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Abstract Background Natural populations inhabiting the rocky intertidal experience multiple ecological stressors and provide an opportunity to investigate how environmental differences influence microbiomes over small geographical scales. However, very few microbiome studies focus on animals that inhabit the intertidal. In this study, we investigate the microbiome of the intertidal barnacle Semibalanus balanoides. We first describe the microbiome of two body tissues: the feeding appendages, or cirri, and the gut. Next, we examine whether there are differences between the microbiome of each body tissue of barnacles collected from the thermally extreme microhabitats of the rocky shores’ upper and lower tidal zones. Results Overall, the microbiome of S. balanoides consisted of 18 phyla from 408 genera. Our results showed that although cirri and gut microbiomes shared a portion of their amplicon sequence variants (ASVs), the microbiome of each body tissue was distinct. Over 80% of the ASVs found in the cirri were also found in the gut, and 44% of the ASVs found in the gut were also found in the cirri. Notably, the gut microbiome was not a subset of the cirri microbiome. Additionally, we identified that the cirri microbiome was responsive to microhabitat differences. Conclusion Results from this study indicate that S. balanoides maintains distinct microbiomes in its cirri and gut tissues, and that the gut microbiome is more stable than the cirri microbiome between the extremes of the intertidal.

Acorn barnacle adults experience environmental heterogeneity at various spatial scales of their circumboreal habitat, raising the question of how adaptation to high environmental variability is maintained in the face of strong juvenile dispersal and mortality. Here, we show that 4% of genes in the barnacle genome experience balancing selection across the entire range of the species. Many of these genes harbor mutations maintained across 2 My of evolution between the Pacific and Atlantic oceans. These genes are involved in ion regulation, pain reception, and heat tolerance, functions which are essential in highly variable ecosystems. The data also reveal complex population structure within and between basins, driven by the trans-Arctic interchange and the last glaciation. Divergence between Atlantic and Pacific populations is high, foreshadowing the onset of allopatric speciation, and suggesting that balancing selection is strong enough to maintain functional variation for millions of years in the face of complex demography. [ABSTRACT FROM AUTHOR]

The mannose-6-phosphate isomerase (Mpi) locus in Semibalanus balanoides has been studied as a candidate gene for balancing selection for more than two decades. Previous work has shown that Mpi allozyme genotypes (fast and slow) have different frequencies across Atlantic intertidal zones due to selection on postsettlement survival (i.e., allele zonation). We present the complete gene sequence of the Mpi locus and quantify nucleotide polymorphism in S. balanoides, as well as divergence to its sister taxon Semibalanus cariosus. We show that the slow allozyme contains a derived charge-altering amino acid polymorphism, and both allozyme classes correspond to two haplogroups with multiple internal haplotypes. The locus shows several footprints of balancing selection around the fast/slow site: an enrichment of positive Tajima's D for nonsynonymous mutations, an excess of polymorphism, and a spike in the levels of silent polymorphism relative to silent divergence, as well as a site frequency spectrum enriched for midfrequency mutations. We observe other departures from neutrality across the locus in both coding and noncoding regions. These include a nonsynonymous trans-species polymorphism and a recent mutation under selection within the fast haplogroup. The latter suggests ongoing allelic replacement of functionally relevant amino acid variants. Moreover, predicted models of Mpi protein structure provide insight into the functional significance of the putatively selected amino acid polymorphisms. While footprints of selection are widespread across the range of S. balanoides, our data show that intertidal zonation patterns are variable across both spatial and temporal scales. These data provide further evidence for heterogeneous selection on Mpi. [ABSTRACT FROM AUTHOR]

The intertidal population of the barnacles Semibalanus balanoides has been explored in the estuarine zone of the Tuloma River at the end of the Kola Bay for the first time. Barnacles S. balanoides are typical species for the littoral zone of the northern seas, they populate both the high and low salinity areas along the coast of the Barents Sea. The ecology and distribution of barnacle population in estuarine intertidal zones of the Barents Sea are not studied previously, and under the critical salinity conditions of any estuary, in particular. The investigation have been carried out on the littoral of the western (left) shore of the southern tribe of the Kola Bay – the estuary section from the Tuloma bridge to the Cape Elovy. The studied site is a sandy-boulder beach with stony bars, its length is about 1 150 m, the area is about 126 500 m2. Counting the number of S. balanoides in clusters has been performed on site (without removal of the copepods from the population) by the standard method for intertidal sampling. Simultaneously with the defining the number of barnacles for the period 2003–2014, the measurements of salinity and temperature of water and air have been made. The peculiarity of the estuarine barnacle population is their complete absence in the upper horizon of the littoral, and in the middle and the lower horizons they are found only within the channels of the littoral streams (3–20 specimens in the sample). For the first time the abundance and biomass of barnacles S. balanoides in this estuarine population has been determined, and the causes of their narrow-local distribution on the estuary littoral of the Tuloma River have been revealed. The main ecological factors determining the abundance and specific distribution of barnacles under the conditions of estuarine littoral have been established. The regular increase in the density of adult S. balanoides settlements along the gradient of water salinity from the Tuloma River to the sea has been shown. The long-term fluctuations in the abundance of barnacle S. balanoides have been found, associated, obviously, with climate changes.

Abstract: Predators can impact competition among prey by altering prey density via consumption or by causing prey to modify their traits or foraging behavior. Yet, differences between these two mechanisms may lead to different cascading impacts on lower trophic levels. Using a crab‐snail‐barnacle rocky intertidal food chain, we tested the effects of predation risk from crabs (top predators) on intraspecific competition among snails (intermediate consumers) and emergent indirect effects on the density of and competition among barnacles (basal resources). The per capita foraging and growth rates of snails declined with high conspecific density. Predation risk from crabs, which caused even larger reductions in snail foraging and growth, weakened competition among snails, whereas a 45% increase in barnacle density had no detectable effect on snail competition. Intraspecific competition between barnacles, however, depended on the interactive effects of barnacle density, snail density, and crab predation risk. Barnacles developed hummocking morphologies as they grew and competed for space. Hummock formation (a proxy for competition) increased as a result of either greater initial barnacle density or reduced snail foraging pressure, but these effects depended on predation risk. The effects of crab predation risk on snail foraging behavior weakened an otherwise strong relationship between barnacle density and hummock development: hummocking increased with barnacle density in the absence of crabs but remained relatively high when crabs were present. In communities with similar final barnacle densities, hummocking was more common in those with crabs than those without crabs. The extent to which predators can drive trophic cascades by suppressing the foraging rates of their prey is highly context‐dependent: the positive trait‐mediated indirect effect of predators on basal resource abundance is stronger when many prey respond simultaneously to the threat of predation. However, our results demonstrate that top predators can also enhance competition among basal resources even when their indirect effect on resource abundance is relatively weak. Hence, the cascading effects of predators on competition within lower trophic levels may play an important but under‐appreciated role in the dynamics of basal resource populations and the communities they support. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Abstract: Understanding the role of larger‐scale processes in modulating the assembly, structure, and dynamics of communities is critical for forecasting the effects of climate‐change and managing ecosystems. Developing this comprehensive perspective is difficult though, because species interactions are complex, interdependent, and dynamic through space and time. Typically, experiments focus on tractable subsets of interactions that will be most critical to investigate and explain shifts in communities, but qualitatively base these choices on experience, natural history, and theory. One quantitative approach to identify the putative forces regulating communities, without reducing system complexity, is estimating transition probabilities among species occupying space (i.e., multispecies Markov chain models). Although not mechanistic, these models estimate the relative frequency and importance of ecological pathways in community assembly and dynamics, and can serve as a framework to identify how pathways change across large scales and which are most important to investigate further. Here, we demonstrate this method in the Gulf of Maine (GOM) intertidal zone, where research has largely focused on the local‐scale processes that influence communities, while the mechanisms responsible for more regional shifts in communities are less clear. Transition probabilities of faunal elements were quantified bimonthly for ~2.5 yr in local intertidal communities at three replicate sites in the southern, mid‐coast, and northern GOM. Transitions related to mortality, colonization, and replacement by mussels, barnacles, red algae, and encrusting corallines differed regionally, suggesting specific pathways related to consumer pressure and recruitment vary across the GOM with shifting intertidal community structure. Combined with species abundance data and insights from previous research, we develop and evaluate the pathways by which communities likely change in the GOM. Species interactions in local communities can be complex, and this complexity should be incorporated into hypothesis building, experiments, theory, interpretations, and forecasts in ecology. Such a comprehensive approach will be critical to understand how regional shifts in local interactions can drive large‐scale community change. [ABSTRACT FROM AUTHOR]

We investigated the nonconsumptive effects (NCEs) of predatory dogwhelks (Nucella lapillus) on intertidal barnacle (Semibalanus balanoides) recruitment through field experiments on the Gulf of St. Lawrence coast and the Atlantic coast of Nova Scotia, Canada. We studied the recruitment seasons (May-June) of 2011 and 2013. In 2011, the Gulf coast had five times more nearshore phytoplankton (food for barnacle larvae and recruits) during the recruitment season and yielded a 58% higher barnacle recruit density than the Atlantic coast at the end of the recruitment season. In 2013, phytoplankton levels and barnacle recruit density were similar on both coasts and also lower than for the Gulf coast in 2011. Using the comparative-experimental method, the manipulation of dogwhelk presence (without allowing physical contact with prey) revealed that dogwhelk cues limited barnacle recruitment under moderate recruit densities (Atlantic 2011/2013 and Gulf 2013) but had no effect under a high recruit density (Gulf 2011). Barnacle recruits attract settling larvae through chemical cues. Thus, the highest recruit density appears to have neutralized dogwhelk effects. This study suggests that the predation risk perceived by settling larvae may decrease with increasing recruit density and that prey food supply may indirectly influence predator NCEs on prey recruitment. [ABSTRACT FROM AUTHOR]

We tested whether the patchiness of barnacle settlement was determined by spatial (local) differences in settling and mortality rates. During the period of 2004-2011, annual photo-registration of a barnacles Semibalanus balanoides population was conducted on six flat rocks in the White Sea intertidal zone (Arctic). The average adult population density was 48 ind dm, and the average barnacle age was 3-4 years. According to age structure, during the first year, the number of recruits dropped by three to fourfold; while in the following 2 years, the population density remained nearly the same. The effect of ice friction influence was not recorded, although the rocks were covered with ice in winter. Settling peaks were recorded in 2005, 2007, and 2009. No correlation was observed between the recruit and adult distribution during 2 years of intense settlement. The death rate was uneven on different quadrates along the transects but did not correlate with population density. Because high settling rates and mortality are not linked to specific sites, this suggests that population patchiness is not spatially stable. [ABSTRACT FROM AUTHOR]

We propose an integrative approach that explains patterns of recruitment to adult populations in sessile organisms by considering the numbers of individuals and their body size. A recruitment model, based on a small number of parameters, was developed for sessile organisms and tested using the barnacle Semibalanus balanoides, a marine invertebrate inhabiting North Atlantic intertidal shores. Incorporating barnacle body size improved model fit beyond that based on density alone, showing that growth played an important role in how resource limitation affected survival. Our approach uncovered the following: First, changes in the shape of the recruitment curve resulted from the balance between individual growth and mortality. Second, recruitment was limited by the least plastic trait used to characterise body size, operculum area. Basal area, a trait that responded to increases in barnacle density, did not contribute significantly to explain patterns of recruitment. Third, some temporal variation is explained by changes in the amount of space occupied by shells of dead barnacles: at high cover barnacles are densely packed and these shells remain long after death. Fourth, seasonal variation and spatial variation in survival can be separated from that resulting from resource limitation; survival was predicted for two different shores and four sampling times using a single recruitment model. We conclude that applying this integrative approach to recruitment will lead to a considerable advance in understanding patterns of mortality of early stages of sessile organisms. [ABSTRACT FROM AUTHOR]

Understanding the patterns of genetic variation within and among populations is a central problem in population and evolutionary genetics. We examine this question in the acorn barnacle, Semibalanus balanoides, in which the allozyme loci Mpi and Gpi have been implicated in balancing selection due to varying selective pressures at different spatial scales. We review the patterns of genetic variation at the Mpi locus, compare this to levels of population differentiation at mtDNA and microsatellites, and place these data in the context of genome-wide variation from high-throughput sequencing of population samples spanning the North Atlantic. Despite considerable geographic variation in the patterns of selection at the Mpi allozyme, this locus shows rather low levels of population differentiation at ecological and trans-oceanic scales (FST ∼ 5%). Pooled population sequencing was performed on samples from Rhode Island (RI), Maine (ME), and Southwold, England (UK). Analysis of more than 650 million reads identified approximately 335,000 high-quality SNPs in 19 million base pairs of the S. balanoides genome. Much variation is shared across the Atlantic, but there are significant examples of strong population differentiation among samples from RI, ME, and UK. An FST outlier screen of more than 22,000 contigs provided a genome-wide context for interpretation of earlier studies on allozymes, mtDNA, and microsatellites. FST values for allozymes, mtDNA and microsatellites are close to the genome-wide average for random SNPs, with the exception of the trans-Atlantic FST for mtDNA. The majority of FST outliers were unique between individual pairs of populations, but some genes show shared patterns of excess differentiation. These data indicate that gene flow is high, that selection is strong on a subset of genes, and that a variety of genes are experiencing diversifying selection at large spatial scales. This survey of polymorphism in S. balanoides provides a number of genomic tools that promise to make this a powerful model for ecological genomics of the rocky intertidal. [ABSTRACT FROM AUTHOR]

Abstract: Amphibalanus (=Balanus) amphitrite is a tropical/sub-tropical barnacle species which is naturally exposed to sea temperatures of 16–27 °C throughout its larval development. An established technique widely employed by investigators of A. amphitrite settlement involves the storage of cyprids at 4–6 °C prior to their use in bioassays. Our study focuses on the effects on ageing temperature on settlement and discrimination of A. amphitrite larvae. Using cyprids aged at 2 temperatures, 6° and 23 °C, we confirmed the general trend in the literature that young d0 cyprids of this species do not appear competent to settle. Performing cyprid settlement assays at 20, 25 or 28 °C, we observed that the proportion of settled cyprids when incubated at 28 °C was greater than that of cyprid incubated at 25 °C and 20 °C. Settlement rates of cyprids aged at 6 °C and 23 °C increased relative to age from d1 to d10 irrespective of temperature. Cyprids lost the ability to undergo attachment and metamorphosis at d14 or d15 when aged at 6 °C or 23 °C respectively. In the choice assays executed in this study, cyprids generally chose to settle on adult extract-treated areas. But localised settlement on adult extract-treated areas decreased with time. Choice settlement assays at set age intervals during the cypris larval stage showed that cyprids were able to discriminate between con- and allospecific adult extracts. The duration and the temperature of cyprids storage influenced selectivity. [Copyright &y& Elsevier]

ABSTRACT Aim Marine sessile inter-tidal invertebrates are expected to undergo shifts in distribution due to climate change. Using a combination of survey and transplant data with thermal modelling, we investigated the role of climate on the poleward contraction of the southern range edge of the north temperate barnacle Semibalanus balanoides. Location Western Atlantic of the United States. Methods Barnacle surveys were conducted along the east coast of the United States in 1963 and 2007. Presence, absence and abundance data were collected and the time periods were compared. Transplant experiments monitoring survival with relation to temperature were conducted upon S. balanoides along the more southerly portion of their range, and modelling predicting barnacle survival with relation to biogeography was completed. Results The southern limit of S. balanoides has contracted approximately 350 km to the north. Main conclusions The changes thus far observed in climate along the east coast of the United States have contributed to the southern limit range contraction of S. balanoides. Further changes in the biogeography of S. balanoides are expected with continued climate warming. [ABSTRACT FROM AUTHOR]

Sex allocation theory for simultaneous hermaphrodites predicts increases in relative allocation to male-specific function as competition for fertilizations increases. Theoretical models developed specifically for competing acorn barnacles predict that the proportional allocation to male function increases toward an asymptote of 50% as the number of competitors for fertilizations increases. Experimental manipulations were used to investigate how mate competition affected both relative and absolute allocation to the sex functions for two species of acorn barnacle: Semibalanus balanoides and Balanus glandula. The ratio of male to female allocation did not increase with the number of competitors for either species. However, both species showed increased allocation to male function (estimated as total mass of sex-specific tissues) with increased crowding. Allocation to female function seemed to be limited by other factors and did not vary with mating group size as predicted. Allocation to male and female function were both positively related to body size, but a trade-off between male and female function, a key assumption of prior models, was not observed. [ABSTRACT FROM AUTHOR]

Population genetic characteristics are shaped by the life-history traits of organisms and the geologic history of their habitat. This study provides a neutral framework for understanding the population dynamics and opportunities for selection in Semibalanus balanoides, a species that figures prominently in ecological and evolutionary studies in the Atlantic intertidal. We used mitochondrial DNA (mtDNA) control region (N = 131) and microsatellite markers (∼40 individuals/site/locus) to survey populations of the broadly dispersing acorn barnacle from 8 sites spanning 800 km of North American coast and 1 site in Europe. Patterns of mtDNA sequence evolution were consistent with larger population sizes in Europe and population expansion at the conclusion of the last ice age, approximately 20 000 years ago, in North America. A significant portion of mitochondrial diversity was partitioned between the continents (ϕST = 0.281), but there was only weak structure observed from mtDNA within North America. Microsatellites showed significant structuring between the continents (FST = 0.021) as well as within North America (FST = 0.013). Isolation by distance in North America was largely driven by a split between populations south of Cape Cod and all others (P < 10−4). The glacial events responsible for generating allelic diversity at mtDNA and microsatellites may also be responsible for generating selectable variation at metabolic enzymes in S. balanoides. [ABSTRACT FROM PUBLISHER]

Predators can initiate trophic cascades by consuming and/or scaring their prey. Although both forms of predator effect can increase the overall abundance of prey's resources, nonconsumptive effects may be more important to the spatial and temporal distribution of resources because predation risk often determines where and when prey choose to forage. Our experiment characterized temporal and spatial variation in the strength of consumptive and nonconsumptive predator effects in a rocky intertidal food chain consisting of the predatory green crab (Carcinus niaenas), an intermediate consumer (the dogwhelk, Nucella lapillus), and barnacles (Sernihalanus halanoides) as a resource. We tracked the survival of individual barnacles through time to map the strength of predator effects in experimental communities. These maps revealed striking spatiotemporal patterns in Nucella foraging behavior in response to each predator effect. However, only the nonconsumptive effect of green crabs produced strong spatial patterns in barnacle survivorship. Predation risk may play a pivotal role in determining the small-scale distribution patterns of this important rocky intertidal foundation species. We suggest that the effects of predation risk on individual foraging behavior may scale up to shape community structure and dynamics at a landscape level. [ABSTRACT FROM AUTHOR]

The article discusses a study on the influence of previous disturbance on the effects of trampling on barnacles. The study was conducted at two sites in Acadia National Park at Mount Desert Island in Maine. The barnacle Semibalanus balanoides dominate the high intertidal zones of both sites. The biotic factors such as predator abundance and human visitation and abiotic factors such as slope and temperature that could possibly influence barnacle cover in each site were compared. The study found a potential relationship between spatial variation in barnacle survivorship and intensity of predation.

The article highlights a novel flume that can describe a barnacle cyprids' swimming behavior in a laboratory downwelling flow. The authors were able to create the homogenous downwelling velocity field they used in the study by pumping seawater through a cylindrical observation chamber with diffusers. They found that larvae of Semibalanus balanoides cyprids swam upward with a maximum vertical velocity of 72.3 millimeter s¹ despite exhibiting different behaviors. They concluded that swimming against downwelling flow may be an adaptive behavior that is used for shoreward transport.

Quantifying species interaction strengths enhances prediction of community dynamics, but variability in the strength of species interactions in space and time complicates accurate prediction. Interaction strengths can vary in response to density, indirect effects, priority effects or a changing environment, but the mechanism(s) causing direction and magnitudes of change are often unclear. We designed an experiment to characterize how environmental factors influence the direction and the strength of priority effects between sessile species. We estimated per capita non-trophic effects of barnacles (Semibalanus balanoides) on newly settled germlings of the fucoid, Ascophyllum nodosum, in the presence and absence of consumers in experiments on rocky shores throughout the Gulf of Maine, USA. Per capita effects on germlings varied among environments and barnacle life stages, and these interaction strengths were largely unaltered by changing consumer abundance. Whereas previous evidence shows adult barnacles facilitate fucoids, here, we show that recent settlers and established juveniles initially compete with germlings. As barnacles mature, they switch to become facilitators of fucoids. Consumers caused variable mortality of germlings through time comparable to that from competition. Temporally variable effects of interactors (e.g. S. balanoides), or spatial variation in their population structure, in different regions differentially affect target populations (e.g. A. nodosum). This may affect abundance of critical stages and the resilience of target species to environmental change in different geographical regions. [ABSTRACT FROM AUTHOR]

Variation in physical factors, such as slope, orientation, and wind exposure, shapes thermal conditions. Variation in substrate size is common in many habitats, but its thermal consequences for organisms are not well characterized. Larger substrates should remain more thermally stable and act as thermal refuges for associated organisms during short, thermally stressful periods such as midday temperature peaks or tidal exposure. In observations and a transplant and thermal integration experiment, we found that larger rock substrates stayed cooler and facilitated greater survival of the barnacle Semibalanus balanoides in the high intertidal relative to small substrates during the hot summer months in southern New England, USA. However, in thermally benign northern New England, rock substrate size had no effect on barnacle distributions, indicating that the thermal effects of substrate size are mediated by regional climate. [ABSTRACT FROM AUTHOR]

Abstract: Larvae of many benthic invertebrates settle on surfaces where they metamorphose into juveniles if suitable substrata are available, and are responsible for the major costs of biofouling. When assessing new formulations or compounds for potential antifouling (AF) application, constraints such as seasonal availability may restrict most bioassays to relatively few taxa and species. For example, amongst barnacles, Amphibalanus amphitrite is popular as a test organism but is it really representative of other barnacle species? In order to test this hypothesis, we have chosen to work with marine natural extracts as a probe. Indeed, one substitution technology to toxic metal-based coatings to control fouling is the development of AF coatings with active compounds derived from marine organisms or analogues of the lead compounds. In this study, the AF activity and toxicity of extracts from 30 algae from the North East Atlantic coast were investigated for their potential anti-settlement activities against larvae of two species of barnacle, A. amphitrite and Semibalanus balanoides. As a trend, most of the active extracts displayed activity towards S. balanoides, only few displayed targeted activity against A. amphitrite, or against both species. In order to better understand if this tendency could be linked to chemical ecology, surface extracts were prepared on a selection of species. The results highlight that surface extracts of algae all displayed highest levels of activity than total extracts when tested on S. balanoides. This difference illustrates that specific compounds in their ecological context can have potentially a better efficacy on target species. [ABSTRACT FROM AUTHOR]

The article presents a study which examines the physiological effects of mannose-6-phospate isomerase (Mpi) and glocuse-6-phospate isomerase (Gpi) polymorphisms in the acorn barnacle Semibalanus balanoides from Narrangansett Bay, Rhode Island. It found that mortality from thermal stress was enhanced under low salinity. It observed that across all low salinity treatments was non-significant trend of decreased fitness associated with the Mpi-SS genotype. It also determined that cirral length variation was correlated with Mpi genotypes in a pattern consistent with survivorship data.

The article presents a study on the effects of spatial oceanographic processes on the growth of interdidal barnacle Semibalanus balanoides in Scotland. The study surveyed populations on 259 sites around the nation in the summers of year 2001 to 2006. Densities of adult and juvenile barnacle were compared through wave fetch, remotely sensed chlorophyll a (chl a) concentration, and seasonal sea surface temperature (SST). Results show that population densities of adults and juveniles have increased with wave fetch while the chl a influenced the density at high shore levels. It concludes that large-scale oceanographic effects on food concentration causes large-scale ecological patterns in the S. balanoides.

Acorn barnacles are important model organisms for the study of sex allocation. They are sessile, nonselfing hermaphrodites that copulate with penises that have been suggested to be phenotypically plastic. On wave-exposed shores, Semibalanus balanoides develop penises with relatively greater diameter whereas in wave-protected sites they are thinner. A reciprocal transplant experiment between wave-exposed and protected sites tested whether these exposure-specific morphologies have adaptive value. Mating success was compared over a range of distances to compare the ability of barnacles to reach mates. Barnacles that grew in the wave-protected site and mated in the wave-protected site fertilized more broods at increasing distances than those transplanted to the wave-exposed site. For barnacles that developed in the wave-exposed site, there was no difference in the ability to fertilize neighbors between sites of differing exposure. This study demonstrates the adaptive value of plasticity in penis morphology. The results suggest a trade-off between development of a penis adapted to wave exposure and the ability to fertilize distant mates. Barnacles in different physical environments are limited by different factors, which may limit numbers of potential mates, constrain optimal sex allocation strategies and alter reproductive behavior. [ABSTRACT FROM AUTHOR]

The strength by which species interact can vary throughout their ontogeny, as environments vary in space and time, and with the density of their populations. Characterizing strengths of interaction in situ for even a small number of species is logistically difficult and may apply only to those conditions under which the estimates were derived. We sought to combine data from field experiments estimating interaction strength of life stages of the barnacle, Semibalanus balanoides, on germlings of Ascophyllum nodosum, with a model that explored the consequences of variability at per capita and per population levels to the abundance of year-old algal recruits. We further simulated how this interaction affected fucoid germling abundance as the timing of their respective settlements varied relative to one another, as occurs regionally across the Gulf of Maine, USA. Juvenile S. balanoides have a weak estimated per capita effect on germlings. Germling populations are sensitive to variation in per capita effects of juvenile barnacles because of the typically large population sizes of the latter. However, high mortality of juvenile barnacles weakens the population interaction strength over time. Adult barnacles probably weakly facilitate fucoid germlings, but greater survival of adults sustains the strength of that interaction at the population level. Germling abundance is positively associated with densities of adult barnacles and negatively associated with that of juvenile barnacles. Metamorphosing cyprid larvae have the strongest per capita effect on germling abundance, but the interaction between the two stages is so short-lived that germling abundance is altered little. Variation in the timing of barnacle and A. nodosum settlement relative to one another had very little influence on the abundance of yearling germlings. Interactions between barnacles and germlings may influence the demographic structure of A. nodosum populations and the persistence of fucoid-dominated communities on sheltered rocky shores in New England. [ABSTRACT FROM AUTHOR]

Biotic interactions will modulate species' responses to climate change. Many approaches to predicting the impacts of climate change on biodiversity so far have been based purely on a climate envelope approach and have not considered direct and indirect species interactions. Using a long-term observational data 1set (>30 years) of competing intertidal barnacle species, we built a hierarchy of age-structured two-taxa population models (Semibalanus balanoides vs. Chthamalus montagui and C. stellatus combined as one taxon) to test if the presence of a dominant competitor can mediate climatic influence on the subordinate species. Models were parameterized using data from populations on the south coast of southwest England and verified by hindcasting using independent north coast population data. Recruitment of the dominant competitor, S. balanoides, is driven by temperature. The mechanisms of competition explored included simple space preemption and temperature-driven interference competition. The results indicate that interspecific competition between juvenile barnacles is important in regulating chthamalid density but not that of the dominant competitor S. balanoides. Simulations were carried out using alternative future climate scenarios to predict barnacle population abundance over the next century. Under all emission scenarios, the cold-water S. balanoides is predicted to virtually disappear from southwest England by the 2050s, leading to the competitive release of Chthamalus throughout the entire region and thereby substantially increasing its abundance and occupied habitat (by increasing vertical range on the shore). Our results demonstrate that climate change can profoundly affect the abundance and distribution of species through both the direct effects of temperature on survival, and also by altering important negative interactions through shifting competitive balances and essentially removing dominant competitors or predators. Climate change impacts on organisms are unlikely to lead only to straightforward, easily predictable changes in population size and distribution. The complex, indirect effects of climate change need to be taken into account if we are to accurately forecast the long-term effects of global warming. [ABSTRACT FROM AUTHOR]

The interplay between local and large spatial scale processes in open systems is often dependent upon ecological context and species specific factors such as longevity, dispersal capability, or vulnerability to predation. When disturbance clears patches in open systems, the successful reestablishment of adult colonizers and the trajectory of succession may depend upon both the scale of the disturbance event and scale of life history characteristics. Here we examine the links between the size of a disturbance event and long. term patterns of variation in recruitment, density, and percent cover in a relatively short-lived but long-range disperser, the acorn barnacle Semibalanus balanoides, and a relatively long lived and short range disperser, the fucoid alga Fucus vesiculosus. Further, a model of linked scale-dependent processes is developed and we explore how long term patterns of scale- dependent recruitment are related to successful establishment of adults. Recruitment, densities, and cover were monitored for five years for both species in a single experiment using clearings of different sizes spread over two north-facing and two south-facing bays on Swan's Island, Maine, USA. Barnacle recruitment was particularly variable in small clearings and was lower overall at more interior sites within bays. While local-scale factors also strongly influenced patterns of variation in fucoid recruitment, fucoids dominate surface cover in large clearings and in south-facing bays. Recruitment was a poor predictor of density, whereas density itself was a good predictor of percent cover for both species. These results indicate that scale dependencies, location-specific factors, and life history traits contribute to patterns of community development on rocky shores and may ultimately determine whether an open patch converges to or diverges from its initial community state. [ABSTRACT FROM AUTHOR]

Intertidal organisms are often assumed to live close to their thermal limits, and have emerged as potential early indicators of the effects of climate change. We compared our survey of the 2004–2006 geographic distribution of the barnacle Semibalanus balanoides to its distribution in 1872, 1955, 1963, 1971, and 1985, from surveys by Fischer, Crisp, Fischer-Piette, Barnes, Powell, and Southward. The southern geographic limit has retreated 300 km in France since 1872, at a rate of 15 to 50 km per decade. We compared our 2006 survey of the geographic distribution of the polychaete Diopatra neapolitana to its distribution in 1893–1923, from surveys by Saint-Joseph and Fauvel, and its distribution in 1969–1976 from surveys by Glémarec. The northern geographic limit of this species has advanced 300 km in France since 1893 at similar rates to Semibalanus. We used NOAA weather reanalysis data and our mechanistic simulation model of intertidal animal body temperatures to hindcast the thermal environmental change near historical geographic limits in Europe for the past 55 years. Results indicate that changes in the southern limit of S. balanoides are due to intolerance of winter body temperatures above 10°C, leading to reproductive failure. Results for Diopatra are ambiguous: based on the northern extension of its range, either cold winters or cool summers limit its range, while gaps in its distribution are consistent with limitation by cooler summer conditions. The parallel shifts of D. neapolitana on sedimentary shores and Semibalanus on rocky shores suggest that similar climatic factors control the geographic limits of both species. The intertidal zone is a model system for examining the effects of climate change on biogeographic change both because of the rapidity of its response, and because the rich historical record allows direct tests of hypotheses. [ABSTRACT FROM AUTHOR]

Abstract: In this paper, we examine metal concentrations in the water and in the crustacean Balanus balanoides from the Huelva estuary, one of the most polluted estuaries in Europe. Metal levels in waters are very high, especially those of Zn, Fe and Cu. Zn presents the highest concentrations, with a mean value of 690μgl−1 in 2001 and 301μgl−1 in 2002. As the water flows down through the estuary toward the sea, the metal concentrations drop sharply and the pH rises. The metal concentrations in balanoides tissues are, in general, very high, undoubtedly due to the high metal pollution of the water where it lives. Metal concentrations in Balanus balanoides tissues behave similarly to those in the water, reaching maximums in the upper part of the estuary and diminishing as we approach the sea. The element that reaches the highest levels in Balanus balanoides is Zn, with a mean value of 54.6gkg−1 in 2001 and 29.9gkg−1 in 2002, followed by Cu and Fe. There is a significant correlation (p<0.01) for concentrations of Cd, Cu, Fe, Mn, Ni and Zn in Balanus balanoides relative to their concentration in waters. Barnacles showed a great capacity to accumulate metals, especially Zn, Cu and Fe. Based on the results obtained, we can conclude that Balanus balanoides is a good tool for monitoring trace metals in the Huelva estuary. [Copyright &y& Elsevier]

In northwest Europe, sheltered rocky shores are dominated by fucoid canopy algae and barnacles are rare, although the latter are extremely abundant on exposed shores. The supply of the intertidal barnacle Semibalanus balanoides (L.) to sheltered, fucoid dominated rocky shores was investigated to determine the importance of larval supply in limiting the abundance of adults in shelter. Larval supply was measured at two spatial scales, at the scale of shore (100s of metres), by comparing larval concentrations at exposed and sheltered sites, and at a smaller spatial scale (m), by examining the role of fucoid canopies in limiting supply to the substratum. Replicate plankton trawls were carried out above the intertidal zone at high water at two sheltered sites and nearby exposed headlands. The concentration of S. balanoides cyprid larvae was significantly higher at the sheltered sites on two out of three sampling occasions with up to 14 times greater larvae on one occasion than the nearby exposed site. The effect of the macroalgal canopy on supply to the substratum was assessed in two ways: directly, by pumping water from the substratum in areas with and without Ascophyllum nodosum (L.) Le Jolis, and indirectly by measuring cyprid settlement in a canopy-manipulation experiment. Pumped plankton samples from mid tide level showed that the A. nodosum canopy did not form a barrier to larval supply and may have had a positive effect on larval concentrations at the substratum. Cyprid settlement was assessed in the mid shore A. nodosum and low shore Fucus serratus L. zones to areas with canopy algae (but protected from the sweeping effects of macroalgal fronds) and without canopy. Settlement over three consecutive 24-h periods showed a consistent pattern; settlement was consistently lower beneath the F. serratus canopy than in cleared areas, suggesting that this algal species forms a barrier, limiting supply of cyprid larvae to the substratum. [ABSTRACT FROM AUTHOR]