Aim Elevated biodiversity is the result of the cradle, museum or sink functions. The contributions of these three functions to species accumulation and their changes through time remain unknown for glacial refugia. Additionally, our understanding of the role these functions played during pre‐glacial periods is limited. We test for changes in contributions of functions through time leading to the current diversity patterns using a model refugium and taxon. Location Anatolia, Western Palaearctic. Taxon Freshwater neritid snails (genus Theodoxus). Methods Assessments were made to define molecular operational taxonomic units (MOTUs) for Theodoxus and reaffirm the genus as a suitable model taxon with elevated interspecific diversity in noted glacial refugia. Thereafter, we constructed a time‐calibrated multilocus Bayesian phylogeny of mtDNA and nDNA by using both fossil data and published substitution rates. Ancestral area estimation was performed on the phylogeny to define the contribution of the functions through time. Results Accumulation of Theodoxus diversity in Anatolia over the Miocene–Pliocene transition is attributed to the museum function, but its contribution was small as only few divergence events occurred. The cradle function dominated during the Pliocene and Early Pleistocene, when most interspecific diversity built up and extant lineages in Anatolia were established. The sink function acted from the Middle Pleistocene to present‐day, but with only a small contribution to the total extant Anatolian interspecific diversity. Main conclusion Our results do not entirely mitigate the role glacial cycles played in species accumulation, but highlight Ice Ages may have been less effective in forcing temperate aquatic interspecific diversity into more opportune areas. The elevated diversity in refugia may rather be the result of earlier in situ diversification. Elevated interspecific diversity attributed to the legacy of glacially forced retreats may need to be re‐evaluated in cases where refugia have long and complex geological histories such as Anatolia. These results highlight the importance of considering species accumulation through a temporal perspective to adequately explain present‐day biodiversity patterns.
The Caspian Sea has been a highly dynamic environment throughout the Quaternary and witnessed major oscillations in lake level, which were associated with changes in salinity and habitat availability. Such environmental pressures are considered to drive strong phylogeographic structures in species by forcing populations into suitable refugia. However, little is actually known on the effect of lake-level fluctuations in the Caspian Sea on its aquatic biota. We compared the phylogeographic patterns of the aquatic Neritidae snail genus Theodoxus across the Pontocaspian region with refugial populations in southern Iran. Three gene fragments were used to determine relationships and divergence times between the sampled populations in both groups. A dated phylogeny and statistical haplotype networks were generated in conjunction with the analyses of molecular variance and calculations of isolation by distance using distance-based redundancy analyses. Extended Bayesian skyline plots were constructed to assess demographic history. Compared with the southern Iranian populations, we found little phylogeographic structure for the Pontocaspian Theodoxus group, with more recent diversification, homogeneity of haplotypes across the Pontocaspian region and a relatively stable demographic history since the Middle Pleistocene. Our results argue against a strong influence of Caspian Sea low stands on the population structure post the early Pleistocene, whereas high stands may have increased the dispersal possibilities and homogenization of haplotypes across the Pontocaspian region during this time. However, during the early Pleistocene, a more dramatic low stand in the Caspian Sea, around a million years ago, may have caused the reduction in Theodoxus diversity to a single lineage in the region. In addition, our results provide new insights into Theodoxus taxonomy and outlooks for regional conservation.
Background The Lake Victoria basin is one of the most persistent hotspots of schistosomiasis in Africa, the intestinal form of the disease being studied more often than the urogenital form. Most schistosomiasis studies have been directed to Schistosoma mansoni and their corresponding intermediate snail hosts of the genus Biomphalaria , while neglecting S. haematobium and their intermediate snail hosts of the genus Bulinus . In the present study, we used DNA sequences from part of the cytochrome c oxidase subunit 1 ( cox 1) gene and the internal transcribed spacer 2 (ITS2) region to investigate Bulinus populations obtained from a longitudinal survey in Lake Victoria and neighbouring systems during 2010–2019. Methods Sequences were obtained to (i) determine specimen identities, diversity and phylogenetic positions, (ii) reconstruct phylogeographical affinities, and (iii) determine the population structure to discuss the results and their implications for the transmission and epidemiology of urogenital schistosomiasis in Lake Victoria. Results Phylogenies, species delimitation methods (SDMs) and statistical parsimony networks revealed the presence of two main groups of Bulinus species occurring in Lake Victoria; B. truncatus / B. tropicus complex with three species ( B. truncatus , B. tropicus and Bulinus sp. 1), dominating the lake proper, and a B. africanus group, prevalent in banks and marshes. Although a total of 47 cox 1 haplotypes, were detected within and outside Lake Victoria, there was limited haplotype sharing (only Haplotype 6 was shared between populations from Lake Victoria open waters and neighbouring aquatic systems) – an indication that haplotypes are specific to habitats. Conclusions The Bulinus fauna of Lake Victoria consists of at least B. truncatus , B. tropicus , Bulinus sp. 1 ( B. trigonus ?) and B. ugandae . The occurrence and wide distribution of Bulinus species in Lake Victoria potentially implies the occurrence of urogenital schistosomiasis in communities living along the shores and on islands of the lake who depend solely on the lake for their livelihood. More in-depth studies are needed to obtain a better picture of the extent of the disease in the Lake Victoria basin.
Hyalomma Koch, 1844 are ixodid ticks that infest mammals, birds and reptiles, to which 27 recognized species occur across the Afrotropical, Palearctic and Oriental regions. Despite their medical and veterinary importance, the evolutionary history of the group is enigmatic. To investigate various taxonomic hypotheses based on morphology, and also some of the mechanisms involved in the diversification of the genus, we sequenced and analysed data derived from two mtDNA fragments, three nuclear DNA genes and 47 morphological characters. Bayesian and Parsimony analyses based on the combined data (2242 characters for 84 taxa) provided maximum resolution and strongly supported the monophyly of Hyalomma and the subgenus Euhyalomma Filippova, 1984 (including H. punt Hoogstraal, Kaiser and Pedersen, 1969). A predicted close evolutionary association was found between morphologically similar H. dromedarii Koch, 1844, H. somalicum Tonelli Rondelli, 1935, H. impeltatum Schulze and Schlottke, 1929 and H. punt, and together they form a sister lineage to H. asiaticum Schulze and Schlottke, 1929, H. schulzei Olenev, 1931 and H. scupense Schulze, 1919. Congruent with morphological suggestions, H. anatolicum Koch, 1844, H. excavatum Koch, 1844 and H. lusitanicum Koch, 1844 form a clade and so also H. glabrum Delpy, 1949, H. marginatum Koch, 1844, H. turanicum Pomerantzev, 1946 and H. rufipes Koch, 1844. Wide scale continental sampling revealed cryptic divergences within African H. truncatum Koch, 1844 and H. rufipes and suggested that the taxonomy of these lineages is in need of a revision. The most basal lineages in Hyalomma represent taxa currently confined to Eurasia and molecular clock estimates suggest that members of the genus started to diverge approximately 36.25 million years ago (Mya). The early diversification event coincides well with the collision of the Indian and Eurasian Plates, an event that was also characterized by large scale faunal turnover in the region. Using S-Diva, we also propose that the closure of the Tethyan seaway allowed for the genus to first enter Africa approximately 17.73Mya. In concert, our data supports the notion that tectonic events and large scale global changes in the environment contributed significantly to produce the rich species diversity currently found in the genus Hyalomma.
The phylogeographic patterns of small mammals in southern Africa are frequently disjunct. This pattern is predominately attributed to vicariant geographical barriers coupled to climate driven diversification. To gain further insights into this hypothesis, we embarked on a comparative mtDNA phylogeographic study of two common rodent species in southern Africa, Mastomys natalensis and Mastomys coucha. Parsimony haplotype networks and SplitsTrees of mtDNA cytochrome oxidase I data showed a large degree of haplotype sharing throughout the sampling range. Within southern Africa, we found no conclusive evidence to support geographic vicariance as a contributing factor towards Mastomys speciation. We proposed that the regional phylogeographic structures detected for M. natalensis and M. coucha are the result of weak isolation by distance coupled to repeated expansions and contractions of suitable habitat. Both species probably survived in multiple refugia during unfavourable periods and mismatch distributions show signs of population expansion. Mitochondrial DNA nucleotide diversity values (π) show marked differences between the two species (M. natalensis: 0.003 and M. coucha: 0.468), and M. coucha also shows a higher level of population differentiation in AMOVA analyses. These differences are most likely due to life history discrepancies between the two species. Mastomys coucha is regarded to be more of a habitat specialist when compared to M. natalensis, and this probably places a higher constraint on M. coucha dispersal abilities. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114, 58–68.
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