Distinct population structure in a phenotypically homogeneous rock-dwelling cichlid fish from Lake Tanganyika

Several lineages of cichlid fishes in the East African Great Lakes display stunning levels of morphological diversification. The rapid evolution of rock-dwelling polygynous mouthbrooders in Lake Malawi, for example, was in part ascribed to their allopatric distribution on disjunct stretches of rocky coast, where even short habitat discontinuities reduce gene flow effectively. However, as seen in other cichlids, ecological barriers do not always prevent gene flow, whereas genetic structure can develop along continuous habitat, and morphological diversification does not necessarily accompany genetic differentiation. The present study investigates the population structure of Variabilichromis moorii, a monogamous substrate-brooding lamprologine of rocky coasts in Lake Tanganyika, which occurs over about 1000 km of shoreline almost without phenotypic variation. Phylogeographic analyses of mitochondrial DNA sequences indicated that dispersal is infrequent and generally occurs between adjacent locations only. Exceptions to this pattern are closely related haplotypes from certain locations on opposite lakeshores, a phenomenon which has been observed in other species and is thought to reflect lake crossing along an underwater ridge in times of low water level. Genetic population differentiation, estimated from mitochondrial DNA and microsatellite data in six adjacent populations, was equally high across localities separated by sandy shores and along uninterrupted stretches of rocky shore. Our results suggest that ecological barriers are not required to induce philopatric behavior in Variabilichromis, and that morphological stasis persists in the face of high levels of neutral genetic differentiation.     

Neolamprologus cancellatus, a new cichlid fish from Lake Tanganyika, Africa

Neolamprologus cancellatus, a new cichlid species, is described on the basis of eight specimens from the Zambian coast of Lake Tanganyika. The new species is characterized by a subtruncate caudal fin and a body that is slender (depth 22.3–25.2% in standard length) and easily distinguishable from its congeners by having 7–8 anal fin spines, 34–37 scales in longitudinal line, 33 total vertebrae, and a gridlike body pattern on a pale brownish body color. This species is only known to inhabit the shallow (2–7 m depth), rocky bottom of Wonzye Point in the southern part of the lake.     

Geometric morphometrics applied to viscerocranial bones in three populations of the Lake Tanganyika cichlid fish Tropheus moorii

Lake Tanganyika comprises the oldest and most diverse species flock of cichlid fishes. Many species are subdivided into numerous populations, often classified as geographical races, colour morphs or sister species. Unlike younger species flocks, in which speciation is accompanied by eco-morphological diversification that of Lake Tanganyika is at a mature stage with little further morphological change, most probably caused by stabilizing selection. This study addresses body shape differences among three genetically distinct but morphologically similar populations of Tropheus  moorii , by focusing on bony structures of the cichlid head. We test by means of geometric morphological methods whether shape changes in the cichlid head are based on particular osteological differences. Specimens were disarticulated enzymatically, and standardized digital images of the disarticulated bony elements were taken. A landmark system was established for the dentary, articular, premaxilla, quadrate and the preopercle. Only the dentary shows significant differentiation among the three populations. Since all three populations live in similar cobble habitats and occupy the same trophic niche, the observed difference in the shape of the dentary can either be explained by different directional selection due to subtle habitat differences, or by neutral drift constrained by borders enforced through stabilizing selection. Lack of difference might indicate stabilizing selection on bone shape.     

Microdontochromis rotundiventralis, a new cichlid fish (perciformes: Cichlidae) from Lake Tanganyika

Microdontochromis rotundiventralis, a new species of cichlid fish, is described on the basis of 13 specimens from Nkumbula Island, Lake Tanganyika (Zambia). It is distinct from its only congener,M. tenuidentatus, in having two (rarely one) rows of teeth on both jaws, a rounded distal margin on the pelvic fin, the outermost pelvic fin soft ray length 1.23–1.43 times the innermost ray, a deeper body (depth 26.8–29.1% standard length) and the anal fin with 9 (rarely 8 or 10) soft rays.     

Description of a new cichlid fish species of the genus Benthochromis (Perciformes: Cichlidae) from Lake Tanganyika

Benthochromis horii , a new cichlid species is described based on 19 type specimens from the deep waters of Lake Tanganyika. It differs from its congeners by having smaller eyes and longer snout (eye length usually shorter than snout length v. equal to or longer than snout length in Benthochromis tricoti and Benthochromis melanoides ), and more dorsal fin rays (total number of spines and soft rays in dorsal fin usually 30 or 31 v. usually 28 or 29 in B. tricoti and B. melanoides ). Large males of the new species differ from those of congeneric species in terms of their body colour pattern and long pelvic fins. This species has been confused with B. tricoti and has been traded as an aquarium fish.     

Validation of daily increment formation in otoliths of juvenile and adult European anchovy

The otoliths of juveniles and adults of European anchovy Engraulis encrasicolus held in aquaria were marked by immersion in oxytetracycline hydrochloride (OTC) at concentrations between 350 and 410 mg l⁻¹ for 12 h. Counts of microincrements between fluorescent bands validated the daily otolith increment formation. The otolith increments were easily readable at ×400 with average increment widths of c . 1·1 µm. Validation was successfully demonstrated in juveniles and adults maintained for short periods in the aquaria in the summer. For European anchovy captured as juvenile and reared to adults, however, increment formation appeared less than daily. The daily periodicity of the otoliths in juvenile European anchovy implies that counting of microincrements can be used to study their birth dates. The application of this technique to adults, however, may lead to the underestimation of actual age and further research needs to be done to clarify the reasons for the apparent loss of the daily rhythm over long periods.     

A revised synonymy of Telmatochromis temporalis (Teleostei, Cichlidae) from Lake Tanganyika (East Africa)

The taxonomic status of the nominal species Telmatochromis temporalis , T. lestradei , T. burgeoni and Julidochromis macrolepis has been reviewed. The synonymy of T. lestradei with T. temporalis is confirmed. A comparison of Telmatochromis burgeoni with Telmatochromis temporalis revealed no significant differences either. Hence T. burgeoni is considered synonymous with T. temporalis . Examination of the type of Julidochromis macrolepis showed it to be conspecific with Telmatochromis dhonti and not with T. temporalis , as had been suggested.     

Validation of daily increment formation in otoliths for three Diplodus species in the Mediterranean sea

The efficacy of two fluorochromes, chlorhydrate of tetracycline (CHTC) and alizarin complexone (ALC), to induce a label on the otoliths of juvenile sparid fishes by immersion techniques was tested in different experimental conditions. CHTC did not mark otoliths in natural sea water and was toxic in divalent cation-free sea water. Immersing fish in a 50-mg 1⁻¹ ALC natural seawater solution for 24 h induced a well-defined mark on the otoliths and had no effect on survival. A daily periodicity of increment formation on otoliths was observed in captivity for Diplodus vulgaris and D. puntazzo , and was conserved in natural environment for D. sargus . The frequency of increment deposition did not vary with the age of juvenile fishes. Thus, otolith microstructure analysis will be a reliable method to give age estimates in these three sparid species during their juvenile life.     

Colour-assortative mating among populations of Tropheus moorii, a cichlid fish from Lake Tanganyika, East Africa

The species flocks of cichlid fishes in the East African Lakes Tanganyika, Malawi and Victoria are prime examples of adaptive radiation and explosive speciation. Several hundreds of endemic species have evolved in each of the lakes over the past several thousands to a few millions years. Sexual selection via colour-assortative mating has often been proposed as a probable causal factor for initiating and maintaining reproductive isolation. Here, we report the consequences of human-mediated admixis among differentially coloured populations of the endemic cichlid fish Tropheus moorii from several localities that have accidentally been put in sympatry in a small harbour bay in the very south of Lake Tanganyika. We analysed the phenotypes (coloration) and genotypes (mitochondrial control region and five microsatellite loci) of almost 500 individuals, sampled over 3 consecutive years. Maximum-likelihood-based parenthood analyses and Bayesian inference of population structure revealed that significantly more juveniles are the product of within-colour-morph matings than could be expected under the assumption of random mating. Our results clearly indicate a marked degree of assortative mating with respect to the different colour morphs. Therefore, we postulate that sexual selection based on social interactions and female mate choice has played an important role in the formation and maintenance of the different colour morphs in Tropheus, and is probably common in other maternally mouthbrooding cichlids as well.     

Male courtship preferences demonstrate discrimination against allopatric colour morphs in a cichlid fish

Whether premating isolation is achieved by male‐specific, female‐specific or sex‐independent assortative preferences often depends on the underlying evolutionary processes. Here we test mate preferences of males presented with females of different allopatric colour variants of the cichlid fish Tropheus sp., a Lake Tanganyika endemic with rich geographical colour pattern variation, in which the strength of sexual isolation varies between populations. We conducted two‐way mate choice experiments to compare behaviour of males of a red‐bodied morph (population Moliro) towards females from their own population with behaviour towards females from four allopatric populations at different stages of phylogenetic and phenotypic divergence. Males courted same‐population females significantly more intensely than females of other populations, and reduced their heteromorphic courtship efforts both with increasing genetic and increasing phenotypic distinctness of the females. In particular, females of a closely related red‐bodied population received significantly more courtship than either genetically distinct, similarly coloured females (‘Kirschfleck’ morph) or genetically related, differently coloured females (‘yellow‐blotch’ morph), both of which were courted similarly. Genetically and phenotypically distinct females (Tropheus polli) were not courted at all. Consistent with previous female‐choice experiments, female courtship activity also decreased with increasing genetic distance from the males’ population. Given successful experimental and natural introgression between colour morphs and the pervasive allopatry of related variants, we consider it unlikely that assortative preferences of both sexes were driven by direct selection during periods of secondary contact or, in turn, drove colour pattern differentiation in allopatry. Rather, we suggest that sexual isolation evolved as by‐product of allopatric divergence.     

Obligate feeding of cichlid eggs by Caecomastacembelus zebratus in Lake Tanganyika

This study examined food habits of four common species of spiny eels (Mastacembelidae) in Lake Tanganyika to find if they eat cichlid eggs and larvae. Diets of Caecomastacembelus zebratus consisted nearly exclusively of eggs of substratum-brooding cichlids, whereas two other species fed on eggs only facultatively and one species never did. More than half individuals of C. zebratus had empty guts, although the mean weight of their gut contents was highest. This spiny eel ate many eggs in the first and second quarters of the lunar cycle when the spawning of substratum-brooding cichlids was active, while it nearly ceased eating in the latter half of the cycle. It is suggested that this fish is an obligate egg-eater and one of the predators that cause the high mortality of early cichlid broods.     

AFLP genome scans suggest divergent selection on colour patterning in allopatric colour morphs of a cichlid fish

Genome scan-based tests for selection are directly applicable to natural populations to study the genetic and evolutionary mechanisms behind phenotypic differentiation. We conducted AFLP genome scans in three distinct geographic colour morphs of the cichlid fish Tropheus moorii to assess whether the extant, allopatric colour pattern differentiation can be explained by drift and to identify markers mapping to genomic regions possibly involved in colour patterning. The tested morphs occupy adjacent shore sections in southern Lake Tanganyika and are separated from each other by major habitat barriers. The genome scans revealed significant genetic structure between morphs, but a very low proportion of loci fixed for alternative AFLP alleles in different morphs. This high level of polymorphism within morphs suggested that colour pattern differentiation did not result exclusively from neutral processes. Outlier detection methods identified six loci with excess differentiation in the comparison between a bluish and a yellow-blotch morph and five different outlier loci in comparisons of each of these morphs with a red morph. As population expansions and the genetic structure of Tropheus make the outlier approach prone to false-positive signals of selection, we examined the correlation between outlier locus alleles and colour phenotypes in a genetic and phenotypic cline between two morphs. Distributions of allele frequencies at one outlier locus were indeed consistent with linkage to a colour locus. Despite the challenges posed by population structure and demography, our results encourage the cautious application of genome scans to studies of divergent selection in subdivided and recently expanded populations.     

Community assembly in Lake Tanganyika cichlid fish: quantifying the contributions of both niche‐based and neutral processes

The cichlid family features some of the most spectacular examples of adaptive radiation. Evolutionary studies have highlighted the importance of both trophic adaptation and sexual selection in cichlid speciation. However, it is poorly understood what processes drive the composition and diversity of local cichlid species assemblages on relatively short, ecological timescales. Here, we investigate the relative importance of niche‐based and neutral processes in determining the composition and diversity of cichlid communities inhabiting various environmental conditions in the littoral zone of Lake Tanganyika, Zambia. We collected data on cichlid abundance, morphometrics, and local environments. We analyzed relationships between mean trait values, community composition, and environmental variation, and used a recently developed modeling technique (STEPCAM) to estimate the contributions of niche‐based and neutral processes to community assembly. Contrary to our expectations, our results show that stochastic processes, and not niche‐based processes, were responsible for the majority of cichlid community assembly. We also found that the relative importance of niche‐based and neutral processes was constant across environments. However, we found significant relationships between environmental variation, community trait means, and community composition. These relationships were caused by niche‐based processes, as they disappeared in simulated, purely neutrally assembled communities. Importantly, these results can potentially reconcile seemingly contrasting findings in the literature about the importance of either niche‐based or neutral‐based processes in community assembly, as we show that significant trait relationships can already be found in nearly (but not completely) neutrally assembled communities; that is, even a small deviation from neutrality can have major effects on community patterns.     

Separated by sand, fused by dropping water: habitat barriers and fluctuating water levels steer the evolution of rock-dwelling cichlid populations in Lake Tanganyika

The conditions of phenotypic and genetic population differentiation allow inferences about the evolution, preservation and loss of biological diversity. In Lake Tanganyika, water level fluctuations are assumed to have had a major impact on the evolution of stenotopic littoral species, though this hypothesis has not been specifically examined so far. The present study investigates whether subtly differentiated colour patterns of adjacent Tropheus moorii populations are maintained in isolation or in the face of continuous gene flow, and whether the presumed influence of water level fluctuations on lacustrine cichlids can be demonstrated in the small-scale population structure of the strictly stenotopic, littoral Tropheus. Distinct population differentiation was found even across short geographic distances and minor habitat barriers. Population splitting chronology and demographic histories comply with our expectation of old and rather stable populations on steeper sloping shore, and more recently established populations in a shallower region. Moreover, population expansions seem to coincide with lake level rises in the wake of Late Pleistocene megadroughts ~100 KYA. The imprint of hydrologic events on current population structure in the absence of ongoing gene flow suggests that phenotypic differentiation among proximate Tropheus populations evolves and persists in genetic isolation. Sporadic gene flow is effected by lake level fluctuations following climate changes and controlled by the persistence of habitat barriers during lake level changes. Since similar demographic patterns were previously reported for Lake Malawi cichlids, our data furthermore strengthen the hypothesis that major climatic events synchronized facets of cichlid evolution across the East African Great Lakes.     

Systematics of the tribe Trematocarini (Perciformes: Cichlidae) from Lake Tanganyika, Africa

 Phylogenetic relationships among eight Trematocara species and a single Telotrematocara species included in the Tanganyikan cichlid tribe Trematocarini were investigated on the basis of morphological features. The monophyly of the tribe is supported by the presence of hypertrophied sensory pores on the head, tendon “c” of adductor mandibulae section 1, a single scale row between the upper lateral line and body axis, great depth of the anteriormost infraorbital (reversed in Trematocara caparti and T. stigmaticum), and the absence of a lower lateral line. Trematocara is paraphyletic unless Telotrematocara is treated as a junior synonym. Received: December 10, 2001 / Revised: March 18, 2002 / Accepted: April 4, 2002     

Four new species of Gomphocythere (Crustacea, Ostracoda) from Lake Tanganyika, East Africa

Four new species of the ostracod genus Gomphocythere are described from Lake Tanganyika (East Africa): Gomphocythere downingi n. sp. G. coheni n. sp., G. wilsoni n. sp., and G. woutersi n. sp. All species are endemic to the lake and are found within a variety of substrates and depths. The addition of these four new species brings the total number of endemic Gomphocythere species in Lake Tanganyika to nine. Other Gomphocythere species are known from water bodies throughout East and South Africa and in the Levant. Brooding is an important, but not a unique, preadaptation for the persistence and taxic prolific speciation of this lineage in Lake Tanganyika.     

The periodicity of primary increment formation in the otoliths of Arctic charr Salvelinus alpinus (L.)

The role of environmental factors in the regulation of sub-annual increment formation in fish otoliths appears to differ markedly between species. To examine the periodicity of primary increment formation in the otoliths of O + Arctic charr, Salvelinus alpinus (L.), and the effects of temperature, and photoperiod on their formation, fish were held under controlled environmental conditions. Primary growth increments were found in the otoliths of fish held at constant temperature (18° C) and at ambient temperature [fluctuating with a circadian and circannual rhythm (4–18° C)]. Consistent and significant disruptions in increment formation occurred however, in experimental groups subjected to rapid change from ambient photoperiod to a 6L: 6D photo-period for 96 h. Disruptions in increment formation were also observed immediately following transportation of fish between holding facilities and following disease treatment. The number of otolith increments formed in fish held on an ambient photoperiod regime, correlated closely with time elapsed in days since checkmark formation ( r = 0.989, P ≤0.001) in fish sampled sequentially over a period of 10 to 105 days. Thus we demonstrate that under conditions of ambient photoperiod, primary increments are formed daily.     

Lunar synchronization of spawning in cichlid fishes of the tribe Lamprologini in Lake Tanganyika

Lunar synchronization of spawning was investigated for eight substrate brooding cichlid fishes belonging to the tribe Lamprologini. Their spawning activities all peaked during the second quarter of the lunar cycle. Comparison between their breeding styles and the degrees of spawning synchronization suggested that the nocturnal guarding-efficiency of eggs, especially exposed ones, is improved by the maximal amount of moonlight during full moon and, in species whose young leave the breeding sites about 2 weeks after spawning, the survival of dispersing young is enhanced by the darkness of moonless nights.     

In situ experiments on the effects of increased sediment loads on littoral rocky shore communities in Lake Tanganyika, East Africa

1. Two in situ experiments investigated the responses to artificially increased sediment loads of the gastropod and fish communities, and of the whole benthos, of the rocky shores of Lake Tanganyika. Sediments were emptied once onto randomly selected quadrats using SCUBA, with control quadrats receiving no addition of sediment. Quadrats were monitored up to 6 months after sediment addition. 2. While the abundance of Lavigeria grandis (Mollusca; Gastropoda) was reduced greatly by the addition of sediment, low abundance and biomass of the whole gastropod community on sediment‐impacted quadrats 6 months after sediment addition suggests more general impacts of inundation in the longer term. 3. Benthic invertebrate communities inundated by sediment had significantly fewer individuals and numbers of taxa than control communities. No significant community recovery was recorded on sediment‐impacted quadrats within the timescale of the experiment. A short‐term period of high turbidity brought about by the emptying of sediments onto sediment‐impacted quadrats also caused severe decreases in benthic invertebrate abundance on controls 10 days after sediment addition. This suggests that even short‐term pulses of sediment may have significant detrimental effects on littoral communities. 4. Although significant impacts of sediment on fish community dynamics were found, and a significant reduction in the number of benthic algivorous species was recorded immediately after sediment addition, littoral fish communities showed strong resistance overall to the addition of sediments. This apparent resistance, however, may be a consequence of the small scale at which the experiment took place. Littoral fish communities were also shown to be more dynamic than indicated previously.     

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.