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.     

Effects of kelp canopies on bleaching and photosynthetic activity of encrusting coralline algae

Canopy-forming algae often coexist with an understorey of encrusting coralline algae that bleach following the loss of canopies. We tested the hypothesis that canopy loss causes a reduction in photosynthetic activity of encrusting coralline algae concomitant with their bleaching. When canopies were experimentally removed, corallines bleached and their photosynthetic activity was rapidly reduced to half their activity observed under canopies. This result prompted us to test, and subsequently accept, the hypothesis that exposure of understorey corallines to enhanced light intensity per se (simulation of canopy loss) acts as a mechanism that causes bleaching and reduced photosynthetic activity. Despite bleaching, encrusting corallines maintained reduced levels of photosynthetic activity, and this may explain why, under certain conditions, bleached corallines can persist in the absence of canopy-forming algae. Nevertheless, our data provide evidence that the positive association between canopy-forming algae and encrusting coralline algae is maintained because of shade provided by the canopy.     

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.     

Predicting understorey structure from the presence and composition of canopies: an assembly rule for marine algae

Assembly rules provide a useful framework for predicting patterns of community assembly under defined environmental conditions. Habitat created by canopy-forming algae (such as kelps) provides a promising system for identifying assembly rules because canopies typically have a large and predictable influence on understorey communities. Across >1,000 km of subtidal South Australian coastline, we identified natural associations between assemblages of understorey algae and (1) monospecific canopies of Ecklonia radiata, (2) canopies comprised of E. radiata mixed with Fucales (Cystophora spp. and Sargassum spp.), and (3) gaps among canopies of algae. We were able to recreate these associations with experimental tests that quantified the assembly of understorey algae among these three habitat types. We propose the assembly rule that understorey communities on subtidal rocky coast in South Australia will be (1) monopolised by encrusting coralline algae beneath monospecific canopies of E. radiata, (2) comprised of encrusting corallines, encrusting non-corallines, and sparse covers of articulated corallines, beneath mixed E. radiata-Fucales canopies, and (3) comprised of extensive covers of articulated corallines and filamentous turfs, as well as sparse covers of foliose algae and juvenile canopy-formers, within gaps. Consistencies between natural patterns and experimental effects demonstrate how algal canopies can act as a filter to limit the subsets of species from the locally available pool that are able to assemble beneath them. Moreover, the subsets of species that assemble to subtidal rocky substrata in South Australia appear to be predictable, given knowledge of the presence and composition of canopies incorporating E. radiata.     

Limited flexibility in resource use in a coral reef grazer foraging on seasonally changing algal communities

Feeding ecology of three life phases of the parrotfish Scarus ferrugineus was studied on a southern Red Sea fringing reef by comparing availability and consumption of benthic algae during the monsoon hot and cool seasons. Dominant biota covering dead carbonate substrates were in decreasing order of importance: turfs on endoliths, turfs on crustose corallines, and crustose corallines. On the reef crest and shallow fore reef, composition of the biota changed seasonally. Cover of turfs on endoliths and turfs on crustose corallines was higher during the hot season, while crustose corallines and macroalgae (only on reef crest) increased during the cool season. Biota in the deep fore reef did not show seasonal variation. All life phases used similar resources and showed selective feeding in all zones. Turfs on endoliths, followed by turfs on crustose corallines, was the primary feeding substrate. These two sources represented over 92% of bites during both seasons. Crustose corallines, macroalgae, and living corals were negligible components being strongly avoided at all zones and seasons. Resource use varied seasonally on the reef crest and shallow fore reef, while it remained unchanged on the deep fore reef. Turfs on endoliths were consistently preferred in both seasons but their contribution increased from 45% in the cool to 70% of bites in the hot season. Electivity for turfs on crustose corallines shifted from random feeding in the hot (27% of bites) to selection in the cool season (47% of bites). Feeding pattern changed diurnally with more bites taken from crustose corallines and turfs on crustose corallines during morning. During the rest of the day, bites from turfs on endoliths predominate. S. ferrugineus shows limited capacity to exploit seasonal increases in the biomass of foliose and canopy forming macroalgae, despite indications of energetic limitation during the cool season.     

Facultative mutualism between an herbivorous crab and a coralline alga: advantages of eating noxious seaweeds

Because encrusting coralline algae rely on herbivory or low light levels to prevent being overgrown by competitively superior fleshy algae, corallines are relatively rare in shallow areas with low rates of herbivory. In contrast to this general trend, the branching coralline alga Neogoniolithon strictum occurs primarily in shallow seagrass beds and along the margins of shallow reef flats where herbivory on macrophytes is low. This alga apparently persists in these habitats by providing refuge to the herbivorous crab Mithrax sculptus at mean densities of 1 crab per 75 g of algal wet mass. When crabs were removed from some host corallines, hosts without crabs supported 9 times the epiphytic growth of hosts with crabs after only 30 days. Crabs without access to a coralline alga were rapidly consumed by reef fishes, while most of those tethered near a host alga survived. These results suggest that the crabs clean their algal host of fouling seaweeds and associate with the host to minimize predation. However, to effectively clean the host, the crab must consume the wide array of macroalgae that commonly co-occur with coralline algae in these habitats, including chemically defended species in the genera Halimeda, Dictyota, and Laurencia. Crabs did readily consume these seaweeds, which were avoided by, and are chemically defended from, herbivorous fishes. Even though crabs readily consumed both Halimeda and Dictyota in whole-plant feeding assays, chemical extracts from these species significantly reduced crab feeding, suggesting that factors other than secondary chemistry (e.g., food value, protein, energy content), may determine whole-plant palatability. Having the ability to use a wide variety of foods, and choosing the most profitable rather than the least defended foods, would diminish foraging time, increase site fidelity, and allow the crab to function mutualistically with the host alga. Despite the obvious benefit of associating with N. strictum, M. sculptus did not prefer it over other habitats offering a structurally similar refuge, suggesting that these crabs are not N. strictum specialists, but rather occupy multiple habitats that provide protection from predators. Structurally complex organisms like N. strictum may commonly suppress competitors by harboring protective symbionts like M. sculptus. It is possible that diffuse coevolution has occurred between these two groups; however, this seems unlikely because both herbivore and host appear to respond most strongly to selective pressures from predators and competitors outside this association.     

Articulated coralline algae of the genus Amphiroa are highly effective natural inducers of settlement in the tropical abalone Haliotis asinina

The initiation of metamorphosis in marine invertebrates is strongly linked to the environment. Planktonic larvae typically are induced to settle and metamorphose by external cues such as coralline algae (Corallinaceae, Rhodophyta). Although coralline algae are globally abundant, invertebrate larvae of many taxa settle in response to a very limited suite of species. This specificity impacts population structure, as only locations with the appropriate coralline species can attract new recruits. Abalone (Gastropoda, Haliotidae) are among those taxa in which closely related species are known to respond to different coralline algae. Here we identify highly inductive natural cues of the tropical abalone Haliotis asinina. In contrast to reports for other abalone, the greatest proportion of H. asinina larvae are induced to settle and metamorphose (92.8% to 100% metamorphosis by 48 h postinduction) by articulated corallines of the genus Amphiroa. Comparison with field distribution data for different corallines suggests larvae are likely to be settling on the seaward side of the reef crest. We then compare the response of six different H. asinina larval families to five different coralline species to demonstrate that induction by the best inductive cue (Amphiroa spp.) effectively extinguishes substantial intraspecific variation in the timing of settlement.     

Natural inducers for coral larval metamorphosis

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

Reef demise and back-stepping during the last interglacial, northeast Yucatan

The elevation of reefs and coastal deposits during the last Interglaciation (MIS-5e) indicates that sea level reached a highstand of as much as 6 m above the present, but it is uncertain how rapidly this level was attained and how it impacted reef development. To investigate this problem, I made a detailed sedimentological analysis of a well-dated reef from the northeast coast of the stable Yucatan Peninsula. Two linear reef tracts were delineated which are offset and at different elevations. The lower reef tract crops out along northern shore for 575 m and extends from below present mean sea level to +3 m. The reef crest facies consists of large Acropora palmata colonies dispersed within a coral boulder-gravel and is flanked by an A. cervicornis-dominated reef-front and a large area of lagoonal framework formed by coalesced patches of A. cervicornis and Montastraea spp. Constituents in the upper centimetre of the lower tract are heavily encrusted by a cap of crustose corallines and, in places, are levelled by a discontinuous marine-erosion surface. The upper reef tract crops out ~150 m inland up to an elevation of +5.8 m and parallels the southern section of shore for ~400 m. It also consist of an A. palmata-dominated crest facies flanked by reef-front, back-reef and lagoonal frameworks. In this case, however, lagoonal frameworks are dominated by a sediment-tolerant assemblage of branching coralline algae. Also different is the lack of encrustation by corallines, and the infiltration of upper tract facies by beach-derived shell-gravels from regressive shoreface deposits above. These results indicate that the lower reef tract and lagoonal patch-reefs formed at a sea level of +3 m. Final capping by crustose corallines and discontinuous marine erosion indicates that the lower tract was terminated by the complete demise of corals on the crest but only patchy demise in the lagoon. Areas of continuous framework accretion between the lagoonal patch reefs and the upper reef-tract, however, require that the demise of this reef was ecologically synchronous with initiation of the upper reef-tract, which had back-stepped 100 m into the lagoon. In this new position, the upper tract developed a reef crest that corresponded to a final sea-level position of +6 m. Reef flat development at +5 m and large in-place colonies of A. palmata at the base of the crest unit indicate, however, that sea level must have risen rapidly from +3 to more than +5 m to accommodate back-stepping. This sea-level jump created a higher energy wave field that mobilized back-reef and lagoonal sediments, and the resulting high sediment flux eroded lagoonal framework and prevented the recovery of the submerged lower reef crest. So this single jump in sea level was responsible not only for reef demise and back-stepping but also for marine erosion and suppression of subsequent reef development—features that elsewhere have been used to support multiple sea-level excursions during the last interglacial.     

Genomics reveals abundant speciation in the coral reef building alga Porolithon onkodes (Corallinales,Rhodophyta)

An essential suite of coral reef ecosystem engineers is coralline red algae. Among these, the smooth, encrusting Porolithon onkodes has historically been considered the most important and common reef building species worldwide. We assess P. onkodes biodiversity by performing a genomic analysis of the lectotype specimen collected in 1892 from the Tami Islands, Gulf of Huon, east of New Guinea. Comparisons of DNA sequences from the lectotype specimen to those deposited in GenBank and to newly generated sequences from both field‐collected and historical specimens demonstrate that at least 20 distinct species are passing under P. onkodes. We hypothesize that there were multiple evolutionary drivers including ecophysiology, hydrodynamic regimes, and biotic interactions as well as historical biogeography, which resulted in this high diversity of smooth, encrusting Porolithon species throughout the tropics. Our results emphasize the need to document the biodiversity, ecophysiology, and habitats of these tropical, reef‐building algae in light of climate change and ocean acidification.     

Dispersion and phagokinesis in the echinoid Evechinus chloroticus (Val.)

Movement in the echinometrid sea urchin Evechinus chloroticus (Val.) was investigated in relation to the availability of a preferred food species, the laminarian alga Ecklonia radiata (C. Ag.). Patterns of microhabitat occupancy, dispersion, and movement were described in two habitats: Coralline Flats, dominated by encrusting coralline algae; and Sediment Flats, occupied by turfing corallines, Ecklonia and the mussel Modiolus areolatus (Gould). In both habitats, sea urchins were positively associated with encrusting coralline algae, despite the fact that encrusting corallines occupied a minor proportion of the substratum in the Sediment Flat habitat. Sea urchins were significantly clumped in both habitat types, and more so in sites within the Sediment Flats habitat. Movement was not directional in either habitat, but the magnitude of movement was almost twice as high in sites from the Coralline Flats habitat, where kelp was absent, as in the Sediment Flats sites, where kelp was present. Behaviour in response to the presence of Ecklonia in both habitats was investigated experimentally. In both habitats, urchins exposed to a “drift” kelp plant at a distance of 1–2 m exhibited net movement approximately twice as great as that shown by control urchins, not exposed to a kelp plant. Despite the relatively small distances the urchins were from the kelp, there was no evidence of directionality in movement. We conclude from the present study that further understanding of the feeding behaviour of urchins requires that a distinction be made between attached and drift algae, and between directionality and magnitude of movement, and that the interaction of these and other factors are best investigated under field conditions.     

Pleistocene reef environments, constituent grains, and coral community structure: Curaçao, Netherlands Antilles

We investigated the degree to which component grains vary with depositional environment in sediments from three reef habitats from the Pleistocene (125?ka) Hato Unit of the Lower Terrace, Curaçao, Netherlands Antilles: windward reef crest, windward back reef, and leeward reef crest. The windward reef crest sediment is the most distinctive, dominated by fragments of encrusting and branching coralline red algae, coral fragments and the encrusting foraminiferan Carpenteria sp. Windward back reef and leeward reef crest sediments are more similar compositionally, only showing significant differences in relative abundance of coral fragments and Homotrema rubrum. Although lacking high taxonomic resolution and subject to modification by transport, relative abundance of constituent grain types offers a way of assessing ancient skeletal reef community composition, and one which is not limited to a single taxonomic group. The strong correlation between grain type and environment we found in the Pleistocene of Curaçao suggests that constituent grain analysis may be an effective tool in delineating Pleistocene Caribbean reef environments. However, it will not be a sufficient indicator where communities vary significantly within reef environments or where evolutionary and/or biogeographical processes lead to different relationships between community composition and reef environment. Detailed interpretation of geological, biological, and physical characteristics of the Pleistocene reefs of Curaçao reveals that the abundance of the single coral species, Acropora palmata, is not a good predictor of the ecological structure of the ancient reef coral communities. This coral was the predominant species in two of the three reef habitats (windward and leeward reef crest), but the taxonomic composition (based on species relative abundance data) of the reef coral communities was substantially different in these two environments. We conclude that qualitative estimates of coral distribution patterns (presence of a key coral species or the use of a distinctive coral skeletal architecture), when used as a component in a multi-component analysis of ancient reef environments, probably introduces minimal circular reasoning into quantitative paleoecological studies of reef coral community structure.     

Settlement and post-settlement processes limit the abundance of the geniculate coralline alga Calliarthron on subtidal walls

Cover of geniculate coralline algae is significantly greater on subtidal platforms than on vertical walls in a central California kelp forest. To determine the cause(s) of this pattern, settlement and growth of Calliarthron were compared on horizontal and vertical substrata attached to platforms or walls. Recruitment on horizontal substrata attached to reef walls indicated that spores were abundant along the walls. Densities of recruits were significantly smaller on vertical substrata attached to walls than on horizontal substrata attached to platforms, indicating that Calliarthron settlement is reduced on vertical substrata. Growth of basal crusts and the densities of fronds were also significantly greater on horizontal substrata attached to platforms than on vertical substrata attached to walls. Irradiance reaching wall surfaces was less than irradiance reaching platform surfaces. We suggest that light reaching wall surface is low enough to affect Calliarthron growth. These results indicate that the low abundance of Calliarthron on subtidal walls is a result of poor recruitment and slow growth.     

The Role of Encrusting Coralline Algae in the Diets of Selected Intertidal Herbivores

Kalk Bay, South Africa, has a typical south coast zonation pattern with a band of seaweed dominating the mid-eulittoral and between two molluscan-herbivore dominated upper and lower eulittoral zones. Encrusting coralline algae were very obvious features of these zones. The most abundant herbivores in the upper eulittoral were the limpet, Cymbula oculus (10.4 ± 1.6 individuals m⁻²; 201.65 ± 32.68 g.m⁻²) and the false limpet, Siphonaria capensis (97.07± 19.92 individuals m⁻²; 77.93 16.02 g.m⁻²). The territorial gardening limpet, Scutellastra cochlear, dominated the lower eulittoral zone, achieving very high densities (545.27 ± 84.35 m⁻²) and biomass (4630.17 ± 556.13 g.m⁻²), and excluded all other herbivores and most seaweeds, except for its garden alga and the encrusting coralline alga, Spongites yendoi (35.93 ± 2.26% cover). In the upper eulittoral zone, encrusting coralline algae were only present in the guts of the chiton Acanthochiton garnoti (30.5 ± 1.33%) and the limpet C. oculus (2.9 ± 0.34%). The lower eulittoral zone limpet, Scutellastra cochlear also had a large percentage of encrusting coralline algae in its gut with limpets lacking gardens having higher (45.1 ± 1.68%) proportions of coralline algae in their guts than those with gardens (25.6 ± 0.8%). Encrusting coralline algae had high organic contents, similar to those of other encrusting and turf-forming algae, but higher organic contents than foliose algae. Radula structure, grazing frequencies as a percentage of the area grazed (upper eulittoral 73.25 ± 3.60% d⁻¹; lower eulittoral 46.0 ± 3.29% d⁻¹), and algal organic content provided evidence to support the dietary habits of the above herbivores. The data show that many intertidal molluscs are actively consuming encrusting coralline algae and that these seaweeds should be seen as an important food source.     

Calcification by Reef-Building Sclerobionts

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

Composition and community structure of the coralline algal reefs from Atol das Rocas, South Atlantic, Brazil

 Coralline-algal frameworks from Atol das Rocas reefs were sampled along windward and leeward transects, and multivariate statistical analysis was used to investigate the composition and community structure of the encrusting community. The following components of the reefs were identified in slabbed and polished reef blocks sampled along each transect: encrusting coralline algae dominated by Porolithon cf. pachydermum, vermetid gastropods, Homotrema rubrum, acervulinid foraminiferans, molluscs, corals, polychaete tubes, clionid sponge borings, polychaete/sipunculid borings, cemented sediments, and growth cavities. Q-mode cluster analysis correctly classified 78% of all windward samples and 69% of all leeward samples, and R-mode separated reef components adapted to high wave energy environments from those adapted to low wave energy. Separate classification and ordination of samples from each transect indicate that seriation breakdown occurs in the windward reef between 25 and 45 m from the reef front. In the leeward reef between 75 and 90 m from the leeward reef front, seriation breakdown was not found to be significant. These results confirm that seriation (zonation) is best developed in the windward reef, as has been documented in coral-dominated reefs. Accepted: 28 February 2000     

Sponge-mediated coral reef growth and rejuvenation

Sponges mediate consolidation of Porites furcata rubble on shallow Caribbean reefs by quickly adhering to rubble and stabilizing it until carbonate secreting organisms can grow and consolidate it to the reef. Experimental investigations demonstrate that the entire cycle from (1) temporary binding of rubble by sponges, through (2) rubble consolidation by encrusting coralline algae, to (3) colonization of consolidated rubble by corals, can be completed within 10 months. Bound rubble both adds to vertical reef growth and also provides stable substrata for colonization by corals. Corals that colonize stabilized rubble are damaged less and survive better than on unstable rubble. Rubble that is not temporarily stabilized by sponges does not become bound to the reef, because continuous movement disturbs the consolidation process, and does not provide suitable substrata for settlement and growth of corals. Sponge-mediated consolidation of rubble may increase rates of reef growth and enhance reef recovery after damage. This new role for sponges in reef growth is not obvious from examination of the internal fabric of a reef frame. Spongemediated consolidation may help to explain geographic and temporal differences in growth and morphology among shallow reefs of ramose corals.     

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.     

The nepionic stage of <Emphasis Type="Italic">Solenomeris</Emphasis> Douvillé, 1924 (Acervulinidae,Foraminiferida): new observations from the uppermost Maastrichtian-early Danian of Austria (Kambühel Formation,Northern Calcareous Alps)

In the shallow-water limestone succession of the Kambühel Formation (upper Maastrichtian to upper Thanetian, Northern Calcareous Alps), the benthic-vagile to encrusting-sessile acervulinid foraminifer Solenomeris Douvillé was identified in upper Maastrichtian and, more commonly, lower Danian rocks. We report on the free nepionic (juvenile) stage of this foraminifer, its relation to the adult encrusting stage, and competitive interaction with coralline algae. Whereas morphology and chamber arrangement of the free nepionts clearly assign the Alpine specimens to Solenomeris, the chamber shape in axial sections differs from the Eocene type-species S. ogormani, accounting for classification in open nomenclature. Nepiont chamber shape can be used as a further specific criterion besides size of adult chambers and, perhaps, growth habit. A two-stage concept of ontogenetic development is presented. Solenomeris flourished in early Danian high-stress environments (perhaps related to cooler waters and/or elevated nutrient level) under conditions less favorable for encrusting corallines that competed for substrate. Absence of Solenomeris in upper Danian limestones and in upper Thanetian coralgal limestones that form the top of the Kambühel Formation may be related to recovery of coralline algae in shallow-marine environments of low to moderate nutrient level.     

Communities of constructional lips and cup reef rims in Bermuda

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