[Show abstract][Hide abstract] ABSTRACT: Species that cross strong environmental gradients are expected to face divergent selective pressures that can act on sexually-selected traits. In the present study, we examine the role of hypoxia and carotenoid availability in driving divergence in two sexually-selected traits, male colour and reproductive behaviour, in the African cichlid Pseudocrenilabrus multicolor victoriae. Low-dissolved oxygen (DO) (hypoxic) environments are expected to be energetically challenging; given that male nuptial colour expression and courtship displays can be costly, we expected fish in low-DO versus high-DO environments to differ in these traits. First, a field survey was used to describe natural variation in male nuptial colour patterns and diet across habitats divergent in DO. Next, using wild-caught fish from a low-DO and high-DO habitat, we tested for differences in reproductive behaviour. Finally, a laboratory rearing experiment was used to quantify the interaction of DO and diet (low- versus high-carotenoid availability) on the expression of male colour during development. In energetically challenging low-DO environments, fish were more red and, in high-DO environments, fish were typically brighter and more yellow. The frequency of reproductive displays in fish of low-DO origin was 75% lower, although this had no consequence for brooding frequency (i.e. both populations produced the same number of broods on average). Our laboratory rearing study showed carotenoid availability to be important in colour production with no direct influence of DO on colour. Additionally, weak patterns of diet variation across wild populations suggest that other factors in combination with diet are contributing to colour divergence.
No preview · Article · Jan 2016 · Biological Journal of the Linnean Society
[Show abstract][Hide abstract] ABSTRACT: Tropical inland fishes are predicted to be especially vulnerable to thermal stress because they experience small temperature fluctuations that may select for narrow thermal windows. In this study, we measured resting metabolic rate (RMR), critical oxygen tension (Pcrit) and critical thermal maximum (CTMax) of the widespread African cichlid (Pseudocrenilabrus multicolor victoriae) in response to short-term acclimation to temperatures within and above their natural thermal range. Pseudocrenilabrus multicolor collected in Lake Kayanja, Uganda, a population living near the upper thermal range of the species, were acclimated to 23, 26, 29 and 32°C for 3 days directly after capture, and RMR and Pcrit were then quantified. In a second group of P. multicolor from the same population, CTMax and the thermal onset of agitation were determined for fish acclimated to 26, 29 and 32°C for 7 days. Both RMR and Pcrit were significantly higher in fish acclimated to 32°C, indicating decreased tolerance to hypoxia and increased metabolic requirements at temperatures only slightly (∼1°C) above their natural thermal range. The CTMax increased with acclimation temperature, indicating some degree of thermal compensation induced by short-term exposure to higher temperatures. However, agitation temperature (likely to represent an avoidance response to increased temperature during CTMax trials) showed no increase with acclimation temperature. Overall, the results of this study demonstrate that P. multicolor is able to maintain its RMR and Pcrit across the range of temperatures characteristic of its natural habitat, but incurs a higher cost of resting metabolism and reduced hypoxia tolerance at temperatures slightly above its present range.
Full-text · Article · Dec 2015 · Conservation Physiology
[Show abstract][Hide abstract] ABSTRACT: This study focused on variation in fish mercury (Hg) concentrations in 185 Nile perch (Lates niloticus) samples collected across four different habitat types in Lake Nabugabo, Uganda, a tropical lake located proximate to Lake Victoria. We quantified the stomach contents of Nile perch using the % index of relative importance, as well as, nitrogen and carbon isotopic concentrations to assess the role of diet and trophic level on Hg concentrations. In each habitat, we also evaluated a suite of chemical and physical characteristics that are commonly associated with variation in Hg bioavailability in temperate systems. Using linear mixed models and ANOVA, we demonstrate that habitat of capture is an important predictor of Hg concentrations in Nile perch from Lake Nabugabo and that the relationship between habitat and Hg is size and diet dependent. Nile perch diet as well as dissolved oxygen concentration and pH were also correlated with observed differences in fish Hg. Overall, Hg concentrations in Nile perch were all well below the WHO/FAO recommended guideline of 500 ng/g (mean 13.6 ± 0.4 ng/g wet weight; range 4.9 and 29.3 ng/g wet weight). This work contributes to a growing awareness of intra-lake divergence in Nile perch, as well as, divergence in Hg concentrations between varying aquatic habitat types, particularly wetlands.
[Show abstract][Hide abstract] ABSTRACT: Human activities, such as species introductions, are dramatically and rapidly altering natural ecological processes, and often result in novel selection regimes. To date, we still have a limited understanding of the extent to which such anthropogenic selection may be driving contemporary phenotypic change in natural populations. Here we test whether the introduction of the piscivorous Nile perch, Lates niloticus, into East Africa's Lake Victoria and nearby lakes coincided with morphological change in one resilient native prey species, the cyprinid fish Rastrineobola argentea. Drawing on prior eco-morphological research, we predicted that this novel predator would select for increased allocation to the caudal region in R. argentea to enhance burst-swimming performance, and hence escape ability. To test this prediction, we compared body morphology of R. argentea across space (nine Ugandan lakes differing in Nile perch invasion history) and through time (before and after establishment of Nile perch in Lake Victoria). Spatial comparisons of contemporary populations only partially supported our predictions, with R. argentea from some invaded lakes having larger caudal regions and smaller heads compared to R. argentea from uninvaded lakes. There was no clear evidence of predator-associated change in body shape over time in Lake Victoria. We conclude that R. argentea have not responded to the presence of Nile perch with consistent morphological changes, and that other factors are driving observed patterns of body shape variation in R. argentea. This article is protected by copyright. All rights reserved.
This article is protected by copyright. All rights reserved.
Full-text · Article · Aug 2015 · Journal of Evolutionary Biology
[Show abstract][Hide abstract] ABSTRACT: The methylated form of mercury (methylmercury) is a potent neurotoxicant and a contaminant of concern for fisheries because of its potential effects on ecosystem and human health. In Africa, inland fisheries are a crucial component of food and economic security, yet little information is available on Hg contamination trends. The authors compiled published data on mercury (Hg) contamination in African freshwater fishes, invertebrates, and plankton, as well as on potential drivers of Hg concentrations in these organisms. From 30 identified studies the authors assembled 407 total Hg concentrations from 166 fish species, 10 types of invertebrates, and various plankton, distributed across 31 water bodies in 12 countries. In fishes, total Hg concentrations, expressed as mean (± standard deviation) per location, averaged 156.0 ± 328.0 ng/g wet weight and ranged from 5.5 ng/g wet weight to 1865.0 ng/g wet weight. Only locations with nearby artisanal and small-scale gold mining operations had mean Hg concentrations above the World Health Organization/Food and Agriculture Organization's recommended guideline for fish (500 ng/g wet wt). The authors used mixed models to detect relationships between fish Hg concentrations and trophic level, mass, latitude, and chlorophyll a. Mass, trophic level, and latitude were all positive predictors of Hg concentration, confirming the presence of Hg bioaccumulation and biomagnification in African fishes. Although strong trends in Hg concentrations were evident, the present study also highlights limited availability of Hg data in Africa. This article is protected by copyright. All rights reserved.
This article is protected by copyright. All rights reserved.
No preview · Article · Feb 2015 · Environmental Toxicology and Chemistry
[Show abstract][Hide abstract] ABSTRACT: Aquatic hypoxia (low oxygen) provides a useful system for exploring ecological and evolutionary consequences of living under extreme conditions. It is also an environmental stressor of accelerating interest due to human activities that have increased the extent of hypoxic waters on a global scale. This chapter characterizes the distribution of hypoxic habitats, reviews key adaptations of fishes to extreme hypoxia, and explores the role of hypoxia as a divergent selective factor. Trade-offs in the costs and benefits of living in hypoxic and normoxic habitats may contribute to faunal diversification by creating spatially divergent selection that leads to specialized phenotypes as illustrated in studies of African fishes from hypoxic swamps and associated normoxic sites. In these systems alternative dissolved oxygen (DO) environments provide a strong predictor of intraspecific variation, particularly in traits related to oxygen uptake efficiency or oxygen limitations, but also in characteristics indirectly affected through trait correlations. Studies of fish persisting under hypoxia highlight the importance of localized extreme habitats as model systems for studying divergent natural selection and more generally for exploring effects of physicochemical stressors on ecological and evolutionary processes.
[Show abstract][Hide abstract] ABSTRACT: External body colour is an important trait contributing to phenotypic diversity and individual fitness in fish species. In this study, we use a combination of experimental techniques and field observations to examine patterns of colour divergence in the introduced Nile perch population of Lake Nabugabo, Uganda. We aim to determine whether the proportion of black–brown body colouration of Nile perch differs over a number of size classes, across ecologically distinct wetland edge and forest edge habitats, and whether these differences are the result of rapid (physiological) or ontogenetic (morphological) colour change. We found substantial colour differences in Nile perch between habitats, but trends were not consistent across size classes. Small Nile perch (<15 cm SL) from wetland edge habitats had darker skin pigmentation than those from forest edge; however, no significant colour differences existed between medium (15–40 cm SL) and large (>40 cm SL) Nile perch. Inter-habitat differences in colour in the small size class, and shifts in colour from juvenile to adult appear to be the result of morphological colour change associated with distinct ontogenetic shifts in resource use.
[Show abstract][Hide abstract] ABSTRACT: Introduced species can have profound direct ecological impacts on native species, yet their potential indirect effects remain relatively unexplored. For instance, introduced predators may directly affect some native species via predation, which may in turn have indirect consequences for other species that are released from competition.We explore this possibility in East Africa's Lake Victoria basin, where the introduction of the predatory Nile perch, Lates niloticus, in the 1950s and 60s contributed to the overall or local extinction of hundreds of native fish by the 1980s. We ask whether this dramatic change in assemblage composition has led to competitive release and niche expansion in Rastrineobola argentea, a resilient native cyprinid that has thrived in this highly perturbed ecosystem.To address this question, we compare the trophic ecology of R. argentea before (1966) and after (2011) the introduction of the Nile perch in Lake Victoria; and across eight satellite lakes that differ in their history of Nile perch invasion. Using this combination of spatial and temporal comparisons, we test for increases in dietary niche breadth (niche expansion) and changes in the level of individual specialisation of R. argentea in invaded versus uninvaded contexts.In our historical comparison, we find good evidence for dietary niche expansion and an increase in interindividual diet variation in R. argentea over time. Across lakes, however, strong bottom-up effects (i.e. variation in prey availability) appear to obscure any potential top-down effects of the Nile perch introduction on the trophic ecology of R. argentea.Overall, we find substantial temporal and spatial variation in the diet and niche breadth of R. argentea, but the underlying drivers remain uncertain, given the complexity of both anthropogenic and natural ecological changes in Lake Victoria over the past century. Understanding both the direct and indirect impacts of introduced species is challenging, but important for successful long-term management of human-altered ecosystems.
No preview · Article · Sep 2014 · Freshwater Biology
[Show abstract][Hide abstract] ABSTRACT: Inland fisheries represent an important source of protein and income for many communities, particularly in the tropics. It has been shown that ectotherms living in climatically stable tropical environments tend to be thermal specialists, and that some of them achieve their optimal metabolic performance at temperatures near their upper tolerance limits. Consequently, tropical freshwater fishes may be exceptionally sensitive to even small changes in temperature anticipated from global climate change. Despite that threat, the thermal biology of tropical freshwater fishes has received little attention. This project aims to assess whether the thermal sensitivity of fish metabolism and energetics can be used to forecast the vulnerability of key tropical freshwater fishes to climate change. Experiments are conducted at three locations (Brazil, Uganda, and Cambodia), where we quantify the metabolic capacities of two species acclimated for 3 weeks to three water temperature treatments (ambient, ambient + 2°C, and ambient + 4°C). The project addresses a time sensitive conservation problem, with a goal to contribute to the protection of culturally and socio-economically important fish species in the tropics.
[Show abstract][Hide abstract] ABSTRACT: The use of non-lethal experiments to elucidate behavioural and physiological thresholds to environmental stressors can provide valuable data for identifying threats to, and critical habitat of, imperilled species. Increased turbidity contributes to population declines and loss of fish diversity globally, but the complex direct and indirect effects of this stressor on imperilled species are not well understood.Response to turbidity was examined in a group of closely related North American fishes (Notropis spp.), including four imperilled shiners: pugnose shiner (N. anogenus), bridle shiner (N. bifrenatus), blacknose shiner (N. heterolepis), blackchin shiner (N. heterodon), and the more common mimic shiner (N. volucellus). Two experiments quantified (i) the behavioural response of each species to progressively increasing turbidity, and (ii) critical swimming speed of fishes acclimated in turbid- and clear-water treatments to provide insights into effects of low turbidity on aerobic performance.Very low levels of turbidity (
Full-text · Article · Aug 2014 · Aquatic Conservation Marine and Freshwater Ecosystems
[Show abstract][Hide abstract] ABSTRACT: African tropical lakes provide vital ecosystem services including food and water to some of the fastest growing human populations, yet they are among the most understudied ecosystems in the world. The consequences of climate change and other stressors on the tropical lakes of Africa have been informed by long-term analyses, but these studies have largely focused on the massive Great Rift Valley lakes. Our objective was to evaluate how recent climate change has altered the functioning and services of smaller tropical lakes, which are far more abundant on the landscape. Based on a paired analysis of 20 years of high-resolution water column data and a paleolimnological record from a small crater lake in western Uganda, we present evidence that even a modest warming of the air (∼0.9°C increase over 20 years) and changes in the timing and intensity of rainfall can have significant consequences on the dynamics of this common tropical lake type. For example, we observed a significant nonlinear increase (R(2) adj = 0.23, e.d.f. = 7, p<0.0001) in thermal stability over the past 20 years. This resulted in the expansion of anoxic waters and consequent deterioration of fish habitat and appears to have abated primary production; processes that may impair ecosystem services for a vulnerable human population. This study on a system representative of small tropical crater lakes highlights the far-reaching effects of global climatic change on tropical waters. Increased research efforts into tropical aquatic ecosystem health and the development of sound management practices are necessary in order to strengthen adaptive capabilities in tropical regions.
[Show abstract][Hide abstract] ABSTRACT: Gene flow among populations in different selective environments should favor the evolution of phenotypic plasticity over local adaptation. Plasticity in development is a common response to long-term hypoxia in some widespread African fishes, including Pseudocrenilabrus multicolor, a cichlid that exploits both normoxic (high oxygen) rivers/lakes and hypoxic (low oxygen) swamps. Previous studies have shown that fish from normoxic and hypoxic sites differ in many traits, including gill size, brain size and body shape, and that much of this variation reflects developmental plasticity. However, these earlier studies focused on areas in Uganda where gene flow between swamp and river or lake populations is high. In this study we tested the hypothesis that P. multicolor from a relatively isolated lake population (Lake Saka, Uganda) exhibit low levels of plasticity in traits related to oxygen uptake. Multiple broods of P. multicolor from Lake Saka were reared under low and high dissolved oxygen, and traits related to gill size, brain mass and body shape were quantified. Surprisingly, both gill size and brain mass showed high levels of developmental plasticity. We suggest that high levels of plasticity, particularly in the gill size of P. multicolor, reflects low costs of maintaining the plastic response, even in relatively isolated populations.
No preview · Article · Jan 2014 · Integrative Zoology
[Show abstract][Hide abstract] ABSTRACT: East Africa’s Lake Victoria provides resources and services to millions of people on the lake’s shores and abroad. In particular, the lake’s fisheries are an important source of protein, employment, and international economic connections for the whole region. Nonetheless, stock dynamics are poorly understood and currently unpredictable. Furthermore, fishery dynamics are intricately connected to other supporting services of the lake as well as to lakeshore societies and economies. Much research has been carried out piecemeal on different aspects of Lake Victoria’s system; e.g., societies, biodiversity, fisheries, and eutrophication. However, to disentangle drivers and dynamics of change in this complex system, we need to put these pieces together and analyze the system as a whole. We did so by first building a qualitative model of the lake’s social-ecological system. We then investigated the model system through a qualitative loop analysis, and finally examined effects of changes on the system state and structure. The model and its contextual analysis allowed us to investigate system-wide chain reactions resulting from disturbances. Importantly, we built a tool that can be used to analyze the cascading effects of management options and establish the requirements for their success. We found that high connectedness of the system at the exploitation level, through fisheries having multiple target stocks, can increase the stocks’ vulnerability to exploitation but reduce society’s vulnerability to variability in individual stocks. We describe how there are multiple pathways to any change in the system, which makes it difficult to identify the root cause of changes but also broadens the management toolkit. Also, we illustrate how nutrient enrichment is not a self-regulating process, and that explicit management is necessary to halt or reverse eutrophication. This model is simple and usable to assess system-wide effects of management policies, and can serve as a paving stone for future quantitative analyses of system dynamics at local scales.
[Show abstract][Hide abstract] ABSTRACT: The African cichlid, Pseudocrenilabrus multicolor victoriae is a eurytopic fish that exhibits high levels of developmental plasticity in response to dissolved oxygen availability. In this study, F1 offspring from three sites in the Mpanga River drainage of Western Uganda characterized by different dissolved oxygen (D.O.) regimes were reared under normoxic or hypoxic conditions. After 1year, enzymes were measured to determine the tissue metabolic capacity of four different tissues: muscle, heart, brain and liver. The enzymes measured were pyruvate kinase [PK], lactate dehydrogenase [LDH], citrate synthase [CS], and cytochrome C oxidase [CCO], and an additional two, malate dehydrogenase (MDH) and fructose 1,6-bisphosphatase (FBPase), were examined in the liver only. Individuals reared under hypoxia exhibited elevated levels of LDH and CCO in the heart; and depressed activity levels of brain CS and liver CCO and MDH relative to normoxia-reared sibs. Results from this study demonstrate that long-term exposure to hypoxia during development can induce changes in the metabolic capacities of P. multicolor. This flexibility may be important in facilitating persistence in variable and/or novel environments, and in the face of increasing global hypoxia.
Full-text · Article · Aug 2013 · Comparative biochemistry and physiology. Part B, Biochemistry & molecular biology
[Show abstract][Hide abstract] ABSTRACT: Population level response to hypoxia has become an issue of global significance because of increased frequency and intensity of hypoxic events worldwide, and the potential for global warming to exacerbate hypoxic stress. In this study, we sequenced two nuclear intronic regions and a single mitochondrial region across seven populations of the African cyprinid, Barbus neumayeri from two river drainages in Uganda: the Rwembaita Swamp-Njuguta River System and the Dura River. We then examined two indices of population structure, G ST and Jost's D, to detect links between oxygen availability and genetic variation and to determine if population genetic structure was associated with (i) dissolved oxygen regime (hypoxia or normoxia), (ii) geographical distance, or (iii) a combination of dissolved oxygen regime and geographical distance. Our results indicate that over a large scale (between drainages), geographical distance significantly affects the genetic structure of populations. However, within a single drainage, dissolved oxygen regime plays a key role in determining the genetic structure of populations. Within the Rwembaita-Njuguta system, gene flow was high between locations of similar oxygen regimes, but low between areas characterized by divergent oxygen regimes. Interestingly, G ST analyses appear to yield less realistic measures of population structure than Jost's D, suggesting that caution must be taken when interpreting and comparing the results from different studies. These results support the idea that aquatic dissolved oxygen can act as a selective force limiting gene flow among populations of aquatic species and therefore should be considered when implementing conservation plans and assessing environmental impact of human activities.
Full-text · Article · Jun 2013 · Ecology and Evolution
[Show abstract][Hide abstract] ABSTRACT: Aquatic hypoxia is generally viewed as stressful for aerobic organisms. However, hypoxia may also benefit organisms by decreasing cellular stress, particularly that related to free radicals. Thus, an ideal habitat may have the minimum O2 necessary to both sustain aerobic metabolism and reduce the need to scavenge free radicals and repair free radical damage. The ability of aquatic organisms to sustain aerobic metabolism relates in part to the ability to maximize gas diffusion, which can be facilitated by small body size when O2 uptake occurs across the body surface, by a large gill surface area, or by the ability to use atmospheric air. We use water-breathing organisms in chronically hypoxic papyrus (Cyperus papyrus) swamps of East Africa to test the hypothesis that cellular-level benefits of hypoxia may translate into increased fitness, especially for small organisms. A review of recent studies of fingernail clams (Sphaerium sp.) shows that clams living in sustained hypoxia have minimized oxidative stress and that these cellular-level benefits may lead to increased fitness. We suggest that organisms in the extreme conditions in the papyrus swamps provide a unique opportunity to challenge the conventional classification of hypoxic habitats as 'stressful' and normoxic habitats as 'optimal.'
No preview · Article · Apr 2013 · Comparative biochemistry and physiology. Part A, Molecular & integrative physiology
[Show abstract][Hide abstract] ABSTRACT: Parallel adaptive radiation events provide a powerful framework for investigations of ecology's contribution to phenotypic diversification. Ecologically driven divergence has been invoked to explain the repeated evolution of sympatric dwarf and normal lake whitefish (Coregonus clupeaformis) species in multiple lakes in eastern North America. Nevertheless, links between most putatively adaptive traits and ecological variation remain poorly defined within and among whitefish species pairs. Here, we examine four species pairs for variation in gill, heart, and brain size; three traits predicted to show strong phenotypic responses to ecological divergence. In each of the species pairs, normals exhibited larger body size standardized gills compared to dwarfs - a pattern that is suggestive of a common ecological driver of gill size divergence. Within lakes, the seasonal hypoxia experienced in the benthic environment is a likely factor leading to the requirement for larger gills in normals. Interestingly, the morphological pathways used to achieve larger gills varied between species pairs from Québec and Maine, which may imply subtle non-parallelism in gill size divergence related to differences in genetic background. There was also a non-significant trend toward larger hearts in dwarfs, the more active species of the two, whereas brain size varied exclusively among the lake populations. Taken together, our results suggest that the diversification of whitefish has been driven by parallel and non-parallel ecological conditions across lakes. Furthermore, the phenotypic response to ecological variation may depend on genetic background of each population.
Full-text · Article · Mar 2013 · Ecology and Evolution