The synergistic effects of increasing temperature and CO2 levels on activity capacity and acid–base balance in the spider crab, Hyas araneus

With global climate change, ocean warming and acidification occur concomitantly. In this study, we tested the hypothesis that increasing CO₂ levels affect the acid–base balance and reduce the activity capacity of the Arctic spider crab Hyas araneus, especially at the limits of thermal tolerance. Crabs were acclimated to projected oceanic CO₂ levels for 12 days (today: 380, towards the year 2100: 750 and 1,120 and beyond: 3,000 μatm) and at two temperatures (1 and 4 °C). Effects of these treatments on the righting response (RR) were determined (1) at acclimation temperatures followed by (2) righting when exposed to an additional acute (15 min) heat stress at 12 °C. Prior to (resting) and after the consecutive stresses of combined righting activity and heat exposure, acid–base status and lactate contents were measured in the haemolymph. Under resting conditions, CO₂ caused a decrease in haemolymph pH and an increase in oxygen partial pressure. Despite some buffering via an accumulation of bicarbonate, the extracellular acidosis remained uncompensated at 1 °C, a trend exacerbated when animals were acclimated to 4 °C. The additional combined exposure to activity and heat had only a slight effect on blood gas and acid–base status. Righting activity in all crabs incubated at 1 and 4 °C was unaffected by elevated CO₂ levels or acute heat stress but was significantly reduced when both stressors acted synergistically. This impact was much stronger in the group acclimated at 1 °C where some individuals acclimated to high CO₂ levels stopped responding. Lactate only accumulated in the haemolymph after combined righting and heat stress. In the group acclimated to 1 °C, lactate content was highest under normocapnia and lowest at the highest CO₂ level in line with the finding that RR was largely reduced. In crabs acclimated to 4 °C, the RR was less affected by CO₂ such that activity caused lactate to increase with rising CO₂ levels. In line with the concept of oxygen and capacity limited thermal tolerance, all animals exposed to temperature extremes displayed a reduction in scope for performance, a trend exacerbated by increasing CO₂ levels. Additionally, the differences seen between cold- and warm-acclimated H. araneus after heat stress indicate that a small shift to higher acclimation temperatures also alleviates the response to temperature extremes, indicating a shift in the thermal tolerance window which reduces susceptibility to additional CO₂ exposure.     

Importance of biotic and abiotic regulators of abundance of the intertidal amphipod Calliopius laeviusculus (Amphipoda: Gammaridae) and assessment of the accuracy and precision of sampling methods for detecting abundance changes

We examined seasonal patterns of abundance for the intertidal amphipod Calliopius laeviusculus (Amphipoda: Gammaridae). Amphipods were sampled with an epibenthic sled during the daytime high tide period from 18 May to 8 August 1988. Amphipod density increased from May to June and reached maxima in both late June and early August. Amphipod density was unrelated to any abiotic component measured in the intertidal community. These variables included sampling location, wave height, water column height, water temperature, salinity and cloud cover. Changes in abundance were related with sampling date and with the onset of capelin (Mallotus villosus) spawning activity in the intertidal. Capelin eggs are an important food item for amphipods. The accuracy of density estimates obtained with the epibenthic sled was assessed through comparison with densities obtained with a more efficient quadrat sampler. Sled samples consistently sampled ca. 1% of the amphipod population. We found that a large portion of the amphipod population burrowed into the sediment and was not effectively sampled by the sled. Sled sampler precision was roughly equivalent to that of quadrats with D (precision) ranging from 0.26 to 0.42 for sled samples and D=0.29 for quadrat samples. Although sample collection with the epibenthic sled was achieved more quickly and under a wider range of weather conditions than was possible with the quadrat sampler, the serious underestimate of amphipod density based on sled samples alone indicates that both sled and quadrat samples, obtained in concert, are required to obtain accurate measures of daily variation in C. laeviusculus abundance.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches oceanographique du Quebec)     

Estimating variation in surface emissivities of intertidal macroalgae using an infrared thermometer and the effects on temperature measurements

Accurate measurements of surface temperatures with an infrared (IR) thermometer require input of the emissivities of the surfaces being measured; however, few determinations of the emissivities of intertidal organisms’ surfaces have been made. Emissivities of intertidal macroalgae were measured to determine whether algal species, measurement angle, hydration, and layering affected them. Emissivities were similar and averaged 0.94 among 11 of 13 species. The species with lower and more variable emissivities (Chondracanthus exasperatus and Desmarestia viridis) differed in morphology from the other species, which were relatively flat thin blades with little surface texture. Measurement angle caused emissivities to decrease significantly in Mazzaella splendens but not in three other species. Hydration and layering of Ulva lactuca also had no effect. At 22 °C, measured temperatures were within 1 °C of actual temperatures when thermometer emissivity settings ranged from 0.75 to 1.00. When emissivities were set lower than actual values, measured temperatures were lower than actual temperatures at 15 °C and higher than actual temperatures at 60 °C. When the IR thermometer was used to measure surface temperatures of nine species of intertidal algae immediately before they were inundated by the incoming tide, temperatures were higher in mid intertidal than low intertidal individuals and higher on a sunnier day than an overcast day. Temperatures of U. lactuca increased with increasing height on the shore, but temperatures of Ulvaria obscura did not. Temperatures were also higher in Fucus distichus blades than receptacles, and lower in U. lactuca and M. splendens occurring in the lower layers of stacks of algae.     

Survival, growth and reproduction of the salt-marsh amphipod Uhlorchestia spartinophila reared on natural diets of senescent and dead Spartina alterniflora leaves

The talitrid amphipod Uhlorchestia spartinophila lives in close association with standing-dead leaves of the smooth cordgrass Spartina alterniflora Loisel in salt marshes along the Atlantic coast of North America. This study probed the strength of the trophic link between the amphipod population and the decomposition process in this detrital-based ecosystem. We measured survival, growth and reproductive output in groups of amphipods reared for 6 wk on five diets derived from sheath and blade portions of S. alterniflora leaves just prior to (senescent) and during (dead) decomposition. In unfed treatments, the daily specific mortality rate was 0.391 and starved amphipods survived no longer than 11 d. Among the fed treatments, a diet of senescent sheaths resulted in the lowest survival (20%) and yielded no offspring. Groups fed senescent blades, dead sheaths, dead blades and unwashed dead sheaths had survival rates of 56 to 84% and produced 5.0 to 12.5 offspring replicate⁻¹. Sex ratio usually favored females, but approached unity in treatments with high overall survival, suggesting that quality of available food resources may influence sex ratio in this species. Mean specific growth rates (mm mm⁻¹ d⁻¹) ranged from 0.013 to 0.016, and matched previous estimates of growth from field populations. Overall ecological performance (survival + growth + reproduction) was similar for all food treatments, except senescent sheaths, which yielded a final mean (±SD) dry biomass (0.4 ± 0.42 mg replicate⁻¹) of amphipods significantly lower than that of other diets (1.7 ± 0.81 to 2.6 ± 0.69 mg replicate⁻¹). Natural diets derived from decomposing cordgrass leaves can fulfill the nutritional requirements of U. spartinophila populations, but variation in initial amounts of living fungal biomass among the five experimental diets only partially explained the responses of amphipods in our experiment. Structural characteristics and variation in rates of fungal occupation within different portions of cordgrass leaves may affect the amphipod's ability to access plant production made available by decomposers. Received: 12 December 1996 / Accepted: 18 December 1996     

Vertical zonation of endosymbiotic zooxanthellae within a population of the intertidal sea anemone,Anthopleura uchidai

Intertidal organisms commonly form zonation bands along the shore. Environmental stressors often determine the vertical position of each zonation band. These stressors may similarly affect the distribution pattern of endogenous species in their intertidal hosts. To evaluate this possibility, we investigated the distribution pattern of endosymbiotic zooxanthellae in the genus Symbiodinium in a population of the intertidal sea anemone Anthopleura uchidai. We used molecular genetics to identify the Symbiodinium clades and found that A. uchidai has two clades of Symbiodinium, clades A and F. These Symbiodinium clades were disproportionally distributed along the vertical gradient of the intertidal shore. Anemones on the upper shore exclusively possessed clade F Symbiodinium while clade A Symbiodinium became dominant in the sea anemones on the lower shore. Photosynthesis activity assays showed that these Symbiodinium clades had similar net productivities at 23.3 and 31.8 °C at all irradiance levels. At 35 °C, however, clade A Symbiodinium exhibited substantially lower net productivities than clade F Symbiodinium, demonstrating that these Symbiodinium clades have distinct tolerances to thermal stress. These results suggest that the thermal gradient across tidal height is a major factor shaping the zonation pattern of Symbiodinium clades in A. uchidai.     

Behaviour moderates climate warming vulnerability in high-rocky-shore snails: interactions of habitat use, energy consumption and environmental temperature

High-rocky-shore intertidal animals are predicted to be generally more vulnerable to climate warming than lower-shore species, because their thermal tolerances lie closer to maximum environmental temperatures (T _e). However, this prediction is based on taxonomically and ecologically limited information. The present study investigated the effect of habitat use on climate warming vulnerability of the tropical high-shore snail, Echinolittorina malaccana (from Brunei Darussalam, 5°N), which aestivates in sun-exposed or shaded habitats. The thermal regimes of these habitats differed vastly, but snails showed similar daily energy consumption in either habitat, due to temperature-insensitive metabolism (TIM) between 35 and 46 °C in the sun-resting snails. However, maximum T _e values in the shade and the sun were 35 and 46 °C, respectively, suggesting that sun-resting snails, which presently experience temperatures near the incipient lethal temperature range (46–56 °C), should be more threatened by further warming than shade-resting snails, which have an 11 °C ‘safety margin’. Thus, vulnerability of high-shore species to climate warming could be moderated by availability of shaded habitat, making predictions for these species more complex than previously realized.     

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.     

Effects of salinity on multiplication and transmission of an intertidal trematode parasite

Salinity levels vary spatially in coastal areas, depending on proximity to freshwater sources, and may also be slowly decreasing as a result of anthropogenic climatic changes. The impact of salinity on host–parasite interactions is potentially a key regulator of transmission processes in intertidal areas, where trematodes are extremely common parasites of invertebrates and vertebrates. We investigated experimentally the effects of long-term exposure to decreased salinity levels on output of infective stages (cercariae) and their transmission success in the trematode Philophthalmus sp. This parasite uses the snail Zeacumantus subcarinatus as intermediate host, in which it asexually produces cercariae. After leaving the snail, cercariae encyst externally on hard substrates to await accidental ingestion by shorebirds, which serve as definitive hosts. We found that at reduced salinities (25 or 30 psu), the cercarial output of the parasite was lower, the time taken by cercariae to encyst was longer, fewer cercariae successfully encysted and encysted parasites had lower long-term survival than at normal seawater salinity (35 psu). The strong effect of salinity on the replication and transmission of this parasite suggests that there may be sources and sinks of transmission to birds along coastal areas, depending on local salinity conditions. Also, unless it evolves to adapt to changing conditions, the predicted reduction in salinity as a consequence of climate change may have negative impact on the parasite’s abundance.     

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.     

Temperature and salinity interactively impact early juvenile development: a bottleneck in barnacle ontogeny

When juvenile mortality or juvenile growth is impacted by temperature and salinity, these factors have a substantial effect on recruitment success and population dynamics in benthic ecosystems. Using freshly settled cyprids of Amphibalanus improvisus, we investigated the combined effects of temperature (12, 20 and 28 °C) and salinity (5, 15 and 30 psu) on early juvenile stage performance. Mortality as well as size (basal diameter, dry weight, and ash-free dry weight) was monitored for a period of 40 days. Mortality was high (42–63 %) during the first week following attachment, regardless of the temperature and salinity treatments. Subsequently, mortality and size were interactively influenced by temperature and salinity. Highest mortality and lowest size of juveniles occurred at lowest temperature (12 °C) and salinity (5 psu). Apparently, low temperature (12 °C) narrowed the barnacles’ salinity tolerance. Juvenile barnacles constructed more shell material compared to body mass at high temperature and high salinity, while a reverse situation was observed at low temperature and low salinity. Our results demonstrate that environmental changes can directly and/or indirectly alter patterns of survivorship and size. Warming and desalination as predicted for the Baltic Sea in the course of climate change may, however, act antagonistically and compensate each other’s isolated effect on barnacles.     

No evidence for homeoviscous adaptation in intertidal snails: analysis of membrane fluidity during thermal acclimation, thermal acclimatization, and across thermal microhabitats

Many eurythermal organisms alter composition of their membranes to counter perturbing effects of environmental temperature variation on membrane fluidity, a process known as homeoviscous adaptation. Marine intertidal gastropods experience uniquely large thermal excursions that challenge the functional integrity of their membranes on tidal and seasonal timescales. This study measured and compared membrane fluidity in marine intertidal snail species under three scenarios: (1) laboratory thermal acclimation, (2) thermal acclimatization during a hot midday low tide, and (3) thermal acclimatization across the vertical intertidal zone gradient in temperature. For each scenario, we used fluorescence polarization of the membrane probe DPH to measure membrane fluidity in individual samples of gill and mantle tissue. A four-week thermal acclimation of Tegula funebralis to 5, 15, and 25°C did not induce differences in membrane fluidity. Littorina keenae sampled from two thermal microhabitats at the beginning and end of a hot midday low tide exhibited no significant differences in membrane fluidity, either as a function of time of day or as a function of thermal microhabitat, despite changes in body temperature up to 24°C within 8 h. Membrane fluidities of a diverse group of snails collected from high, middle, and low vertical regions of the intertidal zone varied among species but did not correlate with thermal microhabitat. Our data suggest intertidal gastropod snails do not exhibit homeoviscous adaptation of gill and mantle membranes. We discuss possible alternatives for how these organisms counter thermal excursions characteristic of the marine intertidal zone.     

Physiological and behavioral responses of temperate seahorses (Hippocampus guttulatus) to environmental warming

The aim of the present study was to evaluate, for the first time, the effect of environmental warming on the metabolic and behavioral ecology of a temperate seahorse, Hippocampus guttulatus. More specifically, we compared routine metabolic rates, thermal sensitivity, ventilation rates, food intake, and behavioral patterns at average spring temperature (18 °C), average summer temperature (26 °C), temperatures that they endure during summer heat wave events (28 °C), and in a near-future warming scenario (+2; 30 °C) in Sado estuary, Portugal. Both newborn juveniles and adults showed significant increases in metabolic rates with rising temperatures. However, newborns were more impacted by future warming via metabolic depression (i.e., heat-induced hipometabolism). In adult stages, ventilation rates also increased significantly with environmental warming, but food intake remained unchanged. Moreover, the frequency of swimming, foraging, swinging, and inactivity did not significantly change between the different thermal scenarios. Thus, we provide evidence that, while adult seahorses show great resilience to heat stress and are not expected to go through any physiological impairment and behavioral change with the projected near-future warming, the early stages display greater thermal sensitivity and may face greater metabolic challenges with potential cascading consequences for their growth and survival.     

Impact of ocean acidification on escape performance of the king scallop, Pecten maximus, from Norway

The ongoing process of ocean acidification already affects marine life, and according to the concept of oxygen and capacity limitation of thermal tolerance, these effects may be intensified at the borders of the thermal tolerance window. We studied the effects of elevated CO₂ concentrations on clapping performance and energy metabolism of the commercially important scallop Pecten maximus. Individuals were exposed for at least 30 days to 4 °C (winter) or to 10 °C (spring/summer) at either ambient (0.04 kPa, normocapnia) or predicted future PCO₂ levels (0.11 kPa, hypercapnia). Cold-exposed (4 °C) groups revealed thermal stress exacerbated by PCO₂ indicated by a high mortality overall and its increase from 55 % under normocapnia to 90 % under hypercapnia. We therefore excluded the 4 °C groups from further experimentation. Scallops at 10 °C showed impaired clapping performance following hypercapnic exposure. Force production was significantly reduced although the number of claps was unchanged between normocapnia- and hypercapnia-exposed scallops. The difference between maximal and resting metabolic rate (aerobic scope) of the hypercapnic scallops was significantly reduced compared with normocapnic animals, indicating a reduction in net aerobic scope. Our data confirm that ocean acidification narrows the thermal tolerance range of scallops resulting in elevated vulnerability to temperature extremes and impairs the animal’s performance capacity with potentially detrimental consequences for its fitness and survival in the ocean of tomorrow.     

Elevated temperature elicits greater effects than decreased pH on the development, feeding and metabolism of northern shrimp (Pandalus borealis) larvae

Climate models predict that the average temperature in the North Sea could increase 3–5 °C and surface-waters pH could decrease 0.3–0.5 pH units by the end of this century. Consequently, we investigated the combined effect of decreased pH (control pH 8.1; decreased pH 7.6) and temperature (control 6.7 °C; elevated 9.5 °C) on the hatching timing and success, and the zoeal development, survival, feeding, respiration and growth (up to stage IV zoea) of the northern shrimp, Pandalus borealis. At elevated temperature, embryos hatched 3 days earlier, but experienced 2–4 % reduced survival. Larvae developed 9 days faster until stage IV zoea under elevated temperature and exhibited an increase in metabolic rates (ca 20 %) and an increase in feeding rates (ca 15–20 %). Decreased pH increased the development time, but only at the low temperature. We conclude that warming will likely exert a greater effect on shrimp larval development than ocean acidification manifesting itself as accelerated developmental rates with greater maintenance costs and decreased recruitment in terms of number and size.     

Energy demand of larval and juvenile Cape horse mackerels, Trachurus capensis, and indications of hypoxia tolerance as benefit in a changing environment

Trachurus capensis is an important fisheries resource in the degraded Namibian upwelling ecosystem. Food supply and shoaling of hypoxic zones are hypothesised to influence the species’ recruitment success. This paper is the first to quantify energy requirements and hypoxia tolerance of larval and juvenile stages of a Trachurus species. We measured normoxic respiration rates of T. capensis with a size range from 0.001 to 20.8 g wet mass (WM) collected off Cape Town (33.9°S, 18.5°E, 12/2009) and in the northern Benguela (17–24°S, 11–15°E, 02/2011). Routine metabolic rate (RMR) and standard routine rate (SRR) (mg O₂ h^?1) followed the allometric functions RMR = 0.418 WM^0.774 and SRR = 0.275 WM^0.855, respectively. Larvae and juveniles had comparatively high metabolic rates, and the energy demand of juveniles at the upper end of the size range appeared too high to be fuelled by a copepod diet alone. T. capensis’ early life stages showed a high tolerance to hypoxic conditions. RMR in larvae did not change until 30 % O₂sat at 18 °C. In juveniles, critical oxygen saturation levels were low (P_C for SRR = 11.2 ± 1.7 % O₂sat and P_C for RMR = 13.2 ± 1.6 % O₂sat at 20 °C) and oxy-regulation effective (regulation index = 0.78 ± 0.09). A high hypoxia tolerance may facilitate the retention of larvae in near-shore waters providing favourable feeding conditions and allowing juveniles to exploit food resources in the oxygen minimum zone. These mechanisms seem to well adapt T. capensis to a habitat affected by spreading hypoxic zones and probably enhance its recruitment success.     

Thermal tolerance of early development in tropical and temperate sea urchins: inferences for the tropicalization of eastern Australia

The thermal envelope of development to the larval stage of two echinoids from eastern Australia was characterized to determine whether they fill their potential latitudinal ranges as indicated by tolerance limits. The tropical sand dollar, Arachnoides placenta, a species that is not known to have shifted its range, was investigated in Townsville, northern Australia (19°20′S, 146°77′E), during its autumn spawning season (May 2012). The subtropical/temperate sea urchin, Centrostephanus rodgersii, a species that has undergone poleward range expansion, was investigated in Sydney, southern Australia (33°58′S, 151°14′E), during its winter spawning season (August 2012). The thermal tolerance of development was determined in embryos and larvae reared at twelve temperatures. For A. placenta, the ambient water temperature near Townsville and experimental control were 24 °C and treatments ranged from 14 to 37 °C. For C. rodgersii, ambient Sydney water temperature and experimental control were 17 °C, and the treatment range was 9–31 °C. A. placenta had a broader developmental thermal envelope (14 °C range 17–31 °C) than C. rodgersii (9 °C range 13–22 °C). Both species developed successfully at temperatures well below ambient, suggesting that cooler water is not a barrier to poleward migration for either species. Both species presently live near the upper thermal limits for larval development, and future ocean warming could lead to contractions of their northern range limits. This study provides insights into the factors influencing the realized and potential distribution of planktonic life stages and changes to adult distribution in response to global change.     

Species-specific dinoflagellate vertical distribution in temperature-stratified waters

Thermal stratification is increasing in strength as a result of higher surface water temperature. This could influence the vertical distribution of vertically migrating dinoflagellates. We studied the diel vertical distribution of the dinoflagellates Heterocapsa triquetra and Prorocentrum minimum using stratified laboratory columns with two thermoclines of different strength (ΔT° = 10 or 17 °C), with below cline temperature of 8 °C. Above the thermocline, nutrient depletion simulated the natural summer conditions in the Baltic Sea. Our study shows that H. triquetra and P. minimum can behave differently in terms of their vertical occurrence, both in space and in time when subjected to thermoclines of different strength. Also, both dinoflagellate species showed species-specific distribution patterns. In the ΔT° = 10 °C treatment, H. triquetra cells performed a diel vertical migration (DVM) behavior just above the thermocline, but not in the ΔT° = 17 °C. In the ΔT° = 17 °C, the cells did not migrate and cell densities in the water column decreased over time. Opposing results were observed for P. minimum, where a DVM pattern was found exclusively below the thermocline of ΔT° = 17 °C, while in the ΔT° = 10 °C treatment, no clear DVM pattern was observed, and the highest number of cells were found in the cold bottom water. These results indicate that an increase in thermal stratification can influence species-specific dinoflagellate distribution, behavior, and survival.     

Behaviour and distribution of the sand-beach amphipod Orchestoidea corniculata

The field distribution of Orchestoidea corniculata Stont a talitrid amphipod inhabiting the upper intertidal zone of sandy beaches, was measured by pitfall trap and subsurface core transects. Variations in the distribution of juveniles and adults were recorded. At Coal Oil Point (Goleta, California, USA), the population was concentrated on the leeward side of the Point and, with increasing distance from the Point, both the number and size of beachhoppers decreased. The amphipods burrowed in a 5 to 7 m wide band during the day, and were active over a 15 to 25 m area at night. When they emerged from their burrows, a net seaward movement of 3 m was observed for the population. Behavioural tests suggested that sand penetrability is an important factor determining the location of burrow sites. Diurnal orientation was investigated, and it was found that the amphipods oriented in a landward direction despite overcast sky conditions, beach slope, moisture gradients, or displacement to a new beach.     

Preferences and tradeoffs in nectar temperature and nectar concentration in the Asian hive bee Apis cerana

Honey bee foragers need to asses and make trade-offs between a number of potentially conflicting floral attributes. Here, we investigate multi-attribute decision making in the eastern honey bee, Apis cerana, when foraging on food sources that varied in warmth and sucrose concentration. We show that foragers prefer warm (30 °C) sucrose solution over cool (10 °C) sucrose solution and concentrated (30 % w/w) sucrose solution over dilute (15 % w/w) sucrose solution. When we offered the preferred sucrose concentration (30 % w/w) at the less-preferred temperature (10 °C), and the less-preferred sucrose concentration (15 % w/w) at the preferred temperature (30 °C), foragers prioritized warmth by choosing the warmer, but lower concentration solution. When the temperature difference was less extreme, bees preferred more concentrated cooler syrup (30 % ww at 15 °C over 15 % 30 °C). However, the addition of a decoy item to the choice set had a significant effect on the bees' preferences. Our results highlight the critical importance of considering context effects when measuring the foraging preferences of animals.     

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.