Grain size distribution of the lagoonal deposits within the South Malé Atoll, Maldives, Indian Ocean

Seismic and multibeam data, as well as sediment samples were acquired in the South Malé Atoll in the Maldives archipelago in 2011 to unravel the stratigraphy and facies of the lagoonal deposits. Multichannel seismic lines show that the sedimentary succession locally reaches a maximum thickness of 15-20 m above an unconformity interpreted as the emersion surface which developed during the last glacial sea-level lowstand. Such depocenters are located in current-protected areas flanking the reef rim of the atoll or in infillings of karst dolinas. Much of the 50 m deep sea floor in the lagoon interior is current swept, and has no or very minor sediment cover. Erosive current moats line drowned patch reefs, whereas other areas are characterized by nondeposition. Karst sink holes, blue holes and karst valleys occur throughout the lagoon, from its rim to its center. Lagoonal sediments are mostly carbonate rubble and coarse-grained carbonate sands with frequent large benthic foraminifers, Halimeda flakes, red algal nodules, mollusks, bioclasts, and intraclasts, some of them glauconitic, as well as very minor ooids. Finer-grained deposits locally are deposited in current-protected areas behind elongated faros, i.e., small atolls which are part of the rim of South Malé Atoll. The South Malé Atoll is a current-flushed atoll, where water and sediment export with the open sea is facilitated by the multiple passes dissecting the atoll rim. With an elevated reef rim and tower-like reefs in the atoll interior it is an example of a leaky bucket atoll which shares characteristics of incipiently drowned carbonate banks or drowning sequences as known from the geological record.

Paleoecology, Diagenesis and Biological Affinity of Tetradiids in the Upper Ordovician (Sandbian) Black River Group of the Kingston Area, Ontario, Canada

Tetradiids are a group of colonial, tubular fossils that occur globally in Middle to Upper Ordovician strata. Tetradiids were first described as a type of tabulate coral; however, based on their four-fold symmetry, division, and presence of a central-sparry canal, they were recently reinterpreted as a florideophyte rhodophyte algae, a reinterpretation that is tested in this thesis. This study focused on understanding the affinity and taphonomy of this order of fossil. Research was conducted by stratigraphic and petrographic analyses of the Black River Group in the Kingston, Ontario region. Tetradiid occurrences were divided into fragment or colonial, with three morphologies of tetradiids described (Tetradium, Phytopsis and Paratetradium). Morphology is specific to depositional environment, with compact Tetradium consistently within ooid grainstones and open branching Phytopsis and chained Paratetradium consistently within mudstones. Two types of patch reefs were recognized: a Paratetradium bioherm, and a Paratetradium, Phytopsis, stromatolite bioherm. The presence of bioherms implies that tetradiids were capable of hypercalcifying. Preservation styles of tetradiids were investigated, and were compared to brachiopods, echinoderms, mollusks, and ooids. Tetradiids were preferentially preserved as molds and demonstrated complete dissolution of skeletal material. Rare specimens, however, demonstrated preserved horizontal partitions, central plates, and a double wall. Skeletal molds were filled with either calcite spar, mud or encrusted by a cryptomicrobial colony. Both calcitic and aragonitic ooids were discovered. The co-occurrence of aragonitic ooids, aragonitic crytodontids, and the evolution of aragonitic, hypercalcifying tetradiids is interpreted as representing the geochemical favoring of aragonite and HMC in a time of global calcite seas. The geochemical favoring of aragonite is interpreted to be independent to global Mg: Ca ratios, but was the result of increased saturation levels and temperature driven by high atmospheric pCO2. Based on the presence of epitheca, tabulae, septa, and the commonality of growth forms, tetradiids are interpreted as an order of Cnidaria. The evolution of an aragonitic skeleton in tetradiids is interpreted to be the result of de novo acquisition of a skeleton from an unmineralized clade.

Paläobiogeographie des Korallenooliths (Mittleres Oxfordium - Unteres Kimmeridgium): Tethyale Faunen- und Florenelement auf höherer Paläobreite

An overview is given here on the palaeobiogeography of the Korallenoolith Formation (middle Oxfordian to early Kimmeridgian) in NW Germany (Lower Saxony Basin). Based on microfacies observations, abundant faunal and floral elements of the tropical tethyan realm are recognized in shallow-marine calcareous sediments of the Korallenoolith Formation. Foraminiferal fauna is both highly diverse and abundant and mostly of mediterranean character. Also, there is a small flora recorded, which includes heavily calcified red algae, aragonitic green algae, and cayeuxiid algae. They display restricted diversity when compared to those of shallow-marine tropical tethyan seas. Chaetetids and diceratids are locally abundant. Lithocodium aggregatum and Bacinella irregularis have been observed in Late Jurassic palaeolatitudes north of the Tethys for the first time. Corals are present in numerous genera and species. Their occurrence is restricted to a few horizons of the Korallenoolith Formation where they build patch reefs, coral biostroms and coral meadows. The overall character of the coral-thrombolite-reefs (florigemma-Bank Member) is very similar to those of the Tethys. The presence of these marine tethyan taxa assigned the position of the Lower Saxony Basin during middle Oxfordian to early Kimmeridgian palaeobiogeographically into the submediterranean province and reflects northward migration of tropical tethyan fauna and flora which reach in the Lower Saxony Basin their northern limit. These biota seem to be biogeo-graphically transitional between communities present in England and the Tethys.

Riffe und fazielle Entwicklung der florigemma-Bank (Korallenoolith, Oxfordium) im Süntel und östlichen Wesergebirge (NW-Deutschland

During the Sedimentation of the platform carbonate deposits of the Korallenoolith Formation (middle Oxfordian to early Kimmeridgian) small buildups ofcorals formed in the Lower Saxony Basin. These bioconstructions are restricted to particular horizons (Untere Korallenbank,ßorigenuna-Bank Member etc.) and represent patch reefs and biostromes. In this study, the development of facies, fossil assemblages, spatial distribution of fossils, and reefs of the ßorigenuna-Bank Member (upper Middle Oxfordian) in the Süntel Mts and the eastern Wesergebirge Mts is described; the formation of reefs is discussed in detail. Twelve facies types are described and interpreted. They vary between high-energy deposits as well winnowed oolites and quiet-water lagoonal mudstones. Owing to the significance of biota, micro- and macrofossils are systematically described. The reefs are preserved in growth position, are characterized by numerous corresponding features and belong to a certain reef type. According to their size, shape and framework, they represent patch reefs, coral knobs (sensu James, 1983), coral thrombolite reefs (sensu Leinfelder et al., 1994) or “Klein- and Mitteldickichte” (sensu Laternser, 2001). Their growth fabric corresponds to the superstratal (dense) pillarstone (sensu Insalaco, 1998). As the top of the ßorigenuna-Bank displays an erosional unconformity (so-called Hauptdiskontinuität), the top of the reefs are erosionally capped. Their maximum height amounts to at least the maximum thickness of the ßorigenuna-Bank which does not exceed 4 metres. The diversity of coral fauna of the reefs is relatively low; a total of 13 species is recorded. The coral community is over- whelmingly dominated by the thin-branched ramose Thamnasteria dendroidea (Lamouroux) that forms aggregations of colonies (77?. dendroidea thickets). Leafy to platy Fungiastrea arachnoides (Parkinson) and Thamnasteria concinna (Goldfuss) occur subordinately, other species are only of minor importance. In a few cases, the reef-core consisting of Th. dendroidea thickets is laterally encrusted by platy F. arachnoides and Th. concinna colonies, and microbial carbonates. This zonation reflects probably a succession of different reef builders as a result of changing environmental conditions (allogenic succession). Moreover, some reefs are overlain by a biostrome made of large Solenopora jurassica nodules passing laterally in a nerinean bed. Mikrobial carbonates promoted reef growth and favoured the preservation of reef organismn in their growth position or in situ. They exhibit a platy, dendroid, or reticulate growth form or occur as downward-facing hemispheroids. According to their microstructure, they consist of a peloidal, clotted, or unstructured fabric (predominately layered and poorly structured thrombolite as well as clotted leiolite) (sensu Schmid, 1996). Abundant endo- and epibiontic organisms (bivalves, gastropods, echinoids, asteroids, ophiuroids, crabs etc) are linked to the reefs. With regard to their guild structure, the reefs represent occurrences at which only a few coral species serve as builder. Moreover, microbial carbonates contribute to both building and binding of the reefs. Additional binder as well as baffler are present, but not abundant. According to the species diversity, the dweller guild comprises by far the highest number of invertebrate taxa. The destroyer guild chiefly encompasses bivalves. The composition of the reef community was influenced by the habitat structure of the Th. dendroidea thickets. Owing to the increase in encrusting organisms and other inhabitants of the thickets, the locational factors changed, since light intensity and hydrodynamic energy level and combined parameters as oxygen supply declined in the crowded habitat. Therefore a characteristic succession of organisms is developed that depends on and responds to changing environmental conditions („community replacement sequence“). The succession allows the differentiation of different stages. It started after the cessation of the polyps with boring organisms and photoautotrophic micro-encrusters (calcareous algae, Lithocodium aggregatum). Following the death of these pioneer organisms, encrusting and adherent organisms (serpulids, „Terebella“ species, bryozoans, foraminifers, thecideidinids, sklerospongid and pharetronid sponges, terebratulids), small mobile organisms (limpets), and microbial induced carbonates developed. The final stage in the community replacement sequence gave rise to small cryptic habitats and organisms that belong to these caves (cryptobionts, coelobites). The habitat conditions especially favoured small non-rigid demosponges (“soft sponges”) that tolerate reduced water circulation. Reef rubble is negligible, so that the reefs are bordered by fossiliferous micritic limestone passing laterally in micritic limestone. Approximately 10% of the study area (outcropping florigemma-Bank) corresponds to reefal deposits whereas the remaining 90% encompass lagoonal inter-reefal deposits. The reef development is a good example for the interaction between reef growth, facies development and sea-level changes. It was initiated by a sea-level rise (transgression) and corresponding decrease in the hydrodynamic energy level. Colonization and reef growth took place on a coarse-grained Substrate composed of oncoids, larger foraminifers and bioclasts. Reef growth took place in a calm marine lagoonal setting. Increasing abundance of spherical coral morphs towards the Northeast (section Kessiehausen, northwestem Süntel Mts) reflects higher turbidity and a facies transition to coral occurrences of the ßorigenuna-Bank Member in the adjacent Deister Mts. The reef growth was neither influenced by stonns nor by input of siliciclastic deposits, and took place in short time - probably in only a thousand years under most probably mesotrophic conditions. The mass appearance of solenoporids and nerineids in the upper part of the ßorigenuna-Bank Member point to enhanced nutrient level as a result of regression. In addition, this scenario of fluctuations in nutrient availability seems to be responsible for the cessation of reef corals. The sea level fall reached its climax in the subaerial exposure and palaeokarst development of the florigemma-Bank. The reef building corals are typical pioneer species. The blade-like, flattened F. amchnoides colonies are characterized by their light porous calcium carbonate skeleton, which is a distinct advantage in soft bottom environment. Thus, they settled on soft bottom exposing the large parts of its surface to the incoming light. On the other hand, in response to their light requirements they were also able to settle shaded canopy structures or reef caves. Th. dendroidea is an opportunistic coral species in very shallow, well illuminated marine environment. Their thin and densely spaced branches led to a very high surface/volume ratio of the colonies that were capable to exploit incoming light due to their small thamasterioid calices characterized by “highly integrated polyps”. In addition, sideward coalescence of branches during colony growth led to a wave-resistant framework and favoured the authochthonous preservation of the reefs. Asexual reproduction by fragmented colonies promoted reef development as Th. dendroidea thickets laterally extend over the sea floor or new reefs have developed from broken fragments of parent colonies. Similar build ups with Th. dendroidea as a dominant or frequent reef building coral species are known from the Paris Basin and elsewhere from the Lower Saxony Basin (Kleiner Deister Mts). These buildups developed in well-illuminated shallow water and encompass coral reefs or coral thrombolite reefs. Intra- and inter-reef deposits vary between well-winnowed reef debris limestone and mudstones representing considerably calmer conditions. Solenoporid, nerineids and diceratides belong to the characteristic fossils of these occurrences. However, diceratides are missing in theflorigemma-Bank Member. Th. dendroidea differs in its colonization of low- to high-energy environment from recent ramose scleractinian corals (e.g., Acropora and Porites sp.). The latter are restricted to agitated water habitats creating coral thickets and carpets. According to the morphologic plasticity of Th. dendroidea, thick-branched colonies developed in a milieu of high water energy, whereas fragile, wide- and thin-branched colonies prevail in low-energy settings. Due to its relatively rapid growth, Th. dendroidea was able to keep pace with increased Sedimentation rates. 68 benthonic foraminiferan species/taxa have been recognized in thin sections. Agglutinated foraminifers (textulariids) predominate when compared with rotaliids and milioliids. Numerous species are restricted to a certain facies type or occur in higher population densities, in particular Everticyclammina sp., a larger agglutinated foraminifer that occurs in rock building amounts. Among the 25 reef dwelling foraminiferal species, a few were so far only known from Late Jurassic sponge reefs. Another striking feature is the frequency of adherent foraminiferal species. Fauna and flora, in particular dasycladaleans and agglutinated foraminifers, document palaeobiogeographic relationships to the Tethys and point to (sub)tropical conditions. Moreover, in Germany this foraminiferan assemblage is yet uncompared. In Southern Germany similar tethyan type assemblages are not present in strata as young as Middle Tithonian.

Epilithic community development on artificial reefs deployed along a cross-shelf environmental gradient off Paraná state, southern Brazil

Concrete modules were deployed on the bottom of the 11, 18 and 30 meters isobaths along a cross-shelf hydrographic gradient off Paraná State, Southern Brazil, with the purpose of studying the colonization of sessile epilithic macroinvertebrates on artificial surfaces. After one year of submersion a total of 63 species of epilithic organisms were identified, dominated by Ostrea puelchana, Chthamalus bisinuatus, Balanus cf spongicola, Astrangia cf rathbuni, Didemnum spp, poryphers and bryozoans. Diversity index and percent cover at reef stations placed at 11, 18 and 30 meters isobaths were respectively 2.28 and 66.7%, 2.79 and 96.6% and 1.66 and 77.4%. Differences of general community structure among the three assemblages were not clearly related to the general environmental conditions at the bottom layers near the reef stations. Turbidity and larval abundance are discussed as important factors affecting colonization processes. Results indicate that depths between 15-20 meters are more suitable for the implementation of large scale artificial reef systems in the inner shelf off Paraná and, possibly, throughout the inner shelves off southern Brazil with similar hydrographic conditions.

Analysis of a Reconfigurable Bandpass Circular Patch Filter

This paper presents an analysis of a reconfigurable patch filter based on a triple-mode circular patch resonator with four radial slots. The analysis has been carried out thanks to the development of a new theoretical approach of the tunable patch filter based on the coupling matrix. The coefficients of the coupling matrix related to the tunable behavior have been identified and some rules for their evolution have been derived. For a proof-of-concept, a bandpass filter has been designed with a continuous tunability obtained with varactors connected across the slots. State-of-the-art results have been obtained, with a frequency tuning range of 27% from 1.95 to 2.43 GHz and a change in fractional bandwidth from 8.5% to 31.5% for the respective frequencies. In the entire tuning range, the return loss is better than 10 dB and the maximum insertion loss is 2 dB. Due to the newly developed coupling matrix, measurements, simulations, and theory showed great agreement.

A TRIPLE-MODE BANDPASS FILTER USING A MODIFIED CIRCULAR PATCH RESONATOR

This article presents a triple-mode bandpass filter using a modified circular patch resonator. Etched slots in the resonator split the TM(1, 1, 0)(z) degenerate fundamental modes and also perturb the TM(2, 1, 0)(z) mode, approximating their resonant frequencies to form a third-order bandpass filter. A 2.42 GHz centered filter was designed and fabricated. Experimental results showed a fractional bandwidth of 29%, return loss better than 16 dB, insertion loss of 0.5 dB, and good second harmonic band rejection. The filter exhibited a size reduction of 51% compared with a filter using an unperturbed circular patch resonator at the same frequency. (C) 2008 Wiley Periodicals, Inc. Microwave Opt Technol Lett 51: 178-182, 2009; Published online in Wiley InterScience (www.interscience.wiley.com). DOI 10.1002/mop.23950

Climate change, coral bleaching and the future of the world's coral reefs

Sea temperatures in many tropical regions have increased by almost 1 degrees C over the past 100 years, and are currently increasing at similar to 1-2 degrees C per century. Coral bleaching occurs when the thermal tolerance of corals and their photosynthetic symbionts (zooxanthellae) is exceeded. Mass coral bleaching has occurred in association with episodes of elevated sea temperatures over the past 20 years and involves the loss of the zooxanthellae following chronic photoinhibition. Mass bleaching has resulted in significant losses of live coral in many parts of the world. This paper considers the biochemical, physiological and ecological perspectives of coral bleaching. It also uses the outputs of four runs from three models of global climate change which simulate changes in sea temperature and hence how the frequency and intensity of bleaching events will change over the next 100 years. The results suggest that the thermal tolerances of reef-building corals are likely to be exceeded every year within the next few decades. Events as severe as the 1998 event, the worst on record, are likely to become commonplace within 20 years. Most information suggests that the capacity for acclimation by corals has already been exceeded, and that adaptation will be too slow to avert a decline in the quality of the world's reefs. The rapidity of the changes that are predicted indicates a major problem for tropical marine ecosystems and suggests that unrestrained warming cannot occur without the loss and degradation of coral reefs on a global scale.

Community structure of Quaternary coral reefs compared with Recent life and death assemblages

This paper assesses the reliability with which fossil reefs record the diversity and community structure of adjacent Recent reefs. The diversity and taxonomic composition of Holocene raised fossil reefs was compared with those of modern reef coral life and death assemblages in adjacent moderate and low-energy shallow reef habitats Of Madang Lagoon, Papua New Guinea. Species richness per sample area and Shannon-Wiener diversity (H') were highest in the fossil reefs, intermediate in the life assemblages, and lowest in the death assemblages. The taxonomic composition of the fossil reefs was most similar to the combination of the life and death assemblages from the modern reefs adjacent to the two fossil reefs. Depth zonation was recorded accurately in the fossil reefs. The Madang fossil reefs represent time-averaged composites of the combined life and death assemblages as they existed at the time the reef was uplifted. Because fossil reefs include overlapping cohorts from the life and death assemblages, lagoonal facies of fossil reefs are dominated by the dominant sediment-producing taxa, which are not necessarily the most abundant in the life assemblage. Rare or slow-growing taxa accumulate more slowly than the encasing sediments and are underrepresented in fossil reef lagoons. Time-averaging dilutes the contribution of rare taxa, rather than concentrating their contribution. Consequently, fidelity indices developed for mollusks in sediments yield low values in coral reef death and fossil assemblages. Branching corals dominate lagoonal facies of fossil reefs because they are abundant, they grow and produce sediment rapidly, and most of the sediment they produce is not exported. Fossil reefs distinguished kilometer-scale variations in community structure more clearly than did the modern life assemblages. This difference implies that fossil,reefs may provide a better long-term record of community structure than modern reefs. This difference also suggests that modern kilometer-scale variation in coral reef community structure may have been reduced by anthropogenic degradation, even in the relatively unimpacted reefs of Madang Lagoon. Holocene and Pleistocene fossil reefs provide a time-integrated historical record of community composition and may be used as long-term benchmarks for comparison with modern, degraded, nearshore reefs. Comparisons between fossil reefs and degraded modern reefs display gross changes in community structure more effectively than they demonstrate local extinction of rare taxa.

Patch dynamics and metapopulation theory: the case of successional species

We present a mathematical framework that combines extinction-colonization dynamics with the dynamics of patch succession. We draw an analogy between the epidemiological categorization of individuals (infected, susceptible, latent and resistant) and the patch structure of a spatially heterogeneous landscape (occupied-suitable, empty-suitable, occupied-unsuitable and empty-unsuitable). This approach allows one to consider life-history attributes that influence persistence in patchy environments (e.g., longevity, colonization ability) in concert with extrinsic processes (e.g., disturbances, succession) that lead to spatial heterogeneity in patch suitability. It also allows the incorporation of seed banks and other dormant life forms, thus broadening patch occupancy dynamics to include sink habitats. We use the model to investigate how equilibrium patch occupancy is influenced by four critical parameters: colonization rate? extinction rate, disturbance frequency and the rate of habitat succession. This analysis leads to general predictions about how the temporal scaling of patch succession and extinction-colonization dynamics influences long-term persistence. We apply the model to herbaceous, early-successional species that inhabit open patches created by periodic disturbances. We predict the minimum disturbance frequency required far viable management of such species in the Florida scrub ecosystem. (C) 2001 Academic Press.

Guadalupian algae-sponge reefs in siliciclastic environments - the reefs at Lengwu (South China) compared with the reef at Iwaizaki (Japan)

Guadalupian reefs occur locally in Guangxi, Guizhou, Yunnan and Western Zhejiang, South China. Two types of Guadalupian reefs can be recognized, one is developed in carbonate platforms, e.g. those in the juncture areas of Guangxi, Yunnan and Guizhou; the other occurs in a littoral clastic shelf. The Lengwu reef in Western Zhejiang is a representative of the latter type, which is a major topic of this paper. Lengwu algae-sponge reef, more than one hundred meters in thickness, are composed mainly of sponges, hydrozoans, algae, bryozoans, microbes and lime mud. Reef limestones sit on the mudstone interbedded with fine sandstone of the proximal prodelta facies and are overlain by coarse clasts of the delta front sediments. Lengwu reef displays a lens-shaped relief, dipping and thinning from the reef core, which is remarkably different from the surrounding sediments, showing a protruding relief. Sponges and microbe/algae form bafflestone, bindstone and framestone of the reef core facies. Fore-reef facies is characterized by lithoclastic rudstone and bioclastic packstone. Reef limestone sequence is composed of three cycles and controlled by sea level changes and sediment influx. Such reef is unique among the Guadalupian reefs in South China, but seems similar in some aspects to lwaizaki reef limestones of south Kitakami in Japan. Algae and microbes growing around sponges to form rigid structure in Lengwu reef are a typical feature, which is distinctly different to Guadalupian reefs in a stable platform facies of Guizhou, Yunnan and Guangxi, South China.

Optimal patch-leaving behaviour: a case study using the parasitoid Cotesia rubecular

1. Parasitoids are predicted to spend longer in patches with more hosts, but previous work on Cotesia rubecula (Marshall) has not upheld this prediction, Tests of theoretical predictions may be affected by the definition of patch leaving behaviour, which is often ambiguous. 2. In this study whole plants were considered as patches and assumed that wasps move within patches by means of walking or flying. Within-patch and between-patch flights were distinguished based on flight distance. The quality of this classification was tested statistically by examination of log-survivor curves of flight times. 3. Wasps remained longer in patches with higher host densities, which is consistent with predictions of the marginal value theorem (Charnov 1976). tinder the assumption that each flight indicates a patch departure, there is no relationship between host density and leaving tendency. 4. Oviposition influences the patch leaving behaviour of wasps in a count down fashion (Driessen et al. 1995), as predicted by an optimal foraging model (Tenhumberg, Keller & Possingham 2001). 5. Wasps spend significantly longer in the first patch encountered following release, resulting in an increased rate of superparasitism.