Processing of allochthonous macrophyte subsidies by sandy beach consumers: estimates of feeding rates and impacts on food resources

Allochthonous subsidies of organic material can profoundly influence population and community structure; however, the role of consumers in the processing of these inputs is less understood but may be closely linked to community and ecosystem function. Inputs of drift macrophytes subsidize sandy beach communities and food webs in many regions. We estimated feeding rates of dominant sandy beach consumers, the talitrid amphipods (Megalorchestia corniculata, in southern California, USA, and Talitrus saltator, in southern Galicia, Spain), and their impacts on drift macrophyte subsidies in field and laboratory experiments. Feeding rate varied with macrophyte type and, for T. saltator, air temperature. Size-specific feeding rates of talitrid amphipods were greatest on brown macroalgae (Macrocystis, Egregia, Saccorhiza and Fucus). Rates for large individuals of both species ranged from ∼40 mg wet wt individual⁻¹12 h⁻¹ on brown macroalgae to negligible feeding by M. corniculata on a vascular plant (surfgrass). Amphipod growth rates were also greatest on Macrocystis and lowest on surfgrass, Phyllospadix. For a Californian beach with substantial inputs of macrophyte wrack (>70 kg wet wt m⁻¹ month⁻¹ in summer), we estimated that the population of talitrid amphipods could process an average of 55% of the palatable Macrocystis input. Our results indicate that talitrid amphipod populations can have a significant impact on drift macrophyte processing and fate and that the quantity and composition of drift macrophytes could, in turn, limit populations of beach consumers.     

Habitat associations and diet of young-of-the-year Pacific cod (<Emphasis Type="Italic">Gadus macrocephalus</Emphasis>) near Kodiak,Alaska

The influence of environmental variables and habitat on growth and survival of juvenile gadoid species in the Atlantic has been clearly demonstrated; conversely, in the North Pacific little is known about the habitat and ecology of juvenile Pacific cod (Gadus macrocephalus Tilesius). The hypothesis that density of young-of-the-year (YOY) Pacific cod in nearshore habitats is predicted by shallow depth and the presence of eelgrass and macroalgae was tested in a variety of nearshore habitats adjacent to commercial fishing grounds near Kodiak Island, AK. From 10 to 22 August 2002, a beach seine and small-meshed beam trawl were used to capture YOY Pacific cod (n = 254) ranging from 42 to 110 mm fork length. Depth, water temperature, salinity, sediment grain size, and percent cover by emergent structure (i.e. tube-dwelling polychaetes, sea cucumber mounds, macroalgae) were measured prior to fishing. Density of YOY Pacific cod was highest in areas of moderate depth (15–20 m) and positively and linearly related to percent cover by sea cucumber mounds and to salinity. No previous studies have documented fish utilizing sea cucumber mounds as habitat. Furthermore, eelgrass and macroalgae were inconsequential to cod distribution. Diets consisted mainly of small calanoid copepods, mysids, and gammarid amphipods and were significantly related to cod length and depth. This work provides important information on previously undocumented factors that affect distribution and feeding of YOY Pacific cod, which ultimately influence growth and survival in this species.     

Responses of fish assemblage structures to sandy beach types in Kyushu Island, southern Japan

To determine differences in fish assemblage structures between beach morphodynamic types on an exposed sandy shore, three beach types (reflective, intermediate, and dissipative) were sampled at Fukiagehama Beach, Kyushu Island, southern Japan, in May, August, and November 2006 and 2007. Distinct differences in physical conditions and the amount of major prey invertebrates, such as copepods, mysids, and amphipods, were recognized between the three beach types. The reflective beach was characterized mainly by turbulent swashes, coarse sediment, and a low abundance of major prey invertebrates. In contrast, the dissipative beach was characterized mostly by benign swashes, fine sediment, and a high abundance of prey invertebrates. The intermediate beach tended toward an intermediate condition between the reflective and dissipative beaches. The fish assemblage structures also differed clearly between the three beach types, with species and individual numbers being greatest on the dissipative beach and lowest on the reflective beach. A similarity index indicated differences in species composition of the assemblages between the beach types. Such differences arose primarily from the differential distributions of two dominant trophic groups, zooplankton (mostly copepods) and epibenthic crustacean (mainly mysids and amphipods) feeders, between the beach types, both groups tending to increase in species and individual numbers from the reflective beach to the dissipative beach. In addition, the mean standard length of all fish collected in each month tended to decrease toward the dissipative beach. These results suggested that beach types strongly influence sandy beach fish assemblage structures.     

Analysis of lipid components for determining the condition of anchovy larvae,Engraulis mordax

It has been presumed that intertidal spawning by Limulus polyphemus minimizes the loss of egges to subtidal predators; however, this strategy involves considerable risks. Massive beach strandings of adults accompany seasonal spawning migrations of crabs along Cape May in Delaware Bay, (USA). At least 190000 horseshoe crabs, approximating 10% of the adult population, died from beach stranding along the New Jersey shore of Delaware Bay during the 1986 (May to June) spawning season. Abnormalities of the telson (which is used in righting behavior) were significantly more common among stranded crabs than among individuals actively spawning on the intertidal beach. The number of stranded crabs per day was not correlated with tidal height or environmental variables (wind speed, wave height) which characterized the conditions at spawning. A complex suite of factors, including the size of the available spawning population, tidal and weather conditions, and beach slope, influence the number stranded during the breeding season. Horseshoe crab stranding results in a large loss of gravid females from the population, and may represent a major input of organic matter to intertidal sandy beaches in certain regions of Delaware Bay.     

Corophium multisetosum (Amphipoda: Corophiidae) in Canal de Mira,Portugal: Some factors that affect its distribution

The population ofCorophium multisetosum (L.) from Canal de Mira, Ria de Aveiro, Portugal, was sampled seasonally from December 1985 to September 1986, as part of a larger survey of the benthic invertebrate macrofauna. Its distribution along the channel exposes the species to a range of salinities from freshwater to above 30. A principal components analysis, using the physical and chemical parameters of the sediment as variables and sampling stations as operational taxonomic units, indicated that abundance is negatively correlated with salinity, depth, and the occurrence of sediments rich in particles below 125µm and rich in organic matter. Abundance is positively correlated with temperature. The distribution of the species does not seem to be affected by the occurrence of sediment grades between 125 and 1 000µm. Paired-choice salinity experiments indicated thatC. multisetosum prefers salinities within the range 2.5 to 10. In multi-choice experiments concerning sediment grade, amphipods did not show any significant preference within the 125 to 500µm range, although the 125µm grade was chosen less frequently. The influence of temperature on the overall distribution ofC. multisetosum is discussed.     

Dispersal,population structure and burrow shape of Ocypode cursor

The distribution of the sand crab Ocypode cursor (L.), as indicated by the number of burrows, was studied for 2 years in a 50×50 m sand beach area in northern Israel. A definite relationship was established between the distribution pattern from the seashore inwards towards the sand dunes, and the degree of sand moisture as it changed seasonally. During autumn, more crabs were found at a distance of 15 to 25 m from the sea where sand moisture was about 14%. At the beginning of winter crabs dispersed evely, disappearing with advancing winter. Crabs reappeared in spring, although in smaller numbers, dispersing in a pattern similar to that in autumn. At the beginning of summer and later on, more crabs appeared and concentrated closer to the sea (5 to 10 m). The population structure was analysed directly by measuring the crab's dimensions, and indirectly by counting burrows and measuring the diameter of their openings. Direct analysis revealed two distinct sizeage groups: smaller crabs 0.5 to 3 cm long, and larger ones over 4 cm long. The smaller burrows were inhabited by the first group and were mostly found closer to the sea; the second group was found more landwards. Three main types of burrow shapes are described.     

Burrowing behaviour inferred from feeding traces of the opheliid polychaete <Emphasis Type="Italic">Euzonus</Emphasis> sp. as response to beach morphodynamics

The distribution of the opheliid polychate Euzonus sp. and the orientation of its feeding traces were examined in a wave-dominated Pacific sandy beach of central Japan by periodical sampling from June to December 2006. This species occurred constantly within sediments at mid-intertidal level, although its horizontal distributions were variable depending on changes of the beach profiles. These facts indicate that Euzonus sp. moved horizontally seaward or landward within the substratum in response to the shifting beach face due to changes of wave conditions from the ocean. By analysing the orientation of feeding traces, the adaptive burrowing behaviour of this species in response to beach topographical changes was revealed: Euzonus worms burrowed horizontally to various directions in a relatively stable condition of beach topography, whereas they moved preferentially landward under a heavy erosion of the beach face by a storm. These findings indicate that the burrowing behaviour prevents Euzonus sp. from excessive burial and washing out due to the beach morphodynamics.     

Local adaptation of immunity against a trematode parasite in marine amphipod populations

Resources allocated to defence against parasites are not available for investment in other functions such as growth or reproduction, resulting in trade-offs between different components of an organism’s fitness. In balancing the cost of infection and the cost of immunity, selection should only favour individuals that allocate more energy to resistance and immune responses in populations regularly exposed to debilitating parasites. Here, we compare the ability of amphipods, Paracalliope novizealandiae, to (1) avoid becoming infected and (2) to respond to infection by encapsulating and melanizing parasites, between two natural populations exposed to different risk of parasitism. One population faces high levels of infection by the debilitating trematode parasite Maritrema novaezealandensis, whereas the other population is not parasitised by this trematode nor by any other parasite. Under controlled experimental conditions, with exposure to a standardized dose of parasites, amphipods from the parasite-free population acquired significantly more parasites than those from the population regularly experiencing infection. Furthermore, a lower frequency of amphipods from the parasite-free population succeeded at melanizing (and thus killing) parasites, and they melanized a lower percentage of parasites on average, than amphipods from the parasitised population. These differences persist when individual factors, such as amphipod sex or body length, are taken into account as potential confounding variables. These results support the existence of local adaptation against parasites: an amphipod population that never experiences trematode infections is less capable of resisting infection, both in terms of its first line of defence (avoiding infection) and a later line of defence (fighting parasites following infection), than a population regularly exposed to infection.     

In vivo marking of shallow-water and deep-sea amphipods by ingestion of bait mixed with fast green

Seven vital stains were mixed with fish muscle and fed to a sublittoral lysianassid (Orchomene sp. A collected in Scripps Canyon off La Jolla, California USA between August 1978 and August 1981) in the laboratory to test the utility of these dyes as feeding labels for scavenging amphipods. Fast-green FCF proved to be the most effective of the stains tested; >90% of starved amphipods fed fast-green-stained bait (Scomber japonicus muscle) for as little as 1 h exhibited a conspicuous green coloration along the digestive tract, which lasted an average of 35 d. The ability to label animals with such a single, short exposure interval makes this dye especially suitable for marking scavengers attracted to bait. Fast green efficiently stained this amphipod over a broad range of concentrations (1 to 8% by weight in water) and feeding regimes, and had no significant effects on survivorship or activity of laboratory-held organisms. Nile blue A also proved to be an acceptable feeding label for Orchomene sp. A for time scales of about 1 wk. Additional laboratory and field tests between August 1978 and August 1979 indicated that fast green is an effective feeding stain for 2 bathyal species (Orchomene sp. B and O. plebs from the Ross Sea, Antarctica) and 3 abyssal species (Eurythenes gryllus, O. gerulicorbis and Paralicella caperesca from the central North Pacific Ocean) of amphipods. Field labeling of amphipods at 5 800 m in the central North Pacific Ocean with fast green demonstrates that feeding stains may be used readily as in situ marking agents for population studies of scavengers in remote environments.     

Scavenging deep-sea amphipods: Effects of food odor on oxygen consumption and a proposed metabolic strategy

In situ respiration rates as a response to the odor of food were measured for two species of scavenging amphipods, Paralicella caperesca from 3 650 m in the western North Atlantic Ocean and Orchomene sp. B from 1 300 m in the Santa Catalina Basin off southern California (USA). In addition, complementary laboratory starvation/respiration rates for a shallow-water species, Orchomene sp. A, were determined. Initial elevated O₂ consumption rates were found for up to 8 h in all deep-sea amphipods exposed to bait odor, followed by a period of lowered respiration equivalent to rates observed in individuals not exposed to bait. Orchomene sp. A revealed a response similar to that observed in the deep-sea species. A metabolic strategy is proposed whereby scavenging amphipods efficiently utilize large episodic organic falls in the food-limited environment of the deep sea. This strategy involves (1) the ability to withstand long periods of starvation, (2) rapid response to an organic fall, (3) rapid location of the organic fall, (4) maximal rate of food consumption with maximal quantity ingested, and (5) efficient utilization of the consumed food. Each of these attributes are explored with the expected and observed mechanisms employed to achieve them.     

Small-scale spatial and seasonal differences in the distribution of beach arthropods on the northwestern Tunisian coast. Are species evenly distributed along the shore?

Spatial distributions of sand beach arthropods were studied at Zouara (Nefza), a beach dune system along the northwestern coast of Tunisia. Two transects with pitfall traps, perpendicular to the shoreline and placed from sea towards land, captured spontaneously crawling arthropods. The pitfalls were used to obtain data on horizontal zonations for 2 days during different seasons (April, October). In April, in order to assess local, long-shore distributions of surface-active beach invertebrates, ten transects with pitfall traps were placed every 100 m, for 2 days. Core samples were also taken in correspondence with each of the ten transects to obtain the distribution of burrowing individuals. Sand samples were taken for successive laboratory analyses (salinity, grain size and organic contents). Local climatic conditions were also recorded. One isopod (Tylos europaeus, subdivided into three different size classes), two amphipods (Talitrus saltator, Talorchestia brito and indistinguishable juveniles) and four coleopteran species (Eurynebria complanata, Scarites laevigatus, Phytosus nigriventris, Phaleria acuminata) were considered. Mean zonations perpendicular to the shoreline showed that crustaceans were found closer to the sea-line limits than coleopterans and that between seasons some species changed their mean zonation more than others. Local, long-shore distributions of active and burrowing individuals showed that most species were unevenly distributed along the shore. These discontinuous distributions differed according to the species and age class and were mainly related to specific microclimatic conditions and to nutrition.     

Niche segregation in sandy beach animals: an analysis with surface-active peracarid crustaceans on the Atlantic coast of Spain

Several species of sandy beach invertebrates regularly switch between burial in the sand during the day and surface activity at night to feed on stranded organic matter. Because all species consume essentially the same type of food (i.e., wrack) deposited over a restricted area, the potential for competition exists. Conversely, spatial and temporal segregation of surface activity behaviour is predicted to allow for niche separation. Here, we tested whether such behavioural niche separation occurs in three species of sympatric crustaceans (the talitrid amphipods Talitrus saltator (Montagu 1808) and Talorchestia brito Stebbing 1891, and the oniscoidean isopod Tylos europaeus Arcangeli 1938). We also assessed whether surface activity is modulated by weather, sea conditions and moon phases. Surface activity was measured over three consecutive moon phases on an exposed beach on the Atlantic Coast of Spain using pitfall traps along three transects from the foredunes to the swash. Adults of the amphipod T. saltator and the isopod T. europaeus overlapped spatially and temporally in their surface activity for most of the night. By contrast, the activity of T. brito was strongly disjunct in time and concentrated into significantly shorter bouts around dawn or dusk. Niche differentiation was also evident along the space axis, where T. brito occurred distinctly lower on the beach. Although several environmental factors (e.g., air and sand temperature, humidity, atmospheric pressure, wind speed and wind direction) were correlated with surface activity, this was highly variable amongst species, life stages, and synodic phases, and did not obscure the fundamental mosaic of spatio-temporal heterogeneity amongst species. Nocturnal feeding on beach wrack creates the potential for competition amongst sympatric crustaceans which can, however, be mitigated by behaviour patterns that separate consumers in time and space.     

Effects of Illegal Harvest of Eggs on the Population Decline of Leatherback Turtles in Las Baulas Marine National Park,Costa Rica

Abstract: Within 19 years the nesting population of leatherback turtles (Dermochelys coriacea) at Parque Nacional Marino Las Baulas declined from 1500 turtles nesting per year to about 100. We analyzed the effects of fishery bycatch and illegal harvesting (poaching) of eggs on this population. We modeled the population response to different levels of egg harvest (90, 75, 50, and 25%) and the effect of eradicating poaching at different times during the population decline. We compared effects of 90% poaching with those of 20% adult mortality because both of these processes were present in the population at Las Baulas. There was a stepwise decline in number of nesting turtles at all levels of egg harvest. Extirpation times for different levels of poaching ranged from 45 to 282 years. The nesting population declined more slowly and survived longer with 20% adult mortality (146 years) than it did with 90% poaching (45 years). Time that elapsed until poaching stopped determined the average population size at which the population stabilized, ranging from 90 to 420 nesting turtles. Our model predicted that saving clutches lost naturally would restore the population when adult mortality rates were low and would contribute more to population recovery when there were short remigration intervals between nesting seasons and a large proportion of natural loss of clutches. Because the model indicated that poaching was the most important cause of the leatherback decline at Las Baulas, protecting nests on the beach and protecting the beach from development are critical for survival of this population. Nevertheless, the model predicted that current high mortality rates of adults will prevent population recovery. Therefore, protection of the beach habitat and nests must be continued and fishery bycatch must be reduced to save this population.     

Further studies on the pressure tolerance of deep-sea crustacea,with observations using a new high-pressure trap

A deep-sea benthic trap is described with which amphipods (Tmetonyx cicada) were both collected and observed at their ambient pressure of 134 atm. T. cicada collected with decompression from the same depth and locality were more sensitive to subsequent recompression than those amphipods brought to the surface at their ambient pressure. T. cicada from 2700 m experience irreversible injury during decompression to atmospheric pressure. The pressure tolerance of the deep-sea mysid Gnathophausia zoea was measured and compared with the tolerance of mid-water decapods and the shallow-water Crangon crangon. G. zoea became more sensitive to high pressure with prolonged exposure to atmospheric pressure. Deep sea animals exhibit a tolerance to high pressure related to their normal ambient pressure; sensitivity to decompression is also related to normal ambient pressure.     

Habitat harshness and morphodynamics: life history traits of the mole crab Emerita brasiliensis in Uruguayan sandy beaches

The habitat harshness hypothesis (HHH) postulates that in reflective beaches the harsh environment forces organisms to divert more energy towards maintenance and they therefore have lower abundance, fecundity, growth and survival rates than in dissipative beaches. Recent investigations have tested this hypothesis through single comparisons of only two beaches, and thus the observed trends in population level variables cannot be attributed incontestably to the beach state, but only to location. Here, abundance, reproduction, recruitment, population structure and body size of the intertidal mole crab Emerita brasiliensis were compared between populations from eight microtidal exposed sandy beaches with contrasting morphodynamics, sampled bimonthly during 22 months throughout the 180 km Uruguayan Atlantic coast. Physical variables and compound indices of the beach state were used to categorize sandy beaches. The results of this bi-annual large-scale analysis were fully consistent with the predictions of the HHH: abundance (total and population components), duration of the reproduction and recruitment seasons and the individual size of megalops and females of the mole crab E. brasiliensis decreased from dissipative to reflective beaches. This was reflected by linear or, mostly, nonlinear relationships between biological and both physical variables and compound indices of beach state. In conclusion, this multi-beach sampling provides compelling evidence of a consistent response of demographic and life history traits of an intertidal beach species to morphodynamic characteristics.     

Population dynamics and growth of <Emphasis Type="Italic">Nassarius reticulatus </Emphasis>(Gastropoda: Nassariidae) in Rhosneigr (Anglesey,UK)

Seasonal changes in catch rate, growth and mortality of Nassarius reticulatus from an intertidal lagoon and a wave-exposed beach at Rhosneigr (Anglesey, North Wales, UK) are described. The number of N. reticulatus caught in baited traps from the lagoon was significantly higher (>125 individuals trap⁻¹) during the summer (>18°C), than at <12°C (<65 individuals trap⁻¹), and the numbers caught in the lagoon were an order of magnitude greater than on the beach, >13 individuals trap⁻¹ in July (>16°C), and <5 individuals trap⁻¹ between December and April (<9.5°C). Predictions of shell growth attained by N. reticulatus annually in the lagoon using graphical modal progression analysis (MPA) of length frequency data, were similar to the growth of marked and recaptured lagoon N. reticulatus. Predictions of shell growth using computerised length frequency distribution analysis (LFDA), however, did not reflect the growth as accurately as MPA. Modal progression analysis demonstrated that N. reticulatus from the lagoon achieved a higher asymptotic maximum shell length (L _∞) and a lower growth constant (K) than animals from the beach. Shell growth was seasonal with growth of the lagoon individuals slowing down towards the end of September and resuming in early April, about a month later than the beach individuals. Mortality of N. reticulatus was greater during the summer, and survival was lower in the lagoon than on the beach. Recruitment patterns were similar in the lagoon and on the beach, and MPA and LFDA predicted that larval N. reticulatus settled between late summer and early autumn, with juveniles (7–8.9 mm) appearing in the population the following year, between February and April. Growth of male and female N. reticulatus in the laboratory was similar and was temperature and size dependent. The different growth patterns between N. reticulatus from the two habitats, predicted using MPA, were maintained when individuals were reared under laboratory conditions for ∼6 months; N. reticulatus <21 mm from the beach grew faster than individuals from the lagoon, although N. reticulatus >21 mm from the lagoon grew faster and attained a larger length (26 mm) than individuals from the beach (24 mm). Low food availability did not affect N. reticulatus survival in the laboratory but significantly suppressed shell growth.     

Use of red algae as hosts by kelp-associated amphipods

To what extent densities of amphipods associated with red algae are related to food value or habitat form and architecture were investigated. Four epiphytic red algae common on kelp stipes (Laminaria hyperborea) were sampled, and the densities of three species of associated amphipods were analysed. The algae were chosen to represent different structures and levels of architectural complexity. Palmaria palmata and Delesseria sanguinea are leaf-shaped, and Ptilota gunneri and Polysiphonia elongata are branched. The algae were later fed to the common epiphyte-associated amphipods Ampithoe rubricata, Jassa falcata and Caprella septentrionalis in no-choice laboratory experiments. Survival and growth were measured. J. falcata was found at the highest densities on P. gunneri and D. sanguinea, C. septentrionalis was found in highest densities on P. elongata, and P. gunneri and A. rubricata were found at the highest densities on P. gunneri. The survival and growth were highest on P. palmata for all amphipods. This indicates that the form and function of the algal host is more important for the distributions of amphipods than the food value. Interspecific distribution differences between amphipods may have been related to differences in their body form and size, as well as to crypsis.Communicated by L. Hagerman, Helsingør     

Comparing macroalgal food and habitat choice in sympatric, tube-building amphipods, Ampithoe lacertosa and Peramphithoe humeralis

For small tube-building amphipods that live on the algae they consume, food and habitat are tightly linked. This study compared two closely related amphipods to determine whether the species’ algal preferences are based on the food value of the algae or on some other aspect of their algal habitat. Ampithoe lacertosa and Peramphithoe humeralis are both abundant on Shannon Point beach (Anacortes, Washington, USA; 48°30.542′ N, 122°41.070′ W) but specialize on different algae. In observations and laboratory experiments conducted July–September 1997, 2007, and 2008, the two species exhibited markedly different choices of food and habitat when offered six common macroalgae. Ampithoe lacertosa ate all algae offered, but preferentially built tubes on the green alga Ulva lactuca. Survival was relatively low among juveniles maintained on single species diets, except when they were fed Mazzaella splendens. Conversely, P. humeralis consumed primarily the brown kelp Saccharina latissima, Alaria marginata, and Desmarestia ligulata and preferred those species for tube building. Juvenile P. humeralis could not survive on a diet of U. lactuca or M. splendens. While A. lacertosa builds simple, temporary tubes and relocates frequently, P. humeralis is a highly thigmotactic species that builds long-term, complex tubes on the alga it prefers to eat. Feeding and habitat preferences of the two species were not clearly linked to nitrogen content of the algae, C:N ratio, or toughness of the algal tissue. Instead, preferences of the species may be related to their mobility and the permanence of the tubes they build. Ampithoe lacertosa and P. humeralis also use different feeding strategies; the former appears to mix algae to produce a high-quality diet, while the latter is more selective and has a capacity for compensatory feeding. The species are abundant on the same protected rocky shores, but specialize on different algae for habitat and food. Results suggest that the nutritional requirements of these sympatric mesograzers differ considerably and even closely related species can exhibit divergent behavioral strategies.     

Spatial structure, sampling design and abundance estimates in sandy beach macroinfauna: some warnings and new perspectives

We discuss methodological aspects directed to quantify the across-shore population structure and abundance of sandy beach macroinfauna. The reliability of estimates derived from design-based (stratified random sampling) and model-based (geostatistics, kriging) approaches is discussed. Our analysis also addresses potential biases arising from environmentally driven designs that consider a priori fixed strata for sampling macroinfauna, as opposed to species-driven sampling designs, in which the entire range of across-shore distribution is covered. Model-based approaches showed, spatially, highly autocorrelated and persistent structures in two intertidal populations of the Uruguayan coast: the isopod Excirolana armata and the yellow clam Mesodesma mactroides. Both populations presented zonation patterns that ranged from the base of the dunes to upper levels of the subtidal. The Gaussian model consistently explained the spatial distribution of species and population components (clam recruits and adults), with a minor contribution (Е%) of unresolved, small-scale variability. The consistent structure of spatial dependence in annual data strongly suggests an across-shore-structured process covering close to 35 m. Kriging predictions through cross-validation corroborated the appropriateness of the models fitted through variographic analysis, and the derived abundance estimates were very similar (maximum difference=7%) to those obtained from linear interpolation. Monthly analysis of E. armata data showed marked variations in its zonation and an unstable spatial structure according to the Gaussian model. The clear spatial structure resulting from species-driven sampling was not observed when data was truncated to simulate an environmentally driven sampling design. In this case, the linear semivariogram indicated a spatial gradient, suggesting that sampling was not performed at the appropriate spatial scale. Further, the cross-validation procedure was not significant, and both density and total abundance were underestimated. We conclude that: (1) geostatistics provides useful additional information about population structure and aids in direct abundance estimation; thus we suggest it as a powerful tool for further applications in the study of sandy beach macroinfauna; and that (2) environmentally driven sampling strategies fail to provide conclusive results about population structure and abundance, and should be avoided in studies of sandy beach populations. This is especially true for microtidal beaches, where unpredictable swash strength precludes a priori stratification through environmental reference points. The need to use adaptive sampling designs and avoid snapshot sampling is also stressed. Methodological implications for the detection of macroecological patterns in sandy beach macroinfauna are also discussed.