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The global decline of brown algal forests along rocky coasts is causing an exceptional biodiversity loss. Regardless of conservation efforts, different techniques have been developed for large-scale restoration strategies in the Mediterranean Sea. In this study we tested ex situ pilot restoration of Gongolaria barbata (=Treptacantha barbata) for the first time in Slovenian coastal waters. Healthy apical fronds of the species were collected and the development of recruits on clay tiles was followed under laboratory conditions for 20 days. Despite the experimental difficulties experienced, especially due to the lack of antibiotics to prevent the growth of the biofilm, G. barbata recruits were outplanted in the sea on two concrete plates with 48 tiles each, protected by purpose-built cages to avoid grazing by herbivorous fish. The high survival rate of juveniles after four months in the field (89% of the tiles on the plate that was constantly protected) suggests that outplanting G. barbata is an operable approach for restoration efforts in the northern Adriatic Sea. Our first experiment in Slovenian coastal waters provides new information for the optimization of the best practices during the laboratory cultivation and addresses the early steps of restoration and introduction of young thalli in the natural environment.

High complexity habitat traits (i.e. high heterogeneity and/or size of discrete habitat units) often promote larger abundances of fauna. Sandy and rhodolith sea bottoms are typically interspersed as mosaics within coastal landscapes. The aim of this study was to experimentally assess the effect of two complexity attributes of rhodolith nodules (i.e. their heterogeneity and size) on the abundance and structure of vagile epibenthic assemblages. An early-colonization experiment was set up (July 2016), where experimental units containing rhodolith nodules of varying heterogeneity and size were deployed at two adjacent recipient habitats: a sandy bottom and a rhodolith seabed. After one month, the abundance of fauna colonizing the experimental units was similar in both habitats, but the assemblage structure (i.e. composition) notably differed. Importantly, the heterogeneity, rather than the size, of experimental rhodolith nodules influenced patterns of vagile macrofauna (>0.5 mm) colonization, despite the habitat type where these experimental units were deployed considerably influenced colonization patterns. This result reinforces the idea of the importance of rhodolith tri-dimensional structure, as key influencer on faunal communities of nearshore habitats.

Rhodolith beds (RBs) are bioconstructions characterized by coralline algae, which provide habitat for several associated species. Mediterranean RBs are usually located in the mesophotic zone (below 40 m), and thus are frequently remote and unexplored. Recently, the importance and vulnerability of these habitats have been recognized by the European Community and more attention has been drawn to their investigation and conservation. This study reports the results of an extensive monitoring program, carried out within the Marine Strategy Framework Directive (2008/56/EC), in six sites off the Campania coast (Italy, Mediterranean Sea). New insights were given into the distribution, cover, vitality (i.e., live/dead rhodolith ratio), structural complexity, and coralline algae composition of RBs. Remotely operated vehicles (ROV) investigations allowed the description of several RBs, and the discovery of a RB with rhodolith cover &gt, 65% offshore the Capri Island. Only two sites (Secchitiello and Punta Campanella) showed a very low mean cover of live rhodoliths (&lt, 10%), hence, not being classifiable as RBs. The collected rhodoliths were mostly small pralines (~2 cm), spheroidal to ellipsoidal, with growth-forms ranging from encrusting/warty to fruticose/lumpy. Coralline algae identification revealed a high diversity within each bed, with a total of 13 identified taxa. The genus Lithothamnion dominated all sites, and Phymatolithon calcareum and Lithothamnion corallioides, protected by the Habitats Directive (92/43/EEC), were detected in all RBs.

International audience; Lithophyllum species in the Mediterranean Sea function as algal bioconstructors, contributing to the formation of biogenic habitats such as coralligenous concretions. In such habitats, thalli of Lithophyllum, consisting of crusts or lamellae with entire or lobed margins, have been variously referred to as either one species, L. stictiforme, or two species, L. stictiforme and L. cabiochiae, in the recent literature. We investigated species diversity and phylogenetic relationships in these algae by sequencing three markers (psbA and rbcL genes, cox2,3 spacer), in conjunction with methods for algorithmic delimitation of species (ABGD and GMYC). Mediterranean subtidal Lithophyllum belong to a well‐supported lineage, hereby called the L. stictiforme complex, which also includes two species described from the Atlantic, L. lobatum and L. searlesii. Our results indicate that the L. stictiforme complex consists of at least 13 species. Among the Mediterranean species, some are widely distributed and span most of the western and central Mediterranean, whereas others appear to be restricted to specific localities. These patterns are interpreted as possibly resulting from allopatric speciation events that took place during the Messinian Salinity Crisis and subsequent glacial periods. A partial rbcL sequence from the lectotype of L. stictiforme unambiguously indicates that this name applies to the most common subtidal Lithophyllum in the central Mediterranean. We agree with recent treatments that considered L. cabiochiae and L. stictiforme conspecific. The diversity of Lithophyllum in Mediterranean coralligenous habitats has been substantially underestimated, and future work on these and other Mediterranean corallines should use identifications based on DNA sequences.

Macroalgal forests are one of the most productive and valuable marine ecosystems, but yet strongly exposed to fragmentation and loss. Detailed large-scale information on their distribution is largely lacking, hindering conservation initiatives. In this study, a systematic effort to combine spatial data on Cystoseira C. Agardh canopies (Fucales, Phaeophyta) was carried out to develop a Habitat Suitability Model (HSM) at Mediterranean scale, providing critical tools to improve site prioritization for their management, restoration and protection. A georeferenced database on the occurrence of 20 Cystoseira species was produced collecting all the available information from published and grey literature, web data portals and co-authors personal data. Data were associated to 55 predictor variable layers in the (ASCII) raster format and were used in order to develop the HSM by means of a Random Forest, a very effective Machine Learning technique. Knowledge about the distribution of Cystoseira canopies was available for about the 14% of the Mediterranean coastline. Absence data were available only for the 2% of the basin. Despite these gaps, our HSM showed high accuracy levels in reproducing Cystoseira distribution so that the first continuous maps of the habitat across the entire basin was produced. Misclassification errors mainly occurred in the eastern and southern part of the basin, where large gaps of knowledge emerged. The most relevant drivers were the geomorphological ones, followed by anthropogenic variables proxies of pollution and urbanization. Our model shows the importance of data sharing to combine a large number of spatial and environmental data, allowing to individuate areas with high probability of Cystoseira occurrence as suitable for its presence. This approach encourages the use of this modelling tool for the prediction of Cystoseira distribution and for supporting and planning conservation and management initiatives. The step forward is to refine the spatial information of presence-absence data about Cystoseira canopies and of environmental predictors in order to address species-specific assessments.

Rhodolith beds represent a key habitat worldwide, from tropical to polar ecosystems. Despite this habitat is considered a hotspot of biodiversity, providing a suite of ecosystem goods and services, still scarce quantitative information is available thus far about rhodolith beds occurrence and ecological role, especially in the Mediterranean Sea. This study reports the composition and patterns of distribution of rhodolith assemblages found in four study areas along ca. 860 km of coast in the Central Mediterranean Sea. These rhodolith beds were studied for the first time and significant differences at all spatial scales have been highlighted, documenting the high variability of this habitat. Rhodolith species composition, morphology and distribution have been discussed considering the potential role of environmental factors in driving these patterns. The need for improving their protection is discussed to complement present conservation and management initiatives, particularly in the frame of the EU Marine Strategy Framework Directive.

Connectivity is a fundamental ecological property affecting stability, resilience and recovery of marine populations, in particular in networks of patchy habitats as the Mesophotic Biogenic Habitats of the Northern Adriatic Sea. Specific information on the dispersal behaviour of many species living in these habitats is lacking, thus the connectivity simulations were performed over different pelagic propagules duration and different reproductive seasons of six consecutive years, covering most of the possible dispersal behaviour variability. The connectivity simulations were analysed with graph analysis methods by characterizing each connectivity graph with several structural measures and by identifying sites with greatest individual and group centrality. We tested the hypothesis that geographic proximity of sites cannot be a proxy for the connectivity and that hydrodynamic connectivity is statistically significantly related to the observed beta diversity among the studied sites. Results showed that species with different pelagic propagules duration follow greatly different dispersal dynamics, and that the network of studied outcrops is only partially connected to coastal benthic populations. Our results also made possible a critical appraisal of the current conservation strategies, evidencing gaps in the existing network of protected Mesophotic Biogenic Habitats in the Northern Adriatic Sea. We suggest possible improvements for an efficient preservation strategy of these unique biodiversity hotspots by including a fundamental ecological process, the dispersal connectivity, into the process of decision making for conservation.

Structure and spatial variability of coralligenous assemblages of the Apulian continental shelf in the southern Adriatic Sea were investigated. The area consists of small coralligenous outcrops distributed between 30 and 100 m of depth on coarse detritic or muddy bottom. Photographic samples, obtained through a Remotely Operated Vehicle, were analyzed to evaluate the abundance of the main taxa or morphological groups of macroalgae and sessile invertebrates, and the deposited sediment. A multifactorial sampling design was used to compare assemblages at two depth ranges, the shallow outcrops (between 35 and 45 m) and the deep outcrops (between 60 and 70 m), and to determine the main scales of spatial variation. Assemblages were dominated by encrusting and erect sponges, encrusting and filamentous algae, scleractinians, encrusting and erect bryozoans and the Zoantharia Parazoanthus axinellae. Deep assemblages were characterized by lower alpha and beta diversity, and by the decrease or disappearance of sensitive organisms, such as coralline macroalgae, scleractinians and erect bryozoans, and by the dominance of encrusting sponges, P. axinellae and hydrozoans. Sedimentation was higher on deep outcrops and was suggested to be the main driver of differences between shallow and deep assemblages. The peculiar composition of assemblages found in the Apulian continental shelf provides evidence that these outcrops represent an uncommon coralligenous habitat.

In the Mediterranean Sea, brown algae belonging to the Cystoseira genus play a valuable role as foundation species. Due to evidences of regression/loss of the habitats of these species caused by the interplay of human and climatic disturbances, active restoration measures have been encouraged by EU regulations. In particular, nondestructive restoration techniques, which avoid the depletion of threatened species in donor populations, are strongly recommended. In the framework of the EU project ROCPOP-Life, the first ex situ outplanting experience of Cystoseira amentacea var. stricta has been implemented in the Cinque Terre Marine Protected Area (northwestern Mediterranean). A total of 400 clay tiles, hosting approximately three mm-long germlings of C. amentacea, were fixed to the rocky shore with screws: the tiles were monitored for the next 2 months by photographic sampling, and survival (presence/absence of juveniles on the tiles), cover and growth were assessed. Additional sampling was performed 6 months after tile deployment, after which an unprecedented storm surge severely affected the restoration performance. After 2 months, over 40% of the tiles were covered with Cystoseira juveniles, which reached approximately eight mm in total length. The tiles that survived the storm hosted three to six cm-long juveniles. The high cover (≥25%), assuring moisture and shading, and the appropriate size of the juveniles, to avert micro-grazing, at time of deployment were key to the survival and growth of the outplanted juveniles, increasing the potential for restoration success. Our findings show that outplanting of midlittoral canopy-forming species is a feasible approach for restoration efforts, with particular attention given to the early phases: (i) laboratory culture, (ii) transport, and (iii) juvenile densities. These results are strongly encouraging for the implementation of restoration actions for C. amentacea on a large scale, in light of EU guidelines.

Herbicides are increasingly recognised as sources of water pollution. Glyphosate-based herbicides (GBHs) are widely used because of their low cost and high effectiveness. By measuring the photosynthetic efficiency of Fucus virsoides fronds exposed to a GBH (Roundup® Power 2.0), we investigated the effect of a continuous exposure (6 days) and the potential of recovery after a short exposure (24 h). Both experiments were carried out combining GBH with and without nutrient enrichment, simulating a runoff event. A factorial experimental design allowed us to assess the potential of interactions between GBH and nutrients, which are likely to co-occur in coastal areas. Our results show deleterious effects of GBH at low concentration on F. virsoides, independently from the duration of exposure and the presence of nutrients. The use of herbicides on land might represent an underestimated threat for coastal systems, as suggested by the irreversible impact on the macroalgae Fucus virsoides observed after a short experimental exposure to a common glyphosate-based herbicide

Due to multiple impacts, Cystoseira forests are experiencing a significant decline, which is affecting the ecosystem services they provide. Despite conservation efforts, there is an urgent need to develop best practices and large-scale restoration strategies. To implement restoration actions, we developed an ex situ protocol for the cultivation of Cystoseira. amentacea var. stricta, aimed at reducing the time needed for laboratory culture, thus avoiding prolonged maintenance and minimizing costs. Specifically, we tested the effects of temperature, light and substratum on settlement and growth of early life stages using a factorial experiment. Temperature (20 and 24°C) and photoperiod (15L:9D) were selected to reflect the conditions experienced in the field during the reproductive period. Two light intensities (125 and 250 μmol photons m-2s-1) were selected to mimic the condition experienced in the absence of canopy (i.e. barren-higher light intensity) or in the understory (lower light intensity) during gamete release. The tested substrata were flat polished pebbles and rough clay tiles. The release of gametes and the successive survival and development of embryo and germlings were followed for two weeks. Regardless of the culture conditions, rougher tiles showed higher zygote settlement, but the substrata did not affect the successive development. Zygote mortality after one week averaged 50% and at the end of the second week, embryonic survival was higher under lower light and temperature conditions, which also determined the growth of larger embryos.

Marine bioconstructions are biodiversity-rich, three-dimensional biogenic structures, regulating key ecological functions of benthic ecosystems worldwide. Tropical coral reefs are outstanding for their beauty, diversity and complexity, but analogous types of bioconstructions are also present in temperate seas. The main bioconstructions in the Mediterranean Sea are represented by coralligenous formations, vermetid reefs, deep-sea cold-water corals, Lithophyllum byssoides trottoirs, coral banks formed by the shallow-water corals Cladocora caespitosa or Astroides calycularis, and sabellariid or serpulid worm reefs. Bioconstructions change the morphological and chemicophysical features of primary substrates and create new habitats for a large variety of organisms, playing pivotal roles in ecosystem functioning. In spite of their importance, Mediterranean bioconstructions have not received the same attention that tropical coral reefs have, and the knowledge of their biology, ecology and distribution is still fragmentary. All existing data about the spatial distribution of Italian bioconstructions have been collected, together with information about their growth patterns, dynamics and connectivity. The degradation of these habitats as a consequence of anthropogenic pressures (pollution, organic enrichment, fishery, coastal development, direct physical disturbance), climate change and the spread of invasive species was also investigated. The study of bioconstructions requires a holistic approach leading to a better understanding of their ecology and the application of more insightful management and conservation measures at basin scale, within ecologically coherent units based on connectivity: the cells of ecosystem functioning.

One of the objectives of the BALMAS project was to conduct Port Baseline Biological Surveys of native and non-indigenous benthic flora in 12 Adriatic ports. Samples of macroalgae growing on vertical artificial substrates were collected in spring and autumn 2014 and/or 2015. A total number of 248 taxa, 152 Rhodophyta, 62 Chlorophyta, and 34 Ochrophyta, were identified. Of these, 13 were non-indigenous seaweeds, mainly filamentous macroalgae, that were probably introduced through hull fouling. Some of these taxa had already been described in the study areas, others were recorded for the first time, a few were no longer detected at sites where they had previously been recorded (e.g. Sargassum muticum). Some other NISS reported for the Adriatic Sea, were not collected at any sampling site (i.e. Caulerpa cylindracea, Codium fragile). Possible reasons for the absence of these species are discussed.

Lithophyllum byssoides is a comm on coralline alga in the intertidal zone of Mediterranean coasts, where it produces biogenic concretions housing a high algal and invertebrate biodiversity. This species is an ecosystem engineer and is considered a target for conservation efforts, but designing effective conservation strategies currently is impossible due to lack of information abo ut its population structure. The morphological and molecular variation of L. byssoides was investigated using morphoanatomy and DNA sequences (psbA and cox2,3) obtained from populations at 15 localities on the Italian and Croatian coasts. Lithophyllum byssoides exhibited a high number of haplotypes (31 psbA haplotypes and 24 cox2,3 haplotypes) in the central Mediterranean. The psbA and cox2,3 phylogenies were congruent and showed seven lineages. For most of these clades, the distribution was limited to one or a few localities, but one of them (clade 7) was widespread across the central Mediterranean, spanning the main biogeographic boundaries recognized in this area. The central Mediterranean populations formed a lineage separate from Atlantic samples; psbA pairwise divergences suggested that recognition of Atlantic and Mediterranean L. byssoides as different species may be appropriate. The central Mediterranean haplotype patterns of L. byssoides were interpreted as resulting from past climatic events in the hydrogeological history of the Mediterranean Sea. The high haplotype diversity and the restricted spatial distribution of the seven lineages suggest that individual populations should be managed as independent units.

Coralline red algae are significant components of sea bottom and up to now considered as exclusively marine species. Here we present the first coralline alga from a freshwater environment, found in the Cetina River (Adriatic Sea watershed). The alga is fully adapted to freshwater, as attested by reproductive structures, sporelings and an inability to survive brackish conditions. Morphological and molecular phylogenetic analyses reveal the species belongs to Pneophyllum and is described as P. cetinaensis sp. nov. The marine-freshwater transition most probably occurred during the last glaciation. The brackish-water ancestor was preadapted to osmotic stress and rapid changes in water salinity and temperature. The particular characteristics of the karst Cetina River, such as hard water enriched with dissolved calcium carbonate and a pH similar to the marine environment, favoured colonization of the river by a marine species. The upstream advance and dispersal is facilitated by exceptionally pronounced zoochory by freshwater gastropods. Pneophyllum cetinaensis defies the paradigm of Corallinales as an exclusively marine group.

The protocols available for sampling and monitoring shallow subtidal rhodolith beds (RBs) are inadequate for the deep Mediterranean analogues, and need calibration in order to attain comparable results. After reviewing the present knowledge of the specificities of Mediterranean RBs, and in the framework of the ongoing international effort for their conservation, a two-step approach is suggested for their definition, identification, delimitation, description, and monitoring. Regional mapping should be improved, and RBs should be identified and delimited as those areas of the sea floor with >10% cover of live rhodoliths over a minimum surface of 500 m2, on 1:10000 scale. More detailed scales (at least 1:1000) should be used for monitoring selected RBs, in order to detect significant changes through time. Beside location and areal extent, the description of RBs should include the occurrence of macroscopic sedimentary structures of the sea floor, thickness of live cover, mean percentage cover of live thalli and surface live/dead ratio, cover of dominant morphologies of rhodoliths (simplified on a ternary diagram), and volumetrically important calcareous algal species. For the purpose of assessment of the ecological status and the evaluation of human-induced impacts, quantitative data about community composition are required. The comparative assessment of ecological status and the identification of RBs of high conservation value for special protection should consider the natural geographic and seasonal/annual variability of RBs. Copyright © 2015 John Wiley & Sons, Ltd.

Phymatolithon calcareum is a species of non-geniculate coralline red algae that commonly forms maerl beds. Tetrasporangial thalli of this species are characterized by unattached growths (rhodoliths), which develop as a branched system without a central nodule, while the attached crustose forms are uncommon and are known only from Brittany (Atlantic coasts of France). In this study we confirm with molecular data the existence of attached crustose tetrasporangial thalli and describe for the first time epilithic, fruticose tetrasporangial specimens from the Mediterranean Sea (Vis Island, Croatia). Since the identification of coralline algal species is difficult due to the high morphological plasticity of the group, molecular analyses based on the nuclear nSSU and the plastidial psbA markers were necessary to correctly identify the specimens. Our study shows that P. calcareum tetrasporangial thalli have the ability to grow both unattached and attached, and we argue this to be in relation to the environmental conditions found at different depths. This ability permits the alga to colonize different habitats of the coastal environment.