Competitive interactions and substratum preferences of two intertidal isopods

The intertidal isopods Gnorimosphaeroma oregonensis oregonensis (Dana) and Exosphaeroma amplicauda (Stimpson) are shown to have differing preferences for substratum particle-size categories. In field sampling, 80.9% of the total number of G. oregonensis oregonensis taken were present on substrata falling within a ?-4.2 to-3.8 diameter size-range, while 60.0% of E. amplicauda occupied the range ?-2.5 to-2.1. These relationships changed upon laboratory manipulation of density ratios, and suggest competitive interaction: when E. amplicauda is tested alone, the resultant preference assumes a greater niche breadth and overlap to that of the field. Introduction of G. oregonensis oregonensis produces a displacement of E. amplicauda, the degree being dependent upon their relative numerical strength. With an experimental ratio of 3:1 (G. oregonensis:E. amplicauda) there is a simulation of field densities, and the laboratory results approximate more closely the overlap in nature.     

Role of peat organic matter on isotopic composition of most abundant benthic organisms in intertidal habitats of SW Iceland

The transfer of peat organic matter (OM) from tidal pools of upper littoral to downstream rocky shores and its potential incorporation into marine biota were investigated using the stable isotope approach. Samplings were carried out in September 2004 in two SW Icelandic sites (Osar and Hvassahraun), where we selected (1) areas with shores where grass gently declined towards rocks and, on reaching the sea line, formed small tidal peat pools and (2) areas where grass and rocky shores were not contiguous, rather the grass lower limits were several hundreds of meters back from the shore. In both grass and no grass areas, in the intertidal zone, all benthic organisms and all potential OM sources were sampled. Dominant macroalgae were Ascophyllum nodosum ([Linnaeus] Le Jolis) and Fucus vesiculosus Linnaeus. In grass sites, organisms partially reflected the isotopic composition of peat OM, while in no grass sites, peat represented only a negligible amount of available OM, or its signal was totally absent from the environment. Intertidal organisms could rely on peat both directly by suspension feeding, grazing or scavenging and indirectly by predation (crabs) on low level consumers (barnacles).     

Structure and temporal fluctuations of two intertidal seagrass-bed communities in New Caledonia (SW Pacific Ocean)

The structure and trophic organization of two intertidal seagrass-bed communities (Halodule uninervis and Thalassia hemprichii) were examined on the southwest coast of New Caledonia (SW Pacific Ocean), from April 1989 to March 1990. Five benthic samples were collected from each site at 2 mo intervals and various environmental parameters were simultaneously monitored. Animal:plant biomass ratios were close to 1 at both sites. Polychaetes dominated in number of species. The suspension-feeding bivalves Gafrarium tumidum and Anadara scapha constituted the greater part of the animal biomass. The evolution of the communities over an annual cycle displayed marked structural and organizational stability, resulting mainly from the absence of distinct recruitment periods for the dominant species, no mortalities during the brief low-salinity periods, and no temporal variations in the granulometry, the organic matter or the chlorophyll a contents of the sediments.     

Ecology of subtropical hermit crabs in SW Madagascar: short-range migrations

Many mobile animals migrate because of the different benefits provided by different localities in time and space. For hermit crabs, such benefits include resource (shell, water, food) acquisition and gamete release. One of the more successful crustacean land-invaders, Coenobita hermit crabs, undertake complex short-range migrations in SW Madagascar. Number of active hermit crabs was inversely related to wind strength and positively related to tidal range, emphasising that movement would conserve water. A circadian component was also recorded in the locomotory activity of Coenobita pseudorugosus and C. rugosus. Path linearity varied with many of the same parameters, but also with beach slope. Movement was primarily perpendicular to shore in small individuals, but the parallel proportion increased with hermit crab size and tidal range, probably driven mostly by shell and food searching. Despite the costs of movement and shell carriage in the terrestrial environment, C. pseudorugosus and C. rugosus were as fast as their marine counterparts. Their speeds varied principally with individual size and were approximately 20% faster without shells and about 20% slower when climbing up a 20° slope, compared to horizontal or downhill travel. Hermit crabs, which are highly numerous and speciose in SW Madagascar, do not seem to partition niches by differential movement patterns. Aside from provision of shells in middens and capturing large adults for bait or pets, human activity may have a profound effect on hermit crab movement: observations at rare uninhabited marine reserves like Nosy Ve show that considerable diurnal activity may take place despite the apparent hostility of the environment to an essentially marine animal.     

The breeding biology of two sympatric species of tropical intertidal hermit crabs,Clibanarius chapini and C. senegalensis

A one-year study of breeding in the two intertidal hermit crabs Clibanarius chapini Schmitt and C. senegalensis Chevreux and Bouvier showed that both species maintain a high reproductive activity throughout the year. Brief low reproductive activities, found in both species, occurred in different months: C. chapini in November, C. senegalensis in May. While these drops cannot be attributed to any recognisable environmental factor, it is suggested that such pattern may result from competition between the two species. Sex-ratio studies indicated slightly more females than males in both species. The maximal size attained by males of both species is considerably greater than that of the females; in C. senegalensis this is very marked. It is suggested that it may be selectively advantageous for a female to attain a smaller maximal size since her brood must also be accomodated within the shell. As expected for continuously breeding species, recruitment of young into the population is also continuous.     

Resource utilization for decorating in three intertidal majid crabs (Brachyura: Majidae)

 For three spider crabs (Tiarinia cornigera, Micippaplatipes and Pugettia quadridens quadridens), patterns of algal utilization for decorating were compared with the dynamics of algae on an intertidal rocky shore reef where the crabs co-occurred. T. cornigera and P. quadridens quadridens were most abundant from autumn to spring when the dominant algae (Boodleacoacta, Sargassum hemiphyllum, S. thunbergii and Corallina pilulifera) occurred in high coverage, while M. platipes was most abundant from spring to autumn. Monthly change of algae used for decorating was not correlated with algae growing in the crab habitat for T. cornigera, but for M. platipes, it was positively correlated for two algal species, and for P. quadridens quadridens, negatively correlated for one algal species. Each species of the spider crabs used some algal species preferentially for decoration. Decorating preference experiments conducted in the laboratory showed that M. platipes and P. quadridens quadridens exhibited similar preference to their algal utilization in the field, whereas for T. cornigera, algal preference in the experiment differed from utilization in the field. Comparisons between materials used for decoration and gut contents revealed that T. cornigera and M. platipes used algal species differently for decorating and feeding, while P. quadridens quadridens used the same algal species for both decorating and feeding. Different tactics for camouflage are discussed in terms of algal utilizations by the three majid species. Received: 25 September 1999 / Accepted: 22 June 2000     

Desiccation stress in two intertidal beachrock biofilms

Chlorophyll a fluorescence was used to look at the effect of desiccation on the photophysiology in two beachrock microbial biofilms from the intertidal rock platform of Heron Island, Australia. The photophysiological response to desiccation differed between the beachrock microbial communities. The black biofilm from the upper shoreline, dominated by Calothrix sp., showed a response typical of desiccation-tolerant cyanobacteria, where photosynthesis closed down during air exposure with a rapid and complete recovery upon rehydration. In contrast, the pink biofilm from the mid-intertidal zone, dominated by Blennothrix sp., showed no distinct response to desiccation stress and instead maintained reduced photosynthesis throughout drying and re-wetting cycles. Spatial differences in photosynthetic activity within the black biofilm were evident with a faster recovery rate of photosynthesis in the surface cyanobacteria than in the deeper layers of the biofilm. There was no variation with depth in the pink biofilm. The photophysiological differences in desiccation responses between the beachrock biofilms exemplify the ecological niche specialisation of these complex microbial communities, where the functional differences help to explain their vertical distribution on the intertidal shoreline.     

Latitudinal patterns of diversity and abundance in North Atlantic intertidal boulder-fields

In order to study taxon richness, biodiversity and abundance patterns in the North Atlantic from temperate latitudes through Arctic to high Arctic latitudes, we recorded the faunas (at ELWS level) colonizing 20 cobbles from three sites at each of seven boulder-field localities (south-west England, 50°N; Wales, 51°N; west Scotland, 56°N; Iceland, 64°N; Tromsø, 70°N; Svalbard, 77°N, 79°N). Inverse correlations were found between latitude and all measures of richness (species, orders, and phyla numbers) and biodiversity (S–W, P). However, these correlations were driven mostly by the consistently impoverished Arctic sites; an even cline of decreasing diversity from south to north did not exist. Multidimensional scaling revealed two communities, temperate–subarctic species-rich and high arctic species-poor. Evenness as measured by Pielou’s index was similar across all latitudes. Abundance data exhibited a similar trend to the species richness and diversity data with a significant negative correlation with latitude, but when Arctic data were excluded the correlation vanished.     

Endolithic invertebrate communities and bioerosion rates in southwestern Atlantic intertidal consolidated sediments

Organisms boring into intertidal consolidated sediments generate bioerosion. It is generally unknown, however, whether they can significantly contribute to coastline retraction. In this paper, we describe endolithic communities and estimate bioerosion and physical erosion rates at three southwestern Atlantic intertidal sites (37, 38, and 42°S; Argentina). In the northernmost site, we have also analyzed spatial variation in species richness and abundance as a function of height within the tidal slope, orientation of the rock surface in relation to breaking waves (i.e., facing or not), and rock hardness. The number of species and the combined abundance of individuals from the different species were larger at the low intertidal level but did not differ between surface orientations. The density of chemically boring organisms increased with increasing rock hardness and calcium carbonate content. In contrast, no correlation was found between rock hardness and the abundance of organisms that bore by mechanical means. Endolithic community composition and bioerosion rates differed among the three sites, the latter being higher at the site with the softer substrate. Bioerosion estimates were two orders of magnitude lower than physical erosion estimates at each site. The bivalve Lithophaga patagonica was the species that contributed the most to bioerosion at all these locations. While results suggest that bioerosion contributes little to overall coastal erosion at the three study sites, boring organisms might still facilitate physical erosion by weakening the rock either via chemical or mechanical means. Besides, their apparently inconsequential direct action as bioeroders can have positive consequences for biodiversity via increased habitat complexity.     

Space partitioning and interactions in an intertidal sand-burrowing amphipod guild

A nine-year study at a moderately exposed, fine sand intertidal beach at Long Sands, Maine, USA identified marked seasonal changes in the foci of sand-burrowing amphipod activities. These included seasonal population movements, degree of interspecific horizontal overlap, life stage segregation within and between species, and sand depth stratification. Behavioral interactions related to competition for available space are considered important in affecting the distribution and abundance of amphipod species, and community structure. Significant negative correlations were shown between the abundance and upper distributional limits of the smaller, lower shore Acanthohaustorius millsi and the abundance of the larger, higher shore species Haustorius canadensis. Tolerance limits to a variety of prevailing physical factors alone, were not considered a serious obstacle to occupation of higher intertidal level sands by A. millsi. Summer brooding females of both species occupied similar 5 to 10 cm sand depths, but were strongly segregated horizontally. Laboratory coexistence experiments during the reproductive period showed increased mortality of adults and negligible reproductive output of A. millsi when in combination with H. canadensis compared to controls where the former species was alone. A third species, Amphiporeia virginiana, occupied primarily very shallow sand (0 to 2.5 cm), and performed seasonal movements opposite in direction to A. millsi and H. canadensis. The presence, amount, and refinement of biological interactions across the wave exposure gradient requires further study. Until then, we consider as inappropriate the blanket designation of all types of intertidal sand communities as physically controlled.     

Comparison of mussel bed community between two intertidal mytilids Septifier virgatus and Hormomya mutabilis

Mussel bed community structure of two intertidal mytilids, Septifer virgatus (Wiegmann) and Hormomya mutabilis (Gould), whose beds were contiguous vertically on a rocky intertidal shore (Wakayama Prefecture, Japan), was compared between 1982 and 1983. In the upper S. virgatus bed, crustaceans and bivalves were dominant in terms of both number of individuals and biomass. There were three barnacle species representative of the epizoans, two isopods and one amphipod as mobile fauna, and two bivalves as infauna. The lower H. mutabilis bed supported virtually no epizoans or mobile fauna. Infaunal free-ranging polychaetes and sipunculids were dominant in terms of both number of individuals and biomass. The H. mutabilis bed contained a much greater amount of sediment than did the S. virgatus bed, and the interstices among individual H. mutabilis and among their byssal threads were filled with sediment. The biomass of six of nine species dominant in the S. virgatus bed was negatively correlated with the amount of sediment. Recruitment of these faunal assemblages into artificial mussel clumps was examined in mussel enclosure experiments, and a negative sediment effect in H. mutabilis clumps was detected for one isopod and one limpet species. Factors causing the differences between two mussel bed communities are discussed, focusing on the effects of sediment.     

Causes and consequences of salt-marsh erosion in an Atlantic estuary in SW Spain

This study reports on the quantification of horizontal erosion by undermining of slopes in the atlantic mesotidal salt marshes of Odiel, SW Spain, and analyses its causes and consequences. Horizontal erosion has produced considerable losses of salt marsh area, including zones of mature salt marsh. Human pressure, such as from water-borne traffic or the exploitation of the slopes for the capture of bait, increases the natural erosion processes. The role of vegetation in protecting the slopes against erosion is studied. Channel banks covered with plants, many of which belong to species with long-living, above-ground creeping stems, were less eroded than those without vegetation cover. The enormous volume of sediments moved (ca. 7000m³ in one year) could contribute to the silting-up of the navigable channels of the estuary, so that continual dredging is necessary to allow access to shipping. These sediments are highly contaminated, and dredging exposes them more directly to the trophic network of the estuary. There is a considerable loss of natural resources. Finally, the integrated management of this coastal ecosystem is discussed.     

Habitat complexity impacts persistence and species interactions in an intertidal whelk

Although experiments have shown that habitat structure may influence the distribution of species and species interactions, these effects are still not commonly integrated into studies of community dynamics. Since habitat structure often varies within and among communities, this may limit our understanding of how various factors influence communities. Here, we examined how mussel bed complexity (the presence and thickness of mussel layers) influenced the persistence of whelks (Nucella emarginata) and interactions with a top predator (ochre sea stars, Pisaster ochraceus) and prey (mussels, Mytilus californianus). Results from a mark?Crecapture experiment indicate that whelk recapture rates are higher in more complex habitats, and laboratory experiments demonstrate that habitat complexity affects whelk feeding, growth, and nonconsumptive interactions with a keystone predator. Habitat complexity therefore has direct effects on species and also may lead to trade-offs among feeding, refuge, and other factors, potentially influencing the distribution of whelks and the effects of both whelks and sea stars on intertidal communities. These results demonstrate that habitat structure may play an important role in intertidal communities and other habitats and should be further considered in the experimental design of future studies of community dynamics.     

Comparison of spatial overlap between the polychaetes Nereis virens and Nephtys caeca in two intertidal estuarine environments

An intensive study of spatial overlap between the polychaetes Nereis virens (Sars) and Nephtys caeca (Fabricius) was conducted in 1992 on two tidal sand flats, which differ by their exposure to dominant winds and residual currents in the Lower St. Lawrence Estuary (Québec, Canada). Results showed that spatial overlap (Lloyd's index and spatial distribution) was higher among adults of both species at the lower tidal elevation of l'Anse-à-l'Orignal (north-east oriented). Results also suggest weaker interspecific interactions among juveniles of both species because of limited spatial overlap. In Baie-du-Ha!Ha! (south-west oriented), spatial overlap was greater than that observed in l'Anse-à-l'Orignal and appeared important in adults as well as in juveniles. Levins' directional measure of competition indicated an asymmetric spatial overlap between N. virens and N. caeca in Baie-du-Ha!Ha! and a symmetric overlap in l'Anse-à-l'Orignal. The intraspecific encounter values, estimated from Lloyd's mathematical expression, were significantly higher than values of interspecific interactions only in l'Anse-à-l'Orignal. Moreover, no vertical stratification was found inside the sediment, with no effect of the densities and individual body weights of the polychaetes. Juveniles of both species mainly inhabited the organic-rich upper portion of the sediments (0 to 12 cm), while adults colonised greater depths (>25 cm) where organic matter content was lower. A complementary field experiment was conducted in 1993 to investigate interspecific interactions (predation and competition) existing between N. virens and N. caeca. Results from this experiment depend on which species was first-introduced and showed an important variation in mortality rates between allopatric and sympatric conditions. The influence of competition and predation on the structure of these populations is also discussed in relation to their spatial overlap.     

Use of artificial holes in studying community development in cryptic marine habitats in a tropical rocky intertidal region

Previous studies on the rocky intertidal shores of the Bay of Panama indicate that for many sessile and mobile organisms holes and crevices are important refuges from consumers. To study the dynamics of these hole-dwelling species assemblages, we devised a method allowing repetitive, non-destructive sampling and species manipulations in artificial holes. These are fiberglass sleeves with flanges at the outer edge that are inserted into receptacles made of quick-setting concrete. Predator and herbivore manipulations include: fences excluding slow-moving benthic consumers, bars across hole mouths restricting entry of large fishes and homogeneous (shelterless) surfaces created around a treatment decreasing abundance of small crabs. This design has withstood the rigors of two wet seasons in Panama. Results from two sites in Panama indicate that colonization occurs rapidly (6–7 months) in holes established in mid dry season. Encrusting fleshy algae (Ralfsia sp.) and ephemeral green algae (Cladophora spp.) colonize first and are succeeded by encrusting species of coralline algae, bryozoans and colonial tunicates and by erect fleshy red algae. Where consumers are present, encrusting corallines predominate; without consumers, bryozoans, tunicates and erect algae are more abundant. Zonation patterns develop within the holes, with the desiccation/heattolerant Ralfsia sp. dominating at the outer edges. With increasing deph, encrusting coralline algae, bryozoans and colonial tunicates reach their respective peak abundances. In contrast to the high variability observed among naturally occurring holes, replicates of each treatment tend to be similar. Between-treatment and between-site comparisons are less so. The naturally-occurring high level of small-scale patchiness is thus presumably due to variation in recruitment and in the local (microspatial) consumer regime. Substratum heterogeneity is therefore directly and indirectly important in maintaining a high local diversity in this community. As in other studies, consumers or disturbances are key factors in regulating patterns of community structure. More experiments of longer duration are necessary to ascertain the relative importances of consumers/disturbance and competition in controlling such assemblages.     

A jellyfish diet for the herbivorous green turtle Chelonia mydas in the temperate SW Atlantic

Feeding ecology of juvenile green turtles (Chelonia mydas) was studied from 2008 to 2011 at Samborombón Bay (35°30′–36°30′S, Argentina), combining data on digestive tract examination and stable isotope analysis through a Bayesian mixing model. We found that animal matter, in particular gelatinous plankton, was consumed in large proportions compared to herbivorous food items such as terrestrial plants and macroalgae. This diet is facilitated by the high abundance of gelatinous plankton in the region, thus confirming the adaptive foraging behaviour of the juveniles according to prey abundance in the SW Atlantic. To our knowledge, this is the first study to employ this combination of techniques and to conclusively demonstrate that animal matter, in particular gelatinous plankton, is important in the diet of the neritic green sea turtles.