Freshwater fish biodiversity restoration in floodplain rivers requires connectivity and habitat heterogeneity at multiple spatial scales.

With a sixth mass extinction looming and freshwater biodiversity declining at unprecedented rates, evaluating ecological efficacy of river restoration efforts is critical in combatting global biodiversity loss. Here, we present a comprehensive study of the functioning for fishes of 46 river restoration projects in the river Rhine, one of the world's most heavily engineered lowland rivers. Floodplains with permanent, either one- or two-sided lateral connectivity to the main channel, favour total fish abundance, and are essential as nursery areas for riverine fishes. Habitat heterogeneity had a strong positive effect on species richness but was negatively related with fish abundances. However, the effects of environmental variables varied between ecological groups and spatial scales. Surprisingly, richness of critical rheophilic fishes declined with large-scale habitat heterogeneity (~1000 m), while it increased at small scales (~100 m), possibly because of the presence of unfavourable habitats for this ecological group at larger scales. Clearly, there is no one-size-fits-all design for river restoration projects. Whether a river section is free-flowing or impounded dictates the scope and efficacy of restoration projects and, within a river section, multiple complementary restoration projects might be key to mitigate freshwater fish biodiversity loss. An essential element for success is that these projects should retain permanent lateral connection to the main channel.

30 years of large river restoration: How long do restored floodplain channels remain suitable for targeted rheophilic fishes in the lower river Rhine?

The ecological efficacy of river restoration projects may change over time, resulting in the loss of their ecological function for targeted species. The goal of this study was to evaluate the rheophilic nursery function of restored floodplain channels over time, by analysing 30 years of monitoring data from 12 restoration projects in the lower river Rhine. We hypothesised that the nursery function would change over time, caused by the combined effects of decreasing flow conditions and succession processes affecting habitat heterogeneity. We found that nursery area suitability for rheophilic fish was almost 4 times higher in two-sided connected channels than in one-sided connected channels, although the response trends of rheophilic fish were similar for both water body types. These response curves showed clear optima with channel age, for rheophilic fish abundance at 13 to 14 years post-restoration, indicating optimal nursery conditions. On the other hand, rheophilic species richness showed a steadily decreasing trend with channel age, suggesting aging channels became less suitable as nursery areas for most rheophilic fish species. The presence of permanent flow was found to be the main driving factor in explaining both rheophilic fish community trends and habitat succession in individual restored channels. We did not observe an effect of habitat heterogeneity on nursery function for rheophilic fish. To create and maintain optimal nursery conditions in restored floodplain channels of strongly anthropogenically influenced rivers such as the river Rhine, we propose a management strategy involving cyclic rejuvenation through human intervention, focusing on restoring permanent flow, with a frequency of on average every 15 years, depending on the rate of aggradation and targeted rheophilic species. We also propose a thorough investigation of the relationship between habitat heterogeneity and nursery success in floodplain channels, as a next step in the identification of suitable nursery areas for rheophilic fishes.

Out of the pot and into the fire: Explaining the vulnerability of an endangered small headwater stream fish to black-bass Micropterus spp. invasion.

Introduced predatory fishes have had consistently severe consequences for native fishes in stream environments around the world, although the drivers of these effects are often unclear. In the Swartkops River headwaters in South Africa, native Eastern Cape redfin Pseudobarbus afer were always absent from sites occupied by non-native black basses Micropterus salmoides and Micropterus dolomieu, but generally co-occurred with the native predators Anguilla marmorata and Anguilla mossambica. A natural experiment provided by flood-mediated recolonization of black-bass occupied sites by P. afer demonstrated depletion in black-bass invaded sites. Field behavioural observations of P. afer indicated that they foraged among benthic cover during the day, but suspended in open water at night. As the nocturnal A. marmorata and A. mossambica foraged actively within structural cover at night and M. dolomieu and M. salmoides are diurnal or crepuscular predators, P .afer is thus optimized to avoid predation by native anguillid predators and not the functionally unique predatory black basses. The integration of distributional, temporal population dynamics and behavioural data suggests that the severe effects of Micropterus spp. are probably a consequence of prey naïveté and behaviour evolved to evade native predators.

Shallow genetic divergence and species delineations in the endemic Labeobarbus species flock of Lake Tana, Ethiopia.

To assess whether the species distinctions of Lake Tana's Labeobarbus spp. are supported by genetic information, microsatellite markers were used. A total of 376 Labeobarbus spp., belonging to 24 populations of 11 species from three regions of the lake (north, south and east), were sampled. Eight microsatellite markers were analysed. In general, differences between conspecific populations were smaller than differences between populations of different species. For six species, conspecific populations from different regions in the lake were consistently more similar than populations of other species from the same region. For four species this was not the case, while for one species two populations were similar, but different from the third population. River-spawning species appeared to be more distinct than presumed lake spawners. On the species level, there was a significant correlation between genetic and morphological differentiation, especially in morphological aspects associated with ecological functioning. This suggests that genetic differentiation arose together with adaptive radiation, although the overall genetic differentiation among the Lake Tana Labeobarbus spp. is small.

Spawning migrations of the endemic Labeobarbus (Cyprinidae, Teleostei) species of Lake Tana, Ethiopia: status and threats.

The reproductive biology of the only known intact species flock of large cyprinids, the 16 Labeobarbus species of Lake Tana (Ethiopia), has been extensively studied for the past two decades. Seven species of Labeobarbus are known to migrate >50 km upstream into tributary rivers for spawning during the rainy season (July to October), whereas eight other species are absent from these rivers and probably developed a new strategy of lacustrine spawning (macro-spatial segregation). One species (L. intermedius) probably spawns in the lake as well as in the rivers. Between the early 1990s and 2000s, the riverine spawners showed a decline of 75% in both biomass and number in both fishery independent surveys and in commercial catches. Reproductive migration makes fishes vulnerable to fisheries and other threats like habitat modifications. Lacustrine spawners are probably more resilient as they are not known to form spawning aggregations that can easily be exploited by fishermen. In addition, upstream rivers and catchments around Lake Tana are highly degraded by erosion and recently subjected to intensive habitat modification for irrigation and hydroelectric power generation. This article reviews results of field studies on the Labeobarbus spawning migration from Lake Tana to spawning rivers, giving emphasis on segregation and homing. It also summarizes existing and emerging threats which form potential causes for the decline of the migratory Labeobarbus species. Knowledge gaps on the reproductive biology are identified for further investigation.

Evolution of MHC class II beta chain-encoding genes in the Lake Tana barbel species flock (Barbus intermedius complex).

Major histocompatibility complex (MHC) class II protein polymorphism is maintained in allelic lineages which evolve in a trans-specific manner, passing from one species to descendant species. Selection pressure on peptide binding residues should be greatest during speciation, when organisms move into new environments and their MHC molecules encounter new pathogens. The isolation of MHC genes from teleost fishes, the most diverse group of vertebrates, has created possibilities for testing this hypothesis. The large barbels of Lake Tana have undergone an adaptive radiation within the last 5 million years, producing 14 morphotypes which inhabit different ecological niches within the lake. We studied the variability in class II beta chain-encoding genes of four of these morphotypes using polymerase chain reaction amplification and DNA sequencing. The sequences obtained were orthologous to four of the known class II genes from the common carp, from which barbels diverged approximately 32 million years ago. When subjected to phylogenetic analysis, the 48 sequences clustered into groups which represent allelic lineages. A comparison of nonsynonymous and synonymous substitutions between the peptide binding region codons and non-peptide binding region codons of these sequences revealed that they are under strong selective pressure.