Spatial and depth-associated distribution patterns of shallow gorgonians in the Algarve coast (Portugal, NE Atlantic)

The ecological role of gorgonians for marine rocky bottoms is worldwide recognized, but the information on the distribution patterns of NE Atlantic temperate species is insufficient, considering current global, regional and local threats. To overcome the lack of information on the spatial distribution patterns of gorgonians in south Portugal, in 2009/2010, the occurrence and abundance of gorgonian species in rocky bottoms were quantified over more than 25 km of coast (37.1°N/8.6°W) down to 30 m depth. Eunicella labiata, Eunicella gazella, Eunicella verrucosa and Leptogorgia sarmentosa were abundant and frequent in the studied area, while Leptogorgia lusitanica was less abundant. All species evidenced a similar depth pattern, that is abundance significantly increased with depth below 15 m. At shallower waters (up to 15 m), the distribution of gorgonians may be constrained by abiotic factors and competition with algae. Indeed, the abundance of gorgonians was negatively correlated with the percentage cover of algae along the depth gradient, but gorgonians and sponges coexist. Competition among gorgonian species also seems to be low in this area because of the similarity in the abundance pattern observed for the most abundant species and also their high association. In NE Atlantic shallow temperate rocky bottoms, the distribution of gorgonians seems to be influenced by environmental factors and biological interactions, namely competition (algae) and coexistence (sponges and other gorgonians).     

Evaluation of the utility of COI and ITS markers as tools for population genetic studies of temperate gorgonians

The aim of this study was to analyse the polymorphism of two molecular markers, mitochondrial COI and nuclear ITS 2, to determine the genetic structure of Mediterranean gorgonian populations. Four gorgonians, Eunicella cavolinii, Eunicella singularis, Paramuricea clavata and Corallium rubrum were analysed for COI, while only E. cavolinii was analysed for ITS. These species have been recently affected by mass mortality events probably linked to climate change. In the context of these strong disturbances, reliable data on the genetic structure of affected populations could be critical in order to propose management and conservation strategies for these species. A hierarchical sampling plan encompassing different spatial scales (from tenths to hundreds of kilometres) was designed to test these markers. Both markers failed to make evident any genetic structure of these populations. The evolutionary features and utility of these markers are critically discussed and future research directions are proposed.     

Structural and taphonomic analysis of a columnar coralline algal build-up from SE Sicily

Unusual “coralligenous” build-ups were found at the infra–circalittoral boundary from SE Sicily. They are columnar, rising up perpendicularly from the bottom, and consist of an inter-growth of encrusting algae and invertebrates. A selected build-up and its longitudinal section, together with some thin sections of selected parts, were studied to detail the growth structures, composition and fabric of the trapped sediments. The coralline algae show a succession of “concentric”, “fruticulose” and “foliaceous” structures. The framework delineates cavities bounded by algal thalli, sometimes filled by sediments. The organisms constituting the framework were grouped into functional guilds (Fagerström, 1991): “primary frame builders” (mainly coralline algae); “secondary frame builders” (vermetids, serpulids and some bryozoans); “bafflers” (erect, flexible bryozoans, some sponges and soft algae); “binders” (encrusting bryozoans, some crustose coralline algae and serpulids); “destroyers” (a few endolithic algae, rare sponges and bivalves); “dwellers” (brachiopods, foraminifers, bivalves, small bryozoans and some motile organisms). The distribution pattern of taphonomic features (composition, grain size and distribution of sediments filling cavities; presence of cement and its location; mineralogical composition and crystal size), together with interactions between growth structures and functional guilds, allowed one to distinguish three taphofacies, each recording different environmental conditions. They have been tentatively related with cyclic sequences testifying to local hydrodynamic regimes, with subsequent slighter and stronger intensity phases, during the last 2200 years.     

The larval and juvenile ecology of the temperate octocoral Alcyonium siderium Verrill. I. Substratum selection by benthic Larvae

The Alcyonacean octocoral Alcyonium siderium Verrill forms dense stands on vertical rock walls (6–18 m subtidal) along the coast of Massachusetts and further north into the Gulf of Maine. Larvae are brooded until their release as crawling demersal planulae in August. Each colony produces several hundred larvae; most crawl onto the nearby substratum although others may be carried away by wave surge.Experimental enclosures were used to examine substratum selection in the field. Planulae metamorphosed in greater numbers than expected by percent cover on the fleshy red crustose alga Waernia mirabilis Wilce (proposed). The crustose coralline algae Phymatolithon rugulosum (Foslie) and Lithothamnium glaciale Kjellman were settled on commonly but less often than expected by their percent cover. Bare rock surfaces were also used, but least frequently. The encrusting invertebrate colonies that cover much of the available substratum were never used as sites of settlement and metamorphosis. These results agree with laboratory evidence that settlement and metamorphosis of Alcyonium planulae can be induced by Waernia and by coralline algae.     

Antarctic patterns of shallow subtidal habitat and inhabitants in Wilke’s Land

Studies of east Antarctic marine assemblages on hard substrata are rare. In relation to sea-ice breakout, we assessed benthic patterns of habitat and inhabitants between islands and bays at each of two depths (6 and 12 m) across the Windmill Islands coast. Island sites experience sea-ice breakout in the austral spring, while bay sites typically retain sea-ice cover into the summer and in some places the cover is virtually permanent. Composition of assemblages differed between sheltered bays and exposed islands. Islands were dominated by macroalgae, which also varied with depth. Immediately below the ice–foot zone at 6 m, substratum space were monopolised by foliose red (Palmaria decipiens) and foliose brown (Desmarestia sp.) algae, whereas at 12 m large canopies of Himantothallus grandifolius was abundant. The understorey consisted of a mixture of turfs and encrusting red algae at 6 m, and coralline algae at 12 m. Sheltered bays had large areas of sediment/algal complex and no canopy-forming macroalgae. We found more sponges and hydroids in bays, and more brittle stars around islands. Experiments testing factors that covary with exposure and depth in Antarctica, such as light, sedimentation and ice scour are necessary to determine processes that maintain these striking patterns.     

Calcification by Reef-Building Sclerobionts

It is widely accepted that deteriorating water quality associated with increased sediment stress has reduced calcification rates on coral reefs. However, there is limited information regarding the growth and development of reef building organisms, aside from the corals themselves. This study investigated encruster calcification on five fore-reefs in Tobago subjected to a range of sedimentation rates (1.2 to 15.9 mg cm⁻² d⁻¹). Experimental substrates were used to assess rates of calcification in sclerobionts (e.g. crustose coralline algae, bryozoans and barnacles) across key reef microhabitats: cryptic (low-light), exposed (open-horizontal) and vertical topographic settings. Sedimentation negatively impacted calcification by photosynthesising crustose coralline algae in exposed microhabitats and encrusting foram cover (%) in exposed and cryptic substrates. Heterotrophs were not affected by sedimentation. Fore-reef, turbid water encruster assemblages calcified at a mean rate of 757 (SD ±317) g m⁻² y⁻¹. Different microhabitats were characterised by distinct calcareous encruster assemblages with different rates of calcification. Taxa with rapid lateral growth dominated areal cover but were not responsible for the majority of CaCO₃ production. Cryptobiont assemblages were composed of a suite of calcifying taxa which included sciaphilic cheilostome bryozoans and suspension feeding barnacles. These calcified at mean rates of 20.1 (SD ±27) and 4.0 (SD ±3.6) g m⁻² y⁻¹ respectively. Encruster cover (%) on exposed and vertical substrates was dominated by crustose coralline algae which calcified at rates of 105.3 (SD ±67.7) g m⁻² y⁻¹ and 56.3 (SD ±8.3) g m⁻² y⁻¹ respectively. Globally, encrusting organisms contribute significant amounts of carbonate to the reef framework. These results provide experimental evidence that calcification rates, and the importance of different encrusting organisms, vary significantly according to topography and sediment impacts. These findings also highlight the need for caution when modelling reef framework accretion and interpreting results which extrapolate information from limited data.     

Natural inducers for coral larval metamorphosis

 Coral gametes from Acropora millepora (Ehrenberg, 1834) and from multi-species spawning slicks provided larvae for use in metamorphosis assays with a selection of naturally occurring inducer chemicals. Four species of crustose coralline algae, one non-coralline crustose alga and two branching coralline algae induced larval metamorphosis. However, one additional species of branching coralline algae did not produce a larval response. Metamorphosis was also observed when larvae were exposed to skeleton from the massive coral Goniastrea retiformis (Lamarck, 1816) and to calcified reef rubble, demonstrating metamorphosis is possible in the absence of encrusting algae. Chemical extracts from these algae and the coral skeleton, obtained using either decalcification or simple methanol extraction procedures, also contained active inducers. These results extend the number of crustose algal species known to induce coral metamorphosis, suggest that some inducers may not necessarily be strongly associated with the calcified algal cell walls, and indicate that inducer sources in reef habitats may be more diverse than previously reported. Accepted: 21 May 1999     

Oxygen consumption in Mediterranean octocorals under different temperatures

Ecosystem resilience to climate anomalies is related to the physiological plasticity of organisms. To characterize the physiological response of some common Mediterranean gorgonians to fluctuations in temperature, four species (Paramuricea clavata, Eunicella singularis, Eunicella cavolinii and Corallium rubrum) were maintained in aquaria, in which the temperature was increased every ten days with increments of 2-3 °C, starting at 14 °C, ending at 25 °C. Oxygen consumption, number of open/closed polyps and percentage of necrotic tissue were monitored. All species showed similar activity patterns with increasing temperature. P. clavata and E. singularis showed the highest respiration rate at 18 °C, E. cavolinii and C. rubrum at 20 °C. Above these temperatures, both oxygen consumption and polyp reactivity decreased in all species. The present data confirm a reduction of the metabolic activity in Mediterranean gorgonians during periods of high temperature. At temperatures above 18 °C, the percentage of open polyps (considered as a parameter to evaluate polyps reactivity) decreased, thus mirroring the trend of oxygen consumption. The average values of Q₁₀ indicated that gorgonians have a definite temperature limit over which the metabolism (oxygen consumption) stop to follow the temperature increase. After three days at 25 °C, metabolic activity in E. cavolinii, C. rubrum and P. clavata further decreased and the first signs of necrosis were observed. At this temperature, activity remained unchanged in E. singularis. This species seems to more resistant to thermal stress. The symbiotic zooxanthellae present in this species are likely to provide an alternative source of energy when polyps reduce their feeding activity.     

Community structure of rhodolith-forming beds on the central Brazilian continental shelf

The community structure of rhodoliths beds in the central Brazilian continental shelf was studied under the hypothesis that nongeniculate coralline algae are the major contributors of the individual rhodoliths. Samples were collected from five localities within a single area at 17–18 m depth. At each locality, rhodoliths were collected in 10 random quadrat samples along a 20-m transect. Our results show that dead cores of rhodoliths were significantly composed by nongeniculate coralline red algae rather than bryozoans, corals, or inorganic material. The live outer layers of the rhodoliths are composed mainly of 7 species of nongeniculate red coralline algae (Lithophyllum coralline, L. johansenii, L. depressum, L. stictaeformis, Neogoniolithon brassica-florida, Spongites fruticosus, and Lithothamnion muellerii) associated with other encrusting organisms such as bryozoans, sponges, corals, barnacles, and Peyssonnelia red algae. Significant differences were found in the proportion of Lithophyllum species in relation to other red coralline algae found in this study. Our results show that on the Brazilian continental shelf, the rhodolith-forming species are quite higher in size than in any other studied areas in the world. There was no difference in the proportion of live-to-dead rhodolith materials, suggesting an old bed deposit. Also, the amount of calcium carbonate material in the specimens is relevant to take in account in terms of the CO₂ balance worldwide.     

Spread of introduced Caulerpa species in macroalgal habitats

A short-term field experiment was designed to identify layers of Mediterranean macroalgal assemblage conducive to successful spread of two introduced Caulerpa species (Bryopsidales, Chlorophyta). By manipulation of species presence, three experimental assemblages were obtained: (1) encrusting algae, having removed the turf and erect species; (2) encrusting and turfing algae, having removed erect species; (3) encrusting, turfing and erect algae, that is, unmanipulated assemblages, which served as a control. Fragments of the two introduced species Caulerpa taxifolia (Vahl) C. Agardh and Caulerpa racemosa (Forsskål) J. Agardh were transplanted in each of the three assemblages. Width of the colony, blade density and percentage of the substratum covered by the two species were measured.

The susceptibility of the indigenous community to the spread of Caulerpa species was related to type of assemblage. Blade density and amount of substratum covered by the two Caulerpa species were different between species and generally greater for C. taxifolia than for C. racemosa. Overall, the spread of these species was strongly dependent on the type but not directly on the complexity of the assemblage. Turf was more favourable than encrusting species alone, while the least advantageous habitat was where the macroalgal assemblage is composed of encrusting, turf and erect species. In other words, increased number of species in the assemblage reduces invasion of the Caulerpa species but the type of algae in the assemblage is likely to be more important than number of species. The presence of turf promotes the spread of Caulerpa species.     

Transient Shifts in Bacterial Communities Associated with the Temperate Gorgonian Paramuricea clavata in the Northwestern Mediterranean Sea

Background

Bacterial communities that are associated with tropical reef-forming corals are being increasingly recognized for their role in host physiology and health. However, little is known about the microbial diversity of the communities associated with temperate gorgonian corals, even though these communities are key structural components of the ecosystem. In the Northwestern Mediterranean Sea, gorgonians undergo recurrent mass mortalities, but the potential relationship between these events and the structure of the associated bacterial communities remains unexplored. Because microbial assemblages may contribute to the overall health and disease resistance of their host, a detailed baseline of the associated bacterial diversity is required to better understand the functioning of the gorgonian holobiont.

Methodology/Principal Findings

The bacterial diversity associated with the gorgonian Paramuricea clavata was determined using denaturing gradient gel electrophoresis, terminal-restriction fragment length polymorphism and the construction of clone libraries of the bacterial 16S ribosomal DNA. Three study sites were monitored for 4 years to assess the variability of communities associated with healthy colonies. Bacterial assemblages were highly dominated by one Hahellaceae-related ribotype and exhibited low diversity. While this pattern was mostly conserved through space and time, in summer 2007, a deep shift in microbiota structure toward increased bacterial diversity and the transient disappearance of Hahellaceae was observed.

Conclusion/Significance

This is the first spatiotemporal study to investigate the bacterial diversity associated with a temperate shallow gorgonian. Our data revealed an established relationship between P. clavata and a specific bacterial group within the Oceanospirillales. These results suggest a potential symbiotic role of Hahellaceae in the host-microbe association, as recently suggested for tropical corals. However, a transient imbalance in bacterial associations can be tolerated by the holobiont without apparent symptoms of disease. The subsequent restoration of the Hahellaceae-dominated community is indicative of the specificity and resilience of the bacteria associated with the gorgonian host.     

ZONATION OF DEEP WATER BENTHIC ALGAE IN THE GULF OF MAINE1

Algal community structure is described for a deep-water rock pinnacle in the Gulf of Maine. Three depth zones of algal dominance were apparent consisting of 1) leathery macrophytes (to 40 m), 2) foliose red algae (to 50 m) and 3) crustose algae (fleshy crusts to 55 m and coralline crusts to 63 m). Microscopic filamentous and erect calcareous algae were also present but inconspicuous. Upright macroscopic filamentous and thin sheet-like forms were not observed on the pinnacle. Sea anemones (Metridium senile) dominated some vertical faces and abrupt prominences in the shallowest regions of the pinnacle (to 24 m) and locally appeared to set the upper vertical limits of kelp and possibly foliose reds. Laminaria sp. formed an open park-like canopy from 24 to 30 m whereas Agarum cribrosum, the deepest kelp, grew as isolated individuals to 40 m. Peyssonnelia sp. and Leptophytum laeve were the deepest occurring fleshy (to 55 m) and calcareous crusts (to 63 m), respectively. The occurrence of these algae at record depths for the Gulf of Maine and for cold water marine environments may be the result of an absence of large herbivores and the high productivity potential of the benthos in these relatively clear waters. By compiling data on depth distribution patterns world-wide, it is evident that the three zone structure of algal morphologies observed in the Gulf of Maine is a global phenomenon.     

Competitive dominance among sessile marine organisms in a high Arctic boulder community

In most hard substrate environments, space is a limiting resource for sessile organisms. Competition for space is often high and is a structuring force within the community. In the Beaufort Sea’s Boulder Patch, crustose coralline red algae are major space occupiers. This research determined if coralline algae were competitively dominant over other sessile organisms. To test this hypothesis, overgrowth was documented in terms of “winners” and “losers” on the contact borders between different species. Crustose corallines occurred in over 80% of the observed interactions but were only winners in approximately half of them. Most frequently, bryozoans, tunicates, and sponges were superior competitors over crustose corallines, while at the same time these invertebrate groups were among the least abundant space occupiers.     

Larval settlement preferences of Acropora palmata and Montastraea faveolata in response to diverse red algae

Settlement specificity can regulate recruitment but remains poorly understood for coral larvae. We studied larvae of the corals, Acropora palmata and Montastraea faveolata, to determine their rates of settlement and metamorphosis in the presence of ten species of red algae, including eight species of crustose coralline algae, one geniculated coralline and one encrusting peyssonnelid. Twenty to forty percent of larvae of A. palmata settled on coralline surfaces of Hydrolithon boergesenii, Lithoporella atlantica, Neogoniolithon affine, and Titanoderma prototypum, whereas none settled and metamorphosed on Neogoniolithon mamillare. Larvae of M. faveolata had 13–25 % settlement onto the surface of Amphiroa tribulus, H. boergesenii, N. affine, N. munitum, and T. prototypum, but had no settlement on the surface of N. mamillare, Porolithon pachydermum, and a noncoralline crust Peyssonnelia sp. Some of these algal species were common on Belizean reefs, but the species that induced the highest rates of larval settlement and metamorphosis tended to be rare and primarily found in low-light environments. The shallow coral, A. palmata, and the deeper coral, M. faveolata, both had increased larval settlement rates in the presence of only a few species of red algae found at deeper depths suggesting that patterns of coral distribution can only sometimes be related to the distribution of red algae species.     

Communities of constructional lips and cup reef rims in Bermuda

Constructional lips and cup reef rims are little studied reef features typical of very turbulent conditions. The relatively low diversity community of these structures consists of three components, and algal mat dominated by Herposiphonia secunda, an encrusting calcareous biota characterised by crustose coralline algae and the vermetid gastropod Dendropoma corrodens, which together are the main hermatypic components, and an endolithic component with very abundant boring sponges, polychaetes, crustaceans and sipunculids. In northerly areas of Bermuda, the hydrozoan Millepora alcicornis is a prominent member of the community while to the south the urchin Echinometra lucunter is abundant. On the south side of Bermuda the community traps sediment which is deposited in the void spaces. The environment shows reduced grazing and this may account for the presence of several relict and rare species.     

Deep-water rhodolith distribution, productivity, and growth history at sites of formation and subsequent degradation

This study provides the first quantitative measures of deep-water (i.e., below scuba depths) rhodolith development, distribution, abundance, and primary productivity at sites of both active formation and breakdown. The 1.27-km² upper platform surface of San Salvador Seamount, Bahamas, ranges in depth from 67 to 91 m and averages 95.8% cover of rhodoliths that contribute an estimated 391 t organic C·yr⁻¹ to deep-sea productivity. The predominant nongeniculate coralline alga of the slope environment has an extremely narrow PI curve (photosynthesis vs. irradiance) of net primary production (0.005) to slightly beyond 0.24 μmol·m⁻²·⁻¹ PAR) suggesting that some deep-water benthic algae may be acclimated to restricted light ranges. Platform areas contain up to fice-deep accumulations (≈45 cm thick) of rhodoliths with their visible, planar (2-D), crustose algal cover (68.5%) composed of 41% Lithophyllum sp., 14.9% average nongeniculate corallines, and 12.6% Peyssonnelia sp. Platform rhodoliths also contain ≈25% average planar cover of the foraminiferan Gypsina sp. overlying the rock-penetrating chlorophyte Ostreobium sp.

On the steep slopes of the seamount, to a depth of 290 m, rhodoliths that have spilled down from the relatively flat platform average 17.4% cover. These nodules tend to be concentrated in fan-shaped deposits that are most prevalent (33.3% cover) on the west side (leeward) of the mount where they are more abundant near the top of the slope than on the other three sides. Cover of living crustose algae on the deeper slope rhodoliths averages only 22.8% and is made up of 14.8% unidentified nongeniculate corallines, 6% Lithophyllum sp., and 2% Peyssonnelia. Gypsina sp. is not an important component of the slope nodules. Biotic overstory on the seamout slopes is greatly reduced relative to the platform, restricted mainly to bedrock, and consists mostly of Halimeda, gorgonians, and sponges along with scattered patches of small frondose algae.

Over platform depths from 67 to 91 m, rhodoliths are fairly uniform in composition and abundance. Ranging from 4 to 15 cm in diameter, with an average of ≈ 9 cm, they are roughly spherical with smooth living surfaces. The rhodoliths spilling down the steep slopes of the seamount to depths below 200 m are characteristically smaller (mean of ≈5 cm diameter), much rougher, and pittend by boring organisms. As shown by cross sections through the centers of the platform nodules, outer, relatively thin (1–3 cm thick), well-preserved envelopes overlie dead laminated crustose layerse. These layers surround much thicker cores of biotically altered carbonate (mostly coralline, foraminiferan, and coral) that have been extensively reworked by boring sponges, algae, polychaetes, and pelecypods. Borings have been infilled with carbonate detritus and are lithified to various degrees ranging from porous to dense and stony.

Radiocarbon dates indicate that the outermost unaltered envelopes that underlie actively growing crusts are 112–880 yr old ( ), while the innermost unaltered layers average 731 ybp (range = 200–1100 ybp). The consistently abrupt transitions from the intact underlying layers of living.     

Competitive dominance by tabular corals: an experimental analysis of recruitment and survival of understorey assemblages

Tabular and staghorn corals of the genus Acropora often form low-diversity stands on shallow coral reefs, presumably due to their rapid growth rate and ability to outcompete understorey assemblages. Coral cover underneath the abundant Indo-Pacific tabular coral, Acropora hyacinthus, was four times lower than on the adjacent substratum on the reef crest at Lizard Island on the northern Great Barrier Reef. We investigated the effect of A. hyacinthus on patterns of recruitment and mortality by placing experimental panels and coral fragments underneath large colonies of A. hyacinthus. After 8 weeks, recruitment of corals, filamentous algae and crustose coralline algae (CCA) underneath A. hyacinthus was 96, 85 and 50% lower, respectively, compared to panels placed in the open. In contrast, recruitment by bivalves and polychaetes was uniform among treatments, while bryozoans recruited four times more abundantly under A. hyacinthus than in the open. Consequently, the low rate of recruitment by corals beneath A. hyacinthus does not appear to be due to a reduction in the delivery of larvae underneath tables. Instead, the disparity between phototrophic and heterotrophic taxa suggests that diminished light levels under A. hyacinthus are partially responsible for the divergence in recruit assemblages. To test the effect of A. hyacinthus on early mortality and growth of established organisms, recruitment panels were placed on the open for 9 weeks then transplanted underneath A. hyacinthus for a further 8 weeks. The survivorship of juvenile corals underneath tables was less than half that of those on control panels on the unshaded reef crest. Furthermore, the abundance of algal turfs and CCA was sharply lower on transplanted panels. In contrast, heterotrophic organisms increased in cover, regardless of treatment. Experimental branch fragments of Acropora intermedia and Pocillopora damicornis also survived poorly following transplantation underneath A. hyacinthus, compared to adjacent, unshaded controls. We conclude that A. hyacinthus is a formidable competitor which can kill neighbouring corals by overgrowing them, and pre-empt future competition by reducing coral recruitment.     

Grazing patterns in Siphonaria gigas (Mollusca,Pulmonata) on the rocky Pacific coast of Panama

The pulmonate limpet Siphonaria gigas is the most abundant molluscan grazer in the mid zone on rocky, wave-exposed shores of the Pacific coast of Panama. Erect macroalgae and sessile invertebrates are rare; crustose algae cover ～90% of the rock. The relative abundance of a common blue-green algal crust (Schizothrix calcicola?) is negatively correlated with Siphonaria's abundance. Large-scale removals of the limpet cause rapid increases in percent cover of Schizothrix and concomitant decreases in other crusts, but no changes in the abundance of erect algae or sessile invertebrates. Removing Siphonaria also (1) increases recruitment of crustose algae and barnacles onto new rock and plexiglass substrata, and (2) decreases the abundance of a calcified form of Schizothrix. Harsh conditions during daytime low tides and foraging by fishes at high tide control the microdistribution of most of this region's mobile and sessile benthic organisms. Wave action and substratum heterogeneity modify these constraints: Siphonaria is rare or absent in sheltered areas, especially on homogeneous surfaces, and is most abundant at wave-exposed sites. However, at extremely wave-beaten sites, Siphonaria and other benthic consumers are rare and ineffective. Crustose algae are reduced in abundance and space is dominated by erect macroalgae and/or barnacles. These normally rare species can outcompete crusts only when thermal or desiccation stress and the effects of benthic consumers and fishes are drastically reduced.     

Alien marine fishes deplete algal biomass in the Eastern Mediterranean

One of the most degraded states of the Mediterranean rocky infralittoral ecosystem is a barren composed solely of bare rock and patches of crustose coralline algae. Barrens are typically created by the grazing action of large sea urchin populations. In 2008 we observed extensive areas almost devoid of erect algae, where sea urchins were rare, on the Mediterranean coast of Turkey. To determine the origin of those urchin-less 'barrens', we conducted a fish exclusion experiment. We found that, in the absence of fish grazing, a well-developed algal assemblage grew within three months. Underwater fish censuses and observations suggest that two alien herbivorous fish from the Red Sea (Siganus luridus and S. rivulatus) are responsible for the creation and maintenance of these benthic communities with extremely low biomass. The shift from well-developed native algal assemblages to 'barrens' implies a dramatic decline in biogenic habitat complexity, biodiversity and biomass. A targeted Siganus fishery could help restore the macroalgal beds of the rocky infralittoral on the Turkish coast.