Eve Caroline Southward (1930–2023) was a multi-talented scientist, motivated by her curiosity and love of nature. Since she was never paid as a scientist, Eve was an amateur, in the best sense of the word. She was highly proficient at transmission electron microscopy and made lasting contributions to polychaete taxonomy, morphology and ecology. Eve was internationally respected, especially for her studies on the Siboglinidae, mouthless and gutless tubeworms (formerly called Pogonophora) that are found worldwide in the deep-sea. She described how the siboglinids obtained nutrition from symbiotic, sulphur-oxidising bacteria and described similar symbiotic relationships in several bivalve species. Eve wrote over 140 scientific publications and described 56 new benthic species, 47 being mouthless and gutless ‘pogonophores’. Eve assisted her husband Alan Southward in starting broad-scale intertidal surveys around the British Isles and Northwest Europe. These surveys formed the foundation for the time-series, later continued by others, that allowed assessments of the influence of climatic fluctuations, using intertidal rocky shore biota as indicators. Eve contributed, with Alan, to what became a 50-year study describing the long-term effects on intertidal communities of the oil pollution and excessive dispersant use resulting from the Torrey Canyon oil spill in 1967. Eve also co-wrote the Linnaean Society Synopsis on Echinoderms of the British Isles and helped complete unpublished work by Alan Southward and others on barnacle taxonomy.

The rocky intertidal zone of sheltered shores in Breiðafjörður, north-west Iceland is dominated by monospecific stands of canopy-forming brown algae Ascophyllum nodosum, which provide habitat for mobile organisms and has been subjected to long-standing rotational harvesting. We investigated the assemblage composition of little-studied mobile brachyuran crabs in this area, to track distributional shifts in a native species responding to climate change and extent of occupancy of the intertidal by a primarily subtidal invasive non-native species. Potential interactive effects of seaweed harvesting were explored. Breiðafjörður was compared with two reference sites in Faxaflói, south-west Iceland. The study revealed clear poleward expansion of the native European green crab Carcinus maenas in the region, displacing the native spider crab Hyas araneus particularly at mid-shore levels. The invasive non-native Atlantic rock crab Cancer irroratus had negligible occupancy in the intertidal zone, indicating limited effects on the intertidal crab assemblage, composition, and abundance. The current harvesting regime of A. nodosum in Breiðafjörður did not affect the composition and abundance of the brachyuran crab assemblage in the rocky intertidal zone. H. araneus is likely being squeezed by displacement subtidally by C. irroratus, and intertidally by C. maenas. Overall, we provide insights into the potential interactions between climate change, invasive species, and human activities in the rocky intertidal zone.

In shell-secreting molluscs, age and growth rate of individuals and hence their performance can normally be measured using growth lines that are deposited in the shell throughout their lives. An annual periodicity of growth line formation of the warm-water limpet Patella depressa was established using marked and recaptured individuals from north Wales, UK. Length at age from suitably prepared shell sections was determined in limpets from non-range-edge populations and at two range edges, where different demographic attributes have been recorded. Individuals collected from their poleward range-edge in north Wales were older when compared with individuals at their range-edge in southern England. Shells collected from southern England were characterized by rapid growth with most individuals reaching >30 mm in maximum length by the fourth or fifth year, contrasting with those from north Wales, where most shells only reached this size at 7–10 years of age. Von Bertalanffy growth coefficients (K-values) were negatively related to P. depressa density, showing faster growth in lower total densities of both P. depressa and Patella vulgata combined. Higher intra-specific effects on K-values were found in P. depressa compared with its congener P. vulgata, with stronger effects in north Wales than in southern England. These results confirm differences in population patterns and individual traits between the two leading edges of P. depressa. Understanding annual growth in P. depressa over large scales could help to disentangle the processes determining differences in shell growth and age structure seen at the two range edges of this limpet species.

Rocky shore ecology has been studied for a long time, starting with qualitative descriptions and becoming more quantitative and experimental over time. Some of the earliest manipulative experimental ecological studies were undertaken on rocky shores. Many, over time, have made considerable contributions to ecological theory, especially highlighting the importance of biological interactions at the community level. The suitability of rocky shores as convenient test systems for ecological experimentation is outlined. Here we consider contributions from rocky shores to the emerging concepts of supply-side ecology, the roles of competition, predation and grazing, disturbance and succession and positive interactions in structuring communities along environmental gradients. We then address alternative stable states, relationships between biodiversity and ecosystem functioning, and bottom-up and top-down control of ecosystems. We briefly consider the feedback and synergies between ecological concepts and experimental work on rocky shores, whilst still emphasizing the traditional values of marine natural history upheld in JMBA since its first publication. The importance of rigorous experimental designs championed by Underwood and co-workers is emphasized. Recent progress taking advantage of new technologies and emerging approaches is considered. We illustrate how experimental studies have shown the importance of biological interactions in modulating species and assemblage-level responses to climate change and informed conservation and management of coastal ecosystems.

Limpets and barnacles are important components of intertidal assemblages worldwide. This study examines the effects of barnacles on the foraging behaviour of the limpet Patella vulgata, which is the main algal grazer in the North-west Atlantic. The behaviour of limpets on a vertical seawall on the Isle of Man (UK) was investigated using autonomous radio-telemetry, comparing their activity patterns on plots characterized by dense barnacle cover and plots from which the barnacles had been removed. Limpet behaviour was investigated at mid-shore level, but two different elevations were considered. This experiment revealed a significant effect of barnacle cover on the activity of P. vulgata. Limpets on smooth surfaces spent a greater proportion of total time active than did limpets on barnacles. Movement activity was also greater in areas that were lower down in the tidal range. In general, limpets were either predominantly active during diurnal high or nocturnal low tides and always avoided nocturnal high tides. Individuals on barnacles at the higher elevation concentrated their activity during nocturnal low water. All the other groups of limpets (smooth surfaces on the upper level and all individuals on the lower shore) had more excursions centred around daylight hours with an equal distribution of activity between periods of low and high water. Inter-individual variability was, however, pronounced.

Cosmopolitan habitat-forming taxa of algae such as the genus Corallina provide an opportunity to compare patterns of biodiversity over wide geographic scales. Nematode assemblages inhabiting Corallina turves were compared between the south coasts of the British Isles and South Korea. A fully nested design was used with three regions in each country, two shores in each region and replicate samples taken from three patches on each shore to compare differences in the taxonomic and biological trait composition of nematode assemblages across scales. A biological traits approach, based on functional diversity of nematodes, was used to make comparisons between countries, among regions, between shores and among patches. The taxonomic and biological trait compositions of nematode assemblages were significantly different across all spatial scales (patches, shores, regions and countries). There is greater variation amongst nematode assemblages at the scale of shore than at other spatial scales. Nematode assemblage structure and functional traits are influenced by the local environmental factors on each shore including sea-surface temperature, the amount of sediment trapped in Corallina spp. and tidal range. The sea-surface temperature and the amount of sediment trapped in Corallina spp. were the predominant factors determining nematode abundance and composition of assemblages and their functional diversity.

Hermaphroditism is thought to be an advantageous strategy common in marine molluscs that exhibit simultaneous, sequential or alternating hermaphroditism. Several species of patellid limpets have previously been shown to be protandrous hermaphrodites. The present study aimed to confirm whether this phenomenon occurs in Patella piperata. Transitional forms of simultaneous protandrous hermaphroditism were found in intermediate size classes of P. piperata, in Madeira (North-eastern Atlantic). Sequential hermaphroditism was confirmed after histological analysis. The overall sex-ratio was biased towards females but approached similar proportions in the larger size classes. Analysis of size at sex change showed that at a shell length of 36 mm 50% of the population probably have changed sex. The results reported confirm the occurrence of sequential hermaphroditism. These findings are of utmost importance to the understanding of the reproductive biology of this species with direct effect on management and conservation of this traditionally harvested limpet.

Canopy-forming fucoid algae have an important role as ecosystem engineers on rocky intertidal shores, where they increase the abundance of species otherwise limited by exposure during low tide. The facilitative relationship between Ascophyllum nodosum and associated organisms was explored using a frond breakage experiment (100%, 50%, 25%, 0% intact-frond treatments) in southern England, to assess the consequences of disturbance. Understorey substratum temperature was on average 3°C higher in 0% and 25% intact-frond treatments than in plots with 50% and 100% intact fronds. Light (as PAR during low tide) doubled in 0% intact-frond treatments in comparison to other treatments (which had similar light levels). Mobile invertebrate species richness declined by on average 1 species per m2 in the treatments with only 25% and 0% intact fronds, and the abundance of Littorina obtusata declined by 2.4–4.2 individuals per m2 in the treatments with 25 and 0% intact fronds. Sessile taxa, including Osmundea pinnatifida and encrusting coralline algae, declined by half on average in the 0% intact-frond treatment. These results suggest that the ability of Ascophyllum to mediate environmental conditions to the understorey is the mechanism responsible for species distributed in the understorey (autogenic ecosystem engineering). The results of this study imply that a pulse disturbance resulting in a 50% breakage of Ascophyllum fronds significantly increases temperature and decreases the abundance of mobile invertebrates usually associated with Ascophyllum. Sessile taxa associated with Ascophyllum can, however, withstand disturbances down to 25% intact Ascophyllum fronds.