Discovery of Dreissena rostriformis bugensis (Andrusov 1897) in Western Europe

Live individuals of Dreissena rostriformis bugensis were found in a trawlnet sample taken in April 2006 near Willemstad in the Hollands Diep, part of the main distributary in the Rhine delta in The Netherlands. This represented the first record of this species in Western Europe and a major westward range extension for this Ponto-Caspian species. Density was very low, with the ratio of Dreissena polymorpha to D. rostriformis bugensis in the order of 100:1. Mean length (n = 5) was 22.5 mm (range 20.4–24.6 mm), suggesting that they were at least two years old and that the original introduction into the Hollands Diep had occurred in 2004 or earlier. Species identity was confirmed by morphological and molecular methods. We speculate that the invasion occurred through the Main-Danube Canal – a pathway that is becoming increasingly recognized as a southern corridor for the potential movement of non-indigenous aquatic species between Eastern and Western Europe.     

Journey into the past: using cryogenically stored samples to reconstruct the invasion history of the quagga mussel (Dreissena rostriformis) in German river systems

Knowledge about the spatial–temporal dynamics of biological invasions often remains incomplete, because precise information about the invaders’ arrival dates is rare. This applies to the quagga mussel, which has become one of the most successful invasive species in Western European freshwaters. We here used cryogenically stored Dreissena samples from the German Environmental Specimen Bank to reconstruct the colonization history of the quagga mussel in German river systems. Our retrospective genetic analysis significantly improved upon previous findings of when the quagga mussel arrived in Germany and can be used as chronological landmarks to reconstruct its range expansion. The discovery of Dreissena rostriformis in 2004 in the Rhine River near Koblenz presented the first record of this species not only in Germany, but also in Western Europe. Our results show that the quagga mussel had already invaded not only large parts of the Rhine and the Danube, but also the Elbe River. This demonstrates the value of cryobanked biological samples for the retrospective analysis of biological ‘pollution’ through alien invasive species.     

Filtration activity of invasive mussel species under wave disturbance conditions

Self-purification is one of the most important ecosystem functions of rivers. Multiple human activities regularly impact this ecosystem service, consequently altering river morphology, hydrology, and the composition of biotic assemblages that contribute to self-purification. However, little quantitative information is available about the importance of such impacts. Hence, we tested how invasive mussel species contribute to self-purification under disturbed riverine conditions. In laboratory experiments, invasive mussel species equipped with magnetic sensors that recorded filtration activity were exposed to artificial waves of varying intensity that simulated the hydraulic effects of inland navigation. Our results suggest that invasive mussel species are more resistant to wave disturbance compared to native species, as estimated threshold values for initiating shell closure are very high (Dreissena rostriformis bugensis) or the duration (Corbicula fluminea) and degree of shell closing (D. rostriformis bugensis, C. fluminea) very low. Also we demonstrated that the invasive species D. rostriformis bugensis and C. fluminea continued filtering during wave impact, whereas Dreissena polymorpha did not behave significantly differently than previously studied native mussel species, based on the studied susceptibility parameters. Thus, D. rostriformis bugensis and C. fluminea appear to be pre-adapted to hydraulic or morphological disturbance, and may compensate against other losses regarding this important ecosystem function in rivers that are intensively used for inland navigation. However, as the dominance of invasive species in river systems may disrupt natural food webs, this compensation of filter-feeding activity may be accompanied by the loss of other ecosystem functions.     

Invasion ecology of quagga mussels (<Emphasis Type="Italic">Dreissena rostriformis bugensis</Emphasis>): a review of evolutionary and phylogenetic impacts

Two invasive freshwater mussels, Dreissena rostriformis bugensis (quagga mussel) and D. polymorpha (zebra mussel), reveal differences in patterns and timing of their invasions in Europe. They belong to different clades in Dreissena phylogenetics: D. rostriformis bugensis genetically is coupled with the brackish water, lacustrine D. r. distincta and the two are believed to represent a single species. As such, the guaqqa mussel has environmental requirements that differ from the congeneric D. polymorpha. D. rostriformis bugensis invasions were confined to reservoirs of the Dnieper, Don and Dniester Rivers of the Black Sea basin. We recorded D. r. bugensis outside the Black Sea basin for the first time between 1992 and 2001, along the Volga River reservoir cascade including the Northern Caspian Sea shallows. This represents a 40-year invasion time lag since an invasion corridor through the Volgo-Don Waterway was established in 1952 (a corridor used extensively by many invertebrate species from the Black Sea region). We attribute the postponed invasion of Europe by D. r. bugensis, including peculiarities in establishment and its absence in fossil records, to its phylogenetically close relationship with D. r. distincta and its recent evolutionary origin. The relatively rapid range expansion of D. r. bugensis in eastern Europe during the past several decades was facilitated by human-mediated ecosystem transformation, notably impoundment of large eastern European rivers, that have allowed this species to utilize newly transformed ecosystems.     

A Review of the Biology and Ecology of the Quagga Mussel (Dreissena bugensis), a Second Species of Freshwater Dreissenid Introduced to North America

SYNOPSIS. North America's Great Lakes have recently been invadedby two genetically and morphologically distinct species of Dreissena.The zebra mussel (Dreissena polymorpha) became established inLake St. Clair of the Laurentian Great Lakes in 1986 and spreadthroughout eastern North America. The second dreissenid, termedthe quagga mussel, has been identified as Dreissena bugensisAndrusov, 1897. The quagga occurs in the Dnieper River drainageof Ukraine and now in the lower Great Lakes of North America.In the Dnieper River, populations of D. polymorpha have beenlargely replaced by D. bugensis; anecdotal evidence indicatesthat similar trends may be occurring in the lower LaurentianGreat Lakes. Dreissena bugensis occurs as deep as 130 m in theGreat Lakes, but in Ukraine is known from only 0–28 m.Dreissena bugensis is more abundant than D. polymorpha in deeperwaters in Dneiper River reservoirs. The conclusion that NorthAmerican quagga mussels have a lower thermal maximum than zebramussels is not supported by observations made of populationsin Ukraine. In the Dnieper River drainage, quagga mussels areless tolerant of salinity than zebra mussels, yet both dreissenidshave acclimated to salinities higher than North American populations;eventual colonization into estuarine and coastal areas of NorthAmerica cannot be ignored.     

Canal construction destroys the barrier between major European invasion lineages of the zebra mussel

Since the mid-1980s the zebra mussel, Dreissena polymorpha, Pallas 1771, has become the protagonist of a spectacular freshwater invasion in North America due to its large economic and biological impact. Several genetic studies on American populations have failed to detect any large-scale geographical patterns. In western Europe, where D. polymorpha has been a classical invader from the Pontocaspian since the early 19th century, the situation is strikingly different. Here, we show with genetic markers that two major western European invasion lineages with lowered genetic variability within and among populations can be discriminated. These two invasion lineages correspond with two separate navigable waterways to western Europe. We found a rapid and asymmetrical genetic interchange of the two invasion lines after the construction of the Main-Danube canal in 1992, which interconnected the two waterways across the main watershed.     

Genetic variability and phylogeographical patterns of a nonindigenous species invasion: a comparison of exotic vs. native zebra and quagga mussel populations

There have been few investigations of the number of founding sources and amount of genetic variability that lead to a successful nonindigenous species invasion, although genetic diversity is believed to play a central role. In the present study, population genetic structure, diversity and divergence patterns were analysed for the zebra mussel Dreissena polymorpha [n=280 samples and 63 putative randomly amplified polymorphic DNA (RAPDs) gene loci] and the quagga mussel D. bugensis (n=136 and 52 loci) from 10 nonindigenous North American and six Eurasian sampling sites, representing their present‐day ranges. Results showed that exotic populations of zebra and quagga mussels had surprisingly high genetic variability, similar to those in the Eurasian populations, suggesting large numbers of founding individuals and consistent with the hypothesis of multiple colonizations. Patterns of genetic relationships indicate that the North American populations of D. polymorpha likely were founded by multiple source populations from north‐western and northcentral Europe, but not from southcentral or eastern Europe. Sampling areas within North America also were significantly divergent, having levels of gene flow and migration about twice those separating long‐established Eurasian populations. Samples of D. bugensis in Lakes Erie and Ontario were significantly different, with the former being more closely related to a native population from the Dnieper River, Ukraine. No evidence for a founder effect was discerned for either species.     

Range expansion of quagga mussels Dreissena rostriformis bugensis in the Volga River and Caspian Sea basin

In 1992, we discovered populations of the nonindigenous quagga mussel Dreissena rostriformis bugensis in the middle reaches of the Volga River. The same species was found in samples collected between 1994 and 1997 in the Volga delta and in shallow areas of the Northern Caspian Sea. D. r. bugensis always co-occurred with its more widespread congener, the zebra mussel D. polymorpha (Pallas 1771). The quagga mussel's contribution to total Dreissena abundance increased over time in the middle Volga reservoirs and Volga River delta. D. r. bugensis was common in the Volga portion of Rybinsk Reservoir during 1997 and, by 2000, it was in Uglich, Rybinsk and Gorky Reservoirs on the Upper Volga River. D. r. bugensis was neither found in Ivankov Reservoir, nor in terminal sections of the Volga-Baltic corridor including the eastern Gulf of Finland. Presently, all but the northern-most regions of the Volga River have been colonized by D. r. bugensis. We hypothesize that its introduction into the Volga River and Caspian basin occurred no later than the late 1980s via commercial shipping that utilized the Volga-Don waterway to navigate between the source Black-Azov Sea region and recipient areas on the Volga River. Larval drift likely contributed to establishment of populations at downstream sites, while human-mediated vectors may be responsible for introductions to upstream locations on the Volga River. We anticipate continued northward dispersal in conjunction with shipping activities.     

Predator‐induced morphological defences in two invasive dreissenid mussels: implications for species replacement

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Low diversity but high differentiation: the population genetics of Aglaope infausta (Zygaenidae: Lepidoptera)

Aim Aglaope infausta is a thermophilous Zygaenid of Atlanto-Mediterranean origin, distributed in Portugal, Spain, France and north-western Italy reaching its north-eastern distribution limit in western Germany. The local, regional and inter-regional genetic structure of this species is studied in this analysis. Location and methods The allozymes of 456 individuals from 12 populations (11 from western Germany and one from southern Portugal) were studied by electrophoresis. Results Six of the 19 loci analysed were monomorphic. Genetic differentiation between populations was high (F_ST: 0.404), while the mean genetic diversity was low (H_e: 3.4%). Most (96.5%) of the genetic variance between populations was between the Portuguese and the German samples, but also the differentiation within Germany was considerable (F_SR: 0.101). In Germany, A. infausta occurs in two major regions (middle Rhine and Nahe) that are geographically separated, and 55.5% of the genetic variance was found between these two regions. The populations of both areas do not differ in their genetic diversity, but those of the middle Rhine have significantly higher genetic distances among them than the Nahe populations (0.020 and 0.015, respectively). F_ST was also higher in the middle Rhine region than in the Nahe region (0.089 and 0.045, respectively). Main conclusions Aglaope infausta shows a very low level of genetic heterogeneity for a lepidopteran species. However, this genetic poverty is not affecting the species’ viability. During the ice ages, differentiation into two genetic lineages occurred, most probably in a south-western and a south-eastern differentiation centre in Iberia. The gap in the distribution range in Germany is clearly reflected in the genetic structure. This differentiation must have developed relatively quickly because western Germany most probably was colonized during the climatic optimum 6000 years ago.     

Natural hybridization of two mussel species <Emphasis Type="Italic">Dreissena polymorpha</Emphasis> (Pallas, 1771) and <Emphasis Type="Italic">Dreissena bugensis</Emphasis> (Andrusov, 1897)

Dreissenids display a high diversity of shell morphology, and it is frequently difficult to ascribe some individuals from mixed populations to one of the two species, Dreissena polymorpha (Pallas, 1771) or D. bugensis (Andrusov, 1897). Presumably, such individuals may be interspecific hybrids. We have analyzed species-specific allozyme loci of the typical representatives of these two mussel species and putative interspecific hybrids. A natural interspecific hybrid between D. polymorpha and D. bugensis was discovered for the first time by genetic methods. It has been demonstrated that D. bugensis was a maternal parent.     

Invasive tall annual willowherb (<Emphasis Type="Italic">Epilobium brachycarpum</Emphasis> C. Presl) in Central Europe originates from high mountain areas of western North America

Identification of the source population of biological invasions has important consequences for the effective control and management of the invader. Tall annual willowherb (Epilobium brachycarpum) is a relatively recent and rapidly spreading neophyte in Europe that was first detected in 1978. Populations of tall annual willowherb from Germany and northern France were analysed by AFLP fingerprinting together with samples from five different localities in its native range in western North America. Three genetically different groups were found corresponding to different altitude zones in the native range. The F_ST is high among all samples indicating a strong genetical separation of the three groups. Invasive populations showed much lower genetic diversity than the native population. Additionally invasive populations revealed genetic affinities to North American specimens originating particularly from high mountain areas. The two large German populations and the population from northern France are genetically distinct while the individuals within the populations are genetically uniform. This suggests multiple introduction events rather than one introduction with consequent spreading across Europe. A third small German population from Treis-Karden in the Mosel valley clusters with North American lowland populations but suffers from frost damage and its permanent establishment is doubtful.     

Comparison of nutrition range in <Emphasis Type="Italic">Dreissena polymorpha</Emphasis> and <Emphasis Type="Italic">Dreissena bugensis</Emphasis> mussels by biochemical markers

Based on biochemical markers, we studied the nutrition ranges of populations of Dreissena polymorpha and Dreissena bugensis mussels that inhabit the Volga reach of the Rybinsk reservoir and differ in the long-term population dynamics. Features of the nutrition ranges of mussels are regarded as the probable cause of changes in the number of mollusca.     

Invasion phylogeography of the Ponto-Caspian crustacean Limnomysis benedeni dispersing across Europe

Aim  Limnomysis benedeni Czerniavsky, 1882 is a mysid crustacean native to the Ponto-Caspian (Black and Caspian Sea) rivers and estuaries, and has recently spread across Europe through intentional and unintentional introductions. We explored the structuring of genetic variation in native and non-native populations with an aim to trace the sources of the invasions, and to infer whether the spread has occurred through a single or multiple invasion waves.
Location  Native estuaries in the Ponto-Caspian basin (Volga, Don, Dnieper, Dniester, Danube) and the recently colonized range along the Danube–Rhine river systems and Lithuania.
Methods  A fragment of the mitochondrial COI gene was sequenced to assess genetic affinities and diversity in native and recently established populations.
Results  The genetic diversity in the native regions is organized into several strongly diverged haplotype groups or lineages, partly allopatric, partly sympatric. All these lineages have also spread beyond the native range. Even the recent rapid dispersal across Europe along the Danube–Rhine system towards the North Sea basin involved several lineages from the Danube delta sector. The structuring of genetic diversity among invaded sites suggests multiple invasion events to the Danube–Rhine drainage. This contrasts with data from some other Ponto-Caspian species, where a single haplotype seems to have occupied most invaded areas. There is no evidence that intentionally stocked reservoirs in the Baltic Sea basin would have contributed to further unintentional spread of L. benedeni.
Main conclusions  Limnomysis benedeni is spreading across Europe using the southern invasion corridor. The invasion most likely involved several waves from differentiated sources in the native Danube delta area.     

Genetic differentiation of European grayling populations across the Main, Danube and Elbe drainages in Bavaria

Analysis of mitochondrial ( ND-1/3/4 genes) and nuclear ( GH-1 gene, four microsatellite loci) DNA markers of European grayling Thymallus thymallus populations identified strong genetic differentiation between the Rhine/Main, Elbe and Danube drainages in Bavaria, as a result of both present processes and past history. The Danube and the Main-Elbe group diverged about five times earlier than the populations of Main and Elbe. These data suggest that exchange and transfer of grayling stocking material between these drainages should be avoided, to maintain the genetic diversity and integrity of populations.     

Transformation of the offshore benthic community in Lake Michigan: recent shift from the native amphipod Diporeia spp. to the invasive mussel Dreissena rostriformis bugensis

1. The native amphipod Diporeia spp. was once the dominant benthic organism in Lake Michigan and served as an important pathway of energy flow from lower to upper trophic levels. Lake‐wide surveys were conducted in 1994/1995, 2000 and 2005, and abundances of Diporeia and the invasive bivalves Dreissena polymorpha (zebra mussel) and Dreissena rostriformis bugensis (quagga mussel) were assessed. In addition, more frequent surveys were conducted in the southern region of the lake between 1980 and 2007 to augment trend interpretation. 2. Between 1994/1995 and 2005, lake‐wide density of Diporeia declined from 5365 to 329 m⁻², and biomass (dry weight, DW) declined from 3.9 to 0.4 g DW m⁻². The percentage of all sites with no Diporeia increased over time: 1.1% in 1994/1995, 21.7% in 2000 and 66.9% in 2005. On the other hand, total dreissenid density increased from 173 to 8816 m⁻², and total biomass increased from 0.4 to 28.6 g DW m⁻². Over this 10‐year time period, D. r. bugensis displaced D. polymorpha as the dominant dreissenid, comprising 97.7% of the total population in 2005. In 2007, Diporeia was rarely found at depths shallower than 90 m and continued to decline at greater depths, whereas densities of D. r. bugensis continued to increase at depths greater than 50 m. 3. The decline in Diporeia occurred progressively from shallow to deep regions, and was temporally coincident with the expansion of D. polymorpha in nearshore waters followed by the expansion of D. r. bugensis in offshore waters. In addition, Diporeia density was negatively related to dreissenid density within and across depth intervals; the latter result indicated that dreissenids in shallow waters remotely influenced Diporeia in deep waters. 4. With the loss of Diporeia and increase in D. r. bugensis, the benthic community has become a major energy sink rather that a pathway to upper trophic levels. With this replacement of dominant taxa, we estimate that the relative benthic energy pool increased from 17 to 109 kcal m⁻² between 1994/1995 and 2005, and to 342 kcal m⁻² by 2007. We project that previously observed impacts on fish populations will continue and become more pronounced as the D. r. bugensis population continues to expand in deeper waters.     

Using DNA barcoding to differentiate invasive Dreissena?species (Mollusca,Bivalvia)

The zebra mussel (Dreissena polymorpha) and the quagga mussel (Dreissena rostriformis bugensis) are considered as the most competitive invaders in freshwaters of Europe and North America. Although shell characteristics exist to differentiate both species, phenotypic plasticity in the genus Dreissena does not always allow a clear identification. Therefore, the need to find an accurate identification method is essential. DNA barcoding has been proven to be an adequate procedure to discriminate species. The cytochrome c oxidase subunit I mitochondrial gene (COI) is considered as the standard barcode for animals. We tested the use of this gene as an efficient DNA barcode and found that it allow rapid and accurate identification of adult Dreissena individuals.     

Molecular ecology of zebra mussel invasions

The invasion of the zebra mussel, Dreissena polymorpha, into North American waters has resulted in profound ecological disturbances and large monetary losses. This study examined the invasion history and patterns of genetic diversity among endemic and invading populations of zebra mussels using DNA sequences from the mitochondrial cytochrome oxidase I (COI) gene. Patterns of haplotype frequency indicate that all invasive populations of zebra mussels from North America and Europe originated from the Ponto-Caspian Sea region. The distribution of haplotypes was consistent with invasive populations arising from the Black Sea drainage, but could not exclude the possibility of an origin from the Caspian Sea drainage. Similar haplotype frequencies among North American populations of D. polymorpha suggest colonization by a single founding population. There was no evidence of invasive populations arising from tectonic lakes in Turkey, while lakes in Greece and Macedonia contained only Dreissena stankovici. Populations in Turkey might be members of a sibling species complex of D. polymorpha. Ponto-Caspian derived populations of D. polymorpha (theta = 0.0011) and Dreissena bugensis (one haplotype) exhibited low levels of genetic diversity at the COI gene, perhaps as a result of repeated population bottlenecks. In contrast, geographically isolated tectonic lake populations exhibited relatively high levels of genetic diversity (theta = 0.0032 to 0.0134). It is possible that the fluctuating environment of the Ponto-Caspian basin facilitated the colonizing habit of invasive populations of D. polymorpha and D. bugensis. Our findings were concordant with the general trend of destructive freshwater invaders in the Great Lakes arising from the Ponto-Caspian Sea basin.     

Tracing recent invasions of the Ponto-Caspian mysid shrimp Hemimysis anomala across Europe and to North America with mitochondrial DNA

The mysid crustacean Hemimysis anomala ('bloody-red shrimp') is one of the most recent participants in the invasion of European inland waters by Ponto-Caspian species. Recently the species also became established in England and the Laurentian Great Lakes of North America. Using information from mitochondrial cytochrome oxidase I (COI) gene sequences, we traced the invasion pathways of H. anomala ; the inferences were enabled by the observed phylogeographical subdivision among the source area populations in the estuaries of the Ponto-Caspian basin. The data distinguish two routes to northern and western Europe used by distinct lineages. One route has been to and through the Baltic Sea and further to the Rhine delta, probably from a population intentionally introduced to a Lithuanian water reservoir from the lower Dnieper River (NW Black Sea area) in 1960. The other lineage is derived from the Danube delta and has spread across the continent up the Danube River and further through the Main–Danube canal down to the Rhine River delta. Only the Danube lineage was found in England and in North America. The two lineages appear to have met secondarily and are now found intermixed at several sites in NW Europe, including the Rhine and waters linked with the man-made Mittellandkanal that interconnects the Rhine and Baltic drainage systems.     

Genetic variation of Eryngium campestre L. (Apiaceae) in Central Europe

In Germany, Eryngium campestre is restricted to dry habitats along the rivers Rhine and Elbe and to a few areas in Central Germany. This distribution pattern is usually regarded as a typical pattern of postglacial immigration. In the present study, we investigated whether these two geographically distinct distribution areas are genetically differentiated and whether conclusions can be drawn regarding colonization history. To analyse the phylogeographic structure of E. campestre in Central Europe, 278 individuals from 29 populations within Germany and from further reference populations within Europe were analysed. We applied amplified fragment length polymorphisms to examine their genetic relatedness. Our analyses revealed three groups: a Mediterranean group additionally including two Rhine populations; a Rhine–Main group which further includes the westernmost population from the central German dry area; and one group which includes all eastern populations. Our results show that the two geographically distinct areas are genetically differentiated. As genetic diversity within the Elbe populations is very low, we conclude that this area, which was strongly affected through the late glacial maximum, was colonized relatively recently. High genetic diversity in the Rhine populations indicates a contact zone where lineages of different origin met. This would imply that today's patterns of genetic variation were caused through glacial range contractions and expansions. The present study is one of the first studies that deal with the postglacial distribution pattern of a dry grassland plant species in Central Europe and the results suggest that a survival of E. campestre at least during the Dryas cold stage might be possible.