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The microsporidian genus Nosema is primarily known to infect insects of economic importance stimulating high research interest, while other hosts remain understudied. Nosema granulosis is one of the formally described Nosema species infecting amphipod crustaceans, being known to infect only two host species. Our first aim was to characterize Nosema spp. infections in different amphipod species from various European localities using the small subunit ribosomal DNA (SSU) marker. Second, we aimed to assess the phylogenetic diversity, host specificity and to explore the evolutionary history that may explain the diversity of gammarid-infecting Nosema lineages by performing a phylogenetic reconstruction based on RNA polymerase II subunit B1 (RPB1) gene sequences. For the host species Gammarus balcanicus, we also analyzed whether parasites were in excess in females to test for sex ratio distortion in relation with Nosema infection. We identified Nosema spp. in 316 individuals from nine amphipod species being widespread in Europe. The RPB1-based phylogenetic reconstruction using newly reported sequences and available data from other invertebrates identified 39 haplogroups being associated with amphipods. These haplogroups clustered into five clades (A-E) that did not form a single amphipod-infecting monophyletic group. Closely related sister clades C and D correspond to Nosema granulosis. Clades A, B and E might represent unknown Nosema species infecting amphipods. Host specificity seemed to be variable with some clades being restricted to single hosts, and some that could be found in several host species. We show that Nosema parasite richness in gammarid hosts is much higher than expected, illustrating the advantage of the use of RPB1 marker over SSU. Finally, we found no hint of sex ratio distortion in Nosema clade A infecting G. balcanicus. This study shows that Nosema spp. are abundant, widespread and diverse in European gammarids. Thus, Nosema is as diverse in aquatic as in terrestrial hosts.

Abstract Mayflies (Ephemeroptera) are among the oldest pterygote insects, with the earliest fossils dating back to the Late Carboniferous. Within mayflies, Leptophlebiidae are a highly diverse and widespread group, with approximately 140 genera and 640 species. Whereas taxonomy, systematics, and phylogeny of extant Leptophlebiidae are in the focus of extensive studies, little is known about leptophlebiid fossil taxa. Because fossil remains of Ephemeroptera in sedimentary rocks are relatively rare, inclusions of mayflies in amber are a unique source of information on their evolution and diversity in the past. Leptophlebiidae found in Cenozoic resins mostly belong to the subfamilies Leptophlebiinae (in Eocene Baltic amber) and Atalophlebiinae (in Miocene Dominican and Mexican ambers). In the present contribution, we confirm the first finding of the genus Calliarcys from Eocene Baltic amber by using Micro-CT, which allowed confirming its generic placement by visualizing diagnostic key characters otherwise hidden by a cloud of turbidity. Additionally, we present first molecular data on the extant species Calliarcys humilis Eaton, 1881 from the Iberian Peninsula and the barcode gap analysis for Leptophlebiinae and Habrophlebiinae.

Abstract The Ponto-Caspian region is an endemicity hotspot that harbours several crustacean radiations, among which amphipods are the most diverse. These poorly known species are severely threatened in their native range, while at the same time they are invading European inland waters with significant ecological consequences. A proper taxonomic knowledge of this fauna is paramount for its conservation within the native region and monitoring outside of it. Here, we assemble a DNA barcode reference library for nearly 60% of all known Ponto-Caspian amphipod species. We use several methods to define molecular operational taxonomic units (MOTUs), based on two mitochondrial markers (COI and 16S), and assess their congruence with current species-level taxonomy based on morphology. Depending on the method, we find that 54–69% of species had congruent morpho-molecular boundaries. The cases of incongruence resulted from lumping distinct morphospecies into a single MOTU (7–27%), splitting a morphospecies into several MOTUs (4–28%), or both (4–11%). MOTUs defined by distance-based methods without a priori divergence thresholds showed the highest congruence with morphological taxonomy. These results indicate that DNA barcoding is valuable for clarifying the diversity of Ponto-Caspian amphipods, but reveals that extensive work is needed to resolve taxonomic uncertainties. Our study advances the DNA barcode reference library for the European aquatic biota, paving the way towards improved taxonomic knowledge needed to enhance monitoring and conservation efforts.

Abstract The region of the Western Carpathians is, among other aspects, very important for survival and diversity of European freshwater fauna due to the presence of a large number of (sub)mountain springs and streams. However, these ecologically and faunistically diversified habitats are still understudied in the context of genetic diversity and population structure of their inhabitants. This study focuses on genetic diversity and distribution patterns of the caddisfly Rhyacophila tristis, common and widespread representative of mountain freshwater fauna. Analysis of the COI mitochondrial marker revealed presence of the western and eastern lineages, with samples from both lineages being grouped in BOLD (Barcode of Life Data System) into separate BINs (Barcode Index Numbers). Our data indicates that eastern lineage (BIN_E) is more closely related to the Balkan populations than to co-occurring western lineage (BIN_W), and that the contact zone of the lineages passes through the W Carpathians. The study revealed phylogeographic and demographic differences between lineages, supporting hypothesis of their evolutionary independence and specific ecological preferences. The obtained genetic data of the R. tristis population from W Carpathians improved our knowledge about population genetics of this aquatic species and can contribute to understanding the state and evolution of biodiversity of freshwater ecosystems in Europe.

Parasites and other symbionts deeply influence host organisms, and no living organism can be considered to have evolved independent of its symbionts. The first step towards understanding symbiotic influences upon host organisms is a strong supporting knowledge of parasite/symbiont diversity. Parasites of freshwater amphipods are diverse, with Microsporidia being a major group. These intracellular parasites impact gammarid fitness in different ways, ranging from reduced fitness to increased fecundity. Many Microsporidia have been recorded using molecular data, with multiple taxa pending formal taxonomic description. While some parasites are common, others are known only through sporadic records of single infections. In this study, we focus on rare/sporadic microsporidian infections within Gammarus balcanicus, a host species complex with a high level of endemism. In addition to enriching our knowledge on Microsporidia parasite diversity in amphipod hosts, we test whether these symbionts are specific to G. balcanicus or if they are the same taxa infecting other gammarid species. Of 2231 hosts from 87 sites, we catalogued 29 sequences of “rare” Microsporidia clustering into 19 haplogroups. These haplogroups cluster into 11 lineages: four pre-described taxa (Cucumispora roeselum, C. ornata, C. dikerogammari and Enterocytospora artemiae) and seven ‘Molecular Operational Taxonomic Units’, which are known from previously published studies to infect other European amphipod species. Our study significantly widens the geographic range of these Microsporidia and expands the known spectrum of hosts infected. Our results suggest that these parasites are ancient infections of European gammarids. For some host-parasite systems, we hypothesize that the common parasite ancestors that infected the hosts' common ancestors, diversified alongside host diversification. For others, we observe Microsporidia taxa with wide host ranges that do not follow host phylogeny.

Abstract Background Although the processes of co-evolution between parasites and their hosts are well known, evidence of co-speciation remains scarce. Microsporidian intracellular parasites, due to intimate relationships with their hosts and mixed mode of transmission (horizontal but also vertical, from mother to offspring), may represent an interesting biological model for investigating co-speciation. Amphipod crustaceans, especially gammarids, are regular hosts of microsporidian parasites, in particular the Dictyocoela spp., which have so far been found limited to these amphipods and are known to use a vertical mode of transmission. The amphipod genus Gammarus has a diversification history spanning the last 50–60 Mya and an extensive cryptic diversity in most of the nominal species. Here, we investigated the degree of co-diversification between Dictyocoela and Gammarus balcanicus, an amphipod with high degrees of ancient cryptic diversification and lineage endemism, by examining the genetic diversity of these parasites over the entire geographic range of the host. We hypothesised that the strong host diversification and vertical transmission of Dictyocoela would promote co-diversification. Results Using the parasite SSU rDNA as a molecular marker, analyzing 2225 host specimens from 88 sites covering whole host range, we found 31 haplogroups of Dictyocoela, 30 of which were novel, belonging to four Dictyocoela species already known to infect other Gammarus spp. The relationships between Dictyocoela and gammarids is therefore ancient, with the speciation in parasites preceding those of the hosts. Each novel haplogroup was nevertheless specific to G. balcanicus, leaving the possibility for subsequent co-diversification process during host diversification. A Procrustean Approach to Co-phylogeny (PACo) analysis revealed that diversification of Dictyocoela was not random with respect to that of the host. We found high degrees of congruence between the diversification of G. balcanicus and that of Dictyocoela roeselum and D. muelleri. However, we also found some incongruences between host and Dictyocoela phylogenies, e.g. in D. duebenum, probably due to host shifts between different G. balcanicus cryptic lineages. Conclusion The evolutionary history of Dictyocoela and Gammarus balcanicus represents an example of an overall host-parasite co-diversification, including cases of host shifts.

Abstract Background Microsporidians are obligate endoparasites infecting taxonomically diverse hosts. Both vertical (from mother to eggs) and horizontal (between conspecifics or between species) transmission routes are known. While the former may promote co-speciation and host-specificity, the latter may promote shifts between host species. Among aquatic arthropods, freshwater amphipod crustaceans are hosts for many microsporidian species. However, despite numerous studies, no general pattern emerged about host specificity and co-diversification. In south-eastern Europe, the gammarid Gammarus roeselii is composed of 13 cryptic lineages of Miocene to Pleistocene age but few genotypes of one lineage have spread postglacially throughout north-western Europe. Based on nearly 100 sampling sites covering its entire range, we aim to: (i) explore the microsporidian diversity present in G. roeselii and their phylogenetic relationships, especially in relation to the parasites infecting other Gammaridae; (ii) test if the host phylogeographical history might have impacted host–parasite association (e.g. co-diversifications or recent host shifts from local fauna). Methods We used part of the small subunit rRNA gene as source of sequences to identify and determine the phylogenetic position of the microsporidian taxa infecting G. roeselii. Results Microsporidian diversity was high in G. roeselii with 24 detected haplogroups, clustered into 18 species-level taxa. Ten microsporidian species were rare, infecting a few individual hosts in a few populations, and were mostly phylogenetically related to parasites from other amphipods or various crustaceans. Other microsporidians were represented by widespread genera with high prevalence: Nosema, Cucumispora and Dictyocoela. Two contrasting host association patterns could be observed. First, two vertically transmitted microsporidian species, Nosema granulosis and Dictyocoela roeselum, share the pattern of infecting G. roeselii over most of its range and are specific to this host suggesting the co-diversification scenario. This pattern contrasted with that of Dictyocoela muelleri, the three species of Cucumispora, and the rare parasites, present only in the recently colonised region by the host. These patterns suggest recent acquisitions from local host species, after the recent spread of G. roeselii. Conclusions Microsporidians infecting G. roeselii revealed two scenarios of host–parasite associations: (i) ancient associations with vertically transmitted parasites that probably co-diversified with their hosts, and (ii) host shifts from local host species, after the postglacial spread of G. roeselii.

In mid-1990s racer goby, Babka gymnotrachelus, penetrated to the Vistula and Western Bug rivers in Poland through the canal connecting the Baltic and the Black Sea basins, namely the Vistula and Dnieper drainages. In early 2000s, the species was reported from Polish section of the Strwiąż River that is an affluent of the Dniester River that drains to the Black Sea basin. According to the Polish legislation, the racer goby has been enlisted in as an alien invasive species that may pose threat to local biota. Our analysis of the mtDNA cytochrome b diversity revealed that the Vistula/Western Bug and Strwiąż populations are different genetic units. First one originated from the Dnieper River, while the second derives from the Dniester River. According to the results of mismatch analysis, both are in the stage of demographic and spatial expansion. The haplotype frequencies in population from the Vistula/Western Bug differ significantly from those in the source population in Dnieper, suggesting founder effect, possibly due to human-mediated introduction of low number of individuals. On the other side, the population in Strwiąż does not differ in structure from the one in Dniester, providing a hint towards spontaneous range expansion. Interpretation of our results in light of historical data lead to the conclusion that presence of racer goby in Strwiąż was probably overlooked in previous, spatially limited, studies. Thus, in Strwiąż the species should not be treated as alien, contrary to its status in the Vistula drainage. This double origin of racer goby populations in Poland creates a peculiar situation for national legislation procedures as one population is alien and invasive, while the other one is not. In light of our findings, the population from Strwiąż should be recognised as a special case. Steps should be undertaken to raise public awareness to prevent translocation of the gobies between the rivers to prevent deterioration of the evolutionarily isolated genetic pools of the Dnieper and of the Dniester basins. Our results illustrate the need for case studies upon genetic population structure, following appearance of new species in previously unoccupied water bodies, even on local scale.

One of the negative impacts of non-native invasive species on trophic interactions in an invaded ecosystem occurs via increased interspecific competition for food resources between the invader and local species of the same food niche. In freshwaters, there are usually several fish species that feed on similar food resources. Ponto-Caspian gobies are amongst the most successful and widespread invaders colonising European waterways. They have a wide food niche and an opportunistic feeding strategy, with a focus on benthic invertebrates and piscivory occurring occasionally mainly in the case of large individuals. Competition with native percids for food resources is predicted on the basis of high dietary overlap. However, studies published so far provide no unequivocal answer. In order to resolve this question, we conducted a comparative taxonomic analysis of gut content, with an emphasis on chironomids and amphipods, of the invasive monkey goby (Neogobius fluviatilis), racer goby (Babka gymnotrachelus) and the native Eurasian perch (Perca fluviatilis) occurring sympatrically in a large lowland European river, the Bug River in Poland. We found that each species forages in slightly different habitats, as indicated by the different composition of prey species in the gut content. This suggests feeding niche partitioning between the studied species facilitating their co-existence and reduction or avoidance of competition for food resources. Resource partitioning regarding prey types and foraging habitats is a mechanism for permitting the co-existence of closely-related alien gobies with similar food preferences in the invaded waters and co-occurrence with local species. This mechanism can contribute to their invasion success, as observed in European waters during the recent decades. We also demonstrate that precise prey identification to the lowest possible taxon is crucial to reveal the dietary overlap between co-occurring fish species and to predict the impact of alien invaders on native species through interspecific competition, as well as to recommend such an approach in studies upon fish foraging strategies.

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 Second Payment Services Directive introduced new services into the European Union legal system—Payment Initiation and Account Information Services. These services are based on payment accounts already opened and maintained for customers by the Account Servicing Payment Service Provider (bank, payment institution, electronic money institution). The Account Services Payment Service provider performs AML/CFT verification of the account holder and applies customer due diligence measures to the account holder, such as identifying beneficial owners, obtaining information on the purpose and intended nature of the business relationship, and ongoing monitoring of the business relationship. Payment Initiation and Account Information services are therefore provided to a previously verified client and based on the payment account currently maintained for him. European Union law does not clearly specify whether a Third-Party Service Provider offering Payment Initiation or Account Information Services is obliged to re-apply financial security measures to customers. The aim of this article was to perform a legal analysis of the regulations and soft law acts in force in the European Union and to answer the question. The purposive (teleological) and linguistic–logical (grammatical) methods of interpretation of regulations were used for the analysis. The structure of the legal system of the European Union as a civil law (code law) system was taken into account. This article shows that in the current legal situation, there is no doubt that Third-Party Service Providers are obliged entities in terms of AML/CFT law and are obliged to apply the AML/CFT to customers using Payment Initiation and Account Information services. However, the degree to which customer due diligence measures have to be applied varies depending on the adopted model of providing Payment Initiation and Account Information services. Third-Party Service Providers will be obliged to apply financial security measures in cases where the relationship between the customer and the service providers will have a continuing character. In the case of occasional provision of services, when the transaction value does not exceed a certain threshold, the supplier may only perform simplified customer verification. In particular, this applies to Payment Initiation service models, where the Payment Initiation Service Provider works for merchants, enabling them to accept payments for goods and services sold. In such a model, the Service Provider has a continuous relationship with the merchant but only performs an occasional transaction for the user. The analysis also allowed for the conclusion that European Union law, including that in the draft phase, does not regulate in a sufficiently precise manner when a given model of Account Services and Payment Initiation Services may be treated as based on an occasional transaction. This made it possible to formulate a de lege ferenda request to include this issue in the proposal for an EU Regulation on the prevention of the use of the financial system for the purposes of money laundering or terrorist financing.

Non-indigenous species (NIS) contribute to the decrease of native species’ diversity on a local and global scale. One of Europe’s most significant donors of freshwater invasions is the Ponto-Caspian Region. Following the construction of artificial canals connecting isolated waterbodies and the resulting heavy boat traffic, the Ponto-Caspian Amphipoda started to spread in Europe. Four amphipod species: Dikerogammarus haemobaphes, Dikerogammarus villosus, Pontogammarus robustoides and Chaetogammarus ischnus have invaded the Masurian Lakeland (North-eastern Poland). Based on literature and our data, we studied their distribution in 22 lakes in the region during the years 2001–2016. We analysed their distribution against several water quality parameters and levels of anthropogenic pressure. Our results also present the first records of two new invaders, D. villosus and C. ischnus, in the studied area. We show that the relative abundance and frequency of these two species rapidly increase and, simultaneously, the populations of the earlier invaders, i.e. D. haemobaphes and P. robustoides, decrease. The native species – Gammarus lacustris – seems to be negatively affected by NIS richness, as well as by the proximity of towns. The spread of NIS in the lakes appears to be facilitated by boating and the lower complexity of the shoreline. Our study shows how anthropogenic pressure, especially tourism, can facilitate bioinvasion, jeopardising native biodiversity unless appropriate regulations are implemented.

We provide an overview of the World Amphipoda Database (WAD), a global species database that is part of the World Register of Marine Species (WoRMS). Launched in 2013, the database contains entries for over 10,500 accepted species names. Edited currently by 31 amphipod taxonomists, following WoRMS priorities, the WAD has at least one editor per major group. All accepted species are checked by the editors, as is the authorship available for all of the names. The higher classification is documented for every species and a type species is recorded for every genus name. This constitutes five of the 13 priorities for completion, set by WoRMS. In 2015, five LifeWatch grants were allocated for WAD activities. These included a general training workshop in 2016, together with data input for the superfamily Lysianassoidea and for a number of non-marine groups. Philanthropy grants in 2019 and 2021 covered more important gaps across the whole group. Further work remains to complete the linking of unaccepted names, original descriptions, and environmental information. Once these tasks are completed, the database will be considered complete for 8 of the 13 priorities, and efforts will continue to input new taxa annually and focus on the remaining priorities, particularly the input of type localities. We give an overview of the current status of the order Amphipoda, providing counts of the number of genera and species within each family belonging to the six suborders currently recognized.

IntroductionPost-glacial climate variation is known to have influenced the distribution of marine species in the North Atlantic. In particular, the Atlantic side of the Arctic has experienced strong fluctuations in both atmospheric and sea surface temperature, as well as seasonal ice coverage since the last glacial maximum (LGM). Here, we aim to unveil the phylogeography and historical demography of three rocky intertidal marine invertebrates showing a trans-Atlantic distribution and presently inhabiting the Arctic: Gammarus oceanicus, Littorina saxatilis and Semibalanus balanoides.MethodsWe used a large amount of mitochondrial DNA barcode data, both newly-obtained and stored in public databases. We performed phylogeographic and demographic analyses on 1119 G. oceanicus, 205 L. saxatilis, and 884 S. balanoides sequences.ResultsOur results show that all three of these boreal species have expanded their effective population sizes in the high Arctic Svalbard Archipelago since the LGM. Analyses investigating the origin of all these populations point to the eastern Atlantic.DiscussionBased on our results we conclude that the expansion of these boreal species to the Arctic possibly happened during an earlier warm cycle of the Holocene era, and is probably not the result of the recent ‘Atlantification’ of the Arctic. We also discuss the effects of dispersal potential on population structure as an important aspect of comparative biogeographical studies.