Effects of height on the shore and complexity of habitat on abundances of amphipods on rocky shores in New South Wales, Australia

Patterns of abundance of epifaunal crustaceans were compared between two common brown algae in intertidal rock pools on a shore near Sydney, New South Wales, Australia. Amphipods were significantly more abundant on Sargassum globulariaefolium (which was most common low on the shore) than on Hormosira banksii (more common at mid-shore levels). Experiments tested the hypotheses that height on the shore and structural complexity of the algae were the main factors influencing patterns of abundance of amphipods. Reciprocal transplants using defaunated plants and similar experiments using artificial plants allowed simultaneous tests of these hypotheses. The abundance of a common intertidal amphipod, Sunampithoe graxon was significantly smaller on Sargassum transferred from low to mid-tidal level of the shore than on any other treatments, indicating that height on the shore affects numbers of this species. There were no Sunampithoe on Hormosira even when transplanted to low shore areas, indicating that other factors, apart from the height on the shore, are also important for this amphipod. Another common amphipod, Hyale maroubrae was generally found in greatest abundances on Hormosira transplanted from mid- to low shore and on Sargassum low on the shore, again indicating the importance of height on the shore. There was no difference in abundance of amphipods on ‘complex’ and ‘simple’ artificial plants indicating that structural complexity (as defined in this study) did not influence these amphipods.     

The mechanics of predation by the shore crab,Carcinus maenas (L.), on the edible mussel,Mytilus edulis L.

Summary Mechanical aspects of predation by the shore crab, Carcinus maenas, on the edible mussel, Mytilus edulis, were examined. The shore crabs from the population studied utilized five distinct, largely size-related, mussel-opening techniques. Crushing the mussel umbone appeared the most successful opening method for medium-sized prey. Small mussels were crushed outright and large mussels could be opened by a slow, uneconomical, boring technique. The strengths of mussels, from an exposed shore, were tested under compression in four separate planes to determine the loads a crab would need to apply to crush the shells outright and the mechanical properties of mussels. Little inter-plane variability in compressive strength was observed, although intra-plane variability appeared high. The compressive strengths of mussels from a sheltered shore were found to be significantly higher than those from the exposed shore in the plane tested. A strain gauge was embedded in a mussel shell enabling the pattern and magnitude of forces produced by crab chelae in opening a mussel to be studied. The crab's chelae did not appear overwhelmingly strong when compared directly to the compressive strength of the crab's preferred mussel sizes. It is, therefore, postulated that crabs usually seek out and exploit weak spots in the umbone of mussels by trial and error, eventually breaking through the shell by a cumulative process of extending minute fractures in the shell substructure.     

Beach morphology and food web structure: comparison of an eroding and an accreting sandy shore in the North Sea

Food web components and inorganic nutrients were studied on two sandy shores of the adjacent barrier islands of Sylt and R?m? in the North Sea, differing in morphodynamics. Implications of high and low wave energy on the food web structure were assessed. The Sylt shore represents a dynamic intermediate beach type, while the R?m? shore is morphologically stable and dissipative. On the steep-profiled, coarse-grained Sylt shore, strong hydrodynamics resulted in erosion and high fluxes of organic material through the beach, but prevented any storage of food sources. In contrast, the flat-profiled, fine-grained R?m? shore, with low wave energy and accretion, accumulated organic carbon from surf waters. At Sylt, oxic nutrient regeneration prevailed, while anoxic mineralization was more important at R?m?. Macrofauna on the Sylt shore was impoverished compared with the community at R?m?. Correspondingly, abundances of epibenthic predators such as shrimps, crabs, fish, and shorebirds were also lower at Sylt. Meiofauna was abundant on both shores, but differed in taxonomic composition. Several major taxa were represented in fairly equal proportions of individual numbers on the well-oxygenated Sylt shore, while nematodes strongly dominated the assemblage at R?m?. Thus, on cold-temperate, highly dynamic intermediate shores with high wave energy and subject to erosion, the "small food web" dominates. Organisms are agile and quickly exploit fresh organic material. Larger organisms and nematodes abound under stable, dissipative and accreting shore conditions, where some food materials may accumulate and zoomass builds up to support numerous visitors from higher trophic levels. Electronic Publication     

Development of on‐shore behavior among polar bears (Ursus maritimus) in the southern Beaufort Sea: inherited or learned?

Polar bears (Ursus maritimus) are experiencing rapid and substantial changes to their environment due to global climate change. Polar bears of the southern Beaufort Sea (SB) have historically spent most of the year on the sea ice. However, recent reports from Alaska indicate that the proportion of the SB subpopulation observed on‐shore during late summer and early fall has increased. Our objective was to investigate whether this on‐shore behavior has developed through genetic inheritance, asocial learning, or through social learning. From 2010 to 2013, genetic data were collected from SB polar bears in the fall via hair snags and remote biopsy darting on‐shore and in the spring from captures and remote biopsy darting on the sea ice. Bears were categorized as either on‐shore or off‐shore individuals based on their presence on‐shore during the fall. Levels of genetic relatedness, first‐order relatives, mother–offspring pairs, and father–offspring pairs were determined and compared within and between the two categories: on‐shore versus off‐shore. Results suggested transmission of on‐shore behavior through either genetic inheritance or social learning as there was a higher than expected number of first‐order relatives exhibiting on‐shore behavior. Genetic relatedness and parentage data analyses were in concurrence with this finding, but further revealed mother–offspring social learning as the primary mechanism responsible for the development of on‐shore behavior. Recognizing that on‐shore behavior among polar bears was predominantly transmitted via social learning from mothers to their offspring has implications for future management and conservation as sea ice continues to decline.     

Spatial variation and temporal stability of littoral water temperature relative to lakeshore morphometry: environmental analysis from the view of fish thermal ecology

Lakeshore developments change the physicochemical properties of the underwater environment by altering shore morphometry, which may have significant effects on spatial variation and temporal stability in water temperature. Spatiotemporal temperature changes are costly to fish in terms of subsequent thermoregulatory behavior and acclimation; therefore, thermal conditions have a heavy impact on the biological function of fishes. Spatiotemporal variation and stability of water temperatures along cross-shore transects in the littoral zone (within 100 m from shore) were monitored and compared on two lakeshores with different cross-shore depth profiles. One shore was associated with a retaining wall and a relatively deep, flat bottom (steep shore), whereas the other extended offshore at a gentle gradient (gentle shore). Water temperature was more spatially variable on the gentle shore than the steep shore [1.44 ± 0.47 and 0.20 ± 0.14°C (mean ± SD), respectively], but a stable temperature range (i.e., the range of temperatures continuously observed on each shore for 48 h) was maintained only on the gentle shore during seasonal temperature decline. These results suggest that gentle shores have higher potential to provide a wider range of thermal options, allowing fish to fine-tune thermoregulatory behavior and acclimate more efficiently to temperature changes.     

Ecology and behavior of Gecarcoidea natalis, the Christmas Island red crab, during the annual breeding migration.

The terrestrial crab Gecarcoidea natalis is endemic to the forests of Christmas Island but must migrate each year to the coast to breed. During 1993 and 1995, radio-tracking, mark and recapture, and counting methods were used to establish the routes, walking speeds, direction of travel, and destinations of migrating crabs, as well as crab numbers and distribution. The density of crabs ranged from 0.09 to 0.57 crabs per square meter, which gave a population estimate of 43.7 million adult crabs on the island. During the dry season the crabs were relatively inactive but on arrival of the wet season immediately began their migration. The crabs generally walked in straight lines, and most crabs from around the Island traveled toward the northwest shore instead of simply walking toward the nearest shore. The maximum recorded distance walked by a red crab in one day was 1460 m, but the mean was 680 m per day in 1993 and 330 m in 1995. Comparing the 1993 and 1995 study seasons, there was a 3-week difference in the timing of the start of the migration, but the spawning date was fixed by the lunar phase and took place 17 to 18 days after mating. In 1993, late rain prompted a "rushed" migration and crabs walked directly to their shore destinations; in contrast, in 1995 most crabs made stops of 1 to 7 days during the downward migration. By giving the crabs a chance to feed along the way and minimizing the time that the population was concentrated near the shore, these stops may be important in ensuring that the animals have enough food after the long dry season. Furthermore, this behavior implies that the crabs are able to judge how far away they are from the shore during the migration.     

Infection of the mollusc Littorina saxatilis with parthenites of trematodes and their impact on a shell form: analysis of populations inhabiting the littoral coast of the White Sea

12 rocky shore populations of Littorina saxatilis from three islands of Chupa Inlet (Kandalaksha Bay, White Sea) were examined for infection with trematodes. Morphometric characters (6 indexes of the shell and aperture shape) of molluscs were investigated for all these populations. Exposed and sheltered sites were considered at every island and high and low littoral samples were fulfilled at every site. Seven species of trematodes, Podocotyle atomon, Cryptocotyle lingua, Renicola sp., Himasthla sp., Microphallus piriformes, M. pygmaeus, M. pseudopygmaeus, were found. Uneven distribution of trematodes was confirmed by log-linear analysis. Sheltered populations of L. saxatilis have the greater infection prevalence than exposed ones. This is due to the heavy infection with M. piriformes and M. pygmaeus. The prevalences by these trematodes are up to 52.97% and 27.16% respectively in sheltered populations of the host. The prevalence of M. piriformes tend to be higher at the upper shore level of sheltered sites. In a contrast, the prevalence of M. pygmaeus is significantly higher at the low part of such sites. Factor analysis shows a significant association of the indices of L. saxatilis shell shape with three factors. The first one is associated with the "elongation" of a shell and reveals L. saxatilis from the exposed rocky shore to be more elongated than the molluscs from sheltered sites. The second one is connected with the "aperture shape" index. There is an association of this factor with the shore level position of samples. The third factor reflects the affect of trematodes on the shell shape. The molluscs infected with M. piriformes show "elongated" shell shape and relatively smaller aperture. Shall peculiarities of the hosts infected with M. piriformes and M. pygmaeus are somewhat different. The results of the factor analysis is justified by the series of analysis of variances on the values of shell indices (MANOVA) according to the factors "exposure", "shore level" and "infection".     

Small- and broad-scale patterns of distribution of the upper-shore littorinid Nodilittorina pyramidalis in New South Wales

Abstract mall-scale (within-shore) and broad-scale (among-shore) patterns of distribution, abundance and size structure of Nodilittorina pyramidalis were measured on a number of shores in New South Wales. This species showed significant patchiness in distribution and size structure. The mean density and mean shell length of snails differed among patches of shore only a few metres apart, among different heights on the shore and from shore to shore. Differences in densities appeared to be strongly correlated to the distribution of particular microhabitats, especially cracks, crevices and pits within the rock surface. This relationship and the presence or absence of these microhabitats appeared to determine the patterns of distribution of this species within and among different shores. The sizes of snails were not correlated with densities of Nodilittorina nor with the coexisting species, Littorina unifasciata, in the populations examined. Processes influencing the distribution and abundance of Nodilittorina were not investigated, but the patterns of distribution found on these shores suggest that recruitment was limited to areas of topographic complexity, ft also seems likely that limited dispersal by the adult snails retain them within restricted patches on the shore.     

Allochthonous aquatic insects increase predation and decrease herbivory in river shore food webs

Rivers produce an abundance of aquatic insects that traverse land, where they can have bottom-up effects on predators, who, in turn, can have top-down effects on terrestrial herbivores. This effect can cascade down to plants. These trophic relationships were demonstrated in a field of stinging nettles, Urtica dioica , along a river in Germany. At the shore compared to similar microhabitats 30–60 m away the abundance and biomass of: midges were highest, spiders were also highest, while herbivorous leafhoppers were lowest. At the shore, nettle plants were less damaged by herbivores and thus had less regrowth. Spiders regularly captured both aquatic midges as well as terrestrial leafhoppers and they captured more individuals of both groups at the shore than further away. Midges supported high densities of shore spiders. This was inferred from correlation of distribution and diet in the absence of other environmental gradients. Removal of spiders from experimental plots caused leafhoppers to increase at the shore, causing more plant damage. These effects were not evident at spider-removal sites away from the shore. This demonstrated that spiders depressed leafhoppers and decreased herbivory on plants only at the shore. It is concluded that aquatic insects had a bottom-up effect on spiders and that this subsidy facilitated a top-down effect that cascaded from spiders to leafhoppers to plants. Similar effects would explain the distribution of arthropods along many rivers. Allochthony connects river food webs with shore food webs, making both components essential for each other.     

Factors shaping the range-size frequency distribution of the endemic fish fauna of the Tropical Eastern Pacific

Aim To assess the effect of habitat fragmentation and isolation in determining the range‐size frequency distribution (RFD) of the shorefish fauna endemic to a discrete biogeographical region. Location The Tropical Eastern Pacific (TEP). Methods Habitat isolation represents the separation between oceanic islands and the continental shore of the TEP and habitat fragmentation the degree of spatial continuity of habitats (i.e. reefs, soft bottom, nearshore waters) along the continental coast of the TEP. The effects of habitat isolation and fragmentation were quantified by comparing the RFDs of (1) the species found on oceanic islands vs. the continental shore, and (2) species on the continental shore that use different habitat types. Results The RFD of the entire TEP fauna was bimodal, with peaks at both small‐ and large‐range ends of the spectrum. The small‐range peak was due almost entirely to island species and the large‐range peak due mainly to species found in both the continental shore and oceanic islands. RFDs varied among species using different habitats on the continental shore: reef‐fishes had a right‐skewed RFD, soft‐bottom species a flat RFD, and coastal‐pelagic fishes a left‐skewed RFD. Main conclusions Variation in dispersal capabilities associated with habitat isolation and fragmentation in the TEP appears to be the main mechanism contributing to differences among RFD structure, although variation in tolerances arising from the dynamic regional environment may contribute to some patterns. Because diversity patterns are strongly affected by RFD structure, it is now evident that the insular and continental components of a fauna should be treated separately when analysing such patterns. Furthermore, contrasts in RFD structure among species using different habitats demonstrate that a full understanding of the causes of diversity patterns requires analyses of complete regional faunas in relation to regional geography.     

Ecology of the intertidal snail Littorina acutispira Smith

Littorina acutispira Smith, a minute gastropod of < 3 mm shell height, lives at great densities in pools and on rock-surfaces at the highest levels of sea-shores in New South Wales. Populations from pools and dry areas were sampled on two shores for 18 months to investigate seasonal changes in density, size-structure, rates of growth and reproductive biology of the snails. Densities of snails increased between February and May, due to an influx of juveniles, and then declined until the following February, when they increased again. The decrease in density was due to the death of the largest snails at the end of summer, and the mortality of medium-sized snails between June and January. Longevity was estimated as 1–2 yr, but most individuals died by ≈ 16 months from settlement on the shore. Newly-settled snails grew to merge in size with those of the previous year's population by winter. During the summer months, the rate of growth of snails from a sheltered shore was greater than that of snails on a shore exposed to wave-action. Laboratory experiments revealed that this could be attributed to the presence of better quality food, or food in greater abundance on the sheltered shore, compared with the exposed shore.During winter months, but not during the summer, snails from dry areas grew more slowly than those from pools. An experiment demonstrated that some snails from dry areas might be able to compensate for reduced periods of feeding by being able to feed faster when submersed. This could not explain the differences in natural rates of growth.L. acutispira bred from October–November to March–April. Spawning in the laboratory was greatest during late summer (January to March). The percentage of mature oocytes in the gonads was small in winter and increased in early summer. Among the largest-sized snails, females outnumbered males. Two experiments, on unsexed and pre-sexed snails, demonstrated that the biased sex-ratio of the largest snails was due to faster growth by females.There was a greater density of snails on the exposed shore, which was correlated with the presence of barnacles. When barnacles were removed from experimental areas, the density of the snails declined within 24 h. This suggested that barnacles provided a refuge from wave-shock, rather than shelter from desiccation or high temperature. In laboratory experiments, snails were exposed to higher temperatures and less humidity than they would normally encounter on the shore. There was negligible mortality of small or large snails after 24 h of these conditions.This minute species grows quickly, recruits annually and has a short life-span. This type of life-history is discussed in comparison with similar small species from other habitats.     

ASYMMETRICAL DEVELOPMENTAL PLASTICITY IN AN INTERTIDAL SNAIL

Field and laboratory experiments were used to investigate the variation and phenotypic plasticity in the adhesive abilities of the intertidal snail Nucella lapillus between high- and low-wave-energy environments. Whelks from an exposed coast produced a larger pedal surface area and were more resistant to dislodgement than were similar-sized individuals from a protected shore. Tenacity (g/cm²) was similar between individuals from exposed and protected shores, indicating that variation in resisting dislodgement was solely a function of pedal surface area. Whelks from exposed and protected shores did not differ in pedal surface area as they emerged from egg capsules or when reared in the laboratory under uniform conditions, suggesting that variation between populations does not represent genetic differentiation. Individuals from high-and low-wave-energy environments reared intertidally produced a larger pedal surface area than did those reared in the laboratory. The extent to which pedal surface area increased corresponded to the intensity of wave action. These findings suggest that pedal surface area is a highly plastic character modulated or induced by the water turbulence accompanying breaking waves. A reciprocal-transplant experiment confirmed this notion but revealed an asymmetry in the plasticity. Snails from the protected site transplanted to an exposed shore formed a much larger pedal surface area than did controls reared on the protected shore. In contrast, whelks from a wave-swept shore transplanted to a protected shore differed little from their controls reared on the exposed shore. The asymmetric response parallels a possible asymmetry in the risks of acclimating to a temporally unpredictable environmental cue, such as wave action.     

The interplay of physical and biological factors in maintaining mid-shore and low-shore assemblages on rocky coasts in the north-west Mediterranean

This study examined the interactive effects of grazing by limpets and inclination of the substratum in maintaining differences between mid-shore and low-shore assemblages of algae in the northwest Mediterranean, at different scales of space and through time. Alternative models leading to different predictions about these effects were proposed and tested. Limpets were excluded by fences from areas of the substratum at mid levels on the shore. The response of algal assemblages to this manipulation was compared with control and enclosure plots at the same level, and with unmanipulated plots in the low shore where limpets are less abundant. The effects of limpets were examined at several replicated sites (0.1–4 km apart) for each slope of the substratum (nearly horizontal vs vertical), at different locations (hundreds of kilometres apart) and at different times. Individual taxa responded differently to limpet exclusion. The percentage cover of the coarsely branched and filamentous algae increased significantly in exclosure plots, in some loser reaching values found on the low shore. These patterns, however, varied greatly from shore to shore and significant effects were found both on horizontal and vertical substrata. Multivariate analyses indicated that grazing by limpets accounted for about 20% of the differences between mid-shore and low-shore assemblages. This effect was independent of substratum inclination and was consistent in space and time, suggesting that physical conditions were not as stressful for macroalgae on vertical substrata as initially supposed. Variable recruitment of algae is proposed as a possible explanation for the lack of consistency in the effects of limpets at the scale of the shore. The results of this study emphasize the need for multiple-scale analyses of the interactive effects of physical and biological factors to understand the organization of natural assemblages. Received: 22 June 1999 / Accepted: 15 November 1999     

Territory size and distribution in the African fish eagle

The fish eagles along 32 km of shoreline were counted regularly from a launch. The territory size was found to be 0.6 km of shore with a density of 3.5 eagles per km. A single aerial count along 323 km of shore returned a density of 1.9 eagles per km with a territory size of 1.0 km of shore. The distribution of fish eagles is related to the relative availability of trees as perches and probably also to the abundance of fish. The quantity of fish taken by the fish eagles is shown to be negligible compared either with the total production or with the yield harvested by man.     

A refutation of critical tidal levels as determinants of the structure of intertidal communities on British shores

Early hypotheses to account for the pattern of zonation and vertical distribution of species on rocky shores in Britain invoked the concept of critical tidal levels. These levels were at heights on the shore where the upper or lower boundaries of distribution of a number of species coincided. Critical levels were correlated with heights on the shore where there were changes in the rate of change, with height, of the annual proportion of the time spent emersed. This was calculated from predicted tide tables. One implications of this was that different assemblages of intertidal species were present between particular levels on the shore. Futhermore, this hypothesis depended on patterns of vertical distribution of several species being identically controlled by physical factors associated with the rise and fall of the tide. This conflicts with more recent hypotheses, based upon experimental evidence, which include the effects of biological interactions among species on their patterns of distribution.The methods of calculation of the annual emersion curve were approximate, with a high degree of extrapolation. The hypothesis that critical levels exist has not been quantitatively tested.New, more accurate calculations of the emersion curve, from predicted tidal heights, indicate a smooth, monotonic curve against height on the shore. This eliminates the possibility of correlation between the coincident boundaries of distribution of a group of species and a height at which the curve changes in slope.The upper and lower boundaries of species were recorded in transects on five shores in different parts of Britain. If critical level exist, the upper or lower boundaries of a set of species must depart from random dispersion up and down the shore, to be clumped or aggregated at the critical levels. A test for non-random dispersion of the boundaries indicated no significant departure from random on any of the shores sampled.There is thus no evidence that critical tidal levels exist, because there is no evidence that the upper, or lower, boundaries of vertical distribution of intertidal species are in any way aggregated. There is no possibility of correlation between changes of slope of the emersion curve at particular heights on the shore and the patterns of distribution of intertidal species on British shores. The hypothesis of critical tidal levels must be abandoned and the rôle of the effects of tidal rise and fall on the vertical distribution of intertidal species must be re-examined in any new hypotheses to account for observed patterns of zonation.     

Size selection by a gape‐limited predator of a marine snail: Insights into magic traits for speciation

The intertidal snail Littorina saxatilis has repeatedly evolved two parallel ecotypes assumed to be wave adapted and predatory shore crab adapted, but the magnitude and targets of predator‐driven selection are unknown. In Spain, a small, wave ecotype with a large aperture from the lower shore and a large, thick‐shelled crab ecotype from the upper shore meet in the mid‐shore and show partial size‐assortative mating. We performed complementary field tethering and laboratory predation experiments; the first set compared the survival of two different size‐classes of the crab ecotype while the second compared the same size‐class of the two ecotypes. In the first set, the large size‐class of the crab ecotype survived significantly better than the small size‐class both on the upper shore and in the laboratory. In the second set, the small size‐class of the crab ecotype survived substantially better than that of the wave ecotype both on the upper shore and in the laboratory. Shell‐breaking predation on tethered snails was almost absent within the lower shore. In the laboratory shore crabs (Pachygrapsus marmoratus) with larger claw heights selected most strongly against the small size‐class of the crab ecotype, whereas those with medium claw heights selected most strongly against the thin‐shelled wave ecotype. Sexual maturity occurred at a much larger size in the crab ecotype than in the wave ecotype. Our results showed that selection on the upper shore for rapid attainment of a size refuge from this gape‐limited predator favors large size, thick shells, and late maturity. Model parameterization showed that size‐selective predation restricted to the upper shore resulted in the evolution of the crab ecotype despite gene flow from the wave ecotype snails living on the lower shore. These results on gape‐limited predation and previous ones showing size‐assortative mating between ecotypes suggest that size may represent a magic trait for the thick‐shelled ecotype.     

Seasonal growth patterns of Saccharina latissima (Phaeophyceae,Ochrophyta) in a glacially‐influenced subarctic estuary

Global climate warming is exacerbating the melting of glaciers in Arctic and subarctic nearshore regions. Glacial discharge causes increases in sedimentation, abrasion of organisms, and sand/silt cover along with lowered light intensity, salinity, nitrate and hard substrate cover. These effects can have deleterious consequences on foundation species, such as the kelps that provide important habitat structure and support tightly‐linked food webs. The purpose of this study was to determine if the kelp, Saccharina latissima, from a glacially‐influenced and an oceanic shore in a subarctic Alaskan estuary exhibits differing seasonal growth patterns in response to its environment. Reciprocal in situ shore transplant studies examined seasonal patterns in growth, physiological competence (as maximum quantum yield), morphology and storage product levels (mannitol) of S. latissima. In situ growth was seasonally different at the two shore locations, with a shorter growing season at the glacially‐influenced shore. During the glacial melt season, the thalli at the two shore locations were morphologically distinct. Mannitol levels were typically higher in thalli from the oceanic site, with generally low mannitol levels at the end and the beginning of the growing season on both shores. Maximum quantum yield was consistently high (≥0.7) at both shore sites and did not vary seasonally. Growth rates of glacially‐influenced transplants to the oceanic shore suggest that the glacially‐influenced population has a different seasonal growth pattern from that of the oceanic shore site, which seems to be genetically fixed or based on differences in gene expression. It appears that S. latissima is a highly resilient species, partly due to high phenotypic plasticity, which may have led to genetic fixation under persistent glacial conditions.     

The use of drinking sites, wallows and salt licks by herbivores in the flooded Middle Zambezi Valley

In the unflooded Middle Zambezi Valley, water was available for drinking in zoogenous pools in the deciduous woodlands of the escarpment hills only during, and for a short period after, the rains. For 5 or 6 months each year these pools were dry. During the dry season the majority of large mammals drank either from remnant pools in stream beds, or migrated to the Zambezi flood plain where water was available in the Zambezi River and in pools on the flood plain. In the part of the valley filled by Lake Kariba the Zambezi and its flood plain have been inundated, and the lake shore now lies permanently in the deciduous woodlands of the escarpment hills. Although the former riverine dry-season water sources have been removed, new sources are available along the lake shore. The suitability of the shore as a drinking site is affected by its exposure to wave action, its pitch, its soil type, and the vegetation type abutting on it, as well as by the preferences of the species using it. Elephant modify drinking sites on the shore by digging water-filled holes at the water's edge in clay soil, and elephant, buffalo and rhinoceros create wallows in similar situations. The holes dug by elephant are much used by antelope, especially impala, for drinking. By analogy with elephant-created pools in the woodlands, these zoogenous pools on the lake shore may have been a source of sodium for species drinking there. Such sources were not widely available in the dry season in the unflooded Zambezi Valley.     

Foraging decisions of North American beaver (<Emphasis Type="Italic">Castor canadensis</Emphasis>) are shaped by energy constraints and predation risk

Foraging herbivores have to trade-off between energy requirements and predator avoidance. We aimed to study the relative roles of these factors in beavers (Castor canadensis) when foraging on land. We hypothesized that beavers were able to assess the risk of predation by using two main cues: the distance from the water and the presence or absence of predator odors. First, we studied the food selection of beavers in relation to distance from the water in natural settings. The transects were made at beaver ponds, and the diameter, species, and distance from the shore of intact and beaver-cut trees were recorded. Secondly, we placed rows of aspen sticks (Populus tremula) perpendicular to the shore around beaver ponds, and treated each row with a neutral, alien, or wolf odor. We found that aspen, downy birch (Betula pubescens), and speckled alder (Alnus incana) were the preferred tree species. More of these species were cut close to the shore, and cut trees were smaller further away from the shore, except in the case of aspen. In the experiment, most of the aspen sticks were taken close to the shore. As predicted, beavers took less aspen sticks in rows treated with wolf odor than water. As the predator odor did not affect the foraging distance from the shore, it is likely that our observation that foraging was the most intense close to shore is due to energetic constraints. However, predation risk probably affects the decision whether to forage on the land in the first place.     

Distribution in relation to life history in the directdeveloping gastropod Batillaria cumingi (Batillariidae) on two shores of contrasting substrata

Population structure and spatial distribution with growth ofthe direct-developing gastropod, Batillaria cumingi, were investigatedon two shores of differing substrata. Sand-mud shore and rockyshore populations differed in size structure; first-year snailswere ca. 7 mm in shell length (SL) in both populations, whereassecond-year snails, merging with older cohorts, measured 15–25mm SL in the sand-mud shore population and ca. 15 mm SL in therocky shore population. Egg distribution matched adult distributionin the sand-mud shore population, but was more restricted thanthat of adults in the rocky shore population. The distributionof newly-hatched juveniles (0–1 mm SL) was restricted inboth populations, but the growth stage at which snails extendedtheir distribution differed between the two populations; 1–2.5mm SL on the sand-mud shore and 5 mm SL on the rocky shore. Floatingachieved by early juveniles (ca. 2 mm SL), was commonly observedin the sand-mud population, but rarely in the rocky shore population.The sudden expansion in distribution of the 1–2.5 mm SLgrowth stage in the sand-mud shore population is consideredto have been caused by floating, while expansion of the distributionof older growth stages (>5 mm SL) in the rocky shore populationprobably occurs by crawling. (Received 13 May 1998; accepted 25 August 1998)