Physiological tolerances across latitudes: thermal sensitivity of larval marine snails (Nucella spp.)

A critical step in understanding how temperature will affect biodiversity in coastal ecosystems is to gain insight into how the tolerances, and ultimately survival, of early life history stages will influence the distribution and abundance of adults. We assessed the thermal tolerance of encapsulated veliger-stage larvae of a common dogwhelk, Nucella ostrina, that occur in the rocky intertidal zone on the west coast of North America. Results showed that veligers collected from northern latitudes in Washington State were less tolerant of heat stress than those from central sites in California. For all sites, we found there to be a subtle difference between the temperatures at which veligers first began to die compared to when veligers reached 100% mortality. On a biogeographic scale, the LT₅₀ temperatures, a measure of larval sensitivity, for N. ostrina veligers displayed a strong latitudinal trend. These findings provide a conservative measurement of the upper thermal limits of encapsulated veligers while illustrating how these early life history stages could be physiologically compromised under future climate warming scenarios.     

Genetic evidence for the cryptic species pair, <Emphasis Type="Italic">Lottia digitalis </Emphasis>and <Emphasis Type="Italic">Lottia austrodigitalis</Emphasis> and microhabitat partitioning in sympatry

It has been proposed that the common West Coast limpet, Lottia digitalis, is actually the northern counterpart of a cryptic species duo including, Lottia austrodigitalis. Allele frequency differences between southern and northern populations at two polymorphic enzyme loci provided the basis for this claim. Due to lack of further evidence, L. austrodigitalis is still largely unrecognized in the literature. Seven additional enzyme loci were examined from populations in proposed zones of allopatry and sympatry to determine the existence of L. austrodigitalis as a sibling species to L. digitalis. Significant allele frequency differences were found at five enzyme loci between populations in Laguna Beach, southern California, and Bodega Bay, northern California; strongly supporting the existence of separate species. Both species exhibit two microhabitat morphotypes, a gooseneck barnacle morph in the mid-intertidal zone and a rock morph in the high-intertidal zone. In sympatry, L. austrodigitalis was more abundant higher in the intertidal on rocks, whereas L. digitalis was more abundant lower in the intertidal on barnacles. This finding supports earlier claims of microhabitat partitioning in this sibling species pair. In addition to this finding, the transition zone between the species was found to have shifted substantially northward in only two decades, from Monterey Peninsula, CA to near Pigeon Point, CA, where L. digitalis previously dominated.     

The influence of temperature and salinity on<Emphasis Type="Italic"> Acartia</Emphasis> (Copepoda: Calanoida) nauplii survival

Adult Acartia congeners, A. bifilosa, A. clausi, A. discaudata and A. tonsa, have distinct seasonal and spatial distribution patterns in Southampton Water (UK), reflecting patterns of temperature and salinity, respectively. The effect of these factors on other life stages, hatch success and naupliar survival was investigated by exposing the congeners to a range of salinity (15.5–33.3) and of temperature (5–20°C). A. clausi is known to prefer more saline waters, and showed highest hatch success at 33.3 salinity. A. tonsa is most tolerant to dilution, and at 15.5 salinity it had the highest hatch success of all the congeners. Hatch success in both A. bifilosa and A. discaudata was similar over the range of salinities investigated, confirming that they are intermediate species in terms of spatial distribution. The nauplii of all species survived well at the higher salinities and best at 33.3, which allows for differential transport of the poorly swimming nauplii to the mouth of the estuary until size and swimming ability increase, after which they can then return to regions of preferred salinity. The summer species, A. clausi and A. tonsa showed higher hatch success at 20°C, whereas A. discaudata, which is present in the water column all year round, showed no significant temperature-related differences in hatch success. A. bifilosa, which diapauses over summer, showed significantly higher hatch success at 10°C than at 20°C. The physiological relationship between temperature and development time was clear; naupliar survival of all species was highest at 20°C and all congeners reached the first copepodite stage (CI) significantly faster at 20°C. However, no consistent pattern was seen for salinity. It would appear that the adult Acartidae in Southampton Water remain in regions of their preferred salinity and lay eggs there which hatch well. However, because the nauplii are not good swimmers, they are swept towards the mouth of the estuary and into areas of higher salinity, where they remain and develop into more advanced stages before moving back up the estuary to take up their adult distribution pattern.Communicated by J.P. Thorpe, Port Erin     

Geographical patterns in reproductive biology of the Pan-American sandy beach isopod<Emphasis Type="Italic"> Excirolana braziliensis</Emphasis>

Eleven populations of the Pan-American sandy beach isopod Excirolana braziliensis, distributed from tropical (9°N) to temperate (39°S) sandy beaches in Atlantic and Pacific Oceans, were analyzed to determine latitudinal variations in: breeding and recruitment patterns, sex ratios, size of ovigerous females and of juveniles and length–fecundity relationships. E. braziliensis exhibited strong latitudinal patterns in all reproductive traits throughout its distribution range. Breeding and recruitment shifted from continuous to seasonal from tropical to temperate beaches, having a predominance of females at higher latitudes. In agreement with the latitudinal gradient hypothesis, ovigerous females inhabiting tropical sandy beaches (low latitudes) were smaller, became sexually mature at smaller sizes and had lower individual fecundity than on temperate beaches. Juveniles were also smallest at low latitudes. Between-ocean comparisons showed very similar reproductive characteristics for roughly the same latitude. These linked reproductive parameters suggest that the intensity of breeding effort is associated with the duration of the breeding season and geographically size-related characteristics of the species. Geographic variations in the breeding and recruitment seasons, as well as in individual fecundity, size structure of mature females and sex ratios, are proposed to have major consequences in explaining local variations in population demography. Our paper also reinforces the notion that sandy beach animals are highly plastic.Communicated by O. Kinne, Oldendorf/Luhe     

Phosphate uptake by intertidal algae in relation to zonation and season

The removal of phosphate from ambient seawater by whole plants of five species of fucoid algae, collected from the east coast of N. Ireland in 1988 and 1989, was followed over 6-h periods. A transient uptake pattern was observed forPelvetia canaliculata (L.) Dcne. et Thuret,Fucus spiralis L.,F. vesiculosus L. andF. serratus L., consisting of an initial period of high uptake, followed by a phase of zero uptake and then a period at an intermediate rate.Ascophyllum nodosum (L.) Le Jolis had a constant slow rate of uptake over 6 h. The initial uptake rate ofF. spiralis was significantly greater than that of any other species. Phosphate uptake over a 2-h period was measured at concentrations ranging from that of ambient seawater to 25µg-at. l^–1 for whole plants ofF. spiralis andF. serratus, using a large scale batch method. A small scale batch method was used for whole plants ofP. canaliculata and sections of the other four species investigated. Uptake abilities of the algae at low concentrations of phosphate were compared using the parameterV ₁ (the uptake rate at 1µg-at. l^–1) and at high concentrations usingV _max, the maximum uptake rate. These kinetic parameters of uptake were calculated using a method that avoids bias and permits statistical evaluation of the results. The fucoid algae studied could be divided into two distinct groups on the basis of their abilities to take up phosphate from seawater.P. canaliculata andA. nodosum had low values ofV ₁ in winter, which were also correlated with their positions on the shore and did not vary between winter and summer. TheFucus species had higher values ofV ₁ in winter, which were also correlated with their positions on the shore. In summer, however,V ₁-values for these species decreased and no longer correlated with their shore heights. TheV _max-value forF. spiralis was higher in winter than in summer but was signifcantly greater than that of any other species at all times of year. The ecological significance ofV _max is discussed in relation to nutrient limitation and the possible occurrence of patches of high nutrient concentration in the intertidal environment.     

Sex-specific temperature dependence of foraging and growth of intertidal snails

Predicting the biological impacts of climate change requires an understanding of how temperature alters organismal physiology and behavior. Given differences in reproductive physiology between sexes, increases in global temperature may be experienced differently by the males and females of a species. This study tested for sex-specific effects of increased air temperature on foraging, growth, and survival of an intertidal snail, Nucella ostrina (San Juan Island, Washington, 48–30′44″N, 123–08′43″W). Snails exhibited periodic peaks in foraging. Subjecting snails to elevated low tide air temperatures did not alter the timing or magnitude of this pattern. Despite similar temporal patterns in foraging, females foraged more than males, even when the risk of thermal stress was high. While males and females appear to have a similar body temperature threshold for optimal growth, females were more likely to cross that threshold resulting in a loss of body mass when exposed to daily increases in air temperature. These results suggest that the consequences of a warming climate in the short term may be different for males and females of N. ostrina, but also imply longer-term costs of reduced reproductive output, abundance, and distribution of this ubiquitous intertidal predator. Generally, this study points to the possible significance of sex-specific responses in an increasingly warm world.     

Differences in resource utilization and behaviour between coexisting <Emphasis Type="Italic">Jassa</Emphasis> species (Crustacea,Amphipoda)

Species of the genus Jassa are an important element of marine fouling communities, several species often co-occurring on a wide range of hard substrates. At Helgoland (North Sea, German Bight), the cosmopolitan J. marmorata occurs associated with the NW European species J. falcata and J. herdmani. Field and laboratory experiments revealed some ecological differences among species which may facilitate their coexistence. Test panels in the field were instantly colonized by the complete spectrum of resident Jassa species, but J. marmorata proved clearly superior to its congeners in rapidly colonizing new habitats. Further differences among species relate to microhabitat selection (differential use of peripheral and central parts of algal thalli) and the behavioural response to mechanical disturbance.     

Latitudinal symbiont zonation in <Emphasis Type="Italic">Stylophora pistillata</Emphasis> from southeast Africa

Studies on latitudinal gradients in Symbiodinium diversity on scleractinian corals are largely restricted to warm-water low latitude locations, and it appears that there is a shift in symbiont distributions with increasing latitude. The Symbiodinium assemblages of high latitude coral communities have largely been undocumented despite occupying an important transitional zone between tropical and temperate regions. Using a combination of the internal transcribed spacer region 2 (ITS2) and denaturing gradient gel electrophoresis (DGGE), we assessed the cladal and subcladal variability of Symbiodinium in the widely distributed species Stylophora pistillata along a latitudinal transect in southeast African waters which extended into high latitude locations. All colonies examined belonged to clade C. Six unique ITS2-DGGE banding profiles (designated C_spa to C_spf) were observed, which showed a latitudinal distribution from north to south, most likely a result of a gradient in water temperature and irradiance driven by riverine input in the southern regions. Sequence analysis revealed that all sequences except one did not match previously identified clade C sub-types, probably due to the lack of regional information in the Western Indian Ocean when compared to the Caribbean and Pacific. This study further supports the applicability of ITS2-DGGE in studies on Symbiodinium diversity, and highlights that potentially ecologically informative biogeographic patterns may be overlooked when only cladal designations are employed.     

Three beetles—three concepts. Different defensive strategies of congeneric myrmecophilous beetles

Myrmecophiles, i.e., organisms associated with ants live in a variety of ecological niches in the vicinity or inside ant colonies and employ different strategies to survive ant encounters. Because different niches are characterized by different encounter rates with host ants, strategies used to avoid ant aggressions should depend on these niches. This hypothesis was studied with three rove beetle species of the genus Pella, which are myrmecophiles of the ant Lasius fuliginosus and the non-myrmecophilous relative Drusilla canaliculata. Behavioral tests in the field revealed that Pella species are better adapted to interactions with ants than D. canaliculata, but that they use species-specific strategies in ant interactions. Pella cognata and Pella funesta avoid encounters with ants by swift movements. Chemical analyses of the defensive tergal gland secretions showed that P. cognata has replaced the aggression inducing undecane by the behaviorally neutral tridecane. P. funesta repels the ants by releasing the panic alarm pheromone sulcatone from its tergal gland resulting in an “ant free space” around the beetles. Finally, Pella laticollis uses a specific and unique appeasing behavior. Behavioral and chemical data did not reveal any indication for the mimicry of the ants' cuticular hydrocarbon profiles by any of the beetle species. It is discussed that the employed strategies correlate with the ecological niches of the beetles. P. cognata and P. funesta are living along ant trails with ample space to escape and the employed strategies are probably sufficient to escape from dangerous conflicts. In contrast, P. laticollis lives in refuse areas of ant nests with frequent ant encounters, and its appeasement strategy allows it to stay at the encounter site.     

Isotopic evidence of distinct feeding ecologies and movement patterns in two migratory predators (yellowfin tuna and swordfish) of the western Indian Ocean

Ecologists primarily use δ¹⁵N values to estimate the trophic level of organisms, while δ¹³C, and even recently δ¹⁵N, are utilized to delineate feeding habitats. However, many factors can influence the stable isotopic composition of consumers, e.g. age, starvation or isotopic signature of primary producers. Such sources of variability make the interpretation of stable isotope data rather complex. To examine these potential sources of variability, muscle tissues of yellowfin tuna (Thunnus albacares) and swordfish (Xiphias gladius) of various body lengths were sampled between 2001 and 2004 in the western Indian Ocean during different seasons and along a latitudinal gradient (23°S to 5°N). Body length and latitude effects on δ¹⁵N and δ¹³C were investigated using linear models. Both latitude and body length significantly affect the stable isotope values of the studied species but variations were much more pronounced for δ¹⁵N. We explain the latitudinal effect by differences in nitrogen dynamics existing at the base of the food web and propagating along the food chain up to top predators. This spatial pattern suggests that yellowfin and swordfish populations exhibit a relatively unexpected resident behaviour at the temporal scale of their muscle tissue turnover. The body length effect is significant for both species but this effect is more pronounced in swordfish as a consequence of their different feeding strategies, reflecting specific physiological abilities. Swordfish adults are able to reach very deep water and have access to a larger size range of prey than yellowfin tuna. In contrast, yellowfin juveniles and adults spend most of their time in the surface waters and large yellowfin tuna continue to prey on small organisms. Consequently, nitrogen isotopic signatures of swordfish tissues are higher than those of yellowfin tuna and provide evidence for different trophic levels between these species. Thus, in contrast to δ¹³C, δ¹⁵N analyses of tropical Indian Ocean marine predators allow the investigation of complex vertical and spatial segregation, both within and between species, even in the case of highly opportunistic feeding behaviours. The linear models developed in this study allow us to make predictions of δ¹⁵N values and to correct for any body length or latitude differences in future food web studies.     

Longitudinal and vertical trends in stable isotope signatures (δ13C and δ15N) of omnivorous and carnivorous copepods across the South Atlantic Ocean

Stable isotope (SI) ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) were measured in omnivorous and carnivorous deep-sea copepods of the families Euchaetidae and Aetideidae across the Atlantic sector of the Southern Ocean to establish their trophic positions. Due to high and variable C/N ratios related to differences in lipid content, δ¹³C was corrected using a lipid-normalisation model. δ¹⁵N signals ranged from 3.0–6.9‰ in mesopelagic species to 7.0–9.5‰ in bathypelagic congeners. Among the carnivorous Paraeuchaeta species, the epi- to mesopelagic species Paraeuchaeta antarctica had lower δ¹⁵N values than the mesopelagic P. rasa and bathypelagic P. barbata. The same trend was observed among omnivorous Aetideidae, but was not significant. In the most abundant species P. antarctica, individuals from the western Atlantic had higher δ¹³C and δ¹⁵N values than specimens at the eastern stations. These longitudinal changes in δ¹³C and δ¹⁵N values were attributed to regional differences in hydrography and sea surface temperature (SST), in particular related to a northward extension of the Antarctic Polar Front (APF) at the easternmost stations. The results indicate that even in a mesopelagic carnivorous species, the changes in surface stable isotope signatures are pronounced.     

Variations in cardiac performance and heat shock protein expression to thermal stress in two differently zoned limpets on a tropical rocky shore

Understanding variation in physiological adaptations to thermal stress is vital when investigating intertidal species?? distribution patterns. The thermal sensitivities of two limpets, Cellana grata and C. toreuma, differed in accordance with their vertical distributions. Cardiac performance was maintained at higher temperatures (~47°C) for the high-zone C. grata than the mid-zone C. toreuma (~42°C). At 40°C, C. grata maintained regular heart function for ~4?h, while heart function of C. toreuma decreased rapidly. Heat shock protein expression revealed that C. toreuma had two constitutive isoforms, Hsp77 and Hsp72, and C. grata one inducible form, Hsp75, which was upregulated at 40°C, suggesting C. grata has a more effective heat shock response than C. toreuma. The temperature-adaptive differences in cardiac thermal tolerance and Hsp expression match observed differences in thermally induced mortalities with the onset of summer and may help predict differential effects of climate change on the two congeners.     

A neutral DNA marker suggests that parallel physiological adaptations to open shore and salt marsh habitats have evolved more than once within two different species of gastropods

Local adaptation is an important mechanism generating physiological diversity and can be especially pronounced in species with restricted dispersal and gene flow such as direct developing snails of the genus Littorina. We compared physiological responses to salinity and desiccation stress in two co-occurring species of northeastern Pacific Littorina (L. subrotundata and L. sitkana) with salt marsh and open shore ecotypes. The animals from salt marsh populations were significantly more tolerant to low salinities and significantly less resistant to desiccation stress than their open shore counterparts. The lower resistance to desiccation in salt marsh animals was not associated with a higher rate of water loss during air exposure or with lower body water reserves, but instead reflected a lower tolerance to high salinities. These habitat-related physiological differences occurred in parallel in the two studied species of Littorina and persisted after prolonged laboratory acclimation, suggesting that they may reflect selection for markedly different local optima in the salt marsh habitats than in the open shore habitats. We used a neutral polymorphic nuclear DNA marker (intron of aminopeptidase N) to estimate the level of gene flow between the populations from different habitats and found isolation by distance regardless of the habitat from which the snails were collected. Our molecular data suggest that physiological cohesiveness of ecotypes can arise despite different genetic backgrounds, and could potentially be due to parallel evolution of convergent phenotypes in similar habitats.Communicated by O. Kinne, Oldendorf/Luhe     

Geographical subdivision,demographic history and gene flow in two sympatric species of intertidal snails, <Emphasis Type="Italic">Nerita scabricosta</Emphasis> and <Emphasis Type="Italic">Nerita funiculata</Emphasis>, from the tropical eastern Pacific

The patchy distribution of rocky intertidal communities in the tropical eastern Pacific (TEP) may impose severe constraints on the genetic connectivity among populations of marine invertebrates associated with this habitat. In this study, we analyzed a portion of the mitochondrial cytochrome c oxidase subunit I (COI) gene in two sympatric species of marine snails, Nerita scabricosta and Nerita funiculata, common inhabitants of the rocky intertidal from the Gulf of California (Sea of Cortez) and outer Pacific coast of the southern Baja California (Baja) peninsula to northern South America, to assess genetic connectivity among populations of each species. One of our aims was to determine whether the morphological, behavioral, and ecological differences observed among populations of both species throughout their range in the TEP corresponded to population genetic differences. In addition, we were interested in elucidating the demographic history of both species. We found no evidence of genetic structure throughout the Gulf of California and outer coast of the Baja peninsula region for either species. Comparisons between Gulf of California/Baja and Panama populations, however, showed significant genetic differentiation for N. scabricosta, but not for N. funiculata. The genetic differences between Mexican and Panamanian populations of N. scabricosta were consistent with previously reported ecological and behavioral differences for this species between these two distant regions. However, previously reported size differences between northern and central/southern Gulf of California individuals of N. scabricosta do not correspond with our findings of genetic connectivity among these populations. Results from neutrality tests (Tajima’s D and Fu’s F _S), the mismatch distribution, and Bayesian skyline analyses suggested that both species have experienced dramatic population expansions dating to the Pleistocene.     

Nutritional status affects the capacity of the snail Concholepas concholepas to synthesize Hsp70 when exposed to stressors associated with tidal regimes in the intertidal zone

Synthesis of the heat shock protein Hsp70 is one of the most important physiological mechanisms that intertidal organisms possess to counteract damage to macromolecules caused by stressors associated with the tidal cycle. However, the synthesis and activity of Hsp70 involves an elevated energetic cost. We evaluated the effect of the nutritional status (fed vs. starved for 2 weeks) of juvenile Concholepas concholepas mollusc on their capacity to synthesize Hsp70 during emersion (i.e. low tide) and immersion (i.e. high tide) at high temperatures (24 °C, e.g. summer conditions) and at low temperatures (7 °C, e.g. winter conditions). In addition, we evaluated whether Hsp70 is induced directly upon exposure to stress (emersion) or during recovery (re-immersion). Starvation decreased the content of stored energy substrates of juveniles as well as their ability to synthesize Hsp70 during emersion under thermal stress, especially at high temperatures. Additionally, analysis of environmental factors associated with laboratory simulation of tidal regimes indicated that juveniles in starvation, in contrast to fed juveniles, did not significantly increase their levels of Hsp70 during cold emersion (7 °C) or warm emersion (24 °C) or upon re-immersion. Induction of Hsp70 occurred during exposure to stress (low-tide conditions) and not when juveniles returned to “normal” conditions (high-tide conditions). Thus, the synthesis of Hsp70 for the juveniles of this intertidal snail species was coordinated and adapted to the tidal cycle, and the species responds in a similar way to hot and cold emersion conditions. The observed levels of Hsp70 reflect the ability of the individual to synthesize these proteins, which is dependent on the nutritional status of the individual.     

Nitrate reductase and glutamine synthetase activity,internal N pools,and growth of<Emphasis Type="Italic"> Ulva lactuca</Emphasis>: responses to long and short-term N supply

Fast-growing macroalgae, including Ulva lactuca Linnaeus, respond rapidly to changes in nutrient conditions, particularly to short-term N supply. This ability to rapidly take up and assimilate N contributes to the increasing occurrence of macroalgal blooms in heavily N loaded coastal ecosystems. To determine whether long-term nutrient histories affect short-term responses in activity of N-assimilating enzymes, including nitrate reductase (NRA) and glutamine synthetase activity (GSA), internal N storage, and macroalgal growth, we conducted an in situ nitrate fertilization experiment between 7 and 22 July 2004, with fronds of U. lactuca collected from estuaries with high and low N loads in Waquoit Bay, Cape Cod, Massachusetts, USA (N 41° and W 70°). Initial NRA, GSA, % N, δ¹⁵N, and growth of U. lactuca fronds were higher in the site where nitrate was in high supply. Differences in NRA persisted even after short-term experimental enrichment. Differences in internal N pools, δ¹⁵N, and growth, in contrast, mirrored the changes of nutrient supply. The rate of turnover of the internal N content of U. lactuca was quite short (<2 d), and turnover of enzyme activity may have been even shorter. N isotopic fractionation by U. lactuca appeared to be of small magnitude, unlike the case of phytoplankton, and similar to that of vascular plants. δ¹⁵N was a better indicator of short-term response to external and internal nutrient supplies in U. lactuca than enzyme activity or N content, and may reliably detect rapid changes in N availability, source, and uptake and assimilation processes.     

Chorioallantoic membrane as a non-lethal sampling method for polychlorinated biphenyls analysis in the northern diamondback terrapin (Malaclemys terrapin terrapin)

Polychlorinated biphenyls (PCBs) are documented endocrine-disrupting chemicals. This study examined the bioaccumulation of PCBs and the utility of the chorioallantoic membrane (CAM) as a non-lethal sampling technique for PCB analysis in the northern diamondback terrapin (Malaclemys terrapin terrapin), a species endemic to US East Coast salt marshes. To determine the viability of the CAM as a surrogate for PCB burdens within this species, maternal transfer of PCBs to CAM and egg samples was studied. Egg and CAM samples were collected from populations in Cape May County, NJ and Jamaica Bay, NY; liver samples were collected only from NJ terrapins. PCB congeners were detected in all sample types, but less-chlorinated congeners preferentially partitioned in the CAM. Liver and egg sample PCB concentrations were strongly correlated (p=0.002, r _s=0.883), but PCB concentrations of CAM were not strongly correlated to liver (p=0.035, r _s=0.733) and egg samples (p=0.036, r _s=0.484). Examination of homologue groups indicated a strong correlation between penta- and hexachlorinated biphenyls for all sample types. This study shows that maternal transfer of PCB contaminants occurs with selective partitioning of congeners into the CAM, and that analysis of the CAM alone may not reflect total PCB body burdens in this species.     

Physiological energetics of the intertidal sea anemone Anthopleura elegantissima

Energy budgets were calculated for individuals of the sea anemone Anthopleura elegantissima (Brandt), collected in 1981 and 1982 from Bodega Harbor, California, USA. Rates of ammonium excretion were measured in high-and low-intertidal, symbiotic and aposymbiotic sea anemones within 24 h of collection. Among symbiotic and aposymbiotic individuals, no differences in excretion rate were found on the basis of intertidal height. However, rates of ammonium excretion in aposymbiotic anemones (2.14 mol NH ₊ ⁴ g^-1 h^-1) were significantly higher than in symbiotic ones (0.288 mol NH ₊ ⁴ g^-1 h^-1). Rates of excretion were used with estimated rates of oxygen uptake to calculate nitrogen quotients (NQ). NQ and RQ values from the literature were used to calculate an oxyenthalpic equivalent [501 kJ (mol O₂)^-1 for R+U], and mass proportions of protein (54%), carbohydrate (44%) and lipid (2%) catabolized during routine metabolism in this species 24 h after feeding. Integrated energy budgets of these experimental anemones were calculated from data on ingestion, absorption and growth, and estimates of translocated energy from the symbiotic algae. Contribution of zooxanthellae to animal respiration based on translocation=90% and RQ=0.97 are 41 and 79% in high-and low-intertidal anemones, respectively. Calculated scope for growth is greater than directly measured growth in both high-and low-intertidal individuals. The deficit, estimated as 30% of assimilated energy in high-intertidal anemones, is attributed to unmeasured costs (specific dynamic effect) or production (mucus). Low-intertidal anemones lost mass during the experiment, implying that the magnitude of the deficit was greater in these anemones than in upper intertidal individuals. Anemones from both shore levels lost zooxanthellae during the experiment, which contributed to energy loss since the contribution of the zooxanthellae is greater in low-intertidal anemones. Scope for growth is preserved in high-intertidal anemones (29% of assimilated energy) because metabolic demands are lower due to aerial exposure, and prey capture rate is higher compared to lowshore anemones. Although possibly underestimated, lower scope for growth in low-shore anemones may result from continuous feeding and digestion processes that are less efficient than those of periodically feeding high-intertidal anemones.     

Analysis of the structure of decapod crustacean assemblages off the Catalan coast (North-West Mediterranean)

We sampled the communities of decapod crustaceans inhabiting the depth zone between 3 and 871 m off the Catalan coast (North-West Mediterranean) from June 1981 to June 1983. The 185 samples comprised 90 species differing widely in their depth distributions. Multivariate analysis revealed four distinct faunistic assemblages, (1) littoral communities over sandy bottoms, (2) shelf communities over terrigenous muds, (3) upper-slope communitics, and (4) lower-slope or bathyal communities. The brachyuran crab Liocarcinus depurator is the most abundant species of the shelf assemblage, although L. vernalis dominates over the shallow sandy bottoms of the shelf. The dominant species of the upper-slope assemblage are nektobenthic species (Solenocera membranacea, Plesionika heterocarpus, Processa canaliculata), pelagic species (Pasiphaea sivado, Sergestes arcticus), and benthic species (Macropipus tuberculatus, Munida intermedia, Nephrops norvegicus). Aristeus antennatus comprise most of the biomass of the lower-slope community, which supports a greater diversity than the other assemblages. The main assemblages appear to be related to different hydrological characteristics, the extent of seasonal fluctuations, and to the changes in sediment structure associated with changes in the steepness of the bottom.     

Spatial and temporal variation in the elemental and stable isotopic content of the seagrasses <Emphasis Type="Italic">Posidonia oceanica</Emphasis> and <Emphasis Type="Italic">Cymodocea nodosa</Emphasis> from the Illes Balears,Spain

Morphology, elemental content and isotopic composition of leaves of the seagrasses Posidonia oceanica and Cymodocea nodosa were highly variable across the Illes Balears, a Spanish archipelago in the western Mediterranean, and varied seasonally at one site in the study area. The data presented in this paper generally expand the reported ranges of nitrogen, phosphorus, iron and arsenic content and δ¹³C and δ¹⁵N for these species. Nitrogen and phosphorus content of P. oceanica leaves also showed significant seasonal variability; on an annual basis, P. oceanica leaves averaged 1.55% N and 0.14% P at this monitoring site. Both N and P were more concentrated in the leaves in winter than in summer, with winter maxima of 1.76% N and 0.17% P and summer minima of 1.34% N and 0.11% P. There was no significant annual pattern observed in the δ¹³C of P. oceanica leaves, but there was a repeated 0.6‰ seasonal fluctuation in δ¹⁵N. Mean annual δ¹⁵N was 4.0‰; δ¹⁵N was lowest in May and it increased through the summer and autumn to a maximum in November. Over the geographic range of our study area, there were interspecific differences in the carbon, nitrogen and phosphorus content of the two species. Posidonia oceanica N:P ratios were distributed around the critical value of 30:1 while the ratios for C. nodosa were lower than this value, suggesting P. oceanica we collected was not consistently limited by N or P while C. nodosa tended toward nitrogen limitation. Nutrient content was significantly correlated to morphological indicators of plant vigor. Fe content of P. oceanica leaves varied by a factor of 5×, with a minimum of 31.1 μg g⁻¹ and a maximum of 167.7 μg g⁻¹. Arsenic was present in much lower tissue concentrations than Fe, but the As concentrations were more variable; the maximum concentration of 1.60 μg g⁻¹ was eight times as high as the minimum of 0.20 μg g⁻¹. There were interspecific differences in δ¹³C of the two species; C. nodosa was consistently more enriched (δ¹³C = −7.8 ± 1.7‰) than P. oceanica (−13.2 ± 1.2‰). The δ¹³C of both species decreased significantly with increasing water depth. Depth related and regional variability in the δ¹³C and δ¹⁵N of both species were marked, suggesting that caution needs to be exercised when applying stable isotopes in food web analyses.