Between-taxon matching of common and rare species richness patterns

Aim: Our primary aim is to understand how assemblages of rare (restricted range) and common (widespread) species are correlated with each other among different taxa. We tested the proposition that marine species richness patterns of rare and common species differ, both within a taxon in their contribution to the richness pattern of the full assemblage and among taxa in the strength of their correlations with each other. Location The UK intertidal zone. Methods: We used high-resolution marine datasets for UK intertidal macroalgae, molluscs and crustaceans each with more than 400 species. We estimated the relative contribution of rare and common species, treating rarity and commonness as a continuous spectrum, to spatial patterns in richness using spatial crosscorrelations. Correlation strength and significance was estimated both within and between taxa. Results: Common species drove richness patterns within taxa, but rare species contributed more when species were placed on an equal footing via scaling by binomial variance. Between taxa, relatively small sub-assemblages (fewer than 60 species) of common species produced the maximum correlation with each other, regardless of taxon pairing. Cross-correlations between rare species were generally weak, with maximum correlation occurring between small sub-assemblages in only one case. Cross-correlations between common and rare species of different taxa were consistently weak or absent. Main conclusions: Common species in the three marine assemblages were congruent in their richness patterns, but rare species were generally not. The contrast between the stronger correlations among common species and the weak or absent correlations among rare species indicates a decoupling of the processes driving common and rare species richness patterns. The internal structure of richness patterns of these marine taxa is similar to that observed for terrestrial taxa.

Seasonal variations in nitrate reductase activity and internal N pools in intertidal brown algae are correlated with ambient nitrate concentrations.

Nitrogen metabolism was examined in the intertidal seaweeds Fucus vesiculosus, Fucus serratus, Fucus spiralis and Laminaria digitata in a temperate Irish sea lough. Internal NO3- storage, total N content and nitrate reductase activity (NRA) were most affected by ambient NO3-, with highest values in winter, when ambient NO3- was maximum, and declined with NO3- during summer. In all species, NRA was six times higher in winter than in summer, and was markedly higher in Fucus species (e.g. 256 ± 33 nmol NO3- min1 g1 in F. vesiculosus versus 55 ± 17 nmol NO3- min1 g1 in L. digitata). Temperature and light were less important factors for N metabolism, but influenced in situ photosynthesis and respiration rates. NO3- assimilating capacity (calculated from NRA) exceeded N demand (calculated from net photosynthesis rates and C : N ratios) by a factor of 0.7–50.0, yet seaweeds stored significant NO3- (up to 40–86 µmol g1). C : N ratio also increased with height in the intertidal zone (lowest in L. digitata and highest in F. spiralis), indicating that tidal emersion also significantly constrained N metabolism. These results suggest that, in contrast to the tight relationship between N and C metabolism in many microalgae, N and C metabolism could be uncoupled in marine macroalgae, which might be an important adaptation to the intertidal environment.

Coastal upwelling drives intertidal assemblage structure and trophic ecology

ecosystems. Coastal oceanic upwelling, for example, has been associated with elevatedbiomass and abundance patterns of certain functional groups, e.g., corticated macroalgae.In the upwelling system of Northern Chile, we examined measures of intertidal macrobenthiccomposition, structure and trophic ecology across eighteen shores varying in theirproximity to two coastal upwelling centres, in a hierarchical sampling design (spatial scalesof >1 and >10 km). The influence of coastal upwelling on intertidal communities was confirmedby the stable isotope values (δ13C and δ15N) of consumers, including a dominantsuspension feeder, grazers, and their putative resources of POM, epilithic biofilm, andmacroalgae. We highlight the utility of muscle δ15N from the suspension feeding mussel,Perumytilus purpuratus, as a proxy for upwelling, supported by satellite data and previousstudies. Where possible, we used corrections for broader-scale trends, spatial autocorrelation,ontogenetic dietary shifts and spatial baseline isotopic variation prior to analysis. Ourresults showed macroalgal assemblage composition, and benthic consumer assemblagestructure, varied significantly with the intertidal influence of coastal upwelling, especiallycontrasting bays and coastal headlands. Coastal topography also separated differences inconsumer resource use. This suggested that coastal upwelling, itself driven by coastlinetopography, influences intertidal communities by advecting nearshore phytoplankton populationsoffshore and cooling coastal water temperatures. We recommend the isotopic valuesof benthic organisms, specifically long-lived suspension feeders, as in situ alternativesto offshore measurements of upwelling influence

Nitrate reductase activity in macroalgae and its vertical distribution in macroalgal epiphytes of seagrasses.

Macroalgal epiphytes within seagrass meadows make a significant contribution to total primary production by assimilating water column N and transferring organic N to sediments. Assimilation of NO3 – requires nitrate reductase (NR, EC 1.6.6.1); NR activity represents the capacity for NO3 – assimilation. An optimised in vitro assay for determining NR activity in algal extracts was applied to a wide range of macroalgae and detected NR activity in all 22 species tested with activity 2 to 290 nmolNO3 – min–1 g–1 frozen thallus. With liquid-N2 freezing immediately after sample collection, this method was practical for estimating NR activity in field samples. Vertical distribution of NR activity in macroalgal epiphytes was compared in contrasting Posidonia sinuosa and Amphibolis antarctica seagrass meadows. Epiphytes on P. sinuosa had higher mass-specific NR activity than those on A. antarctica. In P. sinuosa canopies, NR activity increased with distance from the sediment surface and was negatively correlated with [NH4 +] in the water but uncorrelated with [NO3 –]. This supported the hypothesis that NH4 + released from the sediment suppresses NR in epiphytic algae. In contrast, the vertical variation in NR activity in macroalgae on A. antarctica was not statistically significant although there was a weak correlation with [NO3 –], which increased with distance from the sediment. Estimated capacities for NO3 – assimilation in macroalgae epiphytic on seagrasses during summer (24 and 46 mmolN m–2 d–1 for P. sinuosa and A. antarctica, respectively) were more than twice the estimated N assimilation rates in similar seagrasses. When the estimates were based on annual average epiphyte loads for seagrass meadows in other locations, they were comparable to those of seagrasses. We conclude that epiphytic algae represent a potentially important sink for water-column nitrate within seagrass meadows.

The effects of dispersal mode on the spatial distribution patterns of intertidal molluscs.

1. As many species of marine benthic invertebrates have a limited capacity for movement as adults, dispersal mode is often considered as a determinant of geographical ranges, genetic structure and evolutionary history. Species that reproduce without a larval stage can only disperse by floating or rafting. It is proposed that the colonization processes associated with such direct developing species result in spatial distributions that show relatively greater fine scale patchiness than the distributions of species with a larval dispersal stage. This hypothesis was tested by collecting molluscs at different spatial scales in the Isle of Man. 2. Spatial distribution patterns supported the predictions based on dispersal mode. Estimated variance components for species with larval dispersal suggested that the majority of the spatial variation was associated with variation between shores. In comparison, there was relatively more variability within shores for abundance counts of species with direct development. 3. Multivariate analyses reflected the univariate results. An assemblage of direct developers provided a better discrimination between sites (100 m separation) but the group of species with larval dispersal gave a clearer separation of shores (separated by several km). 4. The fine scale spatial structure of direct developing species was reflected in higher average species diversity within quadrats. Species richness also reflected dispersal mode, with a higher fraction of the regional species pool present for direct developers in comparison to species with larval dispersal. This may reflect the improved local persistence of taxa that avoid the larval dispersal stage.

Non-indigenous marine macroalgae in native communities: a case study in the British Isles

It has traditionally been considered that areas with high natural species richness are likely to be more resistant to non-indigenous species than those with lower numbers of species. However, this theory has been the subject of a debate over the last decade, since some studies have shown the opposite trend. In the present study, a macroalgal survey was carried out at 24 localities in Northern Ireland and southern England, using a quadrat approach in the lower littoral. The two opposing hypotheses were tested (negative versus positive relationship between native and non-indigenous species richness) in this marine environment. The effect of the presence of 'impacts', potential sources of disturbance and species introduction (e.g. marina, harbour or aquaculture), was also tested. A positive relationship was found between the number of non-indigenous species and the native species richness at the three different scales tested (quadrats, sites and localities). At no scale did a richer native assemblage appear to restrict the establishment of introduced species. The analyses revealed greater species richness and different community composition, as well as more non-indigenous species, in southern England relative to Northern Ireland. The presence of the considered impacts had an effect on the community composition and species richness in southern England but not in Northern Ireland. Such impacts had no effect on the non-indigenous species richness in either area.

Do mussel patches provide a refuge for algae from grazing gastropods?

On rocky shores, cover of macroalgae is often greater growing epibiotically on mussels compared to algae growing directly attached to rock. A survey of two shores on the east coast of Ireland confirmed that mussel beds contained greater percentage algal cover and more diverse algal assemblages compared to those on rock. The reasons for this difference are not clear. It has been suggested that mussel beds provide a refuge for algae from grazing gastropods. Surprisingly, we found no evidence to support this. Using wax discs, gastropod grazing patterns were found to be similar within the mussel beds as on rock. The mussel beds do not appear to provide a refuge for algae from grazing activity at this scale and we suggest other possible mechanisms for the prevalence of epibiotic algal cover on mussels. Intertidal grazers may in fact affect the epibiotic algae on mussels and thereby affect indirectly the persistence of mussel beds.

Intertidal molluscan and algal species richness around the UK coast

Geographically referenced databases of species records are becoming increasingly available. Doubts over the heterogeneous quality of the underlying data may restrict analyses of such collated databases. We partitioned the spatial variation in species richness of littoral algae and molluscs from the UK National Biodiversity Network database into a smoothed mesoscale component and a local component. Trend surface analysis (TSA) was used to define the mesoscale patterns of species richness, leaving a local residual component that lacked spatial autocorrelation. The analysis was based on 10 km grid squares with 115035 records of littoral algae (729 species) and 66879 records of littoral molluscs (569 species). The TSA identified variation in algal and molluscan species richness with a characteristic length scale of approximately 120 km. Locations of the most species-rich grid squares were consistent with the southern and western bias of species richness in the UK marine flora and fauna. The TSA also identified areas which showed significant changes in the spatial pattern of species richness: breakpoints, which correspond to major headlands along the south coast of England. Patterns of algal and molluscan species richness were broadly congruent. Residual variability was strongly influenced by proxies of collection effort, but local environmental variables including length of the coastline and variability in wave exposure were also important. Relative to the underlying trend, local species richness hotspots occurred on all coasts. While there is some justification for scepticism in analyses of heterogeneous datasets, our results indicate that the analysis of collated datasets can be informative.

Molluscan assemblages on an intertidal sandflat in Princess Royal Harbour, Westren Austrailia

The molluscan fauna of an intertidal sand beach in Princess Royal Harbour, W.A.. is divided into two associations. The upper association, located between +0.10 and +0.59 m relative to tidal datum, is characterized by Hydrococcus graniformis and Batillariella estuarina. The lower assemblage, between +0.09 and -0.29 m is dominated numerically by Zeacumantus diemenensis and Katelysia scalarina. The assemblages are equivalent to the midlittoral- and sublittoral-fringe groupings described on both sandy and rocky shores by other authors. There is no supralittoral-fringe component in the molluscan fauna ofprincess Royal Harbour. The break between the two assemblages occurs at + 0.1 m, just below the minimum tide level in the area. Possible reasons for the zonation patterns are discussed and compared with other studies of intertidal zonation on sandy shores.

Behavioural, physiological and morphological adaptations of the shanny (Blennius pholis) to the intertidal habitat

Immersed shannies (Blennius pholis) showed peak locomotory activity coincident with daylight high tides. Emersion caused cessation of breathing and bradycardia though Q02 was little affected. Q02 fell, however, when the abdomen was enclosed in an impermeable sheath to block cutaneous respiration. Gulping of air into the extensively vascular oesophagus probably also acts as a means of aerial respiration. Reimmersion of fish caused a transient bradycardia followed by a tachycardia and a fall in Q02 followed subsequently by a rise. The results are discussed in relation to the behavioural, circulatory, respiratory and morphological adaptations of the shanny to the intertidal habitat.