A finite element method to solve a population dynamics stage-structured model of intertidal barnacles

Many marine organisms are fixed or highly sedentary as adults but the adult population may be strongly dependent on the oceanic transport of planktonic larvae. In order to assess interactions between oceanographic and biological processes that determine the population dynamics of marine organisms with a sessile adult phase restricted to the coastline and a planktonic larval phase, we present a stage-structured finite element model for the barnacle Balanus glandula that inhabits the rocky intertidal zone of central California, USA.     

Influence of substratum heterogeneity and settled barnacle density on the settlement of cypris larvae

On the Atlantic coast of Canada, Semibalanus balanoides (L.) is widely distributed in the mid-intertidal zone, whereas in the Gulf of St. Lawrence, this species is mostly limited to crevices. We tested the hypothesis of regional differences in microhabitat selection by barnacle larvae at settlement in 1984 and 1985 at St. Andrews, New Brunswick, Canada. Since larvae settle in microhabitats already colonized by adults, the relative influence of settled barnacle density and of different scales of substratum heterogeneity on settlement were evaluated experimentally at Capucins, Québec, (Gulf of St. Lawrence) and at St. Andrews, New Brunswick (Atlantic coast). On a large scale (>10 cm deep crevices) of heterogeneity, results show that, in the Gulf, cypris larvae settled nearly exclusively (93%) in natural crevices rather than on adjacent horizontal surfaces. On the Atlantic coast, settlement was more important outside than inside of crevices, when the substrata were either natural or artificial. This result is unique and contrasts sharply with all known reports on barnacle settlement in relation to surface contour. The influence of barnacle density on settlement was greater than that of large scale heterogeneity. On a small scale (<1.5 cm deep cracks), the presence of conspecifics had a stronger effect on settlement than heterogeneity in both regions. Field observations showed a relationship between larval settlement density and percentage of adult cover. Settlement increased up to 22 or 30% (Gulf and Atlantic coast) of adult cover and decreased afterwards. The results confirm the hypothesis of larval selection for cryptic habitats in the Gulf and the opposite behaviour (preferences for horizontal surfaces) on the Atlantic coast. This microhabitat selection is apparent at large scales of heterogeneity, whereas at small scales, the presence of conspecifics is the predominant factor.Contribution to the programme of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Inhibition of attachment of larval barnacles,Balanus amphitrite,by bacterial surface films

Films of bacteria on solid substrata can positively or negatively influence the attachment of marine invertebrate larvae. Effects of marine bacteria on the attachment of cypris larvae of the barnacle Balanus amphitrite Darwin were examined in the laboratory. Bacteria, grown to mid-exponential phase and allowed to adsorb irreversibly to polystyrene petri dishes, attached in densities of 10⁷ cells cm^-2. Assays (22h) were used to compare the effects of adsorbed cells of 18 different bacterial species on larval barnacle attachment. Most of the adsorbed bacteria either inhibited or had no effect on larval attachment compared to clean surfaces. Experiments testing the effect of larval age on barnacle attachment were conducted with six species of bacteria and showed that older larvae attached in higher percentages to clean surfaces and that bacterial films generally inhibited larval attaschment. Both the species of bacteria and the in situ age of the adsorbed bacteria affected barnacle attachment response: older films of Deleya (Pseudomonas) marina were more inhibitory. Bacterial extracellular materials may be involved in the inhibitory process.     

Predicting intertidal organism temperatures with modified land surface models

Animals and plants in the marine intertidal zone live at the interface between terrestrial and marine environments. This zone is likely to be a sensitive indicator of the effects of climate change in coastal ecosystems, because of several key characteristics including steep environmental gradients, rapid temperature changes during tide transitions, fierce competition for limited space, and a community of mostly sessile organisms. Here we describe a modular modeling approach using modifications to a meteorological land surface model to determine body temperatures of the ecologically dominant rocky intertidal mussel Mytilus californianus, as a tool that can be used as a proxy for ecological performance. We validate model results against in situ measurements made with biomimetic body temperature sensors. Model predictions lie within the range of variability of biomimetic measurements, based on observations over a 4-year period at sites along 1700 km of the US west coast from southern California (34.5°N) to northern Washington (48.4°N). Our modular approach can be easily applied to many situations in the intertidal zone, including bare rock, mussel, barnacle, and algal beds, salt-marsh grasses, and sand- and mud-flats, by modifying the “vegetation layer” in a standard meteorological land surface model. Biophysical models such as these, which link ecological processes to changing climates through predictions of body temperature, are essential for understanding biogeographic patterns of physiological stress and mortality risk.     

Predators of juvenile sea urchins and the effect of habitat refuges

We evaluated the effects of potential predators from intertidal habitats on Strongylocentrotus purpuratus survival using laboratory experiments and assessed abundances of main predatory species along the Pacific coast of North America. The interactive effects of urchins’ and predators’ sizes in mediating predation were quantified. Habitat complexity (substrate pits, adult spine canopy) was manipulated to examine its effects on predation of most susceptible individuals (<14 mm). Pachygrapsus crassipes was identified as a major predator of urchins up to ≈30 mm. A positive effect of predator size on consumption of progressively larger urchins was detected, probably due to a mechanical limitation on crabs’ ability to consume large prey. Larger claws of males with respect to females of comparable sizes facilitated the handling of larger prey. Substrate refuges significantly reduced mortality on juvenile urchins. These results show that crab predation may be important in organizing intertidal communities, despite multiple ecological mechanisms promoting sea urchin survival.     

Some observations of factors affecting embryonic and larval growth of Pollicipes polymerus (Cirripedia: Lepadomorpha) in vitro

Observations on the effects of several environmental conditions in embryonic and larval cultures are reported for the gooseneck barnacle Pollicipes polymerus (Sowerby, 1833). When the growth rates of control embryos kept in finger bowls at ambient-temperature sea water (ca. 12°–15°C) were compared to those of embryos grown under variously modified circumstances, it was found that growth rates were faster under conditions closely simulating the adult barnacle mantle-cavity (e.g. darkness and aeration). Addition of antimicrobial drugs and reduction of egg-mass size also promoted fast growth and development. Nauplii were fed several species of algae; only three species promoted growth to Stage V or older. Naupliar growth was fastest in larvae fed the combination of Prorocentrum micans/Platymonas sp. Larvae fed Prococentrum micans/pennate diatom grew slowest, and most larvae died at Naupliar Stage III. Interactions between algal species may have affected their nutritional value for Pollicipes polymerus nauplii. Less larval activity, slower growth rates, and higher mortality rates were observed in individuals given small amounts of food. The development of embryos and larvae in culture depends greatly on the culture conditions. These conditions should be described if comparison of timetables and envents are to be made between studies.     

Feeding by larvae of intertidal invertebrates: assessing their position in pelagic food webs

One of the leading determinants of the structure and dynamics of marine populations is the rate of arrival of new individuals to local sites. While physical transport processes play major roles in delivering larvae to the shore, these processes become most important after larvae have survived the perils of life in the plankton, where they usually suffer great mortality. The lack of information regarding larval feeding makes it difficult to assess the effects of food supply on larval survival, or the role larvae may play in nearshore food webs. Here, we examine the spectrum of food sizes and food types consumed by the larvae of two intertidal barnacle species and of the predatory gastropod Concholepas concholepas. We conducted replicated experiments in which larvae were exposed to the food size spectrum (phytoplankton, microprotozoan and autotrophic picoplankton) found in nearshore waters in central Chile. Results show that barnacle nauplii and gastropod veligers are omnivorous grazers, incorporating significant fractions of heterotrophs in their diets. In accordance with their feeding mechanisms and body size, barnacle nauplii were able to feed on autotrophic picoplankton (<5 microm) and did not consume the largest phytoplankton cells, which made the bulk of phytoplankton biomass in spring-summer blooms. Balanoid nauplii exhibited higher ingestion rates than the smaller-bodied chthamaloid larvae. Newly hatched C. concholepas larvae also consumed picoplankton cells, while competent larvae of this species ingested mostly the largest phytoplankton cells and heterotrophic protozoans. Results suggest that persistent changes in the structure of pelagic food webs can have important effects on the species-specific food availability for invertebrate larvae, which can result in large-scale differences in recruitment rates of a given species, and in the relative recruitment success of the different species that make up benthic communities.     

Phenotypic variation in shell form in the intertidal acorn barnacle Chthamalus montagui: distribution,response to predators and life history trade-offs

The acorn barnacle Chthamalus montagui can present strong variation in shell morphology, ranging from flat conic to a highly bent form, caused by a substantial overgrowth of the rostrum plate. Shell shape distribution was investigated between January and May 2004 from geographical to microhabitat spatial scales along the western coast of Britain. Populations studied in the north (Scotland and Isle of Man) showed a higher degree of shell variation compared to those in the south (Wales and south-west England). In the north, C. montagui living at lower tidal levels and in proximity to the predatory dogwhelk, Nucella lapillus, were more bent in profile. Laboratory experiments were conducted to examine behavioural responses, and vulnerability of bent and conic barnacles to predation by N. lapillus. Dogwhelks did not attack one morphotype more than the other, but only 15 % of attacks on bent forms were successful compared to 75 % in conic forms. Dogwhelk effluent reduced the time spent feeding by C. montagui (11 %), but there was no significant difference between conic and bent forms. Examination of barnacle morphology indicated a trade-off in investment in shell structure and feeding appendages associated with being bent, but none with egg or somatic tissue mass. These results are consistent with C. montagui showing an induced defence comparable to that found in its congeners Chthamalus anisopoma and Chthamalus fissus on the Pacific coast of North America, but further work to demonstrate inducibility is required.     

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     

Fertilisation, embryogenesis and larval development in the tropical intertidal sand dollar Arachnoides placenta in response to reduced seawater pH

We examined the response of the tropical sand dollar Arachnoides placenta to reduced seawater pH in experiments spanning ca. 50 % of the planktonic larval duration. A. placenta inhabits intertidal sandy beaches where we observed a minimum in situ pH range 0.06 pH units (pH 8.10–8.16). The responses of gametes and larvae to seawater pH were tested in vitro in ambient (pH 8.14, pCO₂ = 525.7 μatm, total alkalinity = 2,651 μmol kg soln^?1) and three reduced pH seawater treatments (7.8–7.0). Percentage fertilisation decreased significantly with decreasing pH across a range of sperm/egg ratios (4:1 up to 4,000:1). A. placenta reached the advanced pluteus stage in 4 days, and during this time, we saw no difference in survival rate of larvae between the ambient (67 %) and pH 7.79 (72 %) treatments. Four-day survival was, however, reduced to 44 and 11 % in the pH 7.65 and 7.12 treatments, respectively. Larval development and morphometrics varied among pH treatments. Embryos reared in pH 7.12 exhibited arrested development. Larvae reared at pH 7.65 showed delayed development and greater mortality compared with those reared at pH 7.79 and 8.14. When larval morphometrics are compared among larvae of the same size, differences in larval width and total arm length between pH treatments disappear. These results suggest that variation in larval size among the three highest pH treatments at a given time are likely the result of slower development and apparent shrinkage of surviving larvae and not direct changes in larval shape. There were no differences in the percentage inorganic content (a proxy for calcification) in larvae reared in either an ambient or a pH 7.7 treatment. The responses of fertilisation and development to decreased pH/increased pCO₂ in A. placenta are within the range of those reported for other intertidal and subtidal echinoid species from colder latitudes.     

Phylogeography of the European stalked barnacle (Pollicipes pollicipes): identification of glacial refugia

The alternation of glacial and interglacial events during the Pleistocene has produced changes in species distribution ranges leading to bottlenecks and alterations of patterns of gene flow. The European stalked barnacle, Pollicipes pollicipes, is a sessile pedunculate cirripede that inhabits the rocky intertidal frame, from Senegal to the northwestern coast of France. In this work, we have analyzed a fragment of the mitochondrial gene cytochrome c oxidase subunit I for 569 individuals of P. pollicipes in order to investigate whether the shifts in climatic conditions that occurred during the Pleistocene influenced the current pattern of distribution of genetic variation of P. pollicipes. A pre-last glacial maximum pattern of demographic expansion was found, in concordance with many other North Atlantic marine species. On the other hand, three potential glacial refugia were identified: North African coasts, northwestern Iberian Peninsula and English Channel/Brittany.     

Seasonal variation in utilization of biogenic microhabitats by littorinid snails on tropical rocky shores

Mobile species may actively seek refuge from stressful conditions in biogenic habitats on rocky shores. In Hong Kong, the upper intertidal zone is extremely stressful, especially in summer when organisms are emersed for long periods in hot desiccating conditions. As a result, many species migrate downshore between winter and summer to reduce these stressful conditions. The littorinids Echinolittorina malaccana and E. vidua, for example, are found on open rock surfaces high on the shore in winter but the majority migrate downshore in summer to the same tidal height as a common barnacle, Tetraclita japonica. In the laboratory, where environmental conditions could be controlled to approximate those occurring on the shore, we tested whether the downshore migration allowed littorinids to select barnacles as biogenic habitats to reduce stress and if this behaviour varied between seasons. In summer, littorinids demonstrated a strong active preference for the barnacles, which was not observed in the cool winter conditions, when animals were found on open rock surfaces even when barnacles were present. Littorinids, therefore, only actively select biogenic habitats during the summer in Hong Kong when they migrate downshore, suggesting that such habitats may play an important, temporal, role in mitigating environmental stress on tropical shores.     

Activity patterns and predatory behavior of an intertidal nemertean from rocky shores: Prosorhochmus nelsoni (Hoplonemertea) from the Southeast Pacific

Understanding the impact of environmental stressors on predator activity is a prerequisite to understanding the underlying mechanisms shaping community structure. The nemertean Prosorhochmus nelsoni is a common predator in the mid-intertidal zone on rocky shores along the Chilean coast, where it can reach very high abundances (up to 260 ind m⁻²) in algal turfs, algal crusts, barnacle crusts, and mixed substrata. Tidal and diurnal scans revealed that the activity of P. nelsoni is primarily restricted to night and early-morning low tides and is relatively low when air temperatures are high. On average, larger worms crawled faster than smaller worms, with their maximum velocity being influenced by substratum type. Their estimated rate of predation is 0.092 prey items nemertean⁻¹ day⁻¹, just below the laboratory rate of ~0.2 amphipods nemertean⁻¹ day⁻¹ previously estimated for this species. P. nelsoni consumes a diverse spectrum of prey items (i.e., amphipods, isopods, decapods, barnacles, and dipterans) and is possibly exerting a significant influence on its prey populations. We suggest that the opportunistic predatory behavior of this intertidal predator is caused by the trade-off between immediate persistence (e.g., avoidance of desiccation) and long-term survival through successful foraging.     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Are barnacle larvae able to escape from the threat of UV?

Earlier laboratory experiments suggested that environmental levels of UV-B radiation can damage the eyes of barnacle naupliar larvae and impair their phototactic behaviors. However, since barnacle larvae may avoid UV by migrating to deeper waters, it is not known whether such impairment would actually occur under field conditions. For the first time, this study provides both field and laboratory evidences to show that prevailing UV-B in the natural habitat of barnacle larvae could be an important environmental factor affecting natural barnacle populations. We here showed that although barnacle nauplii may avoid UV-B irradiation by downward migration, the amount of UV energy (9.8 × 10⁻⁶ J) received by a naupliar eye during downward migration in the natural water column is within the same order of magnitude as the total energy (7.5 × 10⁻⁶ J) sufficient to cause damages to naupliar eye and impair their phototactic responses. It is possible that solar UV-B prevailing at shallow waters would pose a similar threat to other zooplankton species over large geographic scale.     

Castrating effects of trematode larvae on the reproductive success of a highly parasitized population of Crepipatella dilatata (Caenogastropoda) in Argentina

Crepipatella dilatata (Calyptraeidae) is a common intertidal or shallow subtidal protandric hermaphroditic gastropod. Intertidal snails from Puerto Madryn, Chubut, Argentina (42°46′37″S, 64°54′51″ W), showed a high prevalence, up to 46?%, of a castrating larval microphallid trematode. Reproductive success was studied during a 2-year period from September 2007 to August 2009, and the results were compared to those from an unparasitized subtidal sample. Brooding females and males were never found infected. The proportion of brooding females was higher in months with a higher prevalence of infection. The number of brooding females in the intertidal sample was higher than in the subtidal sample. Generalized linear models were utilized to determine the effects of parasitism on the reproductive success of the host population. Anticipated reproduction in smaller females in the intertidal sample, increased reproductive success in months with higher prevalence, and unparasitized males are interpreted as fitness responses to ensure host population survival when the prevalence of trematode infection is high.     

Physiological tolerances across latitudes: thermal sensitivity of larval marine snails (Nucella spp.)

A critical step in understanding how temperature will affect biodiversity in coastal ecosystems is to gain insight into how the tolerances, and ultimately survival, of early life history stages will influence the distribution and abundance of adults. We assessed the thermal tolerance of encapsulated veliger-stage larvae of a common dogwhelk, Nucella ostrina, that occur in the rocky intertidal zone on the west coast of North America. Results showed that veligers collected from northern latitudes in Washington State were less tolerant of heat stress than those from central sites in California. For all sites, we found there to be a subtle difference between the temperatures at which veligers first began to die compared to when veligers reached 100% mortality. On a biogeographic scale, the LT₅₀ temperatures, a measure of larval sensitivity, for N. ostrina veligers displayed a strong latitudinal trend. These findings provide a conservative measurement of the upper thermal limits of encapsulated veligers while illustrating how these early life history stages could be physiologically compromised under future climate warming scenarios.     

Development of the gooseneck barnacle Pollicipes polymerus (Cirripedia: Lepadomorpha): Fertilization through settlement

Pollicipes polymerus (Sowerby, 1833) is a common barnacle in rocky intertidal zones of the West Coast of North America whose development has been previously undescribed in detail. In the present study, certain aspects of the development of P. polymerus are recorded from fertilization through settlement. In vitro fertilization has been accomplished using already-implanted sperm plus ovulating eggs and oviducal gland fluid. The eggs are small (about 100 μ diameter at fertilization; 90 x 140 μ at first cleavage) and somewhat yolky. Embryos obtained from brooding adults and from in vitro fertilization were reared in glass dishes until hatching. Cleavage was total, asynchronous and unequal. Division of the D macromere lagged behind division of the other blastomeres, and gastrulation occurred by epiboly so that blastoderm surrounded the macromeres. The nauplii hatched an average of 25.4 days (range was 20.3 to 30.1 days) after fertilization, and molted within 1 day to Stage II nauplii. From Stages II through VI the nauplii required food. As they grew from stage to stage, the number of setae per appendage increased. Nauplii have feathery and hispid-type setae, previously described only in Chthamalus aestuarii. In contrast to other pedunculate larvae which have been described, P. polymerus nauplii are small and have specialized feathery setae. Cypris-larva settlement was stimulated only when healthy adult peduncles were available, thus showing an affinity for its own species.     

Biochemical genetic variation and evidence of speciation in Chthamalus barnacles of the tropical eastern Pacific Ocean

Gene-enzyme variation was studied electrophoretically within and between barnacle populations of the genus Chthamalus from 8 intertidal stations from central California to the Pacific coast of Panamá. Horizontal starch-gel electrophoresis separated and resolved 18 enzymes from 512 individual barnacles. A maximum of 25 gene-enzyme systems was interpretable from the resulting zymograms. Electrophoretic phenotypes and patterns of phenotypic variation generally conformed with those observed in other organisms. The amount of genetic variation within barnacle populations varied; average heterozygosity, for example, ranged from a low of 3.4% in a Mexican population of the C. fissus group to a high of 10.4% in C. anisopoma from the Gulf of California. Observed and expected average heterozygosities agreed in all population samples, indicating that these species are outbreeding. In contrast to the prediction stemming from a hypothesis that trophic stability regulates the amount of genetic variation in marine species, average heterozygosity tends to be positively correlated with latitude. Data from these and other barnacle species may support a hybrid environmental heterogeneity-trophic diversity model recently proposed to explain genetic variation in decapod crustaceans. Juxtaposition of individuals from different localities revealed numerous genetic differences among populations of the C. fissus group. At least three partially sympatric sibling species are separated by genetic distances as large as those observed between the C. fissus group and the distinct species C. dalli and C. anisopoma. A cladistic analysis places C. anisopoma close to the Mexican and Panamanian sibling species, with C. fissus from San Diego and C. dalli successively farther removed.     

Mercury concentrations in the surface sediments of the intertidal area along the west coast of Peninsular Malaysia

Total mercury (Hg) levels in the intertidal surface sediments along the west coast of Peninsular Malaysia were analysed by using heat vaporization method. Total Hg levels in these sediments ranged from 3.00 to 201?µg?kg^?1 dry weight with a mean concentration of 60.0?µg?kg^?1 dry weight. More than 90% of the samples studied have the total Hg concentrations less than 150?µg?kg^?1 dry weight. Compare with the regional data and sediment quality guidelines, the Hg contamination in the intertidal area along the west coast of Peninsular Malaysia was not serious, except for a few sites that might have received anthropogenic Hg in the samples collected between 1999 and 2000. It is suggested that a similar study of heavy metals in the sediments of the west coast of Peninsular Malaysia be conducted regularly.