Physiological performance of intertidal coralline algae during a simulated tidal cycle

Intertidal macroalgae endure light, desiccation, and temperature variation associated with sub‐merged and emerged conditions on a daily basis. Physiological stresses exist over the course of the entire tidal cycle, and physiological differences in response to these stresses likely contribute to spatial separation of species along the shore. For example, marine species that have a high stress tolerance can live higher on the shore and are able to recover when the tide returns, whereas species with a lower stress tolerance may be relegated to living lower on the shore or in tidepools, where low tide stresses are buffered. In this study, we monitored the physiological responses of the tidepool coralline Calliarthron tuberculosum (Postels and Ruprecht) E.Y. Dawson and the nontidepool coralline Corallina vancouveriensis Yendo during simulated tidal conditions to identify differences in physiology that might underlie differences in habitat. During high tide, Corallina was more photosynthetically active than Calliarthron as light levels increased. During low tide, Corallina continued to out‐perform Calliarthron when submerged in warming tidepools, but photosynthesis abruptly halted for both species when emerged in air. Surprisingly, pigment composition did not differ, suggesting that light harvesting does not account for this difference. Additionally, Corallina was more effective at resisting desiccation by retaining water in its branches. When the tide returned, only Corallina recovered from combined temperature and desiccation stresses associated with emergence. This study broadens our understanding of intertidal algal physiology and provides a new perspective on the physiological and morphological underpinnings of habitat partitioning.     

The relationship between position on shore and shell ornamentation in two size-dependent morphotypes of Littorina striata,with an estimate of evaporative water loss in these morphotypes and in Melarhaphe neritoides

Shells of small (< 7 mm) Littorina striata are frequently nodulose, but shells of larger individuals are striate. Nodulose L. striata dominated the littoral fringe of a black basalt Azorean shore where daytime rock temperatures rose significantly higher than nearby shores of different rock composition or colour. There was no evidence of intraspecific size-partitioning on the latter shores, where the numbers of striate and nodulose L. striata were approximately equal between high eulittoral (low-shore) and high littoral fringe (high-shore) localities. The prevalence of small L. striata in the littoral fringe is opposite to that usually characteristic of the Littorinidae, where the largest individuals usually occupy the higher positions on the shore. It is hypothesized that small L. striata attain a resting posture better able to minimize heat absorption from the substratum than attained by larger individuals. Smaller individuals also take advantage of both posture and a nodulose shell surface to re-radiate absorbed incident radiant thermal energy more effectively to the atmosphere by convection. Thus, small, nodulose L. striata are especially well adapted to occupy geologically young basaltic rocks commonly found fringing islands of the mid-Atlantic. The rate of evaporative water loss was determined for Melarhaphe neritoides and striate and nodulose L. striata for approximately 11 days emersion. All three groups are exceptionally capable of controlling evaporative water loss. Total percent evaporative water loss by nodulose L. striata (17.9%) was significantly greater than that lost by either striate L. striata (14.1%) or M. neritoides (13.5%) but, among 15 species for which evaporative water loss has been determined by similar methodology, M. neritoides and striate L. striata are the most capable of conserving body water during 11 days of emersion.     

Experiments on factors influencing settlement,survival, and growth of two species of barnacles in new south wales

Adult Tesseropora rosea (Krauss) and Tetraclitella purpurascens (Wood) are mostly found in the eulittoral (barnacle) zone of rocky seashores in New South Wales. Below this zone most space is occupied by the tube-worm Galeolaria caespitosa (Lamarck) or by various species of macroalgae. Within the eulittoral zone, T. rosea are mostly on sunny areas of rock exposed to relatively strong wave-action. T. purpurascens are present mainly in crevices, caves, and under ledges where there is considerable shade.Cyprids of both species settled on sandstone plates and on experimentally cleared areas in the barnacle and Galeolaria zones. Neither species settled where the substratum was already covered by algae or Galeolaria. No spat of T. purpurascens were found in sunny areas of the barnacle zone. T. rosea, however, settled in cleared substrata in sunny and shaded areas. Neither species settled in the littoral fringe above the upper limit of distribution of adults. On boulders transferred to high levels of the shore during a storm, small T. purpurascens died within a few weeks.Barnacles of both species which had settled in experimentally cleared areas in the Galeolaria zone survived and grew. In these areas some T. purpurascens were killed by being smothered by tube-worms which settled after the barnacles. This probably happens to T. rosea, but was not demonstrated experimentally. In the Galeolaria zone, both species of barnacles were very quickly smothered and killed by macroalgae growing over them, except where these were experimentally removed.Within the barnacle zone, all newly-settled spat of T. purpurascens transferred to sunny sites died within two months, whilst many of those in shaded sites survived. In areas where wave-action was strong, spat of T. rosea survived and grew well in sunny areas, but survived better in the shade. Under a ledge, however, where wave-action was reduced, all the T. rosea in sunny sites, and most of those in shaded sites died within two months; many newly-settled T. purpurascens survived in the shade in this area.The grazing limpet Cellana tramoserica (Sowerby) dislodged and crushed some newly-settled T. rosea and reduced survival in some sunny areas. T. rosea settled preferentially on bare rock and were rarely found on the shells of adult barnacles. Thus, the density of spat was greater where adult barnacles were absent. In contrast, many newly-settled T. purpurascens were found on the shells of adults of their own species in shaded areas; they also settled on cleared rock. Because T. purpurascens tended to settle amongst and on adults, and in crevices and confined areas, they were not much affected by limpets. When newly-settled T. purpurascens were in high densities, they had lower survival than in areas with reduced densities, because of squashing and smothering by each other.The upper and lower limits of vertical distribution (zonation) of these two species of barnacles are determined primarily by the settlement of cyprids. Neither species settled at the highest levels on the shore. Whether this was due to the decreasing time of submersion during high tide towards the top of the shore, or a result of preferences for settlement site is unknown. Even if cyprids were to settle in the littoral fringe, the spat would die very quickly probably as a result of desiccation. Below the barnacle zone, the entire substratum is usually occupied by other sessile species, particularly macroalgae, on which the barnacles do not settle. In experimentally cleared areas below the barnacle zone, or in any naturally cleared areas both species settled, and could survive the physical conditions. Newly-settled spat were, however, overgrown and killed by algae and Galeolaria.Within the barnacle zone, T. purpurascens is restricted to shaded areas because of the inability of newly-settled spat to survive the physical stresses of high temperature and desiccation in sunny habitats. T. rosea appears to be excluded from shaded areas by a combination of the lack of suitable substrata on which to settle, and the effects of reduced water-flow in many crevices and under ledges. T. rosea survived better in areas with strong wave-action and can survive in shaded areas where water-flow is not reduced by the topography of the substratum.     

Temporal comparison of the composition and zonation of rocky intertidal organisms at Cocos Island National Park, Pacific, Costa Rica

Several biological and physical factors change the rocky shore communities. The desiccation time and the tolerance of the intertidal species produce the vertical zonation. In many studies around the world, a temporal change in this zonation is presented.In Costa Rica, only studies that include temporal trends were carried out in Punta Mala and Montezuma, Pacific coast in 80's. The rocky intertidal of the Cocos Island National Park, Costa Rica were surveyed photographically. The Chatham bay was sampled in three expeditions (January 2007, October 2007 and April 2008). Photos corresponding to 25x25cm quadrats were taken with the goal to determine diversity and composition differences in rocky shore organisms between sampling dates. The Wafer bay was sampled in January and October 2007. The intertidal of Chatham consists of basaltic rock, while Wafer has basaltic and ignimbrite boulders. The main difference between sites were the higher algae cover (erect-frondose forms) and number of organism bands at Chatham bay. Temporal change was not found in the total cover of sessile fauna and autotrophs. The barnacle Tetraclita stalactifera, that occurs above the algal fringe (lower intertidal), was the invertebrate with the highest coverage. The mobile fauna biodiversity presented no significant trend between sampled months. However, the identity of species, their cover and their abundance showed a moderate temporal change. In October 2007, when the sea surface temperature was 23 degrees C the infralittoral zone had an increase in green algae cover. The red algae (crust and erect-frondose forms) were dominant in January and April. The pulmonate limpet, Siphonaria gigas and a bacterial biofilm at mid littoral showed a negative association. The snails of the high littoral and the supralittoral zone showed a temporal change in their abundance, but with contrasting patterns between sites. The temporal variation in the assemblages increased from the supralittoral to the infralittoral possibly due to changes in the water temperature and climatic conditions, that could influence the intertidal zone during the high and low tide, respectively.     

Influence of shell morphometry,microstructure, and thermal conductivity on thermoregulation in two tropical intertidal snails

Tropical intertidal organisms tolerate large fluctuations in temperature and high desiccation rates when exposed during low tide. In order to withstand the short‐term heat stress, intertidal organisms adopt behavioral responses to maximize their survival. Our previous research showed that tropical littorinids found at the upper and lower intertidal shores in Singapore exhibited different behavioral adaptations during low tide. Most of the upper‐shore Echinolittorina malaccana kept a flat orientation, with the aperture against the substrate and the long axis of the shell towards the sun, whereas a majority of the lower‐shore individuals of Echinolittorina vidua stood with the edge of the aperture perpendicular to the substrate on the rocky shore during low tide. This prompted analyses of the shells of these two species to determine whether the differences in the shell morphometry, microstructure, and thermal conductivity of shells of E. malaccana and E. vidua were associated with their respective behavioral responses to thermal stress. Analyses of shell morphometry and thermal conductivity showed that shells of E. malaccana were more likely to minimize heat gain, despite having a higher thermal conductivity on the outer surface, due to their light‐gray, elongated shell. By contrast, the dark‐colored, globose shells of E. vidua probably gain heat more readily through solar radiation. Scanning electron microscopy images of the shells of both littorinid species further revealed that they have cross‐lamellar structure; however, only individuals of E. vidua showed the presence of disjointed rod layers and a pigmented inner shell surface. Individuals of E. malaccana had a rough outer shell surface with holes that inter‐connect to form water‐trapping channels that probably aid cooling. Individuals of E. vidua, however, had a smooth outer surface with rows of kidney‐shaped depressions as microsculptures which probably help to stabilize shell shape. In both Echinolittorina species, behavioral responses were used to overcome thermal stress during low tide that was associated with shell morphometry and shell thermal conductivity. Such combined adaptations increase survivability of the littorinids at their respective tidal levels.     

A field investigation into the effects of interspecific competition on the distribution and cover of Ulothrix flacca

Ulothrix flacca is a filamentous green alga found in its greatest abundance in the littoral fringe of the rocky intertidal during the winter months. It is much less abundant below this level. This experiment was undertaken tot determine if the lower abundance below the littoral fringe is due to the effects of interspecific competition or physiological limitations. Clearing experiments were performed on the Coast of Maine, USA to eliminate competition. Belts 10 cm wide that ran from the littoral fringe downward into the eulittoral zone were cleared of all attached organisms down to bare rock. A second condition was set up in which only the brown algal canopy was removed from a 60 cm wide belt. Unaltered belts were designated as controls. Squares 0.25 m² in area were cleared in the littoral fringe and eulittoral zone. U. flacca was able to colonize the cleared areas of the eulittoral zone and reach abundances equal to those found in the unaltered control areas of the littoral fringe. This indicates that physiological limitations do not restrict the abundance of U. flacca in the eulittoral zone. However, evidence of grazing in cleared areas adjacent to high concentrations of the barnacle Semibalanus balanoides suggest that competition alone is not responsible for the limited abundance of U. flacca in the eulittoral zone. Herbivores associated with the presence of barnacles may also have an impact on the abundance of U. flacca, even during the winter months.     

Activity rhythms and cyclical changes of microhabitat preferences in the intertidal pipefish Nerophis lumbriciformis (Pisces: Syngnathidae)

As an inhabitant of the intertidal zone, Nerophis lumbriciformis predictably should show a clear rhythmic activity, with its maximum closely correlated with the high tide period, as observed in several other rocky intertidal fish species. We investigated the rhythmic activity patterns of N. lumbriciformis and analysed to what extent specific substratum preferences may be linked to water level and, ultimately, to the particular activity rhythm patterns of the fish. To determine N. lumbriciformis substratum preferences, two different experiments were conducted. The first evaluated substratum preferences from among a group of the four most common substrates found in the sample area. The second experiment evaluated the fish's maintenance of that substratum preference, after alteration of the water level in a simulated ebb tide. From the available substrata, N. lumbriciformis displayed a clear preference for algae until the water level began to decrease. Then, the preference radically shifted from algae to boulders. N. lumbriciformis showed a clear rhythmic activity, affected by both tidal and circadian periodicity, producing asymmetrical activity peaks. Unlike the rhythms of other rocky intertidal fish species, the maximum activity peaks were not centred at the expected high tide period. The adaptive explanation for the apparently peculiar activity peaks appears to be related to the particular substratum preferences of N. lumbriciformis. The detected circatidal period seems, consequently, closely related to small migrations between substrata, whilst the observed increase in diurnal activity, independent of the tidal cycle, may be related to predatory activity, since N. lumbriciformis is a visual feeder. Electronic Publication     

Biotic zonation on rocky shores of Heard Island

Summary The zonation of rocky shore biota on Heard Island is described for the first time, related to a universal zonation scheme and compared with that recorded for other sub-Antarctic localities. The Kelp Zone of holdfasts of the giant kelp, Durvillea antarctica (Chamisso) Hariot (one of the characteristic features of these regions) is confirmed as a sublittoral fringe. The occurrence of a Bare Zone within the littoral zone on sub-Antarctic shores is discussed. The possible roles of submersion, spray, freezing and predation by gulls in influencing the extent and composition of zones are discussed.     

Tide-model analysis of the zonation of intertidal prosobranchs. I. Four species of littorina (L.)

Experiments were carried out in a tide model to determine the patterns of zonation of four species of periwinkles under laboratory conditions. Both upper shore littorinids, Littorina neritoides (L.) and L. saxatilis (Olivi), showed the same zonation in relation to the tidal cycle as observed on the shore at Plymouth. In the model, the animals adopted a zone above the level of high water and then remained inactive. L. littoralis (L.) showed the same distribution unless fucoid algae were provided, in which case the periwinkles clustered on them. L. littorea (L.) showed the ability to adopt a pattern of zonation in the model, which was at, and just below, high tide level, but higher than that normally observed on the shore.     

Transitional ecological requirements for early juveniles of two sympatric stichaeid fishes,Cebidichthys violaceus andXiphister mucosus

Synopsis The distribution patterns, diets, and substratum (refuge) requirements of early juveniles of two sympatric stichaeid fishesCebidichthys violaceus andXiphister mucosus, were investigated in a rocky intertidal habitat at Diablo Canyon, California. Monthly investigations were conducted at low tide for four consecutive months, to assess ontogenetic differences in distribution, diet, and refuge requirements within and between the two species. Distinct differences in vertical zonation were exhibited by both stichaeids throughout the study. Interspecific zonation patterns were similar to those recorded for adults of both species. Diet analyses showed that early juveniles of both stichaeids were zooplanktivorous, differing markedly from the primarily herbivorous diets of adults. Changes in diet were largely due to the selection of larger prey taxa as both fishes, and their mouth size, grew over the study period. Predation by both fishes on water-column planktors (calanoid copepods, zoea and polychaete larvae) was greatest following initial intertidal settlement and habitat establishment by early juvenile fishes. Greater dependence on substrate-oriented and/or benthic prey (harpacticoid copepods, gammarid amphipods and mysid shrimp) was exhibited by both fishes as they grew in size. Affinities for sand, gravel, and pebbles during monthly field surveys were similar for both species throughout the study. As the fishes grew, their substratum preferences changed in relation to the substrata which provided the best refuge. Results from laboratory experiments indicated that young stichaeids select very specific substrata based on fish age (size) and substratum suitability (i.e. adequate refuge). Similarities in diet and substratum preferences, and changes in those preferences over time, appear to be the result of morphological similarities (body size and shape and mouth gape) for both species at a given age.     

Heat production inLittorina saxatilis Olivi andLittorina neritoides L. (gastropoda: Prosobranchia) during an experimental exposure to air

The adaptation of littorinid molluscs to prolonged aerial exposure was investigated by the determination of heat production.Littorina saxatilis, inhabiting the upper eulittoral, reached a maximum metabolic activity during submersion (heat production: 3.26×10⁻³J s⁻¹ (g_adw)⁻¹. On the first three days of desiccation, the heat production was continuously reduced to 40% of the submersed value. A prolonged aerial exposure was lethal for this species. In the supralittoralL. neritoides, three stages of energy metabolism could be observed: An intermediate heat production during submersion (1.97×10⁻³Js⁻¹ (g_adw)⁻¹), an increased metabolism during the first hour of aerial exposure (heat production 204% of submersed value), and a minimal metabolism (39% of the submersed value and 19% of maximum value) during the following days and weeks of desiccation. Recovery depended on water salinity;L. saxatilis proved to be less euryhaline thanL. neritoides. Thus, the metabolic adaptations correlate with the level of littoral habitat; inactivity combined with a drastically reduced energy consumption is a metabolically economic way to survive in periodically dry environments.     

Causes and consequences of thermal tolerance limits in rocky intertidal porcelain crabs, genus petrolisthes

Vertical zonation of intertidal organisms, from the shallowsubtidal to the supralittoral zones, is a ubiquitous featureof temperate and tropical rocky shores. Organisms that livehigher on the shore experience larger daily and seasonal fluctuationsin microhabitat conditions, due to their greater exposure toterrestrial conditions during emersion. Comparative analysesof the adaptive linkage between physiological tolerance limitsand vertical distribution are the most powerful when the studyspecies are closely related and occur in discrete vertical zonesthroughout the intertidal range. Here, I summarize work on thephysiological tolerance limits of rocky intertidal zone porcelaincrab species of the genus Petrolisthes to emersion-related heatstress. In the eastern Pacific, Petrolisthes species live throughouttemperate and tropical regions, and are found in discrete verticalintertidal zones in each region. Whole organism thermal tolerancelimits of Petrolisthes species, and thermal limits of heartand nerve function reflect microhabitat conditions. Speciesliving higher in the intertidal zone are more eurythermal thanlow-intertidal congeners, tropical species have the highestthermal limits, and the differences in thermal tolerance betweenlow- and high-intertidal species is greatest for temperate crabs.Acclimation of thermal limits of high-intertidal species isrestricted as compared to low-intertidal species. Thus, becausethermal limits of high-intertidal species are near current habitattemperature maxima, global warming could most strongly impactintertidal species.     

Intertidal distribution and species composition of brachyuran crabs at two rocky shores in Central Portugal

The objectives of the present study are to describe and compare the brachyuran community of rocky shores within the Central Portuguese coast and to examine the zonation patterns of the most representative species. For this, randomly placed transects were surveyed to obtain crab counts according to microhabitat and intertidal level. Repeated sampling in two different shores during two different seasons provided spatial and temporal replication for zonation analyses. Seven species were registered: Pachygrapsus marmoratus, Eriphia verrucosa, Xantho incisus, Carcinus maenas, Necora puber, Pirimela denticulata and Pilumnus hirtellus. Species density rankings are the same at both localities, but the less exposed shore presents higher diversity. While most species are mainly confined to specific microhabitats in the lower level, P. marmoratus and E. verrucosa can exploit the whole intertidal range. Regardless of shore and season, E. verrucosa is more abundant in the lower intertidal levels, while no such zonation patterns were recorded for P. marmoratus. Initial predictions concerning the effect of wave exposure and temperature on the zonation of those species are not validated after analysing the factorial model proposed. Between-shore contrasts were found instead, with higher densities recorded in the more exposed locality for both species. Possible causes of the observed patterns are discussed.     

Barnacle larval supply to sheltered rocky shores: a limiting factor?

In northwest Europe, sheltered rocky shores are dominated by fucoid canopy algae and barnacles are rare, although the latter are extremely abundant on exposed shores. The supply of the intertidal barnacle Semibalanus balanoides (L.) to sheltered, fucoid dominated rocky shores was investigated to determine the importance of larval supply in limiting the abundance of adults in shelter. Larval supply was measured at two spatial scales, at the scale of shore (100s of metres), by comparing larval concentrations at exposed and sheltered sites, and at a smaller spatial scale (m), by examining the role of fucoid canopies in limiting supply to the substratum. Replicate plankton trawls were carried out above the intertidal zone at high water at two sheltered sites and nearby exposed headlands. The concentration of S. balanoides cyprid larvae was significantly higher at the sheltered sites on two out of three sampling occasions with up to 14 times greater larvae on one occasion than the nearby exposed site. The effect of the macroalgal canopy on supply to the substratum was assessed in two ways: directly, by pumping water from the substratum in areas with and without Ascophyllum nodosum (L.) Le Jolis, and indirectly by measuring cyprid settlement in a canopy-manipulation experiment. Pumped plankton samples from mid tide level showed that the A. nodosum canopy did not form a barrier to larval supply and may have had a positive effect on larval concentrations at the substratum. Cyprid settlement was assessed in the mid shore A. nodosum and low shore Fucus serratus L. zones to areas with canopy algae (but protected from the sweeping effects of macroalgal fronds) and without canopy. Settlement over three consecutive 24-h periods showed a consistent pattern; settlement was consistently lower beneath the F. serratus canopy than in cleared areas, suggesting that this algal species forms a barrier, limiting supply of cyprid larvae to the substratum.     

Maintenance of zonation patterns in two species of flat periwinkle,Littorina obtusata and L. mariae

The zonation patterns of Littorina obtusata (L.) and Littorina mariae Sacchi et Rastelli were shown to be quite distinct on a sheltered rocky shore. L. obtusata was found at all the heights sampled; it reached peak numbers at mid shore on the alga Ascophyllum nodosum L. (Le Jol). There was no difference in the tidal height occupied by adults or juveniles; or in the mean size of L. obtusata along the vertical gradient of the shore. In contrast L. mariae occurred exclusively low on the shore, on Fucus serratus L. Translocation of the two species within their respective levels resulted in random movement after 4 days, although initial movements after 1 and 2 days were sometimes directional. Animals transplanted to the normal level of the other species showed directional movement towards their home zone; this was most pronounced after 4 days. There was no difference in the distance moved by the two species, although the distance moved did vary with tidal height, both species moving further at mid shore than low shore. Distances moved by littorinids at replicate areas in the low shore were similar but those at mid shore did vary. There was an interaction between the species and the different tidal heights which revealed that transplanted species moved further than translocated species at the same tidal level. However, this was only significant in the case of L. mariae. It is suggested that the close relationship between the winkles and their host algae may direct the homing behaviour of displaced individuals.     

Analysis of patterns of dispersion of intertidal prosobranch gastropods in relation to macroalgae and rock-pools

Summary The patterns of dispersion of Nerita atramentosa, Austrocochlea constricta, Bembicium nanum and Cellana tramoserica are analysed from quadrat samples taken between June 1972 and June 1973. In each quadrat, the proportion of animals in each of two substrata, the encrusting alga Peyssonelia gunniana and small pools, was recorded, with the proportional area of cover of each substratum, height on the shore, density of the species and the time of sampling. Multiple regression and covariance analysis showed that, in all cases, there was a linear relationship between proportion in a substratum and proportional area of substratum. No other variables were significant.Juvenile Nerita and Austrocochlea showed preferences for both substrata. Adult Nerita and all Bembicium were dispersed at random with respect to both substrata. Adult Austrocochlea were randomly dispersed with respect to Peyssonelia, but showed a preference for water. Cellana showed a preference for water and against Peyssonelia. These microhabitat preferences are discussed in relation to previously described effects of substrata on the densities of these four species.     

Undaria pinnatifida (Laminariales,Phaeophyta) 12 years after its introduction into the Atlantic Ocean

Since its first introduction into the Atlantic for farming purposes in 1983, the brown alga Undaria pinnatifida has been found at several sites along the European coasts. The present study deals with an ecological experiment that was performed at the island of Ushant (Brittany, France) in order to check the ability of Undaria to recruit to a rocky substratum in situ. On the one hand, the authors were unsuccessful in getting Undaria to colonize a previously denuded area of shore, which was subsequently colonized by other native species, a result that may speak for a competition between Undaria and the local kelps. On the other hand, evidence is presented that Undaria pinnatifida has become a member of the Atlantic flora for good. The sporophytes can be found growing from low water mark of neap tide (+1.5 m) down to at least –15 m below lower water mark of spring tide. Its propagation is thought to occur in at least two ways: either step by step from cultivation sites along the coasts, or from the hulls of ships sailing from harbour to harbour. The preference of Undaria for settlement is on artificial structures.     

Lack of metabolic temperature compensation in the intertidal gastropods,Littorina saxatilis (Olivi) and L. obtusata (L.)

Two intertidal snails, Littorina saxatilis (Olivi, 1972) (upper eulittoral fringe/maritime zone) and Littorina obtusata (Linnaeus, 1758) (lower eulittoral) were collected from a boulder shore on Nobska Point, Cape Cod, Massachusetts, in July and acclimated for 15–20 days at 4 ° or 21 °C. Oxygen consumption rate (Vo₂) was determined for 11–15 subsamples of individuals at 4 °, 11 ° and 21 °C with silver/platinum oxygen electrodes. Multiple factor analysis of variance (MFANOVA) of lo₁₀ transformed values of whole animal Vo₂ with log₁₀ dry tissue weight (DTW) as a covariant revealed that increased test temperature induced a significant increase in Vo₂ in both species (P<0.00001). In contrast, MFANOVA revealed that temperature acclimation did not affect Vo₂ in either L. saxatilis (P= 0.35) or L. obtusata (P= 0.095). Thus, neither species displayed a capacity for the typical metabolic temperature compensation marked by an increase in Vo₂ at any one test temperature in individuals acclimated to a lower temperature that is characteristic of most ectothermic animals. Lack of capacity for metabolic temperature acclimation has also been reported in other littorinid snail species, and may be characteristic of the group as a whole. Lack of capacity for respiratory temperature acclimation in these two species and other littorinids may reflect the extensive semi-diurnal temperature variation that they are exposed to in their eulittoral and eulittoral fringe/maritime zone habitats. In these habitats, any metabolic benefits derived from longer-term temperature compensation of metabolic rates are negated by extreme daily temperature fluctuations. Instead, littorinid species appear to have evolved mechanisms for immediate metabolic regulation which, in L. saxatilis and L. obtusata and other littorinids, appear to centre on a unique ability for near instantaneous suppression of metabolic rate and entrance into short-term metabolic diapause at temperatures above 20–35 °C, making typical seasonal respiratory compensation mechanisms characteristic of most ectotherms of little adaptive value to littorinid species.     

Patterns of organizations of intertidal and shallow subtidal vegetation in wave exposed habitats of central Chile

Wave-exposed rocky intertidal habitats of central Chile exhibit zonation of algal morphologies rather than strict patterns of species zonation. In low shore areas, there is a vertical sequence of perennial belts of calcareous crusts, kelp-like forms and expanded cushions or non-calcareous crusts. The calcareous crusts are represented by species of Mesophyllum, the kelp-like forms include Lessonia nigrescens and Durvillaea antarctica, while the cushions are represented by Gelidium chilense and G. lingulatum and the noncalcareous, expanded crusts by Codium dimorphum. Thin and thick blades, represented by Iridaea laminarioides, Ulva rigida and Porphyra columbia and filamentous forms including Ceramium rubrum, Centroceras clavulatum and Polysiphonia spp. are more patchy than the lower, perennial belts. They may, however, form distinct temporal monocultures at upper intertidal levels. Upper and lower limits of the various zones are set by interactions of several factors, the relative importance of which can change seasonally. When some of the factors restricting species distribution are experimentally removed, other interactions among factors become limiting.Within each zone, species are morphologically similar, with the abundance of species being regulated by symmetric competitive interactions. Competition is often asymmetric at the boundaries of zones except when adults of small-sized forms interact with morphologically similar juveniles of larger forms. Irrespective of their extremely different morphologies, the permanent, zone-forming algal species generally combine escape from grazers or defensive adaptations with clear competitive abilities. Nevertheless, there is a clear competitive hierarchy which is expressed in vertical displacements and zonation. The lowershore habitats could potentially be occupied by any of the different types of algae. Fast growth and large size allow the kelps to occupy this zone pushing the calcareous crust dominated-zone down into shallow subtidal areas and displacing the cushions and fleshy crusts into the low and middle intertidal regions. In turn, these last forms can displace thick and thin foliose forms and filaments to upper levels on the shore. Displaced forms may exist as patches at various levels of the shore.