Genetic isolation and morphological divergence mediated by high-energy rapids in two cichlid genera from the lower Congo rapids

Background  

It is hypothesized that one of the mechanisms promoting diversification in cichlid fishes in the African Great Lakes has been the well-documented pattern of philopatry along shoreline habitats leading to high levels of genetic isolation among populations. However lake habitats are not the only centers of cichlid biodiversity - certain African rivers also contain large numbers of narrowly endemic species. Patterns of isolation and divergence in these systems have tended to be overlooked and are not well understood.     

Speciation within genomic networks: a case study based on Steatocranus cichlids of the lower Congo rapids

Hybridization in animals is a much more common phenomenon as previously thought and may have profound implications for speciation research. The cichlid genus Steatocranus (Teleostei: Cichlidae), a close relative to members of the East African cichlid radiations, radiated under riverine conditions in the lower Congo rapids and produced a small species flock. Previous phylogenetic analyses suggested that hybridization occurred and contributed to speciation in this genus. A re-analysis of an already published 2000 loci-AFLP data set explicitly testing for patterns of ancient gene flow provided strong evidence for a highly reticulate phylogenetic history of the genus. We provide, to our knowledge, the first example of a complex reticulate network in vertebrates, including multiple closely related species connected through ancient as well as recent gene flow. In this context, the limited validity of strictly bifurcating tree hypotheses as a phylogenetic basis for hypothesis testing in evolutionary biology is discussed.     

Molecular and fossil evidence place the origin of cichlid fishes long after Gondwanan rifting

Cichlid fishes are a key model system in the study of adaptive radiation, speciation and evolutionary developmental biology. More than 1600 cichlid species inhabit freshwater and marginal marine environments across several southern landmasses. This distributional pattern, combined with parallels between cichlid phylogeny and sequences of Mesozoic continental rifting, has led to the widely accepted hypothesis that cichlids are an ancient group whose major biogeographic patterns arose from Gondwanan vicariance. Although the Early Cretaceous (ca 135 Ma) divergence of living cichlids demanded by the vicariance model now represents a key calibration for teleost molecular clocks, this putative split pre-dates the oldest cichlid fossils by nearly 90 Myr. Here, we provide independent palaeontological and relaxed-molecular-clock estimates for the time of cichlid origin that collectively reject the antiquity of the group required by the Gondwanan vicariance scenario. The distribution of cichlid fossil horizons, the age of stratigraphically consistent outgroup lineages to cichlids and relaxed-clock analysis of a DNA sequence dataset consisting of 10 nuclear genes all deliver overlapping estimates for crown cichlid origin centred on the Palaeocene (ca 65–57 Ma), substantially post-dating the tectonic fragmentation of Gondwana. Our results provide a revised macroevolutionary time scale for cichlids, imply a role for dispersal in generating the observed geographical distribution of this important model clade and add to a growing debate that questions the dominance of the vicariance paradigm of historical biogeography.     

Trophic niche segregation among herbivorous serrasalmids from rapids of the lower Xingu River,Brazilian Amazon

In the Amazon Basin, several species of herbivorous serrasalmid fishes inhabit rapids, but it is unknown if they partition food resources during the annual low-water period when fish densities are high within greatly reduced volume of aquatic habitat. We investigated the trophic ecology of juveniles and adults of three species, Myleus setiger, Ossubtus xinguense, and Tometes kranponhah, common in rapids of the Xingu River during the low-water period. Diets, stable isotope ratios of muscle tissue, and functional traits were analyzed for 59 specimens of M. setiger, 175 of O. xinguense and 215 of T. kranponhah. The three species overlapped in dietary and isotopic space, with adult O. xinguense being most divergent. Juvenile and adult T. kranponhah and juvenile O. xinguense, two groups with broad diets, had lowest trophic positions estimated from isotopic data. Adult O. xinguense had the highest trophic position despite having large amounts of Podostemaceae in the diet. High trophic overlap during the low-water period suggests that either food resources are not limiting, or niches are partitioned by other means. Differences in functional traits of the three serrasalmids could be associated with differential efficiencies of swimming and feeding within microhabitats that vary according to water velocity and/or structural complexity.

     

Comparative rates of lower jaw diversification in cichlid adaptive radiations

The lower jaw (LJ) provides an ideal trophic phenotype to compare rates and patterns of macroevolution among cichlid radiations. Using a novel phylogeny of four genes (ND2, dlx2, mitfb, and s7), we examined the evolutionary relationships among two of the most phylogenetically disparate cichlid radiations: (i) the Central America Heroines; and (ii) the East African Lake Malawi flock. To quantify jaw morphology, we measured two LJ lever systems in approximately 40 species from each lineage. Using geologic calibrations, we generated a chronogram for both groups and examined the rates of jaw evolution in the two radiations. The most rapidly evolving components of the LJ differed between the two radiations. However, the Lake Malawi flock exhibited a much faster rate of evolution in several components of the LJ. This rapid rate of divergence is consistent with natural selection, promoting unparalleled trophic diversification in Lake Malawi cichlids.     

On the origin and colonization of house mice in the Madeira Islands

The skulls and skins of adult house mice from the Madeira Islands have been studied and compared with those from the Salvage Islands and with material from the neighbouring Portuguese mainland man-associated and wild forms, respectively Mus musclus domesticus Rutty, 1772 and M. sprelus Lataste, 1883. Differences between island and mainland populations were found in some of the analysed features. Insular skins of mice were found to be smaller than those of specimens from the mainland. However, in Madeiran and Salvage mice the toothrows were much more developed than in the mainland house mice. It is considered that the causes of these differences lie in the different characteristics of the habitats, mainly food availability, and also in the isolation of populations. Mus musculus domesticus appears to be the only form of the house mouse to have so far successfully colonized the Madeiras.     

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.     

The origin and colonization history of the barley scald pathogen Rhynchosporium secalis

The origins of pathogens and their past and present migration patterns are often unknown. We used phylogenetic haplotype clustering in conjunction with model-based coalescent approaches to reconstruct the genetic history of the barley leaf pathogen Rhynchosporium secalis using the avirulence gene NIP1 and its flanking regions. Our results falsify the hypothesis that R. secalis emerged in association with its host during the domestication of barley 10,000 to 15,000 years ago in the Fertile Crescent and was introduced into Europe through the migration of Neolithic farmers. Estimates of time since most recent common ancestor (2500-5000 BP) placed the emergence of R. secalis clearly after the domestication of barley. We propose that modern populations of R. secalis originated in northern Europe following a host switch, most probably from a wild grass onto cultivated barley shortly after barley was introduced into northern Europe. R. secalis subsequently spread southwards into already established European barley-growing areas.     

Losing cichlid fish biodiversity: genetic and morphological homogenization of tilapia following colonization by introduced species

Among the many negative impacts of invasive species, hybridization with indigenous species has increasingly become recognized as a major issue. However, relatively few studies have characterized the phenotypic outcomes of hybridization following biological invasions. Here we investigate the genetic and morphological consequences of stocking invasive tilapia species in two water bodies in central Tanzania. We sampled individuals from the Mindu Reservoir on the Ruvu river system, and at Kidatu on the Great Ruaha–Rufiji river system. We screened individuals at 16 microsatellite loci, and quantified morphology using geometric morphometrics and linear measurements. In both the Mindu and Kidatu systems, we identified evidence of hybridization between indigenous Wami tilapia (Oreochromis urolepis) and the introduced Nile tilapia (Oreochromis niloticus) or blue-spotted tilapia (Oreochromis leucostictus). At both sites, purebred individuals could largely be separated using geometric morphometric variables, with hybrids occupying a broad morphospace among the parental species. Our data demonstrate that the gene pools and phenotypic identity of the indigenous O. urolepis have been severely impacted by the stocking of the invasive species. Given the lack of evidence for clear commercial benefits from stocking invasive tilapia species in waters already populated by indigenous congenerics, we suggest further spread of introduced species should be undertaken with considerable caution.     

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.     

Progressive island colonization and ancient origin of Hawaiian Metrosideros (Myrtaceae)

Knowledge of the evolutionary history of plants that are ecologically dominant in modern ecosystems is critical to understanding the historical development of those ecosystems. Metrosideros is a plant genus found in many ecological and altitudinal zones throughout the Pacific. In the Hawaiian Islands, Metrosideros polymorpha is an ecologically dominant species and is also highly polymorphic in both growth form and ecology. Using 10 non-coding chloroplast regions, we investigated haplotype diversity in the five currently recognized Hawaiian Metrosideros species and an established out-group, Metrosideros collina, from French Polynesia. Multiple haplotype groups were found, but these did not match morphological delimitations. Alternative morphologies sharing the same haplotype, as well as similar morphologies occurring within several distinct island clades, could be the result of developmental plasticity, parallel evolution or chloroplast capture. The geographical structure of the data is consistent with a pattern of age progressive island colonizations and suggests de novo intra-island diversification. If single colonization events resulted in a similar array of morphologies on each island, this would represent parallel radiations within a single, highly polymorphic species. However, we were unable to resolve whether the pattern is instead explained by ancient introgression and incomplete lineage sorting resulting in repeated chloroplast capture. Using several calibration methods, we estimate the colonization of the Hawaiian Islands to be potentially as old as 3.9 (-6.3) Myr with an ancestral position for Kaua'i in the colonization and evolution of Metrosideros in the Hawaiian Islands. This would represent a more ancient arrival of Metrosideros to this region than previous studies have suggested.     

Biogeographical origin effects on exotic plants colonization in the insular flora of Japan

Biological Invasions - Understanding the mechanisms of biological invasion is fundamental for biodiversity conservation in the Anthropocene. This study focused on a large-scale colonization pattern...     

Molecular evidence to suggest the origin of a colonization: Drosophila subobscura in America

The recent colonization of America by Drosophila subobscura represents a great opportunity for evolutionary biology studies. Knowledge of the populations from which the colonization started would provide an understanding of how genetic composition changed during adaptation to the new environment. Thus, a 793 nucleotide fragment of the Odh (Octanol dehydrogenase) gene was sequenced in 66 chromosomal lines from Barcelona (western Mediterranean) and in 66 from Mt. Parnes (Greece, eastern Mediterranean). No sequence of Odh fragment in Barcelona or Mt. Parnes was identical to any of those previously detected in America. However, an Odh sequence from Barcelona differed in only one nucleotide from another found in American populations. In both cases, the chromosomal lines presented the same inversion: O(7), and the Odh gene was located within this inversion. This evidence suggests a possible western Mediterranean origin for the colonization. Finally, the molecular and inversion data indicate that the colonization was not characterized by multiple reintroductions.     

Hybrid origin of a cichlid population in Lake Malawi: implications for genetic variation and species diversity

The importance of species recognition to taxonomic diversity among Lake Malawi cichlids has been frequently discussed. Hybridization - the apparent breakdown of species recognition - has been observed sporadically among cichlids and has been viewed as both a constructive and a destructive force with respect to species diversity. Here we provide genetic evidence of a natural hybrid cichlid population with a unique colour phenotype and elevated levels of genetic variation. We discuss the potential evolutionary consequences of interspecific hybridization in Lake Malawi cichlids and propose that the role of hybridization in generating both genetic variability and species diversity of Lake Malawi cichlids warrants further consideration.     

Ancient origin and recent divergence of a haplochromine cichlid lineage from isolated water bodies in the East African Rift system

Phylogenetic analysis identified haplochromine cichlids from isolated water bodies in the eastern branch of the East African Rift system as an ancient lineage separated from their western sister group in the course of the South Kenyan-North Tanzanian rift system formation. Within this lineage, the close phylogenetic relatedness among taxa indicates a recent common ancestry and historical connections between now separated water bodies. In connection with a total lack of local genetic diversity attributable to population bottlenecks, the data suggest cycles of extinction and colonization in the unstable habitat provided by the small lakes and rivers in this geologically highly active area.     

Genetic characterization of black rat (Rattus rattus) of the Canary Islands: origin and colonization

In the Canary Islands two invasive rat species, Rattus rattus and Rattus norvegicus are present, but little is known about the origin and colonization. To this end, a molecular study was performed on R. rattus from the Archipelago and from the nearest continents. Partial cytochrome b gene sequencing offered very low levels of haplotype and nucleotide diversities, with only seven haplotypes identified. All of them belong to the European Lineage I, specifically to the “ship rat” cluster. The haplotype network showed a star-like topology. Haplotype distribution showed a genetic subdivision between eastern and central/western islands, suggesting a double colonization event. This hypothesis is congruent with historical human colonization and it is similar to that proposed for the rodent parasite Hymenolepis diminuta. In addition, a possible role of the Canary Islands as a faunal link with the European and American continents is discussed.     

Time and biosequences: a contribution to the origin of modern man

The origin of modern man is still a highly debated problem. The new contributions of molecular biologists to paleontological data are numerous but discordant as these data obtained with biosequences are based on different methodologies. In order to obtain a more accurate measure of the genetic distance between extant humans and thus the deepest root of the human tree, we have reanalyzed the sequences of the regulatory region of mitochondrial DNA (D-loop) by using an improved version of the Markov Clock model devised in our laboratory. Our analysis, as well as other studies, supports the African origin of modern man. However the quantitative estimate of mtDNA evolution carried out with the corrected Markov Clock model pushes back irradiation time from Africa to 400±100 Kya. Thus according to our model there would be a genetic continuity betweenHomo erectus andHomo sapiens, who would therefore be the result of regional evolution differentiating himself under appropriate environmental pressures.     

Detection of Helicobacter colonization of the murine lower bowel by genus-specific PCR-denaturing gradient gel electrophoresis

Helicobacter genus-specific PCR and denaturing gradient gel electrophoresis can detect and speciate the helicobacters that colonize the lower bowel of laboratory mice. The method's sensitivity is comparable to that of species-specific PCR and may detect unnamed Helicobacter species. This approach should prove useful for commercial and research murine facilities.