During the Early Ordovician Epoch, the Mediterranean brachiopod Province was extensive in the higher-latitude sectors of the globe in the Southern Hemisphere. The latter was much occupied by the massive continent of Gondwana, which stretched from north of the Equator S-wards to cover the South Pole. The Mediterranean Province can be separated into two groups: Group 1, the higher-latitude fauna dominated by large linguliform brachiopods; and Group 2, which is more diverse, particularly in orthides. The large linguliform brachiopod faunas are particularly well known in southern Europe (France, Spain and Bohemia) and North Africa, and the second group in Avalonia, Chile and Argentina. The province is different from, but merges with, more diverse contemporary faunas in the lower latitudes of Gondwana to its north, although the latter contrast with other lower-latitude faunal provinces in South China, Laurentia, Siberia and elsewhere. Since the Rheic Ocean between Avalonia and Gondwana was relatively narrow during the Early Ordovician Epoch, the Avalonian brachiopods were integral parts of the Mediterranean Province, but only until end of the Dapingian Age. This paper focuses on the earlier phases of the Mediterranean Province, although the province continued until near the end of the Ordovician Period. Intermediate-latitude Baltica and some other faunas are included in new principal components and other analyses in order to compare them with the Mediterranean Province faunas. Radiation was very significant for many brachiopod taxa during the period, with first appearances of the Plectambonitoidea (Taffiidae), several orthide families (Euorthisinidae, Tarfayidae and Anamalorthidae) and the earliest endopunctate orthide, the dalmanelloid Lipanorthis.

We present a δ13Ccarb chemostratigraphy for the Late Ordovician Hirnantian Stage based on 208 whole-rock samples from six outcrops in the Oslo–Asker district, southern Norway. Our data include the Norwegian type section for the Hirnantian Stage and Ordovician–Silurian boundary at Hovedøya Island. The most complete record of the Hirnantian Isotope Carbon Excursion (HICE) is identified in a coastal exposure at Konglungø locality where the preserved part of the anomaly spans a c. 24 m thick, mixed carbonate–siliciclastic succession belonging to the upper Husbergøya, Langåra and Langøyene formations and where δ13Ccarb peak values reach c. +6 ‰. Almost the entire HICE occurs above beds containing the Hirnantia Fauna, suggesting a latest Hirnantian age for the peak of the excursion. The temporal development of the HICE in southern Norway is associated with substantial shallowing of depositional environments. Sedimentary facies and erosional unconformities suggest four inferably fourth-order glacio-eustatically controlled sea-level lowstands with successively increased exposure and erosion to the succession. The youngest erosional unconformity is related to the development of incised valleys and resulted in cut-out of at least the falling limb of the HICE throughout most of the Oslo–Asker district. The fill of the valleys contains the falling limb of the HICE, and the postglacial transgression therefore can be assigned to the latest part of the Hirnantian Age. We address the recent findings of the chitinozoan Belonechitina gamachiana in the study area and its relationship to the first occurrence of Hirnantia Fauna in the studied sections, challenging identification of the base of the Hirnantian Stage.

Calcium carbonate biominerals are frequently analysed in materials science due to their abundance, diversity and unique material properties. Aragonite nacre is intensively studied, but less information is available about the material properties of biogenic calcite, despite its occurrence in a wide range of structures in different organisms. In particular, there is insufficient knowledge about how preferential crystallographic orientations influence these material properties. Here, we study the influence of crystallography on material properties in calcite semi-nacre and fibres of brachiopod shells using nanoindentation and electron backscatter diffraction (EBSD). The nano-indentation results show that calcite semi-nacre is a harder and stiffer (H ≈ 3—5 GPa; E = 50–85 GPa) biomineral structure than calcite fibres (H = 0.4—3 GPa; E = 30—60 GPa). The integration of EBSD to these studies has revealed a relationship between the crystallography and material properties at high spatial resolution for calcite semi-nacre. The presence of crystals with the c-axis perpendicular to the plane-of-view in longitudinal section increases hardness and stiffness. The present study determines how nano-indentation and EBSD can be combined to provide a detailed understanding of biomineral structures and their analysis for application in materials science.

The extent of racemization of aspartic acid (Asp) has been used to estimate the ages of 9 shells of the epifaunal calcitic brachiopod Bouchardia rosea and 9 shells of the infaunal aragonitic bivalve Semele casali. Both taxa were collected concurrently from the same sites at depths of 10 m and 30 m off the coast of Brazil. Asp D/L values show an excellent correlation with radiocarbon age at both sites and for both taxa (r2Site 9 B. rosea  = 0.97, r2Site 1 B. rosea = 0.997, r2Site 9 S. casali = 0.9998, r2Site 1 S. casali = 0.93). The Asp ratios plotted against reservoir-corrected AMS radiocarbon ages over the time span of multiple millennia can thus be used to develop reliable and precise geochronologies not only for aragonitic mollusks (widely used for dating previously), but also for calcitic brachiopods. At each collection site, Bouchardia specimens display consistently higher D/L values than specimens of Semele. Thermal differences between sites are also notable and in agreement with theoretical expectations, as extents of racemization for both taxa are greater at the warmer, shallower site than at the cooler, deeper one. In late Holocene marine settings, concurrent time series of aragonitic and calcitic shells can be assembled using Asp racemization dating, and parallel multi-centennial to multi-millennial records can be developed simultaneously for multiple biomineral systems.

In the Saoura Valley (Ougarta Basin, Saharan Algeria), the lower–upper Famennian part of the essentially shally Marhouma Formation is characterized by deep-water facies and includes horizons rich in ammonoids (goniatites and clymeniids) and blind to eye-reduced phacopide trilobites. They are also rich in small-sized and smooth rhynchonellide brachiopods, investigated here for the first time in order to detail their post-Kellwasser recovery. Rhynchonellides clearly predominate in the brachiopod assemblages (representing 90% of the whole assemblage, with 10 species) composed otherwise of athyridides, orthides and spiriferides. Rhynchonellides are mostly represented by relatively flat leiorhynchids and rozmanariids consistent with poor oxygenation on the sea floor. One new species is described (Evanidisinurostrum saouraense sp. nov.); four genera, previously known only from the south-eastern margin of Laurussia, are reported for the first time from the northern margin of Gondwana: the leiorhynchid Sphaeridiorhynchus and the rozmanariids Leptoterorhynchus, Pugnaria and Novaplatirostrum.

Late Ordovician fossils, including the distinctive Hirnantia brachiopod Fauna, have been found in Lower Palaeozoic successions in Istanbul and Bolu (Yığılca), western Pontides, NW Turkey. The Middle Ordovician (Sandbian) faunas belong to the cooler-water Mediterranean Province, and they are followed by Katian brachiopods including Sulevorthis, Nicolella, Hesperorthis, Glyptorthis, Saukrodictya and Kullervo and ostracods such as Piretella, Eochilina and Klimphores, which represent deposition in warmer waters; however, the Mediterranean Province usually cooler-water brachiopods Drabovia and Leptestiina also occur. The Pendik Formation includes thin bryozoan-rich limestones which probably represent the Boda Global Warming Event. The overlying turbidites contain a Hirnantia Fauna, developed within a brachiopod–diplograptid association. Above them there are characteristic Llandovery (Rhuddanian–Aeronian) brachiopods, such as Leangella, Eoplectodonta, Stricklandia and Hindella with the corals Halysites, Paleofavosites and Streptelasma. In the Bolu area, Katian brachiopods such as Mcewanella, Dalmanella, Glyptorthis, Christiania, Oligorhynchia, Nicolella, Howellites and Drabovinella also occur, but there the overlying Hirnantia Fauna is developed within a Hirnantia–Mucronaspis association. The fauna and sediments indicate that the western Pontides were not very cold during the latest Ordovician. Despite Turkey being placed in higher latitudes by previous authors, it seems more probable that the Pontides were at somewhat lower palaeolatitudes, perhaps at about 40°S in those times; however, the precise palaeogeographical position of the terrane remains uncertain: there are no Hirnantian glaciogenic rocks there, such as are found in the Taurides of southern Turkey.

A detailed ecostratigraphical framework is established for the lower Middle Ordovician Kundan regional stage of the East Baltic area corresponding to the Asaphus expansus, A. raniceps and A. eichwaldi trilobite zones (lower Darriwilian). The study is based on approximately 6200 brachiopods collected bed by bed from limestone sections in northern Estonia (Harku Trench and Saka) and western Russia (Putilovo Quarry, Lava River canyon and Lynna River valley) with, in addition, the first detailed systematic assessment of the Kundan brachiopods of the East Baltic. These sections represent an oblique depth transect some 400 kilometres long, deepening eastwards. Five biofacies associations have been recognized using detrended correspondence and cluster analyses: a shallow-water Lycophoria association, a transitional Gonambonites association and two deeper-water associations, the soft-substrate Orthis callactis and the hard-substrate Orthambonites associations. A separate, fifth soft-substrate association is present in the marl beds at the main locality of Putilovo Quarry. The associations reflect a combination of palaeo-water depth and substrate. The biofacies facilitate an ecostratigraphical correlation along the transect, and third and fourth order sea-level curves are reconstructed, reflecting mainly eustasy. The sea-level was relatively low, early in the Kundan, but then rose significantly into the A. raniceps Biozone. This corroborates the recent discovery of possible small early Darriwilian ice caps on Gondwana.

A new species of the lacazelline brachiopod genus Ospreyella is described from shallow cryptic habitats in Palau, western Caroline Islands and Arnd Atoll, Pohnpei, eastern Caroline Islands, north-western Pacific Ocean. This new taxon, here named Ospreyella palauensis, and representing the third discovery of a species of Ospreyella from the Indo-Pacific region, is compared with the other two recently described extant species of the genus, O. depressa Lüter from the Great Barrier Reef, Australia, and O. maldiviana Logan from the Maldive Islands, northern Indian Ocean. An ontogenetic sequence is described for the new species and compared with those previously described for other Holocene lacazellines. Additionally, a study of ontogenetic stages in Thecidellina congregata from Saipan in the Northern Mariana Islands shows early-stage similarities to those of the new species and other lacazellines. These observations on development in extant forms provide important analogies for comparison with ontogenetic stages in ancestral fossil forms and information which may be useful in reconstructing thecideide phylogenetic history.

Forty-one Lower Silurian brachiopods from a single locality in the Newlands Formation of Girvan, Strathclyde, Scotland, are described and illustrated, many for the first time. The age of the fauna is well constrained by overlying magnus Biozone graptolites and underlying cyphus Biozone graptolites, indicating the early Middle Llandovery (Lower Aeronian) and it is the only shelly fauna known of Lower Aeronian age from anywhere in Scotland. A new endemic genus Chronostrophonella (Family Strophonellidae) is described: another new genus, Eopentamerus (Family Pentameridae) is known additionally from Wales. New species are Craniops laurentia, Biparetis caledonia, Eostropheodonta augusta, Chronostrophonella terranova, Strophochonetes newlandensis, Triplesia girvanensis, Resserella praeclara, Camerella? elainae, Eopentamerus inexpectatus and Protatrypa copperi. The Strophochonetes is the earliest-known chonetoid from the Silurian anywhere, although the family originated in the Late Ordovician. Whilst Newlands was on the margin of Laurentia just prior to its Caledonian Orogeny collision in mid-Silurian times with Avalonia–Baltica, the brachiopod fauna there was already similar in the Lower Aeronian, at least at the generic level, to those from the opposite side of the Iapetus Ocean in Wales and the Welsh Borderland (Avalonia) and Norway and other parts of Baltica. The proximity of these and other terranes to each other was the key reason for the relative cosmopolitanism of all lower-latitude Lower Silurian brachiopod faunas.

Eight taxa of marine invertebrates, including two new bivalve species, are described from the Low Head Member of the Polonez Cove Formation (latest early Oligocene) cropping out in the Vauréal Peak area, King George Island, West Antarctica. The fossil assemblage includes representatives of Brachiopoda (genera Neothyris sp. and Liothyrella sp.), Bivalvia (Adamussium auristriatum sp. nov., ?Adamussium cf. A. alanbeui Jonkers, and Limatula (Antarctolima) ferraziana sp. nov.), Bryozoa, Polychaeta (serpulid tubes) and Echinodermata. Specimens occur in debris flows deposits of the Low Head Member, as part of a fan delta setting in a high energy, shallow marine environment. Liothyrella sp., Adamussium auristriatum sp. nov. and Limatula ferraziana sp. nov. are among the oldest records for these genera in King George Island. In spite of their restrict number and diversification, bivalves and brachiopods from this study display an overall dispersal pattern that roughly fits in the clockwise circulation of marine currents around Antarctica accomplished in two steps. The first followed the opening of the Tasmanian Gateway at the Eocene/Oligocene boundary, along the eastern margin of Antarctica, and the second took place in post-Palaeogene time, following the Drake Passage opening between Antarctic Peninsula and South America, along the western margin of Antarctica.

This study describes bioerosion traces ascribed to either predation or endo- and epibiont activity in twenty assemblages from the Mediterranean region and Paratethys, spanning in age from Eocene to Recent. Statistical analysis of the distribution of bioerosion traces among genera and assemblages revealed that there is higher drilling predation intensity on smaller species. Larger species seem to be primarily affected by non-drilling predators. Greatest variety in types of bioerosion could be related to species’ ecology and body size. Both major categories of bioerosion (etchings and traces of predatory activity) vary considerably among samples. Different genera show significant differences in the frequency of different bioerosion types. Shell size seems a major factor contributing to these differences.

With their extensive fossil record and shells of stable low-Mg calcite, rhynchonelliform brachiopods are attractive sources of climate information via seawater temperature proxies such as stable oxygen isotope composition. In Terebratalia transversa (Sowerby) there is a progression towards oxygen isotope equilibrium in the calcite of the innermost secondary layer. This study confirms the lack of any vital effects influencing oxygen isotope composition of T. transversa, even in specialised areas of the innermost secondary layer. Calcite Mg/Ca ratio is another potential seawater temperature proxy, that has the advantage of not being influenced by salinity. Mg concentrations measured by electron microprobe analyses indicate that there is no concomitant decrease in Mg concentration towards the inner secondary layer, associated with the progressive shift towards oxygen isotope equilibrium. Mg distribution is heterogeneous throughout the shell and correlates with that of sulphur, which may be a proxy for organic components, suggesting that some of the Mg may not be in the calcite lattice. It is essential therefore, to determine the chemical environment of the magnesium ions to avoid any erroneous temperature extrapolations in brachiopods or any other calcite biomineral.

A new Middle Permian locality in northern Johore, Peninsular Malaysia, yields a small-sized, but compositionally unique, brachiopod fauna consisting of eight species: Pseudoleptodus sp., Caricula cf. salebrosa Grant, Neochonetes (Nongtaia) aff. arabicus (Hudson & Sudbury), Karavankina sp., Transennatia cf. insculpta (Grant), Hustedia sp., Orthothetina sp., and martiniid indet. The first four genera are new records for Malaysia; in particular, the rare taxa Pseudoleptodus and Caricula characterize the fauna. The brachiopods occur together with the ammonoid Agathiceras sp., the nautiloid Foordiceras? sp., bivalves, and crinoid stems. The locality belongs to the East Malaya terrane of the Cathaysian biotic region, but some affinities to species of the Sibumasu province are recognized. The Malaysian forms of Pseudoleptodus, Caricula and Transennatia are similar to those of the Ratburi Limestone (southern Thailand). A Roadian–early Wordian age is interpreted for the Johore fauna. The similarity of brachiopods reported here with those from the Ratburi Limestone suggests that there was species interchange or one-way migration between shallow waters of East Malaya and Sibumasu across the main Palaeo-Tethys. The Tethyan seaway between the two terranes must have been narrower than previously interpreted by some authors to allow such faunal traffic during the Roadian–Wordian time period.

A new occurrence of the Hirnantia brachiopod fauna is documented from the Criadero Quartzite of Almadén, Ciudad Real Province, Spain. This unit is the regional development of a largely unfossiliferous sandy facies that frequently overlies the typical Late Ordovician diamictitic glaciomarine formations in the Iberian Peninsula and the Armorican Massif. The new occurrence establishes palaeontologically the latest Ashgill age of the quartzite, at least for its lowest horizons, and adds new data on a fauna that, although widespread, has been very rarely documented from peri-Gondwanan Europe. The new collection contains only Hirnantia sagittifera and Plectothyrella crassicosta chauveli. The subspecific status of the latter and its inclusion within Plectothyrella crassicosta is discussed herein, based on the continuous variation in rib thickness of several samples of both forms. The extremely low diversity and the occurrence of the key form P. c. chauveli, are both typical of the Bani Province that developed on the subpolar margins of Gondwana. This contrasts with other occurrences of the Hirnantia Fauna in peri-Gondwanan Europe, such as those from Sardinia and the Carnic Alps, which are characteristic of the more temperate Kosov Province.

A global review of new and existing data on the distribution of uppermost Ordovician (Hirnantian) brachiopods indicates the existence of at least three biogeographically distinct faunas. The typical Hirnantia fauna characterised subtropical and temperate latitudes and comprised a variety of ecological associations; the fauna reached its acme during the bohemicus and uniformis zones. Atypical Hirnantia faunas, developed marginal to Gondwana, are of low diversity and have few species in common with the typical Hirnantia fauna; their spatial distribution probably marked the margin of the polar ice sheets. The extinction of the Hirnantia fauna occurred in response to changes in sea level. Diverse and quite different faunas, including those from the Midcontinent of North America, Kolyma, the Oslo Region and probably Anticosti Island, occupied equatorial latitudes during the latest Ordovician. The Holorhynchus fauna, on evidence to date, predates the Hirnantia fauna.