Seismic characteristics and accretion history of Halimeda bioherms on Kalukalukuang Bank,eastern Java Sea (Indonesia)

Extensive areas of Halimeda bioherms similar to those described by Orme et al. (1978), Orme (1985), Davies and Marshall (1985), and Phipps et al. (1985) from Australia's Great Barrier Reef have formed on Kalukalukuang Bank (K-Bank) 50 km east of the Sunda Shelf margin in the easter Java Sea. K-Bank is an isolated limestone platform whose top slopes southward from a water depth of about 20 m at the north to about 100 m at the south (30 km). It occurs in a bidirectional monsoonal wind regime and a predominantly southerly flowing current from Makassar Strait. The water column around K-Bank has a well developed shallow thermocline (50 m to 150 m). K-Bank has a relatively flat top with marginal banks of suspected Pleistocene origin as interpreted from seismic relationships. A reconnaissancelevel survey grid of high-resolution seismic profiles indicates that Halimeda bioherms are restricted to the bank margins with the exception of the eastern margin. Bioherms either extend to the steep margin of the platforms or are separated from the platform edge by banks of coral and coralline algae. The morphology of the Halimeda bioherms varies from steep-sided, elongate ridges in the northern bank area, through coalescing symmetrical mounds with partly infilled valleys, to broad undulating areas similar to those described by Orme (1985) from the Great Barrier Reef. High-resolution seismic records indicate erosional episodes in the high-relief areas, an interpretation that seems to be supported by accumulation rates calculated from C¹⁴ dates of cores. Thicknesses of Halimeda accumulation above a prominent reflector considered as Pleitocene, vary from around 20 m in the north to a maximum of over 50 m in the southwest. Accumulation rates obtained from dating of two cores to-ward the north of K-Bank, average 0.294 m/100 yr for core VC4 and a maximum of 0.59 m/100 yr in a part of core PC12. Dating of material from the top 30 cm of a deep bioherm (100 m) in the southwest of K-Bank, indicates that the growth rate of these bioherms has slowed markedly, presumably because of increasing water depth (decreasing light) over the Holocene transgression.     

Halimeda bioherms along an open seaway: Miskito Channel,Nicaraguan Rise,SW Caribbean Sea

A recent research cruise to examine small, detached carbonate platforms situated on the Nicaraguan Rise in the SW Caribbean Sea has revealed the presence of numerous Halimeda bioherms. Based upon interpretations from seismic reflection data some exceed 140 m in relief. This is the first documented occurrence of these green-algal buildups in the Caribbean/Bahama Bank region. The Halimeda bioherms form a nearly continuous band that borders the margins of the Miskito Channel—a shallow, open, 125 km long seaway. This 220 m deep channel bisects the Miskito Bank which is a major carbonate shelf. In seismic profile the bioherms appear acoustically soft and reveal a local relief of 20–30 m. Tops of these features lie in about 40–50 m of water. Samples from dredge hauls are coarse, poorly cemented packstones/grainstones which are dominated by largely unbroken, disarticulated Halimeda segments set in a poorly sorted sandy matrix. Exposed surfaces were stained brown. Very little living material was brought up in the dredges. The significance of these bioherms and their full extent in the Caribbean are not understood. Undoubtedly, further study will provide important answers concerning their role in the geologic development of Caribbean carbonate platforms.     

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.     

Hard-coral distribution and cold-water disturbances in South Florida: variation with depth and location

Line transects were used to sample the structure and diversity of the hermatypic coral community (Scleractinia) on four shallow shelf-edge reefs in South Florida (25° 22N to 25° 29N). Low diversities, cover and abundance indicated that this area was a suboptimum habitat for reef-building corals. The lack of acroporids on the shallow fore-reef, the increase in total coral cover with depth and the greater abundance of Montastrea annularis in the deepest zones suggests that cooling of surface water during severe winter cold fronts is a major environmental control on the distribution of hermatypic corals with depth. Such disturbances, occurring more frequently than hurricanes, may preclude the hard-coral community from attaining higher levels of cover and abundance. The shallow zones on the reefs nearest to tidal passes, through which cooled by water enters the reef tract, had the least developed community. In the deeper reef-zones, species richness and abundance increased from north to south over a distance of 13 km.     

Form and seismic stratigraphy of Halimeda banks in part of the northern Great Barrier Reef Province

Twenty-six percent of the total shelf area in the Northern Region of the Great Barrier Reef Province between latitudes 14°27 and 15°02S is occupied by algal (Halimeda) lithofacies. Sea-bed relief over this bankforming algal lithofacies, which dominates the outer shelf, is generally complex and variable over relatively short distances, but lateral continuity of morphological features near Petricola and Stewart shoals has been demonstrated by parallel profiling. The most prominent subbttom reflector is a pre-Holocene erosion surface, regarded as the Holocene/Pleistocene unconformity. In this area, the maximum thickness of Holocene Halimeda banks above the main subbottom reflector is 19 m. Seismic response suggests three main sequences in the Halimeda bank depostis, which probably relate to changes in environmental conditions, and the recognition of similar seismic characteristics in the deposits beneath the Holocene/Pleistocene unconformity indicates that Halimeda banks may have been a feature of the outer shelf of the Great Barrier Reef in Plesitocene times.     

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.     

Devonian carbonate buildup facies in an intra-oceanic island arc (Tamworth Belt, New South-Wales, Australia)

Summary  Biohermal and biostromal buildups were investigated in late Early and Middle Devonian carbonate complexes of the Tamworth Belt. The buildup types and subtypes were studied in three regions (Yarramanbully, Sulcor and, Wyaralong') to clarify their paleo-environmental position. Two stages of development are recognized: Incipient bioherms and bioherms. Incipient bioherms are carbonate buildups with organisms which commonly form true bioherms. They dominate the sediment with small growth forms but are not prolific enough to build large bio-frameworks. Small nodular and globular stromatoporoids characterize the incipient bioherms and are interpreted as stunted growth forms. In one location (‘Wyaralong’) the coarse stromatoporoid calcarenite represents a fore-reef facies, at Sulcor a shallow subtidal setting with moderate water energy can be deduced. The bioherms can be sub-divided into stromatoporoid-, stromatoporoid-Stachyodes-, and stromatoporoid-rugose coral bioherms. Their variable composition probably reflects growth and deposition in different zones of a reef complex and/or different proximity to areas of denundation indicated by high siliciclastic input. In the Tamworth region true bioherms occur only in the Moore Creek Limestone Member (Middle Devonian), and not in older carbonate successions. Biostromes are sub-divided into (1) incipient biostromes with stromatoporoid-heliolitid biostromes and alveolitid biostromes; (2) aggregate biostromes withAmphipora andStachyodes biostromes; (3) stratified biostromes; (4) mixed aggregate/stratified biostromes. The different types of biostromes are not limited to specific time-intervals, but rather to environmental conditions.

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     

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.     

Boring by Macro-organisms in the Coral Montastrea annularis on Barbados Reefs

The fauna boring into Montastrea annularis includes sponges, bivalves, sipunculid and polychaete worms and barnacles. Sponges are most important in hard tissue destruction and account for more than 90% of the total boring in most heads. Bivalves and barnacles are locally important. Sipunculids and sabellids account for less than 4% of the total boring. The volume removed from coral samples by boring ranged from 3–60% and samples from a deeper bank reef were more highly bored than fringing reef samples. An average of 20% of the volume of bank reef corals, and 5% of the volume of fringing reef corals, was removed by boring. The distribution of individual borers is not a function of depth. The density and variety of borers and the extent of boring in coral heads is greater in older heads. The ratio of living coral surface to dead encrusted areas on colonies also influences borer density and the extent of boring.     

Importance of foraminifera for the formation and maintenance of a coral sand cay: Green Island, Australia

 CaCO₃ production by reef-building organisms on Green Island Reef in the Great Barrier Reef of Australia is estimated and compared with the contribution of benthic foraminifera to the sediment mass of the vegetated sand cay. Major constituents of the cay are benthic foraminifera (mainly Amphistegina lessonii, Baculogypsina sphaerulata, and Calcarina hispida), calcareous algae (Halimeda and coralline algae), hermatypic corals, and molluscs. Among these reef-building organisms, benthic foraminifera are the single most important contributor to the sediment mass of the island (ca. 30% of total sediments), although their production of CaCO₃ is smaller than other reef-building organisms. Water current measurements and sediment traps indicate that the velocity of the current around Green Island is suitable for transportation and deposition of foraminiferal tests. Abundant foraminifera presently live in association with algal turf on the shallow exposed reef flat, whose tests were accumulated by waves resulting in the formation and maintenance of the coral sand cay. Accepted: 30 June 1999     

Halimeda bioherms of the northern Great Barrier Reef

The reefless tract directly behind the ribbon reefs on the outer shelf off Cooktown supports a luxuriant growth of Halimeda that, during the Holocene, has developed into bioherms. These mounded biodies of unconsolidated sediment have formed banks that vary in height between 2 and 20 m. Combined shallow, high-resolution seismic reflection profiles and side-scan sonar have diferentiated three areas of biohermal complexes behind the ribbon reefs of Cooktown. Observations by SCUBA and submersible plus the sedimentology of the bioherms indicate that they are in situ accumulations. Evidence from dating of cores suggests that the Halimeda bioherms began to grow about 10 000 years B.P. and their growth has continued to the present time, even though their tops are presently restricted to a depth of -20 m. It is suggested that the origin and morphology of the bioherms are related to a specific hydrodynamic phenomenon, involving jets of nutrient-rich, upwelled oceanic water intruding onto the outer shelf via the narrow passes between the ribbon reefs, and forming eddies behind the ribbons.     

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.     

Reef coral survival and mortality at low temperatures in the Arabian Gulf: new species-specific lower temperature limits

A series of cold fronts passing over the western Arabian Gulf from December 1988 to March 1989 produced the longest period of sustained low water temperatures ever recorded in a coral reef area. Sea water temperatures recorded on two reefs during this period provide new estimates of lower thermal limits for reef coral survival. Severe mortality of the corals Acropora pharaonis and Platygyra daedalea occurred at the northern site where minimum temperatures fell below 11.5°C on four consecutive days and mean daily temperatures were 13°C or less for more than 30 days. However, Porites compressa, the principal reef-former in this area, and various faviid corals initially showed only sub-lethal effects and appeared normal after six months. Corals were not damaged at the southern site, where minimum water temperature fell below 12.5°C for two consecutive days, but mean temperatures were 14°C or less for only 5 non-consecutive days.     

New constraints on the spatial distribution and morphology of the Halimeda bioherms of the Great Barrier Reef,Australia

Halimeda bioherms occur as extensive geological structures on the northern Great Barrier Reef (GBR), Australia. We present the most complete, high-resolution spatial mapping of the northern GBR Halimeda bioherms, based on new airborne lidar and multibeam echosounder bathymetry data. Our analysis reveals that bioherm morphology does not conform to the previous model of parallel ridges and troughs, but is far more complex than previously thought. We define and describe three morphological sub-types: reticulate, annulate, and undulate, which are distributed in a cross-shelf pattern of reduced complexity from east to west. The northern GBR bioherms cover an area of 6095 km², three times larger than the original estimate, exceeding the area and volume of calcium carbonate in the adjacent modern shelf-edge barrier reefs. We have mapped a 1740 km² bioherm complex north of Raine Island in the Cape York region not previously recorded, extending the northern limit by more than 1° of latitude. Bioherm formation and distribution are controlled by a complex interaction of outer-shelf geometry, regional and local currents, coupled with the morphology and depth of continental slope submarine canyons determining the delivery of cool, nutrient-rich water upwelling through inter-reef passages. Distribution and mapping of Halimeda bioherms in relation to Great Barrier Reef Marine Park Authority bioregion classifications and management zones are inconsistent and currently poorly defined due to a lack of high-resolution data not available until now. These new estimates of bioherm spatial distribution and morphology have implications for understanding the role these geological features play as structurally complex and productive inter-reef habitats, and as calcium carbonate sinks which record a complete history of the Holocene post-glacial marine transgression in the northern GBR.

     

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.     

Presence of Symbiodinium spp. in macroalgal microhabitats from the southern Great Barrier Reef

Coral reefs are highly dependent on the mutualistic symbiosis between reef-building corals and dinoflagellates from the genus Symbiodinium. These dinoflagellates spend part of their life cycle outside the coral host and in the majority of the cases have to re-infect corals each generation. While considerable insight has been gained about Symbiodinium in corals, little is known about the ecology and biology of Symbiodinium in other reef microhabitats. This study documents Symbiodinium associating with benthic macroalgae on the southern Great Barrier Reef, including some Symbiodinium that are genetically close to the symbiotic strains from reef-building corals. It is possible that some of these Symbiodinium were in hospite, associated to soritid foraminifera or ciliates; nevertheless, the presence of Symbiodinium C3 and C15 in macroalgal microhabitats may also suggest a potential link between communities of Symbiodinium associating with both coral hosts and macroalgae.     

Distribution,food preference,and trophic position of the corallivorous fireworm Hermodice carunculata in a Caribbean coral reef

The fireworm Hermodice carunculata is a facultative corallivore on coral reefs. It can interact with algal overgrowth to cause coral mortality. However, because of its cryptic nature, little is known about its ecology. We used micropredator attracting devices (MADs) and stable isotope analyses to provide insights into the distribution and diet of H. carunculata in a coral reef on Curaçao, southern Caribbean. MADs consisted of algal clumps inside accessible mesh nets which H. carunculata could use as refuge. To obtain indications on its distribution pattern, MADs filled with Halimeda opuntia were deployed in different reef habitats ranging from 0 to 16 m water depth. Fireworms were found inside MADs in all reef habitats, indicating that they have a widespread horizontal and vertical distribution, ranging from the shoreline to the deeper reef slope. On the reef crest, MADs were filled using different algal species and deployed on dead or live scleractinian corals. MADs hosted more fireworms when placed on live corals, regardless of algal species used, suggesting that algal-induced corallivory may be widespread. To test for food preferences, different food sources were added inside the MADs. Fireworms detected potential prey within 6 h and were significantly more attracted by decaying corals and raw fish than by live corals, hydrozoans, or gorgonians. Stable isotope analyses indicated detritus, macroalgae, and scleractinian corals as potential food sources and revealed an ontogenetic dietary shift toward enriched δ ¹³C and δ ¹⁵N values with increasing fireworm size, suggesting that large-sized individuals feed on food sources of higher trophic levels. Our findings highlight H. carunculata as a widespread, and omnivorous scavenger that has the potential to switch feeding toward weakened or stressed corals, thereby likely acting as a harmful corallivore on degraded reefs.     

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.     

Dynamics and limitations of phytoplankton biomass along a gradient in Mwanza Gulf,southern Lake Victoria (Tanzania)

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