The interpretation of pedicle soft tissue preservation in a unique brachiopod specimen of Wenlock (Silurian) age from Herefordshire, western England, is re-assessed. Bethia serraticulma, assigned originally to the Orthida, is more probably a member of the Strophomenida (Plectambonitoidea). The supposed pedicle structure is more plausibly a weakly mineralised pedicle sheath, which is a common morphological and functional development in the early ontogeny of a number of Palaeozoic brachiopod lineages.

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.

Two Ordovician plectambonitoid genera, Alwynella and Grorudia, occur in drill core sections of Latvia in the East Baltic, and in exposures and loose blocks on the Swedish Island of Öland in the Baltic Sea. The new material confirms differences between the two taxa that are assigned herein to separate families, Alwynellidae fam. nov. and Grorudiidae Cocks & Rong, 1989. In particular, the undercut cardinalia separates Alwynella from Grorudia and indicates its proximity to the sowerbyellids. The genus Grorudia, which is externally similar to Alwynella, is more closely related to the palaeostrophomenines. A new species Grorudia morrisoni sp. nov. is established in the East Baltic. The specimens from Öland are included tentatively within the genus Grorudia due to lack of interiors. Both Alwynella and Grorudia were confined to deeper-water facies in the Baltic palaeobasin, within successions ranging in age from latest Mid (late Llanvirn) to earliest Late Ordovician (mid Caradoc).

The records of columnar shell structures of linguloid brachiopods (Class Lingulata, Order Lingulida, Superfamily Linguloidea) are reviewed in the light of the discovery of two new taxa from the Middle Cambrian Forsemölla Limestone Bed of southern Sweden. The linguloid taxa, described here as Eoobolus? sp. aff. E. priscus (Poulsen) and Canalilatus? simplex sp. nov., are both characterised by a columnar shell structure, a structural type that is representative for acrotretoid brachiopods and that has previously only rarely been reported from the linguloids. Though the two taxa are superficially similar to known genera, i.e., Eoobolus and Canalilatus, their shell structure challenges such affiliations, as the shell structure of the type species of these genera is previously unknown. Linguloid families whose morphological characteristics agree the most with those of the new taxa, i.e., the Zhanatellidae and the Eoobolidae, and from which columnar shell structures have been reported, i.e., the Lingulellotretidae and the Kyrshabaktellidae, are reviewed briefly. Many taxa assigned to these families completely lack shell structure data and are in need of restudy in order to elucidate their systematic position. Knowledge of the representative type of shell structure of the various suprageneric taxa within the Linguloidea is considered crucial, in order to unravel their currently poorly resolved phylogenetic relationships.

The loop configuration, structure and growth are studied in a series of megathiridoid dorsal valves ranging in length from 0·7 to 6 mm. The structure of the posterior and anterior sectors of the descending lamellae and their relationships with the socket ridges, dorsal septa and dorsal valve floor, are studied, illustrated and described. Detailed studies show that internal structures display great variability. In the megathiridoids the lamellae of more typical terebratulide loops are replaced by partially developed loops in early developmental stages, resulting in a mix of free sections of loop together with sections where the lophophore rests directly on the epithelium of the valve floor. Comments on the ecology of the megathiridoid species are included and relate their unusually wide gapes, which position their lophophores fully accessible to the open sea, with their strong lophophore attachments and loss of typical free loops. The new genus Joania (type species Terebratula cordata Risso, 1826) is erected for those Argyrotheca which, although having a typical megathiridoid brachidium, differ in their adult crural development, their narrow hinge line, their prominent cardinal process, their characteristic dorsal median septum and their tuberculate radial ridges terminating anteriorly in tubercles.

The Lower Permian of the Haushi basin, Interior Oman (Al Khlata Formation to Saiwan Formation/lower Gharif member) records climate change from glaciation, through marine sedimentation in the Haushi sea, to subtropical desert. To investigate the palaeoclimatic evolution of the Haushi Sea we used O, C, and Sr isotopes from 31 brachiopod shells of eight species collected bed by bed within the type-section of the Saiwan Formation. We assessed diagenesis by scanning electron microscopy of ultrastructure, cathodoluminescence, and geochemistry, and rejected fifteen shells not meeting specific preservation criteria. Spiriferids and spiriferinids show better preservation of the fibrous secondary layer than do orthotetids and productids and are therefore more suitable for isotopic analysis. δ18O of −3·7 to −3·1℅ from brachiopods at the base of the Saiwan Formation are probably related to glacial meltwater. Above this, an increase in δ18O may indicate ice accumulation elsewhere in Gondwana or more probably that the Haushi sea was an evaporating embayment of the Neotethys Ocean. δ13C varies little and is within the range of published data: its trend towards heavier values is consistent with increasing aridity and oligotrophy. Saiwan Sr isotope signatures are less radiogenic than those of the Sakmarian LOWESS seawater curve, which is based on extrapolation between few data points. In the scenario of evaporation in a restricted Haushi basin, the variation in Sr isotope composition may reflect a fluvial component.

Like most metazoan biomineralisations, the brachiopod shell is the end product of a biologically controlled calcification process. The main agent of the control is the extracellular matrix, which is secreted by the outer mantle epithelium. This matrix mediates the calcification process by allowing crystal nucleation and elongation in specific orientations and finally, by stopping crystal growth. The proteinaceous moiety of brachiopod shell matrices has been extensively studied. Less known are the post-translational modifications that occur in these matrices, in particular glycosylations. In this comparison of five species of Recent articulated brachiopods, the ratio of soluble to insoluble organic matrix varies between the species. Polydisperse macromolecular materials occur in each of these species with discrete proteins of 50 kDa in Notosaria nigricans, Calloria inconspicua and Neothyris lenticularis, 37 kDa in Terebratulina retusa and Gryphus vitreus and 20–25 kDa in N. nigricans. Protein mixtures from all five species respond differently to anionic stains (Stains-All and Alcian Blue). PAS staining results in a positive smear in C. inconspicua and T. retusa and highlights low molecular weight glycoproteins in C. inconspicua. The polysaccharide composition of the soluble matrix of T. retusa is different from the others due to high proportions of arabinose and low proportions of fucose. In all cases, polysaccharide composition of the insoluble matrix is dominated by glucose and glucosamine. Insoluble matrices have more glucose and xylose and less galactosamine and glucosamine than the corresponding soluble matrix. Relatively high amounts of glucosamine may suggest the presence of chitin in the shell matrix of rhynchonelliform brachiopods.

Earliest Silurian (basal Llandovery) brachiopod faunas are surveyed and listed from around the globe, and divided between Lower Rhuddanian and Upper Rhuddanian occurrences. 60 genera are known from the Lower Rhuddanian within 20 superfamilies and there are 87 genera in 25 superfamilies in the Upper Rhuddanian. The 29 areas surveyed span the globe, both latitudinally and longitudinally. Only six superfamilies are Lazarus taxa which are known both from the Ordovician and Middle Llandovery (Aeronian) and later rocks but have not been recorded from the Rhuddanian. These are surprising results, since many previous studies have inferred that the Rhuddanian was a time of very sparse faunas. The global warming that followed the latest Ordovician (Hirnantian) ice age did not proceed quickly, with an ice-cap probably present through at least the Llandovery. There is a marked absence of Lower Rhuddanian bioherms even at low palaeolatitudes; however, the ecological recovery rate was far faster than that following the end-Permian mass extinction event. The partitioning of the Rhuddanian shelf faunas into well-defined benthic assemblages progressed slowly over the interval.

Ultrastructural examination of the brachiopod Thecidellina from three different locations in the Caribbean and the Atlantic revealed that at least three cryptic species are present. One is the type species of the genus, Thecidellina barretti, which may only occur in Jamaica. The other two, previously lumped into T. barretti, are new to science, viz T. bahamiensis Lüter & Logan sp. nov. from the Bahamas and T. williamsi Lüter & Logan sp. nov. from Cape Verde. All three species clearly differ not only in their provenance, but also in specific shell characters, such as spiculation of the ventral valve, presence of a ventral median ridge, shape of two holes in the intrabrachial ridge of the dorsal valve and the structure of the interbrachial lobes. The identification of three instead of a single species and their supposed allopatric speciation is discussed with regard to the thecideide life cycle and independent models of larval dispersal in the Caribbean, based on oceanographic data.

A new rhynchonellide brachiopod has been collected from the Kahuitara Tuff (Campanian–Maastrichtian) of Pitt Island, Chatham Islands, New Zealand. Brachiopods are extremely rare in Cretaceous rocks from New Zealand, and this new genus and species is unlike any other rhynchonellide known from Australasia or elsewhere. Chathamirhynchia kahuitara is distinguished by its small size, strong ribbing, and well-developed sulciplicate folding, and internally by a large, robust cardinal process and raduliform crura. The shell structure of C. kahuitara is shown to be of typical fine fibrous ‘rhynchonellidine’ type. This brachiopod was probably strongly attached to volcanic rock or shells in a shallow-water, high-energy environment. Implications for the biogeography of brachiopods during the Late Cretaceous are briefly discussed.

A review is presented of the genera and species, both living and fossil, comprising the Superfamily Kraussinoidea and of their geographic and stratigraphic distributions. Two new fossil species are described, Kraussina chilensis and Megerlina miracula, from the Miocene of Chile and New Zealand respectively. The ontogenetic development of the brachidium in each of the four kraussinid genera is described and on the basis of recognised differences it is proposed that a new taxonomic subdivision is made that places Megerlia in a new subfamily Megerliinae, distinct from the other three genera in the existing subfamily Kraussininae Dall. The loop ontogeny of kraussinoids is compared with that of other terebratellidines and a phylogenetic linkage between kraussinoids and laqueoids is suggested.

This study explores the implications of different decisions about taxon sampling in studies of phylogeny and macroevolution – how would phylogenetic results differ as decisions about inclusion of taxa differed? The focus is on investigating phylogenetic relationships among families of Devonian terebratulides, and include all 71 named genera in our analyses. Subsets of taxa were experimented with the most complete morphologically from fossil specimens; those that occur earliest in the stratigraphic record; those that include only the name-bearing genera from each family. Including only the most completely known genera produces a result essentially similar to one including all genera, even those for which less than half the characters can be coded. Including only the stratigraphically earliest genera produces a result at odds with the other analyses. Including only name-bearers, representing 13% of all genera, produces a result generally similar to the analysis including all taxa. None of the results of these phylogenetic experiments involving subsets of genera corresponds strongly with the recently revised classification in the Treatise on Invertebrate Paleontology, but general similarities can be discerned. The lack of strong correspondence between classification and several different experimental phylogenetic hypotheses could be ascribed to their different overall goals, and highlights the potential dangers of ascribing evolutionary significance to simple counts of taxa, particularly families, as warned by Alwyn Williams 50 years ago.

A new genus of pentameride brachiopod, Costilamnulella, described herein from the middle Ashgill of Sweden, is particularly interesting in that the ventral valve possesses both an interarea and a palintrope. Brachiopods normally have one or the other, but both structures do occur amongst the pentamerides, e.g. in Clorinda, Anastrophia and Brevilamnulella. The presence of these structures in three separate pentameride families indicates the persistence of the more primitive condition of an interarea whilst the more advanced palintrope is becoming established. Costilamnulella is interpreted as a restricted side line of the main development of the more widely occurring Brevilamnulella, which survived the end-Ordovician event and may have shared its ancestry with the ubiquitous Clorinda.

The lower Meitan Formation (Floian, upper Lower Ordovician) at the Dajiaosi section, Zunyi District, northern Guizhou Province of South China, contains a moderately rich and diverse Sinorthis Fauna, with 22 species attributable separately to 15 families and seven orders of brachiopods. The fauna can be differentiated into three associations: the Paralenorthis serica, the Sinorthis typica, and the Tarfaya intercalare associations. These occupied a relatively wide palaeoecological range from lower BA3 to upper BA2 settings and from silty to clay substrate conditions. In South China, the Early Ordovician brachiopod radiation was marked by the diversification of the orthide-dominated Sinorthis Fauna, which first appeared and diversified in the middle part (relatively deep water) of the Upper Yangtze Platform in the Didymograptellus eobifidus Biozone, but rapidly declined in the succeeding Corymbograptus deflexus and basal Azygograptus suecicus biozones. During the latest Floian and early Dapingian, it expanded into comparatively shallow-water settings in the onshore direction (Changning area, southern Sichuan), offshore carbonate platform (Yichang, Hubei), and areas adjacent to the submergent Qianzhong Arch (such as the Zunyi area). The first appearance datum of the Sinorthis Fauna in shallower-water settings generally postdates that in the deeper-water environment in the central Upper Yangtze Platform, probably as a result of the fauna tracking a favoured BA3 setting during a gradual marine transgression.

Locally abundant and diverse brachiopod faunas, associated with unstable outer shelf and slope environments, occur throughout the Barr and Ardmillan groups (middle Llanvirn–upper Ashgill) in the Girvan district of SW Scotland. A dataset of 350 brachiopod species from 30 horizons through the Middle–Upper Ordovician succession forms the basis for a description of brachiopod diversity through the succession and comparison with global patterns and trends through this time interval. The Middle Ordovician Barr Group incorporates shallow water carbonate and clastic facies, characterised by Valcourea confinis, with deeper-water facies. The trilobite-dominated Albany Group preserves outer shelf biofacies. Deep-water facies occur in the Balclatchie Formation of the lower Ardmillan Group, including an early occurrence of a Foliomena-type species. High diversity brachiopod faunas occur in clastic facies, though many of the biofacies have a transported component to them. Representatives of the deep-water Foliomena fauna occur intermittently throughout the Upper Ardmillan Group, appearing in both the Whitehouse and Drummuck subgroups. This distinctive assemblage of small, thin-shelled brachiopods, including Dedzetina, Christiania, Cyclospira and Foliomena itself, is interbedded with a variety of other less cosmopolitan deep-water assemblages, including the Onniella–Skenidioides and Lingulella–Trimurellina associations. Shallower-water environments in the middle Ashgill Lower Drummuck Subgroup hosted the Fardenia–Eopholidostrophia association in sands, and the Christiania–Leptaena association in muds and silts. The remarkable Lady Burn Starfish Beds in the upper part of the group contain a variety of brachiopod-dominated assemblages, including the Eochonetes and Plaesiomys–Schizophorella associations, transported from various shelf locations, within a very diverse mid Ashgill biota. The upper Ashgill High Mains Formation contains abundant elements of the terminal Ordovician Hirnantia fauna, including Eostropheodonta, Hindella and Hirnantia itself, but also some taxa more typical of the Laurentian Edgewood Province. As a whole, the changing brachiopod biodiversity biofacies reflect environmental fluctuations, on this part of the Laurentian margin, driven by mainly eustatic and tectonic events against a background of global biotic radiation.

We consider the reinterpretation of Bethia by Bassett et al. (2008) to be flawed in several regards. Details of the pedicle rootlets and wrinkles strongly imply a soft-tissue structure rather than calcified sheath. The ornament is unusual, but equally so under either an orthide or a plectambonitoidean model. The reinterpretation of the ‘deltial plates’ as ‘chilidial plates’ is likely correct, but argues against a plectambonitoidean affinity by implying that the pedicle emerges between the valves. Other arguments presented in favour of a plectambonitoidean affinity are also discussed; we consider them unpersuasive. Finally, we contend that Bethia is sufficiently well characterised to deserve a taxonomic name.

Electron backscatter diffraction (EBSD) is used to determine the detailed crystallographic orientation of calcite crystals of craniid brachiopods in the context of shell ultrastructure. Sections of shells of two Recent species, Novocrania anomala and Novocrania huttoni, are analysed to provide 3D crystallographic patterns at high spatial resolution. The c-axis of semi-nacre calcite crystals is oriented parallel to the laminae that define the ultrastructure of the secondary layer. This orientation differs from that of rhynchonelliform calcitic brachiopods where the c-axis is perpendicular to the length of morphological fibres and to the shell exterior.

Siphonotretid brachiopod distribution in the Lower Ordovician of the Prague Basin is described and discussed; the new species Alichovia cometa is established. The preservation and distribution of the group indicate a preference for shallow-water offshore, high-energy environments: siphonotretids are abundant shelly fossils in the shallower-water, mostly marginal sites of the basin but are absent in the deeper parts. Their environmental limits were complementary to those of the graptolites; siphonotretids co-occur with epiplanktonic graptolites at only a few localities. Deeper offshore deposits with biramose dichograptids and diverse, generally delicate benthic dendroids lack siphonotretids. Indeed, siphonotretids were intolerant to dysoxia and preferred a well-aerated siliciclastic, firm sea floor in the shallower parts of the basin. They were tolerant to suspended coarser detritus, using their spines which functioned as an effective sieve. The density and ramification of spines indicate three sifting strategies. Suggested life styles above the seabed on elevated surfaces (algae, sponges, dendroids) are not supported by direct evidence. They more likely lived within more supple and soft matrices (sponges, tufts of algae) stabilised by their spines, with the spinose basket maintaining free space for inhalant and exhalant currents.

The size of the protegulum on the valves of Early Palaeozoic rhynchonelliform brachiopods has previously been used as a criterion to assign various taxa to either a planktotrophic or a lecithotrophic larval mode. A more reliable quantitative character is evaluated in juvenile samples of two genera from the Ordovician Mifflin Formation, the presumed planktotrophic orthid Doleroides pervetus and the presumed lecithotrophic atrypid Protozyga nicolleti, using the coefficient of variation in measurements of protegular size. The Doleroides sample has a significantly higher coefficient of variation than does the Protozyga sample. The results are in accord with observations on living marine invertebrates which indicate that at settlement (when mineralisation of the protegular mantle in brachiopods occurs), planktotrophic larval size is more variable than larval size in lecithotrophs because of fluctuations in food availability and duration of the planktonic phase. This result confirms that protegular size variance can be used to make inferences about larval trophic mode.

This paper describes a terebratelloid articulate brachiopod, Gyrothyris williamsi sp. nov., based on 95 specimens from seamounts on the Lord Howe Rise, Coral Sea, SW Pacific Ocean. The new species is attributed to Gyrothyris on the basis of (a) morphological and growth trajectory similarities; (b) phylogenetic analyses of an alignment of DNA sequence (∼2900-sites) obtained from nuclear-encoded small- and large-subunit ribosomal RNA genes (SSU and LSU); and (c) the presence of a distinctive, two-part deletion in the LSU gene. It is distinguished morphologically from Gyrothyris mawsoni and its subspecies by both internal and external morphology and by its isolated geographical distribution, which extends the patchy, known range of this genus to an area some 2000 km north of its previous northern limit around New Zealand. Phylogenetic analyses of the rDNAs and of mitochondrial cox1 gene sequences (663 sites) confirm previous indications that the New Zealand endemic terebratelloid genera form a clade (Neothyris (Calloria, Gyrothyris, Terebratella), but the position of Terebratella with respect to Calloria and Gyrothyris remains weakly established. These sequences disagree inexplicably about the closeness of the relationship between Neothyris parva and N. lenticularis. Analyses of the first sequences from Calloria variegata, a species restricted to the Hauraki Gulf, New Zealand, are consistent with the possibility that it originated locally, and recently, from C. inconspicua. Magellania venosa from S. America/Falklands joins with Antarctic Magellania fragilis and M. joubini to form an rDNA clade that excludes Terebratalia as the putative sister-group of the New Zealand terebratelloid clade. The cox1 (but not the rDNA) sequences of the New Zealand clade pass a test for clock-like rates of evolution, and maximum likelihood pairwise distances suggest that if genetic isolation between the ancestor of Antarctic Magellania and the last common ancestor of the New Zealand terebratelloid clade was initiated by separation of the Antarctic and New Zealand plates ∼90 Mya, isolation from M. venosa was initiated earlier, perhaps ∼145 Mya. However, in the simple phylogenetic reconstruction presented here from cox1 sequences, S. American and Antarctic Magellania spp. do not yield a well-supported clade, perhaps because of differences in base composition.