African Lakes Research Articles

Ancyrocephalidae (Monogenea) of Lake Tanganyika: III: Cichlidogyrus infecting the world’s biggest cichlid and the non-endemic tribes Haplochromini, Oreochromini and Tylochromini (Teleostei, Cichlidae)

Lake Tanganyika is the deepest and oldest African Great Lake and of economic importance. While the diversity of its endemic cichlid radiations yielded scientific interest, a number of cichlid tribes have few representatives in the lake. Some of those, namely Oreochromini (ex-Tilapiini), Haplochromini and Tylochromini, reach higher species numbers in riverine systems. Conversely, the phylogenetic position of the monospecific and endemic Boulengerochromini is unclear. The oreochromines Oreochromis tanganicae and Oreochromis niloticus, the haplochromine Astatotilapia burtoni, the tylochromine Tylochromis polylepis and the boulengerochromine Boulengerochromis microlepis, the largest cichlid species worldwide, were surveyed for ancyrocephalid monogenean gill parasites. Five new species are proposed. Cichlidogyrus gillardinae sp. n. is described from A. burtoni, Cichlidogyrus mbirizei sp. n. from O. tanganicae and Cichlidogyrus nshomboi sp. n. from B. microlepis. T. polylepis harbours Cichlidogyrus mulimbwai sp. n., Cichlidogyrus muzumanii sp. n. and a third, presently undescribed species. Four species known from outside the Tanganyika Basin were retrieved on the oreochromines. The host species are scientific models or important in the sectors of fisheries or ornamental fish trade. Moreover, their phylogenetic positions render them well-suited to help elucidate the historic relationships between riverine and lacustrine African cichlids. In this framework, their Cichlidogyrus fauna is compared to congeners known from African rivers and to the few Tanganyika representatives described. While the parasites of Oreochromis, A. burtoni and T. polylepis are reminiscent of those infecting related hosts throughout Africa, B. microlepis hosts a Cichlidogyrus morphotype typical of Lake Tanganyika. This supports its placement within an endemic cichlid radiation.

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.

Carbon cycling within an East African lake revealed by the carbon isotope composition of diatom silica: a 25-ka record from Lake Challa, Mt. Kilimanjaro

The carbon cycle of a lake is a balance between supply from the atmosphere and catchment, and the net demand exerted by primary producers, minus losses back to the atmosphere and to sediment storage. Evaluating the sum of these processes and reconstructing them from sediment records of lake history requires a range of methods and a multi-proxy approach. One promising technique is to explore the carbon-isotope composition (δ13Cdiatom) of organic matter incorporated within the silica frustules of diatom algae. Here we present a 25,000-year record of δ13Cdiatom from the sediments of crater Lake Challa on the eastern flank of Mt. Kilimanjaro, and along with other proxy data we make inferences about the three major phases in the history of the lake's carbon cycle. From 25 ka to 15.8 ka years BP, δ13Cdiatom is positively correlated with the δ13C of bulk sediment organic matter (δ13Cbulk), indicating that high diatom productivity, as recorded by high % biogenic silica at this time, was preferentially removing 12C and enriching the δ13C of lake-water dissolved inorganic carbon. From 15.8 to 5.5 ka the correlation between δ13Cdiatom and δ13Cbulk breaks down, suggesting carbon supply to the lake satisfied or exceeded the demand from productivity. From 5.5 ka BP the positive correlation resumes, indicating an increase in the internal demand for carbon relative to external supply. Diatom frustule-bound carbon isotopes offer an original tool in examining long-term fluctuations in a lake's carbon budget and how the balance between supply and demand has changed through time.

Distribution of tetraether lipids in the 25-ka sedimentary record of Lake Challa: extracting reliable TEX86 and MBT/CBT palaeotemperatures from an equatorial African lake

The distribution of isoprenoid and branched glycerol dialkyl glycerol tetraether (GDGT) lipids was studied in the sedimentary record of Lake Challa, a permanently stratified, partly anoxic crater lake on the southeastern slope of Mt. Kilimanjaro (Kenya/Tanzania), to examine if the GDGTs could be used to reconstruct past variation in regional temperature. The study material comprised 230 samples from a continuous sediment sequence spanning the last 25 ka with excellent age control based on high-resolution AMS 14C dating. The distribution of GDGTs showed large variation through time. In some time intervals (i.e., from 20.4 to 15.9 ka BP and during the Younger Dryas, 12.9–11.7 ka BP) crenarchaeol was the most abundant GDGT, whereas at other times (i.e., during the Early Holocene) branched GDGTs and GDGT-0 were the major GDGT constituents. In some intervals of the sequence the relative abundance of GDGT-0 and GDGT-2 was too high to be derived exclusively from lacustrine Thaumarchaeota, suggesting a sizable contribution from methanogens and other archaea. This severely complicated application of TEX86 palaeothermometry in this lake, and limited reliable reconstruction of lake water temperature to the time interval 25–13 ka BP, i.e. the Last Glacial Maximum and the period of post-glacial warming. The TEX86-inferred timing of this warming is similar to that recorded previously in two of the large African rift lakes, while its magnitude is slightly or much higher than that recorded at these other sites, depending on which lake-based TEX86 calibration is used. Application of calibration models based on distributions of branched GDGTs developed for lakes inferred temperatures of 15–18 °C for the Last Glacial Maximum and 19–22 °C for the Holocene. However, the MBT/CBT palaeothermometer reconstructs temperatures as low as 12 °C for a Lateglacial period centred on 15 ka BP. Variation in down-core values of the BIT index are mainly determined by the varying production rate of crenarchaeol relative to in-situ produced branched GDGTs. The apparent relationship of the BIT index with climatic moisture balance can be explained either by the direct influence of lake level and wind strength on nutrient recycling, or by influx of soil nutrients promoting aquatic productivity and nitrification. This study shows that GDGTs can aid in obtaining climatic information from lake records but that the obtained data should be interpreted with care.

A novel primer set for multilocus phylogenetic inference in East African cichlid fishes

The cichlid fishes in the East African Great Lakes are a prime model system for the study of adaptive radiation. Therefore, the availability of an elaborate phylogenetic framework is an important prerequisite. Previous phylogenetic hypotheses on East African cichlids are mainly based on mitochondrial and/or fragment-based markers, and, to date, no taxon-rich phylogeny exists that is based on multilocus DNA sequence data. Here, we present the design of an extensive new primer set (24 nuclear makers) for East African cichlids that will be used for multilocus phylogenetic analyses in the future. The primers are designed to work for both Sanger sequencing and next-generation sequencing with the 454 technology. As a proof of principle, we validate these primers in a phylogenetically representative set of 16 cichlid species from Lake Tanganyika and main river systems in the area and provide a basic evaluation of the markers with respect to marker length and diversity indices.

The impact of the geologic history and paleoclimate on the diversification of East african cichlids.

The cichlid fishes of the East African Great Lakes are the largest extant vertebrate radiation identified to date. These lakes and their surrounding waters support over 2,000 species of cichlid fish, many of which are descended from a single common ancestor within the past 10 Ma. The extraordinary East African cichlid diversity is intricately linked to the highly variable geologic and paleoclimatic history of this region. Greater than 10 Ma, the western arm of the East African rift system began to separate, thereby creating a series of rift basins that would come to contain several water bodies, including the extremely deep Lakes Tanganyika and Malawi. Uplifting associated with this rifting backponded many rivers and created the extremely large, but shallow Lake Victoria. Since their creation, the size, shape, and existence of these lakes have changed dramatically which has, in turn, significantly influenced the evolutionary history of the lakes' cichlids. This paper reviews the geologic history and paleoclimate of the East African Great Lakes and the impact of these forces on the region's endemic cichlid flocks.

Heavy Metals Inflow into Lake Muhazi, Rwanda

Most tropical African lakes are facing pollution problems due to the rapid population growth and industrializa- tion in their riparian communities. Lake Muhazi in Rwanda is one of such lakes which has experienced a dramatic decrease in fish production since the eighties, with also reports of low water transparencies and high turbidities. The lakeshores are now being developed for ecotourism and this requires sound environmental management to make the planned activities viable. The purpose of this study was to assess heavy metal pollution in the tributary rivers of Lake Muhazi. The concentration of heavy metals in major tributaries was monitored for the period July to October 2007. The parameters studied are cadmium, chromium, copper, iron, lead, manganese, zinc, pH and temperature. Fourteen sampling stations on the Inflow Rivers and the spillway were monitored fortnightly. The samples collected for heavy metal analysis were preserved and stored in cooler boxes and analyzed in the laboratory using Standard Methods. Temperature and pH were measured in the field using HACH field testing kits. The concentration (mean ± standard deviation) of Zn was found to be 0.040±0.103 mg/L, Cd 0.031±0.007 mg/L, Pb 0.487±0.452 mg/L, Fe 7.53±13.34 mg/L, Mn 1.01±1.31 mg/L, chromium 0 mg/L and copper 0 mg/L. The concentrations of measured parameters deviate much from the recommended ones thus posing serious problems to aquatic life. It has also been shown that the high metal concentration levels in Lake Muhazi are related to landuse activities in the catchment. It is thus recommended that farming practices and erosion be controlled in the catchment to contain pollutant discharges into the lake. Lead is the main anthropogenic pollutant which has been found in the watershed of Lake Muhazi.

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

Summary1. 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.

Evaluating branched tetraether lipid-based palaeotemperature proxies in an urban, hyper-eutrophic polluted lake in South Africa

We evaluate the application of the branched glycerol dialkyl glycerol tetraether (br GDGT) based palaeotemperature and palaeoenvironmental proxy to a hyper-eutrophic, polluted and shallow oxic lake. Lake Zeekoevlei is the largest freshwater lake in South Africa, located close to Cape Town. We use published lake-based and soil-based calibration equations, and compare the reconstructed mean annual air temperature (MAT) with regional (South African) and local (Cape Town) instrumental temperature records. The distribution of br GDGTs in the lake sediments is influenced by air temperature. The lake-based calibration equation, which uses the methylation index of branched tetraethers/cyclisation ratio of branched tetraethers (MBT/CBT), formulated for African lakes (Tierney et al., 2010), fits well with the instrumental temperature records. Moreover, the CBT-derived pH likely reflects historic socioeconomic changes in catchment. Our results suggest that a polluted/hyper-eutrophic status and shallow water urban setting do not preclude application of the MBT/CBT-MAT proxy. However, further research is necessary to understand the behaviour of br GDGT–producing bacteria in polluted and highly productive lakes.

Ecological opportunity and sexual selection together predict adaptive radiation

A fundamental challenge to our understanding of biodiversity is to explain why some groups of species undergo adaptive radiations, diversifying extensively into many and varied species, whereas others do not. Both extrinsic environmental factors (for example, resource availability, climate) and intrinsic lineage-specific traits (for example, behavioural or morphological traits, genetic architecture) influence diversification, but few studies have addressed how such factors interact. Radiations of cichlid fishes in the African Great Lakes provide some of the most dramatic cases of species diversification. However, most cichlid lineages in African lakes have not undergone adaptive radiations. Here we compile data on cichlid colonization and diversification in 46 African lakes, along with lake environmental features and information about the traits of colonizing cichlid lineages, to investigate why adaptive radiation does and does not occur. We find that extrinsic environmental factors related to ecological opportunity and intrinsic lineage-specific traits related to sexual selection both strongly influence whether cichlids radiate. Cichlids are more likely to radiate in deep lakes, in regions with more incident solar radiation and in lakes where there has been more time for diversification. Weak or negative associations between diversification and lake surface area indicate that cichlid speciation is not constrained by area, in contrast to diversification in many terrestrial taxa. Among the suite of intrinsic traits that we investigate, sexual dichromatism, a surrogate for the intensity of sexual selection, is consistently positively associated with diversification. Thus, for cichlids, it is the coincidence between ecological opportunity and sexual selection that best predicts whether adaptive radiation will occur. These findings suggest that adaptive radiation is predictable, but only when species traits and environmental factors are jointly considered.

Recent speciation between sympatric Tanganyikan cichlid colour morphs

Lake Tanganyika, Africa's oldest lake, harbours an impressive diversity of cichlid fishes. Although diversification in its radiating groups is thought to have been initially rapid, cichlids from Lake Tanganyika show little evidence for ongoing speciation. In contrast, examples of recent divergence among sympatric colour morphs are well known in haplochromine cichlids from Lakes Malawi and Victoria. Here, we report genetic evidence for recent divergence between two sympatric Tanganyikan cichlid colour morphs. These Petrochromis morphs share mitochondrial haplotypes, yet microsatellite loci reveal that their sympatric populations form distinct genetic groups. Nuclear divergence between the two morphs is equivalent to that which arises geographically within one of the morphs over short distances and is substantially smaller than that among other sympatric species in this genus. These patterns suggest that these morphs diverged only recently, yet that barriers to gene flow exist which prevent extensive admixture despite their sympatric distribution. The morphs studied here provide an unusual example of active diversification in Lake Tanganyika's generally ancient cichlid fauna and enable comparisons of speciation processes between Lake Tanganyika and other African lakes.

Introgressive Hybridization between Color Morphs in a Population of Cichlid Fishes Twelve Years after Human-Induced Secondary Admixis

In the extremely species-rich haplochromine cichlid fishes of the East African Great Lakes, prezygotic isolation between closely related species is often maintained by color-assortative mating. In 1998, local fisherman working for the ornamental fish trade released different color morphs of the cichlid genus Tropheus into a small harbor basin in the southern part of Lake Tanganyika. This artificial amalgamation of color morphs provides a unique possibility to study mating patterns in cichlids in a natural environment over time. In a precursor study, we analyzed genotypes and phenotypes of almost 500 individuals sampled between 1999 and 2001 and uncovered a marked degree of color-assortative mating, which depended on the level of color pattern dissimilarity between morphs. Twelve years after introduction of nonindigenous morphs, we again sampled Tropheus individuals from the harbor basin and an adjacent, originally pure population and analyzed phenotypes (coloration) and genotypes (mitochondrial control region and 9 microsatellite loci) to assess the current status of the admixed population. Principal component analyses of color score data and population assignment tests demonstrate an increasing level of introgressive hybridization between morphs but also some ongoing color-assortative mating within morphs. The observed mating pattern might have been influenced by fluctuating environmental conditions such as periodic algal blooms or increased sedimentation causing turbid conditions in an otherwise clear lake.

Seasonal and interannual validation of satellite-measured reservoir levels at the Kainji dam

Although several studies have demonstrated the potential for inland water monitoring using satellite radar altimeters, the technology still carries with it certain limitations. Therefore, the direct application of altimetry data to reservoir management is often preceded by validation of satellite altimetry data with reliable ground-measured data. Kainji reservoir in Nigeria was selected for this study due to its remote geographical location, hydrological behavior, operational requirements, and morphometry. Validation of Kainji reservoir altimetric levels would highlight the applicability of the technology for monitoring one of Africa’s largest reservoirs with a surface area of 1270 km2. We performed and improved the validation exercise by separating each lake level dataset into two separate seasons: dry and wet, to investigate the effect of different seasons on the relationship between ground-measured and satellite-measured data sets. Lake level data from Topex/Poseidon and ERS/ENVISAT satellite altimeters were compared with gage data from 1992 to 2002. The results suggested a significantly higher relationship between gage and satellite data for the wet season (R2=0.93) than for the dry season (R2=0.77). For interannual validation, the trend was the same for both satellite altimeters but the relationship was generally higher for the T/P altimeter (R2=0.95). Root-mean-square errors in water levels ranged between 0.50 to 0.83 m for both altimeters and seasons, agreeing with values expected for lakes of identical physical characteristics. Key words: Satellite altimetry, remote sensing water, reservoir level variation, Kainji dam, African lakes.

The Diplomatic Lead in the United Nations Security Council and Local Actors' Violence: The Changing Terms of a Social Position

ABSTRACT Inspired by interactionist and constructivist sociology, this paper focuses on the United Nations Security Council as an arena with its own social rules and bureaucratic routines. It presents the informal position that security council diplomats delegate to specific members of the council, the position of “leader,” when dealing with specific dossiers. Indeed, a change occurred during the 1990s and the early 2000s in how council diplomats considered the task of leading a United Nations peace operation, notably in regard to the use of offensive military. The analysis of this change is documented with empirical material concerning the French leading position in the African Great Lakes region, from Rwanda in 1993 to Democratic Republic of the Congo in 2003. A second instance of diplomatic leadership considered is the leading role assumed by the British authorities in Sierra Leone (1991–2002). The new “leadership” role calls on United Nations diplomatic leaders to better protect the credibility of “their” peace operations. It sheds a light on the importance of deterring or incentive messages embedded in routine security council decisions and how routine practices may change when facing critical situations.

Microzooplankton feeding behaviour: grazing on the microbial and the classical food web of African soda lakes

We investigated the feeding behaviour of the dominant microzooplankton of saline lakes in the East African Rift Valley. A set of grazing experiments revealed high ingestion rates of the two euryhaline rotifers Brachionus dimidiatus and Brachionus plicatilis and of the large-sized omnivorous ciliates Frontonia sp. and Condylostoma magnum reflecting the unique nature of tropical saline systems. The size spectrum of ingested particles was broad and even included filamentous cyanobacteria such as the commonly dominating Arthrospira fusiformis. Feeding selectivity on cyanobacteria, however, was rather low showing higher values for cryptomonads and small ciliates. Bacterial biomass was favoured by the presence of grazers, as small bacterivorous predators were reduced at an average of 13.9%, showing the cascading effect of large zooplankton on the food web structure. Overall, based on this first-time study of the microzooplankton feeding behaviour in East African soda lakes, a strong structuring effect of rotifers and large ciliates on microbial plankton communities is assumed, especially in times of high consumer biomass.

Luminescence properties and optically (post-IR blue-light) stimulated luminescence dating of limnic sediments from northern Lake Malawi – Chances and limitations

Affected by the shifting Intertropical Convergence Zone, Lake Malawi holds an outstanding limnic sediment archive storing palaeo-environmental information of a climatically highly responsive landscape in the East African Rift Valley. Reliable chronological data are essential to interpret the lake-bottom sediments and optically stimulated luminescence (OSL) dating is an important method to build up the chronology. We report on material-property studies and OSL-dating results for polymineral fine grains extracted from piston cores obtained from the northern basin of Lake Malawi during the 1998 campaign of the International Decade of the East African Lakes (IDEAL). OSL-dating is based on a post-IR blue (pIRB) light stimulation single aliquot regeneration (SAR) protocol. Our results support earlier publications demonstrating the general applicability of OSL-dating on Lake Malawi sediments of Holocene and late Pleistocene age. However, the study also shows that dose rate estimations imply major challenges which need to be investigated in more detail to improve future OSL-dating results.

International Modes of Policy Influence: Does the EU Influence Good-Governance Policies in African, Caribbean and Pacific States?

The European Union is able to transfer its values and influence good governance policies in ACP states, through leverage and transnational networks. This paper tries to establish the leverage-transnational networks nexus in influencing policy and to account for the variance of European Union influence. In a comparative design, it tries to extricate these influences empirically by using a multi-variate regression analysis for the years 1975-2009, to establish whether European Union influences good governance policies in the African Great lakes states of Kenya, Uganda and Tanzania. Though the analysis controls for other influences, still evidence of European Union influence in the three countries is significant. It further indicates that the degree of influence is predicated on the complementary role of the modes of influence, which accounts for the variance.

Past and present mercury flux to a West African crater lake (Lake Bosomtwe/Bosumtwi, Ghana)

Lake sediment cores have been used to reconstruct mercury deposition patterns in many parts of the world; however, no studies to date have used these methods in West Africa, nor are there any published measurements of mercury deposition to this region. We measured mercury in a 210Pb dated sediment core from a meromictic crater lake in West Africa (Lake Bosomtwe, Ghana). Lake Bosomtwe has a very small catchment area to lake area ratio (1.1) and the sediment mercury profile is expected to reflect past and present atmospheric mercury deposition to the lake. Mercury concentrations in sediments as well as mercury flux to the sediments increased from the mid-1800s to latter half of the 1900s, however there has been a sharp decline in mercury flux to Lake Bosomtwe in recent decades. The recent decline in mercury flux to Lake Bosomtwe's sediments does not appear to be consistent with trends in local, regional or global mercury emissions, and may instead reflect declining global atmospheric mercury concentrations or declining European emissions, highlighting the importance of long-range atmospheric transport of mercury.

Divergent hormonal responses to social competition in closely related species of haplochromine cichlid fish

The diverse cichlid species flocks of the East African lakes provide a classical example of adaptive radiation. Territorial aggression is thought to influence the evolution of phenotypic diversity in this system. Most vertebrates mount hormonal (androgen, glucocorticoid) responses to a territorial challenge. These hormones, in turn, influence behavior and multiple aspects of physiology and morphology. Examining variation in competition-induced hormone secretion patterns is thus fundamental to an understanding of the mechanisms of phenotypic diversification. We test here the hypothesis that diversification in male aggression has been accompanied by differentiation in steroid hormone levels. We studied two pairs of sibling species from Lake Victoria belonging to the genera Pundamilia and Mbipia. The two genera are ecologically differentiated, while sibling species pairs differ mainly in male color patterns. We found that aggression directed toward conspecific males varied between species and across genera: Pundamilia nyererei males were more aggressive than Pundamilia pundamilia males, and Mbipia mbipi males were more aggressive than Mbipia lutea males. Males of both genera exhibited comparable attack rates during acute exposure to a novel conspecific intruder, while Mbipia males were more aggressive than Pundamilia males during continuous exposure to a conspecific rival, consistent with the genus difference in feeding ecology. Variation in aggressiveness between genera, but not between sibling species, was reflected in androgen levels. We further found that M. mbipi displayed lower levels of cortisol than M. lutea. Our results suggest that concerted divergence in hormones and behavior might play an important role in the rapid speciation of cichlid fishes.

Cattle, identity and genocide in the African Great Lakes region

Archaeological research into the origins of the cattle-keeping people of southwestern Uganda has overturned long-held beliefs about the separate origins of the pastoral and agricultural populations of the region. Study of indigenous political development and nineteenth-century colonialism shows that the present-day ethnic identities, which fuelled the genocide in Rwanda in 1994, are recent constructions.