L.A.J. Nagelkerke

Zooplankton, fish communities and the role of planktivory in nine Ethiopian lakes

Fish and zooplankton populations of nine Ethiopian freshwater lakes were quantitatively sampled along a North–South gradient. Differences in altitude and latitude resulted in a temperature gradient from North to South. We tested three hypotheses: (1) the degree of zooplanktivory decreases with water temperature, i.e. from North to South; (2) the degree of zooplanktivory increases with the abundance of large-bodied zooplankton; and (3) the pattern of zooplanktivory in eutrophic Ethiopian water bodies differs from other tropical and temperate water bodies. Proportions of zooplanktivory in the fish communities did not show a geographical trend, but mainly depended on fish species, zooplankton density and the availability of large-bodied cladocerans. The degree of zooplanktivory in eutrophic Ethiopian water bodies differs from other eutrophic water bodies, both temperate and tropical. In Ethiopia, the degree of zooplanktivory can be both low and high, in contrast with other tropical water bodies where zooplanktivory is generally low and with temperate eutrophic water bodies where it is generally high. As a result, predation pressure on zooplankton by fish varies dramatically amongst Ethiopian water bodies.

The composition of fish communities of nine Ethiopian lakes along a north-south gradient: threats and possible solutions

Fish populations of nine Ethiopian freshwater lakes were quantitatively sampled with a standardized protocol, using multi-mesh gill nets. In total, 27 species were identified, but only 14 species were common. Based on the common species, the fish communities showed large differences in their species composition, except for Lake Abaya and Lake Chamo which were similar. Most fish species were observed in only one or two lakes. Compared with the information reported in literature the present study generally underestimated the species richness. The empirical model of Amarasinghe and Welcomme (2002) for African lakes was used to estimate fish species richness, which was compared with species presence reported in literature. Biodiversity in the two northern highland lakes is low, but not lower than the model estimate. Lake Tana has a high biodiversity which is close to what is estimated by the model, but three Rift Valley lakes have low biodiversity, lower than estimated by the model. There are also strong indications for the Rift Valley lakes that species richness was higher in the past because the species richness reported in the older literature was generally much higher than those observed by us in the present study and those reported in the more recent literature. Threats like overfishing, high sediment load and degradation of habitats were identified. It is recommended that Ethiopia should develop guidelines for fishery legislation and implement it through an enforcement agency. Moreover, catchments management should be practiced to save the water bodies and their fish communities.

Nile perch distribution in south-east Lake Victoria is more strongly driven by abiotic factors, than by prey densities

We studied the effects of environmental driving factors (maximum depth, visibility, oxygen, temperature, and prey densities) on the distribution and diet composition of Nile perch (Lates niloticus) in south-east Lake Victoria from 2009 to 2011. We tested the hypotheses that (i) Nile perch distribution is regulated by the same environmental factors on a local scale (Mwanza Gulf) and on a regional scale (Mwanza Gulf, Speke Gulf and the open lake in Sengerema district), and (ii) driving factors act differently on different Nile perch size classes. Fish were sampled with gillnets. Nile perch densities were highest in the shallow part of the Mwanza Gulf and during the wet seasons, mainly caused by high densities of juveniles. The environmental driving factors explained Nile perch distributions on both regional and local scales in a similar way, often showing non-linear relationships. Maximum depth and temperature were the best predictors of Nile perch densities. Prey densities of shrimp and haplochromines did not strongly affect Nile perch distributions, but did explain Nile perch diet on a local and regional scale. We conclude that abiotic variables drive Nile perch distributions more strongly than prey densities and that feeding takes place opportunistically.

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.

I ‐ The ecological and evolutionary significance of morphological diversity

Abstract The Lake Tana (northern Ethiopia) ichthyofauna is dominated by ‘large’ barbs (Barbus spp., Cyprinidae). Recently, fourteen Bar‐bus morphs have been described. This paper provides evidence that these morphs are indeed biological species. To investigate the hypothesis that disruptive selection on feeding‐related morphological characters was the evolutionary driving force of speci‐ation, an ecomorphological analysis, investigating the quantitative relationships between morphology and feeding abilities on a number of aquatic food types was investigated. The analysis accurately predicted the potential trophic niches of the Lake Tana barbs and supported the disruptive selection hypothesis. Finally, a hypothetical evolutionary scenario is presented in which disruptive selection led to initial morphological differentiation, and spatial and temporal spawning segregation stabilized the genetic differences among the morphs, resulting in speciation.

Evidence for ontogenetically and morphologically distinct alternative reproductive tactics in the invasive Round Goby Neogobius melanostomus

Alternative reproductive tactics are characterized by the occurrence of discrete alternative morphs that differ in behavioural, morphological and physiological traits within the same sex. Although much effort has been made to describe the behaviour, morphology and physiology of such alternative morphs, less effort has been invested investigating how much overlap there is in the characteristics of such morphs in natural populations. We studied random population samples of the invasive Round Goby Neogobius melanostomus from five different localities in the river Rhine system in the Netherlands. We found two morphologically and physiologically distinct male morphs which likely represent alternative reproductive tactics. Almost all mature males under 9.35 cm total length had a gonadosomatic index > 3%, suggestive of a sneaker tactic, while nearly all males above 9.35 cm has a gonadosomatic index of < 3%, suggestive of a parental tactic. Cheek size and eye diameter alone were sufficient to distinguish the two morphs. Gonads had a different relationship with size in the two morphs, indicating separate growth trajectories. The gonad mass of sneaker morphs would be ca. 7.5 times as high as the gonad mass of parental morphs of the same total length after extrapolation. Few (9%) intermediates were found, suggesting that the expression of alternative reproductive tactics is determined before the first breeding season. This contrasts with studies on other goby species, which show evidence of plastic tactics that can be affected by social circumstances. We conclude that it is possible to distinguish two alternative male morphs in the Dutch Round Goby population using morphological measurements alone. Although behavioural observations are needed to provide conclusive evidence, the difference in GSI between these morphs indicates that these morphs reflect alternative reproductive tactics.

Trophic niche-space imaging, using resource and consumer traits

The strength of trophic (feeding) links between two species depends on the traits of both the consumer and the resource. But which traits of consumer and resource have to be measured to predict link strengths, and how many? A novel theoretical framework for systematically determining trophic traits from empirical data was recently proposed. Here we demonstrate this approach for a group of 14 consumer fish species (Labeobarbus spp., Cyprinidae) and 11 aquatic resource categories coexisting in Lake Tana in northern Ethiopia, analysing large sets of phenotypic consumer and resource traits with known roles in feeding ecology. We systematically reconstruct structure and geometry of trophic niche space, in which link strengths are predicted by the distances between consumers and resources. These distances are then represented graphically resulting in an image of trophic niche space and its occupancy. We find trophic niche to be multidimensional. Among the models we analysed, one with two resource and two consumer traits had the highest predictive power for link strength. Results further suggest that trophic niche space has a pseudo-Euclidean geometry, meaning that link strength decays with distance in some dimensions of trophic niche space, while it increases with distance in other dimensions. Our analysis not only informs theory and modelling but may also be helpful for predicting trophic link strengths for pairs of other, similar species.

Utilization of seagrass habitats by juvenile groupers and snappers in Banten Bay, Banten Province, Indonesia

Coastal development in Banten Bay, Indonesia, decreased seagrass coverage to only 1.5% of its surface area. We investigated the importance of seagrass as habitat for juvenile groupers (Serranidae) and snappers (Lutjanidae), by performing beam trawl hauls on a weekly basis in two seagrass locations and one mudflat area, and monthly trawl hauls in three different microhabitats (dense, mixed and patchy seagrass) in one of the seagrass locations. We studied the effects of location and microhabitat, as well as temporal patterns (diel, weekly and monthly) on the probability of occurrence and abundance of the most abundant grouper (Orange-spotted grouper, Epinephelus coioides) and snapper (Russell’s snapper, Lutjanus russellii). We found that both species were almost exclusively found in seagrass locations, with a preference for microhabitats of high complexity (dense and mixed microhabitats). L. russellii had a higher probability of catch and abundance during the night, most probably because of its ability to avoid the beam trawl during daytime sampling. In addition there was an effect of week and month on the presence and abundance of both species, but patterns were unclear, probably because of high fishing pressure on juvenile groupers and snappers by push net fishermen. Groupers and snappers mainly fed on abundant shrimps, and to a lesser extent on fish. Moreover, juveniles find protection against predators in seagrass, which confirmed the critical role of quantity and quality of seagrass areas for juvenile groupers and snappers in Banten Bay.

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

To assess whether the species distinctions of Lake Tanas 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.

Heterogeneity in food-web interactions of fish in the Mwanza Gulf, Lake Victoria: a quantitative stable isotope study

Stable isotope analyses and derived population-level metrics were used to quantitatively analyse spatial and seasonal heterogeneity in the fish trophic dynamics in relation to environmental variables in Mwanza Gulf, Lake Victoria (Tanzania). The fish community in Lake Victoria, including the top predator Nile perch, is generally omnivorous with a heavy reliance on invertebrates. This is in contrast to findings based on stomach content analyses of Nile perch, which showed a stronger reliance on fish. We tested two hypotheses: (1) during the rainy seasons multiple carbon sources influence the food-web structure inside the Gulf, leading to increased carbon ranges and trophic diversity. (2) During dry periods, the food-web structure mainly relies on pelagic primary production, reducing carbon ranges and trophic diversity. Carbon sources indeed varied seasonally and spatially, affecting the fish community at the highest trophic levels. With the onset of rains, carbon sources became spatially highly differentiated with enriched δ13C values of fish in shallow water inside the Gulf and depleted δ13C values in open waters. Metrics associated with niche size correlated significantly with seasonally varying environmental variables, while δ13C ranges correlated with spatially varying environmental variables.