Reproductive plasticity in mole crabs, Emerita brasiliensis, in sandy beaches with contrasting morphodynamics

Ovarian macroscopical analysis, histological validation and field sampling procedures were used to evaluate the variability in reproductive traits of the mole crab Emerita brasiliensis Schmitt, 1935 (Decapoda: Hippidae) in two exposed sandy beaches of Uruguay with contrasting morphodynamics. All developmental stages involved in the complex life cycle exhibited lower abundance, individual size and temporal occurrence in a harsh reflective beach, compared with a more benign dissipative environment. In addition, this population showed more compressed events of the reproductive cycle (e.g oogenesis, encounter of potential mate and female parental care) and recruitment period. However, the beginning of the vitellogenesis, ovary maturity, male sexual differentiation and ending of spermatogenesis occurred at smaller sizes under dissipative conditions. These results disagree with the recent findings of delayed sexual maturity in dissipative beaches. We postulate that, in dissipative conditions, high food availability might allow an overlapping of reproductive and moulting processes, and thus females may reach optimum size and sexual maturity with fewer moults than in reflective beaches. Hence, reproductive responses must be considered not only in relation to environmental harshness, but also in the context of life history traits and their phylogenetic and allometric constraints. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ecology of sandy beaches in Oman

The benthic macrofauna and physical features of 10 sandy beaches along the coast of Oman were surveyed quantitatively. This is a mesotidal regime mostly subject to low to moderate wave energy but more exposed in the south. Five northern beaches are tide-dominated, with low wave energy, and their profiles consist of a berm, a steep, swash-dominated upper shore and a broad tide-dominated terrace from mid-shore downwards. They are composed of moderately sorted fine to medium sands. Southern beaches experience greater wave energy, particularly during the summer southwest monsoon, and exhibit smoother, concave profiles with fine, fairly well sorted carbonate sand. 58 species and species groups were recorded, with crustanceans, polychaetes and molluscs dominant. In general species richness was high, at least 19–25 species per beach, but dry biomass moderate to low at 26–90g/m shoreline, with one high value of 450g/m. Total abundance was moderate at 3–73×10³ organisms/m of beach. Some zonation was evident with ocypodid crabs andTylos in the supralittoral, cirolanid isopods on the upper shore and a variety of species on the lower shore. The coast of Oman appears to constitute a single zoogeographic region, but with some regional differentiation between north and south due to varying physical conditions. Thus, OmanÕs beaches are characterized by tide-dominated morphodynamics and exceptionally high species richness.     

A preliminary account of two sandy beaches in South West India

Observations were made on 2 beaches during the premonsoon period (January to March) when conditions were relatively stable. The beach below the berm at Cochin is 14 m wide with a slope of 1 in 8.5, and at Shertallai 40 m wide with a slope of 1 in 16. The well-sorted sand has a mean particle size of 275 at Cochin and 175 at Shertallai. The amount of chlorophyll attached to sand grains is very small. Chlorophyll in the water varies between 0.9 and 13.0 g/l. Carbon in the sand amounts to 210 to 630 g/g, and in the water to 355 to 3660 g/l. Throughout observations, the water temperature remained about 31°C, but on the dry sand surface it reached 50°C. Fauna zonation on the 2 beaches is similar to that of other tropical beaches; crabs of the genus Ocypode and the isopod Eurydice sp. occur at high water mark, the polychaetes of the genera Glycera, Onuphis, Scoloplos and Lumbrinereis in the mid-intertidal region and below; the tidal migrants Bullia melanoides (Gastropoda), Donax incarnatus and D. spiculum (Bivalvia) and Emerita holthuisi (Crustacea) inhabit the wash zone of the surf. The average biomass is 0.73 g carbon/m² at Shertallai and 0.85 g C/m² at Cochin, where fewer species are present. This compares with a value of 0.63 g C/m² for a sandy beach on the west coast of Scotland.     

Conservation implications of density-dependent predation by ghost crabs on hatchling sea turtles running the gauntlet to the sea

Protecting eggs from predators is common practice in sea turtle conservation, but routine protection of hatchlings is not. Of 42 loggerhead hatchlings observed emerging from 10 nests on undeveloped Onslow Beach, North Carolina, 24 % were preyed on by ghost crabs. In experimental trials, ghost crabs similarly threatened and captured neonate freshwater sliders, supporting their substitution as proxy for threatened and endangered sea turtle hatchlings in field experiments testing density dependence. Exploiting natural long-shore variation in ghost crab density, we show that a 2.6-fold higher ghost crab density resulted in 5 times more nocturnal threat encounters with sliders and 3.4 times more slider captures. Sliders released in simulated group emergences experienced lower per capita capture risk by ghost crabs than solitary sliders, implying predator dilution. Non-independence of egg and hatchling depredation motivates consideration of merging sea turtle egg and hatchling stages when modeling and managing food web interactions.     

Zonation of the interstitial polychaeteStygocapitella subterranea (Stygocapitellidae) in European sandy beaches

The zonation ofStygocapitella subterranea Knoellner (Polychaeta, Stygocapitellidae) in sandy beaches is compared for several localities along the European coasts. In tidal beaches the species occupies a zone near average high tide level (AHTL), whereas in non-tidal beaches it is found at a fairly long distance from the water line. The regularities observed inS. subterranea indicate that the interstitial fauna may prove well suited for the establishment of a general scheme of zonation in sandy beaches analogous to that for rocky shores proposed byStephenson andStephenson (1949).     

Meio- and macrofauna communities in three sandy beaches of the northern Adriatic Sea protected by artificial reefs

Interactions between meio- and macrofaunal assemblages and food availability were investigated in relation to tourism pressures on three beaches of the northern Adriatic Sea. All beaches were characterised by the presence of artificial reefs, freshwater inputs and tourism pressures. Beach sediments displayed high organic loads and low protein and chlorophyll a contributions to the bulk of the organic matter. Consequently, the organic matter was of mostly refractory composition and, thus, of scarce nutritional quality. Macrofaunal community structure and biomass were related to the quantity and nutritional quality of sedimentary organic matter, whereas meiofauna was apparently not. No relationships were found between physical variables and the metazoan communities. Most likely, the presence of artificial reefs reduced the physical impact of waves on the beach assemblages, whereas the presence of synergistic human impacts, including freshwater inputs and the tourism pressure, negatively influenced both the meio- and the macrofaunal assemblages.     

The ecology of two sandy beaches in south west India. II. Notes on Emerita holthuisi

The mole crab Emerita holthuisi Sankolli occurred on two beaches, at Shertallai and at Cochin, in southwest India. Observations on the size-frequency of samples for Shertallai suggest that there were two main periods of recruitment, one in the pre-monsoon,months of February and March, and one during the monsoon, although there may have been some recruitment throught the year. At Cochin, E. holthuisi was present in the pre-monsoon period, but disappeared from the beach during the monsoon.     

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.     

The ecology of two sandy beaches in south west India. III. Observations on the population of Donax incarnatus and D. spiculum

The biology of two species of Donax, D. incarnatus Gmelin and D. spiculum Reeve on two beaches in south west India is described. Two year groups of D. incarnatus were present on both beaches, from settlements during the monsoon period in 1967 and 1968. At Shertallai, both groups were studied through the year, and data on growth, mortality and production are presented. At Cochin, the rate of growth was slower and the maximum size attained smaller, but mortality during the early monsoon precluded study of D. incarnatus through a full year. D. spiculum occurred at Cochin mainly during the pre-monsoon period, and at Shertallai during the post-monsoon. The species has a shorter life-span than D. incarnatus, but its irregular occurrence did not allow detailed production estimates.     

Characteristics of feeding on dinoflagellates by newly hatched larval crabs

The zoeal larvae of brachyuran crabs must feed soon after hatching on a diet that includes large micro- and mesozooplankton in order to satisfy nutritional requirements. However, newly hatched larvae have been shown to ingest a variety of dinoflagellates, perhaps using microbial carbon sources to sustain them until they encounter more favored prey. Ingestion of dinoflagellates by larval crabs has been documented previously under conditions in which the larvae were exposed to algae provided in monoculture or in defined mixtures of cells. We report here on experiments conducted on the hatching stage of five crab species to determine if ingestion of dinoflagellates occurred when they were provided in combination with Artemia sp. nauplii or after a period of feeding on mesozooplankton. Quantitative measurements of chl a in the larval guts provided evidence of ingestion of algal cells. Active ingestion of the dinoflagellate Prorocentrum micans at specified intervals during an extended feeding period was determined on larvae of two crab species using fluorescently labeled cells provided for brief periods at prescribed time intervals. Stage 1 larvae of four of the five crab species ingested dinoflagellates when they were provided in combination with nauplii and larvae of all five species ingested cells after feeding solely on nauplii for 24 h. Ingestion of algal cells was first evident in the larval guts after 6 h of feeding at both low (200 cell ml⁻¹) and high (1,000 cells ml⁻¹) prey densities. Higher prey densities resulted in higher gut chl a. Larvae continuously exposed to dinoflagellates actively ingested cells at every 3 h interval tested over a 36 h period. Results confirm previous studies that larvae will ingest dinoflagellates even when they are encountered in a mixed prey field or when having previously fed. Ingestion of cells may occur on a continual basis over time.     

The ecology of two sandy beaches in south west India. I. Seasonal changes in physical and chemical factors,and in the macrofauna

Changes during one year in the fauna of two beaches in south India are described in relation to the nutrient content of the water and sand, and the physical changes brought about by the south-west monsoon. At both beaches, the most stable conditions were reached in the pre-monsoon months when the greatest number of species occurred and when there was marked zonation. During the monsoon, erosion took place and only actively migrating species remained to form a permanent element of the beach fauna. Data on population density and biomass indicate markedly seasonal recruitment and rapid growth of individual species, most of which are suspension feeders.     

Impacts of home shopping on vehicle operations and greenhouse gas emissions: multi-year regional study

As a shopping behavior trend, home shopping has become more popular. This shopping method seems to decrease shopping trips, yet more delivery trucks are required to be on the roads. In addition, even more personal trips may occur because saving time on shopping might allow more time for alternate activities. A study investigated the effects of home shopping on vehicle operations and greenhouse gas emissions. The purpose of that study was to identify the home-shopping impacts on transportation system, its net effects on traffic volume of the transportation network, its effects associated with environmental sustainability and then to provide some projections for future condition. Simulation results showed that home shopping will put additional burden on Newark transportation network, as identified through four measures of effectiveness (MOEs) which were travel time, delay, average speed and greenhouse gas (GHG) emissions.     

Body shape and burial behavior of the sand crab Emerita analoga (Stimpson 1857) in a reflective to intermediate morphodynamic range of sandy beaches

Sand crabs belonging to the genus Emerita are the dominant species on sandy beaches of the Pacific and Atlantic coasts of America. The success of this genus has been attributed to the great phenotypic plasticity of life history traits that enables them to cope with swash climate variability from beach to beach. The burrowing rate and variation in carapace shape of Emerita analoga from south-central Chile were studied on a series of five sandy beaches with contrasting reflective-intermediate swash conditions (harsher swash to benign swash). Environmental sampling was carried out monthly from September 2009 to November 2010, while biological experiments were conducted in September and October 2010. A combination of field experiments, geometric morphometric tools and multivariate analysis was used to assess the relationship between burrowing behavior and carapace shape with variables of swash climate. Our results showed that (1) body shape of E. analoga is directly affected by the frequency of effluent line crossing, (2) burrowing rate of E. analoga is affected directly by shape variability, (3) burrowing rate is affected directly and indirectly by the swash climate variables: frequency of effluent line crossing and mean grain size of sediment. Thus, the physical conditions in the intertidal zone may be the main environmental constraints determining cephalothorax shape, with a wider cephalothorax on beaches with finer sand and more benign physical dynamics, and a narrower shape on beaches with coarser sand and harsher physical dynamics. The latter phenotype (narrower shape) probably allows compensating the high difficulty of burying in coarser sediments.     

Effects of shell utilization on reproductive patterns in tropical hermit crabs

Dittoral and sublittoral hermit crabs (Coenobita clypeatus, Clibanarius tricolor, Calcinus dibicen, Petrochirus diogenes, Paguristes grayi, Pagurus miamensis, Paguristes tortugae, Paguristes cadenati and Pagurus sp.) were studied on Lighthouse Atoll, Belize. The Clibanarius tricolor population differed markedly in mean clutch size from congeneric populations at temperate latitudes and from conspecific populations in the Florida Keys. These differences can be attributed primarily to variation in shell utilization pattern, overlap with competitors, and breeding season. Regression analyses indicate that shell weight but not internal shell volume affects the clutch size of C. tricolor in Belize, probably due to selection for optimum volume resulting in the use of broken and encrusted shells with high weight: volume ratios. Both egg size and clutch size differed significantly among egg developmental stages in C. tricolor, Calcinus tibicen, and Pagurus miamensis.     

Spacing of leks in relation to female home ranges,habitat requirements and male attractiveness in the great snipe (Gallinago media)

Summary In an attempt to examine three main hypotheses on the evolution of leks, data on female home ranges, distance between leks and male site fidelity in the great snipe (Gallinago media) were obtained. In a 30 × 40 km area in central Sweden, the locations of 12 leks were identified. Six were within the study area (7 × 11 km) in which probably all leks were known. In the study area, nearest neighbor distances between leks were longer than the predicted drawing area (diameter of female home ranges plus the detection range of leks). Furthermore, in only 1 of 13 radio-tracked females did the estimated home range enclose many leks. These data do not fit the prediction from the hotspot hypothesis on the dispersion of leks, namely, that the distance between leks should be less than the drawing area and hence an average female home range should enclose more than one lek. The observed distance between leks was about the same as predicted by the female preference hypothesis. This hypothesis also predicts females visit mainly one lek; however, we found they sometimes visit two and nest close to a third. We propose that males may first settle according to hotspot rules, but females will resettle according to preferences for certain males and/or larger leks. In this way some hotspot leks are abandoned, and the distance between leks is increased as males become more tightly clumped. However, when all males and females have settled, the leks still existing would be on hotspots. In the female preference hypothesis, males are assumed to aggregate on leks because females prefer clustered males as they can be more easily compared than if they are dispersed. Alternatively, in the attractiveness hypothesis, it is suggested that females prefer certain males, and hence unattractive males surround the attractive are forced to join leks in order to come close to females. Our data on male site fidelity support the attractiveness hypothesis because successful and dominant males return to the same lek and territory, both within and between years, whereas unsuccessful males move to other leks. Attractiveness of certain males may explain why according to hotspot rules in this species males are more clumped than expected. Offprint requests to: J. Höglund     

Rehabilitation of metropolitan beaches by local administrations in Catalonia: new trends in sustainable coastal management

The planning and management of coastal sites in Catalonia has been mainly concerned with rendering beaches functional for mass frequentation. This has caused serious problems for the beaches, including the alteration of dune formation processes and the destruction of beach vegetation and habitats. Municipal capacity to plan and manage beaches is theoretically very limited and relegated to the design of plans related to the maintenance of facilities for beach users. Nonetheless, a singular experience, led by a local council in the metropolitan area of Barcelona demonstrates the crucial role that a local administration can play in achieving a balance between habitat preservation and social use based on sustainable coastal management objectives, in spite of a restrictive legislation.     

Physical damage to coastal dunes and ecological impacts caused by vehicle tracks associated with beach camping on sandy shores: a case study from Fraser Island,Australia

As coastal populations expand, demands for recreational opportunities on beaches and coastal dunes grow correspondingly. Although dunes are known to be sensitive to direct human disturbance and provide irreplaceable ecosystem services (e.g. erosion control, critical habitat and nesting sites), dunes serve as campsites for large numbers of people (∼90,000 p.a.) on the ocean-exposed shores of Fraser Island, Australia. Campsites are located in the established dunes and can only be accessed with 4WD vehicles along tracks cut directly from the beach through the foredunes. Here we quantified the extent of physical damage to foredunes caused by this practice, and tested whether human-induced physical changes to foredunes translate into biological effects. Of the 124 km of ocean-exposed beaches, 122 km (98%) are open to vehicles driven on the beaches, and camping zones cover 28.7 km or 23% of the dunes. A total of 235 vehicle tracks are cut across the foredunes at an average density of eight tracks per km of beach. These tracks have effectively destroyed one-fifth (20.2%) of the dune front in camping zones, deeply incising the dune-beach interface. There is evidence of accelerated erosion and shoreline retreat centred around vehicle tracks, resulting in a “scalloping” of the shoreline. No dune vegetation remains in the tracks and the abundance of ghost crabs (Ocypode spp.) is significantly reduced compared with the abutting dunes. Because current levels of environmental change caused by dune camping may not be compatible with the sustainable use of coastal resources and conservation obligations for the island (listed as a World Heritage Area and gazetted as a National Park), restoration and mitigation interventions are critical. These will require spatial prioritisation of effort, and we present a multi-criteria ranking method, based on quantitative measures of environmental damage and ecological attributes, to objectively target rehabilitation and conservation measures. Ultimately, coastal management needs to develop and implement strategies that reconcile demands for human recreation, including beach camping, with conservation of coastal dune ecosystems.