Hypoxic wetland tributaries as faunal refugia from an introduced predator

Abstract– The introduction of Nile perch ( Lates niloticus ) into the Lake Victoria basin of East Africa has coincided with the decline or disappearance of hundreds of indigenous species. To mitigate additional biodiversity loss, we must learn what limits the spread of Nile perch and what habitats serve as refugia for prey species. Heavily vegetated wetlands may protect fishes from Nile perch predation by providing both structural and low-oxygen refugia for prey species tolerant of hypoxia. To examine the potential of wetlands as refugia we quantified the composition, persistence, and stability of fish assemblages in a wetland tributary of Lake Nabugabo, a satellite lake of Lake Victoria in which Nile perch have been introduced. Nile perch were extremely rare in the wetland, and nine of the 18 species that have disappeared from the open waters of the satellite lake were captured in the tributary in this study. Dissolved oxygen was chronically low in the river and may be important in shaping fish community characteristics. Faunal attenuation occurred as the dry season progressed and oxygen levels dropped; however, the most common species remained through seasonal changes. The chronically low oxygen conditions in the werland tributary may permit persistence of only hypoxia-tolerant species. However, wetland conditions seem to limit exploitation by Nile perch providing critical refugia for a subset of the basin fauna.     

Managed and natural inundation: benefits for conservation of native fish in a semi‐arid wetland system

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Biodiversity value of remnant pools in an intermittent stream during the great California drought

1. In many intermittent streams, remnant pools persist after flow ceases and provide refuge for aquatic organisms able to tolerate stagnant water conditions. The conservation value of these pools may be greatly under‐appreciated, especially in regions with a Mediterranean climate, where perennial streams have been substantially modified or disturbed by human activities.
2. Fish, amphibians, aquatic reptiles, and aquatic invertebrates were sampled from 15 remnant pools and three seeps at Coyote Creek, California, USA, in the late summer of 2014, during the height of the most intense drought that California has experienced in 500 years. Patterns of vertebrate and invertebrate species richness and community composition were compared with abiotic factors (e.g. water quality and habitat size).
3. Thirteen vertebrate species and 172 invertebrate taxa were identified from remnant pools and seeps. Overall vertebrate richness and composition were not correlated with abiotic factors, but fish species richness increased with remnant pool size and depth. Invertebrate taxon richness increased with pool size. Invertebrate community composition differed by habitat type (pool versus seep) and gradients in composition were correlated with several abiotic factors (e.g. pool size, substrate, and canopy cover).
4. Remnant pools at Coyote Creek supported a full assemblage of native fishes and numerous imperilled taxa, including California red‐legged frogs and California floater mussels. Nearly all native fishes and imperilled taxa are absent from artificially perennial and urbanized reaches of Coyote Creek just a few kilometres downstream of the study area.
5. Remnant pools in intermittent streams should be a focus of conservation efforts in regions with a Mediterranean climate, especially during extreme droughts. Native fauna adapted to harsh intermittent flow regimes can thrive in these habitats, whereas non‐native taxa may fare poorly. Furthermore, remnant pools supported by deep groundwater sources, such as those along geological faults, may provide both ecological refuge and evolutionary refugia for freshwater biota.

     

Implications of hypoxia tolerance for wetland refugia use in Lake Nabugabo,Uganda

Aquatic hypoxia can affect predator‐prey interactions by altering the success rate of the predator and/or the vulnerability of prey. For example, in the Lake Victoria basin of East Africa, native prey exploit hypoxic wetlands as refugia from predation by introduced Nile perch (Lates niloticus). Here, it is predicted that species exploitation of wetlands depends on their hypoxia tolerance relative to the heterogeneity of wetland hypoxia. In this study, we compared the hypoxia tolerance of four fish taxa that differ in their use of hypoxic wetlands in Lake Nabugabo, Uganda: the cichlid Pseudocrenilabrus multicolor victoriae that inhabits the dense swamp interior; and three taxa that inhabit wetland ecotones including Nile tilapia (Oreochromis niloticus), L. niloticus and juvenile endemic haplochromine cichlids. We characterised hypoxia tolerance by exposing fish to progressive hypoxia and quantifying variation in aquatic surface respiration (ASR). The effect of body size on tolerance was explored in L. niloticus by quantifying ASR behaviour across a range of size classes. ASR behaviour was also compared between O. niloticus groups from wetland versus open‐water habitats to detect habitat‐associated intraspecific variation. The most tolerant taxon was the swamp specialist P. multicolor, indicated by its low ASR thresholds and small percentage of fish using ASR during the final sample interval. The other three taxa did not differ in ASR behaviour, and no differences were detected between O. niloticus groups. Body size effects were present for L. niloticus suggesting a lower tolerance to hypoxia in larger‐bodied individuals, thus limiting their ability to penetrate wetlands.     

Is current floodplain management a cause for concern for fish and bird conservation in Bangladesh's largest wetland?

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The value of small,natural and man‐made wetlands for bird diversity in the east Colombian Piedmont

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The establishment and impact of Pseudorasbora parva,an exotic fish species introduced into Lake Mikri Prespa (north-western Greece)

• 1. The topmouth gudgeon Pseudorasbora parva Schlegel, 1842, a south-east asian cyprinid, was introduced accidentally in the Danube Delta in Romania in the 1960s and has now achieved a pan-Danubian distribution. P. parva has been introduced into other countries, such as Greece, usually inadvertently included with other species imported for fish farming. In 1984–85, during a study of the fish populations of Lake Mikri Prespa (north-west Greece), several topmouth gudgeon were caught. Our aim was to study the changes in the population size of P. parva over nearly 10 years, to study its life history traits and to assess its potential impacts on native fish species.
• 2. The catch per unit effort (CPUE) in spring increased significantly between 1984 and 1992. We found evidence that P. parva has established a breeding population in the lake. P. parva combines many characteristics likely to favour a successful colonization (resistance to harsh climatic conditions, early sexual maturity, extended breeding season, broad dietary spectrum). Growth in Lake Prespa, where the oldest individuals captured were 3 years old, is very similar to that observed within its native range.
• 3. There is evidence for dietary overlap between P. parva and three endemic species: Paraphoxinus epiroticus prespensis, Cobitis meridionalis and Alburnoides bipunctatus ohridanus. No decline in the populations of these three species has yet been demonstrated. Other possible impacts are discussed. The successful colonization by P. parva was certainly favoured by the absence of a true piscivorous fish in the lake and by the isolation and high level of endemism of fish communities.
• 4. The conservation of the many endemic species in the lake should be a priority and the introduction of exotic species should be banned.

     

Efficacy of artificial refuge to enhance survival of young Barrens topminnows exposed to western mosquitofish

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Potential impacts of agricultural development on freshwater biodiversity in the Lake Victoria basin

1. The Lake Victoria basin (LVB) and its tributary rivers are a major biodiversity hot spot, containing at least 234 native fish species, 135 native aquatic plant species, and 50 native freshwater mollusc species. Lake Victoria itself is home to around 500 fish species, most of which are haplochromine cichlids.
2. The LVB is increasingly under threat from unsustainable land conversion and the intensification of agriculture. High population growth is driving the expansion of agriculture, urbanization, and freshwater abstractions, which have a profound impact on freshwater biodiversity. In addition, increased demand for agricultural crops from domestic and international markets are likely to lead to larger agricultural operations, further threatening freshwater biodiversity.
3. This study explores these potential future impacts on the biodiversity found in freshwater rivers in the LVB as a result of projected future changes in land use.
4. A newly developed database of land‐use impacts on freshwater biodiversity is introduced, with a focus on ecological community composition data from freshwater habitats under human pressures. Impacts on freshwater biodiversity are then projected under four different scenarios of land‐use change.
5. Results show that land use has a significant impact on freshwater biodiversity. Freshwater biodiversity is projected to be at most risk in sub‐basins in the Tanzanian, Rwandan, and Burundian part of the LVB, such as the Kagera and Magoga/Isonga sub‐basins. Local species richness levels are particularly affected in the Magoga/Isonga sub‐basin in Tanzania, with an average loss of 10.8% across all scenarios.
6. Model results show the potential to identify broad spatial patterns of likely threats and pressures on freshwater ecosystems under different socio‐economic futures. The analysis of these patterns, where they are consistent and where they differ, can support the production of policy that strives to find the optimal balance between development and conservation in an uncertain future.

     

Reduction of a native fish fauna by alien species: an example from Brazilian freshwater tropical lakes

The consequences of introducing Cichla cf. monoculus Spix & Agassiz, Astronotus ocellatus (Agassiz) and Pygocentrus nattereri Kner into lakes in the River Doce basin, Brazil, on richness, diversity and efficiency of aquatic macrophytes as natural refugia to native fishes was investigated. Samples were taken from lakes with and without alien fishes in areas with and without aquatic macrophytes. The presence of alien fishes reduced richness and diversity of the native fish community. The refugia function, which could be attributed to the clustering of aquatic macrophytes, does not exist in these lakes probably because the alien fishes exploit such habitats for reproduction. Since introductions threaten the native fish diversity of the region, studies on regional dispersion and factors that minimise the spread of alien fishes are needed.     

Population growth and mass mortality of an estuarine fish,Acanthopagrus butcheri,unlawfully introduced into an inland lake

• 1. In 2006, two periods of hypoxia resulted in the death of approximately 35 tonnes of black bream (Acanthopagrus butcheri) in Lake Indoon, a small inland lake in Western Australia.
• 2. Acanthopagrus butcheri was the first fish species to be recorded in this lake, along with the mosquitofish (Gambusia holbrooki) which was also observed during sampling in 2006. Acanthopagrus butcheri appears to have been introduced to Lake Indoon between 1998 and 2003 and formed a self‐sustaining population. It is believed to have been deliberately introduced for the purpose of creating a recreational fishery, despite the existence of substantial penalties for illegal translocation of fish in Western Australia.
• 3. Recent human‐induced environmental changes, including rising groundwater and salinization, have probably aided the establishment of both species in Lake Indoon. The importance of salinity to recruitment success by A. butcheri was indicated by the presence of only two age classes in 2006, with estimated recruitment dates coinciding with the years of highest recorded salinity in the lake.
• 4. The ‘fish kills’ provided an opportunity to examine aspects of A. butcheri biology in a relatively low salinity environment which is atypical for this estuarine species. In particular, the recruitment period in Lake Indoon was delayed until autumn/winter, rather than spring/summer as seen in other populations. Biological responses in Lake Indoon have implications for natural populations living in estuaries with modified salinity regimes.
• 5. The ecological, social and economic impacts potentially arising from the introduction of fish to Lake Indoon, which is an important migratory bird habitat and a recreational amenity for local residents and tourists, illustrate the complexities of fish translocation and the need for rigorous assessment before stocking to identify potential costs and benefits.

Copyright © 2008 John Wiley & Sons, Ltd.     

Managing invasive carp (Cyprinus carpio L.) for habitat enhancement at Botany Wetlands,Australia

• 1. In Australia, the carp Cyprinus carpio L. is regarded as a threat to the native fish and the aquatic environment. In recent years, Botany Wetlands, a significant coastal wetland in the Sydney region, has been invaded by the undesirable cyprinids, carp and goldfish (Carrasius auratus L.).
• 2. In 1996 a cyprinid removal programme commenced at Botany Wetlands with the objective of managing the invasive species, increasing fish diversity, reducing cyanobacterial blooms and hence enhancing the aquatic habitat. Using electrofishing and gill netting, 4073 carp and 261 goldfish, amounting to 10 117 kg of cyprinid biomass were removed between 1996 and 2004. The captured carp ranged from 60 to 835 mm. Males matured before females. Carp between 350 and 680 mm in length grew at the rate of 1.66±0.38 g day^?1.
• 3. The success of the programme was monitored by assessing four indicators related to carp populations and two related to habitat. The former included the pattern of length–frequency distribution, mean weight per size class, condition factor (CF) and the catch per unit effort (CPUE), and the latter the Secchi disc transparency and cyanobacterial counts. After 8 yr of removal, the plots of carp length–frequency distribution flattened, CF decreased from 2.86 to 1.82 and CPUE decreased from 97 kg day^?1 to 50 kg day^?1. A 10‐fold decrease occurred in cyanobacterial counts and the Secchi disc transparency increased by 20%. More than 20 000 fingerlings of native Australian bass Macquaria novemaculeata (S.) have been introduced to the wetlands, increasing the potential for juvenile carp predation and biodiversity.

Copyright © 2005 John Wiley & Sons, Ltd.     

Diversity of fish and fisheries from the Lake Encantada Environmental Protection Area,Ilhéus,Brazil

• 1. Lake Encantada, in the basin of the Almada River (Bahia State), has an area of 650.5 ha and a perimeter of 13.3 km. In 1993, an area of 11 800 ha around the lake was designated an Environmental Protection Area (Class V category). The main goal of this study was to identify fish species in the lake and to determine possible threats to their conservation.
• 2. Fish samples were obtained with gillnets between May 2002 and February 2004, and used to calculate indices of species diversity Shannon (H′), richness (d) and evenness (J′).
• 3. Eleven fish species comprising nine families were collected during the sampling period. Three introduced species occurred in the lake: Prochilodus argenteus, endemic to the São Francisco river basin, and two African species, Oreochromis niloticus and Clarias gariepinus. In overall ranking, Prochilodus argenteus was first in number of individuals (46%) and weight (59%), followed by Genidens genidens in numbers (31%), and Clarias gariepinus in weight (15%).
• 4. The occurrence and dominance of introduced species is discussed as well as their possible impacts on the trophic structure of this ecosystem.

Copyright © 2007 John Wiley & Sons, Ltd.     

Can management practices in rice fields contribute to amphibian conservation in southern Brazilian wetlands?

• 1. Rice field expansion is one of the activities associated with the disappearance of 90% of the wetlands in southern Brazil. Worldwide, rice agriculture has been recognized as having considerable potential value for many aquatic species. Nevertheless, management practices in such systems must be ameliorated and better investigated.
• 2. This study evaluated the potential role of rice fields as refugia for amphibians, and whether different hydrologic management practices after rice cultivation could contribute to wetland amphibian conservation in southern Brazil.
• 3. Six collections were made in six rice fields with different management practices after cultivation (three dry and three flooded) and three natural wetlands. The amphibians were sampled through six random 15‐min visual transects per collection in each rice field and the natural wetlands.
• 4. In total, 2139 anuran individuals were observed in rice fields (798) and Reserva Lake (1341), comprising 12 anuran species distributed among five anuran families. Anuran richness and abundance varied over the rice cultivating cycle, and they were higher in the growing phases than in the fallow phases. The mean anuran richness and abundance was higher in Reserva Lake than in flooded and dry rice fields.
• 5. The different management practices adopted after the harvesting period (presence or lack of surface water) did not influence the anuran richness and abundance. It did, however, influence species composition.
• 6. The difference in species composition between the management practices adopted is an interesting result in terms of biodiversity conservation. Rice producers could maintain part of their agricultural land flooded during the fallow phase as a strategy to preserve a higher diversity of anurans. These results should be taken into consideration in wetland conservation plans in southern Brazil; however, the percentage of each agricultural land that should be kept flooded should be decided by Brazilian agricultural and conservation policies. Such a strategy would reconcile agricultural/economic needs with the conservation of biodiversity in southern Brazil, where more than 90% of wetland systems have already been lost. Copyright © 2009 John Wiley & Sons, Ltd.

     

A case study on the population ecology of a topmouth gudgeon (Pseudorasbora parva) population in the UK and the implications for native fish communities

• 1. The topmouth gudgeon Pseudorasbora parva is a small Asian cyprinid species that has proved invasive throughout many European countries. Following an initial introduction into the wild in 1996, the species is now proving invasive in the UK, with at least 25 infested waters in England and Wales, of which 10 are known to have direct connection to a major river catchment.
• 2. To demonstrate the threat of P. parva to fisheries in the UK, a case study is presented on a lake located in the Lake District of England where the species was introduced in 2000. The species rapidly established a breeding population that, by 2003, was the dominant species in size classes <70 mm. In 2004, they were the only species in the lake that produced young‐of‐the‐year.
• 3. Individual P. parva adopted the reproductive tactics of early maturity, multiple spawning, male dominance and male nest guarding; sexual dimorphism was manifested in larger body size of males. These traits were in contrast to the resident, native species of the lake, including roach Rutilus rutilus and gudgeon Gobio gobio, which adopted traits of later maturity and single spawning.
• 4. This case study, therefore, revealed relatively rapid establishment of a P. parva population, their subsequent numerical dominance of the fish community, and the impediment of the recruitment of native fish. The implications for UK fisheries are concerning: should P. parva continue to disperse and individuals adopt similar traits as those in this case study, there may be few waters immune from their invasion, numerical dominance and subsequent impacts.

Copyright © 2006 John Wiley & Sons, Ltd.     

To sea or not to sea? Multiple lines of evidence reveal the contribution of non‐diadromous recruitment for supporting endemic fish populations within New Zealand's longest river

1. Otolith microchemistry was used to identify marine‐ versus freshwater‐derived recruitment of three native freshwater fish species belonging to the southern hemisphere family Galaxiidae, in New Zealand's longest river system, the Waikato River.
2. Water chemistry data for trace elements and ⁸⁷Sr/⁸⁶Sr isotope ratios were collected from five lentic and 10 lotic water bodies throughout the lower river floodplain. Potential spawning sites for galaxiids were compared with values obtained by laser ablation inductively coupled mass spectrometry (LA‐ICPMS) depth profiling of young‐of‐the‐year otoliths sampled from fish in nine lower river catchment sites.
3. Otolith chemical signatures from the larval rearing period indicated that catchment‐scale recruitment for two species, Galaxias argenteus (Gmelin, 1789) and Galaxias fasciatus Gray, 1842, was driven predominantly by non‐diadromous recruitment from one lake (Lake Waahi). In contrast, diadromous recruitment appeared to be more common for Galaxias maculatus (Jenyns, 1842); however, non‐diadromous specimens were also identified for the first time from a New Zealand river.
4. Reversing lake outlet flows linked to river stage appears be important in facilitating the dispersal of rheotactic larvae out of lakes, suggesting that lake outflow management at key times could be used to sustain this ecologically important function.
5. This study highlights that some water bodies can supply a disproportionately large number of recruits to support fish populations within the wider riverscape. Identifying these water bodies and managing them to sustain recruitment is key to the conservation of non‐diadromous Galaxiidae in this modified lowland environment

     

Nearshore fish assemblages associated with introduced predatory fishes in lakes

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Status,ecology and conservation of an endemic fish,Oreochromis niloticus baringoensis,in Lake Baringo,Kenya

• 1. The tilapia Oreochromis niloticus baringoensis is a genetically and morphologically distinct sub‐species of Oreochromis niloticus endemic to Lake Baringo, Kenya. In recent years, concern has been expressed as to its status. Recent declines in catch returns suggest the population may be threatened, with conservation action required to safeguard it.
• 2. Catch returns from the Baringo fishery since 1964 have shown considerable fluctuations for all species, but especially O. n. baringoensis. From a peak of 712 t in 1970, their total catch was only 5 t in 2005, despite a 2 year period of closure in 2002 and 2003. Changes in fishery catch and relative abundance were independent of exploitation in the fishery but were significantly correlated with lake level.
• 3. Few individuals were captured at lengths >250 mm, with no fish sampled >284 mm. During periods of high lake level, individuals matured at smaller sizes and were capable of growing to larger ultimate sizes. With maturity at lengths <130 mm and fishery regulations preventing removal of fish <180 mm, there was a relatively large proportion of mature fish that was below exploitation size each year (19 to 44%). In most years, the proportion of fish available for exploitation was <10%.
• 4. Stable isotope analyses revealed O. n. baringoensis was reliant upon planktonic basal resources and zooplankton carbon. There was only minimal trophic overlap with other fish species in the lake, indicating little potential for competition for food resources.
• 5. These data suggest that the population status of O. n. baringoensis is not threatened per se, but subject to an unpredictable and unstable environment that strongly influences their life‐history traits and, ultimately, their population abundance, and should be managed accordingly.

Copyright © 2008 John Wiley & Sons, Ltd.     

Fish movements: the introduction pathway for topmouth gudgeon Pseudorasbora parva and other non‐native fishes in the UK

• 1. The contamination of fish consignments (for stocking or aquaculture) is a major pathway by which non‐native organisms, including fish, are introduced to new areas. One of the best examples of this is the topmouth gudgeon Pseudorasbora parva, which was accidentally imported into Romania and then throughout Europe in consignments of Asian carp species.
• 2. The introduction and spread of topmouth gudgeon in the UK has been linked to imports and movements of the ornamental variety (golden orfe) of ide Leuciscus idus. To examine this hypothesis, relationships between authorized movements of both native and non‐native fish species (in particular ide) and the occurrence in England of topmouth gudgeon were tested at the 10×10 km scale.
• 3. Topmouth gudgeon occurrence in the wild was significantly correlated with the trajectories of movements of ornamental fish species (ide/orfe, sunbleak Leucaspius delineatus) as well as a few non‐ornamental fish species (European catfish Silurus glanis, Atlantic salmon Salmo salar and grass carp Ctenopharyngodon idella).
• 4. These results highlight the mechanism by which non‐native fish species disperse from the point of first introduction, and especially that movements of fish within the country represent an important mechanism for accidental introductions of non‐native species. © Crown copyright 2010. Reproduced with the permission of Her Majesty's Stationery Office. Published by John Wiley & Sons, Ltd.

     

Long‐term changes in the fish assemblage of the Laja River,Guanajuato, central Mexico

• 1. The structure and composition of fish communities in rivers of central Mexico have been altered as a result of water over‐exploitation, habitat fragmentation, introduction of exotic species, and pollution. However, the specific pattern and degree of change are poorly documented.
• 2. Long‐term information from the Laja River (Guanajuato, Mexico) in the Lerma River basin was used to explore trends in fish species richness and community composition (species origin, trophic niche, tolerance, and preferred habitat) from the 1960s to the present in both river and reservoir sites.
• 3. Declines in native, sensitive, benthic native and carnivore species ranging from 11% to 30% per decade, and increases in the number of tolerant and exotic species by 9–20% per decade, are documented. Repeated measures ANOVA and sign tests revealed significant declines in the number of benthic, native, carnivore and sensitive species. Species richness, number of exotics, tolerant species and omnivore species did not change statistically, though statistical power was low. Some important changes occurred in these variables, such as the expansion and establishment of exotics such as Xiphophorus variatus and Micropterus salmoides, which pose a serious potential threat to native species.
• 4. The changes in fish community composition for the Laja portray how the fish communities in other rivers in central Mexico, for which long‐term data do not exist, have changed or could change if environmental deterioration continues.

Copyright © 2006 John Wiley & Sons, Ltd.