, from the Capim to the Mearim. Although this pattern is not exclusively Amazonian, the highland/lowland divide is the most widely discussed of all distributional patterns of Amazonian fishes (see Menezes, 1969, 1976; Kullander, 1986; Jégu, 1992a, 1992b; Lima and Ribeiro, 2011). 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 15 FIG. 6. Amazon Core. A. Yellow area delimits the distribution pattern. B. Boullengerella spp. (data from Vari (1995) with additional records from MZUSP). C. Moenkhausia collettii (records from MZUSP). D. Moenkhau sia oligolepis (records from MZUSP). Lima and Ribeiro (2011) underscored a clear dichotomy between two geomorphological domains in northern cis-Andean South America: lowlands and highlands. Besides differences in historical-geomorphological parameters, highand lowland regions affect their associated biotas differently (Albert et al., 2011; Lima and Ribeiro, 2011). Lowland drainages are more susceptible to hydrogeological changes and are in general more dynamic than highland drainages. Lowland rivers tend to be more directly interconnected than rivers draining other geomorphological areas, due to the action of meanders, anastomoses, megafans and mouth-position changes during sea-level oscillations (Lundberg et al., 1988). All those factors are less intense or nonexistent in highland rivers, which are typically well fitted in valleys of exposed crystalline rock and do not undergo significant lateral movements (Lima and Ribeiro, 2011). Thus, lowland rivers undergo constant and much faster hydrogeographic changes and, as a consequence, congregate more taxa with broad distributions and more cases of sympatry than highland rivers. In addition, habitat stability provided by an enormous living space for lowland Amazonian fish species seems to be an important factor in decreasing the extinction rate of lineages (Lundberg et al., 1988; 2010), which in turn also results in increased diversity. Therefore, those are the factors that explain why most Amazonian fish species fit a South American lowland pattern. There are spatial differences among distinct groups of lowland Amazonian fishes, with at least five different subpatterns as explained below. The examples and subpatterns recognized herein have 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 431 FIG. 7. Amazon and Orinoco Lowlands. A. Yellow area delimits the distribution pattern. B. Moenkhausia lepidura (data from Marinho and Langeani, 2016). C. Potamorhina altamazonica (data from Vari, 1984). D. Vandellia cirrhosa (M.P., unpublished data). a direct relation with the complex geomorphological history of the Western Amazon. Amazon and Orinoco Lowlands Eigenmann (1909) delimited his Amazonian Province from a dispersionist perspective, where present-day connections among drainages provided the explanation for faunal sharing among basins. However, the correct interpretation for most of such massive ichthyofaunal sharing among different lowland South American basins is directly related to a complex historical context that began in the Upper Cretaceous, at least, with the formation of the Sub-Andean Foreland basin (Lundberg et al., 1998) and has little relation to present (and rather ineffective) physical connections (e.g., Casiquiare canal). The sub-Andean Foreland is a series of retroarc depressions lying to the east of the Andean Cordilleras that served as the main drainage axis of South America throughout the Upper Cretaceous to the Paleogene (Cooper et al., 1995; DeCelles and Giles, 1996; Lundberg, 1998; DeCelles and Horton, 2003; Albert and Reis, 2011; Lima and Ribeiro, 2011; Wesselingh and Hoorn, 2011). For much of its existence, the SubAndean Foreland was drained mostly by the proto-Amazon-Orinoco basin (Lundberg et al., 1998), even though the latter has also drained other areas of the South American platform further east. Both the pattern described here and the Amazon Province of Eigenmann (1909) match mostly (exclusive of the La Plata basin included in Eigenmann’s province) the spatial limits of 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 17 FIG. 8. Amazon and Paraguay Lowlands. A. Yellow area delimits the distribution pattern. B. Epapterus dispi lurus (data from Vari and Ferraris, 1998). C. Hemigrammus lunatus (data from Ota et al., 2014). D. Mesonauta festivus (data from Kullander and Silvergrip, 1991, and Schindler, 2005). the proto-Amazon-Orinoco, which was a continuous basin until its fragmentation in the late Miocene (ca. 10 Ma) as a result of the rise of the Vaupes Arch in eastern Colombia that separated the modern Orinoco and Amazon basins (Hoorn, 1994a; Cooper et al., 1995; Harris and Mix, 2002; Albert and Carvalho, 2011; but see Mora et al., 2010, for a more recent estimate). That barrier may have prevented lineages that diversified after its rise from increasing their range throughout all lowland regions and may also have caused the extinction of lineages in some of those basins. Those two factors may explain the absence of some typical Amazonian lowland forms in the Orinoco basin (see examples in Amazon-only Lowland). On the other hand, part of the faunal sharing between the Amazon and Orinoco may result from broad distributions before the modern separation between those basins, i.e., from the proto-Amazon-Orinoco. Still another hypothesis to explain the same pattern is megafan dynamics, geologically more recent (see Wilkinson et al., 2010). As will be discussed in the section Negro and Orinoco, the Canal Casiquiare does not seem to be a relevant dispersal route to explain the extensive list of taxa shared between the Amazon and Orinoco lowlands. Some examples of exclusive taxon sharing between the Amazon and Orinoco lowlands are: Acanthicus hystrix (see Chamon, 2016), Acestro rhynchus heterolepis (see González, 2015), Brachyhypopomus sullivani (see Crampton et al., 2016), Brycon amazonicus (see Lima, 2017), Bou lengerella maculata (see Vari, 1995), Cetopsis coecutiens (see Vari et al., 2005), Cynodon gibbus 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 431 FIG. 9. Amazon-only Lowland. A. Yellow area delimits the distribution pattern. B. Adontosternarchus balae nops (data from Mago-Leccia et al., 1985). C. Cetopsis candiru (data from Vari et al., 2005). D. Curimatella meyeri (data from Vari, 1992a). (see Toledo-Piza, 2000a), Colossoma macropo mum, Lasiancistrus schomburgkii (see Armbruster, 2005), Leptodoras paelongus (see Sabaj Pérez, 2005), Metynnis guaporensis and M. luna (see Ota, 2015), Moenkhausia comma, Moenkhau sia lepidura (fig. 7B, see Marinho and Langeani, 2016), Mylossoma albiscopum (see Mateussi, 2015), Nemadoras cristinae (see Sabaj Pérez et al., 2014), Paragoniates alburnus (see Quevedo, 2006), Peckoltia bachi (see Armbruster, 2008), Potamorhina altamazonica (fig. 7C, see Vari, 1984), Sorubim elongatus (see Littmann, 2007), Trachydoras brevis, T. gepharti, T. microstomus and T. nattereri (see Sabaj and Arce, 2017), Van dellia cirrhosa (fig. 7D), Adontosternarchus spp. (see Mago-Leccia et al., 1985), Brachyrhamdia spp. (see Slobodian, 2013), Chalceus spp. (see Zanata and Toledo-Piza, 2004), Compsaraia (see Bernt and Albert, 2017), Hassar spp. (see Birin- delli et al., 2011), Laemolyta spp. (see Mautari and Menezes, 2006), Liosomadoras spp. (see Birindelli and Zuanon, 2012), Microphilypnus spp. (see Caires and Figueiredo, 2011), Tenellus spp. (sensu Birindelli, 2014; Sabaj Pérez et al., 2014), and Sternarchogiton spp. (see de Santana and Crampton, 2007). Amazon and Paraguay Lowlands There are many Amazonian Lowland fish lineages that also occur in the Paraná-Paraguay basin, which was not permanently connected to the proto-Amazon-Orinoco. The location of the watershed divide between the proto-AmazonOrinoco River basin and the La Plata changed between the end of the Paleogene and the beginning of the Neogene (see Tagliacollo et al., 2015). Initially, it was the Chapare Buttress in 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 19 FIG. 10. Amazonas-Paraguay-Orinoco Lowland. A. Yellow area delimits the distribution pattern. B. Rhaphi odon vulpinus (data from Toledo-Piza, 2000a, with additional records from MZUSP). C. Sorubim lima (data from Littmann, 2007). D. Hypophthalmus oremaculatus (data from Littmann et al., 2015). the Late Oligocene (ca. 30–20 Ma) (Lundberg, 1998) and subsequently the Michicola Arch, starting during the Late Miocene (ca. 11.8–10 Ma) in the area of modern eastern Bolivia (Lundberg et al., 1998; Montoya-Burgos, 2003; Albert and Carvalho, 2011; Carvalho and Albert, 2011a). Several events may have permitted biotic dispersal between the Amazon and Paraguay: upper Paraguay captures of protoAmazonas-Orinoco headwaters (Lundberg et al., 1998), Amazon capture of upper Paraguay headwaters (Lundberg et al., 1998), river megafans involving the upper Río Mamoré and tributaries of the upper Río Paraguay (Wilkinson et al., 2006, 2010; Ota et al., 2014) and capture of upper Rio Paraguay into the upper Rio Guaporé (Ota et al., 2014). Because all possible connections between the Amazon and the Paraná-Par- aguay happened as a result of separate events of different ages, it is very likely that many species shared between those basins, despite their congruent distributions, lack temporal congruence. They correspond instead to cases of pseudocongruence, sensu Donoghue and Moore (2003), and are not biogeographically homologous. Taxa shared between those basins include: Aces trorhynchus abbreviatus (see González, 2015), Acestrorhynchus gr. lacustris (see González, 2015), Brachyhypopomus bombilla (see Crampton et al., 2016), Epapterus dispilurus (see fig. 8B; Vari and Ferraris, 1998), Hemigrammus lunatus (see fig. 8C; Ota et al., 2014), Mesonauta festivus (see fig. 8D; Kullander and Silfvergrip, 1991; Schindler, 2005), Moema spp. (see Costa, 2004), and Prionobrama spp. (see Quevedo, 2006). A complete list of species shared exclu- 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY sively between the Madeira and the Paraguay is presented in Madeira and Paraguay. Amazon-only Lowland Whitewater Amazonian rivers have high sediment and nutrient loads and a neutral pH, draining a relatively young Andean range. Major whitewater tributaries include the Marañón, Purus, Madeira, Juruá, Putumayo, Japurá, and Napo rivers. The whole Rio Amazonas system exhibits whitewater, although it receives other water types from various tributaries. There are few investigations into the impact of such water type changes on the biogeography of Amazonian fishes. Vari (1988) suggested that some curimatids are restricted to whitewater rivers and that their distribution may be more closely linked to ecological rather than historical factors. While the pattern is correct in some cases, we also agree with Lima and Ribeiro (2011: 157) that “some ecological factors that clearly influence fish distribution patterns in northern cisAndean South America, such as water typology, are, as mentioned previously, a consequence of geomorphological processes and, as such, possess a historical component.” Thus, it is possible that lowland species restricted to the Amazon reached such distribution from different causes and histories, either because they diversified after separation of the Orinoco from the protoAmazon-Orinoco basin or because they are whitewater dependent. Evidence suggests that the interpretation of Vari (1988) may be correct for a set of species showing this pattern of distribution. Some of the exclusively Amazonian lowland species are absent in the Rio Tocantins basin, having their distributions limited to the region of the mouth of the Madeira. This may indicate an association with whitewater since tributaries with that type of water become practically nonexistent downstream of that part of the Amazon river. Some examples of the Amazon-only Lowland pattern are: Adon tosternarchus balaenops (see fig. 9B; Mago-Leccia et al., 1985), Agoniates anchovia, Aphanotorulus NO. 431 horridus (see Ray and Armbruster, 2016), Apha notorulus unicolor (see Ray and Armbruster, 2016), Apionichthys nattereri (see Ramos, 2003), Brycon melanopterus (see Lima, 2017), Chalceus erythrurus (see Zanata and Toledo-Piza, 2004), Cetopsis candiru (see fig. 9C; Vari et al., 2005;), Cetopsis oliveirai (see Vari et al., 2005), Chaeto branchopsis orbicularis, Copella stigmasemion (see Marinho and Menezes, 2017), Crenicara punctu latum, Curimata aspera (see Vari, 1989a), C. kneri (see Vari, 1989a), Curimatella meyeri (see fig. 9D; Vari, 1992b), Cyphocharax spiluropsis (see Vari, 1992b), C. notatus (see Vari, 1992b), C. plumbeus (see Vari, 1992b), Denticetopsis seducta (see Vari et al., 2005), Hydrolycus scomberoides (ToledoPiza et al., 1999), Hypostomus pyrineusi (see Armbruster, 2003), Leporinus jamesi (see Garavello et al., 2014), Protocheirodon pi (see Vari et al., 2016), Mylossoma aureum (see Mateussi, 2015), Nemado ras elongatus, N. hemipeltis, N. humeralis (see Sabaj Pérez et al., 2014), Potamorhina latior (see Vari, 1984), Prionobrama filigera (see Quevedo, 2006), Psectrogaster amazonica (see Vari, 1989b), Pseudobunocephalus amazonicus (see Cardoso, 2008), P. bifidus (see Cardoso, 2008), Scoloplax dicra (see Schaefer et al., 1989), Sorubim maniradii (see Littmann, 2007), Steindachnerina bimaculata (see Vari, 1991), S. leucisca (see Vari, 1991), Ster narchella calhamazon (see Lundberg et al., 2013), Trachydoras steindachneri (see Sabaj and Arce, 2017), Aphyolebias spp. (see Costa, 2004), and Chaetobranchopsis spp. The pattern of distribution described herein is repeatedly supported as biogeographically coherent in the analyses of Dagosta and de Pinna (2017). Amazonas-Paraguay-Orinoco Lowland This pattern is the least common one among lowland species in the Amazon basin. Most cases are also present in the Tocantins basin, but not in Guianan drainages. Some examples of this pattern are: Abramites hypselonotus (see Vari and Williams, 1987), Curimatella dorsalis (see Vari, 1992a), Hypophthalmus oremaculatus (see fig. 10D; Littmann et al., 2015), Rhaphiodon vulpinus 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 21 FIG. 11. Amazonas-Guiana-Orinoco Lowland. A. Yellow area delimits the distribution pattern. B. Crenuchus spp. (data from MZUSP). C. Mesonauta spp. (records from Kullander and Silfvergrip, 1991; Schindler, 2003). D. Hemigrammus ocellifer (red dots; records from MZUSP), Hemigrammus unilineatus (yellow dots; data from MZUSP). (see fig. 10B; Toledo-Piza, 2000a), Roeboides affi nis (see Lucena, 2007), Sorubim lima (see fig. 10C; Littmann, 2007), and Mylossoma spp. (see Mateussi, 2015). The Amazonas-Paraguay-Orinoco Lowland pattern comprises areas from the previously described Amazon and Orinoco Lowlands as well as the Amazon and Paraguay Lowlands, and thus the associated geological processes are the same as discussed in the respective headings. Amazonas-Guiana-Orinoco Lowland Species with this pattern of distribution are broadly distributed in the lowlands of the Rio Amazonas and, in some cases, also of the Orinoco and Essequibo, but they are also found in the lowlands of Guiana coastal drainages. The pattern differs from the Amazon Core pattern described above because it is restricted to lowland lineages. Some examples of this pattern here recognized are: Iguanodectinae, Arapaima spp., Brachyplaty stoma spp., Copella spp. (see Marinho and Menezes, 2017), Crenuchus spp. (see fig. 11B; Campanario, 2002; Pires et al., 2016), Electropho rus spp., Heros spp., Hypophthalmus spp. (also in Paraguay basin, see Littmann et al., 2015), Meso nauta spp. (also in Paraguay basin, see fig. 11C; Kullander and Silfvergrip, 1991; Schindler, 2003), Osteoglossum spp., Pachypops spp. (see Casatti, 2002), Ageneiosus dentatus (see Ribeiro et al., 2017), Brachyhypopomus beebei, Brachyhypopo mus brevirostris and Brachyhypopomus regani (see Crampton et al., 2016), Caenotropus labyrinthicus (see Vari et al., 1995), Gnathocharax steindachneri, Hemigrammus unilineatus (fig. 11D), Hemigram mus ocellifer (fig. 11D), and Serrasalmus rhombeus. 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY The geomorphological explanations associated with this pattern as the evolution of the sub-Andean Foreland region (discussed in Amazon and Orinoco Lowlands) and the advance of the freshwater plume associated with marine regressions (discussed in Guiana Mangrove Province), which allowed contact the regions of Amazonas-Orinoco Lowland and the coastal drainages of Guiana. Other Cases of Amazonian Lowland Distribution Fossils of Lowland Amazonian Lineages Several Tertiary fossils belonging to typically lowland Amazonian taxa are present in regions currently lacking any Amazonian connection, such as Magdalena and Caribbean coastal rivers from the northern coast of Venezuela: Arapaima, Brachyplatystoma, Colossoma, Doras, Hydrolycus, and Phractocephalus (G. Ballen, personal communication; Lundberg et al., 1986, 1988, 2010; Lundberg, 1997, 2005; Sabaj Pérez et al., 2007). This demonstrates that such regions were in the past also part of some other distributional patterns described herein. If such fossils were not known, our understanding of the biogeographical history of the region would be severely incomplete or incorrect. Besides, the absence of extant representatives of the listed lineages in the Caribbean coastal rivers from northern coastal Venezuela and in Rio Magdalena basin is a clear demonstration of the dynamic nature of biogeographical phenomena, which change drastically over time and may bear little relationship to present-day physical barriers that determine the distribution of recent taxa. Eastern Lowland Amazon Some lowland distributional patterns comprise basins east of the mouth of the Rio Tocantins, beyond the eastern limit of the Amazon basin itself (i.e., Capim, Gurupi, Turiaçu, Mearim, Itapecuru, and Parnaíba). Interestingly, NO. 431 each of those basins has fewer and fewer Amazonian lineages as they get progressively farther from the mouth of the Amazon. Some of them are poorly sampled, mainly the Gurupi, Turiaçu, and Mearim, resulting in probably artifactual discontinuous distributions between neighboring basins and blurring details of the pattern. Species shared exclusively between the lower Amazon and above-cited eastern basins include: Brachychalcinus parnaibae (lower Tocantins and Parnaíba, see Reis, 1989), Brachyhypopomus pin nicaudatus (Amazon estuary, Capim, and Mearim, see Crampton et al., 2016), Corydoras jullii (lower Tocantins, Mearim, and Parnaíba, see Dagosta and Pinna, 2017); Apistogramma caetei (lower Tocantins, Capim, and Gurupi; see Dagosta and Pinna, 2017) and Nannostomus niti dus (Amazon estuary and Capim; see Dagosta and Pinna, 2017). Most examples of Amazonian lineages that occur in drainages to the east of the Amazon basin are also widespread in lowland waters of the Amazon. Some examples are: Anablepsoides uro phthalmus (Capim, Gurupi and Mearim, see Costa, 2006), Brachyplatystoma spp. (Capim, cf. Lundberg and Akama, 2005; Mearim and Parnaíba; see Ramos et al. Ramos et al., 2014), Cae notropus labyrinthicus (Capim, see Vari et al., 1995; Parnaíba; see Ramos et al., 2014), Curimata spp. (Itapecuru, see Barros et al., 2011; Parnaíba, see Ramos et al., 2014), Cynodon gibbus (Itapecuru, see Barros et al., 2011; Parnaíba, Toledo-Piza, 2000a), Gymnocorymbus thayeri (Gurupi, see Benine et al., 2015; Parnaíba, see Benine et al., 2015), Jupiaba polylepis (Parnaíba, see Ramos et al., 2014), Poptella compressa (Capim, Mearim, Parnaíba, and Turiaçu, see Reis, 1989; Itapecuru, see Barros et al., 2011), Pseudoplatystoma puncti fer (Itapecuru, Barros et al., 2011; Parnaíba, see Buitrago-Suárez and Burr, 2007) and Vandellia cirrhosa (Capim and Turiaçu, fig. 7D). Marine Derived Lineages A lowland Amazonian pattern is also seen in typically marine lineages (peripheral division 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 23 FIG. 12. Distribution of some typical marine lineages which invaded Amazonian waters. A. Achiridae. B. Belonidae. C. Engraulidae. D. Pristigasteridae. E. Tetraodontidae. Data from MZUSP. Map intended to represent general patterns of distribution into Amazonian and adjacent waters, not including marine records and from other basins. sensu Myers (1938) that invaded Amazonian waters, such as Achiridae (fig. 12A), Batrachoididae, Belonidae (fig. 12B), Clupeidae, Engraulidae (fig. 12C), Gobiidae, Hemirhamphidae, Pristigasteridae (fig. 12D), and Tetraodontidae (fig. 12E) (see Bloom and Lovejoy, 2017, for further details). Bloom and Lovejoy (2017) convincingly demonstrate that different marine lineages colonized South American rivers at different ages, influenced by separate events of marine transgression. According to those authors, different groups have biogeographical patterns consistent with invasions during the Oligocene, Eocene, or Miocene marine incursions. The Amazonian halfbeak is the only lineage younger than the Miocene to have invaded Amazonian freshwaters 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY less than a million years ago. These facts make it clear that the lowland Amazonian pattern of marine-derived lineages is pseudocongruent (sensu Donoghue and Moore, 2003). None of the marine-derived lineages is found in the highest parts of Guianese and Brazilian shields (see figs. 12A–E). Such fact suggests that the invasion of peripheral groups in the Amazon occurred subsequently to the establishment of the ichthyofauna in those upland regions. The alternative hypothesis that such lineages went extinct in the higher regions of the shields has no evidential support, either paleontological or geomorphological. Deep Channel Species Another lowland Amazonian pattern is demonstrated by species restricted to the deep channel of large Amazonian rivers, mainly the Amazonas itself (e.g., Sternarchella duccis, see Evans et al., 2017; Sternarchella rex, see Evans et al., 2017; Sternarchella sima, see Evans et al., 2017; Leptodoras juruensis, see Sabaj Pérez, 2005; Pariosternarchus amazonensis, see Albert and Crampton, 2006). Benthic regions of many of these large rivers contain specialized communities, mostly composed of electric fishes. Some of those species are strictly associated with deepwater environments, though there are records also in flooded beaches. Crampton (2007) listed at least 64 species of the Gymnotiformes that inhabit deep waters in the Amazon. Because deepwater Amazonian samples are still few and have been mostly focused on large Amazonian rivers, it is possible that this apparent pattern is the result of a sampling artifact of species actually belonging to some of the other South American lowlands patterns. Guiana Mangrove Province This pattern comprises the lower portions of the Orinoco basin, the Guiana coastal drainages and the lower Amazon (fig. 13A). Its western limit for most taxa is usually the mouth of the NO. 431 Orinoco, but in some cases it extends to the small independent coastal Venezuelan drainages, such as with Polycentrus schomburgkii (see Coutinho and Wosiacki, 2014). The eastern limit of this province is usually the mouth of the Amazon, with some species occurring also in the lower sectors of the Rio Jari, Tocantins, and Xingu and some reaching even further east to the Brazilian State of Maranhão. This pattern was first recognized by Myers (1960), in describing distribution patterns in the subfamily Aspredininae, of Aspredinidae: “They [the Aspredininae] are fishes of the lowland, muddy coast of Guiana and Amazonia, where they occur in the sea, in brackish water, and in the estuaries and tidal portions of rivers. They do not seem to be found far inland anywhere, except in the lower Amazon, where they apparently occur in many (or all) parts of the vast, complicated delta area, where the tides or tidal bores (pororoca) are felt” (Myers, 1960: 133). Myers (1960) describes the limits of the pattern as from the Orinoco delta in Venezuela into the Brazilian state Maranhão, and calls it the “Guyana Mangrove Province,” a name adopted here. The same pattern was later independently described by Vari (1988: 355) on the basis of data from Curimatidae: “The Atlantic slopes of Guyana, Surinam and French Guiana and Amapá in Brazil are another area of endemism....” Vari (1988) notes that some species in that area also occur in the lower Amazon and in the Rio Tocantins, such as Curimata cyprinoides. This pattern is strongly corroborated as a historical unit in the analyses of Dagosta and de Pinna (2017). Examples of Guiana Mangrove Province Pattern include: Anableps spp., Anablepsoides uroph thalmus group (see Nielsen, 2016), Aspredinichthys filamentosus (see Myers, 1960), Aspredinichthys tibicen (see Myers, 1960), Aspredo aspredo (see Myers, 1960), Copella arnoldi (see Marinho and Menezes, 2017), Curimata cyprinoides (see fig. 13B; Vari, 1989a), Cyphocharax helleri (fig. 13C), Hemigrammus rodway, Hemigrammus guyanensis, Nannostomus beckfordi, Piabucus dentatus, Platy stacus cotylephorus (see Myers, 1960), Poecilia 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 25 FIG. 13. Guiana Mangrove Province. A. Yellow area delimits the distribution pattern. B. Curimata cyprinoides (data from Vari, 1989a). C. Cyphocharax helleri (data from Vari, 1992b) with additional records from MZUSP). D. Polycentrus schomburgkii (see data from Coutinho and Wosiacki, 2014). parae, P. picta, Poptella longipinnis, Polycentrus schomburgkii (see fig. 13D; Coutinho and Wosiacki, 2014), Pristella maxillaris, Pseudauchenipterus nodosus, Rhinosardina amazonica (see Whitehead, 1985), and Tomeurus gracilis (see Myers, 1960). It seems unlikely that nonrheophilic species such as those listed above have the pattern of distribution they do as a result of sequential stream capture events or that they represent relictual distributions of ancient and more widely distributed populations. Instead, their distribution pattern seems to be strongly correlated with events of marine transgression and regression. An immense freshwater plume is formed by the discharge of the Amazon in the Atlantic (Goulding et al., 2003; Rocha, 2003). Such volume of freshwater floats above the heavier saltwater and spreads northwestward carried by the southern equatorial current (Jégu and Keith, 1999; Albert et al., 2006), resulting in turbid waters and largely unconsolidated substrates between the mouths of the Amazon and Orinoco (Curtin, 1986; Rocha, 2003). The effect of that plume varies seasonally and according to changes in sea level over time (Rocha, 2003). The first authors to propose the effect of the Amazonian plume on freshwater fish distribution were Jegú and Keith (1999). In their model, the plume serves as a corridor permitting the advance of species from the lower Amazon toward Guiana coastal drainages, thus explaining the common elements between those regions (see the distribution of Brachyhypopomus pinnicaudatus in Crampton et al., 2016, for another example). It is also likely that events of marine transgression and regression have altered the hydrogeological dynamics of the region affected by the plume, isolating or uniting different coastal drainages. 26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY It is not yet known whether the matching distributions of strictly freshwater (as listed above) and marine-tolerant species (e.g., Aspre dinichthys filamentosus, Aspredinichthys tibicen, Aspredo aspredo, Platystacus cotylephorus, Pseu dauchenipterus nodosus, Rhinosardina amazo nica, and Stictorhinus potamius) are congruent or pseudocongruent (sensu Donoghue and Moore, 2003). Myers (1960) notes that Tomeurus and Anableps have the same distribution pattern as Aspredininae, even though the two former taxa are not as tolerant to saltwater. The salt-tolerant species are restricted to lower portions of rivers, close to their mouths. Their tolerance to marine water may have allowed their range expansions without the need of transgression-regression events. Population phylogeographical studies may bring light to this question, by comparing divergence times between populations in the lower Amazon and those in the Guiana coastal drainages, both in strictly freshwater and marine-tolerant lineages. Regardless of salt-tolerance considerations, it is expected that the biogeographical pattern described above is at most ~11 Ma, i.e., as old as the age of the present connection between the Amazonas and the Atlantic (Hoorn, 1994a, 1994b, 1996; Potter, 1997) and also coinciding with the corresponding Andean uplift at that latitude (Hoorn et al., 1995). Eastern Amazon (East of the Purus Arch) Structural arches like the Purus Arch are basement structures located under sediments of different ages that are not exposed superficially in the eastern Amazon (Rossetti et al., 2005). As a consequence, such structures cannot have acted as biogeographical barriers from the end of the Miocene to the present (Campbell et al., 2006; Lima and Ribeiro, 2011). The Purus Arch is thought to have acted as a crucial barrier until the late Miocene, dividing the Eastern and Western Amazon (Figueiredo et al., 2009; Hoorn et al., 2017), although its role in the formation of the Amazon and its influence in the distribution NO. 431 of the biota remain controversial (see Wesselingh and Salo, 2006). Why and how two previously disconnected aquatic systems, the eastern and the western Amazon, merged is still unclear (Hoorn, 1994a, 1994b; Wesselingh, 2006; Figueiredo et al., 2009; Hoorn et al., 2017). Such uncertainty impedes proper understanding of the consequences of the event for the biogeography of Amazonian fishes. It is certain nonetheless that several lineages of fishes display distributions spatially congruent with a western/ eastern Amazon divide, with their limit coinciding exactly with the Purus Arch. The Eastern Amazon pattern may represent the distribution of species historically associated with the region lying east to the Purus Arch, for the most part cratonic and draining clear or black waters (Harris and Mix, 2002; Wesselingh and Hoorn, 2011). This pattern comprises the drainages of the Rio Negro, Orinoco, Essequibo, and Amazonian versants of the Brazilian and Guianan shields (fig. 14A). Some examples of this pattern are: Aphanotorulus emarginatus (see fig. 14B; Ray and Armbruster, 2016), Baryancis trus spp., Bivibranchia fowleri, Colomesus tocan tinensis (see Ruiz, 2015), Caquetaia spectabilis, Geophagus altifrons, Gnathodolus bidens, Hydro lycus tatauaia (see Toledo-Piza et al., 1999), Lep orinus brunneus (see Lima and Ribeiro, 2011), Pachyurus junki (fig. 14C) and Synaptolaemus latofasciatus (see fig. 14D; Britski et al., 2011). There are at least two examples of lineages distributed also in parts of the Atlantic drainages in the Guiana Shield, Brycon gr. pesu (see Planquette et al., 1996) and Hoplias curupira (see Oyakawa and Mattox, 2009). Water type and drainage relief are not enough to explain why the Rio Negro basin, for example, shares such a great number of lineages with highland drainages, despite the Negro´s small shield coverage. The existence of so many shared taxa exclusively between the Negro and shield rivers indicates a shared history. The East of the Purus Arch pattern may be directly related to the hydrogeological dynamics of the Amazon basin during the Miocene. 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 27 FIG. 14. Eastern Amazon. A. Yellow area delimits the distribution pattern (wavy line represents position of Purus Arch). B. Aphanothorolus emarginatus (data from Ray and Armbruster, 2016). C. Pachyurus junki (records from MZUSP). D. Synaptolaemus latofasciatus (data from Britski et al., 2011). The uplift of the central and north portions of the Andes created an overload on the South American plate that caused a lithospheric flexion, which in turn opened space for the formation of a sub-Andean Foreland basin (Sacek, 2014). Between ~24 and 16 Ma, this foreland basin received sediments from rivers draining west of the Purus Arch and east of the Andes, carrying them northward toward the Caribbean (Crampton, 2011; Sacek, 2014). At least since the Eocene, the Purus Arch (fig. 14A) was a divide between the basins draining east (eastern Amazon basin) and those draining westward (Lundberg et al., 1998; Costa et al., 2001; Crampton, 2011; Lujan and Armbruster, 2011). At that time, the eastern Amazon basin was formed by rivers draining shield areas, sediment poor and probably clear- or blackwater (Harris and Mix, 2002; Wesselingh and Hoorn, 2011). That phase was followed by the formation of an immense lacustrine system known as Pebas (~16 to 10.5 Ma), probably separated from the eastern Amazon system also by the Purus Arch (Figueiredo et al., 2009; Crampton, 2011; Sacek, 2014). The accumulation of sediments, mostly Andean in origin, in the foreland basin and the continuing Andean uplift (Crampton, 2011; Lima and Ribeiro, 2011) resulted in a breaching of the Purus Arch and a connection between that drainage and the eastern Amazon, forming a transcontinental basin and starting the Andean sedimentary deposition in the Brazilian equatorial margin, which extends to the present (Figueiredo et al., 2009; Sacek, 2014). Andean sediments in large amounts in the mouth of the Amazon begin approximately by 7 Ma, indicating that the west-east water divide 28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY was effective until that date (Hoorn, 1994b; Figueiredo et al., 2009; Crampton, 2011; for more recent estimates, see Roddaz et al., 2005; Rossetti et al., 2005; Campbell et al., 2006; Espurt et al., 2007). As a consequence, the pattern East of the Purus Arch pattern described herein is at least between 2.6 and 7 Ma, but in reality, it is probably far older because that time interval marks only the last instant before lineages south and north of the Amazonian tributaries were separated. A biogeographical pattern similar to the one described here was mentioned by both Eigenmann (1909) and Lima and Ribeiro (2011) under the highlands designation. However, for both authors relief is the decisive factor explaining the spatial distribution of fish species, rather than the past influence of the Purus Arch as proposed here. It is important to note that the Purus Arch today has no influence as a barrier on species distributions. Its role is relevant as a past barrier, when it formed the water divide between eastern and western Amazon. The reason why most lineages on each side of the divide do not expand their distributions is a mystery, perhaps related to historical-ecological factors independent of any present-day physical remains of the Purus Arch. The fish lineages east of the Purus arch are mostly ecologically restricted to fastflowing and sediment-poor tributaries, not entering the main Amazonian channel. The westward dispersion of those species was once limited by the Purus Arch when it was an effective barrier. The demise of the Purus Arch as a significant barrier is synchronous with the formation of the main channel of the Amazon. Slightly upstream of the mouth of the Rio Negro, approximately at the site of the ancient Purus Arch barrier, the main channel of the Amazon becomes significantly deeper (Geritana and Paiva, 2013). That factor, in combination with the simultaneous massive input of acidic waters from the Rio Negro, probably makes the region impervious to many taxa narrowly adapted to conditions of western Amazon waters. NO. 431 This region then started acting as an ecological barrier to those species west of the former Purus Arch, effectively replacing it. This provides an example that distributional patterns may have been determined by past barriers having no relation to current geographical boundaries, but nonetheless linked by a causal chain of different yet overlapping barriers. Of course, the number of fish species showing this pattern may seem small in view of the potential importance of the Purus Arch. However, the congruent distributions of unrelated lineages despite the absence of any apparent physical or ecological barriers cannot be ignored and may represent the last remnants of a common biogeographical history. Amazon-Core Uplands This pattern comprises species endemic to basins that drain the Brazilian and Guiana shields both in Atlantic and Amazonian versants. As mentioned above in South American Lowlands, Eigenmann (1909) was the first author to identify faunistic differences between South American lowlands and highlands. That author also inferred ages for those regions, implying that they have distinct biogeographical histories. Eigenmann (1909: 318) correctly proposes that both the Guianan and Brazilian shields are older than lowland regions: “The parts that first arose out of the sea and became populated with freshwater fishes were probably two land areas. The one embraces the highlands of Guiana and Northern Brazil, the other the highlands of Brazil east of the Araguay and south of the falls of the Tapajos.” The pattern described here is very similar to the one described by Eigenmann (1909) and differs from the Eastern Amazon pattern in excluding predominantly lowland drainages such as the Rio Negro and by including Guiana coastal basins (fig. 15A). Fishes displaying the Amazon-Core Highlands pattern are in general rheophilic: Acnodon spp., Anostomus ter netzi (see Lima and Ribeiro, 2011), Cetopsidium spp. (also in upper Rio Negro, see fig. 15B; Vari 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 29 FIG. 15. Amazon-core uplands. A. Yellow area delimits the distribution pattern. (B) Cetopsidium spp. (data from Vari et al., 2005, with additional records from MZUSP and LIRP). C. Hemigrammus ora (data from Jerep et al., 2011, with additional records from MZUSP). D. Hoplias aimara (data from Mattox et al., 2006). et al., 2005), Centromochlus schultzi, Hemibrycon surinamensis (see Bertaco and Malabarba, 2010), Hemigrammus ora (see fig. 15C; Jerep et al., 2011), Hoplias aimara (see fig. 15D; Mattox et al., 2006), Jupiaba essequibensis, J. gr. meunieri and J. polylepis, Krobia spp., Leporinus maculatus, Lepo rinus gr. granti, Moenkhausia grandisquamis, Mylesinus spp., Petulanos spp., Retroculus spp., Roeboexodon guianensis (see Lima and Ribeiro, 2011), and Tometes spp. (see Andrade, 2013; Andrade et al., 2016). The five categories described below are sub-patterns within the larger Highland Amazon Core pattern. Amazonian Uplands This pattern comprises exclusively Amazonian rivers draining both shields, Brazilian and Guianan (fig. 16A). Exact limits of this pattern are yet somewhat vague because known examples are species or lineages that occur in very narrow sectors of rivers, forming fragmented distributions based on sparse records. Species following this pattern are typically rheophilic and include: Archolaemus luciae (see Vari et al., 2012), Baryancistrus niveatus, Cetop sidium orientale (see Vari et al., 2005), Doras higu chii (see Sabaj Pérez et al., 2008), Hypomasticus julii, Leporinus britskii (see Feitosa et al., 2011), Leporinus microphysus (see Birindelli and Britski, 2013), Leporinus pachycheilus (also in Rio Araguari basin, see Santos et al., 1996), Metynnis anisurus (also in upper Rio Paraná basin, see Ota, 2015), Moenkhausia celibela (see Marinho and Langeani, 2010), Mylesinus schomburgkii, Sartor spp. (fig. 16B), and Teleocichla spp. (fig. 16C) and Tocantin sia piresi (fig. 16D). 30 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 431 FIG. 16. Amazonian uplands. A. Yellow area delimits the distribution pattern. B. Sartor spp. (records from MZUSP). C. Teleocichla spp. (records from MZUSP). D. Tocantinsia piresi (records from MZUSP). Guiana Shield (Atlantic and Amazonian Versants) This pattern includes lineages shared exclusively between Amazonian and Atlantic versants of rivers draining the Guiana Shield (fig. 17A). It possibly results from ichthyofaunistic exchange caused by stream capture events (see Cardoso and Montoya-Burgos, 2009). The pattern as a whole is probably not temporally congruent, but instead formed by independent events that caused faunistic mixing in the region, a common phenomenon between neighboring headwaters in shield rivers. Nijssen (1970) was the first author to propose that the headwater regions of north and south Guianan rivers might serve as a corridor for fish distribution. Subsequent authors, such as Cardoso and Montoya-Burgos (2009) and Lujan and Armbruster (2011), proposed additional examples of this pattern and its role as a faunistic connection between the Guianas and the Amazon. There are few examples of this pattern, in part as a result of the yet incipient knowledge on the fish fauna of upper reaches of Amazonian versants of the Guiana Shield. Some examples include: Corydoras baderi (Paru do Oeste and Maroni, see Nijssen and Isbrücker, 1980), Cteniloricaria spp. (Paru do Oeste, Maroni, Suriname, Corentyne, and Essequibo, see Covain et al., 2012), Hypomasticus megal epis (Trombetas, Uatumã and Guianese rivers, see Mol et al., 2012; J. Birindelli, personal commun.), Lithoxus spp. (Fisch-Muller, 2003), Microglanis secundus (Trombetas and Saramacca, see Ruiz and Shibatta, 2010), Par odon guyanensis (Paru do Oeste, Maroni, Suriname, C orentyne, and Essequibo), Parotocinclus halbothi (Trombetas and Maroni, 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 31 FIG. 17. Guiana Shield (Atlantic and Amazonian versants). A. Yellow area delimits the distribution pattern. B. Pseudancistrus brevispinis (data fom Cardoso and Montoya-Burgos, 2009). see Lehmann et al., 2014), Pseudancistrus brevispinis (Paru do Oeste, Jari and Guianese rivers, see fig. 17B; Cardoso and MontoyaBurgos, 2009), Stenolicmus ix (Curuá and Maroni, see Wosiacki et al., 2011; G. Dutra, personal commun.), and the clade Hypomasti cus despaxi + H. lineomaculatus (Paru, Jari, and Maroni, see Birindelli et al., 2013). Longitudinal Shield Correspondence among Amazonian Shield Versants This pattern is characterized by lineages that are present in both shields and follow a longitudinal correspondence among basins (fig. 18A). The pattern is expressed as lineages shared among the westernmost and easternmost parts of the cratonic region. In the western basins (Trombetas and Tapajós) examples include Sar tor gr. elongatus (fig. 18B), Bryconexodon spp. (fig. 18C), Laimosemion dibaphus (see Costa, 2006) and Hypoptopoma elongatum (see Aquino and Schaefer, 2010). In the eastern basins, as the Jari, Xingu, and Tocantins, some examples are Acnodon spp., Anablepsoides urophthalmus (see Costa, 2006), Bivibranchia velox (Fig 18D), Hypomasticus multimaculatus (see Birindelli et al., 2016), and Sternarchella sima (Ivanyisky III and Albert, 2014). The first author to recognize this pattern was Jégu (1992a), on the basis of some shared characiform taxa. Brazilian Shield This distribution pattern is defined by lineages occurring exclusively in the area corresponding to Amazon-draining Brazilian Shield rivers, formed by the Tocantins, Xingu, Tapajós basins, and some shield tributaries of the Rio Madeira (fig. 19A). Those are all highland rivers draining the ancient crystalline basement of the Brazilian Shield and most of them possess major rapids and/or waterfalls (Innocencio, 1989; Lima and Ribeiro, 2011). This pattern is recovered, in part, in the analyses of Dagosta and de Pinna (2017). Known examples of this pattern are typically rheophilic species. Probably the ecological conditions prevailing in lowland Amazonian environments act as barriers to their distributions (Géry, 1969). Géry (1962) proposes the circumferential pattern (lateral interbasin migration) for some species, suggesting that this pattern (encircling lowland South American lands, but never entering them) results from the ecological limitations of taxa restricted to fast-flowing rivers with high oxygen levels. This seems to be the explanation for the distributions of many Amazonian taxa restricted to the Brazilian Shield. 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 431 FIG. 18. Longitudinal correspondence among Amazonian Shield versants. A. Red area delimits the distribution pattern of western basins (Trombetas and Tapajós); yellow area the western basins pattern (Jari, Xingu, and Tocantins). B. Sartor gr. elongatus (records from MZUSP). C. Bryconexodon spp. (records from MZUSP). D. Bivibranchia velox (records from MZUSP). Géry (1962) used a dispersalist paradigm to explain lateral movements between basins. The author, however, actually adopted the notion of biotic dispersal (sensu Platnick and Nelson, 1978) rather than a true dispersalist framework. In that sense, his argument was essentially correct, because elements shared between neighboring basins have been associated with rearrangements of the hydrographic network (e.g., stream capture) resulting from neotectonic activity (see Lima and Ribeiro, 2011; Ribeiro et al., 2013). This interpretation of Géry´s hypothesis is clear in the following passage, where he proposes that the suppression of a barrier, even if momentarily, would have allowed the spread of a lineage: “Characids show a tendency to invade laterally their adjacent basins (by means of these temporary or permanent connections)” (Géry, 1962: 68). Another instance that demonstrates that the author did not follow pure dispersalism is: “The speciation (or subspeciation) occurred after the passage of the forms from one basin to another in ‘circumferential’ progression, rather than after having propagated along each great river” (Géry, 1962: 78). Examples of this pattern include: Acestroceph alus nigrofasciatus (Xingu, Juruena, and Jamanxim), Acestrocephalus stigmatus (Tocantins, Xingu, and Tapajós), Ancistrus ranunculus (Tocantins, Xingu, and Tapajós), Baryancistrus longipinnis (Tocantins, Xingu, and Tapajós), Bry conadenos tanaothoros (Xingu, Teles Pires, and Juruena), Caiapobrycon spp. (Tocantins, Xingu, and Tapajós, fig. 19B), Crenicichla acutirostris (Xingu, Tapajós, and shield portions of the Madeira; see Ploeg, 1991), Jupiaba apenima 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 33 FIG. 19. Brazilian Shield. A. Yellow area delimits the distribution pattern. B. Caiapobrycon spp. (records from MZUSP). C. Jupiaba apenima (records from MZUSP). D. Moenkhausia gr. pankilopteryx/pirauba (records from MZUSP). (Tocantins, Xingu, and Tapajós, fig. 19C), Jupi aba iasy (Xingu, Tapajós, and shield portions of the Madeira), Leporinus tristriatus (Tocantins, Xingu, and Tapajós; see Birindelli and Britski, 2013), Moenkhausia gr. pankilopteryx/pirauba (Tocantins, Xingu, Tapajós, and shield tributaries of the Rio Madeira, fig. 19D), Panaque arm brusteri (Tocantins, Xingu, and Tapajós; see Lujan et al., 2010), Petulanos intermedius (Xingu, Tapajós, and shield portions of the Madeira), Thayeria boehlkei (Tocantins, Xingu and, Tapajós; see Lima and Ribeiro, 2011), Rhinopetitia spp. (Tocantins, Xingu, and Tapajós) and Scobi nancistrus spp. (Tocantins, Xingu, and Tapajós). Some species occur both in the Amazonian sector of the Brazilian Shield and in the headwaters of the Rio Paraguay, a pattern discussed by Ribeiro et al. (2013) for Jupiaba acanthogaster. Additional examples include: Hyphessobrycon gr. vilmae, Moenkhausia gr. phaeonota, Moenkhau sia gr. lopesi, and the genus Utiaritichthys. Barring the unlikely possibility that all the taxa listed above became extinct in the Guiana Shield, then their age of diversification is maximally ~12–10 Ma. (Dobson et al., 2001; Figueiredo et al., 2009; Mora et al., 2010), when the Amazon river began depositing sediments on the Brazilian equatorial margin (Sacek, 2014), thus impeding rheophilic lineages from spreading their ranges to Guiana Shield regions. Extreme Shield: Chapada dos Parecis The Chapada dos Parecis is an elevated geomorphological formation located in the western portion of the Brazilian Shield, in central South 34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY America. It includes headwaters of various drainages, such as Rio Machado, Rio Guaporé, Paraguay, and mostly the Juruena. Many papers have proposed the Chapada dos Parecis as an area of endemism (Carvalho and Bertaco, 2006; Britski and Lima, 2007; Lima et al., 2007; Pastana and Dagosta, 2014; Ohara and Lima, 2015a). Yet, there are other noteworthy characteristics that must be noted for the ichthyofauna in that region. The portion of the Chapada dos Parecis drained (mostly) by the Juruena is the extreme case of the pattern expected for shield composition, with an extremely high level of endemism (Carvalho and Bertaco, 2006; Britski and Lima, 2007). Cases of closely related lineages coexisting are rare, diversity is low and there are very few taxa broadly distributed in the rest of the Amazon. East of the Rio Juruena in the Chapada dos Parecis, through the basins of the Rio Arinos, Rio Teles Pires, Rio Xingu, and Rio Tocantins-Araguaia, there is a trend toward reduction in endemism and an increase in the number of sympatric congeneric species, in species diversity, and in widely distributed species. The Rio Juruena, like other basins in that formation, contains no members of typically marine lineages (Myers’ 1938 peripheral division) (figs. 12A–E) and very few lowland Amazonian components. For example, Arapaima, Osteoglossum, Colossoma, and large pimelodids (Brachyplatystoma, see Barthem et al., 2017; Phractocephalus) are all absent. More relevant still is the fact that dozens of lineages present in other Brazilian Shield drainages are absent in the Rio Juruena at Chapada dos Parecis, such as Acestrocephalus acutus, Acestrorhynchus micrope lis (see González, 2015), Anostomoides passionis, Archolaemus luciae (see Vari et al., 2012), Argo nectes robertsi, Astyanax multidens (see Marinho and Birindelli, 2013), Bivibranchia notata, Bry conadenos tanaothoros (present only in Rio Arinos basin), Cyphocharax stilbolepis, Electrophorus electricus, Harttia dissidens, Hemigrammus levis, H. ora (see Jerep et al., 2011), Hyphessobrycon loweae + H. pegeouti clade (see Ingenito et al., 2013), H. moniliger, H. vilmae, H. pulchripinnis, Jupiaba apenima, J. anteroides, J. apenima, J. iasy, NO. 431 J. paranatinga, J. polylepis, Laetacara araguaiae, Leporinus britskii, L. julii, L. microphysus, Lepto doras oyakawai, Leptorhamdia schultzi, Macropso brycon xinguensis, Megadontognathus kaitukaensis, Moenkhausia celibela, M. collettii, Otocinclus hasemani, Panaque armbrusteri, Petu lanos intermedius, Pseudanos spp. (see Birindelli et al., 2012), Pimelodus tetramerus, Rhinopetitia spp., Roeboexodon guyanensis, Serrasalmus rhombeus, Sorubim trigonocephalus, Spectracan thicus murinus, Teleocichla spp. (fig. 16C), and Tocantinsia piresi (fig. 16D). Among all basins of the Amazonian versant of the Brazilian Shield, the portion of the Rio Juruena draining the Chapada dos Parecis has the most rapids and waterfalls. Britski and Lima (2007) suggest this factor as the reason for the high endemism in the region. We add that the same factor may serve as barriers in the opposite direction and explains also the absence of many lineages common in other Brazilian Shield basins. Thus, the abundance of rapids and waterfalls provides a threefold explanation: for the lack of specific lineages, for reduced sympatry and for decreased species richness. The Rio Iriri and Rio Teles Pires draining the Serra do Cachimbo and the upper Tocantins at the Chapada dos Veadeiros are two additional regions that can be classified as Extreme Shield. Both of them also drain the ancient crystalline basement of the Brazilian Shield and are dotted with rapids and waterfalls. Thus, they show pronounced faunal regionalization and are very poor in diversity when compared to other sectors of the Tapajós, Xingu e Tocantins basins. Exclusive Faunal Sharing between Neighboring Basins The sharing of exclusive faunal elements between two basins does not imply that such lineages are broadly distributed in both basins. This fact is evident in the eastern Amazon basins. Geographical distributions tend to be more restricted in highlands (see Albert and Crampton, 2005; Ribeiro, 2006; Maxime and 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Albert, 2009), where species have smaller ranges and most cases of broader distributions involve species or clades shared with neighboring basins. Such faunal similarities are in most instances associated with river captures caused by reactivation of faults or headward erosion (Ribeiro, 2006; Lima, 2017). Below we list and discuss stereotypical cases of exclusive faunal sharing between neighboring basins in the Amazon: Tapajós and Paraguay The fish fauna shared between Tapajós and Paraguay basins has been repeatedly recognized in the literature (see Lima et al., 2007; Carvalho and Albert, 2011a; Ribeiro et al., 2013) and some examples include: Aequidens rondoni (see Lima et al., 2007), Leporinus octomaculatus (fig. 20, see Birindelli and Britski, 2009), and Crenicichla ploegi. Additional examples are: Moenkhausia cosmops (also present in Guaporé basin, fig. 20), Moenkhausia gr. lopesi (also present in Rio Araguaia basin) and the genus Utiaritichthys (also present in shield tributaries of the Madeira). Tapajós and Xingu Taxa shared between the Tapajós and Xingu basins are: Anostomoides passionis, Archolaemus janeae (see Vari et al., 2012), Bryconadenos spp. (fig. 20, see Menezes et al., 2009), Cichla mir ianae (fig. 20, see Kullander and Ferreira, 2006), Creagrutus cracentis (see Dagosta and Pastana, 2014), Hopliancistrus spp., Hyphessobrycon cachimbensis (fig. 20), Hyphessobrycon cyanotae nia (fig. 20, see Dagosta et al., 2016; also in Guaporé basin), Leptodoras oyakawai (see Birindelli et al., 2008), Lebiasina melanoguttata (fig. 20), Leporinus villasboasorum (see Burns et al., 2017), Leptorhamdia schultzi, Megadontognathus kaitu kaensis (see Campos-da-Paz, 1999), Peckoltia feldbergae, Pyrrhulina marilynae (fig. 20, see Netto-Ferreira and Marinho, 2013), Retroculus xinguensis, Spatuloricaria tuira (see Fichberg et al., 2014), and Teleocichla prionogenys. 35 Tapajós and Madeira Different subdrainages that compose the Rio Madeira basin variably share exclusive ichthyofaunistic elements with the Rio Tapajós. Most shared elements are between the Aripuanã and the Juruena, such as Ancistrus parecis (see De Oliveira et al., 2016), Hemigrammus silimoni (fig. 20, see Dagosta, 2016), Inpaichthys spp. (fig. 20, see Dagosta, 2016), Moenkhausia levidorsa (fig. 20, see Dagosta, 2016), the clade Crenicichla chicha + C. hemera (see Varella et al., 2012), and genus Utiaritichthys (also present in upper Paraguay, fig. 20). There are also at least four examples of exclusive sharing between the Rio Juruena and the Rio Guaporé basin (Hyphessobrycon psittacus, fig. 20; Hyphessobrycon hexastichos, fig. 20; Moenkhausia rubra, fig. 20; Moenkhausia uirapuru, fig. 20) and three with the Rio Machado: Bryconops piracolina and Hyphessobrycon melanostichos (see Dagosta, 2016), and Corydoras hephaestus. Some taxa have wider distributions in the Tapajós and the Madeira, but are shared exclusively between the two basins (Steindachnerina fasciata, see NettoFerreira and Vari, 2011). Most importantly, virtually all cases listed above involve shield tributaries of the Madeira, and no known case of a species or clade that occurs in the main channel of the Madeira that is also shared exclusively with the Rio Tapajós basin. Recently, Tencatt and Ohara (2016) proposed a distribution pattern of Amazonian fishes delimited by interfluvial region between the Rio Madeira and the Rio Tapajós. Their arguments on fish species distributed in both systems are the same examples previously listed in Dagosta (2016) as evidence for the historical connections between the Tapajós and Madeira basins. However, Tencatt and Ohara claim the existence of congruence between the distribution of freshwater fishes and terrestrial organisms (birds, butterflies, primates and vascular plants) in the region between the Rio Madeira and the Rio Tapajós. However, freshwater fish distributions are limited by land tracts (save rare exceptions, e.g., Géry, 1964; 1969, for the Rio Amazonas and Goulding et al., 1988, for the Rio 36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 431 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Negro). The ichthyofaunal sharing between the Madeira and Tapajós results from a recent and localized history, influenced by geomorphological processes that resulted in stream capture events across the region that separates those basins and which caused biotic dispersal. Stream capture is a phenomenon entirely independent of the geographical isolation of terrestrial animals as inferred by Tencatt and Ohara. The rivers Tapajós and Madeira are the obvious barriers for the distribution of other terrestrial animals (e.g., birds, see Fernandes et al., 2014; Oppenheimer and Silveira, 2009). Species of fish are limited by waterfalls and land tracts. The patterns result from entirely different biogeographical phenomena and we believe there is no spatial or temporal homology between such apparent coincidences. 37 2015), Hemiancistrus spilomma, Hemiodus tocanti nensis, Hyphessobrycon loweae (see Ingenito et al., 2013), Hypostomus faveolus (see Zawadzki et al., 2008), Laemolyta fernandezi, Melanocharacidium auroradiatum, Mesonauta acora (see Kullander and Silfvergrip, 1991), Moenkhausia loweae (see Marinho, 2009), Moenkhausia pyrophthalma, Rhyn chodoras xingui (see Birindelli et al., 2007), Semaprochilodus brama (see Castro and Vari, 2004), Sternopygus xingu, and Tometes ancylorhyn chus (see Andrade et al., 2016). Tocantins and Paraguay Only Cyphocharax vanderi (see Claro-García and Shibatta, 2013), Hasemania hanseni, and Knodus chapadae (see Ferreira, 2007) are exclusively shared between these basins. Xingu and Paraguay There are few examples of species or clades shared exclusively between these basins: Hypopto poma inexspectatum (see Aquino and Schaefer, 2010), Steindachnerina brevipinna (see Netto-Ferreira and Vari, 2011), and the clade Characidium nupelia + C. xavante (see da Graça et al., 2008). Xingu and Tocantins Examples of this pattern here recognized are: Acnodon normani, Aspidoras poecilus (see Nijssen and Isbrücker, 1976), Astyanax argyrimarginatus, Bivibranchia velox (see Langeani, 1996), Centro mochlus simplex, Creagrutus britskii (see Meza-Vargas, 2015), Creagrutus mucipu (see Meza-Vargas, Tocantins and São Francisco Although examples of taxa shared exclusively between the Tocantins and São Francisco are few, they have received considerable attention in the literature (see Lima and Caires, 2011; Dagosta et al., 2014). The cases recognized here are: Cichla soma sanctifranciscense (see Lima and Caires, 2011; Dagosta et al., 2014), Hyphessobrycon dia statos (see Dagosta et al., 2014), and some lineages of Cynolebias and Hypsolebias (see Costa, 2010). The species Astyanax novae was previously considered as one more example of this pattern (Garutti and Venere, 2009; Lima and Caires, 2011; Dagosta et al., 2014), but in reality has a wider distribution (see Freitas et al., 2015). FIG. 20. Distribution of some lineages in Rio Tapajós basin and neighboring drainages. Dots are records in Rio Tapajós basin; stars are records in neighboring drainages. Data from MZUSP with additional records from literature. Each color represents a different lineage: light blue (Hemigrammus silimoni, see Dagosta et al., 2016); dark blue (Hyphessobrycon cyanotaenia, see Dagosta et al., 2016); light violet (Hyphessobrycon hexas tichos); dark violet (Hyphessobrycon cachimbensis); white (Hyphessobrycon melanostichos); light yellow (Hyphessobrycon psittacus, see Dagosta et al. 2016); dark yellow (Moenkhausia levidorsa, see Dagosta et al., 2016); light green (Bryconadenos tanaothoros); dark green (Inpaichthys spp., see Dagosta et al., 2016); red (Moenkhausia cosmops, see Ohara and Lima, 2015b); orange (Moenkhausia rubra); dark pink (Moenkhausia uirapuru, see Ohara and Lima, 2015b); light pink (Utiaritichthys spp); light brown (Leporinus octomaculatus, see Birindelli and Britski, 2009); dark brown (Pyrrhulina marylinae, see Netto-Ferreira and Marinho, 2013); black (Cichla mirianae, see Kullander and Ferreira, 2006); grey (Lebiasina melanoguttata). 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tocantins and Upper Paraná Some confirmed examples of lineages shared between those regions are: Characidium xan thopterum (see Silveira et al., 2008), Corumbataia spp. (see Britski, 1997; Carvalho, 2008), Hasema nia crenuchoides (see Serra and Langeani, 2015), and Rhinolekos spp. (see Martins and Langeani, 2011; Roxo et al., 2015). Additional species are shared exclusively between the Tocantins and upper Paraná plus the São Francisco: Brycon nat tereri (see Lima, 2017), Moenkhausia aurantia, Hyphessobrycon coelestinus (see Aquino and Carvalho, 2014), and Cetopsorhamdia iheringi. Madeira and Paraguay As discussed above in South American Lowlands, there are many events of biotic dispersal between the Amazon and the upper Paraguay, with all cases involving only part of the Rio Madeira basin. It is therefore not surprising that several taxa are shared between the Madeira and the Paraguay and that such congruent distributions are for the most part temporally decoupled, i.e., pseudocongruences (sensu Donoghue and Moore, 2003). Different subbasins of the Rio Madeira drainage share taxa exclusively with the Paraná-Paraguay, with most of such cases being from the Rio Guaporé and the Rio Mamoré. Many studies have discussed a common biogeographical history between those two regions (see Pearson, 1937; Hubert and Renno, 2006; Carvalho and Albert, 2011b; Ota et al., 2014). Some examples are: Aequidens plagiozonatus, Aphyoch arax anisitsi (see Souza-Lima, 2003), Apisto gramma trifasciata (see Kullander, 2003), Astyanacinus moorii, Astyanax lineatus, Cetopsis starnesi (see Vari et al., 2005), Corydoras polyst ictus, Gymnogeophagus balzanii (see Reis and Malabarba, 1988), Hemigrammus machadoi (see Ota et al., 2014), H. mahnerti (see Ota, 2010), H. tridens, Hyphessobrycon elachys, H. megalopterus (see Lima and Malabarba, 2003), Imparfinis gut tatus (see Queiroz et al., 2013), Laetacara dorsig era (see Linke and Staeck, 1994), Markiana NO. 431 nigripinnis, Megalonema platanum (see Queiroz et al., 2013), Odontostilbe paraguayensis (see Bührnheim, 2006), Oligosarcus pintoi (see Ribeiro and Menezes, 2015), Parodon carrikeri (see Schaefer, 2011), Piabucus melanostomus (see Britski et al., 1999; Queiroz et al., 2013), Pimelo della mucosa (see Queiroz et al., 2013), Psectro gaster curviventris (see Vari, 1989b), Rineloricaria aurata (see Vera-Alcaraz et al., 2012), Scoloplax empousa (see Schaefer et al., 1989), and Trachy doras paraguayensis (see Sabaj and Arce, 2017). Branco and Essequibo Those two basins have a common geomorphological history resulting from a series of capture events of the proto-Berbice by the Rio Branco drainage during the Pleistocene (Crawford et al., 1985; Gibbs and Barron, 1993; Souza et al., 2012). Such events may account for the conspicuous elements shared between the two basins (see Lujan and Armbruster, 2011; Souza et al., 2012). Some examples include: Apistogramma rupununi (see Kullander, 2003), Astyanax rupu nuni (see Souza et al., 2012), Cetopsidium roae (see Souza et al., 2012), Denticetopsis iwokrama (see Souza et al., 2012), Guianacara dacrya (Arbour and López-Fernández, 2011), Hyposto mus macushi (see Armbruster and Souza, 2005), Parodon bifasciatus (see Souza et al., 2012), Pseu dancistrus nigrescens (see Souza et al., 2012), Rhi nodoras armbrusteri (Sabaj Pérez et al., 2008), and Sturisoma monopelte (see Souza et al., 2012). Negro and Orinoco A number of contributions have explored the common biogeographical history of these two basins (see Winemiller et al., 2008; Willis et al., 2010; Winemiller and Willis, 2011). Examples of fish species shared exclusively between the Negro and the Orinoco are numerous and include: Acestridium dichromum (see Retzer et al., 1999), Acestridium martini (see Retzer et al., 1999), Creagrutus phasma (see Vari and Harold, 2001), Creagrutus runa, C. vexillapinnus and C. 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES zephyrus (see Vari and Harold, 2001), Geopha gus abalios and G. dicrozoster (see LópezFernández and Taphorn, 2004), Hemiancistrus subviridis (see Wernecke et al., 2005), Hemi grammus barrigonae, Hemigrammus bleheri (see Géry and Mahnert, 1986), Heterocharax lepto grammus (see Toledo-Piza, 2000b), Hoplarchus psittacus, Hyphessobrycon epicharis (see Weitzman and Palmer, 1997), Hypostomus sculpodon (see Armbruster, 2003), Laetacara fulvipinnis (see Staeck and Schindler, 2007), Leporinus enyae (see Burns et al., 2017), Micro characidium gnomus (see Buckup, 1993), Neblinichthys pilosus (see Ferraris et al., 1986), Odontostilbe pulchra (see Bührnheim and Malabarba, 2007), Phenacogaster prolatus (see Lucena and Malabarba, 2010), Prochilodus mar iae (see Castro and Vari, 2004), Pseudancistrus pectegenitor (see Lujan et al., 2007), Pseudancis trus sidereus (see Armbruster, 2004), Pseudanos varii (see Birindelli et al., 2012), Pseudolithoxus nicoi (see Lujan and Birindelli, 2011), Pterophyl lum altum (see Schultz, 1967), Ptychocharax rhyacophila (see Weitzman et al., 1994), Raceni sia fimbriipinna (see Mago-Leccia, 1994), Rhi nobrycon negrensis (see Lasso et al., 2004), Serrabrycon magoi (see Lasso et al., 2004), and Tometes makue (see Andrade, 2013). The Casiquiare Canal is a portion of the Rio Orinoco that was redirected to flow part of the year to the Rio Negro basin (Albert and Carvalho, 2011) and that now connects the two drainages by a permanent waterway with minimal gradient. Such a connection was mentioned by Eigenmann (1909) in his description of his Amazon Province in a dispersalist context and later proposed by Vari (1988) as the factor responsible for some curimatid species shared between the Amazon and Orinoco. Albert et al. (2006) and Winemiller et al. (2008) questioned the relevance of the physical Casiquiare connection as a species-dispersion route, because there are rapids on both sides of the divide (e.g., in Porto Ayacucho and in São Gabriel da Cachoeira) and possible chemical barriers (pH, temperature, and conductivity). It is possible 39 that part of the fish fauna shared between the Orinoco and Negro is in fact derived from the proto-Amazon-Orinoco and predates their hydrological separation. Negro and Branco The Rio Negro basin provides a clear example that hydrographic limits do not necessarily imply historical connections. Although the Rio Branco is the largest tributary of the Negro and the two are not separated by any physical barriers, each of them shares more species with adjoining nonAmazonian basins than with each other (with the Orinoco in the case of the Negro and with the Essequibo in the case of the Branco). In addition to the different geomorphological history of each basin, markedly different physicochemical parameters may also in part explain the small number of taxa exclusively shared between them. As pointed out by Ferreira et al. (2006), the Rio Negro predominantly drains lowland soils poor in cations with exceptionally low rates of mineral erosion, while the Rio Branco drains highland soils of an older landscape, richer in cations derived from the erosion of relatively stable igneous rocky beds. There are very few examples of species reliably restricted to the Negro and Branco: Physopyxis cristata (see Sousa and Py-Daniel, 2005), Apisto gramma gibbiceps (see Kullander, 1980), and Crenicichla virgatula (see Ito, 2013). Negro to Trombetas This pattern refers to the fish fauna common to the left-bank Amazonian tributaries east of the Rio Negro: Urubu, Uatumã, Nhamundá, and Trombetas. Together, they share some taxa exclusively with the Rio Negro or with the NegroOrinoco: Acestridium discus (Negro, Branco, and Trombetas), Ageneiosus polystictus (Negro, Urubu, and Trombetas; see Ribeiro et al., 2017), Asterophysus batrachus (Orinoco-Negro and Uatumã), Anduzedoras oxyrhynchus (OrinocoNegro, Branco, Urubu, and Trombetas), 40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Auchenipterichthys punctatus (Negro, Branco, and Urubu), Nemuroglanis pauciradiatus (Negro, Branco, Urubu, and Trombetas), Pygidianops amphioxus (Negro and Nhamundá; see de Pinna and Kirovsky, 2011), and Poecilocharax weitzmani (Orinoco-Negro, Branco, Urubu). Poorly Sampled Neighboring Basins in the Guiana Shield Some basins draining the Guiana Shield into the Amazon, such as the Urubu, Uatumã, Trombetas, and Paru, are relatively poorly known as to their ichthyofaunal composition, with comparatively few reported species, rare cases of endemism and few species shared among each other. Such precarious knowledge precludes a clear understanding of the connections of the fish faunas in those basins and few relevant examples deserve note: the Uatumã and Trombetas exclusively share Cetopsidium ferreirai (see Vari et al., 2005) and Cichla vazzoleri (see Kullander and Ferreira, 2006), while the Paru and Jari exclusively have Hypomasticus lineomaculatus (see Birindelli et al., 2013). Cis-Andean Foothills Another pattern related with the circumferential pattern of Géry (1962) is the cis-Andean Foothills distribution. The name refers to the highland region surrounding the cis-Andean lowlands, mostly around the Western Amazon (fig. 21A). As in the Brazilian Shield pattern, the present one comprises rheophilic species, restricted to fastflowing, highly oxygenated waters. This pattern was first identified by Vari (1988: 360): “Other species ranges appear to be associated with the more swiftly flowing piedmont streams of the western margins of the Amazon basin, and those species extend north into the western and northern margins of the Río Orinoco system.” Shortly thereafter, a similar pattern was described by Ibarra and Stewart (1989) for the Rio Napo, where the altitudinal gradient decisively influenced species com- NO. 431 position (see Lujan et al., 2013, for a more complex scenario). The pattern described here differs from the shield patterns in being not only wider, but also associated with rivers draining the eastern versant of the Andean range, and sometimes the western versant as well. Because the examples known are absent in the Amazonian versants of the Brazilian and Guiana shields, this pattern seems to be strictly associated with the history of the foreland basin and with the Andean uplift. Some of the known examples include species with both narrow and wide distributions. Examples in the former category include Acrobrycon ipanquianus (see Arcila et al., 2013), Attonitus (see Vari and Ortega, 2000), Creagrutus flaves cens, C. gephyrus, C. kunturus, and C. muelleri (see Vari and Harold, 2001). Cases of wide distributions in the Andean Foothills pattern comprise the family Astroblepidae (see Schaefer and Arroyave, 2010), Astyanacinus spp. (see fig. 21B; Dagosta, 2011), Ernstichthys spp. (see Stewart, 1985), Rhyacoglanis (see Shibatta and Vari, 2017), Xyliphius spp. (see Carvalho et al., 2017), Lepori nus striatus (see fig. 21C; Birindelli and Britski, 2013), Steindachnerina dobula (fig. 21D) and S. guentheri (see Vari, 1991), a clade composed of Creagrutus muelleri, C. ouranonastes, and C. peruanus (see Vari and Harold, 2001), and putative sister relationship between Brycon hilarii and B. whitei (see Lima, 2017). Another notable example is the entire genus Hemibrycon (excepting H. surinamensis, sole species in the genus with an Amazon-core Highlands pattern; see Bertaco and Malabarba, 2010). Lima and Ribeiro (2011) discuss a pattern similar to the one described here, in which lineages are restricted to upper portions of the foreland basin due to ecological requirements. As done here, those authors also distinguish this highland of the Foreland Basin pattern from that of the Brazilian Shield highland pattern. Wilkinson et al. (2010), in a discussion of the action of the megafans, also propose a pattern similar to the one advanced here, although not distinguishing shield highlands from the foreland-basin highlands. 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES 41 FIG. 21. Cis-Andean foothills. A. Yellow area delimits the distribution pattern. B. Astyanacinus spp. (data from Dagosta, 2011). C. Leporinus striatus (data from Birindelli and Britski, 2013). D. Steindachnerina dobula (data from Vari, 1991). Central Blackwater Amazon The name of this pattern refers to the most common (although by no means exclusive) water type of the rivers within its limits. Its position is approximately at the central portion of the Amazon (fig. 22A), although its western limits are not precisely defined. The distribution of most examples extends to the mouth of the Rio Negro, with some going farther, to the lower Japurá, lake Tefé, or into Peru. To the east, the pattern is almost always delimited by the mouth of the Rio Tapajós. Northward, most examples are restricted to the Negro/Orinoco, with some lineages found also in the Essequibo. To the south, species extend to the middle portion of the Tapajós, but may be more broadly distributed in the Rio Madeira, to tributaries of Mamoré/Guaporé. The first author to propose this pattern of distribution was Kullander (1986), in discussing congruent areas between species of cichlids and characids of the genus Paracheirodon (see Kullander, 1986: figs. 5, 6). Independently, Vari (1988: fig. 7) inferred that a then-undescribed species of Curimata had a pattern of distribution indicative of a preference for acidic waters, not exclusively in the Rio Negro basin, but also in other Central Amazonian localities. This biogeographical pattern also has surfaced occasionally in the literature, where it has been indicative of possible taxonomic problems. The first paper to notice something noteworthy in such distributions was Vari and Harold (2001), in the redescription of Creagrutus maxillaris. That species is broadly distributed in the Orinoco and the upper Rio Negro. The authors then had only a single lot with few poorly preserved 42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 431 FIG. 22. Central Blackwater Amazon. A. Yellow area delimits the distribution pattern. B. Biotoecus spp. (data fom Kullander, 1989). C. Dicrossus spp. (data fom Kullander, 2011). D. Hemigrammus analis (blue dots; records from MZUSP), Hemigrammus coeruleus (red dots; records from MZUSP), Hemigrammus stictus (yellow dots; records from MZUSP). specimens from the Rio Madeira (AMNH 39855) and stated that the presence of C. maxillaris in that basin required confirmation by additional material (later reported by Queiroz et al., 2013). A similar situation happened with Chalceus macrolepidotus in Zanata and Toledo-Piza (2004), whose sole sample from the Madeira basin was considered questionable because of its location widely disjunct from that of other known lots of the species. Likewise, Kullander and Ferreira (2006: 377) disregarded two samples of Cichla temensis from the Rio Madeira, not including them in the map or material examined of the species because “there is nearly no other Cichla material available from the Brazilian portion of the Madeira drainage to permit an understanding of the distribution of C. temensis in this region.” Clearly, in all examples the odd disjunct nature of such distributions influenced the respective authors´ hesitation about their own results. Our recognized pattern, however, shows that such distributions joining the Negro and Madeira are not at all abnormal. This pattern of distribution is recovered, in part, in the analyses of Dagosta and de Pinna (2017). Examples of lineages with a Central Blackwater Amazon pattern include: Aequidens mauesa nus (Madeira and Tapajós, see Kullander, 2003), Acestridium spp. (Orinoco/Negro, Madeira, and Tapajós), Astyanax ajuricaba (Negro, Tapajós, see Marinho and Lima, 2009), Boulengerella lucius (Orinoco/Negro, Tapajós, and Trombetas, see Vari, 1995), Brachyhypopomus hendersoni (Tefé, Negro, and Essequibo, see Crampton et al., 2016), Bryconops inpai (Orinoco/Negro, Tapajós, Madeira, and Trombetas), Chalceus spilogyros 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES (Madeira, Tapajós, and Trombetas, see Zanata and Toledo-Piza, 2004), C. macrolepidotus (Orinoco/Negro/Essequibo and Madeira, see Zanata and Toledo-Piza, 2004), Charax condei (Negro and Tapajós, see Menezes and Lucena, 2014), Cichla temensis (Orinoco/Negro and Madeira, see Kullander and Ferreira, 2006), clade Creagru tus maxillaris + C. cracentis (Orinoco/Negro, Madeira, and Tapajós, see Vari and Harold, 2001), Copella nattereri (Orinoco/Negro, Tapajós, Madeira, Trombetas, and some records in the Amazon above the mouth of Rio Negro, see Marinho and Menezes, 2017), Cynodon septena rius (Essequibo/Orinoco/Negro, Uatumã, Tapajós, Trombetas, and Tefé), Curimata incompta (Orinoco/Negro, Madeira, see Vari, 1988), Cyphocharax abramoides (Negro, Tapajós, and Trombetas, see Vari, 1992b), C. nigripinnis (Negro, Tapajós, and Amazonas, see Vari, 1992b), Elachocharax junki (Negro and Madeira, see Weitzman and Géry, 1981), Biotoecus spp. (Orinoco/Negro, Uatumã, and Trombetas, fig. 22B; see Kullander, 1989), Dicrossus spp. (Orinoco/ Negro, Madeira, Tapajós, and Trombetas, fig. 22C; see Kullander, 2011), Gnathocharax (Orinoco/Negro/Essequibo, Madeira, Tapajós, and Trombetas), Hemigrammus analis (Purus, Jutaí, Negro, Madeira, Tapajós, and Trombetas, fig. 22D), H. coeruleus (Orinoco/Negro/Essequibo, Madeira, Tapajós, and Trombetas, fig. 22D), H. hyanuary (Orinoco/Negro, Madeira, and Tapajós), H. stictus (Orinoco/Negro, Madeira, and Tapajós, fig. 22D), H. vorderwinkleri (Orinoco/ Negro, Madeira, Tapajós, and Trombetas), Het erocharax virgulatus (Orinoco/Negro, Madeira, and Tapajós, see Toledo-Piza, 2000b), Hoploch arax goethei (Orinoco/Negro, Madeira, Tapajós, and Trombetas), Hyphessobrycon sweglesi (lower Purus, Negro, and Madeira), Iguanodectes geisleri (Orinoco/Negro and Madeira), Jupiaba gr. atyp indi (Negro and Madeira), Leporinus altipinnis (Orinoco/Negro, Madeira, and Tapajós, see Britski and Birindelli, 2016), L. aripuanaensis (Branco, Madeira, and Trombetas), Leporinus gomesi (Madeira and Negro), L. klausewitzi (Negro and Madeira), Metynnis hypsauchen 43 (Orinoco/Negro/Essequibo, Madeira, Tapajós, and Trombetas, see Ota, 2015), M. melanogram mus (Orinoco/Negro, Uatumã, Trombetas, Tapajós, and Sucunduri (Madeira), see Ota et al., 2016), Moenkhausia hemigrammoides (Maroni, Suriname, Corentyne, Negro, Madeira, Tapajós, and Trombetas), M. lata (Orinoco/Negro, Madeira, and Tapajós, M. Marinho personal commun.), Nannostomus marilynae (Orinoco/ Negro and Madeira), Oxyropsis acutirostra (Orinoco/Negro and Tapajós), Poecilocharax spp. (Orinoco/Negro and Madeira), Pygidianops spp. (Orinoco/Negro and Madeira), Rhinobrycon neg rensis (Orinoco/Negro and Madeira), Satanoperca lilith (Negro, Uatumã, Trombetas, and Madeira, see Ota, 2013), Steindachnerina planiventris (Negro, Japurá, and Madeira, see Vari, 1991), Symphysodon discus (Negro, Madeira, and Trombetas, see Bleher et al., 2007; Farias and Hrbek, 2008; Amado et al., 2011), Taeniacara candidi (Negro, Tapajós, and Trombetas) and Hypoptopomatinae new genus (Negro, Madeira, and Tapajós, see Delapieve, 2014). Other potential examples are Moenkhausia diktyota (Madeira) and Hemigrammus pretoensis (Amazonas and Negro), which despite their current separate generic assignments are actually close relatives, perhaps even synonyms (F.C.P.D., personal obs.). The sharing of so many lineages clearly indicates strong historical connections among the Orinoco/Negro, Madeira and Tapajós. More importantly, all lineages with this distribution pattern are absent in the Brazilian Shield (except for some rare cases in the middle to lower Tapajós, at the periphery of the Shield, see fig. 22C). Despite such strong signal, no independent geomorphological history was identified that could explain this pattern. While Cretaceous deposits from those regions are well known, the Cenozoic sedimentary history is still very poorly known (Soares, 2007). The lack of such critical data does not allow a more precise evaluation of the biogeographically relevant processes and events in the region. It is clear that the lower sectors of those rivers (Negro, Purus, and Madeira), and even the portion of the Rio Amazonas in that 44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY region, underwent course changes during the Pleistocene as demonstrated by paleocanals (Latrubesse and Franzinelli, 2002; Almeida-Filho and Miranda, 2007; Irion et al., 2010; Teixeira and Soares, 2011; Hayakawa and Rossetti, 2015; but see Albert et al., 2018, for reservations about the accuracy of optically stimulated luminescence, or OSL, method for dating sediments of this type and age). However, details about the dynamics and timing of those events are unavailable at present. One exception is the work of Ruokolainen et al. (2018). The authors present evidence of river captures and avulsions during the late Pleistocene–Holocene in central Amazon, involving rivers Negro, Madeira, Purus and Juruá. Ruokolainen et al. demonstrate that the river network in the region has been anything but stable. According to them, during the past 50,000 years there have been many cases of river avulsions, with consequent changes in the historical connections among major tributary rivers of the central Amazon. The latest major river capture event converted the Japurá from a tributary of the Rio Negro to a tributary of the Amazon, only 1000 years ago. Such broad-scale lability implies that lowland rivers cannot have been efficient biogeographical dispersal barriers to terrestrial biota, and even less so for fishes. In such a scenario, river captures and avulsions in that region may have contributed, at least in part, to the origin of the pattern of distribution discussed here. As another relevant point, the Central Blackwater Amazon pattern follows remarkably closely the range of blackwater Amazonian rivers recently compiled by Venticinque et al. (2016) (see fig. 20A–D). Those authors demonstrate that there are numerous blackwater rivers scattered throughout the central Amazon, confirming Fink and Fink (1979: 18): “the Rio Negro is the major ‘black’ water river in Amazonia; however, similar conditions have a spotty distribution through much of central Amazonia and many igarapés and rios of the terra firma consist of this type of water.” Such a network provides ample opportunity for species NO. 431 restricted to blackwater to inhabit regions of the upper Amazon, approximately up to the mouth of the Rio Marañon in the Ucayali. A hypothesis that blackwater is the determining factor in the pattern herein described must be tested against a refinement of the species’ locality data. The small-scale mosaic physical distribution of blackwater tributaries in that region makes it very difficult to extract such information from usual museum data. For example, the Rio Madeira, although widely recognized as a whitewater river, is abundantly irrigated by tributaries of all water types (fig. 20). Therefore, the provenance of a sample from the Madeira says little about water type preferences unless associated with very precise locality information. Despite such limitations, it is remarkable that many of the species in the Central Blackwater Amazon pattern that occur in the Rio Madeira or Tapajós are restricted to the lower sectors of those basins, exactly where their blackwater tributaries are most abundant. Again, we highlight the fact that water type is not a random variable, but instead closely related to the geological history of the terrain it drains. Therefore, a distribution pattern determined by water type is also indirectly associated with a historical component and cannot be taken at face value as a purely ecological determinant. Allopatric Branco-Tocantins There are few examples of Amazonian fish species with disjunct distributions. Five unrelated species display an intriguing pattern of congruent disjunct distributions: Creagrutus menezesi (see Vari and Harold, 2001), Exodon paradoxus, Lepo racanthicus galaxias, Leporinus desmotes (see Burns et al., 2017), and Leptorhamdia essequiben sis (see Bockmann, 2003). Those species are found in the Branco and Tocantins basins, with some also having records in the Essequibo and Orinoco. All five species are well known in their taxonomy and geographical distribution, thus reducing the possibility of sampling gaps. The Branco and Tocantins basins are widely separated, making such allopatric disjunctions all 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES the more noteworthy, but no geomorphological evidence has been associated with such pattern (the explanations in Eigenmann´s Eastern Highlands [cf. also Albert et al., 2011: 50–52] do not account for the specific pattern discussed here, because in the present case the lineages involved are not present in the rest of the shield, i.e., the Tapajós, Xingu, Madeira, Trombetas, Jari, etc.). The geomorphological history of the Rio Branco is related to the proto-Berbice (Lujan and Armbruster, 2011), where the former had courses preferentially flowing from southwest to northeast toward the Caribbean Sea. Erosion of the rocky basement of the Guiana Shield caused the reorganization of the proto-Berbice drainage network and the southward reversal of its main course, making the Rio Branco a tributary of the Rio Negro (Schaefer and Dalrymple, 1996). The geomorphological history of the Rio Tocantins, in turn, is mostly associated with the geological evolution of the Brazilian Shield and with other large rivers such as the Tapajós, Xingu, Paraná-Paraguay, and São Francisco (Lima and Caires, 2011; Lima and Ribeiro, 2011). Of course, before 10 Ma there were no large whitewater rivers or floodplains separating clear-water tributaries of the Guiana and Brazilian shields, which might seem like a possible explanation. However, we again emphasize that the lineages constituting this pattern are not widely present in shield drainages, thus invalidating this broader paleoscenario as a causal factor. Of course, this scenario would hold in case the present disjunct pattern is a relict of a broader pattern that comprised other shield rivers, a hypothesis for which there is no evidence. The savannahs of the Rio Branco and Essequibo are biogeographically distinct from those of central Brazil, even though they share some fish species (Ferreira et al., 2006). As noted by López-Fernández and Albert (2011), the importance of savannas for the evolution of the modern fish fauna of the Neotropics cannot be overemphasized. In the absence of any geological evidence that might explain the exclusive sharing of species between the Branco and Tocantins, the presence of savannah systems may offer clues for a possible ecological explanation. 45 Absence Patterns Among the most curious distributional phenomena in the Amazon is the absence of some fish taxa in regions where they were expected to occur on the basis of the distribution of their close relatives and higher groups. Those absences are often associated with some clearly identifiable barriers of physical (e.g., waterfalls) or physico-ecological (e.g., water type) nature. The most conspicuous Absence pattern is seen in the upper Juruena river, a pattern described in detail in Chapada dos Parecis: Extreme Shield. Another remarkable example is the upper Rio Tocantins. Upriver from the region of Imperatriz (in the Brazilian state of Maranhão) and Itaguatins (in the Brazilian state of Tocantins), the channel of the Rio Tocantins has rapids in sectors that may help explain the absence of various groups otherwise distributed in the entire Amazon that are present in the Araguaia or in lower Tocantins basins. Some examples are: Acestrorhynchus falcirostris (see González, 2015), Apistogramma spp., Chaetobran chus spp., Hydrolycus tatauaia (see Toledo-Piza et al., 1999), Hypophthalmus marginatus, Hypselecara spp., Mastiglanis asopos, Megalechis thoracata (see Reis, 1997), Moenkhausia cotinho, Mylossoma spp. (see Mateussi, 2015), Ochmacanthus spp. (see Neto, 2014), Pellona spp. (see Melo, 2001), Potamorrha phis spp. (see Collette, 1982), and Semaprochilodus brama (see Castro and Vari, 2004). Other examples of biogeographically isolated Amazonian regions are the mid- and upper Rio Madeira, separated by the rapids in the region of Porto Velho, which block the upriver distribution of, for example, Arapaima. The absence of certain lineages is also influenced by other factors such as tidal effects. Goulding et al. (2003) showed that downstream from the region of Óbidos (in the Brazilian state of Amazonas), the tidal regime starts to influence the circadian rhythm of the Amazon, probably affecting the distribution of fish lineages (Jégu and Keith, 1999; Lima and Ribeiro, 2011), especially those with feeding and breeding periods narrowly associated with drought-flood cycles. Some examples of fishes that 46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY do not occur in the lower Amazon are: Acestrorhyn chus granducolis (see González, 2015), Brycon ama zonicus (see Lima, 2017), Colossoma macropomum (see Araujo-Lima and Goulding, 1997; Lima and Ribeiro, 2011), Copella nattereri (see Marinho and Menezes, 2017), Piaractus brachypomus (see Jégu and Keith, 1999), Potamorhina altamazonica (see Vari, 1984), and Serrasalmus elongatus (see Jégu and Keith, 1999). As seen above, water type has a major influence on biogeochemical processes and on the distribution and dynamics of aquatic habitats and associated biota (Venticique et al., 2016). Expectedly, it is an important factor in the geographical distribution of Amazonian fish lineages. As widely reported in the literature (see Sioli, 1984; Goulding et al., 2003), Amazonian rivers display enormous differences in pH and concentration of dissolved solutes, according to the type of soil they drain. Wallace (1889) was the first to note that water type influenced the composition of fish assemblages in the Amazon (Dagosta and de Pinna, 2018), an observation repeatedly confirmed in subsequent studies (see Roberts, 1972; Kullander, 1986; Goulding et al., 1988; Vari, 1988; Araujo-Lima and Goulding, 1997; Saint-Paul et al., 2000; Lima and Ribeiro, 2011). The extremely acidic water of the Rio Negro, in particular, may be a deterrent to many fish lineages. Some examples of fishes absent in the Negro, yet present in neighboring basins and widely distributed in the Amazon include: the subfamily Stethaprioninae (see Dagosta and Pinna, 2017; Reis, 1989), the genera Galeoch arax (see Giovannetti et al., 2017) and Hypopto poma (see Aquino and Schaefer, 2010), and several species such as Anostomus ternetzi (see Lima and Ribeiro, 2011), Brachyplatystoma juruense, Cheirocerus goeldii (see Stewart and Pavlik, 1985), Copella stigmasemion (see Marinho and Menezes, 2017), Curimatella dor salis (see Vari, 1992a), Hemiodus microlepis (see Langeani, 1996), Hemisorubim platyrhynchos, Jupiaba polylepis, Limatulichthys griseus (see Ohara, 2010), Megalodoras uranoscopus, Oxydo ras niger, Pimelodus blochii, Prochilodus nigri NO. 431 cans (see Castro and Vari, 2004), Pygocentrus nattereri, Semaprochilodus insignis (see Castro and Vari, 2004), and Tympanopleura atronasus (see Walsh et al., 2015). It is possible to go beyond the mere identification of absence biogeographical patterns. Our earlier biogeographic analyses have demonstrated that some absences are the result of extinctions rather than primitive absences (Dagosta and de Pinna, 2017); moreover, we found that the absence of several lineages in the Rio Negro are autapomorphic for the basin, i.e., their ancestral areas (historically related) have the respective taxa. Therefore, their absence in present-day Rio Negro may be the result of extinctions (discarding cases of pseudoabsences). Recently, Ruokolainen et al. (2018) provided convincing evidence that the Rio Japurá was a tributary to the lower Rio Negro and that a river capture event diverted it to flow into the Amazon (Solimões). The connection between the Rio Japurá and the Rio Negro may have been broken as recently as 1000 years ago. According to these authors, until that time the lower Rio Negro was not a blackwater river, as it presently is, and it carried a much larger load of sediments. Such evidence further corroborates the hypothesis of Dagosta and de Pinna (2017) that the lower Rio Negro basin was not always as hostile to some otherwise ubiquitous Amazonian lineages as it is today and may have had a less extreme type of water earlier in its history. At least for the lower part of its course, the Rio Negro did not have waters as acidic and nutrient poor as today, and did not impede the existence of some lineages that are now absent in the basin. Endemism Although a majority of 2716 species of Amazonian fishes examined here occur in more than one subdrainage, there are numerous examples of basin-specific endemics. With the regions defined by Dagosta and de Pinna (2017) as background, at least 831 Amazonian fish species are found in a single drainage or subregion 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES thereof (fig. 4, appendix 1). An additional 196 species are also basin specific, given a wider definition of basin (e.g., species restricted to the Tapajós but occurring in more than one subbasin therein). The latter data are also included in appendix 1, with indications of their respective basins of endemism. CONCLUSIONS Distribution patterns decay over time as new ones are superimposed (Grande, 1985; Hunn and Upchurch, 2001; Upchurch and Hunn, 2002; Upchurch et al., 2002; Morrone, 2009), making the disentanglement of their history a complex procedure. More studies on Amazonian fishes are necessary, both on phylogeny, paleontology, phylogeography, and molecular dating in order to empirically test the temporal congruences of the distributional patterns described here. New data on geological history are needed to better understand the effect of riverine configurations in the biogeography of fishes in the basin. Our findings support the conclusion that the biogeographical history of a river is associated less with its size than with its stability through geological time. The mosaic of patterns shown herein demonstrates that the river network in the Amazon has been anything but stable, and that this instability has been a major factor in fish distributions. Different overlapping geomorphological processes, at different times, have left diffuse marks on the composition and distribution of the fish fauna and this process continues to the present. The recent work by Stokes et al. (2018) directly demonstrates the intense dynamism of the region, showing that the Amazon river is capturing headwaters of the Río Orinoco, another step in the continuing reorganization of South American river systems. Freshwater fishes are physically restricted to hydrographical basins, but in the Amazon basin their distributions often transcend modern hydrographical limits. This is a result of a complex and reticulated history of drainages, a view 47 that has been corroborated by several authors (see Lima and Ribeiro, 2011; Dagosta, 2016; Dagosta and Pinna, 2017). This fact in itself does not disqualify basins as historical agents. Rather, rivers are historically bound areas, even though they are far more complex than hydrographically limited units. Data presented in this paper demonstrate that each hydrographic drainage in the Amazon basin participates simultaneously in various biogeographical patterns and that no single basin is a historically cohesive unit. Likewise, the entire Amazon basin itself does not form a single historical unit. All such conclusions corroborate the hypothesis that hydrographical basins should not be considered a priori as historical units. They are demonstrably reticulate areas that received portions of their biotas at different ages, under the influence of disjunct events. Thus, past geomorphological processes are more informative for understanding the distribution of the Amazonian fishes than present-day basin divides. ACKNOWLEDGMENTS This paper is dedicated to the memory of Richard P. Vari for his epochal contributions to the systematics and biogeography of Neotropical fishes and his generous devotion to the formation of new generations of ichthyologists, of which the authors of this paper are deeply grateful beneficiaries. We thank Cristiano Moreira and Marcelo Britto (MNRJ), Lúcia Py-Daniel and Renildo Oliveira (INPA); Cláudio Oliveira and Ricardo Benine (LBP); André Esguícero and Flávio Bockmann (LIRP); Izaura Maschio and Wolmar Wosiacki (MPEG); Jeff Clayton, Jeff Williams, Kris Murphy, Lynne Parenti, Richard Vari and Sandra Raredon (NMNH) for their help and hospitality during visits to the collections under their care. We are grateful to Eduardo Venticique for providing data on water types of Amazonian rivers for figures 7B–D and 20A–D. Gilberto N. Salvador and Eduardo Baena helped in the editing of the maps. José Birindelli provided photos of Hyphessobrycon psittacus, Lepo rinus octomaculatus, and Utiaritichthys sennaebr 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY agai. Authors were funded by FAPESP (FCPD, 2011/23419-1; 2016/07246-3; MP, 2015/26804-4), CNPq (MP, 201088/2014-2) and CAPES (MP, BEX 5840/2014-07). 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Symbols: *species exclusive to the Amazon basin; ** species poorly known or with vague distribution records. Taxon Species Carcharhiniformes 1 Carcharhinidae 1 Occurrence Lower Xingu, Amazonas main channel, lower Orinoco Carcharhinus leucas Pristiformes 2 Pristidae 2 Pristis pectinata Amazonas Estuary, Essequibo, lower Orinoco, Cauca-Magdalena-Sinu Pristis perotteti Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, Maracaibo, Cauca-Magdalena-Sinu, Atrato Myliobatiformes 27 Potamotrygonidae 27 Heliotrygon gomesi* Endemic – Amazonas main channel Heliotrygon rosai* Purus, Amazonas main channel, Amazonas Estuary Paratrygon aiereba Upper Tocantins, Araguaia, upper Xingu, lower Xingu, Teles Pires, Tapajós, Guaporé, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Plesiotrygon iwamae* Middle-lower Madeira, Napo-Ambyiacu, Amazonas main channel Plesiotrygon nana* Ucayali, Amazonas main channel Potamotrygon adamastor* Endemic – Branco Potamotrygon albimaculata* Restricted to Tapajós basin (Teles Pires, Tapajós) Potamotrygon amandae Mamoré, Guaporé, Paraná-Paraguay Potamotrygon amazona* Juruá, Jutaí, Negro Potamotrygon constellata** Amazonas main channel Potamotrygon falkneri Guaporé, Beni-Madre de Dios, Madeira Shield Tributaries, Trombetas, ParanáParaguay Potamotrygon garmani* Upper Tocantins, Teles Pires Potamotrygon henlei* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Potamotrygon humerosa Tapajós, middle-lower Madeira, Jutaí, Negro, Amazonas main channel, Capim Potamotrygon jabuti* Restricted to Tapajós basin (Teles Pires, Tapajós, Jamanxim) Potamotrygon leopoldi* Restricted to Xingu basin (upper Xingu, lower Xingu) Potamotrygon limai* Endemic – Middle-lower Madeira Potamotrygon motoro Upper Tocantins, Araguaia, lower Tocantins, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Ucayali, Marañon-Nanay, Putumayo, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure, ParanáParaguay Potamotrygon ocellata** 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 65 Occurrence Potamotrygon orbignyi Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Madeira Shield Tributaries, Purus, Tefé, Juruá, Ucayali, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Oiapoque, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Potamotrygon pantanensis Guaporé, Paraná-Paraguay Potamotrygon rex* Endemic – upper Tocantins Potamotrygon schroederi Japurá, Negro, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Potamotrygon scobina* Lower Tocantins, middle-lower Madeira, Madeira Shield Tributaries, UrubuUatumã, Amazonas main channel, Amazonas Estuary Potamotrygon tatianae* Beni-Madre de Dios, Madeira Shield Tributaries Potamotrygon tigrina* Endemic – Amazonas main channel Endemic – Negro Potamotrygon wallacei* Osteoglossiformes 5 Arapaimidae 3 Arapaima agassizii** Arapaima gigas Araguaia, lower Tocantins, lower Xingu, middle-lower Madeira, Purus, Juruá, Putumayo, Japurá, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Essequibo Arapaima leptosoma* Amazonas main channel Osteoglossidae 2 Araguaia, lower Tocantins, lower Xingu, Teles Pires, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Corentyne-Demerara, Essequibo, upper Orinoco Osteoglossum bicirrhosum Branco, upper Orinoco Osteoglossum ferreirai Anguiliformes 1 Ophichthidae 1 Lower Tocantins, lower Orinoco Stictorhinus potamius Clupeifomes 18 Clupeidae 1 Amazonas main channel, Coppename-Suriname-Saramacca, Corentyne-Demerara, lower Orinoco Rhinosardinia amazonica Engraulidae 12 Amazonsprattus scintilla* Lower Xingu, Teles Pires, Tapajós, Madeira Shield Tributaries, Purus, Negro, Branco, Urubu-Uatumã Anchovia surinamensis Lower Tocantins, upper Xingu, lower Xingu, Teles Pires, middle-lower Madeira, Branco, Parnaíba, Itapicuru-Mearim, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco Anchoviella alleni* Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, CoariUrucu, Ucayali Anchoviella carrikeri Araguaia, upper Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Tefé, Ucayali, Marañon-Nanay, Negro, Branco, Amazonas main channel, Capim 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Anchoviella guianensis Araguaia, Anapu, Teles Pires, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Parnaíba, Capim, Maroni-Approuague, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Anchoviella hernanni* Beni-Madre de Dios, Ucayali, Marañon-Nanay Anchoviella jamesi Araguaia, lower Tocantins, Teles Pires, Tapajós, Jamanxim, middle-lower Madeira, Tefé, Negro, Branco, Trombetas, Amazonas main channel, Capim, Essequibo, lower Orinoco, upper Orinoco Anchoviella juruasanga* Lower Tocantins, Tapajós, Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Purus, Negro, Trombetas, Jari, Amazonas main channel Anchoviella nattereri** Jurengraulis juruensis Mamoré, middle-lower Madeira, Juruá, Putumayo, Branco, Trombetas, Amazonas main channel, Essequibo Lycengraulis batesii Araguaia, lower Tocantins, lower Xingu, Tapajós, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Japurá, Branco, Trombetas, Amazonas main channel, Parnaíba, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, lower Orinoco, upper Orinoco, Apure Pterengraulis atherinoides* Lower Tocantins, lower Xingu, Trombetas, Parnaíba, Itapicuru-Mearim, CorentyneDemerara, lower Orinoco, upper Orinoco, Apure Pristigasteridae 5 Ilisha amazonica* Upper Xingu, Iriri, lower Xingu, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Urubu-Uatumã, Trombetas, Amazonas main channel Pellona castelnaeana Araguaia, lower Tocantins, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, lower Orinoco, upper Orinoco Pellona flavipinnis Lower Tocantins, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Coari-Urucu, Tefé, Putumayo, Japurá, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Parnaíba, Coppename-Suriname-Saramacca, Corentyne-Demerara, lower Orinoco, upper Orinoco, Paraná-Paraguay Pristigaster cayana* Upper Tocantins, Araguaia, lower Tocantins, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel Pristigaster whiteheadi Middle-lower Madeira, Purus, Juruá, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Araguari-Macari-Amapá Characiformes Acestrorhynchidae 1063 10 Acestrorhynchus abbreviatus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Javari, Ucayali, Napo-Ambyiacu, Amazonas main channel, Amazonas Estuary, Araguari-Macari-Amapá, Paraná-Paraguay Acestrorhynchus falcatus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, NapoAmbyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, Capim, Araguari-Macari-Amapá, Oiapoque, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 67 Occurrence Acestrorhynchus falcirostris Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Oiapoque, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Acestrorhynchus grandoculis Teles Pires, middle-lower Madeira, Madeira Shield Tributaries, Napo-Ambyiacu, Putumayo, Negro, Branco, Urubu-Uatumã, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Acestrorhynchus heterolepis Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Amazonas main channel, Itapicuru-Mearim, Essequibo, lower Orinoco, upper Orinoco, Apure Acestrorhynchus isalineae* Middle-lower Madeira, Madeira Shield Tributaries Acestrorhynchus maculipinna* Negro, Amazonas main channel Acestrorhynchus microlepis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Coari-Urucu, Tefé, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, AraguariMacari-Amapá, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Acestrorhynchus minimus Upper Xingu, Teles Pires, Tapajós, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure Acestrorhynchus nasutus Teles Pires, Tapajós, middle-lower Madeira, Negro, Branco, Amazonas main channel, Capim, Essequibo, upper Orinoco Alestidae 5 Chalceus epakros Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Chalceus erythrurus* Middle-lower Madeira, Purus, Coari-Urucu, Juruá, Ucayali, Putumayo, Japurá, Negro, Amazonas main channel Chalceus guaporensis* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Amazonas main channel Chalceus macrolepidotus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Japurá, Negro, Urubu-Uatumã, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Tapajós, middle-lower Madeira, Trombetas Chalceus spilogyros* Anostomidae 91 Abramites hypselonotus Upper Tocantins, Araguaia, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Putumayo, Japurá, Amazonas main channel, Amazonas Estuary, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Anostomoides laticeps Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Essequibo, lower Orinoco Anostomoides passionis* Lower Xingu, Teles Pires 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Anostomus anostomus* Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Amazonas main channel, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Anostomus longus** Marañon-Nanay Anostomus ternetzi* Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Madeira Shield Tributaries, Japurá, Branco, Urubu-Uatumã, Trombetas, Capim, Maroni-Approuague, Essequibo, lower Orinoco, upper Orinoco, Apure Gnathodolus bidens Lower Xingu, Teles Pires, Urubu-Uatumã, lower Orinoco, upper Orinoco Hypomasticus julii* Araguaia, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Jamanxim, Trombetas, Jari Hypomasticus lineomaculatus* Paru, Jari Hypomasticus megalepis Upper Tocantins, lower Tocantins, upper Xingu, Iriri, lower Xingu, Branco, Jari, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo Hypomasticus multimaculatus Upper Tocantins, Araguaia, upper Xingu, lower Xingu, Jari, Araguari-MacariAmapá Hypomasticus pachycheilus Upper Tocantins, lower Tocantins, Jamanxim, Madeira Shield Tributaries, UrubuUatumã, Trombetas, Araguari-Macari-Amapá Hypomasticus torrenticola* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Laemolyta fasciata** Endemic – Mamoré Laemolyta fernandezi Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, lower Orinoco, upper Orinoco Laemolyta garmani Araguaia, Guaporé, Madeira Shield Tributaries, Tefé, Jutaí, Javari, Ucayali, NapoAmbyiacu, Putumayo, Japurá, Amazonas main channel, Capim Laemolyta proxima Lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, NapoAmbyiacu, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Corentyne-Demerara, Essequibo Laemolyta taeniata Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure Leporellus vittatus* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Branco, Jari, Essequibo, lower Orinoco, upper Orinoco, Apure, Cauca-MagdalenaSinu, Paraná-Paraguay Leporinus acutidens** Leporinus affinis Araguaia, lower Tocantins, lower Xingu, Japurá, Capim, lower Orinoco Leporinus agassizii Mamoré, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, UrubuUatumã, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, Upper Orinoco Leporinus altipinnis Tapajós, middle-lower Madeira, Negro, Amazonas main channel, upper Orinoco Leporinus amazonicus* Mamoré, Guaporé, middle-lower Madeira, Juruá, Japurá, Amazonas main channel Leporinus arimaspi Branco, lower Orinoco, upper Orinoco Leporinus aripuanaensis* Madeira Shield Tributaries, Branco, Urubu-Uatumã, Trombetas Leporinus bimaculatus* Upper Tocantins, Amazonas main channel Leporinus bistriatus* Restricted to Tocantins basin (upper Tocantins, lower Tocantins) 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 69 Occurrence Leporinus bleheri* Guaporé, Negro, Amazonas Estuary Leporinus britskii* Teles Pires, Jamanxim, Jari Leporinus brunneus Upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Madeira Shield Tributaries, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, lower Orinoco, upper Orinoco Leporinus cylindriformis* Lower Xingu, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel Leporinus desmotes Upper Tocantins, lower Tocantins, Branco, Essequibo Leporinus enyae Negro, lower Orinoco, upper Orinoco Leporinus fasciatus Upper Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Ucayali, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure Leporinus friderici Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Putumayo, Japurá, Negro, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, ParanáParaguay Leporinus geminis* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Leporinus gomesi* Madeira Shield Tributaries, Negro Leporinus granti Upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Madeira Shield Tributaries, Branco, Trombetas, Oiapoque, Maroni-Approuague, CorentyneDemerara, Essequibo Leporinus guttatus* Endemic – Iriri Leporinus jamesi* Mamoré, middle-lower Madeira, Purus, Ucayali, Amazonas main channel Leporinus jatuncochi* Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Marañon-Nanay, Napo-Ambyiacu, Urubu-Uatumã, Jari, Amazonas main channel Leporinus klausewitzi* Middle-lower Madeira, Japurá, Negro, Branco, Urubu-Uatumã, Amazonas main channel Leporinus maculatus Lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Branco, Trombetas, Jari, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Leporinus melanostictus Jari, Oiapoque, Maroni-Approuague Leporinus microphysus* Teles Pires, Jamanxim, Jari Leporinus moralesi Putumayo, Japurá, Amazonas main channel, lower Orinoco Leporinus multifasciatus** Leporinus nattereri Araguaia, middle-lower Madeira, Putumayo, Trombetas, Amazonas main channel, upper Orinoco, Apure Leporinus niceforoi Japurá, upper Orinoco Leporinus nigrotaeniatus Negro, Branco, Corentyne-Demerara, Essequibo Leporinus octomaculatus Teles Pires, Juruena, Tapajós, Paraná-Paraguay Leporinus ortomaculatus Upper Tocantins, Araguaia, Branco, Essequibo, lower Orinoco, upper Orinoco Leporinus pachyurus** 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Leporinus parae Upper Tocantins, Araguaia, lower Tocantins, lower Xingu, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Amazonas main channel, Amazonas Estuary, lower Orinoco, upper Orinoco Leporinus parvulus* Restricted to Tapajós basin (Teles Pires, Tapajós) Leporinus pearsoni* Mamoré, Guaporé, Beni-Madre de Dios, Madeira Shield Tributaries, Purus, Juruá, Ucayali Leporinus pellegrinii Upper Xingu, Branco, Trombetas, Maroni-Approuague, Essequibo Leporinus pitingai** Urubu-Uatumã Leporinus reticulatus* Endemic – Juruena Leporinus santosi* Endemic – lower Tocantins Leporinus sexstriatus* Endemic – Juruena Leporinus steyermarki Putumayo, lower Orinoco, upper Orinoco Leporinus striatus Mamoré, Guaporé, Beni-Madre de Dios, Juruá, Marañon-Nanay, Napo-Ambyiacu, Putumayo, upper Orinoco, Apure, Cauca-Magdalena-Sinu, Atrato, Paraná-Paraguay Leporinus subniger* Japurá, Amazonas main channel Leporinus taeniofasciatus* Endemic – upper Tocantins Leporinus tigrinus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Paraná-Paraguay Leporinus trimaculatus* Endemic – Madeira Shield Tributaries Leporinus tristriatus* Upper Tocantins, upper Xingu, Teles Pires, Juruena, Jamanxim Leporinus unitaeniatus* Upper Tocantins, Araguaia, lower Tocantins, Tapajós Leporinus vanzoi* Restricted to Tapajós basin (Teles Pires, Juruena, Tapajós) Leporinus venerei* Endemic – Araguaia Leporinus villasboasorum* Upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena Leporinus yophorus Mamoré, Guaporé, Beni-Madre de Dios, Purus, Juruá, lower Orinoco, upper Orinoco Megaleporinus trifasciatus* Upper Tocantins, Araguaia, lower Tocantins, Tapajós, Mamoré, Guaporé, BeniMadre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Ucayali, Putumayo, Japurá, Negro, Trombetas, Amazonas main channel Petulanos intermedius* Upper Xingu, lower Xingu, Teles Pires, Jamanxim, Madeira Shield Tributaries Petulanos plicatus Mamoré, Guaporé, Branco, Trombetas, Corentyne-Demerara, Essequibo Pseudanos gracilis Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Pseudanos trimaculatus Lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Coari-Urucu, Juruá, Napo-Ambyiacu, Putumayo, Negro, Branco, Urubu-Uatumã, Jari, Amazonas main channel, Amazonas Estuary, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure Pseudanos varii Negro, upper Orinoco Pseudanos winterbottomi Juruena, Tapajós, Jamanxim, lower Orinoco, upper Orinoco Rhytiodus argenteofuscus* Lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Juruá, Putumayo, Branco, Trombetas, Amazonas main channel Rhytiodus elongatus** Purus Rhytiodus lauzannei* Restricted to Madeira basin (Mamoré, Guaporé, middle-lower Madeira) 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 71 Occurrence Rhytiodus microlepis* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Japurá, Amazonas main channel Sartor elongatus* Teles Pires, Juruena, Trombetas Sartor respectus* Endemic – upper Xingu Sartor tucuriense* Endemic – lower Tocantins Schizodon fasciatus Lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Javari, Ucayali, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Jari, Amazonas main channel, Amazonas Estuary, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, Maracaibo Schizodon vittatus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Essequibo Synaptolaemus latofasciatus Upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Madeira Shield Tributaries, Branco, Urubu-Uatumã, Trombetas, Essequibo, lower Orinoco, upper Orinoco, Apure Characidae 635 Acestrocephalus acutus* Upper Tocantins, Araguaia, lower Tocantins, Teles Pires Acestrocephalus boehlkei* Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo Acestrocephalus maculosus* Endemic – upper Tocantins Acestrocephalus nigrifasciatus* Iriri, Teles Pires, Juruena, Jamanxim Acestrocephalus pallidus* Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries Acestrocephalus sardina Madeira Shield Tributaries, Jutaí, Negro, Branco, Trombetas, Amazonas main channel, upper Orinoco Acestrocephalus stigmatus* Upper Tocantins, Araguaia, upper Xingu, lower Xingu, Teles Pires Acrobrycon ipanquianus UcayaliParaná-Paraguay Acrobrycon starnesi* Endemic – Guaporé Agoniates anchovia Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Japurá, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Agoniates halecinus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Capim, Araguari-Macari-Amapá, Essequibo, upper Orinoco Amazonspinther dalmata* Middle-lower Madeira, Purus Aphyocharacidium bolivianum* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Aphyocharacidium melandetum Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo Aphyocharax agassizii** Amazonas main channel Aphyocharax anisitsi Mamoré, Guaporé, Beni-Madre de Dios, Paraná-Paraguay Aphyocharax avary** Middle-lower Madeira Aphyocharax erythrurus Branco, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Maracaibo Aphyocharax nattereri Mamoré, Guaporé, Beni-Madre de Dios, Amazonas main channel, Paraná-Paraguay Aphyocharax pusillus* Upper Tocantins, Araguaia, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Coari-Urucu, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure 72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Aphyocharax rathbuni Mamoré, Guaporé, Paraná-Paraguay Aphyodite apiaka* Endemic – Madeira Shield Tributaries Aphyodite grammica Mamoré, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Branco, Amazonas main channel, Essequibo Aphyodite tupebas* Tefé, Amazonas main channel Astyanacinus boliviensis* Mamoré, Beni-Madre de Dios Astyanacinus gandhiae* Ucayali, Marañon-Nanay Astyanacinus multidens* Mamoré, Guaporé, Beni-Madre de Dios, Ucayali Astyanacinus villwocki* Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá Astyanax abramis Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Juruá, Ucayali, Putumayo, Japurá, Amazonas main channel, lower Orinoco, upper Orinoco Astyanax ajuricaba* Araguaia, Tapajós, Negro, Amazonas main channel Astyanax anterior* Upper Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, middlelower Madeira, Madeira Shield Tributaries, Urubu-Uatumã, Trombetas, Amazonas main channel Astyanax argyrimarginatus* Araguaia, upper Xingu Astyanax bimaculatus Upper Tocantins, Araguaia, upper Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Putumayo, Japurá, Branco, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, Oiapoque, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Cauca-Magdalena-Sinu, Atrato Astyanax bourgeti* Endemic – Amazonas main channel Astyanax chaparae* Endemic – Mamoré Astyanax clavitaeniatus* Endemic – Branco Astyanax courensis* Endemic – upper Tocantins Astyanax elachylepis* Upper Tocantins, Araguaia, lower Tocantins, Teles Pires Astyanax fasciatus* Mamoré, Guaporé, Beni-Madre de Dios, Ucayali, Putumayo, Branco, Parnaíba, Essequibo, lower Orinoco, upper Orinoco, Cauca-Magdalena-Sinu, Atrato, ParanáParaguay Astyanax goyacensis Araguaia, Paraná-Paraguay Astyanax goyanensis* Endemic – upper Tocantins Astyanax gracilior** Amazonas main channel Astyanax guaporensis* Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries Astyanax guianensis Upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, BeniMadre de Dios, middle-lower Madeira, Negro, Branco, Jari, Capim, Essequibo Astyanax integer Napo-Ambyiacu, Japurá, lower Orinoco, upper Orinoco, Apure, Maracaibo Astyanax joaovitori* Endemic – Araguaia Astyanax kullanderi* Endemic – Araguaia Astyanax lineatus Mamoré, Guaporé, Paraná-Paraguay Astyanax longior* Endemic – Marañon Astyanax maculisquamis* Endemic – Guaporé Astyanax maximus Beni-Madre de Dios, Purus, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Japurá, upper Orinoco 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 73 Occurrence Astyanax multidens* Araguaia, upper Xingu, lower Xingu, Teles Pires, Jamanxim, Amazonas main channel Astyanax novae* Upper Tocantins, Araguaia, upper Xingu Astyanax poetzschkei** Astyanax rupununi Branco, Essequibo Astyanax saltor* Iriri, Teles Pires Astyanax siapae Negro, upper Orinoco, Apure Astyanax symmetricus* Endemic – Amazonas main channel Astyanax unitaeniatus* Endemic – upper Tocantins Astyanax utiariti* Endemic – Juruena Astyanax validus Japurá, Oiapoque, Maroni-Approuague Astyanax xavante* Restricted to Tocantins basin (upper Tocantins, Araguaia) Atopomesus pachyodus* Endemic – Negro Attonitus bounites* Endemic – Beni-Madre de Dios Attonitus ephimeros* Endemic – Ucayali Attonitus irisae* Endemic – Ucayali Axelrodia lindeae* Teles Pires, middle-lower Madeira, Negro, Branco, Amazonas main channel Axelrodia stigmatias* Middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Amazonas main channel Bario steindachneri Tapajós, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Putumayo, Amazonas main channel, Capim Boehlkea fredcochui* Japurá, Amazonas main channel Boehlkea orcesi* Endemic – Marañon Boehlkea weitzmani* Endemic – Japurá Brachychalcinus copei* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Trombetas, Amazonas main channel Brachychalcinus nummus* Mamoré, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá Brachychalcinus parnaibae Lower Tocantins, Parnaíba Brachychalcinus reisi* Endemic – Iriri Brittanichthys axelrodi* Restricted to Negro basin (Negro, Branco) Brittanichthys myersi Negro, Branco, Corentyne-Demerara, Essequibo Brycon amazonicus Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Brycon atrocaudatus** Endemic – Marañon Brycon coxeyi* Endemic – Marañon Brycon falcatus* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Guaporé, Madeira Shield Tributaries, Negro, Branco, Amazonas main channel, Capim, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Brycon gouldingi* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) 74 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Brycon hilarii Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Paraná-Paraguay Brycon melanopterus* Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel Brycon nattereri Upper Tocantins, Parana-Paraguay, São Francisco Brycon pesu* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Guaporé, Madeira Shield Tributaries, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Maroni-Approuague, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Brycon polylepis Upper Tocantins, lower Tocantins, Beni-Madre de Dios, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Amazonas main channel, upper Orinoco, Maracaibo Brycon stolzmanni* Endemic – Marañon Bryconacidnus ellisi* Mamoré, Guaporé, Beni-Madre de Dios, Ucayali, Napo-Ambyiacu Bryconacidnus hemigrammus* Endemic – Beni-Madre de Dios Bryconacidnus paipayensis* Endemic – Marañon Bryconadenos tanaothoros* Upper Xingu, Teles Pires, Juruena Bryconadenos weitzmani* Endemic – Iriri Bryconamericus alfredae* Beni-Madre de Dios, Ucayali Bryconamericus bolivianus* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Bryconamericus carlosi* Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá Bryconamericus deuterodonoides Japurá, lower Orinoco, upper Orinoco, Apure, Maracaibo Bryconamericus diaphanus* Beni-Madre de Dios, Marañon-Nanay Bryconamericus galvisi* Putumayo, Japurá Bryconamericus grosvenori* Beni-Madre de Dios, Ucayali Bryconamericus hyphesson Branco, Essequibo Bryconamericus macrophthalmus Negro, upper Orinoco Bryconamericus novae* Endemic – upper Tocantins Bryconamericus orinocoense Teles Pires, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, upper Orinoco Bryconamericus oroensis* Endemic – Marañon Bryconamericus orteguasae* Beni-Madre de Dios, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá Bryconamericus osgoodi* Endemic – Marañon Bryconamericus pachacuti* Beni-Madre de Dios, middle-lower Madeira, Ucayali, Japurá Bryconamericus pectinatus* Beni-Madre de Dios, Madeira Shield Tributaries, Ucayali Bryconamericus peruanus* Marañon-Nanay, Napo-Ambyiacu Bryconamericus phoenicopterus* Endemic – Napo-Ambyiacu Bryconamericus pinnavittatus* Endemic – Teles Pires Bryconamericus zamorensis* Endemic – Marañon Bryconella pallidifrons* Middle-lower Madeira, Purus, Juruá, Putumayo, Amazonas main channel Bryconexodon juruenae* Endemic – Juruena Bryconexodon trombetasi* Teles Pires, Trombetas 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 75 Occurrence Bryconops affinis Upper Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, Putumayo, Japurá, Branco, Trombetas, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Bryconops alburnoides Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Bryconops caudomaculatus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Guaporé, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Bryconops disruptus* Endemic – Negro Bryconops durbinae** Bryconops giacopinii Araguaia, upper Xingu, lower Xingu, Teles Pires, middle-lower Madeira, Putumayo, Japurá, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Bryconops gracilis* Araguaia, upper Xingu, Trombetas Bryconops humeralis Negro, upper Orinoco Bryconops inpai Middle-lower Madeira, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco Bryconops melanurus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, Putumayo, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo Bryconops munduruku* Endemic – Tapajós Bryconops piracolina* Juruena, Madeira Shield Tributaries Bryconops tocantinensis* Endemic – upper Tocantins Bryconops transitorius** Caiapobrycon tucurui* Restricted to Tocantins basin (upper Tocantins, lower Tocantins) Ceratobranchia binghami* Beni-Madre de Dios, Ucayali Ceratobranchia delotaenia* Endemic – Beni-Madre de Dios Ceratobranchia elatior* Endemic – Marañon Ceratobranchia obtusirostris* Ucayali, Japurá Charax caudimaculatus* Endemic – Beni-Madre de Dios Charax condei* Tapajós, Negro, Amazonas main channel Charax delimai* Endemic – Negro Charax gibbosus Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara Charax hemigrammus Negro, Amazonas main channel, Essequibo Charax leticiae Upper Tocantins, Araguaia, lower Tocantins, Paraná-Paraguay Charax macrolepis* Restricted to Madeira basin (Guaporé, middle-lower Madeira) Charax michaeli * Tefé, Negro, Branco, Jari, Amazonas main channel Charax niger Upper Tocantins, Negro, Araguari-Macari-Amapá Charax pauciradiatus Amazonas main channel, Capim 76 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Charax rupununi Trombetas, Essequibo Charax tectifer* Ucayali, Napo-Ambyiacu, Amazonas main channel Cheirodon luelingi* Endemic – Ucayali Cheirodon ortegai* Endemic – Ucayali Cheirodon stenodon Mamoré, Beni-Madre de Dios, Paraná-Paraguay “Cheirodon” troemneri* Restricted to Madeira basin (Mamoré, middle-lower Madeira) Chrysobrycon eliasi* Endemic – Beni-Madre de Dios Chrysobrycon hesperus* Middle-lower Madeira, Napo-Ambyiacu Chrysobrycon mojicai* Endemic – Amazonas main channel Chrysobrycon myersi* Endemic – Ucayali Chrysobrycon yoliae* Endemic – Ucayali Clupeacharax anchoveoides Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Paraná-Paraguay Creagrutus amoenus* Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá Creagrutus anary* Middle-lower Madeira, Madeira Shield Tributaries Creagrutus atrisignum* Endemic – upper Tocantins Creagrutus barrigai* Purus, Juruá, Javari, Marañon-Nanay, Napo-Ambyiacu, Amazonas main channel Creagrutus beni* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Japurá Creagrutus britskii* Upper Tocantins, lower Tocantins, Iriri Creagrutus cerritulus* Endemic – Marañon Creagrutus changae* Purus, Juruá, Ucayali Creagrutus cochui* Ucayali, Amazonas main channel Creagrutus cracentis* Lower Xingu, Tapajós Creagrutus ephippiatus Negro, lower Orinoco Creagrutus figueiredoi* Restricted to Tocantins basin (upper Tocantins, Araguaia) Creagrutus flavescens* Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá Creagrutus gephyrus* Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá Creagrutus gracilis* Endemic – Marañon Creagrutus holmi* Endemic – Marañon Creagrutus ignotus Upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Jamanxim, Paraná-Paraguay Creagrutus kunturus* Marañon-Nanay, Napo-Ambyiacu Creagrutus manu* Endemic – Beni-Madre de Dios Creagrutus maxillaris Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Negro, Branco, Essequibo, lower Orinoco, upper Orinoco Creagrutus melanzonus Branco, Maroni-Approuague, Corentyne-Demerara, Essequibo, upper Orinoco Creagrutus menezesi* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Negro, Branco Creagrutus molinus* Endemic – Araguaia Creagrutus mucipu* Upper Tocantins, upper Xingu Creagrutus muelleri* Endemic – Marañon 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 77 Occurrence Creagrutus nigrotaeniatus* Restricted to Tapajós basin (Juruena, Tapajós, Jamanxim) Creagrutus occidaneus* Beni-Madre de Dios, Purus Creagrutus ortegai* Ucayali, Marañon-Nanay Creagrutus ouranonastes* Endemic – Ucayali Creagrutus pearsoni* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios, middle-lower Madeira) Creagrutus peruanus* Endemic – Ucayali Creagrutus petilus* Middle-lower Madeira, Madeira Shield Tributaries Creagrutus phasma Negro, lower Orinoco, upper Orinoco, Apure Creagrutus pila* Endemic – Ucayali Creagrutus runa Negro, upper Orinoco Creagrutus saxatilis* Endemic – upper Tocantins Creagrutus seductus* Restricted to Tocantins basin (upper Tocantins, Araguaia) Creagrutus tuyuka* Endemic – Negro Creagrutus ungulus* Endemic – Beni-Madre de Dios Creagrutus vexillapinnus Negro, upper Orinoco Creagrutus yanatili* Endemic – Ucayali Creagrutus yudja* Iriri, lower Xingu Creagrutus zephyrus Negro, upper Orinoco Ctenobrycon hauxwellianus Upper Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, Purus, Juruá, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Branco, Amazonas main channel, Parnaíba Ctenobrycon multiradiatus** Tefé Ctenobrycon spilurus Upper Tocantins, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Purus, Juruá, Ucayali, Putumayo, Japurá, Branco, Trombetas, Jari, Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Ctenocheirodon pristis* Endemic – upper Tocantins Cyanogaster noctivaga* Endemic – Negro Cynopotamus amazonum* Upper Tocantins, Araguaia, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Japurá, Branco, Amazonas main channel Cynopotamus essequibensis Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo Cynopotamus gouldingi* Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries Cynopotamus juruenae* Restricted to Tapajós basin (Teles Pires, Juruena) Cynopotamus tocantinensis* Restricted to Tocantins basin (Araguaia, lower Tocantins) Cynopotamus xinguano* Endemic – upper Xingu Dectobrycon armeniacus** Engraulisoma taeniatum Mamoré, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Napo-Ambyiacu, Putumayo, upper Orinoco, Apure, Paraná-Paraguay Erythrocharax altipinnis* Endemic – Iriri Exodon paradoxus Upper Tocantins, Araguaia, lower Tocantins, Branco, Essequibo, lower Orinoco, upper Orinoco 78 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Galeocharax goeldii* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Galeocharax gulo Upper Tocantins, Araguaia, lower Tocantins, Teles Pires, Juruena, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Gephyrocharax major* Mamoré, Beni-Madre de Dios, Juruá, Ucayali, Marañon-Nanay Gnathocharax steindachneri Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Grundulus bogotensis Putumayo, Cauca-Magdalena-Sinu Grundulus cochae* Endemic – Putumayo Gymnocorymbus flaviolimai* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Gymnocorymbus thayeri Purus, Juruá, Ucayali, Napo-Ambyiacu, Negro, Trombetas, Amazonas main channel, Parnaíba, Gurupi-Turiaçu, Corentyne-Demerara Hasemania crenuchoides Upper Tocantins, Paraná-Paraguay Hasemania hanseni Upper Tocantins, Paraná-Paraguay Hasemania kalunga* Endemic – upper Tocantins Hasemania nambiquara* Endemic – Juruena Hemibrycon beni* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Hemibrycon divisorensis* Endemic – Ucayali Hemibrycon helleri* Ucayali, Marañon-Nanay Hemibrycon huambonicus* Beni-Madre de Dios, Marañon-Nanay Hemibrycon inambari* Beni-Madre de Dios, Madeira Shield Tributaries Hemibrycon jelskii* Mamoré, Beni-Madre de Dios, Purus, Juruá, Ucayali, Marañon-Nanay Hemibrycon metae Japurá, lower Orinoco, upper Orinoco, Apure Hemibrycon mikrostiktos* Endemic – Ucayali Hemibrycon polyodon* Endemic – Marañon Hemibrycon surinamensis Upper Tocantins, lower Tocantins, upper Xingu, lower Xingu, Tapajós, MaroniApprouague, Coppename-Suriname-Saramacca Hemibrycon tridens* Endemic – Ucayali Hemigrammus aguaruna* Marañon-Nanay, Amazonas main channel Hemigrammus analis Teles Pires, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Jutaí, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, upper Orinoco, Apure Hemigrammus arua* Endemic – Amazonas main channel Hemigrammus ataktos* Endemic – upper Tocantins Hemigrammus barrigonae Negro, lower Orinoco, upper Orinoco, Apure Hemigrammus bellottii Tapajós, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Hemigrammus bleheri Negro, upper Orinoco 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 79 Occurrence Hemigrammus coeruleus* Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel Hemigrammus cupreus* Endemic – Amazonas main channel Hemigrammus cylindricus Negro, Branco, Essequibo, upper Orinoco Hemigrammus diagonicus* Jari, Amazonas main channel Hemigrammus durbinae Guaporé, middle-lower Madeira, Purus, Amazonas main channel, Paraná-Paraguay Hemigrammus elegans Amazonas main channel, upper Orinoco, Apure Hemigrammus erythrozonus Branco, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Hemigrammus falsus** Hemigrammus filamentosus* Restricted to Tocantins basin (upper Tocantins, Araguaia) Hemigrammus geisleri* Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Amazonas main channel Hemigrammus gracilis Putumayo, Japurá, Branco, Amazonas main channel, Essequibo, upper Orinoco Hemigrammus guyanensis* Lower Tocantins, middle-lower Madeira, Madeira Shield Tributaries, Amazonas main channel, Amazonas Estuary, Gurupi-Turiaçu, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Essequibo, lower Orinoco, upper Orinoco Hemigrammus haraldi* Purus, Tefé, Ucayali, Japurá, Negro, Amazonas main channel Hemigrammus hyanuary Tapajós, Mamoré, middle-lower Madeira, Negro, Amazonas main channel, upper Orinoco Hemigrammus iota Japurá, Branco, Corentyne-Demerara, Essequibo, upper Orinoco Hemigrammus levis upper Tocantins, Araguaia, Teles Pires, Juruena, Tapajós, middle-lower Madeira, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Essequibo, lower Orinoco, upper Orinoco Hemigrammus luelingi* Ucayali, Putumayo, Japurá, Amazonas main channel Hemigrammus lunatus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel, Corentyne-Demerara, Paraná-Paraguay Hemigrammus machadoi Mamoré, Guaporé, middle-lower Madeira, Paraná-Paraguay Hemigrammus mahnerti Guaporé, Paraná-Paraguay Hemigrammus megaceps* Endemic – Ucayali Hemigrammus melanochrous* Teles Pires, middle-lower Madeira, Juruá, Putumayo, Japurá Hemigrammus microstomus Teles Pires, Putumayo, Japurá, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Hemigrammus mimus Negro, lower Orinoco, upper Orinoco, Apure Hemigrammus neptunus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Branco, Trombetas, Paraná-Paraguay Hemigrammus newboldi Upper Xingu, Essequibo, lower Orinoco, upper Orinoco, Apure Hemigrammus ocellifer Upper Tocantins, Araguaia, lower Tocantins, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Jutaí, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, upper Orinoco Hemigrammus ora Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Teles Pires, Juruena, Tapajós, Amazonas Estuary, Maroni-Approuague Hemigrammus orthus Tapajós, Corentyne-Demerara, Essequibo 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Hemigrammus parana Araguaia, Paraná-Paraguay Hemigrammus pretoensis* Negro, Urubu-Uatumã, Amazonas main channel Hemigrammus pulcher* Putumayo, Amazonas main channel Hemigrammus rodwayi Araguaia, lower Tocantins, Branco, Amazonas Estuary, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Hemigrammus schmardae Juruá, Putumayo, Japurá, Negro, Branco, Amazonas main channel, Essequibo Hemigrammus silimoni* Juruena, Madeira Shield Tributaries Hemigrammus skolioplatus* Endemic – Juruena Hemigrammus stictus Tapajós, Mamoré, Guaporé, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, Apure Hemigrammus tocantinsi* Endemic – upper Tocantins Hemigrammus tridens Guaporé, Paraná-Paraguay Hemigrammus unilineatus Lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Napo-Ambyiacu, Putumayo, Branco, Amazonas main channel, Amazonas Estuary, Gurupi-Turiaçu, Capim, Oiapoque, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Hemigrammus vorderwinkleri Tapajós, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, Trombetas, lower Orinoco, upper Orinoco, Apure Hemigrammus yinyang* Endemic – Negro Heterocharax leptogrammus Negro, upper Orinoco Heterocharax macrolepis Upper Xingu, lower Xingu, Tapajós, middle-lower Madeira, Marañon-Nanay, NapoAmbyiacu, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Heterocharax virgulatus Tapajós, middle-lower Madeira, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Hoplocharax goethei Tapajós, middle-lower Madeira, Negro, Urubu-Uatumã, Amazonas main channel, upper Orinoco Hyphessobrycon agulha* Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Jutaí, Ucayali, Putumayo, Negro, Urubu-Uatumã, Amazonas main channel Hyphessobrycon amandae* Endemic – Araguaia Hyphessobrycon amaronensis* Endemic – Putumayo Hyphessobrycon bentosi Lower Tocantins, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Ucayali, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Hyphessobrycon cachimbensis* Endemic – Teles Pires Hyphessobrycon catableptus Branco, Essequibo Hyphessobrycon clavatus** Hyphessobrycon coelestinus Upper Tocantins, Paraná-Paraguay, São Francisco Hyphessobrycon copelandi Araguaia, lower Xingu, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Jutaí, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Jari, Amazonas main channel, Maroni-Approuague, Coppename-SurinameSaramacca, Essequibo, upper Orinoco Hyphessobrycon cyanotaenia* Juruena, Guaporé, Paraná-Paraguay Hyphessobrycon delimai* Endemic – Jamanxim 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 81 Occurrence Hyphessobrycon diancistrus Upper Xingu, Iriri, Teles Pires, Tapajós, middle-lower Madeira, Purus, Tefé, Japurá, Negro, Amazonas main channel, upper Orinoco Hyphessobrycon diastatos* Upper Tocantins, São Francisco Hyphessobrycon dorsalis* Teles Pires, Negro, Branco Hyphessobrycon eilyos* Endemic – Araguaia Hyphessobrycon elachys Mamoré, Guaporé, Paraná-Paraguay Hyphessobrycon epicharis Negro, upper Orinoco Hyphessobrycon eques Upper Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Amazonas main channel, Paraná-Paraguay Hyphessobrycon ericae* Tapajós, middle-lower Madeira, Trombetas Hyphessobrycon erythrostigma* Purus, Tefé, Marañon-Nanay, Putumayo, Japurá, Branco, Amazonas main channel Hyphessobrycon eschwartzae* Endemic – Beni-Madre de Dios Hyphessobrycon frankei* Endemic – Ucayali Hyphessobrycon gracilior* Beni-Madre de Dios, middle-lower Madeira, Ucayali, Japurá, Amazonas main channel Hyphessobrycon hamatus* Endemic – upper Tocantins Hyphessobrycon haraldschultzi* Restricted to Tocantins basin (Araguaia, lower Tocantins) Hyphessobrycon hasemani* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Hyphessobrycon heliacus* Endemic – Teles Pires Hyphessobrycon heterorhabdus Lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, Capim Hyphessobrycon hexastichos* Juruena, Guaporé Hyphessobrycon inconstans** Hyphessobrycon jackrobertsi* Endemic – Marañon Hyphessobrycon juruna* Restricted to Xingu basin (upper Xingu, lower Xingu) Hyphessobrycon kayabi* Endemic – Teles Pires Hyphessobrycon khardinae* Middle-lower Madeira, Purus Hyphessobrycon krenakore* Endemic – Iriri Hyphessobrycon langeanii* Endemic – Araguaia Hyphessobrycon loretoensis* Purus, Putumayo, Japurá, Amazonas main channel Hyphessobrycon loweae* Araguaia, upper Xingu Hyphessobrycon lucenorum* Endemic – Madeira Shield Tributaries Hyphessobrycon margitae* Endemic – Marañon Hyphessobrycon megalopterus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Paraná-Paraguay Hyphessobrycon melanostichos* Juruena, Madeira Shield Tributaries Hyphessobrycon melasemeion** Hyphessobrycon melazonatus* Japurá, Amazonas main channel Hyphessobrycon minor Branco, Essequibo, lower Orinoco, upper Orinoco Hyphessobrycon moniliger Upper Tocantins, Araguaia, upper Xingu, Teles Pires, Juruena, Paraná-Paraguay Hyphessobrycon montagi* Endemic – Amazonas main channel 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Hyphessobrycon mutabilis* Endemic – upper Xingu Hyphessobrycon myrmex* Endemic – Juruena Hyphessobrycon nigricinctus* Endemic – Beni-Madre de Dios Hyphessobrycon notidanos* Juruena, Madeira Shield Tributaries Hyphessobrycon oritoensis* Endemic – Putumayo Hyphessobrycon paepkei* Endemic – Negro Hyphessobrycon pando* Restricted to Madeira basin (Beni-Madre de Dios, middle-lower Madeira) Hyphessobrycon peruvianus* Putumayo, Japurá, Amazonas main channel Hyphessobrycon petricolus* Endemic – Madeira Shield Tributaries Hyphessobrycon peugeoti* Endemic – Juruena Hyphessobrycon pinnistriatus* Endemic – Teles Pires Hyphessobrycon piranga* Endemic – Juruena Hyphessobrycon platyodus* Endemic – Madeira Shield Tributaries Hyphessobrycon procyon* Endemic – Madeira Shield Tributaries Hyphessobrycon psittacus* Juruena, Guaporé Hyphessobrycon pulchripinnis* Lower Xingu, Teles Pires, Tapajós, Urubu-Uatumã Hyphessobrycon pyrrhonotus* Endemic – Negro Hyphessobrycon robustulus* Endemic – Amazonas main channel Hyphessobrycon rosaceus Trombetas, Capim, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco Hyphessobrycon scholzei* Japurá Hyphessobrycon scutulatus* Endemic – Teles Pires Hyphessobrycon socolofi* Negro, Branco, Amazonas main channel Hyphessobrycon stegemanni* Endemic – upper Tocantins Hyphessobrycon stramineus** Amazonas main channel Hyphessobrycon sweglesi Mamoré, middle-lower Madeira, Purus, Negro, Branco, Amazonas main channel, upper Orinoco, Apure Hyphessobrycon taguae Putumayo, upper Orinoco Hyphessobrycon taphorni* Endemic – Beni-Madre de Dios Hyphessobrycon tenuis* Endemic – Ucayali Hyphessobrycon tropis* Endemic – Negro Hyphessobrycon tukunai* Putumayo, Amazonas main channel Hyphessobrycon vanzolinii* Endemic – Tapajós Hyphessobrycon vilmae Upper Xingu, Teles Pires, Juruena, Madeira Shield Tributaries, Paraná-Paraguay Hyphessobrycon wadai* Endemic – Juruena Hyphessobrycon weitzmanorum* Endemic – Araguaia Hyphessobrycon wosiackii* Endemic – Middle-lower Madeira Iguanodectes adujai Negro, upper Orinoco Iguanodectes geisleri Middle-lower Madeira, Madeira Shield Tributaries, Japurá, Negro, Branco, lower Orinoco, upper Orinoco 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 83 Occurrence Iguanodectes gracilis* Restricted to Negro basin (Negro, Branco) Iguanodectes polylepis* Purus Iguanodectes purusii* Teles Pires, Guaporé, middle-lower Madeira, Amazonas main channel Iguanodectes rachovii Amazonas main channel, Capim Iguanodectes spilurus Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Iguanodectes variatus* Mamoré, Branco Inpaichthys kerri* Juruena, Madeira Shield Tributaries Jupiaba abramoides Iriri, Negro, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco Jupiaba acanthogaster Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Trombetas, Paraná-Paraguay Jupiaba anteroides Lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, middle-lower Madeira, Javari, Japurá, Negro, Trombetas, Amazonas main channel, Capim, lower Orinoco, upper Orinoco Jupiaba apenima* Upper Tocantins, Araguaia, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós Jupiaba asymmetrica* Jutaí, Putumayo, Japurá, Amazonas main channel Jupiaba atypindi Negro, Branco, Trombetas, Essequibo, upper Orinoco Jupiaba citrina* Endemic – Madeira Shield Tributaries Jupiaba elassonaktis* Endemic – upper Tocantins Jupiaba essequibensis Lower Xingu, Madeira Shield Tributaries, Jari, Essequibo Jupiaba iasy* Lower Xingu, Teles Pires, Tapajós, Jamanxim, Madeira Shield Tributaries Jupiaba kurua* Endemic – Iriri Jupiaba meunieri Iriri, Teles Pires, Juruena, Jamanxim, Trombetas, Jari, Araguari-Macari-Amapá, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara Jupiaba minor* Endemic – Teles Pires Jupiaba ocellata Curuá-Paru do Oeste, Oiapoque Jupiaba paranatinga* Restricted to Tapajós basin (Teles Pires, Jamanxim) Jupiaba pinnata Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco Jupiaba pirana* Restricted to Tapajós basin (Teles Pires, Tapajós) Jupiaba poekotero* Endemic – Negro Jupiaba polylepis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Branco, Jari, Parnaíba, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Jupiaba poranga* Teles Pires, Juruena, Madeira Shield Tributaries Jupiaba scologaster Negro, Branco, lower Orinoco, upper Orinoco Jupiaba yarina* Endemic – Juruena Jupiaba zonata Lower Tocantins, middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Amazonas main channel, Capim, lower Orinoco 84 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Knodus borki* Endemic – Amazonas main channel Knodus breviceps** Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Japurá, lower Orinoco, upper Orinoco Knodus chapadae Upper Tocantins, Guaporé, Paraná-Paraguay Knodus delta* Marañon-Nanay, Napo-Ambyiacu, Putumayo Knodus dorsomaculatus* Endemic – Teles Pires Knodus figueiredoi* Endemic – Araguaia Knodus gamma* Beni-Madre de Dios, middle-lower Madeira, Marañon-Nanay, Napo-Ambyiacu Knodus heteresthes Araguaia, Tapajós, Jamanxim, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Negro, Trombetas, Amazonas main channel, MaroniApprouague, upper Orinoco Knodus hypopterus Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Japurá, lower Orinoco, upper Orinoco, Apure Knodus jacunda** Knodus longus** Endemic – Beni-Madre de Dios Knodus megalops* Ucayali, Marañon-Nanay Knodus mizquae* Restricted to Madeira basin (Mamoré, Guaporé) Knodus moenkhausii Iriri, Mamoré, Guaporé, Beni-Madre de Dios, Paraná-Paraguay Knodus pasco* Endemic – Ucayali Knodus savannensis Upper Tocantins, Araguaia, Itapicuru-Mearim, Capim Knodus septentrionalis** Putumayo Knodus shinahota* Endemic – Mamoré Knodus smithi* Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali Leptagoniates steindachneri* Upper Tocantins, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá Leptobrycon jatuaranae* Endemic – Negro Lonchogenys ilisha Putumayo, Negro, Amazonas main channel, upper Orinoco, Apure Macropsobrycon xinguensis* Araguaia, upper Xingu, Teles Pires Markiana geayi Japurá, lower Orinoco, upper Orinoco, Apure Markiana nigripinnis Mamoré, Guaporé, Beni-Madre de Dios, Paraná-Paraguay Microgenys lativirgata* Endemic – Marañon Microgenys weyrauchi** Ucayali Microschemobrycon callops Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Jutaí, Japurá, Negro, Branco, Essequibo, lower Orinoco, upper Orinoco, Apure Microschemobrycon casiquiare Iriri, lower Xingu, Teles Pires, Jamanxim, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Japurá, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Microschemobrycon elongatus Araguaia, upper Xingu, Teles Pires, Beni-Madre de Dios, middle-lower Madeira, Itapicuru-Mearim, Capim, Paraná-Paraguay Microschemobrycon geisleri Araguaia, Teles Pires, Tapajós, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Negro, Capim, Essequibo Microschemobrycon guaporensis* Restricted to Madeira basin (Mamoré, Guaporé, middle-lower Madeira) 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 85 Occurrence Microschemobrycon melanotus Teles Pires, Mamoré, Guaporé, middle-lower Madeira, Coari-Urucu, Jutaí, Japurá, Negro, Amazonas main channel, Essequibo Microschemobrycon meyburgi* Restricted to Negro basin (Negro, Branco) Moenkhausia abyss* Upper Tocantins, Araguaia, lower Tocantins, Tapajós, middle-lower Madeira, Negro, Trombetas, Paru, Jari, Amazonas main channel Moenkhausia affinis* Endemic – Negro Moenkhausia agnesae* Endemic – Amazonas main channel Moenkhausia alesis* Endemic – upper Xingu Moenkhausia atahualpiana** Moenkhausia aurantia* Upper Tocantins, Araguaia, Parará-Paraguay, São Francisco Moenkhausia barbouri* Endemic – Amazonas main channel Moenkhausia beninei* Endemic – Negro Moenkhausia bonita Middle-lower Madeira, Paraná-Paraguay Moenkhausia britskii* Endemic – Putumayo Moenkhausia celibela* Iriri, lower Xingu, Teles Pires, Tapajós, Jamanxim, middle-lower Madeira, Branco, Jari, Amazonas main channel Moenkhausia ceros Teles Pires, Tapajós, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Tefé, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Essequibo Moenkhausia chlorophthalma* Endemic – Iriri Moenkhausia chrysargyrea Lower Tocantins, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Napo-Ambyiacu, Putumayo, Branco, Amazonas main channel, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Moenkhausia collettii Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Moenkhausia comma Lower Tocantins, lower Xingu, Tapajós, Mamoré, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure Moenkhausia conspicua* Endemic – Tapajós Moenkhausia copei Napo-Ambyiacu, Putumayo, Japurá, Branco, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Moenkhausia cosmops Juruena, Paraná-Paraguay Moenkhausia cotinho Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Ucayali, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Moenkhausia crisnejas* Endemic – Marañon Moenkhausia dasalmas* Endemic – upper Tocantins Moenkhausia dichroura Upper Tocantins, Araguaia, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay 86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Moenkhausia diktyota* Middle-lower Madeira, Negro Moenkhausia dorsinuda* Restricted to Madeira basin (Mamoré, Guaporé) Moenkhausia eurystaenia* Endemic – upper Xingu Moenkhausia goya* Endemic – upper Tocantins Moenkhausia gracilima Araguaia, Juruena, Tapajós, middle-lower Madeira, Japurá, Negro, Trombetas, Amazonas main channel, upper Orinoco Moenkhausia grandisquamis* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Branco, Amazonas main channel, Oiapoque, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Moenkhausia hasemani Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, middle-lower Madeira, Negro, Amazonas main channel, Capim Moenkhausia heikoi* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Moenkhausia hemigrammoides Tapajós, Negro, Branco, Trombetas, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara Moenkhausia hysterosticta Upper Tocantins, middle-lower Madeira, Purus, Trombetas, Apure Moenkhausia icae* Putumayo, Japurá, Amazonas main channel Moenkhausia intermedia Upper Xingu, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Amazonas main channel, Maroni-Approuague, lower Orinoco, upper Orinoco, Paraná-Paraguay Moenkhausia ischyognatha* Endemic – upper Tocantins Moenkhausia jamesi* Purus, Juruá, Amazonas main channel, Amazonas Estuary Moenkhausia justae* Araguaia, Japurá, Negro, Amazonas main channel Moenkhausia lata Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Negro, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco Moenkhausia latissima* Endemic – Amazonas main channel Moenkhausia lepidura Araguaia, lower Tocantins, Tapajós, Guaporé, middle-lower Madeira, Purus, Putumayo, Negro, Trombetas, Amazonas main channel, upper Orinoco, Apure Moenkhausia levidorsa* Juruena, Madeira Shield Tributaries Moenkhausia lineomaculata* Endemic – Juruena Moenkhausia lopesi Araguaia, Juruena, Paraná-Paraguay Moenkhausia loweae* Upper Tocantins, Araguaia, upper Xingu, Iriri Moenkhausia margitae* Endemic – Ucayali Moenkhausia megalops Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, Jamanxim, middlelower Madeira, Putumayo, Japurá, Branco, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Moenkhausia melogramma Putumayo, Amazonas main channel, upper Orinoco Moenkhausia miangi Putumayo, Branco, Essequibo, lower Orinoco, upper Orinoco Moenkhausia mikia Lower Xingu, Teles Pires, Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Coari-Urucu, Ucayali, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel, upper Orinoco Moenkhausia monicae* Endemic – Juruena Moenkhausia mutum* Endemic – Juruena 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 87 Occurrence Moenkhausia naponis* Napo-Ambyiacu, Putumayo, Amazonas main channel Moenkhausia newtoni** Upper Xingu Moenkhausia nigromarginata* Endemic – Juruena Moenkhausia oligolepis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Juruá, Jutaí, Ucayali, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Araguari-Macari-Amapá, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Moenkhausia orteguasae* Endemic – Japurá Moenkhausia ovalis* Japurá Moenkhausia pankilopteryx* Restricted to Tocantins basin (upper Tocantins, lower Tocantins) Moenkhausia parecis* Endemic – Madeira Shield Tributaries Moenkhausia petymbuaba* Endemic – Iriri Moenkhausia phaeonota Araguaia, upper Xingu, Teles Pires, Juruena, Paraná-Paraguay Moenkhausia pirauba* Upper Xingu, Iriri, Teles Pires, Juruena, Jamanxim Moenkhausia plumbea* Endemic – Teles Pires Moenkhausia pyrophthalma* Araguaia, upper Xingu Moenkhausia robertsi* Japurá, Amazonas main channel Moenkhausia rubra* Juruena, Guaporé Moenkhausia sthenosthoma* Restricted to Madeira basin (Mamoré, Guaporé, middle-lower Madeira) Moenkhausia surinamensis Trombetas, Araguari-Macari-Amapá, Oiapoque, Coppename-Suriname-Saramacca, Corentyne-Demerara, upper Orinoco Moenkhausia tergimacula* Endemic – upper Tocantins Moenkhausia tridentata** Moenkhausia uirapuru* Juruena, Guaporé Moenkhausia venerei* Endemic – Araguaia Moenkhausia xinguensis* Restricted to Xingu basin (upper Xingu, lower Xingu) Monotocheirodon drilos* Endemic – Beni-Madre de Dios Monotocheirodon kontos* Endemic – Beni-Madre de Dios Monotocheirodon pearsoni* Endemic – Beni-Madre de Dios Odontostilbe dierythrura* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Odontostilbe ecuadorensis* Marañon-Nanay, Napo-Ambyiacu, Putumayo Odontostilbe euspilurus* Beni-Madre de Dios, Purus, Ucayali, Marañon-Nanay, Napo-Ambyiacu Odontostilbe fugitiva* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Amazonas main channel Odontostilbe gracilis Branco, Maroni-Approuague, Corentyne-Demerara, Essequibo, lower Orinoco Odontostilbe nareuda* Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus Odontostilbe paraguayensis Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Paraná-Paraguay 88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Odontostilbe parecis* Endemic – Guaporé Odontostilbe pulchra Negro, lower Orinoco, upper Orinoco, Apure Oligosarcus bolivianus Mamoré, Guaporé, Paraná-Paraguay Oligosarcus pintoi Guaporé, Paraná-Paraguay Oligosarcus schindleri* Endemic – Mamoré Othonocheirodus eigenmanni* Endemic – Marañon Oxybrycon parvulus* Middle-lower Madeira, Ucayali Paracheirodon axelrodi Negro, Coppename-Suriname-Saramacca, upper Orinoco Paracheirodon innesi* Middle-lower Madeira, Purus, Ucayali, Putumayo, Japurá, Amazonas main channel Paracheirodon simulans Negro, Branco, upper Orinoco Paragoniates alburnus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Parapristella aubynei Branco, Corentyne-Demerara, Essequibo, upper Orinoco Parapristella georgiae Araguaia, Branco, lower Orinoco, upper Orinoco, Apure Parecbasis cyclolepis* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali Petitella georgiae* Middle-lower Madeira, Purus, Marañon-Nanay, Negro Phallobrycon adenacanthus* Restricted to Xingu basin (upper Xingu, Iriri) Phallobrycon synarmacanthus* Endemic – lower Xingu Phenacogaster beni* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries Phenacogaster capitulata* Juruá, Ucayali Phenacogaster eurytaenia* Endemic – upper Tocantins Phenacogaster microstictus* Branco, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Phenacogaster naevata* Endemic – upper Tocantins Phenacogaster napoatilis* Endemic – Napo-Ambyiacu Phenacogaster ojitatus* Endemic – Iriri Phenacogaster pectinatus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Putumayo, Japurá, Amazonas main channel Phenacogaster prolatus Negro, upper Orinoco Phenacogaster retropinnus* Upper Xingu, middle-lower Madeira, Madeira Shield Tributaries, Negro Phycocharax rasbora* Endemic – Teles Pires Piabucus caudomaculatus* Restricted to Madeira basin (Mamoré, Guaporé) Piabucus dentatus Amazonas Estuary, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, lower Orinoco Piabucus melanostoma Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Paraná-Paraguay Poptella brevispina Lower Tocantins, Branco, Trombetas, Capim, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Poptella compressa* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Japurá, Branco, Parnaíba, Itapicuru-Mearim, Essequibo, lower Orinoco, upper Orinoco, Apure 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 89 Occurrence Poptella longipinnis Lower Tocantins, Branco, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Priocharax ariel Middle-lower Madeira, Negro, Amazonas main channel, upper Orinoco Priocharax nanus* Endemic – Negro Priocharax pygmaeus* Endemic – Amazonas main channel Prionobrama filigera* Lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Amazonas main channel Pristella maxillaris Lower Tocantins, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Prodontocharax alleni* Mamoré, middle-lower Madeira, Ucayali, Amazonas main channel Prodontocharax howesi* Endemic – Mamoré Prodontocharax melanotus* Mamoré, Guaporé, Beni-Madre de Dios, Purus, Ucayali Protocheirodon pi* Middle-lower Madeira, Purus, Ucayali, Amazonas main channel Ptychocharax rhyacophila Negro, upper Orinoco Rhinobrycon negrensis Negro, upper Orinoco Rhinopetitia myersi* Endemic – Araguaia Rhinopetitia potamorhachia* Endemic – Teles Pires Roeboexodon guyanensis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Juruena, Tapajós, Jamanxim, Jari, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara Roeboides affinis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Jutaí, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Roeboides biserialis* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Amazonas main channel Roeboides descalvadensis Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Trombetas, Amazonas main channel, Amazonas Estuary, Paraná-Paraguay Roeboides dispar* Mamoré, middle-lower Madeira, Juruá, Ucayali Roeboides myersii Araguaia, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Trombetas, Amazonas main channel, lower Orinoco Roeboides oligistos* Tapajós, Negro, Branco, Trombetas Salminus brasiliensis Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Paraná-Paraguay Salminus hilarii Upper Tocantins, Araguaia, lower Tocantins, Guaporé, Beni-Madre de Dios, Putumayo, Japurá, Branco, Paraná-Paraguay “Salminus iquitensis” Upper Tocantins, Araguaia, lower Tocantins, Mamoré, Beni-Madre de Dios, middlelower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Putumayo, Japurá, Branco, upper Orinoco, Apure Schultzites axelrodi Negro, upper Orinoco Scopaeocharax atopodus* Endemic – Marañon Scopaeocharax rhinodus* Endemic – Marañon Serrabrycon magoi Negro, lower Orinoco, upper Orinoco, Apure 90 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Serrapinnus aster* Restricted to Tocantins basin (upper Tocantins, Araguaia) Serrapinnus lucindai* Restricted to Tocantins basin (upper Tocantins, Araguaia) Serrapinnus malabarbai* Endemic – Araguaia Serrapinnus microdon Beni-Madre de Dios, middle-lower Madeira, Paraná-Paraguay Serrapinnus micropterus* Araguaia, Teles Pires, Tapajós, Mamoré, middle-lower Madeira, Amazonas main channel Serrapinnus notomelas Teles Pires, Paraná-Paraguay Serrapinnus sterbai* Endemic – upper Tocantins Serrapinnus tocantinensis* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Stethaprion crenatum* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Amazonas main channel, Amazonas Estuary Stethaprion erythrops* Ucayali, Marañon-Nanay, Amazonas main channel Stichonodon insignis* Purus, Amazonas main channel Tetragonopterus anostomus* Restricted to Tocantins basin (upper Tocantins, Araguaia) Tetragonopterus araguaiensis* Endemic – Araguaia Tetragonopterus argenteus Upper Tocantins, Araguaia, upper Xingu, lower Xingu, Beni-Madre de Dios, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Amazonas main channel, Parnaíba, Essequibo, upper Orinoco, Paraná-Paraguay Tetragonopterus carvalhoi* Endemic – Jari Tetragonopterus chalceus Upper Tocantins, Araguaia, upper Xingu, lower Xingu, Teles Pires, Guaporé, Madeira Shield Tributaries, Napo-Ambyiacu, Negro, Branco, Amazonas main channel, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco Tetragonopterus denticulatus* Restricted to Tocantins basin (upper Tocantins, Araguaia) Tetragonopterus juruena* Restricted to Tapajós basin (Teles Pires, Juruena) Tetragonopterus kuluene* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Tetragonopterus manaos* Tefé, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel Tetragonopterus ommatus* Endemic – Tapajós Tetragonopterus rarus Jari, Maroni-Approuague, Corentyne-Demerara Thayeria boehlkei Araguaia, upper Xingu, Teles Pires, Guaporé, Putumayo, Japurá, upper Orinoco Thayeria obliqua Mamoré, Guaporé, Putumayo, upper Orinoco Thayeria tapajonica* Restricted to Tapajós basin (Teles Pires, Tapajós) Thrissobrycon pectinifer Negro, Essequibo, upper Orinoco Triportheus albus* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary Triportheus angulatus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Essequibo Triportheus auritus Upper Tocantins, Araguaia, lower Tocantins, lower Xingu, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Purus, Javari, Putumayo, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco, Apure 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 91 Occurrence Triportheus brachipomus Upper Xingu, lower Xingu, Juruena, Tapajós, Negro, Branco, Trombetas, Jari, Araguari-Macari-Amapá, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Triportheus culter* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Ucayali, Napo-Ambyiacu Triportheus curtus* Purus, Ucayali, Amazonas Estuary Triportheus pictus* Jutaí, Japurá, Amazonas main channel, Amazonas Estuary Triportheus rotundatus Upper Xingu, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, Purus, Ucayali, Branco, Amazonas main channel, Amazonas Estuary, Corentyne-Demerara, Essequibo Triportheus trifurcatus* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Trochilocharax ornatus** Tucanoichthys tucano* Endemic – Negro Tyttobrycon dorsimaculatus* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Tyttobrycon hamatus* Endemic – Marañon Tyttobrycon marajoara* Endemic – Amazonas Estuary Tyttobrycon spinosus* Endemic – Mamoré Tyttobrycon xeruini* Tapajós, Negro, Branco Tyttocharax cochui* Endemic – Amazonas main channel Tyttocharax madeirae Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Urubu-Uatumã, Amazonas main channel, Capim Tyttocharax tambopatensis* Restricted to Madeira basin (Beni-Madre de Dios, middle-lower Madeira) Xenurobrycon coracoralinae* Endemic – Araguaia Xenurobrycon heterodon* Purus, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Japurá Xenurobrycon polyancistrus* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Xenurobrycon pteropus* Endemic – Amazonas main channel Xenurobrycon varii* Restricted to Tapajós basin (Tapajós, Jamanxim) Chilodontidae 7 Caenotropus labyrinthicus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Coari-Urucu, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, UrubuUatumã, Trombetas, Jari, Amazonas main channel, Parnaíba, Capim, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Caenotropus maculosus Trombetas, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, upper Orinoco Caenotropus mestomorgmatos Japurá, Negro, Amazonas main channel, lower Orinoco, upper Orinoco Caenotropus schizodon* Restricted to Tapajós basin (Teles Pires, Juruena) Chilodus fritillus* Endemic – Beni-Madre de Dios Chilodus gracilis Madeira Shield Tributaries, Putumayo, Negro, Amazonas main channel, upper Orinoco Chilodus punctatus Upper Tocantins, Araguaia, lower Tocantins, Tapajós, Mamoré, Guaporé, BeniMadre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Jutaí, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Capim, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure 92 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Crenuchidae Species NO. 431 Occurrence 47 Ammocryptocharax elegans Teles Pires, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Ammocryptocharax minutus Negro, Branco, lower Orinoco, upper Orinoco Ammocryptocharax vintonae Trombetas, Essequibo, lower Orinoco Characidium boaevistae Branco, lower Orinoco, upper Orinoco, Apure, Maracaibo, Cauca-Magdalena-Sinu Characidium boehlkei* Endemic – Napo-Ambyiacu Characidium bolivianum* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios) Characidium crandellii* Endemic – Branco Characidium declivirostre Negro, lower Orinoco, upper Orinoco Characidium etheostoma* Endemic – Amazonas main channel Characidium fasciatum Mamoré, Guaporé, Beni-Madre de Dios, Juruá, Putumayo, Japurá, Amazonas main channel, Essequibo, Cauca-Magdalena-Sinu, Paraná-Paraguay Characidium hasemani Branco, Essequibo Characidium heinianum* Restricted to Madeira basin (Mamoré, Guaporé) Characidium longum Negro, lower Orinoco, upper Orinoco Characidium mirim* Endemic – Araguaia Characidium nana* Iriri, Tapajós Characidium papachibe* Endemic – Amazonas main channel Characidium pellucidum Branco, Amazonas main channel, Maroni-Approuague, Essequibo, lower Orinoco, upper Orinoco, Apure Characidium pteroides Branco, Corentyne-Demerara, Essequibo, upper Orinoco Characidium purpuratum* Guaporé, Ucayali, Napo-Ambyiacu Characidium roesseli* Endemic – Amazonas main channel Characidium schindleri* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Characidium steindachneri Guaporé, Ucayali, Napo-Ambyiacu, Japurá, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Characidium stigmosum* Endemic – upper Tocantins Characidium summum* Endemic – Middle-lower Madeira Characidium xanthopterum Upper Tocantins, Paraná-Paraguay Characidium xavante* Endemic – upper Xingu Characidium zebra Upper Tocantins, Araguaia, Iriri, Mamoré, Guaporé, Beni-Madre de Dios, Japurá, Branco, Amazonas main channel, Parnaíba, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Crenuchus spilurus* Lower Tocantins, lower Xingu, Tapajós, Mamoré, middle-lower Madeira, Purus, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Elachocharax geryi Negro, Branco, lower Orinoco, upper Orinoco, Apure Elachocharax junki Middle-lower Madeira, Negro, Branco, Amazonas main channel, Essequibo Elachocharax mitopterus Negro, Branco, upper Orinoco 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 93 Occurrence Elachocharax pulcher Teles Pires, Tapajós, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Geryichthys sterbai* Endemic – Ucayali Klausewitzia ritae* Javari, Ucayali, Amazonas main channel Leptocharacidium omospilus Negro, upper Orinoco Melanocharacidium auroradiatum* Araguaia, upper Xingu, Iriri, Teles Pires Melanocharacidium depressum Upper Xingu, Negro, Branco, Urubu-Uatumã, Amazonas main channel, upper Orinoco Melanocharacidium dispilomma Upper Xingu, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Javari, Ucayali, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Melanocharacidium nigrum* Endemic – Branco Melanocharacidium pectorale Mamoré, Guaporé, Madeira Shield Tributaries, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco Melanocharacidium rex* Endemic – Marañon Microcharacidium eleotrioides Mamoré, Guaporé, middle-lower Madeira, Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Microcharacidium geryi* Endemic – Ucayali Microcharacidium gnomus Negro, lower Orinoco, upper Orinoco Microcharacidium weitzmani Lower Tocantins, Tapajós, Mamoré, Guaporé, Napo-Ambyiacu, Negro, Jari, Amazonas Estuary, Capim, upper Orinoco Odontocharacidium aphanes Middle-lower Madeira, Purus, Juruá, Negro, Branco, Amazonas main channel, upper Orinoco Poecilocharax weitzmani Negro, Branco, Urubu-Uatumã, Amazonas main channel, lower Orinoco, upper Orinoco Curimatidae 74 Curimata acutirostris* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Curimata aspera* Mamoré, Guaporé, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Amazonas main channel Curimata cerasina Japurá, lower Orinoco, Apure Curimata cisandina* Ucayali, Branco, Amazonas main channel Curimata cyprinoides Upper Tocantins, lower Tocantins, upper Xingu, lower Xingu, Branco, Trombetas, Jari, Amazonas Estuary, Capim, Araguari-Macari-Amapá, Oiapoque, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Curimata incompta Amazonas main channel, lower Orinoco, upper Orinoco Curimata inornata* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Branco, Trombetas, Amazonas main channel Curimata knerii* Teles Pires, Tapajós, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Negro, Trombetas, Jari, Amazonas main channel 94 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Curimata ocellata Lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, UrubuUatumã, Trombetas, Amazonas main channel, Amazonas Estuary, lower Orinoco, upper Orinoco Curimata roseni Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Curimata vittata Lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, BeniMadre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, UrubuUatumã, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Curimatella alburnus Lower Tocantins, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Essequibo Curimatella dorsalis Araguaia, lower Tocantins, upper Xingu, Teles Pires, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Japurá, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Curimatella immaculata* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Juruá, Ucayali, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Parnaíba, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Curimatella meyeri* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel Curimatopsis cryptica Araguaia, lower Xingu, Tapajós, Mamoré, middle-lower Madeira, Negro, Branco, Trombetas, Amazonas main channel, Capim, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Curimatopsis evelynae Tapajós, Japurá, Negro, Branco, Amazonas main channel, upper Orinoco, Apure Curimatopsis guaporensis* Restricted to Madeira basin (Guaporé, middle-lower Madeira) Curimatopsis jaci* Endemic – Teles Pires Curimatopsis macrolepis Araguaia, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Curimatopsis maculosa* Endemic – Tapajós Curimatopsis melanura* Endemic – Trombetas Curimatopsis microlepis* Purus, Amazonas main channel Curimatopsis pallida* Endemic – Negro Cyphocharax abramoides Lower Xingu, Tapajós, middle-lower Madeira, Branco, Urubu-Uatumã, Trombetas, Capim, upper Orinoco Cyphocharax aninha* Endemic – Paru Cyphocharax biocellatus Teles Pires, Maroni-Approuague Cyphocharax boiadeiro* Endemic – Araguaia Cyphocharax derhami* Ucayali, Marañon-Nanay 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 95 Occurrence Cyphocharax festivus Upper Xingu, lower Xingu, Tapajós, middle-lower Madeira, Marañon-Nanay, NapoAmbyiacu, Putumayo, Branco, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Cyphocharax gangamon* Restricted to Tapajós basin (Juruena, Tapajós) Cyphocharax gouldingi Araguaia, lower Tocantins, lower Xingu, Napo-Ambyiacu, Capim, Araguari-MacariAmapá, Oiapoque, lower Orinoco Cyphocharax helleri Jari, Araguari-Macari-Amapá, Oiapoque, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo Cyphocharax laticlavius* Endemic – Napo-Ambyiacu Cyphocharax leucostictus Upper Tocantins, Araguaia, upper Xingu, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Negro, Branco, Amazonas main channel, Araguari-Macari-Amapá, Essequibo, lower Orinoco, upper Orinoco Cyphocharax mestomyllon* Endemic – Negro Cyphocharax microcephalus Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo Cyphocharax multilineatus Negro, Branco, upper Orinoco Cyphocharax muyrakytan* Endemic – Tapajós Cyphocharax nigripinnis* Tapajós, middle-lower Madeira, Putumayo, Japurá, Negro, Branco, Amazonas main channel Cyphocharax notatus Araguaia, lower Tocantins, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Napo-Ambyiacu, Negro, Amazonas main channel, Capim Cyphocharax pantostictos* Purus, Napo-Ambyiacu, Putumayo, Amazonas main channel Cyphocharax plumbeus* Araguaia, lower Tocantins, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Negro, Branco, Amazonas main channel Cyphocharax sanctigabrielis* Endemic – Negro Cyphocharax signatus* Endemic – Araguaia Cyphocharax spiluropsis* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Negro, Amazonas main channel Cyphocharax spilurus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Jamanxim, Guaporé, Putumayo, Negro, Branco, Trombetas, Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Cyphocharax stilbolepis* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Amazonas main channel Cyphocharax vanderi Upper Tocantins, Paraná-Paraguay Cyphocharax vexillapinnus* Marañon-Nanay, Napo-Ambyiacu, Amazonas main channel Potamorhina altamazonica Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Putumayo, Japurá, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Potamorhina latior* Iriri, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel Potamorhina pristigaster Middle-lower Madeira, Purus, Ucayali, Napo-Ambyiacu, Japurá, Branco, UrubuUatumã, Amazonas main channel, Itapicuru-Mearim 96 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Psectrogaster amazonica* Upper Tocantins, Araguaia, lower Tocantins, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Trombetas, Amazonas main channel Psectrogaster ciliata Negro, Branco, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Psectrogaster curviventris Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Paraná-Paraguay Psectrogaster essequibensis Teles Pires, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Branco, Amazonas main channel, Corentyne-Demerara, Essequibo Psectrogaster falcata* Lower Xingu, Tapajós, Jamanxim, middle-lower Madeira, Amazonas main channel, Amazonas Estuary Psectrogaster rutiloides* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Putumayo, Trombetas, Amazonas main channel, Amazonas Estuary Steindachnerina amazonica* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu Steindachnerina argentea Amazonas main channel, lower Orinoco, upper Orinoco, Apure Steindachnerina bimaculata Upper Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Javari, Ucayali, Marañon-Nanay, Amazonas main channel, Essequibo, lower Orinoco Steindachnerina binotata Mamoré, Beni-Madre de Dios, Amazonas main channel, Itapicuru-Mearim Steindachnerina brevipinna Upper Tocantins, Iriri, Paraná-Paraguay Steindachnerina dobula* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Amazonas main channel Steindachnerina fasciata* Teles Pires, Juruena, Madeira Shield Tributaries Steindachnerina gracilis* Restricted to Tocantins basin (Araguaia, lower Tocantins) Steindachnerina guentheri Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Steindachnerina hypostoma* Beni-Madre de Dios, middle-lower Madeira, Juruá, Javari, Ucayali, Marañon-Nanay, Putumayo, Branco, Amazonas main channel Steindachnerina leucisca* Upper Tocantins, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, CoariUrucu, Juruá, Javari, Ucayali, Marañon-Nanay, Trombetas, Amazonas main channel Steindachnerina notograptos* Endemic – upper Tocantins Steindachnerina planiventris Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Napo-Ambyiacu, Japurá, Negro, Branco, Amazonas main channel, Essequibo Steindachnerina quasimodoi* Javari, Amazonas main channel Steindachnerina seriata* Endemic – Jamanxim Cynodontidae 10 Cynodon gibbus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Cynodon septenarius Teles Pires, Tapajós, Madeira Shield Tributaries, Tefé, Negro, Branco, UrubuUatumã, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 97 Occurrence Hydrolycus armatus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure Hydrolycus scomberoides* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel Hydrolycus tatauaia Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure Hydrolycus wallacei Japurá, Negro, Branco, upper Orinoco Rhaphiodon vulpinus* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Javari, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Roestes itupiranga* Endemic – lower Tocantins Roestes molossus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel, Essequibo Juruá, Marañon-Nanay, Negro, Branco, Essequibo Roestes ogilviei Ctenoluciidae 5 Boulengerella cuvieri Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Javari, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, AraguariMacari-Amapá, Oiapoque, Essequibo, lower Orinoco, upper Orinoco, Apure Boulengerella lateristriga Japurá, Negro, Branco, upper Orinoco Boulengerella lucius Upper Xingu, Tapajós, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Boulengerella maculata Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, lower Orinoco, upper Orinoco Boulengerella xyrekes Tefé, Javari, Putumayo, Japurá, Negro, Branco, Trombetas, lower Orinoco, upper Orinoco, Apure Erythrinidae 5 Erythrinus erythrinus* Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Jutaí, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Hoplerythrinus unitaeniatus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, Capim, Araguari-Macari-Amapá, Oiapoque, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay 98 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Hoplias aimara Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Urubu-Uatumã, Trombetas, Jari, AraguariMacari-Amapá, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco Hoplias curupira Upper Tocantins, lower Tocantins, upper Xingu, lower Xingu, Tapajós, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Capim, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Hoplias malabaricus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, Capim, Maroni-Approuague, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Maracaibo, Cauca-Magdalena-Sinu, Atrato, Paraná-Paraguay Gasteropelecidae 8 Carnegiella marthae Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Putumayo, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Carnegiella myersi Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Juruá, Ucayali, Marañon-Nanay, Putumayo, Amazonas main channel, lower Orinoco Carnegiella schereri* Endemic – Napo-Ambyiacu Carnegiella strigata* Lower Tocantins, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Juruá, Javari, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Capim, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Gasteropelecus levis* Amazonas main channel, Amazonas Estuary Gasteropelecus sternicla Lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Putumayo, Japurá, Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco Thoracocharax securis* Mamoré, Guaporé, Purus, Juruá, Putumayo, Amazonas main channel Thoracocharax stellatus Upper Tocantins, Araguaia, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Negro, Amazonas main channel, lower Orinoco, upper Orinoco, Apure, ParanáParaguay Hemiodontidae 26 Anodus elongatus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Putumayo, Japurá, Negro, Branco, UrubuUatumã, Trombetas, Amazonas main channel Anodus orinocensis Araguaia, lower Tocantins, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Argonectes longiceps Tapajós, middle-lower Madeira, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Maroni-Approuague, Essequibo, lower Orinoco, upper Orinoco, Apure Argonectes robertsi Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Capim Bivibranchia fowleri Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Purus, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 99 Occurrence Bivibranchia notata* Lower Tocantins, Teles Pires, Tapajós, Trombetas Bivibranchia velox* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu Hemiodus amazonum Teles Pires, middle-lower Madeira, Madeira Shield Tributaries, Marañon-Nanay, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Hemiodus argenteus Upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, Japurá, Negro, Branco, Amazonas main channel, Itapicuru-Mearim, Capim, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure Hemiodus atranalis* Middle-lower Madeira, Ucayali, Negro, Branco, Amazonas main channel Hemiodus goeldii Araguaia, lower Tocantins, lower Xingu, Putumayo, Negro, Branco, AraguariMacari-Amapá Hemiodus gracilis Araguaia, Tapajós, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Ucayali, Putumayo, Japurá, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Hemiodus immaculatus Upper Xingu, lower Xingu, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Purus, Jutaí, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Hemiodus iratapuru* Endemic – Jari Hemiodus jatuarana* Endemic – Trombetas Hemiodus langeanii* Endemic – Amazonas main channel Hemiodus microlepis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Teles Pires, Juruena, Jamanxim, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Hemiodus quadrimaculatus Teles Pires, Branco, Trombetas, Araguari-Macari-Amapá, Oiapoque, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco Hemiodus semitaeniatus Araguaia, upper Xingu, Iriri, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Negro, Branco, Urubu-Uatumã, Amazonas main channel, Amazonas Estuary, Capim, Essequibo, lower Orinoco, upper Orinoco, Paraná-Paraguay Hemiodus sterni* Restricted to Tapajós basin (Teles Pires, Juruena, Jamanxim) Hemiodus ternetzi Upper Tocantins, lower Tocantins, upper Xingu, lower Orinoco Hemiodus thayeria Negro, Amazonas main channel, Essequibo, upper Orinoco Hemiodus tocantinensis* Upper Tocantins, Araguaia, lower Tocantins, Iriri, lower Xingu Hemiodus unimaculatus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Hemiodus vorderwinkleri Upper Xingu, Iriri, lower Xingu, Amazonas main channel, lower Orinoco Negro, Urubu-Uatumã, Trombetas, Amazonas main channel Micromischodus sugillatus* Lebiasinidae 49 Copeina guttata* Lower Xingu, Juruá, Putumayo, Trombetas, Amazonas main channel, Amazonas Estuary Copeina osgoodi* Putumayo, Amazonas main channel 100 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Copella arnoldi* Jari, Amazonas main channel, Amazonas Estuary, Capim, Oiapoque, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, Atl. Coastal Drainages of Col. Ven. Copella callolepis* Middle-lower Madeira, Purus, Ucayali, Marañon-Nanay, Putumayo, Trombetas, Jari, Amazonas main channel, Amazonas Estuary Copella compta* Endemic – Negro Copella eigenmanni Putumayo, Japurá, Negro, lower Orinoco, upper Orinoco, Apure, Atl. Coastal Drainages of Col. Ven. Copella nattereri Tapajós, middle-lower Madeira, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, upper Orinoco Copella vilmae* Endemic – Amazonas main channel Lebiasina bimaculata* Endemic – Marañon Lebiasina elongata* Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá Lebiasina erythrinoides Japurá, upper Orinoco, Apure, Maracaibo Lebiasina intermedia** Lebiasina marilynae* Endemic – Iriri Lebiasina melanoguttata* Iriri, Teles Pires Lebiasina minuta* Endemic – Iriri Lebiasina yepezi Negro, Branco, upper Orinoco Nannostomus beckfordi Tapajós, Amazonas main channel, Gurupi-Turiaçu, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Nannostomus bifasciatus Branco, Maroni-Approuague, Coppename-Suriname-Saramacca Nannostomus britskii* Tapajós, Jamanxim, Madeira Shield Tributaries, Urubu-Uatumã, Amazonas main channel Nannostomus digrammus Tapajós, Mamoré, Guaporé, middle-lower Madeira, Purus, Negro, Branco, UrubuUatumã, Trombetas, Amazonas main channel, upper Orinoco Nannostomus eques Lower Tocantins, lower Xingu, Tapajós, middle-lower Madeira, Purus, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure Nannostomus grandis** Nannostomus harrisoni Guaporé, Putumayo, Branco, Trombetas, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Nannostomus limatus* Endemic – Amazonas main channel Nannostomus marginatus Purus, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Nannostomus marilynae Madeira Shield Tributaries, Negro, upper Orinoco Nannostomus mortenthaleri* Endemic – Marañon Nannostomus nitidus Amazonas Estuary, Capim Nannostomus rubrocaudatus** Nannostomus trifasciatus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Amazonas main channel, Capim, Essequibo, upper Orinoco 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 101 Occurrence Nannostomus unifasciatus Lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Pyrrhulina australis Araguaia, upper Xingu, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Jari, Amazonas main channel, Paraná-Paraguay Pyrrhulina beni* Beni-Madre de Dios Pyrrhulina brevis Mamoré, Guaporé, Beni-Madre de Dios, Putumayo, Negro, Urubu-Uatumã, Amazonas main channel, lower Orinoco, upper Orinoco Pyrrhulina eleanorae Ucayali, Napo-Ambyiacu, upper Orinoco Pyrrhulina elongata* Tapajós Pyrrhulina filamentosa Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco Pyrrhulina laeta* Putumayo, Amazonas main channel Pyrrhulina lugubris Japurá, lower Orinoco, upper Orinoco, Apure Pyrrhulina marilynae* Iriri, Teles Pires, Juruena, Tapajós Pyrrhulina maxima* Amazonas main channel Pyrrhulina melanostomus* Mamoré, Beni-Madre de Dios Pyrrhulina obermulleri* Amazonas main channel Pyrrhulina rachoviana** Pyrrhulina semifasciata* Purus, Juruá, Negro, Branco, Amazonas main channel, Amazonas Estuary Pyrrhulina spilota* Mamoré, Guaporé Pyrrhulina stoli Japurá, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, upper Orinoco Pyrrhulina vittata* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel Ucayali Pyrrhulina zigzag* Parodontidae 10 Apareiodon argenteus* Restricted to Tocantins basin (upper Tocantins, Araguaia) Apareiodon cavalcante* Endemic – upper Tocantins Apareiodon machrisi Upper Tocantins, Araguaia, lower Tocantins, Parnaíba Apareiodon tigrinus* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Parodon bifasciatus Branco, Essequibo Parodon buckleyi* Teles Pires, Juruena, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Jari Parodon carrikeri Mamoré, Beni-Madre de Dios, Paraná-Paraguay Parodon guyanensis Curuá-Paru do Oeste, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco Parodon nasus Beni-Madre de Dios, Branco, Paraná-Paraguay Parodon pongoensis* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Juruá, Ucayali, MarañonNanay, Napo-Ambyiacu, Japurá, Amazonas main channel Prochilodontidae Prochilodus britskii* 8 Endemic – Juruena 102 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Prochilodus mariae Negro, lower Orinoco, upper Orinoco, Apure Prochilodus nigricans* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Trombetas, Jari, Amazonas main channel, Amazonas Estuary Prochilodus rubrotaeniatus* Japurá, Negro, Branco, Trombetas, Amazonas main channel, Oiapoque, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Semaprochilodus brama* Araguaia, lower Tocantins, upper Xingu, lower Xingu Semaprochilodus insignis Lower Xingu, Juruena, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Purus, Coari-Urucu, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Essequibo, upper Orinoco Semaprochilodus kneri Japurá, lower Orinoco, upper Orinoco, Apure Semaprochilodus taeniurus* Middle-lower Madeira, Madeira Shield Tributaries, Japurá, Negro, Branco, Trombetas, Amazonas main channel Serrasalmidae 72 Acnodon normani* Upper Tocantins, lower Tocantins, upper Xingu, Iriri, lower Xingu Acnodon oligacanthus Upper Tocantins, lower Tocantins, Maroni-Approuague, Coppename-SurinameSaramacca Acnodon senai* Endemic – Jari Catoprion mento Araguaia, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Colossoma macropomum Lower Tocantins, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Japurá, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Metynnis altidorsalis* Tapajós, middle-lower Madeira, Juruá, Ucayali, Marañon-Nanay, Negro, Branco, Trombetas, Amazonas main channel, Maroni-Approuague, Coppename-SurinameSaramacca, Essequibo, lower Orinoco, upper Orinoco, Apure Metynnis anisurus* Araguaia, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Jari Metynnis cuiaba Araguaia, Mamoré, Guaporé, middle-lower Madeira, Capim Metynnis fasciatus Araguaia, lower Tocantins, lower Xingu, Tapajós, Amazonas main channel, Capim Metynnis guaporensis Araguaia, lower Tocantins, upper Xingu, lower Xingu, Mamoré, Guaporé, BeniMadre de Dios, middle-lower Madeira, Juruá, Marañon-Nanay, Negro, UrubuUatumã, Amazonas main channel, Capim, lower Orinoco, upper Orinoco, Apure Metynnis hypsauchen Teles Pires, Tapajós, middle-lower Madeira, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Metynnis lippincottianus Araguaia, lower Tocantins, Teles Pires, Mamoré, Guaporé, middle-lower Madeira, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, ItapicuruMearim, lower Orinoco, Paraná-Paraguay Metynnis longipinnis Purus, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco Metynnis luna Araguaia, lower Tocantins, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Juruá, Putumayo, Japurá, Branco, Essequibo, lower Orinoco, upper Orinoco, Apure 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 103 Occurrence Metynnis maculatus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, MarañonNanay, Putumayo, Negro, Branco, Trombetas, Essequibo, lower Orinoco, upper Orinoco, Paraná-Paraguay Metynnis melanogrammus Madeira Shield Tributaries, Negro, Branco, Urubu-Uatumã, Trombetas, upper Orinoco Mylesinus paraschomburgkii Urubu-Uatumã, Trombetas, Jari, Araguari-Macari-Amapá Mylesinus paucisquamatus* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Myleus pachyodus* Restricted to Tapajós basin (Teles Pires, Jamanxim) Myleus rhomboidalis Upper Xingu, Putumayo, Branco, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, upper Orinoco Myleus setiger Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Myleus torquatus Upper Tocantins, Araguaia, lower Tocantins, middle-lower Madeira, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Myloplus arnoldi* Araguaia, upper Xingu, Teles Pires, Branco Myloplus asterias Upper Tocantins, lower Tocantins, Teles Pires, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Japurá, Branco, Parnaíba, lower Orinoco, upper Orinoco Myloplus lobatus* Mamoré, middle-lower Madeira, Branco Myloplus lucienae* Endemic – Negro Myloplus rubripinnis Jari, Araguari-Macari-Amapá, Oiapoque, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo Myloplus schomburgkii Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Madeira Shield Tributaries, Marañon-Nanay, Putumayo, Japurá, Negro, Trombetas, lower Orinoco, upper Orinoco, Apure Myloplus zorroi* Endemic – Madeira Shield Tributaries Mylossoma albiscopum Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco Mylossoma aureum* Mamoré, middle-lower Madeira, Purus, Juruá, Amazonas main channel Mylossoma unimaculatum* Restricted to Tocantins basin (Araguaia, lower Tocantins) Ossubtus xinguense* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Piaractus brachypomus Upper Tocantins, Araguaia, lower Tocantins, lower Xingu, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Putumayo, Japurá, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Pristobrycon aureus Upper Xingu, Putumayo, Japurá, Corentyne-Demerara, Essequibo Pristobrycon calmoni Lower Tocantins, Putumayo, Japurá, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Pristobrycon careospinus Putumayo, Japurá, upper Orinoco Pristobrycon maculipinnis Japurá, upper Orinoco Pristobrycon striolatus Upper Xingu, Madeira Shield Tributaries, Putumayo, Japurá, Negro, Branco, Trombetas, Maroni-Approuague, Essequibo, lower Orinoco, upper Orinoco, Apure 104 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Pygocentrus nattereri Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Putumayo, Japurá, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, Essequibo, Paraná-Paraguay Pygopristis denticulata Lower Tocantins, Guaporé, Branco, Trombetas, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, Upper Orinoco, Apure Serrasalmus altispinis* Endemic – Branco Serrasalmus altuvei Japurá, Negro, Branco, lower Orinoco, upper Orinoco, Apure Serrasalmus compressus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Amazonas main channel Serrasalmus eigenmanni Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Mamoré, Guaporé, BeniMadre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Corentyne-Demerara, Essequibo Serrasalmus elongatus Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Serrasalmus geryi* Restricted to Tocantins basin (Araguaia, lower Tocantins) Serrasalmus gibbus* Araguaia, lower Tocantins, Trombetas Serrasalmus gouldingi Putumayo, Negro, Branco, upper Orinoco Serrasalmus hastatus* Negro, Branco, Amazonas main channel Serrasalmus hollandi* Upper Xingu, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Negro, Trombetas, Amazonas main channel Serrasalmus humeralis* Araguaia, upper Xingu, Iriri, lower Xingu, Madeira Shield Tributaries, Putumayo, Japurá, Amazonas main channel Serrasalmus irritans Putumayo, lower Orinoco, upper Orinoco, Apure Serrasalmus maculatus Araguaia, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Urubu-Uatumã, Amazonas main channel, Araguari-Macari-Amapá, Oiapoque, Paraná-Paraguay Serrasalmus manueli Upper Xingu, lower Xingu, Teles Pires, middle-lower Madeira, Madeira Shield Tributaries, Negro, Branco, Urubu-Uatumã, upper Orinoco Serrasalmus marginatus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Paraná-Paraguay Serrasalmus medinai Japurá, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Serrasalmus nigricans** Serrasalmus odyssei* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Serrasalmus rhombeus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Ucayali, Putumayo, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Parnaíba, ItapicuruMearim, Araguari-Macari-Amapá, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Serrasalmus sanchezi* Ucayali, Japurá Serrasalmus serrulatus Lower Tocantins, upper Xingu, lower Xingu, Juruá, Branco, Corentyne-Demerara, Essequibo Serrasalmus spilopleura Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 105 Occurrence Tometes ancylorhynchus* Araguaia, lower Tocantins, lower Xingu, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Putumayo, Japurá, Trombetas, Amazonas main channel, Paraná-Paraguay Tometes camunani* Endemic – Trombetas Tometes kranponhah* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Tometes makue Negro, lower Orinoco, upper Orinoco Tometes siderocarajensis* Endemic – lower Tocantins Tometes trilobatus Jari, Araguari-Macari-Amapá, Oiapoque Utiaritichthys esguiceroi* Endemic – Juruena Utiaritichthys longidorsalis* Endemic – Madeira Shield Tributaries Juruena, Guaporé, Paraná-Paraguay Utiaritichthys sennaebragai* Tarumaniidae 1 Endemic – Negro Tarumania walkerae* Siluriformes 956 Aspredinidae 23 Amaralia hypsiura Lower Tocantins, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, middle-lower Madeira, Napo-Ambyiacu, Japurá, Negro, Branco, Trombetas, Essequibo Aspredo aspredo Trombetas, Parnaíba, Itapicuru-Mearim, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, lower Orinoco Bunocephalus aleuropsis Upper Tocantins, Araguaia, upper Xingu, Teles Pires, Mamoré, Guaporé, BeniMadre de Dios, middle-lower Madeira, Juruá, Ucayali, Japurá, Negro, Amazonas main channel, upper Orinoco, Apure Bunocephalus amaurus* Branco, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Bunocephalus coracoideus Araguaia, lower Tocantins, upper Xingu, lower Xingu, Mamoré, Guaporé, BeniMadre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Ucayali, Marañon-Nanay, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Essequibo Bunocephalus knerii* Iriri, middle-lower Madeira, Marañon-Nanay, Napo-Ambyiacu, Japurá, Negro Bunocephalus verrucosus Tapajós, Purus, Tefé, Juruá, Ucayali, Putumayo, Branco, Amazonas main channel, Maroni-Approuague, Corentyne-Demerara, Essequibo, upper Orinoco Ernstichthys intonsus* Endemic – Napo-Ambyiacu Ernstichthys megistus* Endemic – Marañon Hoplomyzon atrizona Branco, Maracaibo Hoplomyzon papillatus Mamoré, Napo-Ambyiacu, Apure Micromyzon akamai* Middle-lower Madeira, Amazonas main channel, Amazonas Estuary Platystacus cotylephorus Upper Xingu, lower Xingu, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, lower Orinoco Pseudobunocephalus amazonicus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Amazonas main channel Pseudobunocephalus bifidus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Napo-Ambyiacu, Amazonas main channel Pseudobunocephalus iheringii Guaporé, Paraná-Paraguay 106 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Pseudobunocephalus quadriradiatus* Endemic – Ucayali Pterobunocephalus depressus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, NapoAmbyiacu, Negro, Trombetas, Amazonas main channel Pterobunocephalus dolichurus* Marañon-Nanay, Negro, Trombetas, Amazonas main channel Xyliphius anachoretes* Restricted to Tocantins basin (upper Tocantins, Araguaia) Xyliphius lepturus Beni-Madre de Dios, Purus, Ucayali, Marañon-Nanay, Napo-Ambyiacu, upper Orinoco, Apure Xyliphius melanopterus Beni-Madre de Dios, middle-lower Madeira, Purus, Marañon-Nanay, NapoAmbyiacu, Putumayo, upper Orinoco, Apure Endemic – Amazonas main channel Xyliphius sofiae* Astroblepidae 17 Astroblepus boulengeri** Astroblepus caquetae* Endemic – Japurá Astroblepus festae* Endemic – Marañon Astroblepus formosus* Endemic – Ucayali Astroblepus labialis* Endemic – Marañon Astroblepus longiceps* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Astroblepus longifilis Marañon-Nanay, Cauca-Magdalena-Sinu Astroblepus mancoi* Beni-Madre de Dios, Ucayali Astroblepus peruanus* Ucayali, Marañon-Nanay, Napo-Ambyiacu Astroblepus pholeter* Marañon-Nanay, Napo-Ambyiacu Astroblepus praeliorum* Endemic – Marañon Astroblepus prenadillus* Endemic – Marañon Astroblepus sabalo* Endemic – Ucayali Astroblepus supramollis* Ucayali, Marañon-Nanay Astroblepus taczanowskii* Endemic – Ucayali Astroblepus theresiae* Endemic – Marañon Astroblepus vanceae* Endemic – Ucayali Auchenipteridae 79 Ageneiosus akamai* Purus, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel Ageineiosus apiaka* Endemic – Teles Pires Ageneiosus dentatus Araguaia, lower Tocantins, Tapajós, Mamoré, Purus, Negro, Trombetas, Amazonas main channel, Amazonas Estuary, Parnaíba, Araguari-Macari-Amapá, MaroniApprouague, Corentyne-Demerara, Essequibo, lower Orinoco Ageneiosus inermis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Putumayo, Japurá, Negro, Branco, UrubuUatumã, Trombetas, Amazonas main channel, Parnaíba, Itapicuru-Mearim, AraguariMacari-Amapá, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Ageneiosus intrusus Beni-Madre de Dios, middle-lower Madeira, Purus, Negro, Trombetas, Amazonas main channel, Capim, Essequibo 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 107 Occurrence Ageneiosus lineatus Lower Tocantins, lower Xingu, middle-lower Madeira, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel, upper Orinoco Ageneiosus polystictus* Negro, Urubu-Uatumã, Trombetas, Amazonas main channel Ageneiosus ucayalensis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Itapicuru-Mearim, Capim, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Ageneiosus uranophthalmus* Mamoré, middle-lower Madeira, Purus, Ucayali, Negro, Urubu-Uatumã, Amazonas main channel Ageneiosus vittatus Lower Tocantins, upper Xingu, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Negro, Trombetas, Amazonas main channel, lower Orinoco, Apure Asterophysus batrachus Negro, Urubu-Uatumã, lower Orinoco, upper Orinoco, Apure Auchenipterichthys coracoideus Araguaia, lower Tocantins, Mamoré, middle-lower Madeira, Ucayali, MarañonNanay, Branco, Amazonas main channel, Essequibo Auchenipterichthys longimanus Araguaia, lower Tocantins, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Auchenipterichthys punctatus Negro, Branco, Urubu-Uatumã, Amazonas main channel, upper Orinoco Auchenipterichthys thoracatus Lower Tocantins, upper Xingu, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Amazonas main channel, upper Orinoco Auchenipterus ambyiacus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Javari, Marañon-Nanay, Napo-Ambyiacu, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Auchenipterus brachyurus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, UrubuUatumã, Amazonas main channel Auchenipterus britskii* Mamoré, middle-lower Madeira, Purus, Juruá, Amazonas main channel Auchenipterus demerarae Branco, Amazonas main channel, Corentyne-Demerara, Essequibo Auchenipterus fordicei* Endemic – Amazonas main channel Auchenipterus nigripinnis Mamoré, Guaporé, Beni-Madre de Dios, Paraná-Paraguay Auchenipterus nuchalis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Negro, Branco, Trombetas, Amazonas main channel, Capim, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Auchenipterus osteomystax Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Tapajós, Trombetas, Amazonas main channel, Paraná-Paraguay Centromochlus altae* Middle-lower Madeira, Japurá Centromochlus existimatus* Madeira Shield Tributaries, Ucayali, Japurá, Trombetas, Amazonas main channel, Amazonas Estuary Centromochlus heckelii Lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure Centromochlus macracanthus* Endemic – Negro 108 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Centromochlus meridionalis* Endemic – Teles Pires Centromochlus orca* Urubu-Uatumã, Trombetas Centromochlus perugiae Beni-Madre de Dios, middle-lower Madeira, Ucayali, Napo-Ambyiacu, Negro, Amazonas main channel, Paraná-Paraguay Centromochlus punctatus Putumayo, Curuá-Paru do Oeste, Maroni-Approuague, Coppename-SurinameSaramacca, Apure Centromochlus reticulatus Purus, Branco, Amazonas main channel, Essequibo Centromochlus schultzi Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Jari, Essequibo Centromochlus simplex* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu Entomocorus benjamini* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Entomocorus melaphareus* Endemic – Amazonas main channel Epapterus dispilurus Mamoré, Guaporé, middle-lower Madeira, Juruá, Ucayali, Putumayo, Amazonas main channel, Paraná-Paraguay Gelanoglanis nanonocticolus Negro, upper Orinoco Gelanoglanis pan* Endemic – Teles Pires Gelanoglanis travieso* Endemic – Marañon Gelanoglanis varii* Endemic – upper Tocantins Glanidium leopardum Branco, Maroni-Approuague, Essequibo Liosomadoras morrowi* Purus, Jutaí, Ucayali, Putumayo, Japurá, Amazonas main channel Liosomadoras oncinus Negro, Branco, upper Orinoco “Parauchenipterus” porosus Lower Xingu, Tapajós, Mamoré, middle-lower Madeira, Ucayali, Urubu-Uatumã, Jari, Amazonas main channel, Amazonas Estuary, Paraná-Paraguay Pseudauchenipterus nodosus Lower Tocantins, Amazonas Estuary, Itapicuru-Mearim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco Pseudepapterus cucuhyensis* Lower Xingu, Purus, Negro, Trombetas, Amazonas main channel Pseudepapterus hasemani* Middle-lower Madeira, Purus, Negro, Branco, Trombetas, Amazonas main channel Spinipterus acsi* Endemic – Marañon Tatia aulopygia Araguaia, lower Tocantins, upper Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, Apure Tatia brunnea Negro, Branco, Trombetas, Maroni-Approuague, Coppename-Suriname-Saramacca Tatia caxiuanensis* Endemic – Amazonas Estuary Tatia dunni* Purus, Napo-Ambyiacu, Japurá Tatia gyrina Middle-lower Madeira, Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara Tatia intermedia Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Guaporé, Negro, Branco, Trombetas, Jari, Amazonas main channel, Capim, Araguari-Macari-Amapá, Oiapoque, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo Tatia melanoleuca* Endemic – Teles Pires Tatia musaica Urubu-Uatumã, Amazonas main channel, lower Orinoco, upper Orinoco Tatia nigra* Negro, Trombetas 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 109 Occurrence Tatia strigata Negro, Amazonas main channel, upper Orinoco Tetranematichthys barthemi* Endemic – Jari Tetranematichthys quadrifilis Mamoré, Guaporé, middle-lower Madeira, Trombetas, lower Orinoco, upper Orinoco, Apure Tetranematichthys wallacei Lower Tocantins, lower Xingu, Tapajós, Ucayali, Negro, Urubu-Uatumã, Amazonas main channel, Capim, upper Orinoco Tocantinsia piresi* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Jari Trachelyichthys decaradiatus Amazonas main channel, Essequibo Trachelyichthys exilis* Endemic – Marañon Trachelyopterichthys taeniatus Guaporé, middle-lower Madeira, Purus, Javari, Negro, Branco, Urubu-Uatumã, Amazonas main channel, upper Orinoco Trachelyopterus brevibarbis** Trachelyopterus ceratophysus Upper Xingu, Guaporé, Beni-Madre de Dios, Negro, Branco, Trombetas, Essequibo Trachelyopterus coriaceus Araguaia, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, Jari, Amazonas main channel, Amazonas Estuary, Oiapoque, Paraná-Paraguay Trachelyopterus isacanthus** Trachelyopterus galeatus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Ucayali, Marañon-Nanay, NapoAmbyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, GurupiTuriaçu, Araguari-Macari-Amapá, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Trachycorystes menezesi* Endemic – Madeira Shield Tributaries Trachycorystes trachycorystes Araguaia, upper Xingu, middle-lower Madeira, Purus, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Tympanopleura atronasus* Mamoré, Guaporé, middle-lower Madeira, Purus, Juruá, Ucayali, Negro, Branco, Jari, Amazonas main channel Tympanopleura brevis* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, MarañonNanay, Putumayo, Negro, Amazonas main channel Tympanopleura cryptica* Middle-lower Madeira, Ucayali, Amazonas main channel Tympanopleura longipinna* Purus, Ucayali, Amazonas main channel Tympanopleura piperata Middle-lower Madeira, Jutaí, Japurá, Negro, Branco, Amazonas main channel, Essequibo Tympanopleura rondoni* Mamoré, Guaporé, middle-lower Madeira, Amazonas main channel Callichthyidae 133 Aspidoras albater* Restricted to Tocantins basin (upper Tocantins, Araguaia) Aspidoras belenos* Endemic – Araguaia Aspidoras brunneus* Endemic – Araguaia Aspidoras eurycephalus* Endemic – upper Tocantins Aspidoras gabrieli* Endemic – lower Tocantins Aspidoras marianae* Endemic – Iriri Aspidoras mephisto* Endemic – upper Tocantins 110 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Aspidoras microgalaeus* Endemic – upper Xingu Aspidoras pauciradiatus* Endemic – Negro Aspidoras poecilus* Upper Tocantins, Araguaia, upper Xingu Aspidoras velites* Endemic – Araguaia Callichthys callichthys Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Coari-Urucu, Juruá, Ucayali, MarañonNanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Paru, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, ParanáParaguay Callichthys serralabium Negro, upper Orinoco Corydoras acrensis* Endemic – Juruá Corydoras acutus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel Corydoras adolfoi* Endemic – Negro Corydoras aeneus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Maroni-Approuague, Coppename-Suriname-Saramacca, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Corydoras agassizii* Putumayo, Amazonas main channel Corydoras albolineatus* Endemic – Guaporé Corydoras amandajanea* Endemic – Negro Corydoras ambiacus* Napo-Ambyiacu, Amazonas main channel Corydoras amphibelus* Endemic – Napo-Ambyiacu Corydoras apiaka* Restricted to Tapajós basin (Teles Pires, Juruena) Corydoras araguaiaensis* Endemic – Araguaia Corydoras arcuatus* Putumayo, Japurá, Negro, Amazonas main channel Corydoras armatus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Marañon-Nanay Corydoras atropersonatus* Endemic – Marañon Corydoras aurofrenatus Mamoré, Paraná-Paraguay Corydoras baderi Curuá-Paru do Oeste, Maroni-Approuague, Corentyne-Demerara Corydoras benattii* Upper Xingu, lower Xingu, Teles Pires Corydoras bifasciatus* Endemic – Teles Pires Corydoras bilineatus* Endemic – Mamoré Corydoras blochi Branco, Amazonas main channel, Essequibo, lower Orinoco, Apure Corydoras bondi Branco, Corentyne-Demerara, Essequibo, upper Orinoco Corydoras brittoi* Endemic – Madeira Shield Tributaries Corydoras britskii Mamoré, Guaporé, Paraná-Paraguay Corydoras burgessi* Endemic – Negro Corydoras caudimaculatus* Restricted to Madeira basin (Guaporé, middle-lower Madeira) Corydoras cervinus* Restricted to Madeira basin (Guaporé, middle-lower Madeira) 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 111 Occurrence Corydoras cochui* Endemic – Araguaia Corydoras copei* Endemic – Marañon Corydoras coriatae* Endemic – Ucayali Corydoras crimmeni* Endemic – Branco Corydoras cruziensis* Restricted to Madeira basin (Mamoré, Guaporé) Corydoras crypticus* Endemic – Negro Corydoras davidsandsi* Endemic – Negro Corydoras desana* Endemic – Negro Corydoras duplicareus* Endemic – Negro Corydoras elegans* Mamoré, Putumayo, Japurá, Amazonas main channel Corydoras eques* Endemic – Amazonas main channel Corydoras evelynae* Endemic – Amazonas main channel Corydoras eversi* Endemic – Araguaia Corydoras fowleri* Endemic – Amazonas main channel Corydoras geryi* Restricted to Madeira basin (Mamoré, Guaporé) Corydoras gomezi* Endemic – Amazonas main channel Corydoras gossei* Endemic – Mamoré Corydoras gracilis* Endemic – Madeira Shield Tributaries Corydoras guapore* Endemic – Guaporé Corydoras haraldschultzi* Endemic – Guaporé Corydoras hastatus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel, Paraná-Paraguay Corydoras hephaestus* Juruena, Madeira Shield Tributaries Corydoras imitator* Endemic – Negro Corydoras incolicana* Endemic – Negro Corydoras isbrueckeri* Endemic – Guaporé Corydoras julii Lower Tocantins, Parnaíba Corydoras kanei* Restricted to Negro basin (Negro, Branco) Corydoras knaacki* Endemic – Beni-Madre de Dios Corydoras lamberti* Endemic – Marañon Corydoras latus* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Corydoras leopardus** Corydoras leucomelas* Marañon-Nanay, Putumayo, Japurá, Amazonas main channel Corydoras loretoensis* Endemic – Marañon Corydoras maculifer* Endemic – Araguaia Corydoras mamore* Endemic – Mamoré Corydoras melanistius Branco, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Corydoras melini Japurá, Negro, upper Orinoco 112 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Corydoras multiradiatus* Endemic – Napo-Ambyiacu Corydoras napoensis* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Napo-Ambyiacu, Amazonas main channel Corydoras narcissus* Endemic – Purus Corydoras negro* Endemic – Branco Corydoras nijsseni* Endemic – Negro Corydoras noelkempffi* Endemic – Guaporé Corydoras ornatus* Endemic – Tapajós Corydoras orphnopterus* Endemic – Marañon Corydoras osteocarus Branco, lower Orinoco, upper Orinoco, Apure Corydoras ourastigma* Endemic – Purus Corydoras panda* Endemic – Ucayali Corydoras pantanalensis* Restricted to Madeira basin (Mamoré, Guaporé) Corydoras paragua* Endemic – Guaporé Corydoras parallelus* Endemic – Negro Corydoras pastazensis* Marañon-Nanay, Putumayo, Amazonas main channel Corydoras paucerna* Endemic – Guaporé Corydoras pavanelliae* Endemic – Madeira Shield Tributaries Corydoras pinheiroi** Corydoras polystictus Guaporé, Paraná-Paraguay Corydoras potaroensis Negro, Branco, Essequibo Corydoras pulcher* Endemic – Purus Corydoras pygmaeus* Middle-lower Madeira, Amazonas main channel Corydoras rabauti* Javari, Putumayo, Negro, Amazonas main channel Corydoras reticulatus* Japurá, Amazonas main channel Corydoras reynoldsi* Endemic – Japurá Corydoras robineae* Endemic – Negro Corydoras robustus* Endemic – Purus Corydoras sarareensis* Endemic – Guaporé Corydoras schwartzi* Endemic – Purus Corydoras semiaquilus* Endemic – Amazonas main channel Corydoras serratus* Endemic – Negro Corydoras seussi* Endemic – Mamoré Corydoras similis* Endemic – Middle-lower Madeira Corydoras sipaliwini Branco, Maroni-Approuague, Corentyne-Demerara, Essequibo Corydoras sodalis* Endemic – Amazonas main channel Corydoras spectabilis* Endemic – Guaporé Corydoras splendens* Araguaia, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Putumayo, Japurá, Amazonas main channel 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 113 Occurrence Corydoras stenocephalus* Mamoré, Beni-Madre de Dios, Purus, Juruá, Ucayali Corydoras sterbai* Endemic – Guaporé Corydoras sychri* Endemic – Marañon Corydoras trilineatus* Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Putumayo, Japurá, Amazonas main channel Corydoras tukano* Endemic – Negro Corydoras urucu* Endemic – Coari-Urucu Corydoras virginiae* Endemic – Ucayali Corydoras weitzmani* Endemic – Ucayali Corydoras xinguensis* Restricted to Xingu basin (upper Xingu, lower Xingu) Corydoras zawadzkii* Endemic – Madeira Shield Tributaries Corydoras zygatus* Juruá, Marañon-Nanay, Amazonas main channel Hoplosternum littorale Araguaia, Juruena, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Putumayo, Japurá, Negro, Amazonas main channel, Amazonas Estuary, Parnaíba, Capim, Araguari-Macari-Amapá, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Atl. Coastal Drainages of Col. Ven., ParanáParaguay Dianema longibarbis* Lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Ucayali, Marañon-Nanay, Putumayo, Japurá, Jari, Amazonas main channel Dianema urostriatum* Middle-lower Madeira, Amazonas main channel Lepthoplosternum altamazonicum* Purus, Juruá, Ucayali Lepthoplosternum beni* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios, middle-lower Madeira) Lepthoplosternum stellatum* Endemic – Tefé Lepthoplosternum ucamara* Ucayali, Japurá, Amazonas main channel Megalechis picta Araguaia, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Coppename-Suriname-Saramacca, Essequibo, lower Orinoco, upper Orinoco Megalechis thoracata Araguaia, lower Tocantins, upper Xingu, lower Xingu, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Juruá, Ucayali, MarañonNanay, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Itapicuru-Mearim, Capim, Araguari-Macari-Amapá, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Cetopsidae 26 Cetopsidium ferreirai* Urubu-Uatumã, Trombetas Cetopsidium orientale Upper Tocantins, Araguaia, middle-lower Madeira, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Cetopsidium pemon Negro, Branco, lower Orinoco, upper Orinoco Cetopsidium roae Branco, Essequibo Cetopsidium soniae* Endemic – Branco Cetopsis arcana* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) 114 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Cetopsis caiapo* Endemic – upper Tocantins Cetopsis candiru* Upper Tocantins, Araguaia, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Amazonas main channel, Amazonas Estuary Cetopsis coecutiens Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Jari, Amazonas main channel, Amazonas Estuary, lower Orinoco, upper Orinoco, Apure Cetopsis montana* Marañon-Nanay, Napo-Ambyiacu Cetopsis oliveirai* Beni-Madre de Dios, middle-lower Madeira, Napo-Ambyiacu, Putumayo, Amazonas main channel Cetopsis parma* Middle-lower Madeira, Ucayali, Marañon-Nanay, Negro Cetopsis pearsoni* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Cetopsis plumbea Mamoré, Beni-Madre de Dios, middle-lower Madeira, Marañon-Nanay, NapoAmbyiacu, upper Orinoco Cetopsis sandrae* Restricted to Tapajós basin (Teles Pires, Juruena) Cetopsis sarcodes* Endemic – upper Tocantins Cetopsis starnesi Mamoré, Guaporé, Paraná-Paraguay Denticetopsis epa* Endemic – lower Tocantins Denticetopsis iwokrama Branco, Essequibo Denticeptopsis macilenta Negro, Branco, Essequibo, upper Orinoco, Apure Denticetopsis praecox* Negro, Amazonas main channel Denticetopsis royeroi* Endemic – Negro Denticetopsis sauli* Endemic – Negro Denticetopsis seducta* Middle-lower Madeira, Napo-Ambyiacu, Putumayo, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel Helogenes gouldingi* Endemic – Middle-lower Madeira Helogenes marmoratus Upper Xingu, Iriri, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Capim, Oiapoque, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, upper Orinoco Doradidae 73 Acanthodoras cataphractus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Japurá, Negro, Branco, Urubu-Uatumã, Jari, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Acanthodoras depressus* Endemic – Negro Acanthodoras spinosissimus Mamoré, Guaporé, middle-lower Madeira, Coari-Urucu, Juruá, Branco, Trombetas, Amazonas main channel, Essequibo, upper Orinoco Agamyxis albomaculatus Putumayo, Japurá, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Agamyxis pectinifrons* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Putumayo, Japurá, Amazonas main channel 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 115 Occurrence Amblydoras affinis* Araguaia, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Tefé, Ucayali, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Capim, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Amblydoras monitor** Amazonas main channel Amblydoras nauticus** Anadoras grypus* Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, NapoAmbyiacu, Putumayo, Amazonas main channel Anadoras regani** Japurá, Negro, Amazonas main channel Anadoras weddellii Araguaia, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Araguari-MacariAmapá, Oiapoque, Paraná-Paraguay Anduzedoras oxyrhynchus Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco Astrodoras asterifrons* Lower Tocantins, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Jutaí, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel Centrodoras brachiatus* Lower Xingu, middle-lower Madeira, Amazonas main channel, Amazonas Estuary Centrodoras hasemani* Endemic – Negro Doras carinatus Lower Xingu, Trombetas, Oiapoque, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco Doras higuchii* Upper Xingu, Iriri, lower Xingu, Trombetas, Jari Doras phlyzakion* Purus, Juruá, Japurá, Negro, Branco, Amazonas main channel Doras zuanoni* Endemic – Araguaia Hassar gabiru* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Hassar orestis Araguaia, lower Xingu, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Negro, Branco, Trombetas, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Hassar shewellkeimi* Restricted to Tapajós basin (Teles Pires, Juruena) Hassar wilderi* Upper Tocantins, Araguaia, lower Tocantins, Branco Hemidoras morei* Mamoré, Guaporé, Purus, Tefé, Negro, Amazonas main channel Hemidoras morrisi** Middle-lower Madeira, Putumayo, Amazonas main channel Hemidoras stenopeltis* Mamoré, Guaporé, middle-lower Madeira, Purus, Japurá, Negro, Branco, Amazonas main channel Hypodoras forficulatus* Marañon-Nanay, Putumayo, Amazonas main channel Leptodoras acipenserinus* Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Marañon-Nanay, Napo-Ambyiacu Leptodoras cataniai* Upper Tocantins, lower Tocantins, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Ucayali, Napo-Ambyiacu, Negro, Branco, Amazonas main channel Leptodoras copei Negro, Amazonas main channel, lower Orinoco, upper Orinoco Leptodoras hasemani Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, Jamanxim, Negro, Branco, Essequibo, lower Orinoco, upper Orinoco Leptodoras juruensis* Middle-lower Madeira, Putumayo, Amazonas main channel 116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Leptodoras linnelli Mamoré, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Negro, Branco, Urubu-Uatumã, Araguari-Macari-Amapá, Corentyne-Demerara, Essequibo, upper Orinoco Leptodoras marki* Endemic – upper Xingu Leptodoras myersi* Mamoré, middle-lower Madeira, Amazonas main channel Leptodoras oyakawai* Upper Xingu, lower Xingu, Teles Pires, Jamanxim Leptodoras praelongus Upper Tocantins, lower Xingu, Teles Pires, Mamoré, middle-lower Madeira, Negro, Branco, Jari, Amazonas main channel, upper Orinoco Lithodoras dorsalis Purus, Amazonas main channel, Amazonas Estuary, Capim Megalodoras uranoscopus Upper Tocantins, Araguaia, lower Tocantins, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Japurá, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, Apure Nemadoras cristinae Beni-Madre de Dios, middle-lower Madeira, Purus, Marañon-Nanay, Amazonas main channel, upper Orinoco Nemadoras elongatus* Middle-lower Madeira, Purus, Tefé, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Amazonas main channel Nemadoras hemipeltis* Middle-lower Madeira, Ucayali, Urubu-Uatumã, Amazonas main channel Nemadoras humeralis* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Amazonas main channel Opsodoras boulengeri* Mamoré, Guaporé, middle-lower Madeira, Trombetas, Amazonas main channel Opsodoras stuebelii* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Marañon-Nanay, Japurá, Negro, Amazonas main channel Ossancora asterophysa* Tapajós, Mamoré, Guaporé, middle-lower Madeira, Ucayali, Marañon-Nanay, Negro, Trombetas, Amazonas main channel, Amazonas Estuary Ossancora fimbriata* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Tefé, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel Ossancora punctata Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Marañon-Nanay, Amazonas main channel, Paraná-Paraguay Oxydoras niger Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Ucayali, Putumayo, Japurá, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Physopyxis ananas Anapu, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Jutaí, Marañon-Nanay, Negro, Branco, Trombetas, Essequibo, upper Orinoco Physopyxis cristata* Restricted to Negro basin (Negro, Branco) Physopyxis lyra* Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Negro, Urubu-Uatumã, Amazonas main channel Platydoras armatulus Araguaia, lower Tocantins, lower Xingu, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Japurá, Urubu-Uatumã, Amazonas main channel, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Platydoras birindelli* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Platydoras hancockii Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Negro, Branco, Corentyne-Demerara, Essequibo, upper Orinoco Pterodoras granulosus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Juruá, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Amazonas main channel, Paraná-Paraguay 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 117 Occurrence Pterodoras rivasi Amazonas main channel, lower Orinoco, upper Orinoco, Apure Rhinodoras armbrusteri Branco, Essequibo Rhinodoras boehlkei* Upper Tocantins, Araguaia, lower Tocantins, Teles Pires, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Marañon-Nanay, Negro, Amazonas main channel Rhinodoras dorbignyi Upper Tocantins, lower Tocantins, Mamoré, Paraná-Paraguay Rhynchodoras woodsi Mamoré, Ucayali, Marañon-Nanay, Negro, Branco, Amazonas main channel, Essequibo Rhynchodoras xingui* Lower Tocantins, upper Xingu, lower Xingu Scorpiodoras heckelii Tapajós, middle-lower Madeira, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco Scorpiodoras liophysus* Endemic – Middle-lower Madeira Tenellus leporhinus Upper Tocantins, Araguaia, lower Tocantins, Teles Pires, Jamanxim, middle-lower Madeira, Branco, Trombetas, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Tenellus ternetzi Araguaia, lower Tocantins, lower Xingu, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Juruá, Ucayali, Marañon-Nanay, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, lower Orinoco, upper Orinoco Tenellus trimaculatus Upper Tocantins, lower Tocantins, Teles Pires, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Trachydoras brevis Lower Xingu, Tapajós, Mamoré, middle-lower Madeira, Purus, Negro, Branco, Trombetas, Jari, Essequibo Trachydoras gepharti Upper Xingu, Jamanxim, Jutaí, Negro, Branco, Amazonas main channel, lower Orinoco, upper Orinoco Trachydoras microstomus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Negro, Branco, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Trachydoras nattereri Mamoré, Guaporé, middle-lower Madeira, Tefé, Ucayali, Negro, Branco, UrubuUatumã, Trombetas, Amazonas main channel, Essequibo Trachydoras paraguayensis Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Paraná-Paraguay Trachydoras steindachneri* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Trombetas, Amazonas main channel Heptapteridae 87 Brachyglanis frenata Negro, Urubu-Uatumã, Trombetas, Essequibo, upper Orinoco Brachyglanis melas Lower Tocantins, Tapajós, Negro, Branco, Essequibo Brachyglanis microphthalmus* Endemic – Trombetas Brachyglanis nocturna* Endemic – Negro Brachyrhamdia heteropleura Branco, Corentyne-Demerara, Essequibo Brachyrhamdia marthae* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Juruá, Ucayali Brachyrhamdia meesi* Endemic – Amazonas main channel Brachyrhamdia rambarrani* Endemic – Negro Brachyrhamdia thayeria* Endemic – Japurá Cetopsorhamdia filamentosa* Endemic – Ucayali Cetopsorhamdia iheringi Araguaia, Paraná-Paraguay, São Francisco Cetopsorhamdia insidiosa* Lower Tocantins, Iriri, lower Xingu, Negro, Branco 118 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Cetopsorhamdia molinae Upper Tocantins, Beni-Madre de Dios, Apure, Cauca-Magdalena-Sinu Cetopsorhamdia orinoco Napo-Ambyiacu, lower Orinoco, upper Orinoco, Cauca-Magdalena-Sinu Cetopsorhamdia phantasia* Beni-Madre de Dios, middle-lower Madeira, Ucayali, Napo-Ambyiacu, Putumayo Chasmocranus longior Upper Tocantins, upper Xingu, lower Xingu, Beni-Madre de Dios, Negro, Branco, Urubu-Uatumã, Trombetas, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, upper Orinoco Chasmocranus peruanus* Endemic – Marañon Chasmocranus quadrizonatus* Endemic – Marañon Gladioglanis anacanthus* Endemic – Madeira Shield Tributaries Gladioglanis conquistador Middle-lower Madeira, Madeira Shield Tributaries, Purus, Napo-Ambyiacu, Putumayo, Negro, Branco, Amazonas main channel, Capim Gladioglanis machadoi Negro, lower Orinoco, upper Orinoco Goeldiella eques Madeira Shield Tributaries, Juruá, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Horiomyzon retropinnatus* Middle-lower Madeira, Napo-Ambyiacu Imparfinis borodini Upper Tocantins, Paraná-Paraguay Imparfinis cochabambae* Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Ucayali Imparfinis guttatus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Paraná-Paraguay Imparfinis hasemani Negro, Branco, Jari, Amazonas main channel, Maroni-Approuague, Essequibo Imparfinis longicaudus* Endemic – Marañon Imparfinis mirini Araguaia, Paraná-Paraguay Imparfinis nemacheir Amazonas main channel, Apure, Maracaibo, Cauca-Magdalena-Sinu, Atrato Imparfinis pristos Upper Xingu, Negro, Amazonas main channel, upper Orinoco Imparfinis pseudonemacheir Ucayali, lower Orinoco, upper Orinoco Imparfinis stictonotus Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, NapoAmbyiacu, Paraná-Paraguay Leptorhamdia essequibensis Lower Tocantins, Negro, Branco, Urubu-Uatumã, Essequibo Leptorhamdia marmorata Negro, upper Orinoco, Apure Leptorhamdia schultzi* Upper Xingu, lower Xingu, Teles Pires Mastiglanis asopos Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Tapajós, Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Purus, Negro, Branco, UrubuUatumã, Amazonas main channel, Capim, Essequibo, lower Orinoco, upper Orinoco, Apure Myoglanis koepckei* Middle-lower Madeira, Ucayali, Marañon-Nanay, Negro, Amazonas main channel Nannoglanis fasciatus* Napo-Ambyiacu, Japurá Nemuroglanis furcatus* Middle-lower Madeira, Madeira Shield Tributaries Nemuroglanis lanceolatus* Ucayali, Napo-Ambyiacu, Amazonas main channel Nemuroglanis pauciradiatus Negro, Branco, Urubu-Uatumã, Trombetas, upper Orinoco Pariolius armillatus* Endemic – Napo-Ambyiacu Phenacorhamdia boliviana* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Marañon-Nanay Phenacorhamdia nigrolineata* Beni-Madre de Dios, Purus, Ucayali 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 119 Occurrence Phenacorhamdia somnians* Endemic – Araguaia Phreatobius cisternarum Amazonas Estuary, Araguari-Macari-Amapá Phreatobius dracunculus* Restricted to Madeira basin (Guaporé, middle-lower Madeira) Phreatobius sanguijuela* Endemic – Guaporé Pimelodella altipinnis Upper Xingu, Amazonas main channel, Essequibo Pimelodella bockmanni* Endemic – Madeira Shield Tributaries Pimelodella boliviana* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Pimelodella breviceps* Endemic – Negro Pimelodella buckleyi* Mamoré, Beni-Madre de Dios, Marañon-Nanay, Napo-Ambyiacu Pimelodella chaparae* Endemic – Mamoré Pimelodella conquetaensis* Endemic – Japurá Pimelodella cristata Lower Tocantins, Iriri, lower Xingu, Putumayo, Japurá, Branco, Trombetas, Jari, Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Pimelodella cyanostigma* Endemic – Napo-Ambyiacu Pimelodella geryi Japurá, Amazonas main channel, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara Pimelodella gracilis Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Putumayo, lower Orinoco, upper Orinoco, Paraná-Paraguay Pimelodella griffini Mamoré, Beni-Madre de Dios, Paraná-Paraguay Pimelodella hartwelli* Endemic – Ucayali Pimelodella hasemani* Mamoré, Beni-Madre de Dios, middle-lower Madeira, Putumayo Pimelodella howesi* Restricted to Madeira basin (Mamoré, Guaporé, middle-lower Madeira) Pimelodella humeralis* Endemic – Jari Pimelodella leptosoma Branco, Essequibo Pimelodella megalops Branco, Maroni-Approuague, Essequibo, lower Orinoco Pimelodella montana* Endemic – Marañon Pimelodella mucosa Mamoré, Guaporé, Beni-Madre de Dios, Paraná-Paraguay Pimelodella nigrofasciata* Endemic – Beni-Madre de Dios Pimelodella ophthalmica* Endemic – Marañon Pimelodella peruana* Endemic – Ucayali Pimelodella peruensis** Pimelodella roccae* Mamoré, Beni-Madre de Dios, Ucayali Pimelodella serrata* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Pimelodella spelaea* Endemic – upper Tocantins Pimelodella steindachneri* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel Rhamdella montana* Endemic – Ucayali Rhamdella rusbyi* Endemic – Beni-Madre de Dios 120 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Rhamdia foina Lower Tocantins, Negro, Branco, Trombetas, Maroni-Approuague, Essequibo Rhamdia humilis Japurá, Atl. Coastal Drainages of Col. Ven. Rhamdia itacaiunas* Lower Tocantins, upper Xingu Rhamdia laukidi Juruena, middle-lower Madeira, Negro, Branco, Urubu-Uatumã, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Rhamdia muelleri Upper Tocantins, lower Tocantins, middle-lower Madeira, Madeira Shield Tributaries, Napo-Ambyiacu, Negro, Trombetas, Capim, Essequibo, lower Orinoco, upper Orinoco Rhamdia parvus* Endemic – Marañon Rhamdia poeyi* Upper Tocantins, Mamoré, Napo-Ambyiacu Rhamdia quelen Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, Capim, Araguari-Macari-Amapá, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Maracaibo, Atl. Coastal Drainages of Col. Ven., Cauca-Magdalena-Sinu, Atrato, Paraná-Paraguay Pimelodidae 60 Aguarunichthys inpai* Middle-lower Madeira, Amazonas main channel Aguarunichthys tocantinsensis* Restricted to Tocantins basin (upper Tocantins, lower Tocantins) Aguarunichthys torosus* Beni-Madre de Dios, middle-lower Madeira, Marañon-Nanay Brachyplatystoma capapretum Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Negro, Urubu-Uatumã, Amazonas main channel, Amazonas Estuary, Capim Brachyplatystoma filamentosum Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Parnaíba, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Brachyplatystoma juruense Middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Japurá, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Brachyplatystoma platynemum Lower Tocantins, upper Xingu, lower Xingu, Mamoré, Beni-Madre de Dios, middlelower Madeira, Putumayo, Japurá, Branco, Amazonas main channel, Amazonas Estuary, lower Orinoco, upper Orinoco, Apure Brachyplatystoma rousseauxii Lower Tocantins, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Japurá, Branco, Amazonas main channel, CoppenameSuriname-Saramacca, Corentyne-Demerara, lower Orinoco, upper Orinoco, Apure Brachyplatystoma tigrinum* Lower Xingu, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Japurá, Amazonas main channel Brachyplatystoma vaillantii Lower Tocantins, lower Xingu, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Japurá, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Calophysus macropterus Upper Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Putumayo, Japurá, Branco, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Cheirocerus eques* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Napo-Ambyiacu, Putumayo, Amazonas main channel Cheirocerus goeldii* Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Branco, Amazonas main channel 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 121 Occurrence Duopalatinus peruanus Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Trombetas, Amazonas main channel, upper Orinoco, Apure Exallodontus aguanai Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Amazonas main channel, Amazonas Estuary, lower Orinoco, upper Orinoco, Apure Hemisorubim platyrhynchos Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Putumayo, Japurá, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Parnaíba, Itapicuru-Mearim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, ParanáParaguay Hypophthalmus edentatus* Lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Hypophthalmus fimbriatus Lower Xingu, Mamoré, middle-lower Madeira, Japurá, Negro, Trombetas, Amazonas main channel, lower Orinoco Hypophthalmus marginatus* Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Purus, Ucayali, Marañon-Nanay, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, lower Orinoco, upper Orinoco, Apure Hypophthalmus oremaculatus Middle-lower Madeira, Purus, Juruá, Ucayali, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Leiarius marmoratus Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Japurá, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Leiarius pictus Teles Pires, Tapajós, Beni-Madre de Dios, middle-lower Madeira, Purus, Putumayo, Japurá, Urubu-Uatumã, Trombetas, Amazonas main channel, upper Orinoco Megalonema platanum Mamoré, Guaporé, middle-lower Madeira, Paraná-Paraguay Megalonema platycephalum Araguaia, lower Tocantins, lower Xingu, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Megalonema E. amaxanthum Upper Tocantins, lower Tocantins, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Jutaí, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Japurá, Negro, Branco, Amazonas main channel, Essequibo Phractocephalus hemioliopterus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Coari-Urucu, Tefé, Juruá, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Pimelodina flavipinnis Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Putumayo, Japurá, Negro, Trombetas, Amazonas main channel, Capim, lower Orinoco, upper Orinoco Pimelodus albofasciatus Araguaia, lower Xingu, Teles Pires, middle-lower Madeira, Juruá, Negro, Branco, Trombetas, Jari, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Pimelodus altissimus Mamoré, Beni-Madre de Dios, middle-lower Madeira, Juruá, Ucayali, Amazonas main channel, Amazonas Estuary, lower Orinoco, Apure 122 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Pimelodus blochii Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Ucayali, Putumayo, Japurá, Branco, Trombetas, Amazonas main channel, Parnaíba, Itapicuru-Mearim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Cauca-Magdalena-Sinu Pimelodus halisodous* Endemic – upper Tocantins Pimelodus jivaro* Endemic – Marañon Pimelodus joannis* Endemic – upper Tocantins Pimelodus luciae* Endemic – lower Tocantins Pimelodus maculatus Beni-Madre de Dios, Purus, Juruá, Ucayali, Putumayo, Japurá, Parnaíba, ParanáParaguay Pimelodus microstoma Branco, Paraná-Paraguay Pimelodus ornatus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Putumayo, Negro, Branco, Trombetas, Amazonas main channel, Parnaíba, Itapicuru-Mearim, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Pimelodus pictus Beni-Madre de Dios, middle-lower Madeira, Javari, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Amazonas main channel, upper Orinoco, Apure Pimelodus quadratus* Endemic – upper Tocantins Pimelodus speciosus* Endemic – upper Tocantins Pimelodus stewarti* Endemic – upper Tocantins Pimelodus tetramerus* Upper Tocantins, Araguaia, lower Tocantins, Teles Pires, Tapajós Pinirampus pirinampu Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Ucayali, Putumayo, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Platynematichthys notatus Lower Tocantins, lower Xingu, Teles Pires, Tapajós, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, lower Orinoco, upper Orinoco, Apure Platysilurus mucosus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Juruá, Japurá, Branco, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Platysilurus olallae* Endemic – Marañon Platystomatichthys sturio* Lower Tocantins, upper Xingu, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary Propimelodus araguayae* Endemic – Araguaia Propimelodus caesius Lower Tocantins, Mamoré, middle-lower Madeira, Purus, Negro, Amazonas main channel, Amazonas Estuary, Capim Propimelodus eigenmanni Lower Tocantins, Negro, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Oiapoque, Maroni-Approuague Pseudoplatystoma punctifer Araguaia, Iriri, lower Xingu, Teles Pires, Juruena, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Jutaí, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Negro, Parnaíba, Itapicuru-Mearim Pseudoplatystoma reticulatum Negro, Paraná-Paraguay 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 123 Occurrence Pseudoplatystoma tigrinum* Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Marañon-Nanay, Branco, Amazonas main channel Sorubim elongatus Lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Purus, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Sorubim lima Upper Tocantins, Araguaia, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Negro, Branco, Trombetas, Amazonas main channel, Parnaíba, Itapicuru-Mearim, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Sorubim maniradii* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Napo-Ambyiacu, Amazonas main channel Sorubim trigonocephalus* Upper Xingu, lower Xingu, Teles Pires, Juruena Sorubimichthys planiceps Araguaia, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Japurá, Negro, Branco, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Zungaro zungaro Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Zungaropsis multimaculatus** Pseudopimelodidae 16 Batrochoglanis raninus Lower Tocantins, upper Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Putumayo, Japurá, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, MaroniApprouague, Coppename-Suriname-Saramacca, Essequibo, lower Orinoco, upper Orinoco Batrochoglanis villosus Lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Juruá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Itapicuru-Mearim, Capim, Araguari-Macari-Amapá, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure Microglanis lundbergi* Endemic – Amazonas main channel Microglanis maculatus* Endemic – upper Tocantins Microglanis oliveirai* Endemic – Araguaia Microglanis pellopterygius* Endemic – Napo-Ambyiacu Microglanis poecilus Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Branco, Amazonas main channel, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Microglanis robustus* Endemic – lower Tocantins Microglanis secundus Branco, Trombetas, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Microglanis xerente* Endemic – upper Tocantins Microglanis xylographicus* Endemic – Araguaia Microglanis zonatus* Endemic – Marañon 124 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Pseudopimelodus bufonius Upper Tocantins, Araguaia, upper Xingu, Iriri, Teles Pires, Jamanxim, middlelower Madeira, Negro, Branco, Urubu-Uatumã, Trombetas, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Rhyacoglanis epiblepsis* Endemic – Guaporé Rhyacoglanis pulcher* Endemic – Marañon Rhyacoglanis seminiger* Endemic – Juruena Loricariidae 361 Acanthicus adonis* Endemic – lower Tocantins Acanthicus hystrix Upper Tocantins, Araguaia, lower Tocantins, lower Xingu, middle-lower Madeira, Purus, Marañon-Nanay, Japurá, Branco, Amazonas main channel, upper Orinoco Acestridium dichromum Negro, upper Orinoco Acestridium discus* Negro, Branco, Trombetas, Amazonas main channel Acestridium gymnogaster* Endemic – Middle-lower Madeira Acestridium martini Negro, upper Orinoco Acestridium scutatum* Endemic – Middle-lower Madeira Acestridium triplax* Endemic – Tapajós Ancistomus feldbergae* Iriri, lower Xingu, Teles Pires Ancistomus micrommatos* Endemic – upper Tocantins Ancistomus snethlageae* Restricted to Tapajós basin (Teles Pires, Juruena, Tapajós) Ancistomus spilomma* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu Ancistomus spinosissimus* Endemic – upper Tocantins Ancistrus aguaboensis* Endemic – upper Tocantins Ancistrus alga* Marañon-Nanay, Napo-Ambyiacu Ancistrus bolivianus* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Ancistrus brevifilis Amazonas main channel, upper Orinoco, Apure, Atl. Coastal Drainages of Col. Ven. Ancistrus bufonius* Mamoré, Beni-Madre de Dios, Ucayali Ancistrus cryptophthalmus* Endemic – upper Tocantins Ancistrus dolichopterus Beni-Madre de Dios, Madeira Shield Tributaries, Tefé, Branco, lower Orinoco Ancistrus dubius** Ancistrus heterorhynchus* Endemic – Beni-Madre de Dios Ancistrus hoplogenys Lower Tocantins, Purus, Branco, Essequibo, upper Orinoco Ancistrus jataiensis* Endemic – upper Tocantins Ancistrus jelskii* Endemic – Ucayali Ancistrus karajas* Endemic – lower Tocantins Ancistrus krenakarore* Endemic – Tapajós Ancistrus latifrons Marañon-Nanay, upper Orinoco Ancistrus lineolatus* Putumayo, Japurá Ancistrus maculatus* Amazonas main channel Ancistrus malacops* Marañon-Nanay, Napo-Ambyiacu 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 125 Occurrence Ancistrus maximus* Endemic – Branco Ancistrus megalostomus* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios) Ancistrus minutus* Endemic – upper Tocantins Ancistrus montanus* Restricted to Madeira basin (Guaporé, Beni-Madre de Dios) Ancistrus nudiceps* Endemic – Branco Ancistrus occidentalis* Marañon-Nanay, Napo-Ambyiacu Ancistrus occloi* Endemic – Ucayali Ancistrus parecis* Endemic – Juruena Ancistrus ranunculus* Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós Ancistrus reisi* Endemic – upper Tocantins Ancistrus shuar* Endemic – Marañon Ancistrus stigmaticus* Endemic – Araguaia Ancistrus tamboensis* Endemic – Ucayali Ancistrus tombador* Endemic – Juruena Ancistrus variolus* Endemic – Napo-Ambyiacu Ancistrus verecundus* Middle-lower Madeira, Madeira Shield Tributaries Andeancistrus eschwartzae* Endemic – Marañon Andeancistrus platycephalus* Endemic – Marañon Aphanotorulus emarginatus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco, Apure Aphanotorulus horridus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Japurá, Amazonas main channel Aphanotorulus phrixosoma * Juruá, Ucayali Aphanotorulus rubrocauda* Endemic – Madeira Shield Tributaries Aphanotorulus unicolor* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Amazonas main channel Apistoloricaria condei* Napo-Ambyiacu, Putumayo Apistoloricaria laani Mamoré, middle-lower Madeira, upper Orinoco Apistoloricaria ommation* Mamoré, middle-lower Madeira, Amazonas main channel Aposturisoma myriodon* Endemic – Ucayali Araichthys loro* Endemic – Juruena Baryancistrus chrysolomus* Restricted to Xingu basin (Iriri, lower Xingu) Baryancistrus longipinnis* Lower Tocantins, upper Xingu, Tapajós Baryancistrus niveatus* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, Trombetas Baryancistrus xanthellus* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Chaetostoma anale* Endemic – Japurá Chaetostoma branickii* Endemic – Marañon 126 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Chaetostoma breve* Marañon-Nanay, Napo-Ambyiacu Chaetostoma carrioni* Endemic – Marañon Chaetostoma changae* Endemic – Marañon Chaetostoma daidalmatos* Endemic – Marañon Chaetostoma dermorhynchum* Marañon-Nanay, Napo-Ambyiacu Chaetostoma jegui* Endemic – Branco Chaetostoma lineopunctatum* Beni-Madre de Dios, Ucayali Chaetostoma loborhynchos* Endemic – Ucayali Chaetostoma marcapatae* Endemic – Beni-Madre de Dios Chaetostoma marmorescens* Endemic – Marañon Chaetostoma microps* Marañon-Nanay, Napo-Ambyiacu Chaetostoma platyrhynchus* Endemic – Japurá Chaetostoma sericeum* Endemic – Napo-Ambyiacu Chaetostoma spondylus* Endemic – Marañon Chaetostoma stroumpoulos* Endemic – Marañon Chaetostoma taczanowskii* Ucayali, Marañon-Nanay Chaetostoma trimaculineum* Endemic – Marañon Chaetostoma vagum* Endemic – Japurá Corumbataia tocantinensis* Restricted to Tocantins basin (upper Tocantins, Araguaia) Corumbataia veadeiros* Endemic – upper Tocantins Cteniloricaria napova* Endemic – Curuá-Paru do Oeste Curculionichthys itaim* Endemic – Tapajós Curculionichthys karipuna* Endemic – Jari Curculionichthys luteofrenatus* Restricted to Tapajós basin (Teles Pires, Juruena) Curculionichthys sabaji* Restricted to Xingu basin (upper Xingu, Iriri) Curculionichthys tukana* Restricted to Tocantins basin (upper Tocantins, lower Tocantins) Dekeyseria amazonica* Araguaia, lower Tocantins, lower Xingu, Tapajós, middle-lower Madeira, Juruá, Japurá, Trombetas, Jari, Amazonas main channel Dekeyseria picta Negro, upper Orinoco Dekeyseria scaphirhynchus Middle-lower Madeira, Purus, Negro, Branco, upper Orinoco Exastilithoxus fimbriatus Branco, lower Orinoco Exastilithoxus hoedemani* Endemic – Negro Farlowella acus Branco, Essequibo, upper Orinoco, Apure, Atl. Coastal Drainages of Col. Ven. Farlowella altocorpus* Endemic – Beni-Madre de Dios Farlowella amazonum Araguaia, lower Tocantins, lower Xingu, Tapajós, Jamanxim, middle-lower Madeira, Jutaí, Putumayo, Negro, Branco, Trombetas, Jari, Amazonas main channel, GurupiTuriaçu, Capim, Essequibo, Paraná-Paraguay Farlowella gianetii* Endemic – upper Xingu Farlowella gracilis* Endemic – Japurá Farlowella hasemani* Endemic – Amazonas main channel 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 127 Occurrence Farlowella henriquei* Endemic – Araguaia Farlowella knerii* Beni-Madre de Dios, Ucayali, Marañon-Nanay, Napo-Ambyiacu Farlowella nattereri Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Japurá, Negro, Branco, Amazonas main channel, Amazonas Estuary, Essequibo Farlowella odontotumulus Napo-Ambyiacu, upper Orinoco Farlowella oxyrryncha Araguaia, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Napo-Ambyiacu, Japurá, Negro, Branco, Amazonas main channel, Amazonas Estuary, Gurupi-Turiaçu, Capim, lower Orinoco, upper Orinoco, Apure Farlowella platorynchus Coari-Urucu, Marañon-Nanay, Napo-Ambyiacu, Branco, Amazonas main channel, Amazonas Estuary, Capim Farlowella reticulata Branco, Oiapoque, Maroni-Approuague, Essequibo Farlowella rugosa Branco, Maroni-Approuague, Corentyne-Demerara, Essequibo Farlowella schreitmuelleri* Negro, Amazonas main channel, Amazonas Estuary Farlowella smithi* Mamoré, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Ucayali, Japurá, Branco, Amazonas main channel Fonchiiloricaria nanodon* Endemic – Marañon Furcodontichthys novaesi* Tapajós, Purus, Tefé, Juruá, Ucayali, Negro, Trombetas, Amazonas main channel Guyanancistrus brevispinis Curuá-Paru do Oeste, Jari, Araguari-Macari-Amapá, Oiapoque, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara Gymnotocinclus anosteos* Endemic – upper Tocantins Gymnotocinclus canoeiro* Endemic – upper Tocantins Harttia depressa* Endemic – Urubu-Uatumã Harttia dissidens* Restricted to Tapajós basin (Teles Pires, Tapajós, Jamanxim) Harttia duriventris* Endemic – lower Tocantins Harttia panara* Endemic – Iriri Harttia punctata* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Harttia rondoni* Restricted to Xingu basin (upper Xingu, Iriri) Harttia trombetensis* Endemic – Trombetas Harttia tuna* Endemic – Curuá-Paru do Oeste Harttia uatumensis* Endemic – Urubu-Uatumã Harttia villasboas* Endemic – Iriri Hemiancistrus cerrado* Endemic – Araguaia Hemiancistrus guahiborum Negro, lower Orinoco, upper Orinoco Hemiancistrus subviridis Negro, upper Orinoco Hemiodontichthys acipenserinus Araguaia, lower Tocantins, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Itapicuru-Mearim, Essequibo, ParanáParaguay Hisonotus acuen* Endemic – upper Xingu Hisonotus bockmanni* Endemic – Teles Pires Hisonotus chromodontus* Restricted to Tapajós basin (Teles Pires, Juruena) 128 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Hisonotus jumaorum* Endemic – Madeira Shield Tributaries Hopliancistrus tricornis* Upper Xingu, Iriri, lower Xingu, Tapajós, Jamanxim Hypancistrus inspector Negro, upper Orinoco Hypancistrus margaritatus* Endemic – Branco Hypancistrus phantasma* Endemic – Negro Hypancistrus zebra* Restricted to Xingu basin (upper Xingu, lower Xingu) Hypoptopoma baileyi* Mamoré, Guaporé, Beni-Madre de Dios, Amazonas main channel Hypoptopoma bianale* Ucayali, Napo-Ambyiacu Hypoptopoma brevirostratum* Endemic – Amazonas main channel Hypoptopoma elongatum* Teles Pires, Tapajós, Trombetas Hypoptopoma guianense Branco, Corentyne-Demerara, Essequibo Hypoptopoma gulare* Araguaia, lower Tocantins, upper Xingu, Juruá, Ucayali, Marañon-Nanay, NapoAmbyiacu, Amazonas main channel Hypoptopoma incognitum Upper Tocantins, Araguaia, lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Amazonas main channel, Amazonas Estuary, Itapicuru-Mearim Hypoptopoma inexspectatum Upper Xingu, Paraná-Paraguay Hypoptopoma muzuspi* Restricted to Tocantins basin (Araguaia, lower Tocantins) Hypoptopoma psilogaster* Endemic – Marañon Hypoptopoma spectabile Purus, Marañon-Nanay, Napo-Ambyiacu, lower Orinoco, upper Orinoco, Apure Hypoptopoma steindachneri Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Hypoptopoma sternoptychum* Mamoré, Beni-Madre de Dios, middle-lower Madeira, Napo-Ambyiacu, Amazonas main channel Hypoptopoma thoracatum Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Japurá, Branco, Amazonas main channel, Essequibo Hypostomus asperatus* Endemic – Araguaia Hypostomus atropinnis* Endemic – Araguaia Hypostomus bolivianus* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios) Hypostomus carinatus* Beni-Madre de Dios, Juruá, Branco, Urubu-Uatumã, Trombetas Hypostomus dardanelos* Endemic – Madeira Shield Tributaries Hypostomus delimai* Restricted to Tocantins basin (Araguaia, lower Tocantins) Hypostomus ericae* Restricted to Tocantins basin (upper Tocantins, Araguaia) Hypostomus ericius* Endemic – Marañon Hypostomus faveolus* Upper Tocantins, Araguaia, upper Xingu Hypostomus fonchii* Endemic – Ucayali Hypostomus goyazensis* Endemic – Araguaia Hypostomus hemicochliodon Upper Xingu, Juruena, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Negro, Amazonas main channel, upper Orinoco Hypostomus hemiurus Branco, Corentyne-Demerara, Essequibo Hypostomus hoplonites* Mamoré, Guaporé, middle-lower Madeira, Purus, Amazonas main channel 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 129 Occurrence Hypostomus kopeyaka* Endemic – Negro Hypostomus kuarup* Endemic – upper Xingu Hypostomus levis* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios) Hypostomus macushi Branco, Essequibo Hypostomus melanephelis* Endemic – Tapajós Hypostomus niceforoi* Endemic – Japurá Hypostomus oculeus* Ucayali, Marañon-Nanay, Napo-Ambyiacu, Japurá, Branco, Amazonas main channel Hypostomus pantherinus* Restricted to Madeira basin (Mamoré, Guaporé) Hypostomus paucipunctatus* Endemic – lower Tocantins Hypostomus pyrineusi Lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, NapoAmbyiacu, Amazonas main channel, upper Orinoco Hypostomus rondoni* Endemic – Teles Pires Hypostomus sculpodon Negro, upper Orinoco Hypostomus soniae* Restricted to Tapajós basin (Teles Pires, Tapajós, Jamanxim) Hypostomus taphorni Branco, Corentyne-Demerara, Essequibo Hypostomus varimaculosus* Endemic – Japurá Hypostomus weberi* Endemic – Negro Lamontichthys avacanoeiro* Endemic – upper Tocantins Lamontichthys filamentosus* Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Amazonas main channel Lamontichthys parakana* Endemic – lower Tocantins Lamontichthys stibaros* Ucayali, Marañon-Nanay, Napo-Ambyiacu Lasiancistrus heteracanthus* Napo-Ambyiacu, Putumayo, Japurá Lasiancistrus schomburgkii Lower Tocantins, lower Xingu, Jamanxim, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, NapoAmbyiacu, Japurá, Branco, Essequibo, upper Orinoco Leporacanthicus galaxias Upper Tocantins, Araguaia, lower Tocantins, middle-lower Madeira, lower Orinoco, upper Orinoco Leporacanthicus heterodon* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Leporacanthicus joselimai* Restricted to Tapajós basin (Teles Pires, Tapajós) Leptotocinclus ctenistus* Coari-Urucu, Negro, Amazonas main channel Leptotocinclus madeirae* Endemic – Middle-lower Madeira Limatulichthys griseus Upper Tocantins, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Ucayali, Napo-Ambyiacu, Putumayo, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Essequibo, lower Orinoco, upper Orinoco Lithoxus bovallii* Branco, Trombetas Lithoxus jariensis* Endemic – Jari Lithoxus lithoides Branco, Urubu-Uatumã, Trombetas, Corentyne-Demerara, Essequibo Loraxichthys lexa* Endemic – Ucayali 130 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Loricaria birindellii* Restricted to Xingu basin (Iriri, lower Xingu) Loricaria cataphracta Lower Tocantins, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Putumayo, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Essequibo, lower Orinoco, upper Orinoco, Apure Loricaria clavipinna* Ucayali, Amazonas main channel Loricaria lata* Endemic – upper Tocantins Loricaria lundbergi* Endemic – Negro Loricaria pumila* Lower Tocantins, Amazonas main channel, Amazonas Estuary Loricaria simillima Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Branco, Amazonas main channel, upper Orinoco, Atl. Coastal Drainages of Col. Ven., Paraná-Paraguay Loricaria spinulifera* Negro, Amazonas main channel Loricariichthys acutus Araguaia, lower Xingu, Tapajós, middle-lower Madeira, Purus, Negro, Branco, Trombetas, Amazonas main channel, Itapicuru-Mearim, Essequibo Loricariichthys cashibo** Loricariichthys chanjoo** Loricariichthys hauxwelli* Ucayali, Amazonas main channel Loricariichthys maculatus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Negro, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Loricariichthys nudirostris* Lower Tocantins, Tapajós, middle-lower Madeira, Negro, Branco, Trombetas, Amazonas main channel Loricariichthys platymetopon Mamoré, Beni-Madre de Dios, middle-lower Madeira, Jari, Amazonas main channel, Amazonas Estuary, Paraná-Paraguay Loricariichthys stuebelii* Guaporé, Purus, Juruá, Ucayali, Marañon-Nanay, Japurá, Amazonas main channel Loricariichthys ucayalensis** Metaloricaria paucidens Trombetas, Oiapoque, Maroni-Approuague Microplecostomus forestii* Endemic – upper Tocantins Nannoplecostomus eleonorae* Endemic – upper Tocantins Nannoxyropsis acicula* Endemic – Tapajós Neblinichthys pilosus Negro, upper Orinoco Niobichthys ferrarisi* Endemic – Negro Spectracanthicus punctatissimus* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Otocinclus batmani* Ucayali, Japurá, Amazonas main channel Otocinclus caxarari* Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries Otocinclus cocama* Endemic – Ucayali Otocinclus hasemani Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Jamanxim, Parnaíba Otocinclus hoppei* Lower Tocantins, middle-lower Madeira, Madeira Shield Tributaries, Ucayali, Amazonas Estuary Otocinclus huaorani Ucayali, Marañon-Nanay, Napo-Ambyiacu, Upper Orinoco Otocinclus juruenae* Endemic – Juruena 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 131 Occurrence Otocinclus macrospilus* Javari, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Amazonas main channel Otocinclus mangaba* Endemic – Middle-lower Madeira Otocinclus mariae Mamoré, Guaporé, Beni-Madre de Dios, Curuá-Paru do Oeste, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara Otocinclus mura* Mamoré, middle-lower Madeira, Amazonas main channel Otocinclus tapirape* Endemic – Araguaia Otocinclus vestitus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Napo-Ambyiacu, Amazonas main channel, Paraná-Paraguay Otocinclus vittatus Araguaia, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, Marañon-Nanay, Japurá, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Oxyropsis acutirostra Negro, upper Orinoco Oxyropsis carinata* Upper Xingu, Japurá, Negro, Amazonas main channel Oxyropsis wrightiana Teles Pires, middle-lower Madeira, Japurá, Negro, Amazonas main channel, upper Orinoco Panaqolus albivermis* Endemic – Ucayali Panaqolus albomaculatus* Ucayali, Marañon-Nanay, Napo-Ambyiacu Panaqolus changae* Purus, Juruá, Ucayali, Marañon-Nanay Panaqolus claustellifer* Endemic – Branco Panaqolus dentex* Marañon-Nanay, Napo-Ambyiacu Panaqolus gnomus* Marañon-Nanay, Napo-Ambyiacu Panaqolus nix* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios, middle-lower Madeira) Panaqolus nocturnus* Marañon-Nanay, Napo-Ambyiacu Panaqolus purusiensis* Endemic – Purus Panaqolus tankei* Endemic – lower Xingu Panaque armbrusteri* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Jamanxim Panaque bathyphilus* Marañon-Nanay, Amazonas main channel Panaque schaeferi* Middle-lower Madeira, Purus, Ucayali, Marañon-Nanay, Amazonas main channel Panaque titan* Endemic – Napo-Ambyiacu Parancistrus aurantiacus* Araguaia, lower Tocantins, upper Xingu, Iriri, Ucayali Parancistrus nudiventris* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Pareiorhaphis regani* Negro Parotocinclus amazonensis* Middle-lower Madeira, Amazonas main channel Parotocinclus aripuanensis* Endemic – Madeira Shield Tributaries Parotocinclus britskii Araguaia, lower Tocantins, Branco, Amazonas main channel, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo Parotocinclus collinsae* Endemic – Branco Parotocinclus dani* Endemic – Teles Pires Parotocinclus halbothi Trombetas, Maroni-Approuague Parotocinclus longirostris* Endemic – Amazonas main channel 132 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Parotocinclus polyochrus Negro, upper Orinoco Parotocinclus variola* Endemic – Amazonas main channel Parotocinclus yaka* Endemic – Negro Peckoltia braueri* Branco, Amazonas main channel Peckoltia brevis* Lower Tocantins, Mamoré, Guaporé, Purus, Juruá, Amazonas main channel Peckoltia cavatica Tapajós, Essequibo Peckoltia compta* Restricted to Tapajós basin (Tapajós, Jamanxim) Peckoltia ephippiata* Endemic – Madeira Shield Tributaries Peckoltia furcata* Ucayali, Marañon-Nanay, Amazonas main channel Peckoltia multispinis** Peckoltia oligospila Upper Tocantins, lower Tocantins, Capim Peckoltia pankimpuju* Endemic – Amazonas main channel Peckoltia relictum* Endemic – Marañon Peckoltia sabaji Lower Xingu, Teles Pires, Tapajós, Negro, Branco, Jari, Essequibo, upper Orinoco Peckoltia vermiculata** Peckoltia vittata Upper Tocantins, Araguaia, lower Tocantins, Anapu, upper Xingu, Iriri, lower Xingu, Tapajós, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Gurupi-Turiaçu, Capim, upper Orinoco Peckoltichthys bachi Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Amazonas main channel, upper Orinoco Planiloricaria cryptodon* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Putumayo, Amazonas main channel Pseudacanthicus histrix** Lower Tocantins, upper Xingu, lower Xingu, Purus, Japurá, lower Orinoco, upper Orinoco, Apure Pseudacanthicus leopardus Upper Xingu, lower Xingu, Branco, Essequibo Pseudacanthicus major* Restricted to Tocantins basin (upper Tocantins, lower Tocantins) Pseudacanthicus pirarara* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Pseudacanthicus pitanga* Restricted to Tocantins basin (upper Tocantins, lower Tocantins) Pseudacanthicus serratus Upper Tocantins, lower Tocantins, Maroni-Approuague, Coppename-SurinameSaramacca, Essequibo Pseudacanthicus spinosus Lower Tocantins, Anapu, Pacajá, upper Xingu, lower Xingu, Negro, Urubu-Uatumã, Amazonas main channel, Amazonas Estuary, Capim, lower Orinoco Pseudancistrus asurini* Restricted to Xingu basin (Iriri, lower Xingu) Pseudancistrus kayabi* Endemic – Teles Pires Pseudancistrus nigrescens Branco, Essequibo Pseudancistrus pectegenitor Negro, upper Orinoco Pseudancistrus sidereus Negro, upper Orinoco Pseudancistrus zawadzkii* Endemic – Tapajós Pseudohemiodon amazonum* Endemic – Trombetas Pseudohemiodon apithanos* Endemic – Putumayo Pseudohemiodon lamina* Beni-Madre de Dios 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 133 Occurrence Pseudohemiodon thorectes* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Pseudolithoxus kinja* Madeira Shield Tributaries, Urubu-Uatumã, Trombetas Pseudolithoxus nicoi Negro, upper Orinoco Pseudoloricaria laeviuscula Araguaia, lower Xingu, Tapajós, Jamanxim, middle-lower Madeira, Madeira Shield Tributaries, Negro, Branco, Trombetas, Amazonas main channel, upper Orinoco Pseudorinelepis genibarbis* Middle-lower Madeira, Purus, Juruá, Ucayali, Napo-Ambyiacu, Putumayo, Branco, Amazonas main channel Pterosturisoma microps* Mamoré, Guaporé, middle-lower Madeira, Putumayo, Amazonas main channel Pterygoplichthys disjunctivus* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Pterygoplichthys gibbiceps Lower Tocantins, Tapajós, Purus, Putumayo, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Pterygoplichthys joselimaianus* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Pterygoplichthys lituratus* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Juruá, Amazonas main channel Pterygoplichthys multiradiatus Beni-Madre de Dios, lower Orinoco, upper Orinoco, Apure Pterygoplichthys pardalis* Mamoré, Guaporé, middle-lower Madeira, Purus, Ucayali, Napo-Ambyiacu, Putumayo, Amazonas main channel Pterygoplichthys punctatus Lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, Purus, Juruá, Ucayali, Putumayo, Amazonas main channel, Itapicuru-Mearim, Capim Pterygoplichthys scrophus* Purus, Ucayali, Amazonas main channel Pterygoplichthys weberi* Marañon-Nanay, Napo-Ambyiacu, Japurá, Amazonas main channel Pterygoplichthys xinguensis* Restricted to Xingu basin (upper Xingu, lower Xingu) Reganella depressa* Lower Xingu, Tapajós, Guaporé, Negro, Branco, Trombetas, Amazonas main channel Rhadinoloricaria bahuaja* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus Rhadinoloricaria rhami* Juruá, Ucayali, Amazonas main channel Rhadinoloricaria macromystax** Rhinolekos capetinga* Endemic – upper Tocantins Rineloricaria aurata Guaporé, Paraná-Paraguay Rineloricaria beni* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios) Rineloricaria castroi* Branco, Trombetas, Amazonas main channel Rineloricaria daraha* Endemic – Negro Rineloricaria fallax Japurá, Negro, Branco, Trombetas, Corentyne-Demerara, Essequibo Rineloricaria formosa Mamoré, middle-lower Madeira, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Rineloricaria hasemani Upper Tocantins, Araguaia, Branco, Capim Rineloricaria heteroptera* Endemic – Amazonas main channel Rineloricaria jurupari* Endemic – Negro Rineloricaria konopickyi** Rineloricaria lanceolata Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Ucayali, Marañon-Nanay, Negro, Branco, Trombetas, Amazonas main channel, Essequibo, Paraná-Paraguay 134 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Rineloricaria melini* Endemic – Amazonas main channel Rineloricaria microlepidota* Endemic – Juruá Rineloricaria morrowi* Endemic – Ucayali Rineloricaria osvaldoi* Restricted to Tocantins basin (upper Tocantins, Araguaia) Rineloricaria phoxocephala* Purus, Coari-Urucu, Branco, Amazonas main channel Rineloricaria stewarti Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo Rineloricaria teffeana* Endemic – Amazonas main channel Rineloricaria wolfei* Endemic – Ucayali Scobinancistrus aureatus* Restricted to Xingu basin (upper Xingu, lower Xingu) Scobinancistrus pariolispos* Lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós Spatuloricaria caquetae* Putumayo, Japurá Spatuloricaria euacanthagenys** Japurá Spatuloricaria evansii Upper Tocantins, Araguaia, lower Tocantins, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Marañon-Nanay, UrubuUatumã, Trombetas, Paraná-Paraguay Spatuloricaria puganensis** Beni-Madre de Dios, Purus, Juruá Spatuloricaria tuira* Upper Xingu, Iriri, lower Xingu, Teles Pires, Jamanxim Spectracanthicus immaculatus* Endemic – Tapajós Spectracanthicus javae* Endemic – Araguaia Spectracanthicus murinus* Restricted to Tapajós basin (Teles Pires, Tapajós, Jamanxim) Spectracanthicus punctatissimus* Endemic – lower Xingu Spectracanthicus tocantinensis* Endemic – lower Tocantins Spectracanthicus zuanoni* Endemic – lower Xingu Sturisoma brevirostre* Endemic – Amazonas main channel Sturisoma graffini* Endemic – Beni-Madre de Dios Sturisoma guentheri* Marañon-Nanay, Napo-Ambyiacu Sturisoma lyra* Mamoré, middle-lower Madeira, Juruá Sturisoma monopelte Branco, Essequibo Sturisoma nigrirostrum* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Putumayo Sturisoma rostratum* Upper Tocantins, Araguaia, lower Tocantins, Jamanxim Sturisoma tenuirostre Branco, lower Orinoco, upper Orinoco, Apure Sturisomatichthys caquetae* Endemic – Japurá Scoloplacidae 6 Scoloplax baileyi* Negro, Amazonas main channel Scoloplax baskini* Middle-lower Madeira, Madeira Shield Tributaries, Purus, Branco Scoloplax dicra* Mamoré, Guaporé, Beni-Madre de Dios, Negro, Branco, Amazonas main channel Scoloplax distolothrix Araguaia, upper Xingu, lower Xingu, Paraná-Paraguay Scoloplax dolicholophia* Negro, Amazonas main channel Scoloplax empousa Guaporé, Paraná-Paraguay 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Trichomycteridae Species 135 Occurrence 75 Acanthopoma annectens* Mamoré, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Amazonas main channel Ammoglanis amapaensis* Endemic – Jari Ammoglanis diaphanus* Endemic – Araguaia Ammoglanis pulex Amazonas main channel, upper Orinoco Apomatoceros alleni* Middle-lower Madeira, Marañon-Nanay, Amazonas main channel Glanapteryx anguilla Negro, Branco, upper Orinoco Glanapteryx niobium* Endemic – Negro Haemomaster venezuelae Upper Tocantins, Araguaia, upper Xingu, Teles Pires, Juruena, Madeira Shield Tributaries, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Henonemus intermedius* Endemic – Araguaia Henonemus macrops* Endemic – Amazonas main channel Henonemus punctatus Upper Tocantins, Araguaia, lower Xingu, Mamoré, Beni-Madre de Dios, middlelower Madeira, Purus, Juruá, Napo-Ambyiacu, Putumayo, Branco, Amazonas main channel, Essequibo Ituglanis amazonicus Upper Xingu, Jamanxim, Mamoré, middle-lower Madeira, Amazonas main channel, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, lower Orinoco Ituglanis apteryx* Endemic – Iriri Ituglanis bambui* Endemic – upper Tocantins Ituglanis boticario* Endemic – upper Tocantins Ituglanis compactus* Endemic – Jari Ituglanis epikarsticus* Endemic – upper Tocantins Ituglanis goya Upper Tocantins, Paraná-Paraguay Ituglanis ina* Endemic – lower Tocantins Ituglanis macunaima* Endemic – Araguaia Ituglanis mambai* Endemic – upper Tocantins Ituglanis parkoi* Endemic – Javari Ituglanis passensis* Endemic – upper Tocantins Ituglanis ramiroi* Endemic – upper Tocantins Malacoglanis gelatinosus Japurá, upper Orinoco Megalocentor echthrus Mamoré, middle-lower Madeira, Purus, Juruá, Japurá, Negro, Branco, Amazonas main channel, lower Orinoco Miuroglanis platycephalus* Beni-Madre de Dios, middle-lower Madeira, Purus, Urubu-Uatumã, Amazonas main channel Ochmacanthus alternus Lower Tocantins, Guaporé, Putumayo, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, Apure Ochmacanthus orinoco Araguaia, lower Tocantins, lower Xingu, Madeira Shield Tributaries, Negro, Branco, Amazonas main channel, upper Orinoco, Apure Ochmacanthus reinhardtii Upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Juruá, Napo-Ambyiacu, Putumayo, Negro, Branco, Essequibo 136 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Paracanthopoma parva Upper Tocantins, Araguaia, upper Xingu, Teles Pires, Juruena, Tapajós, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Marañon-Nanay, NapoAmbyiacu, Negro, Branco, Trombetas, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Pareiodon microps* Araguaia, middle-lower Madeira, Amazonas main channel Plectrochilus diabolicus* Mamoré, Beni-Madre de Dios, middle-lower Madeira, Juruá, Amazonas main channel Plectrochilus machadoi* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel Plectrochilus wieneri* Endemic – Napo-Ambyiacu Potamoglanis anhanga* Madeira Shield Tributaries, Amazonas main channel Potamoglanis hasemani Anapu, Teles Pires, Juruena, Tapajós, middle-lower Madeira, Purus, Coari-Urucu, Negro, Branco, Amazonas main channel, Amazonas Estuary, Gurupi-Turiaçu, Capim, Essequibo Potamoglanis johnsoni Jari, Amazonas main channel, Paraná-Paraguay Potamoglanis wapixana* Endemic – Branco Pseudostegophilus nemurus Lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Napo-Ambyiacu, Japurá, Negro, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco Pygidianops amphioxus* Negro, Amazonas main channel Pygidianops eigenmanni* Restricted to Negro basin (Negro, Branco) Pygidianops magoi Branco, lower Orinoco, upper Orinoco Sarcoglanis simplex Negro, Branco, upper Orinoco Schultzichthys bondi Middle-lower Madeira, Napo-Ambyiacu, lower Orinoco, upper Orinoco, Apure Stauroglanis gouldingi* Negro, Amazonas main channel Stegophilus panzeri Lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Jamanxim, Madeira Shield Tributaries, Capim Stenolicmus ix Curuá-Paru do Oeste, Maroni-Approuague Stenolicmus sarmientoi* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Trichomycterus barbouri Beni-Madre de Dios, Paraná-Paraguay Trichomycterus bomboizanus* Endemic – Marañon Trichomycterus chaberti* Endemic – Mamoré Trichomycterus dispar* Beni-Madre de Dios, Ucayali Trichomycterus fassli* Endemic – Beni-Madre de Dios Trichomycterus gabrieli* Restricted to Negro basin (Negro, Branco) Trichomycterus guianense Branco, Essequibo, lower Orinoco Trichomycterus megantoni* Endemic – Ucayali Trichomycterus nigromaculatus Putumayo, Cauca-Magdalena-Sinu, Atrato Trichomycterus oroyae* Endemic – Ucayali Trichomycterus punctatissimus* Endemic – Araguaia Trichomycterus rivulatus* Beni-Madre de Dios, Ucayali Trichomycterus taczanowskii* Endemic – Marañon Trichomycterus taeniops* Endemic – Ucayali 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 137 Occurrence Trichomycterus tiraquae** Trichomycterus vittatus* Beni-Madre de Dios, Marañon-Nanay Trichomycterus weyrauchi* Endemic – Ucayali Tridens melanops* Mamoré, Purus, Putumayo, Amazonas main channel Tridensimilis brevis* Mamoré, middle-lower Madeira, Amazonas main channel Tridentopsis pearsoni** Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus Tridentopsis tocantinsi* Upper Tocantins, Araguaia Typhlobelus auriculatus* Endemic – lower Xingu Typhlobelus macromycterus* Lower Tocantins, lower Xingu Typhlobelus ternetzi* Endemic – Negro Vandellia cirrhosa Upper Tocantins, Araguaia, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Marañon-Nanay, NapoAmbyiacu, Japurá, Negro, Branco, Trombetas, Amazonas main channel, GurupiTuriaçu, Capim, upper Orinoco, Apure Vandellia sanguinea Upper Tocantins, Araguaia, Tapajós, middle-lower Madeira, Purus, Branco, Jari, Amazonas main channel, Essequibo, lower Orinoco Gymnotiformes 164 Apteronotidae 62 Adontosternarchus balaenops* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Putumayo, Japurá, Amazonas main channel Apteronotus bonapartii Upper Tocantins, upper Xingu, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Tefé, Ucayali, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Adontosternarchus clarkae Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, NapoAmbyiacu, Negro, Amazonas main channel, upper Orinoco Adontosternarchus duartei* Endemic – Purus Adontosternarchus nebulosus* Middle-lower Madeira, Purus, Tefé, Juruá, Negro, Trombetas, Amazonas main channel Adontosternarchus sachsi Lower Tocantins, Mamoré, Guaporé, Purus, Tefé, Negro, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Apteronotus albifrons Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Ucayali, Branco, Urubu-Uatumã, Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Apteronotus camposdapazi* Endemic – upper Tocantins Apteronotus lindalvae* Endemic – Urubu-Uatumã Apteronotus macrolepis* Endemic – Amazonas main channel Compsaraia compsus Madeira Shield Tributaries, Negro, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Compsaraia iara* Middle-lower Madeira, Branco, Amazonas main channel, Amazonas Estuary Compsaraia samueli* Putumayo, Amazonas main channel Megadontognathus kaitukaensis* Upper Xingu, Teles Pires, Tapajós Melanosternarchus amaru* Lower Xingu, Purus, Negro, Amazonas main channel 138 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Orthosternarchus tamandua* Middle-lower Madeira, Purus, Negro, Branco, Amazonas main channel Parapteronotus hasemani* Endemic – Amazonas main channel Pariosternarchus amazonensis* Middle-lower Madeira, Amazonas main channel Platyurosternarchus crypticus* Endemic – Branco Platyurosternarchus macrostoma Araguaia, lower Tocantins, upper Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Jari, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco, Apure Porotergus duende* Middle-lower Madeira, Ucayali, Japurá, Negro, Amazonas main channel Porotergus gimbeli Lower Tocantins, Anapu, Pacajá, Mamoré, Guaporé, middle-lower Madeira, CoariUrucu, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Essequibo Porotergus gymnotus Mamoré, Guaporé, Maroni-Approuague, Corentyne-Demerara, Essequibo Sternarchorhynchus axelrodi* Restricted to Tocantins basin (upper Tocantins, lower Tocantins) Sternarchorhynchus caboclo* Endemic – Branco Sternarchorhynchus chaoi* Restricted to Madeira basin (Mamoré, middle-lower Madeira) Sternarchorhynchus cramptoni* Mamoré, Marañon-Nanay, Amazonas main channel Sternarchorhynchus curumim* Endemic – Amazonas main channel Sternarchorhynchus curvirostris* Endemic – Marañon Sternarchorhynchus goeldii* Lower Tocantins, Mamoré, middle-lower Madeira, Purus, Negro, Amazonas main channel Sternarchorhynchus hagedornae* Endemic – Beni-Madre de Dios Sternarchorhynchus higuchii* Endemic – Urubu-Uatumã Sternarchorhynchus inpai* Endemic – Trombetas Sternarchorhynchus jaimei* Endemic – Urubu-Uatumã Sternarchorhynchus kokraimoro* Endemic – lower Xingu Sternarchorhynchus mareikeae* Endemic – Trombetas Sternarchorhynchus mesensis* Endemic – upper Tocantins Sternarchorhynchus montanus* Endemic – Amazonas main channel Sternarchorhynchus mormyrus* Purus, Negro, Amazonas main channel Sternarchorhynchus oxyrhynchus Branco, lower Orinoco Sternarchorhynchus retzeri* Negro, Trombetas, Amazonas main channel Sternarchorhynchus schwassmanni* Endemic – Araguaia Sternarchorhynchus severii* Endemic – Branco Sternarchorhynchus starksi* Lower Xingu, Amazonas main channel, Amazonas Estuary Sternarchorhynchus stewarti* Beni-Madre de Dios, Juruá, Marañon-Nanay, Napo-Ambyiacu, Amazonas main channel Sternarchorhynchus taphorni* Ucayali, Marañon-Nanay Sternarchorhynchus villasboasi* Endemic – lower Xingu Sternarchorhynchus yepezi Amazonas main channel, Apure Sternarchella calhamazon* Lower Xingu, middle-lower Madeira, Marañon-Nanay, Napo-Ambyiacu, Negro, Trombetas, Amazonas main channel, Amazonas Estuary 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 139 Occurrence Sternarchella duccis* Purus, Japurá, Negro, Branco, Amazonas main channel Sternarchella orthos Middle-lower Madeira, Napo-Ambyiacu, Amazonas main channel, Amazonas Estuary, lower Orinoco, upper Orinoco, Apure Sternarchella patriciae* Endemic – Beni-Madre de Dios Sternarchella raptor* Jutaí, Japurá, Negro, Branco, Amazonas main channel Sternarchella rex* Endemic – Amazonas main channel Sternarchella schotti* Endemic – Amazonas main channel Sternarchella sima Upper Xingu, Mamoré, Jari, Amazonas main channel, Amazonas Estuary, Capim Sternarchogiton labiatus* Endemic – Amazonas main channel Sternarchogiton nattereri Upper Tocantins, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Purus, Ucayali, Japurá, Negro, Branco, Amazonas main channel, lower Orinoco Sternarchogiton porcinum Guaporé, Branco, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Sternarchogiton preto Lower Tocantins, Anapu, Pacajá, Mamoré, middle-lower Madeira, Purus, Negro, Amazonas main channel, upper Orinoco, Apure Sternarchogiton zuanoni* Endemic – lower Xingu Sternarchorhamphus muelleri Middle-lower Madeira, Juruá, Ucayali, Negro, Branco, Trombetas, Amazonas main channel, Capim, Araguari-Macari-Amapá, lower Orinoco, upper Orinoco, Apure Gymnotidae 26 Electrophorus electricus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Gymnotus arapaima* Tefé, Amazonas main channel Gymnotus capanema* Endemic – Amazonas Estuary Gymnotus carapo Upper Tocantins, Araguaia, lower Tocantins, Iriri, lower Xingu, Teles Pires, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Gymnotus cataniapo Negro, Amazonas Estuary, Maroni-Approuague, Essequibo, upper Orinoco Gymnotus chaviro* Beni-Madre de Dios, Ucayali Gymnotus coatesi* Lower Tocantins, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Amazonas main channel Gymnotus coropinae Lower Tocantins, Tapajós, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Japurá, Negro, Trombetas, Amazonas main channel, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Gymnotus curupira* Mamoré, Beni-Madre de Dios, middle-lower Madeira, Tefé, Ucayali, NapoAmbyiacu, Putumayo Gymnotus diamantinensis* Restricted to Tapajós basin (Teles Pires, Juruena) Gymnotus eyra* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios, Ucayali) Gymnotus inaequilabiatus Guaporé, Paraná-Paraguay Gymnotus javari* Javari, Ucayali, Napo-Ambyiacu, Amazonas main channel Gymnotus jonasi* Middle-lower Madeira, Ucayali, Amazonas main channel 140 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Gymnotus mamiraua* Endemic – Amazonas main channel Gymnotus melanopleura* Endemic – Amazonas main channel Gymnotus obscurus* Endemic – Amazonas main channel Gymnotus onca* Endemic – Amazonas main channel Gymnotus pantanal Mamoré, Guaporé, Beni-Madre de Dios, Paraná-Paraguay Gymnotus pedanopterus Madeira Shield Tributaries, Negro, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Gymnotus riberalta* Restricted to Madeira basin (Guaporé, Beni-Madre de Dios) Gymnotus stenoleucus Negro, Amazonas main channel, lower Orinoco, upper Orinoco Gymnotus tigre* Marañon-Nanay, Amazonas main channel Gymnotus tiquie* Endemic – Negro Gymnotus ucamara* Endemic – Ucayali Gymnotus varzea* Ucayali, Marañon-Nanay, Amazonas main channel Hypopomidae 24 Brachyhypopomus alberti* Restricted to Madeira basin (Mamoré, Guaporé) Brachyhypopomus arrayae* Restricted to Madeira basin (Beni-Madre de Dios, middle-lower Madeira) Brachyhypopomus batesi* Negro, Amazonas main channel Brachyhypopomus beebei Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Purus, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Araguari-Macari-Amapá, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure Brachyhypopomus belindae* Endemic – Amazonas main channel Brachyhypopomus benjamini* Ucayali, Marañon-Nanay Brachyhypopomus bennetti* Beni-Madre de Dios, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Amazonas main channel, Amazonas Estuary Brachyhypopomus bombilla Mamoré, Guaporé, Beni-Madre de Dios, Paraná-Paraguay Brachyhypopomus brevirostris Araguaia, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Marañon-Nanay, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, MaroniApprouague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Brachyhypopomus bullocki Negro, Branco, Essequibo, lower Orinoco, upper Orinoco, Apure Brachyhypopomus cunia* Restricted to Madeira basin (Guaporé, middle-lower Madeira) Brachyhypopomus flavipomus* Javari, Amazonas main channel Brachyhypopomus hamiltoni* Middle-lower Madeira, Marañon-Nanay, Japurá, Negro, Urubu-Uatumã, Amazonas main channel Brachyhypopomus hendersoni Tefé, Japurá, Negro, Amazonas main channel, Essequibo Brachyhypopomus pinnicaudatus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Amazonas main channel, Amazonas Estuary, Itapicuru-Mearim, Capim, Oiapoque Brachyhypopomus provenzanoi* Upper Negro, upper Orinoco 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 141 Occurrence Brachyhypopomus regani Araguaia, Tapajós, Jamanxim, middle-lower Madeira, Purus, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Maroni-Approuague, lower Orinoco, upper Orinoco, Apure Brachyhypopomus sullivani Teles Pires, Juruena, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Napo-Ambyiacu, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Essequibo, upper Orinoco Brachyhypopomus verdii* Endemic – Marañon Brachyhypopomus walteri Upper Tocantins, Araguaia, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Negro, Branco, Urubu-Uatumã, Amazonas main channel, Capim, Essequibo, Paraná-Paraguay Hypopomus artedi Mamoré, Guaporé, Putumayo, Amazonas main channel, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Microsternarchus bilineatus Araguaia, Tapajós, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Capim, lower Orinoco, upper Orinoco, Apure Procerusternarchus pixuna* Endemic – Negro Racenisia fimbriipinna Negro, upper Orinoco Rhamphichthyidae 23 Gymnorhamphichthys bogardusae Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco, Apure Gymnorhamphichthys hypostomus Araguaia, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Marañon-Nanay, Napo-Ambyiacu, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco Gymnorhamphichthys rondoni Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Coari-Urucu, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, upper Orinoco Gymnorhamphichthys rosamariae Lower Xingu, Tapajós, Jutaí, Negro, Amazonas main channel, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco, Paraná-Paraguay Hypopygus benoneae* Endemic – Anapu Hypopygus cryptogenes Negro, upper Orinoco Hypopygus hoedemani* Endemic – Negro Hypopygus lepturus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Paraná-Paraguay Hypopygus minissimus Negro, upper Orinoco Hypopygus neblinae Negro, Corentyne-Demerara, lower Orinoco, upper Orinoco, Apure Hypopygus nijsseni* Tefé, Amazonas main channel 142 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Hypopygus ortegai* Ucayali, Amazonas main channel Hypopygus varii* Endemic – Trombetas Iracema caiana* Endemic – Amazonas main channel Rhamphichthys drepanium Mamoré, Tefé, Negro, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Rhamphichthys heleios* Mamoré, Amazonas main channel Rhamphichthys lineatus* Ucayali, Amazonas main channel Rhamphichthys longior** Trombetas, Curuá-Paru do Oeste Rhamphichthys pantherinus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Parnaíba, ItapicuruMearim, Araguari-Macari-Amapá, upper Orinoco Rhamphichthys rostratus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Tefé, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Araguari-Macari-Amapá, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, upper Orinoco Steatogenys duidae Middle-lower Madeira, Negro, Branco, Curuá-Paru do Oeste, Amazonas main channel, Capim, lower Orinoco, upper Orinoco Steatogenys elegans* Lower Tocantins, Tapajós, middle-lower Madeira, Purus, Putumayo, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure Steatogenys ocellatus Anapu, Pacajá, Tapajós, middle-lower Madeira, Purus, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco Sternopygidae 29 Archolaemus blax* Upper Tocantins, lower Tocantins, Branco Archolaemus ferreirai Branco, Capim Archolaemus janeae* Upper Xingu, Iriri, lower Xingu, Jamanxim Archolaemus luciae Upper Xingu, Teles Pires, Tapajós, Jamanxim, Trombetas, Jari, Araguari-MacariAmapá Archolaemus santosi* Endemic – Middle-lower Madeira Distocyclus conirostris Lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Negro, Trombetas, Amazonas main channel, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco, Apure Eigenmannia antonioi* Endemic – Anapu Eigenmannia limbata Middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, MarañonNanay, Napo-Ambyiacu, Negro, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Eigenmannia loretana* Ucayali, Marañon-Nanay Eigenmannia macrops Araguaia, upper Xingu, Iriri, Mamoré, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Branco, Parnaíba, Essequibo, lower Orinoco, upper Orinoco, Apure Eigenmannia matintapereira* Endemic – Negro Eigenmannia muirapinima* Endemic – Amazonas main channel Eigenmannia nigra Araguaia, lower Tocantins, Napo-Ambyiacu, Negro, Amazonas main channel, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco, Apure 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 143 Occurrence Eigenmannia vicentespelaea* Endemic – upper Tocantins Eigenmannia virescens Upper Tocantins, Araguaia, upper Xingu, lower Xingu, Guaporé, Beni-Madre de Dios, Ucayali, Putumayo, Japurá, Branco, Trombetas, Amazonas main channel, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, CaucaMagdalena-Sinu, Paraná-Paraguay Eigenmannia waiwai* Endemic – Trombetas Rhabdolichops caviceps Upper Tocantins, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, NapoAmbyiacu, Japurá, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Rhabdolichops eastwardi Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Napo-Ambyiacu, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Rhabdolichops electrogrammus Middle-lower Madeira, Negro, Branco, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Rhabdolichops lundbergi* Endemic – Amazonas main channel Rhabdolichops navalha* Tefé, Amazonas main channel Rhabdolichops nigrimans* Tefé, Negro, Amazonas main channel Rhabdolichops stewarti Teles Pires, Tapajós, lower Orinoco, upper Orinoco Rhabdolichops troscheli Lower Tocantins, Mamoré, Guaporé, middle-lower Madeira, Purus, Tefé, Negro, Branco, Trombetas, Amazonas main channel, lower Orinoco Sternopygus astrabes Tefé, Negro, Urubu-Uatumã, Amazonas main channel, upper Orinoco Sternopygus branco* Tefé, Japurá, Negro, Trombetas, Amazonas main channel Sternopygus macrurus Upper Tocantins, Araguaia, lower Tocantins, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Tefé, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, UrubuUatumã, Trombetas, Jari, Amazonas main channel, Parnaíba, Itapicuru-Mearim, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Cauca-Magdalena-Sinu, ParanáParaguay Sternopygus obtusirostris* Lower Tocantins, Tefé, Negro, Branco, Amazonas main channel Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu Sternopygus xingu* Batrachoidiformes 2 Batrachoididae 2 Potamobatrachus trispinosus* Restricted to Tocantins basin (Araguaia, lower Tocantins) Thalassophryne amazonica* Lower Xingu, Marañon-Nanay, Putumayo, Negro, Amazonas main channel Cyprinodontiformes Anablepidae 166 1 Mamoré, Guaporé, Paraná-Paraguay Jenynsia alternimaculata Cynolebiidae 141 Anablepsoides amanan* Endemic – Amazonas main channel Anablepsoides atratus* Middle-lower Madeira, Purus, Juruá, Amazonas main channel Anablepsoides beniensis* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Anablepsoides cajariensis* Endemic – Amazonas Estuary Anablepsoides chapare* Endemic – Mamoré 144 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Anablepsoides christinae* Endemic – Beni-Madre de Dios Anablepsoides derhami* Endemic – Ucayali Anablepsoides elongatus* Endemic – Ucayali Anablepsoides erberi* Endemic – Napo-Ambyiacu Anablepsoides gamae* Endemic – Jari Anablepsoides henschelae* Endemic – Negro Anablepsoides hoetmeri* Endemic – Purus Anablepsoides intermittens* Endemic – Amazonas main channel Anablepsoides iridescens* Endemic – Ucayali Anablepsoides jari* Endemic – Jari Anablepsoides jucundus* Endemic – Marañon Anablepsoides limoncochae* Napo-Ambyiacu, Japurá, Amazonas main channel Anablepsoides lineasoppilatae* Endemic – Beni-Madre de Dios Anablepsoides luitalimae* Endemic – Middle-lower Madeira Anablepsoides micropus* Negro, Amazonas main channel Anablepsoides monticola* Endemic – Marañon Anablepsoides ophiomimus* Napo-Ambyiacu, Japurá, Amazonas main channel Anablepsoides ornatus* Middle-lower Madeira, Negro, Branco, Trombetas, Amazonas main channel Anablepsoides ottonii* Endemic – Negro Anablepsoides parlettei* Endemic – Beni-Madre de Dios Anablepsoides peruanus** Anablepsoides roraima* Endemic – Branco Anablepsoides rubrolineatus* Ucayali, Putumayo, Amazonas main channel Anablepsoides speciosus* Endemic – Amazonas main channel Anablepsoides stagnatus Branco, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Anablepsoides taeniatus* Purus, Japurá, Negro, Amazonas main channel Anablepsoides tocantinensis* Endemic – upper Tocantins Anablepsoides urophthalmus Upper Tocantins, lower Tocantins, lower Xingu, Putumayo, Jari, Amazonas main channel, Itapicuru-Mearim, Gurupi-Turiaçu, Capim Anablepsoides urubuiensis* Endemic – Urubu-Uatumã Anablepsoides xanthonotus** Anablepsoides xinguensis* Endemic – lower Xingu Aphyolebias boticarioi* Endemic – Purus Aphyolebias claudiae* Endemic – Guaporé Aphyolebias manuensis* Endemic – Beni-Madre de Dios Aphyolebias obliquus* Endemic – Mamoré Aphyolebias peruensis* Napo-Ambyiacu, Amazonas main channel Aphyolebias rubrocaudatus* Endemic – Beni-Madre de Dios Aphyolebias schleseri* Endemic – Amazonas main channel 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species Occurrence Aphyolebias wischmanni* Endemic – Ucayali Austrofundulus rupununi* Endemic – Branco Austrolebias accorsii* Endemic – Mamoré Austrolebias monstrosus Guaporé, Paraná-Paraguay Austrolebias vandenbergi Guaporé, Paraná-Paraguay Cynolebias griseus* Endemic – upper Tocantins Hypsolebias brunoi* Endemic – upper Tocantins Hypsolebias flammeus* Endemic – upper Tocantins Hypsolebias marginatus* Endemic – upper Tocantins Hypsolebias multiradiatus* Endemic – upper Tocantins Hypsolebias notataus* Endemic – upper Tocantins Hypsolebias radiosus* Endemic – upper Tocantins Hypsolebias tocantinensis* Endemic – upper Tocantins Laimosemion amanapira* Endemic – Negro Laimosemion dibaphus* Tapajós, Trombetas Laimosemion gili* Endemic – Negro Laimosemion kirovskyi* Endemic – Urubu-Uatumã Laimosemion jauaperi* Endemic – Negro Laimosemion leticia* Endemic – Amazonas main channel Laimosemion rectocaudatus* Endemic – Amazonas main channel Laimosemion romeri* Endemic – Negro Laimosemion tecminae Negro, upper Orinoco Laimosemion uakti* Endemic – Negro Laimosemion uatuman* Endemic – Urubu-Uatumã Laimosemion ubim* Endemic – Japurá Maratecoara formosa* Restricted to Tocantins basin (upper Tocantins, Araguaia) Maratecoara gesmonei* Endemic – upper Xingu Maratecoara lacortei* Endemic – Araguaia Maratecoara splendida* Endemic – upper Tocantins Melanorivulus britzkei* Endemic – Teles Pires Melanorivulus canesi* Endemic – Iriri Melanorivulus crixas* Endemic – Araguaia Melanorivulus ignescens* Endemic – Araguaia Melanorivulus imperatrizensis* Endemic – upper Tocantins Melanorivulus jalapensis* Endemic – upper Tocantins Melanorivulus javahe* Endemic – Araguaia Melanorivulus karaja* Endemic – Araguaia Melanorivulus kayabi* Upper Xingu, Teles Pires, Juruena 145 146 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Melanorivulus kayapo* Endemic – Araguaia Melanorivulus kunzei* Endemic – Araguaia Melanorivulus litteratus* Endemic – Araguaia Melanorivulus megaroni* Endemic – upper Xingu Melanorivulus modestus* Endemic – Juruena Melanorivulus petrisecundi* Endemic – Araguaia Melanorivulus pictus Araguaia, Paraná-Paraguay Melanorivulus pindorama* Endemic – upper Tocantins Melanorivulus planaltinus* Endemic – upper Tocantins Melanorivulus rubromarginatus* Endemic – Araguaia Melanorivulus rubroreticulatus* Endemic – lower Xingu Melanorivulus salmonicaudus* Endemic – Araguaia Melanorivulus schuncki* Endemic – Jari Melanorivulus spixi* Endemic – Araguaia Melanorivulus ubirajarai* Endemic – Araguaia Melanorivulus violaceus* Endemic – Araguaia Melanorivulus wallacei* Endemic – upper Tocantins Melanorivulus zygonectes* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Juruena Moema apurinan* Endemic – Purus Moema beucheyi* Endemic – Guaporé Moema hellneri* Endemic – Napo-Ambyiacu Moema kenwoodi* Endemic – Beni-Madre de Dios Moema nudifrontata* Endemic – Branco Moema pepotei* Restricted to Madeira basin (Mamoré, Guaporé) Moema portugali* Endemic – Branco Moema quiii* Endemic – Beni-Madre de Dios Moema staecki* Endemic – Amazonas main channel Neofundulus guaporensis* Endemic – Guaporé Neofundulus ornatipinnis Mamoré, Paraná-Paraguay Neofundulus splendidus* Endemic – Mamoré Papiliolebias ashleyae* Endemic – Guaporé Papiliolebias francescae* Endemic – Guaporé Papiliolebias habluetzeli* Endemic – Mamoré Pituna compacta* Restricted to Tocantins basin (upper Tocantins, Araguaia) Pituna obliquoseriata* Endemic – Araguaia Pituna poranga* Endemic – Araguaia Pituna xinguensis* Restricted to Xingu basin (upper Xingu, lower Xingu) Plesiolebias altamira* Endemic – lower Xingu 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species Occurrence Plesiolebias aruana* Endemic – Araguaia Plesiolebias canabravensis* Endemic – upper Tocantins Plesiolebias filamentosus* Endemic – upper Tocantins Plesiolebias fragilis* Endemic – Araguaia Plesiolebias lacerdai* Endemic – Araguaia Plesiolebias xavantei* Endemic – upper Tocantins Pterolebias longipinnis Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel, Amazonas Estuary, Paraná-Paraguay Rivulus staecki* Endemic – Negro Simpsonichthys cholopteryx* Endemic – Araguaia Spectrolebias bellidoi* Endemic – Mamoré Spectrolebias brousseaui* Restricted to Madeira basin (Mamoré, Guaporé) Spectrolebias costae* Endemic – Araguaia Spectrolebias filamentosus* Restricted to Madeira basin (Mamoré, Guaporé) Spectrolebias gracilis* Endemic – upper Tocantins Spectrolebias inaequipinnatus* Endemic – upper Tocantins Spectrolebias pilleti* Endemic – Mamoré Spectrolebias reticulatus* Endemic – lower Xingu Spectrolebias semiocellatus* Endemic – Araguaia Trigonectes macrophthalmus* Endemic – Guaporé Trigonectes rogoaguae* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Trigonectes rubromarginatus* Endemic – Araguaia Trigonectes strigabundus* Endemic – upper Tocantins Cyprinodontidae 7 Orestias agassizii* Beni-Madre de Dios, Ucayali Orestias empyraeus* Endemic – Ucayali Orestias gymnota* Endemic – Marañon Orestias jussiei* Endemic – Ucayali Orestias munda* Endemic – Ucayali Orestias pentlandii* Endemic – Ucayali Orestias polonorum* Endemic – Ucayali Poeciliidae 17 Cnesterodon septentrionalis* Endemic – Araguaia Fluviphylax obscurus Negro, lower Orinoco, upper Orinoco Fluviphylax pygmaeus Middle-lower Madeira, Amazonas main channel, upper Orinoco Fluviphylax simplex* Endemic – Amazonas main channel Fluviphylax zonatus* Endemic – Negro Pamphorichthys akroa* Endemic – upper Tocantins Pamphorichthys araguaiensis* Upper Tocantins, Araguaia, upper Xingu, lower Xingu, Trombetas 147 148 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Pamphorichthys minor* Upper Xingu, lower Xingu, Guaporé, Amazonas main channel Pamphorichthys scalpridens* Upper Tocantins, lower Tocantins, Teles Pires, Juruena, Tapajós, Amazonas main channel Phalloceros leticiae* Endemic – Araguaia Poecilia bifurca** Poecilia caucana** Poecilia branneri** Poecilia parae Lower Xingu, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, lower Orinoco Poecilia picta Amazonas Estuary, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, lower Orinoco Poecilia waiapi* Endemic – Jari Tomeurus gracilis Amazonas Estuary, Capim, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco Atheriniformes 1 Atherinopsidae 1 Mamoré, Paraná-Paraguay Odontesthes bonariensis Beloniformes 9 Belonidae 8 Belonion apodion Guaporé, middle-lower Madeira, Purus, Japurá, Branco, Trombetas, Amazonas main channel, upper Orinoco Belonion dibranchodon Japurá, Negro, upper Orinoco Potamorrhaphis eigenmanni Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel, Paraná-Paraguay Potamorrhaphis guianensis Araguaia, lower Tocantins, upper Xingu, Tapajós, middle-lower Madeira, Javari, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Potamorrhaphis labiatus* Juruá, Ucayali, Amazonas main channel Potamorrhaphis petersi Negro, Trombetas, lower Orinoco, upper Orinoco, Apure Pseudotylosurus angusticeps Lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Branco, Amazonas main channel, Amazonas Estuary, Paraná-Paraguay Pseudotylosurus microps Upper Tocantins, Araguaia, lower Tocantins, lower Xingu, middle-lower Madeira, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Parnaíba, Maroni-Approuague, CoppenameSuriname-Saramacca, Essequibo, lower Orinoco, upper Orinoco, Apure Hemiramphidae 1 Amazonas main channel Hyporhamphus brederi Synbranchiformes 3 Synbranchidae 3 Synbranchus lampreia* Endemic – Amazonas Estuary Synbranchus madeirae* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Juruá 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 149 Occurrence Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, Purus, Tefé, Juruá, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, ItapicuruMearim, Gurupi-Turiaçu, Capim, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure, Maracaibo, Atl. Coastal Drainages of Col. Ven., Cauca-Magdalena-Sinu, Atrato, ParanáParaguay Synbranchus marmoratus* Perciformes 286 Cichlidae 258 Acarichthys heckelii Lower Tocantins, lower Xingu, Tapajós, middle-lower Madeira, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Essequibo Acaronia nassa Lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, BeniMadre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Jutaí, Ucayali, Marañon-Nanay, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Oiapoque, Corentyne-Demerara, Essequibo Acaronia vultuosa Madeira Shield Tributaries, Purus, Negro, Branco, lower Orinoco, upper Orinoco, Apure Aequidens chimantanus Japurá, Amazonas main channel, lower Orinoco Aequidens diadema Juruá, Ucayali, Putumayo, Negro, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Aequidens epae Teles Pires, Tapajós, Capim Aequidens gerciliae* Endemic – Madeira Shield Tributaries Aequidens mauesanus* Middle-lower Madeira, Amazonas main channel Aequidens metae Putumayo, Japurá, upper Orinoco Aequidens michaeli* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Aequidens pallidus Middle-lower Madeira, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Capim Aequidens patricki* Endemic – Ucayali Aequidens plagiozonatus Guaporé, Paraná-Paraguay Aequidens potaroensis Putumayo, Branco, Essequibo Aequidens rondoni Juruena, Paraná-Paraguay Aequidens superomaculatum Negro, upper Orinoco Aequidens tetramerus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Javari, Ucayali, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Gurupi-Turiaçu, Capim, Oiapoque, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Aequidens tubicen* Endemic – Trombetas Aequidens viridis* Restricted to Madeira basin (Mamoré, Guaporé) Apistogramma acrensis* Middle-lower Madeira, Purus Apistogramma agassizii Lower Tocantins, Tapajós, Mamoré, Guaporé, middle-lower Madeira, Purus, Javari, Ucayali, Marañon-Nanay, Japurá, Trombetas, Amazonas main channel, Capim 150 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Apistogramma aguarico* Endemic – Napo-Ambyiacu Apistogramma alacrina Japurá, upper Orinoco Apistogramma allpahuayo* Endemic – Amazonas main channel Apistogramma amoena** Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios) Apistogramma angayuara* Endemic – Trombetas Apistogramma arua* Endemic – Amazonas main channel Apistogramma atahualpa* Endemic – Amazonas main channel Apistogramma baenschi* Endemic – Marañon Apistogramma barlowi* Endemic – Napo-Ambyiacu Apistogramma bitaeniata* Marañon-Nanay, Putumayo, Japurá, Amazonas main channel Apistogramma brevis Negro, upper Orinoco Apistogramma cacatuoides* Ucayali, Marañon-Nanay, Amazonas main channel Apistogramma caetei Lower Tocantins, Gurupi-Turiaçu, Capim Apistogramma cinilabra* Endemic – Amazonas main channel Apistogramma commbrae Mamoré, Paraná-Paraguay Apistogramma cruzi* Napo-Ambyiacu, Putumayo, Japurá Apistogramma diplotaenia Negro, Amazonas main channel, upper Orinoco Apistogramma eleutheria* Endemic – Iriri Apistogramma elizabethae* Endemic – Negro Apistogramma eremnopyge* Endemic – Amazonas main channel Apistogramma erythrura* Endemic – Mamoré Apistogramma eunotus* Araguaia, Javari, Ucayali, Marañon-Nanay, Putumayo, Japurá, Amazonas main channel Apistogramma flavipedunculata* Endemic – Iriri Apistogramma geisleri* Endemic – Trombetas Apistogramma gephyra* Lower Xingu, Mamoré, middle-lower Madeira, Japurá, Negro, Branco, Amazonas main channel Apistogramma gibbiceps* Restricted to Negro basin (Negro, Branco) Apistogramma hippolytae* Negro, Branco, Amazonas main channel Apistogramma huascar* Endemic – Marañon Apistogramma inconspicua Mamoré, Guaporé, Beni-Madre de Dios, Japurá, Paraná-Paraguay Apistogramma iniridae Putumayo, Japurá, Negro, upper Orinoco Apistogramma juruensis* Endemic – Juruá Apistogramma kullanderi* Endemic – Iriri Apistogramma linkei* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Apistogramma luelingi* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Apistogramma martini* Endemic – Marañon Apistogramma megastoma* Endemic – Amazonas main channel Apistogramma meinkeni* Restricted to Negro basin (Negro, Branco) 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species Occurrence Apistogramma mendezi* Endemic – Negro Apistogramma moae* Endemic – Juruá Apistogramma nijsseni* Endemic – Ucayali Apistogramma norberti* Endemic – Amazonas main channel Apistogramma ortegai* Endemic – Napo-Ambyiacu Apistogramma ortmanni Japurá, Branco, Amazonas main channel, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Apistogramma panduro* Endemic – Amazonas main channel Apistogramma pantalone* Endemic – Marañon Apistogramma paucisquamis* Negro, Amazonas main channel Apistogramma paulmuelleri* Endemic – Amazonas main channel Apistogramma payaminonis* Endemic – Napo-Ambyiacu Apistogramma personata* Endemic – Negro Apistogramma pertensis* Tapajós, Tefé, Japurá, Negro, Trombetas, Jari, Amazonas main channel Apistogramma playayacu* Endemic – Napo-Ambyiacu Apistogramma pulchra* Endemic – Middle-lower Madeira Apistogramma regani* Lower Xingu, Negro, Branco, Amazonas main channel Apistogramma resticulosa Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Negro, Amazonas main channel, Araguari-Macari-Amapá Apistogramma rositae* Endemic – Marañon Apistogramma rubrolineata* Restricted to Madeira basin (Beni-Madre de Dios, middle-lower Madeira) Apistogramma rupununi Branco, Essequibo Apistogramma salpinction* Endemic – Trombetas Apistogramma similis* Restricted to Madeira basin (Mamoré, Beni-Madre de Dios) Apistogramma sororcula* Endemic – Guaporé Apistogramma staecki* Restricted to Madeira basin (Mamoré, Guaporé, middle-lower Madeira) Apistogramma steindachneri Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo Apistogramma taeniata* Endemic – Tapajós Apistogramma trifasciata Mamoré, Guaporé, Beni-Madre de Dios, Paraná-Paraguay Apistogramma tucurui Lower Tocantins, Capim Apistogramma uaupesi Negro, upper Orinoco Apistogramma urteagai* Endemic – Beni-Madre de Dios Apistogramma wapisana* Endemic – Branco Apistogrammoides pucallpaensis* Marañon-Nanay, Amazonas main channel Astronotus crassipinnis Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Branco, Amazonas main channel, Amazonas Estuary, Paraná-Paraguay Astronotus ocellatus Upper Tocantins, Araguaia, lower Tocantins, Tapajós, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, lower Orinoco 151 152 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Biotodoma cupido Araguaia, lower Tocantins, Teles Pires, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Corentyne-Demerara, Essequibo Biotodoma wavrini Japurá, Negro, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Biotoecus dicentrarchus Negro, lower Orinoco, upper Orinoco Biotoecus opercularis* Middle-lower Madeira, Purus, Negro, Branco, Trombetas, Amazonas main channel Bujurquina apoparuana* Endemic – Ucayali Bujurquina cordemadi* Restricted to Madeira basin (Beni-Madre de Dios, middle-lower Madeira) Bujurquina eurhinus* Restricted to Madeira basin (Beni-Madre de Dios, middle-lower Madeira) Bujurquina hophrys* Endemic – Ucayali Bujurquina huallagae* Marañon-Nanay, Putumayo Bujurquina labiosa* Endemic – Ucayali Bujurquina mariae Putumayo, Japurá, Amazonas main channel, lower Orinoco, upper Orinoco Bujurquina megalospilus* Endemic – Ucayali Bujurquina moriorum* Ucayali, Amazonas main channel Bujurquina ortegai* Marañon-Nanay, Putumayo Bujurquina pardus* Endemic – Marañon Bujurquina peregrinabunda* Putumayo, Japurá, Amazonas main channel Bujurquina robusta* Juruá, Ucayali Bujurquina syspilus* Ucayali, Amazonas main channel Bujurquina tambopatae* Endemic – Beni-Madre de Dios Bujurquina vittata Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Paraná-Paraguay Bujurquina zamorensis* Endemic – Marañon Caquetaia myersi* Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Amazonas main channel Caquetaia spectabilis Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Capim, Araguari-Macari-Amapá Chaetobranchopsis orbicularis* Lower Tocantins, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Negro, Branco, Amazonas main channel, Amazonas Estuary Chaetobranchus flavescens Araguaia, lower Tocantins, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Napo-Ambyiacu, Putumayo, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Oiapoque, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Chaetobranchus semifasciatus* Middle-lower Madeira, Purus, Tefé, Japurá, Amazonas main channel Cichla jariina* Endemic – Jari Cichla kelberi* Upper Tocantins, Araguaia, lower Tocantins, Amazonas Estuary Cichla melaniae* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Cichla mirianae* Upper Xingu, lower Xingu, Teles Pires, Juruena, Jamanxim Cichla monoculus Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Ucayali, NapoAmbyiacu, Putumayo, Negro, Branco, Amazonas main channel, Amazonas Estuary, Araguari-Macari-Amapá, Oiapoque, Coppename-Suriname-Saramacca, CorentyneDemerara, upper Orinoco 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 153 Occurrence Cichla nigromaculata* Endemic – Negro Cichla ocellaris Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo Cichla orinocensis Negro, Branco, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Cichla pinima Lower Tocantins, lower Xingu, Teles Pires, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Amazonas Estuary, Capim Cichla piquiti* Restricted to Tocantins basin (upper Tocantins, Araguaia, lower Tocantins) Cichla pleiozona* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Amazonas main channel Cichla temensis Negro, Branco, Urubu-Uatumã, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Cichla thyrorus* Endemic – Trombetas Cichla vazzoleri* Urubu-Uatumã, Trombetas Cichlasoma amazonarum* Upper Tocantins, lower Tocantins, Mamoré, Guaporé, middle-lower Madeira, Juruá, Ucayali, Marañon-Nanay, Putumayo, Amazonas main channel, Amazonas Estuary Cichlasoma araguaiense* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu Cichlasoma bimaculatum Lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, Branco, Amazonas main channel, Oiapoque, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco Cichlasoma boliviense* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus Cichlasoma sanctifranciscense Upper Tocantins, Parnaiba, São Francisco Crenicara latruncularium* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios) Crenicara punctulata Lower Tocantins, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Branco, Amazonas main channel, Essequibo Crenicichla acutirostris Upper Xingu, lower Xingu, Teles Pires, Tapajós, Madeira Shield Tributaries, Branco, Amazonas main channel, Essequibo Crenicichla adspersa* Lower Tocantins, Tapajós, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Urubu-Uatumã, Amazonas main channel Crenicichla albopunctata Jari, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara Crenicichla alta Putumayo, Japurá, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco, upper Orinoco Crenicichla anamiri* Endemic – lower Xingu Crenicichla anthurus* Ucayali, Marañon-Nanay, Putumayo, Japurá, Branco Crenicichla cametana* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu Crenicichla chicha* Endemic – Juruena Crenicichla cincta* Middle-lower Madeira, Madeira Shield Tributaries, Purus, Napo-Ambyiacu, Putumayo, Amazonas main channel, Amazonas Estuary Crenicichla compressiceps* Endemic – lower Tocantins Crenicichla cyanonotus Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Ucayali, Amazonas main channel, Itapicuru-Mearim, Apure Crenicichla cyclostoma* Endemic – lower Tocantins Crenicichla dandara* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Crenicichla heckeli* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Trombetas 154 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Crenicichla hemera* Endemic – Madeira Shield Tributaries Crenicichla hummelincki* Endemic – Trombetas Crenicichla inpa Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, middle-lower Madeira, Negro, Amazonas main channel, Amazonas Estuary, Capim Crenicichla isbrueckeri* Endemic – Madeira Shield Tributaries Crenicichla jegui* Endemic – lower Tocantins Crenicichla johanna Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Tapajós, Jamanxim, Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Tefé, Juruá, Ucayali, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Araguari-Macari-Amapá, Oiapoque, Corentyne-Demerara, Essequibo, upper Orinoco Crenicichla labrina Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Capim, AraguariMacari-Amapá Crenicichla lenticulata Putumayo, Japurá, Negro, Branco, Trombetas, Essequibo, upper Orinoco Crenicichla lepidota Araguaia, Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Capim, Paraná-Paraguay Crenicichla lucius* Ucayali, Marañon-Nanay, Napo-Ambyiacu, Amazonas main channel Crenicichla lugubris Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Guaporé, middle-lower Madeira, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas main channel, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco Crenicichla macrophthalma* Upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Negro, Trombetas, Amazonas main channel Crenicichla marmorata* Lower Tocantins, Iriri, lower Xingu, middle-lower Madeira, Madeira Shield Tributaries, Trombetas, Jari, Amazonas main channel Crenicichla monicae* Endemic – Negro Crenicichla notophthalmus Negro, Amazonas main channel, Capim Crenicichla pellegrini* Endemic – Madeira Shield Tributaries Crenicichla percna* Restricted to Xingu basin (upper Xingu, lower Xingu) Crenicichla phaiospilus* Endemic – upper Xingu Crenicichla ploegi Juruena, Paraná-Paraguay Crenicichla proteus* Mamoré, Beni-Madre de Dios, Purus, Juruá, Ucayali, Marañon-Nanay, NapoAmbyiacu, Putumayo, Japurá, Amazonas main channel Crenicichla pydanielae* Endemic – Trombetas Crenicichla regani Araguaia, lower Tocantins, Teles Pires, Tapajós, Mamoré, Guaporé, middlelower Madeira, Madeira Shield Tributaries, Purus, Tefé, Branco, Urubu-Uatumã, Trombetas, Curuá-Paru do Oeste, Jari, Amazonas main channel, Amazonas Estuary, Capim Crenicichla reticulata Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Napo-Ambyiacu, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Gurupi-Turiaçu, Essequibo, upper Orinoco Crenicichla rosemariae* Endemic – upper Xingu Crenicichla santosi* Mamoré, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Amazonas main channel 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 155 Occurrence Crenicichla saxatilis Upper Xingu, lower Xingu, Teles Pires, Putumayo, Japurá, Branco, Amazonas main channel, Araguari-Macari-Amapá, Oiapoque, Maroni-Approuague, CoppenameSuriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Crenicichla sedentaria Juruá, Ucayali, Marañon-Nanay, Napo-Ambyiacu, upper Orinoco Crenicichla semicincta* Teles Pires, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá Crenicichla stocki* Endemic – lower Tocantins Crenicichla strigata Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Capim, Essequibo Crenicichla tigrina* Endemic – Trombetas Crenicichla urosema* Restricted to Tapajós basin (Tapajós, Jamanxim) Crenicichla virgatula* Restricted to Negro basin (Negro, Branco) Crenicichla vittata Mamoré, Paraná-Paraguay Crenicichla wallacii Lower Tocantins, Negro, Branco, Essequibo, lower Orinoco, upper Orinoco Dicrossus filamentosus Negro, Branco, lower Orinoco, upper Orinoco Dicrossus foirni* Endemic – Negro Dicrossus maculatus* Middle-lower Madeira, Branco, Amazonas main channel Dicrossus warzeli* Restricted to Tapajós basin (Teles Pires, Tapajós) Geophagus abalios Negro, lower Orinoco, upper Orinoco, Apure Geophagus altifrons Upper Tocantins, lower Tocantins, upper Xingu, Iriri, lower Xingu, Tapajós, middlelower Madeira, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Capim Geophagus argyrostictus* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Geophagus dicrozoster Negro, lower Orinoco, upper Orinoco, Apure Geophagus megasema* Guaporé, Beni-Madre de Dios, middle-lower Madeira, Japurá Geophagus mirabilis* Endemic – Madeira Shield Tributaries Geophagus neambi* Upper Tocantins, Araguaia, lower Tocantins, lower Xingu, Teles Pires, Tapajós Geophagus proximus* Araguaia, lower Tocantins, upper Xingu, lower Xingu, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Ucayali, Japurá, Negro, Branco, Trombetas, Amazonas main channel Geophagus surinamensis Putumayo, Japurá, Branco, Maroni-Approuague, Coppename-Suriname-Saramacca, Essequibo Geophagus sveni* Restricted to Tocantins basin (upper Tocantins, Araguaia) Geophagus winemilleri Japurá, Negro, upper Orinoco Guianacara dacrya Branco, Essequibo Guianacara sphenozona Branco, Corentyne-Demerara, Essequibo Gymnogeophagus balzanii Guaporé, Paraná-Paraguay Heroina isonycterina* Marañon-Nanay, Napo-Ambyiacu, Japurá Heros efasciatus Lower Tocantins, upper Xingu, lower Xingu, Tapajós, Madeira Shield Tributaries, Tefé, Juruá, Ucayali, Urubu-Uatumã, Trombetas, Amazonas main channel, Oiapoque Heros notatus Upper Tocantins, Araguaia, Negro, Branco, Amazonas main channel, Capim, Araguari-Macari-Amapá, Maroni-Approuague, Corentyne-Demerara, Essequibo 156 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Species NO. 431 Occurrence Heros severus Negro, upper Orinoco Heros spurius* Restricted to Madeira basin (Mamoré, Guaporé, middle-lower Madeira) Hoplarchus psittacus Middle-lower Madeira, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Hypselecara coryphaenoides Tapajós, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Hypselecara temporalis Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Ucayali, Putumayo, Japurá, Branco, UrubuUatumã, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Capim, Oiapoque Ivanacara adoketa* Endemic – Negro Krobia xinguensis* Restricted to Xingu basin (upper Xingu, lower Xingu) Laetacara araguaiae Upper Tocantins, Araguaia, upper Xingu, Teles Pires, Tapajós, Paraná-Paraguay Laetacara curviceps* Trombetas, Amazonas main channel, Capim Laetacara flavilabris* Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Amazonas main channel Laetacara fulvipinnis Negro, upper Orinoco Laetacara thayeri* Middle-lower Madeira, Purus, Coari-Urucu, Tefé, Marañon-Nanay, Negro, Amazonas main channel Mesonauta acora Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, lower Xingu, Capim Mesonauta festivus Araguaia, lower Tocantins, upper Xingu, lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Putumayo, Japurá, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary, Araguari-Macari-Amapá, Essequibo, upper Orinoco, ParanáParaguay Mesonauta guyanae Lower Tocantins, Tapajós, middle-lower Madeira, Negro, Branco, Amazonas main channel, Amazonas Estuary, Capim, Oiapoque, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Mesonauta insignis Guaporé, Beni-Madre de Dios, middle-lower Madeira, Putumayo, Negro, Branco, Urubu-Uatumã, Amazonas main channel, lower Orinoco, upper Orinoco, Apure Mesonauta mirificus* Ucayali, Putumayo, Japurá, Amazonas main channel Mikrogeophagus altispinosus* Restricted to Madeira basin (Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira) Pterophyllum altum Negro, upper Orinoco Pterophyllum leopoldi Lower Tocantins, Putumayo, Negro, Branco, Amazonas main channel, Amazonas Estuary, Gurupi-Turiaçu, Essequibo Pterophyllum scalare Araguaia, lower Tocantins, lower Xingu, middle-lower Madeira, Purus, Javari, Ucayali, Putumayo, Japurá, Branco, Jari, Amazonas main channel, Amazonas Estuary, Capim, Oiapoque, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo Retroculus acherontos* Endemic – upper Tocantins Retroculus lapidifer Upper Tocantins, Araguaia, lower Tocantins, Capim Retroculus xinguensis* Upper Xingu, Iriri, lower Xingu, Tapajós, Jamanxim Rondonacara hoehnei* Endemic – Araguaia Satanoperca acuticeps* Beni-Madre de Dios, middle-lower Madeira, Purus, Japurá, Negro, Branco, UrubuUatumã, Trombetas, Amazonas main channel Satanoperca curupira* Endemic – Madeira Shield Tributaries Satanoperca daemon Putumayo, Japurá, Negro, lower Orinoco, upper Orinoco, Apure 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 157 Occurrence Satanoperca jurupari Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Mamoré, Guaporé, Beni-Madre de Dios, middlelower Madeira, Madeira Shield Tributaries, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Paru, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Capim, Araguari-Macari-Amapá, Oiapoque, Essequibo, upper Orinoco Satanoperca leucosticta Middle-lower Madeira, Madeira Shield Tributaries, Purus, Juruá, Branco, UrubuUatumã, Amazonas main channel, Corentyne-Demerara, Essequibo Satanoperca lilith* Middle-lower Madeira, Madeira Shield Tributaries, Negro, Branco, Trombetas, Amazonas main channel Satanoperca pappaterra Mamoré, Guaporé, Beni-Madre de Dios, Japurá, Amazonas main channel, ParanáParaguay Symphysodon aequifasciatus Upper Xingu, lower Xingu, Tapajós, Coari-Urucu, Tefé, Putumayo, Japurá, Negro, Trombetas, Amazonas main channel, Amazonas Estuary, Capim Symphysodon discus* Middle-lower Madeira, Negro, Trombetas Symphysodon haraldi* Middle-lower Madeira, Purus, Amazonas main channel Symphysodon tarzoo* Middle-lower Madeira, Tefé, Juruá, Jutaí, Amazonas main channel Taeniacara candidi* Lower Xingu, Tapajós, middle-lower Madeira, Negro, Branco, Trombetas, Amazonas main channel Tahuantinsuyoa chipi* Endemic – Ucayali Tahuantinsuyoa macantzatza* Endemic – Ucayali Teleocichla centisquama* Restricted to Xingu basin (Iriri, lower Xingu) Teleocichla centrarchus* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Teleocichla cinderella* Restricted to Tocantins basin (Araguaia, lower Tocantins) Teleocichla gephyrogramma* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Teleocichla monogramma* Restricted to Xingu basin (upper Xingu, lower Xingu) Teleocichla preta* Restricted to Xingu basin (upper Xingu, Iriri, lower Xingu) Teleocichla prionogenys* Iriri, Tapajós, Jamanxim Teleocichla proselytus* Upper Xingu, Tapajós Teleocichla wajapi* Endemic – Jari Uaru amphiacanthoides* Lower Xingu, Tapajós, middle-lower Madeira, Purus, Tefé, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Amazonas Estuary Eleotridae 9 Dormitator maculatus* Lower Tocantins, Amazonas Estuary Eleotris pisonis Lower Tocantins, lower Xingu, Amazonas main channel, Amazonas Estuary, Maroni-Approuague, Coppename-Suriname-Saramacca, lower Orinoco Leptophilypnion fittkaui* Endemic – Negro Leptophilypnion pusillus* Endemic – Tapajós Microphilypnus acangaquara* Endemic – Tapajós Microphilypnus hypolyrasimeion* Endemic – Negro Microphilypnus macrostoma Negro, Amazonas main channel, lower Orinoco, upper Orinoco Microphilypnus tapajosensis* Endemic – Tapajós Microphilypnus ternetzi Lower Tocantins, Teles Pires, Tapajós, Mamoré, middle-lower Madeira, Madeira Shield Tributaries, Purus, Japurá, Negro, Branco, Amazonas main channel, Capim, lower Orinoco, upper Orinoco 158 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Taxon Gobiidae Species Occurrence 1 Lower Tocantins, lower Xingu, Maroni-Approuague, Corentyne-Demerara, Essequibo, lower Orinoco, Cauca-Magdalena-Sinu, Atrato Awaous flavus Polycentridae NO. 431 3 Monocirrhus polyacanthus Lower Xingu, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Coari-Urucu, Tefé, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Capim, lower Orinoco, upper Orinoco, Apure Polycentrus jundia* Endemic – Negro Polycentrus schomburgkii Jari, Amazonas Estuary, Capim, Araguari-Macari-Amapá, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco, Apure Sciaenidae 15 Pachypops fourcroi Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Guaporé, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Itapicuru-Mearim, Araguari-Macari-Amapá, Oiapoque, Maroni-Approuague, Coppename-SurinameSaramacca, Corentyne-Demerara, Essequibo, lower Orinoco, upper Orinoco, Apure Pachypops pigmaeus* Middle-lower Madeira, Madeira Shield Tributaries, Branco, Trombetas Pachypops trifilis Mamoré, Guaporé, Purus, Tefé, Juruá, Japurá, Negro, Urubu-Uatumã, Trombetas, Amazonas main channel, Maroni-Approuague, Essequibo Pachyurus calhamazon* Branco, Trombetas Pachyurus gabrielensis Purus, Ucayali, Marañon-Nanay, Negro, Branco, upper Orinoco Pachyurus junki* Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Jamanxim, Madeira Shield Tributaries, Negro, Branco, Amazonas main channel Pachyurus paucirastrus* Upper Tocantins, Araguaia, lower Tocantins, Mamoré, middle-lower Madeira, Trombetas, Amazonas main channel Pachyurus schomburgkii Araguaia, lower Tocantins, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Juruá, Putumayo, Japurá, Negro, Branco, Urubu-Uatumã, Trombetas, Jari, Amazonas Estuary, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, lower Orinoco, upper Orinoco Pachyurus stewarti* Beni-Madre de Dios, Purus, Napo-Ambyiacu Petilipinnis grunniens Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, middlelower Madeira, Madeira Shield Tributaries, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel, Essequibo Plagioscion auratus Lower Tocantins, Amazonas main channel, Capim, Maroni-Approuague, Coppename-Suriname-Saramacca, Corentyne-Demerara, Essequibo, lower Orinoco Plagioscion magdalenae Lower Tocantins, Iriri, lower Xingu, Trombetas, Amazonas main channel, Amazonas Estuary, Capim, Araguari-Macari-Amapá, Cauca-Magdalena-Sinu Plagioscion montei* Middle-lower Madeira, Purus, Juruá, Ucayali, Negro, Branco, Urubu-Uatumã, Trombetas, Amazonas main channel Plagioscion squamosissimus Upper Tocantins, Araguaia, lower Tocantins, upper Xingu, Iriri, lower Xingu, Teles Pires, Juruena, Tapajós, Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Madeira Shield Tributaries, Purus, Tefé, Juruá, Marañon-Nanay, NapoAmbyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Parnaíba, Itapicuru-Mearim, Capim, Araguari-MacariAmapá, Maroni-Approuague, Coppename-Suriname-Saramacca, CorentyneDemerara, Essequibo, lower Orinoco, upper Orinoco, Apure 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Taxon Species 159 Occurrence Trombetas, Essequibo, Cauca-Magdalena-Sinu Plagioscion surinamensis Pleuronectiformes 9 Achiridae 9 Achirus achirus* Lower Tocantins Apionichthys asphyxiatus* Araguaia, Amazonas main channel Apionichthys dumerili Lower Tocantins, Japurá, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Itapicuru-Mearim, Coppename-Suriname-Saramacca, CorentyneDemerara, lower Orinoco Apionichthys menezesi Napo-Ambyiacu, Negro, lower Orinoco Apionichthys nattereri* Mamoré, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Javari, Marañon-Nanay, Napo-Ambyiacu, Negro, Amazonas main channel Apionichthys rosai* Middle-lower Madeira, Ucayali, Amazonas main channel, Amazonas Estuary Apionichthys seripierriae* Middle-lower Madeira, Napo-Ambyiacu, Branco, Amazonas main channel Hypoclinemus mentalis Araguaia, lower Tocantins, upper Xingu, lower Xingu, Teles Pires, Tapajós, Guaporé, Beni-Madre de Dios, middle-lower Madeira, Purus, Juruá, Javari, Ucayali, Marañon-Nanay, Napo-Ambyiacu, Putumayo, Japurá, Negro, Branco, Trombetas, Jari, Amazonas main channel, Amazonas Estuary, Essequibo, lower Orinoco, upper Orinoco Soleonasus finis Mamoré, middle-lower Madeira, Purus, Juruá, Ucayali, Marañon-Nanay, Branco, Amazonas main channel, Essequibo Tetraodontiformes 2 Tetraodontidae 2 Colomesus asellus Araguaia, lower Tocantins, upper Xingu, lower Xingu, middle-lower Madeira, Napo-Ambyiacu, Putumayo, Japurá, Branco, Trombetas, Amazonas main channel, Essequibo, lower Orinoco Colomesus tocantinensis* Lower Tocantins, Teles Pires Ceratodontiformes 1 Lepidosirenidae 1 Lepidosiren paradoxa Mamoré, Guaporé, Beni-Madre de Dios, Amazonas main channel, Amazonas Estuary, upper Orinoco, Paraná-Paraguay 160 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 431 APPENDIX 2 Test for “Amazon Core” distribution Statistical Test for Species with Distribution Maps A, Boullengerella spp., 176 records Expected inside: 88; observed inside: 176; expected outside: 88; observed outside: 0; Χ2 = 176. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Moenkhausia collettii, 113 records Expected inside: 56.5; observed inside: 113; expected outside: 56.5; observed outside: 0; Χ2 = 113. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Moenkhausia oligolepis, 104 records Expected inside: 52; observed inside: 104; expected outside: 52; observed outside: 0; Χ2 = 104. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. H0: The species is randomly distributed. H1: The species is not randomly distributed. Χ2 = Σ [ (O - E)2 / E ], where: E is the expected frequency (species can occur inside or outside of the defined limits of the distribution pattern. Thus, E is the number of records of a species divided by 2 possibilities [inside or outside]); andO is the Observed frequency (observed number of records of the species inside the defined limits of the distribution pattern). Degrees of freedom (2 - 1 =1). Conventionally accepted significance level of 0.05 (Chi square distribution table 3.841). Test for species with “Broadly distributed lineages” A, Hoplias malabaricus, 249 records Expected inside: 124.5; observed inside: 249; expected outside: 124.5; observed outside: 0; Χ2 = 249. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Erythrinus erythrinus, 41 records Expected inside: 20.5; observed inside: 41; expected outside: 20.5; observed outside: 0; Χ2 = 41. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Hoplerythrinus unitaeniatus, 95 records Expected inside: 47.5; Observed inside: 95; Expected outside: 47.5; Observed outside: 0; Χ2 = 95. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. D, Synbranchus marmoratus, 144 Expected inside: 72; observed inside: 144; expected outside: 72; observed outside: 0; Χ2 = 144. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. E, Callichthys callitchthys, 110 records Expected inside: 55; observed inside: 109; expected outside: 55; observed outside: 1; Χ2 = 106,03. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. Test for “Amazon and Orinoco Lowlands” distribution A, Moenkhausia lepidura, 51 records Expected inside: 25.5; observed inside: 51; expected outside: 25.5; observed outside: 0; Χ2 = 51. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Potamorhina altamazonica, 29 records Expected inside: 14.5; observed inside: 29; expected outside: 14.5; observed outside: 0; Χ2 = 29. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Vandellia cirrhosa, 75 records Expected inside: 37.5; observed inside: 74; expected outside: 37.5; observed outside: 1; Χ2 = 71. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. Test for “Amazon and Paraguay Lowlands” distribution A, Hemigrammus lunatus, 21 records Expected inside: 10.5; observed inside: 21; expected outside: 10.5; observed outside: 0; Χ2 = 21. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES B, Epapterus dispilurus, 12 records Expected inside: 6; observed inside: 12; expected outside: 6; observed outside: 0; Χ2 = 12. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Mesonauta festivus, 33 records Expected inside: 16.5; observed inside: 32; expected outside: 16.5; observed outside: 1; Χ2 = 29.12. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. Test for “Amazonas-Paraguay-Orinoco Lowland” distribution A, Rhaphiodon vulpinus, 152 records Expected inside: 76; observed inside: 143; expected outside: 76; observed outside: 9; Χ2 = 118.13. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Sorubim lima, 40 records Expected inside: 20; Observed inside: 39; Expected outside: 20; Observed outside: 1; Χ2 = 36.1. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Hypophthalmus oremaculatus, 39 records Expected inside: 19.5; Observed inside: 38; Expected outside: 19.5; Observed outside: 1; Χ2 = 35.1. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. 161 Expected inside: 7; observed inside: 14; expected outside: 7; observed outside: 0; Χ2 = 14. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. D, Hemigrammus ocellifer, 36 records Expected inside: 18; observed inside: 36; expected outside: 18; observed outside: 0; Χ2 = 36. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. Test for “Guyana Mangrove Province” distribution A, Curimata cyprinoides, 19 records Expected inside: 9.5; observed inside: 19; expected outside: 9.5; observed outside: 0; Χ2 = 19. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Cyphocharax helleri, 14 records Expected inside: 7; observed inside: 14; expected outside: 7; observed outside: 0; Χ2 = 14. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Polycentrus schomburgkii, 8 records Expected inside: 4; observed inside: 8; expected outside: 4; observed outside: 0; Χ2 = 8. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. Test for “Eastern Amazon” distribution Test for “Amazonas-Guyana-Orinoco Lowland” distribution A, Crenuchus spp., 44 records Expected inside: 22; observed inside: 44; expected outside: 22; observed outside: 0; Χ2 = 44. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Mesonauta spp., 64 records Expected inside: 32; observed inside: 55; expected outside: 32; observed outside: 9; Χ2 = 33. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Hemigrammus unilineatus, 14 records A, Synaptolaemus latofasciatus, 12 records Expected inside: 6; observed inside: 12; expected outside: 6; observed outside: 0; Χ2 = 12. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Aphanothorolus emarginatus, 46 records Expected inside: 23; observed inside: 42; expected outside: 23; observed outside: 4; Χ2 = 28. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Pachyurus junki, 12 records Expected inside: 6; observed inside: 12; expected outside: 6; observed outside: 0; Χ2 = 12. Χ2 >3.841, 162 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY rejects the null hypothesis, i.e., the species is not randomly distributed. Test for “Amazon-core uplands” distribution A, Cetopsidium spp., 41 records Expected inside: 20.5; observed inside: 32; expected outside: 20.5; observed outside: 9; Χ2 = 12.9. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Hemigrammus ora, 16 records Expected inside: 8; observed inside: 16; expected outside: 8; observed outside: 0; Χ2 = 16. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Hoplias aimara, 41 records Expected inside: 20.5; observed inside: 41; expected outside: 20.5; observed outside: 0; Χ2 = 41. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. Test for “Amazonian uplands” distribution A, Sartor spp., 4 records Expected inside: 2; observed inside: 4; expected outside: 2; observed outside: 0; Χ2 = 4. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Teleocichla spp., 24 records Expected inside: 12; observed inside: 24; expected outside: 12; observed outside: 0; Χ2 = 24. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Tocantinsia piresi, 11 records Expected inside: 5.5; observed inside: 11; expected outside: 5.5; observed outside: 0; Χ2 = 11. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. NO. 431 Test for “Longitudinal correspondence among Amazonian Shield versants” distribution A, Sartor gr. elongatus, 4 records Expected inside: 2; observed inside: 4; expected outside: 2; observed outside: 0; Χ2 = 4. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Bryconexodon spp., 14 records Expected inside: 7; observed inside: 14; expected outside: 7; observed outside: 0; Χ2 = 14. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Bivibranchia velox, 18 records Expected inside: 9; observed inside: 18; expected outside: 9; observed outside: 0; Χ2 = 18. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. Test for “Brazilian Shield” distribution A, Caiapobrycon spp., 18 records Expected inside: 9; observed inside: 18; expected outside: 9; observed outside: 0; Χ2 = 18. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Jupiaba apenima, 28 records Expected inside: 14; observed inside: 28; expected outside: 14; observed outside: 0; Χ2 = 28. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Moenkhausia gr. pankilopteryx/pirauba, 19 records Expected inside: 9.5; observed inside: 19; expected outside: 9.5; observed outside: 0; Χ2 = 19. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. Test for “cis-Andean foothills” distribution Test for “Guyana Shield” distribution A, Pseudancistrus brevispinis, 18 records Expected inside: 9; observed inside: 18; expected outside: 9; observed outside: 0; Χ2 = 18. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. A, Astyanacinus spp., 23 records Expected inside: 11.5; observed inside: 23; expected outside: 11.5; observed outside: 0; Χ2 = 23. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. B, Leporinus striatus, 71 records 2019 DAGOSTA & DE PINNA: DISTRIBUTION & BIOGEOGRAPHICAL PATTERNS OF AMAZON FISHES Expected inside: 35.5; observed inside: 50; expected outside: 35.5; observed outside: 21; Χ2 = 11.8. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. C, Steindachnerina dobula, 25 records Expected inside: 12.5; observed inside: 25; expected outside: 12.5; observed outside: 0; Χ2 = 25. Χ2 >3.841, rejects the null hypothesis, i.e., the species is not randomly distributed. Test for type of the water affecting distribution Ho: The distribution of the species is not affected by the type of the water. H1: The distribution of the species is affected by the type of the water. Χ2 = Σ [ (O - E)2 / E ], where: E is the expected frequency (species can occur in black-, white- or clear water). Thus, E is the number of records of a species divided by 3 possibilities (black-, white- or clear water); O is the observed frequency (number of records of the species observed in a determined water type). Degrees of freedom (3 - 1 = 2). Conventionally accepted significance level of 0.05 (Chi square distribution table 5.991). Test for “Amazon-only Lowland” distribution A, Cetopsis candiru, 9 records Expected black: 3; observed black: 0; expected white: 3; observed white: 8; expected clear: 3; observed clear: 1; Χ2 = 12.6. Χ2 > 5.991, rejects the null hypothesis, i.e., the distribution of the species is affected by the type of the water. B, Curimatella meyeri, 38 records Expected black: 12; observed black: 2; expected white: 12; observed white: 33; expected clear: 12; observed clear: 3; Χ2 = 51.8. Χ2 > 5.991, rejects the null hypothesis, i.e., the distribution of the species is affected by the type of the water. C, Adontosternarchus balaenops, 21 records Expected black: 7; observed black: 0; expected white: 7; observed white: 20; expected clear: 7; observed clear: 1; Χ2 = 21.16. Χ2 > 5.991, rejects the null hypothesis, i.e., the distribution of the species is affected by the type of the water. 163 Test for “Central Blackwater Amazon” distribution A, Biotoecus spp., 19 records Expected black: 6.33; observed black: 14; expected white: 6.33; observed white: 5; expected clear: 6.33; observed clear: 0; Χ2 = 8.38. Χ2 > 5.991, rejects the null hypothesis, i.e., the distribution of the species is affected by the type of the water. B, Dicrossus spp., 25 records (collecting points in Orinoco were not considered for the water type classification) Expected black: 8.33; observed black: 16; expected white: 8.33; observed white: 3; expected clear: 8.33; observed clear: 6; Χ2 = 7.72. Χ2 > 5.991, rejects the null hypothesis, i.e., the distribution of the species is affected by the type of the water. C, Hemigrammus analis, 18 records (collecting points in Orinoco were not considered for the water type classification) Expected black: 6; observed black: 13; expected white: 6; observed white: 1; expected clear: 6; observed clear: 4; Χ2 = 6.5. Χ2 >5.991, rejects the null hypothesis, i.e., the distribution of the species is affected by the type of the water. D, Hemigrammus coeruleus, 15 records Expected black: 5; observed black: 12; expected white: 5; observed white: 2; expected clear: 5; observed clear: 1; Χ2 = 6.16. Χ2 > 5.991, rejects the null hypothesis, i.e., the distribution of the species is affected by the type of the water. E, Hemigrammus stictus, 18 records (collecting points in Orinoco were not considered for the water type classification) Expected black: 6; observed black: 16; expected white: 6; observed white: 3; expected clear: 6; observed clear: 6; Χ2 = 10.16. Χ2 > 5.991, rejects the null hypothesis, i.e., the distribution of the species is affected by the type of the water. Scientific Publications of the American Museum of Natural History American Museum Novitates Bulletin of the American Museum of Natural History Anthropological Papers of the American Museum of Natural History Publications Committee Robert S. Voss, Chair Board of Editors Jin Meng, Paleontology Lorenzo Prendini, Invertebrate Zoology Robert S. Voss, Vertebrate Zoology Peter M. Whiteley, Anthropology Managing Editor Mary Knight Submission procedures can be found at http://research.amnh.org/scipubs All issues of Novitates and Bulletin are available on the web (http://digitallibrary.amnh. org/dspace). Order printed copies on the web from: http://shop.amnh.org/a701/shop-by-category/books/scientific-publications.html or via standard mail from: American Museum of Natural History—Scientific Publications Central Park West at 79th Street New York, NY 10024 This paper meets the requirements of ANSI/NISO Z39.48-1992 (permanence of paper). On the cover: Distribution patterns of Amazonian fishes: (upper left) Guiana Mangrove Province; (upper right) Amazononly Lowland; (lower left) Cis-Andean Foothills; (lower right) Eastern Amazon.

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THE FISHES OF THE AMAZON: DISTRIBUTION AND BIOGEOGRAPHICAL PATTERNS, WITH A COMPREHENSIVE LIST OF SPECIES

THE FISHES OF THE AMAZON: DISTRIBUTION AND BIOGEOGRAPHICAL PATTERNS, WITH A COMPREHENSIVE LIST OF SPECIES

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