Physiological tolerance predicts species composition at different scales in a barnacle guild

This study examined how the species composition of an intertidal barnacle guild varied according to physical gradients in the environment at small scales governed by microclimates, medium scales of wave exposure and large scales of latitude. Barnacle distributions at small and medium scales were sampled in Ireland between 51°29′ and 52°44′N and 6°50′ and 10°08′W. Sampling on European shores spanned ~18° latitude from 37°05′ to 55°16′N. Barnacle surveys mainly took place in 2003–2004. An index of wave fetch was calculated along the wave exposure gradient using a digital coastline-based model that was supported by a biological exposure scale. A ‘dryness’ index was defined according to mean monthly wind speed, fetch along the average wind direction and mean monthly air or sea surface temperatures for 2 years (January 2001–December 2002) which is the period when the most recent adults in the barnacle community would have settled and grown to adulthood. The proportion of the dry-loving barnacle Chthamalus montagui Southward increased within the barnacle guild at all scales as the habitat became warmer and drier. Barnacle densities were high in all habitats, mean densities ranged from a minimum of 4.16 cm⁻² on moderately exposed shores to a maximum of 6.27 cm⁻² in sunlit or south-facing microclimates. Percentage cover of barnacles across the gradient of latitudes was usually >70%. The results suggest that the distribution and abundance of interacting barnacle species on European coasts is strongly controlled by abiotic factors, most likely temperature and desiccation.     

Reproductive development of the barnacle Chthamalus malayensis in Hong Kong: implications for the life-history patterns of barnacles on seasonal, tropical shores

Hong Kong, lying just below the Tropic of Cancer (22°17′N, 114°09′E), experiences a strongly seasonal environment, with a cool almost temperate winter and a hot, tropical, summer. Histological sectioning of the gonads of the high-shore barnacle, Chthamalus malayensis Pilsbry, showed a seasonal trend in the development of its reproductive organs. Four stages of female gonad development were identified according to the cell types present: post-spawning, resting, growth and mature stages. The female gonad was mature from April to November, which was related to seawater temperatures, and entered a resting phase from December to March. Although the male gonad showed a seasonal developmental trend and reached maximum maturity in summer, the seminal vesicles were full of spermatozoa and functional throughout the year. The reproductive season of this species is therefore solely dependant upon the maturity of the female gonad. The estimated maximum number of broods per year was up to 10 and the maximum number of eggs produced per brood can reach 3,000 eggs. The minimum size for female gonad maturity was 6 mm rostro-carinal diameter (RCD) at which size, the barnacles were ~6-month old. Sperm production occurred at a smaller size (2 mm=2-month old). Compared with Chthamalus montagui and Chthamalus stellatus from temperate regions, C. malayensis produced a greater number of broods per year, had a longer reproductive period and faster gonad development. Chthamaloid barnacles in tropical regions may, therefore, invest more energy per year in reproduction during their life span. Contrary to the seasonal gonad developmental pattern of C. malayensis in the present study, however, C. malayensis in Singapore (which experiences only slight seasonal variation) had mature female and male gonads throughout the year, further supporting the strong role of climatic conditions effecting the reproductive biology of barnacles.     

Function of the shell spine in the predaceous rocky intertidal snail Acanthina spirata (Prosobranchia: Muricacea)

Feeding observations revealed that Acanthina spirata (Blainville, 1832) uses the shell spine in a ramming and prying motion, resulting in the fracturing and/or forcing open of opercular plates in the barnacles Balanus glandula (Darwin, 1854) and Chthamalus fissus (Darwin, 1854). Attacking barnacles by using the shell spine may represent a second widespread feeding mechanism (in addition to drilling) in muricacean gastropods. Specialization of the shell spine for attacking barnacles is advantageous over drilling within the context of an optimal foraging theory. Rockey intertidal field and laboratory experiments at Palos Verdes and Los Angeles, California, respectively, in 1981–1982 revealed, that snails with a shell spine have lower handling times and feed at a greater rate than spineless snails which drill their prey. Spine penetration of prey as opposed to drilling takes considerably less time than a tidal cycle. This is expected to increase the success rate for completion of feeding, since during tidal flux snails risk being dislodged. Plasticity in attack mechanisms allows A. spirata with broken spines to penetrate prey by drilling while undergoing spine repair. Differences exist in spine-feeding by A. spirata on B. glandula and C. fissus. The attack process takes more than twice as long when snails attack B. glandula versus C. fissus. However, there is no difference in the mean number of spine thrusts required to penetrate the opercular plates of the two species. When feeding on the larger barnacle B. glandula, A. spirata ingests significantly greater dry weight per unit handling time than with C. fissus.     

Population dynamics of the barnacle <Emphasis Type="Italic">Chthamalus montagui</Emphasis> at two spatial and temporal scales in northern Spain

This study describes spatial and temporal patterns of variability in population parameters in the barnacle Chthamalus montagui Southward in three localities of northern Spain and evaluates whether density-dependent settlement may regulate population dynamics. The sampling design considered two spatial scales, localities and sites within localities, and two temporal scales, years and six month intervals. Density, amount of free space, mortality, growth rate and magnitude of settlement (both absolute and per unit of free space) were obtained from photographs of permanent quadrats and from direct counts in the field. The number of settlers in scraped and untouched quadrats was used to estimate the importance of the presence of conspecifics in settlement. Significant variation at the two spatial and temporal scales was found for most parameters. Large spatial and temporal variations in adult mortality rate, density, and settlement were observed. Patterns of mortality were not consistent with differences in density among localities. Differences in settlement among localities were maintained through time. We suggest that magnitude of settlement is regulated by persistent features such as topography or local water circulation. We assume that early post-settlement mortality does not differ among localities. In the absence of differential mortality, settlement determines average population density within localities. Within localities, settlement was independent of density and free space. No consistent evidence was found on preferential settlement at the vicinity of conspecifics. The main conclusion is that density-dependent settlement is not relevant for the regulation of the populations of C. montagui in the northern Spain. Regulation might occur by density-dependent processes within the adult fraction of the population and/or the larval phase before settlement.     

The effect of water temperature on drilling and ingestion rates of the dogwhelk Nucella lapillus feeding on Mytilus edulis mussels in the laboratory

In highly seasonal intertidal habitats, changes in temperature through the year may drive substantial shifts in feeding and growth rates of organisms. For the dogwhelk Nucella lapillus, attacking and consuming Mytilus edulis mussels can take hours or days, depending on temperature. Handling time of dogwhelks feeding on mussels is therefore greatly affected by ocean temperature. I recorded attack time in the laboratory, partitioned into drilling and consumption time, for juvenile dogwhelks across a range of seawater temperatures representative of field seawater temperatures during the main growing seasons of summer and autumn. The combined length of a drilling attack and subsequent ingestion time tripled across the 10 °C decline in water temperatures from July through November, driven primarily by an increase in ingestion time. The observed reduction in handling time, coupled with projected sea surface warming in New England by the end of the twenty-first century, could extend the length of the growing season for Nucella and subsequently have cascading effects on the prey community.     

Effects of settlement density on spatial arrangement in four intertidal barnacles

Balanus amphitrite was studied in an estuary (Halifax River) in central Florida from October to November 1990, while three other barnacle species were studied on the central coast (Monterey Bay) of California from April 1988 to July 1989. Mean nearest-neighbor distances indicated that in the majority of cases the spatial arrangement of settlers was random for three different balanomorph barnacles-B. amphitrite, B. glandula, and Chthamalus dalli. In Pollicipes polymerus, a lepidomorph species, strong aggregation among settlers was almost always observed. In the three balanomorph species there was no apparent relationship between settler density and the degree of aggregation. In P. polymerus there was a significant positive correlation between settler density and the degree of aggregation. Morphological differences between balanomorph and lepidomorph barnacles may influence the evolution of settlement behavior.     

Tolerance of juvenile barnacles (Amphibalanus improvisus) to warming and elevated pCO2

We investigated the impacts of warming and elevated pCO₂ on newly settled Amphibalanus improvisus from Kiel Fjord, an estuarine ecosystem characterized by significant natural pCO₂ variability. In two experiments, juvenile barnacles were maintained at two temperature and three pCO₂ levels (20/24 °C, 700–2,140 μatm) for 8 weeks in a batch culture and at four pCO₂ levels (20 °C, 620–2,870 μatm) for 12 weeks in a water flow-through system. Warming as well as elevated pCO₂ hardly affected growth or the condition index of barnacles, although some factor combinations led to temporal significances in enhanced or reduced growth with an increase in pCO₂. While warming increased the shell strength of A. improvisus individuals, elevated pCO₂ had only weak effects. We demonstrate a strong tolerance of juvenile A. improvisus to mean acidification levels of about 1,000 μatm pCO₂ as is already naturally experienced by the investigated barnacle population.     

Sex in the tropics: reproduction of<Emphasis Type="Italic"> Chthamalus malayensis</Emphasis> Pilsbury (Class Cirripedia) at the equator

The reproduction of a key intertidal space occupier, the barnacle, Chthamalus malayensis, was examined over a 13-month period. The reproductive cycle of C. malayensis was studied at three shores (Singapore, east and west coast of Malaysia) near the centre of its geographical range. Testes, vesiculae seminales, ovaries and embryos were scored for their stages of development. Although, in general, C. malayensis contained gonads throughout the year, the timing and level of peak development varied between the three shores. Embryos were present throughout the year in west Malaysia but were absent for 4–5 months in east Malaysia (June–August and October) and Singapore (February, April and August). Variations in the reproductive cycles and brooding were not clearly linked to the timing of monsoons. The lack of seasonal peaks in breeding may be attributed to the relatively constant climatic conditions experienced near the equator.Communicated by M. S. Johnson, Crawley     

Shell selection in the hermit crab Pagurus bernhardus

Three shell variables were deemed to be important in the selection of a shell by the hermit crab Pagurus bernhardus (L.). These were the weight, volume and aperture width of a shell. Each of these variables were used in multiple regressions against crab weight for 25 Littorina littorea and 25 Thais lapillus shells chosen when both shell species were together in a tank, and for a further 25 L. littorea and 25 T. lapillus when the two shell species were separate. Principal component analysis was performed on the four groups of 25 shells selected, and multiple-regression equations were calculated using the principal components as the new variables. It was found that crabs chose a shell of suitable general dimensions rather than solely on the basis of one shell variable. The advantages of using statistical techniques developed in this paper over previous approaches to shell selection are discussed.     

Remote Antarctic feeding ground important for east Australian humpback whales

Understanding the dynamics of population recovery is particularly complex when an organism has multiple, remote breeding and feeding grounds separated by one of the longest known migration routes. This study reports on the most comprehensive assessment of humpback whale (Megaptera novaeangliae) movements between remote Antarctic waters south of New Zealand and east Australia (EA), and the migratory corridors and breeding grounds of Australia and Oceania. A total of 112 individual whales were identified; 57 from microsatellites and 61 by fluke with 23 % (n = 26) matched to sites outside Antarctica. Despite large datasets from other southern regions being included in the comparison, the whales were predominantly linked to EA (n = 24). Only two matches to the Oceania catalogues directly north was surprising; therefore the primary feeding grounds of these endangered whales still remain unknown. The confirmation of the Balleny Islands as an important feeding ground for EA whales could provide an insight into reasons behind the rapid recovery of this population. Determining the feeding grounds of Oceania’s whales may explain whether prey energetics or migration length are limiting factors to their recovery and will allow an understanding of future ecosystem changes in these whales.     

Primary production and phytoplankton community structure during a winter shelf-scale phytoplankton bloom off Western Australia

Regions of high primary production along the oligotrophic west coast of Australia between 34 and 22°S in May–June 2007 (midway through the annual phytoplankton bloom) were found around mesoscale features of the Leeuwin Current. At 31°S, an anticyclonic eddy-forming meander of the Leeuwin Current had a mixed layer depth of >160 m, a depth-integrated chlorophyll a (Chl a)-normalised primary production of 24 mg C mg Chl a ^?1 day^?1 compared to the surrounding values of <18 mg C mg Chl a ^?1 day^?1. In the north between 27 and 24°S, there were several stations in >1,000 m of water with a shallow (<100 m) and relatively thin layer of high nitrate below the mixed layer but within the euphotic zone. These stations had high primary production at depths of ~100 m (up to 7.5 mg C m^?3 day^?1) with very high rates of production per unit Chl a (up to 150 mg C mg Chl a ^?1 day^?1). At 27–24°S, the majority of the phytoplankton community was the ubiquitous tropical picoplankters, Synechococcus and Prochlorococcus. There was a decline in the dominance of the picoplankters and a shift towards a more diverse community with more diatoms, chlorophytes, prasinophytes and cryptophytes at stations with elevated production. Photosynthetic dinoflagellates were negligible, but heterotrophic dinoflagellate taxa were common. Haptophytes and pelagophytes were also common, but seemed to contribute little to the geographical variation in primary production. The mesoscale features in the Leeuwin Current may have enhanced horizontal exchange and vertical mixing, which introduced nitrate into the euphotic zone, increasing primary production and causing a shift in phytoplankton community composition in association with the annual winter bloom.     

Consequences of maternal isolation from salinity stress for brooded embryos and future juveniles in the estuarine direct-developing gastropod Crepipatella dilatata

At reduced salinities, brooding females of some gastropods and bivalves may isolate their mantle cavities from the environment for several days, maintaining internal osmotic concentration but causing severe declines in dissolved oxygen and pH, and increases in ammonia and other toxic substances in the mantle fluid. This study in November–December examined the immediate consequences of such stresses for brooded embryos of Quempillén estuary gastropod Crepipatella dilatata, in terms of time to juvenile emergence and rates of embryonic growth [measured as shell length (SL)]. Juveniles were also monitored for latent effects on feeding rates, oxygen consumption, and growth for the first 4 weeks after emergence into normal salinity seawater. An acute salinity stress lasting 3 days applied to females that were brooding pre-shelled or intermediate-shelled stages increased embryonic incubation periods, but without affecting SL at emergence. Growth rates were reduced for encapsulated embryos regardless of the stage at which the salinity stress was applied. Latent effects on juvenile development included slower shell growth and reduced rates of oxygen consumption and feeding. These effects were sustained for the first month after release from the female. The results suggest that marked reductions in salinity lasting for several days indirectly but negatively affect the development of brooded embryos of C. dilatata and also affect the juveniles for at least several weeks following their release, even after salinity has returned to normal.     

Environmental and structural control of trace elements in barnacle shells

The shell composition of 5 species of barnacles [Balanus balanoides (L.), B. crenatus Bruguière, B. hameri Ascanius, Elminius modestus Darwin and Chthamalus stellatus (Poli)] collected from four different sites on the coast of North Wales and one site in the Irish Sea were studied using chemical, X-ray diffraction and electron-microprobe analyses. All the shells consisted of calcite. The trace substances studied were Mg, Sr, Na, Mn, PO₄, and SO₄. Other trace elements detected included Si, Li, Ba, Zn, Cl, and Al. For each species, Mn was the only element in the shell which varied significantly from one locality to another. Variation of concentration of Mn in the shells may be related to the variations of the concentration of Mn in the water, since concentrations in the shells were highest in (estuarine) waters containing high concentrations of Mn. In Balanus balanoides L. and Elminius modestus Darwin, the Mn:Ca and Mg:Ca ratios decreased with decreasing shore level and Sr:Ca varied in the opposite direction. When B. crenatus Bruguière, a subtidal and lower littoral species, and B. hameri Ascanius, a Continental-Shelf species, were compared with the intertidal, species B. balanoides and E. modestus, the Mn:Ca and Mg:Ca ratios followed the same decreasing trend, and the Sr:Ca ratios the same increasing trend with increasing depth. These variations may be related either to increased growth rate of the shells of each species or maxium temperature to which the species is exposed, or both. The differences between species are more consistently reflected, in the concentration of Mn. However, since the concentration, of Mn is correlated with the amount of organic matter in the shell, the specific variations may reflect only the variations in the amount of organic material in the shell of each species.     

Marine turtle threats in Uruguayan waters: insights from 12 years of stranding data

We present the first study conducted in a wide spatio-temporal scale on marine turtles strandings (N = 1,107) over a 12-year period (1999–2010) in Uruguay. Five species were recorded Chelonia mydas (N = 643; 58.1 %), Caretta caretta (N = 329; 29.7 %), Dermochelys coriacea (N = 131; 11.8 %), Eretmochelys imbricata (N = 3; 0.3 %), and Lepidochelys olivacea (N = 1; 0.1 %). The first three species stranded throughout the Uruguayan coast, but differences in distribution patterns were detected among species. Although occurring year round, stranding records show a clear seasonal pattern with variation in monthly distribution among species, but with a peak of records in austral summer. Strandings provide indirect evidence of threats to marine turtles in Uruguayan and surrounding waters, particularly fisheries and marine debris. Our results demonstrate that Uruguayan coastal waters likely serve as a foraging or development area for at least three endangered marine turtle species in temperate waters.     

Feeding rates and prey selectivity of planktonic decapod larvae in the Western English Channel

Meroplankton are seasonally important contributors to the zooplankton, particularly at inshore sites, yet their feeding ecology is poorly known relative to holoplankton. While several studies have measured feeding in decapod larvae, few studies have examined the feeding rates of decapod larvae on natural prey assemblages throughout the reproductive season. We conducted 8 feeding experiments with Necora puber, Liocarcinus spp. and Upogebia spp. zoea larvae collected from the L4 monitoring site off Plymouth (50°15.00′N, 4°13.02′W) during spring–summer 2009 and 2010. This period spanned moderate-to-high food availability (0.5–1.6 µg chl-a L^?1), but a great range in food composition with small cells <20 µm dominating in 2010. Daily rations averaged 17, 60 and 22 % of body C for the 3 respective decapod species. Clearance rates differed according to prey type, and all 3 decapod genera showed evidence of selection of dinoflagellates. Importantly, small cells including nano- and pico-plankton were ingested, this being demonstrated independently by flow cytometric analysis of the feeding experiments and molecular analysis. PCR-based analysis of the haptophyte portion of the diet revealed ingestion of Isochrysis galbana by decapod larvae in the bottle incubations and Isochrysis galbana and Phaeocystis globosa by decapod larvae collected directly from the field. This study has shown that pico- and nano-sized plankton form an important supplement to the diverse and variable diet of decapod larvae.     

Genetic structure of populations of two species of Chthamalus (Crustacea: Cirripedia) in the north-east Atlantic and Mediterranean

Protein electrophoresis on starch gels was used to investigate population genetic structure of the barnacles Chthamalus montagui Southward and C. stellatus (Poli) over their north-east Atlantic and Mediterranean ranges. In each species, a single locus exhibited marked differentiation of allele frequencies between Atlantic and Mediterranean localities; in C. stellatus, genetic differentiation between the two basins had not previously been noted. In both species, mean heterozygosity per locus appeared higher in the Mediterranean samples than in the Atlantic, and Mediterranean populations had more alleles at the loci studied. Possible explanations for the differentiation between the Atlantic and the Mediterranean populations are discussed. Received: 30 May 1996 / Accepted: 17 September 1996     

Hypoxic life of intertidal acorn barnacles

Oxygen levels were monitored within the mantle cavities of three barnacle species ( Chthamalus stellatus, Semibalanus balanoides, Elminius modestus), using optode microsensors. Conditions were always hypoxic, even when barnacles were actively using the prosoma and cirri to pump aerated seawater into the mantle cavity. Mantle fluid oxygen concentrations were extremely variable; behaviour and oxygen concentrations were not closely coupled. Ventilation of the mantle cavity depended partially on external water flow, with higher and more stable mantle fluid oxygen concentrations being sustained when the water around barnacles was agitated. During emersion, barnacles initially pumped seawater between the mantle cavity and the cone above the opercular plates to achieve ventilation. As water was lost it was replaced by air bubbles, eventually resulting in an air-filled mantle cavity. In S. balanoides and E. modestus, once the mantle cavity was filled with air, the barnacle usually used up the oxygen within the bubble within 2–3 h and did not regain oxic conditions until the barnacle was reimmersed in seawater. In C. stellatus, the air bubble was repeatedly refreshed for many hours by pneumostome formation. In response to low environmental salinity, all three species closed the opercular plates firmly and rapidly used up oxygen within the mantle fluid.     

Habitat,movements and environmental preferences of dusky sharks,Carcharhinus obscurus,in the northern Gulf of Mexico

The dusky shark (Carcharhinus obscurus) is the largest member of the genus Carcharhinus and inhabits coastal and pelagic ecosystems circumglobally in temperate, subtropical and tropical marine waters. In the western North Atlantic Ocean (WNA), dusky sharks are overfished and considered vulnerable by the International Union for the Conservation of Nature. As a result, retention of dusky sharks in commercial and recreational fisheries off the east coast of the United States (US) and in the northern Gulf of Mexico is prohibited. Despite the concerns regarding the status of dusky sharks in the WNA, little is known about their habitat utilization. During the summers of 2008–2009, pop-up satellite archival tags were attached to ten dusky sharks (one male, nine females) at a location where they have been observed to aggregate in the north central Gulf of Mexico southwest of the Mississippi River Delta to examine their movement patterns and habitat utilization. All tags successfully transmitted data with deployment durations ranging from 6 to 124 days. Tag data revealed shark movements in excess of 200 km from initial tagging locations, with sharks primarily utilizing offshore waters associated with the continental shelf edge from Desoto Canyon to the Texas/Mexican border. While most sharks remained in US waters, one individual moved from the northern Gulf of Mexico into the Bay of Campeche off the coast of Mexico. Sharks spent 87 % of their time between 20 and 125 m and 83 % of their time in waters between 23 and 30 °C. Since dusky sharks are among the most vulnerable shark species to fishing mortality, there is a recovery plan in place for US waters; however, since they have been shown to make long-distance migrations, a multi-national management plan within the WNA may be needed to ensure the successful recovery of this population.     

A new method for high-resolution bivalve growth rate studies in hydrothermal environments

The age and shell growth rate of deep-sea hydrothermal bivalves were investigated for the first time using in situ chemical staining combined with high-resolution micro-increment analysis. A staining chamber developed for this purpose was applied to a patch of Bathymodiolus thermophilus mussels at 2,500 m depth at the 9°47′N vent field on the East Pacific Rise (EPR) in May 2010. This approach minimizes disturbance of the mussels in their habitat. Bathymodiolus thermophilus grows according to a circalunidian rhythm, with one increment formed each day, and displays tide-related growth rate variability. Based on the von Bertalanffy growth rate model, the largest shell collected (SL = 20.5 cm) would be 10.0 year old, with a growth rate of 4.2–1.1 cm year^?1 as the shell ages. This fast growth rate is consistent with the instability of the environment in this section of the EPR and observed recolonization rates and could reflect a specific adaptation of this species.