Return of the native facilitated by the invasive? Population composition,substrate preferences and epibenthic species richness of a recently discovered shellfish reef with native European flat oysters (Ostrea edulis) in the North Sea

After being ecologically extinct for almost a century, the discovery of a shellfish reef with native European flat oysters (Ostrea edulis) in the Dutch coastal area of the North Sea by the authors of this study called for an extensive survey to better understand some of the key requirements for the return of the native oyster in coastal waters. We assessed habitat conditions, its potential for increasing biodiversity, and the role of substrate provision by other bivalves such as the invasive alien Pacific oyster (Crassostrea gigas). Using underwater visual census, O. edulis size-frequency distributions and attachment substrate was investigated, as well as the composition of the epibenthic community and substrata types inside quadrats that were distributed across the reef. This reef was found to be composed of native European flat oysters, invasive alien Pacific oysters and blue mussels (Mytilus edulis), alternated with sandy patches. The O. edulis population (6.8?±?0.6 oysters m^?2) consisted of individuals of different size classes. In quadrats with native and non-native oysters the number of epibenthic species was 60% higher compared to adjacent sand patches within the reef. Notably, our results showed that the native oyster predominantly used shell (fragments) of the invasive Pacific oyster as settlement substrate (81% of individuals). Our results optimistically show that conditions for native oyster restoration can be suitable at a local scale in the coastal North Sea area and suggest that the return of native oysters may be facilitated by novel substrate provided by invasive oysters at sites where their distribution overlap.     

The European oyster <Emphasis Type="Italic">(Ostrea edulis)</Emphasis> and its epibiotic succession

This study aimed to investigate intra- and inter-site differences in the epibiota on the European oyster Ostrea edulis of different ages and on hard substrata to assess their contribution to biodiversity. This research has shown that shells of O. edulis do show higher species diversity than non-living hard substrata and as oysters grow larger epibiotic diversity will increase. This investigation has revealed that O. edulis possesses biogenic engineering qualities and the value of O. edulis as a vehicle for increasing biodiversity should not be underestimated. Relatively few studies have examined the shell surfaces of Ostrea edulis, which, together with their associated epifauna provide a large portion of the natural hard substrata for sessile filter-feeding organisms, including oysters, in coastal and estuarine ecosystems. It can be considered a keystone species which has an influence on intertidal and subtidal community structures.     

Strategies for the retention of high genetic variability in European flat oyster (<Emphasis Type="Italic">Ostrea edulis</Emphasis>) restoration programmes

The native European flat oyster Ostrea edulis is listed in the OSPAR Convention for the Protection of the Marine Environment of the North-East Atlantic (species and habitat protection) and in the UK Biodiversity Action Plan. Once extremely abundant in the nineteenth century, European stocks of O. edulis have declined during the twentieth century to rare, small, localised populations due to overexploitation, habitat degradation and, most recently, the parasitic disease bonamiosis. Selective breeding programmes for resistance to bonamiosis have been initiated in France and Ireland. High genetic diversity and bonamiosis-resistance would be important features of any sustainable restoration programmes for O. edulis. Oysters were sampled across Europe from four hatchery sources, four pond-cultured sources and four wild, but managed fisheries and were genotyped at five microsatellite loci. Hatchery-produced populations from small numbers of broodstock showed a significant loss of genetic diversity relative to wild populations and pedigree reconstruction revealed that they were each composed of a single large full-sib family and several small full-sib families. This extremely low effective population size highlights the variance in reproductive success among the potential breeders. Pond-cultured oysters were intermediate in genetic diversity and effective population size between hatchery and wild populations. Controlled hatchery production allows the development of bonamiosis-resistant strains, but at the expense of genetic diversity. Large scale pond culture on the other hand can provide a good level of genetic diversity. A mixture of these two approaches is required to ensure a healthy and sustainable restoration programme for O. edulis in Europe.     

Ecosystem engineering by a canopy‐forming kelp facilitates the recruitment of native oysters

Ecosystem engineers are species that influence the abiotic and biotic environment around them and may assist the restoration of associated species, including other habitat‐forming species. We deployed an array of 28 artificial reefs with transplanted Ecklonia radiata, the dominant canopy‐forming kelp species across southern Australia, to investigate how the patch size and density of E. radiata influenced the establishment of the associated communities of plants and animals. Many of the reefs were rapidly colonized by Ostrea angasi, a critically depleted reef‐forming oyster. Over the 24‐month deployment of the reefs, thick oyster mats formed across the entire surface of many of the reefs with estimated biomass densities exceeding 5 kg of live oysters/m²; however, oyster density was dependent on E. radiata patch size and density. Increasing patch size and the presence of kelp resulted in significantly higher densities of oysters 5 months after the reefs were deployed and at the end of the experiment, where oysters were approximately three times more numerous on reefs with kelp compared to those without kelp. E. radiata appeared to facilitate the establishment of O. angasi largely through its capacity to reduce benthic light and thus suppress competition from turfing algae. These results may inform the development of novel approaches to tackle recruitment bottlenecks affecting the restoration of O. angasi reefs.     

Evaluation of Long‐Term Response of Intertidal Creek Nekton to Phragmites australis (Common Reed) Removal in Oligohaline Delaware Bay Salt Marshes

In the oligohaline Alloway Creek watershed of the upper Delaware Bay, invasive Phragmites australis (Common reed; hereafter Phragmites) has been removed in an attempt to restore tidal marshes to pre‐invasion form and function. In order to determine the effects of Phragmites on nekton use of intertidal creeks and to evaluate the success of this restoration, intertidal creek nekton assemblages were sampled with weirs from May to November for 7 years (1999‐2005) in three marsh types: natural Spartina alterniflora (Smooth cordgrass; hereafter Spartina), sites treated for Phragmites removal (hereafter referred to as Treated), and invasive Phragmites marshes. Replicate intertidal creek collections in all three marsh types consisted primarily of resident nekton and were dominated by a relatively low number of ubiquitous intertidal species. The Treated marsh nekton assemblage was distinguished by greater abundances of most nekton, especially Fundulus heteroclitus (Mummichog). Phragmites had little impact on nekton use of intertidal creeks over this period as evidenced by similar nekton assemblages in the Spartina and Phragmites marshes for most years. Long‐term assemblage‐level analyses and nekton abundances indicated that the Treated marsh provided enhanced conditions for intertidal creek nekton. The response of intertidal creek nekton suggests that the stage of the restoration may influence the results of comparisons between the marsh types and should be considered when evaluating marsh restorations.     

Settlement and Survival of the Oyster Crassostrea virginica on Created Oyster Reef Habitats in Chesapeake Bay

Efforts to restore the Eastern oyster (Crassostrea virginica) reef habitats in Chesapeake Bay typically begin with the placement of hard substrata to form three‐dimensional mounds on the seabed to serve as a base for oyster recruitment and growth. A shortage of oyster shell for creating large‐scale reefs has led to widespread use of other materials such as Surf clamshell (Spisula solidissima), as a substitute for oyster shell. Oyster recruitment, survival, and growth were monitored on intertidal reefs constructed from oyster and Surf clamshell near Fisherman’s Island, Virginia, U.S.A. and on a subtidal Surf clamshell reef in York River, Virginia, U.S.A. At the intertidal reefs, oyster larvae settlement occurred at similar levels on both substrate types throughout the monitoring period but higher levels of post‐settlement mortality occurred on clamshell reefs. The oyster shell reef supported greater oyster growth and survival and offered the highest degree of structural complexity. On the subtidal clamshell reef, the quality of the substrate varied with reef elevation. Large shell fragments and intact valves were scattered around the reef base, whereas small, tightly packed shell fragments paved the crest and flank of the reef mound. Oysters were more abundant and larger at the base of this reef and less abundant and smaller on the reef crest. The availability of interstitial space and appropriate settlement surfaces is hypothesized to account for the observed differences in oyster abundance across the reef systems. Patterns observed emphasize the importance of appropriate substrate selection for restoration activities to enhance natural recovery where an underlying habitat structure is destroyed.     

The importance of predation and competition in organizing the intertidal epifaunal communities of Barnegat Inlet,New Jersey

Summary Community organization was studied by experiment and observation from October 1972–October 1974 in the marine epifaunal assemblages at each end of Barnegat Inlet, New Jersey. The rock jetty at the wave-exposed eastern end of the inlet possesses an intertidal community with the following attributes: (1) a high intertidal zone dominated by the barnacle, Balanus balanoides, but also occupied by the blue mussel, Mytilus edulis, in rock crevices, (2) a mid and low intertidal zone with usually <10% free space and extreme numerical dominance by Mytilus edulis (usually >85% cover) during summer and fall, and (3) almost no intertidal predators or herbivores. The predatory seastar, Asterias forbesi, is abundant subtidally. Controlled removal experiments indicate that in the mid and low intertidal underlying barnacles perish as a consequence of the establishmentof extensive secondary cover by Mytilus, probably because Mytilus outcompetes Balanus through suffocation or starvation. Mytilus transplants demonstrate that the mussels do not survive outside of crevices in the high intertidal, which thus may represent for Balanus a refuge from competition by Mytilus.The pilings on docks at the protected western end of Barnegat Inlet possess an intertidal epifaunal community with the following characteristics: (1) a high intertidal zone that includes Balanus balanoides, a second barnacle, Balanus eburneus, and an herbivorous gastropod, Littorina littorea, (2) a mid and low intertidal zone with usually >40% free space in the summer and fall and the remaining area covered by several abundant species with no extreme dominant, and (3) abundant predators, chiefly the oyster drill, Urosalpinx cinerea, the blue crab, Callinectes sapidus, and a mud crab, Neopanope texana sayi. Asterias forbesi, while abundant subtidally, is also occasionally present on intertidal surfaces. Controlled exclusion of predators by caging several replicate pilings at the western end of the inlet reveals that predation prevents monopolization of mid and low intertidal space by the apparent competitive dominant, Mytilus. Predation appears to be a direct cause of the relatively great temporal and spatial heterogeneity in the mid and low intertidal of these pilings.Thus, although the Barnegat Inlet intertidal system appears to follow closely the patterns of community organization described for several other rocky intertidal coastlines, this organizational pattern is noteworthy because it is repeated here in a far more seasonal environment and with a new cast of interacting competitors and predators. That crabs play an important role as predators is novel for North America, but only perhaps because all previous North American studies have ignored the rocky intertidal zones of quiet, estuarine waters where in Europe predatory crabs are known to be extremely significant.This paper is dedicated to the memory of Robert H. MacArthur     

江苏海门蛎岈山牡蛎礁生态现状评价

基于2013—2014年间的生态调查结果,评价了江苏海门蛎岈山牡蛎礁的生态现状。无人机航拍结果显示,江苏海门蛎岈山分布有750个潮间带区牡蛎礁斑块,总面积约为201519.37 m2;与2003年相比,海门蛎岈山牡蛎礁面积约下降了38.8%。活体牡蛎的平均盖度约为66%,2013年5和9月熊本牡蛎Crassostrea sikamea的平均密度分别为(2199±363)个/m2和(2894±330)个/m2。2013年5月海门蛎岈山熊本牡蛎种群的平均肥满度(CI)和性腺指数(GI)分别为(9.76±0.95)%和(1013±82)mg/g,均显著低于浙江象山港养殖的熊本牡蛎种群(P0.05)。海门蛎岈山熊本牡蛎的单倍体多样性和核苷酸多样性指数分别为0.119和0.00028,均高于长江口野生种群和浙江象山港养殖种群。海门蛎岈山熊本牡蛎种群受到尼氏单孢子虫(Haplosporidium nelson)的轻度浸染,其感染率(17.2%)低于浙江象山港养殖群体(47.3%)。泥沙沉积和人类捕捞是江苏海门蛎岈山牡蛎礁面临的主要胁迫因子,今后牡蛎礁恢复的重点是增加附着底物的数量。     

Evaluating Intertidal Oyster Reef Development in South Carolina Using Associated Faunal Indicators

Eastern oyster (Crassostrea virginica) habitat is increasingly being restored for the ecosystem services it provides rather than solely as a fishery resource. Community‐based projects with the goal of ecological restoration have successfully constructed oyster reefs; however, the habitat benefits of these restoration efforts are usually not assessed or reported. In this study, we examined oyster habitat development at five community‐based oyster restoration sites in South Carolina using oyster population parameters, resident fauna densities, and sedimentation (percent sediment coverage) as assessment metrics. All sites included multiple‐aged reefs (1–3 years old) at the time of the fall 2004 sampling. Resident crabs and mussels were abundant at all five sites and crab assemblages were related to the size structure of the oyster microhabitat. Scorched mussel (Brachidontes exustus) abundances were most frequently correlated with oyster and other resident species abundances. Associations among oysters and resident crabs and mussels were not evident when analyses were conducted with higher level taxonomic groupings (e.g., total number of crabs, mussels, or oysters), indicating that species‐level identifications improve our understanding of interactions among reef inhabitants and oyster populations. Community‐based restoration sites in South Carolina provide habitat for mussels and resident crabs, in some cases in the absence of dense populations of relatively large oysters. Monitoring programs that neglect species‐level identifications and counts of mussels and crabs may underestimate the successful habitat provision that can arise independent of large, dense oyster assemblages.     

Rapid development of a restored oyster reef facilitates habitat provision for estuarine fauna

Oyster reef restoration has become a principal strategy for ameliorating the loss of natural Crassostrea virginica populations and increasing habitat provision. In 2014, a large‐scale, high‐relief, 23‐ha subtidal C. virginica reef was restored at the historically productive Half Moon Reef in Matagorda Bay, TX, using concrete and limestone substrates. Encrusting and motile fauna were sampled seasonally until 17 months postrestoration at the restored reef and at adjacent unrestored sites. Restored oysters developed rapidly and were most abundant 3 months postrestoration, with subsequent declines possibly due to interacting effects of larval settlement success on new substrate versus post‐settlement mortality due to competitors and predators. Oyster densities were 2× higher than in a restored oyster population in Chesapeake Bay that was reported to be the largest reestablished metapopulation of native oysters in the world. Resident fauna on the restored reef were 62% more diverse, had 433% greater biomass, and comprised a distinct faunal community compared to unrestored sites. The presence of three‐dimensional habitat was the most important factor determining resident faunal community composition, indicating that substrate limitation is a major hindrance for oyster reef community success in Texas and other parts of the Gulf of Mexico. There were only minor differences in density, biomass, and diversity of associated fauna located adjacent (13 m) versus distant (150 m) to the restored reef. The two substrate types compared had little influence on oyster recruitment or faunal habitat provision. Results support the use of reef restoration as a productive means to rebuild habitat and facilitate faunal enhancement.     

Local grassland restoration affects insect communities

1. It is hypothesised that ecological restoration in grasslands can induce an alternative stable state shift in vegetation. The change in vegetation influences insect community assemblages and allows for greater functional redundancy in pollination and refuge for native insect species. 2. Insect community assemblages at eight coastal California grassland sites were evaluated. Half of these sites had undergone restoration through active revegetation of native grassland flora and half were non‐restored. Insects were collected from Lupinus bicolor (Fabaceae) within 2 × 2‐m² plots in spring 2017. Lupinus bicolor is a common native species that is used in California restoration projects, and home and state landscaping projects. 3. Ordination demonstrated that insect community assemblages were different between restored and non‐restored sites. These differences were seen in insect functional groups as well as taxa‐specific differences and were found to be driven by environmental characteristics such as non‐native forb cover. 4. Functional redundancy of herbivores decreased at restored sites, while pollinators became more redundant compared with non‐restored sites. The assemblages of the common species found at restoration sites contained more native insects than those found at non‐restored sites, including species such as Bombus vosnesenskii. 5. Local grassland restoration has the potential to induce an alternative stable state change and affect insect community assemblages. Additionally, it was found that grassland restoration can be a potential conservation tool to provide refugia for bumblebees (Bombus), but additional studies are required to fully understand its broader applicability.     

Cuing oyster recruitment with shell and rock: implications for timing reef restoration

Reducing uncertainty surrounding the biological responses of degraded habitat is key to providing confidence and efficiency in its restoration. Many coastal habitats are so extensively degraded that organismal responses to restoration interventions are entirely unknown. Among the most degraded coastal ecosystems are oyster reefs, whose restoration typically occurs where oysters are degraded to the point of functional extinction. This loss creates uncertainty on the fundamental processes for reef recovery; the timing of oyster recruitment and their preferred substratum for settlement. Such knowledge can inform restoration strategies to accelerate habitat recovery. Near the site of Australia's largest restoration of native oyster reef, we quantified temporal variability in recruitment of the native flat oyster (Ostrea angasi) and assessed their preference between the settlement substrata deployed for the reef restoration. Combining half a decade of environmental data with oyster recruitment data, we provide a model that identified distinct peaks in oyster recruitment which correlate with food availability and seawater temperature. In addition, oysters preferentially settled on oyster shell relative to other materials used in local restoration. In combination, these results suggest that there are opportunities to augment recruitment through addition of shell substratum synced with recruitment peaks. Our recruitment model likely represents a minor investment with large returns, providing opportunities to capture peak recruitment and greater confidence in utilizing natural recruitment as a restoration resource.     

Historical human activities reshape evolutionary trajectories across both native and introduced ranges

The same vectors that introduce species to new ranges could move them among native populations, but how human‐mediated dispersal impacts native ranges has been difficult to address because human‐mediated dispersal and natural dispersal can simultaneously shape patterns of gene flow. Here, we disentangle human‐mediated dispersal from natural dispersal by exploiting a system where the primary vector was once extensive but has since ceased. From 10th to 19th Centuries, ships in the North Atlantic exchanged sediments dredged from the intertidal for ballast, which ended when seawater ballast tanks were adopted. We investigate genetic patterns from RADseq‐derived SNPs in the amphipod Corophium volutator (n = 121; 4,870 SNPs) and the annelid Hediste diversicolor (n = 78; 3,820 SNPs), which were introduced from Europe to North America, have limited natural dispersal capabilities, are abundant in intertidal sediments, but not commonly found in modern water ballast tanks. We detect similar levels of genetic subdivision among introduced North American populations and among native European populations. Phylogenetic networks and clustering analyses reveal population structure between sites, a high degree of phylogenetic reticulation within ranges, and phylogenetic splits between European and North American populations. These patterns are inconsistent with phylogeographic structure expected to arise from natural dispersal alone, suggesting human activity eroded ancestral phylogeographic structure between native populations, but was insufficient to overcome divergent processes between naturalized populations and their sources. Our results suggest human activity may alter species' evolutionary trajectories on a broad geographic scale via regional homogenization and global diversification, in some cases precluding historical inference from genetic data.     

Plant-soil feedback as a mechanism of invasion by Carpobrotus edulis

Invasive plant species have been suggested to change the composition of the soil community in a way that results in a positive feedback for them and a negative feedback for the native plant community. Carpobrotus edulis, a species native to South Africa, is one of the most aggressive exotic species in Mediterranean Europe. Although several aspects of its invasion biology have been studied, the occurrence of plant-soil feedback has been scarcely investigated. We first checked for the existence of biotic resistance in soils from two invaded sites of Mediterranean Europe and one site in the native area. Secondly, we evaluated the effects of soil conditioning on the germination and plant growth of C. edulis and two key species of native dunes. Finally, we tested the effects of short- and long-term soil conditioning on the performance and reproductive effort of C. edulis. Our results show that at first there is a natural resistance to invasion by the soil biota. Later, biotic resistance in invaded soil is suppressed by the establishment of a soil community that enhances the growth of C. edulis and that negatively influences the growth and survival of the native plants. Long-term soil conditioning in the field resulted in shifts in the balance between vegetative growth and sexual reproduction. Long-term invasion was also reflected in high levels of endophyte colonization by chytrids in roots, although the physiological consequences of this colonization remain unknown. The results obtained illustrate a mechanism that explains how C. edulis breaks the initial biotic resistance of newly-invaded landscapes. Finally, this study highlights the importance of studying plant-soil interactions on different members of the plant community and temporal stages in order to fully understand invasion.     

The value and opportunity of restoring Australia's lost rock oyster reefs

Recognizing the historical loss of habitats and the value and opportunities for their recovery is essential for mobilizing habitat restoration as a solution for managing ecosystem function. Just 200 years ago, Sydney rock oysters (Saccostrea glomerata) formed extensive reef ecosystems along Australia's temperate east coast, but a century of intensive harvest and coastal change now confines S. glomerata to encrusting the hard‐intertidal surfaces of sheltered coastal waters. Despite the lack of natural reef recovery, there appears enormous potential for the restoration of intertidal S. glomerata ecosystems across Australia's east coast, with large anticipated benefits to water quality, shoreline protection, and coastal productivity. Yet, no subtidal reefs remain and the potential for subtidal restoration is a critical knowledge gap. Here, we synthesize historical, ecological, and aquaculture literature to describe a reference system for the traits of S. glomerata reefs to inform restoration targets, and outline the barriers to, and opportunities and methods for, their restoration. These reefs support extremely biodiverse and productive communities and can ameliorate the environmental stress experienced by associated communities. Rock oyster restoration, therefore, provides an ecosystem‐based strategy for assisting the adaptation of marine biodiversity to a changing climate and intensive human encroachment. Though an estimated 92% of S. glomerata ecosystems are lost, there remains great potential to restore these valuable and resilient ecosystems.     

Restoration of Arthropod Assemblages in a Spartina Salt Marsh following Removal of the Invasive Plant Phragmites australis

Invasive plants are one of the most serious threats to native species assemblages and have been responsible for the degradation of natural habitats worldwide. As a result, removal of invasive species and reestablishment of natural vegetation have been attempted in order to restore biodiversity and ecosystem function. This study examined how native arthropod assemblages, an abundant and functionally important group of organisms in many ecosystems, are affected by the incursion of the invasive wetland plant Phragmites australis and if the restoration of the native vegetation in brackish Spartina alterniflora marshes results in the reestablishment of the arthropod community. The invasion of Phragmites into a coastal Spartina marsh in southern New Jersey seriously altered arthropod assemblages and trophic structure by changing the abundance of trophic groups (detritivores, herbivores, carnivores) and their taxonomic composition. Herbivore assemblages shifted from the dominance of external free‐living specialists (e.g., planthoppers) in Spartina to concealed feeders in Phragmites (stem‐feeding cecidomyiids). Moreover, free‐living arthropods in Phragmites became dominated by detritivores such as Collembola and chironomids. The dominant marsh spiders, web‐building linyphiids, were significantly reduced in Phragmites habitats, likely caused by differences in the physical environment of the invaded habitats (e.g., lower stem densities). Thus, trophic structure of arthropod assemblages in Phragmites, as seen in the large shifts in feeding guilds, was significantly different from that in Spartina. Removal of Phragmites with the herbicide glyphosate resulted in the rapid return of Spartina (≤5 yrs). Moreover, return of the dominant vegetation was accompanied by the recovery of most original habitat characteristics (e.g., live and dead plant biomass, water flow rate). The arthropod assemblage associated with Spartina also quickly returned to its preinvasion state and was not distinguishable from that in uninvaded Spartina reference sites. This study provides evidence that the reestablishment of native vegetation in areas previously altered by an invasive plant can result in the rapid recovery of the native arthropod assemblage associated with the restored habitat.     

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.     

Variability of haemocyte and haemolymph parameters in European flat oyster Ostrea edulis families obtained from brood stocks of different geographical origins and relation with infection by the protozoan Bonamia ostreae

A research project to compare productive traits (growth and mortality), disease susceptibility and immune capability between Ostrea edulis stocks was performed. This article reports the results on the immune capability and its relation with infection by the intrahaemocytic protozoan Bonamia ostreae. Four to five oyster spat families were produced from each of four European flat oyster populations (one from Ireland, one from Greece and two from Galicia, Spain) in a hatchery. The spat were transferred to a raft in the Ría de Arousa (Galicia) for on growing for 2 years. Total haemocyte count (THC) and differential haemocyte count (DHC) were estimated monthly through the second year of growing-out. Three types of haemocytes were distinguished: granulocytes (GH), large hyalinocytes (LHH) and small hyalinocytes (SHH). Significant correlations between the mean relative abundance of GH and SHH of the families and the mean prevalence of B. ostreae, the overall incidence of pathological conditions and the cumulative mortality of the families were found; these correlations supported the hypothesis that high %GH and low %SHH would enhance oyster immune ability and, consequently, would contribute to lower susceptibility to disease and longer lifespan. Infection by B. ostreae involved a significant increase of circulating haemocytes, which affected more markedly the LHH type. The higher the infection intensity the higher the %LHH. This illustrates the ability of B. ostreae to modulate the immune responses of the O. edulis to favour its own multiplication. A significant reduction of the phenoloxidase activity in the haemolymph of oysters O. edulis infected by B. ostreae was observed. Nineteen enzymatic activities in the haemolymph of O. edulis and Crassostrea gigas (used as a B. ostreae resistant reference) were measured using the kit api ZYM^®, Biomerieux. Qualitative and quantitative differences in enzyme activities in both haemocyte and plasma fractions between B. ostreae noninfected O. edulis from different origins were recorded. However, no clear positive association between enzyme activity and susceptibility to bonamiosis was found. The only enzyme detected in the resistant species C. gigas that was not found in the susceptible one O. edulis was β-glucosidase (in plasma). B. ostreae infected O. edulis showed significant increase of some enzyme activities and the occurrence of enzymes that were not detected in noninfected oysters. These changes could be due to infection-induced enzyme synthesis by the host or to enzyme synthesis by the parasite.     

Alien barnacle Amphibalanus amphitrite epizoic on two native oyster species in the southern Gulf of Mexico: spatio-temporal variability and current status of its epibiosis

The study examines the current invasive status of the barnacle Amphibalanus amphitrite on natural populations of two native oyster species (the eastern oyster Crassostrea virginica and the mangrove oyster C. rhizophorae) in a estuary of the southern Gulf of Mexico. The spatio-temporal variability of the frequency of occurrence, density and coverage of A. amphitrite on shells of each oyster species was examined and compared with those of other native congeners (A. eburneus, A. venustus and A. improvisus) with which it coexists. These three variables were significantly higher in C. rhizophorae than in C. virginica and were within the range recorded for its congeners. The density of this invasive barnacle varied significantly at spatial and temporal scales. Its highest overall mean density occurred in February (at temperatures 26–27°C and salinities 29–32). Multivariate analyses showed a separation of Amphibalanus assemblages into three main groups according to variations in their composition and abundance among sampling sites and dates. It was also determined that salinity and sedimentation rate were the variables that best explain the distribution of the Amphibalanus species within the estuary. The limited presence of A. amphitrite on C. virginica reefs is likely due to (amongst other factors) the relatively higher sedimentation loads (370–5709?g?m^?2?d^?1) and lower salinities (9–23) recorded in these sites. This study can act as a reference point for the subsequent monitoring of this invasive species, since possible increases in its population density and competitive ability could threaten the integrity of oyster populations and other organisms in the southern Gulf of Mexico.