The Messinian reef complex of the Salento Peninsula (southern Italy): Stratigraphy, facies and paleoenvironmental interpretation

Summary An integrated study of the early Messinian reef complex cropping out along the eastern coast of the Salento Peninsula (southern Italy), including stratigraphy, facies analysis and paleoecological aspects, is here presented. Fourteen facies types belonging to three main facies associations (back reef and shelf, shelf-edge, slope) have been recognized. They document a wide spectrum of depositional environments, reef building organisms and growth fabrics, in response to depth and other environmental factors in different parts of the reef complex. The biotic structure of the reef is also described and discussed in detail. It consists of different types of reef building organisms and of their bioconstructions (mainlyPorites coral reefs,Halimeda bioherms and vermetidmicrobial “trottoirs”), that differ in composition and structure according to their position on the shelf edge-toslope profile. Results indicate that the reef complex of the Salento Peninsula has strong similarities with the typical early Messinian reefs of the Mediterranean region. However, the recognition of some peculiar features, i.e. the remarkable occurrence ofHalimeda bioherms and of vermetid-microbial “trottoirs”, gives new insights for better understanding reef patterns and development of the reef belt during the Late Miocene in the Mediterranean.     

Reef to basin sediment transport using Halimeda as a sediment tracer,Grand Cayman Island,West Indies

Fragments of the calcareous green alga Halimeda form a large part of the sediment in the fringing reef system and adjacent deep marine environments of Grand Cayman Island, West Indies. Nine species combine to form three depth-related assemblages that are characteristic of the major reef-related environments (lagoonpatch reef, reef terraces, and deep reef). These modern plant assemblages form the basis of the use of Halimeda as a sediment tracer. Halimeda-based tracer studies of Holocene sediments indicate that only sediments containing deep reef species of Halimeda are presently being transported through the reef system by sediment creep and being deposited at the juncture of the upper and lower island slope. Sediments containing shallow reef Halimeda are retained within the reef and lithified by marine carbonate cements. Tracer studies of Pleistocene sediment indicate large amounts of reef-derived carbonate sand containing deep water Halimeda were produced during interglacial high stands of sea level. Much of this material was removed by turbidity currents moving out of the reef system to the island slope down submarine channels perpendicular to the reef trend. These channels may still be identified on bathymetric profiles, but are no longer receiving coarse reef debris and are veneered with a blanket of pelagic carbonate mud.     

Distributional patterns of mobile fauna associated with Halimeda on the Tiahura coral-reef complex (Moorea,French Polynesia)

This study deals with the mobile fauna living associated with the turfs of three Halimeda species [H. incrassata, H. opuntia (three forms) and H. macroloba] from the Tiahura Reef complex. Ten 0.05 m² test areas of each Halimeda form have been randomly sampled from each geomorphological reef unit in order to obtain the specific abundance and biomass of different faunistic groups and species living within Halimeda populations. Then the raw data have been classified using the correspondence factor analysis to highlight the relationships between faunistic groups, or species, and the different Halimeda forms studied. The distributional patterns of the faunal communities seems to be controlled by the morphological features of the host-alga. The complex arborescent structure of the rhyzophytic H. incrassata species shelters fauna three times higher in abundance than other Halimeda heads. This fauna is mostly composed of a highly rich community of microcrustaceans and a very diversified community of Syllidae polychaetes. H. macroloba living on the outer reef flat retains a lot of small Nereidae polychaetes and a rich microgastropod assemblage. Dense H. opuntia tussocks (type A) on the fringing-reef and isolated H. opuntia fronds of pendulous chains (type B) on the barrier reef as well as beds of scattered H. opuntia (type C) on the outer reef flat provide environment for larger animals, and according to their zone of life, they retain very different faunal associations. Some ophiuroids, and some fish, collected in H. opuntia A and C, are newly reported from the Society Islands, French Polynesia, and the Pacific province. One Brachyurid species is presumably undescribed.     

Sedimentation on three caribbean atolls: Glovers reef,lighthouse reef and turneffe Islands,belize

Summary The chief mode of carbonate sedimentation on the Belizean atolls Glovers Reef, Lighthouse Reef and Turneffe Islands is the accumulation of organically-derived particles. Variations in the distribution of the composition and grain-sizes of surface sediments, collected along transects across the atolls, are environmentally controlled. Two major sediment types may be distinguished. (1) Reef and fore reef sediments are dominated by fragments of coral, coralline algae andHalimeda. Mean grain-sizes range from 1–2 mm. (2) Back reef sediments contain more mollusk fragments, more fine-grained sediment (<125 μm) and appear to have fewerHalimeda fragments. In addition, sediments from inner platforms and shallow lagoonal parts of Glovers and Lighthouse Reefs comprise non-skeletal grains, namely fecal pellets. Sediments from lagoonal patch reefs may contain up to 20% coral fragments. Mean grain-sizes range from 0.1–1 mm and are finest on the inner platform and lagoon floor of the back reef environment. Within the reef and fore reef environments, it is not possible to distinguish sub-environments on the basis of textural and compositional differences of the sediments. Sediments from patch reefs contrast with those from back reef lagoons and inner platforms and are similar in terms of grain-sizes and compositions to reef and fore reef surface sediments. Non-skeletal grains forming in shallow parts of the back reef in Glovers and Lighthouse Reefs are interpreted to be indurated by interstitial precipitation of calcium carbonate from warm, supersaturated water flushing the sediment. The lack of hardened non-skeletal particles in the back reef sediments of Turneffe Islands is most probably due to the abundance of muddy, organic-rich sediment in the well-protected lagoon. Fine sediment is less permeable and organic films prevent cement overgrowth on particles.     

Significance of <Emphasis Type="Italic">Halimeda</Emphasis> bioherms to the global carbonate budget based on a geological sediment budget for the Northern Great Barrier Reef,Australia

Since the correlation between carbon dioxide (CO₂) levels and global temperatures was established in the ice core records, quantifying the components of the global carbon cycle has become a priority with a view to constraining models of the climate system. The marine carbonate budget is still not adequately constrained and the quantitative significance of the calcareous green alga Halimeda still remains particularly poorly understood. Previously, it has been suggested that Halimeda bioherms on the shelf of the Great Barrier Reef may contain a volume of carbonate equal to or greater than that contained within the shelf edge coral reefs. This study uses published datasets to test this hypothesis in the Northern Great Barrier Reef (NGBR) province. It is estimated that Halimeda bioherms on the outer shelf of the NGBR contain at least as much (and up to four times more) CaCO₃ sediment as the adjacent ribbon reef facies. Globally, if these findings are even only partially applicable, the contribution of shallow water carbonate sediments to the global carbon budget based on coral reefs alone is currently substantially underestimated.     

Growth rate,ultrastructure and sediment contribution of Halimeda incrassata and Halimeda monile,Nonsuch and Falmouth Bays,Antigua, W.I.

Halimeda incrassata and Halimeda monile, the two dominant rhipsalian Halimeda, were evaluated behind a bank barrier reef, in a fringing reef lagoon and in an open lagoon. Growth was calculated in number of segments, weight of segments and turnover rate. More than 1800 plants were stained with Alizarin Red-S dye, yielding average number of segments/plant/day and g CaCO₃/m²/year for each of the above areas of 2.17/114, 1.43/65.7 and 1.6/56.9, respectively. Average weight CaCO₃/segment was 4 mg. SEM revealed ultrastructure of short and long unoriented aragonite crystals forming in new segments within 24 h and an effective holdfast system with filaments partially coated with carbonate fragments. Greatest growth occurred within thin to medium density grass beds. In Nonsuch Bay sediment production from these two species alone was 0.057 mm/year or 1 1/2 orders of magnitude less than estimates of the total production from all Halimeda species (1.01 mm/year) over the past 6745 years.     

Studies on Halimeda

Large areas of the inter-reefal seabed in the Great Barrier Reef are carpeted with vegetation composed almost entirely of the green calcareous alga Halineda. These meadows occur principally in the northern sections between 11°30 and 15°35S at depths of 20 to 40 m, but there are also some in the central and southern sections, where they have been found at depths down to 96 m. The vegetation is dominated by the same sprawling Halimeda species that are common on coral reefs in this region. However, on reefs these species grow on solid substrata, not soft sediments like the Halimeda-rich gravels that underlie the meadows. A total of 12 Halimeda species, together with two Udotea and one Penicillus species, are characteristic components of the shallow meadows. Below 50 m depth, species composition is restricted to only two major components. One, H. copiosa, is also important shallower, but the other is an unusually large and heavily calcified form of H. fragilis, a species that is normally a minor, fragile component of the shallow meadows. The maximum biomass found in these meadows was 4637 gm² of calcareous algae, although the thean for vegetated areas was 525 gm². These meadows are confined to the nutrient-depleted waters of the outer continental shelf just inside the outer barrier reefs, and are usually associated with distinct shoaling of the seabed caused by accumulation of thick deposits of calcareous Halimeda segments. The meadows are probably supported by very localized upwelling of nutrients from the adjacent Coral Sea onto the shelf, where they enrich the otherwise nutrient-depleted waters.Contribution No.367 from the Australian Institute of Marine Science     

Halimeda growth and diversity on the deep fore-reef of Enewetak Atoll

The deep fore-reef at Enewetak has been examined from the submersible Makali'i. Green algae grow to about-150 m at photon flux densities of approximately 1 Em^-2s^-1. Halimeda cover is 50% at many sites down to-90 m. Halimeda populations are important within the zone of scleractinian corals down to about-65 m, while a Halimeda zone with low coral cover or lacking corals between-65 m and-150 m probably is an important source of reef carbonate. Halimedas of the deep fore-reef, like those of the lagoon, constitute an important structural component in reef building. Other calcareous green algae such as Tydemania are less important on the deep fore-reef, but growth of coralline red algae continues to over-200m. Halimeda diversity is high down to near the base of the euphotic zone.     

Leeward bank margin Halimeda meadows and draperies and their sedimentary importance on the western Great Bahama Bank slope

Bryopsidalean algal meadows in water depths of 20–40 m on the leeward side of western Great Bahama Bank (WGBB) lie between non-skeletal-dominated sand flats on the bank top to the east and a cemented steep escarpment to the west. The meadows contain dense populations of rhipsalian Halimeda species, as well as Udotea and Rhipocephalus. Extensive populations of other Halimeda species (opuntioids) occur at greater depths on the cemented rocky escarpment, growing as drapes or vines rather than as upright thalli. These meadows and draperies are important sources of coarse-grained carbonate sediments. This is shown by (1) deeper bank-edge sediments (30–60 m) containing considerably more Halimeda fragments than do the bank top, non-skeletal sands, and (2) the coarser fraction of slope sediments (down to 200 m) dominated by Halimeda plates, partly or extensively altered and internally cemented by magnesian calcite and aragonite. A transect across the bank margin from bank top (<10 m) to lower slope (300 m) provides a useful comparison for the locus of sediment production and accumulation. The production of Halimeda in these bank-edge habitats approximates that in the Great Barrier Reef or off Indonesia and Nicaragua in similar water depths. The apparent lack of thick sediment accumulation in WGBB compared to that seen elsewhere may reflect the high rates of downslope transport off Great Bahama Bank.     

Phytoplankton biomass and production in shelf waters off NW Spain: spatial and seasonal variability in relation to upwelling

Summary Chlorophyll-a and primary production on the euphotic zone of the N-NW Spanish shelf were studied at 125 stations between 1984 and 1992. Three geographic areas (Cantabrian Sea, Rías Altas and Was Baixas), three bathymetric ranges (20 to 60 m, 60 to 150 m and stations deeper than 200 m), and four oceanographic stages (spring and autumn blooms, summer upwelling, summer stratification and winter mixing) were considered. One of the major sources of variability of chlorophyll and production data was season. Bloom and summer upwelling stages have equivalent mean and maximum values. Average chlorophyll-a concentrations approximately doubled in every step of the increasing productivity sequence: winter mixing — summer stratification — high productivity (upwelling and bloom) stages. Average primary production rates increased only 60% in the described sequence. Mean (± sd) values of chlorophyll-a and primary production rates during the high productivity stages were 59.7 ± 39.5 mg Chl-a m^–2 and 86.9 ± 44.0 mg C m^–2 h^–1, respectively. Significant differences in both chlorophyll and primary production resulted between geographic areas in most stages. Only 27 stations showed the effects of the summer upwelling that affected coastal areas in the Cantabrian Sea and Rías Baixas shelf, but also shelf-break stations in the Rías Altas area. The Rías Baixas area had lower chlorophyll than both the Rías Altas and the Cantabrian Sea areas during spring and autumn blooms, but higher during summer upwelling events. On the contrary, primary production rates were higher in the Rías Baixas area during blooms in spring and autumn. Mid-shelf areas showed the highest chlorophyll concentrations during high productivity stages, probably due to the existence of frontal zones in all geographic areas considered. The estimated phytoplankton growth rates were comparable to those of other coastal upwelling systems, with average values lower than the maximum potential growth rates. Doubling rates for upwelling and stratification stages in the northern and Rías Altas shelf areas were equivalent, despite larger biomass accumulations during upwelling events. Low turnover rates of the existing biomass in the Rías Baixas shelf in upwelling stages suggests that the accumulation of phytoplankton was due mainly to the export from the highly productive rías, while the contribution of in situ production to these accumulations was relatively lower.     

Trace and minor element ratios inHalimeda aragonite from the Great Barrier Reef

The calcareous green algaHalimeda can be a substantial contributor to aragonite sediment in reef ecosystems. In contrast to coral aragonite, little is known about the trace and minor element composition ofHalimeda aragonite, so it is difficult to test oceanographic hypotheses about factors controlling its past growth. We investigated adapting trace element cleaning protocols for modern and HoloceneHalimeda aragonite, modern and HoloceneHalimeda trace and minor element compositions, and the potential utility ofHalimeda aragonite for paleoceanographic investigations. We successfully adapted and applied sample treatment protocols developed for measuring trace elements in coral aragonite (generally less than 500 y old) toHalimeda aragonite (modern to approximately 5000 y old in this study). ModernHalimeda aragonite from John Brewer Reef in the Central GBR had mean Cd/Ca ratios of 5.19 ± 1.68 nmol/mol forHalimeda micronesica and 2.35 ± 0.38 nmol/mol for three closely related species important in bioherm accumulationHalimeda copiosa, Halimeda hederacea, andHalimeda opuntia. Mn/Ca ratios, with means from 89–239 nmol/mol for these four species, showed both intra-and inter-specific variability. Sr/Ca ratios (10.9 ± O.1 mmol/mol) and Mg/Ca ratios (1.35 ± 0.26 mmol/mol) were similar for all samples. HoloceneHalimeda aragonite samples from cores of two bioherms in the northern GBR seemed well preserved on the basis of mineralogy and Sr/Ca and Mg/Ca ratios similar to those in modernHalimeda aragonite. Cd/Ca ratios (overall mean 0.96 ± 0.15 nmol/mol) were lower than those measured in the modernHalimeda from the central GBR location. However, Mn/Ca ratios in both cores were substantially higher than in modernHalimeda aragonite. While it may be possible to extract paleoceanographic information fromHalimeda aragonite, substantial care is needed to evaluate and avoid the effects of post-depositional alteration.     

Tectonic stability since the last interglacial offsets the Glorieuses Islands from the nearby Comoros archipelago

The four fossil terraces of the Glorieuses Islands are described, and new dates are provided to resolve their stratigraphy, depositional setting, and tectonic behavior. Most outcrops consist of a single sedimentary unit that represents the remains of an extensive reef flat dominated by Isopora palifera corals. At Lys Island, this unit is locally overlain by dipping layered beds composed of Halimeda segments, tentatively interpreted as storm overwash. Reliable U/Th dates obtained from corals sampled from the fossil outcrops mostly fall between 127 and 123 kyr, suggesting that these reefs formed exclusively during the first MIS-5e sea-level highstand, when sea level reached +3 m. The mean elevation of these terraces being +4.5 m, an uplift of 0.012 ± 0.002 mm year^?1 is inferred. This relative tectonic stability contrasts with the subsidence reported from Mayotte Island, suggesting a different geologic setting for the nearby Comoros and Glorieuses archipelagoes.     

Diversity of Halimeda (Bryopsidales,Chlorophyta) in New Caledonia: a Combined Morphological and Molecular Study

Halimeda is a genus of calcified and segmented green macroalgae in the order Bryopsidales. In New Caledonia, the genus is abundant and represents an important part of the reef flora. Previous studies recorded 19 species that were identified using morphological criteria. The aim of this work was to reassess the diversity of the genus in New Caledonia using morpho‐anatomical examinations and molecular analyses of the plastid tufA and rbcL genes. Our results suggest the occurrence of 22 species. Three of these are reported for the first time from New Caledonia: Halimeda kanaloana, H. xishaensis, and an entity resembling H. stuposa. DNA analyses revealed that the species H. fragilis exhibits cryptic or pseudocryptic diversity in New Caledonia. We also show less conclusive evidence for cryptic species within H. taenicola     

Morphology and sedimentology of Halimeda bioherms from the eastern Java Sea (Indonesia)

Halimeda bioherms, occurring primarily along the western and southern margins of Kalukalukuang Bank in the eastern Java Sea, display a wide variety of thicknesses and shapes. In general, high-frequency forms of the northern bank are replaced by thicker and lower frequency forms along the deeper southern margin. Sidescan sonar data suggest aperiodic reworking of shallow bioherm crests of the northern bank into features suggestive of bedforms. These features are not associated with deeper bioherms of the southern bank. Cores from the bioherms indicate that they consist mostly of disarticulated Halimeda plates set in a lime mud matrix composed largely of Halimeda fragments and foraminifera tests. Carbon-14 dating shows that, with exception of some deep southern bank examples, bioherms are actively accreting. Results of mineralogy and elemental chemistry on piston core PC-12 suggest cyclic variations in Mg-calcite (cement in Halimeda utricles), which may be related to periodic excursions of cold Pacific throughflow water onto the bank. Composition of the Mg-calcite (8.6 mole-%) suggests a temperature of formation of about 22 °C, which is 7 °C below average surface water temperatures. The carbon and oxygen isotope compositions of both aragonite and Mg-calcite phases are remarkably homogeneous, but were inconclusive with regard to the cold-water intrusion hypothesis. However, a lack of reef-building corals below a depth of 15 m, abundance of Halimeda bioherms on the western margin of K-Bank, where upwelling is predicted, extensive boring of sedimentary particles by endolithic boring algae, and high nutrient values of water at the thermocline (50–70 m deep) all support the incursion of cold, nutrient-rich water onto the bank. Upwelling and nutrient overloading are suggested as explanations for remarkable algal growth at the expense of reef-building corals.     

Comparison between community structures of fishes in <Emphasis Type="Italic">Enhalus acoroides</Emphasis>- and <Emphasis Type="Italic">Thalassia hemprichii</Emphasis>-dominated seagrass beds on fringing coral reefs in the Ryukyu Islands,Japan

To clarify differences in community structures and habitat utilization patterns of fishes in Enhalus acoroides- and Thalassia hemprichii-dominated seagrass beds on fringing coral reefs, visual censuses were conducted at Iriomote and Ishigaki islands, southern Japan. The numbers of fish species and individuals were significantly higher in the E. acoroides bed than in the T. hemprichii bed, although the 15 most dominant fishes in each seagrass bed were similar. Cluster and ordination analyses based on the number of individuals of each fish species also demonstrated that fish community structures were similar in the two seagrass beds. Species and individual numbers of coral reef fishes which utilized the seagrass beds numbered less than about 15% of whole coral reef fish numbers, although they comprised about half of the seagrass bed fishes. Of the 15 most dominant species, 5 occurred only in the two seagrass beds, including seagrass feeders. Ten other species were reef species, their habitat utilization patterns not differing greatly between the two seagrass beds. Some reef species, such as Lethrinus atkinsoni and L. obsoletus, showed ontogenetic habitat shifts with growth, from the seagrass beds to the coral areas. These results indicate that community structures and habitat utilization patterns of fishes were similar between E. acoroides- and T. hemprichii-dominated seagrass beds, whereas many coral reef fishes hardly utilized the seagrass beds.     

Skeletal composition of modern lagoon sediments in New Caledonia: Coral,a minor constituent

The skeletal composition of 273 sediment samples, collected within 14615 km² of lagoon habitat in New Caledonia (Ouvea and Chesterfield atolls and eastern and northern lagoons of the main island), was analyzed. Major constituents were molluscs (bivalves and gastropods), foraminifers, andHalimeda plates. The quantitative examination showed that, even in a pure coralline structure such as the two atolls studied, coral debris and calcareous algae, potentially produced within the barrier reef, never constituted a dominant element in the lagoonal sediments. Distribution of coral debris showed that coral is significant only close to the barrier reef (i.e. passes and back-reef slope). From the point of view of sedimentology, this suggests that the major role of the barrier reef is to provide a physical barrier that allows the development and preservation of lagoon sediments. Sedimentation within the lagoon of grains coarser than 63 µm is the result of in situ organic production combined with low hydrodynamic control.     

Ben Franklin temperate reef and deep sea `Agassiz Coral Hills' in the Blake Plateau off North Carolina

This paper deals with two deep water coral ecosystems off the North Carolina coast, both under the depositional, erosional, and biological influence of the Gulf Stream and its eddies and episodic upwelling processes. The first coral community, Ben Franklin temperate reef, is located at 20 meters in Onslow Bay and is characterized by the ahermatypic coral Oculina arbuscula Verrill and unusually high abundance of the small predatory isopod Eurydice bowmani George and Longerbeam. The deep sea coral community, Agassiz Coral Hills, is located over the Blake Plateau at 650 meters, dominated by the fossilized, dead and living coral Bathypsammia tintinnabulum, with rare occurrence of the deep sea solitary coral Flabellum goodei. The deep sea coral sites were sampled with an otter trawl, outfitted with `Benthos' flotation spheres. The results suggest that the fish and shrimp fauna exhibits high species richness in the Agassiz Coral Hills off North Carolina in comparison with a control site over the Blake Plateau off Florida. Species richness may be linked to episodic upwelling events along the Gulf Stream meanders. The fish fauna in the Agassiz Coral Hills includes the following two commercially important deep sea fish species: the wreck fish Polyprion americanus and the eel Synaphobranchus koupi. The Ben Franklin reef and the Agassiz Coral Hills are recommended as Marine Protected Areas.     

Calcium carbonate productivity by Halimeda macroloba in the tropical intertidal ecosystem: The significant contributor to global carbonate budgets

Halimeda is a potential carbon sink species and an important player in the global carbonate budget. The objectives of this study were to: (i) examine the CaCO₃ and sediment productions of H. macroloba by measuring the density, growth rate, and recruitment; (ii) quantify the numbers of aragonite crystals; (iii) document reproductive events; and (iv) determine the life‐span. This study was carried out at Lidee Lek Island, Satun, Thailand during July 2015 to April 2016. The density was measured using quadrats (0.25 m²) and three 50 m line transects. Alizarin Red‐S marking technique was used for the growth rate and CaCO₃ accumulation rate assessments. The recruitment, reproduction and life‐span were measured by tagging 500 individuals. Tagged individuals and new plants were counted. In this study, mean and the highest density of Halimeda were 44.42 ± 13.95 and 138.22 ± 11.68 thalli m^?2, respectively, and Halimeda produced 1–2 new segments.thallus^?1 day^?1 or 0.021 ± 0.001 g dry weight.thallus^?1.day^?1. The annual biomass production was 1910–5950 g m^?2 year.^?1. There was a low rate of occurrence of sexual reproduction, observed in late July to September, ranging from 0.17% to 1.92%. For the mortality and recruitment rates, approximately 70–80% of individuals were lost during July to September 2015, probably from sexual reproduction and the recruitment rate varied from 5.36 ± 0.79% to 21.03 ± 2.33%. The highest density of new recruits was found in September 2015 right after the sexual reproductive event occurred. New recruits have been found up to April 2016 without any reproductive events, suggesting that both sexual and asexual reproduction helped maintain the population. The life span of Halimeda was 8–12 months. In addition, Halimeda accumulated CaCO₃ at approximately 0.018 g CaCO₃ thallus^?1 day^?1 and produced CaCO₃ at approximately 291.94–908.11 g m^?2 year^?1, indicating that Halimeda contributes to CaCO₃ and helps to sink carbon through calcification. The results in terms of the density, growth rate, and CaCO₃ accumulation rate can be used to calculate the mass of carbonate sediment contributed by Halimeda.     

Development and eustatic control of an Upper Jurassic reef complex (Saint Germain-de-Joux,Eastern France)

Summary This study consists of a sedimentological and diagenetical analysis of reef facies from the Upper Kimmeridgian (sensu gallico). The investigated deposits are situated in eastern France, about fifty kilometres west of the city of Geneva (Switzerland). The reef complex is a fine example of vertical development and facies differentiation. It is subdivided into two distinct sequences by a perforated hardground horizon and sand shoals. The onset of the first reef sequence is characterized by a pioneer growth stage followed by up to 20 m of reef-core and-flank facies. Corals forming the reef-core are typically the ramose variety ofCalamophylliopsis flabellum. The second reef sequence has a reef-core with an average thickness of about 5 m. Corals, however, display much more varied morphologies, and in some areas massive rudist (Heterodiceras) build-ups occur. Development of the second reef sequence was seriously weakened by a storm which produced a 2 m thick accumulation of coral rubble. A shallowing-upwards trend gradually leads to the formation of beach deposits, followed by a newly detected black-pebble horizon. Diagenesis is an important aspect of the reef complex. Especially noteworthy is the dolomitization of certain horizons. At the base of the reef formation, the passage of the phreatic mixing zone provoked invasive dolomitization in large irregular patches (probably deposits richer in Mg-calcite). Some of the beds above the black-pebble horizon, in particular a deposit of accumulated microbial mats, are also dolomitized. In this case, dolomitization is stratiform and is interpreted as having precipitated under conditions of evaporative pumping. The sedimentary record clearly shows the imprint of eustasy. The reef complex was initiated during a transgressive cycle and the hardground found between the two reef sequences is interpreted as a maximum flooding surface (mfs). At the top of the sequence, the horizon overlain by the black-pebble conglomerate is believed to represent the new sequence boundary SB140. Other significant features identified from the St. Germain-de-Joux deposits include the discovery of a new foraminifera,Troglotella incrustans, which is only marginally covered here but is the topic of another paper (Wernli & Fookes, 1992); the subdivision of the first coralligenous level defined byPelletier (1953) into two reef sequences; and a proposition to redefine the ‘Calcaires de la Semine’ (Bernier, 1984). The investigations carried out in the past on the Kimmeridgian deposits in the area of St. Germain-de-Joux were mostly based on stratigraphy and palaeontology. These reefs are among the finest known in the Jura Mountains, but no thorough study had been made on their sedimentological aspects. The aim of this study is to fill this void and also to clarify the more confusing aspects of local stratigraphy (paper based onFookes, 1991).     

The ploys of sex: relationships among the mode of reproduction,body size and habitats of coral-reef brittlestars

Observations were made of 33 species of brittlestars (3980 specimens) from specific substrata collected in four zones on the Belize Barrier Reef, Caribbean Sea. The body size of most species of brittlestars with planktonic larvae differs significantly among different substrata. Generally, individuals from the calcareous alga Halimeda opuntia are smallest, those found in corals (Porites porites, Madracis mirabilis, and Agaricia tenuifolia) are larger, and those from coral rubble are the largest. This suggests that brittlestars with planktonic larvae move to new microhabitats as they grow. In contrast, most brooding and fissiparous species are relatively small and their size-distributions are similar among all substrata. Halimeda harbours denser concentrations of brittlestars and more small and juvenile individuals than the other substrata. Juveniles of the brooding and fissiparous species are most common in Halimeda on the Back Reef whereas juveniles developing from planktonic larvae are most common in Halimeda patches in deeper water. Fissiparity and brooding may be means for individuals (genomes) of small, apomictic species to reach large size (and correspondingly high fecundities) in patchy microhabitats that select for small body sizes. Small brittlestar species and juveniles are most numerous in the microhabitats called refuge-substrata, such as Halimeda, which may repel predators and reduce environmental stress. Whether young brittlestars are concentrated in refuge-substrata through settlement behavior, migration, or differential survival remains unknown. Experiments revealed that coral polyps kill small brittlestars, perhaps accounting for the rarity of small and juvenile brittlestars in coral substrata.