Body shape vs. colour associated initial divergence in the Telmatherina radiation in Lake Matano, Sulawesi, Indonesia

Highly polymorphic colouration patterns are often associated with sexual selection in fish and can be the initial cause of divergence among closely related taxa. Here we use genetic, body colour and geometric morphometric data collected on 118 fish from Lake Matano, Sulawesi, Indonesia to test if colouration is the initial cause of divergence in the radiating Telmatherina genus. Results reveal that all Telmatherina previously described in this system can be categorized into three mitochondrial lineages and that colouration is only weakly associated with early divergence. Clade-specific body shapes, however, likely adapted to microenvironments are key to the initial divergence in this system. Data also show that although colourations were not likely instrumental in seeding divergence in these fish, they appear to have developed in parallel within each clade. Our results are consistent with an emerging pattern repeated in many vertebrate radiations, whereby divergence by colouration or other display traits is preceded by specialization to environmental adaptive peaks.     

Genetic and morphological data supporting the hypothesis of adaptive radiation in the endemic fish of Lake Matano

Adaptive radiation resulting from differential selection acting on functional features is believed to be an important source of biodiversity. In this study, morphometric measures and mitochondrial DNA are used to test for adaptive radiation within four fish genera (Glossogobius, Oryzias, Dermogenys and Telmatherina) endemic to an ancient island lake (Lake Matano, Sulawesi, Indonesia), using the framework proposed by Schluter (The Ecology of Adaptive Radiation, Oxford University Press, 2000). We demonstrate common ancestry and rapid divergence in one genus (Telmatherina) based on 560 bp of 16S sequence data. We found higher levels of variation in feeding-related traits (N = 8) for Telmatherina relative to the other genera, while no differences were found for sexual display traits (N = 8) or neutral morphological traits (N = 8). Telmatherina also had the highest number of distinct colouration patterns among the four genera. These data, combined with the very low productivity of the lake, are indicative of selection driving adaptive radiation. The morphometric divergence in the Telmatherina likely results from selection acting on feeding traits in this low productivity lake, leading to trophic specialization among closely related morphotypes. These results provide indirect but compelling data supporting the adaptive radiation of Telmatherina in this system.     

Dispersal and population genetic structure of Telmatherina antoniae,an endemic freshwater Sailfin silverside from Sulawesi,Indonesia

Population genetic structure in the presence of substantial dispersal provides a unique perspective on the evolution of reproductive isolation. We sampled Telmatherina antoniae, an endemic fish species, at 10 sites in Lake Matano, Indonesia. Significant genetic structure (F_ST = 0.03) was found, despite a migration rate of 10.2% and a mean dispersal distance of 13.6 km, estimated by genotype assignment. Neither dispersal distance nor direction differed from random expectations, indicative of no dispersal barrier in Lake Matano. However, Bayesian genotype cluster assignment identified a population structure consisting of four to six clusters that did not coincide with sample site distribution, but explained two to three times more genetic variance than sample site. The mechanism for continued isolation of those genetic clusters is unknown, but assortative mating and temporal isolation are obvious candidates. Our results resolve the apparent paradox of population genetic structure coupled with frequent dispersal, and highlight the importance of considering cryptic genetic structure.     

The plankton community of Lake Matano: factors regulating plankton composition and relative abundance in an ancient,tropical lake of Indonesia

Recent evidence reveals that food webs within the Malili Lakes, Sulawesi, Indonesia, support community assemblages that are made up primarily of endemic species. It has been suggested that many of the species radiations, as well as the paucity of cosmopolitan species in the lakes, are related to resource limitation. In order to substantiate the possibility that resource limitation is playing such an important role, a study of the phytoplankton and zooplankton communities of Lake Matano was implemented between 2000 and 2004. We determined species diversity, relative abundances, size ranges, and total biomass for the phytoplankton and zooplankton, including the distribution of ovigerous individuals throughout the epilimnion of Lake Matano in three field seasons. The phytoplankton community exhibited very low biomass (<15 μg l^?1) and species richness was depressed. The zooplankton assemblage was also limited in biomass (2.5 mg l^?1) and consisted only of three taxa including the endemic calanoid Eodiaptomus wolterecki var. matanensis, the endemic cyclopoid, Tropocyclops matanensis and the rotifer Horaella brehmi. Zooplankton were very small (<600 μm body length), and spatial habitat partitioning was observed, with Tropocylops being confined to below 80 m, while rotifer and calanoid species were consistently observed above 80 m. Less than 0.1% of the calanoid copepods in each year were egg-bearing, suggesting very low population turnover rates. It was concluded that chemical factors as opposed to physical or biological processes were regulating the observed very low standing crops of phytoplankton which in turn supports a very minimal zooplankton community restricted in both species composition and abundance. As chemical factors are a function of the catchment basin of Lake Matano, it is predicted that resource limitation has long played an important role in shaping the unique endemic assemblages currently observed in the food web of the lake.     

Evolution of the tribe Tropheini from Lake Tanganyika: synchronized explosive speciation producing multiple evolutionary parallelism

One of the most surprising outcomes of recent molecular studies on cichlid fishes of the three Great East African Lakes Victoria, Malawi and Tanganyika, was the stunning rapidity of speciation and cladogenesis at early stages of adaptive radiation. Despite their rapid pace, speciation events were so far intuitively assumed to proceed in a bifurcating and tree-like fashion, even if they could not be resolved by gene phylogenies due to a lack of resolution. On the basis of phylogenetic analyses of the Tropheini, a lineage of endemic rock-dwelling cichlid fishes from Lake Tanganyika, we suggest a pathway of explosive speciation that accounts for a non-bifurcating manner of cladogenesis. This pattern is likely to be the result of the contemporaneous origin of a multitude of founder populations in geographically isolated rock habitats among which gene flow was interrupted simultaneously by a major change of the lake habitat in the form of a rapid rise of the lake level. As a consequence, all new species arising from that vicariance event must exhibit almost equal genetic distances to each other, within the scope of genetic diversity of the founder population(s), even if the actual processes of subsequent speciation and eco-morphological diversification followed independent routes. Our phylogeny also suggests a high frequency of parallel evolution of equivalent trophic specialization in the Tropheini. This phenomenon seems to be an inherent feature of this pathway of speciation, due to the action of similar selective forces on the same set of species colonizing isolated habitats of the same type. Explosive speciation via synchronization of genetic divergence triggered by rapid environmental changes seems to be particularly likely to occur at advanced stages of adaptive radiation, when species are already adapted to particular habitats and have a reduced ability for dispersal.     

Changes in fish species composition in Lake Naivasha,Kenya

The fish species composition in Lake Naivasha has been modified during recent years. Out of seven introduced and one indigenous species reported in 1962, only three introduced ones are found today. The elimination of species has been brought about by predation and changing ecological conditions. The most common fish species today is Sarotherodon leucosticuus. The hybridization found by previous workers between S. leucostictus x S. spilurus niger has stopped, as S. s. niger is no longer found in the lake. Most of the hybrid characters have been lost by the few surviving hybrids due to backcrossing between hybrids and between hybrids and S. leucostictus. The hybrids now mostly resemble S. leucostictus.     

Genetic divergence in morphology-performance mapping between Misty Lake and inlet stickleback

Different environments should select for different aspects of organismal performance, which should lead to correlated divergence in morphological traits that influence performance. The result should be genetic divergence in aspects of performance, morphology and associations ('maps') between morphology and performance. Testing this hypothesis requires quantifying performance and morphology in multiple populations after controlling for environmental differences, but this is rarely attempted. We used a common-garden experiment to examine morphology and several aspects of swimming performance within and between the lake and inlet populations of threespine stickleback (Gasterosteus aculeatus) from the Misty system, Vancouver Island, Canada. Controlling for body size, lake stickleback had shallower bodies, larger caudal fins and smaller pelvic girdles. With or without morphological covariates, lake stickleback showed greater performance in both sustained and burst swimming. In contrast, inlet stickleback showed greater manoeuverability than did lake stickleback in some analyses. Morphology-performance relationships were decoupled when considering variation within vs. between populations. Moreover, morphology-performance mapping differed between the two populations. Based on these observations, we advance a hypothesis for why populations adapting to different environments should show adaptive genetic divergence in morphology-performance mapping.     

Organochlorine residues in fish from Lake Victoria, Kenya

Eighty-two samples of either nile perch fish fat or muscle were collected from the Kenyan region of Lake Victoria for detection of organochlorine residues. Nine organochlorine residues were detected in the following percentages: α-BHCIHCB-40%; P-BHC/HCB-40%; y-BHC/HCB/lindane-4%; aldrin-9%; dieldrin-1%; p, p'-DDE-73%; p, p'-DDD-9%; o, p'-DDT-170; and p, p'-DDT-11%. All levels of organochlorine residues were below the Extraneous Residue Limit (ERL), apart from just one sample of fish fat which had 4.51 ppm of sum DDT above ERL. There was a higher level of organochlorines from Mbita than from Luanda fishing point.

Résumé

Eighty-two échantillons de graisse ou de muscle de perche du Nil ont été récoltés dans la région kényane du lac Victoria pour rechercher des résidus organochlorés. On a détecté 9 résidus organochlorés, avec les pourcentages suivants: α-BHC/HCB-40%; β-BHC/HCB-40%;γ-BHC/HCB/lindane-4%;aldrine-9%; dieldrine-1%; p, p'-DDE-73%; p, p'-DDD-9%; o, p'-DDT-1%; and p, p'-DDT-11%. Tous les taux de résidus organochlorés ktaient infkrieurs a la 'Extraneous Residue Limit' (ERL) sauf pour un seul échantillon de graisse de poisson qui contenait 4,51 ppm de DDT total au-dessus de 1'ERL. I1 y avait un taux plus élevé d'organochlorés a Mbita qu'au point de pêche de Luanda.     

Adaptive speciation and sexual dimorphism contribute to diversity in form and function in the adaptive radiation of Lake Matano's sympatric roundfin sailfin silversides

The utility of traits involved in resource exploitation is a central criterion for the adaptive character of radiations. Here, we test for differentiation in morphology, jaw mechanics and nutrition among species and sexes of Lake Matano's sympatric 'roundfin' sailfin silversides. The three incipient fish species differ significant in several candidate traits for adaptation following ecological selection pressure, corresponding to contrasting jaw mechanics and distinct patterns in food resource use. These findings are consistent with functional adaptation and suggest divergence following alternative modes of feeding specialization. Further, intersexual resource partitioning and corresponding adaptation in jaw mechanics is evident in two of the three incipient species, demonstrating that sexual dimorphism contributes to the ecomorphological and trophic diversity of the emerging radiation. This is, to the best of our knowledge, the first study reporting interspecific as well as intersexual adaptation by alternative modes of form and function in an evolving fish species flock.     

Testing the stages model in the adaptive radiation of cichlid fishes in East African Lake Tanganyika

Adaptive radiation (AR) is a key process in the origin of organismal diversity. However, the evolution of trait disparity in connection with ecological specialization is still poorly understood. Available models for vertebrate ARs predict that diversification occurs in the form of temporal stages driven by different selective forces. Here, we investigate the AR of cichlid fishes in East African Lake Tanganyika and use macroevolutionary model fitting to evaluate whether diversification happened in temporal stages. Six trait complexes, for which we also provide evidence of their adaptiveness, are analysed with comparative methods: body shape, pharyngeal jaw shape, gill raker traits, gut length, brain weight and body coloration. Overall, we do not find strong evidence for the ‘stages model’ of AR. However, our results suggest that trophic traits diversify earlier than traits implicated in macrohabitat adaptation and that sexual communication traits (i.e. coloration) diversify late in the radiation.     

Genetic and ecological divergence of a monophyletic cichlid species pair under fully sympatric conditions in Lake Ejagham, Cameroon

Although there is mounting evidence that speciation can occur under sympatric conditions, unambiguous examples from nature are rare and it is almost always possible to propose alternative allopatric or parapatric scenarios. To identify an unequivocal case of sympatric speciation it is, therefore, necessary to analyse natural settings where recent monophyletic species flocks have evolved within a small and confined spatial range. We have studied such a case with a cichlid species flock that comprises five Tilapia forms endemic to a tiny lake (Lake Ejagham with a surface area of approximately 0.49 km2) in Western Cameroon. Analysis of mitochondrial D-Loop sequences shows that the flock is very young (approximately 10(4) years) and has originated from an adjacent riverine founder population. We have focused our study on a particular pair of forms within the lake that currently appears to be in the process of speciation. This pair is characterized by an unique breeding colouration and specific morphological aspects, which can serve as synapomorphic characters to prove monophyly. It has differentiated into a large inshore and a small pelagic form, apparently as a response to differential utilization of food resources. Still, breeding and brood care occurs in overlapping areas, both in time and space. Analysis of nuclear gene flow on the basis of microsatellite polymorphisms shows a highly restricted gene flow between the forms, suggesting reproductive isolation between them. This reproductive isolation is apparently achieved by size assortative mating, although occasional mixed pairs can be observed. Our findings are congruent with recent theoretical models for sympatric speciation, which show that differential ecological adaptations in combination with assortative mating could easily lead to speciation in sympatry.     

Sixty years of environmental change in the world's largest freshwater lake – Lake Baikal, Siberia

High-resolution data collected over the past 60 years by a single family of Siberian scientists on Lake Baikal reveal significant warming of surface waters and long-term changes in the basal food web of the world's largest, most ancient lake. Attaining depths over 1.6 km, Lake Baikal is the deepest and most voluminous of the world's great lakes. Increases in average water temperature (1.21 °C since 1946), chlorophyll a (300% since 1979), and an influential group of zooplankton grazers (335% increase in cladocerans since 1946) may have important implications for nutrient cycling and food web dynamics. Results from multivariate autoregressive (MAR) modeling suggest that cladocerans increased strongly in response to temperature but not to algal biomass, and cladocerans depressed some algal resources without observable fertilization effects. Changes in Lake Baikal are particularly significant as an integrated signal of long-term regional warming, because this lake is expected to be among those most resistant to climate change due to its tremendous volume. These findings highlight the importance of accessible, long-term monitoring data for understanding ecosystem response to large-scale stressors such as climate change.     

Post-impoundment changes in the fish fauna of Lake Itezhi-tezhi, Zambia

The fish fauna of the newly inundated Lake Itezhi-tezhi, Zambia was observed between 1980 and 1985. Marked changes in the community structure were identified. The most obvious were a decline in species diversity and a shift in species composition from acommunity with a preponderance of Alesles lateralis (Boulenger) (Characidae), to one dominated by cichlids. These changes were primarily due to members of the families Cyprinidae, Mormyridae and Schilbeidae being unable to adapt to the new environmental conditions, such as spawning and feeding grounds, and their inability to cope with fluctuating water levels in the lake. It was suggested that a sound fishery data collection system be implemented to provide reliable information on which to base management policy.     

Molecular phylogeny and diversification of freshwater shrimps (Decapoda, Atyidae, Caridina) from ancient Lake Poso (Sulawesi, Indonesia)--the importance of being colourful

Ancient Lake Poso on the Indonesian island Sulawesi hosts a highly diverse endemic fauna, including a small species flock of atyid Caridina shrimps, which are characterized by conspicuous colour patterns. We used a mtDNA based molecular phylogeny to test the assumption of a monophyletic origin and intralacustrine radiation of the species flock and to assess the species specificity of some colour morphs. Our data reveal a rapid radiation of Caridina in the entire Poso drainage system, but provide no strong evidence for a monophyletic radiation of the lake species. Nevertheless each lacustrine species shows a varying degree of substrate or trophic specialization, usually considered a hallmark of adaptive radiation. Two distinct colour forms previously attributed to a single species, C. ensifera, lack distinguishing qualitative morphological characters, but are shown to be two different species. In contrast, morphologically rather distinct lake species lacking specific colour patterns may be hybridizing with riverine taxa. These results suggest that colour may play a similar role in species recognition and possibly speciation in ancient lake Caridina as hypothesized, e.g. for some African cichlids.     

Phenotypic and genetic divergence among harbour porpoise populations associated with habitat regions in the North Sea and adjacent seas

Determining the mechanisms that generate population structure is essential to the understanding of speciation and the evolution of biodiversity. Here, we investigate a geographical range that transects two habitat gradients, the North Sea to North Atlantic transition, and the temperate to subpolar regions. We studied the harbour porpoise (Phocoena phocoena), a small odontocete inhabiting both subpolar and temperate waters. To assess differentiation among putative populations, we measured morphological variation at cranial traits (N = 462 individuals) and variation at eight microsatellite loci for 338 of the same individuals from Norwegian, British and Danish waters. Significant morphological differentiation reflected the size of the buccal cavity. Porpoises forage in relatively shallow waters preying mainly on benthic species in British and Danish waters, and on mesopelagic and pelagic fish off the coast of Norway. We suggest that the observed differentiation may be explained by resource specialization and either adaptation or developmental responses to different local habitats.     

Acoustic assessment of the number of pelagic fish in Lake Kinneret,Israel

The pelagic fish stock (mostly lavnun, Mirogrex terraesanctae) of Lake Kinneret was assessed using HADAS, a newly developed echo-counting analysis system. Total fish numbers (during Mar–Apr) were estimated to be 61 million in 1987, 218 million in 1988, 120 million in 1989, and 69 million in April 1990. The total fish number for March 1981 was recalculated from published data and estimated at 170 million. The decline in total numbers between 1988 and 1990 resulted primarily from a decrease in the relative abundance of the smallest size classes of fish. Since fish abundance apparently declined between 1981 and 1990, the decline in zooplankton standing stocks documented during these years cannot be attributed to higher fish predation caused by larger fish stocks. The HADAS estimates of lavnun population size are large enough to account for the known catch of lavnun.Contribution from Israel Oceanographic & Limnological Research, Ltd. This research was supported in part by the Israeli Water Commissioner, the Israeli Electric Company, and by a grant from the U.S. Agency for International Development.