Indications for microbial clues for bryozoans when settling

Summary A correlation between skeletal growth of recent bryozoans and external environmental factors (e.g. abrasion, suspended sediment load) exists in ahermatypic tropical shallow water associations of the Philippines. Yet, the spatial zonation pattern of bryozoan species in a modern Philippine reef exhibits an amazingly high level of stability and order which cannot be readily correlated with external environmental factors. This is due to the observation that zonation correlates with the bryozoan taxa but not with their growth form categories (e.g. flexible-erect). Bryozoans are closely linked with their reef (micro)environment and reef microbiota by feedback control: Interacting biofilms, microbial mats, bryozoans and bryozoan larvae illustrate how such internal functions in a Philippine reef may stabilize the biotope structure as a whole     

A modern-type Kimmeridgian reef (Ota Limestone,Portugal): Implications for Jurassic reef models

Upper Jurassic reefs rich in microbial crusts generally appear in deeper (sponge—‘algal’ crust reefs) or in very shallow but protected settings (coral or coral-coralline sponge meadows with ‘algal’ crusts). Upper Jurassic high-energy reefs (coral reefs and coral-stromatoporoid reefs) normally lack major participation of microbial crusts but rather represent huge bioclastic piles with only minor framestone patches preserved. An exception to this rule is represented by the high-energy, coral-‘algal’ Ota Reef from the Kimmeridgian of the Lusitanian Basin (Portugal). The narrow Ota Reef tract rims a small intra-basinal carbonate platform exhibiting perfect facies zonation (from W to E: Reef tract, back reef sands, peritidal belt, low-energy shallow lagoon). The reef is dominated by massive corals (Thamnasteria, Microsolena, Stylina). Complete preservation of coral framework is rare: like other Upper Jurassic high-energy reefs, the Ota Reef is very rich in debris; however, this debris is largely stabilized by algal and microbial crusts, what contrasts the other examples and gives the Ota Reef the appearance of a typical modern high-energy coral-melobesioid algal reef. Further similarities to modern reefs are the likely existence of a spur-and-groove system, the perfect sheltering of inner platform areas and the occurrence of small islands, as indicated by local blackenings and early vadose and karstic features.     

Anisian (middle triassic) buildups of the Northern Dolomites (Italy): The recovery of reef communities after the permian/triassic crisis

Summary After the end-Permian crisis and a global ‘reef gap’ in the early Triassic, reefs appeared again during the early Middle Triassic. Records of Anisian reefs are rare in the Tethys as well as in non-Tethyan regions. Most Anisian reefs are known from the western part of the Tethys but there are only very few studies focused on biota, facies types and the paleogeographical situation of these reefs. From the eastern part of the Tethys, Anisian reefs, reefal buildups or potential reef-building organisms have been reported from different regions of southern China. Most of the Anisian reefs known from western and central Europe as well as from southern China seem to be of middle and late Pelsonian age. The study area is situated in the northern Dolomites (South Tyrol, Italy) southeast of Bruneck (Brunico). It comprises the area between Olang (Valdaora) and Prags (Braies). The study is based on detailed investigations of the regional geology, stratigraphy and lithofacies (R. Zühlke, T. Bechst?dt) as well as on a comprehensive inventory of Anisian reef organisms (B. Senowbari-Daryan, E. Flügel). These data are used in the discussion of the controls on the recovery of reefs during the early Middle Triassic. Most late Anisian reef carbonates studied are represented by allochthonous talus reef blocks of cubicmeter size. Small biostromal autochthonous mounds are extremely rare (Piz da Peres). The reef mounds as well as most of the reef blocks occur within the middle to late Pelsonian Recoaro Formation. They were formed on the middle reaches of carbonate ramps in subtidal depths, slightly above the storm wave base with only moderate water energy. Most lithotypes observed in the reef blocks correspond to sponge and/or algal bafflestones. Low-growing sessile organisms (Olangocoelia (sponge, alga?), sphinctozoan sponges, bryozoans, soleno-poracean algae, corals) and encrusting epibionts (sponges, porostromate algae, cyanophycean crusts, foraminifera, worms, microproblematica) created low cm-sized biogenic structures (bioconstructions) which baffled and bound sediment. Organic framework was only of minor importance; it is restricted to theOlangocoelia lithotype. Framework porosity was small in these reef mounds. Submarine carbonate cements, therefore, are only of minor importance s compared with Permian or Ladinian reefs. The relatively high number of lithotypes encountered in the reef blocks indicates a high biofacies diversity. Regarding the relative frequency, the diverse biota consist in descending order ofOlangocoelia, sponges (sphinctozoans, inozoans, siliceous sponges), bryozoans, porostromate algae and worm tubes. The sphinctozoans are characterized by small, mostly incrusting forms. The numerical diversity (species richness) is low compared with late Permian or Ladinian and late Triassic sphinctozoan faunas occurring within reefs. Following the sponges, monospecific bryozoans (Reptonoditrypa cautica Sch?fer & Fois) are the most common organisms in the reef limestones. Porostromate algae were restricted to areas within the bioconstructions not inhabited by sponges. The low-diverse corals had no importance in the construction of an organic framework. Surprisingly, microbial crusts are rare or even lacking in the investigated Anisian bioconstructions. This is in contrast to late Permian and Ladinian as well as Carnian reefs which are characterized by the abundance of specific organic crusts. The same comes true for‘Tubiphytes’ which is a common constituent in Permian, Ladinian and Carnian reef carbonates but is very rare in the Anisian of the Olang Dolomites. Instead of‘Tubiphytes’ different kinds of worm tubes (spirorbid tubes, Mg-calcitic tubes and agglutinated tubes) were of importance as epifaunal elements. Macrobial encrustations consisting of characteristic successions of sponges, bryozoans, algae, worm tubes and microproblematica seem to be of greater quantitative importance than in Ladinian reefs. Destruction of organic skeletons (predominantly of bryozoans) by macroborers (cirripedia?) is a common feature. The Anisian reef organisms are distinctly different from late Permian and from most Ladinian reef-builders. No Permian Lazarus taxa have been found. New taxa: Sphinctozoan sponges—Celyphia? minima n.sp.,Thaumastocoelia dolomitica n. sp.,Deningeria tenuireticulata n. sp.,Deningeria crassireticulata n. sp.,Anisothalamia minima n.g. n.sp., Inozoan sponges-Meandrostia triassica n.sp. Microproblematica-Anisocellula fecunda n.g. n.sp., Porostromate alga-Brandneria dolomitica n.g. n.sp. Most of our data are in agreement with the model described byFois & Gaetani (1984) for the recovery of reef-building communities during the Ansian but the biotic diversity seems to be considerably higher than previously assumed. Anisian deposition and the formation of the reef mounds within the Pelsonian Recoaro Formation of the Dolomites were controlled by the combined effects of synsedimentary tectonics and eustatic changes in sea-level. During several time intervals, especially the early Anisian (northern and western Dolomites: tectonic uplift), the early Pelsonian (eastern Dolomites: drowning) and the late Illyrian (wide parts of the Dolomites: uplift and drowning), the sedimentation was predominantly controlled by regionally different tectonic subsidence rates. The amount of terrigenous clastic input associated with synsedimentary tectonics (tectonic uplift of hinterlands) had a major influence on carbonate deposition and reef development. The re-appearance of reef environments in the Olang Dolomites was controlled by a combination of regional and global factors (paleogeographic situation: development of carbonate ramps; decreasing subsidence of horst blocks; reduced terrigenous input; moderate rise in sea-level).     

Assessing the ‘deep reef refugia’ hypothesis: focus on Caribbean reefs

Coral reefs in shallow-water environments (<30 m) are in decline due to local and global anthropogenic stresses. This has led to renewed interest in the ‘deep reef refugia’ hypothesis (DRRH), which stipulates that deep reef areas (1) are protected or dampened from disturbances that affect shallow reef areas and (2) can provide a viable reproductive source for shallow reef areas following disturbance. Using the Caribbean as an example, the assumptions of this hypothesis were explored by reviewing the literature for scleractinian corals—the reef framework builders on tropical reefs. Although there is evidence to support that deep reefs (>30 m) can escape the direct effects of storm-induced waves and thermal bleaching events, deep reefs are certainly not immune to disturbance. Additionally, the potential of deep reefs to provide propagules for shallow reef areas seems limited to ‘depth-generalist’ coral species, which constitute only ~25% of the total coral biodiversity. Larval connectivity between shallow and deep populations of these species may be further limited due to specific life history traits (e.g., brooding reproductive strategy and vertical symbiont acquisition mode). This review exposes how little is known about deep reefs and coral reproduction over depth. Hence, a series of urgent research priorities are proposed to determine the extent to which deep reefs may act as a refuge in the face of global reef decline.     

Norian-Rhaetian reefs in Argolis Peninsula,Greece

Summary Upper Triassic to Lower Jurassic shallow-water carbonate sequences of the ‘Pantokrator limestones’ are widely distributed in the Argolis Peninsula, southern Greece. Within this sequence are some reef or reefal structures. In the Mavrovouni Mountains, near Sarmeika, 6 km SE of the ancient theatre of Epidavros (Argolis Peninsula), a Norian-Rhaetian reef complex has been identified. This is the first well-documented Norian-Rhaetian reef in Greece. The main reef builders are coralline sponges (‘sphinctozoans,’ ‘inozoans’, and sclerosponges), followed by dendroid, cerioid, and solitary corals, and algae. The reef type corresponds to a ‘sponge-coral reef’.     

Strengths abd weaknesses of the reef guild concepts and quantitative data: Application to the upper Capitan-massive community (Permian), Guadalupe Mountains, New Mexico-Texas

Summary Analyses of large acatate sheet tracings, close-up photos and 105 sub-horizontal quadrat surfaces at four localities near the base of the Guadalupe Mountains Escarpment indicate that the biotic framework of the upper Capitan reef was built by about 35 species: one codiacean (Eugonophyllum sp.), 17 calcisponges, 9 bryozoans, one richthofenid brachipod, some crinoid (known only from columns), 4 Problematica and microbes. This widespread fossil community included members of the Constructor, Baffler and Binder Guilds. A re-evaluation of the Guild Concept (Fagerstrom, 1987, 1991) highlights the validity of the functional roles of the Constructor and Binder Guilds for reef construction. Members of the Baffler Guild, however, need to be revised and an interpretation of microbial micrite and cryptic biota remains controversial. Open surface phylloid algal and cryptic sponge-bryozoan dominated sub-communities were of only local importance. The upper Capitanmassive differs from its Permian conterparts in the low diversity and areal cover of the frame-building biota, low micrite content and abundant micro-frameworks, i.e, intergrown small sponges, Problematica and syndepositional cements (botryoidal and isopachous, fibrous calcite). Quantitative areal cover data were assessed at various scales. Large acetate sheets generally have low coverage of macro-biota (5.4%). By contrast, analysis of small areas of local high areal cover (selected acetate sheet quadrats, subvertical photographs, and quadrat samples: 15–21%) provide detailed insights into clustered patches forming the inital reef framework. Both data sets provide useful clues for an integrated approach to framework assessment. Mean acetate sheet data are limited by their somewhat generalized pattern, while small investigation areas may overemphasize local variation. Erect and pendant sponges with solitary, sub-cylindrical and multi-branche/clonal forms, were the predominant initial frame-builders in both open surface and cryptic habitats. Selective larval recruitment of erects sponges to firm substrates produced continous upward accretion of the initial framework. On open surfaces and in pores formed by tabular sponges and fenestrate bryozoans, erect and pendant sponges were supported in their hydrodynamically unstable growth position by encrusters, chieflyArchaeolithoporelle hidensis, Shamovella obscura, an unnamed tubular organism, and microbes. Subsequent growth ofArchaeolithoporella hidensis, microbial crusts and syndepositional cements on the outer walls of live sponges would have impeded ambient water circulation and may have led to ‘creeping sponge death by suffocation’ or complete encrustation after death. Filling of pores in the initial and encrusted reef framework by internal sediment (packstone-grainstone; derived from the framework and the back-reef shelf/platform) and voluminous syndepositional marine-phreatic cements completed the framebuilding process.     

Biomineralization in bryozoans: present,past and future

Many animal phyla have the physiological ability to produce biomineralized skeletons with functional roles that have been shaped by natural selection for more than 500 million years. Among these are bryozoans, a moderately diverse phylum of aquatic invertebrates with a rich fossil record and importance today as bioconstructors in some shallow‐water marine habitats. Biomineralizational patterns and, especially, processes are poorly understood in bryozoans but are conventionally believed to be similar to those of the related lophotrochozoan phyla Brachiopoda and Mollusca. However, bryozoan skeletons are more intricate than those of these two phyla. Calcareous skeletons have been acquired independently in two bryozoan clades – Stenolaemata in the Ordovician and Cheilostomata in the Jurassic – providing an evolutionary replicate. This review aims to highlight the importance of biomineralization in bryozoans and focuses on their skeletal ultrastructures, mineralogy and chemistry, the roles of organic components, the evolutionary history of bimineralization in bryozoans with respect to changes in seawater chemistry, and the impact of contemporary global changes, especially ocean acidification, on bryozoan skeletons. Bryozoan skeletons are constructed from three different wall types (exterior, interior and compound) differing in the presence/absence and location of organic cuticular layers. Skeletal ultrastructures can be classified into wall‐parallel (i.e. laminated) and wall‐perpendicular (i.e. prismatic) fabrics, the latter apparently found in only one of the two biomineralizing clades (Cheilostomata), which is also the only clade to biomineralize aragonite. A plethora of ultrastructural fabrics can be recognized and most occur in combination with other fabrics to constitute a fabric suite. The proportion of aragonitic and bimineralic bryozoans, as well as the Mg content of bryozoan skeletons, show a latitudinal increase into the warmer waters of the tropics. Responses of bryozoan mineralogy and skeletal thickness to oscillations between calcite and aragonite seas through geological time are equivocal. Field and laboratory studies of living bryozoans have shown that predicted future changes in pH (ocean acidification) combined with global warming are likely to have detrimental effects on calcification, growth rate and production of polymorphic zooids for defence and reproduction, although some species exhibit reasonable levels of resilience. Some key questions about bryozoan biomineralization that need to be addressed are identified.     

Factors influencing initial larval settlement: temporal, spatial and surface molecular components

The impact of initial surface chemistry on settlement of barnacle, bryozoan, and hydroid larvae was assessed. Temporal (on a scale of weeks to months) and fine scale spatial (centimeter to meter) variation in settlement were quantified. Four arrays of silanized glass surfaces, deployed at ≈2-wk intervals, were monitored after 1 and 3 days of immersion. Settlement of all larval types exhibited strong temporal variation. There was a 25, 22- and 18-fold difference between the highest and lowest Day 1 settlement for barnacles, bryozoans and hydroids, respectively. Bryozoan and hydroid settlement was spatially variable, barnacle settlement was not. Barnacle and bryozoan settlement was influenced by inttial surface chemistry, hydroid settlement was not. For barnacles, there was a 2-fold difference between total settlement over the four arrays on untreated glass and diphenyl-silanized surfaces; for bryozoans, there was a 51-fold difference, and in the opposite direction. There was a negative correlation between barnacle and bryozoan settlement with respect to surface. This result was independent of the presence of the other species. Bryozoan settlement was also spatially and temporally quantified in a separate, single surface 56-h array, and total bryozoan settlement ·h⁻¹ was found to be correlated with light intensity. Patterns of initial colonization were strongly influenced by surface chemistry and spatial temporal variation in larval supply. These effects on initial colonization may influence subsequent community development.     

Microbial mats associated with bryozoans (Coorong Lagoon, South Australia)

Summary Bryostromatolites are laminated carbonate rocks composed of bryozoan zoarial laminae. The laminated texture is frequently caused by patterns of bryozoan self overgrowth as a regular defensive tactic against microbial fouling. In the Coorong Lagoon (South Australia), another type of bryostromatolite is present where the laminated growth of the weakly calcifying bryozoan speciesConopeum aciculata is postmortally stabilized by cyanobacterial mats at the surface, and fungal mats settling in the zooecial cavities. A tough extracellular slime network produced by benthic cyanobacteria is a trap for sediment particles, provides a method of adhesion to the bryozoan substrate, and produces a biological lamination by the vertical stratification of dead bryozoan skeletons. These slimes are also important for the preservation of cell structures and for their fossilization. Seasonal fluctuations in salinity and water level are the most important regional control factors, causing a phase displacement in the growth optima of microbial mats and bryozoans, thereby resulting in a rigid bryostromatolitic fabric.     

New estimates of global and regional coral reef areas

 Global and regional coral reef area statistics are of considerable value in fields ranging from global environmental change to fisheries to conservation. Although widely quoted, Smith’s 1978 figure of 600 000 km² is only an approximate calculation. The World Conservation Monitoring Centre has prepared a new estimate of reef coverage by mapping emergent reef crest and very shallow reef systems. These data were rasterised, using 1 km grid squares, as a means of reducing errors arising from variation in scale. Global and regional reef coverages were calculated from the resultant grid. The total global area is estimated at 255 000 km², considerably lower than many previous estimates. Variation in reef area estimates is, in part, a function of variation in reef definition. Accepted: 3 April 1997     

Upper Triassic reefs of the Oman Mountains: Data from the South Tethyan margin

Summary The Upper Triassic reefal limestones of the Oman Mountains were investigated with respect to their microfacies, palaeontology and community structure. The reef fauna described and figured for the first time occurs in parautochthonous slope deposits of the Arabian platform (Sumeini Group) and in allochthonous reefal blocks (‘Oman Exotics’, Hawasina Complex). The ‘Oman Exotics’ are tectonically dislocated blocks, derived from isolated carbonate platforms on seamounts in the Hawasina basin or in the South Tethys Sea. The lithofacies and fauna of these blocks comprise a cyclic platform facies with megalodonts, reef and reef debris facies. The reefal limestones are dated as Norian/Rhaetian by benthic foraminiferal associations (Costifera, Siculocosta, Galeanella) and typical encrusting organisms (Alpinophragmium, Microtubus). Some small ‘Oman Exotics’ are of Carnian age. The shallow-marine organisms include scleractinian corals of different growth forms, ‘sphinctozoans’, ‘inozoans’ chaetetids, spongiomorphids, disjectoporids and solenoporacean algae as the main reef builders, various encrusters like microbes, foraminifers, sponges and many different problematical organisms for the stabilisation of the reef framework and a group of dwellers including benthic foraminifers, gastropods, bivalves and a few dasycladacean algae. The reef communities are characterized by the coverage of organisms and distributional pattern. Analogies with the coeval reef deposits from the European part of the Tethys have been recognized. Some species, now collected in Oman, were also reported from American and Asian localities.     

Bryozoans and microbial communities of cool-temperate to subtropical latitudes—paleoecological implications

The environmental distribution of encrusting bryozoans settling on disarticulated and living bivalve shells has been recorded from five stations in Japan and New Zealand. Some insight into the observed distribution patterns emerges from information on the interaction of bryozoans with microbial mats. Advancing existing classifications, we have subdivided the encrusting bryozoan morphotypes into seven different growth types that largely reflect the biological potentials of bryozoans in competition for space on substrate surfaces. The frequency distribution of these types (s-/c-/m-/z-laminae, runners, spots, bryostromatolites) reveals the influence of microbial mats as a control factor of bryozoan substrate coverage. Microbial mats in turn are correlated with latitudinal gradients in Japan and New Zealand from cool-temperate to subtropical and tropical waters. Unlike erect bryozoans, laminar ones are probably underrated as facies fossils. Accordingly, laminar bryozoan growth types are reconsidered as a tool for paleoecological interpretation of marine hard substrate communities.     

Interaction between the invasive macroalga <Emphasis Type="Italic">Lophocladia lallemandii</Emphasis> and the bryozoan <Emphasis Type="Italic">Reteporella grimaldii</Emphasis> at seagrass meadows: density and physiological responses

Invasive epiphyte Lophocladia lallemandii macroalga induces changes in the erect bryozoan Reteporella grimaldii at shallow Posidonia oceanica meadows at a Mediterranean pristine location. Bryozoan densities at noninvaded seagrass plots (88.32 ± 3.11 colonies m⁻²) are higher than those at invaded plots (13.39 ± 1.09 colonies m⁻²) with a fourfold decrease in number of colonies. Activation of enzymatic pathways (catalase, superoxide dismutase, glutathione peroxidase) and increase in lipid peroxidation malondialdehyde (MDA) [0.80 ± 0.06 nmol/mg prot at Posidonia oceanica plots to 1.08 ± 0.04 nmol/mg prot at L. lallemandii (P < 0.05)] is observed on sessile bryozoans as response to anoxia caused by L. lallemandii. δ¹³C of bryozoan isotopic composition differed among treatments, covering a broad range (−19.30‰ invaded to −2.84‰ at noninvaded plots), suggesting modification of food sources. Induced shifts of a filter-feeding erect bryozoan by dense algal turfs at invaded seagrasses are demonstrated, highlighting the need to further address interaction across natural communities and alien species invaded systems before further cascade effects are driven.     

Taphonomy of larger foraminifera: Relationships between living individuals and empty tests on flat reef slopes (Sesoko Island, Japan)

Summary The depth distributions of larger foraminifera (27 species) were investigated along two transects in the fore reef areas of a NW Pacific fringing reef. One transect is distinguished by a strong flattening below the steep reef slope (−30 m), whereas further steepening characterizes the equivalent part in the other transect. According to the different taphonomic processes affecting foraminiferal tests before final sedimentation, empty tests were classified into the three categories ‘optimally’, ‘well’ and ‘poorly’ preserved. The depth distribution of each preservation state was compared with living individuals. While distributions of optimally preserved tests almost coincide with living individuals, well-preserved tests are characterized by significant depth shifts that are stronger at the upper-most slope compared with the deeper parts. Since the time-averaged traction forces are similar in both investigated transects, differences between the distributions of living individuals and well-preserved tests are more intensive on steep versus flat slopes. Poorly preserved tests signalize allochthonous origin or reworking of relict sediments.     

Response of triassic reef coral communities to sea-level fluctuations, storms and sedimentation: Evidence from a spectacular outcrop (Adnet, Austria)

Summary The Upper Rhaetian coral limestone of Adnet, southeast of Salzburg Austria has been repeatedly referred to as one of the most spectacular examples of an ancient ‘autochthonous’ coral reef structure. The ‘Tropfbruch’ quarry is probably the best outcrop for interpreting the distributional patterns of biotic successions and communities of a late Triassic patch reef. Our study is based on the interpretation of a) outcrop photographs, b) reef maps resulting from quadrat transects, and c) the analysis of quantitative data describing the distribution and frequency of reef organisms and sediment. A new methodological approach (combination of reef mapping and photo-transects) is used to obtain quantitative field data which can be compared in greater detail with data from modern coral reefs investigated by corresponding quantitative surveys. Three unconformities and three well-defined ‘reef growth stages’ reflecting the vertical and lateral development of the reef structure were differrentiated using transects: Stage 1, representing the reef growth optimum, is characterized by laterally differentiated coral reef knobs with corals in growth position. Criteria supporting this interpretation are the extraordinary size of the corals, their preservation in situ and the great thickness of this interval. The massive coralPamiroseris grew under higher energy conditions at the rim of the reef knob, whereas branchingRetiophyllia colonies preferred less agitated water in the center. Vertical changes are reflected by an increase in frequency of the dasycladacean algaDiplopora adnetensis and by the decreasing size ofRetiophyllia. These sedimentological and biological criteria together with the unconformity above indicate a fall in the sea level as a major control mechanism. Stage 2, separated from stage 1 by an unconformity caused by partial subaerial exposure and karstification, is characterized by vertically stacked coral successions with diverse reef debris. Facies heterogeneity is reflected by differences in the diversity, taphonomy and packing density of reef-building organisms as well as by differences in sediment input from the platform. Water depths and accommodation space were lower, therefore minor sea level fluctuations had a stronger effect on the biotic composition. The high percentage of coral debris and corals reworked by storms and the increase in the input of platform sediment led to a reduction of reef growth. Stage 3, again separated at the base by an unconformity associated with karstification, is characterized by bioclastic sediments with isolated reefbuilders forming a level-bottom community. The distribution of different coral morphotypes suggests that sea level fluctuations were not the only controlling factor. Variations in the substrate were caused by differences in the input of platform sediment. The three-step development seen in Adnet documents the response of low-diverse coral associations to variations caused by small-scale sea level changes, storm activity and sedimentation. The vertical changes in reef community structures correspond to a sequence of ‘allogenic replacements’. The Adnet reef structure should not be regarded as a general model of Alpine Upper Rhaetian reefs, because of the particular setting of the patch reef. Only the ‘capping beds’ of the Upper Rhaetian Reef Limestone of the Steinplatte exhibit criteria similar to Adnet. Potential modern analogues of features seen in the coral communities of Adnet are the internal structure of theRetiophyllia thickets, the key role of branching corals within the communities, the scattered distribution and low and even diversity of corals subsequent to breaks in settlement, segration patterns of corals indicating ‘contact avoidance’, toppling of large coral colonies by intensive boring, and decreasing coral coverage from deeper and sheltered settings to more shallower water depths.     

Bryozoaires et Brachiopodes des “Calcaires bajociens a Bryozoaires” des Causses (France) et leur paléoécologie

Two Lower Bajocian new localities, with many silicified bryozoans and brachiopods have been discovered in Causses (France): Saint-Rome-de-Cernon (Aveyron) and Mende (Lozère). The study of the faunas allows to increase the anterior knowledges supplied by the study of localities in the South of Causses (neighbourhood of Alzon and Sumène = North of the Seuil sud-caussenard).The interest is, at first, the distance between the localities: Mende in the North, Saint-Rome-de-Cernon in the West. Moreover, in the second, well-preserved faunas were fossilized without transport. Bryozoans and brachiopods are very numerous. Large and complete bryozoan colonies have been get out by action of HCl. Fine internal structures like rhynchonellid Nannirhynchia or like thecidean transversarium and hemispondylium have been studied. At last, observations and paleoecological hypothesis are drawn, likewise comparison between three localities or locality-groups known in the Causses basin.The fauna from Saint-Rome-de-Cernon is biocoenosis, the substratum was soft, the hydrodynamism low and the depth, according to lamellibranchs, about 200 meters. The beds with bryozoans and small brachiopods contain a fauna with two predominant bryozoan species: Ceriocava straminea and Mesenteripora wrighti. These two species could built large colonies, in spite of fine granulometry of the sedimentary bottom. These colonies provided supports to less adapted bryozoans, to thecidids and to lamellibranch Lopha. The beds with terebratulids and bryozoans contain lesser bryozoans and thecidids. Terebratulids are the main group and are fixed with their peduncle to organic support above the irregular deposits. Beside these benthic animals, the rhynchonellid Nannirhynchia campestriensis is epibiontic on gorgonids or epiphytic on floating sea-weeds.On the contrary, all fossils from Mende have been more or less carried, excepted large sponges perhaps. The most numerous bryozoans (Ceriocava straminea, Mesenteripora wrighti) and numerous Lenticulines (Foraminifera) lived on hard bottom with moving water in community with crinoids. It is possible that a smaller part of bryozoans, requiring lower hydrodynamism, lived, either in sheltered parts of the crinoid community or closer to the sedimentation zone. Thecidids were fixed either on bryozoans in crinoid communities or on shelly remains and on bryozoans nearer to the sedimentation zone. Like in Saint-Rome-de-Cernon, Nannirhynchia campestriensis was epibiontic, or epiphytic, or both.In comparison with the whole localities from the Seuil sud-caussenard, we notice the simplicity of Saint-Rome-de-Cernon and Mende. The localities from the Seuil sud-caussenard were assemblages proceeding from 2 or 3 biotops. On the contrary, at Saint-Rome-de-Cernon we have a biocoenosis and at Mende a thanatocoenosis supplied by one biotop almost.     

What would Darwin have written now?

We often wonder how many of the pristine places left on Earth we can protect from deterioration before it is too late. The assumption that remote regions remain pristine plays a key role in directing policies for regional environmental management and conservation, and affects the local and global financial impetus to do so. In this paper, we use Argentinean Patagonia and the SW Atlantic as examples to argue that the assumption ‘remote region = pristine region’ is unjustified and based on a lack of information rather than on scientific evidence. We also discuss the major existing environmental threats to this supposedly ‘pristine’ region, and use emblematic examples to provide a more realistic picture of the regional environmental integrity and to set recommendations directed to improve environmental management and conservation within this context.     

Beta diversity of a Gulf of California rocky-shore fish community

Synopsis Reef fish community composition in three segments of a peninsular rocky shoreline in the Gulf of California was estimated over four periods by visual observation. ‘Point’ and ‘bay’ segments had regular and distinct species compositions over most periods while a ‘middle’ segment was least distinct but consistently had the greatest number of species. Compositional change along the peninsula was least regular during the coldest sea temperature period (April). Mean species turnover between segments was highest between point and bay. Within segments, the point had greater compositional predictable composition (lowest species turnover). When species with regular frequency of encounter were classified into ‘point’, ‘middle’, ‘bay’, and ‘no trend’ groups it was found that comparatively more ‘point’ species had pelagic eggs and comparatively more ‘bay’ species had demersal eggs. Beta diversity of rocky-shore fishes along the physical gradient of the Punta Doble peninsula reflects a transition between exposed and protected rocky shoreline communities. The correlated physical environmental characteristics associated with exposed and protected habitats are discussed in relation to diversity regulation and dispersal strategies in reef fishes.     

The adaptive significance of larval dispersal in coral reef fishes

Synopsis Coral reef fishes almost universally disperse over relatively great distances during a pelagic larval phase. Barlow (1981) suggested that this dispersal is adaptive because adult fishes inhabit a patchy, uncertain environment. This reiterated an older idea that the random extinction of local populations necessarily favours dispersal, since ultimately all populations of non-dispersers will disappear. Whereas this view is based on adult survival, we emphasize a less frequent view that substantial larval dispersal may be adaptive when offspring experience patchy and unpredictable survival in the pelagic habitat. We do not address the question of why these animals ‘broadcast’ rather than ‘brood’, but suggest that species committed to pelagic offspring will be under selection to disperse siblings to spread the risk of failure among members of a cohort. Our arguments are supported by a heuristic computer simulation.     

Mineralogy of Arctic bryozoan skeletons in a global context

Bryozoans are major carbonate producers in some ancient and Recent benthic environments, including parts of the Arctic Ocean. Seventy-six species of bryozoans from within the Arctic Circle have been studied using XRD to determine their carbonate mineralogies and the Mg content of the calcite. The majority of species were found to be calcitic, only four having bimineralic skeletons that combined calcite and aragonite, and none being entirely aragonitic. In almost all species, the calcite was of the low- (<4 mol% MgCO₃) or intermediate-Mg (4–11.99 mol% MgCO₃) varieties. Previous regional studies of bryozoan biomineralogy have found higher proportions of bimineralic and/or aragonitic species in New Zealand and the Mediterranean, with a greater number of calcitic species employing intermediate- and high-Mg calcite. The Antarctic bryozoan fauna, however, has a similar mineralogical composition to the Arctic. The lesser solubility of low-Mg calcite compared to both Mg calcite and aragonite in cold polar waters is most likely responsible for this latitudinal pattern. However, it is unknown to what extent environmental factors drive the pattern directly through eliciting an ecophenotypic response from the bryozoans concerned or the pattern reflects genetic adaptations by particular bryozoan clades.