Calcareous Red Algae Research Articles

Controls on mesophotic carbonate facies and sediment distribution across the Maltese shelf, central Mediterranean Sea

Although ~ 20% of global carbonate production occurs on extra-tropical carbonate depositional systems, our understanding of these environments still lags behind that of tropical ones. The Maltese shelf in the central Mediterranean offers an opportunity to study in situ facies distribution and the factors controlling it in a light-dominated setting. The investigated region of the Maltese shelf visually exhibits three main depositional environments: seagrass meadows, sand flats and rhodolith and maerl beds. While visually distinctive, the grain composition of the sediments does not provide a clear differentiation of the three environments but rather a gradient. This gradient is marked by increasing grain size with water depth, a transition from green to red calcareous algae and an increase in the fraction of low magnesium calcite of total carboantes. While some of these features can be explained by changes in light availability, other factors are also in play. Baffling by seafloor vegetation and currents, storms and internal waves inducing sediment reworking appear to play important roles in governing the sediment texture and composition across the Maltese shelf. The role of seagrass meadows in regulating production and accumulation rates of carbonates appears to be of greater importance in Mediterranean C-type carbonate factories than in southern Atlantic ones and this could be an important marker to identify them in the geological record.

How the Ecology of Calcified Red Macroalgae is Investigated under a Chemical Approach? A Systematic Review and Bibliometric Study.

Characteristics such as calcareous morphology and life cycle are used to understand the ecology of calcified rhodophytes. However, there is limited information regarding their chemical profiles and biological activities. Therefore, a systematic review (PRISMA) was conducted to assess the influence of the chemistry of calcareous rhodophytes on ecological interactions in the marine environment. The keywords used were: ["Chemical AND [Ecology OR Interaction OR Response OR Defense OR Effect OR Cue OR Mediated OR Induce]"] AND ["Red Seaweed" OR "Red Macroalgae" OR Rhodophy?] AND [Calcified OR Calcareous] in Science Direct, Scielo, PUBMED, Springer, Web of Science, and Scopus. Only English articles within the proposed theme were considered. Due to the low number of articles, another search was conducted with three classes and 16 genera. Finally, 67 articles were considered valid. Their titles, abstracts, and keywords were analyzed using IRaMuTeQ through factorial, hierarchical and similarity classification. Most of the studies used macroalgae thallus to evaluate chemical mediation while few tested crude extracts. Some substances were noted as sesquiterpene (6-hydroxy-isololiolide), fatty acid (heptadeca5,8,11-triene) and dibromomethane. The articles were divided into four classes: Herbivory, Competition, Settlement/Metamorphosis, and Epiphytism. Crustose calcareous algae were associated with studies of Settlement/Metamorphosis, while calcified algae were linked to herbivory. Thus, the importance of chemistry in the ecology of these algae is evident,and additional studies are needed to identify the substances responsible for ecological interactions. This study collected essential information on calcified red algae, whose diversity appears to be highly vulnerable to the harmful impacts of ongoing climate change.

Enhancing strawberry quality and resistance to Botrytis cinerea using calcareous seaweed-derived Ca2+/carrageenan extracts

The dilute HCl pretreatment of the calcareous red alga, Liagora viscida, either at room temperature or 100 °C, produced Ca2+/carrageenan-containing extracts. Dipping strawberries in these extracts for 10 s demonstrated their ability to maintain various quality and nutritional traits during cold storage. The weight loss of the fruits after 7 days at 4 °C was reduced to 2.48–2.63 % in the treatments, compared to 3.26 % in the control. The treatments were effective in retaining the contents of vitamin C, titratable acidity, and phenolics, compared to marked reductions in the control. Additionally, the levels of anthocyanins, flavonoids, and the total antioxidant capacity were enhanced during cold storage. The treated fruits also showed higher chitinase activities, while the activities of cellulase, polygalacturonase, and pectin methylesterase (PME) showed remarkable reductions. The application of the extract derived at 100 °C reduced the decay caused by Botrytis cinerea decay to 34 % and 63.26 % after 3 and 4 days of infection, respectively, compared to 55.09–97 % in the untreated control. This reduction was concomitant with higher anthocyanins, flavonoids, and chitinase, and lower cellulase and PME in this treatment. The levels of malondialdehyde indicated less stressed fruits during both cold storage and fungal infection. Overall, the treatments promoted systemic resistance in the fruits, enhancing their ability to defend against pathogens. These results suggest the promising utilization of calcareous macroalgae as a sustainable biomass for developing a cost-effective and eco-friendly preservative to extend the shelf-life of strawberries during cold storage and enhance their resistance to pathogens.

Morphological insights into the three-dimensional complexity of rhodolith beds

The ecological importance of rhodolith beds stems from their role as structurally complex three-dimensional habitat formed by free-living red calcareous algae. Their structural singularity is due to the great variety of complex and branching morphologies exhibited by rhodoliths that create interstitial spaces and increase their surface area. This increases the ecological niches for cryptofauna and provide refuge for a high number of organisms, which is why rhodolith beds are considered biodiversity hotspots. In this work, we studied a rhodolith bed located in the Menorca Channel, formed by several species of red calcareous algae that exhibit a great variety of morphologies and form an extensive and heterogeneous habitat. This study explored the morphological diversity of the rhodolith bed, comparing the ‘Core Habitat’ (within the center of the bed with the highest densities of rhodoliths) with the boundaries or ‘Adjacent Habitat’ where rhodolith density was lower. Our results show that all rhodolith growth forms (branched, pralines and boxwork) in the Core Habitat had higher interstitial space and were larger than the ones from adjacent zones. Moreover, we explored the three-dimensional techniques to study the morphological characteristics that have historically been studied in two dimensions. This study contributes to the knowledge of morphological diversity in well-preserved rhodolith beds from continental shelves in the western Mediterranean Sea and reinforces the use of three-dimensional measurements, specifically the interstitial space of branched rhodoliths, to provide more accurate data on habitat complexity.

Crustose red algae in deep time environments: Palaeoecological insights from northeastern India and Türkiye (Turkey)

Crustose red calcareous algae are key organisms in benthic ecosystems worldwide with critical functions like reef-building and substrate stabilization. Coralline algae thrived as major carbonate producers, with corals and/or larger benthic foraminifera (LBF) in numerous shallow-marine Tethyan carbonate platforms from the early Palaeogene to the Neogene. The Palaeocene–Eocene limestone successions in the Jaintia and Khasi Hills, Meghalaya, northeastern India, encompass two principal community types — algal-foraminiferal and coralline algal, with no associated colonial corals, while the Oligocene–Miocene carbonates from the Sivas basin and Siirt province in Türkiye are characterized by an algal-foraminifera assemblage with bryozoans, corals, and molluscs as other noteworthy components pertinent to the carbonate platform environment. We found that the diversity and ecology of these algal communities were influenced by systematic structure of the algal assemblage, sediment input, substrate stability, water depth, and hydrodynamic energy. The algal assemblages from different epochs and varying localities diverge based on the sedimentary regime, environmental settings, and the predominant taxa. The climatic transition from the Palaeocene–Eocene greenhouse to the Oligocene–Miocene icehouse corresponded to a shift from crustose algal assemblages dominated by Sporolithales to assemblages dominated by Hapalidiales.

Coral Primary Polyp Settlement on Crustose Coralline Algae from India

Crustose coralline algae (CCA) are calcareous red algae belonging to the diverse group Rhodophyta. They are known to play numerous important ecological roles in the coral reef ecosystems such as various invertebrate larval settlement, cementing the reef together, participating effectively in primary productivity. One of the most important ecological roles these algae play, are the settlement and metamorphosis of coral larvae. Though lots of observations have been made on a juvenile coral being settled on CCA, there has not been any study on the initial settlement of coral larvae on CCA. The present study was carried out in the intertidal regions of South Andaman district. During intertidal survey, a single thin layered CCA was observed to house four different coral recruits which had settled and metamorphosed to form a single polyp coral.

Shallow-water carbonate facies herald the onset of the Palaeocene-Eocene Thermal Maximum (Hazara basin, Northern Pakistan)

We investigate the Palaeocene succession of the Hazara Basin (Northern Pakistan) to better understand the impact of climate change on marine carbonate-producing organisms. These shallow-water carbonates, deposited during the Late Palaeocene, before the onset of the Palaeocene-Eocene Thermal Maximum, were studied using a quantitative approach to highlight changes in the skeletal assemblage. We recognise a decrease in the abundance of colonial corals and green calcareous algae and an increase in larger benthic foraminifera and red calcareous algae from the early Thanetian to the late Thanetian. Increasing temperatures may represent a plausible cause for the decline of the more sensitive colonial corals in favor of the more tolerant larger benthic foraminifera. A similar pattern is observed in most successions deposited along the margins of the Neotethys Ocean, suggesting a connection with the Late Palaeocene environmental changes that heralded the PETM hyperthermal event. Our stratigraphic analysis of the Hazara Basin strata suggests that the biotic turnovers occurred during the Palaeocene – Eocene transition started already before the onset of the Palaeocene Eocene Thermal Maximum as recorded by the geochemical proxies.

Distribution of rhodolith beds and their functional biodiversity characterisation using ROV images in the western Mediterranean Sea

Underwater biogenic habitats composed of unattached calcified red algae, named as rhodolith or maërl beds, may extant either alive or dead, over the seabed. The accumulation of rhodoliths constitute three-dimensional structured biogenic habitats that harbour high diversity of benthic organisms. In the Mediterranean Sea, rhodolith beds can be found between ca. 50 and 100 m, increasing diversity of the continental shelf habitats and their conservation value. Despite the remarkable relevance of these habitats, information regarding their spatial distribution in the western Mediterranean is scarce. In addition, these habitats are threatened by a range of anthropogenic activities and by climate-driven changes. In this study, we explored areas with rhodoliths' occurrence from the north to the south of the east coast of Spain. By feeding video recording data into a spatial distribution model, we assessed which biophysical drivers: (i) shape the spatial variation in the abundance of rhodoliths and (ii) define the assemblages of the biological traits and functional richness of the associated epibenthic fauna recorded by video. In addition, we examined the impact of fishing activities on these habitats. Our results evidenced that ‘Depth’ and ‘Temperature’ were important environmental factors explaining rhodoliths variation, and we defined their optimal distribution range in the western Mediterranean. The biological traits approach showed significant effects of the geographical location of rhodolith beds and their small-scale spatial heterogeneity on the functional richness index. Indeed, the lowest functional richness value was observed in the area closer to trawl fishing which could be related to habitat degradation due to trawling. This study contributes to the knowledge of deep-water rhodolith beds in the western Mediterranean and reinforce the importance of rhodolith beds in continental shelves as these constitute heterogeneous seabed habitats that harbour a high species and functional diversity.

An Overview of Rhodoliths: Ecological Importance and Conservation Emergency.

Red calcareous algae create bio-aggregations ecosystems constituted by carbonate calcium, with two main morphotypes: geniculate and non-geniculate structures (rhodoliths may form bio-encrustations on hard substrata or unattached nodules). This study presents a bibliographic review of the order Corallinales (specifically, rhodoliths), highlighting on morphology, ecology, diversity, related organisms, major anthropogenic influences on climate change and current conservation initiatives. These habitats are often widespread geographically and bathymetrically, occurring in the photic zone from the intertidal area to depths of 270 m. Due to its diverse morphology, this group offers a special biogenic environment that is favourable to epiphyte algae and a number of marine invertebrates. They also include holobiont microbiota made up of tiny eukaryotes, bacteria and viruses. The morphology of red calcareous algae and outside environmental conditions are thought to be the key forces regulating faunistic communities in algae reefs. The impacts of climate change, particularly those related to acidification, might substantially jeopardise the survival of the Corallinales. Despite the significance of these ecosystems, there are a number of anthropogenic stresses on them. Since there have been few attempts to conserve them, programs aimed at their conservation and management need to closely monitor their habitats, research the communities they are linked with and assess the effects they have on the environment.

New data on red Visean calcareous algae of the Dnipro-Donetsk Depression: systematic composition and facial confinement


 
 
 As a result of the study of the core material of 540 thin sections from the boreholes of the northern marginal part and the paraxial part of the Dnieper-Donetsk Depression (DDD), a complex of fossil red calcareous algae including 14 species of 7 genera, as well as 2 species belonging to 2 genera identified for the first time for the Visean deposits of the DDD has been discovered. The studied carbonate deposits represent the Hlybokynskyi, Sukhynskyi, Stylskyi, Donetskyi and lower part of Mezhivskyi horizons of the Don-Dnipro Rift (DDR). It has been found that the greatest diversity of the taxonomic composition of fossil red calcareous algae in the DDD is observed in the deposits of the Sukhynskyi, Stylskyi and the most part of Donetskyi horizons, (XIII, XIIa, XII microfaunistic horizons of the DDD). It has been established that the limestones of the DDD paraxial zone dated to the uppermost Donetskyi and lower Mezhivskyi horizons (XI MFH of the DDD) are represented mainly by foraminiferous-polydetrital and algal wackestones, less often by wacke-packstones. The association of calcareous algae with dark ferruginous fragments indicatesthat the deposits formed in relatively deep conditions of the outer shelf. Carbonate sediments of the most part Stylskyi and Donetskyi horizons (XIIa, XII MFH of the DDD) of the northern marginal zone of the DDD are represented mainly by polydetrital wackestones, sometimes they are spicular, sometimes iron-bearing, which makes it possible to assume that these deposits, like the previous ones, belong to the relatively deep-water facies of the open platform, with moderate and reduced water circulation. Carbonate deposits of the Sukhynskyi and lowermost Stylskyi horizons (XIII MFH of the DDD) of the paraxial and northern rift side zones are represented mainly by polydetrital and fine-detrital-algal wackestones with remains of ostracods, mosses, spicules, brachiopod shells, fragments of Stacheaceae, green tubular algae of the family Palaeoberesellaceae Mamet et Roux. The predominance of green siphonocladal algae in the algal association indicates a calm hydrodynamic regime below the tide level as well as the depth of the basin, which mostly does not exceed 3-5 m, a small fluctuation in the depth of the sea basin, good illumination, and thermal conductivity of the sea basin. Carbonate deposits of the Hlybokynskyi horizon (XIV MFH of the DDD) exposed by the well 10 Skorobahatky in the northern marginal part of the DDD, are represented by organogenic, detritus-muddy packstones and wackestones with remains of bivalves, foraminifera, crinoids, ostracods, green, tubular algae of the family Palaeoberesellaceae Mamet et Roux with a transition in the upper part to sandy grainstone. This indicates that sediments were formed in the shallow water conditions under constant wave action, near the base of this action that led to the washout of clay mass.
 
 

A “cool-water”, non-tropical, mixed volcaniclastic–carbonate ramp from the Early Cretaceous of southern Chile (45°40’S)

The Aysén-Río Mayo Basin was a back-arc/marginal basin developed in southwestern South America (43°–47°S) between the Tithonian–Aptian. Its sedimentary fill corresponds to the Coyhaique Group, which represents a transgressive–regressive succession. Six lithofacies and five microfacies were defined for three outcrops exposed south of Coyhaique (45°40’S). The outcrops have a mixed calcareous–volcaniclastic composition and were assigned to the early transgressive Toqui Formation, i.e., lowermost part of the Coyhaique Group. These mixed rocks comprise bioclastic–volcaniclastic conglomerate, gravelly allochemic sandstone, and gravelly–sandy allochem limestone. Bedding is sharp to amalgamated, sometimes rippled, depicting a wave- and storm-influenced, mixed inner- to mid-ramp. The ramp developed over a Valanginian, active volcanic terrain (Foitzick Volcanic Complex), source of the volcaniclastic sediments. Limestones are rich in reworked bioclasts, and controlled by calcitic organisms including gryphaeid oysters, non-geniculate red algae, and echinoid fragments, defining a heterozoan association (“maerl”-like sediments); less frequent are ahermatypic corals, serpulids, and carbonized wood. Based on their inferred paleolatitude (south of 45°–50°S), fossil assemblage (heterozoan), and kind of carbonate platform (ramp-type), these calcareous rocks of the Toqui Formation depict a “cool-water” (sensu lato), non-tropical setting. The fossil assemblage includes oysters (Aetostreon spp.), and abundant calcareous red algae attributed to Archamphiroa jurassica Steinmann (1930), a taxon previously known from the upper Tithonian Cotidiano Formation of Argentina. A. jurassica is here reported for the first time for the Lower Cretaceous of Chile, suggesting a broader upper Tithonian—Valanginian-Hauterivian? range for the species. The facies model presented here contrasts with the depositional environments depicted for correlative reefal rocks in Argentina (Tres Lagunas Formation), which reflect a “warm-water” setting. In the Aysén-Río Mayo Basin, the influence of sea-water key physical variables in the carbonate sedimentation, as well as the position and hydraulic regime of the carbonate platforms within the basin, and their interaction with the volcanism are still unclear.

Morpho-anatomical, and chemical characterization of some calcareous Mediterranean red algae species

Climatic changes are anticipated to have a detrimental effect on calcifying marine species. Calcareous red algae may be especially vulnerable to seasonal variations since they are common and essential biologically, but there is little research on the morpho-anatomical, and chemical characterization of such species. This study conducted the seasonal investigation of the three dominant Mediterranean calcified red algae. Morphological and 18S rRNA analysis confirmed the identification of collected species as Corallina officinalis, Jania rubens, and Amphiroa rigida. In general, C. officinalis was represented in the four seasons and flourishing maximum in autumn (70% of total species individuals). While J. rubens species was represented in winter, autumn, and spring and completely absent in summer. A. rigida was abundant only in the summer season by 40%. A full morphological and anatomical description of these species were examined, and their chemical compositions (carbohydrate, protein, lipid, pigments, and elements content) were assessed in different seasons, where carbohydrates were the dominant accumulates followed by proteins and lipids. Pearson correlation analysis confirmed a positive correlation between salinity level and nitrogenous nutrients of the seawater with the pigment contents (phycobiliproteins, carotenoids, and chlorophyll a) of the studied seaweeds. The results proved that calcified red algae were able to deposit a mixture of calcium carbonates such as calcite, vaterite, calcium oxalate, calcite-III I calcium carbonate, and aragonite in variable forms depending on the species.

Seasonality and phenology of an epiphytic calcareous red alga, Hydrolithon boreale, on the leaves of Posidonia oceanica (L) Delile in the Turkish water.

Epiphytes on Posidonia oceanica play a crucial role for determination of the ecological status of marine environment in time and space besides the seagrasses alone. The study was aimed to estimate the spatiotemporal ecological status linked to variation in biometry of an epiphytic micro-calcareous red alga, Hydrolithon boreale, found on leaves of the meadow with the exclusive environmental parameters along the entire Turkish coast of the Mediterranean Sea. Collection of Posidonia oceanica samples was conducted at 64 stations in winter (December 2018-January 2019) and 112 stations in summer (June-July 2019) by SCUBA (0.4 × 0.4m of a quadrate frame) in the infralittoral zone along the entire Turkish Mediterranean coast surrounded by the siliciclastic Taurus Mountain Range which favor growth of epiphytic micro-calcareous red algae. Percent occurrence of the epiphyte changed seasonally-lower in winter (25%) than in summer (44%). The epiphyte which is an indicator and sensitive to undisturbed marine area grew up well to 5mm in diameter, 0.35mm in thickness of the crust size, and was populated up to 1006 ind/m2 in summer owing to the increased utilization of the carbonate by the epiphyte with the increased water temperature. The size was contrasted to the density (abundance and biomass) in space. The biometry was significantly dependent on the siliciclastic-carbonate deposition as inferred from SiO4-Si of the water in relation to the leaf area index (LAI) of P. oceanica. Therefore, this deposition induced specimens to grow in size, followed by the reduced density concerning the N-based nutrient of the water. Further major environmental parameters which negatively affected the biometry were pH and total suspended matter of the water, analogous to turbidity. Of the trace elements, Ni was negatively correlated with the biometry, whereas the LAI was however positively correlated with all the anthropogenic-sourced trace elements (V, Cu, Zn, As, Cd, and Pb) in the leaves. Of the bottom types, the calcite rock had a higher density than the other soft bottoms in contrast to the size of the epiphyte. Future studies could be based on the present study for determination of the ecological status regarding two dominant epiphytes on leaves of two seagrasses (H. boreale on P. oceanica and partly Pneophyllum fragile on Cymodocea nodosa) found in the different environments and substrates in space and time.

Calcification traits for cryptic species identification: Insights into coralline biomineralization.

Calcareous red algae are foundation species and ecosystem engineers with a global distribution. The principles governing their calcification pathways are still debated and the morphological characters are frequently unreliable for species segregation, as shown by molecular genetics. The recent description of the new species Lithophyllum pseudoracemus, previously undetected and morphologically confused with Lithophyllum racemus, offered a challenging opportunity to test the effectiveness of microanatomy and ultrastructural calcification traits as tools for the identification of these two species, for integrative taxonomy. High resolution SEM images of molecularly identified samples showed that the different size of the perithallial cells and the features of the asexual conceptacle chambers may contribute to the separation of the two species. The two species share the same crystallite morphology in the primary and secondary cell-wall calcification, as previously described in other species belonging to the same clade. However, the perithallial secondary calcification was significantly thicker in L. racemus than in L. pseudoracemus. We described a granular calcified layer in the innermost part of the cell wall, as a putative precursor phase in the biomineralization and formation of the secondary calcification. The hypothesis of different pathways for the formation of the primary and secondary calcification is supported by the observed cell elongation associated with thicker and higher Mg/Ca primary calcification, the inverse correlation of primary and secondary calcification thickness, and the absence of primary calcification in the newly formed wall cutting off an epithallial cell from the meristem.

Palaeocene to Miocene southern Tethyan carbonate factories: A meta‐analysis of the successions of South‐western and Western Central Asia

AbstractOne hundred and forty‐four published successions of shallow‐water carbonates, deposited between the Palaeocene and the Miocene, from the Levant to the Himalayas, have been re‐analysed using a standardised approach to investigate the distribution of carbonate facies and carbonate‐producing organisms. Large benthic foraminifera were found to be the volumetrically most important group of carbonate producers during the whole period, with a peak in abundance during the Eocene. Colonial corals are relatively abundant during the Palaeocene and Miocene, their abundance peaks during the Oligocene and has a minimum during the Eocene. Red calcareous algae have a similar pattern although their peak in abundance covers both the Oligocene and Miocene. Green calcareous algae decrease from the Palaeocene onward. Facies related to very shallow and/or restricted marine conditions peak during the Miocene and in particular during the Aquitanian. Both the pattern of large benthic foraminifera and of colonial corals seems to be related to temperature, with warm periods favouring the former group and cool periods the latter group. Red calcareous algae display a pattern similar to that of colonial corals suggesting that the periods favourable for one group are, on a large scale, also favourable for the other. The progressive decrease of green calcareous alga could be tentatively related to a preservation bias connected to the transition from Palaeogene assemblages that included presumably calcitic taxa of green algae to Neogene assemblages entirely constituted by aragonitic taxa with limited preservation potential. The Aquitanian peak in facies related to very shallow and/or restricted marine conditions is most likely connected to the progressive narrowing of the Tethys related to the collision between Arabia and Eurasia. These results denote an overall agreement between the abundance of the various types of shallow‐water carbonate facies and large‐scale environmental and geological processes, highlighting the potential for palaeoenvironmental reconstruction locked in the shallow‐water record.

The Main Builders of Mediterranean Coralligenous: 2D and 3D Quantitative Approaches for its Identification

Along the Mediterranean Sea shelf, algal reefs made of crustose coralline algae and Peyssonneliales are known as Coralligenous. It ranks among the most important ecosystems in the Mediterranean Sea because of its extent, complexity, and heterogeneity, supporting very high levels of biodiversity. Descriptive approaches for monitoring purposes are often aimed at assessing the surficial ephemeral canopy, which is sustained and controlled by the occurrence of the long-lasting rigid structure at the base. This practice led to the non-univocal definition of Coralligenous, sometimes indicated as “animal Coralligenous” because of the surficial dominance of these components. The quantitative assessment of the builders that actively build up the persistent structure through geological time is therefore a fundamental topic. We collected two discrete coralligenous samples in front of Marzamemi village (Sicily, Ionian Sea), the first from an area of a dense coralligenous cover (- 37 m) and the second one from an area with sparse build-ups (- 36 m). By using image analysis and computerized axial tomography, we distinguished and quantified the different components both on the surface and inside the framework. In both cases, our results confirm the primary role of crustose coralline algae as major builders of the Mediterranean Coralligenous, this aspect matching with the evidence from the Quaternary fossil record. We suggest that the role of encrusting calcareous red algae in the Coralligenous should be considered in conservation and management policies.

Macrofauna Associated With a Rhodolith Bed at an Oceanic Island in the Eastern Tropical Pacific (Isla del Coco National Park, Costa Rica)

Rhodoliths are round calcareous red algae that form extensive beds and associated with them are a diverse suite of species. Rhodolith beds are among the least known coastal–marine ecosystems, and even less is understood about their associated flora and fauna. Here, we present an evaluation of the biodiversity larger than 500 μm associated with rhodoliths at Isla del Coco National Park, Costa Rica, an oceanic island in the Eastern Tropical Pacific, 500 km offshore of the mainland. This research determined the influence of rhodolith degree of aggregation (distance among individual rhodolith) as well as rhodolith complexity, volume, and mass in relation to the diversity, composition, and biomass of the associated fauna. A total of 145 taxa were collected in 60 rhodolith samples. Arthropods, polychaetes, and mollusks were the dominant taxa in terms of richness, and crustaceans + acari represented >50% of the total abundance. Five potentially new species were collected in this study. Collections identified 31 new records, with 20 of them being newly reported genera for Isla del Coco. Many of the organisms found were juveniles as well as adult stages bearing eggs, demonstrating the importance of this ecosystem. The faunal composition changed along the gradient of rhodolith aggregation. Moderately aggregated rhodoliths (separated by 5 to 10 cm) had the highest diversity, with the highest averages of taxon richness and total numerical abundance and the highest faunal biomass. There were more organisms in more complex rhodoliths; nevertheless, the complexity of the rhodolith did not affect the number of taxa or total organism biomass. Larger rhodoliths, in size and mass, favored higher amounts of taxa and organisms. The rhodolith bed studied is an example of the intermediate disturbance hypothesis because the highest value of biodiversity of invertebrates was at the moderate aggregation level of rhodoliths. In this density of rhodoliths, the fauna was less exposed to hard environmental conditions and prevented dominant species.

The Eocene carbonate platform of the central-western Malaguides (Internal Betic Zone, S Spain) and its meaning for the Cenozoic paleogeography of the westermost Tethys

The Eocene Peñicas (Almería) and Harania (Málaga) stratigraphic sections from the Malaguide Complex (Betic Cordillera, South Spain) belonging to the Mesomediterranean Microplate from the westernmost Tethys (about-35°N and 0° to 5°E according to the Eocene coordinates) have been studied. The Eocene sections cover the Cuisian to middle Lutetian deposits, which show several lithofacies representing shallow marine platform realms. Based on the fossiliferous assemblage, texture and fabrics, eight microfacies related to inner to outer ramp settings were defined. In the inner ramp of the Harania section abundant colonial corals have been recognized. The Eocene deposits are arranged into a transgressive succession composed by three minor transgressive-regressive sedimentary cycles. The Eocene fossiliferous assemblage shows a mixture of photozoan (Larger Bentic Foraminifera –LBF–, green and red calcareous algae and corals) and heterotrophic (mollusks, echinoids, bryozoans, small benthic and planktic foraminifers) elements, suggesting euphotic to mesophotic conditions in oligo-mesotrophic marine warm-waters at low-middle latitudes. This assemblage indicates a transition from photozoan to heterozoan carbonates and in particular a shift towards outer marine ramp settings. During the Early Eocene, the widespread distribution of LBF leads in the Tethyan domains to disappearance or extreme reduction of coral constructions. Nevertheless, abundant corals associated to inner ramp realms have been observed in the Harania stratigraphic section indicating that corals could continue to develop in the westernmost Tethys at the transition to the Atlantic Ocean, in contrast with respect to other Tethyan sectors. Therefore, the Ypresian-Lutetian time-span is a transitional period for the global temperature during which corals locally survived only where optimal ecologic conditions occurred, preferably in marginal contexts, as it seems to have happened in the studied area. • Westernmost Tethys (about-30° N and 0° to 5°E according to the Eocene coordinates) • Cuisian to Middle Lutetian deposits • Shallow marine platform realms • Photozoan elements (LBF, green and red calcareous algae and corals) • Abundant corals associated to inner ramp realms

Characterization of rhodolith beds-related backscatter facies from the western Pontine Archipelago (Mediterranean Sea)

Rhodoliths (nodular calcareous red algae) are considered one of the most important bioengineers in the Mediterranean Sea, making rhodolith beds ecologically relevant ecosystems. On the insular shelf surrounding the western Pontine Archipelago (depth from 43 to 112 m), rhodolith beds were identified through the analysis of an extensive dataset of grab samples and videos to ground-truth the backscatter acoustic facies. Six acoustic facies (low backscatter, dishomogeneous low-backscatter, dishomogeneous high-backscatter, high-backscatter, rocks and high backscatter, and rocks and medium backscatter) were recognized. We studied how rhodoliths characteristics (density, morphotype, size and structure) differently influence the backscatter signature. At the western Pontine Archipelago, rhodolith beds are mainly represented by facies dishomogeneous high backscatter, high backscatter, high backscatter with rocks, and medium backscatter with rocks. The obtained results increase both the knowledge on the heterogeneous structure of such ecologically relevant benthic habitat and highlight the use of distinctive acoustic facies for their identification. Finally, the used approach could be considered a useful method for indirect detection and mapping of rhodolith beds.

Global biodiversity and biogeography of rhodolith-forming species

Unattached nodules of calcareous red algae (Rhodophyta), known as rhodoliths, are widely reported and studied in places that extend from the tropics to polar latitudes. Factors controlling the distribution of the rhodolith-forming species remain poorly understood. A review of the global distribution of present-day rhodolith beds was undertaken, collating information on 106 rhodolith-forming species from 10 families, representing 21 genera distributed through 11 realms: 1) Arctic, 2) Temperate Northern Atlantic, 3) Temperate Northern Pacific, 4) Tropical Atlantic, 5) Western Indo-Pacific, 6) Central Indo-Pacific, 7) Eastern Indo-Pacific, 8) Tropical Eastern Pacific, 9) Temperate South America, 10) Temperate Australasia, and 11) Southern Ocean. The Central Indo-Pacific and Temperate Australasia proved to be the most diverse realms. Of 62 provinces across these realms, the Tropical Southwestern Atlantic, the Mediterranean Sea and the Tropical East Pacific feature the highest diversity of rhodolith-forming species. A significant proportion of the 106 species (14.2%; 15 species) are endemic to a single biogeographic province. Species richness is weakly related to sampling effort (r2=0.573) and unrelated to littoral area (r2=0.012). Even when high latitude provinces are excluded from the analysis, no correlation between species richness and littoral was found (r2 = 0.0005). A wider, evolutionary-time framework revealed that the existence of marine barriers and the geological age of their final emplacement are key elements to explaining compositional differences between the rhodoliths of former contiguous areas (e.g., Pacific versus Atlantic shores of Panama and Costa Rica, in the Central America; eastern Mediterranean Sea versus Red Sea and Gulf of Aden). Finally, we propose that the lower diversity of the rhodolith-forming species in the tropical Pacific Ocean when compared to the Atlantic Ocean (23 versus 33 spp.), may be linked to the higher abundance of corals and coral reefs in the Pacific, which act as competitors with coralline algae for space.