Introductions of non-native fishes into a heavily modified river: rates,patterns and management issues in the Paranapanema River (Upper Paraná ecoregion,Brazil)

Understanding the pathways and impacts of non-native species is important for helping prevent new introductions and invasions. This is frequently challenging in regions where human activities continue to promote new introductions, such as in Brazil, where aquaculture and sport fishing are mainly dependent on non-native fishes. Here, the non-native fish diversity of the Paranapanema River basin of the Upper Paraná River ecoregion, Brazil, was quantified fully for the first time. This river has been subject to considerable alteration through hydroelectric dam construction and concomitant development of aquaculture and sport fishing. Through compilation of a non-native fish inventory by literature review, with complementary records from recent field studies, analyses were completed on the timings of introduction, and the taxonomy, origin and introduction vectors of the non-native fishes. A total of 47 non-native fishes are now present across the basin. Of these, 24 invaded from the Lower Paraná River following construction of Itaipu Dam that connected previously unconnected fish assemblages. Activities including fish stocking, aquaculture and sport angling continue to result in new introductions. Discounting Itaipu invasions, the introduction rate between 1950 and 2014 was approximately one new introduction every 3 years. Introduced fish were mainly of the Cichlidae and Characidae families; most species were from other South American ecoregions, but fishes of African, Asian, North American and Central American origin were also present. These introductions have substantially modified the river’s fish fauna; when coupled with altered lentic conditions caused by impoundment, this suggests that the river’s native fishes are increasingly threatened.     

The use of an integrative approach to improve accuracy of species identification and detection of new species in studies of stream fish diversity

Genetica - In this study, we made an inventory of the stream and headwater ichthyofauna of the left bank of the Itaipu Dam Reservoir, located in the lower part of the Upper Paraná River basin,...     

Length‐weight relationships for 55 freshwater fish species from the Iguatemi River,Upper Paraná River basin,Brazil

This study reports the length‐weight relationship estimatives (LWRs) for 55 fish species caught in the Iguatemi River drainage, Upper Paraná River Basin, Mato Grosso do Sul State, Brazil. The LWRs of 15 of the species are provided for the first time.     

Organization patterns of headwater-stream fish communities in the Upper Paraguay–Paraná basins

We compared environmental influences on the assemblages of stream fishes of contiguous but biogeographically distinct parts of the Upper Paraguay and Paraná basins in Brazil, aiming to describe the main distribution patterns of fish species in the headwater streams. We analyzed bimonthly samples in 10 streams in each basin from January to November 2004. Sixty fish species were collected, including 40 species in streams of the Paraguay Basin and 42 species in streams of the Paraná Basin. The species abundance rank did not differ between the basins. We found a clear differentiation between the Paraguay and Paraná stream fish assemblages. There were significant differences in species composition among streams, but no seasonal differences in species composition. Connectivity between streams explains the differences in species composition for the Paraguay streams, but not for the Paraná streams. Hydrological characteristics were the main factors determining species distribution in both basins, suggesting that the migratory capacity of each species and biogeographical barriers may act to filter the local species composition from the total species pool of colonizers. Handling editor: S. M. Thomaz     

Influence of large South American rivers of the Plata Basin on distributional patterns of tropical snakes: a panbiogeographical analysis

Aim The main drainages of the Plata Basin – the Paraná, Paraguay and Uruguay rivers – begin in tropical latitudes and run in a north–south direction into subtropical–temperate latitudes. Consequently, the biota of these rivers has tropical elements that contrast with temperate biomes through which the rivers run. We apply a panbiogeographical approach, to test whether the large rivers of the Plata Basin have a differential influence on distributional patterns of tropical snakes in subtropical and temperate latitudes of South America. Location Subtropical and temperate sections of the major Plata Basin rivers, South America. Methods We compared the individual tracks of 94 snake taxa. The track analysis consisted of: (1) plotting the localities of each taxon on maps, (2) connecting the localities of each taxon using a minimal geographical proximity determinant of the ‘individual tracks’, and (3) superimposing the individual tracks to determine generalized tracks. To detect tropical snakes that reach higher latitudes through the rivers we used the preferential direction of distribution concept. For each taxon we measured the angular deviations between the line of its individual track and the course of the rivers in a 100 × 100 km scaled grid. Average angular values < 45° indicated a positive association with the rivers. Results Thirty‐five of 94 taxa showed distributions associated with the major rivers of the Plata Basin, including fauna from distinct biogeographical lineages, supported by the occurrence of five generalized tracks as follows: (1) the Paraguay–Middle Paraná, (2) the Paraguay–Paraná fluvial axis, Upper Paraná and Middle Paraná to Upper Delta, (3) the Lower Paraguay, Paraná and Uruguay rivers, excluding the sectors High Paraná and High Uruguay, (4) the Uruguay River and Upper Paraná, and (5) the High Paraná. The Atlantic species occurred with significantly higher frequency in the Uruguay River and High Paraná river sections, the Amazon species were found with significantly higher frequency in the Paraguay and Middle Paraná sections, and the species with a Pantanal distribution were found in all sections. Main conclusions The observed distributional patterns may be explained by the interaction of ecological, geographical and historical factors. Previous authors have developed ecological (hydrological or environmental similarity) or dispersalist (effect of rivers as migration routes) explanations. The coincidence between generalized tracks and past geomorphological events that caused displacements and changed relationships between the Paraguay, Paraná and Uruguay river sections supports hypotheses involving the strong influence of historical factors in the present configuration of tropical snake distribution in temperate latitudes.     

A test of the riverine barrier hypothesis in the largest subtropical river basin in the Neotropics

The riverine barrier hypothesis proposes that large rivers represent geographical barriers to gene flow for terrestrial organisms, leading to population differentiation and ultimately allopatric speciation. Here we assess for the first time if the subtropical Paraná–Paraguay River system in the Del Plata basin, second in size among South American drainages, acts as a barrier to gene flow for birds. We analysed the degree of mitochondrial and nuclear genomic differentiation in seven species with known subspecies divided by the Paraná–Paraguay River axis. Only one species showed genetic differentiation concordant with the current river channel, but another five species have an east/west genetic split broadly coincident with the Paraná River's dynamic palaeochannel, suggesting this fluvial axis has had a past role in shaping present‐day genetic structure. Moreover, dating analyses show that these splits have been asynchronous, with species responding differently to the riverine barrier. Comparisons informed by the geological history of the Paraná River and its influence on the ecological and climatic differences among ecoregions in the study area further bolster the finding that responses to this geographical barrier have been species‐specific.     

Changes in taxonomy and species distributions and their influence on estimates of faunal homogenization and differentiation in freshwater fishes

Aim To assess how changing taxonomy and distribution data affect estimates of faunal homogenization and differentiation as agents of global change in freshwater fishes. Location Provinces and territories of Canada. Methods Species presence–absence data were collated in 2000 and 2005 from regional and national lists, and faunal homogenization and differentiation were calculated using Jaccard’s faunal similarity index. Differences between time periods and areas were summarized using principal coordinate analysis. Differences in faunal assemblages between native and total faunas were assessed via Whittaker’s (1960) beta diversity (β_w) index and tests of differences in multivariate dispersion of fish species compositions. Results Among aquatic ecoregions in one province (British Columbia) there were four taxonomic changes and 18 distributional changes between the 2000 and 2005 databases. Pairwise Jaccard’s faunal similarity index between the eight aquatic ecoregions declined by an average of 4.8% from 35.9% in native faunas to 31.1% in total faunas (introductions – extinctions/extirpations) indicating overall faunal differentiation. Average pairwise similarity declined by 0.9% between 2000 and 2005. Across thirteen provinces and territories of Canada, there were five taxonomic changes and 61 distributional changes between the 2000 and 2005 databases. Generally, faunal homogenization increased; pairwise Jaccard’s increased by an average of 1.8% from 27.1% in native faunas to 28.9% in total faunas or an average of 0.6% per comparison. Main conclusions Despite changing taxonomy and fish distribution information, comparative analysis of 2000 and 2005 databases consistently show overall faunal differentiation at the smallest (provincial) spatial scale and homogenization at the largest scale (across Canada) and that these trends continued between time periods. Homogenization and differentiation followed expectations from conceptual models based on the relative prevalence of species invasions and extinctions within communities. General conclusions of the onset and extent of homogenization and differentiation were relatively insensitive to our changing understanding of taxonomy and distribution.     

Spatial and temporal variations among fish with similar strategies: patterns of reproductive guilds in a floodplain

Seasonal flooding regimes are closely related to the life history of neotropical fish, especially with regard to their reproduction. The classification of fish into reproductive guilds serves to identify broad patterns in reproductive types, which are important in developing management and conservation measures. We tested the hypothesis that the fish reproductive guilds in the Upper Paraná River floodplain exhibit spatial and temporal distributions. Samples were taken each quarter in 2010 from several environments (i.e., biotopes) and rivers of the plain. Fish were categorized into four reproductive guilds based on migration, external fertilization, and parental care. Moreover, the abundance and species richness within each guild were used to evaluate their patterns. Guilds were spatially structured, possibly due to species dispersion as well as the influence of the hydrological and limnological characteristics of the biotopes and rivers of the plain. The Paraná River presented lower analyzed metric values, which demonstrates the negative effects of the dam built upstream of the study area. The guilds also presented temporal structure. During the flood season, the guilds presented similar spatial structures, and the local environmental characteristics led to spatial differentiation in the structure of guilds (i.e., flooding promoted the homogenization of the reproductive guilds).     

Patterns of change over time in darter (Teleostei: Percidae) assemblages of the Arkansas River basin,northeastern Oklahoma,USA

Rivers and streams are among the most threatened ecosystems worldwide, and their fish assemblages have been modified by anthropogenic habitat alteration and introductions of non‐native species. Consequently, two frequently observed patterns of assemblage change over time are species loss and biotic homogenization. In the present study, we compared contemporary (2006–2007) and historical (1948–1955) assemblages of darters, a group of small benthic fishes of the family Percidae, in the Arkansas River drainage of northeastern Oklahoma, USA. Results showed species loss between the two sampling periods, with historical estimates of overall species diversity across the study area exceeding contemporary estimates by five to eight species. Assemblages showed a low degree of darter similarity based on species presence and absence, with pairwise site comparisons (Jaccard's similarity index) between historical and contemporary samples averaging < 0.35. No significant homogenization or differentiation of assemblages occurred. Range expansion of widespread species, one of the primary mechanisms of biotic homogenization, was not observed; rather, all species occurred at a smaller proportion of sites in contemporary samples. Our results highlight the threat posed by anthropogenic habitat alteration to taxonomic groups such as darters, most of which are habitat specialists. However, our results suggest that biotic homogenization is unlikely to occur in the absence of immigration, especially if assemblages are subjected to ‘novel disturbances’ such as dam construction and watershed‐scale habitat degradation which negatively affect all components of the assemblage.     

First record of non‐native hybrid catfish Pseudoplatystoma reticulatum × Leiarius marmoratus in the Upper Paraná River basin,Brazil

The aim of this study is to report the presence of a three non‐native hybrid long‐whiskered catfishes (family Pimelodidae) in the Upper Paraná River basin, Brazil. Genetic analyses demonstrated that the three presumptive hybrids were a result of the crossbreeding of Pseudoplatystoma reticulatum (central Amazonas River basin and Lower Paraná River) and Leiarius marmoratus (Amazonas, Essequibo and Orinoco rivers), producing a hybrid commonly known in Brazil as cachandiá. The potential threat to biodiversity, due to possible genetic contamination, competition and predation of wild stocks, of such artificially produced hybrid fishes is discussed.     

Morphological and genetic divergence of a small stream fish species along a watershed

Piabina argentea is a species of Characidae family that is widely distributed in the hydrographic basins of the São Francisco and Paraná Rivers and in some Atlantic coastal drainages of South America. The genus contains only one other species, which is restricted to the Tietê River headwaters (the Upper Paraná River Basin) and is considered monophyletic within the Stevardiinae subfamily. Despite the apparent morphological conservation of at least six well-structured clades, great genetic distance revealed by DNA barcoding has already been reported. In an attempt to evaluate the effect of the watershed of the two river basins in the Arc of Upper Paranaíba region (Upper Paraná and São Francisco River basins), we analyzed the populations from both basins using mitochondrial DNA sequences and morphometry and constructed a distribution scenario for the species in both basins.     

Taxonomic and functional homogenization of an endemic desert fish fauna

Aim The highly endemic fishes of the arid Southwest USA have been heavily impacted by human activities resulting in one of the most threatened fish faunas in the world. The aim of this study was to examine the patterns and drivers of taxonomic and functional beta diversity of freshwater fish in the Lower Colorado River Basin across the 20th century. Location Lower Colorado River Basin (LCRB). Methods The taxonomic and functional similarities of watersheds were quantified to identify patterns of biotic homogenization or differentiation over the period 1900–1999. Path analysis was used to identify the relative influence of dam density, urban land use, precipitation regimes and non‐native species richness on observed changes in fish faunal composition. Results The fish fauna of the LCRB has become increasingly homogenized, both taxonomically (1.1% based on β_sim index) and functionally (6.2% based on Bray–Curtis index), over the 20th century. The rate of homogenization varied substantially; range declines of native species initially caused taxonomic differentiation (?7.9% in the 1960s), followed by marginal homogenization (observed in the 1990s) in response to an influx of non‐native species introductions. By contrast, functional homogenization of the basin was evident considerably earlier (in the 1950s) because of the widespread introduction of non‐native species sharing similar suites of biological traits. Path analysis revealed that both taxonomic and functional homogenization were positively related to the direct and indirect (facilitation by dams and urbanization) effects of non‐native species richness. Main conclusions Our study simultaneously examines rates of change in multiple dimensions of the homogenization process. For the endemic fish fauna of the LCRB, we found that the processes of taxonomic and functional homogenization are highly dynamic over time, varying both in terms of the magnitude and rate of change over the 20th century.     

Taxonomic homogenization in ungulates: patterns and mechanisms at local and global scales

Aim The aim of this paper is to examine taxonomic homogenization in ungulates globally and at the local scale in South Africa. Specifically, we aim to examine the roles of distance, scale, time, extinctions vs. introductions, and extralimital vs. extraregional introductions in the homogenization of ungulate biotas, and to determine pathways of introduction of ungulate species globally and the proximate explanatory variables of ungulate introductions in South Africa. Location Forty‐one countries globally and three spatial resolutions in South Africa. Methods Indigenous, extirpated and established introduced ungulate species data were obtained for countries globally, and at a quarter‐degree grid‐cell resolution in South Africa. Homogenization was calculated using Jaccard’s index of similarity (JI) for countries globally and for three spatial resolutions in South Africa. Zoo holdings and transfer data from the International Species Information System database were used to investigate the relationship between non‐indigenous ungulate species introductions and the number of non‐indigenous ungulate species in zoos. Relationships between JI and species richness, and between numbers of introductions and several environmental and social factors were examined using generalized linear models. Results Homogenization in ungulates was 2% for countries globally and 8% at the coarsest resolution in South Africa. Homogenization increased with increasing resolution and with time, but it decreased with increasing percentage change in species richness. Globally, introductions contributed more to homogenization than did extinctions. Within South Africa, extralimital introductions contributed more to the homogenization of ungulate assemblages than did extraregional ones, and ungulates were typically introduced to high‐income areas with high human population and livestock densities. The same was not true in the past, when ungulates were introduced to ungulate species‐poor areas. The number of non‐indigenous ungulate species established in a country is significantly related to the number of non‐indigenous ungulate species in zoos in the country, possibly owing to sales of surplus animals from zoos. Main conclusions Ungulate faunas are homogenized at both the global scale and in South Africa, with extralimital introductions being of considerable significance regionally. In consequence, increasing attention will have to be given to the conservation consequences of ungulate translocations, both within particular geopolitical regions and across the globe.     

Temporal and spatial distributions of the fish larval assemblages of the Ivinheima River sub-basin (Brazil)

Fish, like other animals, choose environments for reproduction that are favorable for their offspring’s initial development. For example, these environments may be chosen to provide shelter and food. This study examined the fish larvae inhabiting the Ivinheima River sub-basin. The study aimed to (i) characterize the taxonomic composition of the larval fish fauna, (ii) analyze the structure of the larval assemblage, and (iii) verify the relationship between the larval assemblages and environmental variables. The sampling areas included three environments: the Ivinheima River, the Finado Raimundo Lagoon and the Patos Lagoon. Sampling was performed between October 2002 and March 2006 during four reproductive periods (RPs). The data were analyzed with correspondence analysis (CA) and canonical correlation analysis (CCA). A total of 120 619 larvae were collected. These larvae belonged to several species characterized by different reproductive strategies. Our results showed differences between the composition of the Ivinheima River’s assemblages, where there were more larvae of migratory fish, and the lagoons, where larvae of sedentary species were predominant. In addition, differences were found among the RPs evaluated in the Ivinheima River: the first and second RPs were characterized by milder floods and by a greater abundance of migratory piscivore larvae, whereas the third and fourth RPs had more outflow and a greater abundance of larvae of detritivore and omnivore species. We conclude that the Ivinheima River plays a significant role in the maintenance of native migratory fish stocks in the upper Paraná River basin and that the environment shows a segregation of the spatial and temporal composition and abundance of larvae related to running and flooding/standing water.     

Seasonal variation in genetic population structure of sábalo (Prochilodus lineatus) in the Lower Uruguay River

Prochilodus lineatus is a highly migratory fish species that sustains the most important commercial fishery of Paraná-Paraguay basin. Migratory patterns are poorly known and only few population genetic studies are available for this species in the Upper Paraná. To assess genetic population structure, we genotyped a sample of 93 individuals from the Lower Uruguay River close to Gualeguaychú city (Entre Ríos, Argentina) at three different times, July 2008 (Winter), September 2008 (Spring) and May 2009 (Fall). All individuals were genotyped for 12 microsatellite loci previously found to be informative to assess populations of P. lineatus. Our results show seasonal variation of the genetic sub-structuring at this locality that may be related to the presence of different migratory stocks throughout the year. The Fall sample includes an additional genetic cluster of individuals not detected in Winter and Spring, suggesting that this species should be considered a mixed stock fishery.     

Length–weight and length–length relationships for 23 fish species of Porto Primavera reservoir,Upper Paraná River,Brazil

The study presents length–weight relationships (LWR) and length–length relationships (LLR) for 23 fish species captured in the Porto Primavera Reservoir, Upper Paraná River. Seventeen of the LWRs and 20 of the LLRs are reported for the first time. New maximum standard lengths are presented for 17 species as well as weights for three species and new total weight records for 19 species.     

Migratory movements of pacu, Piaractus mesopotamicus, in the highly impounded Paraná River

A mark‐recapture study was conducted in 1997–2005 to investigate movements of stocked pacu, Piaractus mesopotamicus, in the Paraná River Basin of Brazil, Paraguay, and Argentina. Fish raised in cages within the Itaipu Reservoir and in ponds were tagged externally (n = 2976) and released in the Itaipu Reservoir (53.2%) and bays of its major tributaries (46.8%). In total, 367 fish (12.3%) were recaptured. In all, 91% of the pacu moved away from the release site; upstream movements were more extensive than downstream movements. Pacu traveled upstream a maximum of 422 km (average of 41.3 km) at a maximum rate of 26.4 km day⁻¹ (av. 0.8). Downstream movements were limited in terms of number of individuals and distance moved. Fish released during the wet season moved farther than those released during the dry season, and feeding rather than spawning might have been the compelling reason for movement. Although fish passed downstream through dams, none of the marked fish were detected to have moved upstream through the passage facilities. Pacu showed movement patterns not radically different from those of other neotropical migratory species, but their migratory movements may not be as extensive as those of other large migratory species in the basin.     

A panbiogeographical model to prioritize areas for conservation along large rivers

Aim The large rivers of the Neotropics are considered areas of high diversity and endemism, which play an important role in the distribution patterns and evolution of Neotropical biota. Several methods have been proposed for prioritizing terrestrial conservation areas, but there has been little effort to develop models for river systems. We propose a panbiogeographical approach to identify priority areas for conservation along rivers. Location The Plata Basin rivers. Methods We compared the individual tracks (IT) of 96 snake taxa and identified the species associated with rivers using the concept of preferential direction of distribution. For each taxon, we measured the angular deviations between the line of its IT and the course of the rivers on a 100 × 100 km scaled grid. Average angular values < 45° indicated a positive association with rivers. We detected 35 taxa associated with rivers, and their IT were used to determine the generalized tracks (GT) and nodes. We applied a complementarity algorithm to identify the minimum set of nodes required to represent all species. Results Six nodes were found. The region where the High and Upper Paraná Rivers converge (Node 1) is of first priority, with 60 of 96 species. The second priority is the Lower Paraguay River and northern section of Middle Paraná River (Node 2). The third is the High Paraná, which together with Nodes 1 and 2, comprises 94% of the total species. The fourth and fifth are the High and Middle Uruguay, and the western section of the Upper Paraná and Iberá Marsh system, respectively. These five nodes include all species. Main conclusions Our results highlight the areas of particular interest for the conservation of rivers and provide a biogeographical algorithm for detecting priority conservation areas. The nodes are a biogeographical approach that allows evolutionary and ecological traits to be included in conservation assessment.     

Taxonomic homogenization and differentiation across Southern Ocean Islands differ among insects and vascular plants

Aim To investigate taxonomic homogenization and/or differentiation of insect and vascular plant assemblages across the Southern Ocean Islands (SOI), and how they differ with changing spatial extent and taxonomic resolution. Location Twenty‐two islands located across the Southern Ocean, further subdivided into five island biogeographical provinces. These islands are used because comprehensive data on both indigenous and non‐indigenous insect and plant species are available. Methods An existing database was updated, using newly published species records, identifying the indigenous and non‐indigenous insect and vascular plant species recorded for each island. Homogenization and differentiation were measured using Jaccard’s index (JI) of similarity for assemblages across all islands on a pairwise basis, and for island pairs within each of the biogeographical provinces. The effects of taxonomic resolution (species, genus, family) and distance on levels of homogenization or differentiation were examined. To explore further the patterns of similarity among islands for each of the taxa and groupings (indigenous and non‐indigenous), islands were clustered based on JI similarity matrices and using group averaging. Results Across the SOI, insect assemblages have become homogenized (0.7% increase in similarity at species level) while plant assemblages have become differentiated at genus and species levels. Homogenization was recorded only when pairwise distances among islands exceeded 3000 km for insect assemblages, but distances had to exceed 10,000 km for plant assemblages. Widely distributed non‐indigenous plant species tend to have wider distributions across the SOI than do their insect counterparts, and this is also true of the indigenous species. Main conclusions Insect assemblages across the SOI have become homogenized as a consequence of the establishment of non‐indigenous species, while plant assemblages have become more differentiated. The likely reason is that indigenous plant assemblages are more similar across the SOI than are insect assemblages, which show greater regionalization. Thus, although a suite of widespread, typically European, weedy, non‐indigenous plant species has established on many islands, the outcome has largely been differentiation. Because further introductions of insects and vascular plants are probable as climates warm across the region, the patterns documented here are likely to change through time.