The Atlantification of the European Arctic has been an increasingly discussed topic in polar science over the past two decades. The alteration of local marine ecosystems towards a more temperate ...state and the appearance/range expansion of subarctic-boreal species at higher latitudes is a complex phenomenon induced mainly by the changing properties of Atlantic water (AW) transported from the south. Areas under the direct influence of AW experience biological Atlantification of their communities on all trophic levels, resulting in the growing complexity of arctic food webs. Here, besides summarising the main documented messages of biological Atlantification, we take a critical view on the threat posed on Arctic marine communities. We take into account the formation of the Arctic marine fauna, as well as the nature of (re)colonisation of Arctic sites by boreal organisms when evaluating the extent of the issue. We take a look at the history of Arctic colonisations by boreal organisms in an attempt to identify ‘neonative taxa returning home’. We also highlight the role of floating plastic debris as an ‘instrument from the toolbox of the Anthropocene’ aiding the distribution of marine taxa.
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•Marine ecosystems of the European Arctic homogenise with those of the North Atlantic.•The biological messages of the Atlantification are expressed on all trophic levels.•The change is rather slow and quantitative towards more complicated food webs.•Neonative taxa are reappearing at sites, from where they retreated during the LGM.•Floating plastic debris may serve as a new way of transport for expanding boreal taxa.
The severe climatic changes during the Pleistocene ice ages have shaped the genetic structure and distribution of biota in Europe. We aimed to reveal in detail the genetic diversity, geographical ...population structure, historical and present demography, migration patterns, and the presence of possible glacial refugia within the nominative subspecies Asellus aquaticus aquaticus.
We analysed DNA sequences of the mitochondrial cytochrome c oxidase subunit I gene and nuclear noncoding internal transcribed spacer II region, from populations inhabiting post‐glacial Europe (from the British Isles and Scandinavia to the northern Mediterranean and Black Sea coasts).
The origin of the taxon, including establishment in the Dinaric Western Balkans, dates to the Middle/Late Pliocene, but most of its genetic diversity emerged during the Middle/Late Pleistocene before the Last Glacial Maximum.
Despite the general absence of spatial genetic structure with population growth, we discovered two different phylogeographic stories across 11 clusters revealed by a coalescent approach. Firstly, the periglacial cluster group—spatially restricted mainly to the northern Balkans, Pannonian Basin, and Pontic Region—is older and more divergent. It apparently retained a relatively stable population size during the glacial‐interglacial cycles. Conversely, the proglacial cluster group—widely distributed in areas close to the glacier margins (north of the Alps, Sudetes, Carpathians) and in the Pannonian Basin—is younger and composed of closely related individuals. It originated in Pleistocene and lasted continuously through the Last Glacial Maximum in numerous high latitude refugia. This was probably due to the vast network of proglacial lakes and rivers, which played a crucial role in the maintenance of genetic diversity, population growth, and high dispersal rate.
The evolutionary history of A. a. aquaticus reveals unexpected patterns and is an important lesson when making predictions for other aquatic taxa. Our results suggest that we should stop perceiving the proglacial habitats as lifeless ice desert.
In this paper we derive mathematical description of TCP (Transmission Control Protocol) retransmission probability based on Jacobson’s smoothing algorithm that belongs to EWMA (Exponentially Weighted ...Moving Average) category. This description is parametrized on the probability density function (pdf) of RTT (Round Trip Time) samples and α, β – two primary parameters of Jacobson’s algorithm. Although it is not a close form expression, it is formulated as an effective algorithm that let us to explicitly calculate the values of RTO (Retransmission Time Out) probability as a function of α, β and the pdf of RTT samples. We achieve the effectiveness of this approach by applying smart discretization of the state space and replacement of continuous functions with discrete approximate equivalents. In this way, we mitigate the cardinality of discrete distributions we deal with that results in linear (n+m) instead of multiplicative (n⋅m) growth of computational complexity. We provide the evaluation of RTO probability for a wide set of α, β parameter values and differently shaped Normal and Laplace pdfs the RTT samples are taken from. The obtained numerical results let us to draw some conclusions regarding the choice of optimal values of α, β parameters as well as the impact of pdf the RTT samples are taken from.
The Carpathians are one of the key biodiversity hotspots in Europe. The mountain chain uplifted during Alpine orogenesis and is characterised by a complex geological history. Its current biodiversity ...was highly influenced by Pleistocene glaciations. The goal of the current study was to examine the phylogenetic and demographic history of Gammarus balcanicus species complex in the Carpathians using multiple markers as well as to delimit, using an integrative approach, and describe new species hidden so far under the name G. balcanicus. Results showed that divergence of the studied lineages reaches back to the Miocene, which supports the hypothesis of their survival in multiple micro refugia. Moreover, the increase of their diversification rate in the Pleistocene suggests that glaciation was the driving force of their speciation. The climatic changes during and after the Pleistocene also played a major role in the demography of the local Carpathian lineages. Comparison of diversity patterns and phylogenetic relationships of both, the mitochondrial and nuclear markers, provide evidence of putative hybridisation and retention of ancient polymorphism (i.e., incomplete lineage sorting). The morphological examination supported the existence of two morphological types; one we describe as a G. stasiuki sp. nov. and another we redescribe as a G. tatrensis (S. Karaman, 1931).
The formation of continental Europe in the Neogene was due to the regression of the Tethys Ocean and of the Paratethys Sea. The dynamic geology of the area and repetitious transitions between marine ...and freshwater conditions presented opportunities for the colonization of newly emerging hydrological networks and diversification of aquatic biota. Implementing mitochondrial and nuclear markers in conjunction with a large‐scale sampling strategy, we investigated the impact of this spatiotemporal framework on the evolutionary history of a freshwater crustacean morphospecies. The Gammarus balcanicus species complex is widely distributed in the area previously occupied by the Paratethys Sea. Our results revealed its high diversification and polyphyly in relation to a number of other morphospecies. The distribution of the studied amphipod is generally characterized by very high local endemism and divergence. The Bayesian time‐calibrated reconstruction of phylogeny and geographical distribution of ancestral nodes indicates that this species complex started to diversify in the Early Miocene in the central Balkans, partially in the shallow epicontinental sea. It is possible that there were several episodes of inland water colonization by local brackish water lineages. Subsequent diversification within clades and spread to new areas could have been induced by Alpine orogeny in the Miocene/Pliocene and, finally, by Pleistocene glaciations. The present distribution of clades, in many cases, still reflects Miocene palaeogeography of the area. Our results point out that investigations of the historical aspect of cryptic diversity in other taxa may help in a general understanding of the origins of freshwater invertebrate fauna of Europe.
Our main aim was to investigate the diversity, origin and biogeographical affiliations of freshwater gammarids inhabiting the Aegean Islands by analysing their mtDNA and nDNA polymorphism, thereby ...providing the first insight into the phylogeography of the Aegean freshwater gammarid fauna. The study material was collected from Samothraki, Lesbos, Skyros, Evia, Andros, Tinos and Serifos islands as well as from mainland Greece. The DNA extracted was used for amplification of two mitochondrial (COI and 16S) and two nuclear markers (28S and EF1-alpha). The multimarker time-calibrated phylogeny supports multiple origins and different diversification times for the studied taxa. Three of the sampled insular populations most probably represent new, distinct species as supported by all the delimitation methods used in our study. Our results show that the evolution of freshwater taxa is associated with the geological history of the Aegean Basin. The biogeographic affiliations of the studied insular taxa indicate its continental origin, as well as the importance of the land fragmentation and the historical land connections of the islands. Based on the findings, we highlight the importance of studying insular freshwater biota to better understand diversification mechanisms in fresh waters as well as the origin of studied Aegean freshwater taxa.
Effective identification of species using short DNA fragments (DNA barcoding and DNA metabarcoding) requires reliable sequence reference libraries of known taxa. Both taxonomically comprehensive ...coverage and content quality are important for sufficient accuracy. For aquatic ecosystems in Europe, reliable barcode reference libraries are particularly important if molecular identification tools are to be implemented in biomonitoring and reports in the context of the EU Water Framework Directive (WFD) and the Marine Strategy Framework Directive (MSFD). We analysed gaps in the two most important reference databases, Barcode of Life Data Systems (BOLD) and NCBI GenBank, with a focus on the taxa most frequently used in WFD and MSFD. Our analyses show that coverage varies strongly among taxonomic groups, and among geographic regions. In general, groups that were actively targeted in barcode projects (e.g. fish, true bugs, caddisflies and vascular plants) are well represented in the barcode libraries, while others have fewer records (e.g. marine molluscs, ascidians, and freshwater diatoms). We also found that species monitored in several countries often are represented by barcodes in reference libraries, while species monitored in a single country frequently lack sequence records. A large proportion of species (up to 50%) in several taxonomic groups are only represented by private data in BOLD. Our results have implications for the future strategy to fill existing gaps in barcode libraries, especially if DNA metabarcoding is to be used in the monitoring of European aquatic biota under the WFD and MSFD. For example, missing species relevant to monitoring in multiple countries should be prioritized for future collaborative programs. We also discuss why a strategy for quality control and quality assurance of barcode reference libraries is needed and recommend future steps to ensure full utilisation of metabarcoding in aquatic biomonitoring.
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•DNA barcode representation in public databases of 28,000 aquatic species is analysed.•Gaps in barcode reference libraries are largest for diatoms and invertebrates.•Sequence coverage varies considerably among invertebrate groups.•Species monitored by one or few countries more frequently lack reference barcodes.•Strategies should be implemented to maintain quality of barcode reference libraries.
The amphipod Dikerogammarus villosus has colonized most of the European main inland water bodies in less than 20 years, having deteriorating effect on the local benthic communities. Our aim was to ...reveal the species phylogeography in the native Black Sea area, to define the source populations for the colonization routes in continental Europe and for the newly established UK populations. We tested for the loss of genetic diversity between source and invasive populations as well as along invasion route. We tested also for isolation by distance. Thirty three native and invasive populations were genotyped for mtDNA (COI, 16S) and seven polymorphic nuclear microsatellites to assess cryptic diversity (presence of deeply divergent lineages), historical demography, level of diversity within lineage (e.g., number of alleles), and population structure. A wide range of methods was used, including minimum spanning network, molecular clock, Bayesian clustering and Mantel test. Our results identified that sea level and salinity changes during Pleistocene impacted the species phylogeography in the Black Sea native region with four differentiated populations inhabiting, respectively, the Dnieper, Dniester, Danube deltas and Durungol liman. The invasion of continental Europe is associated with two sources, i.e., the Danube and Dnieper deltas, which gave origin to two independent invasion routes (Western and Eastern) for which no loss of diversity and no isolation by distance were observed. The UK population has originated in the Western Route and, despite very recent colonization, no drastic loss of diversity was observed. The results show that the invasion of the killer shrimp is not associated with the costs of loosing genetic diversity, which may contribute to the success of this invader in the newly colonized areas. Additionally, while it has not yet occurred, it might be expected that future interbreeding between the genetically diversified populations from two independent invasion routes will potentially even enhance this success.