Morphological, reproductive, and genetic variability among three populations of Crucibulum quiriquinae (Gastropoda: Calyptraeidae) in northern Chile

Crucibulum quiriquinae (Lesson, 1830) is the only species of Crucibulum currently recognized in northern Chile. Recent analysis of three Crucibulum populations obtained in northern Chile demonstrates the existence of morphological, genetic, and reproductive differences among populations. Two populations present in Bahía Tongoy (30°15'S), one inhabiting the shells of the snail Turritella cingulata and the other shells of the pectinid Argopecten purpuratus, showed morphological differences. However, both had planktonic larval development and show low genetic divergence (D=0.002). A third population from Bahía La Herradura (29°58'S), which also inhabits the shells of T. cingulata, did not show morphological differences compared with its counterpart from Bahía Tongoy. However the Bahía La Herradura population had intracapsular development and metamorphosis, and a larger genetic distance (D=0.06) from both Tongoy populations. The results of the reproductive and genetic analyses strongly suggested that the two Tongoy populations, although showing morphological differences, are biologically the same species, Crucibulum quiriquinae, whereas the La Herradura population is a new species.     

Morphological,gravimetric, and biochemical changes in<Emphasis Type="Italic"> Crepidula fecunda</Emphasis> (Gastropoda: Calyptraeidae) egg capsule walls during embryonic development

Walls of egg capsules of the gastropod Crepidula fecunda Gallardo, 1979 were examined at different developmental stages during the period of intracapsular development of embryos. The weight, biochemical composition, and structural features (using scanning and transmission electron microscopy) of the capsule walls were examined at several intervals during development of the embryos to the hatching stage. Biochemical analyses were also carried out on the intracapsular fluid to identify the possible transfer of organic material from the capsule wall to the intracapsular fluid. The capsule wall is composed principally of organic matter, primarily protein (91%), plus minor lipid and carbohydrate components. The capsule wall consists of a thin, fibrous external layer, which overlies a thicker, spongy inner layer. The spongy layer has almost disappeared by the end of the developmental period, losing about 90% of its thickness. The 40% loss in weight of the capsule walls over the developmental period is due to loss of organic matter as protein. This suggests that the inner layer of the capsules dissolves and/or disintegrates as larval development advances. The levels of dissolved and/or particulate proteins in the intracapsular fluid are much higher than those typical of the seawater surrounding the capsules. This suggests that, as embryonic development proceeds, the inner capsule walls could potentially provide extra nutrients to the embryos.Communicated by J.P. Grassle, New Brunswick     

Changes in feeding mechanisms during early ontogeny in juveniles of <Emphasis Type="Italic">Crepidula fecunda</Emphasis> (Gastropoda,Calyptraeidae)

Feeding in early life stages of the sedentary snail Crepidula is effected both by the use of the radula and by filtering with the gill. The present study is a contribution to the knowledge of the mechanisms of feeding by early juvenile snails of Crepidula fecunda. Experimental observations were made on specimens of known ages as they were fed constant concentrations of microalgae in the presence of a primary biological film on a glass substrate. Feeding activity was filmed under the microscope, and images were digitized for the identification and quantification of feeding structures. A morphological analysis was made of the structures associated with this process. Results showed that the radula was functional beginning in 1-day-old juveniles, and the gill functioned in respiration. Gill function in filter-feeding began in juveniles of 9 days old when dorsal and ventral ciliation had developed, as well as the food pouch. The latter structure begins activity as soon as the gill starts food collection. Osphradia appeared simultaneous with development of the filter feeding capacity by the gill. In their earliest stages after metamorphosis, the young snails begin life by radular scraping of primary biofilms, gradually shifting to filter feeding as the gill developed a critical number of filaments and cilia.     

Variation in size at sex-change among natural populations of the protandrous hermaphrodite, <Emphasis Type="Italic">Crepidula fornicata</Emphasis> (Gastropoda,Calyptraeidae)

Protandrous hermaphrodites are predicted to change sex from male to female when relative reproductive fitness of females surpasses that of males. How size at sex transition varies with population, mating group and individual parameters was investigated for five populations of the protandrous hermaphrodite slipper snail, Crepidula fornicata. The populations varied for density, size distribution, average mating group size and sex ratio. Size at sex-change was correlated with the population sex ratio. Comparisons of multiple hypotheses revealed that variables predicting the sex of a snail vary among positions in the mating group. The variables included body size, the relative size of the snail sitting atop the focal snail and population density. Our data support the conclusions that size at sex-change (and by inference, the size at which one sex has relatively greater fitness) is not fixed for these hermaphrodites and that individual size, social conditions and population differences all influence variation in relative fitness.     

Aspects of the ecology and growth of an intertidal juvenile population of Concholepas concholepas (Mollusca: Gastropoda: Muricidae) at Las Cruces,Chile

The habitat, density and growth rate for an intertidal population of Concholepas concholepas (Bruguière, 1789) were studied at Las Cruces, Chile (Lat. 33°30′S; Long. 71°38′W) during 1977–1978. The growth rate (3.67 mm month^-1) was determined in a newly settled group, whose average length was 11.3 mm, and whose age was estimated at 3 months. The densities found ranged between 1.1 and 107.3 “locos” m^-2. Based on these results, the time of settlement of C. concholepas was calculated; capsule deposition, maximum maturity and recruitment reported by other authors for different localities are discussed.     

A molecular phylogeny of the Patellogastropoda (Mollusca: Gastropoda)

Phylogenetic analyses of partial 18S rDNA sequences from species representing all living families of the order Patellogastropoda, most other major gastropod groups (Cocculiniformia, Neritopsina, Vetigastropoda, Caenogastropoda, Heterobranchia, but not Neomphalina), and two additional classes of the phylum Mollusca (Cephalopoda, Polyplacophora) confirm that Patellogastropoda comprises a robust clade with high statistical support. The sequences are characterized by the presence of several insertions and deletions that are unique to, and ubiquitous among, patellogastropods. However, this portion of the 18S gene is insufficiently informative to provide robust support for the monophyly of Gastropoda, or to address the division of the Gastropoda into the subclasses Eogastropoda (= Patellogastropoda + hypothetical coiled ancestors) and Orthogastropoda. These sequence data invariably group Patellogastropoda in a weakly supported clade with cocculiniform limpets, despite greater sequence divergences between Patellogastropoda and “Cocculiniformia” than between the Patellogastropoda and Orthogastropoda. Partial 18S sequences support the inclusion of the family Neolepetopsidae within the superfamily Acmaeoidea, and refute its previously hypothesized position as sister group to the remaining living Patellogastropoda. This region of the 18S rDNA gene diverges at widely differing rates, spanning an order of magnitude among patellogastropod lineages, and therefore does not provide meaningful resolution of the relationships among higher taxa of patellogastropods. Data from one or more genes that evolve more uniformly and more rapidly than the 18S rDNA gene (possibly one or more of the mitochondrial genes) seem more likely to be informative about relationships within Patellogastropoda. Received: 5 February 1999 / Accepted: 16 May 2000     

Imposex in Hexaplex trunculus (Gastropoda: Muricidae): first results from biomonitoring of tributyltin contamination in the Mediterranean

Imposex (the occurrence of penis and vas deferens in females) in Hexaplex trunculus collected from Malta (Central Mediterranean) in 1992 is reported. This phenomenon is related to the levels of tributyltin (TBT) in the organisms as well as in sediments. Imposex indices (relative penis size and vas deferens sequence) are used to biomonitor coastal contamination by TBT for the first time in the Mediterranean. In its response to TBT, H. trunculus is ranked as one of the most sensitive neogastropod species studied so far. Accumulation of TBT and its metabolites in the digestive glands, gonads and the rest of the body are described, and sex-related differences are shown. No preferential female mortalities are recorded in populations exposed to high levels of TBT. However, a reported shift in the size frequency distribution of H. trunculus in contaminated sites, towards bigger snails, may suggest reduced reproductive potential.     

Chemical defense and evolution in the Sacoglossa (Mollusca: Gastropoda: Opisthobranchia)

Summary. Recent advances in both the systematics and the natural products chemistry of the order Sacoglossa (=Ascoglossa) in the gastropod subclass Opisthobranchia suggest a revised and improved historical account of the evolution of the group. Although the algal genus Caulerpa makes a suitable model for the ancestral food of the order, other siphonaceous algae are consistent with both morphological and chemical data. At an early evolutionary stage terpenoids are sequestered from the food, and used defensively, often with modification. With an evolutionary switch to different kinds of algal food, there is often a shift to other, related defensive chemicals. A switch to new food source sometimes leads to the abandonment of chemical defense, but in other cases there is de-novo synthesis of defensive metabolites. The synthesis of polypropionates, which are used defensively, occurs in some other gastropods, but otherwise is known only in fungi. The systematic distribution of the defensive polypropionates suggests that their defensive use has evolved several times among gastropods. Failure to detect them may mean that synthetic capacity has evolved more than once, or it may mean that they exist at low levels, perhaps having a non-defensive function. Received 9 February 1998; accepted 20 March 1998.     

Evolution and systematics in Haliotidae (Mollusca: Gastropoda): inferences from DNA sequences of sperm lysin

Abalone taxonomy and systematics have remained unresolved: neither stable species-level nomenclature nor a cladistic hypothesis of relationships among species have been established. To infer the phylogeny of the genus Haliotis and to identify species using molecular data, we compared complementary DNA (cDNA) sequences of sperm lysin from 27 species-group taxa from California, Japan, Australia, New Zealand, Taiwan, Borneo, Madagascar, South Africa, Greece, France, Italy and the Azores. The lysin cDNA sequences reveal that 22 of the 27 taxa are clearly distinguishable by >20 nucleotide differences. Of the remaining 5, H. coccinea from the Azores may be a subspecies of H. tuberculata, if not a sibling species (10 nucleotide differences). The other four taxa are most probably the same species as one of the 22 taxa: the lysin sequences are almost identical between H. madaka and H. discus hannai, H. conicopora and H. rubra, H. diversicolor supertexta and H. diversicolor aquatilis, and H. tuberculata lamellosa and H. tuberculata tuberculata. The phylogeny of lysin cDNA suggests that there are three groups among the 27 species-group taxa: (1) all California species and 3 Japanese species (H. gigantea, H. discus hannai, and H. madaka): (2) 1 New Zealand species (H. iris); (3) 1 Japanese species (H. diversicolor aquatilis), Indo-West Pacific species and European species. These groups can be assigned to three previously recognized subgenera (Nordotis, Paua and Padollus) in the genus Haliotis. Two historical hypotheses are proposed to explain the biogeography and evolution within these abalone: (1) Tethyan distribution of the ancestral abalone, during the Cretaceous, followed by extinction in most of the habitat, but radiation in California and Southeast Asia which later spread to the other areas; (2) North Pacific rim distribution of the ancestral abalone, followed by dispersal to the other areas during the Paleogene.     

Density-dependent mortality of recruits of the abaloneHaliotis rubra (Mollusca: Gastropoda)

In three wild populations ofHaliotis rubra surveyed off southeastern Australia during the three years 1987–1989, 0 to 10% post-settlementH. rubra survived for 5 mo. Sequential censuses (February, May and December in each year) provided evidence that mortality of post-settlementH. rubra increased with the density of settlement. The occupation of cryptic habitat byH. rubra soon after settlement may well be an adaptation for avoiding predators or grazers and dislodgement by violent water movement.     

Types of larval development in Littorinidae (Gastropoda: Prosobranchia) of the World Ocean,and ecological patterns of their distribution

Relationships between the types of larval development in 4 species of Littorina from British shores and their inhabitancy of different tidal levels were established by Woodward (1909). His data have led to the view that the Littorinidae present a “text-book” example illustrating obligate interrelationships between the distribution of marine bottom invertebrates in certain biotopes and the possession by them of definite types of larval development. Analysis of data describing larval development of 39 species of littorinids from various regions of the World Ocean covering all its climatic zones, from the tropics to the Arctic, accumulated in the literature (Tables 1-5) demonstrates, however, that there exists only a relationship between viviparity and the inhabitancy of supralittoral zones, and between direct development and the inhabitancy of littoral zones. In 34 out of 39 littorinid species studied pelagic development, in various modifications, is present at all tidal levels; it is most common in the supralittoral zone, but not in the sublittoral zone, as has been generally assumed. The data presented in Tables 1-5 confirm the author's opinion (Mileikovsky, 1971, 1973) that the concept of obligate interrelationships between definite types of larvel development in marine benthic invertebrate and their existence in definite biotopes, based on Woodward's scheme, is false and must be rejected.     

Distribution of the soft-bottom macroinfauna in an estuary of Southern Chile

In April 1981, a quantitative survey (20 stations) of the soft-bottom macroinfauna of the littoral and sublittoral was conducted in a Southern Chilean estuary. A principal component analysis rendered three groups of stations with different substrate conditions. A total of 16 775 animals was collected (40 taxa), with Oligochaeta, Polychaeta and Ostracoda representing 83% of the individuals. Most of the biomass was attributed to Bivalvia (43.7%) and Polychaeta (33.6%). Factor analysis using abundance data of the 21 most abundant taxa rendered three groups of stations in the Q-mode (a cluster analysis rendered 4) and three groups of taxa in the R-mode. One group mainly inhabited the sandy outlet area of the estuary where salinity was highest. Another group of organisms was mainly found in the upper zone of the estuary where salinity was low and the substrate was muddy. The third group was principally distributed in the central part of the estuary and spread from there either to the outlet area or to the upper zone. Some ungrouped taxa were distributed relatively homogeneously within the estuary. It was found that besides substrate properties salinity and turbulence also played an important role in the distribution of organisms in the Río Lingue Estuary.     

Ontogenetic change in the radula of Conus magus (Gastropoda)

Specimens of the Indo-Pacific piscivorous gastropod Conus magus Linnaeus, 1758 were either collected by SCUBA in the field in Palau (1983–1984) or raised in the laboratory from egg capsules. C. magus captures adult fish with a specially modified radula tooth. Radula morphology was analyzed in a size range of individuals from 4.1 to 43.7 mm shell length, which encompassed early postmetamorphic juveniles to adults. Post-metamorphic C. magus juveniles are too small to consume fish, and all individuals below 9 mm possess a juvenile radula tooth totally different from that of the adult and resembling that of some vermivorous Conus species. The only food remains found in the digestive tracts of juveniles were the setae of syllid polychaetes. All individuals above 10.5 mm possessed adult teeth and had only fish remains in the gut. Two specimens, 10.1 and 9.2 mm in shell length, had intermediate-type radula teeth.     

The nature of heat coma in Littorina littorea (Mollusca: Gastropoda)

 The nature of heat coma was examined in the edible periwinkle Littorina littorea (L.). Duration of acclimation did not influence heat-coma temperature at 12 °C, although other acclimation temperatures were important in influencing thermal tolerance, with positive shifts in coma temperature observed in response to elevated temperatures. Previous thermal history also influenced heat-coma temperatures. Individuals subjected to repeat heat-coma events on a daily basis showed significant declines (P < 0.05) in coma-temperature; in contrast individuals exposed to repeat heat-coma events on a weekly basis showed no decline in thermal tolerance. Size-effects occurred at selected sites, where decreased heat-coma temperatures were recorded in large individuals. Received: 16 August 1999 / Accepted: 13 June 2000     

Factors affecting fertilization success in two species of patellid limpet (Mollusca: Gastropoda) and development of fertilization kinetics models

Laboratory experiments were undertaken to examine fertilization success in the intertidal prosobranch limpets Patella ulyssiponensis and Patella vulgata. Alkalization of eggs (10 min in pH 9.0 sea water) prior to fertilization trials improved fertilization rates greatly. Fertilization success was found to be a function of sperm concentration, gamete age and contact time. Sperm concentration needed for optimum fertilization success in vivo ranged between 10⁵ and 10⁷ sperm ml⁻¹ for both species although at higher concentrations the number of normally developing trochophore larvae decreased. For P. vulgata, sperm longevity (at a concentration of 10⁶ sperm ml⁻¹) did not exceed 6 h, whereas eggs of both species were fertilizable for up to 12 h. Maximum fertilization success occurred after 15–30 min gamete contact time. The Vogel et al. (Math Biosci 58:189–216, 1982) fertilization kinetics model is developed to allow for non-complete fertilizations under optimal sperm concentrations, and a new parameter fitting technique is developed to improve estimates of fertilization success for short gamete contact times.     

Recruitment and juvenile survival in the sea hare Aplysia juliana (Gastropoda: Opisthobranchia)

The specificity in settling response of larval Aplysia juliana (Quoy and Gaimard, 1832) on species of its food algae, Ulva spp., provided a rare opportunity for the quantitative study of recruitment and early survival rates in a non-sessile marine invertebrate. Post-metamorphic juveniles spend about 3 weeks feeding and growing on the algae before moving to the rocky habitat of the adults. Recruitment and survival were studied by collecting weekly samples of algae and holding them in the laboratory until the recruits were large enough to be seen and reliably counted. Recruitment occurred throughout the year, although the rates were relatively low in late winter and spring. Recruitment was monitored relative to algal weight and bottom area. Variation in larval abundance and algal standing crop influenced recruitment to the study area. Mortality during the juvenile stage was very high in all cases, although the rates and shapes of the survivorship curves varied between algal species and location.     

Reproductive cycle and gametogenesis in the black abalone Haliotis cracheroidii (Gastropoda: Prosobranchiata)

Seasonal changes in gonald size and stages of gametogenesis in the black abalone Haliotis cracheroidii were related to changes in environmental parameters. H. cracheroidii showed an annual reproductive cycle terminating in a synchronized spawning in late summer. Gametogenesis was initiated immediately after spawning. Gametes were present in the gonad through the winter months. Gametogenesis was initiated a second time in the spring months. Maximal gonad growth (to a gonad index of 20%) occurred during summer months prior to spawning. Changes in gonad size andperiods of initiation of gametogenesis revealed no apparent correlation with changes in seasonal water temperature. Changes in gonad size showed no apparent relation to change in day length. Total polysaccharide levels in foot tissue changed seasonally, indicating that food availability is probably not a factor in directly regulating gonad growth. Gonad index data for the chiton Katharina tunicata (collected over a 10 year period) showed no apparent correlation to seasonal change in water temperature.