Larval settlement is an important process that drives population and community dynamics of marine invertebrates. Barnacles are frequently used to investigate settlement mechanisms of marine invertebrate larvae. Adult barnacles induce settlement of conspecific larvae nearby which ultimately facilitates copulation with neighbouring individuals. A significant factor involved in the larval induction process is the proteinaceous waterborne settlement pheromone (WSP), which is purified from adult barnacles. A previous study suggested that the concentration of WSP informs barnacle cyprid larvae about the abundance of adult barnacles in the environment nearby but it is unclear whether WSP works in a species-specific or non-species-specific manner. In this study, we conducted settlement assays using recombinant WSPs and cyprids of two congenic barnacle species, Amphibalanus amphitrite and A. improvisus, to investigate the species specificity of WSPs. We found that A. amphitrite and A. improvisus cyprids responded similarly to con-specific WSPs as to allo-specific WSP stimuli indicating that WSPs are not species-specific. Our findings suggest that cyprids approach potential settlement sites using WSPs, non-species-specific settlement pheromones, before performing a closer search of the substratum using a species-specific pheromone.

While parasites serve as prey, it is unclear how the spatial distribution of parasite predators provides transmission control and influences patterns of parasitism. Because many of its organisms are sessile, the rocky intertidal zone is a valuable but little used system to understand spatial patterns of parasitism and elucidate the underlying mechanisms driving these patterns. Sea anemones and barnacles are important space competitors in the rocky intertidal zone along the Pacific coast of North America. Anemones are voracious, indiscriminate predators; thus, they may intercept infectious stages of parasites before they reach a host. We investigate whether a sea anemone protects an associated barnacle from parasitism by Hemioniscus balani, an isopod parasitic castrator. At Coal Oil Point, Santa Barbara, California USA, 29% of barnacles were within 1 cm from an anemone at the surveyed tidal height. Barnacles associated with anemones had reduced parasite prevalence and higher reproductive productivity than those remote from sea anemones. In the laboratory, anemones readily consumed the transmission stage of the parasite. Hence, anemone consumption of parasite transmission stages may provide a mechanism by which community context regulates parasite prevalence at a local scale. Our results suggest predation may be an important process providing parasite transmission control.

Identification of marine invertebrate larvae using morphological characters is laborious and complicated by phenotypic plasticity. Balanus amphitrite is a dominant barnacle, important in the context of intertidal ecology and biofouling of manmade structures. Morphological identification of barnacle larval forms in a mixed population is difficult because of their intricacy and similarity in size, shape and developmental stages. We report the development and application of a nucleic acid-based Polymerase Chain Reaction (PCR) method for the specific identification of the barnacle, B. amphitrite, from the heterogeneous zooplankton sample. This method is reliable and accurate thereby overcoming taxonomic ambiguity. Sequence alignment of the 18S rRNA gene region of selected species of barnacles allowed the design of B. amphitrite-specific PCR primers. Assay specificity was evaluated by screening DNA obtained from selected species of barnacles. The oligonucleotide primers used in the study flanked a 1600 bp region within the 18S rRNA gene. The primer is specific and can detect as few as 10 individuals of B. amphitrite larvae spiked in a background of ~186 mg of zooplankton. This technique facilitates accurate identification and the primer can be used as a marker for enumeration of B. amphitrite larvae in the plankton.

Our study tracked three non-native, marine, sessile, invertebrate species which have recently colonized the south-eastern United States Atlantic coastlines and estuaries: Mytella charruana, the charru mussel; Perna viridis, the Asian green mussel; and Megabalanus coccopoma, the titan acorn barnacle. Along the eastern Floridian coast, Mytella charruana was discovered in the Indian River Lagoon, Florida (FL) in 2004. Perna viridis was found in Ponce de Leon Inlet, FL in 2002, after being introduced to Tampa Bay, FL in 1989. Megabalanus coccopoma was documented in St Augustine, FL and Brunswick, Georgia in 2006. From 2006 through to 2011, a biannual survey of the south-eastern Atlantic coastlines and estuaries was conducted from Jupiter, Florida to Charleston, South Carolina (894 km) to document the distribution and range expansion of M. charruana, P. viridis and M. coccopoma. Field surveys were conducted in June and December each year at 82 sites (e.g. docks, boat ramps, jetties and mangrove roots). The ranges of the three invertebrates have expanded and retracted along the Atlantic coast since the survey began. Mytella charruana had spread as far north as Savannah, Georgia in 2007, but the species was absent from northern Georgia in 2010 and 2011. Perna viridis and M. coccopoma were present in Georgia in 2007, but were absent in the 2010 and 2011 surveys. These range fluctuations may be explained by extreme cold temperatures, which occurred during the 2009/2010 and 2010/2011 winters. The patterns observed with these three sessile invertebrates provide information regarding how non-native species disperse and establish in new locations.

Acrothoracican barnacles of the genus Trypetesa are obligate symbionts of hermit crabs that burrow into the gastropod shells occupied by their hosts. In the present study, hermit crabs were examined for the presence of trypetesids, based on collections from the United States, Jamaica, and the Philippines made between 1997 and 2008. Shells from Jamaica and New York contained Trypetesa lateralis, a trypetesid previously documented from central California. Trypetesa lateralis is redescribed based on light and scanning electron microscopy, showing the presence of an external mantle flap and asymmetrical opercular bars diagnostic for this species. The mean prevalence of trypetesids in Jamaica was 8.3% and most barnacles were associated with Calcinus tibicen; in New York the barnacles were found in 1.6% of shells occupied by Pagurus longicarpus. Specimens from the Philippines were identified as Trypetesa spinulosa (formerly known only from Madagascar) based on the presence of their diagnostic orificial palps. The mean prevalence of T. spinulosa in the Philippines was 3.7% and most barnacles were associated with Calcinus spp. Hermit crab eggs were observed in the guts of T. lateralis from Jamaica and T. spinulosa from the Philippines. In both of these regions the trypetesids were found significantly more often in shells occupied by female hermit crab hosts (80–87% with females). These findings suggest the barnacles be classified as transient parasites. The biology of trypetesids is reviewed and a key to the family is provided. Further studies are needed to determine if egg predation occurs in all trypetesids and the impacts on hosts.

The antifouling activity of a series of hexane, ethyl acetate, ethanol and aqueous extracts from 11 species of marine sessile organisms collected from the south-east coast of China was investigated. Settlement inhibition of cyprid larvae of the barnacle Balanus albicostatus was used to evaluate their antifouling efficacy. Screening of the 44 extracts showed antifouling activity in 90.9% of the hexane extracts followed by 90.9% of the ethyl acetate, 72.7% of the ethanol and 36.4% of the aqueous extracts. The hexane extracts of Tubularia mesembryanthemum, Notarcus leachii cirrosus and Styela canopus, the ethyl acetate extracts of Bugula neritina and N. leachii cirrosus, and the ethanol extracts of B. neritina and Anthopleura sp. were the most active in inhibiting the settlement of B. albicostatus, with EC50 values all below 50 μg/ml. At least one of the four extracts of each tested species exhibited antifouling activity, suggesting that all 11 marine sessile organisms contained antifouling substances and they may have evolved chemical defences against biofouling on their surfaces.

Some specimens of the blue crab are bearing barnacles which have been identified as Chelonibia patula. Although generally known in the Mediterranean Sea this is the first report of C. patula from the Turkish coasts. It has been photographed and some morphometric measurements are provided.

Settlement of the fouling community is generally influenced by the physical, chemical and biological properties of the settling surface. The colour of the substratum may also exert an influence on larval settlement. In the present study, the influence of surface colour on the recruitment of fouling communities was investigated by suspending acrylic panels (red, green, blue, white and yellow) in coastal waters. Results showed that the colour of the substratum plays an important role in the recruitment of sessile species. Recruitment was comparatively high on red and blue coloured surfaces. Barnacle and tubeworm recruitment showed significant variation (Tukey test, P < 0.05) between different coloured surfaces. The significant conclusion of this study is that the colour of the artificial substrata should be taken into consideration when interpreting results from short-term biofouling studies.

The distribution of acetylcholinesterase (AChE) in the lateral eye and brain of the horseshoe crab was investigated with histochemical means using standard controls to eliminate butyrylcholinesterase and nonspecific staining. Intense staining was observed in the neural plexus of the lateral compound eye, in the lateral optic nerve, and in various neuropils of the brain. Nerve fibers with moderate to weak staining were widespread in the brain. No sornata were stained in either the lateral eye or the brain. The distribution of acetylcholinesterase in the supraesophageal ganglia and nerves of the giant barnacle was also investigated for comparison. Although both the median optic nerve of the barnacle and the lateral optic nerve of the horseshoe crab appear to contain the fibers of histaminergic neurons, only the lateral optic nerve of the horseshoe crab shows AChE staining. Other parts of the barnacle nervous system, however, showed intense AChE staining. These results along with the histochemical controls eliminate the possibility that some molecule found in histaminergic neurons accounted for the AChE staining but support the possibility that acetylcholine might be involved as a neurotransmitter in lateral inhibition in the horseshoe crab retina. Two reasonable neurotransmitter candidates for lateral inhibition, histamine and acetylcholine, must now be investigated.

We tested the hypothesis that histamine is the transmitter released by barnacle photoreceptors. Median and lateral ocelli were incubated with 3H-histidine and found to synthesize 3H-histamine, identified by high-voltage electrophoresis. Synthesis could be blocked by the histidine decarboxylase inhibitor (S)-α-fluoromethylhistidine. Histamine was applied to 1-cells either by superfusion or by pressure ejection from a pipette (100 µM or 1 mM histamine) positioned close to the I-cell's soma. When bath-applied at concentrations ranging from 5–100 µM, histamine hyperpolarized the I-cell in a dose-dependent fashion and increased its conductance. At 100 µM, histamine abolished the I-cell's response to light. The response to a pulse of pressure-applied histamine was a hyperpolarization whose amplitude was graded with dose (determined by the duration of the pulse). This response persisted in concentrations of Co2+ and Cd2+ that blocked synaptic transmission from the photoreceptors. Cimetidine, an antagonist of mammalian H2 receptors, markedly decreased the cell's responses both to HA and to light at 100 µM and blocked both responses at 1 mM. Pyrilamine and triprolidine, H1 antagonists, had a complex effect on the I-cell's responses to histamine and to light. Neither H1 nor H2 antagonists, nor histamine itself, affected the voltage or light responses recorded in the presynaptic terminal region, or any phase of calcium-dependent action potentials induced in the terminal in the presence of tetraethylammonium ion. Thus, biochemical, immunocytochemical, and physiological evidence suggests that HA is the transmitter from these photoreceptors to the I-cells. Although gammaaminobutyric acid (GABA) is also present in the photoreceptors, it did not affect the I-cell's responses to light or to histamine when bath-applied at 100 µM. Thus, GABA does not appear to modulate transmission from the photoreceptor to the I-cell.

We tested the hypothesis that gamma-aminobutyric acid (GABA) is the transmitter released by barnacle photoreceptors onto postsynaptic cells (I-cells). GABA was applied to I-cells either by superfusion or by ejecting it with pressure from a pipette positioned close to the I-cell's soma. The I-cell's response to GABA was compared with its response to light (i.e. to the photoreceptors' transmitter) by recording intracellularly from its soma. Bath-applied (100 µm to 10 mM) and pressure-applied GABA (10 mM in pipette) hyperpolarizes I-cells by increasing their conductance, as does the photoreceptors' transmitter. The response to pressure-applied GABA consists of two components; both persist when Co2+ or Cd2+ are added to the saline to block synaptic transmission in the preparation, indicating that GABA affects the I-cell directly rather than affecting a presynaptic cell. GABA hyperpolarizes the I-cell when applied to the cell over the soma and ipsilateral arbor or over the contralateral arbor. The I-cells' responses to GABA and to light both depend on extracellular K+ and are affected by changes in intracellular and extracellular Cl−. However, picrotoxin and β-guanidinopropionic acid block the response to pressure-applied GABA but do not block the response to light even at an order of magnitude higher concentration. Thus, GABA is not likely to be the transmitter that causes the hyperpolarizing response of the I-cell. It may be a neuromodulator or the transmitter of an unknown input to the I-cell.

The distribution and abundance of the Australasian barnacle Austrominius modestus around the Isle of Cumbrae, Scotland, were examined in October 2007. The species was found at all of the nineteen sites examined, which included highly exposed to highly sheltered sites. Its abundance varied from common to occasional. Over fifty years after its initial appearance as a single specimen (1954), Austrominius now appears to be well established on the island. Factors that may have influenced its spread and establishment are described, however, at this stage the relative importance of these cannot be determined. Consequently, further research is planned here, to collect quantitative data on the abundance of Austrominius and the other barnacle species that occur intertidally and how their relative abundances vary both within and between shores.

Previous observations in the St Lawrence Estuary (eastern Canada) suggested that larvae of intertidal barnacles (Semibalanus balanoides) would settle almost exclusively inside crevices on shores that are scoured by sea ice every winter. It was suggested that the strong ice scour in winter on that coast (which removes organisms outside of crevices) would select for such a larval behaviour. We tested the generality of this pattern by sampling other ice-scoured shores within the Gulf of St Lawrence system. In particular, we surveyed a shore in Nova Scotia where exposed habitats (subjected to strong ice scour in winter) are interspersed with sheltered habitats (which suffer milder ice scour). Such a topographical complexity might allow for the coastal larval pool to contain a proportion of larvae that have no particular settlement preference for crevices, as selective pressures for such a behaviour would be minimal in ice-sheltered habitats. Consistently with this notion, barnacle recruits (which appear after the winter ice melts) occurred abundantly both inside and outside of crevices across the shore in the spring seasons of 2005 and 2006. Average recruit density on rocky surfaces ranged between 337 and 588 recruits dm−2, depending on the habitat. It is therefore possible that barnacle recruitment patterns on ice-scoured shores may be affected indirectly by the structural complexity of the coast.

The sexuality of large (non-dwarf) individuals and the characteristics of eggs, larvae and dwarf males were investigated in the pedunculate barnacle Scalpellum stearnsii. All 103 large individuals collected in this study were not hermaphrodites but females. The major length axis of the egg was on average 0.50 mm, which was larger than that in most thoracican barnacles with planktotrophic larvae. The larvae hatched as nauplii and metamorphosed into cyprids without feeding (i.e. lecithotrophic development). There was a positive relationship between body weight and egg mass weight in ovigerous females. The number of males attached per female ranged from 0 to 35 (average: 5.9) and there was a positive relationship between the number of males and female body weight. The distribution of dwarf males was skewed significantly towards the lower part of the occludent margin, which is near the fertilization site.

Moulting and copulation by the barnacle Tetraclita japonica attached to an intertidal concrete pier were observed continuously with a video camera housed in a plastic case in summer 1996 and 1997. In a total of 779 individual-day observations, 10 individuals became sexually receptive as functional females and 14 moulted. Copulation occurred not only immediately after, but also before moulting. Duration of receptivity varied from 2 min to nearly 12 h, during which functional females accepted between two to 582 penis insertions by between one to 11 surrounding males. Each insertion time was short, ranging from 0.6 to 42 s, with a median of 2.4 s. Sex role changes were observed between two animals located close to each other, the shortest interval for which was 3.3 h. No significant size differences were detected between functional males and females. A temporal pattern of copulation, with a high frequency of penis insertions of short durations, is discussed in relation to natural habitat conditions.