Spatial variation of reef fishes and the relative influence of biotic and abiotic habitat traits

Patterns of distribution of reef fishes were examined across three spatial scales and related to habitat traits along 25 km of the northern Portuguese coast. Response variables included the multivariate assemblage structure, the total number of taxa and individuals, and the abundance of single groups categorized according to their preference for the benthic, proximo-benthic or pelagic environment, feeding and reproductive behaviour. Habitat traits included topographic elements (small and large ‘drops’ like cracks and crevices) and the extent of dominant morpho-functional types of macroalgae (kelp, large foliose, small erect, turf-forming filamentous, and encrusting). All fish responses were characterized by the largest variance at the smallest scale (among transects tens m apart), followed by that among reefs (hundreds m to 1 km apart) and almost null variance among sites (some km apart). Small and large ‘drops’ of the substratum explained, respectively, considerable variation of assemblage structure and the total abundance of individuals, while the extent of bare rock influenced the richness of taxa and that of benthic fishes, fishes feeding on sessile invertebrates and fishes laying benthic eggs or having nesting behaviour. Combinations of abiotic and biotic structural attributes of reefs influenced proximo-benthic fishes, the predators of mobile animals and fishes releasing pelagic eggs. The here reported associations between patterns of distribution of reef fishes and habitat traits have implications for the design of future protection schemes suitable to guarantee the conservation of reef fish communities and of the processes responsible for their variation. Within the SLOSS (single-large vs. several-small) debate in the design of marine reserves, for example, effective protection to the studied reef fishes would be provided by a set of small reserves, rather than a single large which might be appropriate for fishes having wider home ranges

Potential effects of kelp species on local fisheries

1. Kelp species are ecosystem engineers in temperate coasts, where they provide valuable services to humans. Evidence of the declines of kelp forests exists from several regions, but their effects on fisheries still need to be elucidated. More effective management strategies for sustainable fisheries require a synthesis of research findings and an assessment of how research could be improved to fill current gaps. 2. This review aimed to: (i) summarize the available evidence on the influence of changes in kelp density and/or area on the abundance and diversity of associated fisheries and (ii) examine how research on kelp–fisheries interactions could better support effective management. 3. Most studies (67%) reported data ascribable, directly or indirectly, to a positive relationship between kelp and fishery-relevant variables, 11% provided evidence of a negative relationship, 15% indicated species-specific findings and the remaining found unclear or ‘neutral’ relationships. 4. Important shortcomings were identified, including the paucity of experimental studies suitable to test for unequivocal cause–effect relationships, the disproportion between North America, which is well-studied, and other regions and between the large number of fish-based investigations and the small number of those focusing on other commercially important organisms, and the general lack of studies carried out over spatial and temporal scales comparable to those of global processes driving patterns of distribution of both kelps and fisheries. 5. Synthesis and applications. The consistency of most studies in showing a positive kelp–fishery relationship supports the protection of kelp habitats stated by current environmental directives. However, achieving their goals requires that the limitations we detect are addressed through better connections between research, management practice and policy. This would require the following: (i) researchers to combine multiple approaches (large-scale experimental studies and modelling) for the analysis of kelp–fisheries relationships; (ii) funding agencies to provide resources needed to fill the existing gaps; and (iii) researchers and institutions from less studied regions to strengthen collaborations with those from regions where there have been more investigations into kelp–fishery systems. This is essential under present and predicted environmental changes, with the ultimate aim of conserving and allowing the sustainable use of critically important habitats and of fishery resources relying on these

Compounded perturbations in coastal areas: contrasting responses to nutrient enrichment and the regime of storm-related disturbance depend on life-history traits

1. Natural systems are exposed to compounded perturbations, whose changes in temporal variance can be as important as those in mean intensity for shaping the structure of assemblages. Specifically, climate-related physical disturbances and nutrient inputs due to natural and/or anthropogenic activities occur concomitantly, but experimental tests of the simultaneous effects of changes in the regime of more than one perturbation are generally lacking. Filling this gap is the key to understand ecological responses of natural assemblages to climate-related change in the intensity and temporal patterning of physical disturbance combined with other global stressors. 2. Responses to factorial manipulations of nutrient enrichment, mean intensity and temporal variability in storm-like mechanical disturbance were examined, using benthic assemblages of tide-pools as model system. 3. Response variables were mean abundance values and temporal variances of taxa with different life-traits. Consistent negative effects of disturbance intensity were observed for the mean cover of long-living taxa (algal canopies and the polychaete Sabellaria alveolata), whose temporal fluctuations were also reduced by more severe mechanical stress. More resilient taxa (ephemeral algae, mostly green of the genus Ulva) increased under enriched conditions, particularly when low-intensity events were irregularly applied over time. Opposite effects of disturbance intensity depending on nutrient availability occurred on filamentous algae (e.g. red of the genus Ceramium). This was probably due to the fact that, although nutrient enrichment stimulated the abundance of both algal groups, when this condition was combined with relatively mild physical disturbance the competitively superior ephemeral green algae tended to become dominant over filamentous red algae. The same did not occur under high intensity of disturbance since it likely damaged large, foliose fronds of Ulva-like forms more than small, filamentous fronds of Ceramium-like forms. Grazers were positively affected by nutrients, likely responding indirectly to more food available. 4. A direct relationship between the mean abundance of most organisms and their temporal fluctuations was documented. However, all organisms persisted throughout the study, even under experimental conditions associated to the largest temporal variation in their abundance, likely due to their ability to resist to/quickly recover from, the applied perturbations. Therefore, in systems with great recovery abilities of dominant organisms (e.g. rocky intertidal, grasslands), effects of traits of the regime of disturbance and nutrient enrichment may modulate the fluctuations of populations not through the elimination and substitution of species, but through changes in relative abundances of the same species. This contrasts with the theory that temporal variation in abundance would be directly related to the risk of local extinction. Present findings enable more accurate predictions of the consequences of climatic and non-climatic scenarios on the biodiversity of marine and terrestrial systems sharing analogous functional traits of organisms. Future more intense physical disturbances are expected to exert negative effects on slow-growing/recovering species (e.g. habitat-formers) irrespectively of the temporal patterning of the same disturbances and nutrient inputs. On the contrary, more resistant species (e.g. encrusting algae on rocky shores or below-ground vegetation in grasslands) are expected to benefit from intense physical disturbance. Species whose abundance is more directly related to the availability of nutrients (e.g. filamentous and ephemeral algae or herbs) are expected to generally increase under enriched conditions, but their ability to eventually become dominant would depend on their ability to grow fast and attain cover large enough to overwhelm any possible control of concomitant disturbance intensity on their abundance. If, such as in the present examined system, virtually all organisms can persist, over the temporal scale of the experiment, under any combination of physical disturbance and nutrient availability, the resulting overall diversity is not predicted to change drastically. Nevertheless, low-intensity events evenly distributed and high-intensity events irregularly distributed appear as the conditions supporting the highest richness of taxa, independently of the availability of nutrients

The amphipod assemblages of Sabellaria alveolata reefs from the NW coast of Portugal: An account of the present knowledge, new records, and some biogeographic considerations

Amphipod assemblages associated with the biogenic reefs built by the honeycomb worm Sabellaria alveolata were studied at two sites (Praia da Aguda and Belinho) along the northwestern coast of Portugal. A total of 3909 specimens were collected, comprising 14 different amphipod species. A first record from the northeastern Atlantic coast was registered here for the species Caprella santosrosai, which was, up to now, recorded only along the Mediterranean coast of the Iberian Peninsula. A male specimen collected from the Sabellaria-reef located in Belinho allowed an update to the known distribution of C. santosrosai, thus altering its previous status as an endemic Mediterranean species. The most common species collected during the study were Microdeutopus chelifer (n = 1828), Jassa ocia (n = 1426), and Hyale stebbingi (n = 452). Forty-three percent of the total recorded species were encountered in both study sites, whereas the remaining 57 % were restricted to a single site (Belinho). The majority of the collected species (93 %) showed an Atlantic-Mediterranean distribution, confirming the close affinity between eastern Atlantic and Mediterranean amphipod assemblages and the role of the Portuguese coast as a transition zone through which numerous warm-water species, coming from North Africa and the Mediterranean Sea, could enter into the Atlantic and possibly get mixed with species coming from the North Sea and the Arctic, typically having affinity for colder waters

Connell and Slatyer's models of succession in the biodiversity era

Understanding how species interactions drive succession is a key issue in ecology. In this study we show the utility of combining the concepts and methodologies developed within the biodiversity–ecosystem functioning research program with J. H. Connell and R. O. Slatyer’s classic framework to understand succession in assemblages where multiple interactions between early and late colonists may include both inhibitory and facilitative effects. We assessed the net effect of multiple species interactions on successional changes by manipulating the richness, composition, and abundance of early colonists in a low-shore assemblage of algae and invertebrates of the northwestern Mediterranean. Results revealed how concomitant changes in species richness and abundance can strongly alter the net effect of inhibitory vs. facilitative interactions on succession. Increasing richness of early colonists inhibited succession, but only under high levels of initial abundance, probably reflecting the formation of a highly intricate matrix that prevented further colonization. In contrast, increasing initial abundance of early colonists tended to facilitate succession under low richness. Thus, changes in abundance of early colonists mediated the effects of richness on succession

Herbivory drives kelp recruits into ‘hiding’ in a warm ocean climate

Assessing effects of herbivory across broad gradients of varying ocean climate conditions and over small spatial scales is crucial for understand- ing its influence on primary producers. Effects of her- bivory on the distribution and abundance of kelp re- cruits were examined experimentally at two regions under contrasting ocean climate. Specifically, the abundance and survivorship of kelp recruits and the abundance of macro-herbivores were compared be- tween a ‘cool’ and a ‘warm’ region in northern and central Portugal, respectively. In each region, the abundance of kelp recruits and the intensity of grazing were compared between habitats of different topography within reefs (open reef vs. crevices). Com- pared to the ‘warm’ region, the abundance of kelp re- cruits was 3.9 times greater in the ‘cool’ region, where 85% of recruits were found in open reef habitats. In contrast, 87% of recruits in the ‘warm’ region were re- stricted to crevices. The ‘warm’ region had 140 times greater abundances of sea urchins, 45 times more herbivorous fish and 4.1 times more grazing marks on kelp recruits than the ‘cool’ region. Grazing assays showed ca. 50 times higher rates of kelp biomass con- sumption, mainly by fishes, and zero survivorship of kelp recruits in the ‘warm’ relative to the ‘cool’ region. This study suggests both temperature and herbivores affect abundances of kelp recruits across latitudes, and demonstrates how herbivores affect their distri- bution at local scales, driving kelp recruits into ‘hiding’ in crevices under intense herbivory. Conse- quently, where net recruitment success is compro- mised by herbivory, the persistence of kelps will be contingent on availability of topographical refuges

Reducing the data-deficiency of threatened European habitats: Spatial variation of sabellariid worm reefs and associated fauna in the Sicily Channel, Mediterranean Sea

Biogenic reefs, such as those produced by tube-dwelling polychaetes of the genus Sabellaria, are valuable marine habitats which are a focus of protection according to European legislation. The achievement of this goal is potentially hindered by the lack of essential empirical data, especially in the Mediterranean Sea. This study addresses some of the current knowledge gaps by quantifying and comparing multi-scale patterns of abundance and distribution of two habitat-forming species (Sabellaria alveolata and S. spinulosa) and their associated fauna along 190 km of coast on the Italian side of the Sicily Channel. While the abundance of the two sabellariids and the total number of associated taxa did not differ at any of the examined scales (from tens of centimetres to tens-100 of kilometres), the structure (composition in terms of both the identity and the relative abundance of constituting taxa) of the associated fauna and the abundance of several taxa (the polychaetes Eulalia ornata, Syllis pulvinata, S. garciai, Nereis splendida and Arabella iricolor, and the amphipods Apolochus neapolitanus, Tethylembos viguieri and Caprella acanthifera) varied among locations established ~50e100 km apart. Syllis pulvinata also showed significant variation between sites (hundreds of metres apart), analogously to the other syllid polychaetes S. armillaris and S. gracilis, the nereidid polychaete Nereis rava, and the amphipod Gammaropsis ulrici. The largest variance of S. spinulosa, of the structure of the whole associated fauna and of 56% of taxa analysed individually occurred at the scale of replicates (metres apart), while that of the dominant bio-constructor S. alveolata and of 25% of taxa occurred at the scale of sites. The remaining 19% and the total richness of taxa showed the largest variance at the scale of locations. Present findings contribute to meet a crucial requirement of any future effective protection strategy, i.e., identifying relevant scales of variation to be included in protection schemes aiming at preserving representative samples not only of target habitats and organisms, but also of the processes driving such variabilit

The regime of climate-related disturbance and nutrient enrichment modulate macroalgal invasions in rockpools

Patterns of invasion by the seaweeds Grateloupia turuturu Yamada and Sargassum muticum (Yendo) Fensholt under crossed combinations of the regime (mean intensity and temporal variability) of climate-related mechanical disturbance and constant nutrient enrichment were experimentally examined in rockpools in north Portugal. The cover of both species was larger under high compared to low intensity of disturbance, but this effect was enhanced by events more heterogeneously distributed over a period of 19 months. The invasion by G. turuturu was also larger in enriched pools, but only when disturbance was applied at high intensity. The richness of native taxa was increased by high intensity events of disturbance evenly distributed over time and by low intensity events heterogeneously distributed, while no differences were documented for other treatments. Temporal variability of disturbance and nutrients interactively affected the total cover of native taxa and the availability of bare rock in different directions. Enriched conditions increased the space occupancy by natives and reduced substratum availability only when associated to heterogeneous events of disturbance. At the same time, relatively more variable disturbances caused a reduced cover by native taxa and an increased availability of free space, but only under natural nutrient levels. Present findings contribute to understand the conditions that would be more likely to facilitate the spread of G. turuturu and S. muticum under current and predicted scenarios of compounded environmental changes and in relation to traits of recipient systems that are considered relevant for the success of invasions, including the native richness and the degree of usage of resources, i.e. primarily space