On the Cyprideis species flock (Crustacea, Ostracoda) in Lake Tanganyika, with the description of four new species

Four new species, belonging to three genera, of ostracods from Lake Tanganyika are described: Mesocyprideis nitida sp.nov., Romecytheridea belone sp.nov., R. plegma sp. nov. and Cyprideis loricata sp. nov. The Cyprideis species flock of Lake Tanganyika now consists of 17 formally described species (16 endemic) in 6 genera (4 endemic). The geographical and ecological distribution of the four species is briefly discussed and some supplementary remarks on the radiation of the Cyprideis species flock in Lake Tanganyika are offered.     

On the age and origin of the species flock of haplochromine cichlid fishes of Lake Victoria

Recent suggestions concerning the age and origin of the flock of haplochromine cichlid fishes in Lake Victoria (East Africa) are considered. These accept as proven the suggestion that Lake Victoria dried out completely in the Late Pleistocene, was dry for several thousand years, and refilled ca. 12400 years ago. Apart from the fact that other geophysical evidence contradicts this claim, its biological implications, which do likewise, have never been considered by those who have accepted it. Like those of all previous authors who have seized on the presence of the haplochromine flock of perhaps more than 600 species as evidence of extremely rapid evolution since the lake allegedly refilled, the account completely overlooks the fact that any such desiccation must have eliminated not only the haplochromine cichlids but the entire biota of the lake. Nevertheless, its present fauna not only includes the haplochromines but many other endemic organisms that would not be expected, and whose presence and history demand an explanation if the lake did indeed dry out. No such explanation has been offered, nor does such seem possible. The recent interpretation of events is questioned and rejected.     

Biparental mouthbrooding of the catfish Phyllonemus filinemus in Lake Tanganyika

This article reports biparental mouthbrooding of the bagrid catfish Phyllonemus filinemus in Lake Tanganyika, based on analysis of specimens collected during SCUBA diving. This catfish was nocturnally active, and in the daytime it was concealed singly or in pairs beneath rocks. Within a breeding pair, the male or female alone incubated all the brood in the mouth until the offspring attained 12 mm or so in total length, but thereafter joint mouthbrooding or guarding by both parents took place. Most females of nonbrooding pairs showed high values of gonadosomatic index (GSI), whereas all females of brooding pairs and most single females showed low GSI values. This fact indicated that a pair is formed at a time near the gonadal maturation of the female and separates after the brood is reared. No significant difference in body condition among parents of different reproductive states was observed, which suggested that their condition does not deteriorate markedly as the result of foraging by an off-duty parent. Received: September 16, 2000 / Revised: November 18, 2000 / Accepted: January 23, 2001     

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.     

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.     

Evolution in isolation: the Gyraulus species flock from Miocene Lake Steinheim revisited

The Miocene Steinheim Basin in SW Germany is an ancient (long-lived) palaeo-lake that has existed over some hundreds of thousands of years. It is an iconic fossil site, because the historically oldest phylogenetic tree of extinct organisms was based on specimens described from this locality. Today the basin contains 30–40 m thickness of lake sediments with planorbid snails of the genus Gyraulus occurring in rock-forming quantities. The shells are morphologically highly disparate with forms ranging from the tiny, planispiral founder species Gyraulus kleini, to fragile corkscrew-like uncoiled forms and to large trochiform morphs with thick shells. In total, this presumably monophyletic species flock contains 17 species distributed in time and space, all of which are endemic, except for the founder species. Up to nine of them occur in a single sedimentary level and are inferred to have lived together. Such an extreme rate of endemism makes fossil Lake Steinheim special among extant and fossil lakes. This review article summarises and discusses the species concept(s), indications for endemism, speciation processes, the phylogenetic concept(s) and factors controlling evolution. It also provides directions for future research.     

Haematology and habits in catfish of the genus Synodontis

African catfish of the genus Synodontis vary in their habits, from inverted plankton eaters to normally oriented benthos eaters. Haemoglobin concentrations and red cell counts vary between populations of one species, and between species of differing habits. Synodontis schall from the well-aerated White Nile had lower haemoglobin concentrations than members of the same species from Lake Kainji, Nigeria. Synodontis membranaceus , an inverted plankton eater, had lower haemoglobin concentrations than the bottom dwelling S. filamentosus and S. nigrita. Species with intermediate habits, such as S. batensoda and S. ocellifer had intermediate concentrations of haemoglobin.
It is suggested that these differences in haemoglobin concentration are a result of differing habits. Those species that enter poorly aerated water are stimulated to make more haemoglobin.     

Ecological correlates of species differences in the Lake Tanganyika crab radiation

The endemic crabs of Lake Tanganyika include a phenotypically diverse clade that exhibits recent divergence and low phylogenetic species resolution. There are indications that ecological niche segregation has played a prominent role in the divergence of this clade. We used habitat surveys, gut content analyses and stable isotope analyses to test the extent to which morphological species are ecologically different. Our data show some interspecific segregation in depth, substrate type and mean stable isotope signatures. At the same time, a considerable level of ecological niche overlap is evident among species of Platythelphusa that coexist in rocky littoral habitats. We consider these results in the framework of adaptive radiation theory, and we discuss general ramifications for the maintenance of species diversity in Lake Tanganyika.     

Molecular analyses of ostracod flocks from Lake Baikal and Lake Tanganyika

Ancient lakes are excellent laboratories for evolutionary research, where species can be studied in the cradle where they originated. In this article, we investigate two endemic ostracod species flocks from the two oldest lakes in the world, Lake Baikal (LB) (ca. 28 myr) and Lake Tanganyika (LT) (ca. 12 myr), with DNA sequence data. Nuclear ITS1 failed to resolve the phylogeny of both flocks. Whilst most phylogenetic relationships of the Tanganyika flock are resolved with mitochondrial COI, the Baikalian tree contains multifurications of up to seven different clades. The Tanganyikan Cyprideis flock shows higher genetic variability, which matches its higher morphological variability. A significant deviation from a constant divergence rate through time indicates that the Cytherissa flock most likely experienced explosive speciation events during its earlier history. Comparative analyses of substitution rates furthermore revealed that they are not clock-wise for COI. Ancestral Cytherissa probably radiated in LB 5–8 myr ago, around the time when the cold, oxygenated abyss was formed in LB. The Tanganyikan Cyprideis flock is almost twice as old as the Baikalian Cytherissa flock, and possibly older than LT itself, ca. 15 myr. The Cyprideis flock has survived drastic lake level changes and resulting salinity crises during its entire history.     

Retinofugal and retinopetal connections in the upside-down catfish (Synodontis nigriventris)

Summary The retinofugal and retinopetal connections in the upside-down catfish Synodontis nigriventris were studied by use of the horseradish-peroxidase (HRP) techniques, autoradiography, and degeneration-silver methods. An unusual retinal projection to the torus semicircularis as well as projections to the retina from three different sources in the brain are described. After intra-ocular injections of HRP, labeled cells were found in the optic tectum, the dorsomedial optic nucleus and one of the pretectal nuclei. These new findings support the basic hypothesis (i) that neuronal connections are more extensive in primitive brains, and (ii) that the evolutionary development of more complex brains involves the loss of some selected connections.     

Nuclear gene variation and molecular dating of the cichlid species flock of Lake Malawi

The cichlid fishes of Lake Malawi are famously diverse. However, evolutionary studies have been difficult because of their recent and uncertain phylogenetic history. Portions of 12 nuclear loci were sequenced in nine rock-dwelling species (mbuna) and three representatives of pelagic nonmbuna species. In contrast to the pattern of variation at mitochondrial genes, which do provide phylogenetic resolution at the level of mbuna versus nonmbuna, and among some genera, the nuclear loci were virtually devoid of phylogenetic signal. Only a small minority of variable positions were phylogenetically informative, and no phylogenetic branches are supported by more than one site. From the nuclear gene perspective the Malawian radiation appears to be a star phylogeny, as if the founding of the lake was accompanied by a partial bottleneck. The pattern is different from that found in Lake Victoria, in which nuclear loci share large amounts of ancestral variation. In the case of nuclear genes of Lake Malawi, the absence of phylogenetically informative variation suggests a relative absence of ancestral variation. Nuclear genes also differed from the mitochondria in having nearly twice the amount of divergence from Oreochromis (tilapia). An approximate splitting time between mbuna and nonmbuna lineages was estimated as 0.7 Myr. Oreochromis is estimated to have diverged from the cichlids in Lake Malawi and Lake Tanganyika about 18 MYA.     

Diatoms from surface sediments of the northern part of Lake Tanganyika

227 Diatom taxa were observed in the surface sediments of the northern part of Lake Tanganyika, including 1 new to science: Amphora tanganyikae. The diatom community of these sediments is mainly composed of benthic organisms while planktonic diatoms are rather rare. Many brackish-water and a few marine organisms were observed. Cosmopolitan organisms (77.1%) dominate the diatom flora but tropical, tropical African and African taxa are also well represented (22.9%)Deceased.Deceased.     

Abundance,distribution, and territory areas of rock-dwelling Lake Tanganyika cichlid fish species

Lake Tanganyika, the second-oldest and second-deepest lake in the world, harbors an impressive cichlid fish fauna counting about 250 endemic species that are characterized by a great level of ecological, morphological, and behavioral specialization. This study describes and compares cichlid fish communities at two rocky shores with differential human impact in the south of Lake Tanganyika. Species inventories and depth-dependent abundances were elaborated. About 41 and 46 sympatric cichlid species were recorded in the two study sites, respectively. Variabilichromis moorii was the most abundant species (29–60% of total number of fishes), followed by Aulonocranus dewindti (3–19%), Tropheus moorii (12%), Ophthalmotilapia ventralis (4–10%), Eretmodus cyanostictus (6–11%), and Cyathopharynx furcifer (0.01–9%). All other species had abundances below 5%. It further emerged that large cichlids such as Petrochromis species, Cyathopharynx furcifer, and Lobochilotes labiatus were very rare at one location, with frequencies of 0.55% or less. Territorial sizes of three particularly abundant species, Variabilichromis moorii, Aulonocranus dewindti, and Tropheus moorii, were assessed by behavioral observations. We distinguished between territorial core areas and total defended area, yielding average core areas between 0.4 (V. moorii) and 1.6 m² (T. moorii), and total defended areas averaging for each species between 1.6 (V. moorii) and 5.0 m² (A. dewindti) with no significant differences between the two study sites. The data on individual densities are also relevant for evolutionary studies, in that they allow more accurate calculations of effective population sizes.     

Population changes within a group of fish species in Lake Tanganyika following their exploitation

Pelagic fish populations in Lake Tanganyika consist mainly of two small clupeid species and four centroponiid species which prey on them. Exploitation with purse-seines began in the southeast arm in 1962 and catches were sampled up to early 1969. Inshore clupeid populations were also sampled with a scoop-net (lusenga of the kind used in the traditional inshore fishery.
Clupeid biomass reached annual maxima around September. One clupeid species, Stolothrissa tanganicae Regan, dominated pelagic catches except in 2 years when about equal quantities of the other clupeid, Limnothrissa miodon Blgr. were caught. Numbers of each predator (three species of Lates and Luciolates stappersii Blgr.) decreased from 1963 to 1966 and remained low thereafter. The clupeid catch rose from 1964 to 1967 and remained high in 1968. The average nightly catch weight per year of all species together altered relatively little. Changes in population size distributions occurred.
Sampling methods were concluded to be valid for clupeid populations, but probably indi-cated only general changes in predator populations. Clupeid life-cycles are mostly accomplished within a year, and appear closely related to the periodic and spatial variations in plankton production. The two species are competitive and can replace one another in the pelagic zone. Certain distribution patterns of the clupeids and of the predator young are believed to be adaptations to severe predation. Low replacement rates under fishing pressure account for the decline of predator species, and the clupeid increase resulted from reduction in predation. A fairly stable exploited phase has apparently been reached in which clupeid biomass is much greater and predator biomass much less than in the natural state. The data encourage certain predictions. Similar major trends occurred in the much larger fishery in the Burundi sector of the lake.     

Drivers of phytoplankton diversity in Lake Tanganyika

In keeping with the theme of this volume, the present article commemorates the 50 years of Hutchinson’s (Am Nat 93:145–159, 1959) famous publication on the ‘very general question of animal diversity’, which obviously leads to the more important question regarding the driving forces of biodiversity and their limitation in various habitats. The study of phytoplankton in large lakes is a challenging task which requires the use of a wide variety of techniques to capture the range of spatial and temporal variations. The analysis of marker pigments may provide an adequate tool for phytoplankton surveys in large water bodies, thanks to automated analysis for processing numerous individual samples, and by achieving sufficient taxonomic resolution for ecological studies. Chlorophylls and carotenoids were analysed by HPLC in water column samples of Lake Tanganyika from 2002 through 2006, at two study sites, off Kigoma (north basin) and off Mpulungu (south basin). Using the CHEMTAX software for calculating contributions of the main algal groups to chlorophyll a, variations of phytoplankton composition and biomass were determined. We also investigated selected samples according to standard taxonomic techniques for elucidating the dominant species composition. Most of the phytoplankton biomass was located in the 0–40 m layer, with maxima at 0 or 20 m, and more rarely at 40 m. Deep chlorophyll maxima (DCM) and surface ‘blooms’ were occasionally observed. The phytoplankton assemblage was essentially dominated by chlorophytes and cyanobacteria, with diatoms developing mainly in the dry season. The dominant cyanobacteria were very small unicells (mostly Synechococcus), which were much more abundant in the southern basin, whereas green algae dominated on average at the northern site. A canonical correspondence analysis (CCA) including the main limnological variables, dissolved nutrients and zooplankton abundance was run to explore environment–phytoplankton relations. The CCA points to physical factors, site and season as key determinants of the phytoplankton assemblage, but also indicates a significant role, depending on the studied site, of calanoid copepods and of nauplii stages. Our data suggest that the factors allowing coexistence of several phytoplankton taxa in the pelagic zone of Lake Tanganyika are likely differential vertical distribution in the water column, which allows spatial partitioning of light and nutrients, and temporal variability (occurring at time scales preventing long-term dominance by a single taxon), along with effects of predation by grazers.     

Endemic diversification of the monophyletic cottoid fish species flock in Lake Baikal explored with mtDNA sequencing

In the ancient Lake Baikal in East Siberia, cottoid fishes have diversified into an endemic flock of 33 species. From an ancestral shallow-water, benthic life-style, Baikalian cottoids have shifted to deep-water life in environments even below 1500 m, and also colonized the pelagic habitat. We examined phylogenetic relationships among 22 Baikalian and 10 extra-Baikalian cottoid taxa using a total of 2822 bp of mitochondrial DNA sequence, from complete sequences of ATPase 8 and 6 and cytochrome b genes and the control region. Unlike in earlier studies, we found strong support for a monophyly of the whole endemic Baikalian cottoid diversity. The Baikalian clade, currently assigned to three families and 12 genera, appears to be nested within the Holarctic freshwater genus Cottus. In the molecular phylogeny, all but one of the current Baikalian genera formed well-supported monophyletic groups. However, the topology was inconsistent with the present morphology-based familial subdivision; particularly in positioning the genus Batrachocottus of Cottidae within Abyssocottidae. The branching order of the Baikalian genera could not be resolved completely, however; short basal branches indicate rapid diversification early in the history of the species flock. Using synonymous divergence rates from other fish species for calibration, the diversification of the Baikalian cottoids seems to have started in the Pliocene or early Pleistocene.     

Intraspecific brood-mixing of the cichlid fishPerissodus microlepis in Lake Tanganyika

Synopsis The frequency and origin of intraspecific brood-mixing in the biparental cichlid fishPerissodus microlepis were investigated by the cohort analysis of schooling young and the underwater observation of guarding parents. The cohort analysis showed that brood-mixing started from the early guarding state when the young were smaller than 10 mm standard length and nearly all schools of young larger than 16 mm contained alien young from up to 6 broods. Brood farming-out of this fish, which was originally proposed to be a way adopted only by a deserted parent, was performed also by paired parents. We suggest that brood-mixing inP. microlepis is attributed mainly to brood farming-out by paired and deserted parents.     

Ancient lakes as evolutionary reservoirs: evidence from the thalassoid gastropods of Lake Tanganyika

Ancient lakes are often collectively viewed as evolutionary hot spots of diversification. East Africa's Lake Tanganyika has long been the subject of scientific interest owing to dramatic levels of endemism in species as diverse as cichlid fishes, paludomid gastropods, decapod and ostracod crustaceans and poriferans. It is the largest and deepest of the African rift lakes, and its endemic fauna has been presented with a stable inland environment for over 10 Myr, offering unique opportunities for within-lake diversification. Although astonishing diversification has been documented in the endemic cichlid fauna of the lake, similar patterns of rapid diversification have long been assumed for other groups. In contrast to this hypothesis of rapid speciation, we show here that there has been no acceleration in the rate of speciation in the thalassoid gastropods of the lake following lake colonization. While limited within-lake speciation has occurred, the dramatic conchological diversity of gastropods presently found within the lake has evolved from at least four major lineages that pre-date its formation by as much as 40 Myr. At the same time, a widespread group of African gastropods appears to have evolved from taxa presently found in the lake. While Lake Tanganyika has been a cradle of speciation for cichlid fishes, it has also been an important evolutionary reservoir of gastropod lineages that have been extirpated outside the basin.