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Baltic Ordovician compound sponges as erratics on Gotland (Sweden), in northern Germany and the eastern Netherlands
Abstract and Figures
Compound orchocladine sponges are unusual in the Early Palaeozoic. In Europe, silicified material of Late Ordovician age has hitherto been referred to as Aulocopium aurantium Oswald, 1847 and the invalid Aulocopium compositum Conwentz, 1905. An examination of new material has resulted in the recognition of a new genus, Hydraspongia, with two new species, H. polycephala and H. erecta, and a third new species, Perissocoelia megahabra, to which most specimens can now be assigned. These taxa form part of rich erratic sponge assemblages, which originate from unknown source areas in the Baltic, and have been collected in northern and western Europe from fluvial sandy deposits of the Eridanos River system, which drained the Baltic area from the Middle Miocene to Early Pleistocene.
... Rhebergen & van Kempen (2002) summarized a discussion between Lindstr€ om, Schmidt (1891) and Stolley (Stolley 1900(Stolley , 1929 on the age and lithostratigraphical origin of the sponges. Lindstr€ om probably intended to describe at least part of the sponge assemblage, because he had drawings made by Lilljeval, but did not complete the work (Rhebergen 2007). Schl€ uter (1884) reported on a remarkable sponge specimen, Astylospongia gothlandica, which has been figured by Rauff (1894) as an aberrant form of the Ordovician species Caryspongia diadema. ...
... Hydraspongia Rhebergen, 2007 includes compound sponges that usually form bulbous oscular heads and show conspicuously distinct growth stages. Postperissocoelia n. gen. ...
... Canals that achieved their maximum length induced the formation of a new cluster of excurrent canals, in order to drain the newly formed parts. This continuous growth apparently caused the development of new oscular heads, while the formation of stacked skeletal trabs series continued uninterrupted (Rhebergen 2007). ...
A diverse assemblage of silicified sponges, including orchocladines, rhizomorines, stromatoporoids, hexactinellids and non-lithistid demosponges, has been collected from scree accumulations in a restricted area along the western coast of the Island of Gotland, Sweden. The assemblage comprises 29 species in 20 genera, 18 species of which have not previously been recorded from Baltica, together with several taxa in open nomenclature. This chapter discusses this sponge assemblage in detail. It first gives a brief introduction on the geological settings, and the material, repository, methods and terminology used for the study of the assemblage. The chapter discusses the sponge assemblage under various families, classes and orders such as Hexactinellida, Orchocladina, Stromatoporoidea, Rhizomorina and Streptosolenida.
... During the 20 th Century, sponge research in Estonia and Russia decreased to a small number of papers in which sponge taxa were listed, such as those by Asatkin (1931Asatkin ( , 1949 and Rõõmusoks (1970), but has undergone a revival in Germany and the Netherlands since about 1980. An extensive and diverse sponge fauna has been revealed by a series of works on material from fluvial deposits in the Netherlands and adjacent parts of westernmost Germany (Van Kempen, 1978, 1990Von Hacht, 1985, 1994Reitner & Kohring, 1990;Von Hacht & Rhebergen, 1996, 1997Rhebergen, 1997Rhebergen, , 2007Rhebergen et al., 2001). ...
A new species of complex hexactinellid sponge, Haljalaspongia inaudita, is described from fluvially transported blocks found in northwestern Germany, near the Dutch border, but which probably originated from the eastern Baltic region. The heavily folded wall is composed of multiple spicule layers, including dermal and gastral layers of acanthohexactines, and a central layer of sub-parallel monaxons. The monaxial layer is lined on one side by a reticulate array of smooth monaxons. The sponge is difficult to assign to any known fossil or recent group, but shares some features with the dictyospongioid family Docodermatidae.
There are few sponges known from the end-Ordovician to early-Silurian strata all over the world, and no records of sponge fossils have been found yet in China during this interval. Here we report a unique sponge assemblage spanning the interval of the end-Ordovician mass extinction from the Kaochiapien Formation (Upper Ordovician-Lower Silurian) in South China. This assemblage contains a variety of well-preserved siliceous sponges, including both Burgess Shale-type and modern type taxa. It is clear that this assemblage developed in deep water, low energy ecosystem with less competitors and more vacant niches. Its explosion may be related to the euxinic and anoxic condition as well as the noticeable transgression during the end-Ordovician mass extinction. The excellent preservation of this assemblage is probably due to the rapid burial by mud turbidites. This unusual sponge assemblage provides a link between the Burgess Shale-type deep water sponges and the modern forms. It gives an excellent insight into the deep sea palaeoecology and the macroevolution of Phanerozoic sponges, and opens a new window to investigate the marine ecosystem before and after the end-Ordovician mass extinction. It also offers potential to search for exceptional fossil biota across the Ordovician-Silurian boundary interval in China.
Sponges have an unrealized potential importance in biogeographic analysis. Biogeographic patterns determined from our analysis of all published data on distribution of Ordovician genera indicate Early Ordovician sponge faunas have relatively low diversity and are completely dominated by demosponges. Early Ordovician (Ibexian) faunas are characterized by the widespread co-occurrence of Archaeoscyphia and the problematic Calathium. This association is commonly found in biohermal structures. Middle Ordovician faunas show an increase in diversity, and two broad associations are differentiated: Appalachian faunas (including Southern China and the Argentine Precordillera) and Great Basin faunas.
Late Ordovician faunas show important changes in diversity and provincialism. Hexactinellid and calcareous sponges became important and new demosponge families appeared. Four Mohawkian-Cincinnatian associations are recognized here, including: 1) Midcontinent faunas; 2) Baltic faunas; 3) New South Wales faunas; and 4) Western North American (California and Alaska) faunas. However, two separate biogeographic associations are differentiated based on faunal differences. These are a Pacific association (western North American and New South Wales) and an Atlantic association (Midcontinent Laurentia and Baltica).
Distribution of sponge genera and migration patterns are utilized to consider paleogeographic dispositions of the different continental plates, climatic features, and oceanic currents. Such an analysis points to close paleogeographic affinities between the Argentine Precordillera and Laurentian Appalachian faunas. However, significant endemicity and the occurrence of extra-Laurentian genera suggest a relative isolation of the Precordillera terrane during the Late Ibexian-Whiterockian. The study also shows a faunal migration from the Appalachian region to South China during the Middle Ordovician and the migration of faunas from Baltica to Laurentia in the Late Ordovician. The occurrence of Laurentian migrants in New South Wales during the Late Ordovician could be related to inferred oceanic current circulation between these two areas, although other paleogeographic features may be involved.
S ponges include the most primitive multicellular organisms and have a record commencing in the Late Precambrian. All major sponge groups are repre-sented as Cambrian fossils. Ordovician sponges are only moderately well known when compared with other groups of fossils, though they are known from most continents and 275 species have been described. This relatively limited known diversity may be due in part to the scarcity of taxonomic studies and in part to the phylum's conservative history. Ordovician diversification is marked by the ex-tensive radiation of some groups, whereas others remained limited and little diversified. Even though several sponge groups are moderately well known in the Ordovician, their origins from Cambrian ele-ments are obscure, largely because of significant breaks between Middle Cambrian and Lower Ordovician sponge records. Ordovician sponge distribution is known in vary-ing degrees of detail for different areas. The limited stratigraphic information given in the older literature makes the assignment of some ranges very difficult. In addition, a particular problem is the correlation of northern European sponge associations, which are largely known only from Quaternary glacial erratic boulders. Microfossils found in the associated matrix indicate that most are of early Ashgill age (Hacht and Rhebergen 1997; Rhebergen et al. 2001). However, some limestone boulders in the same deposits are mid to late Caradoc in age. As a consequence, accurate stratigraphic positions cannot be determined for this probable carbonate platform association, limiting the detailed analysis. We adopted two alternatives: a system in which the first appearances (FAD) of European sponges are allocated a range from time slice TS.5c to the last appearances (LAD) during TS.6a. This system po-tentially overestimates the real diversity values. The other alternative is to exclude the European associa-tion from the database, which considerably under-estimates the values. The resulting diversity curves, constructed using normalized diversity measures, are shown in figure 12.1. The curve showing relative dis-tribution for the time intervals for the major sponge groups was calculated based on the first of the alter-natives data (figure 12.2). The sponge database has been incorporated into the database managed by Arnold I. Miller (University of Cincinnati).
A modular organization confers many ecological advantages in reef-building settings. The Archaeocyatha, an early Cambrian group of calcified sponges, were the first skeletal metazoans to develop a modular habit and to be associated with reefs. Modular archaeocyaths survived beyond the extinction of most solitary forms, suggesting a relative immunity to extinction presumably on account of ecological response. The solitary nature of many archaeocyaths explains many of their previously supposed apomorphies. -from Authors
ABSTRACT The Eridanos fluvio-deltaic system, draining most of north-western Europe, developed during the Late Cenozoic as a result of simultaneous uplift of the Fennoscandian shield and accelerated subsidence in the North Sea Basin. This seismo-stratigraphic study aims to reconstruct the large-scale depositional architecture of the deltaic portion of the basin fill and relate it to external controls. A total of 27 units have been recognized. They comprise over 62×103 km3 in the Southern North Sea Basin alone, and have an average delta surface area of 28×103 km2, which suggests that the size of the drainage area was about 1.1×106 km2. Water depth in the depocentre is seen to decrease systematically over time. This trend is interrupted by a deepening phase between 6.5 and 4.5 Ma that can be correlated with the simultaneous occurrence of increased uplift of the Fennoscandian shield, increased subsidence of the Southern North Sea Basin, and a long-term eustatic highstand. All these observations point to a tectonic control on long-term average rates of accommodation and supply. Controls on short-term variations are inferred from variations in rates of sediment supply and bifurcation of the delta channel network. Both rates were initially low under warm, moist, relatively stable climate conditions. The straight wave-dominated delta front gradually developed into a lobate fluvial-dominated delta front. Two high-amplitude sea-level falls affected the Pliocene units, which are characterized by widespread delta-front failures. Changes in relative sea level and climate became more frequent from the late Pliocene onward, as the system experienced the effects of glacial–interglacial transitions. Peaks in sedimentation and bifurcation rates were coeval with cold (glacial) conditions. The positive correlation between rates of supply and bifurcation on the one hand, and climate proxies (pollen and δ18O records) on the other hand is highly significant. The evidence presented in this study convincingly demonstrates the control of climate on time-averaged sediment supply and channel-network characteristics, despite the expected nonuniformity and time lags in system response. The presence of a clearly discernible climate signal in time-averaged sediment supply illustrates the usefulness of integrated seismo-stratigraphic studies for basin-wide analysis of delta evolution on geological time scales.
Lithistid sponges are an important component of many Ordovician faunas and form some of the earliest reef communities of the Paleozoic Evolutionary Fauna, yet they have received relatively little attention. Fossiliferous strata with abundant sponges occur at many horizons and localities within the Lower and Middle Ordovician Pogonip Group of the Great Basin. One of the best-known occurrences is near the middle of the Antelope Valley Limestone in central Nevada, where sponge-bearing units occur within the lower Whiterockian Stage. Lithistid sponges and associated species were also examined within the Shingle Limestone of eastern Nevada, where the sponge communities occur as small reef mounds about 55 m below the top of the formation in beds corresponding to the uppermost Ibexian Series. To acquire a more regional view of the environments and faunas present in Nevada during the Ordovician, contemporaneous units in the Pogonip Group were examined elsewhere in Nevada and in the Ibex area of western Utah.
The general depositional setting of the Great Basin at this time was that of a carbonate ramp in the upper Ibexian developing into a carbonate platform during the Whiterockian. The sponge mounds in the Shingle Limestone, forming in water 1 to 15 m deep, may have participated in this development by interfering with water circulation along the outer portion of the carbonate ramp. The mounds are several meters in size and are often stacked, with grainstone-filled channels cutting through the mounds in several places. The mounds themselves have muddy cores and are flanked by well-sorted echinoderm grainstones. The reef trend extends for nearly two kilometers before being faulted out, and was probably even more extensive during its development. Besides sponges, the assemblage contains abundant calcareous algae and cyanobacteria, as well as trilobites, brachiopods, echinoderms, and other groups characteristic of the Paleozoic Fauna. These mounds are very similar to those found in the Wahwah and Juab Limestones of the Pogonip Group, suggesting that these latter buildups also formed in very shallow water.
The younger sponge communities in the Antelope Valley Limestone at Ikes Canyon probably formed in about 20 to 40 m of water along the uppermost slope of a carbonate platform. The sediments consist of interbedded wackestones, mudstones, and shales with occasional hardgrounds. Some of the sponges have root tufts, indicating that they settled on soft substrate. Besides lithistid and heteractinid sponges, the unit contains a diverse assemblage of metazoans and occasional receptaculitids and other algae.
Estimates of relative species abundances were calculated for random samples from the sponge-bearing units. The Shannon Diversity Index for the Antelope Valley Limestone fauna is 3.24 with an equitability value of 0.68, while the Shingle Limestone community has values of 3.44 and 0.74, respectively. Thus, not only were both communities diverse, but the species within each community were comparable in abundance. These values compare favorably with those of modern marine communities, especially when the effects of preservational bias are considered, and suggest that well-integrated ecological communities with substantial amounts of biotic interaction had evolved by the Middle Ordovician.
The sponge communities of the Antelope Valley Limestone and Shingle Limestone are thus quite similar, yet only in the latter did the organisms build bioherms. The sucession of communities within these buildups indicate that the greater abundance of cyanobacteria and calcareous algae in the Shingle Limestone probably helped to bind and stabilize the sediment, creating a firm substrate. This was then settled by cylindrical and branching lithistid sponges and other species that not only acted as bafflers, causing mud to accumulate, but produced additional sediment when they died. As the mounds grew, the communities diversified, with cup-shaped lithistids becoming dominant as the bioherms approached sea level. The importance of calcareous algae and cyanobacteria in determining whether mud mounds developed in this depositional setting suggests that factors controlling the distribution of algae, primarily substrate, turbidity, and water depth, also control the distribution of mud mounds. This is probably true not only for these sponge mounds, but for mud mounds throughout the Paleozoic.
Abundant hexactinellid and lithistid sponges occur in Ludlow (Silurian) platform slope strata of the Cape Phillips Formation, Canadian Arctic Islands. The following are new: in the Anthaspidellidae Miller, 1889, Rhodesispongia simplex n. gen and sp., Climacospongia snowblindella n. sp., and Climacospongia undulata n. sp.; in the Sphaerocladina Schrammen, 1910, Caryospongia tuberosa n. sp.; in the Hexactinellida Schmidt, 1870, Malumispongium? cornwallisi n. sp., of the monospecific superfamily Malumispongium Rigby, 1967. Other previously described sponges recognized in the study assemblage include Hindia sphaeroidalis Duncan, 1879 and Caryospongia juglans Rauff, 1894. These occur in association with abundant Archaeoscyphia sp. in thin, laterally extensive lithistid sponge biostromes which throve on platform slope sediments during the Late Silurian. Based on the dominant dendroclone forms in the skeletons of these and other genera, Anthaspidellidae can be divided informally into two groups: (i) the polyclonids, possessing complex skeletons built of variable dendroclones types, including I- , X- , and Y-shape dendroclones, and accessory spicules, including rhizoclones, chiastoclones, and monactines; and (ii) the monoclonids, possessing simpler parenchymal canal systems and skeletons constructed mainly of I-shaped (amphiarborescent) dendroclones and lacking accessory spicules. The classification, based chiefly on skeletal architecture, suggests two main evolutionary trends in the family. The monoclonids generally maintained structural simplicity, inherited from the probable ancestral monactinellid, whereas the polyclonids evolved a varied and complex skeletal architecture that was more successful.
One of the most diverse Late Ordovician sponge faunas known, and certainly the most diverse Ordovician sponge fauna known from Australasia, is from the Malongulli Formation of central New South Wales. These fossils occur in blocks within breccias embedded in the graptolitic and spiculitic siltstone succession. They occur at three major localities along the Belubula River 200 km west of Sydney where a total of 44 species (39 new) represent (26 new) of demosponges, calcareous sponges and hexactinellid sponges. Lithistid demosponges are the most common and diverse. The New South Wales sponge assemblage provides a unique record of deeper-water carbonate environments at the margin of an 'island-arc' shelf sequence and certainly suggests that deeper-water assemblages are far more diverse than currently documented. -from Authors
The Silurian sponge fauna described here from Baille-Hamilton and Cornwalls islands is one of the most diverse known from North America. The fossils are from Wenlockian-Ludlovian age deposits of the Cape Phillips Formation, which accumulated on the slope and in the basin of the Franklinian Geosyncline. The Cape Phillips Formation consists of interbedded shale, calcareous shale, and limestone, and most of the fossils are found in allochthonous limestone beds, mass movements having transported much of the fossil material downslope. Silurian sponges described here are representative of a distinct sponge biogeographic province that occurs north and northwest of a line along the Transcontinental Arch and contrasts with one to the southeast where sphaerocladine lithistid and heteractinid calcareous sponges are dominant. -from Authors
The oldest fossil lithistid sponges (Demospongiae, ‘lithistids’) are known from the Cambrian. Lithistids are very common in the fossil record since the Ordovician, but their distribution is very punctuated: large faunas of lithistid demosponges are known from the Ordovician, Silurian, Devonian, Permian, Upper Jurassic, Upper Cretaceous and the Eocene. The Paleozoic faunas are dominated by Orchocladina (extinct in the Permian) and Sphaerocladina (continuing up to the Recent); Mesozoic faunas are dominated by Tetracladina and Rhizomorina, which are also common in the Tertiary and Recent lithistid faunas. Phylogeny of fossil lithistids, especially relationships between the Paleozoic and later forms are still poorly understood, partly because only desmas are usually preserved in the fossil state, thus no microscleres or rarely ectosomal spicules are known. These megascleres are crucial diagnostic characters in Recent lithistid taxonomy. Given their good fossilization potential this fauna is diverse, with 13 suborders, 34 families and over 200 genera discussed in the present work, although many more nominal genera of fossil lithistids are known.
Late Cenozoic deposits in the North-West European Basin with a typical gravel composition, called the Baltic Gravel Assemblage, have been studied from the literature and in the field. The gravel indicates a provenance from the Fennoscandian area and is characterized by a high proportion of translucent quartz and the presence of silicified Palaeozoic sediments. From the available data it is concluded that one river system, called the Baltic River System, was the transporting agent of the Miocene, Pliocene and Pleistocene deposits containing the Baltic Gravel Assemblage. In the Miocene the sedimentation was to the S and W. From the Late Miocene into the Early Pleistocene the main transport was towards the W. The Baltic River System was destroyed by the inland-ice of the Menapian glacial.-Author
Abundant hexactinellid and lithistid sponges occur in Ludlow (Silurian) platform slope strata of the Cape Phillips Formation, Canadian Arctic Islands. The following are new: in the Anthaspidellidae Miller, 1889, rhodesispongia simplex n.gen and sp., Climacospongia snowblindella n.sp., and Climacospongia undulata n.sp.; in the Sphaerocladina Schrammen, 1910, Caryospongia tuberosa n.sp.; in the Hexactinellida Schmidt, 1870 Malumispongium? cornwallisi n.sp., of the monospecific superfamily Malumispongium Rigby, 1967. Other previously described sponges recognized in the study assemblage include Hindia sphaeroidalis Duncan, 1879 and Caryospongia juglans Rauff, 1894. These occur in association with abundant Archaeoscyphia sp. in thin, laterally extensive lithistid sponge biostromes which throve on platform slope sediments during the Late Silurian. Based on the dominant dendroclone forms in the skeletons of these and other genera, Anthaspidellidae can be divided informally into two groups: (i) the polyclonids, possessing complex skeletons built of variable dendroclones types, including I-, X-, and Y-shape dendroclones, and accessory spicules, including rhizoclones, chiastoclones, and monactines; and (ii) the monoclonids, possessing simpler parenchymal canal systems and skeletons constructed mainly of I-shaped (amphiarborescent) dendroclones and lacking accessory spicules. The classification, based chiefly on skeletal architecture, suggests two main evolutionary trends in the family. The monoclonids generally maintained structural simplicity, inherited from the probable ancestral monactinellid, whereas the polyclonids evolved a varied and complex skeletal architecture that was more successful.
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Die fossile Fauna der schlesischen Diluvialgeschiebe von Sadewitz bei Oels in Nieder-Schlesien
- Roemer