Analysis of patterns of dispersion of intertidal prosobranch gastropods in relation to macroalgae and rock-pools

Summary The patterns of dispersion of Nerita atramentosa, Austrocochlea constricta, Bembicium nanum and Cellana tramoserica are analysed from quadrat samples taken between June 1972 and June 1973. In each quadrat, the proportion of animals in each of two substrata, the encrusting alga Peyssonelia gunniana and small pools, was recorded, with the proportional area of cover of each substratum, height on the shore, density of the species and the time of sampling. Multiple regression and covariance analysis showed that, in all cases, there was a linear relationship between proportion in a substratum and proportional area of substratum. No other variables were significant.Juvenile Nerita and Austrocochlea showed preferences for both substrata. Adult Nerita and all Bembicium were dispersed at random with respect to both substrata. Adult Austrocochlea were randomly dispersed with respect to Peyssonelia, but showed a preference for water. Cellana showed a preference for water and against Peyssonelia. These microhabitat preferences are discussed in relation to previously described effects of substrata on the densities of these four species.     

Comparative studies on the biology of nerita atramentosa Reeve, bembicium nanum (Lamarck) and cellana tramoserica (Sowerby) (gastropoda: Prosobranchia) in S.E. Australia

Populations of three coexisting intertidal gastropods, Nerita atramentosa Reeve, Bembicium nanum (Lamarck) and Cellana tramoserica (Sowerby), were sampled from a shore in Botany Bay, New South Wales, from July 1972 to September 1973. The recruitment and growth rates of each species were analysed from size frequency distributions. Mortality of each age cohort, and longevity, were estimated from analyses of the densities of Nerita atramentosa and Bembicium nanum.Nerita atramentosa showed no significant mortality during the first two years on the shore, but high mortality (at an instantaneous rate of 0.084 deaths/individual/month) after reaching the age of reproductive maturity, which was 20 months from settlement and at a mean shell-length of 13.5 mm. No growth could be detected after reproductive maturity was reached. Longevity of N. atramentosa was estimated as 3–5.5 years. Bembicium nanum juveniles showed higher mortality (at an instantaneous rate of 0.233 deaths/individual/month) than reproductively mature animals (0.060 deaths/individual/month). Reproductive maturity was reached at a mean shell-breadth of 11.0mm, i.e., about ten months after settlement on the shore. Longevity was estimated as from 4–8 years. Cellana tramoserica showed different growth patterns depending on the time of settlement.The three species showed different patterns of growth and life history relating to variable recruitment (which was demonstrated in all three species) and different rates of mortality of age cohorts.     

Movements of intertidal gastropods

Movements of marked individuals of Nerita atramentosa Reeve, Austrocochlea constricta (Lamarck), Bembicium nanum (Lamarck), and Cellana tramoserica (Sowerby) were recorded at various sites in New South Wales in 1974 and 1975. A model of random movements during feeding at high tide was proposed, i.e., that animals wandered in random directions and the distances moved were exponentially distributed. Appropriate statistical tests for departures from such a model are given. The types of departure tested were non-random directions, non-exponentially distributed dis- tances, different distances in different directions, and correlated distances or directions of movement between days. With two exceptions, the movements of the animals over short periods were adequa- tely described by the assumption of random movements. One exception was A. constricta in one locality where movements of the sample were limited by it being in a large rock-pool and the other was the limpet C. tramoserica which displayed homing behaviour, so that the population could be divided into homers and non-homers. Movements of non-homing limpets were random.The mean distance moved by N. atramentosa was not significantly different from that of A. con- stricta. The mean distances moved by these two species were greater than those of B. nanum and the non-homing C. tramoserica, which were not different from each other. Adult and one-year old N. atramentosa did not differ significantly in mean distance moved, but both moved more than juveni- les. These patterns of movement are discussed with respect to hypotheses about feeding in these species.The model of movement was used to predict the effect of different percentage cover in small pools on the movement of A. constricta, which shows a preference for this micro-habitat. Increased availability of small pools reduced the mean distance moved by this species.     

Field experiments with “cageless” methods to manipulate grazing gastropods on intertidal rocky shores

Excluding particular species or guilds of grazing gastropods from experimental areas on intertidal rocky shores is a manipulation frequently used by marine ecologists. Manual removals, cages or fences are classical methods, but they usually require large areas or multiple procedural controls to avoid experimental artefacts. In order to minimise some of those constraints, alternative methods have been developed and are increasingly being used. The effects and limitations of these methods are, however, generally assumed and hardly ever tested. The experiment described here was designed to compare the effectiveness of manual removals, copper paint and sticky barriers in excluding intertidal grazing gastropods from experimental areas on a natural sandstone platform in South-eastern Australia. The response of the grazing assemblage was evaluated for individual species and collectively (integrating all species) by direct counts of individuals and from radular marks on wax discs. The observed patterns varied considerably among species. As predicted, barriers of copper paint effectively excluded the limpet Cellana tramoserica, for several weeks, while having no apparent effect on co-occurring species of grazing gastropods. Sticky barriers were also effective in excluding C. tramoserica, but for shorter periods of time. Contrary to expectations, however, sticky barriers were generally incapable of deterring the coiled snails Bembicium nanum and Nerita atramentosa. Barriers of copper paint are, therefore, a cost-effective way to manipulate diversity (densities of individuals, number and composition of species) and function (intensity of grazing) in this type of assemblage. Care must be taken to clarify the underlying mechanisms and to identify adequate procedural controls.     

Effects of body size and water temperature on grazing rates of four intertidal gastropods

Abstract Rates of radular activity were measured in four species of Australian gastropods. The species were Austrocochlea constricta, Bembicium nanum, Cellana tramoserica, and Nerita atramentosa. Work was done in Botany Bay at Cape Banks, NSW, Australia from September 1989 to June 1990. A hydrophone was used to record radular raspings of snails in situ. Averages based on all observations showed Cellana had the fastest rate with 80 rasps/min, and was followed by Austrocochlea (71 rasps/min), Bembicium (57), and Nerita (39). Faster rates were associated with warmer water temperature and smaller body sizes in all species except Bembicium. The potential for these differences in radular activity affecting competitive interactions is discussed.     

Vertical and seasonal patterns in competition for microalgae between intertidal gastropods

Grazing by the snail Nerita atramentosa and the limpet Cellana tramoserica caused similarly great reductions in abundance of microalgae — measured by direct counts and by estimation of chlorophyll analyses. A smaller snail, Bembicium nanum, caused smaller reductions of microalgal resources, compared with ungrazed areas. These results were consistent with the competitive abilities of these three species. Chlorophyll concentrations in samples of grazed rocks were reliable estimates of the nature and abundance of food available to the grazers.Inter- and intra-specific competition amongst Nerita and Cellana were investigated at various densities in experimental cages. To examine the effects of different availability of food resources, the experiments were repeated at three heights on the shore (abundance of food decreases with height) and during autumn/winter and spring/summer periods of the year (less food is available during summer).Density-dependent mortality of Cellana was caused by the presence of other limpets, or of Nerita. Mortality was greatest at higher levels and during the spring/summer and was significantly, negatively correlated with mean chlorophyll concentration in the experimental cages. The only exception was that all limpets suddenly died in cages at the highest level during the summer period, which cannot be explained solely by competitive interactions. Nerita showed no density-dependent mortality during the short periods of these experiments. Tissue-weights of Nerita declined with increased density, but the effect of Cellana was not as great as the intraspecific effect of Nerita. Snails retained weight better at lower than at higher levels, and during the autumn/winter which is consistent with the availability of food. Tissue weights of both Nerita and Cellana were positively correlated with chlorophyll concentrations inside the cages in both seasons investigated.These experimental results demonstrate that intensity of competition will vary from place to place and time to time according to the densities and mixtures of the grazers, and according to the availability of microalgal food.     

Competition between Cellana tramoserica (Sowerby) (gastropoda) and Patiriella exigua (Lamarck) (asteroidea), and their influence on algal standing stocks

Caging experiments were undertaken in the field to test if competition occurs within and between the limpet Cellana tramoserica (Sowerby) and the starfish Patiriella exigua (Lamarck). Both species suffer from intraspecific competition, even at natural densities, and increased density reduces body weight and survival in Cellana tramoserica and growth rate in Patiriella exigua. Cellana tramoserica depresses the growth of Patiriella exigua, while the starfish unexpectedly seems beneficial to Cellana tramoserica, increasing its body weight. The difference in these effects may be due to the divergent methods of feeding used by the two species. C. tramoserica digs into the substratum and removes most available microflora, while Patiriella exigua everts its stomach onto the rock face and can only remove superficial or loosely attached microflora and detritus, and leaves a mucous web which may supplement the diet of Cellana tramoserica. Even at low densities C. tramoserica can completely prevent macroalgae from developing, while Patiriella exigua is unable to do so, although it shows the rate of development. Macroalgae grew in all the cages which contained only P. exigua, but in spite of this the starfish still suffered from intraspecific competition, probably because they cannot feed on macroalgae. Cellana tramoserica also competes by interference, and on contact with Patiriella exigua it extends its mantle and pallial tentacles, both of which may release mucus. P. exigua retreats from Cellana tramoserica, and those parts of its body that have been touched by the limpet may temporarily be immobilised and become wrinkled up. It is suggested that this interference behaviour by C. tramoserica is derived from a similar but more intense reaction that it has to invertebrate predators. Patiriella exigua lacks a pelagic larva and does not have a refuge outside the range of Cellana tramoserica. Other factors must thus be sought to explain the continued coexistence of Patiriella exigua with Cellana tramoserica. It is suggested that although C. tramoserica has a competitive effect on Patiriella exigua, it (and other grazers) may be necessary for P. exigua, preventing macroalgal growth and maintaining a surface suitable for P. exigua to feed on.     

Effects of interactions between algae and grazing gastropods on the structure of a low-shore intertidal algal community

Summary At low levels on shores in New South Wales, foliose algae are abundant and often occupy all substrata; microalgal grazing gastropods are rare or absent. At higher levels, foliose algae are sparse or absent and grazing gastropods are abundant. Hypotheses for the causes of the lower vertical limits of distribution of these grazers include the effects of increased predation or the deleterious physiological effects of increased period of submergence at lower levels on the shore. Alternatively, the presence of the algae, because they occupy space and deprive the grazers of substratum for feeding, may prevent the downward movement, or survival of the grazers at low levels. Under the first two of these hypotheses, algae are able to colonize and grow in low-shore areas as an indirect result of factors which remove grazers. Under the third hypothesis, the algae are directly responsible for the lack of grazers.Experimental clearings of the low-shore algae and introductions of the mid-shore limpets Cellana tramoserica and Siphonaria denticulata were used to test these hypotheses. C. tramoserica grazes microalgae and removes them from the substratum, preventing colonization. S. denticulata, in contrust, crops the algae, leaving a visible cover of algae on the substratum, which can grow rapidly. Because of its method of feeding, S. denticulata had no measurable impact on the rates of colonization, nor on the dry weights of algae, compared with those of ungrazed areas. C. tramoserica could keep cleared areas tree from foliose algae, but only when the limpets were mainfained in great density (10 per 900 cm²). They were less effective where wave-action was greater.Neither species of limpets could survive when placed onto beds of mature algae, because they had no substratum on which to cling and were swept away by the waves. C. tramoserica did not invade clearings below their lower limit of distribution where they had to move over a bed of foliose algae. Few C. tramoserica moved directly downshore into cleared areas. When placed on bare rock within low-shore beds of algae of different ages, S. denticulata remained amongst the algae and maintained their tissue-weights. Few C. tramoserica remained in areas with well-developed algae, compared with areas having sparse algal growth. Those Cellana which remained amongst well-developed algae lost weight, whereas limpets in areas with less algal growth mammtained their weights. In experimental cages in low-shore beds of algae, where the limpets were inaccessible to potential predators, C. tramoserica lost weight and died. On cleared areas they survived for many weeks, but lost weight and died as algae grew and covered the substratum. In the absence of predation, the micro-algal grazer C. tramoserica could not survive in lowshore areas because algae grew too fast and occupied the substratum, making it inaccessible for the limpets to graze; the algae, once grown beyond small sporelings, are not a suitable food-source for C. tramoserica, and the loss of weight and death of these limpets is attributable to starvation.The lower limit of distribution of C. tramoserica is not due to the direct effects of physical factors associated with prolonged submersion, nor to the impact of predators, but is apparently determined by the presence of rapidly growing, extensive beds of foliose algae at low levels on the shore. The cause of the limit of distribution of S. denticulata is not yet known and predation may prove to be important. Removal of S. denticulata from low-shore algal beds would not, however, affect the domination of substrata by algae. Grazing by S. denticulata at very great density had no effect on algal cover nor weight. In the intertidal community studied, the persistence of a low-shore algal zone, bounded above by abundant grazers is not influenced by the activities of predators, but is a direct result of interactions between the grazers and the algae.     

The grazing effect of limpets on the macroalgal community of a rocky intertidal shore

Abstract Abundance of macro-algae in the mid-littoral zone on a Victorian intertidal rocky shore varied seasonally, algae being rare in Summer and common in Winter and Spring. Field experiments demonstrated that, of two species of grazing limpets (Cellana tramoserica and Siphonana diemenensis), only Siphonaria had a major effect on the abundance of foliose algae (e.g. Scytopsiphon lomeniarid) and neither species had a great effect on the encrusting algae (Ralfsia spp.). There was no evidence of competition for food between the two species of limpets, in contrast with results found for similar species in New South Wales.     

Population dynamics and resource utilization by larval Chironomidae (Diptera) in a backwater area of the River Danube

1. The five most abundant species of the larval chironomid community in a backwater area of the River Danube were analysed in terms of population dynamics and utilization of space from July 1985 to July 1986. The life histories of Prodiamcsa olivacea (Meigen), Cryptochironomus defectus (Kieffer), Harnischia curtilamellata (Malloch), Polypedilum laetum (Meigen) and Polypedilum scalaenum Schrank were described; the number of estimated generations per year was two for P. olivacea, H. curtilamellata, P. laetum and P. scalaenum and three for C. defectus. A decline in larval densities of P. olivacea and P. scalaenum was significantly correlated with increasing cumulative water discharge in an area close to a dam (P<0.01). 2. Spatial overlap and resource width were estimated for these chironomid species across four ranges of water depth, in three sections of the River Danube. An increasing mean spatial overlap between larval species coincided significantly with higher mean resource utilization in all cross-sections during the sampling period. Simulations of spatial overlap between randomized species associations were conducted for analyses of coexistence and dispersion between larval populations. On the basis of comparison with a neutral model, in which patches of individuals were randomly reassigned to habitat units, spatial segregation was significantly (P<0.05) different from chance for seventy-three out of 263 species pairs examined through the year. 3. An analysis of the spatial dispersion of each chironomid species (using the standardized Morisita index) suggested temporally and spatially varying patterns of dispersion, whereas the mean spatial segregation between patches of larval species might have resulted from temporary formations of random patches in this backwater area of the River Danube. 4. Spatial stability estimated as the minimum positive eigenvalue of the species matrix was significantly positively related to mean spatial segregation, indicating that a lower habitat segregation between patches of larval species pairs may lead to a destabilization of the larval species assemblage.     

Sex‐related spatial patterns of Poa ligularis in relation to shrub patch occurrence in northern Patagonia

Abstract. Poa ligularis is a dioecious species and a valuable forage plant which is widespread in the arid steppe of northern Patagonia (Argentina). The vegetation in these areas consists of a system of perennial plant patches alternating with bare soil areas defining contrasting micro‐environments. We hypothesized that (1) male and female individuals of P. ligularis are spatially segregated in different micro‐environments, (2) the intensity of spatial segregation of sexes depends on plant structure and (3) spatial segregation of sexes is enhanced by competitive interactions between the sexes within the vegetation patches. We analysed the spatial distribution of female and male individuals in relation to the spatial pattern of vegetation in two areas differing in their vegetation structure. The location of P. ligularis within patches where either male, female or both sexes occurred was also analysed. The results indicate that different patterns of spatial distribution of sexes of P. ligularis may be found at the community level depending on the dominant life forms and geometric structure of plant patches. Where patches are of a lower height, with a high internal patch cover, individuals of both sexes are concentrated within patch canopies. In sites characterized by large, tall patches and less internal patch cover suitable microsites for female and male P. ligularis occur both within and outside the patch with males located at further distances from the patch edge. Where the patch is large and tall enough to allow the establishment of males and females at relatively high numbers, males occupy the patch periphery or even colonize the interpatch bare soil. These spatial patterns are consistent with selective traits in which females better tolerate intraspecific competition than males, while males tolerate wider fluctuations in the physical environment (soil moisture, nitrogen availability, wind intensity, etc.).     

Phenotypic flexibility in the metabolic response of the limpet Cellana tramoserica to thermally different microhabitats

Temperature determines all physiological responses by limiting cellular reaction rates. Daily temperature variation differs between microhabitats, which means that subpopulations of the same species may respond differently to temperature. The aim of this study is to determine how physiological responses to temperature of the limpet Cellana tramoserica differ between limpets from variable and from stable thermal environments. Oxygen consumption and anaerobic and aerobic metabolic capacities were measured over a range of temperatures in limpets from thermally stable and variable field sites in summer and winter, and in laboratory acclimation treatments. Limpets from both variable and stable sites, showed acclimatisation of anaerobic and aerobic potentials. Limpets from stable environments, but not from variable environments, showed increased oxygen consumption in winter. Comparison of field and laboratory data showed that temperature was the signal for acclimatisation. The physiological response of C. tramoserica to temperature depends on season and microhabitat. Care must therefore be taken when conducting interspecies comparisons of response to temperature to address the confounding effects of phenotypic plasticity. Differences in physiological response to temperature in phenotypically flexible species like C. tramoserica may simply reflect individual reactions to immediate environmental conditions.     

Effects of prey density,prey size and predator size on rates of feeding by an intertidal predatory gastropod Morula marginalba Blainville (Muricidae), on several species of prey

As a prerequisite for models of foraging behaviour of the whelk, Morula marginalba Blainville (Muricidae), the effects of variation in density of prey on the rate of feeding of the predator were examined in field conditions for three coexisting species of prey. Densities of prey used were those at which the prey, two limpets and a barnacle, occurred naturally in the rocky intertidal habitat.Large limpets, Cellana tramoserica (Sowerby) can resist attacks by predatory gastropods by raising the mantle over the outside of the shell. These experiments showed that no C. tramoserica were killed by Morula marginalba even at very great densities and with no alternative prey present. For the small limpet Patelloida latistrigata (Angas), one of the whelk's most highly preferred prey, juveniles were eaten 1.4 times as fast as adults. Fitting the random predator equation gave greater attack coefficients and shorter handling times for juvenile than adult limpets.Sizes of both predator and prey affected rates of eating barnacles, Tesseropora rosea (Krauss), but not in a simple way. Whelks of 15-mm aperture length ate adult barnacles 4.2 times faster than did 12-mm whelks, but there was no significant difference in the rates at which the two sizes of snail ate juvenile barnacles.Rates of feeding on T. rosea and Patelloida latistrigata increased significantly with prey density. These results form a basis for including the density of prey in models of spatial dispersion of the predatory gastropod Morula marginalba.     

Patterns of coexistence between two mesocarnivores in northern Patagonia in the presence of invasive hares and anthropogenic disturbance

We analysed coexistence patterns between two mesocarnivores, Geoffroy's cat (Leopardus geoffroyi: Felidae) and culpeo fox (Pseudalopex culpaeus: Canidae), in northern Patagonia, Argentina. We examined spatial distribution influenced by land cover, anthropogenic disturbance and invasive hare presence, and analysed temporal activity patterns and dietary composition. If competitive exclusion accounts for carnivore coexistence in this system, we predicted segregation would occur in one or more of these aspects as a mechanism for coexistence. We performed camera trapping in Nahuel Huapi National Park, from February to May 2012 and January to April 2013. Using camera detections, we analysed spatial patterns with co‐occupancy modelling and temporal patterns by fitting kernel density estimates and measuring overlap. We performed a dietary meta‐analysis using available literature and performed a discriminant function analysis of diet categories between species. We observed high spatial and temporal overlap between Geoffroy's cats and culpeo foxes. We found no evidence of segregation in relation to land cover occupancy, anthropogenic disturbance, invasive hare occurrence or activity patterns. Though both species consumed predominantly small and medium mammals, Geoffroy's cats consumed more birds, reptiles and amphibians, and culpeo foxes consumed more large mammals, carrion and plant material; coexistence between these two carnivores in this Patagonian protected area appears to be facilitated by diet segregation.     

Randomness and aggregation: analysis of dispersion in an epiphytic chironomid community

1. Patterns of dispersion in a chironomid community on the submersed macrophyte Myriophyllum spicatum were analysed. 2. Random dispersion commonly occurred throughout the year, with an average of 40% of all species being observed with random spatial patterns. The frequency of occasions with random dispersion varied among chironomid species, ranging from 3.3% in Rheotanytarsus curtistylus to 56% in Thienemanniella majuscula. 3. Estimates of the negative binomial parameter k show that 26% of all cases demonstrate strong aggregation (0 < k < 1.0) while nearly half (47%) have quasi-random dispersion. Interspecific variation in k was not significant statistically when all the species were considered together, although a pairwise comparison between two abundant species Tvetenia calvescens and Rheotanytarsus curtistylus demonstrated a marginally significant difference. When different instars were compared, the percentage frequency of strong aggregation (0 < k < 1.0) declined from first instars (49%) to later instars (II-38%, III-24% and IV-27%). 4. Variance/mean and m*—m regressions (m* is Lloyd's mean crowding statistic and m is the sample mean) both fitted the data well, but there was little indication of significant interspecific variation in parameter values, particularly the slope of regression. 5. Dispersion patterns were examined along with the analysis of spatial overlap in this community. Forty-two per cent of species-pairs with reduced spatial overlap (spatially ‘segregated’ pairs) contained one or both species with random dispersion, while the corresponding value for spatially unsegregated pairs was 57%. This suggests that spatial segregation is not necessarily caused by strong, independent aggregation of both species. Comparing spatially segregated vs. unsegregated pairs, the former tend to have one species with a stronger tendency of aggregation than species of the latter. 6. Patterns of dispersion observed were considered in the light of ‘random patch formation’. Random patch formation emphasizes the stochasticity of patch-forming processes as well as the stochastically dynamic nature of resultant patches. Unlike terrestrial drosophilid assemblages, where strong aggregation is a predominant pattern, this chironomid community demonstrates widely varying degrees of dispersion with high occurrence of randomness, reflecting the stochasticity of dispersal and recolonization processes. It is suggested that, in terms of species coexistence, more emphasis should be placed on stochasticity rather than on aggregation in this type of community.     

Spatial indicators of plant community assembly from a 453-year sand-dune chronosequence

Background : We explored evidence for spatial patterning in vegetation across a sand-dune chronosequence spanning 453 years of primary succession to test for indications of a temporal signature of niche versus neutral processes during community assembly.

Methods : The study was conducted on a series of dune-capped beach ridges located in Wilderness State Park in Emmet County of northern Lower Michigan, United States (45° 43′ N, 84° 56′ E). Nearest-neighbour survey data were subjected to species dispersion and association analyses.

Results : Both species richness and species evenness showed the classic ‘hump-shaped’ relationship across successional time, suggesting that competition is an important force in structuring this community. However, the only evidence of intra-specific aggregation occurred in the youngest dune. Older dunes displayed random spatial structure and the oldest dunes tended toward a mix of random patterns and intra-specific segregation, i.e. with nearest neighbours more likely than by chance to be inter-specific. In addition, the frequency of species having lower neighbour-species richness than expected by chance was higher in younger, not older dunes. Species with significantly low neighbour richness (based on absolute nearest neighbours) on later dunes tended to have a high proportion of nearest neighbours that were intra-specific.

Conclusions : The data provided no evidence that resident species in later succession have greater intra-specific aggregation or more niche differentiation based on horizontal niche space. The data are more consistent with theories of community assembly involving competitive equivalence and/or effects of facilitation.     

Intertidal zonation of prosobranch gastropods: Analysis of densities of four co-existing species

The densities of populations of Nerita atramentosa Reeve, Austrocochlea constricta Lamarck, Bembiciuin nanum (Lamarck), and Cellana tramoserica (Sowerby) were recorded in 0.25 m² quadrats on landward and seaward halves of a sheltered and a moderately exposed rock platform from June 1972 to June 1973. The data have been subjected to variance and multiple regression analysis. Differences between densities of each species on the two shores and in the two areas of each shore are discussed with respect to the regressions on littoral height. Densities of Nerita on the sheltered shore, Austrocochlea and Bembicium on both shores, and Cellana, on the exposed shore, increased with height. The density of Cellana on the sheltered shore decreased as height increased. The density differences in the landward and seaward areas on the two shores were attributable to the effect of height for Nerita and Austrocochlea. Height was the major variable affecting the density of these two species on the sheltered shore. Bembicium and Cellana densities on both shores, and Austrocochlea density on the exposed shore, are not primarily determined by height.On the sheltered shore, Nerita and Cellana increased in density with increasing cover of water in rock pools. On the exposed shore, Bembicium and Cellana became sparser with increasing water cover. Bembicium on both shores increased in density with increasing cover of the alga Peyssonelia, as did Cellana on the exposed shore. Cellana and Austrocochlea tended to increase in density with Nerita on the sheltered shore. Cellana tended to be sparse where Bembicium was dense on the exposed shore. The biological effects of substrata and other species on the densities of each species remain unexplained.Significant time effects were found for variations in the densities of Nerita and Cellana on the sheltered shore and of Austrocochlea and Cellana on the exposed shore. The seasonal change in density of Nerita was parabolic with a minimum in summer, and is consistent with the known period of recruitment of this species. The lack of significant time effects on density of Austrocochlea on the sheltered shore is consistent with the known continuous recruitment of this species. The declining density of Austrocochlea on the moderately exposed shore was found, by size-frequency analysis, to be due to reduced recruitment compared with surrounding shores. The density of Cellana on both shores showed seasonal changes, with a maximum in summer, which is consistent with the known period of recruitment in Cellana. The lack of seasonal changes in density of Bembicium is briefly discussed.This type of multifactor analysis identifies biological interactions affecting variation in density of these species. It also corroborates the results of other ecological investigations and determines the relative importance of a variety of physical and biological factors in the distribution of density of intertidal gastropods.     

Effects of floral neighborhood on seed set and degree of outbreeding in a high-alpine cushion plant

Plants flowering together may influence each other’s pollination and fecundity over a range of physical distances. Their effects on one another can be competitive, neutral, or facilitative. We manipulated the floral neighborhood of the high-alpine cushion plant Eritrichium nanum in the Swiss Alps and measured the effects of co-flowering neighbors on both the number of seeds produced and the degree of inbreeding and outbreeding in the offspring, as deduced from nuclear microsatellite markers. Seed set of E. nanum did not vary significantly with the presence or absence of two Saxifraga species growing as near neighbors, but it was higher in E. nanum cushions growing at low conspecific density than in those growing at high density. In addition, floral neighborhood had no detectable effect on the degree of selfing of E. nanum, but seeds from cushions growing at low conspecific density were more highly outbred than seeds from cushions at high density. Thus, there was no evidence of either competition or facilitation between E. nanum and Saxifraga spp. as mediated by pollinators at the spatial scale of our experimental manipulation. In contrast, the greater fecundity of E. nanum cushions at low density was consistent with reduced intraspecific competition for pollinators and might also represent a beneficial effect of highly outbred seeds as brought about by more long-distance pollinator flights under low-density conditions.     

Reproductive effort in two subtidal populations of the limpet,Patelloida mufria

Summary The reproductive cycle and reproductive effort (RE) of 2 populations of the acmaeid limpet Patelloida mufria were investigated in a subtidal region of N.S.W., Australia. These two populations had vastly different densities, rates of growth and mortality. Nonetheless, their patterns of spawning were remarkably similar with 5 spawning periods during the year of monitoring, and their values for RE were very similar; individuals from each population spawned about 35% of their somatic weight each 6 months. This does not, therefore, conform with theories that the RE of populations should be correlated with their rates of mortality. The results for P. mufria were compared with those of 4 populations of Cellana tramoserica and 2 of P. alticostata which had been studied previously at this location. There was still no general correlation between RE and mortality, but by only using certain combinations of these 8 populations (i.e. only one from each species at a time), positive or negative correlations could be obtained. This illustrated the need to ensure that intraspecific variability is accounted for in any analysis of life-history strategies. Finally, it was concluded that instead of the absolute vale of RE being important, the amount of flexibility in this and other traits may represent the real difference in strategies amongst species.     

Distribution and abundance of the acmaeid limpet,Patelloida latistrigata,and its interaction with barnacles

Summary At Cape Banks, New South Wales, adults of the small intertidal limpet, Patelloida latistrigata occur exclusively in the barnacle zone, and are primarily associated with the barnacle, Tesseropora rosea. Limpet density increases with barnacle density. Juvenile limpets can be found throughout the barnacle zone, and on patches of bare rock that may be temporarily available at lower levels on the shore. The failure of juveniles to survive and grow in places other than among Tesseropora is due to a combination of factors. These include desication at high levels on the shore, smothering by rapidly growing algae low on the shore, and the grazing activities of the larger limpet, Cellana tramoserica. These latter two factors also reduce the survival of experimentally transplanted adult Patelloida: algae by covering the substratum and smothering the limpets, and Cellana by outcompeting them for food. The density of Cellana is greater on patches of bare rock than among barnacles, and these large limpets may be unable to move and feed effectively over the irregular surface created by Tesseropora. Patelloida, however, is small enough to feed over and among these barnacles, and hence has a refuge from competition with Cellana. Barnacles may also provide shelter from the effects of desiccation and strong wave action, and thus increase the survival of juvenile Patelloida. By being associated with barnacles, however, Patelloida becomes vulnerable to intermittent predation by the whelk, Morula marginalba. This effect may be serious enough to eliminate small local populations of limpets, either by direct predation or by removing the refuge-providing barnacles. The association with barnacles may also limit the maximum size to which Patelloida can grow. Patelloida is not always found with Tesseropora, and adults in different localities can be found in association with other sessile organisms. It may be argued that small species of limpets require a spatial refuge from physical and/or biological pressures. To examine this hypothesis, the relationship between Patelloida and Tesseropora is compared to other published accounts of limpets with specialised modes of life.