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.     

Dispersal of the Ponto-Caspian amphipod Echinogammarus ischnus: invasion waves from the Pleistocene to the present

The geographical range of the amphipod crustacean Echinogammarus ischnus has expanded over the past century from the Ponto-Caspian region to Western Europe, the Baltic Sea, and the Great Lakes of North America. The present study explores the phylogeographic patterns of this amphipod across its current distribution, based on an examination of nucleotide diversity in the mitochondrial cytochrome c oxidase subunit I (COI) gene. Marked genetic divergence exists among populations of E. ischnus from the Black and Caspian Seas, as well as those from the drainage system of the Black Sea. This divergence suggests the prolonged geographic isolation of these native populations, reflecting the limited dispersal capability of E. ischnus. By contrast, invading populations are characterized by a lack of genetic variation; a single mitochondrial genotype of Black Sea origin has colonized sites from the Rhine River to North America. The dispersal pattern in E. ischnus is very similar to that in the Ponto-Caspian cladoceran Cercopagis pengoi. Despite their contrasting life history strategies, these invading species followed the same route of invasion from the northern Black Sea to the Baltic Sea region, and subsequently to North America.     

Human-mediated lineage admixture in an expanding Ponto-Caspian crustacean species <Emphasis Type="Italic">Paramysis lacustris</Emphasis> created a novel genetic stock that now occupies European waters

Identifying the origins and colonization routes of alien species is critical to understanding the mechanisms, consequences and management of invasions. This study aims to demonstrate and explain conflicting genetic signals in a widespread invasive Ponto-Caspian mysid Paramysis lacustris using mitochondrial DNA and microsatellite markers. Native Black Sea populations of P. lacustris are divided into three distinct phylogeographic groups: Danube–Dniester region, coastal Danube lakes, and Dnieper River delta. In the 1950s–1970s the species has been extensively used for relatively well documented transplantations in Eastern Europe. However, the populations in the newly established distribution area do not match any of the known native groups, and their genetic signal does not correspond with the recorded transplantation history. Using Bayesian Approximate Computation we explore alternative scenarios that could explain the unexpected mitochondrial and nuclear signals in the introduced populations. We suggest that reservoir construction and break-up of natural dispersal barriers on the River Dnieper have brought about a novel stock, probably admixed from a previously isolated and unknown “relict” population in the middle Dnieper and the genetically distinct Dnieper delta lineage. Through further intentional transfers this new P. lacustris lineage is now widely present in Eastern Europe and is likely to have new gene and trait combinations. The conflicting mitochondrial-nuclear signal in non-native populations also highlights the importance of using multiple genetic markers when tracing invasion sources and pathways.     

Postglacial colonization of Northern Europe by distinct phylogeographic lineages of the bullhead, Cottus gobio

Three major phylogeographic lineages of the cottid fish Cottus gobio (bullhead) were identified in northern Europe from mitochondrial DNA sequences and allozyme data. The largely separate freshwater distributions of the lineages demonstrate distinct postglacial colonization histories. West of the Baltic Sea, Swedish lakes were invaded from the southwest (Germany). Another, eastern lineage has colonized the inland waters northeast and east of the Baltic, from refugia in northwest Russia; this lineage comprises a distinct subgroup found only from Estonia. The third lineage, found south and southeast of the Baltic, probably descended from rivers draining to the Black Sea from the north (e.g. Dnepr). In coastal waters of the Baltic Sea, and in near-coast inland waters, the lineages are now found intermixed in various combinations. The alternating fresh- and saltwater phases of the Baltic basin have variously enabled and disabled the use of coastal waters as colonization routes. Hypotheses on the chronology of dispersal and lineage mixing can be based on the distribution of the marker genes and the paleohydrographical record. The diversity of the Fennoscandian bullhead thus comprises anciently diverged (probably mid-Pleistocene) refugial lineages that in their freshwater range constitute distinct evolutionarily significant units. The thorough mixing of the various genomic origins in and around the Baltic, however, refutes the controversial view of distinct species status for the western and eastern ('Cottus koshewnikowi') bullheads. The postglacial contact of the lineages has created new diversity that cannot be interpreted in a conventional hierarchical framework of taxonomic or conservation units.     

Distribution and invasion genetics of the quagga mussel (<Emphasis Type="Italic">Dreissena rostriformis bugensis</Emphasis>) in German rivers

The quagga mussel Dreissena rostriformis bugensis, native to the Dnieper and the northern Black Sea, has become a major invasive species in both the Volga River and the North American Great Lakes since the early 1990s. Findings in the Netherlands (2006) and Germany (2007) mark the start of its establishment in Western Europe. We investigated the current distribution, time of first arrival and population structure of D. rostriformis bugensis from the rivers Rhine, Main and in the Main-Danube canal in Germany. Two putative sources of the German populations were analysed by genetically comparing these populations to older invasive populations from North America and the southeast Danube. Dreissena rostriformis bugensis was abundant in the Main and in three Rhine harbours, but rare in the actual Rhine river and absent south of the Main-Danube canal. Mussels found in the Rhine harbours were significantly smaller than in the Main. Population genetic analyses found no sign of founder effects and minimal differentiation between German, North American and southeast Danube populations. The genetic data suggest that these invasive populations derive from a common and rapidly expanding source. Based on the non-continuous distribution and shell size differences of Rhine harbour and Main populations, our results indicate that expansion in Germany involved at least two independent settling events, one of which happened before 2005, and most likely was caused by jump dispersal.     

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.     

Phylogeography and postglacial dispersion of the chub (Leuciscus cephalus) in Europe

A phylogeographic analysis of mitochondrial DNA variation was performed in order to test the hypothesis of a postglacial recolonization of mid- and north-European rivers from a Danubian refuge. Over 345 chub specimens from European rivers covering most of the species' native range were investigated using 600 bp of the cytochrome b gene. Chub in European rivers belong to four highly divergent mitochondrial groups (lineages) differing by mean divergence estimates from 5.2% to 7.89%. These four lineages have a largely allopatric distribution, implying four geographical sets: two Mediterranean, and two north-European sets. This pattern provided strong evidence for: (i) the eradication of this species from most of Europe during maximum ice extent; (ii) its survival in four refugia (Adriatic side of the Balkans, eastern Greece (Aegean Rivers), southern tributaries of the Danube, and periphery of Black and Caspian Seas); (iii) a differential postglacial recolonization of mid- and northern Europe from the last two refugia only; (iv) the occurrence of this recolonization in two steps for the Danubian (western) lineage that entered western Europe (Rhine-Rhone-Loire drainages) during the Riss-Würm interglacial period and survived the last glaciation there before colonizing Garonne, UK and German drainages up to the Elbe during the Holocene; and (v) the occurrence of this recolonization in a single step for the Ponto-Caspian (eastern) lineage that entered the Baltic area as far as the Oder in the Holocene. Both lineages came into contact in the River Elbe without evident mixing.     

An eDNA Assay to Monitor a Globally Invasive Fish Species from Flowing Freshwater

Ponto-Caspian gobies are a flock of five invasive fish species that have colonized freshwaters and brackish waters in Europe and North America. One of them, the round goby Neogobius melanostomus, figures among the 100 worst invaders in Europe. Current methods to detect the presence of Ponto-Caspian gobies involve catching or sighting the fish. These approaches are labor intense and not very sensitive. Consequently, populations are usually detected only when they have reached high densities and when management or containment efforts are futile. To improve monitoring, we developed an assay based on the detection of DNA traces (environmental DNA, or eDNA) of Ponto-Caspian gobies in river water. The assay specifically detects invasive goby DNA and does not react to any native fish species. We apply the assay to environmental samples and demonstrate that parameters such as sampling depth, sampling location, extraction protocol, PCR protocol and PCR inhibition greatly impact detection. We further successfully outline the invasion front of Ponto-Caspian gobies in a large river, the High Rhine in Switzerland, and thus demonstrate the applicability of the assay to lotic environments. The eDNA assay requires less time, equipment, manpower, skills, and financial resources than the conventional monitoring methods such as electrofishing, angling or diving. Samples can be taken by untrained individuals, and the assay can be performed by any molecular biologist on a conventional PCR machine. Therefore, this assay enables environment managers to map invaded areas independently of fishermen’s’ reports and fish community monitorings.     

Cryptic species in marine polychaete and their independent introduction from North America to Europe

The vast body of ballast water carried across oceans by freight ships represents a major source for the introduction of foreign species into marine ecosystems. The worm Marenzelleria viridis, originally found only in North America, appeared in estuaries of the North Sea in 1979 and 6 years later also in the Baltic, where it has developed into a major faunal element. Two competing hypotheses are discussed here: either both populations owe their presence to a single introductory event in the North Sea, or each population originated from a separate introduction. Our phylogeographic analysis of Baltic, North Sea and American Marenzelleria, based on mitochondrial 16S rDNA sequences (326- bp segment) of 98 individuals from 17 localities on the North American, North Sea, and Baltic coasts not only favors the two-event hypothesis, but also separates the locations of origin for the introductions. Eighteen mitochondrial genotypes were identified altogether. In agreement with allozyme data, three lineages were identified: genotypes assigned to the same lineage differed from each other by up to 5 point mutations, and those assigned to different lineages differed by up to 17. The existence of three morphologically indistinguishable, and thus cryptic, species is therefore suggested. The individuals from the Baltic Sea probably originated from the Atlantic coast of the United States between Chesapeake Bay and Georgia, and the North Sea populations may stem from the U.S. coast region north of Chesapeake Bay to Nova Scotia. Despite their similar morphologies, the two European Marenzelleria species may differ ecologically with respect to their preference for habitat salinity. Assuming that transport via ballast water occurs quite frequently, we hypothesize that both European cryptic species of Marenzelleria may originally have been introduced to both the North Sea and the Baltic Sea but that neither of them was able to proliferate in both water bodies owing to their differential physiological performances at high and low salinities.      

Restoration strategies for river floodplains along large lowland rivers in Europe

1. Most temperate rivers are heavily regulated and characterised by incised channels, aggradated floodplains and modified hydroperiods. As a consequence, former extensive aquatic/terrestrial transition zones lack most of their basic ecological functions.
2. Along large rivers in Europe and North America, various floodplain restoration or rehabilitation projects have been planned or realised in recent years. However, restoration ecology is still in its infancy and the literature pertinent to river restoration is rather fragmented. (Semi-) aquatic components of floodplains, including secondary channels, disconnected and temporary waters as well as marshes, have received little attention, despite their significant contribution to biological diversity.
3. Many rehabilitation projects were planned or realised without prior knowledge of their potential for success or failure, although, these projects greatly contributed to our present understanding of river–floodplain systems.
4. River rehabilitation benefits from a consideration of river ecosystem concepts in quantitative terms, comparison with reference conditions, historical or others, and the establishment of interdisciplinary partnerships.
5. We present examples from two large European rivers, the Danube and the Rhine, in which the role of aquatic connectivity has been extensively studied. The Danube delta with its diversity of floodplain lakes across an immense transversal gradient (up to 10 km) serves as a reference system for restoration projects along lowland sections of large rivers such as the Rhine in the Netherlands.     

The round goby (Neogobius melanostomus) in the Gulf of Gdańsk – a species introduction into the Baltic Sea

In recent years, information concerning the awareness of organisms accidentally introduced into the Baltic Sea has substantially improved. Non-indigenous Estuarine and Marine Organisms (NEMO's) are hazardous for the Baltic ecosystem. Currently, about one hundred species are identified as accidentally or intentionally introduced into the Baltic Sea. Ballast waters and escape from aquaculture are the most important invasion vectors. During the last decade, an invasion of the round goby (Neogobius melanostomus) – a Ponto-Caspian fish species has been observed in the Gulf of Gdańsk. The first record of this fish in the Baltic Sea is from 1990. Early detection of the invader enabled the study of population growth and changes in the area. The first years of invasion were characterized by low numbers of individuals and a limited distribution. Later, the round goby gradually colonized all shallow waters in the western part of the Gulf of Gdańsk. Initially the fish inhabited stony and rocky habitats, but later it also occupied sandy bottoms. The round goby is now the dominating fish species in most of the shallow waters of the Gulf of Gdańsk. Two main factors account for the successful invasion of this fish in the region: the state of ecosystem at the time of the invasion and the biological features of N. melanostomus. In the late 1980s, the shallow waters of the Gulf of Gdańsk were almost devoid of piscivorous fishes. Concurrently, bivalves (a preferred prey of the round goby) have increased. Important is also parental care of laid eggs and reproductive strategy. Population growth potential enables the colonization of nearby regions. The first round gobies in the Vistula Lagoon were collected in 1999 and colonization of other Baltic Sea areas is anticipated.     

First report of the oriental shrimp <Emphasis Type="Italic">Palaemon macrodactylus</Emphasis> Rathbun, 1902 (Decapoda,Caridea, Palaemonidae) from German waters

The native East Asian shrimp Palaemon macrodactylus has become a common inhabitant of estuaries along the Pacific coast of North America. More recently (documented since 1999), the species has also been colonising European waters and has been reported from Spain, England, Belgium and the Netherlands. In this study, we present a chronology of the reported introductions of this species and provide the first detailed report of its occurrence in German waters. P. macrodactylus was found in the Geeste river mouth (Weser Estuary) as well as in Hooksiel, north of Wilhelmshaven between 2004 and 2005. We assume its presence in other estuarine habitats of the North Sea and predict its introduction into the Baltic Sea.     

Distribution and dispersal capacity of the Ponto-Caspian tubificid oligochaete Potamothrix moldaviensis Vejdovský et Mrázek, 1903 in the Baltic Sea Region

Over the last few centuries, several Ponto-Caspian tubificid oligochaetes have gradually dispersed from the Black Sea – Caspian Sea region to the north-west and west over Europe. The present world distribution comprising also the Great Lakes of North America clearly demonstrates that anthropochorous vectors of dispersal are involved. Passive transportation in the ballast water of ships has radically changed the possibilities of dispersal for many invertebrate species and has even made dispersal between continents possible. The construction of navigable canals has furthermore facilitated the crossing of watersheds and continents. Other likely vectors of longway dispersal for oligochaetes, as well as for other small-size aquatic invertebrates, are birds and mammals. The dispersal of the Potamothrixspecies is likely to have taken place in successive waves (three) with front-lines still on the move from the east to the west over the Baltic States and Scandinavia. The rheophilous species Potamothrix moldaviensishas presently reached – apart from the large rivers of Russia and many Central-European water bodies – also the Baltic States and south-eastern Sweden. Trajectories of dispersal demonstrate routes across the Baltic Sea – via the ballast water of ships. In the largest rivers of the Eastern Baltic Region (Neva, Daugava, Nemunas), downstream dispersal is the most likely way of transportation. P. moldaviensis together with P. heuscheri(second wave) and P. vejdovskyi(third wave) are presently forming front-lines running obliquely from the north-east to the south-west over south Sweden. In mesotrophic-eutrophic basins of eastern Lake Mälaren, the abundance as well as the species diversity of oligochaetes is particularly high wherever Ponto-Caspian Potamothrixspecies – often several species together – are involved.     

Microsatellites reveal origin and genetic diversity of Eurasian invasions by one of the world’s most notorious marine invader,Mnemiopsis leidyi (Ctenophora)

Marine invasions are taking place at an increasing rate. When occurring in blooms, zooplanktivorous comb jellies of the genus Mnemiopsis are able to cause pelagic regime shifts in coastal areas and may cause the collapse of commercially important fish populations. Using microsatellites, developed for the first time in the phylum Ctenophora, we show that Mnemiopsis leidyi has colonized Eurasia from two source regions. Our preliminary data set included four sites within the putative source region (US East Coast and Gulf of Mexico) and 10 invaded locations in Eurasian waters. Bayesian clustering and phylogeographic approaches revealed the origin of earlier invasions of the Black and Caspian Sea in the 1980s/1990s within or close to the Gulf of Mexico, while the 2006 invasion of the North and Baltic Seas can be directly traced to New England (pairwise F_ST = 0). We found no evidence for mixing among both gene pools in the invaded areas. While the genetic diversity (allelic richness) remained similar in the Baltic Sea compared to the source region New England, it was reduced in the North Sea, supporting the view of an initial invasion of Northern Europe to a Baltic Sea port. In Black and Caspian Sea samples, we found a gradual decline in allelic richness compared to the Gulf of Mexico region, supporting a stepping‐stone model of colonization with two sequential genetic founder events. Our data also suggest that current practices of ballast water treatment are insufficient to prevent repeated invasions of gelatinous zooplankton.     

Timing of downstream migration and food uptake of juvenile North Sea houting stocked in the Lower Rhine and the Lippe (Germany)

In 1992 a stocking programme was established to re‐introduce North Sea houting Coregonus oxyrinchus , which disappeared from the River Rhine in the 1940s, in the Lower Rhine. Juvenile North Sea houting have been stocked since 2001 in a gravel pit lake permanently connected to the Lower Rhine, and in the Lippe, a potamal tributary of the Rhine. Monitoring studies showed that the majority of the small North Sea houting left the gravel pit lake within 4 days after stocking. In the Lippe, some of the juveniles were found in the drift immediately after stocking, generally preferring the middle surface areas of the river. Other North Sea houting waited until dawn before they started their downstream migration. Juveniles immediately started to feed on the zooplankton resources in both waters. Pond studies revealed high growth rates of juvenile North Sea houting, ranging from 0·44 to 0·94 mm day⁻¹. This study suggests that the stocking strategies at both waters of the Lower Rhine comply with the requirements of the migration behaviour of juvenile North Sea houting.     

European colonization by the spined loach (Cobitis taenia) from Ponto-Caspian refugia based on mitochondrial DNA variation

In the last 20 years, new species, asexual reproduction, polyploidy and hybridization have all been reported within the genus Cobitis. An understanding of the current distribution and baseline phylogeographical history of 'true' nonhybrid Cobitis species is crucial in order to unravel these discoveries. In the present work, we investigated the phylogeography of the spined loach, Cobitis taenia, using 1126 bp of the mitochondrial cytochrome b gene from 174 individuals collected at 47 sites. In total, 51 haplotypes that differed at 49 positions (4.35%) were detected. We deduce that C. taenia survived European glaciations in at least three refugees in the Ponto-Caspian area. Two of these refugees each provided a major lineage that recolonized Europe in separate directions: one westward to England and the other spreading north into Russia before moving west. A third (minor) lineage that contributed little to the recolonization of Europe was also revealed--remaining near its Black Sea refuge. However, more recent history was difficult to resolve with colonization from a more western refugium during the last glacial maximum (LGM) a distinct possibility. Nested clade analysis indicates a pattern of restricted gene flow with isolation by distance at the first two levels and overall. Unlike many other European freshwater fish species, the Danube is not part of the current distribution of C. taenia, nor was it used as either a refuge or a source of colonization of Europe. Low genetic diversity within C. taenia suggests that its colonization of Europe is relatively recent. Demographic analyses revealed a history of recent expansion and isolation by distance.     

Reappraisal and range extension of non-indigenous Mysidae (Crustacea, Mysida) in continental and coastal waters of eastern France

A survey of species belonging to the family Mysidae, conducted in June 2007 in fresh- to brackish waters of eastern France, revealed a recent range extension of the invasive Ponto-Caspian species Hemimysis anomala to the Moselle, Saône, and Rhône rivers. In the estuary of the Grand Rhône it reached for the first time the Mediterranean coast. The network of navigation canals in NE France was likely a key element of its north to south pathway starting from the Rhine River. Important range extensions were also noted for Limnomysis benedeni in this network and in the Moselle River. The euryhaline species Neomysis integer, endemic in coastal waters of the NE Atlantic, was found in the Rhône delta, thus confirming previous very rare records in the 1930–1950s from the Mediterranean coast of France. Invasion mechanisms and pathways, expansion potential, and establishment conditions of the species are discussed.     

Molecular phylogeny of the genus Gobio Cuvier, 1816 (Teleostei: Cyprinidae) and its contribution to taxonomy

The phylogenetic relationships among gudgeons that represent most nominal taxa within Gobio gobio sensu lato were examined by mitochondrial and nuclear genome sequencing. The molecular analyses confirmed the separate generic status of Gobio as a monophyletic group and revealed 15 Eurasian lineages divided into two main clades, the Northern European and the Ponto-Caspian. The validity of eleven nominal taxa as distinct species was confirmed, gudgeons from the Volga River basin were described as a new species G. volgensis, and three revealed phylogenetic lineages were submitted for a comprehensive revision as "species-in-waiting". The species G. gobio showed a wide range extending from the British Isles to the Black Sea coast and overlapped the areas of several other species. Four pure lineages were detected in the middle Danube River basin. The Crimean Peninsula was found to be a region with the occurrence of individuals of hybrid origin. This region will require special investigation to define species participating in hybridization events, and to establish further steps for the conservation of endemic native gudgeon species. A simple diagnostic method, based on different lengths of the PCR products, called "S7indel diagnostics" is presented for further taxonomic investigations in the genus Gobio.     

Where did Marenzelleria spp. (Polychaeta: Spionidae) in Europe come from?

Abundant populations of Marenzelleria spp. were reported for the first time in the North Sea during the late 1970s and then in the Baltic Sea in 1985. Genetic analysis by means of allozyme electrophoresis and sequencing of a segment of mitochondrial 16srDNA showed that two different genetic types or sibling species of Marenzelleria were present in Europe. Marenzelleria Type I is found only in the North Sea, whereas Type II has been found in both the North Sea and the Baltic Sea. The North Sea animals (Type I) correspond to Type I specimens found in North American coastal waters between Nova Scotia and Cape Henlopen (Delaware), while Marenzelleria Type II from the Baltic Sea correspond to Marenzelleria type II animals from the Arctic (Tuktoyaktuk Harbor, Northwest Territories, Canada), New Hampshire and coastal waters between Chesapeake Bay southward to the Ogeechee River (Georgia). Human mediated introduction (by shipping) and natural range expansion are discussed as the possible causes of these virtually simultaneous invasions by two sibling species. Marenzelleria Type I appears to colonize habitats with a higher salinity and/or in which salinities tend to fluctuate considerably. The osmolality of the coelomic fluid after acclimation to various salinities between 0.25 and 18 is the same for both sibling species/genetic types. Although the two types do not differ in respect of hyperosmoregulation (<10), differences may exist in their cell volume regulation or its time course in the almost isoosmotic range at salinities >10.     

An invasive lineage of sculpins, Cottus sp. (Pisces, Teleostei) in the Rhine with new habitat adaptations has originated from hybridization between old phylogeographic groups

Fish abundance surveys in the Rhine system have shown in the past two decades that there is a rapid upriver invasion of a freshwater sculpin of the genus Cottus. These fish are found in habitats that are atypical for the known species Cottus gobio, which is confined to small cold streams within the Rhine drainage. Phylogeographic analysis based on mitochondrial haplotypes and diagnostic single nucleotide polymorphisms indicates that the invasive sculpins are hybrids between two old lineages from the River Scheldt drainage and the River Rhine drainage, although it is morphologically more similar to the Scheldt sculpins. Most importantly, however, the invasive population possesses a unique ecological potential that does not occur in either of the source populations from the Rhine or the Scheldt, which allows the colonization of new habitats that have previously been free of sculpins. Microsatellite analysis shows that the new lineage is genetically intermediate between the old lineages and that it forms a distinct genetic group across its whole expansion range. We conclude that hybridization between long separated groups has lead to the fast emergence of a new, adaptationally distinct sculpin lineage.