Bryozoan-cnidarian mutualism triggered a new strategy for greater resource exploitation as early as the Late Silurian.

Bryozoans were common benthic invertebrates in the Silurian seas. The large biodiversity among Silurian benthic organisms prompted diversified interactions, and as a result bryozoans hosted many other organisms as symbionts. Here we analyse the cystoporate bryozoan Fistulipora przhidolensis and unidentified trepostomes intergrown with auloporid tabulate corals and putative hydrozoans. The material comes from the uppermost Prídolí Series (Late Silurian) of the Sõrve Peninsula, Saaremaa, Estonia. Our analysis shows that the interaction was beneficial for both organisms-cnidarians benefited from feeding currents created by the host bryozoan, while the latter benefited from the protection from predators by cnidae, it can thus be classified as mutualism. Such associations are common in modern seas. The analysed organisms are typically encrusting when the symbiosis is absent, when intergrown they display erect, branching morphologies, raised over the substratum, thus exploiting a higher suspension-feeding tier. While similar associations were known from the Devonian, we demonstrate that this novel ecological strategy for greater resource exploitation started as early as the latest Silurian.

Functional consequences of Palaeozoic reef collapse.

Biogenic reefs have been hotspots of biodiversity and evolutionary novelty throughout the Phanerozoic. The largest reef systems in Earth's history occurred in the Devonian period, but collapsed during the Late Devonian Mass Extinction. However, the consequences for the functional diversity of Palaeozoic reefs have received little attention. Here, we examine changes in the functional diversity of tabulate coral assemblages over a 35 million year period from the middle Devonian to the Carboniferous, straddling the multiphase extinction event to identify the causes and ecological consequences of the extinction for tabulate corals. By examining five key morphological traits, we show a divergent response of taxonomic and functional diversity to the mass extinction: taxonomic richness peaked during the Givetian (~ 388-383 Ma) and coincided with peak reef building, but functional diversity was only moderate because many species had very similar trait combinations. The collapse of taxonomic diversity and reef building in the late Devonian had minimal impact on functional richness of coral assemblages. However, non-random shifts towards species with larger corallites and lower colony integration suggest a shift from photosymbiotic to asymbiotic taxa associated over the study period. Our results suggest that the collapse of the huge Devonian reef systems was correlated with a breakdown of photosymbiosis and extinction of photosymbiotic tabulate coral taxa. Despite the appearance of new tabulate coral species over the next 35 million years, the extinction of taxa with photosymbiotic traits had long-lasting consequences for reef building and, by extension, shallow marine ecosystems in the Palaeozoic.

Early development and coloniality in Oligophylloides from the Devonian of Morocco-Are Heterocorallia Palaeozoic octocorals?

Heterocorals represent an enigmatic group of Palaeozoic corals, known from relatively short time intervals in the Devonian and Carboniferous periods. The major differences between Heterocorallia and other Palaeozoic corals are the lack of an external theca (epitheca), lack of calices and the presence of dichotomously dividing septa-like structures. Heterocoral skeleton was presumably externally covered by the soft tissue and each branch of their skeleton has, until now, been regarded as a corallite-a skeleton of a single polyp. We investigated upper Famennian Oligophylloides from Morocco, focussing on branching processes, wall structure, previously poorly known initial growth stages and the growing tip, described here for the first time. We demonstrate that Oligophylloides shows a unique colony development not known in any group of anthozoans possessing a septate-like architecture and suggest that the previously postulated homology between true septa in hexa- and rugose corals on one hand, and Oligophylloides on the other, must be rejected. Based on the skeleton structure and branching patterns, we postulate, contrary to former ideas, that the stem and branches of heterocorals represent the skeleton of a multi-polyp colonial coral, similar to many extant octocorals. We found numerous potential homologies with octocoral skeletons (notably the Keratoisidinae within the Isididae) and, as a result, we propose the inclusion of the order Heterocorallia within the subclass Octocorallia. This suggestion requires, however, further research on the other taxa of heterocorals. We also propose some changes to the morphological terminology for the Heterocorallia.

Fossil tabulate corals reveal outcrops of Paleozoic sandstones in the Atlantic Coastal Plain Province, Southeastern USA.

The geologic history of the Southeastern United States of America is missing nearly 350-million-years of rocks, sediments, and fossils. This gap defines the Fall Line nonconformity where Upper Ordovician consolidated rocks are directly overlain by Upper Cretaceous unconsolidated sediments of the Atlantic Coastal Plain Province. Here we begin to fill in the missing geologic record by reporting the discovery of fossils of lower-to-middle Paleozoic tabulate corals (Syringophyllidae) in angular, quartz-rich, ferruginous sandstones that crop out in the Carolina Sandhills Physiographic Province that forms the updip margin of the Atlantic Coastal Plain Province near the Fall Line. These fossils of extinct tabulate corals are the first evidence that Paleozoic (Upper Ordovician-Lower Silurian) sandstones crop out amidst the mostly Mesozoic-to-Cenozoic deposits of the Atlantic Coastal Plain Province of the United States of America. This discovery of Paleozoic fossils and strata in a region in which they were previously entirely unknown offers a more complete insight into the geologic history of the Southern Appalachian Mountains Region, Carolina Sandhills and updip margin of the Atlantic Coastal Plain Province and extends the previously identified range of Syringophyllidae in North America.

Tabulate Corals after the Frasnian/Famennian Crisis: A Unique Fauna from the Holy Cross Mountains, Poland.

Famennian tabulate corals were very rare worldwide, and their biodiversity was relatively low. Here we report a unique tabulate fauna from the mid- and late Famennian of the western part of the Holy Cross Mountains (Kowala and Ostrówka), Poland. We describe eight species (four of them new, namely ?Michelinia vinni sp. nov., Thamnoptychia mistiaeni sp. nov., Syringopora kowalensis sp. nov. and Syringopora hilarowiczi sp. nov.); the whole fauna consists of ten species (two others described in previous papers). These corals form two assemblages-the lower, mid-Famennian with Thamnoptychia and the upper, late Famennian with representatives of genera ?Michelinia, Favosites, Syringopora and ?Yavorskia. The Famennian tabulates from Kowala represent the richest Famennian assemblage appearing after the F/F crisis (these faunas appear some 10 Ma after the extinction event). Corals described here most probably inhabited deeper water settings, near the limit between euphotic and disphotic zones or slightly above. At generic level, these faunas show similarities to other Devonian and Carboniferous faunas, which might suggest their ancestry to at least several Carboniferous lineages. Tabulate faunas described here represent new recruits (the basin of the Holy Cross mountains was not a refuge during the F/F crisis) and have no direct evolutionary linkage to Frasnian faunas from Kowala. The colonization of the seafloor took place in two separate steps: first was monospecific assemblage of Thamnoptychia, and later came the diversified Favosites-Syringopora-Michelinia fauna.

New Famennian colonial coral (Rugosa) from the Holy Cross Mountains (Poland): an example of local evolution after Frasnian-Famennian extinction.

Colonial rugose corals are extremely rare in the fossil record after the Late Devonian (Frasnian-Famennian) extinction event. Here, we report a new genus and species, Famastraea catenata, from the late Famennian of the western part of the Holy Cross Mountains (Kowala) in Poland. Although this taxon is colonial, it displays many morphological characters very close to the typically late Famennian solitary species Palaeosmilia aquisgranensis (Frech, 1885), described earlier from the same locality. Hence, we postulate that F. catenata is derived from P. aquisgranensis. In contrast to other Famennian colonial rugose corals, the new taxon represents an example of local evolution within the group of so-called 'Strunian' corals. Consequently, we postulate that the new taxon represents a new colonial rugose fauna, which, however, did not survive the subsequent Late Devonian crisis (i.e. Hangenberg event). F. catenata most probably inhabited deeper water settings, possibly near the boundary between the euphotic and dysphotic zones, as inferred from many other benthic taxa described from this locality.

Enigmatic Fossils from the Lower Carboniferous Shrimp Bed, Granton, Scotland.

The Lower Carboniferous (Visean) Granton Lagerstätte (Edinburgh, Scotland) is principally known for the discovery of the conodont animal, but has also yielded numerous crustaceans and other faunas. Here we report on small branching colonies, reaching 10 mm in length. They are small, erect, arborescent, and irregularly branched with predominant monopodial and dichotomous growth. They bud in a single plane. In one specimen the wall microstructure is well preserved and it is composed of evenly spaced, linear fibers, running parallel to the axis of the stems, and connected by transverse bars. We discuss possible biological affinities of these organisms; we consider algal, poriferan, hydrozoan and bryozoan affinities. The general pattern of branching, presence of fan-like structures (interpreted here as possible gonophores) and microstructure suggests affinity to Hydrozoa, affinity to non-calcifying algae is less likely. Assuming hydrozoan nature; the microstructure might suggest affinities with the extant family Solanderiidae Marshall, 1892 that possess an internal chitinous skeleton. The EDS analysis shows that fossils discussed here are preserved as phosphates. The skeletons were probably not mineralized, the presence of phosphorus suggests that the colonies were originally composed of chitin. We describe these organisms as Caledonicratis caridum gen. et sp. nov. (Solanderiidae?, Capitata?). Colonies of C. caridum gen et. sp. nov. sometimes encrust the exuviae of crustaceans, which very probably lived in fresh to brackish water thus indicating a likely habitat of Caledonicratis.

Evidence of photosymbiosis in Palaeozoic tabulate corals.

Coral reefs form the most diverse of all marine ecosystems on the Earth. Corals are among their main components and owe their bioconstructing abilities to a symbiosis with algae (Symbiodinium). The coral-algae symbiosis had been traced back to the Triassic (ca 240 Ma). Modern reef-building corals (Scleractinia) appeared after the Permian-Triassic crisis; in the Palaeozoic, some of the main reef constructors were extinct tabulate corals. The calcium carbonate secreted by extant photosymbiotic corals bears characteristic isotope (C and O) signatures. The analysis of tabulate corals belonging to four orders (Favositida, Heliolitida, Syringoporida and Auloporida) from Silurian to Permian strata of Europe and Africa shows these characteristic carbon and oxygen stable isotope signatures. The δ(18)O to δ(13)C ratios in recent photosymbiotic scleractinians are very similar to those of Palaeozoic tabulates, thus providing strong evidence of such symbioses as early as the Middle Silurian (ca 430 Ma). Corals in Palaeozoic reefs used the same cellular mechanisms for carbonate secretion as recent reefs, and thus contributed to reef formation.

First fossil record of parasitism in Devonian calcareous sponges (stromatoporoids).

INTRODUCTION: Palaeozoic calcareous sponges (stromatoporoids) are common bio-constructing fossils; they are sometimes found in association with helicoidal structures of unknown biological affinities. The interaction between the tube-forming organisms has usually been classified as commensalism. METHODS: About 260 stromatoporoid skeletons from the Middle Devonian (Givetian) of the Mont d'Haurs section near Givet (Champagne-Ardenne, France) were thin-sectioned and analysed under transmitted light. RESULTS: Approximately 10% of the examined stromatoporoids (mainly belonging to the genera Actinostroma, Stromatopora and Stromatoporella) contain tubes classified as Torquaysalpinx sp. The Torquaysalpinx organisms penetrated the skeletons of stromatoporoids in vivo (as evidenced by skeletal overgrowths); around the infesting organisms, growth bands are bent down. CONCLUSION: Diminished growth rates around the infesting organism demonstrate a negative influence on the host, similar to that seen in the modern demosponge-polychaete association of Verongia-Haplosyllis. This is demonstrated by changes in growth bands. As in the above-mentioned association, the endosymbiont might have been feeding directly upon the tissues of the host. The Torquaysalpinx organisms were gaining habitat and possibly also food resources - for them this interaction was clearly positive. This long-term association can therefore be classified as parasitism. This is the first evidence for parasitism in Palaeozoic sponges.