Zinc phosphate granules in tissue surrounding the midgut of the barnacle Balanus balanoides

The chemical composition of inorganic granules found in parenchyma cells surrounding the midgut of adult Balanus balanoides (L.) was determined. X-ray microanalysis indicated the presence of magnesium, phosphorus, potassium, calcium, iron and zinc in the granules. A quantitative analysis of granule-rich pellets isolated from the midguts of adult barnacles showed that the granules were composed mainly of zinc phosphate, the remaining elements being minor constituents only.     

Importance of physical and biological settlement cues used at different spatial scales by the larvae of Semibalanus balanoides

The settling cues used by larvae of Semibalanus balanoides (L.) were examined at large (1 m), medium (1 mm), and small (<300 m) spatial scales, corresponding roughly to choices made during broad and close exploration and final inspection within the settling zone. The experiments were carried out at two locations in Canada in May and June 1984. Samples of substratum surfaces from above (n=87 sites unselected by cyprids) and below (n=84 sites selected by cyprids) the upper limit of the barnacle zone were characterized according to 13 quantitative and semi-quantitative variables (diatom cover, macroalgae, detritus) using scanning electron microscopy (SEM). The data were analysed using a stepwise logistic regression. At the large scale, the presence of an alga (Urospora wormskjoldii Mertens) in the upper uncolonized zone and a diatom (Achnantes parvula Kützing) within the settling zone are potential settlement cues. At the medium scale in the settling zone, larvae select clean sites, devoid of detritus and diatoms or on which detrital matter, when present, is of finer texture than on the unchosen sites within the settling zone. At the smallest scale (<300 m, equal to the average maximum width of the larvae), we tested the hypothesis that the larva seeks optimal microheterogeneity (presumably providing good adhesion). Microheterogeneity of the surface was measured immediately under the larva, around the settling organs, the antennules. Our results show that such surface microheterogeneity is significantly greater and the vertical center of gravity of the surface significantly lower on the sites selected by the cyprids than on the unselected contiguous sites. Fractal measurements carried out on selected and contiguous unselected surfaces indicate that larvae discriminate microheterogeneity levels below a step length of 35 m, a value approximately the size of the diameter of the antennular discs (40 m), which are used to explore the substratum and attach.Contribution to the programme of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Fatty acid composition of the triacylglycerols of the cypris larva of the barnacle Balanus balanoides during metamorphosis

In the free-swimming cypris larva of Balanus balanoides, the mean level of triacylglycerols (triglycerides) was 8% of the ash-free dry weight. During metamorphosis into the juvenile barnacle, the triacylglycerol level in the newly settled non-feeding cypris larva fell to a mean of 2% of the ash-free dry weight. The ratio of fatty acids in the triacylglycerols remained constant throughout metamorphosis, indicating that all species of fatty acids were used as energy sources. There was no evidence for the selective utilisation of particular fatty acids.     

Participation of the C-type hemolymph lectin in mineralization of the acorn barnacle Megabalanus rosa

BRA-2, a major C-type lectin in the hemolymph of the acorn barnacle Megabalanus rosa was detected in shell-associated proteins. Immobilized BRA-2 promoted CaCO₃ crystallization as determined by an in vitro pH-drift mineralization assay. In nature, calcification occurs as wound repair in M. rosa after breakage of the shell plate. To observe this phenomenon, we inserted a rubber cube inside the mantle cavity of M. rosa through the basal plate, and kept the organisms in a tank for several weeks. The rubber cube was covered first with brownish-colored material, probably a melanin, and then with CaCO₃ 5 weeks after the insertion. In the CaCO₃-associated proteins, BRA-3, a minor component of M. rosa hemolymph lectins, was present in addition to BRA-2. The presence of both the lectins and their ligands on the hemocyte was also observed. These results suggest that the C-type lectins of the M. rosa hemolymph participate in mineralization as well as defense in these organisms.     

Seasonal changes in the free glycerol level in the body parts of the adult barnacle Balanus balanoides

Seasonal changes in the free glycerol level in body parts of the adult barnacle Balanus balanoides have been followed during the 1969 season. Glycerol levels were highest in winter when the barnacles were most cold-tolerant and lowest in summer when they were least cold-tolerant. It is suggested that glycerol, and possibly other solutes, may be concentrated in the blood of B. balanoides, thus promoting supercooling and reducing the amount of ice formed at sub-zero temperatures.     

Population structure and phylogeography of an acorn barnacle with induced defense and its gastropod predator in the Gulf of California

Using sequence variation in the mitochondrial cytochrome oxidase I locus, we estimated the population structure and phylogeography of the intertidal acorn barnacle, Chthamalus anisopoma, and its gastropod predator, Mexacanthina lugubris angelica. Both are endemic to the Gulf of California, being derived from taxa on the Pacific coast of the Baja peninsula, and both exhibit phenotypic plasticity for traits affecting their coevolutionary interactions. Consistent with expectations based on differences in dispersal, C. anisopoma populations generally lack geographic structure, while those of M. l. angelica are more strongly structured. However, the variable degree of differentiation in both species suggests that the extent of reciprocal selection and local adaptation in the species will vary geographically, a result consistent with the concept of a geographic mosaic of coevolution. The pattern of variation in C. anisopoma shows clear evidence of recent spatial expansion, possibly due to increased habitat availability following the last glacial maximum. Phylogeographic analyses suggest that M. l. angelica diversified into three distinct clades after the colonization of the Gulf. Overall, our results illustrate how dispersal potential, geological and climatic events, and recent population growth have impacted the pattern of sequence variation in the two species.     

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.     

Effects of salinity and adult extract on settlement of the oligohaline barnacle Balanus subalbidus

Balanus subalbidus (Henry) has the most oligohaline distribution of three congeneric barnacles in Chesapeake Bay and tolerates prolonged exposure to fresh water. We studied larval settlement (i.e., permanent attachment and metamorphosis) of B. subaldius in the laboratory, over a 3 yr period, May 1989 to March 1992, under the following conditions: (1) across an array of salinities at 25°C in the presence and absence of settlement factor consisting of adult B. subalbidus extract; (2) in the presence of conspecific or congeneric settlement factors; and (3) cyprids which were, and were not, induced to delay metamorphosis were compared in their capacities to settle in a range of salinities. Discrepancies between salinity profiles of larval settlement in the laboratory and adult oligohaline distribution in the estuary were striking, and there was a significant interaction between salinity and settlement factor. Averaging results of four different batches of larvae, although peak settlement (87±9%) of B. subalbidus occurred at 2 ppt salinity in the presence of adult cue, substantial settlement also occurred at higher salinities: >70% at 5, 10 and 15 ppt; and 47% at 20 and 25 ppt. In addition, settlement in the absence of settlement factor was relatively high (>50%) and peaked at mid-salinity ranges (e.g. 56±10% at 15 ppt). Variation observed in settlement among larval batches reflected detailed differences in settlement between adjacent test salinities. No difference in settlement occurred between replicate aliquots of cyprids within a batch. Cyprids of B. subalbidus settled most abundantly in the presence of settlement factor extracted from conspecifics, followed in decreasing order by settlement factor extracted from B. improvisus and B. eburneus. Delay of metamorphosis produced by keeping B. subalbidus cyprids for 8 d at 5°C resulted in a decreased level of settlement, but settlement frequency patterns of delayed and non-delayed cyprids were indistinguishable relative to salinity. These results indicate that the oligohaline distribution of adult B. subalbidus is probably not determined by larval behavior at settlement. We suggest that pre-settlement behavior, resulting in larval retention in low saline waters, could be an important factor in determining distribution of this species.     

Effects of crowding due to habitat loss on species assemblage patterns

Terrestrial animals are negatively affected by habitat loss, which is assessed on a landscape scale, whereas secondary effects of habitat loss, such as crowding, are usually disregarded. Such impacts are inherently hard to address and poorly understood, and there is a growing concern that they could have dire consequences. We sampled birds throughout a deforestation process to assess crowding stress in an adjacent habitat remnant in the southern Brazilian Atlantic Forest. Crowding is expected of highly mobile taxa, especially given the microhabitat heterogeneity of Neotropical forests, and we hypothesized that the arrival of new individuals or species in refuges shifts assemblage patterns. We used point counts to obtain bird abundances in a before-after-control-impact design sampling of a deforestation event. Temporal changes in taxonomic and functional diversity were examined with metrics used to assess alpha and beta diversity, turnover of taxonomic and functional similarity, and taxonomic and functional composition. Over time increased abundance of some species altered the Simpson index and affected the abundance-distribution of traits in the habitat remnant. Taxonomic composition and functional composition changed in the remnant, and thus bird assemblages changed over time. Taxonomic and functional metrics indicated that fugitives affected resident assemblages in refuges, and effects endured >2 years after the deforestation processes had ceased. Dissimilarity of taxonomic composition between pre- and postdeforestation assemblages increased, whereas functional composition reverted to preimpact conditions. We found that ecological disruptions resulted from crowding and escalated into disruptions of species’ assemblages and potentially compromising ecosystem functioning. It is important to consider crowding effects of highly mobile taxa during impact assessments, especially in large-scale infrastructure projects that may affect larger areas than is assumed.     

Effects of temperature and food availability on growth and reproduction in the neustonic pedunculate barnacle Lepas anserifera

To elucidate the life history of neustonic animals, growth and reproductive patterns were investigated in the hermaphroditic pedunculate barnacle Lepas anserifera in field and laboratory experiments in Wakayama, western Japan from 2006 to 2008. The effects of temperature (19, 24 or 29°C) and food availability (once or twice a week) on growth and reproduction were also studied in the laboratory. The barnacles grew and matured rapidly, especially in the field: individuals on the average grew from 3 mm to more than 12 mm in capitulum length within 15 days and some were brooding. High temperature and high food availability resulted in greater growth. High temperature also resulted in earlier maturation of both testes and ovaries, whereas the effect of food availability was less clear. The rapid growth and maturation, together with earlier maturation at higher temperatures, may be an adaptation to ephemeral floating objects to which they attach.     

Allelic esterase isozymes,their variation with season,position on the shore and stage of development in the cirripede Balanus balanoides

Samples of Balanus balanoides (L.) in the Menai Straits were taken from their upper limit of distribution on the shore, from mean low water (MLW) and at three different times of the year, January, May and August. Individual prosomae were homogenised, electrophoresed on polyacrylamide gels and esterase isozymes determined. Group BbEI esterase isozymes fitted the Hardy-Weinberg predicted values for 4 co-dominant alleles in all samples. A comparison of BbEI phenotype frequency distribution of the upper shore and MLW samples showed no significant difference at the 5% level. The phenotype frequency of samples taken at different times of the year were significantly different at the 5% level, due to an excess of allele A in winter and an excess of allele C in summer. It is argued that the seasonal variation of phenotype frequency is probably not due to selection operating on a balanced polymorphism, but that the expression of the alleles through the synthesis of esterase isozymes is dependent on some environmental agent, postibly temperature or day length.     

Variability of shell growth and morphology of the wall-plate junctions of the intertidal barnacle Tesseropora rosea (Cirripedia: Tetraclitidae)

Variability in shell growth and morphology of the wallplate junctions of Tesseropora rosea (Krauss) were studied in 1983 in New South Wales, Australia, at two sites with differing wave exposure. The wall-plate junctions, a combination of dado (geometrically interlocking) and lap bints, increase in complexity with age, but do not differ between sites. The number of parietal tubes is fixed at settlement and does not differ between sites, but differs among wall plates in a manner consistent with their evolutionary history (rostrum > laterals > carina). The number of secondary septa increases with age, but does not differ between sites. In epilithic T. rosea, growth of the alae contributes to orifice enlargement. Epizoic T. rosea (on limpets) have conspicuous radii and display diametric growth. At the more exposed site, the shell wall was sleeper, and the aperture length, height and weight of the shell increased at a faster rate than at the less exposed site, but the maximum height of the shell was less. These differences appear to be due to a combination of greater feeding rates and greater erosion rates at the exposed site. The greater maximum height attained at the more sheltered site may be due to greater individual longevity. Juveniles at the sheltered site had thinner septa in the laterals than those at the exposed site. Septa in adults were rhinner at the exposed site, but this is due to age differences between sites in the adult populations.     

Testing the effects of wave exposure,site, and behavior on intertidal mussel body temperatures: applications and limits of temperature logger design

The rocky intertidal mussel Mytilus californianus is exposed to potentially damaging thermal conditions during low tide. However, because the temperatures of ectothermic organisms are driven by multiple climatic factors, we do not fully understand what the body temperatures of intertidal invertebrates are under field conditions, or how thermal stress varies between intertidal sites. We designed a temperature logger that thermally matches (similar size, color, morphology and thermal inertia) living mussels of the species M. californianus, and tested its ability to provide realistic measurements of body temperature in the field. As part of these tests, we examined the propensity of M. californianus to gape, a behavior in which the mussel opens its shell valves, and which may permit evaporative cooling. Because our instruments were unable to mimic this behavior, we tested the degree to which gaping contributes to animal cooling by exposing M. californianus to a range of climatic conditions while recording body temperatures, gaping behavior and water loss. Results indicated no significant influence of gaping on body temperature. Tests comparing temperatures of loggers to those of real mussels under laboratory and field conditions showed that thermally matched loggers recorded temperatures within a few degrees of living animals and that unmodified loggers regularly incurred errors of up to 14°C. We then deployed a series of thermally matched loggers at two sites in central Oregon (Boiler Bay and Strawberry Hill) previously hypothesized to display site-specific differences in aerial temperature, and at various wave-exposure regimes within each site. Significant differences were demonstrated between sites using a subset of temperature metrics in a multivariate analysis. Yearly peaks in maximum temperature, average daily maximum temperature, and degree hours were useful in discriminating between sites. In 2001, wave-exposed sites at Strawberry Hill displayed fewer degree–hours than wave-protected sites, but an equivalent or greater maximum temperature. In 2002 both of these metrics were significantly lower in wave-exposed sites. Boiler Bay and Strawberry Hill differed in thermal regime, but not in a way that would indicate one was hotter than the other.Communicated by P.W. Sammarco, Chauvin     

Effects of diet,ultraviolet exposure,and gender on the ultraviolet absorbance of fish mucus and ocular structures

Ultraviolet (UV) radiation can be damaging to fish skin and ocular components. Coral reef fishes are regularly exposed to potentially harmful radiation. It was recently discovered that tropical marine fishes possess UV-absorbing compounds in their mucus. This experiment demonstrates significant effects of both diet and ultraviolet exposure on the UV-absorbing compounds in the mucus of a tropical wrasse, Thalassoma duperrey. Fish that are exposed to UV radiation increase the UV absorbance of their mucus only if UV-absorbing compounds are provided in their diet. Fish that are protected from UV radiation decrease the UV absorbance of their mucus regardless of diet. Mucus from female T. duperrey absorbed less UV and females had higher rates of skin damage than males. Females sequester UV-absorbing compounds in their pelagic eggs as well as their epithelial mucus, whereas males do not sequester these compounds in the testes. Spectral transmission through the whole eye was not affected by diet or UV manipulations, but corneal tissue transmission decreased significantly in the UV-exposed individuals. These results demonstrate that coral reef fish can adapt to UV exposure, so long as UV-absorbing compounds are available in the diet.Communicated by P.W. Sammarco, Chauvin     

The histology,histochemistry and ultrastructure of the cement apparatus of three adult sessile barnacles,Elminius modestus,Balanus balanoides and Balanus hameri

The histology, histochemistry and ultrastructure of the cement gland cells of 3 species of sessile barnacles are described. Cells of both Elminius modestus Darwin and Balanus balanoides (L.) (membranous bases) are similar histologically and ultrastructurally, and within the cells there are 2 distinct regions. One is the site of secretion accumulation, the other the site of secretion synthesis. In Balanus hameri Ascanius (calcareous base) the cells are different in that the cytoplasm is more homogeneous. In all 3 species protein is the final secretion product (cement) of the cement cells. The mode of discharge of secretion from the cells differs between the membrane based and calcareous based barnacles. In the former, there is a series of collecting canals within the cytoplasm of the cement cells (intracellular canals) which join with the larger extracellular cement ducts. Secretion passes into the intracellular canals and is moved along to the larger cement ducts, which have an inner chitin lining. In Balanus hameri (calcareous base) there are no intracellular collecting canals. Secretions are thought to pass from the cement cells into the cement duct cells direct. The cement ducts in this species are not chitin lined.     

Interactive effects of wave exposure and tidal height on population structure of the mussel Perna perna

The influence of wave exposure and of tidal height on mussel (Perna perna Linnaeus) population structure (size, density, biomass and adult/juvenile correlations) was examined at 18 sites along the south coast of South Africa. Sites were classified as exposed or sheltered prior to sampling, without reference to the biota, on the basis of aspect, topography and wave regime. A single set of samples was collected from each site during three spring tide cycles. Adult mussels on these shores almost always attach directly to the rocks, and layering of mussels is virtually absent. Shore height always had a strong influence on population structure, but exposure had significant effects only lower on the shore, and almost exclusively on mussel sizes. Principal component analysis (PCA), based on size distribution data for each population, revealed a general upshore decrease in the modal size of the adult cohort. The effects of exposure on size distribution, however, varied with tidal height. PCA separated exposed zones, with larger mussels, from sheltered zones on the low-shore. Farther upshore the two shore types were increasingly confounded. The maximum size of mussels showed a similar pattern, with significant differences (ANOVA, p < 0.05) between exposed and sheltered sites only on the low- and mid-shores. Density was calculated from randomly placed quadrats (i.e. not necessarily from areas of 100% cover) and showed a different pattern. Adult (>15 mm) densities decreased up the shore, with low-, mid- and high-shore zones being significantly different from one another (ANOVA, p < 0.0001; followed by multiple range tests). However, exposure had no significant effect on density, nor was there a significant interaction with zone. Recruit (<15 mm) densities were positively correlated with adult (>15 mm) densities in all zones and for both exposure regimes ( p < 0.05 in all cases), but there was considerable variability and extremely low predictability in these relationships (r ² generally <0.2). Predictability tended to be greater towards the high-shore, where adults were more clumped. As with density, biomass was not affected by exposure, but decreased upshore as mean size and density decreased. A reduction in the influence of exposure farther upshore may be caused by greater emersion overriding the effects of exposure. The presence of free space within mussel beds and significant correlations between recruit and adult densities suggest that these mussel populations are recruit limited. Received: 7 January 2000 / Accepted: 6 July 2000     

Material interests, moral reputation, and crowding out species protection on private land

We consider how the reputation of being socially responsible works as an effective source of motivation in protecting a public good, such as endangered species. This paper investigates the mechanism design for endangered species protection on private land under asymmetric information about reputation and land quality. We examine optimal monetary transfer by designing an efficient mechanism which takes into account the crowding out effect of monetary rewards on socially responsible behavior. We find landowners who have good reputation contribute more than the optimum level. Landowners with poor reputation sacrifices information rent; rather they buy reputation.     

Abundance,population structure and claw morphology of the semi-terrestrial crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis> (Fabricius, 1787) on shores of differing wave exposure

Wave action is known to influence the abundance and distribution of intertidal organisms. Wave action will also determine the duration and suitability of various foraging windows (high-tide and low-tide, day and night) for predation and can also affect predator behaviour, both directly by impeding prey handling and indirectly by influencing prey abundance. It remains uncertain whether semi-terrestrial mobile predators such as crabs which can access intertidal prey during emersion when the effects of wave action are minimal, are influenced by exposure. Here, we assessed the effect of wave action on the abundance and population structure (size and gender) of the semi-terrestrial intertidal crab Pachygrapsus marmoratus on rocky shores in Portugal. The activity of P. marmoratus with the tidal cycle on sheltered and exposed shores was established using baited pots at high-tide to examine whether there was activity during intertidal immersion and by low-tide searches. Because prey abundance varies along a wave exposure gradient on most Portuguese shores and because morphology of crab chelipeds are known to be related to diet composition, we further tested the hypothesis that predator stomach contents reflected differences in prey abundance along the horizontal gradient in wave exposure and that this would be correlated with the crab cheliped morphology. Thus, we examined phenotypic variation in P. marmoratus chelipeds across shores of differing exposure to wave action. P. marmoratus was only active during low-tide. Patterns of abundance and population structure of crabs did not vary with exposure to wave action. Stomach contents, however, varied significantly between shores of differing exposure with a higher consumption of hard-shelled prey (mussels) on exposed locations, where this type of prey is more abundant, and a higher consumption of barnacles on sheltered shores. Multivariate geometric analysis of crab claws showed that claws were significantly larger on exposed shores. There was a significant correlation between animals with larger claws and the abundance of mussels in their stomach. Variation in cheliped size may have resulted from differing food availability on sheltered and exposed shores.     

Markets and the crowding out of conservation-relevant behavior

Markets are increasingly being incorporated into many aspects of daily life and are becoming an important part of the conservation solution space. Although market-based solutions to environmental problems can result in improvements to conservation, a body of social science research highlights how markets may also have unforeseen consequences by crowding out or displacing 3 key types of behaviors potentially relevant to conservation, including people's willingness to engage in collective action and civic duty; tolerance for inflicting harm on others (third-party externalities); and desire for equity. Better understanding of the contexts and mechanisms through which this crowding out occurs and whether specific market-based instruments are more prone to different types of crowding out will be crucial to developing novel conservation initiatives that can reduce or prevent crowding out.