Shell-size selection by intertidal sympatric hermit crabs

This study evaluated selection for shell size by three species of tropical intertidal hermit crabs, Clibanarius antillensis, C. sclopetarius, and C. vittatus, from species of shells which are frequently used in nature. Crab size and weight were strongly and significantly related to all measured parameters of the selected shells. The strength of these relationships (r² values) depended neither on the crab nor on the shell variables taken into account. The relationships between crab size and the dimensions of the selected shells showed higher r² values than the corresponding relationships with the shells that the crabs had occupied when they were collected (0.482–0.903 in comparison to 0.091–0.652, respectively), indicating that the crabs were occupying sub-optimal shells in nature. Negative allometry was frequently found in the relationships between crab and shell variables, indicating that large crabs select and use proportionally lighter shells than do small crabs. This negative allometry was stronger for the shells used in nature (except for C. antillensis), i.e. larger crabs tended to select heavier shells in the laboratory than in nature. Different allometric relationships were also recorded among the dimensions of shells used in nature and those selected by the hermit crabs in free-access experiments: as shell length increased, the selected shells were heavier and had larger apertures than the shells used in nature. The relationships between crab size and the length and weight of the selected shells did not depend on the species of crab or species of shell, but only on crab size. Therefore, analyses using these variables can be performed without taking the species of crab or shell into account, i.e. data from different crab or shell species can be pooled. The influence of crab and/or shell species was recorded only in the models fitted for aperture length and width, variables which were more related to shell architecture than did shell length or weight. In contrast, if crab weight is used as an independent variable, different crab or shell species can be analyzed together independently of the particular shell parameter. This indicates that crab weight may be less susceptible than crab shield length to shell morphological constraints. Finally, the results indicate that the preferred size of a given shell type chosen by a given hermit crab will depend more on crab size or weight, than on the crab or shell species under consideration, i.e. crab shell-size relationships are not species specific.Communicated by P.W. Sammarco, Chauvin     

The chromosomes of Trypetesa lampas (Cirripedia,Acrothoracica)

Observations were made of the chromosomes of the burrowing barnacle Trypetesa lampas (Hancock). A method of squash preparation was used, which incorporated staining the material in a solution of 2% orcein in 45% acetic acid. The diploid number of chromosomes of T. lampas was found to be 12 and the haploid number 6, in both males and females. No obvious chromosomal mechanism of sex determination was found. There was, therefore, no cytogenetic confirmation of Kühnert's view (1934) that the sex of the larva is predetermined genetically. During mitosis in the females and embryos of Trypetesa lateralis Tomlinson and Kochlorine floridana Wells and Tomlinson, the diploid number of chromosomes was found to be about 14. The numbers of chromosomes in the 3 species of Acrothoracica studied were approximately half those observed in all 12 species of Thoracica and all 5 species of Rhizocephala previously investigated.     

Morphologie,Entwicklung und Ökologie digener Trematoden-Larven der StrandschneckeLittorina littorea

More than 6000 individuals of the snailLittorina littorea L., from 6 different localities on the coast of the German Bay (North Sea), were examined for larval trematodes by crushing their shells, an additional 6000 by isolation in small quantities of sea water. Six species of larval trematodes were found inL. littorea and described: Notocotylid larva —Cercaria lebouri Stunkard, 1932; Cercaria ofHimasthla elongata (Mehlis, 1831); Cercaria ofPodocotyle atomon (Rudolphi, 1802); Cercaria ofRenicola roscovita (Stunkard, 1932) n. comb.; Metacercaria ofMicrophallus pygmaeus (Levinsen, 1881); Cercaria ofCryptocotyle lingua (Creplin, 1825). Considering the large number of investigated snails, the trematode fauna ofL. littorea in this area is assumed to be completely known. The life cycles ofHimasthla elongata andRenicola roscovita are described. Larvae of the different trematode species show differential preferences for distinct size groups of snail hosts. Juvenile snails are not infected. No correlation exists between infection rates and sex of the hosts. Quantitative aspects of multiple infections are calculated and discussed. The incidence of larval trematodes in the investigated localities differs. Information is presented on seasonal variations of infestations and their possible causes.     

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.     

Large-scale patterns of shell variation in Littorina striata, a planktonic developing periwinkle from Macaronesia (Mollusca: Prosobranchia)

Littorina striata King and Broderip, 1832 is a strictly Macaronesian, intertidal periwinkle with planktonic development. The species displays a high degree of shell variation involving size and sculpture (nodulose vs smooth shells). The present work provides a preliminary account of some aspects of this shell variation on wave-exposed shores over the entire geographical range of the species. Based on morphological patterns observed among other prosobranchs it was predicted that southern specimens of L. littorina should on the average be larger, heavier, more nodulose, and should show more shell repair marks, than northern specimens. These expectations were confirmed for shell size and weight. In contrast, there was no consistent pattern in nodulosity between archipelagos, even though there were differences at much smaller scales. Shell repair marks were more prevalent in northern populations, but this trend was only due to a significant N–S difference among nodulose shells. This is surprising as nodulose shells displayed significantly fewer shell repair marks than smooth shells. These observations were tentatively interpreted as a function of presumed differential N–S patterns of wave action and ambient temperatures. In this context, wave action in Macaronesia seems to increase in the south (contrary to what current theories predict). This atypical situation may confound the interpretation of morphological patterns in L. striata so that firm conclusions cannot be drawn without further experimental work at different spatial scales. Nevertheless, it seems that generalisations about macrogeographic shell morphology patterns, based on interspecific comparisons, are not directly applicable to intraspecific patterns, and may strongly depend on local conditions which make adequate sampling and data treatment very difficult. Received: 10 September 1997 / Accepted: 15 January 1998     

Lethal and sub-lethal effects of phototrophic endoliths attacking the shell of the intertidal mussel Perna perna

Animals that bore into calcareous material can cause considerable damage to molluscan shells. In contrast, smaller microbial phototrophic endoliths have until recently been thought of as relatively benign. Phototrophic endoliths (primarily cyanobacteria) infest the shells of 50 to 80% of midshore populations of the mussel Perna perna (L.) in South Africa. This infestation causes clearly visible shell degradation, and we record here ecologically important lethal and sub-lethal effects (e.g. changes in growth and reproductive output) of the endoliths on their mussel hosts. Endolith infestation reduced the strength of shells significantly and also affected shell growth. In situ marking of shells, using the fluorochrome calcein, showed that infested and non-infested mussels increased in shell length at the same rate. However, the rate of increase in shell thickness (associated with shell repair) was significantly faster in infested than in uninfested individuals. This increase in the rate of shell thickening was not sufficient to compensate for rapid endolith-induced shell degradation and, around the site of adductor muscle attachment, infested shells were thinner than their uninfested counterparts. The shells of 18% of recently dead mussels had holes induced by endolith erosion. This effect was highly size dependent, and the proportion of mortality due to endoliths rose to almost 50% for the largest mussels. The re-routing of energy due to shell repair had important sub-lethal effects on the reproductive rates of mussels. During the reproductive period, mean dried flesh mass for large (>70 mm), non-infested P. perna was substantially higher than for infested individuals. This difference was almost entirely due to differences in gonad mass, which was approximately 100% higher for non-infested mussels. We conclude that, by attacking the shell, phototrophic endoliths reduce both the longevity and reproductive output of large mussels on the midshore. Received: 26 January 1999 / Accepted: 17 August 1999     

Precipitation of aragonite by calcitic bivalves in Mg-enriched marine waters

To understand the relative importance of biological versus physicochemical control over biomineralization, we have tested if the chemical composition of the medium (i.e., the Mg/Ca ratio) can change the mineralogy of mollusk shells. The shells of mollusks are made of calcite and/or aragonite, which are by far the most common CaCO₃ polymorphs. Several species of bivalves with predominantly calcitic shells have been cultivated in artificial seawater with a Mg/Ca molar ratio within the range of 8.3–9.2, well above the present value for seawater (5.2). Four out of six species used (the scallop Chlamys varia, the oyster Ostrea edulis, the saddle oyster Anomia ephippium and the mussel Mytilus edulis) survived long enough to secrete significant amounts of calcium carbonate. The deposits (sometimes extensive) formed on the interior shell surfaces were predominantly aragonitic. Three individuals of C. varia also increased their length by adding new shell at the margin. Contrary to the internal shell deposits, these margins were high-Mg calcite. This implies that the marginal mantle is able to exert a more strict control on the secreted mineral phase than the mantle facing the internal shell surface. This is the first report on an in vivo experimentally forced switch in bivalve shell mineralogy, from calcite to aragonite due to a change in water chemistry.     

Influence of occupant microhabitat on the composition of encrusting communities on gastropod shells

The exploitation of microhabitats is widely considered to increase biodiversity in marine ecosystems. Although intertidal hermit crabs and gastropods may inhabit the same shell type and shore level, their microhabitat may differ depending on the state of the tide. On the south coast of Wales the hermit crab Pagurus bernhardus mainly inhabits the shells of Nucella lapillus (84%). Hermit crab shells had a significantly different encrusting community compared with live N. lapillus shells. At low tide the live gastropods were found on exposed rock surfaces whereas hermit crabs were restricted to tidal pools. Communities encrusting live gastropod shells were characterised by lower species richness and abundance compared with shells inhabited by hermit crabs (12 species found in total). A greater abundance and richness of epibionts was recorded from both shell types during the summer compared with winter. Differences in community composition between shell occupant types were attributed to microhabitats used by gastropods and hermit crabs and the associated desiccation pressures, rather than competitive interactions or shell characteristics. This contradicts earlier studies of subtidal shells where biological processes were considered more important than physical factors in controlling species abundance and richness patterns. The use of rockpool microhabitats by hermit crabs increases the biodiversity of rocky shores, as some species commonly found on hermit-crab-inhabited shells were rare in other local habitats.Communicated by J.P. Thorpe, Port Erin     

Incidence and distribution of phototrophic shell-degrading endoliths of the brown mussel Perna perna

The incidence, distribution and infestation sequences of four endolithic cyanobacteria in the shells of the brown mussel Perna perna (L.) were studied along the south coast of South Africa. The incidence of endolith-infested shells varied significantly among the 21 study sites (∼23 to 95%), with highest infestation rates occurring on promontories and headlands as compared to sites within bays. At a smaller scale, the incidence of infested shells also varied with tidal height, being high at the upper tidal levels of mussel distribution and low or absent on the lowshore. The observation that small-scale variability in infestation rates was positively related to water movement suggested that physical damage to the outer protective periostracum of mussels may have facilitated colonisation by endoliths. This hypothesis was supported by the fact that shells with artificially damaged periostraca became infested at a greater rate than did control shells. Once colonisation by the filamentous cyanobacteria Plectonema terebrans, Hyella caespitosa and Mastigocoleus testarum had taken place, endoliths spread throughout the shell, causing progressive shell deformation and damage. Only older shells that were infested by the late successional, cavity-forming Pleurocapsa sp., however, exhibited severe shell deformations, became brittle and eventually fractured near the structurally important site of adductor muscle attachment. Heterotrophic endoliths typically associated with shell degradation in previous studies were extremely rare and if present did not contribute to shell disintegration. Although it is generally perceived that endolithic algae and cyanobacteria are restricted to the surface layers of shells by light limitation within the substratum, it is clear from this study that the interaction of a combination of factors (i.e. erosion of the periostracum, successional sequence of colonists and mechanical properties of the shell) may result in phototrophic endoliths causing severe shell degradation and eventually mussel mortality. Received: 18 December 1998 / Accepted: 18 June 1999     

Observations on shell form and its ecological significance in thaisid gastropods of the genus Lepsiella in New Zealand

This paper is concerned with the ecological significance of variation in shell form within the thaisid gastropod genus Lepsiella in New Zealand. Shell form has been investigated by measurement of shell height and breadth, aperture length and width, the diameters of consecutive whorls, apical angle, shell weight, and shell capacity, although in many cases shell height and shell breadth could not be measured because of erosion. L.albomarginata has been studied intensively at 4 stations in the South Island, and L. scobina less intensively at 6 stations in the North and South Islands. Comparisons of pairs of characters between stations have been tested by regression analysis and analysis of covariance where appropriate. Shells of L. albomarginata are relatively taller and narrower, and have a thicker wall, at a very sheltered station (Hakahaka Bay) that at more wave-exposed stations. L. scobina (sensu stricto), characterised by the presence of spiral ribs on the shell, exhibits less striking but comparable differences in shell shape. In laboratory tests in a tidal tank the thicker-shelled L. albomarginata from a sheltered station (Hakahaka Bay in Port Underwood) was much better able to resist attack by the shore crab Hemigrapsus edwardsi than was L. albomarginata from a nearby wave-exposed station (Whites Bay, near Cape Campbell, South Island). L. scobina from both stations was resistant to attack. H. edwardsi abounds at sheltered stations, but is missing from wave-exposed rock reefs such as those at Whites Bay, so that the ability to survive encounters with shore crabs is ecologically important to L. albomarginata inhabiting sheltered stations. L. scobina occupies a lower zone on the shore, where it is probably liable to encounter other more powerful predators. Its spiral ribs probably strengthen the shell. We do not know to what extent differences in shell form and thickness depend on environmental factors, and to what extent they originate genetically. Thin shells are associated with an abundance of mussels (Mytilus edulis ssp. aoteanus or Modiolus neozelanicus). There is an interesting possibility that a scarcity of mussels or other food caused by superior nonspecific predators might result in the production of better-protected Lepsiella.     

Goniadella gracilis,a polychaete new to British seas

Polychaetes of the genus Goniadella Hartman were found in Liverpool Bay, England, in sand and sandy gravel below depths of 15 m. The worms had most of the characteristics of G. gracilis (Verrill), a species previously recorded only from eastern North America and South Africa.     

Consequences of shell-species preferences for female reproductive success in the hermit crab Pagurus bernhardus

Littoral hermit crabs, Pagurus bernhardus, show a strong preference for Littorina obtusata shells rather than those of Gibbula species. The fitness consequences, in terms of fecundity, for this shell preference is examined for female crabs. Females in the preferred species produced eggs earlier in the season, produced more eggs in the first brood, and produced a second brood more often than did females in the less preferred species. The smaller brood for Gibbula spp. was not a consequence of egg loss from the pleopods due to an unfavourable shape of shell, but rather reflected lower egg production. It is not clear, however, if this differential reproduction is due to direct costs of carrying an unfavourable shell, i.e. the shell impedes reproduction, or whether crabs compete aggressively for favoured shells so that only crabs of low quality inhabit lowquality shells.     

Some direct observations on the ecology and behaviour of the Norway lobster Nephrops norvegicus

A detailed investigation of a small area of sea bed occupied by the Norway lobster Nephrops norvegicus (L.) was carried out by diving and television observations at depths of 30 m in Loch Torridon, Scotland. The density of burrows was 1/2 m², but only a proportion of these were occupied by N. norvegicus. Although about 70% of the larger burrows were occupied by N. norvegicus, giving a density of 1 lobster/8 m², very few juveniles (carapace length less than 30 mm) were found in the area. Many of the small burrows were occupied by the gobiid fish Lesueurigobius friesii (Collett). There was evidence that N. norvegicus frequently change their burrows, and fighting for burrows was observed. N. norvegicus remain within their burrows during the day, emerge around sunset to forage for food during the night, and then return to their burrows at dawn. This, and other aspects of their burrowing behaviour, have a marked effect on the commercial trawl catches of N. norvegicus which show large seasonal and diurnal variations in size and sex composition.     

Oxygen and carbon isotope profiles from Nassa mutabilis shells (Gastropoda): accretion rates and biological behaviour

Six shells of the Gastropoda species Nassa mutabilis L. were collected in the Adriatic Sea at a depth of a few metres off the central Italian coast during 1996 and 1997. The oxygen and carbon isotope composition of spot samples collected from the shell apex to the top was measured according to well-established techniques. The results obtained allow the following conclusions: (1) the CaCO₃ of the N. mutabilis shells is essentially formed by aragonite as proved by X-ray diffraction measurements; (2) the CaCO₃ is precipitated under isotope equilibrium conditions or very close to equilibrium with the environmental water; (3) shell accretion takes place all year round, although the warm period is preferred since the accretion rates are higher during the warm half of the year and considerably reduced in winter; (4) the carbon isotope composition changes according to the age of the specimen, probably in relation to changes in its nutritional regimen; and (5) the alternation of warm and cold periods recorded by the shells allows the evaluation of their age and of the season of their birth and death. Received: 3 December 1998 / Accepted: 7 September 1999     

Desiccation as a factor in the intertidal zonation of barnacles

Four species of balanomorph barnacles, Balanus crenatus Brugière, B. balanoides (L.), Elminius modestus Darwin and Chthamalus stellatus (Poli), were studied to assess the susceptibility of intertidal barnacle species to desiccation. Known sized samples of barnacles were exposed to controlled desiccating conditions and subsequent survival and water loss were determined. It is clear that the ability to live high on the shore is dependent on a reduction of the overall permeability to water loss. Because of greater surface area to volume ratios, small stages are particularly prone to desiccation. In normal intertidal emersion periods, small stages of B. crenatus particularly, and also of B. balanoides and E. modestus which are similar in their desiccation resistance, would be susceptible to desiccation at normal temperatures and low humidities. Large barnacles would be more prone to death from high temperatures when the tide is out. The spat of C. stellatus, although surviving much longer than spat of larger dimensions of the other species, must also be prone to prolonged emersion conditions at high shore levels.     

Molecular phylogeny and evolution of the pelagic copepod genus Neocalanus (Crustacea: Copepoda)

Portions of the mitochondrial genome (ca. 4 kb), encoding three protein-coding (COI, ND4L, ND6) and two ribosomal RNA (srRNA, lrRNA) genes, were sequenced for all six currently recognized species, plus one form, of the pelagic calanoid copepod genus Neocalanus. In Neocalanus gracilis, the ND6 gene was not found in the sequenced portion of the mitochondrial genome. Unambiguously aligned sequences were subjected to Bayesian, maximum-likelihood, maximum-parsimony, and neighbor-joining analyses using Eucalanus bungii as an outgroup. The resultant tree topologies from these four methods were congruent, robust, and all nodes were supported by high bootstrap values and posterior probabilities of 92–100%. Two tropical and subtropical species (N. gracilis and N. robustior) occupied the most basal position, and a subantarctic (N. tonsus) and three subarctic Pacific species (N. cristatus, N. plumchrus, and N. flemingeri) diverged subsequently. Transequatorial dispersal of the ancestral population during glaciations is suggested for this pattern of speciation, in which sister clades exhibited antitropical distributions. Although the area of ocean is much broader in the subantarctic than the subarctic Pacific, a higher number of species occur in the subarctic Pacific (three) than the subantarctic (one). The possibility that marginal seas, such as Japan Sea and Okhotsk Sea, function as natal areas for the divergence of species is discussed.     

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.     

Shell parameters in relation to zonation in mediterranean limpets

The relation of shell form to habitat in the Mediterranean limpets Patella caerulea (L.) and P. lusitanica (Gmelin) was investigated. P. caerulea shells were significantly longer, broader, but lower in height than P. lusitanica shells. The regression of the ratio (length plus breadth)/2 on the height of the shell was found to characterize the two limpet populations, with a small degree of overlap reflecting the distribution of the two limpets on the shore.     

Morphological and habitat divergence in the intertidal limpet Patelloida pygmaea

In the intertidal limpet Patelloida pygmaea, two distinctive morphs, the forms pygmaea and conulus, have been recorded. The former possesses a flat elongated shell, and the latter has an extremely high round shell. It has been observed in the field that pygmaea is found on oyster shells Crassostrea gigas. The form conulus uses an unusual substrate for attachment. It is found on the living shells of the intertidal gastropod Batillaria cumingi. Although conulus is normally found only on shells of Batillaria, it can also be found on oyster shells when pygmaea and Batillaria shells are not present in nature. An electrophoretic analysis of allozymes showed that these two forms are reproductively isolated from each other and coexist without gene exchange on the same mudflat. Laboratory experiments showed that pygmaea prefers oyster shells and conulus prefers Batillaria shells as substrates for attachment when both oyster and Batillaria shells are present. The form pygmaea did not attach to Batillaria shells, even when only Batillaria shells were available. However, conulus also attached to oyster shells when Batillaria shells were not available. The proportion of individuals of conulus that attached to oyster shells decreased significantly when pygmaea was attaching to the oyster shells. These results suggest that pygmaea is ecologically more specialized to living on oyster shells than conulus.     

Influence of size,species and damage on shell selection by the hermit crabPagurus longicarpus

Shell preferences were examined inPagurus longicarpus Say, collected from Wakulla Beach, Florida, USA in 1987. Relative shell size was a more important shell characteristic in shell selection than either shell species or shell damage. Hermit crabs rarely selected relatively large shells but often selected relatively small shells over damaged shells. Preferences for a particular shell species were offset by varying the relative size or condition of the preferred shell species. Because specific environmental pressures are linked to particular shell characteristics, an assessment of the importance of various shell features may indicate which environmental pressures are ameliorated via the gastropod shell. Predation and desiccation are linked to relatively small shells and particular shell species; these stresses may be minimized by alternative behavioral mechanisms.