Effects of Polinices duplicatus (Gastropoda: Naticidae) on infaunal community structure at Barnstable Harbor,Massachusetts, USA

Effects of the predatory naticid snail Polinices duplicatus (Say) on species composition, diversity and density, of the infauna of intertidal sand-flats at Barnstable Harbor, Massachusetts, USA, were studied using field experiments. Responses of molluscs (prey of P. duplicatus) and nonmolluscs (nonprey) were considered separately to distinguish between the effects of feeding and sediment disturbance during foraging. The fauna of 0.25 m² predator-exclusion cages, coarse-mesh cages, sieved areas, and controls was followed for 1 yr. Species associations within cages from which predators were excluded were denser, more diverse, and richer in molluscs than those in other treatments. Larger areas (3×3 m) of natural bottom were fenced and maintained as snail exclosures and enclosures during two feeding seasons. For both molluscs and nonmolluscs, diversity (H'), number of species, evenness (SD), and density all decreased with increasing snail density. Intense predation pressure on molluscs and selective feeding on thin-shelled bivalves, which were rare, removed individuals and species of molluscs from the community. Comparison of samples taken inside and outside trails made by snails showed that disturbance of the surface sediment layers by snails decreased the abundance of spionid polychaetes and total nonmolluscs. The sipunculan Phascolopsis gouldi and the bivalve Gemma gemma dominated the community in all experimental treatments and were little affected by the activity of Polinices duplicatus. Predation and disturbance by snails lowered community diversity by removing individuals of the less abundant species, and generally maintained population densities below the level where strong competition would occur.     

Energy metabolism in the foot of the marine gastropod Nassa mutabilis during environmental and functional anaerobiosis

This study concerns the effects of oxygen deprivation due to incubation in oxygen free sea water (environmental anoxia) or exercise (functional anoxia) and of exposure to air on the mode of energy production in the foot of the whelk Nassa mutabilis. Additionally, energy metabolism of the foot muscle was investigated during exercise after different anoxia periods and during the subsequent recovery period. During environmental anoxia, phosphoarginine, glycogen and aspartate are broken down as substrates and alanine and succinate are formed as products. There was no production of D-lactate or octopine. The energy charge value fell after 24-h anoxia. Exposure to air resulted in only small changes in phosphoarginine and alanine levels, suggesting that oxygen uptake was impaired in the first phase of air exposure but that, later, aerial respiration kept pace with the energy demand. Exercise caused a dramatic decrease of phosphoarginine concentration, coupled with glycolytic ATP production via octopine formation. In the recovery period (after exercise), the level of phosphoarginine was rapidly restored. An anaerobic component was evident during recovery as shown by the accumulation of D-lactate. Thus, both terminal dehydrogenases, octopine- and lactate dehydrogenase, are active in the muscle, but under different physiological conditions. Octopine formation also took place when the whelks were subjected to exercise after 4 or 24 h of anoxia. In this case, glycolysis provided between 70 and 90% of the energy required since the phosphagen store had already been depleted during the anoxic period. When the work load was increased (greater number of leaps), it became evident that the action of arginine kinase and octopine dehydrogenase are not closely linked. First there was an increase of arginine and then later a condensation of arginine with pyruvate to form octopine.     

Encapsulated development of the marine prosobranch gastropod Nucella lapillus

Nucella (=Thais) lapillus (L.) develops to the juvenile stage within an egg capsule. This study was undertaken to investigate nutritive and bacteriostatic properties of the intracapsular fluid for this species. Capsules were collected intertidally at Nahant, Massachusetts, USA, in the summers of 1982 and 1983. The average capsule contained only 1.1 l of intracapsular fluid per embryo. A statistically significant correlation was observed between capsule height and number of developing embryos contained therein, although the number of embryos per capsule was more closely related to capsule volume. Total dry weight of encapsulated, shelled embryos increased exponentially as a function of shell length. However, weight increases may be entirely due to shell calcification; the average dry tissue weight of shelled embryos was significantly below that for pre-shelled individuals, suggesting a net loss of biomass during development. Aliquots of intracapsular fluid failed to inhibit the growth of the 13 bacterial strains tested. Despite the above results, pre-shelled embryos generally survived only several days following their premature removal from egg capsules at 12°C. Shelled veligers were reared outside of the capsules with no apparent ill effects. An explanation for these survivorship results is elusive.     

Attraction to prey and stimulus to attack in the predatory gastropod Urosalpinx cinerea

Urosalpinx cinerea (Say), accustomed to feeding on Balanus balanoides, were strongly attracted in a choice chamber by the effluents of B. balanoides and B. eburneus, were indifferent to the effluents of Crassostrea virginica, Crepidula fornicata and Mytilus edulis, and responded negatively to the effluent of their own species. Oyster drills from a population feeding on Crassostrea virginica were attracted to oyster effluent; when these snails were offered a choice between Crassostrea virginica and B. balanoides effluents, they responded in equal numbers to the two effluents. Concentrations of NH₄Cl-ammonia in the range 18.2 to 73.3 mol did not attract the snails, and their responses to animal effluents were not correlated with the ammonia and amino-acid concentrations of the effluents, which ranged from 11.8 to 21.0 mol. It is argued that these results deny ammonia the role of a nonspecific distance attractant. Confined separately with various potential prey species, Urosalpinx drilled Balanus balanoides, B. eburneus, Crassostrea virginica, Crepidula fornicata, C. plana, Mercenaria mercenaria, Modiolus demissus, Mya arenaria, Mytilus edulis, Spisula solidissima, and Yoldia limatula, but not Anomia simplex. The prey species that were the least attractive in running water were generally rendered attractive and subject to attack in standing water. Freshly shucked Crepidula fornicata shells were scarcely drilled unless continuously perfused by C. fornicata effluent, and then only in small numbers. Altering the texture of living C. fornicata shells by wrapping them in nylon netting, by polishing, or by roughening, did not make them immune to drilling. Oyster drills in contact with C. fornicata attacked only if they received the effluent of the living prey, proving that tactile stimuli alone are not adequate to induce drilling.     

Defenses and refuges: alternative mechanisms of coexistence between a predatory gastropod and its ascidian prey

Only five of the 14 species of solitary ascidians in the San Juan Islands, Washington, USA commonly co-occur with an abundant predator of the rocky subtidal, the gastropod Fusitriton oregonensis. None of the common subtidal species is defended by vanadium or sulfuric acid; concentrations of these substances are highest in species that are eaten readily by F. oregonensis and are uncommon in the rocky subtidal. Experimental manipulations indicate that Halocynthia igaboja are protected by stiff spines of the tunic. The defense mechanism of Pyura haustor consists of a tough, leathery tunic with inorganic inclusions, and microscopic spines around the siphons. Boltenia villosa and Styela gibbsii are eaten by snails in the laboratory, but live in areas where the predator is abundant. Both find refuge as epizooites on P. haustor and H. igaboja. The mechanisms by which Cnemidocarpa finmarkiensis persists in the presence of F. oregonensis is unknown.     

Isolation and population divergence of a widespread Indo-West Pacific marine gastropod at Easter Island

Oceanic islands represent excellent systems for studying the link between geographic isolation and population divergence. Easter Island is the world’s most isolated island and exhibits a high level of endemicity in the nearshore marine environment. Yet few studies have examined the effect of such extreme isolation on the divergence of populations of widespread species that occur at Easter Island. Conus miliaris, a marine gastropod distributed throughout much of the Indo-West Pacific, occurs at Easter Island where the population is ecologically and morphologically distinct from other populations of the species. To determine whether these phenotypic differences are associated with genetic isolation of the Easter Island population, we investigated the phylogeography of this species by examining mitochondrial COI sequences obtained from 141 individuals from eight localities occurring predominantly in the western, central and southeastern Pacific. Results from our analyses show that C. miliaris at Easter Island differs genetically from other populations. We estimate that C. miliaris colonized Easter Island shortly after the origin of the island ≤0.7 million years ago and that since population founding, gene flow has occurred predominantly from Easter Island to the west and that little migration has occurred into Easter Island.     

Accumulation and regulation of heavy metals by the intertidal snail Polinices sordidus

The gastropod Polinices sordidus was collected from an uncontaminated area in Quibray Bay, New South Wales, Australia, in 1990. The snails were exposed for 2 wk to polluted sediments collected from Port Kembla Harbour, Blackwattle Bay, Lake Illawarra, Lake Macquarie in New South Wales, Australia, and the Derwent River in Tasmania, Australia. Metal accumulation and regulation by this species were evaluated. Metal concentrations in snail tissues and total, EDTA and HCl-extractable metals in the sediments were compared. Copper concentrations were extremely varied in snails exposed to the same sediment. This was not the case for other matals tested. No accumulation of copper was found in snails exposed to different sediments. There was no zinc accumulation from sediments containing less then 10 mg Zn/g. P. sordidus could accumulate lead, manganese and iron from some of the sediments. Manganese concentrations in the snail tissues correlated with total, HCl-extractable and, more significantly, EDTA-extractable Mn in the sediments. P. sordidus was not considered to be a good bioindicator of copper and zinc contaimination in sediments; however, this species could be used as an indicator of lead and manganese contamination.     

Variability and its effect on the 24h chlorophyll budget of a small marine basin

In an intensive study (lasting 25 h) of the production, export and grazing of phytoplankton in a small marine basin, it was found that 58% of the production (11% of the total standing stock) was lost by exchange with the sea and 34% was consumed by grazing of zooplankton. The measured production of phytoplankton could be balanced, to within a few percent, against grazing, export, and a small, measured, net change in the total standing stock of the basin. Large variations were observed in concentrations of chlorophyll and zooplankton at the mouth of the basin over the 25 h period. These variations were associated with changes in the height of the tide, but were about 2^1/2 h out of phase with it. Strong negative correlations were observed between chlorophyll and transport, such that only 35% of the chlorophyll exported was exchanged via the mean flow, while 65% was exchanged via the fluctuations. The correlation was even more striking with zooplankton, for which virtually all the export was associated with the fluctuations in the transport. Time series observations in the centre of the basin revealed considerable short-term variability in both chlorophyll and zooplankton, but the variations were smaller than those observed at the mouth of the basin, and the phase lag with the tide was longer. The variability studies enable suggestions to be made about more economical design of sampling programs, but illustrate the difficulty of providing verification data for any continuous model of primary production in such a basin.Bedford Institute Contribution No. 231.Canadian Contribution to IBP No. 97.     

Lack of genetically-subdivided population structure in Bullia digitalis, a southern African marine gastropod with lecithotrophic development

Larval shell morphology in fossil and present-day gastropods is often used to infer modes of larval development and levels of dispersal. Dispersal ability influences not only genetic population structure, but also is thought to influence a species' geographical range and evolutionary duration. We tested these predictions in Bullia digitalis, a sandy-beach whelk, by examining genetic variability at 33 protein-coding loci in nine samples (N=739) taken in 1984 to 1985 at localities extending over about three-quarters of the geographical range of this species in southern Africa. Females of this species deposit eggs into benthic or brooded capsules in which larvae develop through the trochophore and veliger stages to emerge as crawling juveniles. Scanning electron micrographs confirmed a protoconch morphology typical for gastropods with lecithotrophic larval development. Contrary to expectations, subpopulations of B.␣digitalis had high levels of variability (H=0.102) and lacked a genetically-fragmented structure (=0.013). The lack of a genetically-subdivided population structure would not have been correctly inferred, if this species were known only from well-preserved fossil shells. Indirect estimates of migration between populations based on and the island model of migration, which assumes drift-mutation equilibrium, ranged between 19 and 23 individuals per generation. Either an undescribed mechanism of dispersal facilitates gene flow between populations, or the geographical range of this species has recently expanded to produce the appearance of high levels of gene flow. Gene-frequency distributions showed that relative to four other species of Bullia, populations of B. digitalis were in mutation-drift disequilibrium, with a significant excess of low-frequency alleles that is consistent with a recent rapid expansion from a small population. Also contrary to expectations, this species has a large geographical range (2 400 km) and an apparently long evolutionary history extending 5 to 20 million years, as estimated from an allozyme phylogeny with four other species of Bullia. Received: 15 January 1997 / Accepted: 28 January 1997     

Age and growth of the naticid gastropod <Emphasis Type="Italic">Polinices pulchellus</Emphasis> (Gastropoda: Naticidae) based on length frequency analysis and statolith growth rings

In Red Wharf Bay, UK the naticid gastropod, Polinices pulchellus, was more abundant and more highly aggregated during the summer months (June–August 2001) than during the winter (December 2000). Whilst small numbers of juvenile P. pulchellus (4–6 mm shell length) were present throughout the year the population consisted mainly of individuals of 12–14 mm shell length. Juvenile snails grew rapidly in size during the winter and early spring; growth then virtually ceased between May and June, following which there was a further period of rapid growth between August and February. Densities ranged between 57 and 4,073 ha⁻¹ and the largest individual collected during this investigation measured 16.2 mm in shell length. Statoliths from adult P. pulchellus revealed the presence of a settlement ring and two prominent growth rings (rings 1 and 2). A curvilinear relationship exists between statolith diameter and shell length in snails up to 16 mm in length. Settlement rings ranged in diameter from 19.7 to 45.2 μm (mean 29.8 μm; SE=0.41) giving an estimated shell length of the settled juvenile of 1.1 mm. The diameter of ring 1 and ring 2 were significantly correlated indicating that rapid growth during the first year is maintained during year 2. Shell lengths estimated from the diameters of the prominent statolith rings and those obtained from length frequency data analysis (LFDA), were broadly congruent strongly suggesting an annual periodicity to the statolith rings. The largest snails (>15 mm) present within this population were estimated to be between 2 and 3 years old. Von Bertallanfy seasonal growth curves obtained from the LFDA predicted values of L∞, K and t ₀ of 14.32 mm, 1.54 and −0.14 years, respectively, suggesting that P. pulchellus rapidly attains its maximum asymptotic size.     

Yolk utilization in the gastropod Crepidula fornicata

The increases in constituents per unit weight of eggs during embryonic development of the gastropod Crepidula fornicata amounted to 14.6% for ash, 1.0% for protein, and 0.3% for non-protein nitrogen. During the same stages, fat content decreased from 33.7 to 20.3%, carbohydrate from 10.2 to 7.7% and energy content from 6209 to 5298 cal/g dry organic substance. The cumulative efficiencies for yolk utilization were 83.8% for dry weight, 61.0% for energy, 85.1% for protein, 50.7% for fat, and 63.6% for carbohydrate. A single egg contained 0.0269 cal, a single veliger 0.0164 cal. Of the 0.0105 cal expended on metabolic processes of the embryo, oxidation of fat contributed as much as 65.3%, while that of protein and carbohydrate amounted only to 18.8 and 6.3%, respectively. On the basis of ecophysiological considerations, a new classification of eggs is proposed.This paper is based on a lecture presented during the first Convention of the Indian Association of Biological Sciences in Madras, December, 1968.     

Role of herbivory in controlling phytoplankton abundance: annual pigment budget for a temperate marine fjord

An annual pigment budget was constructed for Dabob Bay, Washington (USA) by comparing the downward vertical loss of phytoplankton pigments (chlorophyll and pheopigments) to the production of chlorophyll within the euphotic zone. The vertical flux of pigments was measured with sediment traps deployed at intervals of 1 to 6 wk over a 2.5 yr period yielding 763 d of trap exposure (28 November 1978–16 June 1981). The production rate of chlorophyll was calculated from measurements of algal specific growth rates, chlorophyll (chl) crops, primary production (as carbon) and appropriate C: chl ratios. Sixty one to 77% of the annual chlorophyll production was accounted for by the vertical flux of pheopigments resulting from herbivorous zooplankton grazing (macrozooplankton). Algal sinking, represented by downward chlorophyll flux, accounted for only 5 to 6% of the annual chlorophyll production. The remaining fraction of chlorophyll production was estimated to be consumed by small herbivores (microzooplankton), whose fecal material contributes to the suspended pool of pheopigments found in the euphotic zone. The suspended pheopigments are continuously removed by photodegradation. In Dabob Bay, the major process controlling phytoplankton abundance is zooplankton grazing and it appears that the ultimate fate of most phytoplankton is to be consumed by herbivores.     

Life history of the temperate subtidal gastropod Umbonium costatum

The life history and reproductive biology of the trochid sand snail Umbonium costatum (Kiener) were investigated on a subtidal sandy shore in Hakodate Bay, Japan between 1988 and 1991. Female U. costatum mature at 1 yr of age (shell diameter-11 mm), reproduce twice (in June-July and September-October) in successive years, grow to a maximum size (shell diameter=22 mm) at age 8 yr, increase annual fecundity with age from 2000 (age 1 yr) to 37000 (age 8 yr), and show a maximum monthly gonad somatic index of 8% which is constant among ages. In comparison to a previously studied life history of a tropical Umbonium vestiatium, temperate U. costatum shows more sustained growth and a longer life span after maturation. This could be explained by: (1) the optimal size model concerned with resource investment, in gametes (Sebens 1987); and by (2) bet hedging to compensate large variability in larval success at high latitudes. These two hypotheses are not mutually exclusive, but both are based on season-related extremes of environment at high latitudes where the period suitable for reproduction is short.     

Nitrate uptake in marine phytoplankton: Energy sources and the interaction with carbon fixation

Field studies of whole natural phytoplankton communities from Knight Inlet, B. C., Canada and laboratory cultures of the diatom Skeletonema costatum indicate inorganic carbon fixation may be temporarily suppressed following 10 to 15% enrichment with NO ₃ ^- or NH ₄ ⁺ . (This effect is suggested to be due to competition between inorganic carbon and nitrogen for adenosine triphosphate (ATP), and is reduced when chlorophyll a is increased intracellularly after 6 to 8 h.) Results imply that the source of ATP for nitrate uptake is primarily from Photosystem I (cyclic photophosphorylation) in the presence of light. It would appear that a transient nutrient-adaptive response occurs upon addition of extracellular nitrogen.     

Age-and sex-dependent patterns of aggregation in the tropical gastropod Strombus luhuanus

The spatial dispersion of the tropical coral sand-dwelling mesogastropod Strombus luhuanus Linné, 1758 was studied at several sites in two localities (Australian Great Barrier Reef and southern Papua New Guinea), from September 1980 to February 1983. S. luhuanus usually occurred in local aggregations, many of which were relatively dense with discrete boundaries: Four types of aggregation are described: mixed age-class, juvenile, mating, and clusters. The former two are termed colonies, because they persisted over time and exhibited some coordinated movements. Mixed age-class colonies contain individuals of all ages, but within them the different ageclasses are frequently spatially segregated from each other. This was sometimes due to the younger age-classes' preference for shallower areas, but also occurred in areas of constant depth. Juvenile colonies are relatively smaller in area but higher in density, and are dominated by a single size-class. Mating aggregations are concentrations of copulating individuals, in which competition between males for access to females occurs. Clusters are concentrations of inactive individuals (juvenile and adult) piled together in close contact. All types of aggregation were concentrated in only a part of what appeared to be the suitable habitat. Most aggregations' boundaries shifted over time within the habitat. We conclude that intraspecific attraction must be an important factor which maintains the spatial structure of all aggregations, although habitat specificity sets the larger area in which they move. Processes which may underlie this pattern, and some of its implications, are discussed.     

Evaluation of the significance of metal-binding proteins in the gastropod Littorina littorea

Measurements of metals in Littorina littorea from clean and contaminated sites in Great Britain confirmed that body concentrations of Ag, Cd and Hg vary according to environmental contamination, while the essential elements Cu and Zn are regulated. Investigation of the cytosolic distribution of metals in L. littorea, using Sephadex G-75 gel-permeation chromatography, revealed important distinctions in the partitioning of elements. In contrast to Ag and Hg, which were mainly associated with high molecular weight ligands, Cd was bound predominantly to an intermediate molecular weight, soluble protein (CdBP-I, apparent mol. wt=20 000). However, although the main function of CdBP-I, both in clean as well as contaminated individuals, was sequestration of Cd, significant quantities of other non-essential metals (Ag, Hg) were similarly bound. Laboratory experiments confirmed the role of CdBP-I in complexing cadmium, and also revealed the induction of a second cadmium-binding protein (CdBP-II) in response to high cadmium levels. The apparent molecular weight (10 000), absorbance characteristics and high (inducible)-SH content of CdBP-II suggest similarities with metallothionein. A dose-related increase in the-SH content of very low molecular weight (>3 000) fractions was also observed in cadmium-exposed L. littorea, although no cadmium was associated with these ligands. A detoxifying role is tentatively proposed for the metal-binding proteins CdBP-I and II in L. littorea. However, some spillover of cadmium to the high molecular weight protein pool was observed in individuals exposed to cadmium in the field and laboratory. The use of L. littorea in biochemical assays of environmental contamination is discussed.     

Aspects of the feeding strategy of the predatory crab Scylla serrata

Scylla serrata in a South African estuary occurred more frequently in that part which had the highest number of prey organisms. Food location was by contact chemoreception, using the dactyls of the walking legs. Major prey groups were burrowing bivalves, attached bivalves and small crabs. s. serrata showed a preference for small crabs as prey. Because of their larger mass and higher energy content compared with other prey organisms, these crabs represented the major energy source of S. serrata in this area.     

Feeding biology of a predatory and a facultatively predatory nematode (Enoploides longispiculosus and Adoncholaimus fuscus)

This paper reports on the feeding biology of a predatory and of a facultatively predatory nematode, Enoploides longispiculosus and Adoncholaimus fuscus, respectively. Both species represent genera which are common and abundant in the littoral of the North Sea and in adjacent estuaries. Observations on the foraging behaviour of both species are given, and for the former species, a range of prey from its natural habitat is identified. Respiration was determined using a polarographic oxygen electrode technique and compared to consumption determined as predation rates on the monhysterid nematode Diplolaimelloides meyli. The daily C-loss due to respiration accounted for 15% of the measured C-consumption in E. longispiculosus and for 111% in A. fuscus, proving the observed feeding rates in the latter species to have been inadequate for the maintenance of its aerobic metabolism. Daily respiration rates at an average environmental temperature were 219 ng C ind⁻¹d⁻¹ for adults of A. fuscus and 21.9 ng C ind⁻¹ d⁻¹ for adults of E. longispiculosus. Using radiotracer techniques, no uptake of bacterial cells or of organic matter in the dissolved phase was demonstrated for E. longispiculosus. In A. fuscus, however, a significant drinking of label in the dissolved or volatile fraction occurred; bacterial cells were taken up at a level insignificant to the nematode's daily C-ration. It is concluded that E. longispiculosus has a fairly strict predatory feeding strategy, while A. fuscus gains a majority of C from additional foraging strategies, among which the uptake of dissolved material and scavenging on macrofauna carcasses (as reported in the literature) may be of particular importance. Received: 28 August 1998 / Accepted: 8 March 1999