Climate change drives a shift in peatland ecosystem plant community: Implications for ecosystem function and stability

The composition of a peatland plant community has considerable effect on a range of ecosystem functions. Peatland plant community structure is predicted to change under future climate change, making the quantification of the direction and magnitude of this change a research priority. We subjected intact, replicated vegetated poor fen peat monoliths to elevated temperatures, increased atmospheric carbon dioxide (CO₂), and two water table levels in a factorial design to determine the individual and synergistic effects of climate change factors on the poor fen plant community composition. We identify three indicators of a regime shift occurring in our experimental poor fen system under climate change: nonlinear decline of Sphagnum at temperatures 8 °C above ambient conditions, concomitant increases in Carex spp. at temperatures 4 °C above ambient conditions suggesting a weakening of Sphagnum feedbacks on peat accumulation, and increased variance of the plant community composition and pore water pH through time. A temperature increase of +4 °C appeared to be a threshold for increased vascular plant abundance; however the magnitude of change was species dependent. Elevated temperature combined with elevated CO₂ had a synergistic effect on large graminoid species abundance, with a 15 times increase as compared to control conditions. Community analyses suggested that the balance between dominant plant species was tipped from Sphagnum to a graminoid‐dominated system by the combination of climate change factors. Our findings indicate that changes in peatland plant community composition are likely under future climate change conditions, with a demonstrated shift toward a dominance of graminoid species in poor fens.     

Climate change impact on seaweed meadow distribution in the North Atlantic rocky intertidal

The North-Atlantic has warmed faster than all other ocean basins and climate change scenarios predict sea surface temperature isotherms to shift up to 600 km northwards by the end of the 21st century. The pole-ward shift has already begun for many temperate seaweed species that are important intertidal foundation species. We asked the question: Where will climate change have the greatest impact on three foundational, macroalgal species that occur along North-Atlantic shores: Fucus serratus, Fucus vesiculosus, and Ascophyllum nodosum? To predict distributional changes of these key species under three IPCC (Intergovernmental Panel on Climate Change) climate change scenarios (A2, A1B, and B1) over the coming two centuries, we generated Ecological Niche Models with the program MAXENT. Model predictions suggest that these three species will shift northwards as an assemblage or “unit” and that phytogeographic changes will be most pronounced in the southern Arctic and the southern temperate provinces. Our models predict that Arctic shores in Canada, Greenland, and Spitsbergen will become suitable for all three species by 2100. Shores south of 45° North will become unsuitable for at least two of the three focal species on both the Northwest- and Northeast-Atlantic coasts by 2200. If these foundational species are unable to adapt to the rising temperatures, they will lose their centers of genetic diversity and their loss will trigger an unpredictable shift in the North-Atlantic intertidal ecosystem.     

Potential for adaptation in response to thermal stress in an intertidal macroalga

Understanding responses of marine algae to changing ocean temperatures requires knowledge of the impacts of elevated temperatures and the likelihood of adaptation to thermal stress. The potential for rapid evolution of thermal tolerance is dependent on the levels of heritable genetic variation in response to thermal stress within a population. Here, we use a quantitative genetic breeding design to establish whether there is a heritable variation in thermal sensitivity in two populations of a habitat‐forming intertidal macroalga, Hormosira banksii (Turner) Descaisne. Gametes from multiple parents were mixed and growth and photosynthetic performance were measured in the resulting embryos, which were incubated under control and elevated temperature (20°C and 28°C). Embryo growth was reduced at 28°C, but significant interactions between male genotype and temperature in one population indicated the presence of genetic variation in thermal sensitivity. Selection for more tolerant genotypes thus has the ability to result in the evolution of increased thermal tolerance. Furthermore, genetic correlations between embryos grown in the two temperatures were positive, indicating that those genotypes that performed well in elevated temperature also performed well in control temperature. Chlorophyll a fluorescence measurements showed a marked decrease in maximum quantum yield of photosystem II (PSII) under elevated temperature. There was an increase in the proportion of energy directed to photoinhibition (nonregulated nonphotochemical quenching) and a concomitant decrease in energy used to drive photochemistry and xanthophyll cycling (regulated nonphotochemical quenching). However, PSII performance between genotypes was similar, suggesting that thermal sensitivity is related to processes other than photosynthesis.     

Thermal tolerance,evaporative water loss,air-water oxygen consumption and zonation of intertidal prosobranchs: a new synthesis

Duration of emergence increases with tidal height on rocky shores therefore, emergence adaptations in intertidal species such as littorine and other prosobranch gastropods have been considered correlated with zonation patterns; temperature tolerance, desiccation resistance and aerial respiration rate all commonly assumed to increase progressively with increasing zonation level. Such direct correlations are rarely observed in nature. Maximal aerial gas exchange occurs in mid-shore, not high shore species. Temperature tolerance and desiccation resistance do not increase directly with shore height. Thus, hypotheses regarding physiological correlates of zonation require revaluation. A new hypothesis is presented that the high tide mark presents a single major physiological barrier on rocky shores. Above it, snails experience prolonged emergence and extensive desiccation; below it, predictable submergence and rehydration with each tidal cycle. Thus, desiccation stress is minimal below the high tide mark and maximal above it. Therefore, species restricted below high tide (the eulittoral zone) should display markedly different adaptive strategies to emergence than those above it (the eulittoral fringe). A review of the literature indicated that adaptations in eulittoral species are dominated by those allowing maintenance of activity and foraging in air including: evaporative cooling; low thermal tolerance; elevated aerial O₂ uptake rates; and high capacity for radiant heat absorption. Such adaptations exacerbate evaporative water loss. In contrast, species restricted to the eulittoral fringe display adaptive strategies that minimize desiccation and prolong survival of emergence including: foot withdrawal, preventing heat conduction from the substratum; aestivation in air; elevated thermal tolerance reducing necessity for evaporative cooling; position maintenance by cementation to the substratum and increased capacity for heat dissipation. In order to test of this hypothesis the upper thermal limits, tissue and substratum temperatures on emergence in direct sunlight and evaporative water loss and tissue temperatures on emergence in 40 °C were evaluated for specimens of six species of eulittoral and eulittoral fringe gastropods from a granite shore on Princess Royal Harbour near Albany, Western Australia. The results were consistant with adaptation to the proposed desiccation barrier at high tide. The eulittoral species, Austrocochlea constricta, Austrocochlea concamerata, Nerita atramentosa and Lepsiella vinosa, displayed adaptations dominated by maintenance of activity and foraging during emergence while the eulittoral fringe littorine species, Bembicium vittatum and Nodilittorina unifasciata displayed adaptations dominated by minization of activity and evaporative water loss during emergence. The evolution of adaptations allowing tolerance of prolonged desiccation have allowed littorine species to dominate high intertidal rocky shore gastropod faunas throughout the world's oceans.     

The limpet Patella vulgata L. at night in air: effective feeding on Ascophyllum nodosum monocultures and stranded seaweeds

Opportunistically, the limpet Patella vulgata will switch frommicrophagous grazing to feeding on attached macroalgae or, anovel finding, on accumulations of stranded seaweeds. Macroalgalfood is ingested primarily at night during emersion, when thethalli come to lie on the rock surface and are sufficientlydamp. Monocultures of Ascophyllum nodosum are not immune tothe destructive effects of limpets as previously thought. Patellavulgata can graze them down to complete disappearance, thuspushing Ascophyllum back to refuges where P. vulgata is absentor rare. Such refuges were found along a wide range from verysheltered to strongly wave-exposed rocky shores in Brittany,France. By a combination of its macrophagous and microphagousfeeding habits, P. vulgata can control mid-tidal macroalgalassemblages in a similar fashion to the action of sea urchinsupon kelp communities inhabiting the lower intertidal and shallowsublittoral zones of rocky shores. (Received 30 October 2006; accepted 25 May 2007)     

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.     

Physical effects of habitat‐forming species override latitudinal trends in temperature

Latitudinal and elevational temperature gradients (LTG and ETG) play central roles in biogeographical theory, underpinning predictions of large‐scale patterns in organismal thermal stress, species' ranges and distributional responses to climate change. Yet an enormous fraction of Earth's taxa live exclusively in habitats where foundation species modify temperatures. We examine little‐explored implications of this widespread trend using a classic model system for understanding heat stresses – rocky intertidal shores. Through integrated field measurements and laboratory trials, we demonstrate that thermal buffering by centimetre‐thick mussel and seaweed beds eliminates differences in stress‐inducing high temperatures and associated mortality risk that would otherwise arise over 14° of latitude and ~ 1 m of shore elevation. These results reveal the extent to which physical effects of habitat‐formers can overwhelm broad‐scale thermal trends, suggesting a need to re‐evaluate climate change predictions for many species. Notably, inhabitant populations may exhibit deceptive resilience to warming until refuge‐forming taxa become imperiled.     

Microhabitats choice in intertidal gastropods is species-, temperature- and habitat-specific

Understanding how behavioural adaptations can limit thermal stress for intertidal gastropods will be crucial for climate models. Some behavioural adaptations are already known to limit desiccation and thermal stresses as shell-lifting, shell-standing, towering, aggregation of conspecifics or habitat selection. Here we used the IRT (i.e. infrared thermography) to investigate the thermal heterogeneity of a rocky platform, with four different macrohabitats (i.e. bare rock, rock with barnacles, mussels and mussels incrusted by barnacles) over four thermally contrasted months. We investigated the body temperature of Littorina littorea and Patella vulgata found on this platform and the temperature of their microhabitat (i.e. the substratum within one body length around of each individual). We also considered the aggregation behaviour of each species and assessed the percentage of thermal microhabitat choice (i.e choice for a microhabitat with a temperature different than the surrounding substrate). We did not find any aggregation of L. littorea on the rocky platform during the four studied months. In contrast, P. vulgata were found in aggregates in all the studied periods and within each habitat, but there was no difference in body temperature between aggregated and solitary individuals. These two gastropods species were preferentially found on rock covered by barnacles in the four studied months. The presence of a thermal microhabitat choice in L. littorea and P. vulgata is habitat-dependent and also season-dependent. In June, July and November the choice was for a microhabitat with temperatures lower than the temperatures of the surrounding substrate whereas in December, individuals choose microhabitats with higher temperatures than the temperatures of their substratum. Taken together, these results suggest that gastropods species are able to explore their environment to find sustainable thermal macrohabitats and microhabitats and adapt this behaviour in function of the conditions of temperatures.     

Loss of thermal refugia near equatorial range limits

This study examines the importance of thermal refugia along the majority of the geographical range of a key intertidal species (Patella vulgata Linnaeus, 1758) on the Atlantic coast of Europe. We asked whether differences between sun‐exposed and shaded microhabitats were responsible for differences in physiological stress and ecological performance and examined the availability of refugia near equatorial range limits. Thermal differences between sun‐exposed and shaded microhabitats are consistently associated with differences in physiological performance, and the frequency of occurrence of high temperatures is most probably limiting the maximum population densities supported at any given place. Topographical complexity provides thermal refugia throughout most of the distribution range, although towards the equatorial edges the magnitude of the amelioration provided by shaded microhabitats is largely reduced. Importantly, the limiting effects of temperature, rather than being related to latitude, seem to be tightly associated with microsite variability, which therefore is likely to have profound effects on the way local populations (and consequently species) respond to climatic changes.     

Response of two species of amphibious stichaeoid fishes to temperature fluctuations in an intertidal habitat

The temperature regime of the intertidal microhabitat of two species of co-existing amphibious stichaeoid fishes, Anoplarchus purpurescens and Pholis ornata, were compared with experimentally determined tolerances to elevated temperatures. Studies of the critical thermal maxima of the two species revealed only slight differences in temperature tolerance but exposure to a cycled pattern of high temperatures sharply differentiated the resistance times of the two species with P. ornata capable of tolerating greater cumulative exposure to thermal stress when administered in a cycled program of temperature fluctuations with peak temperature at 27 °C. While the experimentally determined temperature tolerances exceeded those measured in the field, the greater tolerance of P. ornata may facilitate habitation of intertidal mudflats during summer months.     

Differences in leaf thermoregulation and water use strategies between three co‐occurring Atlantic forest tree species

Given anticipated climate changes, it is crucial to understand controls on leaf temperatures including variation between species in diverse ecosystems. In the first study of leaf energy balance in tropical montane forests, we observed current leaf temperature patterns on 3 tree species in the Atlantic forest, Brazil, over a 10‐day period and assessed whether and why patterns may vary among species. We found large leaf‐to‐air temperature differences (maximum 18.3 °C) and high leaf temperatures (over 35 °C) despite much lower air temperatures (maximum 22 °C). Leaf‐to‐air temperature differences were influenced strongly by radiation, whereas leaf temperatures were also influenced by air temperature. Leaf energy balance modelling informed by our measurements showed that observed differences in leaf temperature between 2 species were due to variation in leaf width and stomatal conductance. The results suggest a trade‐off between water use and leaf thermoregulation; Miconia cabussu has more conservative water use compared with Alchornea triplinervia due to lower transpiration under high vapour pressure deficit, with the consequence of higher leaf temperatures under thermal stress conditions. We highlight the importance of leaf functional traits for leaf thermoregulation and also note that the high radiation levels that occur in montane forests may exacerbate the threat from increasing air temperatures.     

Large-scale coexistence and small-scale segregation of key species on rocky shores

Biodiversity is defined by the scales at which organisms coexist. Coexistence at larger spatial scales may be underpinned by segregation at smaller spatial scales or temporal variability in behaviour and/or distribution. Limpets of the genus Patella are keystone grazers controlling the growth of macro-algae on many rocky shores. This study examined the distribution and abundance of P. vulgata and P. ulyssiponensis in relation to habitat and to each other at a range of spatial scales. Limpets were sampled on four shores on each of the east and west coasts of Ireland, in two different transects (10 m apart) at each of three shore heights in winter 2003 and summer 2004. P. vulgata and P. ulyssiponensis coexist around the coast of Ireland, but on shores where they co-occur, the pattern of coexistence begins to break down at the level of shore height: P. ulyssiponensis were more abundant on the low-shore than mid-shore, and P. vulgata were distributed evenly throughout the low and the mid-shore. At the level of habitat (pools vs. open rock), the two species exhibited a high degree of segregation: P. vulgata of all sizes were more common on open rock and P. ulyssiponensis of all sizes were more common in pools. The presence or absence of P. ulyssiponensis had no effect on the distribution of juvenile P. vulgata with respect to pools. Unexpectedly, however, a greater proportion of adult P. vulgata were found in pools at sites where P. ulyssiponensis were present in the pools than at sites where they were absent. The results of this study emphasize the need for multiple-scale analyses of the distribution and abundance of organisms to understand the organisation of natural assemblages. Handling editor: K. Martens     

Intertidal community composition along rocky shores in South-west Greenland: a quantitative approach

The intertidal communities on rocky shores are directly subjected to climatic changes in air and water temperatures and to derived effects of climate change, such as changes in freshwater run-off and ice dynamics. Global warming occurs at elevated rates in Greenland and results in changing species distributions with range expansions to the north and new species entering terrestrial habitats from the south. There is, however, no quantitative knowledge of past or present species distribution in the littoral zone of Southern Greenland, an area which represents an important gateway for northern range expansions of temperate species. This study provides baseline information on abundances of macroorganisms in the eulittoral Southern Greenland. This knowledge will pave the way for future studies on the impact of climate change and anthropogenic activities on these communities. Nine sites, situated at different exposure levels, were investigated. A total of 22 taxa were recorded, suggesting low species richness. Patellid limpets and predators such as dogwhelks, starfish and crabs were absent. Total standing stock ranged from 0 to 31,898 g m^?2. Species composition and biomasses were related to locally generated wave exposure, oceanic swells and ice scouring. The high standing stock at sheltered sites indicated that neither light, temperature nor nutrients, constrained buildup of biomass in this environment. Inshore seasonal measurements of water and air temperatures were recorded for the first time in the region, displaying low water temperatures and high variation in air temperatures, indicating lack of insulating stable sea ice in the area. Possible impacts of ongoing temperature changes are discussed based on recorded temperatures and meteorological data from the past 30 years.     

Phenological changes in intertidal con‐specific gastropods in response to climate warming

There is substantial evidence from terrestrial and freshwater systems of species responding to climate change through changes in their phenology. In the marine environment, however, there is less evidence. Using historic (1946–1949) and contemporary (2003–2007) data, collected from rocky shores of south‐west Britain, we investigated the affect of recent climate warming on the reproductive phenology of two con‐specific intertidal limpet grazers, with cool/boreal and warm/lusitanian centres of distribution. Reproductive development in the southern limpet, Patella depressa, has advanced, on average, 10.2 days per decade since the 1940s, with a longer reproductive season and more of the population reproductively active. The peak in the proportion of the population in advanced stages of gonad development was positively correlated with sea surface temperature (SST) in late spring/early summer, which has increased between the 1940s and 2000s. The advance in peak reproductive development of this species is double the average observed for terrestrial and freshwater systems and indicates, along with other studies, that marine species may be responding faster to climate warming. In contrast, the northern limpet, Patella vulgata, has experienced a delay in the timing of its reproductive development (on average 3.3 days per decade), as well as an increase in reproductive failure years and a reduction in the proportion of the population reaching advanced gonad stages. These results are the first to demonstrate a delay in the reproductive development of a cool‐temperate, winter spawner, towards cooler more favourable environmental conditions in response to climate warming. Such a delay in spawning will potentially lead to trophic miss‐matches, resulting in a rapid nonlinear decline of this species.     

Upward ant distribution shift corresponds with minimum,not maximum,temperature tolerance

Rapid climate change may prompt species distribution shifts upward and poleward, but species movement in itself is not sufficient to establish climate causation. Other dynamics, such as disturbance history, may prompt species distribution shifts resembling those expected from rapid climate change. Links between species distributions, regional climate trends and physiological mechanism are needed to convincingly establish climate‐induced species shifts. We examine a 38‐year shift (1974–2012) in an elevation ecotone between two closely related ant species, Aphaenogaster picea and A. rudis. Even though A. picea and A. rudis are closely related with North American distributions that sometimes overlap, they also exhibit local‐ and regional‐scale differences in temperature requirements so that A. rudis is more southerly and inhabits lower elevations whereas A. picea is more northerly and inhabits high elevations. We find considerable movement by the warm‐habitat species upward in elevation between 1974 and 2012 with A. rudis, replacing the cold‐habitat species, A. picea, along the southern edge of the Appalachian Mountain chain in north Georgia, USA. Concomitant with the distribution shifts, regional mean and maximum temperatures remain steady (1974–2012), but minimum temperatures increase. We collect individuals from the study sites and subject them to thermal tolerance testing in a controlled setting and find that maximum and minimum temperature acclimatization occurs along the elevation gradient in both species, but A. rudis consistently becomes physiologically incapacitated at minimum and maximum temperatures 2 °C higher than A. picea. These results indicate that rising minimum temperatures allow A. rudis to move upward in elevation and displace A. picea. Given that Aphaenogaster ants are the dominant woodland seed dispersers in eastern deciduous forests, and that their thermal tolerances drive distinct differences in temperature‐cued synchrony with early blooming plants, these climate responses not only impact ant‐ant interactions, but might have wide implications for ant‐plant interactions.     

Temperature,invaders and patchy habitat interact to limit the distribution of a vulnerable freshwater fish

Interacting global‐change drivers such as invasive species and climate warming are likely to have major and potentially unexpected influences on aquatic ecosystems. In river networks, modified water temperature combined with patchy physical conditions will likely cause shifts in the amount and distribution of suitable habitat, with influential invasive species further altering habitat availability. We examined how distributions of a thermally sensitive galaxiid fish native to the alpine rivers of New Zealand, Galaxias paucispondylus, were influenced by these drivers using spatially extensive presence–absence electrofishing surveys of 46 sites spread over four subcatchments. A unimodal response to water temperature and an interaction with substratum size meant G. paucispondylus were limited to streams with average summer water temperatures between 10.6 and 13.8 °C and were absent when average substratum sizes were <36 mm, regardless of temperature. In addition, non‐native trout >150 mm long excluded G. paucispondylus, but were only found in streams with average summer water temperatures <10.6 °C. These influences of trout likely strengthened the unimodal temperature response of G. paucispondylus and led to a very small G. paucispondylus realized niche. When predicted temperature increases were applied to catchment models, G. paucispondylus distributions were patchy and variable across subcatchments. Moreover, local physical characteristics of river networks were particularly important because of the non‐linear and interactive influences of temperature and substratum size on the outcome of species interactions. Therefore, substratum sizes, water temperature and a non‐native predator combined to influence the distribution of this thermally sensitive fish, illustrating how the effects of climate warming will likely be strongly context‐dependent and interactive.     

Caribbean mesophotic coral ecosystems are unlikely climate change refugia

Deeper coral reefs experience reduced temperatures and light and are often shielded from localized anthropogenic stressors such as pollution and fishing. The deep reef refugia hypothesis posits that light‐dependent stony coral species at deeper depths are buffered from thermal stress and will avoid bleaching‐related mass mortalities caused by increasing sea surface temperatures under climate change. This hypothesis has not been tested because data collection on deeper coral reefs is difficult. Here we show that deeper (mesophotic) reefs, 30–75 m depth, in the Caribbean are not refugia because they have lower bleaching threshold temperatures than shallow reefs. Over two thermal stress events, mesophotic reef bleaching was driven by a bleaching threshold that declines 0.26 °C every +10 m depth. Thus, the main premise of the deep reef refugia hypothesis that cooler environments are protective is incorrect; any increase in temperatures above the local mean warmest conditions can lead to thermal stress and bleaching. Thus, relatively cooler temperatures can no longer be considered a de facto refugium for corals and it is likely that many deeper coral reefs are as vulnerable to climate change as shallow water reefs.     

Evidence for a host role in thermotolerance divergence between populations of the mustard hill coral (Porites astreoides) from different reef environments

Studying the mechanisms that enable coral populations to inhabit spatially varying thermal environments can help evaluate how they will respond in time to the effects of global climate change and elucidate the evolutionary forces that enable or constrain adaptation. Inshore reefs in the Florida Keys experience higher temperatures than offshore reefs for prolonged periods during the summer. We conducted a common garden experiment with heat stress as our selective agent to test for local thermal adaptation in corals from inshore and offshore reefs. We show that inshore corals are more tolerant of a 6‐week temperature stress than offshore corals. Compared with inshore corals, offshore corals in the 31 °C treatment showed significantly elevated bleaching levels concomitant with a tendency towards reduced growth. In addition, dinoflagellate symbionts (Symbiodinium sp.) of offshore corals exhibited reduced photosynthetic efficiency. We did not detect differences in the frequencies of major (>5%) haplotypes comprising Symbiodinium communities hosted by inshore and offshore corals, nor did we observe frequency shifts (‘shuffling’) in response to thermal stress. Instead, coral host populations showed significant genetic divergence between inshore and offshore reefs, suggesting that in Porites astreoides, the coral host might play a prominent role in holobiont thermotolerance. Our results demonstrate that coral populations inhabiting reefs <10‐km apart can exhibit substantial differences in their physiological response to thermal stress, which could impact their population dynamics under climate change.     

Carry‐over effects of multiple stressors on benthic embryos are mediated by larval exposure to elevated UVB and temperature

Damaging effects of UVB in conjunction with other stressors associated with global change are well‐established, with many studies focused on vulnerable early life stages and immediate effects (e.g., mortality, developmental abnormalities). However, for organisms with complex life cycles, experiences at one life stage can have carry‐over effects on later life stages, such that sublethal effects may mediate later vulnerability to further stress. Here, we exposed embryos in benthic egg masses of the New Zealand intertidal gastropod Siphonaria australis to treatments of either periodic stress (e.g., elevated UVB, salinity, and water temperature mimicking tidepool conditions in which egg masses are commonly found during summer) or control conditions (low UVB, ambient salinity, and water temperatures). Although there was high mortality from stressed egg masses, 24% of larvae hatched successfully. We then exposed the hatching larvae from both egg mass treatments to different combinations of water temperature (15 or 20 °C) and light (high UVB or shade) 12 h per day for 10 days. The most stressful larval conditions of 20 °C/high UVB resulted in low survival and stunted growth. Carry‐over effects on survival were apparent for shaded larvae exposed to elevated temperature, where those from stressed egg masses had 1.8× higher mortality than those from control egg masses. Shaded larvae were also larger and had longer velar cilia if they were from control egg masses, independent of larval temperature. These results demonstrate that previous experience of environmental stress can influence vulnerability of later life stages to further stress, and that focus on a single life stage will underestimate cumulative effects of agents of global change.     

Effects of thermal history on the responses to thermal stress of a large benthic foraminifera,Calcarina gaudichaudii

Marine ecosystems, particularly coastal environments, are rapidly changing due to anthropogenic impacts resulting in increased global climate change (ocean warming), ocean acidification, hypoxia, and eutrophication. On coral reefs, symbiont-bearing large benthic foraminifera (LBFs) can play a key role as reef constituents and carbonate producers, contributing up to 5% of reef-scale carbonate budgets. However, projected climate change, particularly ocean warming, has the potential to significantly alter the conditions in which marine organisms persist. While the response of LBFs to elevated thermal stress is well documented in laboratory studies, the potential influence of adaptation or acclimatization through prior environmental thermal history on this response remains largely unknown. In this study, specimens of Calcarina gaudichaudii, an LBF from the Penghu Islands, Taiwan, were collected from thermally variable intertidal and thermally stable subtidal (~ 6 m depth) environments representing thermal history. LBFs were then acclimated to laboratory conditions at ambient (25 °C) and elevated (28 °C) temperatures for three weeks, and subsequently exposed to control and heat stress treatments (25 °C, 28 °C, 30 °C, 33 °C) for an additional one week. Photosynthetic rates (determined through oxygen flux measurements) of C. gaudichaudii significantly decreased in specimens collected at subtidal depths acclimated at 25 °C when compared to those acclimated at 28 °C, whereas there was no effect of thermal history on respiration, indicating that symbiont and holobiont responses may differ in LBFs. Additionally, maximum photochemical efficiency (Fv/Fm) significantly decreased as a result of heat stress, although bleaching was not visually observed after one week. These results highlight the plastic responses of the algal microbiome and indicate that thermal history, acclimatization temperature, and heat stress interact to affect the physiological status of C. gaudichaudii. This study adds to the growing literature which highlights the larger implications of understanding thermal history as an important factor to consider to better understand how ecosystem processes (e.g., carbonate production) are altered on modern coral reefs.