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.     

Multiple regression analysis of the sources of variation in orientation of two sympatric sandhoppers, <Emphasis Type="Italic">Talitrus saltator</Emphasis> and <Emphasis Type="Italic">Talorchestia brito</Emphasis>, from an exposed Mediterranean beach

Two sympatric species from an exposed sandy beach in north-western Tunisia, the sandhoppers Talitrus saltator (Montagu) and Talorchestia brito Stebbing, were compared as regards orientation under natural conditions. Sets of experiments were carried out on the beach using two experimental arenas, one permitting the view of both the sky and landscape and the other of the sky only. Replicates were made at different hours of the day, in different days and period of the year and with various climatic conditions. Multiple regression models were fitted to the angular data obtained. These models allow an estimate of the trends of variation depending on a combination of factors. The simultaneous effects of environmental factors influencing orientation were quantified for each species. The sun was confirmed as the major orientation cue in these populations, but the concentration of directional choices was enhanced by the landscape vision. The climatic factors affected orientation in different ways depending on the time of day and year. When the two species were compared, Talitrus saltator showed a higher flexibility of response than Talorchestia brito under the same environmental conditions, supporting the hypothesis of a higher level of terrestrialization of the former species.     

Temporal and spatial use of stranded wrack by the macrofauna of a tropical sandy beach

During two field studies the macrofauna associated with wrack stranded on a tropical sandy shore was analysed. During the first period all animals present in small wrack deposits were randomly collected with cores. During the second field study active animals were sampled in larger wrack deposits for an entire lunar period, and tube traps were used. After a comparison between the two studies, only the data from the tube traps were examined from a temporal and spatial point of view. The aim of the present work was to gain information on how, when and where wrack was colonised by invertebrates, keeping in mind cyclical aspects such as lunar, diel and tidal phases. Predatory taxa, such as Staphylinoidea and the amphipods Talorchestia martensii, were most abundant, whereas Diptera larvae and Tenebrionidae were scarce. Successional changes of beach wrack colonisation were evident throughout a semi-lunar period, with molluscs invading wrack during the first days of deposition and histerids during the last ones. Strictly nocturnal or diurnal surface-active species were found to be active in the wrack during both the day and the night. The analysis of the tidal component of species activity in the wrack showed that some species moved at ebbing tide, and others moved at rising tide. Differences were found also in the mean hours of tidal activity, calculated separately during day and night periods. The study of the zonation of each species showed that in some cases wrack deposits were closely followed by the fauna as their position changed during the semi-lunar phase. In other cases differences occurred between species zonation during day and night periods and tidal phases. Cluster analysis indicates that most species exploit the wrack in different ways (as refuges and/or feeding site), both in space and in time. Received: 11 July 1999 / Accepted: 20 January 2000     

Temporal and spatial distribution of three supralittoral amphipod species on a sandy beach of central Italy

The distribution of three talitrid species—Talitrus saltator (Montagu, 1808), Orchestia gammarella (Pallas, 1766), Platorchestia platensis (Kroyer, 1845)—in the beach-dune system at the mouth of the Mignone River (central Italy) was analysed. It was related to the variations of the following abiotic factors: temperature, penetrability, pH, conductivity and moisture of the sediment. The beach-dune system is influenced by human impact and natural erosion. All species showed a maximum capture frequency in November, while the abundance decreased to a minimum in July. T. saltator was dominant on the beach, the other two along the riverbank. Juveniles were abundant along the riverbank; they were most abundant in November and almost disappeared in summer. T. saltator was more abundant near the waterline during the hottest months and occupied the inner beach in winter, with occasional presences on the dune. Regression analysis between the abiotic factors and species abundance showed a positive relationship with pH and temperature for T. saltator, while O. gammarella was negatively related to pH. Canonical correspondence analysis (CCA) showed that penetrability, moisture and temperature had the greatest influence on the species. T. saltator was almost entirely confined to the beach transects and strongly associated with penetrability, temperature and pH values. O. gammarella was mostly associated with the riverbank and P. platensis only found there and on a nearby pool. The two species were mainly influenced by moisture and variations in grain sizes of the sediment.     

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.     

Physiological capacity and environmental tolerance in two sandhopper species with contrasting geographical ranges: Talitrus saltator and Talorchestia ugolinii

We investigated the physiological plasticity and environmental tolerance of two phylogenetically closely related, ecologically similar and co-occurring species of supralittoral amphipods differing drastically in the size of their geographical ranges. A series of physiological traits were characterised for the Corsican-endemic Talorchestia ugolinii Bella-Santini and Ruffo and the widespread Talitrus saltator Montagu. The effect of body mass, temperature and salinity on heart rate (used as proxy for metabolic activity and stress), the effect of temperature on oxygen consumption and the tolerance to salinity exposure were investigated in both species, together with the characterisation of haemolymph osmoregulation in T. ugolinii. Our results showed that there is a clear difference in the resting metabolic rates and physiological capacity, as well as environmental tolerance, between T. saltator and T. ugolinii, with T. saltator overall showing a broader physiological niche. Although T. ugolinii showed a relatively good ability to regulate its haemolymph osmotic concentration (similar to that previously described for T. saltator), it demonstrated a lower tolerance to exposure to hypo-osmotic stress. In addition, a consistent picture emerged between the ability to control the cardiac function and the capacity to actively respond to osmotic stress. The physiological findings are discussed in relation to the known ecology and geographical distribution of T. ugolinii.     

Astronomical, landscape and climatic factors influencing oriented movements of Talitrus saltator in nature

 The present study was conducted on a Mediterranean beach (Burano, southern Tuscany, Italy) to examine the timing, orientation, and motivating and directing factors of the spontaneous movements of the sandhopper Talitrus saltator (Montagu, 1808). In April 1994, October 1994 and June 1995, during different moon phases, traps were positioned in the eulittoral zone, which intercepted sandhoppers when walking on the sand surface. At the same time environmental parameters were registered. Contemporaneously, orientation tests were carried out on active individuals using two arenas, one of which permitted a view of both sky and landscape and one which prohibited the landscape view. The results show landward migration after sunset, for juveniles later than for adults, nonoriented activity for 2 or 3 h after midnight and seawards zonal recovery before and after sunrise. The main microclimatic factor modulating activity was sand temperature, while the nonoriented activity between the two migratory movements seems to be endogenously determined. Both sky and landscape cues are used by sandhoppers for orienting their course, but the landscape view is sufficient and necessary on new moon nights. These results contribute to solution of the controversy on the mechanisms actually motivating sandhoppers in nature. Received: 24 September 1996 / Accepted: 25 October 1996     

Supralittoral amphipod abundances across habitats on Mediterranean temperate beaches

Community complexity and abiotic conditions are key components of environmental heterogeneity that affect the abundance and distribution of species. In this study we evaluated how environmental conditions affect abundances of supralittoral amphipods (Talitridae) in four habitats (sandy beach, rivermouth, wrack and lakeshore), along the Italian peninsula in the Mediterranean Sea. All samplings covered a 12 month period, and used the same sampling methodology thereby enabling comparison of abundances and species composition and richness. Four species (Talitrus saltator (Montagu, 1808), Orchestia gammarellus (Pallas, 1766), O. montagui Audouin 1826, O. cf. cavimana Heller 1865) were collected in the different habitats, but most species were found or were abundant in only one of the four habitats. Abundances of talitrids (numbers per sampling hour) differed significantly among the habitats with highest abundances found in the wrack and on the riverbank in proximity to an estuary, and lowest abundances observed on four sandy beach sites. Environmental conditions (temperature, moisture, substrate penetrability) differed among the habitats and were associated with some of the among-site variability in abundances. Our findings demonstrate that talitrids thrive better in some supralittoral habitats than others, and that some habitats could be considered to be “hotspots” of talitrid ecology and biodiversity.     

Spatial and temporal strategies in the surface activity of some sandy beach arthropods living along the French Atlantic Coast

During a field study on the eco-ethology of sandy beach arthropods conducted along the French Atlantic Coast it emerged thatTylos europaeus, Phaleria cadaverina andLabidura riparia were among the most abundant species that lived in the eulittoral. Tetradirectional pitfall traps, placed from the base of the dune to the tidal limits, were used. The traps intercepted surface-active arthropods, and the number of traps varied according to the levels of the tides. Data were recorded hourly during two periods corresponding to a spring tide and the following neap tide. At the saine time the most important environmental parameters were registered each hour. The results indicate that all three species show prevalently nocturnal surface activities. ForT. europaeus andPh. cadaverina an influence of the synodic and tidal phases was found on the zonation of the surface activity and on the extent of movement. The latter was due to the greater or smaller distances that the isopods and tenebrionids had to move from the diurnal burrowing sites to the foraging zones. ForL. riparia no variations in the total mean zonations were found between the two synodic phases, and only at neap tide were seaward, followed by landward movements found. Multiple regression analysis showed that the environmental parameters registered in this period were never limiting, as threshold values in this case probably were not reached.     

Environmental factors influencing the surface activity and zonation of Tylos europaeus (Crustacea: Oniscidea) on a Tyrrhenian sandy beach

The present study considers a population of Tylos europaeus Arcangeli, 1938 living on a Tyrrhenian sandy beach (Burano, GR, Italy). Monthly surveys were carried out between April 1986 and March 1987 with directional pitfall traps so as to intercept the isopods moving in four directions on the beach surface. In addition a bimonthly study was performed from March 1991 to January 1992 using two methods of capture: pitfall traps joined by 10-cm high strips of fibreglas for surface-active individuals and sieving for those burrowed in the sand. Capture frequencies allowed analysis of annual abundance, daily activity and zonation of juveniles and male and female adults. Variations of these spatio-temporal data were correlated by means of multiple regression with many environmental parameters: temperature and relative humidity of the air and sand, evaporation, wind direction and speed, global radiation, atmospheric pressure, rainfall, the sand salinity and granulometric parameters. T. europaeus was found to be mainly active in summer and autumn and during the night, and was zoned along the eulittoral. The surface activity was influenced by almost all the environmental factors when they were limiting but especially by the relative humidity of the air. The mean zonation of active specimens, however, varied hourly according to the sand temperature. On the other hand, the zonation of the buried individuals depended on the mean grain size, which involves many other parameters, such as moisture and oxygen contents.     

Genetic differentiation between populations of Talitrus saltator and Talorchestia deshayesii (Crustacea: Amphipoda) from coastal areas of the north-western European continent

Electrophoretic studies of gene-enzyme variation in the littoral talitrids Talitrus saltator (Montagu) and Talorchestia deshayesii (Audouin) were undertaken to estimate the amount of divergence among geographically separated populations. Samples of both species were taken from sandy beaches over a transect of approximately 3 500 km along the coast of the European continent including Baltic, North Sea and Atlantic locations. A total of 22 T. saltator and 15 T. deshayesii populations were analysed for genetic variation at various enzyme loci. Both amphipods revealed relatively low levels of polymorphism and heterozygosity. Among the loci studied, phosphoglucose isomerase (Pgi) and phosphoglucomutase (Pgm) were highly polymorphic. Patterns of micro- and macrogeographic variation in terms of distributions of allele frequencies at these particular loci are compared. Interpopulation allozymic variation was shown to be lower in T. deshayesii than in T. saltator. As demonstrated by T. saltator populations sampled in coastal sites ranging from Denmark to western France, clinal variation in frequencies of two alleles became evident at the PGI locus, exhibiting a steady increase in the level of polymorphism with decreasing latitudes. It is argued that limited gene flow and, to some extent, random genetic drift may account for the gene pool structure of the talitrid species investigated.     

Lunar rhythms of vertical migrations coded in otolith microstructure of North Atlantic lanternfishes,genus Hygophum (Myctophidae)

Otoliths of five Hygophum species were examined by means of light and scanning electron microscopy. In otoliths of four species (H. benoiti, H. macrochir, H. reinhardtii and H. taaningi) a strong cyclic pattern of the incremental structure was observed. In the fifth species (H. hygomii) such a pattern did not exist. An analysis of archival data on mesopelagic collections suggested three types of Hygophum spp. migratory behavior in relation to the lunar cycle which corresponded with the otolith microstructure. In H. hygomii only limited influence of moon phase on the uppermost range of night-vertical migration toward the surface was observed. The abundant nighttime occurrence of this species moved from the 0 to 50 m into the 50 to 100 m depth strata at full moon. In H. benoiti, a great part of the population, mainly juveniles, showed a tendency toward cessation of the vertical migrations during the first and fourth quarters of the lunar cycle. H. macrochir and H. taaningi had the strongest correlation of behavior with the lunar cycle. Both species showed arrested vertical migrations at the new moon phase, staying at day depths during the night, i. e., below 400 m. Thus, sequences of clear growth increments in otoliths represented a fast-growth period associated with the night migration to the warm surface layers, while bands without easily distinguishable incremental structure were interpreted as a period of slow growth in deep, cold waters due to limitation of the upward migration range occurring approximately at new moon.     

<Emphasis Type="Italic">Sargassum</Emphasis> on Santa Rosa Island,Florida: faunal use and beachgoer perception

The yearly influx of Sargassum onto the beaches of northwest Florida is considered a nuisance to some and a necessity to others. In Pensacola Beach, the Santa Rosa Island Authority rakes the wrack with mechanical beach cleaners to improve the aesthetic quality for beachgoers. The purpose of this study was three-fold: to evaluate the local faunal use of Sargassum wrack, to gauge public perception of Sargassum on the beach, and to test whether public perception of the beauty of the beach, the necessity of raking, and the likelihood of visiting could be influenced by a simple educational sign. A two-part methodology consisted of 1) systematic observation of faunal use, and 2) interviews of 200 beachgoers via a detailed pre-post/post only public use survey. Results showed that 11 of the 22 species of shorebirds documented, including two uncommon migrants, were observed using Sargassum wrack to forage, rest, and hide. Public survey results demonstrated that although beachgoers generally considered themselves to be “ecofriendly”, their perceptions of Sargassum wrack can be positively influenced through environmental education such as informative signage on the beach. In conclusion, Sargassum wrack provides valuable additional habitat to shorebirds and other critters, and that leaving the beach wrack to naturally become part of the ecosystem would not deter most beachgoers (70%) from visiting Pensacola Beach. This research contributes valuable information to coastal managers and other stakeholders for improved ecosystem protection and management.     

Variation in zonal recovery in four Talitrus saltator populations from different coastlines: a comparison of orientation in the field and in an experimental arena

Under stressful conditions (e.g. finding themselves on dry or moisture-saturated substrates) littoral talitrids (Crustacea, Amphipoda) demonstrate zonal orientation, in which they must promptly reach the optimal zone of the beach, the wet fringe near the shoreline. A relationship might therefore exist between the use of orientation and the frequency of such stressful conditions in the natural environment. Moreover, the efficiency of orientation toward the sea could be related to the possibility of using strategies other than zonal orientation in order to avoid stress. This study analysed the actual use and efficiency of orientation under natural conditions of four Talitrus saltator (Montagu, 1808) populations from Mediterranean and northern European Atlantic coasts with different ecological features. Orientation tests were carried out on the beach with all natural cues available. Then the same individuals underwent control experiments to study their sun orientation far from the sea in an experimental arena. The following results emerge from the comparison of the circular distributions: (1) marked differences among populations in the precision of zonal recovery under natural conditions; (2) a common solar orientation capacity in the control tests far from the sea; (3) different orientation choices of the same individuals according to the test conditions, natural or controlled. The habitat diversity of the four populations (amount, distribution and kind of detritus and wrack on the beach, degree of coastal erosion, orientation of the shoreline, human use of the beach) provides an ecological interpretation for the differences in orientation observed among populations. Received: 13 October 1997 / Accepted after revision: 26 April 1998     

Crustacean epifauna of seagrass and macroalgae in Apalachee Bay,Florida, USA

Epifaunal crustaceans on turtlegrass (Thalassia testudinum) and five dominant macroalgae (Anadyomene stellata, Digenia simplex, Halimeda incrassata, Laurencia poitei and Penicillus lamourouxii) were quantitatively sampled bimonthly over a one-year period from September 1979 to September 1980 in a subtropical seagrass meadow in Apalachee Bay, Florida (northeastern Gulf of Mexico). These plant species exhibited a wide range of morphologies, with surface area-to-biomass ratios differing by over 2.5 times. A similar suite of crustaceans occurred on all macrophytes despite differences in shape or architecture among plant species. Relative abundances of many crustaceans, however, varied among plant hosts. Similarity analysis indicated that the epifaunal associates of T. testudinum were distinct from those of the macroalgae. Species richness was generally higher on turtlegrass than on any of the macroalgae. Abundances of total crustaceans per plant biomass or per plant surface area, on the other hand, were greater on all macroalgal species compared to the seagrass. Abundances (per plant biomass or plant surface area) of 14 of the 16 numerically dominant epifaunal species differed significantly among macrophytes. Twelve of the 16 species had greater abundance on one or more macroalgae, while only two species were more abundant on T. testudinum. Almost half of the dominant species had greatest abundances on the branching red alga L. poitei. Although abundances per plant biomass and plant surface area were greater on macroalgae relative to turtlegrass, densities (individuals per meter square of bottom) of animals associated with T. testudinum were significantly greater than those associated with macroalgae, primarily because of the greater abundance of turtlegrass in the grass bed. Both surface area-to-biomass ratios and degree of branching were poorly correlated with epifaunal abundance and number of species. Neither structural feature is an adequate predictor of faunal abundance and species richness among plant species, especially when macrophytes with very different morphologies are compared.     

The trophic ecology of two abundant mesopredators in south-east coastal waters of Tasmania, Australia

Investigating predator–prey relationships is an important component for identifying and understanding the factors that influence the structure and function of ecosystems. Mesopredators, defined as mid-level predators, have a profound effect on ecosystem structure by contributing an important link between apex predators and lower trophic levels. The diet of two elasmobranch mesopredators, Squalus acanthias and Mustelus antarcticus, was investigated in three locations in south-east Tasmania. Squalus acanthias consumed predominantly pelagic teleosts and cephalopods, while M. antarcticus predominantly consumed benthic crustaceans. As a result, there was low dietary and niche overlap between the two species. There was however evidence of intra-specific dietary variations between locations for both the species. This study has contributed to a better understanding of the top-down dynamics of the food web in coastal Tasmania, by providing important dietary information of two abundant mesopredators. In addition, the similar dietary patterns for S. acanthias and other Mustelus species over much of their global range suggest they may be consistent in their trophic roles across systems, with limited competition between these two sympatric mesopredators to be expected.     

The spatiotemporal expression pattern of trypsinogen and bile salt-activated lipase during the larval development of red porgy (<Emphasis Type="Italic">Pagrus pagrus</Emphasis>, Pisces,Sparidae)

A restoration programme was introduced in the Mondego Estuary (Portugal) to recover seagrass beds of Zostera noltii endangered by eutrophication. A long-term survey of 10 years was used to assess the development of the processes involved, focusing one of the key species (Cyathura carinata, Isopoda). The mitigation measures implemented since 1998 (nutrient loading reduction, freshwater circulation improvement and seagrass bed protection) enhanced water quality and seagrass recovery, thus preventing the development of macroalgal blooms. C. carinata was resilient to the occurrence of floods and macroalgal blooms, although both events caused dispersion of individuals. This isopod was not much influenced by the changes occurring in the estuary, showing an unalterable population structure during the entire study period. After 1998, its density and biomass became more stable at an inner unvegetated sand flat area, where this isopod was most abundant; its population slightly increased in a bare mud flat at the middle section of the estuary; but it could not establish successfully in a downstream Z. noltii bed, contrarily to other common estuarine species. Apart from other unknown reasons, the disrupted balanced between trematodes and their hosts, caused by the eutrophication processes, may have an important role in the discontinuity of C. carinata at the Z. noltii bed. If the intertidal areas become fully restored to the original seagrass coverage, high prevalence and intensity trematodes may prevent this isopod and other crustaceans from recovering within the intervened areas, by enhancing host mortality and recruitment failure. In order to avoid this kind of situation, it may be necessary to survey the levels of parasite infestation within the target hosts and safeguard areas where crustaceans present healthy populations.     

Biomonitoring of polychlorinated byphenyls contamination in the supralittoral environment using the sandhopper Talitrus saltator (Montagu)

We evaluated the use of Talitrus saltator as biomonitor of polychlorinated byphenyls (PCBs) contamination of the supralittoral zone of Mediterranean sandy shores, an area not yet investigated about the presence of these pollutants. Twenty of 22 PCB congeners analysed were detected and quantified in sand and sandhoppers' samples collected in 6 different sites along the Tyrrhenian coast of Central Italy. Among the congeners analysed, PCB 153 and PCB 138 were the most abundant both in amphipods and sand samples and, regarding the congener pattern analysis, hexa-, hepta- and penta-PCBs were the most abundant congeners in sandhoppers. Though hydrophobic compounds, such as PCBs, bind strongly to sand and sediments, PCB concentrations were higher in T. saltator than in the sand revealing a good ability of this species to accumulate this class of contaminants. The concentration of each PCB congener recorded in tissues of sandhoppers showed significant differences among sampling sites revealing a good ability of these species to accumulate PCBs at different concentrations depending on the contamination levels of the sampling site. Therefore, our results suggest the possible utilisation of T. saltator as a biomonitor of PCB contamination of the supralittoral zone of Mediterranean sandy shores.     

Surface activity,zonation and migrations of Talitrus saltator on a Mediterranean beach

The present study considers a population of Talitrus saltator (Montagu, 1808) living along a Mediterranean beach (Burano, Grosseto, Italy). From April 1986 to March 1987 the annual and daily patterns of amphipod activity were analysed together with zonation and the direction and extent of movements. An annual bimodal activity pattern was established with a first peak of activity in late spring and another one in autumn. Daily activity was predominantly nocturanl with two peaks, one of which could be suppressed or depressed in accordance with particular climatic conditions. The species studied was generally limited to the eu- and supralittoral part of the beach, but the extent of zonation could vary according to climate. Differences between juveniles and adults appeared both for activity patterns and for zonation. Movements were predominantly (ca. 85%) along the sea/land axis with a landward migration always followed by seaward one. Under conditions of high relative humidity and rain, the migrations towards land could extend to the dune. Movements parallel to the shoreline were demonstrated in a separate experiment (in June and October 1987 and June 1988) using the capture-mark-recapture method. The maximum distance covered registered in one night was 200 m.