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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 million years 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.

Pelagic larvae of benthic organisms comprise a substantial part of the coastal Arctic zooplankton community in spring–summer. We studied the timing, growth, and pelagic duration of Cirripedia larvae in Adventfjorden, a high-Arctic fjord in Spitsbergen, Svalbard. Two distinct abundance peaks were found: one in early May (~ 25.000 ind. m−3) and another one in late May (~ 35,580 ind m−3). DNA barcoding based on the COI gene was used to identify the barnacle larvae to species. Whereas both Balanus balanus and Semibalanus balanoides were present, the first one dominated (50–100%) the barnacle abundance. High resolution sampling and size measurements of Cirripedia larvae revealed that these larvae most likely originated from a single spawning event. Development of the larvae suggested a pelagic residence time of roughly 2 months for B. balanus and at least 1 month for S. balanoides in the Arctic. Long pelagic residence time, large potential for biofouling on ships and larger plastic debris, combined with the disappearance of landfast sea ice and less ice scouring opens up new opportunities for barnacles to colonize the high-Arctic littoral zone. In a future warmer Arctic, we therefore expect establishment of new, more temperate Cirripedia species in Svalbard.

Environmental controls on the biogeographic distribution of species are becoming increasingly relevant under the present climate change conditions. The reproductive cycle of the acorn barnacle Semibalanus balanoides at its southernmost European distribution limit in Galicia (Northwest Iberian Peninsula) was studied in two locations (42.57°N 8.96°W, 42.61°N 8.89°W) with different temperatures during a 4-year time series (2012–2016), where an isolated population breeds even though the temperature thresholds documented for northern populations of the species are exceeded. Ovary production, total number of embryos and pre-hatching developmental stage were significantly higher in the colder location than in the warmer one. Fecundity was higher in colder reproductive seasons mediated by upwelling regime and food supply 3 months prior to fertilization. Investment in ovary tissue was favored by high temperatures and food availability, whereas suspended sediment was negatively correlated with investment in ovary tissue. Larvae release was associated with high temperatures and suspended sediment. Phenology of reproductive events was similar to what was described in farther north populations. Results indicated that reproductive output of the species at its southernmost European distribution limit is comparable to that reported in northern latitudes and suggest adaptation or acclimatization to local environmental conditions as maturation of gonads and fertilization was successfully carried out at higher temperatures than documented in northern European populations.

The northern acorn barnacle (Semibalanus balanoides) is a robust system to study the genetic basis of adaptations to highly heterogeneous environments. Adult barnacles may be exposed to highly dissimilar levels of thermal stress depending on where they settle in the intertidal (i.e., closer to the upper or lower tidal boundary). For instance, barnacles near the upper tidal limit experience episodic summer temperatures above recorded heat coma levels. This differential stress at the microhabitat level is also dependent on the aspect of sun exposure. In the present study, we used pool-seq approaches to conduct a genome wide screen for loci responding to intertidal zonation across the North Atlantic basin (Maine, Rhode Island, and Norway). Our analysis discovered 382 genomic regions containing SNPs which are consistently zonated (i.e., SNPs whose frequencies vary depending on their position in the rocky intertidal) across all surveyed habitats. Notably, most zonated SNPs are young and private to the North Atlantic. These regions show high levels of genetic differentiation across ecologically extreme microhabitats concomitant with elevated levels of genetic variation and Tajima's D, suggesting the action of non-neutral processes. Overall, these findings support the hypothesis that spatially heterogeneous selection is a general and repeatable feature for this species, and that natural selection can maintain functional genetic variation in heterogeneous environments. publishedVersion

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.

Human growth has caused an unprecedented increase in artificial light at night (ALAN). In coastal habitats, many species rely on day/night cycles to regulate various aspects of their life history and these cycles can be altered by this stressor. This study assessed the influence of ALAN on the early (cyprid) and late (spat) settlement stages of the acorn barnacle Semibalanus balanoides, a species widely distributed in natural and man-made coastal habitats of the North Atlantic. A newly designed settlement plate, originally for studies in rocky intertidal habitats in the southeast Pacific, was adapted to measure settlement rates on man-made habitats -wharf seawalls- located in Atlantic Canada. Plates equipped with a small LED diode powered by an internal battery (ALAN plates) were used to quantify settlement rates in comparison to plates lacking a light source (controls). These plates were deployed for 6 d in the mid-intertidal levels, where adult barnacles were readily visible. ALAN and control plates collected large number of settlers and showed to be suitable for this type of man-made habitats. The number of early settlers (cyprids) did not differ between plates but the number of late settlers (spat) was significantly lower in ALAN plates than in controls. These results suggest that light pollution has little influence on the early stages of the acorn barnacle settlement but is clearly detrimental to its late stages. As barnacles dominate in many natural and man-made hard substrates, it is likely that ALAN also has indirect effects on community structure.

Climate change has already altered the environmental conditions of the world’s oceans. Here we report declines in gastropod abundances and recruitment of mussels (Mytilus edulis) and barnacles (Semibalanus balanoides) over the last two decades that are correlated with changes in temperature and ocean conditions. Mussel recruitment is declining by 15.7% per year, barnacle recruitment by 5.0% per year, and abundances of three common gastropods are declining by an average of 3.1% per year (Testudinalia testudinalis, Littorina littorea, and Nucella lapillus). The declines in mussels and the common periwinkle (L. littorea) are correlated with warming sea temperatures and the declines in T. testudinalis and N. lapillus are correlated with aragonite saturation state, which affects rates of shell calcification. These species are common on shores throughout the North Atlantic and their loss is likely to lead to simplification of an important food web on rocky shores., Petraitis and Dudgeon report a decline in mussel and gastropod abundance and recruitment on rocky intertidal shores of the Gulf of Maine over a period of 20 years. These declines are correlated with increased water temperatures and aragonite saturation state, which affects rates of shell calcification, and suggests environmental impacts on the food webs of rocky shores.

A critical question in marine ecology is understanding how organisms will cope with environmental conditions under climate change. Increasing temperatures not only have a direct effect on marine organisms but may also lead to food limitation through for example trophic mismatches, or by the increased metabolic demands imposed by developing at high temperatures. Using barnacles from a population of North Wales, we studied the combined effect of temperature and food density on the survival, settlement success, developmental time and body size of larvae of the native barnacle Semibalanus balanoides and its exotic competitor, the barnacle Austrominius modestus. Larvae were reared at similar food levels but at temperature ranges which varied among species reflecting their different phenology and tolerances. For S. balanoides (spring larval release) we used a lower temperature of 9 °C, reflecting spring temperatures from N Wales to SW England, and 15 °C representing warmer conditions; for A. modestus (summer larval release) a typical summer temperature for this geographic range of 15 °C was used with a raised temperature of 18 °C. Larvae were reared under controlled conditions in automated, computer programmable incubators and fed diatoms (Skeletonema costatum) at three food levels. We found stress effects of food limitation on larval performance of S. balanoides. While survival during naupliar development was little affected by food and temperature, low food levels strongly depressed survival and settlement during the cyprid stage of S. balanoides at both tested temperatures, but especially at 15 °C. By contrast, at the tested temperatures little effects were found on survival and settlement success in the exotic A. modestus. Both species delayed development in response to low food levels while S. balanoides cyprids showed decreased body size at the high tested temperature. The main impact occurred as a delayed effect, at the time when cyprids attempt to settle, rather than as an effect on naupliar survival or metamorphosis to the cyprid stage. Response in body size and developmental time may have costs at the time of metamorphosis (delayed settlement) or after metamorphosis. Overall, our experiments suggest that as temperature increases, settlement success of S. balanoides larvae (but not that of its competitor A. modestus) will become more sensitive to conditions of food limitation, imposed for instance by phenological mismatches with periods of phytoplankton peak.

Population connectivity plays a major role in the ecology and evolution of marine organisms. In these systems, connectivity of many species occurs primarily during a larval stage, when larvae are frequently too small and numerous to track directly. To indirectly estimate larval dispersal, ocean circulation models have emerged as a popular technique. Here we use regional ocean circulation models to estimate dispersal of the intertidal barnacle Semibalanus balanoides at its local distribution limit in Southwest England. We incorporate historical and recent repatriation events to provide support for our modeled dispersal estimates, which predict a recolonization rate similar to that observed in two recolonization events. Using graph theory techniques to describe the dispersal landscape, we identify likely physical barriers to dispersal in the region. Our results demonstrate the use of recolonization data to support dispersal models and how these models can be used to describe population connectivity.

A twenty-five year (1994–2018) survey at 10 rocky sites within Lough Hyne Marine Reserve (SW Ireland) showed that, despite considerable spatial and temporal patchiness, three barnacle species (Chthamalus stellatus, Semibalanus balanoides, and Austrominius modestus) declined in overall abundance in the early 2000s, particularly at three sites. There was no clear response of A. modestus or C. stellatus to the cold winters of 2010 and 2011. In 2004, however, S. balanoides declined suddenly, but showed a slight recovery after the cold winter of 2010 to maintain sparse populations in the lough, but remained abundant outside the lough. A fourth species, Chthamalus montagui, showed a different pattern, with wide fluctuations in abundance but no overall decline. Changes in the Atlantic Multidecadal Oscillation may have influenced the overall composition of barnacle populations. Rising maximum air temperatures, resulting in high body temperatures recorded for A. modestus even during mild days, are suggested to have impacted the barnacle populations as a result of the prolonged aerial exposure times caused by the lough's asymmetric tidal pattern.

In intertidal communities, consumers (especially carnivorous gastropods) have historically been thought to exert strong top-down control on community composition by regulating competitively dominant mussels and barnacles. This paradigm was formulated based on wire mesh cage exclusion experiments, which have well-known artifacts such as altering hydrodynamics and excluding non-target, but potentially important, consumer species. Recent research highlights the potential importance of multiple consumers that are often subtle and transient, as well as the modifications of consumer pressure by spatial and temporal environmental variability and bottom-up processes. Manipulative experiments that target individual taxa will be essential to more clearly identify their roles in complex ecological communities. The predatory gastropod Nucella lapillus has long been considered an important consumer controlling the structure and dynamics of intertidal communities in the Gulf of Maine. To test the role of N. lapillus in shaping community structure, we manually reduced its densities for 2.33 years. Species composition, stable stage community composition (based on a Markov model), and transition probabilities (as measures of ecological pathway strength) were compared between treatments (control vs. density reductions), and among seasons (spring, summer, fall, winter). In contrast to previous studies, exclusions had no effect on community composition or long-term Markov model predictions of stable stage community structure. Reducing N. lapillus abundance increased the persistence and reduced the mortality of the barnacle Semibalanus balanoides in the model, but did not affect blue mussels, Mytilus edulis , in a similar way. Reducing N. lapillus abundance had additional indirect effects of increasing Ralfsia spp. mortality and increasing hydroid persistence. Despite differences in transition probabilities among treatments, compensatory changes in direct and indirect pathways led to communities that converged over the long-term. Our results contrast previous estimates of the role of N. lapillus in intertidal communities and might reflect low mussel recruitment, predation by alternate consumers (e.g., C. maenas , C. borealis , Tautogolabrus adspersus , birds), or spatial and/or temporal environmental variation that influenced the role of N. lapillus in structuring these communities. The role of N. lapillus may be more limited, or variable, than often assumed in the Gulf of Maine, and the methods manipulating broad functional groups such as “consumers”, though useful in developing basic conceptual models, blur the effects of individual species in community assembly and dynamics.

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.

In the high Arctic, recruitment of hard-bottom benthic organisms has been studied at single locations, but little is known about how it varies spatially or temporally, or how it is influenced by abiotic factors. In this study, settlement plates were simultaneously deployed at five locations in three Svalbard (Norway) fjords at depths ranging from 7 m to 215 m. Recruitment was significantly different among fjords and among locations within a fjord. Recruits at each site co-occurred randomly even though interspecific overgrowth was observed. This finding provides further evidence that there is not necessarily a relationship between non-random co-occurrence and interspecific competition, such as is traditionally assumed for other isolated, island-like habitats. We found significantly lower recruitment in an Arctic-influenced fjord than in more Atlantic-influenced fjords. The abundance and richness of recruits was significantly lower in fall-winter than in spring-summer, but the spirorbid Circeis armoricana recruited in high abundance in fall-winter. Both the abundance and taxonomic richness of recruits declined exponentially with depth, with the hydroid Stegopoma plicatile dominating at 215 m in an Atlantic-influenced fjord. The most abundant recruiting taxa (C. armoricana, Semibalanus balanoides, Harmeria scutulata, Celleporella hyalina) can be described as pioneer (early-succession) species. Crustose coralline algae, a slow-growing superior competitor, also recruited at one location. Recruitment in Svalbard is highly variable, both spatially and temporally, and our results show the influence of local factors such as adult species composition.

Space is one of the primary limiting resources for organisms on the intertidal rocky shore. This paper examined the effect of reduced density on key traits (mortality and growth) on the intertidal barnacles, Chthamalus montagui and Semibalanus balanoides, on the mid-shore in Plymouth, UK. Intra- and interspecific treatments comprising of C. montagui and S. balanoides were manipulated to reduce densities at two similar sites. Changes in mortality and operculum growth were assessed over an 8-week period using digital photography. Covariates of growth included nearest neighbour distance, competition between closest pairs and initial size. Conflicting patterns were observed when comparing growth rates between treatments and sites. At Site 1, interspecific treatments had a lower growth rate than intraspecific treatments, whereas at Site 2, interspecific growth rates were higher. ANCOVA showed that nearest neighbour distance had no significant effect on growth, but when comparing differences in growth of closest neighbouring pairs, C. montagui treatment showed evidence of competition whereas S. balanoides did not. ANCOVA analysis indicated no difference in growth between each outcome of pair competition, suggesting winners are initially bigger than losers. Comparisons of mortality between treatments indicated mortality over time with no significant differences observed between treatments, but response surface methodology (RSM) revealed no effects of competition on mortality of S. balanoides, but negative effects of both intra- and interspecific competition on C. montagui survivorship. Examination of natural populations of barnacles in the mid-shore indicated there was strong spatial variation in growth rates, perhaps driven by small-scale differences within sites.

In species with complex life cycles, laboratory studies have shown that variations in the traits of settling larvae can affect post-settlement survival and influence recruitment and benthic− pelagic coupling. However, we still know little about the magnitude and spatial scale of natural trait variation. We studied spatial variation in body size and nutritional reserves (carbon, nitrogen and lipids) of settled cyprids of the barnacle Semibalanus balanoides along the coast of West Scotland. We quantified variation among regions (north vs. south: range ~700 km), locations (~50 km), shores (~10 km) and within shores (~10 m). We also evaluated trait responses to gradients in chlorophyll and shore openness and compared swimming vs. settled cyprids in order to infer the likely influence of costs of substratum search on trait variation. Variability between regions was large, with higher trait values (e.g. carbon cyprid−1: 35 to 50% higher) in the north. Most traits correlated negatively with pelagic chlorophyll a (a proxy for larval/juvenile food availability); this counter-gradient pattern suggests an adaptive role of increased reserves, buffering benthic juveniles from low food availability during the critical early post-settlement period. Body size and nitrogen content correlated positively with shore openness; lower than expected carbon content suggest increased costs of substratum search on open shorelines. Higher nitrogen content but lower percent carbon was found in settled vs. swimming larvae, suggesting costs of sub - stratum search at the time of settlement. Overall, we uncovered the spatial scales at which trait variation, shaped by pelagic processes, can affect post-metamorphic survival, recruitment and benthic−pelagic coupling.

Marine benthic populations are dependent on early life-history stages surviving multiple population bottlenecks. Failure at one or several of these bottlenecks can alter species’ patterns of distribution and abundance. The barnacleSemibalanus balanoidesis found along temperate and sub-arctic shorelines of the Atlantic and Pacific Oceans. Over the past century the southern range limits ofS. balanoideshave shifted hundreds of kilometres poleward on both coasts of the Atlantic. Here we tested if temperature limits fertilization and used these data, along with those from previous studies, to create mechanistic biogeographic models to understand which potential population bottlenecks in the early life-history ofS. balanoidesinfluence its distribution and abundance. In the western Atlantic survival of new recruits is probably more important in setting the southern range limit than the effects of temperature on early life-history stages because fertilization, brooding and the probability of larval release matching phytoplankton availability were all predicted to be high near the historical range edge. Phytoplankton mismatch may partially explain the ephemeral nature ofS. balanoidesin some parts of the English Channel. Further south along the coast of France predicted brooding success was reduced in a pattern consistent with historical range shifts in this region. Within Galicia, Spain fertilization was predicted to be low near the southern limit, and likely plays an important role in setting this range edge. Mismatches between phytoplankton abundance and larval release in Galicia may further limit reproductive success within this region.

Barnacle recruitment is often studied in rocky intertidal habitats due to the relevant role that barnacles can play in intertidal communities. In 2014, barnacle (Semibalanus balanoides) recruitment was measured at high elevations in wave-exposed intertidal habitats on the NW Atlantic coast in Nova Scotia, Canada. Values were considerably lower than previously reported for middle elevations in wave-exposed intertidal habitats on the NE Atlantic and NE Pacific coasts. To determine if such differences in recruitment may have resulted from elevation influences, I did a field experiment in 2019 in wave-exposed intertidal habitats in Nova Scotia to test the hypothesis that recruitment is higher at middle than at high elevations, based on known environmental differences between both elevation zones. Based on data from three locations spanning 158 km of the Nova Scotia coast, barnacle recruitment was, on average, nearly 200 % higher (and recruits were larger) at middle than at high elevations. However, even with this increase, barnacle recruitment on this NW Atlantic coast is still lower than for comparable habitats on the NE Atlantic and NE Pacific coasts, and also lower than previously reported for wave-exposed locations farther south on the NW Atlantic coast, in Maine, USA. Therefore, barnacle recruitment in wave-exposed intertidal environments in Nova Scotia appears to be only moderate relative to other shores. This difference in the supply of barnacle recruits might influence the intensity of interspecific interactions involving barnacles.

Correlative imaging combines information from multiple modalities (physical–chemical–mechanical properties) at various length scales (centimetre to nanometre) to understand the complex biological materials across dimensions (2D–3D). Here, we have used numerous coupled systems: X-ray microscopy (XRM), scanning electron microscopy (SEM), electron backscatter diffraction (EBSD), optical light microscopy (LM) and focused ion beam (FIB-SEM) microscopy to ascertain the microstructural and crystallographic properties of the wall-plate joints in the barnacle Semibalanus balanoides . The exoskeleton is composed of six interlocking wall plates, and the interlocks between neighbouring plates (alae) allow barnacles to expand and grow while remaining sealed and structurally strong. Our results indicate that the ala contain functionally graded orientations and microstructures in their crystallography, which has implications for naturally functioning microstructures, potential natural strengthening and preferred oriented biomineralization. Elongated grains at the outer edge of the ala are oriented perpendicularly to the contact surface, and the c -axis rotates with the radius of the ala. Additionally, we identify for the first time three-dimensional nanoscale ala pore networks revealing that the pores are only visible at the tip of the ala and that pore thickening occurs on the inside (soft bodied) edge of the plates. The pore networks appear to have the same orientation as the oriented crystallography, and we deduce that the pore networks are probably organic channels and pockets, which are involved with the biomineralization process. Understanding these multiscale features contributes towards an understanding of the structural architecture in barnacles, but also their consideration for bioinspiration of human-made materials. The work demonstrates that correlative methods spanning different length scales, dimensions and modes enable the extension of the structure–property relationships in materials to form and function of organisms.

Barnacles are conspicuous members of rocky intertidal communities and settlement of the final larval stage, the cyprid, is influenced by the presence of biofilms. While modulation of cyprid settlement by biofilms has been studied extensively, the acquisition of a specific microbiome by the settling larva has not. This study investigated settlement in the field of Semibalanus balanoides in two consecutive years when the composition of the benthic bacterial community differed. In both years, settling cyprids adopted a specific sub-set of benthic bacteria that was distinct from the planktonic cyprid and the benthos. This microbiome was consistent, regardless of annual variability in the benthic community structure, and established within hours of settlement. The results imply that a natural process of selection occurs during the critical final transition of S. balanoides to the sessile form. The apparent consistency of this process between years suggests that optimal growth and survival of barnacles could depend upon a complex inter-kingdom relationship, as has been demonstrated in other animal systems.

Benthic–pelagic coupling refers to the ecological relationships between benthic and pelagic environments. Studying such links is particularly useful to understand biological variation in intertidal organisms along marine coasts. Filter-feeding invertebrates are ecologically important on marine rocky shores, so they have often been used to investigate benthic–pelagic coupling. Most studies, however, have been conducted on eastern ocean boundaries. To evaluate benthic–pelagic coupling on a western ocean boundary, we conducted a 5-year study spanning 415 km of the Atlantic coast of Nova Scotia (Canada). We hypothesized that the summer size of intertidal barnacles (Semibalanus balanoides) recruited in the preceding spring would be positively related to the nearshore abundance (biomass) of phytoplankton, as phytoplankton constitutes food for the nauplius larvae and benthic stages of barnacles. Every year between 2014 and 2018, we measured summer barnacle size in clearings created before spring recruitment on the rocky substrate at eight wave-exposed locations along this coast. We then examined the annual relationships between barnacle size and chlorophyll-a concentration (Chl-a), a proxy for phytoplankton biomass. For every year and location, we used satellite data to calculate Chl-a averages for a period ranging from the early spring (when most barnacle larvae were in the water) to the summer (when barnacle size was measured after weeks of growth following spring benthic recruitment). The relationships were always positive, Chl-a explaining nearly half, or more, of the variation in barnacle size in four of the five studied years. These are remarkable results because they were based on a relatively limited number of locations (which often curtails statistical power) and point to the relevance of pelagic food supply to explain variation in intertidal barnacle size along this western ocean boundary coast.

The abundant-centre hypothesis (ACH) predicts that the density of a species should peak at its distribution centre and decrease similarly towards distribution margins. The ACH has been deduced from a theory that postulates that environmental conditions should be most favourable for a species at the centre of its distribution. This idealised density pattern, however, has been supported by limited field studies, as natural patterns are often more complex. It is thus of interest to examine under what conditions compliance with the ACH could be favoured. Such conditions could be smooth environmental gradients with limited habitat patchiness throughout the distribution range of a species. Thus, we tested the ACH by measuring the density of an intertidal barnacle (Semibalanus balanoides) across its full vertical distribution range (from low to high elevations) on a rocky shore with similar substrate properties across elevations. To do a reliable test, we surveyed eight elevation zones applying an equal sampling effort in each zone. Average barnacle density conformed to the ACH, as it peaked at the middle of the vertical distribution range of this species. The same underlying theory predicts a similar unimodal pattern for maximum body size, but this trait was decoupled from density, as maximum barnacle size increased from low to high elevations. Overall, although the ACH is not a universal predictive tool as once envisioned, it may predict some cases well, as shown by this study. Therefore, the ACH should not be discarded completely, and its domain of application should be further evaluated.

1.AbstractCorrelative imaging combines information from multiple modalities (physical-chemical-mechanical properties) at various length-scales (cm to nm) to understand complex biological materials across dimensions (2D-3D). Here, we have used numerous coupled systems: X-ray microscopy (XRM), scanning electron microscopy (SEM), electron backscatter diffraction (EBSD), optical light microscopy (LM), and focused-ion beam (FIB-SEM) microscopy to ascertain the microstructural and crystallographic properties of the wall-plate joints in the barnacle Semibalanus balanoides. The exoskeleton is composed of six interlocking wall-plates, and the interlocks between neighbouring plates (alae) allow barnacles to expand and grow whilst remaining sealed and structurally strong. Our results indicate that the ala contain functionally-graded orientations and microstructures in their crystallography, which has implications for naturally functioning microstructures, potential natural strengthening, and preferred oriented biomineralisation. Elongated grains at the outer edge of the ala are oriented perpendicularly to the contact surface, and the c-axis rotates with the radius of the ala. Additionally, we identify for the first time three-dimensional nano-scale ala pore networks revealing that the pores are only visible at the tip of the ala, and that pore thickening occurs on the inside (soft-bodied) edge of the plates. The pore networks appear to have the same orientation as the oriented crystallography, and we deduce that the pore networks are probably organic channels and pockets which are involved with the biomineralisation process. Understanding these multi-scale features contributes towards an understanding of the structural architecture in barnacles, but also their consideration for bioinspiration of human-made materials. The work demonstrates that correlative methods spanning different length-scales, dimensions and modes enable the extension of structure-property relationships in materials to form and function of organisms.

Biofouling by marine organisms can result in a variety of negative environmental and economic consequences, with decontamination procedures remaining problematic, costly and labour-intensive. Here, we examined the efficacy of direct steam exposure to induce mortality of selected biofouling species: Mytilus edulis; Magallana gigas; Semibalanus balanoides; Fucus vesiculosus; and an Ulva sp. Total mortality occurred at 60-sec of steam exposure for M. edulis and juvenile M. gigas, at 30-sec for S. balanoides, while 300-sec was required for adult M. gigas. Application of steam reduced the biomass of F. vesiculosus and significantly reduced Ulva sp. biomass, with complete degradation being observed for Ulva sp. following 120-sec of exposure. Accordingly, it appears that steam exposure can cause mortality of biofouling organisms through thermal shock. Although preliminary, our novel and promising results suggest that steam applications could potentially be used to decontaminate niche areas and equipment.

This study used morphometric analyses to compare the structure of the third antennular segment, also called the attachment organ, in cyprid larvae from cirripede species representing a diverse set of taxonomic groups. The aim was to investigate the degree of morphological variation in view of the diversity of habitats, settlement substrata, and modes of life found in the Cirripedia. In all cyprids the third segment features a flat surface (the attachment disc) covered with small cuticular villi thought to function in adhesion. The parameters analyzed were the angle of this disc relative to the long axis of the antennule, its shape (outline), the density of cuticular villi, and the type of cuticular structure encircling the disc. The 10 species studied came from most major groups of cirripedes, and comprised shallow-water forms inhabiting hard bottoms (Capitulum mitella, Pollicipes pollicipes, Semibalanus balanoides, Austrominius modestus, Megabalanus rosa), sublittoral forms (Verruca stroemia, Scalpellum scalpellum), epibiotic forms settling on live, soft tissues (Balanus spongicola, Savignium crenatum), and a parasite (Peltogaster paguri). Significant structural variation was found among the species, but due to limited taxon sampling it was unclear whether the differences relate to ecological factors or phylogenetic affiliation. The disc perimeter is guarded by either a series of long and thin cuticular fringes overreaching the rim of the disc (= a velum) or a few low, but very broad cuticular flaps (= a skirt). The presence of a velum (in all rocky-shore species) or a skirt (all other species) around the attachment disc was the only parameter that was clearly correlated with habitat. The shape of the third antennular segment varied from a symmetrical bell shape with a distally facing attachment disc having a circular disc outline, to segments that were elongated in side view, with a very tilted ventral disc surface having an elliptical disc outline. The bell shape may be most common in forms from rocky shores, but in our test of morphometric parameters only Scalpellum scalpellum (sublittoral), Savignium crenatum (epibiotic in corals), and Peltogaster paguri (parasitic) had shapes that differed significantly from the other species. The density of villi on the attachment disc varied significantly, but also showed no clear-cut correlation with substratum or habitat. Attachment organ structure is clearly the most variable feature in cirripede cyprids. To evaluate the degree to which attachment organ structure is correlated with habitat, settlement substratum, and mode of life, future studies should employ a more refined statistical analysis on an enlarged dataset, with much increased taxon sampling and a more multifaceted definition of ecological variables.

The saccate Asiatic brown seaweed Colpomenia peregrina was first recorded within the NW Atlantic in 1960 at Atkins Point, Nova Scotia. By 2014 it was recorded from 57 sites ranging from the Grand Barachois Lagoon, Newfoundland to South Wellfleet, MA. We first found extensive populations in New England at Fort Foster, Kittery, Maine where we subsequently conducted floristic and ecological studies of the alga. Colpomenia peregrina is superficially similar to another saccate brown alga, Leathesia marina, although they have differences in morphology, anatomy, and reproduction. The species is a common epiphyte on at least 33 seaweeds within the NW Atlantic, with Corallina officinalis being its most common host. It also grows on the seagrass Zostera marina and on at least two common invertebrates, Mytilus edulis and Semibalanus balanoides. Colpomenia peregrina is an annual somewhat like Petalonia fascia, with some specimens bearing plurilocular gametangia year-round. Based upon collection of samples fr...

The risk of predation can drive trophic cascades by causing prey to engage in antipredator behavior (e.g. reduced feeding), but these behaviors can be energetically costly for prey. The effects of predation risk on prey (nonconsumptive effects, NCEs) and emergent indirect effects on basal resources should therefore depend on the ecological context (e.g. resource abundance, prey state) in which prey manage growth/predation risk tradeoffs. Despite an abundance of behavioral research and theory examining state-dependent responses to risk, there is a lack of empirical data on state-dependent NCEs and their impact on community-level processes. We used a rocky intertidal food chain to test model predictions for how resources levels and prey state (age/size) shape the magnitude of NCEs. Risk cues from predatory crabs Carcinus maenas caused juvenile and sub-adult snails Nucella lapillus to increase their use of refuge habitats and decrease their growth and per capita foraging rates on barnacles Semibalanus balanoides. Increasing resource levels (high barnacle density) and prey state (sub-adults) enhanced the strength of NCEs. Our results support predictions that NCEs will be stronger in resource-rich systems that enhance prey state and suggest that the demographic composition of prey populations will influence the role of NCEs in trophic cascades. Contrary to theory, however, we found that resources and prey state had little to no effect on snails in the presence of predation risk. Rather, increases in NCE strength arose because of the strong positive effects of resources and prey state on prey foraging rates in the absence of risk. Hence, a common approach to estimating NCE strength – integrating measurements of prey traits with and without predation risk into a single metric – may mask the underlying mechanisms driving variation in the strength and relative importance of NCEs in ecological communities.

Spatial patterns in aggregations form as a result of the interplay between costs and benefits experienced by individuals. Such self-organisation of aggregations can be explained using a zonal model in which a short-range zone of repulsion and longer-range zone of attraction surrounding individuals leads to emergent pattern properties. The signal of these processes can be detected using spatial pattern analyses. Furthermore, in sessile organisms, post-settlement mortality reveals the relative costs and benefits of positions within the aggregation. Acorn barnacles are known to require contact with conspecifics for reproduction and are therefore believed to aggregate for this purpose; isolated individuals may also be more susceptible to abiotic stress and predation. At short distances, however, competition for space and resources is likely to occur. In this study spatial patterns of barnacles (Semibalanus balanoides L.) were analysed using pair-correlation functions. Individuals were dispersed at distances below 0.30 cm, but peak relative density occurred at a distance of 0.36 cm from conspecifics. This is much closer than required for reproductive access, implying a strong aggregative drive, up to the point of physical contact with neighbours. Nevertheless, analysis of dead barnacles illustrated that such proximity carries a cost as barnacles with many neighbours were more likely to have died. The inferences obtained from these patterns are that barnacles aggregate as closely as they can, and that local neighbourhood competition is a powerful determinant of mortality. These processes give rise to the observed pattern properties.

Introduced bioengineering organisms may fundamentally change native coastal ecosystems by modifying existing benthic habitat structures and thereby habitat-specific species interactions. The introduction of the Pacific oyster Magallana gigas into the sedimentary coastal area of the south-eastern North Sea and its preferred settlement on native blue mussel shells caused a large-scale shift from monospecific Mytilus edulis beds to current mixed reefs of mussels and oysters. To investigate whether the newly developed biotic habitat affects the occurrence of associated native key organisms and their ecological functions, we studied the long-term density trajectory of the gastropod Littorina littorea and its grazing activity on barnacles attached to Pacific oyster reefs in the northern Wadden Sea. We found no significant correlation between oyster and snail densities on blue mussel beds in the last two decades, which spans a time-period from the beginning of Pacific oyster establishment to today's oyster dominance. A manipulative field experiment revealed that snail density significantly affects the recruitment success of barnacles Semibalanus balanoides on oyster shells with the highest number of barnacle recruits at snail exclusion. Thus, density and grazing activity of the snail L. littorea may control barnacle population dynamics on epibenthic bivalve beds in the Wadden Sea. This interspecific interaction was already known for blue mussel beds before the oyster invasion and, therefore, we conclude that despite the strong modifications that non-native ecosystem engineers cause in native biotic habitats, the ecological functions of associated key species can remain unchanged.

Barnacle recruitment is often studied in rocky intertidal habitats due to the relevant role that barnacles can play in intertidal communities. In 2014, recruitment of the barnacle Semibalanus balanoides was measured at high elevations in wave-exposed intertidal habitats on the NW Atlantic coast in Nova Scotia, Canada. Values were considerably lower than previously reported for middle elevations in wave-exposed intertidal habitats on the NE Atlantic and NE Pacific coasts. To determine if such differences in recruitment may have resulted from elevation influences, I did a field experiment in 2019 in wave-exposed intertidal habitats in Nova Scotia to test the hypothesis that recruitment is higher at middle than at high elevations, based on known environmental differences between both elevation zones. Based on data from 3 locations spanning 158 km of the Atlantic coast of Nova Scotia, barnacle recruitment was, on average, nearly 200% higher (and recruits were larger) at middle than at high elevations. However, even with this increase, barnacle recruitment on this NW Atlantic coast is still lower than for comparable habitats on the NE Atlantic and NE Pacific coasts, and also lower than previously reported for wave-exposed locations farther south on the NW Atlantic coast, in Maine, USA. Therefore, barnacle recruitment in wave-exposed intertidal habitats on the Atlantic coast of Nova Scotia is moderate relative to other shores. This difference in the supply of barnacle recruits might influence the intensity of interspecific interactions involving barnacles.

The ability to survive freezing has repeatedly evolved across multiple phyla. This suggests that the mechanisms of freeze tolerance must be readily evolvable from basal physiological traits. While several biochemical correlates to freeze tolerance have been described, the mechanism that confers freeze tolerance is still not well understood. To understand both the basic biochemical mechanisms of freeze tolerance as well as their role in local adaptation at the poleward range edge, we conducted a transcriptomic study on two populations (one from the poleward range margin in the White Sea, Russia, and one from the central coast of British Columbia, Canada) of the cold water acorn barnacleSemibalanus balanoideson a time series following a freezing event. We found that the British Columbia population (at the equatorward range margin) was significantly less freeze tolerant than the White Sea population (at the poleward range margin). After assembling and annotating ade novotranscriptome forS. balanoides, we found that the patterns of differential transcript expression following freezing were almost entirely non-overlapping between the two populations, with the White Sea population expressing a series of heat shock proteins in response to freezing stress as well as several aquaporins, while the British Columbia population expressed a series of proteases instead, indicating severe protein damage. We found strong evidence of purifying selection on the significantly upregulated transcripts in the White Sea population, suggesting local adaptation to freezing threat. Taken together, this shows the importance of freeze tolerance to population survival at the poleward range margin, and highlights the central roles of aquaporins and heat shock proteins to the trait of freeze tolerance across taxa.

Ice can be an important structuring factor physically removing intertidal flora and fauna. At high latitudes in particular, the removal of canopy-forming algae by ice scour may be important as their canopy may serve to modify the extreme environment for marine organisms at low tide. We simulated the effect of ice scouring by manipulating the biomass of the canopy-forming algae Ascophyllum nodosum in a sub-Arctic fjord [“Full canopy,” “Reduced canopy,” “Bare (start),” “Bare (annual)”]. Over a three-year period, we quantified key physical parameters and the recolonization of flora and fauna to test the hypothesis that A. nodosum and rock rugosity facilitate recolonization of sub-Arctic intertidal fauna and that potential facilitation could rely on an ability of A. nodosum canopy to modify air temperature and ice scour. Finally, we estimated the recovery period of A. nodosum canopy height to pre-disturbance levels based on estimated early growth rates. We found that A. nodosum canopy facilitated higher species richness and recolonization of dominating faunal species (Littorina saxatilis, Littorina obtusata, Mytilus edulis, and Semibalanus balanoides), and also significantly reduced the high temperatures in summer and raised the low temperatures in winter. The abundance of M. edulis and A. nodosum recolonization increased significantly with rock rugosity and the recovery of A. nodosum canopy height was estimated to a minimum of 15 years. We conclude that algal canopy and rock rugosity play key roles in structuring sub-Arctic intertidal communities, likely by modifying environmental stress such as extreme temperature, desiccation, and by increasing the settling surface and the habitat complexity. As the distribution of canopy-forming algae is expected to shift northward, they may act as a key habitat facilitating a northward colonization of intertidal fauna in the Arctic. We highlight the importance of considering scales relevant to biological communities when predicting impacts of climate change on distributional patterns and community structure in the Arctic intertidal.

Range shifts due to annual variation in temperature are more tractable than range shifts linked to decadal to century long temperature changes due to climate change, providing natural experiments to determine the mechanisms responsible for driving long-term distributional shifts. In this study we couple physiologically grounded mechanistic models with biogeographic surveys in 2 years with high levels of annual temperature variation to disentangle the drivers of a historical range shift driven by climate change. The distribution of the barnacle Semibalanus balanoides has shifted 350 km poleward in the past half century along the east coast of the United States. Recruits were present throughout the historical range following the 2015 reproductive season, when temperatures were similar to those in the past century, and absent following the 2016 reproductive season when temperatures were warmer than they have been since 1870, the earliest date for temperature records. Our dispersal dependent mechanistic models of reproductive success were highly accurate and predicted patterns of reproduction success documented in field surveys throughout the historical range in 2015 and 2016. Our mechanistic models of reproductive success not only predicted recruitment dynamics near the range edge but also predicted interior range fragmentation in a number of years between 1870 and 2016. All recruits monitored within the historical range following the 2015 colonization died before 2016 suggesting juvenile survival was likely the primary driver of the historical range retraction. However, if 2016 is indicative of future temperatures mechanisms of range limitation will shift and reproductive failure will lead to further range retraction in the future. Mechanistic models are necessary for accurately predicting the effects of climate change on ranges of species.

Increasing glacial discharge may influence future recruitment patterns and growth rates of marine benthos in glacial fjords as changes in surface-water temperature and salinity as well as an increase in ice-scouring events. We deployed settling plates at three spatial and temporal scales in a sub-Arctic glacial fjord system to quantify spatial and temporal variation in the recruitment rates and early growth of subtidal marine benthos, and to determine potential drivers of recruitment. We found significant spatial variation in recruitment of benthos (flora and fauna) at both the small (20 m) and large (> 30 km) scale, indicating that both abiotic and biotic factors may be important. We observed that substrate modification by bushy macroalgae facilitated Mytilus spp. settlement, yet limited Semibalanus balanoides recruitment, underlining that timing in settlement of different species is important in structuring benthic communities. Spatial variation in physical parameters, such as temperature and salinity, likely affected growth and recruitment patterns of benthos (S. balanoides). Finally, we found large spatial and temporal variation in kelp recruitment varying from none to 5866 ind. M−2, with an annual production of more than 6000 g dw m−2 year−1. This large growth potential of kelp suggests recruitment as a bottleneck in kelp population dynamics. Conclusively, variation in sub-Arctic benthic recruitment and growth patterns may respond to changes in temperature and salinity, yet further effort is needed to elucidate potential drivers of benthic recruitment in a fast changing Arctic environment.

The northern acorn barnacle (Semibalanus balanoides) is a robust system for the study of evolutionary processes in the intertidal. S. balanoides has a well-characterized ecology, a wide circumboreal distribution, and a life history characterized by tractable environmental stressors at various ecological scales. In this chapter, we discuss a variety of topics concerning the development of S. balanoides as a model in ecological genomics as well as inferences of demography and historical phylogeography. In addition, we introduce two novel genomic tools for S. balanoides: the complete mtDNA sequence and the second draft of the nuclear genome (Sbal2). Using these tools, we conducted a reanalysis of previously described mtDNA haplotypes, a and b, as well as genome-wide levels of variation and population structure across the North Atlantic using pooled sequencing approaches. Analyses of sequence data from older and more recent Illumina platforms revealed the effects of technical bias in the estimates of population genomic metrics. We found concordant levels of nuDNA and mtDNA genetic variation with no evidence of demographic bottlenecks. We observed low genome-wide FST values across the Atlantic, suggesting a large number of ancestral polymorphisms and shared standing variation across the basin. Comparisons of genome-wide estimates of FST with those derived from a discriminant analysis of principal components uncovered population-structure-informative SNPs. This suggests the existence of latent population structure across broad scales, despite the capacity for extensive planktonic dispersal. Noticeably, our samples collected in Iceland displayed higher similarity to North American populations than to the rest of Europe. We hypothesize this is consistent with a periglacial refugium in Iceland concomitant with a barrier to gene flow caused by the North Atlantic current. Lastly, we discuss challenges and opportunities for the improvement of genomic tools in barnacles. Our reflections in this area are easily generalizable to most natural populations.

A new method to autonomously collect larvae and environmental variables such as temperature, salinity, and circulation is described. A large volume in situ pumping system, recently developed for discrete biogeochemical sample collection in deep-sea environments, was adapted and mounted to the autonomous underwater vehicle (AUV) REMUS 600 for autonomous larval and environmental sampling in coastal waters. To assess the performance of this system, the distribution of barnacle larvae was assessed in March 2014 with two transects perpendicular to the coastline (~ 9.9 and 11.2 km) in Buzzards Bay, Massachusetts, USA. The second transect included a complex sampling mission through a relatively deeper channel, and sampling at discrete depth intervals. In this deployment, nearshore and surface waters were fresher, with distinct vertical stratification due to salinity. Barnacle larvae were classified into early nauplii, late nauplii, and cyprid stages, and the mitochondrial COI barcode marker was sequenced to identify individual larvae of different stages. In an analysis of 164 barcode sequences, larvae belonging to Amphibalanus sp., Semibalanus balanoides, and Chthamalus fragilis were found, with Amphibalanus sp. the most abundant larval taxon overall. In the second deployment, early and late nauplii were relatively more abundant near the bottom. However, there was no obvious pattern relative to depth with cyprids, and there were no clear cross-shore distributional patterns for nauplii and cyprids. While additional deployments are necessary to corroborate these observations, the results demonstrate the feasibility of this approach. Autonomous vehicle based sampling has the potential to collect larvae of other invertebrates as well as zooplankton, and together with genetic identifications, overcomes many existing limitations and will provide valuable new insight in understanding larval distributions and transport dynamics.

A new species of tardigrade in the family Echiniscoididae, Echiniscoides wyethi, was found on the barnacle Semibalanus balanoides (L.) at several intertidal sites around Allen Island, Maine, USA. The new species is named after the American artist Andrew Wyeth and the Wyeth family who own the island on which this species was discovered. The new species is separated from all other Echiniscoides by the smooth cuticle, long flexible buccal tube, and the number of claws on the legs of the adult.

The invasive barnacle Austrominius (= Elminius) modestus has been present in Europe since the 1940s, and has recently been recorded to outnumber native barnacle species at some locations, including an Irish marine nature reserve. It has been suggested that these increases in abundance following a lag phase since establishment, represent the awakening of an ‘ecological sleeper’, due to changes in environmental conditions. Austrominius modestus was first recorded on the Isle of Cumbrae, Scotland in 1955, and was reported to be well established on the island by 2007. Since this location is close to the northern limit of the invaded range of A. modestus , it has been chosen as a site for the long term monitoring of this species. Quantitative and semi-quantitative surveys of the abundance of A. modestus , together with the native barnacle species Chthamalus montagui and Semibalanus balanoides , have been made on the island on a biannual basis since 2009. This study examined changes in the abundances of these three species from 2009 to 2013, and reports on the present status of this invasive species on the island. Austrominius modestus was found at all sites surveyed, but did not outnumber native barnacle species at any site. Semibalanus balanoides , a cold water boreal species, was the most abundant barnacle species at most sites from 2009 to 2013. All three barnacle species underwent a decline in 2011, but had increased in abundance by 2013. Despite undergoing the smallest increase in abundance between 2011 and 2013, S. balanoides remains the dominant barnacle species on the Isle of Cumbrae. Nearly sixty years after its initial discovery on the island, A. modestus is widespread, however it has not outnumbered native species, which continue to recruit at high densities. This competitive pressure makes it unlikely that A. modestus will outnumber native barnacle species at this location, close to its northern limit, in the near future.

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.

Recruitment is a key demographic process for population persistence. This paper focuses on barnacle (Semibalanus balanoides) recruitment. In rocky intertidal habitats from the Gulf of St. Lawrence coast of Nova Scotia (Canada), ice scour is common during the winter. At the onset of intertidal barnacle recruitment in early May (after sea ice has fully melted), mostly only adult barnacles and bare substrate are visible at high elevations in wave-exposed habitats. We conducted a multiannual study to investigate if small-scale barnacle recruitment could be predicted from the density of pre-existing adult barnacles. In a year that exhibited a wide adult density range (ca. 0–130 individuals dm−2), the relationship between adult density and recruit density (scaled to the available area for recruitment, which excluded adult barnacles) was unimodal. In years that exhibited a lower adult density range (ca. 0–40/50 individuals dm−2), the relationship between adult and recruit density was positive and resembled the lower half of the unimodal relationship. Overall, adult barnacle density was able to explain 26–40% of the observed variation in recruit density. The unimodal adult–recruit relationship is consistent with previously documented intraspecific interactions. Between low and intermediate adult densities, the positive nature of the relationship relates to the previously documented fact that settlement-seeking larvae are chemically and visually attracted to adults, which might be important for local population persistence. Between intermediate and high adult densities, where population persistence may be less compromised and the abundant adults may limit recruit growth and survival, the negative nature of the relationship suggests that adult barnacles at increasingly high densities stimulate larvae to settle elsewhere. The unimodal pattern may be especially common on shores with moderate rates of larval supply to the shore, because high rates of larval supply may swamp the coast with settlers, decoupling recruit density from local adult abundance.

To explore how climate may affect the structure of natural communities, we quantified the role of thermal stress in setting the high intertidal borders of the acorn barnacle, Semibalanus balanoides. At sites north and south of Cape Cod, a major faunal and thermal boundary on the east coast of North America, we examined the interacting effects of thermal stress and recruit density on individual survivorship. At hotter southern sites, particularly in bays, high intertidal barnacle survivorship was enhanced by experimental shading or by neighbors which ameliorate heat and desiccation stresses. In contrast, at cooler northern bay and coastal sites, neither shading nor group benefits increased barnacle survival, and mortality patterns were driven primarily by predators with largely boreal distributions. Our field results, like recent laboratory microcosm studies, suggest that predicting even simple community responses to climate change may be more complex than is currently appreciated.

Experimental tests of the hypothesis that ecological communities can exist in “multiple stable states” are rare, and some argue, impossible, because of the unlikelihood that any system will meet the necessary criteria. These are that alternative states (1) are in the same location, (2) experience the same environment, (3) persist for multiple generations, (4) resist repeated perturbations, and (5) result from a pulse manipulation. In 1974, we initiated an experiment testing the ability of Ascophyllum nodosum-dominated rocky intertidal communities to recover from complete canopy removal. Manipulations were monitored frequently for 5 yr after clearance and resurveyed again after a 35-yr hiatus. After clearance, Ascophyllum was replaced immediately by another fucoid alga, Fucus spp, which continued to dominate the space through 1979 despite regular annual recruitment by Ascophyllum. Observations in 2009 revealed that Fucus spp. still dominated the cleared plots. Surveys in 2011 and 2013 demonstrated that Fucus persisted in experimental plots and Ascophyllum persisted in adjacent, unmanipulated plots. All criteria for testing an alternative state were met. Fucus persisted through multiple generations of both fucoids, a steady annual rain of recruits of both species, a high frequency of storm-driven perturbations, and it resulted from a pulse manipulation. Likely mechanisms include poor Ascophyllum recruitment directly under the Fucus spp. canopy despite abundant recruitment in adjacent areas, self-maintenance by Fucus spp. through high recruitment and fast growth, and recruitment facilitation of Fucus by the barnacle Semibalanus balanoides. Several lines of evidence indicate that other possible mechanisms including indirect facilitation of fucoids through predation on competitors (mussels), and positive or negative effects of littorine grazing, are unlikely. Although prior results in Maine suggested that the mussel Mytilus edulis was an alternative stable state, the new results suggest that Fucus spp. was the alternative state to Ascophyllum. During the nearly four decades of this experiment, a number of important species including fucoids, littorine grazers, and mussels all declined in abundance, most likely due to climate change. The presumed impact of climate change makes prediction of the long-term response of this system difficult, but it already differs dramatically from its structure in the 1970s.

Studies identifying interactions between biological invasions and other stressors have generally focussed on quantifying their cumulative effects on mature species assemblages. In benthic systems, however, early life history processes are key determinants of assemblage structure and functioning. This study tested whether the presence of an invasive species affected early life history processes of two common barnacle species and whether this was affected by a second common stressor, nutrient enrichment. The results of a field experiment identified and characterised the effects of an invasive oyster, Crassostrea gigas, on the early life history processes of the two barnacle species under ambient and enriched nutrient conditions. In the presence C. gigas, the invasive barnacle Austrominius modestus, had a lower recruitment rate, however, there was no effect of the presence of C. gigas on native barnacle, Semibalanus balanoides, recruitment. Nutrient enrichment also reduced the recruitment rate of A. modestus, however, there was no evidence of synergistic or antagonistic interactions between these stressors, indicating their cumulative effects were additive. There was no effect of nutrient enrichment on native barnacle recruitment. Our results show that the presence of an invasive oyster and nutrient enrichment altered the recruitment of another non-native benthic species. These findings emphasise the importance of considering early life history processes when assessing effects of multiple stressors on communities.

Geographic distributions of species are frequently influenced by temperature and as a result, there is considerable interest in predicting the effect of climate change on biogeography. Previous studies have found strong correlations between sea surface temperature and recruitment in the acorn barnacle Semibalanus balanoides, an important member of temperate intertidal communities in the Northern Hemisphere. The underlying mechanism that drives this relationship is, however, unknown. The hypothesis tested here is that variation in recruitment is determined by temperature-induced variation in survival of brooded embryos. Brooding adults were held under several environmentally-relevant temperature conditions from 7 °C to 13 °C until eggs were ready to hatch. There was no significant effect of temperature on embryonic development rate. Brooding individuals held at colder temperatures had more than twice the reproductive mass of those held at warmer temperatures. This mass difference is caused by an over three-fold larger number of embryos surviving in the coldest treatment (7 °C) compared to the warmest treatment (13 °C). Temperature-induced variation in number of surviving embryos likely contributes to differences in recruitment following cold winters vs. warm winters. Such temperature-dependent variation in recruitment has been shown to cause year to year fluctuations in the positions of the geographic limits of the species in Southwest England, France, and Spain. This study illustrates the key role of adult input into the larval pool in generating variation in recruitment and its influence on biogeography.

Few links have been established between the Atlantic Multidecadal Oscillation and long-term dynamics of marine systems due to the scarcity of sustained biological time-series with sufficient multi-decadal coverage. The abundances of co-occurring boreal and Lusitanian species of barnacle have been recorded annually at a rocky shore in Devon, southwest England since 1953. Multidecadal cycles in relative abundances of the cold-water Semibalanus balanoides and warm-water Chthamalus spp. are strongly correlated with both local sea surface temperatures, and a ‘Warm Index’ of barnacle abundance shows strong links to the basin-scale Atlantic Multidecadal Oscillation. In contrast there are weak or no observed relationships with the North Atlantic Oscillation for either species. The shorter lifecycle of S. balanoides compared to the chthamalids and the increase in spring and summer temperatures to which newly settled S. balanoides recruits have been exposed during the last decade are likely mechanisms by which barnacle densities are responding to low-frequency temperature variability expressed in the Atlantic Multidecadal Oscillation.

It has increasingly been recognized that organisms can interfere with parasitic free-living stages, preventing them from infecting their specified host and thus reducing infection levels. This common phenomenon in freshwater and terrestrial systems has been termed the ‘dilution effect’ and, so far, is poorly studied in marine systems. Ten common intertidal organisms found in the Dutch Wadden Sea (North Sea) were tested to establish their effects on the free-living cercarial stages of the trematode parasite Himasthla elongata. Most species tested resulted in a significant reduction in cercariae over a 3 hr time period. The amphipod Gammarus marinus removed 100% of the cercariae, while other effective diluters were Crangon crangon (93%), Sargassum muticum (87%), Semibalanus balanoides (71%), Crassostrea gigas (67%), Hemigrapsus takanoi (>54%), Crassostrea gigas shells (44%) and Idotea balthica (24%). In contrast, mixed shells (Cerastoderma edule, Mytilus edulis, Ensis americanus and Littorina littorea) and Fucus versiculosus had no significant effect. These results suggest that dilution effects are widespread in the trematode of H. elongata, with potentially strong effects on its population dynamics.

Predators may have nonconsumptive effects (NCEs) on prey populations mediated by chemical cues detected by prey. We experimentally investigated dogwhelk (Nucella lapillus) NCEs on intertidal barnacle (Semibalanus balanoides) recruit density in Nova Scotia, Canada. Under a moderate abundance of coastal phytoplankton (food for barnacle larvae and recruits), barnacle recruitment was moderate and the nearby presence of dogwhelks limited barnacle recruit density at the end of the recruitment season. Under a high phytoplankton abundance, barnacle recruitment was high and neutralized dogwhelk NCEs on barnacle recruit density, likely through the chemical attraction that recruits exert on larvae seeking settlement.

Barnacles are dominant sessile invertebrates on many rocky shores worldwide. Hence, investigating the factors that affect their recruitment is important. Through field experiments done on the Atlantic coast of Canada, we investigated interspecific and intraspecific relationships affecting intertidal barnacle recruitment. Specifically, we evaluated the effects of seaweed canopies (Ascophyllum nodosum) and adult barnacles (Semibalanus balanoides) on the density of barnacle recruits at the end of the recruitment season. The effects of three canopy treatments on barnacle recruitment and understory environmental conditions allowed us to identify positive and negative effects of canopies. At mid-intertidal elevations subjected to a moderate wave action, we found that, during high tides, the flexible algal fronds damage wire sensors established on the substrate (whiplash effect) and limit barnacle recruitment. However, at low tide, algal canopies limit water loss and temperature extremes and enhance barnacle recruitment in understory habitats. The net effects of algal canopies on barnacle recruitment, however, were neutral, as the positive and negative influences balanced out. By manipulating the abundance of adult barnacles under the seaweed canopies, we found that adult barnacles enhance barnacle recruitment, likely due to the known attraction that adults exert on larvae seeking settlement and to the absence of post-settlement events that could otherwise have blurred such effects by the adults. The presence of adult barnacles, however, did not protect developing recruits from canopy whiplash effects. By understanding the contrasting influences that intertidal algal canopies have on understory abiotic conditions and barnacle recruitment, our ability to predict net canopy effects depending on the relative degree of physiological (e.g., high vs. low intertidal zone) and physical (e.g., sheltered vs. exposed shores) stresses should increase. This study also suggests that recruitment, considered as an important external factor in environmental models of community organization, can also be affected by components of the community itself, potentially triggering local feedbacks.