Distribution and faunal richness of Cladocera in western Uganda crater lakes

In this study, we analyse the distribution and species richness of epibenthic and planktonic Cladocera (Crustacea: Branchiopoda) in 62 Uganda crater lakes, spread across the climatic gradient between the sub-humid shoulder and semi-arid floor of the East African Rift Valley. Together, these lakes cover large environmental gradients in salinity, trophic conditions and depth. In total, 36 species of Cladocera were encountered in the freshwater lakes (<1,500 μS/cm), whereas only a single species was found in the true saline lakes (>10,000 μS/cm). Cladoceran species richness in individual lakes was found to be determined primarily by the presence of a well-developed littoral belt of submerged and emergent aquatic macrophytes, pH and salinity. The highest species richness occurred in fresh but eutrophic shallow waters, with relatively low pH (6.5–7) and dense aquatic macrophyte growth. As identified by multivariate statistical analysis, the distribution of Cladocera species among the Uganda lakes was most strongly determined by nutrient availability (measured as total phosphorus), the presence and diversity of aquatic macrophyte habitat, pH, mean annual temperature and the fraction of the crater catchment that is currently under agriculture. Since Cladocera play an important role in aquatic food webs, and as such contribute to the ecological integrity of aquatic ecosystems, an increased understanding of the environmental controls underlying their distribution provides valuable information on aquatic ecosystem functioning needed for management and conservation. The significant turnover of cladoceran species composition along the sampled environmental gradients demonstrates their potential as biological indicators for water quality and ecosystem health in East African lakes. Our results suggest that changes in land use are the greatest threat to natural ecosystem functioning in these African lakes, and particularly so in the shallower lakes.     

Early Human Speciation,Brain Expansion and Dispersal Influenced by African Climate Pulses

Early human evolution is characterised by pulsed speciation and dispersal events that cannot be explained fully by global or continental paleoclimate records. We propose that the collated record of ephemeral East African Rift System (EARS) lakes could be a proxy for the regional paleoclimate conditions experienced by early hominins. Here we show that the presence of these lakes is associated with low levels of dust deposition in both West African and Mediterranean records, but is not associated with long-term global cooling and aridification of East Africa. Hominin expansion and diversification seem to be associated with climate pulses characterized by the precession-forced appearance and disappearance of deep EARS lakes. The most profound period for hominin evolution occurs at about 1.9 Ma; with the highest recorded diversity of hominin species, the appearance of Homo (sensu stricto) and major dispersal events out of East Africa into Eurasia. During this period, ephemeral deep-freshwater lakes appeared along the whole length of the EARS, fundamentally changing the local environment. The relationship between the local environment and hominin brain expansion is less clear. The major step-wise expansion in brain size around 1.9 Ma when Homo appeared was coeval with the occurrence of ephemeral deep lakes. Subsequent incremental increases in brain size are associated with dry periods with few if any lakes. Plio-Pleistocene East African climate pulses as evinced by the paleo-lake records seem, therefore, fundamental to hominin speciation, encephalisation and migration.     

The Great Lakes in East Africa: biological conservation considerations for species flocks

The three largest water bodies of East Africa, Lake Victoria, Tanganyika, and Malawi contain an estimated number of 2,000 endemic cichlid fish species, in addition, to a mostly uncounted wealth of invertebrates. While the terrestrial diversity is reasonably well protected, as economic and touristic interests coincide with biological conservation strategies, this is not the case for most African lakes and rivers. Nonetheless, it must be promoted that these aquatic ecosystems also deserve protection. Conservation strategies for aquatic biota have so far been the same as for terrestrial environments, i.e., by declaring biodiversity hotspots national parks. Such parks also contain rivers and lake shores. Here, I argue that it seems questionable that this strategy will work, given strong micro-geographic structure of the species flocks and the great degree of local endemism. I suggest a novel strategy for protecting African Lake communities that accounts for local endemism, derived from recent molecular phylogenetic and phylogeographic studies on East African cichlid fishes. While connectivity is the major problem for terrestrial and marine national parks, to ensure a large enough effective population size of the protected animals, this is not the case in most taxa of African rivers and lakes, where local endemism prevails. For example, most littoral cichlid species are subdivided into numerous distinct “color morphs” with restricted distribution, and unlike marine fishes with planktonic larvae display brood care with small offspring numbers. It is argued that the establishment of “micro-scale protected areas,” a large number of small stretches of strictly protected coast line, each only some hundreds of meters long, is likely to work best to preserve the littoral communities in African lakes. Such protected zones can sustain a reasonably effective population size of littoral species, serve as protected spawning ground or nursery area for pelagic species, and at the same time re-seed neighboring populations that are exploited continuously. As long-term stability of littoral fishing grounds is in the immediate interest of village communities, such small protected areas should be managed and controlled by the local communities themselves, and supervised by governmental institutions.     

Phylogeography and speciation in the <Emphasis Type="Italic">Pseudocrenilabrus philander</Emphasis> species complex in Zambian Rivers

Haplochromine cichlids form the most species-rich lineage of cichlid fishes that both colonized almost all river systems in Africa and radiated to species flocks in several East African lakes. The enormous diversity of lakes is contrasted by a relatively poor albeit biogeographically clearly structured species diversity in rivers. The present study analyzed the genetic structure and phylogeographic history of species and populations of the genus Pseudocrenilabrus in Zambian rivers that span two major African drainage systems, the Congo- and the Zambezi-system. The mtDNA phylogeny identifies four major lineages, three of which occur in the Congo-system and one in the Zambezi system. Two of the Congo-clades (Lake Mweru and Lunzua River) comprise distinct albeit yet undescribed species, while the fish of the third Congo-drainage clade (Chambeshi River and Bangweulu swamps), together with the fish of the Zambezi clade (Zambezi and Kafue River) are assigned to Pseudocrenilabrus philander. Concerning the intraspecific genetic diversity observed in the sampled rivers, most populations are highly uniform in comparison to lacustrine haplochromines, suggesting severe founder effects and/or bottlenecking during their history. Two bursts of diversification are reflected in the structure of the linearized tree. The first locates at about 3.9% mean sequence divergence and points to an almost simultaneous colonization of the sampled river systems. Subsequent regional diversification (with about 1% mean sequence divergence) occurred contemporaneously within the Kafue River and the Zambezi River. The clear-cut genetic biogeographic structure points to the dominance of geographic speciation in this lineage of riverine cichlid fishes, contrasting the importance of in situ diversification observed in lake cichlids.     

The bounty of minor lakes: the role of small satellite water bodies in evolution and conservation of fishes in the Lake Victoria Region,East Africa

The Lake Victoria Region (LVR) encompasses the large lakes Victoria, Kyoga, Edward, George and Kivu, as well as scores of small satellite lakes within the parent catchments. Taken as a whole, the LVR originally harbored a unique fish fauna that included in excess of 600 endemic species of cichlid fishes. As a result of human influence, including a commercial fishery and the introduction of several exotic species, nearly 200 cichlid species and several endemic genera have become extinct from lakes Victoria and Kyoga. Recently, we have discovered that some of the apparently extinct taxa survive as extant representatives in the satellite lakes. Here, we summarize the findings of our ecological survey of the fish species of the satellite lakes. We also discuss the results of some preliminary genetic analyses, and highlight major genetic and ecological changes in the fish fauna that have taken place in the regional fishery. Minor lakes now play a crucial role in conserving the endangered species of the entire region, and also as living museums of East African ecological history. Our findings allude to the historical importance of minor satellite lakes as natural refugia for the fishes of the Lake Victoria Region, a region characterized by a history of geological and climatic instability.     

Explosive speciation in cichlid fishes of the African Great Lakes: a dynamic model of adaptive radiation

In the African cichlid species flocks several corresponding ecotypes and species communities have evolved independently in each of the lakes. These flocks can be viewed as reiterations of a process, induced by the same type of external event (the formation of a lake) and seeded by the same group of organisms, equipped with the same innovative potential. The East African lakes differ in their ages, so that different stages of adaptive radiation of a single group of fishes can be studied simultaneously. This review combines findings on various African cichlid species flocks and derives a generalized model of cichlid adaptive radiation. The model constitutes four stages, defined by the most influential habitat characteristics and interactions among members of the species community at each phase. It is an attempt to provide an explicit hypothesis for a dynamic evolutionary process, as a basis to future studies.     

Taxonomy and diversity of Afroalpine Chironomidae (Insecta: Diptera) on Mount Kenya and the Rwenzori Mountains, East Africa

Aim Anthropogenic climate change is expected to result in the complete loss of glaciers from the high mountains of tropical Africa, with profound impacts on the hydrology and ecology of unique tropical cold‐water lakes located downstream from them. This study examines the biodiversity of Chironomidae (Insecta: Diptera) communities in these scarce Afroalpine lake systems, in order to determine their uniqueness in relation to lowland African lakes and alpine lakes in temperate regions, and to evaluate the potential of Afroalpine Chironomidae as biological indicators to monitor future changes in the ecological integrity of their habitat. Location Mount Kenya (Kenya) and Rwenzori Mountains (Uganda). Methods The species composition of Afroalpine chironomid communities was assessed using recent larval death assemblages extracted from the surface sediments of 11 high‐mountain lakes between 2900 and 4575 m. Results were compared with similar faunal data from 68 East African lakes at low and middle elevation (750–2760 m), and with literature records of Chironomidae species distribution in sub‐Saharan Africa, the Palaearctic region and elsewhere. All recovered taxa were fully described and illustrated. Results The 11‐lake analysis yielded 1744 subfossil chironomid larvae belonging to 16 distinct taxa of full‐grown larvae, and three taxa of less differentiated juveniles. Eleven of these 16 are not known to occur in African lakes at lower elevation, and eight taxa (or 50% of total species richness) appear restricted to the specific habitat of cold lakes above 3900 m, where night‐time freezing is frequent year‐round. The faunal transition zone coincides broadly with the Ericaceous zone of terrestrial vegetation (c. 3000–4000 m). Snowline depression during the Quaternary ice ages must have facilitated dispersion of cold‐stenothermous species among the high mountains of equatorial East Africa, but less so from or to the Palaearctic region via the Ethiopian highlands. Main conclusions Chironomid communities in glacier‐fed lakes on Africa's highest mountains are highly distinct from those of lowland African lakes, and potentially unique on a continental scale. By virtue of excellent preservation and their spatial and temporal integration of local community dynamics, chironomid larval death assemblages extracted from surface sediments are powerful biological indicators for monitoring the hydrological and ecological changes associated with the current retreat and loss of Africa's glaciers.     

Phylogeny of the Lake Tanganyika cichlid species flock and its relationship to the Central and East African haplochromine cichlid fish faunas

Lake Tanganyika, the oldest of the East African Great Lakes, harbors the ecologically, morphologically, and behaviorally most complex of all assemblages of cichlid fishes, consisting of about 200 described species. The evolutionary old age of the cichlid assemblage, its extreme degree of morphological differentiation, the lack of species with intermediate morphologies, and the rapidity of lineage formation have made evolutionary reconstruction difficult. The number and origin of seeding lineages, particularly the possible contribution of riverine haplochromine cichlids to endemic lacustrine lineages, remains unclear. Our phylogenetic analyses, based on mitochondrial DNA sequences of three gene segments of 49 species (25% of all described species, up to 2,400 bp each), yield robust phylogenies that provide new insights into the Lake Tanganyika adaptive radiation as well as into the origin of the Central- and East-African haplochromine faunas. Our data suggest that eight ancient African lineages may have seeded the Tanganyikan cichlid radiation. One of these seeding lineages, probably comprising substrate spawning Lamprologus-like species, diversified into six lineages that evolved mouthbrooding during the initial stage of the radiation. All analyzed haplochromines from surrounding rivers and lakes seem to have evolved within the radiating Tanganyikan lineages. Thus, our findings contradict the current hypothesis that ancestral riverine haplochromines colonized Lake Tanganyika to give rise to at least part of its spectacular endemic cichlid species assemblage. Instead, the early phases of the Tanganyikan radiation affected Central and East African rivers and lakes. The haplochromines may have evolved in the Tanganyikan basin before the lake became a hydrologically and ecologically closed system and then secondarily colonized surrounding rivers. Apparently, therefore, the current diversity of Central and East African haplochromines represents a relatively young and polyphyletic fauna that evolved from or in parallel to lineages now endemic to Lake Tanganyika.     

Lake level fluctuations synchronize genetic divergences of cichlid fishes in African lakes

Water level fluctuations are important modulators of speciation processes in tropical lakes, in that they temporarily form or break down barriers to gene flow among adjacent populations and/or incipient species. Time estimates of the most recent major lowstands of the three African Great Lakes are thus crucial to infer the relative timescales of explosive speciation events in cichlid species flocks. Our approach combines geological evidence with genetic divergence data of cichlid fishes from the three Great East African Lakes derived from the fastest-evolving mtDNA segment. Thereby, we show for each of the three lakes that individuals sampled from several populations which are currently isolated by long geographic distances and/or deep water form clusters of equally closely related haplotypes. The distribution of identical or equally closely related haplotypes in a lake basin allows delineation of the extent of lake level fluctuations. Our data suggest that the same climatic phenomenon synchronized the onset of genetic divergence of lineages in all three species flocks, such that their most recent evolutionary history seems to be linked to the same external modulators of adaptive radiation. A calibration of the molecular clock of the control region was elaborated by gauging the age of the Lake Malawi species flock through the divergence among the utaka-cichlid and the mbuna-cichlid lineages to minimally 570,000 years and maximally 1 Myr. This suggests that the low-lake-level period which established the observed patterns of genetic relatedness dates back less than 57,000 years, probably even to 17,000-12,400 years ago, when Lake Victoria dried up and Lakes Malawi and Tanganyika were also low. A rapid rise of all three lakes about 11,000 years ago established the large-scale population subdivisions observed today. Over that period of time, a multitude of species originated in Lakes Malawi and Victoria with an impressive degree of morphological and ecological differentiation, whereas the Tanganyikan taxa that were exposed to the same habitat changes hardly diverged ecologically and morphologically. Our findings also show that patterns of genetic divergences of stenotopic organisms provide valuable feedback on geological and sedimentological time estimates for lake level changes.     

Molecular Characterization of Two Endothelin Pathways in East African Cichlid Fishes

The adaptive radiations of cichlid fishes in East Africa have been associated with the acquisition of evolutionary novelties as well as the ecological opportunities existing in the East African Great lakes. Two remarkable evolutionary innovations are the pharyngeal jaw apparatus, found in all cichlid species, and the anal fin egg-spots of mouthbrooding cichlids. Based on their conserved functions during the development of both the jaw apparatus and pigmentation, the endothelin ligands and receptors form a putative link between these naturally and sexually selected traits. Here we study the evolutionary history of four members of two endothelin pathways (Edn1/EdnrAa and Edn3b/EdnrB1a) to elucidate their possible roles during the evolution and development of key innovations in East African cichlids species. The analyses performed on partial sequences (ca. 6,000 bp per taxon) show that all four endothelin family members evolved under purifying selection, although both ligands are characterized by an accelerated rate of protein evolution in comparison to the receptors. In accordance with earlier findings, we show that the mature protein sequence of Edn1 and Edn3 are highly conserved, also in cichlids, whereas the preproendothelin parts are variable indicating relaxed selective constraints. In the receptors, nonsynonymous substitutions were mainly found in the ligand-binding domains suggesting functional divergence. Gene expression assays with Real-Time PCR indeed reveal that the two studied endothelin pathways are expressed in the cichlid pharyngeal jaw and in the haplochromine egg-spot (among other pigment-cell containing tissues), suggesting their involvement during morphogenesis of naturally and sexually selected traits in cichlids.     

The oldest fossil cichlids (Teleostei: Perciformes): indication of a 45 million-year-old species flock

Five closely related species of fossil cichlids collected from an Eocene site in Tanzania, East Africa, represent the oldest known cichlids. The specimens are whole-body, articulated fishes that are extremely well preserved and, therefore, have the potential to add to our knowledge of the history of this family. Modern cichlids are particularly well known for the numerous species flocks of the East African Great Lakes. A great deal of research is ongoing regarding all aspects of the fishes in these flocks, including their evolutionary history The new collection of fossils reported here is interpreted as representing a species flock that arose in a small crater lake. These fossils indicate that cichlids' ability to form species flocks evolved early in the history of this family.     

Oligochaeta and Aphanoneura in ancient lakes: a review

By their antiquity, history, rarity, great depth in many instances and the presence of highly diverse faunas with many endemics, ancient lakes constitute ecosystems of a special nature, clearly apart from the large majority of extant lakes. While the fauna of these lakes is becoming better and better known for various animals groups, the Oligochaeta are still poorly known. Tubificidae and Naididae are found in each ancient lake. On the other hand, some families are restricted to only one lake, such as Aeolosomatidae and Proppapidae in Lake Baikal or Eudrilidae and Ocnerodrilidae (megadriles) in Lake Tanganyika, but such a distribution is probably due to a lack of knowledge or sampling biases. All ancient lakes have an endemic oligochaete fauna except Lake Kinneret (Israël). The oldest, Lake Baikal (20–25 Ma), holds the most abundant and diverse oligochaete fauna, in which species flocks are even recognizable or suspected. In contrast, the oligochaete fauna of the slightly younger Lake Tanganyika is very scarce. This is partly due to an obvious lack of studies, as the oligochaete fauna of other great African lakes is virtually unknown, but this might be the result of an environment in these lakes less favourable to oligochaetes. Some factors likely to interact with speciation in oligochaetes are discussed but nothing can be concluded to date. A recent interest in African great lakes revealed a more diverse oligochaete fauna than previously assumed but a better study of this fauna is still badly needed.     

Tectonics, orbital forcing, global climate change, and human evolution in Africa: introduction to the African paleoclimate special volume

The late Cenozoic climate of Africa is a critical component for understanding human evolution. African climate is controlled by major tectonic changes, global climate transitions, and local variations in orbital forcing. We introduce the special African Paleoclimate Issue of the Journal of Human Evolution by providing a background for and synthesis of the latest work relating to the environmental context for human evolution. Records presented in this special issue suggest that the regional tectonics, appearance of C(4) plants in East Africa, and late Cenozoic global cooling combined to produce a long-term drying trend in East Africa. Of particular importance is the uplift associated with the East African Rift Valley formation, which altered wind flow patterns from a more zonal to more meridinal direction. Results in this volume suggest a marked difference in the climate history of southern and eastern Africa, though both are clearly influenced by the major global climate thresholds crossed in the last 3 million years. Papers in this volume present lake, speleothem, and marine paleoclimate records showing that the East African long-term drying trend is punctuated by episodes of short, alternating periods of extreme wetness and aridity. These periods of extreme climate variability are characterized by the precession-forced appearance and disappearance of large, deep lakes in the East African Rift Valley and paralleled by low and high wind-driven dust loads reaching the adjacent ocean basins. Dating of these records show that over the last 3 million years such periods only occur at the times of major global climatic transitions, such as the intensification of Northern Hemisphere Glaciation (2.7-2.5 Ma), intensification of the Walker Circulation (1.9-1.7 Ma), and the Mid-Pleistocene Revolution (1-0.7 Ma). Authors in this volume suggest this onset occurs as high latitude forcing in both Hemispheres compresses the Intertropical Convergence Zone so that East Africa becomes locally sensitive to precessional forcing, resulting in rapid shifts from wet to dry conditions. These periods of extreme climate variability may have provided a catalyst for evolutionary change and driven key speciation and dispersal events amongst mammals and hominins in Africa. In particular, hominin species seem to differentially originate and go extinct during periods of extreme climate variability. Results presented in this volume may represent the basis of a new theory of early human evolution in Africa.     

Populations genetically rifting within a complex geological system: The case of strong structure and low genetic diversity in the migratory freshwater catfish,Bagrus docmak,in East Africa

The complex geological history of East Africa has been a driving factor in the rapid evolution of teleost biodiversity. While there is some understanding of how macroevolutionary drivers have shaped teleost speciation in East Africa, there is a paucity of research into how the same biogeographical factors have affected microevolutionary processes within lakes and rivers. To address this deficiency, population genetic diversity, demography, and structure were investigated in a widely distributed and migratory (potamodromous) African teleost species, Ssemutundu (Bagrus docmak ). Samples were acquired from five geographical locations in East Africa within two major drainage basins; the Albertine Rift and Lake Victoria Basin. Individuals (N  = 175) were genotyped at 12 microsatellite loci and 93 individuals sequenced at the mitochondrial DNA control region. Results suggested populations from Lakes Edward and Victoria had undergone a severe historic bottleneck resulting in very low nucleotide diversity (π = 0.004 and 0.006, respectively) and negatively significant Fu values (?3.769 and ?5.049; p  < .05). Heterozygosity deficiencies and restricted effective population size (N _eLD) suggested contemporary exposure of these populations to stress, consistent with reports of the species decline in the East African Region. High genetic structuring between drainages was detected at both historical (?_ST = 0.62 for mtDNA; p  < .001) and contemporary (microsatellite F _ST = 0.460; p  < .001) levels. Patterns of low genetic diversity and strong population structure revealed are consistent with speciation patterns that have been linked to the complex biogeography of East Africa, suggesting that these biogeographical features have operated as both macro‐ and micro‐evolutionary forces in the formation of the East African teleost fauna.     

Time and origin of cichlid colonization of the lower Congo rapids

Most freshwater diversity is arguably located in networks of rivers and streams, but, in contrast to lacustrine systems riverine radiations, are largely understudied. The extensive rapids of the lower Congo River is one of the few river stretches inhabited by a locally endemic cichlid species flock as well as several species pairs, for which we provide evidence that they have radiated in situ. We use more that 2,000 AFLP markers as well as multilocus sequence datasets to reconstruct their origin, phylogenetic history, as well as the timing of colonization and speciation of two Lower Congo cichlid genera, Steatocranus and Nanochromis. Based on a representative taxon sampling and well resolved phylogenetic hypotheses we demonstrate that a high level of riverine diversity originated in the lower Congo within about 5 mya, which is concordant with age estimates for the hydrological origin of the modern lower Congo River. A spatial genetic structure is present in all widely distributed lineages corresponding to a trisection of the lower Congo River into major biogeographic areas, each with locally endemic species assemblages. With the present study, we provide a phylogenetic framework for a complex system that may serve as a link between African riverine cichlid diversity and the megadiverse cichlid radiations of the East African lakes. Beyond this we give for the first time a biologically estimated age for the origin of the lower Congo River rapids, one of the most extreme freshwater habitats on earth.     

Repeated trans-watershed hybridization among haplochromine cichlids (Cichlidae) was triggered by Neogene landscape evolution

The megadiverse haplochromine cichlid radiations of the East African lakes, famous examples of explosive speciation and adaptive radiation, are according to recent studies, introgressed by different riverine lineages. This study is based on the first comprehensive mitochondrial and nuclear DNA dataset from extensive sampling of riverine haplochromine cichlids. It includes species from the lower River Congo and Angolan (River Kwanza) drainages. Reconstruction of phylogenetic hypotheses revealed the paradox of clearly discordant phylogenetic signals. Closely related mtDNA haplotypes are distributed thousands of kilometres apart and across major African watersheds, whereas some neighbouring species carry drastically divergent mtDNA haplotypes. At shallow and deep phylogenetic layers, strong signals of hybridization are attributed to the complex Late Miocene/Early Pliocene palaeohistory of African rivers. Hybridization of multiple lineages across changing watersheds shaped each of the major haplochromine radiations in lakes Tanganyika, Victoria, Malawi and the Kalahari Palaeolakes, as well as a miniature species flock in the Congo basin (River Fwa). On the basis of our results, introgression occurred not only on a spatially restricted scale, but massively over almost the whole range of the haplochromine distribution. This provides an alternative view on the origin and exceptional high diversity of this enigmatic vertebrate group.     

Punctuated equilibria, morphological stasis and the palaeontological documentation of speciation: a biological appraisal of a case history in an African lake

The present-day faunas of the great African lakes present some of the world's best examples of 'explosive speciation'. Lakes Victoria and Malawi each probably have several hundred endemic species of cichlid fishes. Much can be inferred about the evolution of these fishes from morphology, behaviour and intra-lacustrine distribution and from the fact that they include taxa ranging from local races, through sibling species, to forms that display extensive differentiation. The time taken to acquire specific distinctness can sometimes be accurately defined, but fossil lineages are unknown. A recent study of a fossil sequence of molluscs in the Turkana basin throws new light on the history of African lake faunas. It also claims to have resolved events during speciation. While critical analysis based on our knowledge of living molluscs in this area fails to substantiate this claim, the fossil molluscs complement information provided by the biology of extant fishes and invertebrates and emphasize the importance of these lakes in the study of evolution in living and extinct populations.     

Does the decline of gastropods in deep water herald ecosystem change in Lakes Malawi and Tanganyika?

1. Ancient, deep lakes have traditionally been considered as stable, ecological islands, well buffered from environmental change because of their great depth. However, they are not immune to anthropogenic and climatic stress. Ecosystems of the permanently stratified warm Lakes Malawi and Tanganyika in the Great East African Rift are particularly delicate. Their stratification regime has historically limited the distribution of benthic biota to a ‘bathtub ring of biodiversity’, namely substrata in the upper, oxygenated water layer. 2. We use historical data on the endemic deep‐water molluscs of these lakes to assess present‐day stress on their benthic ecosystems. During the 20th century, these molluscs have probably decreased in abundance and migrated to shallower water. 3. These apparent trends have a significance beyond species‐based conservation, foremost because deep‐water organisms heavily rely on the position and temporal stability of the oxycline and therefore provide an early warning of large‐scale changes in the distribution of dissolved oxygen. Oxygen demands have increased in the East African Great Lakes over the last century whereas ventilation of deep water has remained the same or declined. 4. The combination of these factors is resulting in a narrowing of the ring of biodiversity and a changed nutrient flux through this ring. Reduction in the habitat available inevitably puts biota at risk, whereas changes in nutrient flux may cause shifts in the entire ecosystem or the collapse of parts of it. 5. Considering the socioeconomic value of these lakes and the potentially grave implications for their faunal biodiversity and entire ecosystems, existing evidence of faunal decline, especially in taxa that depend strongly on the stratification regime, is of great concern. Moreover, because the factors responsible are widespread and include surface‐water warming, increased run‐off and eutrophication, respiration stress may also develop in other tropical and subtropical lakes.     

The role of physiology in the divergence of two incipient cichlid species

Sexual selection on male coloration has been implicated in the evolution of colourful species flocks of East African cichlid fish. During adaptive radiations, animals diverge in multiple phenotypic traits, but the role of physiology has received limited attention. Here, we report how divergence in physiology may contribute to the stable coexistence of two hybridizing incipient species of cichlid fish from Lake Victoria. Males of Pundamilia nyererei (males are red) tend to defeat those of Pundamilia pundamilia (males are blue), yet the two sibling species coexist in nature. It has been suggested that red males bear a physiological cost that might offset their dominance advantage. We tested the hypothesis that the two species differ in oxidative stress levels and immune function and that this difference is correlated with differences in circulating steroid levels. We manipulated the social context and found red males experienced significantly higher oxidative stress levels than blue males, but only in a territorial context when colour and aggression are maximally expressed. Red males exhibited greater aggression levels and lower humoral immune response than blue males, but no detectable difference in steroid levels. Red males appear to trade off increased aggressiveness with physiological costs, contributing to the coexistence of the two species. Correlated divergence in colour, behaviour and physiology might be widespread in the dramatically diverse cichlid radiations in East African lakes and may play a crucial role in the remarkably rapid speciation of these fish.