The influence of coastal upwelling on the functional structure of rocky intertidal communities

Summary Relationships between organisms at all trophic levels are influenced by the primary productivity of the ecosystem, and factors which enhance rates of primary production may modify trophic relationships and community structure. Nutrient enrichment of intertidal and nearshore waters leads to enhanced production by intertidal algae, and it was hypothesized that where rocky shores are washed by nutrient-rich upwelled waters, the intertidal communities should show a characteristic functional structure, based on the effects of enhanced primary production. Study sites were chosen on rocky shores in southern Africa, central Chile and the Canary Islands, in areas with and without coastal upwelling, and mid-shore community structure at these sites was analysed in terms of the abundance of certain functional guilds of organisms.It was found that algal cover and the biomass of herbivorous limpets supported per unit area on rocky shores were significantly greater in regions of coastal upwelling than in regions where upwelling did not occur. Ground cover by sessile filter-feeding organisms was significantly greater on shores in non-upwelled areas. However, correspondence analysis showed no functional aspect of intertidal community structure that was characteristic of coasts washed by upwelled waters. Primary reasons for this are probably the large variations in the nature of nutrient enrichment that accompanies upwelling, and in the nutrient status of non-upwelled areas. Other factors are man's exploitation of intertidal organisms and differences in the genetic origins of the intertidal species involved.     

Spatial patterns of growth in the mussel, Mytilus californianus, across a major oceanographic and biogeographic boundary at Point Conception, California, USA

The Point Conception, California, USA region (hereafter PC) is one of the most important biogeographic and oceanographic discontinuities on the US west coast. Here we address how mesoscale oceanographic variability in the region around PC affects the growth of the competitively dominant species in the rocky intertidal assemblage: the mussel Mytilus californianus. Strong upwelling and high wave exposure dominate the California coast north of PC, and weak, seasonal upwelling and warmer water temperatures are characteristic of the region south/east of PC. We hypothesized that the oceanographic gradients in temperature, upwelling, wave exposure and productivity around PC would exert strong bottom–up influences on growth rates of mussels, potentially underlying large-scale differences in community structure around the PC region. We evaluated these predictions by measuring mussel growth rates across the PC region both in the intertidal and offshore on moorings. Intertidal mussels grew at much higher rates at sites south relative to north of PC and growth rates decreased in a gradient from south to north. The gradient in intertidal mussel growth around PC was uncorrelated with inshore concentrations of chlorophyll-a, and was most strongly correlated with the alongshore gradient in wave exposure and intertidal temperature. Mussels on moorings offshore from the intertidal sites grew at much higher rates than those in the corresponding intertidal areas, and mussel growth rates did not differ significantly among moored locations around PC. The gradient of increasing temperature from north to south among mooring sites was correlated with a decreasing gradient in productivity in the same direction, potentially contributing to equal and opposite effects on mussel growth at offshore moorings. This study suggests that environmental factors such as cold temperatures and high wave exposure contribute to the spatial pattern of decreasing mussel growth rates from south to north around PC, underlying large-scale patterns of community structure in this region.     

Divergent succession and implications for alternative states on rocky intertidal shores

It has been hypothesized that rockweed stands and mussel beds in sheltered bays in the Gulf of Maine, USA, are alternative community states. As a test of this hypothesis, experimental clearings of different sizes were established in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis to determine if successional changes in large clearings developed species assemblages distinctly different from the surrounding A. nodosum stands. Clearings ranging from 1 to 8 m in diameter were created at 12 sites in 4 bays on Swan's Island, Maine, in 1996 to mimic the effects of ice scour, and abundances of gastropods, barnacles, mussels and fucoid algae were monitored until 2002. ANOVAs and MDS showed strong effects of clearing size and divergent successional changes in large clearings. Large clearings were quickly filled in and remained dominated by the alga Fucus vesiculosus L. and the barnacle Semibalanus balanoides (L). There was no evidence for site-specific effects, and Mantel tests showed clearing size was a better predictor of species composition than geographic distances among sites. Results suggest that large pulse disturbances using clearings of 8 m in diameter can initiate divergent successional pathways and have a protracted effect on species composition. Results are also consistent with the hypothesis that mussel beds and rockweed stands in sheltered bays may be alternative community states.     

Distribution, abundance, size and recruitment of the mussel, Mytilus californianus, across a major oceanographic and biogeographic boundary at Point Conception, California, USA

The Point Conception, California, USA region (hereafter PC) is one of the most important biogeographic and oceanographic discontinuities on the US west coast. Here we address how mesoscale oceanographic and environmental variability in the region around PC, CA may influence the distribution, abundance and size of the mussel Mytilus californianus, a competitively dominant species in rocky intertidal assemblages along the northeast Pacific. Strong upwelling and high wave exposure dominate the California coast north of PC, and weak, seasonal upwelling and warmer water temperatures are characteristic of the region south/east of PC. We hypothesized that the gradient in temperature, upwelling, and wave exposure around PC would greatly influence patterns of recruitment and abundance of mussels, potentially underlying large-scale differences in community structure. We evaluated these predictions by surveying intertidal community structure, mussel distribution, size, abundance and recruitment at a range of intertidal sites around PC. We found that intertidal communities north of PC were dominated mainly by macrophytes, while mussels and barnacles were relatively scarce. Intertidal communities south of PC were dominated by mussels and barnacles, with a low abundance of macrophytes. Mussels were larger and mussel beds were more expansive and extended lower in elevation at sites ranging from north to south around PC. At northern sites, high abundances of sea star predators and elevated wave exposure effectively displaced the entire mussel zone upwards. We found no differences in the numbers of mussel recruits to sites around PC, suggesting that spatial patterns of mussel abundance were not driven by differential recruitment. These results suggest that unlike other well-studied systems, supply of benthic larvae does not underly the large-scale gradient in community structure around PC. We suggest that environmental conditions favorable to macroalgal growth north of PC, and conditions favorable to filter-feeder growth south of PC may underly mesoscale patterns of intertidal community structure in this region.     

Community stability: effects of limpet removal and reintroduction in a rocky intertidal community

Summary The stability of a high rocky intertidal community was assessed in a controlled field experiment in which the most common consumers, limpets, were temporarily removed. Compared to the unmanipulated plots, the exclusion plots developed greater algal abundance and altered species composition of both algae and barnacles. The community was not perturbed beyond its capacity to recover, since the community structure of the limpet-removal plots converged on the structure of the unmanipulated plots following limpet reintroduction. Different components of the community recovered at different rates, depending on whether or not the species had a size-related escape from the limpets. Algae had no size-related escape from limpets. The difference in algal abundance between manipulated and unmanipulated plots lasted less than six months after limpet reintroduction. Barnacles, however, had a size-related escape from limpets and therefore recovered more slowly. The difference in barnacle species composition between the perturbed and unperturbed plots lasted for 17 months after limpet reintroduction. The length of the limpet removal period (16 or 28 months) did not appear to affect the rate of community recovery.     

Climate-driven interactions among rocky intertidal organisms caught between a rock and a hot place

To explore how climate may affect the structure of natural communities, we quantified the role of thermal stress in setting the high intertidal borders of the acorn barnacle, Semibalanus balanoides. At sites north and south of Cape Cod, a major faunal and thermal boundary on the east coast of North America, we examined the interacting effects of thermal stress and recruit density on individual survivorship. At hotter southern sites, particularly in bays, high intertidal barnacle survivorship was enhanced by experimental shading or by neighbors which ameliorate heat and desiccation stresses. In contrast, at cooler northern bay and coastal sites, neither shading nor group benefits increased barnacle survival, and mortality patterns were driven primarily by predators with largely boreal distributions. Our field results, like recent laboratory microcosm studies, suggest that predicting even simple community responses to climate change may be more complex than is currently appreciated. Received: 26 January 1999 / Accepted: 5 April 1999     

A test of the Menge-Sutherland model of community organization in a tropical rocky intertidal food web

Summary Menge and Sutherland (1976) predicted that in physically benign habitats: (1) community structure will be most strongly affected be predation, (2) the effect of predation will increase with a decrease in trophic position in the food web, (3) trophically intermediate species will be influenced by both predation and competition, and (4) competition will occur among prey species which successfully escape consumers. These predictions were tested in a tropical rocky intertidal community on the Pacific coast of Panama. The most abundant mobile species included fishes and crabs, which occupied the top trophic level, and predaceous gastropods and herbivorous molluscs, which occupied intermediate trophic levels. The most abundant sessile organisms were encrusting algae, foliose algae, barnacles, and bivalves. Diets were broad and overlapping, and 30.3% of the consumers were omnivorous. Each consumer group had strong effects on prey occurring at lower trophic levels: (1) Fishes and crabs reduced the abundance of predaceous snails, herbivorous molluscs, foliose algae, and sessile invertebrates. (2) Predaceous gastropods reduced the abundance of herbivorous molluscs and sessile invertebrates. (3) Herbivorous molluscs reduced the abundance of foliose algae and young stages of sessile invertebrates, and altered relative abundances of the encrusting algae. The encrusting algae, although normally the dominant space occupiers, proved to be inferior competitors for space with other sessile organisms when consumers were experimentally excluded. However, the crusts escaped consumers by virtue of superior anti-herbivore defenses and competed for space despite intense grazing. Observations do not support the hypothesis that the trophically intermediate species compete. Hence, with the exception of this last observation, the predictions of the Menge and Sutherland model were supported. Although further work is needed to evaluate other predictions of the model in this community, evidence from this study joins an increasing body of knowledge supporting the model. Contradictory evidence also exists, however, indicating that aspects of the model require revision.     

Tidal reduction and its effects on intertidal hard-substrate communities in the Oosterschelde estuary

Developments of intertidal hard-substrate communities in the Oosterschelde estuary were examined in perpendicular transects between high-water line and low-water line in the period 1982–1992. Prior to the beginning of the Oosterschelde estuary works a typology of communities was established and an overall survey of the estuary was carried out. The communities contain flora (algae) as well as fauna. Due to asphalting of dikes in 1986, much of the surface of several communities has been destroyed. The originally well developed communities with large species-richness have not returned. The small reduction in tidal amplitude due to the construction of the storm-surge barrier had a minor effect on the zonation of communities. In the upper part of the intertidal zone the boundaries of the communities moved 0.5–1.0 m downward in the transects along the dike-slopes. At an average inclination of 18° this means a vertical shift of about 15–30 cm. This reflects the reduction of the tidal amplitude: the high-water line shifted ca 22 cm downward. In a number of places sedimentation has caused a reduction in the number of smaller seaweed species in the lower eulittoral zone. At monitoring locations presence of the original communities is rather unchanged. Rare species like Pelvetia canaliculata, Actinia equina and Gelidium pusillum have been able to maintain quite successfully.     

Temporal variation of rocky intertidal resident fish assemblages - patterns and possible mechanisms with a note on sampling protocols

In this paper we review the temporal patterns of variation of rocky intertidal resident fish assemblages and discuss possible mechanisms that may explain these patterns. These assemblages tend to be highly resilient and stable on an inter-annual basis, but tend to undergo marked seasonal fluctuations, as different species recruit and reach different phases of their life cycles. Larval supply is usually not a limiting factor suggesting that post-recruitment deterministic mechanisms exert a strong influence on assemblage organization. At methodological level, it is argued that traditional destructive sampling techniques should be avoided whenever possible. It is proposed that a deeper understanding of these assemblages requires more detailed information on intraspecific interactions between members of each constituent species, and information on the interactions between limited numbers of species for which mutually influences are particularly strong. It is argued that size, topography and biotic cover of a pool may provide a limited number of favourable sites for fishes of a given species and class size so that intraspecific competition, and possibly predation of the individuals less able to get access to best sites, may explain to a considerable extent the inter-annual stability and resilience of these assemblages.     

Changes in the macrozoobenthos of the intertidal zone at Helgoland (German Bight, North Sea): a survey of 1984 repeated in 2002

Changes in the presence and absence of invertebrates as well as in species conspicuousness were documented in a rocky intertidal community based on surveys in 1984 and 2002. In 2002 six vertically and/or morphologically different stations of an intertidal platform were sampled. Five of these six habitats had already been surveyed in 1984. Replicating precisely the method of the first assessment, presence/absence changes as well as changes in species conspicuousness of 83 invertebrate species were documented, indicating that this intertidal community changed considerably during the 18-year interval. Compared with the study in 1984, 27 species newly appeared, whereas 32 species disappeared. Furthermore, 16 species increased in conspicuousness, whereas eight invertebrates decreased. The total number of species in 2002 was 154 versus 158 in 1984. Although algal species were not recorded as thoroughly as invertebrates, a massive decline in cover of Halidrys siliquosa was noted. Conversely, two invasive algal species became established after 1984, Sargassum muticum (since 1988), a cosmopolitan fucoid alga that prefers shallow subtidal areas for colonization, and Mastocarpus stellatus (introduction in the 1980s) that particularly colonized areas in the mid intertidal. In 1984 the mid intertidal zone was dominated by the brown alga Fucus serratus, whereas in 2002 the blue mussel Mytilus edulis and the periwinkle Littorina littorea were the most conspicuous organisms. Annual mean sea surface temperature (BAH measurements) warmed by 1.1°C over the past four decades. Range-related community shifts, introductions of non-indigenous species and the input of pollutants, are considered to explain long-term ecological changes in the invertebrate community at Helgoland.     

The distribution of Oligochaeta on an exposed rocky shore in southeast Ireland

The distribution of oligochaetes was studied on a wave-exposed, granite shore at Carnsore Point, County Wexford. Habitats sampled were crevices and shallow surface cracks in rock, Lichina pygmaea turf, a Mytilus edulis bed, barnacles, Laurencia pennatifida turf, Corallina officinalis turf and pools. Lichina, mussels, barnacles greater than 8 mm in height, Laurencia and Corallina were sampled quantitatively.Three species of Tubificidae and nine species of Enchytraeidae were recorded including two new species described from this locality and five which are undescribed. Oligochaetes were present in all the habitats sampled but were rare in pools and numbers were low on barnacle-covered rock except where barnacles were more than 8 mm in height. The number of species increased with decreasing tide level and was highest in lower shore crevices. Highest densities were recorded from Corallina turf and in barnacles > 8 mm. Lumbricillus semifuscus was the most widespread species, occurring in crevices and mats at all tide levels; other species had more restricted distributions. Grania species were confined to Corallina turf. Only three species of oligochaete were recorded from cracks where they were almost the only fauna present. Species diversity and density were influenced by physical structure of the habitat, particularly the amount of retained sediment.The oligochaetes are members of a rich cryptofauna in habitats which provide them with organic matter and moisture and protect them from environmental extremes and wave damage and from predators during immersion. Reasons for the scarcity of oligochaete records from exposed rocky shores and the high proportion of new species in this study are discussed.     

Macrofaunal diversity and habitat structure in intertidal boulder fields

Low-midshore boulder fields in Europe are increasingly subject to degradation related to beach fishing for crabs and abalones. The aim of the study was to better understand the structure and species diversity of macrofaunal assemblages living in a low-midshore boulder field in order to define conservation strategies for this biotope. Sampling strategy involved different spatial scales (macro- and microstrata) relative to the complexity and heterogeneity of the habitat. Assemblages of species and the corresponding habitats were identified by multifactorial analysis and compared by ANOVAs. The results show a hierarchical organization of the macrofauna within the boulder field, corresponding to three spatial levels of habitat structure: (i) eight microhabitats at the lowest level of spatial organization, each defined by a specific assemblage (subcommunity); (ii) three habitats at a middle level combining these microhabitats, each associated with a specific community (open rock, protected rock and sediment); and (iii) three macrohabitats at the uppermost level (corresponding to the scale of the entire boulder field and including the main geomorphological features of the beach), each defined by a specific assemblage of species (boulders on boulders, boulders on bedrock, and boulders on sediment). Two microhabitats with particularly high species diversity were regarded as the most important ecological units of the field in terms of conservation of specific and functional biodiversity. Comparison of habitat/community parameters showed (i) that habitat heterogeneity was not an accurate indicator of faunal diversity, and (ii) that habitat complexity enhanced the species richness of the fauna, but only above a threshold value. This enhancement was due to semi-sheltered microhabitats, which were found only in the most complex areas of the boulder field. It is likely that this complexity affects species richness qualitatively more than by the diversity of microhabitats. In other words, a cross-scale effect is apparent in which high habitat complexity on the middle spatial scale creates microhabitats on the lowest spatial scale that are characterized by stable semi-sheltered environmental conditions conducive to a maximum of species.     

Sediment on rocky intertidal reefs: Effects on early post-settlement stages of habitat-forming seaweeds

Modification of the coastal environment by human activities often leads to an increase in sedimentation of nearshore waters, with potential impacts on benthic marine assemblages. Here we assess the relationships between the levels of sedimentation, wave exposure and benthic organisms on rocky intertidal platforms around the Kaikoura Peninsula in southern New Zealand. We designed and tested five sediment traps to provide a tool for measuring the relative abundance of sediment across sites. Using field- and laboratory-based experiments, we tested hypotheses concerning whether different levels of sedimentation affected algal germling survival and algal zygote attachment, and whether the interactions of grazers and sediments affected germling survival. Levels of sediment and exposure were inversely related across seven sites. The fucoid alga Hormosira banksii characterized the more sedimented wave-sheltered and intermediately wave-exposed sites, with up to 80% cover in the lower mid-tidal zone, while the bull kelp Durvillaea antarctica characterized the three most wave-exposed sites. Grazing molluscs were found across all sites but species abundances varied by sediment and exposure levels. We did two 11-day trials testing the effects of different levels of sediment and different species of molluscan grazers on the survival of 1-week-old Hormosira germlings. Generally, there was no significant treatment effect of grazers, but mortality varied considerably among sites. In particular, one site had very high levels of sediment, which resulted in 100% mortality of germlings across all grazer treatments. Removing sediment at 1-week, 2-week and 4-week intervals made no difference to the survival of Hormosira germlings. In laboratory-based experiments, a light dusting of sediment reduced the percentage of zygotes of Hormosira by 34% and Durvillaea by 71% that attached to primary substratum, and a complete cover of sediment prevented attachment altogether. Overall, the effects of sediments and its interaction with molluscan grazing were highly variable but often large, particularly on the attachment of zygotes to primary substratum.     

Predation by gulls on crabs in rocky intertidal and shallow subtidal zones of the Gulf of Maine

Most organisms in intertidal areas are marine in origin; many have distributions that extend into the subtidal zone. Terrestrial predators such as mammals and birds may exploit these animals during low tide and can have considerable effects on intertidal food webs. Several studies have shown that avian predators are capable of reducing densities of sessile and slow-moving intertidal invertebrates but very few studies have considered avian predation on mobile invertebrate predators such as crabs. In this study, we investigated predation by Great Black-backed Gulls (Larus marinus Linnaeus) on three species of crabs (Cancer borealis Stimpson, Cancer irroratus Say, and Carcinus maenas Linnaeus). The study was at Appledore Island, ME (a gull breeding island) and 8 other sites throughout the Gulf of Maine, including breeding islands and mainland sites. On Appledore Island, intertidal and subtidal zones provided over one-third of prey remains found at gull nests, and crabs were a substantial proportion (∼ 30% to 40%) of the total remains. Similarly, collections of prey remains from intertidal areas indicated that crabs were by far the most common marine prey. C. borealis was eaten far more often and C. irroratus and C. maenas less often than expected at each site. Comparing numbers of carapaces to densities of crabs in low intertidal and shallow subtidal zones at each site, we estimated that gulls remove between 15% and 64% of C. borealis during diurnal low tides. The proportion of C. borealis eaten by gulls was independent of proximity to a gull colony. Approximately 97% of the outer coast of Maine is within 20 km of a breeding island. Thus, a lot of gull predation on crabs may occur throughout the Gulf of Maine during summer months. Crabs are important predators of other invertebrates; if predation by gulls reduces the number of crabs in intertidal and shallow subtidal areas, gulls may have important indirect effects on intertidal food webs.     

Thermal constraints on daily patterns of aggregation and density along an intertidal gradient in the periwinkle Echinolittorina peruviana

Variations in environmental temperature have both direct and indirect effects that affect organisms at levels ranging from intra-cellular physiological processes to ecological patterns. These variations are especially important for intertidal marine ectotherms such as littorinids since they alternate between periods of immersion in seawater, and must also experience long periods of emersion. In central Chile, Echinolittorina peruviana is one of the most conspicuous species on rocky intertidal shores, occurring at high tidal levels and in the splash zone. The species is known to resist direct exposure to the sun for long periods, although juveniles tend to be restricted to protected microhabitats. Adults show seasonal variations in abundance and vertical distribution and may form aggregations that have been shown to help reduce water loss and body temperature. In this study, we evaluate the relationship between daily thermal variations throughout the vertical distribution of this species and how these affect the patterns of density and aggregation. Our results suggest that one of the leading determinants of the spatio-temporal variation of density in E. peruviana may be operative temperature (T_O: the amount of stored heat resulting from the balance between heat fluxes into and out of the body, measured with taxidermic mounts mimicking heat transfer properties of the snail). T_O showed a strong negative relationship with density and a strong positive relationship with aggregation in the highest intertidal level monitored. The strength of these relationships decreased in importance at lower levels. While T_O alone cannot explain the abundance of E. peruviana throughout its range of distribution, our results show that it does have a strong influence that should be considered in addition to other ecological factors affecting the density of intertidal littorinids.     

The vertical distribution and seasonal abundance of intertidal microalgae on a rocky shore in New South Wales

The abundance of intertidal microalgal food resources was estimated on shores at Cape Banks (Botany Bay, New South Wales) by indirect assays of the concentration of chlorophyll on and in the surface of sandstone rocks. Examination of the relationships between chlorophyll concentration and weight and area of rock samples allowed valid assays to be made despite uncontrollable variations from replicate to replicate in the amount of rock sampled. Microalgal cells were also sampled directly by brushing them from the surface of the rock. The relationship between amount of chlorophyll in samples, and the number of microalgal cells in immediately adjacent brushed areas indicated a relatively constant amount of chlorophyll per cell at different heights and on different parts of the shore throughout different seasons of the year. These preliminaries determined that assays of chlorophyll provided reliable, repeatable estimates of the number of microalgal cells present.Replicated sampling at five heights on a transect spanning the mid-shore regions of a sheltered shore dominated by grazing gastropods, at approximately monthly intervals from 1977 to 1979, revealed a vertical gradient of increasing abundance of microalgae towards the bottom of the shore, except during summer. This trend was not correlated with decreasing abundance of microalgal grazers. During summer, the abundance of microalgae declined abruptly at all levels. The decrease was greatest at the lower levels, resulting in little or no difference among the abundances of microalgae at the five heights during summer. The seasonal changes were again unrelated to any major changes in densities of grazers. These trends confirm the general patterns of distribution of intertidal microalgae in previous studies in other parts of the world. The present data are discussed with respect to problems of interpretation of microalgal food resources for intertidal grazers.     

The transfer of cadmium, mercury, methylmercury, and zinc in an intertidal rocky shore food chain

We examined the transfer of cadmium (Cd), inorganic mercury [Hg(II)], methylmercury (MeHg), and zinc (Zn) in an intertidal rocky shore food chain, namely from marine phytoplankton to suspension-feeding rock oysters (Saccostrea cucullata) and finally to predatory whelks Thais clavigera. The uptake of metals from the dissolved phase was also concurrently quantified in the oysters and the whelks. Metal uptake by the oysters was not directly proportional, whereas metal uptake by the whelks was directly proportional to metal concentration in the water. The order of uptake was MeHg>Hg(II)>Zn>Cd, and was much higher in the oysters than in the whelks. The relative uptake of Zn and Cd was comparable between oysters and whelks, whereas MeHg and Hg(II) showed disproportionally higher uptake in oysters than in whelks as compared to Zn and Cd. The assimilation efficiencies (AEs) were in the order of MeHg>Zn>Cd=Hg(II) in oysters, whereas the AEs were highest for MeHg and comparable for Zn, Cd, and Hg(II) in the whelks. Pre-exposure of the oysters to different dissolved concentrations of Cd significantly elevated the AEs of Cd and Hg(II) but not of Zn, in association with the induction of metallothioneins in the oysters. The whelks significantly assimilated Cd and Zn from various prey (barnacles, oysters, mussels, and snails) with contrasting strageties of metal sequestration and storage. There was no significant relationship between the metal AE and the metal partitioning in the soluble fraction (including metallothionein-like proteins, heat stable protein, and organelles). The insoluble fraction of metals was also available for metal assimilation. Our calculations show that the dietary uptake of metals can be dominant in the overall bioaccumulation in the oysters and whelks, and the trophic transfer factor was >1 for all metals. Thus, the four metals have a high potential of being biomagnified in the intertidal rocky shore food chain. MeHg possessed the highest and Hg(II) and Cd the lowest potential of trophic transfer among the four metals considered.     

On the edge: The use of infrared thermography in monitoring responses of intertidal organisms to heat stress

Monitoring changes in the environment and the corresponding effects on biological systems still represents a major challenge in many marine and terrestrial ecological studies. Infrared thermography (IRT), and its application within the marine environment, represents an effective non-invasive tool for measuring the temperatures of organisms and their surrounding environment in situ. The use of IRT within the intertidal zone is particularly useful since habitat and organismal temperatures are highly variable across both fine spatial and temporal scales. We review the growing number of intertidal studies that utilise IRT to investigate the role of small-scale temperature variability in contributing to various demographic and ecological processes. In particular, we introduce two indicators of the thermal quality of intertidal habitats that can be readily used by ecologists but also management and conservation policy makers to assess the suitability of a given habitat for a range of species under actual and predicted climatic conditions. We also outline a range of potential applications involving IRT that have yet to be explored for monitoring coastal environments. These include combining photogrammetry, unmanned aerial vehicles and IRT to large-scale three-dimensional thermal maps of intertidal habitats. We also suggest ways in which this technology could facilitate environmental management objectives in a warming world, such as the identification and quantification of thermal refugia across various spatial and temporal scales. We affirm with previous studies that such thermal refugia are vital for the adaptation of intertidal communities to climate change and that IRT could facilitate more effective management and conservation of these areas. The IRT applications outlined in this review are by no means exhaustive or limited to rocky intertidal environments. We envision that IRT will become increasingly popular as environmental management agencies become increasingly concerned about global climate change and how to combat its negative consequences on ecosystems.