A comparison of spatial learning and memory capabilities in intertidal gobies

For the majority of animals, the ability to orient in familiar locations is a fundamental part of life, and spatial memory allows individuals to remember key locations such as food patches, shelter, mating sites or areas regularly occupied by predators. This study determined if gobies collected from rocky platforms and sandy beaches differ in their ability to learn and memorise the locations of tide pools in a simulated rocky intertidal zone. Intertidal rock pool gobies show acute homing abilities and, therefore, should be expected to display superior learning and memory capabilities. In contrast, it is unlikely that natural selection would favour similar learning skills in sandy shore fishes because they simply shift back and forth with the tides. The learning abilities of gobies were tested using small replica rock platforms, containing four tide pools that retained varying depths of water at simulated low tide. Gobies were categorised as having learnt the task if they were able to consistently locate the tide pool that retained the most water at simulated low tide as the pool with the most favourable conditions. Rock pool species were able to locate the deepest pool to wait out low tide for ~95 % of the trials, while species from sandy shores were found in the deepest pool ~10 % of trials. Despite repeated stranding, sandy shore fish continued to follow the tide out. We propose that rock pool species memorised the location of rock pools during simulated high tide enabling them to relocate the best refuge for low tide.     

Movements and site fidelity of grass shrimp (<Emphasis Type="Italic">Palaemonetes pugio</Emphasis> and <Emphasis Type="Italic">P. vulgaris</Emphasis>) in salt marsh intertidal creeks

Grass shrimp are abundant residents of shallow-water habitats in US Atlantic and Gulf coast estuaries. Mark-recapture events determined the extent and direction of their movements in meso-tidal salt marsh creeks at North Inlet, South Carolina. Microwire-tagged Palaemonetes vulgaris demonstrated high fidelity for a dock in a subtidal channel. In the 48 days following release, 11 % of the 422 tagged shrimp were recaptured and 99 % of all recaptures were made there an average of 14 days post-release. Palaemonetes pugio, which favored the use of shallow intertidal areas, exhibited moderate fidelity for pools within creek beds with 53 % of the recaptured shrimp collected at the release site. Higher fidelity was observed for confluences of intertidal creeks and the subtidal channels. Of the 5565 shrimp released with coded microwire tags, 7.5 % were recaptured and 98 % of those recaptures were at release sites up to 53 days later. During another event, P. pugio released 235 m from their origin demonstrated relatively low fidelity for the new site and a lack of homing behavior. Of the 544 tagged shrimp, 3 % were recaptured; only 40 % of those recaptured were from the release site. Site fidelity is a mechanism by which previously discovered large and consistent differences in grass shrimp use of neighboring intertidal creek basins can be sustained over weeks, seasons, and years. Spatial variations and the tendency of these keystone organisms to resist relocation by tidal currents and occupy certain areas for extended periods should be considered in decisions about proposed alterations to creeks and shorelines.     

Corallina and Ellisolandia (Corallinales,Rhodophyta) photophysiology over daylight tidal emersion: interactions with irradiance,temperature and carbonate chemistry

The photophysiology of three geniculate coralline algal species (Corallina officinalis, C. caespitosa and Ellisolandia elongata) was determined in intertidal rock pools in the south-west UK at Combe Martin (51°12′31N 4°2′19W) and Heybrook Bay (50°31′66N 4°11′41W), at the start, middle and end of summer (September 1 and 2) and winter (February 9 and 10) daylight tidal emersion periods, in relation to prevailing irradiance, temperature and carbonate chemistry conditions. Algal photophysiology was assessed from rapid light curves performed using pulse amplitude modulation fluorometry. Corallina and Ellisolandia experienced significant fluctuations in irradiance, temperature and carbonate chemistry over seasonal and tidal cycles. Rock pool carbonate chemistry was predictable (R ² = 0.82, P < 0.0001) by photodose (summed irradiance) plus water temperature, but not significantly related to photophysiology. In contrast, Corallina and Ellisolandia relative maximum electron transfer rate showed a significant negative relationship (R ² = 0.65, P < 0.0001) with irradiance plus water temperature. At a seasonal resolution, photoacclimation to maximize both light harvesting during winter months and photoprotection during summer months was observed for all species. Dynamic photoinhibition was apparent over both summer and winter tidal emersion, in relation to irradiance fluctuations. More effective photoinhibition was apparent during summer months, with greater sensitivity to irradiance and slower recovery in F _v/F _m, observed during winter. With sustained high irradiance over tidal emersion, the establishment of high pH/low inorganic carbon conditions may impact photochemistry. This study represents the first assessment of C. officinalis, C. caespitosa and E. elongata photophysiology underpinned by clear species concepts and highlights their ability to adapt to the dramatically fluctuating conditions experienced in intertidal rock pools.     

Host preference,site fidelity,and homing behavior of the symbiotically luminous cardinalfish,Siphamia tubifer (Perciformes: Apogonidae)

The sea urchin cardinalfish, Siphamia tubifer (Perciformes: Apogonidae), is unusual among coral reef fishes for its use of bioluminescence, produced by symbiotic bacteria, while foraging at night. As a foundation for understanding the relationship between the symbiosis and the ecology of the fish, this study examined the diel behavior, host urchin preference, site fidelity, and homing of S. tubifer in June and July of 2012 and 2013 at reefs near Sesoko Island, Okinawa, Japan (26°38′N, 127°52′E). After foraging, S. tubifer aggregated in groups among the spines of the longspine sea urchin, Diadema setosum, and the banded sea urchin, Echinothrix calamaris. A preference for D. setosum was evident (P < 0.001), especially by larger individuals (>25 mm standard length, P < 0.01), and choice experiments demonstrated the ability of S. tubifer to recognize and orient to a host urchin and to conspecifics. Tagging studies revealed that S. tubifer exhibits daily fidelity to a host urchin; 43–50 and 26–37 % of tagged individuals were associated with the same urchin after 3 and 7 days. Tagged fish also returned to their site of origin after displacement; by day two, 23–43 and 27–33 % of tagged individuals returned from displacement distances of 1 and 2 km. These results suggest that S. tubifer uses various environmental cues for homing and site fidelity; similar behaviors and cues might be used by larvae for recruitment to settlement sites and for the acquisition of luminous symbiotic bacteria.     

Site fidelity and homing behavior in juvenile loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>)

To investigate site fidelity and homing behavior in juvenile loggerheads ( Caretta caretta, L.), a mark-recapture study spanning four years (1998–2001) was conducted in Core Sound, N.C., USA. Each year of the study, approximately half of the turtles captured were tagged and released near the capture sites ( n=207), while the remaining turtles were displaced 15–20 km and released ( n=198). Loggerheads in both groups were recaptured in equal proportions near the original capture sites and many individuals were also recaptured in subsequent years. These data imply that juvenile loggerheads often returned to their capture sites following displacement, because if turtles dispersed randomly or remained near their release sites, then fewer displaced turtles should have been caught again. Moreover, because turtles migrate out of North Carolina sounds each winter, turtles recaptured at the same locations in different years evidently returned to specific sites following long migrations. To further investigate homing behavior, a small number of displaced turtles ( n=28) were tracked using radio telemetry following their release. Although transmitters detached from most turtles within a few days, analyses of initial headings showed strong orientation in the direction of the capture site. In addition, four turtles successfully tracked for longer periods of time all returned rapidly to the vicinity of the capture location and remained in the area. Taken together, the results of this study indicate that juvenile loggerheads exhibit fidelity to specific areas during summer months and possess the navigational abilities to home to these areas following forced displacements and long-distance migrations.     

Monitoring spatio-temporal variation in molluscan grazing pressure in seasonal, tropical rock pools

In Hong Kong, low-shore rock pools support a patchy mosaic of different successional stages of crustose coralline algae. Echinoid and molluscan grazers are abundant in these low-shore rock pools and spatial and temporal variation in grazing pressure is thought to play an important role in structuring these assemblages. Direct records of graze marks using a modified wax-disc method were used to monitor spatial (different pools) and temporal (periods, 3 or 4 days each; spring vs. neap tides; summer vs. winter) variations in grazing pressure. Radula marks of Chlorostoma argyrostoma, Lunella coronata and Nerita albicilla were common in all pools. Grazing pressure was highly variable with tidal state and season but also within these two factors. Lunella and Nerita exerted highest grazing pressure at neap tides during summer, whilst Chlorostoma exerted highest grazing pressure in winter. Although overall patterns of temporal variation were similar and consistent in all pools, there was between pool variation in grazing pressure and individual species showed different patterns. Within the seasonal and tidal variations, large period-to-period differences were found for all species, illustrating small-scale temporal variation. Fine-scale spatial and temporal patterns of variation in grazing pressure of different species, therefore, contribute to the highly dynamic nature of these low-shore rock pools.     

A quantitative study of intertidepool movement of the wooly sculpin Clinocottus analis

Movements of the wooly sculpin Clinocottus analis between all pools in a designated area of a southern California (USA) intertidal zone were studied. Fish were captured during low tides, finclipped or bead-tagged, and released in the pool of capture. Percent recaptured in pool of first capture declined from 30 after 2 weeks to about 5 after 20 weeks. Percent recaptured in any pool in the study area declined from 50 after 2 weeks to 20 after 8 weeks, but remained at 20 through 16 weeks. A higher fidelity to an area of the intertidal zone than to a specific pool was suggested. Large fish exhibited higher probability of recapture than smaller individuals, indicating they become more sedentary with age. Changes in pool topography influenced percent recurrence in a pool, most likely as a consequence of changes in quality and quantity of cover available. Such responses to changes in specific pools and fidelity to intertidal areas containing several pools appear to be advantageous behavioral patterns in an environment where topographic change is frequent and often massive.     

Size and spatial arrangement of home range of checkered snapper <Emphasis Type="Italic">Lutjanus decussatus</Emphasis> (Lutjanidae) in an Okinawan coral reef determined using a portable GPS receiver

Although snappers (Lutjanidae) are commercially important fishery resources in tropical and subtropical waters, their home range size and its spatial arrangement have not been sufficiently clarified. In the present study, the size and spatial arrangement of the home range of the reef-associated checkered snapper Lutjanus decussatus (Lutjanidae) were investigated by use of a portable GPS receiver. In a 120 m × 120 m quadrat established on a section of the fringing reef of Ishigaki Island, 29 individuals of the species were identified by their color pattern and individuals were divided into five arbitrary size classes (class 1 <10 cm TL ≤ class 2 <15 cm TL ≤ class 3 <20 cm TL ≤ class 4 <25 cm TL ≤ class 5). Fish tracking by snorkeling was conducted with the portable GPS receiver. Site fidelity of this species was high. Home range size ranged from 93.0 to 3638.4 m², and there was a significant positive correlation between the home range size and fish total length. Home ranges of the same-sized individuals abutted each other (8.8% area overlap), whereas those of different-sized individuals overlapped (44.0% area overlap). Agonistic behavior (attack and agonistic display) was more frequently found among same-sized individuals (times of agonistic behavior/times of all encounters × 100 = 71.3%), whereas such agonistic behavior was rarely found among different-sized individuals (times of agonistic behavior/times of all encounters × 100 = 6.9%). These results suggest that home ranges of Lutjanus decussatus can be regarded as territories against same-sized individuals, but not different-sized individuals. The usefulness of the fish tracking by snorkeling using a portable GPS receiver for home range size estimation and the function of the overlapping territory of the species are discussed.     

Population genetic structure of protected allis shad (<Emphasis Type="Italic">Alosa alosa</Emphasis>) and twaite shad (<Emphasis Type="Italic">Alosa fallax</Emphasis>)

Determining the magnitude of homing behaviour within migratory fish species is essential for their conservation and management. We tested for population genetic structuring in the anadromous alosines, Alosa alosa and A. fallax fallax, to establish fidelity of stocks to spawning grounds in the United Kingdom and Ireland. Considerable genetic differences were present among populations of both species, suggesting strong fidelity to breeding grounds and compatible with homing to natal origins. Moreover, the genetic structure of A. fallax fallax showed a clear pattern of isolation-by-distance, consistent with breeding populations exchanging migrants primarily with neighbouring populations. Spatial genetic differences were on average much greater than temporal differences, indicating relatively stable genetic structure. Comparing anadromous A. fallax fallax populations to the landlocked Killarney shad subspecies, A. fallax killarnensis (Ireland), demonstrated a long history of separation. These results demonstrating regional stock structure within the British Isles will inform practical management of stocks and their spawning habitats.     

Effects of grazer richness and composition on algal biomass in a closed and open marine system

Most natural local systems exchange organisms with a regional pool of species through migration and dispersal. Such metacommunity processes of interconnected multispecies assemblages are likely to affect local dynamics of both species and processes. We present results from an artificial marine outdoor rock pool system in which we investigated the factors of (1) local grazer richness and composition, and (2) connectivity of local patches to a regional species pool, and their effects on algal biomass. Local species richness of six grazers was manipulated in both open and closed pools, which were embedded in a regional species pool containing all six grazers. Grazer richness showed significant net biodiversity effects on grazing in the closed, but not in the open, system. Grazer composition, on the other hand, showed significant effects on grazing in both open and closed systems, depending on which species were initially present. The two most efficient grazers were able to compensate for less efficient grazers in species mixtures, hence ensuring the function of grazing. The efficiency of top-down control of algal biomass in open systems thus depends on which particular species are lost. Further, differences in grazing between the open and closed system changed over time due to temporal dynamics in grazer composition. The results emphasize the importance of including system connectivity in experimental designs to allow an extrapolation of biodiversity ecosystem-functioning relationships to natural systems.     

Ecology of Echinometra mathaei (Echinoidea: Echinodermata) at Diani Beach,Kenya

Studies were carried out on the inner and outer coral reefs at Diani Beach on the Kenya coast to assess the distribution, density and behaviour of Echinometra mathaei (de Blainville). Transects 1 m wide were run on the two reefs in April, June and September, 1970. Test measurements on representative samples from the animal populations on both reefs were also taken. Direct observations on specimens of E. mathaci in selected rock pools on the outer reef were made to determine their movement, gregariousness, homing and feeding behaviour. Population density was higher on the inner reef furthest from the sea at low tide than on the outer reef. On the submerged inner coral reef at low tide, E. mathaei occurred mainly exposed on the seaweeds, but, on the exposed outer reef, its main niches were crevices in rock pools and under coral ledges. Sizefrequency distributions revealed that smaller individuals occurred on the inner reef and larger ones on the outer reef. The growth rate of E. mathaei was estimated from the positions of modal values, calculated from size-frequency distributions. No gregarious or homing behaviour was observed and, once settled in a suitable crevice, E. mathaei showed little movement.     

Strange parental decisions: fathers of the dyeing poison frog deposit their tadpoles in pools occupied by large cannibals

Parents may increase the probability of offspring survival by choosing suitable rearing sites where risks are as low as possible. Predation and competition are major selective pressures influencing the evolution of rearing site selection. Poison frogs look after their clutches and deposit the newly hatched tadpoles in bodies of water where they remain until metamorphosis. In some species, cannibalism occurs, so parents deposit their tadpoles singly in very small pools. However, cannibalism also occurs in species that deposit tadpoles in larger pools already occupied by heterospecific or conspecific larvae that could be either potential predators or competitors. Here, I test the hypothesis that, given the choice, males of Dendrobates tinctorius would deposit their newly hatched tadpoles in low-risk sites for their offspring. I characterised the pools used by D. tinctorius for tadpole deposition, conducted experiments to determine the larval traits that predict the occurrence of and latency to cannibalism, and tested whether parents deposit their tadpoles in low-risk pools. I found that (1) neither pool capacity nor the presence of other larvae predict the presence/absence or number of tadpoles; (2) cannibalism occurs often, and how quickly it occurs depends on the difference in size between the tadpoles involved; and (3) the likelihood of males depositing their tadpoles in occupied pools increases with the size of the resident tadpole. I suggest that predation/cannibalism is not the only factor that parents assess when choosing deposition sites, and that the presence of larger conspecifics may instead provide information about pool quality and stability.     

Home range and movement patterns of the Redlip Parrotfish (Scarus rubroviolaceus) in Hawaii

The activity and movement of the Redlip Parrotfish, Scarus rubroviolaceus, near Wawaloli reef, Hawaii (19°43′06″N, 156°03′11″W), was investigated with active and passive tracking techniques from April through November 2007. Site fidelity was determined and home range sizes were measured for 21 individuals. All individuals showed strong diurnal activity patterns and relatively stable home ranges. Terminal phase individuals had larger home range areas than initial phase fish, and home range size increased with depth. Occasional long forays from the home range were observed in many individuals. Forays often occurred in crepuscular hours and may represent migrations to/from nocturnal sheltering sites. Findings from this study underscore the importance of depth and nighttime shelter migrations as design considerations for effective marine protected areas (MPA) for large parrotfishes.     

Tidal-linked behaviour of the shrimp Ligur uvea in a land-enclosed marine pool

At Aldabra Atoll, the shrimp Ligur uvea inhabits a land-locked marine pool which is connected to the sea by a subterranean fissure. The shrimps migrate daily with the tide from a hole in the subterranean labyrinth to a feeding area at the bottom of the pool. The timing of the migration is tidal-linked and independent of light and temperature. With each incoming tide, tagged individuals return to a definite feeding ground. Such homing has not been reported from other L. uvea pools. Feeding activity was quantitatively measured, differences between day and night activity are interpreted as adaptations against predators (terrestrial wading birds). Compared to other L. uvea pools, the shrimps at Aldabra Atoll are exposed to a more rigorous environmental situation, which probably influences the tidal-linked behaviour of the local shrimp population.     

Long-term trends in invertebrate–habitat relationships under protected and fished conditions

Few studies examine the long-term effects of changing predator size and abundance on the habitat associations of resident organisms despite that this knowledge is critical to understand the ecosystem effects of fishing. Marine reserves offer the opportunity to determine ecosystem-level effects of manipulated predator densities, while parallel monitoring of adjacent fished areas allows separating these effects from regional-scale change. Relationships between two measures of benthic habitat structure (reef architecture and topographic complexity) and key invertebrate species were followed over 17 years at fished and protected subtidal rocky reefs associated with two southern Australian marine reserves. Two commercially harvested species, the southern rock lobster (Jasus edwardsii) and blacklip abalone (Haliotis rubra) were initially weakly associated with habitat structure across all fished and protected sites. The strength of association with habitat for both species increased markedly at protected sites 2 years after marine reserve declaration, and then gradually weakened over subsequent years. The increasing size of rock lobster within reserves apparently reduced their dependency on reef shelters as refuges from predation. Rising predation by fish and rock lobster in the reserves corresponded with weakening invertebrate–habitat relationships for H. rubra and sea urchins (Heliocidaris erythrogramma). These results emphasise that animal–habitat relationships are not necessarily stable through time and highlight the value of marine reserves as reference sites. Our work shows that fishery closures to enhance populations of commercially important and keystone species should be in areas with a range of habitat features to accommodate shifting ecological requirements with ontogenesis.     

How hot for how long? The potential role of heat intensity and duration in moderating the beneficial effects of an ecosystem engineer on rocky shores

Studies of the importance of ecosystem engineers have focused on their benefit to biodiversity through ameliorating environmental stress, without understanding the exact benefits gained by associated organisms. On monsoonal tropical shores, species experience strong seasonality in environmental conditions from almost temperate winters to tropical summers when mass mortalities can occur during daytime emersion. The limpet, Cellana grata, associates with an ecosystem engineer, the barnacle Tetraclita japonica, in summer, but not in winter. To understand the benefits of this association, physiological responses (heart rates and osmotic responses) of the limpet either amongst Tetraclita or on open rock surfaces were investigated under three environmental conditions in a laboratory mesocosm: awash (non-stressed), low (30 °C) and high thermal stress (40 °C) of varying durations (3 or 6 h). In general, at 30 °C, limpets showed similar physiological responses under all conditions for both exposure durations. After 6 h at 40 °C, however, all limpets on open rock surfaces died, whereas those associated with barnacles survived. The surviving limpets experienced similar levels of stress as those exposed to 40 °C on open rock surfaces for half the time (3 h), showing that both the level and duration of stress were important. Limpets, therefore, gain benefits from engineering species when conditions are extreme or stress is prolonged. Under low temperatures or for short durations (e.g. winter), associating with barnacles does not provide physiological benefits. Understanding how, and to what extent, associates benefit from ecosystem engineers highlights how close the margin between survival and mortality can be when conditions are extreme.     

Annual,semilunar, and diel reproductive rhythms in the Hawaiian labrid Thalassoma duperrey

In September 1978, 5 034 juvenile western rock (spiny) lobsters (Panulirus cygnus George) were tagged on a shallow coastal reef near Cliff Head in Western Australia. Of these, 26% were recaptured and removed from the population during the subsequent commercial fishery between 15 November 1978 and 30 June 1979. A resurvey of the tagging site in September 1979 found that size structure of the population was similar to the tagged population in September 1978 but that the 1979 population was composed of almost completely different individuals and only 2% of the rock lobsters tagged in September 1978 were recaught at this time. A wide range of movement was indicated by the recapture positions of 354 rock lobsters caught off the tagging site. Most travelled almost directly offshore but some also travelled offshore in a northwesterly direction. Both trends were shown in the data from the recaptures of 18 rock lobster released farther offshore from Cliff Head in September 1981, and of 19 released at a close inshore site in earlier years. Maximum offshore rates of movement of up to 622 m d^-1 were recorded for distances up to 37 nautical miles (ca. 68 km). Growth rates for the tagged individuals which moved off, and those that were caught on, the tagging site were similar and close to growth rate under optimal conditions for this species. Under the present high levels of exploitation few, if any, of the rock lobsters remain to become permanent residents in the coastal reefs. It is believed that the north-westerly movements of some of the pre-adult rock lobsters, on their way to the breeding grounds on the outer edge of the continental shelf, are due to the upstream movement of these rock lobsters in a current from the north. The north-westerly movements allow the rock lobster population increased utilization of the food supplies of the coastal area.     

Does the effect of low temperature on osmoregulation by the prawn Palaemon elegans Rathke, 1837 explain winter migration offshore?

Osmoregulation by the European glass prawn Palaemon elegans Rathke, 1837 was impaired at low temperature (2°C) and high salinities (full strength sea water 35 PSU). Under these conditions, small individuals showed stronger osmoregulatory ability than larger individuals. This observation could partially explain differences in winter migrations off the UK coast where mainly large P. elegans go offshore, leaving smaller individuals in high shore rock pools, a finding consistent with winter experiments in situ.     

Delimiting soil chemistry thresholds for nickel hyperaccumulator plants in Sabah (Malaysia)

Nickel hyperaccumulator plants have been the focus of considerable research because of their unique ecophysiological characteristics that can be exploited in phytomining technology. Comparatively little research has focussed on the soil chemistry of tropical nickel hyperaccumulator plants to date. This study aimed to elucidate whether the soil chemistry associated with nickel hyperaccumulator plants has distinctive characteristics that could be indicative of specific edaphic requirements. The soil chemistry associated with 18 different nickel hyperaccumulator plant species occurring in Sabah (Malaysia) was compared with local ultramafic soils where nickel hyperaccumulator plants were absent. The results showed that nickel hyperaccumulators in the study area were restricted to circum-neutral soils with relatively high phytoavailable calcium, magnesium and nickel concentrations. There appeared to be a ‘threshold response’ for the presence of nickel hyperaccumulator plants at >20 μg g^?1 carboxylic-extractable nickel or >630 μg g^?1 total nickel, and >pH 6.3 thereby delimiting their edaphic range. Two (not mutually exclusive) hypotheses were proposed to explain nickel hyperaccumulation on these soils: (1) hyperaccumulators excrete large amounts of root exudates thereby increasing nickel phytoavailability through intense rhizosphere mineral weathering; and (2) hyperaccumulators have extremely high nickel uptake efficiency thereby severely depleting nickel and stimulating re-supply of Ni from diffusion from labile Ni pools. It was concluded that since there was an association with soils with highly labile nickel pools, the available evidence primarily supports hypothesis (2).     

Carbon emission associated with respiration and calcification of nine gastropod species from the intertidal rocky shore of Western Europe

Intertidal rocky shores are characterized by vertical zonation that results from the interplay between environmental conditions, organism physiology, and species interactions. Metabolism of intertidal organisms is highly variable between species and it changes with vertical position along the intertidal gradient. The present study aimed to quantify the carbon metabolism of nine intertidal rocky shore gastropods, in order to clarify their respective roles in carbon production during emersion and immersion. The influences of monthly temperature variation and tidal level were tested for each species. Analyses were performed in the laboratory using the infrared gas analyzer method for measuring aerial respiration rates, and the dissolved inorganic carbon and total alkalinity technique for measuring aquatic respiration rate and calcification. Hourly carbon fluxes were calculated for the mean annual temperature of 13 °C measured in both air and underwater in the study area. Respiration rates were similar for emersion (8–25 μmol CO₂ g AFDW^?1 h^?1) and immersion (10–23 μmol DIC g AFDW^?1 h^?1). For all species, underwater respiration fluxes were more influenced by monthly temperature variation than by air fluxes, probably as an adaptation to the rapid changes occurring during emersion. Calcification was an important factor influencing annual carbon fluxes for all studied species; every species showed different calcification rates according to its size and position on the intertidal zone. Annual carbon emissions were calculated using the mean immersion/emersion time of each species. Intertidal gastropod carbon emission was primarily influenced by body biomass and their vertical position within the intertidal zone.