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.     

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 red algal limestones of the late Eocene alpine Foreland Basin in Upper Austria: Component analysis, facies and palecology

Summary Late Eocene sediments of the Upper Austrian Alpine Foreland Basin discordantly overlie Mesozoic and crystalline rocks, which are deeply eroded and form a distinct pre-Eocene relief. Late Eocene deposits contain red algal limestones with a remarkable lateral extent and a high diversity of sedimentary facies. Towards the south the algal limestones change into more clastic sediments, which are characterized by larger foraminifera and bryozoans. Main components are coralline algal branches and detritus, coralline crusts, rhodoliths, peyssonneliacean aggregates and crusts, nummulitid and orthophragminid foraminifera, corals, bryozoans, as well as terrigenous components. Rank correlation and factor analysis were calculated in order to obtain informations about relations between components. Hierarchical cluster analysis allowed the designation of 17 facies, most of them are dominated by coralline algae. Actualistic comparisons and correlations obtained from statistical analyses allowed the reconstruction of the depositional environments. Main features of the northern area are huge accumulations of unattached coralline algae (branches, rhodoliths, detritus), which are comparable to the present-day “Maerl”-facies. They formed loose frameworks cut by sand channels. The frequency of coralline detritus decreases upsection. Peyssonneliacean algae in higher parts of the profiles show growth-forms that are comparable to peyssonneliaceans of the Mediterranean circalittoral soft bottoms. This succession can be interpreted by an increasing relative sea level. Besides, crustose coralline algal frameworks were growing on morphological highs which are partially comparable to the present-day “Coralligéne de Plateau” of the Mediterranean Sea. In contrast to the northern area, sedimentation rate of the southern area is too low to keep up with rising sea level. The typical succession from nummulitid- to orthophragminid-and bryozoan-dominated facies can be interpreted by an increasing water depth from shallowest subtidal to the deeper photic zone and finally to the aphotic zone.     

Coralline algal facies and their palaeoenvironments in the Late Eocene of Northern Italy (Calcare di Nago, Trento)

Summary The Calcare di Nago is a carbonate unit of Middle-Late Eocene (Bartonian and Priabonian) age which is well exposed at the north-eastern end of Lake Garda, on the western margin of the Lessini Shelf (Southern Alps). This unit is highly fossiliferous as far as the coralline red algae and large foraminifera are concerned. Corals, bryozoans, echinoderms, and molluscs are also present. The present study deals with the relationships among the coralline taxa, the coralline growth-forms, and their facies development in the Priabonian part of the type section of the Calcare di Nago. The taxonomic investigation led to the identification of 15 coralline red algal species belonging to 7 non-geniculate and 2 geniculate genera. One species of Peyssonneliacean (red alga) and one of Halimedacean (green alga) were also recognized. The quantitative and qualitative analyses based on coralline red algae and large foraminifera enabled five facies to be distinguished: Algal crust-branch rudstone, Algal/Discocyclina packstone, Coralalgal boundstone, Rhodolith mound wacke/packstone, and Rhodolith pavement. According to the coralline assemblages, coralline growth-forms, and large foraminiferal associations, the five facies reflect solid and soft substrate types. Some of these facies are dominated byin situ rhodoliths, others by reworked algal debris. In the architecture of an interpreted prograding carbonate ramp, shallow water facies are dominated by members of the subfamily Mastophoroideae, while deeper water facies are dominated by those of the subfamily Melobesioideae and family Sporolithaceae. There is a significant increase both in size and in constructional voids of the rhodoliths with depth. A concomitant decrease in algal species diversity with depth has been also recognized. LargeDiscocyclina assemblages are localized across the inner and mid ramp boundary.Pellatispira andBiplanispira are present only in the uppermost mid-ramp.Nummulites, Assilina, andSpiroclypeus are dominant together with small orthophragminids both in the mid- and uppermost outer ramp facies.     

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.     

The occurrence of Ezo epiyessoense Adey,Masaki & Akioka (Rhodophyta,Corallinaceae) in England with a summary of parasitism and endophytism in nongeniculate Corallinaceae

The parasitic, nongeniculate, coralline red alga Ezo epiyessoense (Rhodophyta, Corallinaceae) was described in 1974 by Adey, Masaki & Akioka on the basis of specimens growing on Lithophyllum yessoense in Japan. The authors considered Ezo to be an adelphoparasite because it resembled its host taxonomically in being a member of the coralline subfamily Lithophylloideae. The species had not been recorded outside Japan until the present observation in England where it was found growing on another lithophylloid species, Titanoderma pustulatum. The structure of the English material of E. epiyessoense is described and shown to closely resemble that of the type material despite its occurrence on a different host species. Tetrasporangia and trisporangia are recorded for the first time in Ezo. A summary is given of known nongeniculate coralline parasites, semi-endophytes and outgrowths.     

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.     

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.     

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.     

Calcified epibionts as palaeoecological tools: examples from the Recent and Pleistocene reefs of Barbados

Calcified epibionts (crustose coralline algae, bryozoans, foraminiferans and serpulid worms) which colinize primary framebuilders of Recent Barbados reefs exhibit a well-defined zonation of species and morphological growth forms in response to environmental factors such as water turbulence and light. Exposed environments are characterized by thick crusts of coralline algae whereas cryptic environments are dominated by thin crusts of algae, bryozoans, foraminiferans and serpulid worms. A model, based on this zonation, was used to decipher the environments of growth and early burial of Pleistocene reefs. Lagoonal corals possess an assemblage of encrusters which document prolonged growth in a uniform environment. Reef crest corals support a mixed succession of shallow water encrusters which record a gradual decrease in light as substrates are smothered by accumulating debris. Sequences such as these represent growth under stable conditions. The model can also be used to interpret sequences formed by catastrophic events and fluctuations in sea level.     

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.     

Ecological Shifts in Mediterranean Coralligenous Assemblages Related to Gorgonian Forest Loss

Mediterranean gorgonian forests are threatened by several human activities and are affected by climatic anomalies that have led to mass mortality events in recent decades. The ecological role of these habitats and the possible consequence of their loss are poorly understood. Effects of gorgonians on the recruitment of epibenthic organisms were investigated by manipulating presence of gorgonians on experimental panels at 24 m depth, for Eunicella cavolinii, and at 40 m depth, for Paramuricea clavata, at two sites: Tavolara Island (Tyrrhenian Sea) and Portofino Promontory (Ligurian Sea). After 4 months, the most abundant taxa on the panels were encrusting green algae, erect red algae and crustose coralline algae at 24 m depth and encrusting brown algae and erect red algae at 40 m depth. Assemblages on the panels were significantly affected by the presence of the gorgonians, although effects varied across sites and between gorgonian species. Species diversity and evenness were lower on panels with gorgonian branches. Growth of erect algae and recruitment of serpulid polychaetes were also affected by the presence of the gorgonians, primarily at Tavolara. Crustose coralline algae and erect sponges were more abundant on E. cavolinii panels at 24 m depth, while encrusting bryozoans were more abundant on P. clavata panels at 40 m depth. Effects of gorgonians on recruited assemblages could be due to microscale modification of hydrodynamics and sediment deposition rate, or by a shading effect reducing light intensity. Gorgonians may also intercept settling propagules, compete for food with the filter-feeders and/or for space by producing allelochemicals. Presence of gorgonians mainly limits the growth of erect algae and enhances the abundance of encrusting algae and sessile invertebrates. Therefore, the gorgonian disappearances may cause a shift from assemblages characterised by crustose coralline algae to filamentous algae assemblages, decreasing complexity and resilience of coralligenous bioconstructions.     

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.     

Photosynthesis and respiration of a deep-water periphyton community (Macclesfield Bank,South China Sea)

The production parameters of the deep-water periphyton were investigated on the Macclesfield Bank (South China Sea) at depths down to 240 m. At 70–90 m depths the substrate is composed of the remains of dead corals and fragments of porous calcareous material overgrown with red coralline and articulated thalline algae Halimeda sp. At 220–240 m depths the substrate consists of rhodolites, which are individual unattached nodules encrusted mainly with red calcareous coralline algae Lithophyllum sp., Porolithon sp. and red lamellar alga Hypoglossum sp. At all depths the substrate is pierced with green filiform algae Ostreobium sp. and covered with benthic diatom algae. The zooperiphyton in the studied area is comprised of bryozoans, sponges, foraminifers and drilling mollusks. Respiration (R) and the gross (Pg) and net (Pn) primary production of the deepwater periphyton community were determined using the oxygen technique (polarographic electrode) at values of photosynthetically active radiation (PAR) close to natural levels. At 70–90 m depths the Pg of the photoperiphyton was 23.3 mg O₂/(m² h), and the Pn of the community had negative values due to respiration of phytoand mainly zooperiphyton and bacteria. At 220–240 m depths, where less than 0.001% of the surface PAR penetrates, the Pg of the phytoperiphyton decreased to 9.6 mg O₂/(m² h), and the R of the community was 25.9 mg O₂/(m² h). A high efficiency of light energy utilization by the phytoperiphyton was found. The apparent quantum yield (Fa) for algae collected at 70–90 m depths was close to the maximum level of 0.097. The values of Fn that we obtained for 220–240 m depths and calculated from identical data [27], were much higher than theoretical values, which cannot be explained based on modern views on the mechanisms of photosynthesis. Possible sources of errors are discussed. It is proposed that we observed additional evolution of oxygen as the result of H₂O₂ degradation.     

COMPOSITION,DISTRIBUTION, AND ABUNDANCE OF DEEP‐WATER (>30 m) MACROALGAE IN CENTRAL CALIFORNIA1

The deep‐water macroalgal assemblage was described at 14 sites off the central California coast during 1999 and 2000 from SCUBA and remotely operated vehicle sampling. The stipitate kelp Pleurophycus gardneri Setchell & Gardner, previously thought to be rare in the region, was abundant from 30 to 45 m, forming kelp beds below the well‐known giant kelp forests. Macroalgae typically formed three broadly overlapping zones usually characterized by one or a few visually dominant taxa: 1) the upper “Pleurophycus zone” (30–45 m) of stipitate kelps and Desmarestia spp. with a high percent cover of corallines, low cover of uncalcified red algae, and rare green algae; 2) a middle “Maripelta zone” (40–55 m) with other uncalcified red algae and infrequent corallines and green algae; and 3) a zone (55–75 m) of infrequent patches of nongeniculate coralline algae. The green alga Palmophyllum umbracola Nelson & Ryan, not previously reported from the Northeast Pacific, was found over the entire geographical range sampled from 35 to 54 m. Year‐round profiles of water column irradiance revealed unexpectedly clear water with an average K₀ of 0.106·m ^{? 1} Received 18 January 2002. Accepted 16 December 2002. . The low percent surface irradiance found at the average lower macroalgal depth limits in this study (0.56% for brown algae, 0.12% for uncalcified red algae, and 0.01% for nongeniculate coralline algae) and lack of large grazers suggest that light controls the lower distributional limits. The ubiquitous distribution, perennial nature, and similar lower depth limits of deep‐water macroalgal assemblages at all sites suggest that these assemblages are a common persistent part of the benthic biota in this region.     

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.     

Reef‐building coralline algae from the Southwest Atlantic: filling gaps with the recognition of Harveylithon (Corallinaceae,Rhodophyta) on the Brazilian coast

The Southwest Atlantic is notable for having extensive reef areas cemented by nongeniculate coralline red algae. Based on an analysis of four genetic markers and morpho‐anatomical features, we clarify the species of Harveylithon in the tropical and warm temperate Southwest Atlantic. Species delimitation methods (mBGD, ABGD, SPN, and PTP), using three markers (psbA, rbcL, and COI), support the recognition of three new species: H. catarinense sp. nov., H. maris‐bahiensis sp. nov., and H. riosmenum sp. nov., previously incorrectly called Hydrolithon samoënse. Our findings highlight the importance of using an approach with several lines of evidence to solve the taxonomic status of the cryptic species.