Stromatoporoid-coral intergrowths in a Silurian biostrome

Four stromatoporoid species from a stromatoporoid biostrome in the middle Ludlow Hemse Beds, Gotland, Sweden, show intergrowths with syringoporid tabulate and rugose corals, and indicate close relationships between particular coral and stromatoporoid species. The stromatoporoid Clathrodictyon convictum always contains ?Syringopora and this tabulate is rarely found in the other stromatoporoids. C. convictum is also closely associated with Tryplasma flexuosum (rugosa) while Petrozium pelagicum (rugosa) occurs only in the stromatoporoids Plectostroma intermedium and Parallelostroma typicum. The microstructure of ?Syringopora within the stromatoporoids is composed of an inner lamellar layer and an outer radial layer of calcite crystals. Diagenetic alteration has affected the microstructure which differs from recently described Devonian forms having only a radial layer. This shows variability in the structure of the tabulates within stromatoporoids. Information is sparse on the range of such variation and assessment of the relative importance of taxonomic, palaeoenvironmental and diagenetic effects is not possible in the present sample. No evidence is found to prove the precise nature of the relationships; they were not parasitic but may have been mutually symbiotic, or (most probably) commensal. The results suggest that the corals selected the most suitable stromatoporoid species for their requirements. Stromatoporoid morphology may have had an important influence on the association, where corals are more abundantly associated with those stromatoporoid species which adopted a high profile. Overall the associations appear to have allowed the corals to explore higher energy habitats otherwise unavailable to their delicate branching structure.     

Diagenesis of growth bands in fossil scleractinian corals: identification and modes of preservation

Light and dark bands in fully recrystallized fossil hermatypic corals are generally interpreted to represent annual growth increments reflecting a photosymbiotic life style—an interpretation of far reaching significance in palaeoecology. In this paper we describe annual growth bands in the colonial coral Porites in a perfect (aragonite and microstructures retained) and fully recrystallized (sparry calcite mosaic) style of preservation from sediments of Late Miocene age (Crete, Greece). Analysis of a continuous spectrum of transitional preservational stages shows that in Miocene Porites preservation of the growth banding was controlled by preferential dissolution of the high-density band associated with cementation by drusy calcite spar during freshwater diagenesis/shallow burial diagenesis. Marine precipitates (pelletoidal Mg-calcite) preferentially accumulated along tabulate dissepiments producing an additional growth rhythmicity. Massive Porites had annual growth rates of ∼4.0 mm, whereas in ramose branching Porites, a conspicuous banding is formed by concentrations of marine micropelletoidal cement along dissepiments at ∼1.8 mm spacing. If taken as annual growth increments, these bands represent very low extension rates, however, they may rather reflect subannual forcing functions (i.e., lunar cycles). An identical scenario of precipitation and concentration of pelletoidal carbonate along dissepiments and dissolution-controlled documentation of growth bands can be inferred for Late Jurassic microsolenids. Therefore, growth bandings in fossil corals potentially reflect both, monthly and annual cycles. Consequently, care must be taken when using coral growth bands in palaeoecology and palaeoclimatology.     

The history of Devonian-Carboniferous reef communities: Extinctions,effects, recovery

Summary Analysis of the taxonomic composition, diversity and guild structure of five “typical” reef and mud mound communities ranging in age from Late Devonian-Early Carboniferous indicates that each of these aspects of community organization changed dramatically in relation to three extinction events. These events include a major or mass extinction at the end of the Frasnian; reef communities were also effected by less drastic end-Givetian and mid-late Famennian extinctions of reef-building higher taxa. Peak Paleozoic generic diversities for reef-building stromatoporoids and rugose corals occurred in the Eifelian-Givetian; reef-building calcareous algal taxa were longranging with peak diversity in the Devonian. These three higher taxa dominated all reef-building guilds (Constructor, Binder, Baffler) in the Frasnian and formed fossil reef communities with balanced guild structures. The extinction of nearly all reef-building stromatoporoids and rugose corals at the end of the Frasnian and the survival of nearly all calcareous algac produced mid-late Famennian reef communities dominated by the Binder Guild. Despite the survival of most calcareous algae and tabulate corals, the mid-late Famennian extinction of all remaining Paleozoic stromatoporoids and nearly all shelf-dwelling Rugosa brought the already diminished Devonian reef-building to a halt. These Devonian extinctions differ from mass extinctions by the absence of a statistically significant drop in taxonomic diversity and by their successional and cumulative effects on reef communities. Tournaisian mud mounds contain communities markedly different from the frame-building communities in Late Devonian and Visean reefs. Mound-building biotas consist of an unusual association dominated by erect, weakly skeletonized members of the Baffler Guild (chiefly fenestrate Bryozoa; Pelmatozoa) and laterally expanded, mud-binding algae/stromatolites and reptant Bryozoa. The initial recovery to reefs with skeletal frameworks in the Visean was largely due to the re-appearance of new species of abundant colonial rugose corals (Constructor Guild) and fenestrate Bryozoa. This Frasnian-Visean evolution in the taxonomic composition and structure of the reef-building guilds is also expressed by abrupt changes in biofacies and petrology of the reef limestones they produced. Thus, “typical” Frasnian reef limestones with balanced guild structures are framestones-boundstones-bafflestones, Famennian reefs are predominantly boundstones, Tournaisian mud mounds are bafflestones and Visean reefs are bafflestones-framestones.     

Environmental influences on skeletal banding in eastern Pacific (Panama) corals

The timing of skeletal band formation and concomitant changes in calcification rates and linear skeletal extension were investigated in Pavona corals growing under two distinct thermal regimes along the Pacific coast of Panama: fluctuating, marked by seasonal upwelling (Gulf of Panama) and stable, nonupwelling (Gulf of Chiriqui). The purpose of this study was to test the hypothesis that banding in corals is largely mediated by seasonal variations in temperature (Highsmith 1979). Our results indicate that the timing of band formation is synchronous at these two environmentally distinct locations. The low density (LD) portion of the annual band is accreted over a five month period (January–June) and represents an increase in linear skeletal extension (mm/mo.) as well as a marked increase in calcification rate (g CaCO₃ · cm^-2 · mo^-1) relative to the high density portion which forms over the remaining seven month period (July through December). In contrast to the predictions of the Highsmith model these findings indicate that variations in light levels rather than fluctuation in temperature is a better correlate to changes in skeletal density. Qualitatively, banding patterns were similar at the two sites; however, higher growth rates (particularly with respect to the LD band) for Pavona clavus in the Gulf of Panama indicate that lower water temperatures and higher productivity, or both, may be responsible for quantitative differences in banding between sites. We found that formation of the HD band corresponds to lower light levels and the production of gametes. We propose that banding in corals is a complex phenomenon governed by endogenous processes (e.g. reallocation of energy from growth to reproduction) which may be mediated by exogenous factors (e.g light and productivity).     

Size and shape distribution of level-bottom tabulate corals and stromatoporoids (Silurian)

Baarli, B. G., Johnson, M. E. & Keilen, H. B. 1992 07 15: Size and shape distribution of level-bottom tabulate corals and stromatoporoids (Silurian).
Size and shape distribution of tabulate coral and stromatoporoid faunas were studied at two Lower Silurian sections in Norway on the Baltic platform and one section in eastern Iowa on the Laurentian platform. All the sections are dominated by storm beds with predominantly shallowing upward features; they differ from one another in clastic content and proximity to land. Most of the fauna lived below normal wave-base, but within the photic zone, on fine grained and firm bottoms. In all three localities, size clearly increases up-section for stromatoporoids, favositids, and heliolitids but it decreases for halysitids in the Oslo region. Sediment influx worked negatively on size but generally not so much as to affect average size. lurbulence in combination with decreasing sediment influx affected the size decrease in heliolitids up section. Nutrients and turbidity were of minor importance as growth controlling factors. Light intensity is the only factor that shows a pattern consistent with increased size up-section. Sediment influx alfected fauna composition and shape: heliolitids had the highest sediment tolerance and stromatoporoids had the lowest. Ragged skeletons were linked to tempestites. Corals and stromatoporoids reveal a population-specific response to environmental change, largely independent of benthic assemblage zones or communities. Tabulate corals, stromaroporaids, bathymetry, size variation. clastic input, benthic assemblages .     

Fine-Scale Skeletal Banding Can Distinguish Symbiotic from Asymbiotic Species among Modern and Fossil Scleractinian Corals

Understanding the evolution of scleractinian corals on geological timescales is key to predict how modern reef ecosystems will react to changing environmental conditions in the future. Important to such efforts has been the development of several skeleton-based criteria to distinguish between the two major ecological groups of scleractinians: zooxanthellates, which live in symbiosis with dinoflagellate algae, and azooxanthellates, which lack endosymbiotic dinoflagellates. Existing criteria are based on overall skeletal morphology and bio/geo-chemical indicators—none of them being particularly robust. Here we explore another skeletal feature, namely fine-scale growth banding, which differs between these two groups of corals. Using various ultra-structural imaging techniques (e.g., TEM, SEM, and NanoSIMS) we have characterized skeletal growth increments, composed of doublets of optically light and dark bands, in a broad selection of extant symbiotic and asymbiotic corals. Skeletons of zooxanthellate corals are characterized by regular growth banding, whereas in skeletons of azooxanthellate corals the growth banding is irregular. Importantly, the regularity of growth bands can be easily quantified with a coefficient of variation obtained by measuring bandwidths on SEM images of polished and etched skeletal surfaces of septa and/or walls. We find that this coefficient of variation (lower values indicate higher regularity) ranges from ~40 to ~90% in azooxanthellate corals and from ~5 to ~15% in symbiotic species. With more than 90% (28 out of 31) of the studied corals conforming to this microstructural criterion, it represents an easy and robust method to discriminate between zooxanthellate and azooxanthellate corals. This microstructural criterion has been applied to the exceptionally preserved skeleton of the Triassic (Norian, ca. 215 Ma) scleractinian Volzeia sp., which contains the first example of regular, fine-scale banding of thickening deposits in a fossil coral of this age. The regularity of its growth banding strongly suggests that the coral was symbiotic with zooxanthellates.     

Paleoecology of Upper Eifelian and Lower Givetian coral limestones in the northwestern Sauerland (Devonian; Rhenish Massif)

Summary The prevailing sandy/silty lower part of the Middle Devonian in the northwestern Sauerland includes two coral limestone horizons, which contain a rich fauna of corals, stromatoporoids, and calcareous algae. The Ihmert-Formation is subdvided into three parts. The older coral limestone horizon is the Grünewiese-Member of the Ihmert-Formation (uppermost Eifelian), the younger is in the Bredenbruch-Member of the Unterhonsel-Formation (lower Lower Givetian). Conclusions about the environmental constraints are drawn from the sedimentology and the fossil content of the coral limestones. Predominant biostromes are built between storm wave base and normal wave base. Only the few bioherms grew above the normal wave base. These coral limestones were deposited in a tropical or subtropical normal marine environment in the shallow euphotic zone. Among the reef-builders epoecism is very frequent, and until now this phenomenon has not been investigated in detail. Fragile rugose and tabulate corals lived as commensals with stromatoporoids. Some other aspects of paleoecology are concisely presented.     

Paleoecologic,temporal, and spatial analysis of early Silurian Reefs of the Chicotte Formation,Anticosti Island,Quebec, Canada

Summary Reefs of the Lower Silurian Chicotte Formation are the largest and most faunally diverse known on Anticosti Island, Quebec. They reach up to 25 m in thickness and 250 m in diameter and are present predominantly at two intervals, forming a lower and upper reef cluster. Remnants of bioherms are represented on the present-day wave-cut terrace as 60 to 100 m diameter, subcircular erosional depressions known as Philip structures or as outcrop. The bioherms were relatively low structures, with approximately 3 to 5 m maximum synoptic relief, some of which developed on hardgrounds and possible paleokarst surfaces of crinoidal wackestone and packstone. Dominant skeletal framework builders and sediment producers within all of the reefs are laminar to low domical stromatoporoids, colonial cerioid and fasciculate rugose corals, colonial tabulate corals, and cryptostome bryozoans. Vertical zonation of reef biota is evident within well-exposed reefs of the lower reef cluster. Three to four stages are recognizable:1) a low-diversity tabulate coral-dominatedpioneering community including large tabulate coral colonies (halysitids and favositids), and few stromatoporoids (clathrodictyids, ecclimadictyids), fasciculate rugosans, large generally monotypic stalked crinoids, and shelly benthos (brachiopods, few ostracodes and trilobites);2) an intermediate- to high-diversity, mixed tabulate coral-stromatoporoid-dominatedreef-core community;3) a slightly lower diversity stromatoporoid-tabulate coral-dominatedclimax community with laminar coenitids and alveolitids; and,4) in a few localities, a capping, low-diversity tabulatecoral-dominated (alveolitid and coenitid), and stromatoporoid-bearing community comprising laminar forms. Amelioration of Early Silurian climates, following Late Ordovician glaciation, allowed gradual reestablishment of extensive shallow-water reef growth, by mainly new and increasingly diverse genera and species of metazoans. Reef development within the Chicotte Formation coincided with global, widespread development of latest Llandovery and earliest Wenlock reefs in subtropical to tropical areas. Chicotte reefs have broad characteristics, in terms of overall biotic composition, vertical successions recognized, and paleogeographic setting, similar to those of equivalent and slightly younger age from intracratonic settings in Baltica (Gotland, Sweden and Estonia) and central and northern Laurentia (Midcontinent, U.S.A.; Hudson Bay, Canada; and North Greenland, Denmark).     

Early Devonian reefs at Reefton, New Zealand: guilds, origin and paleogeographic significance

Study of 35 systematically collected and 10 semi‐random samples (about 100 cm 2 each) from one outcrop of the Lankey Limestone (Emsian) near Reefton, New Zealand indicates that it is a reef framestone built by tabulate (4? spp.) and clonal rugose (one sp.) corals, two species of stromatoporoids, a few bryozoans (2? spp.) and crinoids. The guild structure of the reef community and data on skeleton orientation and growth direction further indicate that there was relatively minor biostratonomic alteration of the original community. Skeletons are either in growth position or tipped, toppled or even overturned, but they have remained in situ. Other clone‐rich Lankey Limestone outcrops in the Reefton area support the notion of an upper shelf reef system and stratigraphic and sedimentologic data suggest that it was located near the Gondwana margin at a paleolatitude of about 35°S.     

The skeletal structure of Desmophyllum cristagalli: the use of deep-water corals in sclerochronology

Skeletal banding has been found in the deep-water scleractinian coral Desmophyllum cristagalli , an important animal in studies of climate change. This banding pattern sheds light on skeletogenesis and suggests methods by which the record of climate change contained within the coral skeletons may be interpreted. A central wall built of trabeculae forms the interior of the septa and rings the theca. Lamellae form a sheath over the trabecular frame, showing continuity from thecal edge to septum. Skeletal bands are added by the tissue layer, which overlaps and seals the internal coral and upper portion of the outer theca. Truncated inner bands on the outer theca indicate a pattern of skeletal deposition and dissolution dependent on the presence or absence of the live tissue layer. A long-term record will be difficult to collect from D. cristagalli since lamellae are less than 10 μm thick and band position is unpredictable. Density banding in shallow-water coral skeletons has long been recognized as a valuable paleo-oceanographic tool, and deep-water corals are now being used to reconstruct deep-ocean environments. Pattern of skeletal growth must be carefully considered if deep-water corals are to be used as proxy climate recorders.     

Seasonal growth bands in Famennian rugose coral Scruttonia kunthi and their environmental significance

Large colonies of rugose coral Scruttonia kunthi occurring in the upper Famennian of Sudetes (southern Poland) reveal distinct growth banding in their skeletons. They were investigated for internal structural characteristics and stable isotopic composition. The skeletal tissue consists of alternating light and dark bands which differ in thickness, density and morphology of structural elements, and in occurrence of corallite contraction and rejuvenescense. Darker parts with densely arranged thick skeletal elements are thin in comparison to lighter parts. In addition, they include frequently offsets and contraction of corallites. A couplet of dense and less dense bands is interpreted to represent most probably an annual cycle. The calculated growth rate for Scruttonia kunthi varied from 6 mm/yr to 12 mm/yr. Growth-band formation was influenced environmentally. Oxygen isotopic data provide an evidence that high-density bands were formed in the season of higher environmental stress, with relatively warmer temperatures and higher sedimentation rates. Carbon isotopic signatures are very uniform, and thus enigmatic. They indicate that at least growth rate of the skeleton and seawater temperature had no influence on the coral δ¹³C.     

Upper Permian (Murghabian) rugose corals from Oman (Ba’id area,Saih Hatat): Community structure and contributions to reefbuilding processes

Summary Coral-dominated communties are rare in Upper Permian reefs. The study of Murghabian rugose and tabulate corals from allochtonous carbonates (‘Oman exotics’) of the Hawasina Complex and autochthonous carbonates of the Saih Hatat area/Arabian Platform (Oman) provides evidence for a significant contribution of rugose corals to the formation of Late Permian reefs. The corals are described with respect to taxonomy, microfacies and community structure. 8 genera and 7 species were recognized.Monothecalis minor n.sp.,Praewentzelella regulare n.sp. andWentzelella katoi magna n.ssp. are new. The corals represent three communities: (1)Praewentzelella community (Hawasina Complex), (2) cerioid coral community (Hawasina Complex), and (3)Waagenophyllum community (Hawasina Complex and Saih hatat). The corals from the Hawasina Complex and the Saih Hatat flourished in significantly different environments: Rugosa from the Hawasina Complex are representatives of reefs, whereas their counterparts from the Saih Hatat lived in level-bottom communities. Coral-bearing reefal boundstones are characterized by a diverse assemblage of sphinctozoans, inozoans, chaetetids, bryozoans, crinoids,Tubiphytes, Archaeolithoporella and algae. These communities produced bafflestones or framestones and were part of a sponge reef complex. The level-bottom community of the Saih Hatat is low-diverse only comprising rugose and tabulate corals. These of isolated colonies locally acted as bafflers.     

Fluorescent and skeletal density banding in Porites lutea from Papua New Guinea and Indonesia

Shallow water Porites lutea corals were collected along two transects normal to mainland shorelines, parallel to gradients in water quality: one, 7 km long, near Motupore Island in South Papua New Guinea, the other, 70 km long, from Jakarta Bay along the Pulau Seribu chain in the Java Sea. The corals were slabbed and studies were made of skeletal density bands as revealed by X-ray photography and fluorescent bands as revealed by ultraviolet light. Water quality measurements and rain-fall data were assembled for the two areas and related to skeletal banding patterns. For both areas, with increasing distance form mainland there is a decrease in overall brightness of fluorescence in corals and an increase in the contrast between bright and dull fluorescent bands. Fluorescence is bright, but seasonal banding is obscure in corals within about 2 km of stream mouths at Motopure and about 5 km of the coast in Jakarta Bay; this suggests that, despite low freshwater run-off during dry seasons, there are sufficient organic compounds which cause fluorescence in coral skeletons, to swamp seasonal effects. During the wet seasons, deluges of freshwater consequent on mainland rainfall of greater than about 150 mm/ month extend at least 7 km offshore in the Motupore area and perhaps tens of kilometres into Java Sea, giving distinctive bright and dull fluorescent banding in off-shore corals. The fluorescent banding pattern within corals on the Motupore reefs is similar in most corals along the transect and it correlates well with the Port Moresby monthly rainfall data. This relationship suggests that the same body (or bodies) of freshwater affect all reefs of the area during the wet season. The fluorescent banding in Java Sea corals does not show a precise correlation with either mainland or island monthly rainfall data; indeed the pattern of fluorescent banding on Pulau Seribu can only be matched in corals from reefs less than about 25 km apart. This suggests that in this area discrete water bodies carrying the relevant organic acids for coral fluorescence affect the fringing reefs on the chain of islands. Comparisons of fluorescent and density banding have revealed that for these areas, in general, periods of high freshwater run-off are times of deposition of less dense skeleton in Porites lutea corals.     

Coral growth rates and environmental control of density banding

Linear and mass growth rates are compared for the massive coral species Favia pallida (Dana), Goniastrea retiformis (Lamarck), and Porites lutea Milne Edwards & Haime at Enewetak Atoll. Marshall Islands. Goniastrea retiformis is the densest of the three species and has an intermediate growth rate; Porites lutea grows more rapidly. All three grow indeterminately at a declining rate with increasing depth.The high-density portion of annual band couplets is produced during the late summer and fall when water temperatures are highest and possibly the availability of light is reduced, and the low density portion is formed during periods of seasonally lower water temperatures and possibly higher availability of light. A similar pattern is found in three massive coral species from Belize.The high density portions of annual bands account for a greater proportion of linear and mass growth in deeper water and in corals with relatively slow growth rates. I predict that linear growth rates will be highest where conditions are most favorable for deposition of the low density portion.Geographical patterns of coral density banding based on the literature are discussed and a model is proposed relating the interplay of light availability and water temperature to the production of high and low density skeletal bands.     

Densitometry from digitized images of X-radiographs: Methodology for measurement of coral skeletal density

Skeletons of massive coral colonies contain annual density bands that are revealed by X-radiography of slices cut along growth axes. These bands allow measurement of skeletal growth parameters such as annual extension rate and annual calcification rate. Such measurements have been important in understanding coral growth, in assessing environmental impacts and in recovering proxy environmental information. Measurements of coral calcification rate from annual density banding require measurements of skeletal density along tracks across skeletal slices and, until now, such density measurements have depended upon specialized and expensive equipment. Here, we describe a straightforward, inexpensive and accurate technique for measuring skeletal density from digitized images of X-radiographs of coral skeletal slices. An aragonitic step-wedge was included in each X-radiograph of a coral slice together with two aluminium bars positioned along the anode-cathode axis. Optical density was measured along tracks across the X-ray images of these different objects. The aragonite step-wedge provided a standard for converting optical density to skeletal density. The aluminium bars were used to correct for the heel effect—a variation in the intensity of the X-ray beam along the anode-cathode axis that would, otherwise, introduce large errors into measurements of skeletal density. Exposure was found to vary from X-radiographs to X-radiograph, necessitating the inclusion of the calibration standards in each X-radiograph of a coral slice. Results obtained using this technique compared well with results obtained by direct gamma densitometry of skeletal slices.     

Structure and growth rates of the high-latitude coral: <Emphasis Type="Italic">Plesiastrea versipora</Emphasis>

The high-latitude coral species Plesiastrea versipora was investigated to identify growth rates in colonies over 1 m in diameter. Six colonies from two temperate gulfs (latitudes of 33°–35°S) in Southern Australia were examined using X-ray, luminescence and ²³⁸U/²³⁰Th dating techniques. Annual density bands were present in each coral but varied in width and definition, suggesting different linear extension and calcification rates. Differences in density band width were observed at the local scale (amongst colonies on the same reef) and regional scales (between the two gulfs). Extension rates of the P. versipora colonies examined in this study varied between 1.2 and 7 mm per year, which are amongst the slowest growth rates reported for hermatypic corals. As only one of the six P. versipora colonies had obvious luminescent banding, we conclude that luminescent banding is not an accurate chronological marker in this species of temperate water coral. Coral age estimates derived from counting density bands in X-radiographs ranged from 90 to 320 years for the six colonies studied. U-Th ages from the same colonies determined by high-precision multi-collector inductively coupled plasma mass spectrometer established radiometric ages between 105 and 381 years. The chronological variation in absolute ages between the two techniques varied between 2 and 19% in different colonies, with the lowest growth rates (~1 mm) displaying the greatest variation between density band age and radiometric U-Th age. This result implies that the age of P. versipora and other slow-growing corals cannot be determined accurately from density bands alone. The outcome of this research demonstrates that P. versipora may be useful as a paleoclimate archive, recording several centuries in a single colony in high-latitude environments (corals found in latitudes greater than 30° in either hemisphere), where other well-established coral climate archives, such as Porites, do not occur.     

Gender-related differences in the apparent timing of skeletal density bands in the reef-building coral Siderastrea siderea

Density banding in skeletons of reef-building corals is a valuable source of proxy environmental data. However, skeletal growth strategy has a significant impact on the apparent timing of density-band formation. Some corals employ a strategy where the tissue occupies previously formed skeleton during as the new band forms, which leads to differences between the actual and apparent band timing. To investigate this effect, we collected cores from female and male colonies of Siderastrea siderea and report tissue thicknesses and density-related growth parameters over a 17-yr interval. Correlating these results with monthly sea surface temperature (SST) shows that maximum skeletal density in the female coincides with low winter SSTs, whereas in the male, it coincides with high summer SSTs. Furthermore, maximum skeletal densities in the female coincide with peak Sr/Ca values, whereas in the male, they coincide with low Sr/Ca values. Both results indicate a 6-month difference in the apparent timing of density-band formation between genders. Examination of skeletal extension rates also show that the male has thicker tissue and extends faster, whereas the female has thinner tissue and a denser skeleton—but both calcify at the same rate. The correlation between extension and calcification, combined with the fact that density banding arises from thickening of the skeleton throughout the depth reached by the tissue layer, implies that S. siderea has the same growth strategy as massive Porites, investing its calcification resources into linear extension. In addition, differences in tissue thicknesses suggest that females offset the greater energy requirements of gamete production by generating less tissue, resulting in differences in the apparent timing of density-band formation. Such gender-related offsets may be common in other corals and require that environmental reconstructions be made from sexed colonies and that, in fossil corals where sex cannot be determined, reconstructions must be duplicated in different colonies.     

Paleoecology of a low-diversity silurian community from the tofta beds of Gotland

The Silurian (Wenlockian) Tofta Beds at Galgberget 1, Gotland, Sweden, formed in a protected intertidal setting. Massive fenestral limestone at this locality contains a low diversity community dominated by stromatoporoids, calcareous algae, and ostracods, with less common rugose corals, bryozoans, brachiopods, and trilobites. Abundance of stromatoporoids, which form about 40% of sediment volume, suggests reef-like conditions. The Tofta community differs from typical Silurian reef communities, however, in its low diversity, very limited tiering, and absence of groups such as crionozoans and tabulates. These differences are possibly due to intertidal conditions which precluded upward growth of a mound structure and subjected the community to periodic desication.     

Paleoecology of Middle Ordovician stromatoporoid mounds in Vermont

Fossiliferous mounds of carbonate mud are a distinctive facies in the middle Chazy Group (Crown Point Formation) at Isle La Motte, Lake Champlain. The mounds are surrounded by bedded calcarenite of spar-cemented pelmatozoan debris. Channels which cut into the mounds during mound growth are filled with the same calcarenite. The mud-free intermound rocks and the mound biota suggest agitated, normal marine shallow-water environments. The principal lime-secreting organisms within the mounds are stromatoporoids, calcareous algae, tabulate corals, sponges, and bryozoans. Each mound is dominated in terms of biomass by one of three groups: stromatoporoids, calcareous algae, and bryozoans. Most of the mound biota first appear at the base of the Crown Point Formation. In the lower Crown Point Formation the organisms increase in number and species. Both changes in the biota are related to periods of shallowing of the Chazy sea which are also reflected in the character of the carbonate sands.     

Chaetetid growth form and its controlling factors

Chactetid sponge morphology is examined to provide details on growth styles and their controlling factors. Chactetid growth forms range from laminar to domical. bulbous. columnar and complex branching in a variety of sizes. The chactetid skeleton began as a laminar unit comprising growth of many calicles across the substrate at the same time. Several styles of early growth. involving differential calicle growth rates and varying directions of adjacent calicle growth. are recognized. and result in complex arrangements of caliclcs in the skeleton. Despite this. the cross-sectional protile of the gross morphology at any stage of growth is usually a simple outline. implying that internal complexities of calicle development are modulated to produce an optimum cross-sectional outline for the individual chactetid. The morphological range of chactetids is similar to stromatoporoids. some tabulate. heliolitid and colonial rugose corals. some bryozoans. stromatolites. encrusting foraminifera and calcareous algae: the common environmental controlling factors of sedimentation and turbulence profoundly influenced growth form in all these organisms by virtue of their common sessile shallow marine habit. Chactetid growth forms show a general relationship to the environment: columnar and branching forms grew in quiet water. while laminar and domical were better adapted to environments of higher energies. The environmental adaptations of laminar forms. however. remain problematic. because they are found in both high and low energy facies. and interpretation depends strongly on facies study. Also. interpretation of all growth forms is suspected to relate to taxonomic aspects. as has been recognized for other groups. Unfortunately. chaetetid taxonomy is in need of revision. and at present no certain relationship has been demonstrated between taxonomy and growth form. Some modern calcareous sponges with a chaetetiform architccture also show similarities in growth form to fossil types. and may be subject to similar controls. □Chaetetid. calcified sponge. growth form. Pennsyloanian. North America.