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.     

A population genetic study of the hybrid zone of Mytilus trossulus Gould, 1850 and an introduced species,M. galloprovincialis Lamarck, 1819, (Bivalvia: Mytilidae) in peter the great bay in the Sea of Japan

This paper examines the genetic variability of the Pacific mussel Mytilus trossulus and an introduced Atlantic species, M. galloprovincialis, in the northwestern Sea of Japan (Peter the Great Bay and Kievka Bay). The genotyping of individuals from eight populations was carried out using eight polymorphic enzyme loci and two nuclear DNA markers (Me-5 and ITS-1,2); the occurrence frequency of parent species and their hybrids was determined. The enzyme and nuclear markers demonstrated concordant genetic variation. The genotypes of the native species M. trossulus were predominant in the samples studied. The frequency of the introduced species M. galloprovincialis in the total material was relatively low; however, it reached 42 ± 2% in samples that were collected in Possjet Bay near the town of Zarubino in a zone of active international navigation. In this area the greatest number of hybrids was found as well. It is concluded that the invasion of M. galloprovincialis in the northwestern Sea of Japan is continuing; permanent populations of this mussel appeared in Possjet Bay that were not recorded here previously.     

Three species of Mytilus and their hybrids identified in a Scottish Loch: natives, relicts and invaders?

Three species of the mussel, Mytilus, occur in the North Atlantic region, M. edulis, M. galloprovincialis and M. trossulus, and hybrid zones are present where their distributions overlap. M. edulis is a native species in the UK. M. galloprovincialis originated in the Mediterranean and its distribution extends northwards along the Atlantic seaboard to Scotland. Baltic Sea mussels have a M. trossulus ancestry but are highly introgressed by M. edulis. In recent decades, farming of mussels on long-line rope culture systems has been introduced into Scotland. On farms in Loch Etive, a form of mussel with a fragile shell and a different shape to either M. edulis or M. galloprovincialis has been increasing in frequency over recent years. Samples of fragile shelled, normal strong shelled and intermediate mussel types were sampled from two farms in 2006 and compared with samples of M. edulis, M. galloprovincialis and M. trossulus from other sources where their species identity is well established. Abundance relative to depth, shell strength, condition index and shell morphology were analysed together with 5 allozyme loci and one nuclear DNA genetic marker (Me 15/16). The fragile shelled mussels, and many of those classed as intermediate, were identified as a mixture of M. trossulus and M. trossulus x M. edulis hybrids. This identification was strongly supported by both morphological and genetic data and is the first record of the presence of M. trossulus in UK waters. M. trossulus in Loch Etive are most likely to be a post-glacial relict population restricted to the low salinity area of the Loch that has recently increased in abundance due to commercial mussel growing activity. In addition, individual mussels of all three species and their hybrids were detected amongst Loch Etive mussels. This is the first genetic demonstration of all three species and their hybrids occurring together in one location in the Atlantic region and provides a unique opportunity to study the processes of speciation, divergence, and introgression in the genus Mytilus.     

The thermal environment of nests of the Australian stingless bee, Austroplebeia australis

The greatest diversity of stingless bee species is found in warm tropical regions, where brood thermoregulation is unnecessary for survival. Although Austroplebeia australis (Friese) naturally occurs in northern regions of Australia, some populations experience extreme temperature ranges, including sub-zero temperatures. In this study, the temperature was monitored in A. australis colonies’ brood chamber (n = 6) and the hive cavity (n = 3), over a 12-month period. The A. australis colonies demonstrated some degree of thermoconformity, i.e. brood temperature although higher correlated with cavity temperature, and were able to warm the brood chamber throughout the year. Brood production continued throughout the cold season and developing offspring survived and emerged, even after exposure to very low (?0.4 °C) and high (37.6 °C) temperatures. Austroplebeia australis, thus, demonstrated a remarkable ability to survive temperature extremes, which has not been seen in other stingless bee species.     

Study of Two Species of Mussels, Mytilus trossulusand Mytilus galloprovincialis(Bivalvia, Mytilidae), and Their Hybrids in Peter the Great Bay of the Sea of Japan with the Use of PCR Markers

Bivalve mollusks of the genus Mytilus(M. trossulusand M. galloprovincialis) occurring in Peter the Great Bay of the Sea of Japan were first studied in Russia. A region of nonrepetitive sequences of the gene encoding the polyphenolic adhesive protein bissus was used as a species-specific genetic marker. After amplification using specific primers, a 126-bp fragment was found to amplify in all representatives ofM. galloprovincialiscollected from driftwood in the gulf Posset (the southwestern part of Peter the Great Bay). M. trossulusspecimens from the same region were shown to have a 168-bp fragment. In Vostok Gulf (the eastern part of Peter the Great Bay), both artificially grown mussels and those from natural habitats contained a 168-bp fragment or two fragments (126- and 168-bp) that corresponded to a hybrid form between the above species. The possibility of using this genetic marker to identify closely related Mytilusstrains and their hybrids in similar habitats, near the Primorye coast in particular, was demonstrated. The presence of approximately 9% of hybrid specimens confirms that a zone of hybridization between M. trossulusand M. galloprovincialismay exist in this region.     

Historical changes in the distributions of invasive and endemic marine invertebrates are contrary to global warming predictions: the effects of decadal climate oscillations

Aim We tested whether a hybrid zone that has formed between an endemic and an invasive species of marine mussel has shifted poleward as expected under a general hypothesis of global warming or has responded instead to decadal climate oscillations. Location We sampled 15 locations on the coast of California, USA, that span the distributions of the two species of marine mussels and their hybrids. Methods Mussels were sampled in 2005–08 and analysed at three nuclear gene loci using methods identical to those used in a study a decade earlier in order to document the genetic architecture of this system. Change in the system was determined by comparing the frequency of species‐specific alleles and multi‐locus genotypes over the intervening decade. Climate variation over the same period was examined by comparing the Pacific Decadal Oscillation (PDO), El Niño/Southern Oscillation (ENSO), upwelling indices and sea surface temperature (SST) during and prior to the study period. Results Contrary to the general expectations of global warming we show that the highly invasive warm‐water mussel Mytilus galloprovincialis and the hybrid zone formed with the endemic species Mytilus trossulus has rapidly contracted southwards. Mytilus galloprovincialis declined in abundance over the northern third of its geographic range (c. 540 km) and has become rare or absent across the northern 200 km of the range it previously colonized during its initial invasion. The distribution of the native species M. trossulus has remained unchanged over the same time period. Main conclusions The large‐scale range shift in the warm‐water invasive species M. galloprovincialis and the hybrid zone it forms with M. trossulus has been exceptionally rapid and is in the opposite direction to that predicted by the global warming hypotheses. This shift, however, is consistent with decadal climate variation associated with the ENSO and the PDO. Since the biogeography of this system was first described in 1999, the PDO has shifted from a warm phase, dominated by frequent and large El Niño events, to a cold‐phase period, with minimal ENSO activity. Thus recent decadal climate variation can oppose global trends in average temperature and this study illustrates the need to integrate the effects of climate change across multiple time‐scales.     

A historical perspective of the genus Mytilus (Bivalvia: Mollusca) in New Zealand: multivariate morphometric analyses of fossil, midden and contemporary blue mussels

The taxonomic status of smooth shelled blue mussels of the genus Mytilus has received considerable attention in the last 25 years. Despite this, the situation in the southern hemisphere remains uncertain and is in need of clarification. Recent work suggests that contemporary New Zealand mussels from two cool/cold temperate locations are M. galloprovincialis. However, the distribution of Mytilus in New Zealand ranges from 35 ° to 52 ° south (～ 1800 km), meaning that large areas of the subtropical/warm temperate north and the subantarctic south remain unsampled, an important consideration when species of this genus exhibit pronounced macrogeographical differences in their distributions which are associated with environmental variables such as water temperature, salinity, wave action and ice cover. This study employed multivariate morphometric analyses of one fossil, 83 valves from middens, and 92 contemporary valves from sites spanning the distributional range of blue mussels to determine a historical and contemporary perspective of the taxonomic status of Mytilus in New Zealand. The findings indicated that all fossil and midden mussels are best regarded as M. galloprovincialis and confirmed that contemporary mussels, with one possible regional exception, are also best regarded as M. galloprovincialis. Contemporary mussels from the Bay of Islands (warm temperate/subtropical) exhibited much greater affinity to M. edulis than they did to M. galloprovincialis, indicating that mussels from this area require detailed genetic examination to determine their taxonomic status. The analyses revealed a significant difference between the fossil/midden mussels and the contemporary mussels, consistent with levels of present day differentiation among intraspecific populations and not thought to reflect any substantive temporal change between mussels of the two groups. The continuous distribution of M. galloprovincialis in New Zealand from the warm north to the subantarctic south indicates that the physiology of this species is adapted to a wide range of water temperature conditions. Therefore, the distribution of this species on a worldwide scale is unlikely to be restricted by its adaptation to warm water alone, as has previously been widely assumed. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 329–344.     

Larval settlement can explain the adult distribution of Mytilus californianus Conrad but not of M. galloprovincialis Lamarck or M. trossulus Gould in Moss Landing, central California: Evidence from genetic identification of spat

We investigated the spatial distribution of adult and newly settled mussels (Mytilus galloprovincialis Lamarck, Mytilus trossulus Gould and Mytilus californianus Conrad) on the shore at Moss Landing, California to test the hypothesis that adult distributions are a result of settlement patterns. Adult M. californianus were most abundant on a wave-exposed rocky jetty and adults of Blue mussels (M. trossulus and M. galloprovincialis) were more abundant inside the protected Moss Landing harbor. Using taxon-specific polymerase chain reactions, we monitored recruitment during continuous 1-2 week intervals on fibrous scrubbing pads for 12 months in 2002-2003. All mussel species settled in greatest numbers on the exposed jetty, and Blue mussels settled in greater numbers there than did M. californianus. Because Blue mussels settled abundantly where their adults were rare, post-settlement mortality appeared to be the strongest influence on adult distribution. In contrast, M. californianus settled mostly in their adult habitat.     

SEASONAL DIFFERENCES IN THE TEMPERATURE RANGES OF GROWTH OF VIRGINIA ALGAE1

One hundred and fifteen clonal, unialgal strains were isolated and tested for their ability to grow over a range of temperatures from 2 to 40° C. Responses of 63 strains isolated from habitats that were 6° C when sampled and 52 strains isolated from habitats that were 20° C when sampled showed trends toward increasing adaptation to cold or warm temperatures commensurate with their seasonal in situ temperatures. Based on temperature-growth responses alone, 24% of the plankton isolates and 17% of the periphyton isolates could be perennial within the natural habitats. At 5° C, 56% of the warm water plankton isolates and 48% of the warm water periphyton isolates were incapable of growth and, therefore, probably could not be important components of the winter algal community. Likewise at 25° C, 25% of the cold water plankton isolates and 13% of the cold water periphyton isolates were incapable of growth. Thus, temperature alone probably is an important variable regulating seasonal changes in algal community structure. Pollution of these habitats by a thermal enrichment averaging + 5° C year-round could effect a pronounced change in algal species composition because many more taxa could be perennial and more taxa would be incapable of growth during naturally warm periods.     

Physiological performance of the cold-water coral Dendrophyllia cornigera reveals its preference for temperate environments

Cold-water corals (CWCs) are key ecosystem engineers in deep-sea benthic communities around the world. Their distribution patterns are related to several abiotic and biotic factors, of which seawater temperature is arguably one of the most important due to its role in coral physiological processes. The CWC Dendrophyllia cornigera has the particular ability to thrive in several locations in which temperatures range from 11 to 17 °C, but to be apparently absent from most CWC reefs at temperatures constantly below 11 °C. This study thus aimed to assess the thermal tolerance of this CWC species, collected in the Mediterranean Sea at 12 °C, and grown at the three relevant temperatures of 8, 12, and 16 °C. This species displayed thermal tolerance to the large range of seawater temperatures investigated, but growth, calcification, respiration, and total organic carbon (TOC) fluxes severely decreased at 8 °C compared to the in situ temperature of 12 °C. Conversely, no significant differences in calcification, respiration, and TOC fluxes were observed between corals maintained at 12 and 16 °C, suggesting that the fitness of this CWC is higher in temperate rather than cold environments. The capacity to maintain physiological functions between 12 and 16 °C allows D. cornigera to be the most abundant CWC species in deep-sea ecosystems where temperatures are too warm for other CWC species (e.g., Canary Islands). This study also shows that not all CWC species occurring in the Mediterranean Sea (at deep-water temperatures of 12–14 °C) are currently living at their upper thermal tolerance limit.     

Variation in phenotypic plasticity for native and invasive populations of Bromus tectorum

Phenotypic plasticity is often considered important for invasive plant success, yet relatively few studies have assessed plasticity in both native and invasive populations of the same species. We examined the plastic response to temperature for Bromus tectorum populations collected from similar shrub-steppe environments in the Republics of Armenia and Georgia, where it is native, and along an invasive front in California and Nevada. Plants were grown in growth chambers in either ‘warm’ (30/20 °C, day/night) or ‘cold’ (10/5 °C) conditions. Invasive populations exhibited greater adaptive plasticity than natives for freezing tolerance (as measured by chlorophyll a fluorescence), such that invasive populations grown in the cold treatment exhibited the highest tolerance. Invasive populations also exhibited more rapid seedling emergence in response to warm temperatures compared to native populations. The climatic conditions of population source locations were related to emergence timing for invasive populations and to freezing tolerance across all populations combined. Plasticity in growth-related traits such as biomass, allocation, leaf length, and photosynthesis did not differ between native and invasive populations. Rather, some growth-related traits were very plastic across all populations, which may help to dampen differences in biomass in contrasting environments. Thus, invasive populations were found to be particularly plastic for some important traits such as seedling emergence and freezing tolerance, but plasticity at the species level may also be an important factor in the invasive ability of B. tectorum.     

The temperature dimension of the seed germination niche in fen wetlands

The seed germination niche partly determines adaptation, ecological breadth and geographic range in plant species. In temperate wetlands, environmental temperature is the chief regulator of germination timing, but the ecological significance of high and low temperatures during dormancy break and germination is still poorly understood. Our aim was to characterize the temperature dimension of the germination niche in mountain base-rich fens, determining (1) the effect of different temperatures on dormancy break and germination, and (2) whether different germination strategies may be identified at the species level. We conducted laboratory germination experiments with seeds of 15 species from these habitats, collected in 18 fen sites in the Cantabrian Mountains (Spain) for two consecutive years. In all the species, the seeds were totally or conditionally dormant at dispersal and stratification produced a significant increase of germination. In most cases, there was not an obligatory requirement for cold temperatures during dormancy break, since warm stratification promoted germination as well. Although the optimal germination thermoperiod was generally high (30/20 °C), most species could also germinate at lower temperatures after cold-stratification. We also identified a group of species associated to cold-water springs that germinated only at low temperatures. Our results demonstrate that dormancy break in mountain base-rich fens does not obligatorily depend on cold temperatures during overwintering. Furthermore, germination at cool temperatures may be more widespread in wetland habitats than previously thought. The existence of two distinctive germination strategies, ‘warm’ and ‘cool’, can potentially give rise to divergent species responses to climate change.     

The effect of temperature on the development times of eggs,naupliar and copepodite stages of five species of cyclopoid copepods

The duration times of eggs, combined naupliar instars and of the different copepodite stages of five species of cyclopoid copepods — Acanthocyclops robustus, Cyclops vicinus, Diacyclops bicuspidatus, Mesocyclops leuckarti, and Thermocyclops crassus — were investigated at five different temperatures. The five species can be divided in two groups: two species, C. vicinus and D. bicuspidatus, adapted to cold water conditions and three species, A. robustus, M. leuckarti and T. crassus adapted to warm water conditions. The cold water species showed a faster egg development than M. leuckarti and T. crassus at 5–15 °C. The eggs and instars of the warm water species M. leuckarti tend to develop faster than those of the former two species at higher temperatures. A. robustus showed the shortest egg and instar development at 10–25 °C. The warm water species T. crassus produced no eggs at 10 °C and temperatures below. At higher temperatures (20, 25 °C) the egg and instar duration times were similar or longer than those of the other species. When cultured in total darkness a great part of the C_IV respectively C_V copepodites of the summer forms entered arrest and the percentage of copepodites that showed an arrest of development was highest at lowest temperatures. The present results are compared with data from literature and differences are discussed.     

Predictors of invasion success by Daphnia species: influence of food, temperature and species identity

Non-indigenous North American Daphnia ‘pulex’ has recently invaded lakes and reservoirs in South Island, New Zealand, that formerly contained only native Daphnia carinata. New Zealand is characterised by a wide range of freshwater ecosystems and low species diversity of planktonic crustaceans, particularly Cladocera. The potential success of a species to invade and establish in a new community is likely to be predicted more accurately when reproductive response norms and fitness of key resident species, as well as the non-indigenous species, have been established under a range of relevant environmental conditions. Based on the results of experiments to test aspects of reproduction and fitness of the invader, D. ‘pulex’, and D. carinata when grown together at a range of relevant temperatures (8–23 °C) and related photoperiods, I predict the species-specific potential of D. ‘pulex’ to be dispersed and colonise New Zealand lentic habitats, and the potential of D. carinata to persist with the invader in these habitats. Larger population densities of D. ‘pulex’ compared to D. carinata at higher temperatures and food level, and larger densities of D. carinata at low temperatures, imply a potential for both species to coexist in New Zealand lakes, facilitated by seasonal succession; increased water temperature and nutrient input associated with climate and land use changes appear likely to promote the wider establishment of D. ‘pulex’, with both negative and positive implications for the conservation and management of New Zealand’s freshwater ecosystems.     

The effect of overwintering temperature on the body energy reserves and phenoloxidase activity of bumblebee Bombus lucorum queens

Warming winters and changes in species composition related to the estimated global warming may cause a threat to bumblebees adapted to cold winters. During the overwintering period, their intermediary and respiratory metabolism decreases but metabolism remains responsive to temperature. The effect of temperature on diapause survival, phenoloxidase (PO) activity, and energy expenditure of the white-tailed bumblebee (Bombus lucorum) after a 4-month diapause were studied by manipulating the diapause temperature. Two overwintering temperatures were used, cold (1.8 °C) and warm (9 °C). Body fat content was used as an estimate of the remaining energy resources and PO activity as an immune function parameter of overwintering bumblebee queens. The baseline levels of PO activity were used to measure the differences in B. lucorum queen responses after overwintering in either temperature. We found a 0.4 g pre-diapause threshold weight of survival in B. lucorum. Large queens had more fat left and a higher PO activity compared to small ones after overwintering in warm conditions, but in the cold there was no effect of size on the remaining fat in the fat body of queens or their PO activity. The observed difference in energy usage appears to relate to normal size-dependent metabolism and variation in energy allocation between basic metabolism and immune functions.     

Chemosensory and thermal cue responses in the sub-Antarctic moth Pringleophaga marioni: Do caterpillars choose Wandering Albatross nest proxies?

On the South Indian Ocean Province Islands of the sub-Antarctic, most nutrients are processed through a detritus-based food web. On Marion Island, larvae of the moth Pringleophaga marioni are one of the key decomposers. Abundance of these caterpillars is higher in newly abandoned Wandering Albatross (Diomedea exulans) nests than other habitats, and this observation has been explained by hypotheses regarding the thermal and nutrient advantages of nests. These hypotheses require a mechanism for increasing the abundance of caterpillars, since nests are an ephemeral resource, and here, we determine whether caterpillars respond to chemosensory and thermal cues using a laboratory choice chamber approach. Caterpillars show no significant preference for newly abandoned nest material over no other choice, old nest material, and the common mire moss Sanionia uncinata. Caterpillars that are acclimated to warm (15 °C) conditions do prefer lower (5 °C) to higher (15 °C) temperatures, perhaps reflecting negative effects of prolonged exposure to warm temperatures on growth. Caterpillars also show significant avoidance of conspecifics, possibly because of incidental cannibalism previously reported in this species. Thus, we find no empirical support for nest-finding ability in caterpillars based on chemosensory or thermal cues. It is possible that adult females or very early instar caterpillars show such ability, or high caterpillar density and biomass in nests are an incidental consequence of better conditions in the nests or deposition by the birds during nest construction.     

A common strategy of cold hardiness in ants of the genus <Emphasis Type="Italic">Myrmica</Emphasis> (Formicidae,Hymenoptera) in Northeast Asia

The biotopic distribution, nest structure, wintering conditions, and cold hardiness of four ecologically contrasting ant species (Myrmica angulinodis, M. kamtschatica, M. bicolor, and M. transsibirica) are considered. The cold hardiness of these species is typical of the genus: the supercooling points vary from −28 to −31°C; cold hardiness levels (LT50%) are higher by 5°C. At this level of cold hardiness, ants can be practically ubiquitous across the whole Hypoarctic (Berman et al., 2007). However, the above Myrmica species are strictly segregated (M. kamtschatica occurs in moss bogs, M. angulinodis and M. transsibirica, on dry and warm south slopes, and M. bicolor, in sandy-gravel floodplains), probably due to different requirements for weather conditions in summer and depth of ground thawing. At present, the excess cold hardiness common to the species in question (exceeding the nest temperature by 5–10°C in different years) is not adaptive and may be considered as preadaptive. It could have been acquired during ancient cold epochs or inherited by the genus as a concomitant result of adaptation not to low temperatures but, for instance, to aridity. These Myrmica species do not undergo selection for resistance to negative temperatures since their current level of cold hardiness is excessive, considering the possible wintering temperatures.     

A first report on coexistence and hybridization of <Emphasis Type="Italic">Mytilus trossulus</Emphasis> and <Emphasis Type="Italic">M. edulis</Emphasis> mussels in Greenland

Semi-sessile Mytilus mussels are used as indicators of climate changes, but their geographic distribution is not sufficiently known in the Arctic. The aim of this study was to investigate the taxonomic status and genetic differentiation of Mytilus populations in a Northwest Greenlandic fjord at Maarmorilik, impacted by contaminations from a former mine. In this study, mussels were collected at three sites differing in exposure to environmental factors. A total of 54 polymorphic SNPs found in the Mytilus EST and DNA sequences analyzed were successfully applied to 256 individuals. The results provided the first evidence for the existence of M. trossulus in Greenland. The mussel from M. trossulus and M. edulis taxa are shown to coexist and hybridize in the fjord. The three studied sites were found to differ significantly in the distribution of taxa with a higher prevalence of M. trossulus in the inner fjord. The identified M. edulis × M. trossulus hybrids mostly had a hybrid index score of about 0.5, indicating a similar number of alleles characteristic for M. trossulus and M. edulis. There was a low number of backcrosses between ‘pure’ taxa and hybrids. This newly discovered hybrid zone between the two taxa is unique in comparison with the Canadian populations. As Mytilus mussels in Greenland hitherto have been regarded as the one taxon M. edulis, the results have importance for biogeography and future monitoring and environmental studies.     

Introgression between invasive and native blue mussels (genus Mytilus) in the central California hybrid zone

The ecological and genetic factors determining the extent of introgression between species in secondary contact zones remain poorly understood. Here, we investigate the relative importance of isolating barriers and the demographic expansion of invasive Mytilus galloprovincialis on the magnitude and the direction of introgression with the native Mytilus trossulus in a hybrid zone in central California. We use double‐digest restriction‐site‐associated DNA sequencing (ddRADseq) to genotype 1337 randomly selected single nucleotide polymorphisms and accurately distinguish early and advanced generation hybrids for the first time in the central California Mytilus spp. hybrid zone. Weak levels of introgression were observed in both directions but were slightly more prevalent from the native M. trossulus into the invasive M. galloprovincialis. Few early and advanced backcrossed individuals were observed across the hybrid zone confirming the presence of strong barriers to interbreeding. Heterogeneous patterns of admixture across the zone of contact were consistent with the colonization history of M. galloprovincialis with more extensive introgression in northern localities furthest away from the putative site of introduction in southern California. These observations reinforce the importance of dynamic spatial and demographic expansions in determining patterns of introgression between close congeners, even in those with high dispersal potential and well‐developed reproductive barriers. Our results suggest that the threat posed by invasive M. galloprovincialis is more ecological than genetic as it has displaced, and continues to displace the native M. trossulus from much of central and southern California.