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.

Antagonism among bacteria is widespread and plays an important role in structuring communities. Inhibitory compounds can confer competitive advantage, but energetic trade-offs can result in non-transitive (i.e. ‘rock-paper-scissors’) interactions, ultimately allowing co-existence and community stability. Competition in sedimentary habitats is especially keen given high densities and attachment to inorganic particles. Because measuring trade-offs between bacterial species is challenging, much of our understanding of competitive interactions is based on theoretical modelling and simplified in vitro experiments. Our objectives were to determine (1) if interference competition occurs in microcosms mimicking in situ conditions; (2) whether the presence of sediment influences antagonistic interactions; and (3) if more complex assemblages alleviate or synergize interactions. Four sedimentary isolates, including antibiotic-producing, resistant and susceptible strains were incubated in porewater microcosms in 1-, 2- and 3-species combinations, both with and without natural sediments. Microcosms were sampled over 72 h to generate growth curves using quantitative PCR. Multiple growth attributes (growth rate, maximum density, lag time) were used to assess effects of treatment (species combinations) and environment (sediment vs porewater alone). Antimicrobial producers were more effective at inhibiting target species in microcosms that included sediment, in agreement with theory. We observed growth inhibition by antimicrobial-producing bacteria in both 2- and 3-species microcosms. However, the expected protection of sensitive bacterial strains by resistant strains was observed in only one (of four) 3-species combinations, thus the ‘rock-paper-scissors’ prediction was not fully supported. These results reinforce the notion that interspecies interactions are context-dependent, reliant on environmental conditions and the species involved.

The Antarctic Peninsula is undergoing rapid change due to global warming, including air and water temperature increases. Fauna inhabiting the intertidal zone are particularly exposed to warming impacts, as they are subjected to high variations in both terrestrial and marine environmental settings. This study aimed to assess intertidal macrofaunal and meiofaunal biodiversity, tidal height-related assemblage structural patterns and their responses to variability in environmental parameters on King George Island. A total of 39 macrofaunal taxa were identified, with polychaetes and amphipods being the most diverse groups and gastropods, amphipods and bivalves being the most abundant. In the case of meiofauna, 16 taxa were found, mainly nematodes, copepod nauplii and harpacticoids. There was a significant decrease in the number of species, abundance and biomass for both macrofauna and meiofauna with increasing tidal height. Our investigation documented highly diverse and abundant fauna in the Antarctic intertidal zone. With its thriving life, it could serve as a perfect model system for detecting climate change impacts on local biodiversity. Therefore, we propose the Antarctic intertidal zone as a suitable habitat for monitoring these changes.

The Order Suberitida is defined as a group of marine sponges without an obvious cortex, a skeleton devoid of microscleres, and with a deletion of a small loop of 15 base pairs in the secondary structure of the 28S rDNA as a molecular synapomorphy. Suberitida comprises three families and 26 genera distributed worldwide, but mostly in temperate and polar waters. Twenty species were reported along the entire Brazilian coast, and although the north-eastern coast of Brazil seems to harbour a rich sponge fauna, our current knowledge is concentrated along the south-eastern Atlantic coast. A survey was implemented along the northern coast of Brazil, and the collection allowed the identification of six species belonging to the Order Suberitida. Two of them are considered new to science: Suberites purpura sp. nov., Hymeniacidon upaonassu sp. nov., and four, Halichondria (Halichondria) marianae Santos, Nascimento & Pinheiro, 2018, Halichondria (H.) melanadocia de Laubenfels, 1936, Suberites aurantiacus (Duchassaing & Michelotti, 1864), and Terpios fugax Duchassaing & Michelotti, 1864, are re-described. Taxonomic comparisons are made for Tropical Western Atlantic species and type species of the four genera. Finally, an identification key for the Western Atlantic Suberites species is provided.

The establishment of permanent animal communities on land was a defining event in the history of evolution, and one for which the ichnofauna and facies of the Tumblagooda Sandstone of Western Australia have been considered an archetypal case study. However, terrestrialization can only be understood from the rock record with conclusive sedimentological evidence for non-marine deposition, and original fieldwork on the formation shows that a marine influence was pervasive throughout all trace fossil-bearing strata. Four distinct facies associations are described, deposited in fluvial, tidal and estuarine settings. Here we explain the controversies surrounding the age and depositional environment of the Tumblagooda Sandstone, many of which have arisen due to the challenges in distinguishing marine from non-marine depositional settings in lower Palaeozoic successions. We clarify the terminological inconsistency that has hindered such determination, and demonstrate how palaeoenvironmental explanations can be expanded out from unambiguously indicative sedimentary structures. The Tumblagooda Sandstone provides a unique insight into an early Palaeozoic ichnofauna that was strongly partitioned by patchy resource distribution in a littoral setting. The influence of outcrop style and quality is accounted for to contextualize this ichnofauna, revealing six distinct low-disparity groups of trace fossil associations, each related to a different sub-environment within the high-ichnodisparity broad depositional setting. The formation is compared with contemporaneous ichnofaunas to examine its continued significance to understanding the terrestrialization process. Despite not recording permanent non-marine communities, the Tumblagooda Sandstone provides a detailed picture of the realm left behind by the first invertebrate pioneers of terrestrialization.

Macroalgae play important ecological roles, including as hosts for a wide range of epifauna. However, the diversity relationships between macroalgae and epifauna are poorly understood for most tropical host species and algal morphologies. This study aims to characterize and analyse the diversity of invertebrates present amongst macroalgae with three distinct morphologies (three-dimensional, filamentous and foliose) across different tropical intertidal sites in Singapore. Morphological and DNA barcoding tools were employed for epifaunal species identification, and ordination statistics and multiple linear regression were used to test the effects of algal morphology, species and site on community structure and diversity of epiphytic invertebrates. Overall, epifaunal communities were distinct among sites and algal morphologies, and diversity was affected significantly by algal morphology. In particular, filamentous macroalgae hosted the highest abundance of epifauna dominated mainly by amphipods, which were able to take advantage of the high surface area to volume ratio in filamentous algal mats as a consequence of their thinner forms. Foliose species showed a significantly negative effect on invertebrate diversity. Our findings highlight the diverse associations between intertidal macroalgae and invertebrates with high turnover between algal morphology and sites that contribute to the high biodiversity of tropical shores. Future studies should consider the effects of the host habitat, seasonality and more algal species on epifaunal 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.

Where two species occupy the same habitat and similar niches, competition is likely to drive small-scale spatial niche separation or resource partitioning that may not be immediately apparent. A stable isotope approach was used to investigate potential trophic niche separation between co-existing rocky shore crabs in the North-West (NW) Arabian Gulf. Leptodius exaratus and Pilumnopeus convexus which occupy similar shore height on the same rocky intertidal habitats. We also investigated conspecific differences between males vs females and adults vs juveniles. δ15N results indicated that adults of both species occupy a high trophic level in the rocky shore community, suggesting similar functional roles and potential for competition for food resources, while significant differences in δ13C values indicated differences in dietary sources between the two species, and also changes in diet between juveniles and adults in both species. MixSIAR analysis of δ15N and δ13C data confirmed field observations that both species are generalist omnivores, with potential for direct competition including adult predation on juveniles, including conspecifics. Differentiation in isotopic niches (SIBER analysis) was mainly driven by the significant differences in δ13C values, suggesting that co-existence of the two crab species is at least in part mediated by trophic niche separation or dietary resource partitioning, with some (unquantified) potential for spatial resource partitioning at the microhabitat level.

The spoon-billed sandpiper Calidris pygmaea, a migratory Arctic-breeding shorebird, is one of the rarest birds and its population has declined since the 1970s. We surveyed its most important known wintering area in the Upper Gulf of Mottama in Myanmar to estimate recent (2009–2016) changes in its numbers there. The total number of small shorebirds present in the Upper Gulf was counted and the proportion of them that were spoon-billed sandpipers was estimated from sample scans. These two quantities were multiplied together to give the estimated number of spoon-billed sandpipers in each of 4 years. Total numbers of combined small shorebird species tripled from 21,000 to 63,000 between 2009 and 2016, coincident with efforts to reduce hunting pressure on waterbirds. However, the proportion of small shorebirds that were spoon-billed sandpipers declined and their estimated absolute numbers fell by about half, from 244 to 112 individuals. It is probable that loss of intertidal habitat and shorebird hunting elsewhere on the migration route of the spoon-billed sandpipers wintering at Mottama is causing a continued decline, although this is occurring at a less rapid rate than that recorded from Arctic Russia before 2010. The number of spoon-billed sandpipers wintering on the Upper Gulf of Mottama remains the highest single-site total for this species from any known wintering site. Preventing resurgence of illegal shorebird hunting and ensuring long-term protection of the intertidal feeding habitats and roost sites in the Gulf are high priorities if extinction of this species is to be averted.

We describe and illustrate a new genus and species of sea anemone from the intertidal in the Gulf of California (Mexico). Tenactis gen. nov. is characterized by mesenteries decamerously arranged, a single pair of directives attached to a single and strong siphonoglyph, small conchula, endodermal marginal sphincter muscle, pedal disc with basilar muscles, cinclides in pedal disc and proximal column, column with verrucae and pseudoacrorhagi, and a cnidom with p-mastigophores A2 and p-mastigophores B1 and B2a. Tenactis riosmenai gen. et sp. nov. is characterized by having a column not divisible into regions with longitudinal rows of verrucae distally and pseudoacrorhagi forming spherical structures at the tip, only 10 pairs of perfect mesenteries, diffuse endodermal marginal sphincter muscle and p-mastigophores A2 and p-mastigophores B1 and B2a in the filaments. Because of the decamerous arrangement of the mesenteries Tenactis gen. nov. most closely resembles some members of Actiniidae, Haloclavidae, Minyadidae, Oractiidae and Limnactinidae (Actinioidea), Halcampidae (Metridioidea) and Halcuriidae (Actinernoidea). However, the combination of having a decamerous arrangement of mesenteries, single siphonoglyph, conchula and cinclides indicate this new genus belongs to Haloclavidae. Additionally, the presence of a diffuse endodermal marginal sphincter muscle, basilar muscles, a single pair of directives, column with longitudinal rows of verrucae distally and pseudoacrorhagi, p-mastigophores A2, and p-mastigophores B1 and B2a in the filaments distinguish Tenactis riosmenai gen. et sp. nov. within Haloclavidae. This is the first record of the family Haloclavidae for the Gulf of California.

Gnathiid isopods are one of the most common fish ectoparasites, and are found in both temperate and tropical oceans. On coral reefs, gnathiids are most active at dusk and dawn, and contribute significantly to trophic dynamics, as they are a prey resource for cleaner fish and parasitize numerous fishes. Gnathiids also inhabit temperate intertidal waters, but their activity patterns and contribution to intertidal trophic dynamics remain unstudied. To provide the first ecological data on temperate intertidal gnathiid activity patterns, 172 gnathiid-free Clinus superciliosus were set in an intertidal system in Tsitsikamma National Park, South Africa, during early morning, morning, afternoon, early evening, and evening, high and low tide, and within the inter- and infra-tidal zone to examine gnathiid infestation levels. After exposure, gnathiids from each fish were identified to the species level, counted, and their developmental stage was recorded. All gnathiids were identified as Gnathiia africana. On average, 1 ± 5SD gnathiids were collected from each fish, and the majority of gnathiids collected were stage 1. Significantly more gnathiids were collected during morning and afternoon compared with all other time periods. The number of gnathiids collected was not influenced by the fish's exposure to high or low tide, or placement within the tide zone. Although G. africana is free from cleaner fish predation because cleaner fish do not reside in temperate intertidal habitat, G. africana abundance is surprisingly small. Future studies should examine what regulates G. africana population size and the role they play in temperate intertidal food webs.

Six benthic biotic indices (Shannon–Wiener H′log2, W-statistics, BOPA, BENTIX, AMBI and M-AMBI), based on different ecological principles, were applied to assess the health of variously disturbed tropical intertidal habitats of the Port Blair coastline. A total of 243 replicate samples were collected during the dry period (January, February and March) of 2014–2016. Temperature, salinity, dissolved oxygen, sediment pH, total organic matter content and texture were analysed. A high mean abundance of opportunistic species (Orbinia sp. 748, Capitella singularis 237 and Armandia sp. 114 ind. m−2) was observed at Phoenix Bay, a gradual decline in diversity at Junglighat and a comparatively high diversity and moderate biomass at Wandoor, reflecting a human pressure gradient. Results showed an annual decline of benthic quality from 2014 to 2016 (good to moderate). Overall BOPA failed to distinguish the magnitude of disturbances, while the rest of the indices classified the benthic quality from undisturbed/high (WD), slightly disturbed/good (JG), to moderately disturbed/moderate to poor (PB). The subjective analysis demonstrated that the urban centres corresponded to disturbed benthic communities of dominant first and second order opportunistic species, while sensitive (EGI) and indifferent (EGII) were associated with the least disturbed or undisturbed site. The study successfully demonstrated the performance of temperate indices in intertidal habitats against the mild organic enrichment. However, for an effective assessment, setting natural reference conditions and sampling in stable dry periods (strong seasonality in tropics) is desirable. In order to test the performance of biotic indices, a long-term monitoring approach of taking abiotic and biotic descriptors into account is recommended.

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.

This study tested for social monogamy in Alpheus brasileiro Anker, 2012. We also analysed egg production in this species. Sampling was conducted bimonthly from March 2013 to January 2014, and specimens were collected manually in the intertidal estuarine zone of Cananéia, São Paulo, Brazil. A total of 186 specimens (92 males and 94 females) were captured, and ~46% of them were found living in heterosexual pairs. The carapace length (CL mm) of paired males and females was positively correlated. We analysed 35 females carrying eggs underneath the abdomen: 28 of these females had eggs in stage I of development, four had eggs in stage II, and three had eggs in stage III. The average (±SD) mean fecundity for females carrying early (stage I) eggs was 149 (±93) eggs. Egg volume differed significantly among developmental stages, with a 35.4% increase in egg volume between stages I and III. The egg volume and size (CL) of females correlated positively. A positive correlation between the number of eggs and carapace length was found, with a proportional increase in the number of eggs according to the size of females. Our results suggest that the studied population of A. brasileiro is socially monogamous and provide the first insights into egg production in this species.

We resolve taxonomic confusion regarding brooding sea anemones in the genus Epiactis Verrill 1869a in the North Pacific Ocean based on newly collected material from Hokkaido (Japan), Haida Gwaii (British Columbia, Canada), and Kodiak and Adak Islands (Alaska, USA), and museum specimens collected from the Kurile Islands (Russia), Alaska, British Columbia, Oregon (USA), and California (USA). We find that the internally brooding individuals identified by Hand & Dunn (1974) as Cnidopus ritteri (Torrey, 1902) and placed in the genus Epiactis by Fautin & Chia (1986) belong to a new species which we describe and name Epiactis handi sp. nov. Epiactis handi and E. ritteri can be differentiated by morphological and behavioural features including ornamentation and structure of the column and mode of brooding offspring. To highlight and clarify these differences, we redescribe E. ritteri based on specimens from Alaska. We provide the first account of external brooding in E. ritteri, which necessitates a clarification of the differences between E. ritteri and another externally brooding species from the North Pacific, E. japonica Verrill, 1869b. Epiactis ritteri and E. japonica differ in sex allocation, ornamentation of the column and details of external brooding: members of E. ritteri are gonochoric with a smooth column and brood groove which tightly closes, whereas those of E. japonica are hermaphroditic and have mid-column spherules.