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.     

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.     

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.     

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.     

Sedimentary history of Upper Silurian biostromes of Podolia (Ukraine) based on stromatoporoid morphometry

The sedimentary history of stromatoporoid biostromal accumulations reflecting various depositional conditions (autoparabiostromes and parabiostromes) is studied in two isochronous, Late Silurian carbonate sections of the Malynivtsy Formation from Podolia (western Ukraine, Kam'janec' Podil'skyj area). This study focuses on morphometrical analysis of massive stromatoporoids. Various stromatoporoid attributes, such as growth form, volume, surface character etc., are interpreted in terms of growth environments. Attributes of redeposited specimens are also analysed in terms of their susceptibility to exhumation and redeposition, and new criteria are presented in this matter. The exposed facies succession, which can be subdivided into three units: an oncolitic–fenestral complex and the stromatoporoid–coral complexes that underlie and cover it, represents the belt of shoals located at a considerable distance from shore, and its transition to a narrow zone of back-shoal tidal flats. The facies patterns proved to be strongly obscured by an intensive process of onshore redeposition of material during high energy episodes. These events caused exhumation and landward transport of stromatoporoids inhabiting soft-sediment bottoms of outer shelf areas, which were afterwards accumulated in parabiostromes in calm waters on lee side of a zone of shoals. The main process governing the distribution of redeposited stromatoporoids is fractional (weight) segregation. The high energetic events had less effect on stromatoporoid–coral autoparabiostromes that formed the zone of shoals, which were inhabited by stromatoporoids better adapted to permanent wave action, but nonetheless, they caused their partial reworking and depletion from those forms that did not resist redeposition, on one hand, and supplementation by specimens derived from offshore areas, on the other.     

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.     

Identification of calcified coccoid cyanobacteria forming stromatoporoid stromatolites

A simple technique is described enabling the identification in scanning electron micrographs (SEM) of combined Iractured and etched stromatoporoid sections, the cyanobacteria-hkc coccoid aggregates building the entire stromatoporoid structure. The coccoid aggregates from stromatoporoids are closely related to extant calcified coccoid cyanobacteria (Plcurocapsales) forming stromalolitic mats in Laguna Mormona (Baja California) and Sabkha Gavish (Sinai). Stromatoporoids, stromatolites, cyanobacteria.     

Indigenous demosponge spicules in a Late Devonian stromatoporoid basal skeleton from the Frasnian of Belgium

This paper records the first example of a demosponge spicule framework in a single specimen of a Devonian stromatoporoid from the Frasnian of southern Belgium. The small sample (2.5 × 2 cm) is a component in a brecciated carbonate from a carbonate mound in La Boverie Quarry 30 km east of Dinant. Because of the small size of the sample, generic identification is not confirmed, but the stromatoporoid basal skeleton is similar to the genus Stromatopora. The spicules are arranged in the calcified skeleton, but not in the gallery space, and are recrystallized as multi‐crystalline calcite. The spicules fall into two size ranges: 10–20 μm diameter and 500–2000 μm long for the large ones and between 5–15 μm diameter and 50–100 μm length for the small ones. In tangential section, the spicules are circular, they have a simple structure, and no axial canal has been preserved. The large spicules are always monaxons, straight or slightly curved styles or strongyles. The spicules most closely resemble halichondrid/axinellid demosponge spicules and are important rare evidence of the existence of spicules in Palaeozoic stromatoporoids, reinforcing the interpretation that stromatoporoids were sponges. The basal skeleton may have had an aragonitic spherulitic mineralogy. Furthermore, the spicules indicate that this stromatoporoid sample is a demosponge.     

Early symbiotic rugosan endobionts in stromatoporoids from the Rhuddanian of Estonia (Baltica)

The earliest known symbiotic rugosan endobionts occur in stromatoporoids from the early Rhuddanian of Estonia. The stromatoporoid Ecclimadictyon nikitini from the Tamsalu Formation contains the rugosan Donacophyllum middendorffii endobiont. A stromatoporoid Clathrodictyon boreale from the Varbola Formation contains Streptelasma estonica and Bodophyllum sp. endobionts. There are up to three endobiotic rugosans per stromatoporoid host. Stromatoporoid hosts were beneficial for symbiotic rugosans as elevated substrates on a seafloor that offered a higher tier for feeding; they also offered enhanced substrate stability. Stromatoporoids of the end‐Ordovician mass‐extinction recovery fauna hosted a diverse fauna of symbiotic endobionts. There were few if any negative effects of this mass extinction on the symbiotic endobionts.     

Occurrence of Giant Borings of Osprioneides kampto in the Lower Silurian (Sheinwoodian) Stromatoporoids of Saaremaa,Estonia

The distribution of Osprioneides is more environmentally limited than that of Trypanites in the Silurian of Baltica. Osprioneides probably occurred only in large hard substrates of relatively deepwater muddy bottom open shelf environments. Osprioneides were relatively rare, occurring in 4.7% of all stromatoporoid specimens in that environment, in contrast to small Trypanites-Palaeosabella borings, which occur in 88.4% of stromatoporoids and 88.9% of heliolitid corals. Osprioneides is reported only from the lower Sheinwoodian stromatoporoids of the exposed Silurian of Saaremaa (Wenlock to Pridoli). Osprioneides borings probably played a minor role in the general bioerosion in the Silurian of Baltica.     

High-energy sedimentary events in lagoonal successions of the Upper Silurian of Podolia, Ukraine

Three Late Silurian carbonate profiles of the Malynivtsy and Skala Formations from Podolia (western Ukraine) are discussed in terms of sedimentation dynamics. Their common feature is the appearance of thick, stromatoporoid-rich beds within fine-grained peritidal deposits. These intercalations are composed of fossils typical of offshore sedimentary environments. In the most spectacular case, a channel, several tens of metres wide and infilled with stromatoporoids, is incised in a peritidal cyclic complex. The successions investigated exhibit sedimentary features that are diagnostic of onshore redeposition. Independently of the scale of the recorded sedimentary events, the onshore redeposition was caused by factors with energy levels exceeding those of average storms, probably by hurricanes or even tsunami waves. The dynamic nature of some of the stromatoporoid beds has to be taken into account when constructing the curves illustrating bathymetrical and facies development of the Silurian succession of Podolia. The genesis of lens-shaped stromatoporoid beds, elongated depending on their origin, either parallel or perpendicular to the facies belts, should be considered an important factor in reconstructions of the depositional architecture of sedimentary hydrocarbon collectors.     

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.     

CLASSIFICATION AND CONTROLS OF INTERNAL BANDING IN PALAEOZOIC STROMATOPOROIDS AND COLONIAL CORALS

Abstract:  Palaeozoic corals and stromatoporoids exhibit a variety of internal banding phenomena, many of which have been commonly interpreted as annual growth bands. We evaluate bands through analysis of colonial corals and stromatoporoids from three stratigraphic intervals: Upper Ordovician of Manitoba Canada, and Llandovery–Wenlock and Ludlow of Gotland, Sweden. Banding features are divided into four categories: (1) absence of banding; (2) density banding formed by variation in density or form of elements; (3) growth-interruption banding indicating growth cessation and regeneration; and (4) post-mortem banding caused by compaction or diagenesis. For discrimination of band types, it is essential to examine internal structures and skeletal margins in thin sections or acetate peels. Species vary considerably in degree and type of banding; each has a distinct pattern of variation. We propose criteria to determine if banding is consistent with seasonally induced growth variation: (1) consistency in band character and thickness; (2) continuity of skeletal growth; (3) marginal features; and (4) evidence of diagenetic alteration. Density bands in tabulate and rugose corals probably represent annual growth variations, but results for stromatoporoids are more ambiguous; although stromatoporoids commonly show banding, unequivocal density banding is poorly developed and growth interruption generated most stromatoporoid banding. Cerioid rugose and tabulate corals possess the thickest density bands; the thinnest bands are in stromatoporoids and heliolitid tabulates.     

Cavities and cryptic faunas beneath non-reef stromatoporoids

Kershaw, Stephen 1980 10 15: Cavities and cryptic faunas beneath non-reef stromatoporoids. Lethaia , Vol. 13 , pp. 327–338. Oslo. ISSN 0024–1161.
Stromatoporoids from level-bottom shales and argillaceous limestones in the Silurian of Gotland, Sweden, form substrates for a variety of encrusting and boring organisms. Overturned stromatoporoids have encrusters and borers on both upper and lower surfaces, while coenostea preserved in situ have encrusters on both surfaces but borings on upper surfaces only. This suggests that cavities, now infilled, existed below coenostea. The stromatoporoids are isolated and not part of a reef framework where growth alone could have created overhangs and cavities. The scouring activity of currents removing sediment from around and beneath the edges of coenostea, and small current-controlled movements leaving stromatoporoids imperfectly settled on the uneven substrate or partly overlying skeletal debris, are invoked to explain the presence of cavities and hence encrusters on stromatoporoid lower surfaces. Both processes probably operated on many specimens. The lower surfaces of these stromatoporoids also show basal concavities which range from shallow to deep and reflect the topography of the substrate and the success of stromatoporoids growing on positive features from which they could shed sediment easily. Overturned stromatoporoids and coenostea with deep, encrusted, basal concavities, point to violent environmental events, such as storms, more powerful than currents producing scour and small movements of coenostea.     

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.     

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 .     

Parameterization of stromatoporoid shape

Kershaw, Stephen & Riding, Robert 1978 07 15: Parameterization of stromatoporoid shape. Lethaia , Vol. 11. pp. 233–242. Oslo. ISSN 0024–1164.
Stromatoporoid cross-sectional shape can be considered as a product of the interaction of three morphological variables: the relative proportions of basal, vertical and diagonal dimensions. The gross arrangement of internal lamination can be superimposed upon the resulting outlines. These shapes are taken to represent cross sections in any vertical plane through the centre of the coenosteum. This simple parameterization scheme is presented in triangular arrays which include the stromatoporoid morphotypes laminar, domical and bulbous, and varieties of them together with forms not utilized by stromatoporoids. Dendroid and irregular forms are too complex to be readily included in the scheme. Smooth and ragged varieties of laminar and domical forms are distinguished and related to sedimentation on the flanks of the coenosteum. Two types of mutual arrangement of latilaminae within the coenosteum are recognized: enveloping, where they completely overlap previous latilaminae, and non-enveloping, where they do not. Laminar-domical-bulbous forms represent a series generated mainly by reduction of the basal dimension. They commonly exhibit enveloping latilaminae except in ragged varieties and some extended domes. Measurement of defined dimensions in the field allows stromatoporoid morphotypes to be plotted onto the triangular arrays and provides a rapid method of displaying the range of forms present.     

Ein Spongienmodell für die Favositidae

The opinion that the Favositidae are a group of the Madreporaria is based on a paper by Edwards & Haime 1849. These authors mentioned no reasons for their assumption. A critical review of the sceletal material, microstructure and morphology shows that it is difficult to compare this group with the true Madreporaria. On the other hand, the discovery of the Sclerospongiae has shown that there are recent organisms which are similar in their material and morphology to the Favositidae. The only difference is the occurrence of wall pores in the Favositidae and of astrorhizae in the Sclerospongiae. Like the primary wall grooves of the upper edge of the walls of the Favositidae, which are the primary stadium of the pores, the astrorhizae are initial grooves on the surface of the Sclerospongiae. Therefore the formation of the pores and of the astrorhizae is similar. If the Favositidae are interpreted as Porifera, the wall grooves would analogically be the impressions of excurrent canals of the soft part on the surface of the skeleton. The main difficulty in an interpretation of the Favositidae as a new subphylum of the Porifera is that we are not yet informed about the formation of the basal skeleton of the Sclerospongiae.     

The oldest labechiid stromatoporoids from intraskeletal crypts in lithistid sponge–Calathium reefs

Early Ordovician (early Floian) reefs of South China include lithistid sponge–Calathium reefs with a three‐dimensional skeletal framework. These structures are among the first post‐Cambrian skeletal‐dominated reef structures and provides an opportunity to test how the novel metazoan builders changed the environments and increased topographic complexity within benthic communities. We document the oldest labechiid stromatoporoid (Cystostroma) in a lithistid sponge–Calathium reef of the Hunghuayuan Formation in southeastern Guizhou, South China. These earliest stromatoporoids may have originated in reefs, and we argue that the complex topography created by the hypercalcified sponge Calathium facilitated the emergence of stromatoporoids. Beyond Cystostroma, keratose sponges, Pulchrilamina (hypercalcified sponge) and bryozoans have also inhabited in the micro‐habitats (cavities and hard substrates) provided by Calathium. These findings suggest that ecosystem engineering by Calathium played an important role in the further diversification of reefs during the Ordovician.