Natural fragmentation in river networks as a driver of speciation for freshwater fishes

Although habitat fragmentation fosters extinctions, it also increases the probability of speciation by promoting and maintaining divergence among isolated populations. Here we test for the effects of two isolation factors that may reduce population dispersal within river networks as potential drivers of freshwater fish speciation: 1) the position of subdrainages along the longitudinal river gradient, and 2) the level of fragmentation within subdrainages caused by natural waterfalls. The occurrence of native freshwater fish species from 26 subdrainages of the Orinoco drainage basin (South America) was used to identify those species that presumably arose from in‐situ cladogenetic speciation (i.e. neo‐endemic species; two or more endemic species from the same genus) within each subdrainage. We related subdrainages fish diversity (i.e. total, endemic and neo‐endemic species richness) and an index of speciation to our two isolation factors while controlling for subdrainages size and energy availability. The longitudinal position of subdrainages was unrelated to any of our diversity measures, a result potentially explained by the spatial grain we used and/or the contemporary connection between Orinoco and Amazon basins via the upstream Casiquiare region. However, we found higher neo‐endemic species richness and higher speciation index values in highly fragmented subdrainages. These results suggest that habitat fragmentation generated by natural waterfalls drives cladogenetic speciation in fragmented subdrainages. More generally, our results emphasize the role of history and natural waterfalls as biogeographic barriers promoting freshwater biodiversity in river drainage basins.     

黄河流域淡水鱼类多样性和保护

本文介绍了黄河流域鱼类研究的历史、淡水鱼类的物种组成、整体分布格局、特有性、濒危性, 以及鱼类多样性在黄河上、中、下游等的特点。历史上对于黄河鱼类多样性的研究, 历经了四个阶段, 从最初的零星记录一直到现阶段的深入研究。综合历史记录和野外调查, 已知黄河流域分布的淡水鱼类共计147种, 隶属于12目21科78属, 其中鲤形目种类占据绝对优势。另外, 全流域黄河特有种计有27种, 受危物种24种, 分别占总数的18.37%和16.32%。同中国主要江河相比, 黄河鱼类在高级分类阶元上的多样性较高, 但物种多样性则处在较低水平; 尽管黄河特有鱼类和受危物种比例低于全国平均水平, 但上游特有鱼类和珍稀濒危鱼类的占比很高。目前黄河鱼类多样性大幅降低, 现状调查仅能采获历史记录种类的53.06%。梯级水电开发、水资源过度利用、外来物种、水域污染和过度捕捞都是威胁鱼类多样性的重要因素, 但对各河段和支流的影响不一, 应做出有针对性的保护部署。     

新疆阿勒泰地区额尔齐斯河和乌伦古河流域鱼类多样性演变和流域健康评价

为了解阿勒泰地区额尔齐斯河和乌伦古河流域的鱼类多样性现状和历史演变, 本研究于2013-2016年间在该流域的鱼类多样性进行了连续调查, 并结合历史资料和标本, 以Margalef丰富度指数、Shannon-Wiener多样性指数、Pielou均匀度指数分析评估了流域内鱼类的多样性水平和时空变化。该流域历史上分布有土著鱼类23种, 当前记录到19种, 流域内还有外来鱼类15种。阿勒泰鱼类的区系组成以鲤科种类为主, 其中特有和珍稀濒危物种占比高, 具有重要的保护价值。多样性指数计算结果显示, 2013-2016年鱼类多样性情况整体稳定, 额尔齐斯河鱼类物种数多于乌伦古河。研究还基于鱼类生物完整性指数(Fish Index of Biological Integrity, F-IBI)对34个采集点的河流健康状况进行了评价, 结果显示额尔齐斯河流域大多数调查点的健康状况处于“亚健康”或“一般”水平, 乌伦古河流域多数调查点的健康状况处于“健康”水平。水利工程、外来物种、过度捕捞是影响阿勒泰地区鱼类多样性的重要因素。未来应通过水利工程的联合调度、下泄合理生态流量、布设鱼类通道、规范养殖渔业、严控外来物种、本地土著鱼类的人工增殖放流, 以及合理的就地保护措施对阿勒泰地区的鱼类多样性加以保护, 提升水体健康程度。     

Identifying climatic niche shifts using coarse‐grained occurrence data: a test with non‐native freshwater fish

Aim We tested whether coarse‐grained occurrence data can be used to detect climatic niche shifts between native and non‐native ranges for a set of widely introduced freshwater fishes. Location World‐wide. Methods We used a global database of freshwater fish occurrences at the river basin scale to identify native and non‐native ranges for 18 of the most widely introduced fish species. We also examined climatic conditions within each river basin using fine‐grained climate data. We combined this information to test whether climatic niche shifts have occurred between native and non‐native ranges. We defined climatic niche shifts as instances where the ranges of a climatic variable within native and non‐native basins exhibit zero overlap. Results We detected at least one climatic niche shift for each of the 18 studied species. However, we did not detect common patterns in the thermal preference or biogeographic origin of the non‐native fish, hence suggesting a species‐specific response. Main conclusions Coarse‐grained occurrence data can be used to detect climatic niche shifts. They also enable the identification of the species experiencing niche shifts, although the mechanisms responsible for these shifts (e.g. local adaptation, dispersal limitation or physiological constraints) have yet to be determined. Furthermore, the coarse‐grained approach, which highlights regions where climatic niche shifts have occurred, can be used to select specific river basins for more detailed, fine‐grained studies.     

Environmental correlates of species richness for native freshwater fish in Oregon, U.S.A.

Aim To better understand how environmental factors affect fish species richness across the state of Oregon. Location Oregon, U.S.A. Methods A database showing collection locations of 4911 fish specimens in the Oregon State University museum was modified by the Oregon Natural Heritage Program to include probable occurrences, and mapped within a grid of 375 hexagons that cover the state. The individual species maps of freshwater fish in Oregon were reviewed and revised by thirty regional fish biologists and then synthesized into a single map of native species richness. We used regression tree analysis (RTA) and multiple linear regression (MLR) to assess patterns of fish species richness with twenty environmental, three anthropogenic, and two historical variables. Results RTA explained 66% of the variation in native species richness, associating richness with annual air temperature range, minimum January temperature, introduced species richness, and stream density. MLR explained 68% of native species richness variation and associated richness with maximum July temperature, air temperature range, standard deviation of monthly temperature, stream density, introduced species richness, and basin connectivity. Main conclusions We conclude that for these data and at this scale, native fish species richness in Oregon is associated with annual climatic extremes, spatial variability of climate, stream density, basin connectivity, and introduced fishes.     

BIODIVERSITY RESEARCH: Native and introduced fish species richness in Mediterranean streams: the role of multiple landscape influences

Aim To examine the role of multiple landscape factors on the species richness patterns of native and introduced freshwater fish. Location Mediterranean streams, south‐western Iberian Peninsula, Europe (c. 87,000 km²). Methods We used a dataset of fish occurrences from 436 stream sites. We quantified the incremental explanatory power of multiple landscape factors in native, introduced, and overall local species richness using regression analysis. First, we related variation in local species richness across river basins to regional species richness (here, the basin species pool), area and factors of climate and topography. Second, we related within‐river basin local species richness to site’s climate and topography, and spatial structure derived from Principal Coordinates of Neighbour Matrices approach, after testing for species richness spatial autocorrelation; predicted local richness was mapped. Results Patterns of local species richness across river basins were strongly associated with regional species richness for overall, native and introduced species; annual rainfall showed a significant incremental contribution to variation in introduced species richness only. Within river basins, environmental factors were associated with local richness for the three species groups, though their contributions to the total explained variation were inferior to those of spatial factors; rainfall seasonality and stream slope were the most consistent environmental correlates for all species groups, while the influence of spatial factors was most prevalent for native species. Main conclusions Landscape factors operating among and within river basins seem to play a relevant role in shaping local species richness of both native and introduced species, and may be contingent on basin‐specific contexts. Nevertheless, local factors, such as habitat characteristics and biotic interactions and human‐induced disturbances may also be at play. Multiscale approaches incorporating a multitude of factors are strongly encouraged to facilitate a deeper understanding of the biodiversity patterns of Mediterranean streams, and to promote more effective conservation and management strategies.     

Fish-AMAZBOL: a database on freshwater fishes of the Bolivian Amazon

The Bolivian part of the Amazon Basin contains a mega diverse and well-preserved fish fauna. Since the last decade, this fish fauna has received an increasing attention from scientists and the national authorities as fishes represent one of the most important sources of proteins for local human communities. However, this fish fauna still remains poorly documented. Here, we present a database for fishes from the Bolivian Amazon. To build the database, we conducted an extensive literature survey of native and non-native (exotic) fishes inhabiting all major sub-drainages of the Bolivian Amazon. The database, named Fish-AMAZBOL, contains species lists for 13 Amazonian hydrological units, covering 100% of the Bolivian Amazon and approximately 65% (722,137 km²) of the all territory. Fish-AMAZBOL includes 802 valid species, 12 of them being non-native, that have been checked for systematic reliability and consistency. To put this number in perspective, this represents around 14% of the all Neotropical ichthyofauna and around 6% of all strictly freshwater fishes inhabiting the planet. This database is currently the most comprehensive database of native and non-native fish species richness available so far for the Bolivian Amazon.     

Impending extinctions of North American freshwater mussels (Unionoida) following the zebra mussel (Dreissena polymorpha) invasion

1. Freshwater mussels (Order Unionoida) are the most imperiled faunal group in North America; 60% of described species are considered endangered or threatened, and 12% are presumed extinct. Widespread habitat degradation (including pollution, siltation, river channelization and impoundment) has been the primary cause of extinction during this century, but a new stress was added in the last decade by the introduction of the Eurasian zebra mussel, Dreissena polymorpha , a biofouling organism that smothers the shells of other molluscs and competes with other suspension feeders for food. Since the early 1990s, it has been spreading throughout the Mississippi River basin, which contains the largest number of endemic freshwater mussels in the world. In this report, we use an exponential decay model based on data from other invaded habitats to predict the long-term impact of D. polymorpha on mussel species richness in the basin.
2. In North American lakes and rivers that support high densities (>3000 m⁻²) of D. polymorpha , native mussel populations are extirpated within 4–8 years following invasion. Significant local declines in native mussel populations in the Illinois and Ohio rivers, concomitant with the establishment of dense populations of D. polymorpha , suggest that induced mortality is occurring in the Mississippi River basin.
3. A comparison of species loss at various sites before and after invasion indicates that D. polymorpha has accelerated regional extinction rates of North American freshwater mussels by 10-fold. If this trend persists, the regional extinction rate for Mississippi basin species will be 12% per decade. Over 60 endemic mussels in the Mississippi River basin are threatened with global extinction by the combined impacts of the D. polymorpha invasion and environmental degradation.     

Changes in fish assemblages in catchments in north‐eastern Spain: biodiversity,conservation status and introduced species

1. North‐eastern Spain is a hot spot for the introduction of alien fish species, and its native fish fauna is one of the most endangered worldwide. We used an extensive data set from 2002 to 2003 and historical information from the area to characterize fish diversity and establish conservation priorities in river catchments. 2. Diversity indices were used to characterize fish diversity at the basin scale. An index of conservation status was applied for each species, which considers the occurrence, abundance and endemicity of each taxon. We used indirect ordination methods to test the relationship among basin features and to identify those variables most correlated with each other. To identify physical, biotic and environmental characteristics that seem to make a basin particularly susceptible to invasion, we performed a step‐wise multiple regression to examine the relationship between the number of native, translocated and introduced fish species (including the original native species richness of each basin), and landscape variables. 3. Over a period of approximately 50 years, the mean range size of native fish species has decreased by 60%. The greatest decline occurred in Gasterosteus gymnurus, Anguilla anguilla and Salaria fluviatilis, for which species over 75% of the original distribution area has been lost. The species with the highest conservation index were Gasterosteus gymnurus and Salaria fluviatilis. 4. Basin area and the catchment type explained 70% of variation in native species richness, whereas the number of dams and basin area accounted for more than 80% of variation in the number of introduced species. 5. The original native species richness and the number of introduced species at basin scale were not related, and thus there was no evidence of “biotic resistance” to invasion. The restoration of natural hydrologic processes and the development of specific management tools to protect native species, such as the prioritization of areas for fish conservation and the eradication of local populations of exotic species, are required to restore native fish fauna in these catchments.     

Global species richness of hydrobiid snails determined by climate and evolutionary history

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Fish Invasions in the World's River Systems: When Natural Processes Are Blurred by Human Activities

Because species invasions are a principal driver of the human-induced biodiversity crisis, the identification of the major determinants of global invasions is a prerequisite for adopting sound conservation policies. Three major hypotheses, which are not necessarily mutually exclusive, have been proposed to explain the establishment of non-native species: the “human activity” hypothesis, which argues that human activities facilitate the establishment of non-native species by disturbing natural landscapes and by increasing propagule pressure; the “biotic resistance” hypothesis, predicting that species-rich communities will readily impede the establishment of non-native species; and the “biotic acceptance” hypothesis, predicting that environmentally suitable habitats for native species are also suitable for non-native species. We tested these hypotheses and report here a global map of fish invasions (i.e., the number of non-native fish species established per river basin) using an original worldwide dataset of freshwater fish occurrences, environmental variables, and human activity indicators for 1,055 river basins covering more than 80% of Earth's surface. First, we identified six major invasion hotspots where non-native species represent more than a quarter of the total number of species. According to the World Conservation Union, these areas are also characterised by the highest proportion of threatened fish species. Second, we show that the human activity indicators account for most of the global variation in non-native species richness, which is highly consistent with the “human activity” hypothesis. In contrast, our results do not provide support for either the “biotic acceptance” or the “biotic resistance” hypothesis. We show that the biogeography of fish invasions matches the geography of human impact at the global scale, which means that natural processes are blurred by human activities in driving fish invasions in the world's river systems. In view of our findings, we fear massive invasions in developing countries with a growing economy as already experienced in developed countries. Anticipating such potential biodiversity threats should therefore be a priority.     

From richer to poorer: successful invasion by freshwater fishes depends on species richness of donor and recipient basins

Evidence for the theory of biotic resistance is equivocal, with experiments often finding a negative relationship between invasion success and native species richness, and large‐scale comparative studies finding a positive relationship. Biotic resistance derives from local species interactions, yet global and regional studies often analyze data at coarse spatial grains. In addition, differences in competitive environments across regions may confound tests of biotic resistance based solely on native species richness of the invaded community. Using global and regional data sets for fishes in river and stream reaches, we ask two questions: (1) does a negative relationship exist between native and non‐native species richness and (2) do non‐native species originate from higher diversity systems. A negative relationship between native and non‐native species richness in local assemblages was found at the global scale, while regional patterns revealed the opposite trend. At both spatial scales, however, nearly all non‐native species originated from river basins with higher native species richness than the basin of the invaded community. Together, these findings imply that coevolved ecological interactions in species‐rich systems inhibit establishment of generalist non‐native species from less diverse communities. Consideration of both the ecological and evolutionary aspects of community assembly is critical to understanding invasion patterns. Distinct evolutionary histories in different regions strongly influence invasion of intact communities that are relatively unimpacted by human actions, and may explain the conflicting relationship between native and non‐native species richness found at different spatial scales.     

Fish Invasions in the World's River Systems: When Natural Processes Are Blurred by Human Activities

Because species invasions are a principal driver of the human-induced biodiversity crisis, the identification of the major determinants of global invasions is a prerequisite for adopting sound conservation policies. Three major hypotheses, which are not necessarily mutually exclusive, have been proposed to explain the establishment of non-native species: the “human activity” hypothesis, which argues that human activities facilitate the establishment of non-native species by disturbing natural landscapes and by increasing propagule pressure; the “biotic resistance” hypothesis, predicting that species-rich communities will readily impede the establishment of non-native species; and the “biotic acceptance” hypothesis, predicting that environmentally suitable habitats for native species are also suitable for non-native species. We tested these hypotheses and report here a global map of fish invasions (i.e., the number of non-native fish species established per river basin) using an original worldwide dataset of freshwater fish occurrences, environmental variables, and human activity indicators for 1,055 river basins covering more than 80% of Earth's surface. First, we identified six major invasion hotspots where non-native species represent more than a quarter of the total number of species. According to the World Conservation Union, these areas are also characterised by the highest proportion of threatened fish species. Second, we show that the human activity indicators account for most of the global variation in non-native species richness, which is highly consistent with the “human activity” hypothesis. In contrast, our results do not provide support for either the “biotic acceptance” or the “biotic resistance” hypothesis. We show that the biogeography of fish invasions matches the geography of human impact at the global scale, which means that natural processes are blurred by human activities in driving fish invasions in the world's river systems. In view of our findings, we fear massive invasions in developing countries with a growing economy as already experienced in developed countries. Anticipating such potential biodiversity threats should therefore be a priority.     

Patterns and drivers of fish extirpations in rivers of the American Southwest and Southeast

Effective conservation of freshwater biodiversity requires spatially explicit investigations of how dams and hydroclimatic alterations among climate regions may interact to drive species to extinction. We investigated how dams and hydroclimatic alterations interact with species ecological and life history traits to influence past extirpation probabilities of native freshwater fishes in the Upper and Lower Colorado River (CR), Alabama‐Coosa‐Tallapoosa (ACT), and Apalachicola‐Chattahoochee‐Flint (ACF) basins. Using long‐term discharge data for continuously gaged streams and rivers, we quantified streamflow anomalies (i.e., departure “expected” streamflow) at the sub‐basin scale over the past half‐century. Next, we related extirpation probabilities of native fishes in both regions to streamflow anomalies, river basin characteristics, species traits, and non‐native species richness using binomial logistic regression. Sub‐basin extirpations in the Southwest (n = 95 Upper CR, n = 130 Lower CR) were highest in lowland mainstem rivers impacted by large dams and in desert springs. Dampened flow seasonality, increased longevity (i.e., delayed reproduction), and decreased fish egg sizes (i.e., lower parental care) were related to elevated fish extirpation probability in the Southwest. Sub‐basin extirpations in the Southeast (ACT n = 46, ACF n = 22) were most prevalent in upland rivers, with flow dependency, greater age and length at maturity, isolation by dams, and greater distance upstream. Our results confirm that dams are an overriding driver of native fish species losses, irrespective of basin‐wide differences in native or non‐native species richness. Dams and hydrologic alterations interact with species traits to influence community disassembly, and very high extirpation risks in the Southeast are due to interactions between high dam density and species restricted ranges. Given global surges in dam building and retrofitting, increased extirpation risks should be expected unless management strategies that balance flow regulation with ecological outcomes are widely implemented.     

Invasion of North American drainages by alien fish species

1. Data from the literature were used to document colonization patterns by introduced freshwater fishes in 125 drainages across temperate North America. We analysed this data set to quantify susceptibility to invasion, success of the invaders and changes in species richness.
2. Drainages with a high number of impoundments, large basin area and low native species diversity had the greatest number of introduced species. Those drainages containing few native fishes exhibited great variation in the number of invaders, while waters with a rich native fauna contained few introduced species. However, this pattern did not differ significantly from random simulations because the pool of potential invaders is greater for drainages with low species richness.
3. In most drainages, there were more introduced than imperilled or extirpated species, suggesting that invaders tend to increase overall species richness.
4. These patterns suggest that North American fish communities are not saturated with species, but instead, are capable of supporting higher levels of diversity if the pool of potential colonists and the rate of colonization from that pool is increased.     

Evaluating the relationship between basin‐scale fish species richness and ecologically relevant flow characteristics in rivers worldwide

1. River flow alterations due to climate change and increasing water usage affect freshwater biodiversity including fish species richness. Here, we statistically explored the relationships of fish species richness to 14 ecologically relevant flow metrics as well as basin area and latitude in 72 rivers worldwide. 2. The statistical models best supported by the data included three variables with positive coefficients (mean river discharge, basin area and the maximum proportion of no‐flooding period) and three variables with negative coefficients (latitude, coefficients of variation in the frequency of low flow and the Julian date of annual minimum flow). 3. The model outputs have provided the first empirical indication that specific low‐ and high‐flow characteristics may be important in explaining variations in basin‐scale fish species richness. Our findings can be useful in identifying high‐risk basins for conservation of fish species diversity. 4. The results not only support the adoption of mean discharge as a predictor, but also suggest the importance of basin area in predicting basin‐scale fish species richness around the world.     

Contrasting patterns and mechanisms of spatial turnover for native and exotic freshwater fish in Europe

Aim We compare the distribution patterns of native and exotic freshwater fish in Europe, and test whether the same mechanisms (environmental filtering and/or dispersal limitation) govern patterns of decrease in similarity of native and exotic species composition over geographical distance (spatial species turnover). Locations Major river basins of Europe. Methods Data related to geography, habitat diversity, regional climate and species composition of native and exotic freshwater fish were collated for 26 major European river basins. We explored the degree of nestedness in native and exotic species composition, and quantified compositional similarity between river basins according to the beta‐sim (independent of richness gradient) and Jaccard (dependent of richness gradient) indices of similarity. Multiple regression on distance matrices and variation‐partitioning approaches were used to quantify the relative roles of environmental filtering and dispersal limitation in shaping patterns of decreasing compositional similarity over geographical distance. Results Native and exotic species exhibited significant nested patterns of species composition, indicating that differences in fish species composition between river basins are primarily the result of species loss, rather than species replacement. Both native and exotic compositional similarity decreased significantly with increasing geographical distance between river basins. However, gradual changes in species composition with geographical distance were found only for exotic species. In addition, exotic species displayed a higher rate of similarity decay (higher species turnover rate) with geographical distance, compared with native species. Lastly, the majority of explained variation in exotic compositional similarity was uniquely related to geography, whereas native compositional similarity was either uniquely explained by geography or jointly explained by environment and geography. Main conclusions Our study suggests that large‐scale patterns of spatial turnover for exotic freshwater fish in Europe are generated by human‐mediated dispersal limitation, whereas patterns of spatial turnover for native fish result from both dispersal limitation relative to historical events (isolation by mountain ranges, glacial history) and environmental filtering.     

Biogeography of freshwater fishes of the Balkan Peninsula

Delineating biogeographical regions is a critical step towards the establishment and evaluation of conservation priorities. In the present study, we analysed the distribution patterns of the freshwater fish of an understudied European biodiversity hotspot, the Balkan Peninsula. Based on the most extensive available database of native freshwater fish species distributions, we performed a hierarchical clustering analysis to identify the major biogeographical regions of the Balkan Peninsula. We also highlighted the ‘hottest hotspots’ of freshwater fish diversity across the delimited biogeographical regions by describing the patterns of species richness, endemic and vulnerable species; indicator species were also determined. The bioregionalisation scheme consisted of eight groups of drainage basins that correspond to distinct regions of the Balkan Peninsula. Overall, the delineated biogeographical regions varied in terms of species richness, endemism, vulnerability (i.e. extinction threats) and indicator species composition. From a conservation perspective, this study emphasises the prioritisation of areas characterised by high levels of irreplaceability (endemism) and vulnerability (i.e. the Attikobeotia region, Ionian Sea and Prespa Lakes) and stresses the necessity of implementing a network of protected freshwater areas across the Balkan Peninsula.     

Non-native fishes and native species diversity in freshwater fish assemblages across the United States

The proliferation of non-native species in North American freshwater ecosystems is considered a primary threat to the integrity of native community structure. However, a general understanding of consistent and predictable impacts of non-native species on native freshwater diversity is limited, in part, because of a lack of broad-scale studies including data from numerous localities across multiple drainages. This study uses data from 751 localities collected during the United States Geological Survey (USGS) National Water Quality Assessment (NAWQA) program to examine the influence of non-native fish species on native freshwater fish assemblages across the United States. In general, no significant differences in native fish richness and diversity measures were detected between sites with only native species and sites containing non-native species. However, at sites with non-native species, the number of non-native species present was negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Non-native piscivores were negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Native piscivores were positively correlated with native richness and diversity and negatively correlated with native evenness at sites with only native species. Our results suggest that from a superficial perspective, native species richness and diversity are not different among sites with and without non-native species. However, when patterns of native species richness and diversity are examined at sites containing non-native species, correlations between non-native and native species richness and diversity imply the expected negative effect of invasive taxa. Additionally, non-native piscivores appear to have a significant negative effect on native taxa and possibly represent a novel selective force on naive native prey.     

Distribution of fish in the Virgin River,a tributary of the lower Colorado River

Synopsis The Virgin River, a major tributary of the lower Colorado River, contains a unique, though impoverished, native ichthyofauna (six species) that is largely intact despite the introduction of exotic fishes (13 species) and alterations of the river. The main conclusions of the study are: (1) The longitudinal distribution pattern of the native species does not follow theoretical expectations of a downstream increase in species richness; (2) for the size of the basin (>15,000 km²), the native ichthyofauna is extremely depauperate; and (3) most of the exotic fish are not established in the river, but persist through repeated invasions from downstream reservoirs.