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.     

Range extension and morphological characterization of rhodolith-forming species (Corallinales,Rhodophyta) from shallow water in the Mexican South Pacific

Living rhodolith beds are widely distributed along the Eastern Pacific ocean. Despite their widespread distribution, little is known about the rhodolith-forming species from shallow water in the Mexican South Pacific. Many taxonomic and morphological studies about rhodoliths have been carried out in the Gulf of California, where the forming species belong to the Hapalidiaceae and Corallinaceae families. This paper is the first report on the occurrence of the rhodolith-forming Hapalidiaceae species Lithothamnion muelleri and Phymatolithon repandum at three sites in the Mexican South Pacific. The branch density, maximum length and sphericity were measured for each determined species. Rhodoliths were distributed between 4 and 6 m depth, but differences in the branch density between species and sites were not found. Finally, the present record of L. muelleri fills the gap in the species distribution along the Eastern Pacific ocean, while the record of P. repandum is the first of the species in the region.     

Arctic rhodolith beds and their environmental controls (Spitsbergen, Norway)

Coralline algae (Corallinales, Rhodophyta) that form rhodoliths are important ecosystem engineers and carbonate producers in many polar coastal habitats. This study deals with rhodolith communities from Floskjeret (78°18′N), Krossfjorden (79°08′N), and Mosselbukta (79°53′N), off Spitsbergen Island, Svalbard Archipelago, Norway. Strong seasonal variations in temperature, salinity, light regime, sea-ice coverage, and turbidity characterize these localities. The coralline algal flora consists of Lithothamnion glaciale and Phymatolithon tenue. Well-developed rhodoliths were recorded between 27 and 47 m water depth, while coralline algal encrustations on lithoclastic cobbles were detected down to 77 m water depth. At all sites, ambient waters were saturated with respect to both aragonite and calcite, and the rhodolith beds were located predominately at dysphotic water depths. The rhodolith-associated macrobenthic fauna included grazing organisms such as chitons and echinoids. With decreasing water depth, the rhodolith pavements were regularly overgrown by non-calcareous Polysiphonia-like red algae. The corallines are thriving and are highly specialized in their adaptations to the physical environment as well as in their interaction with the associated benthic fauna, which is similar to other polar rhodolith communities. The marine environment of Spitsbergen is already affected by a climate-driven ecological regime shift and will lead to an increased borealization in the near future, with presently unpredictable consequences for coralline red algal communities.     

Heterozoan carbonate-enriched beach sand and coastal dunes—with particular reference to rhodoliths,Dirk Hartog Island,Shark Bay,Western Australia

This is a preliminary interdisciplinary study on the enrichment of heterozoan carbonates on Dirk Hartog Island, Shark Bay, Western Australia, with particular reference to rhodolith (free-living non-geniculate) coralline algae. The current study aims to investigate the geological impact of shallow-water rhodoliths in Shark Bay, as well as fill critical information gaps on the biogeographical distribution of rhodoliths in Australia. We analyzed the composition of sand from eight sites (totaling 21 beach and sand dune samples) on the eastern (windward) shore of the island, and investigated the origin of the coralline algal grains. Heterozoan carbonates (shell, geniculate coralline algae grains, and rhodolith grains) together comprised 3–84% of the carbonate-enriched beach and dune sand samples. While shell fragments often comprised the highest percentage (up to 73%), rhodolith grains (up to 27%) were found in 12 of 21 samples, with rhodolith grains also occurring in two dune samples. Geologically, the study has shown that rhodoliths and rhodolith beds are important shallow-marine habitats in Shark Bay, with a proven capacity to enrich beach/dune sands in Shark Bay and potentially other areas along the Australian coast. Biogeographically, the study confirmed the presence of a previously undescribed shallow rhodolith bed in Shark Bay (the first bed documented on the Western shore) with the possibility of a third bed near Sandy Point on Dirk Hartog Island. It also confirmed the presence of rhodolith forming Neogoniolithon brassica-florida and Lithophyllum sp. in Shark Bay, and is the first record of Hydrolithon reinboldii rhodoliths in Australia.     

Variation in Photosynthetic Performance Relative to Thallus Microhabitat Heterogeneity in Lithothamnion australe (Rhodophyta,Corallinales) Rhodoliths

Rhodoliths are free‐living, coralline algae that create heterogeneous structure over sedimentary habitats. These fragile ecosystems are threatened by anthropogenic disturbances that reduce their size and three‐dimensional structural complexity. We investigated how physical disturbance from boat moorings affects photosynthetic performance in the rhodolith Lithothamnion australe. Photosynthetic parameters were measured for intact rhodoliths and crushed rhodolith fragments of two sizes (ca. 1 and 2 cm diameter), while chlorophyll fluorescence was measured at the surface of rhodoliths of these two sizes, between the interior branches of the larger rhodoliths, and at the surface of 52 various sized (0.4–3.5 cm diameter) rhodoliths. Gross productivity and net productivity were 15% and 36% higher, respectively, in the smaller L. australe, while respiration was 10% higher in the larger individuals. Thallus crushing reduced gross productivity by 20% and 41%, and net productivity by 9% and 14% in the smaller and larger rhodoliths, respectively. It also reduced respiration by 33% and 60% in the smaller and larger rhodoliths, respectively. Fluorescence parameters were all greater at the surface of the larger L. australe than the smaller individuals, and greater at the surface than in the interior parts of the larger individuals. Across a range of rhodolith sizes, surface fluorescence parameters were at their maxima in 1.54 to 2.32 cm diameter individuals. These results show that L. australe’s complex structure creates heterogeneity in photosynthesis and respiration between their surface and interior parts and among rhodolith sizes. This information can help predict how rhodoliths may respond to disturbance and environmental stressors.     

High CO2 decreases the long‐term resilience of the free‐living coralline algae Phymatolithon lusitanicum

Mäerl/rhodolith beds are protected habitats that may be affected by ocean acidification (OA), but it is still unclear how the availability of CO₂ will affect the metabolism of these organisms. Some of the inconsistencies found among OA experimental studies may be related to experimental exposure time and synergetic effects with other stressors. Here, we investigated the long‐term (up to 20 months) effects of OA on the production and calcification of the most common mäerl species of southern Portugal, Phymatolithon lusitanicum. Both the photosynthetic and calcification rates increased with CO₂ after the first 11 months of the experiment, whereas respiration slightly decreased with CO₂. After 20 months, the pattern was reversed. Acidified algae showed lower photosynthetic and calcification rates, as well as lower accumulated growth than control algae, suggesting that a metabolic threshold was exceeded. Our results indicate that long‐term exposure to high CO₂ will decrease the resilience of Phymatolithon lusitanicum. Our results also show that shallow communities of these rhodoliths may be particularly at risk, while deeper rhodolith beds may become ocean acidification refuges for this biological community.     

New records of rhodolith-forming species (Corallinales, Rhodophyta) from deep water in Espírito Santo State, Brazil

Little is known about the diversity of non-geniculate coralline red algae (Rhodophyta, Corallinophycidae) from deep waters in Brazil. Most surveys undertaken in this country have been carried out in shallow waters. In 1994, however, the REVIZEE program surveyed the sustainable living resources potential of the Brazilian exclusive economic zone to depths of 500?m. In the present study, the rhodolith-forming coralline algae from the continental shelf of Espírito Santo State were identified. Samples were taken from 54 to 60?m depth by dredging during ship cruises in 1997. Three rhodolith-forming species were found: Spongites yendoi (Foslie) Chamberlain, Lithothamnion muelleri Lenormand ex Rosanoff and Lithothamnion glaciale Kjellman. These records extend the distribution ranges of these species into Brazilian waters and extend the depth distribution of non-geniculate coralline red algae into Brazilian water to 58?m.     

GENETIC DIFFERENCES BETWEEN TWO GROWTH‐FORMS OF LITHOPHYLLUM MARGARITAE (RHODOPHYTA) IN BAJA CALIFORNIA SUR,MEXICO1

Unattached, nongeniculate, coralline algae or rhodoliths exhibit a range of morphological variability seemingly dependent on environmental factors. Rhodoliths have an extensive fossil record, and environmentally dependent characteristics make them potentially reliable paleoindicators. Species of the rhodolith‐forming genus Lithophyllum Philippi in Baja California Sur, Mexico were recently consolidated into one species. Under the new classification, L. margaritae (Hariot) Heydrich consists of several growth forms presumably reflecting local environmental conditions. We examined the genetic structure of four populations of this species using amplified fragment length polymorphisms (AFLP) to characterize the extent of genetic variation associated with foliose and fruticose growth forms. AFLP band sharing analysis revealed that foliose growth forms exhibited consistently higher intrapopulation similarities (0.75–0.85) than fruticose growth forms (similarity range, 0.55–0.67). This trend was also evident in comparisons of geographically isolated populations. These data indicate that the two morphologies are genetically distinct and that genetic exchange between foliose and fruticose growth forms of L. margaritae may be limited. Consequently, rhodolith growth forms appear to be the result of an interplay between both genetic makeup and environmental conditions.     

Coralline algae as depth indicators in the Sommières Basin (early Miocene,Southern France)

The aim of this work is to demonstrate the benefits of digital surface-quantification and species identification in the study of coralline algae. Within the well-constrained context of the Miocene Sommières Basin (Southern France), whose sedimentological and paleontological setting has been reconstructed by previous researches, the abundance of coralline algae is estimated through quantification of the area occupied by the different species, which are thoroughly described in order to make future taxonomic-revisions as easy as possible. Coralline distribution patterns are studied along a bathymetric gradient to assess the response of the major groups and of the different species to variations in water depth. Surface quantification highlights clear trends both at the species level and higher taxonomic ranks, showing that coralline algae can be a quite accurate instrument in paleoenvironmental reconstructions. Among the clearly identified species, Lithophyllum sp. A, Sporolithon sp. A, and Spongites fruticulosus prefer shallow-water; Mesophyllum roveretoi dominates at intermediate depth together with Lithothamnion ponzonense, while Phymatolithon sp. A occurs at the lower limit of the photic zone. The approach presented here can be applied to a variety of different contexts, providing a powerful instrument for the interpretation of coralline-algal associations.     

Tracking paleoenvironmental changes in coralline algal-dominated carbonates of the Lower Oligocene Calcareniti di Castelgomberto formation (Monti Berici, Italy)

Lower Oligocene, shallow-water carbonates of the Calcareniti di Castelgomberto formation (Monti Berici, Italy, Southern Alps) are studied in detail with respect to fabric and component distributions in order to trace paleoecological changes along a monotonous sedimentary stacking pattern. The carbonates are dominated by coralline algal rudstones with a packstone to wackestone matrix. Non-geniculate coralline algae include six genera: Lithoporella melobesioides, Mesophyllum, Neogoniolithon, Spongites, Sporolithon, and Subterraniphyllum. The algae are found in the form of encrusting thalli, rhodoliths, and coralline debris. Non-algal components include larger, small benthic, and planktonic foraminifera associated with bryozoans, zooxanthellate corals, and echinoderms. Four carbonate facies are distinguished: (1) coralline algal facies, (2) coralline algal-coral facies, (3) coralline algal-larger foraminiferal facies, and (4) coralline algal debris facies. Marly horizons also occur in the section. The facies and coralline algal content are interpreted with respect to light intensity, hydrodynamic energy, biotic interactions, and substrate stability. Facies development along the studied section shows systematic variations, suggesting asymmetric sea-level changes with rapid regressions and gradual transgressions.     

Coralline-algal assemblages of a Burdigalian platform slope: implications for carbonate platform reconstruction (northern Sardinia,western Mediterranean Sea)

The rhodolithic slope deposits of a Burdigalian carbonate platform in Sardinia near Sedini were analyzed to reconstruct facies and palaeobathymetry. There is a distinct red-algal growth zonation along the platform slope. The clinoform rollover area consists of coralline-algal bindstones, which downslope change into a zone where rhodoliths are locally fused by progressive encrustation. Mid-slope rhodoliths are moderately branched, and downslope rhodoliths have fruticose protuberances, resulting in branching rhodolith growth patterns. There is a sharp change from the rhodolitic rudstones to the basinal, bivalve-dominated rudstones at the clinoform bottomsets. Red-algal genera identified include Sporolithon, Lithophyllum, Spongites, Hydrolithon, Mesophyllum, Lithoporella, Neogoniolithon, and other mastophoroids and melobesioids. Genera and subfamilies show a zonation along the clinoforms, allowing palaeobathymetric estimates. The clinoform rollovers formed at a water depth of around 40 m and the bottomsets around 60 m. Results from geometrical reconstruction show that coral reefs in the inner platform formed at water depths of around 20 m. Therefore, the Sedini carbonate platform is an example of a reef-bearing platform in which the edge or the platform-interior reefs do not build up to sea level.     

Biomineralization of calcium carbonate in the cell wall of Lithothamnion crispatum (Hapalidiales,Rhodophyta): correlation between the organic matrix and the mineral phase

Over the past few decades, progress has been made toward understanding the mechanisms of coralline algae mineralization. However, the relationship between the mineral phase and the organic matrix in coralline algae has not yet been thoroughly examined. The aim of this study was to describe the cell wall ultrastructure of Lithothamnion crispatum, a cosmopolitan rhodolith‐forming coralline algal species collected near Salvador (Brazil), and examine the relationship between the organic matrix and the nucleation and growth/shape modulation of calcium carbonate crystals. A nanostructured pattern was observed in L. crispatum along the cell walls. At the nanoscale, the crystals from L. crispatum consisted of several single crystallites assembled and associated with organic material. The crystallites in the bulk of the cell wall had a high level of spatial organization. However, the crystals displayed cleavages in the (104) faces after ultrathin sectioning with a microtome. This organism is an important model for biomineralization studies as the crystallographic data do not fit in any of the general biomineralization processes described for other organisms. Biomineralization in L. crispatum is dependent on both the soluble and the insoluble organic matrix, which are involved in the control of mineral formation and organizational patterns through an organic matrix‐mediated process. This knowledge concerning the mineral composition and organizational patterns of crystals within the cell walls should be taken into account in future studies of changing ocean conditions as they represent important factors influencing the physico‐chemical interactions between rhodoliths and the environment in coralline reefs.     

Sequencing type material resolves the identity and distribution of the generitype Lithophyllum incrustans,and related European species L. hibernicum and L. bathyporum (Corallinales,Rhodophyta)

DNA sequences from type material in the nongeniculate coralline genus Lithophyllum were used to unambiguously link some European species names to field‐collected specimens, thus providing a great advance over morpho‐anatomical identifi‐cation. In particular, sequence comparisons of rbcL, COI and psbA genes from field‐collected specimens allowed the following conclusion: the generitype species, L. incrustans, occurs mostly as subtidal rhodoliths and crusts on both Atlantic and Mediterranean coasts, and not as the common, NE Atlantic, epilithic, intertidal crust reported in the literature. The heterotypic type material of L. hibernicum was narrowed to one rhodolith belonging in Lithophyllum. As well as occurring as a subtidal rhodolith, L. hibernicum is a common, epilithic and epizoic crust in the intertidal zone from Ireland south to Mediterranean France. A set of four features distinguished L. incrustans from L. hibernicum, including epithallial cell diameter, pore canal shape of sporangial conceptacles and sporangium height and diameter. An rbcL sequence of the lectotype of Lithophyllum bathyporum, which was recently proposed to accommodate Atlantic intertidal collections of L. incrustans, corresponded to a distinct taxon hitherto known only from Brittany as the subtidal, bisporangial, lectotype, but also occurs intertidally in Atlantic Spain. Specimens from Ireland and France morpho‐anatomically identified as L. fasciculatum and a specimen from Cornwall likewise identified as L. duckerae were resolved as L. incrustans and L. hibernicum, respectively.     

Deep-water rhodolith productivity and growth in the southwestern Atlantic

Rhodolith beds are an important source of marine calcium carbonate worldwide. Dense aggregations are found in deep water in the tropical southwestern Atlantic. In order to understand the distribution of coralline algae that build rhodoliths, algae responses were measured in light levels over their depth range. Qualitative samples were obtained by dredging at 90–100 m depth 80 km offshore of Cabo Frio Island, southeastern Brazil. Histological sections indicate that Mesophyllum engelhartii (Foslie) Adey was the most frequent coralline algae in 22 sampling stations. Its lumpy and thin thallus is characterized by raised multiporate sporangial conceptacles, with cells of similar size and shape around the pore canal. Accretion growth rates were below detectable levels for two rhodolith species. Photosynthetic peak was reached at 0.5–1.5 % of the maximum surface irradiance (10–30 μmol photons m^?2 s^?1) showing an extremely narrow P–I curve of net primary production. Readings of primary production at irradiance close to saturation and lower light levels (5 μmol photons m^?2 s^?1) showed no significant difference, suggesting that rhodoliths are acclimated to restricted light ranges in deep water.     

There is more to maerl than meets the eye: DNA barcoding reveals a new species in Britain,Lithothamnion erinaceum sp. nov. (Hapalidiales,Rhodophyta)

Due to the high plasticity of coralline algae, identification based on morphology alone can be extremely difficult, so studies increasingly use a combination of morphology and genetics in species delimitation. A DNA barcoding study was carried out on maerl-forming coralline algae using the mitochondrial cytochrome oxidase 1 gene, CO1, and the plastid gene, psbA, on field specimens from Falmouth and Oban together with herbarium specimens from the Natural History Museum, UK, and the Smithsonian Institution, Washington, USA. Results revealed the presence in the north of Britain of a new species, Lithothamnion erinaceum Melbourne & J. Brodie, sp. nov., which was previously misidentified as Lithothamnion glaciale. The results also indicated that Lithothamnion lemoineae, which had earlier been recorded from Britain, was not present. One of the biggest concerns at present is how organisms will respond to climate change and ocean acidification, and it is imperative that investigations are put on a firm taxonomic basis. Our study has highlighted the importance of using molecular techniques to aid in the elucidation of cryptic diversity.     

Rhodolith formation induced by reef erosion in the Red Sea,Egypt

Along the northwestern margin of Safaga Island (Northern Bay of Safaga, Red Sea, Egypt) a small fringing reef (several hundred meters long, up to 2 m high) and small patch reefs are developed due to the local current regime which is favorable for coral growth. Corals and reef rock are encrusted by coralline algae, predominantly by branchedLithophyllum kotschyanum. Owing to destructional processes dominated by sea urchin activities, fragmentation of (1) corals, (2) reef rock, and (3) coralline algae takes place resulting in the formation of almost mono-specific, branchedLithophyllum kotschyanum rhodoliths. Rhodolith formation takes place in various reef environments: (1) in depressions on the reef flat where ellipsoidal rhodoliths develop, with interlocking and fusing branches leading to a coralline algal framework; (2) in discharge channels where smaller elongated rhodoliths occur; (3) in leeward positions between reef flat and seagrass meadows, where a dense belt of spheroidal to ellipsoidal rhodoliths is formed; scattered rhodoliths occur in adjacent seagrass beds. The formation and preservation of rhodoliths requires a complex interplay of destruction, growth, transportation, movement, and stabilization.     

Experimental effects of kelp canopies on subtidal coralline algae

Plants are often grouped as canopy species or understorey species because it is thought that that these sets of taxa interact in predictable ways. Mensurative experiments in southern Australia demonstrated that the percentage cover of encrusting coralline algae was greater, and articulated (branching) coralline algae less, on boulders under a canopy of dense kelp (>7 plants per m²), Ecklonia radiata, than on boulders without kelp. Experimental clearances of kelp and reciprocal transplants of boulders between patches of E. radiata and patches without kelp showed that canopies maintained and facilitated the growth of encrusting coralline algae and reduced the cover of articulated coralline algae. Potential artefacts associated with clearing kelp and transplanting boulders were not detected when tested with a series of translocation controls. These results reject the model that the co‐occurrence of E. radiata and encrusting corallines is just an assemblage of plants caused by spatial and temporal coincidence. Instead, they support the model that kelp facilitates the growth and survival of understorey algae.     

3-D distribution of nongeniculate corallinales: a case study from a reef crest of South Sinai (Red Sea,Egypt)

An innovative technique for the estimation of species and growth-form abundance of coralline algae, including information on their vitality, was adopted on the reef crest of Ras Nosrani and Coral Bay, South Sinai. Data of coralline abundance from visual census and collection of voucher specimens were plotted on a 3-D sketched representation of the horizontal and vertical planes of the reef crest and of its crannies. Coralline dominance at the two investigated sites was not significantly different, with values ranging from 8.55 and 10.06% on the vertical plane and from 5.3 to 7.17% on the horizontal plane. About one-third of total corallines of the South Sinai reef crest was located in crannies, where the algae are completely overlooked by routine field surveys. Pink to violet, healthy corallines with encrusting growth-form, mainly belonging to Hydrolithon onkodes and Neogoniolithon, with subordinate fruticose Lithophyllum kotschyanum dominated the reef crest at both sites. The fruticose growth form, usually associated with L. kotschyanum, was more common in the horizontal than in the vertical plane. Purple, healthy, encrusting Sporolithon uncommonly occurred. Whitish, gray or green unhealthy or dead corallines were more common on the horizontal plane at both sites, possibly resulting from excessive solar radiation.