Holocene vegetation and climate change on the Colorado Great Plains,USA, and the invasion of Colorado piñon (Pinus edulis)

Aim To reconstruct the last c. 7000 years of vegetation and climate change in an unusual region of modern Great Plains grassland and scarp woodland in south‐east Colorado (USA), and to determine the late Holocene biogeography of Colorado piñon (Pinus edulis) at its easternmost extent, using a series of radiocarbon‐dated packrat (Neotoma sp.) middens. Location The West Carrizo Canyon drains the Chaquaqua Plateau, a plateau that projects into the western extent of the southern Great Plains grasslands in south‐eastern Colorado, USA. Elevations of the study sites are 1448 to 1525 m a.s.l. Today the plateau is mostly Juniperus scopulorum–P. edulis woodland. Methods Plant macrofossils and pollen assemblages were analysed from 11 ¹⁴C‐dated packrat middens. Ages ranged from 5990 yr bp (6839 cal. yr bp ) to 280 yr bp (485 cal. yr bp ). Results The results presented here provide information on the establishment and expansion of Juniperus–P. edulis woodland at its eastern limits. The analysis of both plant macrofossils and pollen from the 11 middens documents changes in plant communities over the last 7000 years, and the establishment of P. edulis at its easternmost limit. Though very minor amounts of P. edulis pollen occur as early as the middle Holocene, plant macrofossils were only recovered in middens dating after c. 480 cal. yr bp . Main conclusions Originally, midden research suggested a late glacial refuge to the north‐east of the Carrizo Canyon site, and a middle Holocene expansion of P. edulis. Results reported here are consistent with a late Holocene expansion, here at its eastern limits, but noted elsewhere at its northern and north‐eastern limits. In general, this late Holocene expansion is consistent with pollen data from sediments in Colorado and New Mexico, and suggests that P. edulis is still expanding its range at its present extremes. This has implications for further extension of its range due to changing climatic conditions in the future.     

Bioinvasion threatens the genetic integrity of native diversity and a natural hybrid zone: smooth‐shelled blue mussels (Mytilus spp.) in the Strait of Magellan

Smooth‐shelled blue mussels of the Mytilus edulis species complex are widely distributed bivalve molluscs whose introductions threaten native marine biodiversity (non‐indigenous species – NIS). The aim of the present study was to identify the species and hybrids of Mytilus present in the Magellan Region (southern Chile). Results indicate that three mussel species of the Mytilus edulis complex are found in the region – M. edulis, M. chilensis (or the Southern Hemisphere lineage of Mytilus galloprovincialis), and M. galloprovincialis of Northern Hemisphere origin. For the first time, alleles of the introduced M. trossulus are reported from the Southern Hemisphere. In the Strait of Magellan the native Pacific blue mussel, Mytilus chilensis and the native Atlantic blue mussel, Mytilus edulis, meet and mix at a natural hybrid zone (about 125 km in length). This is the first record of a natural Mytilus hybrid zone in the Southern Hemisphere and is also the first record of the co‐occurrence of genes from all four Mytilus species in any one region. These results contribute to the knowledge of the biodiversity and delimitation of mussel species in southern South America, and highlight how introduced species may threaten the genetic integrity of native species through hybridization and introgression.     

Genetic traits in the bivalve Mytilus from Europe, with an emphasis on Arctic populations

Genetic and some ecophysiological traits of mussels collected in the European Arctic, up to their northeastern distribution limit in the Barents Sea, were studied and compared with traits of mussels from the Mediterranean, Atlantic and Baltic. The genetic traits of these populations were analysed by isoenzyme electrophoresis on seven loci in order to assess the Mytilus complex to which populations in the Arctic region belong. Ecophysiological variables, the weight-index and glycogen were analysed to assess the physiological fitness of the populations. Three distinct groups were recognised: (1) Mytilus (edulis) galloprovincialis in the Mediterranean and Spain, (2) M. (edulis) edulis along the Atlantic coast from the Netherlands northwards into Russia, and (3) the Baltic Mytilus (edulis) trossulus. The mussels from populations in the Russian Arctic all belong to the Atlantic Mytilus (edulis) edulis group. The genetic variability and ecophysiological measures indicated that the sub-Arctic White Sea mussel populations have a relatively lower performance capacity, whereas those in the Arctic at the edge of their northern distribution showed a surprisingly strong performance. Accepted: 14 June 2000     

Post-glacial changes in spatial patterns of vegetation across southern New England

Aim We analysed lake‐sediment pollen records from eight sites in southern New England to address: (1) regional variation in ecological responses to post‐glacial climatic changes, (2) landscape‐scale vegetational heterogeneity at different times in the past, and (3) environmental and ecological controls on spatial patterns of vegetation. Location The eight study sites are located in southern New England in the states of Massachusetts and Connecticut. The sites span a climatic and vegetational gradient from the lowland areas of eastern Massachusetts and Connecticut to the uplands of north‐central and western Massachusetts. Tsuga canadensis and Fagus grandifolia are abundant in the upland area, while Quercus, Carya and Pinus species have higher abundances in the lowlands. Methods We collected sediment cores from three lakes in eastern and north‐central Massachusetts (Berry East, Blood and Little Royalston Ponds). Pollen records from those sites were compared with previously published pollen data from five other sites. Multivariate data analysis (non‐metric multi‐dimensional scaling) was used to compare the pollen spectra of these sites through time. Results Our analyses revealed a sequence of vegetational responses to climate changes occurring across southern New England during the past 14,000 calibrated radiocarbon years before present (cal yr bp ). Pollen assemblages at all sites were dominated by Picea and Pinus banksiana between 14,000 and 11,500 cal yr bp ; by Pinus strobus from 11,500 to 10,500 cal yr bp ; and by P. strobus and Tsuga between 10,500 and 9500 cal yr bp . At 9500–8000 cal yr bp , however, vegetation composition began to differentiate between lowland and upland sites. Lowland sites had higher percentages of Quercus pollen, whereas Tsuga abundance was higher at the upland sites. This spatial heterogeneity strengthened between 8000 and 5500 cal yr bp , when Fagus became abundant in the uplands and Quercus pollen percentages increased further in the lowland records. The differentiation of upland and lowland vegetation zones remained strong during the mid‐Holocene Tsuga decline (5500–3500 cal yr bp ), but the pattern weakened during the late‐Holocene (3500–300 cal yr bp ) and European‐settlement intervals. Within‐group similarity declined in response to the uneven late‐Holocene expansion of Castanea, while between‐group similarity increased due to homogenization of the regional vegetation by forest clearance and ongoing disturbances. Main conclusions The regional gradient of vegetation composition across southern New England was first established between 9500 and 8000 cal yr bp . The spatial heterogeneity of the vegetation may have arisen at that time in response to the development or strengthening of the regional climatic gradient. Alternatively, the differentiation of upland and lowland vegetation types may have occurred as the climate ameliorated and an increasing number of species arrived in the region, arranging themselves in progressively more complex vegetation patterns across relatively stationary environmental gradients. The emergence of a regional vegetational gradient in southern New England may be a manifestation of the increasing number of species and more finely divided resource gradient.     

Identification of Mytilus spp. and Pecten maximus in Irish Waters by Standard PCR of the 18S rDNA Gene and Multiplex PCR of the 16S rDNA Gene

Two molecular protocols for the identification of mussel and scallop have been developed using specific primers targeting the mitochondrial 16S ribosomal DNA gene and the nuclear 18S ribosomal DNA gene. Primers for the mitochondrial 16S ribosomal DNA gene in multiplex polymerase chain reaction (PCR) protocols yielded diagnostic DNA fragments for the mussels Mytilus edulis, Mytilus galloprovincialis, and the hybrid Mytilus edulis/galloprovincialis (335 bp), the king scallop Pecten maximus (382 bp) and the black scallop Mimachlamys varia (398 bp). DNA from the queen scallop Aequipecten opercularis showed no consistent PCR amplification of the 16S rDNA gene. Primers for the nuclear 18S rDNA gene in standard PCR protocols yielded similar-sized, diagnostic DNA fragments (approx. 190 bp) for the mussels Mytilus edulis, Mytilus galloprovincialis, and the hybrid Mytilus edulis/galloprovincialis, the king scallop Pecten maximus, the black scallop Mimachlamys varia, and the queen scallop Aequipecten opercularis. Both protocols have been tested with Mytilus spp., P. maximus, and 6 other bivalve species from a wide range of locations in Irish and European waters. Cross reaction of the specific primers with DNA template from any of the 6 other bivalve species was not observed. Rapid DNA extraction using FTA Card technology and the16S rDNA primers allowed for the detection of at least 10 mussel larvae in a subsample of natural plankton.     

A 40,000-year woodrat-midden record of vegetational and biogeographical dynamics in north-eastern Utah, USA

Aim A conspicuous climatic and biogeographical transition occurs at 40–45° N in western North America. This pivot point marks a north–south opposition of wet and dry conditions at interannual and decadal time‐scales, as well as the northern and southern limits of many dominant western plant species. Palaeoecologists have yet to focus on past climatic and biotic shifts along this transition, in part because it requires comparisons across dissimilar records [i.e. pollen from lacustrine sediments to the north and plant macrofossils from woodrat (Neotoma) middens to the south]. To overcome these limitations, we are extending the woodrat‐midden record northward into the lowlands of the central Rocky Mountains. Location Woodrat middens were collected from crevices and rock shelters on south‐facing slopes of Dutch John Mountain (2000–2200 m, 40°57′ N, 109°25′ W), situated on the eastern flanks of the Uinta Mountains in north‐eastern Utah. The site is near the regional limits for Pinus ponderosa, P. edulis, P. contorta, Cercocarpus ledifolius var. intricatus, Abies concolor, Ephedra viridis and other important western species. Methods We analysed pollen and plant macrofossils from the 40,000‐year midden sequence. The middens represent brief, depositional episodes (mostly years to decades). Four middens represent the early to full‐glacial period (40,000–18,000 cal‐yr bp ), eight middens are from the late‐glacial/early Holocene transition (13,500–9000 cal yr bp ), and 33 middens span the mid‐to‐late Holocene (last 7500 years). Temporal density of our Holocene middens (one every c. 210 years) is comparable with typical Holocene pollen sequences from lake sediments. Results Early to full‐glacial assemblages are characterized by low diversity and occurrence of montane conifers (Picea pungens, Pseudotsuga menziesii, P. flexilis, Juniperus communis) absent from the site today. Diversity increases in the late‐glacial samples with the addition of J. scopulorum, J. horizontalis, C. montanus, C. ledifolius var. intricatus and mesic understory species. The coniferous trees and J. communis declined and J. osteosperma appeared during the late‐glacial/Holocene transition. Juniperus osteosperma populations have occupied the site throughout the Holocene. Pinus ponderosa was established by 7500 cal‐yr bp , and has occurred at least locally ever since. Montane conifers and J. horizontalis persisted until c. 5500 cal‐yr bp . The signature events of the late Holocene were the invasions of P. edulis and Ephedra viridis and establishment of pinyon–juniper woodland in the last 800 years. Main conclusions The Dutch John Mountain midden record adds to an emerging picture in which mid‐elevation conifers (P. flexilis, Pseudotsuga menziesii, Picea pungens, J. scopulorum, J. communis) dominated vegetation over a wide area of the Colorado Plateau and adjacent Rocky Mountains. Rather than being fragmented, as often assumed in phylogeographical studies, these species had broader and more‐connected distributions than they do in the region today. Paradoxically, subalpine conifers (Picea engelmannii, A. lasiocarpa) occurred at higher elevations to the south, possibly representing declining precipitation from south to north owing to southward displacement of the polar jet stream. The Dutch John Mountain record displays a series of extinction and invasion events. Most of the extinctions were local in scale; nearly all constituents of fossil midden assemblages occur within a few kilometres of Dutch John Mountain, and some occur at least locally on its slopes. The sole exception is J. horizontalis, which is regionally extinct. In contrast to extinctions, Holocene invasions were regional in scale; J. osteosperma, P. ponderosa, P. edulis and Ephedra viridis immigrated from glacial‐age source populations far to the south.     

Species status and population structure of mussels (Mollusca: Bivalvia: Mytilus spp.) in the Wadden Sea of Lower Saxony (Germany)

Three species of mussel (genus Mytilus) occur in Europe: M. edulis (Linnaeus 1758), M. galloprovincialis (Lamarck 1819) and M. trossulus (Gould, Boston Society of Natural History 3: 343?C348, 1850). Although these species are indigenous to the North Sea, the Mediterranean and the Baltic Sea, respectively, they form an extended patchy species complex along the coasts of Europe (??the Mytilus edulis complex??) and are able to hybridize where their distributions overlap. Recent studies examining the taxonomic status and genetic composition of Mytilus populations in the Netherlands and the British Isles have revealed introgressive hybridization processes within this species complex, with hints of an invasion of nonindigenous M. galloprovincialis into the North Sea. Furthermore, an extensive international mussel fishery industry in Europe (i.e., Great Britain, the Netherlands, Denmark, and Germany) is also in discussion for a possibly anthropogenically induced bioinvasion of nonindigenous Mytilus traits into the Wadden Sea area. Although it is assumed that the Wadden Sea of Germany comprises M. edulis only, this has never been confirmed in a molecular genetic study. To assess the situation for the Wadden Sea of Lower Saxony, we conducted the first molecular study of the Mytilus genus in the region. Taxonomic identification of 504 mussels from 13 intertidal mussel banks using the nDNA marker Me15/16 revealed a population composition of 99% M. edulis and 1% M. edulis X M. galloprovincialis hybrids. Hence, the Wadden Sea population is unaffected by range expansion of nonindigenous Mytilus traits. The genetic structure of the M. edulis populations was investigated using the phylogenetic and population genetics analyses of the mitochondrial DNA cytochrome-c-oxidase subunit I (COI) and the first variable domain of the control region (VD1), which were sequenced for >120 female individuals. These results showed a heterogeneous, panmictic population due to unrestricted gene flow. This can be attributed to extensive larval dispersal linked to the tidal circulation system in the back barrier basins of the Wadden Sea.     

Biogenic and taphonomic processes affecting the development of shell assemblages: an actuopaleontological case study from mussel banks on North Sea tidal flats

Summary From spring 1997 to winter 2000, the composition and structure of epibenthic bivalve clusters and patches on a backbarrier tidal flat of the East Frisian Wadden Sea were continoously analyzed. The studied bivalve aggregates developed from intense and approximately simultaneous spatfalls of Mytilus edulis and Cerastoderma edule in 1996, after a severe winter (1995/96) with 64 days of driftice coverage. Spatfall occurred on a bare sandy tidal flat, whereas adjacent shell layers situated at the southern margin of the spatfall site were not colonized. Usually, such shell layers are the most common secondary hard substrates in the backbarrier area favorable for initial settlement of Mytilus larvae. The shell layers predominantly originate from erosion of former Mya arenaria and C. edule populations. So far, the studied bivalve aggregates are composed of approximately equal proportions of living specimens of both species, M. edulis and C. edule. The incorporated cockles were prevented from burrowing into the rediment by the coherent byssal mesh. In contrast to the results of previous studies, the great bulk of epibenthic living C. edule did survive for several years within the clusters. Therefore, the common terms ‘Mytilus banks’, ‘Mytilus patches’ or ‘Mytilus clumps’ may be too general and may erroneously imply a dominance of (or mono-specific composition by) M. edulis. In the fossil record, i.e. in diagenetically consolidated shell layers, the enforced epibenthic life-mode of C. edule and the remarkable species composition as well as the temporal development of the aggregates would not be recognizable.     

Holocene persistence of wooded steppe in the Great Hungarian Plain

Aim We used a combination of new and previously published palaeoecological data to test three hypotheses: (1) that wooded steppe persisted in the Great Hungarian Plain throughout the Holocene; (2) that wooded steppe and steppe were most extensive between c. 9900 and 8300 cal. yr bp (the ‘Boreal steppe’ period); and (3) that Southern Continental, Pontic and Eastern Sub‐Mediterranean steppe species reached the region during the early Holocene via the ‘Lower Danube Corridor’. Location Sarló‐hát oxbow lake, Hungary and the Eastern European wooded steppe zone. Methods Holocene sediments deposited in the Sarló‐hát oxbow lake were subjected to pollen and microcharcoal analyses. Twelve radiocarbon age estimates were obtained to determine sediment chronology. In addition, previously published palaeoecological data from the Great Hungarian Plain were compiled, analysed and compared with previous studies in other regions of steppe and wooded steppe in eastern Europe. Results Palynological data from two sediment cores extending to c. 11,400 cal. yr bp indicate the persistent dominance of the landscape by temperate deciduous wooded steppe throughout the Holocene, although with varying canopy composition. Warm‐continental steppe grasslands and saline tall‐grass meadows developed on edaphically constrained areas, which remained steppe‐dominated throughout the Holocene. The extent of steppe grasslands did not increase between 9900 and 8300 cal. yr bp . After c. 3100 cal. yr bp , anthropogenic activities led to the development of cultural steppe. Thermophilous steppe species of the Southern Continental, Pontic and Sub‐Mediterranean floristic elements probably reached the Great Hungarian Plain principally via the Lower Danube Corridor during the late glacial interstadial and Holocene. Eurythermic members of these elements, however, probably survived the Last Glacial Maximum in favourable microsites, extending their ranges during the Holocene from these local sources. Main conclusions Our results confirm the Holocene persistence of wooded steppe in the Great Hungarian Plain, disprove the ‘Boreal steppe’ theory, and suggest an Early Holocene period of greater vegetation openness between 11,400 and 9900 cal. yr bp . Evidence for the post‐glacial immigration of south‐eastern steppe elements into the Carpathian Basin is equivocal: the last glacial/interglacial presence of several southern steppe species suggests that the Hungarian Plain hosted suitable habitats for them during warm and cold phases alike.     

Recombination in Mitochondrial DNA of European Mussels <Emphasis Type="Italic">Mytilus</Emphasis>

Mitochondrial DNA was long believed to be purely clonal and free from recombination. Major phylogenetic studies still depend on such assumptions. The peculiar genetic system of marine mussels Mytilus in which two divergent mitochondrial genomes exist provides a unique opportunity to study mtDNA recombination. Previous reports showed the existence of a few haplotypes having very strong recombination signal in the control region of mtDNA. Those recombinant variants have been found in a Baltic Sea population of Mytilus trossulus as well as in Mytilus galloprovincialis from the Black Sea. In both cases the mosaic genomes switched their transmission route and have been inherited paternally. In the present study rearranged mtDNA genomes found in all three European Mytilus species are described. The structure of their control region is a result of intra- and intermolecular recombination between mitochondrial genomes. Together with the phylogenetic reconstruction and geographic distribution, this suggests that two interlineage recombination events have occurred in the control region of mtDNA of European mussels Mytilus. Contrary to earlier observations, some of the mosaic genomes do not show any gender bias, which has important implications regarding the transmission and evolution of blue mussel mitochondrial genomes.     

Polar bear (<Emphasis Type="Italic">Ursus maritimus</Emphasis>) maternity den distribution in Svalbard,Norway

The aim of the current study was to identify maternity denning areas and to describe aspects of denning behaviour in polar bears (Ursus maritimus) in Svalbard, Norway. Den observations made during aerial and ground surveys (n = 553), observations of recently emerged family groups made during capture and tagging of bears (n = 206), and dens recorded by satellite telemetry (n = 76 dens) were used to describe the distribution of dens in the Barents Sea region. Data were collected during March, April, and May from 1972 to 2010, and while most dens were abandoned during the first 2 weeks of April, there was substantial variation. The highest number of dens was recorded on the islands in the eastern and northern parts of Svalbard with fewer dens found further west on the island Spitsbergen. Females with cubs-of-the-year or their tracks indicated some denning in southern, eastern, and northern parts of Spitsbergen. Telemetry also revealed that females captured in Svalbard and the Barents Sea den in Franz Josef Land, Russia. The majority of dens (62%) in Svalbard were located on land within ca. 1 km of the shore with the most inland den 19 km from shore. Heavy exploitation of polar bears before 1973, when a hunting moratorium was implemented, likely led to reduced denning in some areas. Our observations of den distribution indicate that denning is now more widespread in the archipelago than 50 years ago and reflects a reestablishment of denning areas following years of protection.     

Glacial history of the European marine mussels Mytilus,inferred from distribution of mitochondrial DNA lineages

Mussels of the genus Mytilus have been used to assess the circumglacial phylogeography of the intertidal zone. These mussels are representative components of the intertidal zone and have rapidly evolving mitochondrial DNA, suitable for high resolution phylogeographic analyses. In Europe, the three Mytilus species currently share mitochondrial haplotypes, owing to the cases of extensive genetic introgression. Genetic diversity of Mytilus edulis, Mytilus trossulus and Mytilus galloprovincialis was studied using a 900-bp long part of the most variable fragment of the control region from one of their two mitochondrial genomes. To this end, 985 specimens were sampled along the European coasts, at sites ranging from the Black Sea to the White Sea. The relevant DNA fragments were amplified, sequenced and analyzed. Contrary to the earlier findings, our coalescence and nested cladistics results show that only a single M. edulis glacial refugium existed in the Atlantic. Despite that, the species survived the glaciation retaining much of its diversity. Unsurprisingly, M. galloprovincialis survived in the Mediterranean Sea. In a relatively short time period, around the climatic optimum at 10 ky ago, the species underwent rapid expansion coupled with population differentiation. Following the expansion, further contemporary gene flow between populations was limited.     

Late‐glacial and Holocene vegetation,climate and fire dynamics in the Serra dos Órgãos,Rio de Janeiro State,southeastern Brazil

We present a high‐resolution pollen and charcoal record of a 218 cm long sediment core from the Serra dos Órgãos, a subrange of the coastal Serra do Mar, located at 2130 m altitude in campos de altitude (high elevation grass‐ and shrubland) vegetation near Rio de Janeiro in southeastern Brazil to reconstruct past vegetation, climate and fire dynamics. Based on seven AMS ¹⁴C ages, the record represents at least the last 10 450 ¹⁴C yr bp (12 380 cal years bp ), The uppermost region was naturally covered by campos de altitude throughout the recorded period. Diverse montane Atlantic rain forest (ARF) occurred close to the studied peat bog at the end of the Late‐glacial period. There is evidence of small Araucaria angustifolia populations in the study area as late as the early Holocene, after which point the species apparently became locally extinct. Between 10 380 and 10 170 ¹⁴C yr bp (12 310–11 810 cal yr bp ), the extent of campos de altitude was markedly reduced as montane ARF shifted rapidly upward to higher elevations, reflecting a very wet and warm period (temperatures similar to or warmer than present day) at the end of the Younger Dryas (YD) chronozone. This is in opposition to the broadly documented YD cooling in the northern Hemisphere. Reduced cross‐equatorial heat transport and movement of the Intertropical Convergence Zone over northeastern Brazil may explain the YD warming. Markedly extended campos de altitude vegetation indicates dry climatic conditions until about 4910 ¹⁴C yr bp (5640 cal yr bp ). Later, wetter conditions are indicated by reduced high elevation grassland and the extension of ARF into higher elevation. Fire frequency was high during the early Holocene but decreased markedly after about 7020 ¹⁴C yr bp (7850 cal yr bp ).     

Mytilus galloprovincialis Lmk. in Southern Africa

The Mediterranean mussel, Mytilus galloprovincialis Lmk. has been identified on the west coast of southern Africa using morphological and biochemical-genetic comparisons with samples of “pure” M. edulis L. from Denmark and M. galloprovincialis from the Mediterranean coast of Spain. Our results show that the southern African Mytilus more closely resembles M. galloprovincialis in anterior adductor, posterior adductor and hinge size. It also has a more ventrally located maximal shell width like the Mediterranean mussel. The electrophoretic examination of 23 proteins showed that many alleles diagnostic for M. galloprovincialis appeared in the southern African Mytilus. Nei's genetic distance between the southern African Mytilus and M. galloprovincialis was 0.010 ± 0.004 and the average genetic distance between these samples and M. edulis was 0.162 ± 0.077.The presence of M. galloprovincialis in southern Africa could be explained by biogeographic dispersal along the west coast of Africa during Pleistocene cooling, or by a recent inadvertant or intentional introduction by man. To test these two hypotheses, we examined ≈750 mussel shells collected from Koi-san middens and ≈350 shells from a raised-beach deposit all located on the west coast of Cape Province, South Africa. The ages of these deposits range from 1500 yr to 120000 yr and pre-date the arrival of Europeans to the Cape. This examination did not reveal any shells of Mytilus. We, therefore, conclude that the introduction of M. galloprovincialis to southern Africa was a recent event. We suggest that, since the first confirmed reference to Mytilus in southern Africa was made from a 1972 collection, the introduction occurred within the last two decades.     

Late Weichselian and Holocene sedimentary environments and ice rafting in Isfjorden, Spitsbergen

Multi-proxy analyses of two sediment cores from central Isfjorden were used to reconstruct the glacial history in central Spitsbergen during the Late Weichselian and the Holocene, and to relate iceberg rafting and sea-ice rafting to climatic and oceanographic changes in the north Atlantic region. A basal till was deposited beneath an ice stream prior to 12,700 cal. years BP (calendar years before the present). Several tidewater glaciers influenced the sedimentary environment during the Younger Dryas and probably already during the Allerød. The Younger Dryas cooling might be reflected by enhanced sea-ice formation and suspension settling, as well as reduced iceberg rafting. The final deglaciation was dominated by intense iceberg rafting. It terminated around 11,200 cal. years BP. Optimum Holocene climatic and oceanographic conditions with significantly reduced ice rafting occurred between c. 11,200 and 9000 cal. years BP. Ice rafting occurred almost exclusively from icebergs after 10,200 cal. years BP. The icebergs most probably originated from tidewater glaciers on east Spitsbergen, indicating the presence of a strong east–west temperature gradient at this time. An increase of iceberg rafting around 9000 cal. years BP, followed by enhanced sea-ice rafting, reflects the onset of a general cooling in the western Barents Sea–Svalbard region. Comparatively intense rafting from icebergs and sea ice between 9000 and 4000 cal. years BP is related to this cooling. A general reduction in ice rafting after 4000 cal. years BP is most probably the result of the enhanced formation of shore-fast and/or more permanent sea-ice cover, reducing the drift of icebergs and sea ice in the fjord. The results indicate that the palaeoenvironmental conditions on central Spitsbergen to a large degree depended on the oceanographic conditions in the western Barents Sea and west off Spitsbergen. However, climatic trends affecting Greenland and the entire north Atlantic region can also be identified.     

Development of a real-time PCR assay for detection of Mytilus species specific alleles: Application to a sampling survey in Scotland

Shellfish aquaculture is a growing industry in Scotland, dominated by the production of the mussel Mytilus edulis, the native species. Recently the discovery of Mytilus galloprovincialis and Mytilus trossulus together with M. edulis and all 3 hybrids in cultivation in some Scottish sea lochs led to questions regarding the distribution of mussel species in Scotland. The establishment of an extensive sampling survey, involving the collection of mussels at 34 intertidal sites and 10 marinas around Scotland, motivated the development of a high-throughput method for identification of Mytilus alleles from samples. Three Taqman®-MGB probes and one set of primers were designed, based on the previously described Me 15/16 primers targeting the adhesive protein gene sequence, and samples were screened for the presence of M. edulis, M. galloprovincialis and M. trossulus alleles using real-time PCR. Mytilus edulis alleles were identified in samples from all 44 sites. Mytilus galloprovincialis alleles were found together with M. edulis alleles extensively in northern parts of the west and east coasts. Mytilus trossulus alleles were identified in samples from 6 sites in the west and south-west of Scotland. Because M. trossulus is generally undesirable in cultivation and therefore preventing the geographical spread of this species across Scotland is considered beneficial by the shellfish aquaculture industry, these 6 samples were further analysed for genotype frequencies using conventional PCR. Although distribution of the non-native species M. galloprovincialis and M. trossulus have proven to be more widespread than previously thought, there is no evidence from our study of either M. trossulus or M. galloprovincialis acting as an invasive species in Scotland. The real-time PCR method developed in this study has proven to be a rapid and effective tool for the identification of M. edulis, M. galloprovincialis and M. trossulus alleles from samples and should prove useful in future surveys, ecological or aquaculture management related studies in both unispecific and mixed species areas of these species.     

Aspects of the population genetics of Mytilus (Mytilidae; Mollusca) in the British Isles

Starch gel electrophoresis was used to study variation at 11 loci in mussels sampled mainly from British coastal sites. Two types of mussel were identified, Mytilus edulis, the common mussel and its southern relative Mytilus galloprovincialis. Several partially diagnostic loci were used to map the distribution of the two forms. Mytilus edulis was present at all sites sampled in Britain and Ireland but was at low frequency in SW England; M. galloprovincialis was detected in SW England, the south and west of Ireland. Scotland and NE England, but was absent from south Wales, the Irish sea coasts of Wales and Ireland, and SE England. Apart from the occurrence of M. galloprovincialis in NE England, this distribution conforms with the results of studies using morphological characters and parallels the distribution of many other southern species in Britain. At the microgeographical level, M. edulis was found to prefer more sheltered and estuarine conditions than M. galloprovincialis. Analysis using the best diagnostic loci showed that hybridization is occurring between M. edulis and M. galloprovincialis at all localities where they occur sympatrically but that the extent of hybridization varies considerably between localities. The distribution of localities having high proportions of hybrid individuals is best interpreted by assuming that hybrids have higher fitness than parental types at these localities. A study was made of variation within and between those localities where only M. edulis individuals were observed. Little significant geographic variation in allele frequency was detected, but significant deficits of heterozygotes compared with Hardy-Weinberg expectations were seen for most loci. Analysis suggests that the Wahlund effect is not involved and that the most likely cause of the deficit is low frequencies of null alleles. In M. edulis no differences in phenotypic variance in shell height and width were observed between samples of multiply heterozygous and multiply homozygous individuals and no genetic differences were found between juveniles and adults. Overall little evidence was found that balancing selection is responsible for maintenance of the polymorphisms studied in M. edulis. The pattern of geographic variation in gene frequencies in Mytilus in the British Isles is discussed in relation to variation in the south and north of Europe and North America. It is concluded that steep clines in gene frequencies in M. edulis observed by other workers in the Baltic and in Long Island Sound cannot be attributed to the presence of M. galloprovincialis.     

Recently established <Emphasis Type="Italic">Crassostrea</Emphasis>-reefs versus native <Emphasis Type="Italic">Mytilus</Emphasis>-beds: differences in ecosystem engineering affects the macrofaunal communities (Wadden Sea of Lower Saxony,southern German Bight)

Since 1998 the non-indigenous Pacific oyster Crassostrea gigas (Thunberg 1793) has been invading the Wadden Sea of Lower Saxony, southern German Bight. C. gigas settles predominantly on intertidal Mytilus-beds (M. edulis L.) and subsequently create rigid reef-like structures. Both bivalve species are ecosystem engineers in sedimentary tidal flats. They provide hard substrate for sessile species, mobile organisms find refuge within the habitat matrix of dense suspension feeders, and biodeposits enrich the sediments with organic matter. The transformation of Mytilus-beds into Crassostrea-reefs gives rise to the question whether the invader may affect the native community. We investigated two parts of a changing bivalve bed in the backbarrier area of the island of Juist in March 2005. One part was still dominated by M. edulis whereas the other part was already densely colonized by C. gigas. Crassostrea-reefs compensate for the conceivable loss of Mytilus-beds in the intertidal of the Wadden Sea by replacing the ecological function of M. edulis. There was no indication of a suppression of indigenous species. This even applied to M. edulis, which persisted at the site invaded by C. gigas. The associated macrofaunal community showed increased species richness, abundance, biomass, and diversity in the Crassostrea-reef. The latter particularly favored sessile species like anthozoans, hydrozoans, and barnacles. Higher abundance and biomass for vagile epizoic species like the shore crab Carcinus maenas and the periwinkle Littorina littorea also occurred among oysters. Abundance of deposit feeding oligochaetes was enhanced by oysters as well. More opportunistic, facultative filter-feeding polychaetes occurred in the Crassostrea-reef.     

Mytilid mussels: global habitat engineers in coastal sediments

Dense beds of mussels of the family Mytilidae occur worldwide on soft-bottoms in cold and warm temperate coastal waters and have usually been considered hot spots of biodiversity. We examined intertidal mussel beds at four distant locations around the globe with the same sampling method, to find out whether this “hot spot” designation holds universally. We studied species assemblages within the matrices of byssally interconnected mussels engineered by Mytilus edulis in the North Sea, by mixed Perumytilus purpuratus and Mytilus chilensis at the southern Chilean coast, by Musculista senhousia in the Yellow Sea and by Xenostrobus inconstans at the coast of southern Australia. In all cases, species assemblages inside mussel beds were significantly different from those outside with many species being restricted to one habitat type. However, species richness and diversity were not generally higher in mussel beds than in ambient sediments without mussels. In the North Sea (M. edulis) and at the Chilean coast (P. purpuratus, M. chilensis), mussel beds have markedly higher species numbers and diversities than surrounding sediments, but this was not the case for mussel beds in Australia (X. inconstans) and the Yellow Sea (M. senhousia) where numbers of associated species were only slightly higher and somewhat lower than in adjacent sediments, respectively. In conclusion, although soft bottom mytilid mussels generally enhance habitat heterogeneity and species diversity at the ecosystem level, mussel beds themselves are not universal centres of biodiversity, but the effects on associated species are site specific.     

Combining hydrodynamic modelling with genetics: can passive larval drift shape the genetic structure of Baltic Mytilus populations?

While secondary contact between Mytilus edulis and Mytilus trossulus in North America results in mosaic hybrid zone formation, both species form a hybrid swarm in the Baltic. Despite pervasive gene flow, Baltic Mytilus species maintain substantial genetic and phenotypic differentiation. Exploring mechanisms underlying the contrasting genetic composition in Baltic Mytilus species will allow insights into processes such as speciation or adaptation to extremely low salinity. Previous studies in the Baltic indicated that only weak interspecific reproductive barriers exist and discussed the putative role of adaptation to environmental conditions. Using a combination of hydrodynamic modelling and multilocus genotyping, we investigate how oceanographic conditions influence passive larval dispersal and hybrid swarm formation in the Baltic. By combining our analyses with previous knowledge, we show a genetic transition of Baltic Mytilus species along longitude 12°‐13°E, that is a virtual line between Malmö (Sweden) and Stralsund (Germany). Although larval transport only occurs over short distances (10–30 km), limited larval dispersal could not explain the position of this genetic transition zone. Instead, the genetic transition zone is located at the area of maximum salinity change (15–10 psu). Thus, we argue that selection results in weak reproductive barriers and local adaptation. This scenario could maintain genetic and phenotypic differences between Baltic Mytilus species despite pervasive introgressive hybridization.