Improving the parameterization of stromatoporoid shapes – a detailed approach to stromatoporoid morphometry

Eight specimens of Devonian stromatoporoids with well visible latilaminae arrangement exposed on polished slabs were subjected to a detailed morphometric analysis. The studies have revealed that the so far used stromatoporoid parameterization method leaves a broad field of uncertainty concerning the exact definitions of particular parameters. The possible ways of making the measurements and the propositions of more precise definitions of the parameters used in the method, both of the growth form above the sea bottom and of the whole skeleton are therefore presented in this paper. The B_m - basal length of the skeleton, has been defined as a straight line joining the two ends of the basal surface, which have been pointed basing on the stromatoporoid's overall shape, latilaminae arrangement and angular relation to the main growth axis position in ontogenetical development, length. The B_n - basal length of the growth form above the sea bottom, is to be measured along a straight line joining the ends of the last visible latilamina. From among a number of possible definitions of the V_m - vertical height of the whole skeleton, a straight line joining the initial growth nucleus and the highest point on the specimen's surface above the B_m line has been selected. The V_n - vertical height of the growth form above the sea bottom is defined as the height of the point on the stromatoporoid surface highest above the B_n measured perpendicularly to it. A parameter of burial ratio, BR=(V_m-V_n)/V_m, is introduced.     

Facies and paleoecology of the late ordovician (Caradoc-Ashgill) stromatoporoid bioherms of Tasmania, Australia

Summary Stromatoporoids, together with other sedentary organisms, form bioherms in the Ordovician Gordon Group which were deposited on a carbonate platform of the Western Tasmanian Terrane. The shallow marine carbonates of the older formations show monotonous lithofacies and biota. The variety of the lithofacies and the diversity of sedentary organisms increases in the younger formations which exhibit evidence of subaerial exposure (fissure fillings and mud-cracks). These phenomena partly reflect the tectonic history of the Western Tasmania Terrane, and probably indicate a general increase in amplitude of sea-level change during the late Ordovician (Caradoc-Ashgill). The bioherms are most frequent in the uppermost horizons (the Den Formation—late Caradoc to early Ashgill?), where the outerops exhibit floatstone and bindstone fabrics. Stromatoporoids and corals construct generally small-scale (less than several meters in width and less than 1 m in height) binding structure. Based on growth forms, stromatoporoid genera are assigned to two morphotypes. Morphotype A generally shows laminar to low domical forms (low height/width ratio) exhibiting ragged margins and sediment inclusions within skeletons. In contrast, morphotype B consists of high domical growth forms (high height/width ratio) and lacks sediment inclusions. These differences in growth forms are interpreted to reflect different modes of biomineralization, together with environmental preferences of the individual organisms. Results of this study and previous publications, overall suggest a progressive development and diversification of biohermal biota in the middle to upper Ordovician of Tasmania. The scale and diversity of the Tasmanian bioherms are probably much smaller than the bioherms and reefs of the younger ages (Silurian and Devonian), and for stromatoporoids, the tendency of diversification is consistent with those of the other Ordovician sections. The Gordon Group provides useful information regarding the early evolutionary history of the Ordovician-Devonian reef-forming communities.     

Stromatoporoid palaeoecology in the Frasnian (Upper Devonian) Belgian platform,and its applications in interpretation of carbonate platform environments

Abstract: Stromatoporoid faunas in the Frasnian of southern Belgium are abundant in the carbonate platform environments present in this area. Stromatoporoids dominate the large skeletal organisms and occur principally in biostromes. The stromatoporoid assemblage is represented by a small number of taxa. Stromatoporoid genera include Actinostroma, Amphipora, Atelodictyon, Clathrocoilona, Salairella, Stachyodes, Stictostroma, Stromatopora and Trupetostroma which are present in environments ranging from the outer, outer intermediate, inner intermediate and inner zones and associated biostromes. Most large skeletal stromatoporoids are low profile, which reinforces the conclusions of previous studies that low‐profile growth forms were the most successful stromatoporoid forms. These low‐profile forms are likely to have been important sediment stabilisers that may have led to expansion of the carbonate factory. Growth forms vary between facies, indicating some degree of environmental control on form; for example, laminar in the intermediate zone, bulbous and domical in the inner and outer zones. Stromatoporoid taxa vary in occurrence across the environmental gradient from shallow to deep. There is some taxonomic control on growth forms, with some taxa showing more variability than others in different environments.     

Intracoenosteal variation in a Devonian stromatoporoid

Fagerstrom, J. A. & Saxena, K. M. L.: Intracoenosteal variation in a Devonian stromatoporoid.
Statistical analysis of four different features of coenosteal morphology measured in sixteen randomly distributed vertical and tangential thin sections from a specimen of Syringostroma sherzeri (Grabau) from the Detroit River Group near Ingersoll, Ontario, indicates that although the degree of variation is very high, the variation in a single section is representative of the variation in the entire coenosteum. This discovery may therefore be regarded as a partial vindication of current practice in stromatoporoid taxonomy of baaing identifications on examination of one vertical and one tangential sectior. Furthermore, this study indicates that certain morphological features are less variable than others and are therefore of greater value as taxonomic criteria. In cases where the number of measurements per section is small, tests of the significance of differences based on a comparison of the medians, rather than means, is preferred because such tests do not assume that the data are normally distributed.     

Sedimentology and stromatoporoid palaeoecology of Frasnian (Upper Devonian) carbonate mounds in southern Belgium

Da Silva, A.‐C., Kershaw, S. & Boulvain, F. 2011: Sedimentology and stromatoporoid palaeoecology of Frasnian (Upper Devonian) carbonate mounds in southern Belgium. Lethaia, Vol. 44, pp. 255–274. Stromatoporoids are the most abundant large skeletal organisms in middle Frasnian carbonate mound environments of southern Belgium. They occur in environments ranging from flank and off‐mound, mound core, shallow mound and restricted mound. A detailed log and comprehensive sampling of stromatoporoids in a single section cutting through all middle Frasnian mound levels in La Boverie–Rochefort Quarry, near Rochefort and Dinant reveals a stromatoporoid assemblage comprising 10 genera; 472 samples, containing an overall total of 3079 stromatoporoids (including complete and fragmented specimens) have been studied. The following list gives abundance using numbers of specimens and areas of total stromatoporoid area on outcrop surfaces (% number; % area in cm²): Actinostroma (0.4; 9.2), Amphipora (15.5; 1.7), Atelodictyon (0.2; 4.4), Clathrocoilona (0.3; 0.5), Euryamphipora (13.7; 0.7), Idiostroma (2; 1.9), Salairella (1.2; 9.6), branching Stachyodes (43.2; 59.1), laminar Stachyodes australe (1.9; 1.3), Stictostroma (4.8; 13.1) and Trupetostroma (0.2; 0.8), showing that Stachyodes is approximately half of the total assemblage. Deeper environments contain more abundant low profile forms, shallow water facies contain more domical and bulbous forms; branching forms are ubiquitous. Low profile stromatoporoids are likely to have been important sediment stabilizers that may have led to expansion of the carbonate factory, and they may have therefore contributed to the structural building of the mounds. Stromatoporoid‐coral intergrowths are observed in only Stictostroma suggesting that there is a close biological relationship between them; however, stromatoporoid skeletons in almost all cases appear to be unaffected by the presence of intergrown corals, suggesting they were commensals. □Frasnian, Late Devonian, mounds, palaeoecology, stromatoporoid.     

Growth forms and palaeoenvironmental interpretation of stromatoporoids in a Middle Devonian reef, southern Morocco (west Sahara)

Growth forms of well-preserved stromatoporoids, including genera Actinostroma, Stachyodes, and Stromatopora, are described for the first time from the Devonian Sabkhat Lafayrina reef complex of southern Morocco (west Sahara), one of the best exposed Middle-Devonian stromatoporoid-dominated fossil reefs. Three facies types representing the well illuminated fore-reef, reef-core and transition to back-reef facies display the distribution and growth of stromatoporoids in a high latitude setting at 40–50° south of the palaeoequator. Stromatoporoids are largely in growth position and reflect the well-preserved reef architecture. Although outcrops are low topography, the reef's prominent profile is indicated by presence of spur and groove form and a clearly defined reef margin. Stromatoporoids are mostly laminar and domical forms, with little evidence of ragged margins, and indicate normal turbulence shallow waters, with low sediment deposition.     

Early ordovician reefs from Argentina: Stromatoporoid vs stromatolite origin

Summary Late Arenigian biohermal reef mounds and biostromes within the shallow-marine platform facies of the upper San Juan Formation of the Precordillera (Western Argentina) represent a new Early Ordovician reef type. The meter-sized reefs are dominated byZondarella communis n.g. n. sp. The new taxon is characterized by domical, bulbous and laminar morphotypes exhibiting growth layers and thin horizontal and vertical as well as intermingled skeletal elements included within different sets. The fossil maybe compared with stromatolites and stromatoporoids but an interpretation as primitive stromatoporoids is favoured.     

Euzkadiella erenoensis n. gen. n. sp. ein stromatopore mit spikulärem skelett aus dem oberapt von ereño (prov. guipuzcoa,nordspanien) und die systematische stellung der stromatoporen

Euzkadiella erenoensis n. gen. n. sp. from the Early Cretaceous of Ereño (Prov. Guipuzcoa, northern Spain) is the first known »stromatoporoid« with a spicular skeleton and a basal skeleton consisting of calcific spherulites. The stromatoporoid sponge shows subtylostyle, oxea and strongyle megascleres. Microscleres are unknown. The sclere arrangement appears as disorganized bundles connected by horizontal sclere bridges. This sclere arrangement is characteristic of the order Haplosclerida (Class Demospongiae, Subclass Ceractinomorpha). The new species is compared with the modern coralline spongesCalcifibrospongia andAstrosclera, and with the Mesozoic stromatoporoids of the families Milleporellidae and Actinostromariidae. In both subclasses of the Demospongiae and within the Class Calcarea stromatoporoid basal skeletons are observed. Therefore the subclass Stromatoporoidea does not exist as a true systematic unit.     

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.     

THE ENIGMATIC EARLY CAMBRIAN SALANYGOLINA– A STEM GROUP OF RHYNCHONELLIFORM CHILEATE BRACHIOPODS?

Abstract:  New material of the enigmatic brachiopod Salanygolina obliqua Ushatinskaya from the Early Cambrian of Mongolia shows that it has a colleplax – a triangular plate – in the umbonal perforation, which is enlarged by resorption. This structure is otherwise only known from the equally enigmatic Palaeozoic orders Chileida and Dictyonellida (Rhynchonelliformea, Chileata). The colleplax in Salanygolina is here considered to be homologous with that of the chileates. Salanygolina is also provided with a ridge-like pseudodeltidium, which is another chileate feature. Other characters of Salanygolina , like the radial arrangement of adductor muscle scars and postero-medially placed internal oblique muscles are characteristic of chileates, but also found in the paterinates. In contrast, mixoperipheral dorsal valves with low rudimentary interareas are well known in paterinates, but not yet recorded from chileates. Thus, Salanygolina shows a mosaic combination of morphologic characters, known both from the paterinates and chileates, indicating that it may represent a stem group of the rhynchonelliform chileate brachiopods. The laminar phosphatic secondary shell of Salanygolina is composed of closely packed and nearly identical hexagonal prisms, oriented with their long axis normal to the laminae in a honeycomb pattern. The prism walls appear to have originally been composed of organic membranes and might represent precursors of the organic sheaths of calcite fibers that are typical of calcitic shells with a fibrous microstructure.     

The stromatoporoid animal

Stromatoporoids were a subphylum of the Porifera whose soft parts can be reconstructed by comparisons with the living sclerosponges Merlia and Astrosclera . The living tissue was confined to the upper surface and penetrated only short distances into the coenosteum. Astrorhizae are traces of an excurrent water canal system that interfered with the secretion of the skeleton in some stromatoporoids but was entirely above the hard tissue in others. The stromatoporoid skeleton was composed of trabecular or spherulitic aragonite. Calcitization and dissolution of the aragonite proceeding from the centers of calcification outward account for the microstructures (fibrous, compact, tripartite, ordinicellular, cellular, melanospheric) commonly observed in the calcite skeletons of fossil stromatoporoids. Reconstructions showing the proposed relationship of the soft tissue to the hard tissue of Labechia, Stictostroma, Actinostroma and Stromatopora are presented.     

The relationship of the stromatoporoids to the sclerosponges

Similarities of the extinct strornatoporoids to the sponges of the recently established order Sclerospongia have strengthened arguments that these fossils are closer to the Porifera than to the Coelenterata. Major features favouring the affinity of the stromatoporoids to the sclerosponges include: (1) lack of evidence of colonialism in the strornatoporoids, (2) similarity of gross structure of some stromatoporoids to that of one sclerosponge ( Astrosclera ), (3) fibrous microstruc-ture of sclerosponges, Mesozoic stromatoporoids, and some Paleozoic stromatoporoids, (4) similarity of stromatoporoid astrorhizae to the excurrent canals of sclerosponges. Points of dissimilarity include: (1) the solid aragonitic skeletons of most sclerosponges, (2) the presence of dissepiments, laminae, and latilaminae in stromatoporoids, (3) the absence of siliceous spicules in stromatoporoids.
These comparisons suggest that the stromatoporoids were basically encrusting filter feeders like the sclerosponges but had progressed by loss of spicules and periodic introduction of dissepirnents and laminae toward a secretion of a skeleton of the coelenterate type. They cannot be placed with confidence in either the sclerosponges or the hydrozoans and should be recognized us a separate subphylum of the Porifera.     

Ultrastructure of Draparnaldia glomerata (Chaetophorales, Chlorophyceae). 1. The flagellar apparatus of the zoospore

The fine structure of the quadriflagellate zoospores of Draparnaldia glomerata (Vauch.) Agardh is described with emphasis on the flagellar root system and compared with the flagellar apparatus of related green algae. It is demonstrated that the flagellar root system in Draparnaldia is similar to that of the zoospore of Uronema belkae. Common features include presence of a cruciate root system (formula 2–5–2–5), prominent striated distal fibre connecting opposite basal bodies, a system I striated root component associated with the 2–stranded root, association of electron dense material with the 5–stranded root, mode of arrangement of the basal bodies in the absolute configuration model, and presence of four striated peripheral fibres interconnecting adjacent basal bodies. Differences exist in the shape of the striated peripheral fibres, the origin of the 2– and 5– stranded roots in the proximal part of the flagellar apparatus, and the architecture and striation pattern of the proximal part of the system I fibre that detaches from the 2–stranded root between adjacent basal bodies. Both the 2– and 5–stranded roots originate near the basal bodies and descend deeply into the zoospore. One of the 5–stranded roots passes near the eyespot of the chloroplast. The implications of these findings for the taxonomic position of the genus Draparnaldia are discussed. In addition, an evaluation is given of the present status of the order Chaetophorales. Suggestions are given to standardize some aspects of the current terminology of the cruciate flagellar root system in green algae.     

Significance of stromatoporoids in Jurassic reefs and carbonate platforms—concepts and implications

Although many case studies describe stromatoporoid-rich Jurassic reefs, there are only few reliable data as to their distribution pattern. This is in part due to a largely taxonomic and systematic focus on the enigmatic stromatoporoids which now are interpreted as a polyphyletic informal group of demosponges by most specialists. The common co-occurrence of Jurassic scleractinian corals and stromatoporoids might, at first hand, point to very similar environmental demands of both organismic groups, but autecological considerations as well as evaluation of stromatoporoid distribution patterns should allow for a much more refined interpretation. This study concludes that Jurassic corals and stromatoporoids show a relatively broad overlap of environmental demands but their maximum ecological tolerances appear to differ considerably. Jurassic corals were dominating in mesotrophic to mildly oligotrophic, slightly deeper settings, where they largely outcompeted stromatoporoids. On the other hand, stromatoporoid growth was particularly favoured in very shallow water, strongly abrasive, high-energy settings as well as in possibly overheated waters. Many taxa and growth forms were very tolerant towards frequent reworking and redistribution, a feature which is compatible with the sponge nature of the stromatoporoids. As such, stromatoporoid facies may be common in low-accommodation regimes, giving rise to frequent “shelf shaving” and redistribution across wide shelf areas. The mixed coral-stromatoporoid reefs from the margins of isolated Intra-Tethys platforms are interpreted to be indicative of oligotrophic normal marine waters. This is corroborated by statistical cluster analysis of stromatoporoid taxa from representative areas. In addition, Arabian stromatoporoid occurrences might have been adapted to overheated and slightly hypersaline waters. There also are a few exceptional stromatoporoid taxa which might have had environmental tolerances different from the bulk tolerances of other Jurassic stromatoporoids. Part of our interpretations are preliminary and should stimulate further research. However, the present results already help explain the observed compositional differences between Jurassic North Tethys/North Atlantic, Intra-Tethys, and South Tethys shallow-water reefs and platforms.     

Improved negative staining of microfilament arrangements in detergent-extracted Physarum amoeboflagellates

A motile, lamellipodium-like structure, the ridge, forms as amoeboflagellate cells of Physarum polycephalum release from a substratum and begin swimming in fluid. Actin microfilaments form a distinct laminar core within the ridge; they are seen as a sparse, disordered meshwork in cytoskeletons prepared by conventional methods using uranyl acetate negative staining [10]. Preservation and visualization of these filaments and their arrangements improved considerably when cytoskeletons were imaged with phosphotungstic acid buffered with ammonium hydroxide (PTA(NH4]. Microfilaments within ridge cytoskeletons were found to form loose bundles and criss-crossing, 'meshwork' arrays several layers deep. Differences could be detected in morphology and detailed arrangement of microfilaments within cytoskeletons prepared in the presence of phalloidin. PTA(NH4) may be useful for studies of cytoskeletal elements and their rearrangements in dynamic, motile regions of cells.     

Control of shape and pattern during the assembly of a large microtubule bundle

Microtubules are packed and linked together in a well defined hexagonal arrangement in the cytopharyngeal microtubule bundles of the ciliate Nassula. Early stages in the morphogenesis of these bundles have been examined. Elements which nucleate assembly of bundle microtubules are apparently closely associated before tubule assembly commences. These nucleating elements seem to be bound together in highly ordered arrays to form microtubule-nucleating-templetes. Each array of elements is attached to the proximal end of a basal body and appears to establish the pattern of tubule packing and cross-sectional shape of a tubule bundle. A self-assembly procedure which accounts for the anisometric growth and shaping of a template and its microtubule bundle is proposed.     

Ultrastructural analysis of the midaxial extracellular matrix in spinal dysraphism.

Ultrastructural aspects of the extracellular matrix (ECM) in the midaxial region of dysraphic embryos of the loop-tail (Lp) mutant mouse were analyzed by means of electron microscopy. In 17-23 somite embryos, ultrastructural differences in the ECM occurred with respect to the presence of a pair of long trailing basal laminar strands extending continuously from the ventral notochordal cells to the gut in abnormal (Lp/Lp) embryos, in contrast to short, ragged, discontinuous strands in normal (+/+; Lp/+) embryos. The ultrastructural localization and configuration of fibronectin (FN) and laminin (L) associated with these strands, however, were similar in normals and abnormals. In addition, FN occurred over interstitial bodies, fibrils, and sporadically along the basal laminae of the neural tube (or folds), notochord, gut, and vessels, whereas L was largely confined to the basal laminae. The results indicate that although the ultrastructural pattern of FN and L reactivity are similar in normal and abnormal embryos, a disturbance in the manner whereby the notochord detaches from the gut in dysraphic embryos may be of causal significance in the etiology of dysraphism in this mutant.     

Embryogenesis in mouth-breeding cichlids (Osteichthyes,Teleostei) structure and fate of the enveloping layer

The eggs of African mouth-brooders are of unusual size and shape. Studying their development may help to more clearly understand epiboly, gastrulation, and the relation between enveloping layer (periderm) and epidermis. When epiboly has progressed over just one fifth of the yolk mass, the germ ring and embryonic shield are already well established. Behind the germ ring very few deep cells are present at this early stage of epiboly, except in the embryonic shield. When the blastodisc covers the animal half of the yolk mass, the future body is already well established with notochord, somites and developing neural keel. Apart from these structures, no deep cells can be detected between enveloping layer and yolk surface; not even a germ ring remains behind the advancing edge of the enveloping layer. Epiboly over the greater part of the yolk is achieved only by the enveloping layer and the yolk syncytial layer. As the margin of the enveloping layer begins to reduce its circumference when closing around the vegetal pole, groups of cells in the advancing edge become spindle-shaped, with a single cell in between of each of these groups broadening along the edge. The enveloping layer (called periderm after epiboly) remains intact until after hatching, when, together with the underlying ectoderm, it forms the double-layered skin of the larval fish. Thereafter, cells deriving from the subperipheral ectoderm gradually replace the decaying periderm cells to form the final epidermis. Thus, in the cichlids studied, the enveloping layer alone forms the yolk sac to begin with, and it covers the larval body until some days after hatching.     

Structure of reciprocal connections in visual cortical areas 17 and 18 in cats

Single cortical columns of areas 17, 18 in the cat were microiontophoretically injected with horseradish peroxidase. Spatial and laminar distributions of retrogradell labelled cells in both areas were investigated. Following injections in area 17 or in area 18 the labelled cells' region in area 17 was elongated (in a tangential plane) along the representation of visual field horizontal meridian. However the labelled cells' region in area 18 was elongated along the representation of vertical meridian. Such projection patterns appear to be common in these cortical areas throughout the central 10 degrees on various elevations (from -40 degrees to +10 degrees) of the visual field representation. Thus the spatial arrangement of intrinsic and extrinsic connections in each area coincides, at the same time in area 17 they are orthogonal to area 18. The following visual information exchange scheme may be suggested. Area 17 may supply the area 18 with more detailed information on the horizontal component of the visual image, and in the opposite direction the information on the vertical component of the same image may be supplied.