Implications of regional fault distribution and kinematics for the uplift of rift flanks around the Rwenzori Mountains, Southwestern Uganda

Das Ziel dieser Arbeit bestand in der Untersuchung der Störungsverteilung und der Störungskinematik im Zusammenhang mit der Hebung der Riftschultern des Rwenzori Gebirges.rnDas Rwenzori Gebirge befindet sich im NNE-SSWbis N-S verlaufenden Albertine Rift, des nördlichsten Segments des westlichen Armes des Ostafrikanischen Grabensystems. Das Albertine Rift besteht aus Becken unterschiedlicher Höhe, die den Lake Albert, Lake Edward, Lake George und Lake Kivu enthalten. Der Rwenzori horst trennt die Becken des Lake Albert und des Lake Edward. Es erstreckt sich 120km in N-S Richtung, sowie 40-50km in E-W Richtung, der h¨ochste Punkt befindet sich 5111 ü. NN. Diese Studie untersucht einen Abschnitt des Rifts zwischen etwa 1°N und 0°30'S Breite sowie 29°30' und 30°30' östlicher Länge ersteckt. Auch die Feldarbeit konzentrierte sich auf dieses Gebiet.rnrnHauptzweck dieser Studie bestand darin, die folgende These auf ihre Richtigkeit zu überprüfen: ’Wenn es im Verlauf der Zeit tatsächlich zu wesentlichen Änderungen in der Störungskinematik kam, dann ist die starke Hebung der Riftflanken im Bereich der Rwenzoris nicht einfach durch Bewegung entlang der Graben-Hauptst¨orungen zu erklären. Vielmehr ist sie ein Resultat des Zusammenspiels mehrerer tektonische Prozesse, die das Spannungsfeld beeinflussen und dadurch Änderungen in der Kinematik hervorrufen.’ Dadurch konzentrierte sich die Studie in erster Linie auf die Störungsanalyse.rnrnDie Kenntnis regionaler Änderungen der Extensionsrichtung ist entscheidend für das Verständnis komplexer Riftsysteme wie dem Ostafrikanischen Graben. Daher bestand der Kern der Untersuchung in der Kartierung von Störungen und der Untersuchung der Störungskinematik. Die Aufnahme strukturgeologischer Daten konzentrierte sich auf die Ugandische Seite des Rifts, und Pal¨aospannungen wurden mit Hilfe von St¨orungsdaten durch Spannungsinversion rekonstruiert.rnDie unterschiedliche Orientierung spr¨oder Strukturen im Gelände, die geometrische Analyse der geologischen Strukturen sowie die Ergebnisse von Mikrostrukturen im Dünnschliff (Kapitel 4) weisen auf verschiedene Spannungsfelder hin, die auf mögliche Änderungen der Extensionsrichtung hinweisen. Die Resultate der Spannungsinversion sprechen für Ab-, Über- und Blattverschiebungen sowie für Schrägüberschiebungen (Kapitel 5). Aus der Orientierung der Abschiebungen gehen zwei verschiedene Extensionsrichtungen hervor: im Wesentlichen NW-SE Extension in fast allen Gebieten, sowie NNE-SSW Extension im östlichen Zentralbereich.rnAus der Analyse von Blattverschiebungen ergaben sich drei unterschiedliche Spannungszustände. Zum Einen NNW-SSE bis N-S Kompression in Verbindung mit ENE-WSW bzw E-W Extension wurde für die nördlichen und die zentralen Ruwenzoris ausgemacht. Ein zweiter Spannungszustand mit WNW-ESE Kompression/NNE-SSW Extension betraf die Zentralen Rwenzoris. Ein dritter Spannungszustand mit NNW-SSE Extension betraf den östlichen Zentralteil der Rwenzoris. Schrägüberschiebungen sind durch dazu schräge Achsen charakterisiert, die für N-S bis NNW-SSE Kompression sprechen und ausschließlich im östlichen Zentralabschnitt auftreten. Überschiebungen, die hauptsächlich in den zentralen und den östlichen Rwenzoris auftreten, sprechen für NE-SW orientierten σ2-Achsen und NW-SE Extension.rnrnEs konnten drei unterschiedliche Spannungseinflüsse identifiziert werden: auf die kollisionsbedingte Bildung eines Überschiebungssystem folgte intra-kratonische Kompression und schließlich extensionskontrollierte Riftbildung. Der Übergang zwischen den beiden letztgenannten Spannungszuständen erfolgte Schrittweise und erzeugte vermutlich lokal begrenzte Transpression und Transtension. Gegenw¨artig wird die Störungskinematik der Region durch ein tensiles Spannungsregime in NW-SE bis N-S Richtung bestimmt.rnrnLokale Spannungsvariationen werden dabei hauptsächlich durch die Interferenzrndes regionalen Spannungsfeldes mit lokalen Hauptst¨orungen verursacht. Weitere Faktoren die zu lokalen Veränderungen des Spannungsfeldes führen können sind unterschiedliche Hebungsgeschwindigkeiten, Blockrotation oder die Interaktion von Riftsegmenten. Um den Einfluß präexistenter Strukturen und anderer Bedingungen auf die Hebung der Rwenzoris zu ermitteln, wurde der Riftprozeß mit Hilfe eines analogen ’Sandbox’-Modells rekonstruiert (Kapitel 6). Da sich die Moho-Diskontinuität im Bereich des Arbeitsgebietes in einer Tiefe von 25 km befindet, aktive Störungen aber nur bis zu einer Tiefe von etwa 20 km beobachtet werden können (Koehn et al. 2008), wurden nur die oberen 25 km im Modell nachbebildet. Untersucht und mit Geländebeobachtungen verglichen wurden sowohl die Reihenfolge, in der Riftsegmente entstehen, als auch die Muster, die sich im Verlauf der Nukleierung und des Wachstums dieser Riftsegmente ausbilden. Das Hauptaugenmerk wurde auf die Entwicklung der beiden Subsegmente gelegt auf denen sich der Lake Albert bzw. der Lake Edward und der Lake George befinden, sowie auf das dazwischenliegende Rwenzori Gebirge. Das Ziel der Untersuchung bestand darin herauszufinden, in welcher Weise das südwärts propagierende Lake Albert-Subsegment mit dem sinistral versetzten nordwärts propagierenden Lake Edward/Lake George-Subsegment interagiert.rnrnVon besonderem Interesse war es, in welcherWeise die Strukturen innerhalb und außerhalb der Rwenzoris durch die Interaktion dieser Riftsegmente beeinflußt wurden. rnrnDrei verschiedene Versuchsreihen mit unterschiedlichen Randbedingungen wurden miteinander verglichen. Abhängig vom vorherrschenden Deformationstyp der Transferzone wurden die Reihen als ’Scherungs-dominiert’, ’Extensions-dominiert’ und als ’Rotations-dominiert’ charakterisiert. Die Beobachtung der 3-dimensionalen strukturellen Entwicklung der Riftsegmente wurde durch die Kombination von Modell-Aufsichten mit Profilschnitten ermöglicht. Von den drei genannten Versuchsreihen entwickelte die ’Rotationsdominierten’ Reihe einen rautenförmiger Block im Tranferbereich der beiden Riftsegmente, der sich um 5−20° im Uhrzeigersinn drehte. DieserWinkel liegt im Bereich des vermuteten Rotationswinkel des Rwenzori-Blocks (5°). Zusammengefasst untersuchen die Sandbox-Versuche den Einfluss präexistenter Strukturen und der Überlappung bzw. Überschneidung zweier interagierender Riftsegmente auf die Entwicklung des Riftsystems. Sie befassen sich darüber hinaus mit der Frage, welchen Einfluss Blockbildung und -rotation auf das lokale Stressfeld haben.

Evolution und historische Biogeographie von Senecio L. mit Fokus auf die Alte Welt

Senecio L., eine der größten Angiospermengattungen, entstand im mittleren Miozän in trockenen oder Winterregengebieten des südlichen Afrikas. Von dort ausgehend wurden die anderen Kontinente mehrmals unabhängig voneinander besiedelt. Die hohe Artenanzahl, das junge Alter und die was weltweite Verbreitung der Gattung stellt eine Besonderheit dar. Die meisten anderen jungen und schnellen Radiationen sind nicht so weit verbreitet. Es scheint, als wäre die hohe Artenanzahl und die weite Verbreitung das Ergebnis von einer guten Ausbreitungsfähigkeit, einem hohe Samenansatz sowie einer Präferenz für offene und gestörte Standorte.rnDie Paläarktis wurde zweimal unabhängig voneinander kolonisiert, beide Linien kommen in etwa in der gleichen Region vor, unterscheiden sich aber in ihrer Habitatpräferenz und Lebensdauer. Die eine Linie kommt zumeist entlang des Mittelmeeres vor und stammt am wahrscheinlichsten von annuellen Tieflandarten aus den trockenen oder Winterregengebieten des südlichen Afrikas ab. Die andere Linie stammt wahrscheinlich von Drakensbergarten ab und besiedelt auch bevorzugt bergige Regionen in der Paläarktis. Senecio ist außerdem die artenreichste Gattung in den tropisch afroalpinen Regionen Ostafrikas. Die Arten sind während des Pliozäns oder Pleistozäns entstanden und das Ergebnis von mehrfacher Einwanderung aus anderen temperaten Regionen, in situ Artbildung spielte nur eine untergeordnete Bedeutung. rn

Investigations into the degassing and eruption mechanisms of Nyamuragira volcano, Democratic Republic of the Congo (Africa)

One of two active volcanoes in the western branch of the East African Rift, Nyamuragira (1.408ºS, 29.20ºE; 3058 m) is located in the D.R. Congo. Nyamuragira emits large amounts of SO2 (up to ~1 Mt/day) and erupts low-silica, alkalic lavas, which achieve flow rates of up to ~20 km/hr. The source of the large SO2 emissions and pre-eruptive magma conditions were unknown prior to this study, and 1994-2010 lava volumes were only recently mapped via satellite imagery, mainly due to the region’s political instability. In this study, new olivine-hosted melt inclusion volatile (H2O, CO2, S, Cl, F) and major element data from five historic Nyamuragira eruptions (1912, 1938, 1948, 1986, 2006) are presented. Melt compositions derived from the 1986 and 2006 tephra samples best represent pre-eruptive volatile compositions because these samples contain naturally glassy inclusions that underwent less post-entrapment modification than crystallized inclusions. The total amount of SO2 released from the 1986 (0.04 Mt) and 2006 (0.06 Mt) eruptions are derived using the petrologic method, whereby S contents in melt inclusions are scaled to erupted lava volumes. These amounts are significantly less than satellite-based SO2 emissions for the same eruptions (1986 = ~1 Mt; 2006 = ~2 Mt). Potential explanations for this observation are: 1) accumulation of a vapor phase within the magmatic system that is only released during eruptions, and/or 2) syn-eruptive gas release from unerupted magma. Post-1994 Nyamuragira lava volumes were not available at the beginning of this study. These flows (along with others since 1967) are mapped with Landsat MSS, TM, and ETM+, Hyperion, and ALI satellite data and combined with published flow thicknesses to derive volumes. Satellite remote sensing data was also used to evaluate Nyamuragira SO2 emissions. These results show that the most recent Nyamuragira eruptions injected SO2 into the atmosphere between 15 km (2006 eruption) and 5 km (2010 eruption). This suggests that past effusive basaltic eruptions (e.g., Laki 1783) are capable of similar plume heights that reached the upper troposphere or tropopause, allowing SO2 and resultant aerosols to remain longer in the atmosphere, travel farther around the globe, and affect global climates.

Evolution of the Southern Kenya Rift from Miocene to present with a focus on the Magadi area

The Kenya (a.k.a., Gregory) Rift is a geologically active area located within the eastern branch of the larger East African Rift System (EARS). The study area is located in the southern Kenya Rift between 1° South and the Kenya-Tanzania border (covering approximately 1.5 square degrees, semi-centered on Lake Magadi) and is predominantly filled with extrusive igneous rocks (mostly basalts, phonolites and trachytes) of Miocene age or younger. Sediments are thin, less than 1.5Ma, and are confined to small grabens. The EARS can serve both as an analogue for ancient continental rifting and as a modern laboratory to observe the geologic processes responsible for rifting. This study demonstrates that vintage (as in older, quality maps published by the Kenya Geological Survey, that may be outdated based on newer findings) quarter-degree maps can be successfully combined with recently published data, and used to interpret satellite (mainly Landsat 7) images to produce versatile, updated digital maps. The study area has been remapped using this procedure and although it covers a large area, the mapping retains a quadrangle level of detail. Additionally, all geologic mapping elements (formations, faults, etc.) have been correlated across older map boundaries so that geologic units don't end artificially at degree boundaries within the study area. These elements have also been saved as individual digital files to facilitate future analysis. A series of maps showing the evolution of the southern Kenya rift from the Miocene to the present was created by combining the updated geologic map with age dates for geologic formations and fault displacements. Over 200 age dates covering the entire length of the Kenya Rift have been compiled for this study, and 6 paleo-maps were constructed to demonstrate the evolution of the area, starting with the eruption of the Kishalduga and Lisudwa melanephelinites onto the metamorphic basement around 15Ma. These eruptions occurred before the initial rift faulting and were followed by a massive eruption of phonolites between 13-10 Ma that covered most of the Kenya dome. This was followed by a period of relative quiescence, until the initial faulting defined the western boundary of the rift around 7Ma. The resulting graben was asymmetrical until corresponding faults to the east developed around 3Ma. The rift valley was flooded by basalts and trachytes between 3Ma and 700ka, after which the volcanic activity slowed to a near halt. Since 700ka most of the deposition has been comprised of sediments, mainly from lakes occupying the various basins in the area. The main results of this study are, in addition to a detailed interpretation of the rift development, a new geologic map that correlates dozens of formations across old map boundaries and a compilation of over 300 age dates. Specific products include paleomaps, tables of fault timing and displacement, and volume estimates of volcanic formations. The study concludes with a generalization of the present environment at Magadi including discussions of lagoon chemistry, mantle gases in relation to the trona deposit, and biology of the hot springs. Several biologic samples were collected during the 2006 field season in an attempt to characterize the organisms that are commonly seen in the present Lake Magadi environment. Samples were selected to represent the different, distinctive forms that are found in the hotsprings. Each sample had it own distinctive growth habit, and analysis showed that each was formed by a different cyanobacterial. Actual algae was rare in the collected samples, and represented by a few scattered diatoms.

Spatial and Temporal Evolution of the Volcanics and Sediments of the Kenya Rift

New volumetric and mass flux estimates have been calculated for the Kenya Rift. Spatial and temporal histories for volcanic eruptions, lacustrine deposition, and hominin fossil sites are presented, aided by the compilation of a new digital geologic map. Distribution of volcanism over time indicates several periods of southward expansion followed by relative positional stasis. Volcanism occurs throughout the activated rift length, with no obvious abandonment as the rift system migrated. The main exception is a period of volcanic concentration around 10 Ma, when activity was constrained within 2° of the equator. Volumes derived from seismic data indicate a total volume of c. 310,000 km3 (2.47 x 1010 kg/yr ), which is significantly more than the map-derived volumes found here or published previously. Map-based estimates are likely affected by a bias against recognizing small volume events in the older record. Such events are, however, the main driver of erupted volume over the last 5 Ma. A technique developed here to counter this bias results in convergence of the two volume estimation techniques. Relative erupted composition over time is variable. Overall, the erupted material has a mafic to silicic ratio of 0.9:1. Basalts are distinctly more common in the Turkana region, which previously experienced Mesozoic rifting. Despite the near equal ratio of mafic to silicic products, the Kenya Rift otherwise fits the definition of a SLIP. It is proposed that the compositions would better fit the published definition if the Turkana region was not twice-rifted. Lacustrine sedimentation post-dates initial volcanism by about 5 million years, and follows the same volcanic trends, showing south and eastward migration over time. This sedimentation delay is likely related to timing of fault displacements. Evidence of hominin habitation is distinctly abundant in the northern and southern sections of the Kenya Rift, but there is an observed gap in the equatorial rift between 4 and 0.5 million years ago. After 0.5 Ma, sites appear to progress towards the equator. The pattern and timing of hominid site distributions suggests that the equatorial gap in habitation may be the result of active volcanic avoidance.

Aerological observations in the Tropics in the Early Twentieth Century

In the first decades of the 20th century, aerological observations were for the first time performed in tropical regions. One of the most prominent endeavours in this respect was ARTHUR BERSON’s aerological expedition to East Africa. Although the main target was the East African monsoon circulation, the expedition provided also other insights that profoundly changed meteorology and climatology. BERSON observed that the tropical tropopause was much higher and colder than that over midlatitudes. Moreover, westerly winds were observed in the lower stratosphere, apparently contradicting the high-altitude equatorial easterly winds that were known since the Krakatoa eruption (‘‘Krakatoa easterlies’’). The puzzle was only resolved five decades later with the discovery of the Quasi-Biennial Oscillation (QBO). In this paper we briefly summarize the expedition of BERSON and review the results in a historical context and in the light of the current research. In the second part of the paper we re-visit BERSON’s early aerological observations, which we have digitized. We compare the observed wind profiles with corresponding profiles extracted from the ‘‘Twentieth Century Reanalysis’’, which provides global three-dimensional weather information back to 1871 based on an assimilation of sea-level and surface pressure data. The comparison shows a good agreement at the coast but less good agreement further inland, at the shore of Lake Victoria, where the circulation is more complex. These results demonstrate that BERSON’s observations are still valuable today as input to current reanalysis systems or for their validation.

Evaluating watershed service availability under future management and climate change scenarios in the Pangani Basin

Watershed services are the benefits people obtain from the flow of water through a watershed. While demand for such services is increasing in most parts of the world, supply is getting more insecure due to human impacts on ecosystems such as climate or land use change. Population and water management authorities therefore require information on the potential availability of watershed services in the future and the trade-offs involved. In this study, the Soil and Water Assessment Tool (SWAT) is used to model watershed service availability for future management and climate change scenarios in the East African Pangani Basin. In order to quantify actual “benefits”, SWAT2005 was slightly modified, calibrated and configured at the required spatial and temporal resolution so that simulated water resources and processes could be characterized based on their valuation by stakeholders and their accessibility. The calibrated model was then used to evaluate three management and three climate scenarios. The results show that by the year 2025, not primarily the physical availability of water, but access to water resources and efficiency of use represent the greatest challenges. Water to cover basic human needs is available at least 95% of time but must be made accessible to the population through investments in distribution infrastructure. Concerning the trade-off between agricultural use and hydropower production, there is virtually no potential for an increase in hydropower even if it is given priority. Agriculture will necessarily expand spatially as a result of population growth, and can even benefit from higher irrigation water availability per area unit, given improved irrigation efficiency and enforced regulation to ensure equitable distribution of available water. The decline in services from natural terrestrial ecosystems (e.g. charcoal, food), due to the expansion of agriculture, increases the vulnerability of residents who depend on such services mostly in times of drought. The expected impacts of climate change may contribute to an increase or decrease in watershed service availability, but are only marginal and much lower than management impacts up to the year 2025.

Metabolic divergence between sibling species of cichlids Pundamilia nyererei and Pundamilia pundamilia

This study compared Pundamilia nyererei and Pundamilia pundamilia males in routine metabolic rate (RR ) and in the metabolic costs males pay during territorial interactions (active metabolic rate, RA ). Pundamilia nyererei and P. pundamilia males housed in social isolation did not differ in RR . In contrast to expectation, however, P. nyererei males used less oxygen than P. pundamilia males, for a given mass and level of agonistic activity. This increased metabolic efficiency may be an adaptation to limit the metabolic cost that P. nyererei males pay for their higher rate of aggressiveness compared to P. pundamilia males. Thus, the divergence between the species in agonistic behaviour is correlated with metabolic differentiation. Such concerted divergence in physiology and behaviour might be widespread in the dramatically diverse cichlid radiations in East African lakes and may be an important factor in the remarkably rapid speciation of these fishes. The results did not support the hypothesis that higher metabolic rates caused a physiological cost to P. nyererei males that would offset their dominance advantage.

Weathering geochemistry and Sr-Nd fingerprints of equatorial upper Nile and Congo muds

This study investigates processes of sediment generation in equatorial central Africa. An original, complete and integrated mineralogical-geochemical database on silt-sized sediments derived from different parent rocks (basalt, granite, gneiss, metapsammite, sandstone) along the East African Rift from 5°S in Tanzania to 5°N in Sudan is presented and used to assess the incidence of diverse factors controlling sediment composition (source-rock lithology, geomorphology, hydraulic sorting, grain size, recycling), with particular emphasis on chemical weathering.

Adaptive Molecular Evolution in the Opsin Genes of Rapidly Speciating Cichlid Species

Cichlid fish inhabit a diverse range of environments that vary in the spectral content of light available for vision. These differences should result in adaptive selective pressure on the genes involved in visual sensitivity, the opsin genes. This study examines the evidence for differential adaptive molecular evolution in East African cichlid opsin genes due to gross differences in environmental light conditions. First, we characterize the selective regime experienced by cichlid opsin genes using a likelihood ratio test format, comparing likelihood models with different constraints on the relative rates of amino acid substitution, across sites. Second, we compare turbid and clear lineages to determine if there is evidence of differences in relative rates of substitution. Third, we present evidence of functional diversification and its relationship to the photic environment among cichlid opsin genes. We report statistical evidence of positive selection in all cichlid opsin genes, except short wavelength–sensitive 1 and short wavelength–sensitive 2b. In all genes predicted to be under positive selection, except short wavelength–sensitive 2a, we find differences in selective pressure between turbid and clear lineages. Potential spectral tuning sites are variable among all cichlid opsin genes; however, patterns of substitution consistent with photic environment–driven evolution of opsin genes are observed only for short wavelength–sensitive 1 opsin genes. This study identifies a number of promising candidate-tuning sites for future study by site-directed mutagenesis. This work also begins to demonstrate the molecular evolutionary dynamics of cichlid visual sensitivity and its relationship to the photic environment.

Nuclear markers reveal unexpected genetic variation and a Congolese-Nilotic origin of the Lake Victoria cichlid species flock

Phylogenetic analyses based on mitochondrial (mt) DNA have indicated that the cichlid species flock of the Lake Victoria region is derived from a single ancestral species found in East African rivers, closely related to the ancestor of the Lake Malawi cichlid species flock. The Lake Victoria flock contains ten times less mtDNA variation than the Lake Malawi radiation, consistent with current estimates of the ages of the lakes. We present results of a phylogenetic investigation using nuclear (amplified fragment length polymorphism) markers and a wider coverage of riverine haplochromines. We demonstrate that the Lake Victoria–Edward flock is derived from the morphologically and ecologically diverse cichlid genus Thoracochromis from the Congo and Nile, rather than from the phenotypically conservative East African Astatotilapia. This implies that the ability to express much of the morphological diversity found in the species flock may by far pre–date the origin of the flock. Our data indicate that the nuclear diversity of the Lake Victoria–Edward species flock is similar to that of the Lake Malawi flock, indicating that the genetic diversity is considerably older than the 15 000 years that have passed since the lake began to refill. Most of this variation is manifested in trans–species polymorphisms, indicating very recent cladogenesis from a genetically very diverse founder stock. Our data do not confirm strict monophyly of either of the species flocks, but raise the possibility that these flocks have arisen from hybrid swarms.

Molecular epidemiology of tuberculosis in Cambodian children

SUMMARY We analysed Mycobacterium tuberculosis strains from children, hospitalized from January 2004 to July 2008 in the largest paediatric hospital complex in Cambodia. Specimens were tested for drug susceptibility and genotypes. From the 260 children, 161 strains were available. The East African-Indian genotype family was the most common (59·0%), increasing in frequency with distance from the Phnom Penh area, while the frequency of the Beijing genotype family strains decreased. The drug resistance pattern showed a similar geographical gradient: lowest in the northwest (4·6%), intermediate in the central (17·1%), and highest in the southeastern (30·8%) parts of the country. Three children (1·9%) had multidrug-resistant tuberculosis. The Beijing genotype and streptomycin resistance were significantly associated (P < 0·001). As tuberculosis in children reflects recent transmission patterns in the community, multidrug resistance levels inform about the current quality of the tuberculosis programme.

Evidence of trepanation in Late Iron Age Switzerland (420–240BC)

In Europe the procedure of trepanation is known since the Neolithic and is still practiced today in East African native tribes. Trepanation is the oldest known surgical intervention and is defined as the intentional penetration of the cranial vault with removal of piece of a skull bone. Therefore, it is believed that neurosurgery is one of the world’s oldest professions. In this study two skulls with lesions from the Late Iron Age cemetery of Münsingen (420–240 BC), Switzerland, are presented. Aim of the study was to analyse the lesions and to assess whether they were caused by surgical interventions. Sex and age of the individuals were determined by current morphologic-anthropological methods. Radiological examinations were performed with a multislice CT-scanner. Different trepanation methods and signs of healing are discussed. In Switzerland about 33 skulls with possible trepanations dating from Neolithic to medieval times are known and are presented in a short review. Studies of ancient surgical interventions provide important information of pre- and early historic populations. Trepanations of the skull such as the presented cases prove a profound understanding of the human body in ancient times.

Explosive speciation rates and unusual species richness in haplochromine cichlid fishes: Effects of sexual selection

Ancient lakes are often unusually species rich, mostly as a result of radiation and species-flock formation having taken place in only one or a few of many taxa present. Understanding why some taxa radiate and others do not is at the heart of understanding biodiversity. In this chapter I discuss possible explanations for disproportionally large species numbers in some cichlid fish lineages in East African Great Lakes: the halochromine cichlid fishes in Lakes Victoria and Malawi. I show that speciation rates in this group are higher than in any other lacustrine fish radiation. Against this background, I review hypotheses put forward to explain diversity in cichlid species flocks. The evolution of species diversity requires three processes: speciation, ecological radiation and anatomical diversification, and it is wrong to consider hypotheses that are relevant to different processes as alternatives to each other. The African cichlid species flocks show unusually high ecological species packing in several phylogenetic groups and unusually high speciation rates in haplochromines. Therefore, it maybe concluded that at least two evolutionary models are required to explain the difference between cichlid diversity and other fish diversity in East African Lakes: one for speciation in haplochromines and one for coexistence. Subsequently I review work on speciation in haplochromines, and in particular studies aimed at testing the hypothesis of speciation by sexual selection. Haplochromines have a polygynous mating system, conducive to sexual selection, but other polygynous cichlids are not particularly species rich. This suggests that more than just strong sexual selection is required to explain haplochromine species richness. Recent palaeoecological evidence undermines the previously popular hypotheses that explained the species richness of Lake Victoria in terms of speciation under varying natural or sexual selection regimes in satellite lakes or in isolated lake basins. I summarize experimental and comparative studies, which provide evidence for two mechanisms of sympatric speciation by disruptive sexual selection on polymorphic coloration. Such modes of speciation may explain (i) the high speciation rates in colour polymorphic lineages of haplochromine cichlids under conditions where colour variation is visible in clear water, and (ii) in combination with factors that affect population survival, the unusual species richness in haplochromine species flocks. I argue that sexual selection, if disruptive, can accelerate the pace of adaptive radiation because the resultant genetic population fragmentation allows a much increased rate of differential response to disruptive natural selection. Hence, the ecological pattern of diversity resembles that produced by disruptive natural selection, with the difference that disruptive sexual selection continues to cause (gross) speciation even after niche space is saturated. This may explain the unusually high numbers of very closely related and ecologically similar species in haplochromine species flocks. The role of disruptive sexual selection is twofold: it not only causes speciation, but also maintains reproductive isolation in sympatry between species that have evolved in sympatry or allopatry. Therefore, the maintenance of diversity in species flocks that originated through sexual selection depends on the persistence of the selection regime within the environmental signal space under which that diversity evolved.

Can sympatric speciation by disruptive sexual selection explain rapid evolution of cichlid diversity in Lake Victoria?

Rapid speciation can occur on ecological time scales and interfere with ecological processes, resulting in species distribution patterns that are difficult to reconcile with ecological theory. The haplochromine cichlids in East African lakes are an extreme example of rapid speciation. We analyse the causes of their high speciation rates. Various studies have identified disruptive sexual selection acting on colour polymorphisms that might cause sympatric speciation. Using data on geographical distribution, colouration and relatedness from 41 species endemic to Lake Victoria, we test predictions from this hypothesis. Plotting numbers of pairs of closely related species against the amount of distributional overlap between the species reveals a bimodal distribution with modes on allopatric and sympatric. The proportion of sister species pairs that are heteromorphic for the traits under disruptive selection is higher in sympatry than in allopatry. These data support the hypothesis that disruptive sexual selection on colour polymorphisms has caused sympatric speciation and help to explain the rapid radiation of haplochromine species flocks.