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A remarkable new Middle Sandbian (Ordovician) hexactinellid sponge in Baltic erratics
Abstract and Figures
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.
... The fossil record of sponges is incomplete, complex, and increasingly important for understanding early animal evolution . One of the key indicators of our limited state of knowledge in Ordovician and Silurian sponge diversification is the large proportion of highly problematic taxa being described (e.g., Botting & Rhebergen 2011;Botting & Zhang 2013). This implies long phylogenetic gaps between known taxa, which severely hinders a full understanding of how the diversification occurred. ...
... The new species further supports previous suggestions that dramatic diversification was occurring among sponges in shallow-water habitats during the Ordovician (Botting 2005;Muir et al. 2013), and highlights the limitations of our current knowledge of sponge evolution within these groups. The differences between the new species and any potential ancestors or close relatives, as with Amphispongia (see Botting 2004), Haljalospongia (Botting & Rhebergen 2011), and others (e.g., Rigby & Webby 1988), implies that we currently have no knowledge of large parts of sponge evolution from this interval, especially regarding the stem group (and perhaps early crown group) of Hexactinellida. If we are correct in interpreting the specimen as a plausible crown-group rossellid allied to Caulophacus, or at least crown-group lyssacinosidan, then this builds on other results from China ) that exacerbate the problems of the missing record of sponge diversification, as it implies that many modern groups must have arisen cryptically, and left almost no trace in the fossil record. ...
The Pentland Hills sponge fauna (Llandovery, Telychian) consists of an unusual, aberrant assemblage, but of low diversity. A new specimen of a unique sponge, Eoghanospongia carlinslowpensis gen. et sp. nov., is described from the classic locality of R82. The mushroom-shaped, probably stalked body (peduncle attachment to body not exposed) resembles that of some living rossellids, especially Caulophacus . The sponge also shows prominent projecting pentactins and monaxon derivatives – a spicule type diagnostic of the Rossellidae among living taxa, albeit in a robust form not described from extant sponges, including Caulophacus or other pedunculate lyssacinosidans. Certain attribution to the Rossellidae is not possible from the single specimen, largely because of weak preservation of the primary spicule skeleton, but no other fossil or modern sponges show any significant similarity to it. Although similarly early relatives of the Rossellidae have recently been described from elsewhere, the new sponge is even more unexpected in being from a shallow-water environment, making the absence of rossellids through the rest of the Palaeozoic much more problematic.
... Later Ordovician assemblages of non-lithistid sponges are relatively rare (Figs. 3, 5), but several diverse faunas are now being uncovered in Avalonia (Muir and Botting, 2015) and South China (Li et al., 2015;Botting et al., 2017), together with the well-known Malongulli fauna of Australia (Rigby and Webby, 1988), and isolated articulated sponges from other areas (e.g., Beecher, 1889;Botting and Rhebergen, 2011). The most commonly recurring genera in this time interval are the Cambrian survivors Cyathophycus, Choiaella and Pirania ...
Non-lithistid spiculate sponges were major components of Cambrian and Ordovician communities, but currently there is limited understanding of how sponge distribution patterns developed during the Cambrian Explosion and the Great Ordovician Biodiversification Event (GOBE). We compiled data on Cambrian and Ordovician sponge occurrences from the literature and our own research. Sponge genera were widely distributed during the Cambrian Period, with broadly consistent assemblages between palaeocontinents, from different palaeolatitudes, and over a timespan of approximately 25 Myr. Sponges exhibited a marked shift from cosmopolitan to highly endemic genera during the Tremadoc and later Ordovician. Surviving Cambrian genera continued to be very widespread, but taxa that originated during the Ordovician had much narrower geographic and environmental ranges, perhaps related to the origination of modern-style, patchy ecology. The cosmopolitanism of the Cambrian faunas is suggestive of rapid dispersal of recently evolved groups without competition from pre-existing sponges, suggesting that sponges had not diversified during the Precambrian. In contrast, the Ordovician record indicates local diversification against a complex ecological backdrop. Based on the ecological record of sponges, the Cambrian Explosion was a fundamentally different process to the GOBE.
... Fossils consist of seafloor debris because trilobites, crinoids and machaeridia are nearly always disintegrated. Post mortem overgrowth of fossils and marks of etching and burrowing organisms, which are abundant in type 2, are less frequent in type 1. Trace fossils (e.g., Chondrites) and other signs of bioturbation are common but do not occur as strongly concentrated as in type 2. All this suggests that highenergy conditions, with periodical minor mud flows burying the seafloor, prevailed (Botting & Rhebergen 2011). There are no signs of winnowing. ...
This study seeks to describe ‘baksteenkalk’, an erratic silicified bioclastic carbonate of the Upper Sandbian from the eastern part of the Netherlands. To date, baksteenkalk has received little attention among palaeontologists. This is to be regretted on two grounds. First, baksteenkalk contains a varied fossil flora and fauna comprising many species, several of which are not or only rarely found in coeval rocks. Second, owing to a complicated silicification process, fossils, in particular algae, have preserved exceptional anatomical details. The primary aim of this study is to arouse the interest of professional, in particular, Estonian, palaeontologists in baksteenkalk. Based on lithology and fossil assemblage (most conspicuously, with regard to the algal flora), two basic types of baksteenkalk are distinguished. A list of species, differentiated for both types, is provided. It is argued that baksteenkalk reflects the ecology of a shallow, subtropical, epicontinental sea. The distribution of erratics, facies and fossil content point to an origin within the North Estonian Confacies Belt, probably west of Estonia. Baksteenkalk survived as an erratic because it was already silicified at its place of origin. A potential source of silica may have been Upper Ordovician bentonite layers. The causes and mechanisms of the silicification process which gave shape to baksteenkalk are not yet understood, however. The palaeontology of baksteenkalk is compared with that of two other erratic Sandbian silicified carbonates of Baltic origin: German Backsteinkalk and ‘Lavender-blue Hornstein’.
... Spectacular fossils such as Xenusion auerswaldae Pompeckj, 1927 (probably originating from the Lower Cambrian Kalmarsund Sandstone of southern Sweden: Jaeger and Martinsson 1967; Hauschke and Kretschmer 2015) have been found in glacial erratics, and new taxa are even now being described from glacial(?) erratics originating from the Palaeozoic succession of Scandinavia (e.g. Botting and Rhebergen 2011). ...
Ordovician and Silurian glacial erratics of the Laerheide area (Lower Saxony, north-western Germany) bear well-preserved graptolites. The faunas provide important information on the origin and transport direction of the sediments preserved in a kame, representing the Drenthe stadial of the Saalian glaciation. The faunas even include species not commonly encountered in the successions of mainland Sweden, from where the erratics presumably originated. The most common graptolites are from Upper Ordovician (Sandbian to Katian) limestones and from Katian black shales. More common, however, are greenish limestones, sand- and siltstones, often combined in the term ‘Gru¨nlich-Graues Graptolithengestein’, in which upper Wenlock to Ludlow (upper Silurian) graptolites are common.
... Occasionally, these faunas may include thin-walled taxa in addition to the normal assemblage (Rigby, 1971). Many of the described non-lithistid sponges are of isolated occurrences, often involving single species (Beecher, 1889;Botting and Rhebergen, 2011;Botting and Zhang, 2013;Rigby et al., 1981), although other undescribed species occur in at least some of these faunas (pers. obs.). ...
Sponges form a significant component of the diversity of the Tremadocian (Early Ordovician) Fezouata Biota in the Anti-Atlas of Morocco, but are distributed intermittently and have so far received only limited attention. New material reveals a high diversity of undescribed taxa (a total of at least 27 species), including numerous representatives of protomonaxonid groups such as the Leptomitidae, Piraniidae and “Choiidae”. Some of these taxa show unusually complex skeletal architecture, and represent derived variations of their lineages relative to those in the Cambrian Burgess Shale-type faunas, although most species are assignable to described families. Reticulosan sponges are rare in the Fezouata Biota, and usually fragmentary. The palaeoecology of the sponge fauna is unusual, with most species known only from single sites (frequently single bedding planes), but are often abundant where they occur. It is very rare for two species of sponge to be found on the same slab, or at precisely the same horizon. With some species of protomonaxonid, particular bedding planes are crowded with individuals of species that are rarely, if ever, found isolated. Only two species (Pirania auraeum Botting, 2007 and an undescribed hazeliid) are found at numerous levels, and these are not known from crowded assemblages; this may relate to differences in reproductive strategy. The dense, usually monospecific populations can best be explained through repeated colonisation events in a frequently hostile environment, rather than representing a range of different stable communities.
... Heteractenigma falls into a growing number of Cambrian-Silurian hexactinellid-like sponges (e.g. Botting 2004, Wu et al. 2005, Botting & Rhebergen 2011 that are difficult to assign to previously known groups, or to place into a phylogenetic framework. The continued discovery of new, morphologically unique taxa also implies that the known record is very incomplete even at family level. ...
A new sponge, Heteractenigma yui gen. et sp. nov., is described from the Corymbograptus deflexus Biozone (Floian) of the Ningkuo Formation in western Zhejiang Province, China. The tall, thin-walled species possessed unique features including a combination of robust longitudinal monaxons, fine sub-transverse monaxons, and minute hexactine and heteractine body wall spicules. The heteractine spicules have five rays in one plane, and closely resemble those of the problematic Burgess Shale sponge Petaloptyon danei. The monaxon organisation and body form, in contrast, accord with those of leptomitids. Although it is not currently possible to establish the precise relationships of this species, due to the major differences from other known taxa, it includes features characteristic of heteractinids, reticulosans and leptomitid protomonaxonids. This may indicate that the species belongs to the stem-group of either Porifera or Silicea. The unique architecture of the species also confirms that there remain large gaps in our knowledge of early sponges, even at high taxonomic level.
... (M'Coy, 1850;Rhebergen et al. 2001), which most probably should be re-interpreted as an indeterminate root-tuft. Recently,Botting & Rhebergen (2011) described Haljalaspongia inaudita, collected in the westernmost part of Germany; this is the first hexactinellid sponge known from the Ordovician (Sandbian; Upper Ordovician) erratic blocks of silicified limestone originating in Baltic region. ...
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.
... Reworked fossils can occur either as clasts in modern sedimentary environments or reinterred in younger sedimentary rocks. The possible range of reworked taxa is as diverse as the fossil record itself, but common macrofossils may include sponges, brachiopods and trilobites (Ulrich, 1927;Botting and Rhebergen, 2011), echinoderms (e.g., Donovan, 2012;Donovan and Pawson, 2013), molluscs (such as Chubb, 1971, p. 195, pl. 40, fig. ...
The Late Oligocene oyster Hyotissa antiguensis (Brown) is locally common in the Antigua Formation of Hughes Point, eastern Antigua, Lesser Antilles; it was not commonly bored at that time. Its valves and shells are robust, and reworked into the shallow water near-shore environment in Antigua; it could potentially be incorporated into younger rocks. Its neoichnology includes clues that would facilitate identification of these oysters as reworked fossils. The suite of modern borings found in these specimens includes common Caulostrepsis taeniola Clarke, Gastrochaenolites isp. cf. G. turbinatus Kelly and Bromley and Entobia isp., and rare Oichnus simplex Bromley and Rogerella? isp. The latter three taxa are limited to oyster shell substrates. Of the common ichnotaxa, Caulostrepsis and Gastrochaenolites are particularly prominent in limestone clasts and limestone cemented to oyster shells, which would be an indicator of reworking if found in a post-Oligocene lithified deposit. Caulostrepsis and Gastrochaenolites are relatively less common in oyster shells and valves, and in many specimens are seen to terminate against the shell. Entobia is the only common boring limited to the shell substrate. The fidelity of preservation of modern borings is also superior in limestone clasts. This suite of borings is comparable with those found in the Neogene of the Antillean region.
An abundant sponge collection from basal Caradoc mudstones of the Llanfawr Quarries, Llandrindod, Powys is the most diverse fine‐sediment sponge fauna recorded from the British Palaeozoic. It is dominated by reticulosans, but includes some lyssakids and demosponges. Eight new genera and 11 species are erected and one new family proposed: Xylochos palindromica gen. et sp. nov., Pirania llanfawrensis sp. nov., Heminectere minima gen. et sp. nov., Asthenospongia cambria sp. nov. (Asthenospongiidae nov.), Acutipuerilis spinosus gen. et sp. nov., Granulispongia obscura gen. et sp. nov., Hemidiagoniella tenax gen. et sp. nov., H. caseus sp. nov., Solusrectus rosetta gen. et sp. nov., Cyathophycus loydelli sp. nov., and Dilatispongia tumidus gen. et sp. nov. Some specimens are preserved with organic or pyritic films representing soft tissue and most have high‐fidelity preservation of spicule external moulds, although iron oxide spicule replacement obscures details prior to cleaning. D. tumidus shows a thickened wall with specialised dermalia, while retaining some degree of quadruling and represents a lineage that may have given rise to the brachiospongioids. Others, such as S. rosetta, appear close to the ancestral state of the modern Lyssacinosida. The hex‐actinellid taxa are interpreted in a reasoned phylogenetic framework that stresses the importance of root tuft structure, the presence of zero‐order spicules and the number and nature of spicule layers. It also attempts to link extant and Lower Palaeozoic palaeontological classifications. The parallel evolutionary development of thick walls and strong attachment structures is interpreted as being due to an offshore‐onshore expansion, contrary to many other groups.
Fluvial deposits of Miocene to Early Pleistocene age in Germany and the Netherlands were laid down in the delta of the Eridanos River System, but the exact provenance of this material continues to be a subject of discussion. The aim of the present study is twofold. Firstly, a comparison of Ordovician sponges in these deposits with those from northern Estonia and the St Petersburg region (Russia) demonstrates that these erratics originated from the drainage area of the Pra Neva, a tributary of the Eridanos. Secondly, the importance of Late Ordovician silicified boulders, which yield forms of preservation that are unknown in comparable fossils, preserved in situ, is outlined. Some recommendations for future studies are made.
Platform-margin strata of the Avalanche Lake area in the Mackenzie Mountains of the Northwest Territories have yielded a fabulous diversity of silicified and non-silicified fossils, including Silurian (Wenlock) sponges. The demosponge assemblages comprise the epipolasid Belemnospongia epiradiatus n. sp.; the rhizomorine Haplistion(?) frustrum Rigby and Chatterton, and an abundant and diverse group of anthaspidellid orchocladines, including Archaeoscyphia annulata(?) (Rigby), Archaeoscyphia attenuata de Freitas, Archaeoscyphia alternata de Freitas, Archaeoscyphia gislei de Freitas, Archaeoscyphia rectilinearis de Freitas, and Archaeoscyphia scalaria de Freitas, along with Somersetella(?) digitata Rigby and Dixon. The orchocladines are also represented in the sponge fauna by Patellispongia alternata Rigby and Chatterton, Patellispongia mackenziensis n. sp.,Patellispongia rugosa n. sp., Patellispongia(?) attenuata n. sp., and one coarsely canalled anthaspidellid of uncertain generic assignment. The astylospongiid sphaerocladines in the collection are: Astylospongia praemorsa(?) (Goldfuss), Palaeomanon cratera (Roemer), and Carpospongia castanea (Roemer), and a small, thin-walled, globe-shaped sponge of uncertain generic assignment. The sole tricranocladine lithistid is Hindia sphaeroidalis Duncan. Small initially calcareous and more rare permosphinctid demosponges respectively comprise the questionable aporate aphrosalpingid Palaeoschada(?) sp., and a porate cryptocoeliid of uncertain generic affinity. Unfortunately, some skeletal details have been obscured by coarse siliceous replacement so that details of microstructure are uncertain in some species. Gross skeletal fabrics, canal patterns, and growth forms, however, are commonly sufficient to identify the species. This is one of the most diverse Silurian assemblages known from North America, and certainly the most diverse assemblage from northwest of the Transcontinental Arch in the western United States and northwestern Canada. Diverse assemblages dominated by orchocladine lithistid sponges characterize faunas from northwest of the arch, while those from southeast of the arch are dominated by sphaerocladine and tricranocladine lithistids and astraeosponge heteractinids.
Mit 12 Abbildungen und 9 Tafeln Zusammenfassung Es konnte erstmals nachgewiesen werden, daß Mikroskleren vom Typus Hemihexaster, Hexaster, Spir-und Echinhexaster, sowie Clavule und Paraclavule bereits ab dem tiefen Ordovizium auftreten. Sie sind ein Beleg hiefür, daß die Hexasterophora spätestens mit Beginn des Ordoviziums einsetzten; sehr wahr-scheinlich schon im Laufe des Kambriums entstanden sind. Skleren vom Typus Hemi-und Amphidisk sind aus oberkambrischen Schichtfol-gen überliefert, was sicherstellt, daß deren Entstehung im Laufe des Kambriums vonstatten ging. Die hier vorgestellten Neuergebnisse ermöglichen es, an die Systematik rezenter Hexactinellida anzuschließen und aufzuzeigen, daß die Vertreter beider Unterklas-sen sich im Altpaläozoikum, unabhängig von den Protospongioidea und Brachio-spongioidea, entfaltet haben. Daraus muß gefolgert werden, daß sich die Hexactinel-lida in mehreren eigenständigen Linien entwickelt haben, und somit polyphyletisch entstanden sind. Summary For the first time it could be proved that microscleres of the hemihexaster-, hexaster-, spir-and echinhexaster-type, as well as clavules and paraclavules occurred already in the Lower Ordovician. They are a mark that the Hexasterophora appeared not later than with the beginning of the Ordovician, very probably they were developed in the Cambrian. Scleres of the hemi-and amphidisc-type are known from Upper Cambrian carbonate sequences, which proves that their origin was in the Cambrian. The new results, introduced in this paper, allow to attach to the systematics of recent Hexactinellida and to show that the members of both subclasses evolved in the early Paleozoic without dependence on the Protospongioidea and Brachiospon-gioidea. This leads to the conclusion that the Hexactinellida developed in several independent lineages; this means that they are of polyphyletic origin.
Cambrian spicular sponge faunas are dominated by a distinctive assemblage of demosponges and hexactinellids that are known from Burgess shale-type faunas worldwide. Most of these are previously unknown outside the Lower–Middle Cambrian (and perhaps Tremadoc) and have no obvious close relatives in later sequences. This paper describes examples of Choia sp., Pirania auraeum sp. nov. and Hamptonia christi sp. nov. from the Arenig of Morocco, associated with isolated hexactinellid spicules. A summary of the stratigraphic ranges of the major Cambrian sponge lineages is provided. These indicate an environmental contrast in the Lower Palaeozoic evolution of hexactinellids and non-lithistid demosponges, with demosponges probably undergoing cryptic diversification in nearshore environments during the Upper Ordovician.RésuméLes éponges siliceuses cambriennes se distinguent par des assemblages dominés par les démosponges et les hexactinellidés. Ces assemblages sont connus dans les faunes de type Burgess partout dans le monde. La plupart des éponges étaient auparavant inconnues après le Cambrien inférieur–moyen, et peut-être le Trémadocien, et n’ont pas de formes apparentées dans des séries plus jeunes. Cet article décrit quelques exemples de Choia sp., Pirania auraeum sp. nov. et Hamptonia christi sp. nov. dans l’Arenig du Maroc, en association avec des spicules d’hexactinellidés isolées. Un résumé de l’extension stratigraphique des plus importantes lignées d’éponges est fourni. Ceci indique un contraste environnemental dans l’évolution des hexactinellidés et des démosponges non-lithistidés au Paléozoïque. Les démosponges ont probablement subit une diversification cryptique dans des environnements sub-littoraux à l’Ordovicien supérieur.
A diverse fauna of three-dimensionally preserved sponges is described from nearshore volcanogenic sandstones near Llandrindod, Mid-Wales. The fauna was preserved through early marginal silicification, in rare examples with silicification of soft tissue, and includes aspicular and spicular demosponges, hexactinellids, and a heteractinid. The fauna is largely endemic, with the following new species, genera and families erected: Onerosiconcha gregalia gen. et sp. nov., Ordinisabulo quadragintaforma gen. et sp. nov., Miritubus erinaceus gen. et sp. nov., Vadosifistula milvus gen. et sp. nov., Polycornua trescelestus gen. et sp. nov. (Pseudolanciculidae fam. nov.), P. entropus sp. nov., Palaeocallyoides improbabilis gen. et sp. nov., Reticulicymbalum tres gen. et sp. nov., Triactinella rigbyi gen. et sp. nov. (Triactinellidae, fam. nov.), Spissiparies minuta gen. nov., Brevicirrus arenaceus gen. et sp. nov., Pyritonema scopula sp. nov. (Pyritonemidae, fam. nov.) and Microastraeum tenuis gen. et sp. nov. In addition, Pseudolancicula Webby and Trotter is recorded outside Australia for the first time, although only as isolated spicules. The environment represented by this fauna has not previously yielded articulated sponges, and thus the apparent endemism may be misleading.
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.
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