Low densities of sea urchins influence the structure of algal assemblages in the western Mediterranean

Numerous studies of interactions between urchins and algae in temperate areas have shown an important structuring effect of sea urchin populations. These studies focused almost wholly on the effect of high urchin densities on laminarian forests. In contrast, algal communities below 5–6 m depth in the northwestern Mediterranean are characterised by low sea urchin densities (<5 ind m⁻²) and the absence of laminarian forests. No previous research has addressed sea urchin/algal interactions in this type of community. To determine the effect of the most abundant echinoid species, Paracentrotus lividus, on well-established algal communities in this area, we performed a removal–reintroduction experiment in rocky patches located between 13 and 16 m depth in the northwestern Mediterranean, where sea urchin densities ranged between 0.9 and 3.4 ind m⁻². After 6 months, the cover of non-crustose algae was significantly higher in the plots from which sea urchins had been removed than in control plots (84 vs 67% cover). These removal plots reverted to their original state upon reintroduction of sea urchins. The non-crustose algae consisted of turfing and frondose forms, with the former representing some 70% of the non-crustose algal cover. Change in the cover of turfing algae was responsible for the significant increase in algal development in the sea urchin removal plots. The response of frondose algae to the treatment varied between algal species. It is concluded that grazing by P. lividus exerts a significant effect on habitat structure, even in communities with low sea urchin densities, such as those found in vast areas of the Mediterranean sublittoral.     

Demographic Characteristics of Lytechinus variegatus (Echinoidea: Echinodermata) from Three Habitats in a North Florida Bay, Gulf of Mexico

Abstract. The population densities, spatial distributions, size frequencies, growth rates, longevity and reproductive activities of sub‐populations of the sea urchin Lytechinus variegatus were investigated over a two‐year period. Sea urchins were examined in three habitats in Saint Joseph Bay, Florida, which is within the northern limits of their distribution. Densities of sea urchins, which ranged as high as 35 individuals ·^?2, fluctuated seasonally at all sites and were higher in seagrass beds comprised of Thalassia testudinum than Syringodium filiforme or on a sand flat. A cold front caused large‐scale, catastrophic mortality among adult, and especially juvenile, sea urchins in nearshore habitats of the Bay in the spring of 1993, leading to a dramatic decline in sea urchin densities at the Thalassia seagrass site. The population recovered over 6 months at this site and was attributable to immigration of new adults. Juvenile recruitment displayed both interannual and site‐specific variability, with recruitment being highest in seagrass habitats in fall and spring. The most pronounced recruitment event occurred in fall 1993 at the Thalassia site. Spatial distributions of adult individuals ascertained monthly never varied from random in the seagrass beds (T. testudinum and S. filiforme) or during spring, summer or fall months on the sand flat. Nonetheless, aggregations of adult sea urchins were observed on the sand flat in the winter months and were associated with patchy distributions of plant food resources. Juvenile sea urchins (< 25 mm test diameter) exhibited aggregations at all sites and 67 % of all juveniles under 10 mm test diameter (91 of 165 individuals observed) were found under the spine canopies of adults. Measurements of the inducibility of spawning indicated peak gametic maturity in all three sub‐populations in spring and summer. Gonad indices varied between habitats and years, but distinct maxima were detected, particularly in spring 1993 and late summer 1994. The mean gonad index of individuals at the Syringodium seagrass site was 2‐ to 4‐fold higher than the other sites in spring 1993 and gonad indices were much higher at all sites in spring of 1993 than 1994. Estimates of growth based on changes in size frequency cohorts coupled with measurements of growth bands on lantern demipyramids indicated that L. variegatus in three habitats of Saint Joseph Bay have similar growth rates and attain a mean test diameter of approximately 35 mm in one year. In contrast to populations within the central biogeographical range of the species, which may attain test diameters up to 90 mm, the largest individuals recorded in Saint Joseph Bay were 60 mm in test diameter, and almost all individuals were no more than 45 mm in test diameter or two years of age. The demographics of L. variegatus in the northern limits of their distribution appear to be strongly influenced by latitudinally driven, low‐temperature events and secondarily by local abiotic factors, especially springtime low salinities, which may negatively impact larval development and recruitment.     

Evidence for a positive relationship between juvenile abalone Haliotis midae and the Sea urchin Parechinus angulosus in the South-Western Cape,South Africa

Previous field observations have suggested an association between the urchin Parechinus angulosus and juveniles of the abalone Haliotis midae. To test the generality and nature of this association, surveys were carried out at five sites between Cape Point and Danger Point in the kelp beds of the South-Western Cape, South Africa. These showed that both species occupy primarily hard substrata, showing preferences for encrusting coralline algae. They also confirmed a strong, positive relationship between urchins and juvenile abalone. Of the juvenile abalone sampled, more than 98% were found beneath sea urchins. All small (3–10 mm) and medium-sized(11–20 mm) juvenile abalone were under urchins, whether on flat or vertical reef, or in crevices. A small proportion (～10%) of larger juveniles(21–35 mm) was not found under urchins, and in these instances they occupied crevices instead. These findings are of particular importance in terms of their implications for the lucrative commercial abalone fishery in South Africa, indicating that urchins are of critical importance to the continued survival of viable abalone populations. There has been a dramatic decrease in natural populations of sea urchins over the past five years in the heart of the abalone fishing grounds, and the present findings suggest that this will lead to recruitment failure of abalone, because juvenile abalone seem dependent on the urchins. The long-term consequences for the industry may be crucial.     

Evaluating the effects of protection on fish predators and sea urchins in shallow artificial rocky habitats: a case study in the northern Adriatic Sea

Man-made defence structures (e.g., breakwaters, jetties) are becoming common features of marine coastal landscapes all around the world. The ecology of assemblages of species associated with such artificial structures is, however, poorly known. In this study, we evaluated the density and size of fish predators of echinoids (i.e., Diplodus sargus, Diplodus vulgaris, Sparus aurata), and the density of sea urchins (i.e., Paracentrotus lividus) at defence structures (i.e., breakwaters) inside and outside the marine protected area of Miramare (northern Adriatic Sea) in order to: (1) assess possible differences in fish predator density and size between protected and fished breakwaters; (2) assess whether fish predation may have the potential to affect sea urchin density in artificial rocky habitats. Surveys were carried out at four random times over a period of two years. Total density, and density of medium- and large-sized individuals of the three predatory fishes were generally greater at the protected than at the fished breakwaters, whereas no differences were detected in the density of small-sized individuals. Density of the sea urchin P. lividus did not show any difference between protected and fished breakwaters. The results of this study suggest that: (1) protection may significantly affect predatory fishes in artificial rocky habitats; (2) differences in predatory fish density, and size may be unrelated with the density of the sea urchin P. lividus; (3) protected artificial structures such as breakwaters, originally planned for other purposes, could represent a potential tool for fish population recovery and enhancement of local fisheries.     

Effect of two different habitats on spine and gonad colour in the purple sea urchin Paracentrotus lividus

This study investigates the effect of different Mediterranean habitats on spine and gonad colouring and gonadal index in the purple sea urchin Paracentrotus lividus (Echinodermata: Echinoidea). Specimens were collected in winter 2012 in Posidonia oceanica meadows and rocky bottom areas at two sites in Southern Sardinia (Italy, Tyrrhenian Sea). Colorimetric analyses of spines and gonads were conducted with a digital colorimeter using CIELAB colour spaces, which defines the colour in terms of three coordinate values: lightness (L*), redness (a*) and yellowness (b*). Lightness and redness of the spines differed significantly between habitats, whereas colour spaces of the gonads did not. A careful analysis of the gonad index showed it to be significantly affected by habitat, registering higher values in rocky bottom locations. In conclusion, it seems that habitat does affect spine colour but not gonad colour. Spine colour seems to be site‐dependent, whereas the gonad index depends solely on the habitat.     

Sedentary urchins influence benthic community composition below the macroalgal zone

Sea urchins are important ecosystem engineers in subtidal ecosystems worldwide, providing biogenic structure and altering nutrient dynamics through intensive grazing and drift algal capture. The current work evaluates red urchin (Strongylocentrotus franciscanus) density on fixed transects through time, individual displacement, and urchin‐associated benthic community composition using a field‐based approach at multiple depths (in and outside of the macroalgal zone) and replicated across sites in the San Juan Archipelago, Washington. Urchins exhibited no large‐scale, temporal or directional changes in density among depths. Furthermore, 87% of individual urchins observed in repeated small‐scale surveys over 3 weeks exhibited no change in position. Individual displacement was negatively correlated to drift algal capture. Evidence of sedentary behavior from the displacement surveys was supported by the sessile and mobile community composition in areas directly under versus adjacent to (control) urchins. The benthos under urchins had a higher percentage of bare space, crustose coralline algae, and increased density of snails, crabs and shrimp relative to associated control plots. Abundance of mobile organisms associating with urchins increased relative to control plots at the deepest survey depth (30 m), indicating a greater strength of interaction with distance from macroalgal production. This work presents evidence of food availability‐related behavior in red urchins and indicates that even when sedentary, urchins have a strong influence on ecosystem structure through increasing availability of shelter and macroalgal detritus to the benthos.     

Persistent alternate abundance states in the coral reef sea urchin Diadema savignyi: evidence of alternate attractors

Alternate attractors have been shown to exist in a variety of terrestrial and aquatic systems, e.g. temperate forests, savannas, shallow lakes, wetlands, coral reefs, kelp forests. The shift from one attractor to another, also referred to as a regime shift, is thought to occur when a system passes some critical threshold such that the trajectory of the system changes direction. Alternate attractors in population dynamics can also exist, leading to alternate stable states in the population abundance of a species. This study explored alternate attractors in the population dynamics of the Indo‐Pacific sea urchin Diadema savignyi and the potential underlying mechanisms that promote its bi‐stability. In Moorea, French Polynesia, the local abundance of D. savignyi, a functionally important herbivore in lagoon habitats, occurs in two states: (i) solitary individuals that occupy crevices in low densities and (ii) aggregations of tens to hundreds of individuals. These different states are temporally stable and are not explained by spatial differences in recruitment rates of juveniles. A field experiment revealed that the per capita mortality rate of adult D. savignyi was substantially lower at sites where urchins occurred in aggregations compared with sites at which they were solitary individuals. An additional experiment showed that per capita mortality decreased with increasing aggregation size. Individuals in high‐density aggregations, however, had significantly smaller test diameters than solitary individuals, indicating that individuals in aggregations may be food limited. Collectively, the evidence suggests that the two different local abundance states of D. savignyi result from negative feedback loops where high local density can be maintained by aggregative behavior that greatly reduces per capita risk of predation when the local number of adult sea urchins is sufficiently large; sites with few sea urchins remain at low density because individuals are more susceptible to predation when crevices are occupied but there are not enough individuals to form large aggregations. Thus, there may be alternate attractors in the population dynamics of D. savignyi that can produce either persistently low or high local population densities.     

A mass mortality of subtropical intertidal populations of the sea urchin Paracentrotus lividus: analysis of potential links with environmental conditions

An outbreak of bald sea urchin disease was detected affecting intertidal populations of the edible temperate species Paracentrotus lividus at its southernmost geographical limit. The mortality event was detected in three sites (Güímar, Palmar and Alcalá) off Tenerife, Canary Islands (Northeastern Atlantic), in October 2003, coinciding with the highest sea surface temperatures (SST) and the lowest wave heights of the year. The prevalence of infection reached 100% but did not affect equally all studied sites or the different size classes. The disease was more prevalent in large sea urchins (>40 mm). The prevalence appeared to be positively correlated with the environmental variable SST and negatively correlated with wave height. In October 2004, only two infected urchins were found (prevalence 1%) at one of the studied sites. However, during 2004 no such extreme values of temperature and wave height were observed. We conclude that high SSTs and low wave heights such as those observed during 2003 are conditions that may trigger outbreaks of urchin disease.     

Community structure and functional traits of mollusks associated with coastal reef macroalgae in Northeastern Brazil

This study investigated the community structure and functional traits of the mollusk fauna associated with macroalgae with different thallus morphologies in a reef environment in Northeastern Brazil. A total of 15 individuals of each species of macroalgae adhered to natural substrate and 15 individuals of Padina gymnospora detached from the substrate were collected. The structural complexity of algal habitats was measured and the associated malacofauna screened and identified. All three macroalgae differed significantly in the complexity of their habitat, with Sargassum polyceratium being the most complex. A total of 823 specimens of mollusks belonging to 22 species and 11 families were recorded, of which Columbellidae was the most represented with six species. The functional trait “size” revealed that micromollusks smaller than 10 millimeters were predominant in the community; however, individuals of larger sizes (up to 24.54 millimeters) belonging to young stages of other species were also present. Eight functional trophic groups were identified, of which “carnivorous” stood out with seven species. Six functional groups of microhabitats were identified, with intra‐specific variation in habitats, while habitat expansion was documented for species not yet recorded in association with macroalgae. The structure of the molluskan community differed among the three algae species with the greatest richness, abundance, and diversity of mollusks and functional traits occurring with S. polyceratium. Community structure differed between algae adhered to natural substrate and detached algae, with the latter having lower mollusk richness and diversity, but with greater abundance of some species that remained on algal fronds after release from the reef environment. This study reinforces the importance of algal habitat for marine invertebrate fauna, especially for micromollusks that spend their entire life cycle, or part of it, in association with macroalgae.     

Comparative study on the fauna composition of intertidal invertebrates between natural and artificial substrata in the northeastern coast of Jeju Island

This study was carried out to learn about differences in the sessile macrobenthic fauna communities between the artificial and natural habitats. There were some differences in terms of species composition and dominant species and community structure between two habitat types. The dominant species include Pollicipes mitella and Granuilittorina exigua in natural rocky intertidal zones; Monodonta labio confusa, Ligia exotica, Tetraclita japonica in the artificial rocky intertidal zones. Among all the species, L. exotica and T. japonica occurred only in the artificial rocky intertidal zone. The results of cluster analysis and nMDS analysis showed a distinct difference in community structure between artificial and natural rocky intertidal zones. The fauna in the natural rocky intertidal zones were similar to each other and the fauna in the artificial rocky intertidal zones were divided depending on the slope of the substratum. In the case of a sloping tetrapod, M. labio confusa and P. mitella were dominant, but at the vertical artificial seawall, Cellana nigrolineata, L. exotica T. japonica were dominant. The analysis of the species presented in natural and artificial rocky intertidal areas showed the exclusive presence of 10 species on natural rocks and 12 species on artificial rocks. The species in the natural rocky intertidal area included mobile gastropods and cnidarians (i.e. rock anemones), and the species in the artificial rocky intertidal area mostly included non-mobile attached animals. The artificial novel structure seems to contribute to increasing the heterogeneity of habitats for marine invertebrate species and an increase the species diversity in rocky coastal areas.     

Welcome mats? The role of seagrass meadow structure in controlling post-settlement survival in a keystone sea-urchin species

Processes acting on the early-life histories of marine organisms can have important consequences for the structuring of benthic communities. In particular, the degree of coupling between larval supply and adult abundances can wield considerable influence on the strength of trophic interactions in the ecosystem. These processes have been relatively well described in rocky systems and soft-sediment communities, and it is clear that they are governed by very different bottlenecks. Seagrass meadows make interesting study systems because they bear structural affinities to both soft sediments as well as rocky substrates. We examined the early-life history of Paracentrotus lividus, one of the dominant herbivores in Mediterranean seagrass meadows, to identify the drivers of population dynamics in this species. We measured spatial and temporal variability in sea urchin post-settlement in 10 Posidonia oceanica meadows in the North-Western Mediterranean over a period of two years, and compared the numbers with the one-year old cohort a year later (i.e. the new population recruitment) as well as between successive size–age groups. Urchin post-settlers differed substantially between meadows but were present in both years in all meadows surveyed, suggesting that larval supply was not limiting for any of the studied sites. However, in six of the studied meadows, the one-year cohort of urchins was absent in both years, indicating that post-settlement processes strongly affected urchins in these meadows. In contrast, in four of the studied meadows, there was a strong coupling between post-settlers and one-year cohort individuals. These meadows were structurally different from the others in that they were characterised by an exposed matrix of rhizomes forming a dense seagrass mat. This mat apparently strongly mediates post-settlement mortality, and its presence or absence dictates the successful establishment of urchin populations in seagrass meadows. As the population aged, the relationship between size–age groups decreased evidencing the action of other processes. Yet, these results indicate that differences in physical structure are a vital bottleneck for sea urchin populations in seagrass meadows. Exploring the interaction between ecosystem structure and early-life history may provide a broader and more unified framework to understand the dynamics of a range of benthic habitats, including rocky substrates, soft sediments and seagrass meadows.     

Patterns of Activity of the Sea Urchin Paracentrotus lividus in the Bay of Port-Cros (Var, France, Mediterranean)

Abstract. Tagged individuals of the sea urchin Paracentrotus lividus followed in Port Cros Island showed mostly nocturnal movements, with individual peaks of activity which were not coordinated for the population during the night period. The straight line distance travelled in 24 h has a mean value of 49 cm, with a range of 0 to 240 cm. Neither sea urchin size nor depth affect activity. The urchins living on rocky substrate moved significantly more than those on seagrass beds. From one day to another, the distance travelled by the population of urchins varies significantly; this is not related to season. The activity is negatively correlated with the mean wind speed; during a period of turbulence lasting several hours, movement was significantly oriented to the deep. Periods of inactivity of 1 or 2 days were observed; they are not coordinated for the individuals followed, and the relation between feeding and movement is not clear. Urchins seem to stay in a quite small area, at least for some months.     

Biotic consequences of a shift in invertebrate ecosystem engineers: Invasion of New Zealand rocky shores by a zone‐forming ascidian

The pervasive effects of invasive ecosystem engineers, that is those species that modify their environment, are well documented, but rarely have the broader impacts of one foundation invertebrate species being replaced by another been examined. In New Zealand, green‐lipped mussels, Perna canaliculus, commonly dominate wave‐exposed rocky shores. The recent appearance of an invasive ecosystem engineer, the ascidian Pyura doppelgangera, at the very northern tip of New Zealand now threatens to exclude these bivalves from this habitat. Here, we report major shifts in assemblages associated with the invader and chronicle its continued spread. We examined epibiota associated with clumps of mussels and clumps of Pyura from two rocky shore habitats—pools and emergent substrata at two locations. We detected some differences in species richness in biota associated with the two foundation species, but faunal abundance only differed between the locations. These minor changes were dwarfed by the shift in species composition within clumps of each foundation species. Molluscs, particularly gastropods, and crustaceans dominated the assemblage within mussels. In contrast, tubicolous polychaetes dominated the fauna associated with the ascidian. Sessile epifauna, notably barnacles and calcareous tube‐dwelling polychaetes, were common on mussels, but never encountered on the ascidian. Multivariate analysis revealed marked dissimilarity (>80%) between the characteristic mussel and ascidian faunas with virtually no overlap. This biotic shift overshadowed any differences between habitats and locations. The broader implications of these faunal shifts for local and regional patterns of biodiversity, as well as ecosystem function, remain unclear, but deserve further attention.     

Relationships among predatory fish, sea urchins and barrens in Mediterranean rocky reefs across a latitudinal gradient

Previous studies conducted on a local scale emphasised the potential of trophic cascades in Mediterranean rocky reefs (involving predatory fish, sea urchins and macroalgae) in affecting the transition between benthic communities dominated by erected macroalgae and barrens (i.e., bare rock with partial cover of encrusting algae). Distribution patterns of fish predators of sea urchins (Diplodus sargus sargus, Diplodus vulgaris, Coris julis and Thalassoma pavo), sea urchins (Paracentrotus lividus and Arbacia lixula) and barrens, and fish predation rates upon sea urchins, were assessed in shallow (3-6m depth) sublittoral rocky reefs in the northern, central and southern sectors of the eastern Adriatic Sea, i.e., on a large spatial scale of hundreds of kilometres. No dramatic differences were observed in predatory fish density across latitude, except for a lower density of small D. sargus sargus in the northern Adriatic and an increasing density of T. pavo from north to south. P. lividus did not show any significant difference across latitude, whereas A. lixula was more abundant in the southern than in the central Adriatic. Barrens were more extended in the southern than in the central and northern sectors, and were related with sea urchin density. Fish predation upon adult sea urchins did not change on a large scale, whereas it was slightly higher in the southern sector for juveniles when predation rates of both urchins were pooled. Results show that: (1) assemblages of predatory fish and sea urchins, and barren extent change across latitude in the eastern Adriatic Sea, (2) the weak relations between predatory fish density and predation rates on urchins reveal that factors other than top-down control can be important over large scale (with the caveat that the study was conducted in fished areas) and (3) patterns of interaction among strongly interacting taxa could change on large spatial scales and the number of species involved.     

Macrofauna associated to Mycale microsigmatosa (Porifera, Demospongiae) in Rio de Janeiro State, SE Brazil

The macrofauna (endo- and epi-biotic) associated to the sponge Mycale (Carmia) microsigmatosa Arndt, 1927 was studied at three sites in Rio de Janeiro State, Brazil (Arraial do Cabo, Niterói, and Rio de Janeiro). A total of 2235 individuals (over 1 mm long) of 75 invertebrate species were found associated to 19 specimens of the sponge. The most abundant and diverse taxa were the crustaceans (83%, 31 spp.), polychetes (10%, 18 spp.), and molluscs (3.7%, 15 spp.). Cnidarians, platyhelminthes, ascidians, echinoderms, pycnogonids, bryozoans, and sponges were also represented. Amphipod crustaceans were the dominant group, comprising 61% of all individuals collected. Species richness and abundance of associated fauna were highly correlated with sponge volume, but diversity and evenness were not. The site of collection influenced the species composition of the fauna associated to M. microsigmatosa but did not change significantly its diversity, abundance, richness, and dominance patterns of higher taxa. Pregnant females and juvenile stages of 29% of the species associated, including crustaceans, molluscs, echinoderms, and pycnogonids were frequently found inside M. microsigmatosa. Although many of these organisms do occur and reproduce in other habitats outside the sponge as well, M. microsigmatosa is also important for their reproduction and survivorship, thus contributing for the maintenance of biodiversity in Southwestern Atlantic sublittoral rocky shores.     

Size-dependent distribution, abundance and diurnal rhythmicity patterns in the short-spined sea urchin Anthocidaris crassispina

Little is known about the ecology and behaviour of the sea urchin Anthocidaris crassispina, particularly in relation to changes in its surrounding environment. In Cape d'Aguilar Marine Reserve, Hong Kong Island, urchins ranged from a high abundance of 16 ind m⁻² on steeply inclined rocky outcrops, which were exposed to strong onshore waves surges, to a complete absence on gravel and sandy substrata. Using a generalized additive model (GAM) the abundance and spatial extent of A. crassispina was predicted within the Bay. Predictions were strongly associated with the surface complexity of the habitat. On steep rocky slopes A. crassispina exhibited a size-dependent gradient where the mean test diameter increased in a down-shore direction as water depth increased. Since the mechanisms for maintaining this size-dependent distribution are unknown, a translocation experiment was conducted on two different size-classes of urchins. Following translocation, both large and small size-classes of A. crassispina were able to re-establish their original size-gradient within 3–5 days. Size-dependent distribution in A. crassispina may indicate resource partitioning, although the influence of hydrodynamic conditions on test size may also mediate this segregation. Anthocidaris crassispina was predominately nocturnal with almost 100% of the population moving between dusk and dawn. Locomotory activity patterns of urchins were strongly correlated with changes in seawater depth and changes in the direction of water-flow during tidal cycles. Anthocidaris crassispina exhibited an endogenously controlled locomotory activity pattern that was synchronized with changes in the tidal cycle, and which remained free-running for 13 h under constant laboratory conditions.     

Effects of point-source nutrient addition and mussel removal on epibiotic assemblages in Perumytilus purpuratus beds

The objective of this study was to evaluate the effects of point-source nutrient addition and mussel removal on species diversity of the epibenthic assemblage of the purple mussel Perumytilus purpuratus. It was hypothesised that mussels cushion the effects of disturbance on the associated organisms and thereby favour species diversity in patches with added nutrients. In order to test this hypothesis, a two-factorial field experiment was conducted at an exposed rocky shore of northern-central Chile in which nutrient addition and mussel removal were manipulated with two levels. Experimental units consisted of 100-cm² mussel plots that were sampled after two months of experimental manipulations. Local nutrient addition was done with coated slow-release fertilisers. Mussel removal treatments consisted in three destructive events applied throughout the experimental phase, and each event consisted in the removal of 20% of the mussel biomass from each plot. While point-source nutrient addition had only minor effects on diversity, mussel removal had significant and negative effects on faunal species richness and the abundance of suspension-feeders and sessile organisms. The abundance of predators/scavengers significantly decreased with mussel removal in the presence of nutrient addition. Grazers and mobile organisms were very abundant in the mussel assemblage but their abundance remained unaffected by treatments. Finally, species richness and total abundance of algae were also not affected by the treatments. Although the duration of our experiment was relatively short, we suggest that the effects of local nutrient addition are of minor importance at the study site in comparison to the effects of mussel removal. Based on our results and previous studies conducted in soft-bottom systems, we suggest that physical and biological stress acting on exposed hard-bottom communities overshadows the potential effects of local nutrient addition.     

Sponge epibionts on ecosystem-engineering ascidians: The case of Microcosmus sabatieri

The study of epibionts on habitat engineering ascidians is of increasing interest because changes in the population structure of the latter may affect associated communities, especially in the case of commercially exploited species. The solitary ascidian Microcosmus sabatieri lives on rocky cliffs in the Eastern Mediterranean and is harvested in certain Aegean areas. Its hard, wrinkled tunic is usually fouled by various epibionts both sessile and motile. Sponges are an important component of this complex and their biomass may be higher than that of the ascidian itself, strongly affecting diversity and abundance of the motile epifauna. The aim of this study was to examine in detail the structure of the epibiotic sponge assemblage on ascidians collected from their main fishing grounds in the South Aegean Sea. A rich (41 species) and taxonomically diverse sponge assemblage was found, while only eight species contributed 80% of the total sponge cover. Most of the epibiotic sponges commonly grow on the surrounding sublittoral cliffs. The encrusting sponge growth form prevailed in cover of the ascidian tunic, while two massive species dominated in terms of frequency of appearance and abundance. Ascidian dimensions, weight and volume were significantly correlated with sponge diversity, abundance and cover area, thus structuring the epibiotic sponge assemblage. Spatial patterns in sponge cover were not clear, but a general declining NW to SE trend in sponge richness, abundance and cover appeared in accordance with previous records. Sponge distribution on the ascidian tunic presented a clear pattern related with characteristic features of the ascidian: the posterior zone supported the richest and most expansive sponge fauna. The ecosystem-engineering process performed by the ascidian is enhanced by the diverse epibiotic sponge assemblage, thus further increasing habitat complexity in this space-limited, temperate, sublittoral, rocky environment.     

Effects of Paracentrotus lividus (Lamark, 1816) harvesting on benthic assemblages. An experimental approach

The effect of harvesting on the diversity and structure of benthic assemblages is nowadays a relevant component of fishery management. Invertebrate harvesting is increasing worldwide and sea urchins are one of the main invertebrate target species. However, current knowledge about the effects of sea urchin harvesting on benthic assemblages is still poor. The aim of this study is to evaluate the influence of Paracentrotus lividus (Lamark, 1816) harvesting on tide pool assemblages from intertidal rocky shores. To achieve this objective, a field removal experiment was done for 10 months. Results of univariate and multivariate analyses showed that P. lividus harvesting had a significant effect on the number of taxa and structure of the benthic assemblage. Assemblages from pools where P. lividus were removed showed significant differences against controls; namely, where P. lividus were eliminated, pools showed higher number of taxa than control pools. This result contrasts with previous studies assessing effects of invertebrate harvesting that generally have found a biodiversity reduction in the exploited habitat. Despite this apparently positive effect of P. lividus harvesting, we should be cautious because effects of P. lividus harvesting on higher trophic levels or during long periods of time remain unexplored.     

Spatial pattern of coastal fish assemblages in different habitats in North‐western Mediterranean

Variability of fish assemblages across habitat structures can depend on spatial scales. A hierarchical sampling design was used to assess the spatial variability of temperate fish assemblages in different habitats and at multiple scales. Underwater visual censuses were carried out along the coasts of Elba Island (NW Mediterranean) on Posidonia oceanica beds, rocky algal reefs and sandy habitat at three spatial scales, namely tens of metres (individual replicates), hundreds of metres (sites) and tens of kilometres (locations). At the assemblage level, there was a clear relationship between fish and habitat type and the observed habitat‐related differences were largely dependent on species identity. Fish assemblages on P. oceanica beds and rocky reefs shared a high number of species, whereas overlap with sandy assemblages was negligible. Multivariate analyses revealed significant differences in fish assemblages among habitats, although there was also a significant habitat × site interaction. These differences relied mainly upon assemblage composition and species richness. Assemblages on rocky reefs and P. oceanica meadows usually harboured a higher number of species and individuals compared with sandy assemblages. Nevertheless, the patterns of habitat‐related differences in species richness and, especially, in the total number of fish, changed significantly from site to site. Eight species showed significant differences over habitats, but they were not consistent due to the interaction of habitat with site. Predictability of fish at both assemblage and population levels decreased with the scale of observation, and the spatial pattern of fish observed at the smallest scale was likely dependent on factors other than habitat type.