Early Ordovician bryozoans from north-western Russia

ABSTRACT. A bryozoan assemblage from the Billingen Stage of north-western Russia (Ingria) is apparently the oldest known in the world. It consists of six species distributed among the trepostomate genera Esthoniopora Dianulites, Revalotrypa Phragmophora and Hemiphragma . Two species are new: Phragmophora lavaense Pushkin, sp. nov. and Hemiphragma priscum Pushkin, sp. nov. The bryozoans are characteristic of the medium diversity benthic fauna of uncertain origin which migrated into the Baltic Basin during Billingen time (late Prioniodus elegans — early Oepikodus evae zones). The Billingen bryozoans have a close similarity to the late Arenig bryozoan faunas of Baltoscandia that are also dominated by trepostomates, but differ significantly from the low diversity late Arenig bryozoan assemblages of North America, Ireland, North China, Vajgach and Novaja Zemlja.     

Zooidal anatomy in Ordovician and Carboniferous trepostome bryozoans

Fossil bryozoans sometimes contain fossilised brown bodies which remain after polypide degeneration. Position and shape of brown bodies as well as different skeletal diaphragms within living chambers allow outlining of autozooid anatomy in Palaeozoic trepostome bryozoans. In some trepostome bryozoans short autozooids were restricted by complete basal diaphragms to distal parts of autozooecia. In species with alternating hemiphragms, as in the specimen ofHemiphragma sp. described here, long autozooids occupied the entire autozooecial chambers. Their short polypides were positioned in distal parts of the autozooid. Both anatomic types correspond to the progressive polypide cycle afterBoardman. For species with ring septa, non-alternating hemiphragms (studied in an example ofNipponostenopora karatauensis) as well as without any diaphragms the stationary type of polypide cycle seems also to be possible. In that case, the polypides should be as long as the cystid.      

Horizon of the oldest known bryozoans (Ordovician)

Among the six species of fossil bryozoans described by Xia et al. [Xia, F.S., Zhang, S.G., Wang, Z.Z., 2007. The oldest bryozoans: new evidence from the late Tremadocian (Early Ordovician) of East Yangtze Gorges. Journal of Paleontology 81 (6), 1308–1326] from the Fenghsiang Formation of the Chenjiahe section of Yichang and the Guanzhuangping section, Liujiachang Town of Songzi, the Yangtze Gorges, Nekhorosheviella nodulifera occurs in the pristinus Conodont Subzone (lower Subzone of deltifer Conodont Zone) and the five other species are associated with the conodont Paltodus deltifer deltifer and the graptolites Acanthograptus sinensis and A. erectoramus. Based on the conodonts and graptolites, this bryofauna stratigraphically corresponds to the A. sinensis Graptolite Zone or P. deltifer Conodont Zone. The bryofauna is of Tremadocian Age (Early Ordovician) and represents the oldest bryozoans so far as known.     

Stratigraphic and paleogeographic distribution of the Upper Ordovician bryozoans of Mongolia

The taxonomic diversity and distribution of the Upper Ordovician bryozoans of Mongolia are analyzed. Five heterochronous assemblages of bryozoans are established: Chigertei and Tsagaan del in the Sandbian Stage and Bairim Ovoo, Uuregnur, and Sairin in the Katian Stage. The bryozoans are shown to be important for solving complex problems of stratigraphy of the Ordovician of Mongolia and for paleogeographic reconstructions.     

Ecological consequences of Late Quaternary extinctions of megafauna

Large herbivorous vertebrates have strong interactions with vegetation, affecting the structure, composition and dynamics of plant communities in many ways. Living large herbivores are a small remnant of the assemblages of giants that existed in most terrestrial ecosystems 50 000 years ago. The extinction of so many large herbivores may well have triggered large changes in plant communities. In several parts of the world, palaeoecological studies suggest that extinct megafauna once maintained vegetation openness, and in wooded landscapes created mosaics of different structural types of vegetation with high habitat and species diversity. Following megafaunal extinction, these habitats reverted to more dense and uniform formations. Megafaunal extinction also led to changes in fire regimes and increased fire frequency due to accumulation of uncropped plant material, but there is a great deal of variation in post-extinction changes in fire. Plant communities that once interacted with extinct large herbivores still contain many species with obsolete defences against browsing and non-functional adaptations for seed dispersal. Such plants may be in decline, and, as a result, many plant communities may be in various stages of a process of relaxation from megafauna-conditioned to megafauna-naive states. Understanding the past role of giant herbivores provides fundamental insight into the history, dynamics and conservation of contemporary plant communities.     

Symbiosis between gastropods and bryozoans in the late Ordovician of Cumbria, England

Five examples of symbiosis between gastropods and trepostome bryozoans are described from the Ashgill (late Ordovician) of Cumbria (England). The gastropods are invariably found associated with the bryozoans. whereas the bryozoans may be free-living. Encrustation is considered to have taken place, in most cases, on mature, living gastropod shells, resulting in the trepostome utilizing the shell as a surrogate basal disc. Three specimens show encrustation on an empty gastropod shell, causing the trepostome to develop a basal disc similar to non-encrusting forms. The bryozoan colony probably benefited from the symbiotic relationship by increased water flow over the colony, caused by gastropod locomotion, whereas the gastropod was afforded protection from predators. The new species Spiroecus nidhoeggi. Diplotrypa hvergelmi and Monotrypa fontinalis are described.     

Paleoenvironment of Late Mississippian fenestrate bryozoans, eastern United States

Fenestrate bryozoans in Chesterian rocks of the eastern United States are common to abundant in diverse carbonate platform lithologies but are not common in clastic deposits. Fenestella and Archimedes are the most abundant and widespread. Fenestella apparently tolerated a range of water agitation and lived on shoal margins as well as in more protected areas. Archimedes required quiet bottoms and was locally extremely abundant immediately leeward of shoals. Polypora and Septopora were less common than, but had a similar distribution to, Fenestella except that they apparently did not live on shoals. Lyroporella inhabited zones of pronounced current activity and fine quartz sand influx. Penniretepora and Ptylopora were least often encountered, restricted to fine-grained sediments deposited in protected regions.     

Upper Ordovician bryozoans from the Xiazhen Formation of Yushan,northeastern Jiangxi,East China

The bryozoan fauna from the Xiazhen Formation (Katian, Upper Ordovician) of northeast Jiangxi Province, southeast China is reported here. Seventeen species of bryozoans belonging to fifteen genera and four orders are identified: Homotrypa yushanensis, Homotrypa sp., Prasopora yushanensis, Trematopora sp., Monotrypella sp., Rhombotrypa sp., Orbignyella sp., Constellaria jiangxiensis, Constellaria sp., Stictopora nicholsoni, Trigonodictya parvula, Ptilodictya ensiformis, Stictoporella sp., Pseudopachydictya sp., Nematopora sp., Arthrostylidae sp. indet., and Chasmatoporidae sp. indet. Four of these genera have been reported previously but nine genera (Trematopora, Monotrypella, Rhombotrypa, Orbignyella, Trigonodictya, Ptilodictya, Stictoporella, Pseudopachydictya, and Nematopora), one rhabdomesine and one fenestrate are found for the first time in the Late Ordovician strata of South China. Our palaeogeographical analysis suggests that the bryozoan association is typical for the Katian, which is mostly widespread in Laurentia, Siberia, Baltica and Mediterranean, and displays palaeobiogeographical relationships to the Laurentia–Siberia Province.     

Late Ordovician extinction of North American and British crinoids

After reaching a diversity peak in the Caradocian, North American Ordovician crinoids underwent a gradual decline to a nadir in the early Ashgillian (Maysvillian). This interval, recording extinction of the Cleiocrinidae, Merocrinidae, Ottawacrinidae, Hybocystitidae, and several lineages of camerate crinoids, was apparently caused by major environmental shifts in seas of eastern North America resulting from a westward-prograding wedge of terrigenous clastics derived from the Taconic Highlands, possibly coupled with a marine transgression in the Maysvillian that allowed colder water slope biofacies to invade the craton. Crinoids suffered a major episode of extinction in the late Ashgillian (late Richmondian/Rawtheyan). This event, preceding the end of the Ordovician by at least one stage or 2 to 4 million years, resulted in extinction of 12 families of crinoids including the Xenocrinidae, Tanaocrinidae, Reteocrinidae, Archaeocrinidae, Anthracocrinidae, Cincinnaticrinidae, Iocrinidae, Anomalocrinidae, Carabocrinidae, Cupulocrinidae, Porocrinidae, and Hybocrinidae. Glacio-eustatic lowering of sea level may have triggered this crisis by partially draining the North American craton, resulting in changes in oceanic circulation, salinity, and temperature. Latest Ordovician (Hirnantian) carbonates of the North American mid-continent region contain pelmatozoan assemblages from which Silurian crinoids radiated. These taxa were largely unaffected by a minor extinction event at the Ordovician/Silurian boundary.     

Stepwise mass extinctions and impact events: Late Eocene to early Oligocene

Species ranges and relative abundances of dominant planktonic foraminifers of eight late Eocene to early Oligocene deep-sea sections are discussed to determine the nature and magnitude of extinctions and to investigate a possible cause-effect relationship between impact events and mass extinctions.Late Eocene extinctions are neither catastrophic nor mass extinctions, but occur stepwise over a period of about 1–2 million years. Four stepwise extinctions are identified at the middle/late Eocene boundary, the upperGlobigerapsis semiinvoluta zone, theG. semiinvoluta/Globorotalia cerroazulensis zone boundary and at the Eocene/Oligocene boundary. Each stepwise extinction event represents a time of accelerated faunal turnover characterized by generally less than 15% species extinct and in itself is not a significant extinction event. Relative species abundance changes at each stepwise extinction event, however, indicate a turnover involving > 60% of the population implying major environmental changes.There microtektite horizons are present in late Eocene sediments; one in the upperG. semiinvoluta zone (38.2 Ma) and two closely spaced layers only a few thousand years apart in the lower part of theGloborotalia cerroazulensis zone (37.2 Ma). Each of the three impact events appears to have had some effect on microplankton communities. However, the overriding factor that led to the stepwise mass extinctions may have been the result of multiple causes as there is no evidence of impacts associated with the step preceding, or the step following the deposition of the presently known microtektite horizons.     

Mass extinctions

Factors influencing the evolution of complex traits such as body size are notoriously difficult to study but a new review of work on marine iguanas in the Galapagos islands suggests an answer may lie in the interplay of natural and sexual selection.     

Trace fossils on a Late Ordovician glacially striated pavement in Algeria

An exceptional exposure of Late Ordovician glaciogenic sediments crops out in Dider, SE Algeria, within the Tassili N'Ajjer region. The sediments consist of sandstones and diamictites sandwiched between a Mid Ordovician fluvial and tidal sandstone (In Tahouite Formation) below and Early Silurian shale (Oued Imirhou Formation) above. Stratigraphic discontinuities within the Late Ordovician glaciogene succession include palaeovalley incisions and glacially striated pavements. Striation and fluting of a soft-bedded sediment beneath an ancient ice sheet is supported by abundant dewatering structures and soft-sediment gouges interpreted to have been produced by the action of stone ploughing. In Dider, two types of previously undescribed circular structures sit in negative relief on this glacial pavement, namely 1) paired thumb-shaped impressions 2 cm in diameter and 3 mm in depth, and 2) a 5 cm wide impression with 3 segmented nested cycles. A framboid or aggregate origin may be appropriate for the smaller of the features but the larger impression is interpreted as biogenic: internal complexity is characteristic, discounting concretion moulds and water escape structures. A biogenic origin as a coelenterate resting trace is proposed with speculation on conditions of exceptional preservation in an ancient periglacial environment.     

Paleozoic bryozoans of Mongolia

Stratigraphic assemblages characteristic for the Ordovician, Silurian, Devonian, Carboniferous, and Permian are presented based on the analysis of newly refined and supplemented data on the composition and distribution of Paleozoic bryozoans in Mongolia. Thirty-four auxiliary biostratigraphic units ranked as beds with bryozoans are established in order to subdivide the Paleozoic strata of Mongolia.     

The maintenance of evolutionary equilibrium in Late Ordovician benthic marine invertebrate faunas

An upper evolutionary limit on species equilibrium number (S of MacArthur & Wilson) has been postulated but never demonstrated. Results of the present study indicate that an S of about 10 was attained very rapidly in a Late Ordovician benthic marine invertebrate faunal succession. This S value was then maintained for perhaps five million years within a marine environment that did not remain uniform but changed progressively from relatively deep to shallow water. Most importantly, significant taxonomic and presumed trophic changes that accompanied the gradual habitat alteration seem to have had a negligible effect on the plateau-like limit to species equilibrium number. We conclude from these preliminary results that the maintenance of evolutionary equilibrium in an open ecosystem strongly suggests a long-term component of biotic resiliency, at least within this one marine ecosystem, and perhaps in many others. The reasons for this resiliency remain to be explored.     

Permian bryozoans of the NW-tethys

Summary Permian bryozoan faunas from the Lower Permian sequences of the Carnic Alps (UpperPseudoschwagerina Formation and Trogkofel Formation) and from some other Permian units of the NW-Tethys (Sicily, Tunisia) include cystoporid, trepostomid, fenestellid, rhabdomesid, and timanodictyid taxa. Fenestellids and cystoporids species dominate. The Lower Permian bryozoan fauna of the Carnic Alps displays close relations to faunas of Sakmarian-Artinskian age of the Russian Platform and Pamir as well as of the Lower Permian of Australia. Bryozoans from Permian sequences of Sicily and Tunisia display relations to the Permian faunas of Indonesia and Australia.     

New observations on the skeletons of the earliest bryozoans from the Fenhsiang Formation (Tremadocian,Lower Ordovician), Yichang,China

The earliest unequivocal bryozoans occurred in the Fenhsiang Formation (upper Tremadocian, Lower Ordovician) of Yichang, South China, providing important clues about the early evolution of primitive bryozoans. In this study, SEM and EDAX were used to analyze the microstructure of the walls of two bryozoan genera, Nekhorosheviella and Orbiramus, from the Fenhsiang Formation. All walls of Nekhorosheviella and endozonal walls of Orbiramus show poor preservation in thin sections, with a granular appearance reflecting extensive neomorphism, whereas exozonal walls of Orbiramus are distinctly laminated. These preservational differences may reflect skeletal chemistry, particularly the magnesium content of the calcite. Phosphatic skeletal linings were found to be distributed unevenly in the autozooecia of Nekhorosheviella, but were absent in Orbiramus.     

A molecular phylogeny of bryozoans

We present the most comprehensive molecular phylogeny of bryozoans to date. Our concatenated alignment of two nuclear ribosomal and five mitochondrial genes includes 95 taxa and 13,292 nucleotide sites, of which 8297 were included. The number of new sequences generated during this project are for each gene:ssrDNA (32), lsrDNA (22), rrnL (38), rrnS (35), cox1 (37), cox3 (34), and cytb (44). Our multi-gene analysis provides a largely stable topology across the phylum. The major groups were unambiguously resolved as (Phylactolaemata (Cyclostomata (Ctenostomata, Cheilostomata))), with Ctenostomata paraphyletic. Within Phylactolaemata, (Stephanellidae, Lophopodidae) form the earliest divergent clade. Fredericellidae is not resolved as a monophyletic family and forms a clade together with Plumatellidae, Cristatellidae and Pectinatellidae, with the latter two as sister taxa. Hyalinella and Gelatinella nest within the genus Plumatella. Cyclostome taxa fall into three major clades: i. (Favosipora (Plagioecia, Rectangulata)); ii. (Entalophoroecia ((Diplosolen, Cardioecia) (Frondipora, Cancellata))); and iii. (Articulata ((Annectocyma, Heteroporidae) (Tubulipora (Tennysonia, Idmidronea)))), with suborders Tubuliporina and Cerioporina, and family Plagioeciidae each being polyphyletic. Ctenostomata is composed of three paraphyletic clades to the inclusion of Cheilostomata: ((Alcyonidium, Flustrellidra) (Paludicella (Anguinella, Triticella)) (Hislopia (Bowerbankia, Amathia)) Cheilostomata); Flustrellidra nests within the genus Alcyonidium, and Amathia nests within the genus Bowerbankia. Suborders Carnosa and Stolonifera are not monophyletic. Within the cheilostomes, Malacostega is paraphyletic to the inclusion of all other cheilostomes. Conopeum is the most early divergent cheilostome, forming the sister group to ((Malacostega, Scrupariina, Inovicellina) ((Hippothoomorpha, Flustrina) (Lepraliomorpha, Umbonulomorpha))); Flustrina is paraphyletic to the inclusion of the hippothoomorphs; neither Lepraliomorpha nor Umbonulomorpha is monophyletic. Ascophorans are polyphyletic, with hippothoomorphs grouping separately from lepraliomorphs and umbonulomorphs; no cribrimorphs were included in the analysis. Results are discussed in the light of molecular and morphological evidence. Ancestral state reconstruction of larval strategy in Gymnolaemata revealed planktotrophy and lecithotrophy as equally parsimonious solutions for the ancestral condition. More comprehensive taxon sampling is expected to clarify this result. We discuss the extent of non-bryozoan contaminant sequences deposited in GenBank and their impact on the reconstruction of metazoan phylogenies and those of bryozoan interrelationships.     

Frasnian bryozoans (Late Devonian) from the Khoshyeilagh Section,Alborz Mountains (northern Iran)

A bryozoan fauna containing seven species is described from the Upper Frasnian (Upper Devonian) rocks in the Khoshyeilagh Section, Alborz Mountains (northern Iran). The studied bryozoan assemblage includes one new trepostome species, Eridotrypella alborzensis sp. nov., an additional four species identified at species level: two trepostomes, Minussina akkayaensis Volkova, 1974 and Leptotrypella inaudita Morozova, 1961, and two rhabdomesine cryptostomes, Bigeyella mariae (Morozova, 1961) and Saffordotaxis multispinata (Morozova, 1955). Furthermore, two species are described in open nomenclature: the trepostome Schulgina sp. and the rhabdomesine cryptostome Nicklesopora sp. The studied fauna shows a close similarity at the species level between northern Iran and the Altai-Sayan Folded Belt (Russia), south China, and Transcaucasia.