Marine biodiversity and climate change: assessing and predicting the influence of climatic change using intertidal rocky shore biota. Occasional Publication of the Marine Biological Association 20

In the last 60 years climate change has altered the distribution and abundance of many seashore species. Below is a summary of the findings of this project. The MarClim project was a four year multi-partner funded project created to investigate the effects of climatic warming on marine biodiversity. In particular the project aimed to use intertidal species, whose abundances had been shown to fluctuate with changes in climatic conditions, as indicator species of likely responses of species not only on rocky shores, but also those found offshore. The project used historic time series data, from in some cases the 1950s onwards, and contemporary data collected as part of the MarClim project (2001-2005), to provide evidence of changes in the abundance, range and population structure of intertidal species and relate these changes to recent rapid climatic warming. In particular quantitative counts of barnacles, limpets and trochids were made as well as semi-quantitative surveys of up to 56 intertidal taxa.Historic and contemporary data informed experiments to understand the mechanisms behind these changes and models to predict future species ranges and abundances.

The influence of mussel-modified habitat on Fucus serratus L. a rocky intertidal canopy-forming macroalga

The influence of habitat modification by Mytilus edulis L. on the settlement and development of Fucus serratus populations was investigated on rocky shores of the Isle of Anglesey, North Wales. Settlement of fucoids was higher inside mussel habitat than outside on one of two shores studied. The effect of microhabitat on survival of fucoid germlings was examined by transplanting the germlings into and outside mussel habitats, each with and without the exclusion of grazers. Observation showed that periwinkles and top shells were abundant in mussel habitat, while limpets dominated bare rock. Exclusion of grazers greatly enhanced the survival of fucoid germlings in both habitats, indicating that while mussel habitat supports a different grazer assemblage to bare rock, both assemblages are important in limiting fucoid recruitment. The risk of dislodgement was assessed and compared between fucoids growing on mussel shells and bare rock. In situ pull-tests showed that less force was required to detach large fertile thalli growing on mussel shells than those growing on the rock. Adhesion was generally broken between the mussel and the rock rather than between the holdfast and the mussel. These observations indicate that mussels provide an unstable substrate for mature fucoids. Overall results suggest that a negative effect of mussel-modified habitat on fucoids is profound in adults; but the effect is context-dependent in juveniles and can be positive at settlement. Results from a survey on population structure of fucoids across two shores showed that there were greater numbers of large fertile fucoids growing directly attached to rock than on mussel shells, while there was no difference for juvenile fucoids confirming the experimental results. Moreover thalli larger than 60 cm were found only on the rock but not on shells. This finding suggests that a mussel dominated habitat may have a significant impact on reproductive output in fucoid populations.

The influence of mussel-modified habitat on Fucus serratus L. a rocky intertidal canopy-forming macroalga

The influence of habitat modification by Mytilus edulis L. on the settlement and development of Fucus serratus populations was investigated on rocky shores of the Isle of Anglesey, North Wales. Settlement of fucoids was higher inside mussel habitat than outside on one of two shores studied. The effect of microhabitat on survival of fucoid germlings was examined by transplanting the germlings into and outside mussel habitats, each with and without the exclusion of grazers. Observation showed that periwinkles and top shells were abundant in mussel habitat, while limpets dominated bare rock. Exclusion of grazers greatly enhanced the survival of fucoid germlings in both habitats, indicating that while mussel habitat supports a different grazer assemblage to bare rock, both assemblages are important in limiting fucoid recruitment. The risk of dislodgement was assessed and compared between fucoids growing on mussel shells and bare rock. In situ pull-tests showed that less force was required to detach large fertile thalli growing on mussel shells than those growing on the rock. Adhesion was generally broken between the mussel and the rock rather than between the holdfast and the mussel. These observations indicate that mussels provide an unstable substrate for mature fucoids. Overall results suggest that a negative effect of mussel-modified habitat on fucoids is profound in adults; but the effect is context-dependent in juveniles and can be positive at settlement. Results from a survey on population structure of fucoids across two shores showed that there were greater numbers of large fertile fucoids growing directly attached to rock than on mussel shells, while there was no difference for juvenile fucoids confirming the experimental results. Moreover thalli larger than 60 cm were found only on the rock but not on shells. This finding suggests that a mussel dominated habitat may have a significant impact on reproductive output in fucoid populations.

Larval development of the intertidal barnacles Chthamalus stellatus and Chthamalus montagui

Two recently-distinguished species of Chthamalus (Cirripedia) are found on rocky shores in the north-eastern Atlantic: C. stellatus predominant on islands and headlands and C. montagui more abundant in bays. Larvae of the two species were produced in laboratory cultures to describe and compare the morphology and to allow identification in plankton samples. Nauplius larvae of C. stellatus are up to 30% larger than those of C. montagui. Differences in setation are minor. The two species are easily distinguishable from the size and shape of the cephalic shield. Chthamalus stellatus has a subcircular shield with longer body processes in later stages while C. montagui is more ovoid. The former develop more slowly in culture than the latter. Chthamalus stellatus larvae in a culture at 19 °C reached stage VI in 16 d compared to 11 d for larvae of C. montagui at the same temperature. The morphology and longer development time of C. stellatus larvae suggests adaptation to a more oceanic lifestyle and wider dispersal to reach more fragmented habitats than larvae of C. montagui. --------------------------------------------------------------------------------

Recovery from TBT pollution in English Channel environments: A problem solved?

Following recognition of effects in the 1980s, tributyltin (TBT) has been monitored at sites in the English Channel to evaluate the prognosis for biota – spanning the introduction of restrictions on TBT use on small boats and the recent phase-out on the global fleet. We describe how persistence and impact of TBT in clams Scrobicularia plana has changed during this period in Southampton Water and Poole Harbour. TBT contamination (and loss) in water, sediment and clams reflects the abundance and type of vessel activity: half-times in sediment (up to 8y in Poole, 33y in Southampton) are longest near commercial shipping. Recovery of clam populations – slowest in TBT-contaminated deposits – provides a useful biological measure of legislative efficacy in estuaries. On rocky shores, recovery from imposex in Nucella lapillus is evident at many sites but, near ports, is prolonged by shipping impacts, including sediment legacy, for example, in the Fal.

Shore Shapers: Introducing children and the general public to biogeomorphological processes and geodiversity

Coastal processes and wildlife shape the coast into a variety of eye-catching and enticing landforms that attract people to marvel at, relax and enjoy coastal geomorphology. These landforms also influence biological communities by providing habitat and refuge. There are very few field guides to explain these processes to the general public and children. In contrast, there is a relative wealth of resources and organised activities introducing people to coastal wildlife, especially on rocky shores. These biological resources typically focus on the biology and climatic controls on their distribution, rather than how the biology interacts with its physical habitat. As an outcome of two recent rock coast biogeomorphology projects (detailed at: www.biogeomorph.org/coastal) a multi disciplinary team produced the first known guide to understanding how biogeomorphological processes help create coastal landforms. The ‘Shore Shapers’ guide (shoreshapers.org) is designed to: a. bring biotic geomorphic interactions (how animals, algae and microorganisms protect and shape rock) to life and b. introduce some of the geomorphological and geological controls on biogeomorphic processes and landform development. The guide provides scientific information in an accessible and interactive way – to help sustain children’s interest and extend their learning. We tested a draft version of the guide with children,the general public and volunteers on rocky shore rambles using social science techniques and present the findings, alongside initial results of an evaluation of a newer version of the guide and interactive workshops taking place throughout 2014.

The relationship between the distribution of genetically distinct inbred lines and upper lethal temperature in Lasaea rubra

Lasaea rubra is an inbreeding bivalve species, living at most heights on rocky shores. Freshly collected animals from different shore heights showed significantly different upper median lethal temperatures (MLTs), with upper shore animals having higher MLTs than lower shore specimens. Experiments with animals acclimated for at least one month to a single temperature (15°C) demonstrated that these differences in upper MLT were unaffected by thermal acclimation. Electrophoretic investigation showed that the differences in thermal response had a genetic basis. Homogeneous populations of the high-water inbred line (‘Inbred line A’) had a higher MLT than homogeneous populations of ‘Inbred line C’ which was found on the middle and lower shore. No differences were detected between the MLTs of separate populations of Inbred lines A or C. A third inbred line (‘Inbred line B’) was found on the middle shore, but no homogeneous populations were found. However, indirect evidence suggests that Inbred line B has a thermal response intermediate between those of Inbred lines A and C. Study of populations made up of mixtures of inbred lines confirmed the relationship between upper MLTs and genetic composition of the population.

Macroalgae contribute to the diet of Patella vulgata from contrasting conditions of latitude and wave exposure in the UK

Analysis of gut contents and stable isotope composition of intertidal limpets (Patella vulgata) showed a major contribution of macroalgae to their diet, along with microalgae and invertebrates. Specimens were collected in areas with limited access to attached macroalgae, suggesting a major dietary component of drift algae. Gut contents of 480 animals from 2 moderately wave-exposed and 2 sheltered rocky shores in each of 2 regions (western Scotland, 55-56°N; and southwest England, 50°N), were analysed in 2 years (n = 30 site-1 yr-1). The abundance of microalgae, macroalgae and invertebrates within the guts was quantified using categorical abundance scales. Gut content composition was compared among regions and wave exposure conditions, showing that the diet of P. vulgata changes with both wave exposure and latitude. Microalgae were most abundant in limpet gut contents in animals from southwestern sites, whilst leathery/corticated macroalgae were more prevalent and abundant in limpets from sheltered and northern sites. P. vulgata appears to have a more flexible diet than previously appreciated, and these keystone grazers consume not only microalgae, but also large quantities of macroalgae and small invertebrates. To date, limpet grazing studies have focussed on their role in controlling recruitment of macroalgae by feeding on microscopic propagules and germlings. Consumption of adult algae suggests that P. vulgata may also directly control the biomass of attached macroalgae on the shore, whilst consumption of drift algae indicates that the species may play important roles in coupling subtidal and intertidal production.

Macroalgae contribute to the diet of Patella vulgata from contrasting conditions of latitude and wave exposure in the UK

Analysis of gut contents and stable isotope composition of intertidal limpets (Patella vulgata) showed a major contribution of macroalgae to their diet, along with microalgae and invertebrates. Specimens were collected in areas with limited access to attached macroalgae, suggesting a major dietary component of drift algae. Gut contents of 480 animals from 2 moderately wave-exposed and 2 sheltered rocky shores in each of 2 regions (western Scotland, 55-56°N; and southwest England, 50°N), were analysed in 2 years (n = 30 site-1 yr-1). The abundance of microalgae, macroalgae and invertebrates within the guts was quantified using categorical abundance scales. Gut content composition was compared among regions and wave exposure conditions, showing that the diet of P. vulgata changes with both wave exposure and latitude. Microalgae were most abundant in limpet gut contents in animals from southwestern sites, whilst leathery/corticated macroalgae were more prevalent and abundant in limpets from sheltered and northern sites. P. vulgata appears to have a more flexible diet than previously appreciated, and these keystone grazers consume not only microalgae, but also large quantities of macroalgae and small invertebrates. To date, limpet grazing studies have focussed on their role in controlling recruitment of macroalgae by feeding on microscopic propagules and germlings. Consumption of adult algae suggests that P. vulgata may also directly control the biomass of attached macroalgae on the shore, whilst consumption of drift algae indicates that the species may play important roles in coupling subtidal and intertidal production.

Assessment of the potential impacts of coasteering on rocky intertidal habitats in Wales

The relatively new recreational pursuit of coasteering, which has developed in the St David's area of Pembrokeshire, appears to be expanding rapidly. The majority of local commercial recreation providers (outdoor pursuit centers etc.) now appear to offer this pursuit. The majority of the rocky coastlines where it takes place lie within Pembrokeshire Marine Special Area of Conservation (SAC), and are also Sites of Special Scientific Interest (SSSI). No assessment has yet been undertaken of coasteering's potential impact on the intertidal habitats. Therefore the Countryside Council for Wales (CCW) commissioned the Marine Life Information Network (MarLIN) to undertake a desk study of the likely environmental effects of coasteering on rocky intertidal habitats within the Pembrokeshire marine SAC. The desk study was based on a review of the available literature, and in particular the effects of trampling on rocky intertidal communities. Communities (as biotopes) within the Pembrokeshire marine SAC likely to be exposed to coasteering activities were identified from Phase I biotope data for the area, provided by CCW. Where possible, existing research by MarLIN into the intolerance, recoverability and sensitivity of the biotopes identified, was used to identify their potential vulnerability to trampling. The literature review revealed that: - foliose canopy forming algae (e.g. fucoids) were particularly intolerant and sensitive to trampling impacts; - trampling damaged erect coralline turfs, barnacles, and resulted in an increase in bare space; in some cases paths across the shore were visible; - on brown algae dominated shores, understorey algae could suffer due to increased desiccation but algal turf species, opportunists and gastropod grazers (e.g. limpets) could increase in abundance as an indirect effect of trampling, and that - trampling impacts resulted from physical contact and wear and were dependant on the intensity, duration, and frequency of trampling, and even the type of footwear used. A total of 19 intolerant rocky intertidal biotopes were identified as potentially vulnerable to trampling and hence coasteering within the Pembrokeshire marine SAC, of which six are of Welsh importance and eight are nationally rare or scarce. Trampling is a highly localized impact and it was not possible to identify biotopes, and hence communities, actually impacted by coasteering activities in the Pembrokeshire marine SAC. In addition, the majority of the literature addresses the impacts of trampling on wave sheltered or moderately exposed brown algal dominated shores, while coasteering occurs on more wave exposed, steeply inclined shores. Therefore, direct survey of the routes used by coasteering groups within the Pembrokeshire marine SAC is required to identify the intensity, duration and frequency of trampling impact, together with the communities impacted. Given the paucity of data concerning trampling effects in the rocky intertidal in the UK, a survey of the impacts of coasteering would provide an opportunity to examine the effects of trampling and visitor use in steep rocky, wave exposed shores. The report recognizes the potential to engage coasteerers in contributing to the development of strategies for minimizing adverse impacts, recording impacts and collecting information of use in identifying climate change and the occurrence of non-native species.

Using historical data to detect temporal changes in the abundances of intertidal species on Irish shores

An historical data set, collected in 1958 by Southward and Crisp, was used as a baseline for detecting change in the abundances of species in the rocky intertidal of Ireland. In 2003, the abundances of each of 27 species was assessed using the same methodologies (ACFOR [which stands for the categories: abundant, common, frequent, occasional and rare] abundance scales) at 63 shores examined in the historical study. Comparison of the ACFOR data over a 45-year period, between the historical survey and re-survey, showed statistically significant changes in the abundances of 12 of the 27 species examined. Two species (one classed as northern and one introduced) increased significantly in abundance while ten species (five classed as northern, one classed as southern and four broadly distributed) decreased in abundance. The possible reasons for the changes in species abundances were assessed not only in the context of anthropogenic effects, such as climate change and commercial exploitation, but also of operator error. The error or differences recorded among operators (i.e. research scientists) when assessing species abundance using ACFOR categories was quantified on four shores. Significant change detected in three of the 12 species fell within the margin of operator error. This effect of operator may have also contributed to the results of no change in the other 15 species between the two census periods. It was not possible to determine the effect of operator on our results, which can increase the occurrence of a false positive (Type 1) or of a false negative (Type 2) outcome

Facing the future: the importance of substratum features for ecological engineering of artificial habitats in the rocky intertidal

Coastal defences are proliferating in response to climate change, leading to the creation of more vertical substrata. Efforts are being made to mitigate their impacts and create novel habitats to promote biodiversity. Little is known about the effect of aspect (i.e. north–south directionality) and inclination on intertidal biodiversity in artificial habitats. Artificial and natural habitats were compared to assess the role of aspect and substratum inclination in determining patterns of biodiversity at two tidal heights (high and mid). We also compared grazing activity between north- and south-facing surfaces in natural habitats to examine the potential for differential grazing pressure to affect community structure and functioning. Results were variable but some clear patterns emerged. Inclination had no effect on biodiversity or abundance. There was a general trend towards greater taxon richness and abundance on north-facing than south-facing substrata in natural and artificial habitats. On natural shores, the abundance and grazing activity of ‘southern’ limpets (i.e. Patella depressa) was greater on south-facing than north-facing substrata, with possible implications for further range-expansion. These results highlight the importance of incorporating shaded habitats in the construction of artificial habitats. These habitats may represent an important refuge from grazing pressure and thermal and desiccation stress in a warming climate.