Lucky rugose corals on crinoid stems: unusual examples of subepidermal epizoans from the Devonian of Morocco

Berkowski, B & Klug, C. 2011: Lucky rugose corals on crinoid stems: unusual examples of subepidermal epizoans from the Devonian of Morocco. Lethaia, Vol. 45, pp. 24–33. In the fossil record, evidence for true epizoans, i.e. living animals inhabiting other living host‐animals, is rather rare. A host reaction is usually needed to proof the syn vivo‐settling of the epizoan. Herein, we provide a first report of such an epizoan biocoenosis from various strata of the Early Devonian of Hamar Laghdad, the world‐renowned Moroccan mud‐mound locality. In this case, solitary rugose corals settled as larvae on crinoid stems, perhaps at a spot where the epidermis was missing for some reason (injury, disease). Both the crinoid and the coral began to grow around each other. By doing so, the affected crinoid columnals formed a swelling, where ultimately only an opening slightly larger than the coral orifice remained. We discuss both macroecological and small‐scale synecological aspects of this biocoenosis. The coral profited from its elevated home because it reached into more rapid currents providing the polyp with more food than at the densely populated seafloor, which was probably covered by a coral‐meadow around the mounds and hydrothermal vents. □Corals, crinoids, Early Devonian, epizoans, Morocco, Rugosa.     

New offset-associated structures in some Carboniferous rugose corals

Tubular structures termed ducts, which extend from the base of offsets axially and upwards through the disscpimental tissue of the parent corallite, are described from some Carboniferous rugose corals. Two ducts are usually associated with each offset, one from each extremity where the offset wall meets that of the parent. The ducts may merge to form a tangentially elongate slit at higher levels in the disscpimcntarium of the parent. The structures served to prolong gastric and nervous communication between parent and daughter polpys following budding. Ducts and slits are well developed in cerioid Lonsdaleia but are absent in fasciculate Lonsdaleia , a distinction lending credence to the recognition of the former group of species as a distinct genus, Actinocyathas , within the Axophyllidac. In addition, less well developed ducts are present in at least some cerioid species of Lithostrotion , whereas they are absent from fasciculate species, supporting separation of the latter from Lithostrotion sensu stricto and their assignment to the genus Siphonodendron. The form and distribution of duets also confirms polyphyly in Thysanophyllum. The separation of T. pseudovermiculare (as Dorlodotia ), without ducts, from Thysanophyllum sensu stricto , with weak ducts similar to those in some Lithostrotion spp., is supported. Both arc assigned to the T. pmedictum has well developed ducts, however, which reinforces other evidence suggesting its removal from Thysanophyllum and classification in the Axophyllidac in close affinity with Actinocyathus, Rugosa, offsets, ducts, functional morphology, taxonomy, Carboniferous.     

Principles of early ontogeny in the rugose corals: a critical review

It has been suggested that septal furrows in the Palaeozoic rugose corals are secondary in comparison both to growth lines and to the septal blades inside calices. The leading role of furrows as an indicator of septal increase is thus questioned. Septa are always grouped in quadrants but their increase is generally towards the cardinal septum. The arrangement of the contratingent minor septa seems to be independent of the increase in septal furrows. Hence, the growth of the polyp's skeleton may have started from an aseptal cup.     

Some aspects of life strategies of Early Palaeozoic rugose corals

Examination of some well-preserved specimens of Silurian rugose corals from Gotland reveals that the following categories of life strategies can be distinguished: ambitopic, liberosessile, fixosessile, rhizosessile and possibly limited vagile. Most solitary rugose corals appear to have been liberosessile and are characterized by initial attachment to a small sediment grain but subsequently becoming recumbent on a soft substrate. Detailed studies of Phaulactis angelini, Holophragma calceoloides, Laccophyllum lindstroemi, Rhegmaphyllum conulus and Rhabdocyclus ocksaroensis n. sp., and comparison with earlier described species, provide some new aspects on different life strategies for solitary corals.     

Restudy of the Early Devonian rugose coral Xystriphylloides from South China

Early Devonian rugose coral Xystriphylloides in South China is restudied from the perspective of its taxonomic definition, biostratigraphy, paleoecology, and implication for the Yujiang Event. The diagnostic characters of the genus are clarified and its taxonomic position is also revised. In particular the septa of Xystriphylloides are shown to be fibrous rather than trabeculate. Xystriphylloides distinctus Yu n. sp. is described. Species of Xystriphylloides are widely distributed in South China and North Vietnam, so they are useful for regional biostratigraphic correlation and regional geological mapping. As an important biostrome builder, Xystriphylloides became extinct below the base of the nothoperbonus Conodont Zone, so it can be considered as one of the important pieces of evidence for recognizing the first episode of the Early Devonian Emsian Yujiang Event, which is also significant for studying paleoecology and event stratigraphy.     

Upper Permian coral reef and colonial rugose corals in northwest Hunan, South China

Summary The roles of Permian colonial corals in forming organic reefs have not been adequately assessed, although they are common fossils in the Permian strata. It is now known that colonial corals were important contributors to reef framework during the middle and late Permian such as those in South China, northeast Japan, Oman and Thailand. A coral reef occurs in Kanjia-ping, Cili County, Hunan, South China. It is formed by erect and unscathed colonies ofWaagenophyllum growing on top of one anotherin situ to form a baffle and framework. Paleontological data of the Cili coral reef indicates a middle to late Changhsing age (Late Permian), corresponding to thePalaeofusulina zone. The coral reef exposure extends along the inner platform margin striking in E-S direction for nearly 4 km laterally and generally 35 to 57 m thick. The Cili coral reef exhibits a lateral differentiation into three main reef facies; reef core facies, fore-reef facies, and marginal slope facies. The major reef-core facies is well exposed in Shenxian-wan and Guanyin-an sections where it rests on the marginal slope facies. Colonial corals are dispersed and preserved in non-living position easward. Sponges become major stabilizing organisms in the eastern part of Changhsing limestone outcrop in Kanjia-ping, but no read sponge reefs were formed. Coral reefs at Cili County in Human are different distinctly from calcisponge reefs in South China in their palaeogeography, lithofacies development, organic constitutuents, palaeoecology and diagenesis. The Cili coral reef also shows differences in age, depositional facies association, reef organisms and diagenesis from coral reefs in South Kitakami of Japan, Khorat Plateau of Thailand, and Saih Hatat of Oman. Although some sponge reefs and mounds can reach up to the unconformable Permian/Triassic boundary, coral reef at Kanjia-ping, Cili County, is the latest Permian reef known. This reef appears to had been formed in a palaeoenvironment that is different from that of the sponge reefs and provides an example of new and unique Permian reef type in South China, and could help us to: 1) understand the significance of colonial corals in Permian carbonate buildups; 2) evaluate the importance of coral community evolution prior to the collapse of reef ecosystems at the Permian/Triassic boundary; 3) better understand the effects of the biotic extinction events in Palaeotethys realm; 4) look for environmental factors that may have controlled reefs through time and space, and 5) provide valuable data for the study of Permian palaeoclimate and global evolutionary changes of Permian reefs and reef community.     

Late Moscovian (mid-Pennsylvanian) rugose corals from Wadi Araba (Egypt,Eastern Desert): Taxonomy,palaeoecology and palaeobiogeography

An important late Moscovian rugose coral association is described from the Rod El Hamal Formation of the Wadi Araba area (northern Eastern Desert, Egypt). The upper part of the formation yielded a moderately abundant but poorly diversified coral fauna composed of large dissepimented solitary rugose corals. In total, six species belong to the Bothrophyllidae and the Geyerophyllidae, including three new species: Bothrophyllum suezensis, Bothrophyllum cylindricum, and Amygdalophylloides omarai. Three undeterminable Bothrophyllidae taxa are left in open nomenclature. The corals were attached to the soft substrate by talons and rootlets, either rooted and growing upward as mud-stickers or as secondary recliners. They show frequent rejuvenations and constrictions. Encrustation and/or bioerosion are scarce. Carbonate microfacies indicate a habitat in littoral and inner neritic zones adjacent to a low terrigenous hinterland. Colonial taxa are missing and tabulate corals are rare. The corals from the Rod El Hamal Formation are the only example of a late Moscovian coral fauna on the southern margin of the Palaeotethys, that is to say along the northern margin of Gondwana. The association shows similarities with northwestern Spain (Cantabrian Mountains), the Donets and Moscow basins, and thus a general attribution to the Palaeotethys realm.     

Early Silurian (Rhuddanian) rugose corals and sponges from the Ak-Kerme Peninsula,Kazakhstan

An assemblage of earliest Silurian (Llandovery, Rhuddanian) fossils from South Kazakhstan (Ak-Kerme Peninsula, Lake Balkhash) contains solitary rugose corals (Calostylis denticulata, Streptelasma? sp., and Cystipaliphyllum sp.) and the demosponge Calycocoelia typicalis, which are described here. This assemblage occurs with previously described brachiopods and constitutes a post extinction survival fauna; such faunas are poorly known and this study fills a gap in our knowledge. All three genera of Rugosa were transitional across the Ordovician–Silurian boundary and are also reported from other parts of Kazakhstan, Uzbekistan, and Tajikistan during the Llandovery. Streptelasma had already been present in Kazakhstan during the Ordovician, while during the Rhuddanian Calostylis immigrated from Baltica or China, and Cystipaliphyllum from the Australian part of Gondwana. Demosponges are rare during the Llandovery but probably had a cosmopolitan distribution. Calycocoelia typicalis marks the first Rhuddanian record of lithistid demosponges, and the first record of Silurian demosponges from Kazakhstan.     

Habitats of colonial rugose corals: the Mississippian of western Europe as example for a general classification

Aretz, M. 2010: Habitats of colonial rugose corals: the Mississippian of western Europe as example for a general classification. Lethaia, DOI: 10.1111/j.1502‐3931.2010.00218.x. Colonial rugose corals are a major constituent of shallow‐water marine benthic communities in Mississippian times. The study of western European rugose coral habitats from the base of the Tournaisian stage to the Serpukhovian stage allows the recognition of four basic habitat types, which can be divided into a total of 11 subtypes. The classification is mainly based on field data, and thus rapidly applicable. Level‐bottom communities in which large colony distances are characteristic (type A) represent the most basic community type; polyspecific (subtype A1) and monospecific (subtype A2) subtypes occur. Reduced colony distances result in the formation of coral meadows (type B), which either show homogenous coral distribution (subtype B1) or the development of patches (subtype B2). Coral biostromes (type C) represent a spectrum between hydrodynamically controlled biostromes (nothing in place, subtype C1) and biologically constructed and controlled biostromes (subtype C2). The bulk of the biostromes represent mixtures of those two subtypes (subtype C3). Colonial rugose corals are widely encountered in Mississippian bioherms where they are dwellers (subtype D1), form capping beds (subtype D2), support framework building along with other organisms (subtype D3) and form coral framework (subtype D4). The latter is probably the most uncommon of all subtypes in Mississippian times. The classification is widely applicable to other groups. □Classification, habitats, Mississippian, palaeoecology, palaeoenvironment, rugose corals.     

Epizoic bryozoans and corals as indicators of life and post-mortem orientations of the Devonian brachiopod Meristella

The in-life and post-mortem orientations of the Lower Devonian brachiopod Meristella atoka from the Haragan Formation (Lower Devonian; south-central Oklahoma) are inferred from the distribution of epizoic bryozoans and the orientations of base plates of epizoic corals. Three bryozoans, Cyphotrypa corrugata, Fistuliporella maynardi and Leioclema pulchellum, and one coral, Favosites conicus, are considered. Most zoaria that contact the commissure terminate at the commissurc, and a few zoaria terminate at growth lines. This suggests that the bryozoans were primarily life associates of Meristella atoka. Collectively, these three bryozoan species most extensively encrusted marginal sectors of the brachial valve and are very rare on the posteromedian sector (the umbo) of the pedicle valve. This distributional pattern indicates that the preferred living orientation of Meristella atoka was umbo-down (posteromedian sector of the pedicle valve resting on or buried in the substratum) with the commissure steeply inclined to the sediment water interface. Most coralla of Favosites conicus that contacted the commissure encrusted over the commissure. This indicates that Favosites conicus either preferentially encrusted Meristella atoka post-mortem or colonized living brachiopods but subsequently caused them to die. Furthermore, Favosites conicus most extensively encrusted anterior sectors of the brachial valve, especially the fold. The lateral and anterior orientations of the commissure with respect to the base plates (holothecae) of Favosites conicus indicates that the brachiopods were oriented approximately horizontal with respect to the base plates. This suggests that the preferred post-mortem orientation of Meristella atoka was resting nearly horizontally on the substratum. These data and interpretations confirm previously inferred in-life orientations of Meristella atoka and are consistent with post-mortem orientations hypothesized for elongate-oval athyrid brachiopods. □Brachiopoda, Athyridacea, Meristella atoka , palaeoecology, autecology, epizoans, bryozoans, corals, sedimentology, Lower Devonian, North America, Oklahoma.     

Large dwellers of the Silurian Halysites biostrome: rhizosessile life strategies of cystiphyllid rugose corals from the Llandovery of Gotland

An exceptionally well‐preserved, unusual biostrome composed of the framebuilding cateniform tabulate coral Halysites catenularius (Linnaeus, 1767) bears an assemblage of the relatively large solitary cystiphyllid rugosan Cystiphyllum visbyense Wedekind, 1927. The corallites of solitary cystiphyllids are embedded within the ranks of the halysitid colonies, which developed on a soft, muddy substrate and in relatively turbid water. The cystiphyllid larvae successively settled mostly on the ranks of halysitid colonies and on colonies of the tiny phaceloid rugose coral Nanophyllum ramosum Johannessen, 1995, whereas calice‐in‐calice recruitment was not successful for these cystiphyllid corals. Further growth of C. visbyense was supported by rhizoid structures, which were most frequently developed on the cardinal (convex) side of the corallite. The process of formation of the rhizoid structures is here discussed and explained in detail, showing that they were formed by the extension of the basal ectodermal tissue of the polyp. The cystiphyllids, which settled on the walls of living corallites of halysitid colonies, used sweeper tentacles to kill the smaller polyps of the colony to maintain the space around them and expand. Hence, they ultimately used the halysitid colonies only as a hard substrate to stabilize their position on the soft muddy sediment.     

Vertical migrations of fish larvae: Eulerian and Lagrangian observations in the Eastern English Channel

The Eastern English Channel is known for its strong hydrodynamics.Tidal and residual currents are reinforced by the south-westerndominant wind and drift waters from the English Channel to theNorth Sea. Previous spatial studies have shown that the advectionof larvae could differ from one species to another. Flounder(Pleuronectes flesus) larvae were found offshore, drifting tothe north until the fins were formed; then they were found nearthe coast. However, sole (Solea solea) larvae remained in coastalwaters during their development. The difference in larval spatialdistribution is assumed to be related to the interaction betweenvertical migration and advection by alternative tidal currents,leading to a selective tidal stream transport. To describe thevertical distribution of these larvae, two strategies were used.First, a Eulerian study was carried out with samples taken atthe same geographical location every 1.5 h for 41 h. Ichthyoplanktonwere collected in the water mass using a Bongo net and witha suprabenthic multi-net sledge, at four layers above the seabed, between 0.1 and 1.4 m. Secondly, to enable water movementto be disregarded, a Lagrangian study was carried out by usinga Bongo net every 3 h, following a drifting buoy for 3 days.The results show that even during the youngest stages, solelarvae are able to perform tidal and diel vertical migration.We assume that they may limit their advection to the North Seabecause of their upward migration during ebb and at night, whichmay enable them to remain in the same area dealing with thecurrents. Flounder larvae begin their vertical migration atthe stage of notochord flexion, which ends their drift to thenorth. The larvae reach the bottom of the water column, particularlyduring ebb when they are concentrated in the first 40 cm abovethe bottom. This behaviour favours their advection during flow,leading to efficient and fast transport towards the coast.     

Pleistocene to Recent scleractinian deep-water corals and coral facies in the Eastern Mediterranean

Recent investigations of the Eastern Mediterranean Sea carried out during the GECO cruise with RV Urania provided a substantial number of new cold-water coral (CWC) records, including branching and solitary scleractinian species. These new sites are located along steep escarpments and on topographic highs along the margins of Crete, Karpathos, and Rhodes. The majority of the corals represent fossil occurrences, predominantly Late Pleistocene assemblages. Our research documents that the Eastern Mediterranean Basin has been colonized by CWC at favorable times during the Last Glacial, in particular during the Younger Dryas. Schizocyathus fissilis is reported for the first time for the Mediterranean, while the finding of Ceratotrochus magnaghii represents the first record for the Eastern Mediterranean. Various coral facies occur on the southerly island slopes of Crete, Karpathos, and Rhodes, including hardgrounds and loose skeletal sediments. Hardgrounds occur on steep topographies between ca. 500 and 1,700 m, and can conveniently be subdivided as (1) Neopycnodonte-Desmophyllum framestone, (2) Desmophyllum-Caryophyllia framestone, (3) Madrepora-Lophelia rudstone, (4) Pelagic mudstone and wackestone, and (5) Siliciclastic-carbonate conglomerate and breccia. Unconsolidated skeletal sediments containing corals mainly occur on gentler topographic situations between ca. 140 and 600 m and can be subdivided as: (A) Lophelia-Madrepora rubble, (B) Dendrophyllia rubble, (C) Stenocyathus rubble, (D) Caryophyllia calveri rubble, and (E) fine-grained sediment with octocoral axes. Many of these facies types are also present in the western part of the Mediterranean and have fossil representatives from the Pleistocene to the Recent. Radiocarbon dating (AMS-¹⁴C) reveals Younger Dryas ages between 12.4 and 12.0 ka cal BP for Lophelia pertusa and Madrepora oculata. Desmophyllum dianthus occurs during the Last Glacial Maximum (21.8 ka cal BP) and the Younger Dryas (11.7 ka cal BP), as well as during the Late Holocene and subrecent times (4.4?C0.6 ka cal BP). Caryophyllia sarsiae occurs during the Late Glacial (15.5 ka cal BP), while Caryophyllia calveri occurs during the Early Preboreal (10.8 ka cal BP). The ages for the framework-constructing corals L. pertusa and M. oculata are coherent with their temporal predominance during the Younger Dryas in other parts of the Mediterranean.     

Isolation of potential fungal pathogens in gorgonian corals at the Tropical Eastern Pacific

A major environmental problem in the ocean is the alarming increase in diseases affecting diverse marine organisms including corals. Environmental factors such as the rising seawater temperatures and terrestrial microbial input to the ocean have contributed to the increase in diseased organisms. We isolated and identified the fungal agents that may be leading to a disease in the Pacific sea fan Pacifigorgia eximia (Gorgoniidae, Octocorallia) in the Tropical Eastern Pacific. We isolated thirteen fungal genera in healthy and diseased colonies including Aspergillus sydowii. Aspergillus has been previously identified as responsible for the mortality of gorgonian corals in the Caribbean. This disease was observed in the Eastern Pacific affecting a completely different set of species nearly 30 years after the Caribbean outbreak, which concur with rising seawater temperatures and thermal anomalies that have been observed in the last 4 years.     

Lower Devonian biostratigraphic correlation in Eastern Australia and Western North America

Savage, N. M.: Lower Devonian biostratigraphic correlation in Eastern Australia and Western North America.
Correlation of Lower Devonian marine strata of Eastern Australia and Western North America can be attempted using several fossil groups. In general the faunas are similar, with many species common to both areas. Graptolites occur infrequently, but are of great value because of their precise zonal significance. Several of these graptolites occur in the classic type sequences of Bohemia but are absent from the Rhineland stages. For this reason, and because of the Bohemian affinities of the shelly faunas, the Bohemian stage names are preferable to the Rhenish names for use in Eastern Australia and Western North America. Cono-donts are less precise index fossils than graptolites, but are considerably more abundant and therefore have greater application. Brachiopods are valuable if viewed as assemblages. In terms of the Rhenish stages, most Eastern Australian and Western North American early Lower Devonian formations appear to have been placed too high. Recent evidence from northern Canada throws new light on the problem of correlating Rhenish and Bohemian classic early Lower Devonian stages.     

Extinctions in simple and complex communities

Disagreement exists between the results of theoretical and empirical exploration into the effect of increasing community complexity on the stability of multispecies ecosystems. A recent return to interest in this area suggests previous results should be re-assessed, from both experimental studies and models, to understand where this discrepancy arises from. Here we propose various simple extensions to a standard multispecies community model that each increase the complexity of the system in a different way. We find that increasing the number of species in a community leads to a decrease in community persistence after the system is perturbed, and go on to show that increasing the dynamical diversity of the community members leads to an increase in stability through a reduction in extinction events, relative to the less complex form of the model. Our results suggest that different forms of complexity lead to different outcomes in the stability properties of the community. While aspects of this work agree with previous empirical findings that more complex communities are more robust to perturbation, we stress that the type of complexity included and the measure of stability used in community models must be properly defined, to allow objective comparisons to be made with previous and future work.     

Influence of temperature and nutrients on species richness of deep water corals from the western coast of the Americas

Studies on biogeography of stony corals from the eastern Pacific have been conducted in detail only for reef species, and to date there have been no attempts to explain the differences of regional species richness on the basis of oceanographic conditions. The objective of this work was to determine the relationship between deep-water (<200 m) scleractinian species richness along the western coast of the Americas, and four oceanographic variables (temperature, nitrates, phosphates and silicates), and the feasibility to use this information to model effects of global warming on those associations. Data on coral distribution were gathered from bibliography and museum collections, while information on oceanographic conditions from 200 to 2000 m depth was obtained from NOAA atlases. Species richness, estimated for intervals of 5 degrees of latitude, was correlated with abiotic factors using the Spearman rank coefficient. In the Northern Hemisphere, total species richness was positively influenced by temperature, but negatively by nutrients. In contrast, there was no effect of those factors on coral diversity in the Southern Hemisphere. At the family level, high temperatures favored species richness of Caryophylliidae and Dendrophylliidae north of the Equator, but diversity was reduced in areas of high concentration of silicates. In the Southern Hemisphere, temperature was not associated with deep-water coral richness, but correlated negatively with diversity of Caryophylliidae. Nutrients also showed an inverse relationship with richness of the latter family. In the rest of the families analyzed, there was no apparent effect of oceanic conditions on species richness in the Southern Hemisphere. The results indicated that richness may be influenced by changes in oceanographic factors (especially temperature and silicate concentration). Then, it is feasible to develop numerical models to predict possible changes in deep-water coral diversity on the basis of scenarios from global warming models.     

Middle and Early Upper Devonian Miospore Zonations in Eastern Yunnan and the Significance of Their Stratigraphy

The Devonian marine and non marine deposits were widely distributed in East Yunnan, where rich marine faunas and floras were present. The Middle and early Upper Devonian miospores of Eastern Yunnan were also abundant and usually well- preserved. The present paper provides both analysis of qualitative and quantitative composition development of Middle and early Upper Devonian spore assemblage in Eastern Yunnan, and a palynological zonatlon containing four successive assemblage zones: 1.Zone of Calyptosporites velatus-Rhabdosporites langii (VL); 2. Zone of Archaeozonotriletes variabilis-Calyptosporites proteus (VP); 3. Zone of Geminospora lemurata-Cristatisporites triangulatus (LT); 4. Zone of Archaeoperisaccus ovalis-Lagenicula bullosum (OB). The Middle and early Upper Devonian spore assemblages of Eastern Yunnan may be compared with those in South China and West Qinling. The spore zonation possesses stratigraphical dating of the Devonian megafloras of the region, particularly those from the Middle and Upper Devonian. The proposed spore zonation is closely compared with that erected for the Middle and early Upper Devonian of Old Sandstone Continent and adjacent Region. Based on palynological data, the reconstruction of Paleogeography and Paleoecology are discussed.     

New species of spiriferids (Brachiopoda) from the Devonian and Carboniferous of Eastern Europe

In a revision of previously published materials, two new spiriferid species from the Late Tournaisian, Unispirifer subtornacensis sp. nov. and Mesochorispira ussuilensis sp. nov., and Atylephorus nalivkini sp. nov. from the Lytva Horizon of the Upper Devonian of Bashkiria are described. Based on the original collection, a new Serpukhovian subspecies, Podtsheremia duplicicosta triplicicosta, from the southern Ural Mountains and a new Late Carboniferous species, Purdonella kalashnikovi, from the polar Ural Mountains and adjacent islands are established.