A Neolithic stone circle at Waun Mawn, in the Mynydd Preseli, west Wales, has been proposed as the original location of some dolerite megaliths at Stonehenge, including one known as Stone 62. To investigate this hypothesis, in-situ analyses, using a portable XRF, have been obtained for four extant non-spotted doleritic monoliths at Waun Mawn, along with two weathered doleritic fragments from a stonehole (number 91). The data obtained have been compared to data from spotted and non-spotted dolerite outcrops across the Mynydd Preseli, an area known to be the source of some Stonehenge doleritic bluestones, as well as data from in-situ analysis of Stone 62 (non-spotted dolerite) and ex-situ analysis of a core taken from Stone 62 in the late 1980′s. Recently, Stone 62 has been identified as coming from Garn Ddu Fach, an outcrop some 6.79 km to the east-southeast of Waun Mawn. None of the four dolerite monoliths at Waun Mawn have compositions which match Stonehenge Stone 62, and neither do the weathered fragments from stonehole 91. Rather the data show that the Waun Mawn monoliths, and most probably the weathered stonehole fragments, can be sourced to Cerrig Lladron, 2.37 km southwest of Waun Mawn, suggesting that a very local stone source was used in construction of the Waun Mawn stone circle. It is noted that there is evidence that at least eight stones had been erected and subsequently removed from the Waun Mawn circle but probability analysis suggests strongly that the missing stones were also derived, at least largely, from Cerrig Lladron.
Middle to Late Ordovician (late Darriwilian –early Katian) trilobites from trinucleid, raphiophorid and cyclopygid biofacies are for the first time documented from the West Balkhash and North Betpak‐Dala regions in south‐central Kazakhstan. Twenty taxa are described; six of them, that is, Miaopopsis sokyrensis sp. nov., Pricyclopyge keralensis sp. nov., Kongqiaoheia sarytumensis sp. nov., Shumardia karasaiensis sp. nov., Triarthrus akkermensis sp. nov. and Mynaralaspis perforata gen. et sp. nov. are new to science. Six taxa are reported in open nomenclature and two are not identified on a genus level. A new trinucleid Subfamily Nanshanaspinae is designated. Newly discovered faunas show strong links to contemporaneous faunas of Tarim and, to a lesser degree, to those of South China and the Australian sector of Gondwana. Based on results of cluster analysis (Dice similarity), trilobite associations of raphiophorid biofacies from the Chu‐Ili and Stepnyak terranes form a single first‐order cluster with little similarity to the trilobite associations characteristic of the nielid and illenid–cheirurid biofacies from the Stepnyak and Tarbagatai regions. Also, there is no distinct interaction between the pelagic Degamella–Pricyclopyge association and other trilobite faunas. The geographical distribution of raphiophorid and trinucleid trilobites described in this paper, including species of genera Malongullia?, Miaopopsis, Nanshanaspis and Kongqiaoheia, supports Ordovician palaeogeographical reconstructions showing the Kazakh terranes as part of a huge archipelago situated on both sides of the equator in proximity to the west coast of Gondwana and positioned a considerable distance from the Siberia continent and its associated terranes.
Animal biodiversity is greatly underestimated in nontemperate marine regions, especially for intertidal benthic organisms such as oysters. Recent surveys in the northern Arabian Gulf suggest the presence of numerous unidentified species, some of which form shallow reef ecosystems while others are cryptic and found under rocks. In this study, we focused on small oysters from Kuwait, which show typical characteristics in common with the genus Ostrea except for the presence of lophine chomata that would link it to the genera Lopha, Dendostrea, and Alectryonella. Phylogenetic analyses based on mitochondrial and nuclear markers unambiguously placed the Kuwait oyster within the Ostreinae as a sister to the Japanese species Ostrea futamiensis. The hypothesis that the Kuwait oyster represents a new species was assessed with phylogenetic and species delimitation methods combined with a morphological assessment. Results corroborated the Kuwait oysters as a new species herein described as Ostrea oleomargarita Oliver, Salvi, and Al-Kandari, sp. nov. The phylogeny of the Ostreinae shows extensive disagreement between morphology-based genera and phylogenetic clades. The genus Ostrea is polyphyletic, and the form and distribution of taxonomic characters such as chomata are not as definitive as suggested in previous studies. This study, along with other recent investigations, confirmed the Arabian Gulf as a key region for discovering marine animal diversity and suggested a possible biogeographic divide between the Eastern and Western Indo-Pacific. A pattern that has been documented in a growing number of taxa and that warrants further research attention.
This is the fourth Active Healthy Kids (AHK) Wales Report Card. The 2021 card produced grades on children and young people’s physical activity (PA) using pre-COVID-19 data that were not used in previous versions. Eleven quality indicators of PA were graded through expert consensus and synthesis of the best available evidence. Grades were assigned as follows: Overall PA—F; Organised Sport and PA—C; Active Play—C+; Active Transportation—C−; Sedentary Behaviours—F; Physical Fitness—C−; Family and Peer Influences—D+; School—B−; Community and the Built Environment—C; National Government and Policy—C; and Physical Literacy—C−. All but three grades remained the same or decreased from the 2018 AHK-Wales Report Card (Active Play increased from C− to C+; Active Transportation, D+ to C−; Family and Peers, D to D+). This is concerning for children’s health and well-being in Wales, particularly given recent evidence that PA has further decreased during the COVID-19 pandemic. The results from the Report Card should be used to inform the decision making of policy makers, practitioners and educators to improve children and young people’s PA levels and opportunities and decrease PA inequalities.
We introduce a new method of estimating accepted species diversity by adapting mark-recapture methods to comparisons of taxonomic databases. A taxonomic database should become more complete over time, so the error bar on an estimate of its completeness and the known diversity of the taxon it treats will decrease. Independent databases can be correlated, so we use the time course of estimates comparing them to understand the effect of correlation. If a later estimate is significantly larger than an earlier one, the databases are positively correlated, if it is significantly smaller, they are negatively correlated, and if the estimate remains roughly constant, then the correlations have averaged out. We tested this method by estimating how complete MolluscaBase is for accepted names of terrestrial gastropods. Using random samples of names from an independent database, we determined whether each name led to a name accepted in MolluscaBase. A sample tested in August 2020 found that 16.7% of tested names were missing; one in July 2021 found 5.3% missing. MolluscaBase grew by almost 3,000 accepted species during this period, reaching 27,050 species. The estimates ranged from 28,409 ± 365 in 2021 to 29,063 ± 771 in 2020. All estimates had overlapping 95% confidence intervals, indicating that correlations between the databases did not cause significant problems. Uncertainty beyond sampling error added 475 ± 430 species, so our estimate for accepted terrestrial gastropods species at the end of 2021 is 28,895 ± 630 species. This estimate is more than 4,000 species higher than previous ones. The estimate does not account for ongoing flux of species into and out of synonymy, new discoveries, or changing taxonomic methods and concepts. The species naming curve for terrestrial gastropods is still far from reaching an asymptote, and combined with the additional uncertainties, this means that predicting how many more species might ultimately be recognized is presently not feasible. Our methods can be applied to estimate the total number of names of Recent mollusks (as opposed to names currently accepted), the known diversity of fossil mollusks, and known diversity in other phyla.
The doleritic bluestone monoliths at Stonehenge have long been known to have been sourced from the Mynydd Preseli area in west Wales, some 225 km away. On geochemical grounds, based on a range of major and trace elements determined by laboratory-based X-ray fluorescence spectrometry, they have been divided into three groups (Groups 1–3). Subsequently, rare earth element data obtained by solution nebulization ICP-MS showed Group 2 Stone 45 to have been sourced from Cerrigmarchogion with Group 2 Stone 62 showing similarities to the outcrops of Carn Ddafad-las and Garn Ddu Fach. In order to test this possible link, portable XRF (pXRF) analyses were obtained in situ from Stone 62 at Stonehenge and from key outcrops in the Preseli. To obtain reliable results from heterogeneous coarse-grained igneous rocks using pXRF to enable comparisons between orthostats and possible sources, a robust analytical strategy was developed. For outcrops this involved a series of horizontal traverses through individual outcrops comprising 10 to 15 analyses a few centimetres apart, giving 60–175 independent analyses per outcrop. This gives a large, analysed surface area per outcrop providing representative data and also has potential to show any vertical variation in source outcrops. Analyses were taken from similarly weathered surfaces on orthostats and outcrops to minimise compositional changes from surface weathering, and a minimum of 20 analyses were taken from orthostats at various locations across the stone. This approach provides a set of well-determined elements which can reliably be used for provenance studies (including K, Fe, Mn, Zn, Rb, Sr, Zr, Nb, Ba). Nickel, Ti and V are affected slightly more by weathering but still prove useful in comparisons, but Cr, used in earlier studies as a compositional discriminant with Ni, here has poor accuracy by pXRF and is not used. The pXRF analyses show that Stone 62 sits within the same ‘compositional space’ (for multiple element concentrations and ranges) as analyses from Carn Ddafad-las and Garn Ddu Fach, two outcrops of the same intrusion which differ from all other analysed outcrops. More specifically, analyses from Stone 62 and Garn Ddu Fach overlap, these forming a subset of the analyses from Carn Ddafad-las, which shows a more extensive range of those same elements. On this basis it is suggested that Garn Ddu Fach is the likely source of Stonehenge non-spotted dolerite Stone 62, a suggestion supported by indistinguishable petrographic characteristics between Stone 62 and dolerite sample PGDF24 from Garn Ddu Fach.
Chu‐Ili, now in Kazakhstan, was a substantial independent equatorial microcontinental terrane in Ordovician times, with a small Precambrian core fringed by several island arcs. Its mid‐Ordovician (late Darriwilian to early Katian) faunas were a major evolutionary hotspot within an equatorial archipelago at a period when Palaeozoic sea levels and temperatures were at their highest. As well as reviewing the previously described brachiopods from elsewhere in Chu‐Ili, the mid‐Ordovician brachiopods of the West Balkhash Region, which outcrops west of Lake Balkhash within Chu‐Ili, are newly described here, mainly from the Berkutsyur and Baigara formations. Many represent the earliest occurrence of their lineages, notably the oldest member of the Order Atrypida. More than twelve brachiopod associations are defined, many for the first time and together hosting 73 genera and over 91 species. The new family Kellerellidae is erected within the superfamily Lissatrypoidea. New genera are Aploobolus (Obolidae), Doughlatomena (Rafinesquinidae), and Altynorthis , Lictorthis , and Baitalorthis (all Plectorthidae), Baitalorhynchus (Sphenotretidae), Lydirhyncha (Ancistrorhynchidae) and Costistriispira (Kellerellidae). Eleven new species, including Aploobolus ? tenuis , Doughlatomena splendens , Bimuria karatalensis , Apatomorpha akbakaiensis , Lepidomena betpakdalenis , Sonculina baigarensis , Altynorthis betpakdalensis , Altynorthis vinogradovae , Phaceloorthis ? corrugata , Batailorhyncha rectimarginata and Costistriispira proavia , and one new subspecies Sowerbyella ( Sowerbyella ) verecunda baigarensis are also erected. The global palaeogeographical affinities of all the Chu‐Ili brachiopod faunas are discussed, as well as Chu‐Ili's place within the peri‐Gondwanan archipelago. Newly named stratigraphical units are the Berkutsyur (Darriwilian to early Sandbian) and overlying Kopkurgan (Sandbian to Katian) formations within West Balkhash, and the Tastau (Darriwilian) and Takyrsu (Darriwilian to early Sandbian) formations within the northern Betpak‐Dala desert.
Two massive precipitation events of polymetallic ore deposits, encrusted by a mixture of authigenic carbonates, are documented from the Cambrian of the semi-enclosed Baltoscandian Basin. δ34S (‒9.33 to ‒2.08‰) and δ33S (‒4.75 to ‒1.06‰) values from the basal sulphide breccias, sourced from contemporaneous Pb–Zn–Fe-bearing vein stockworks, refect sulphide derived from both microbial and abiotic sulphate reduction. Submarine metalliferous deposits were triggered by non-buoyant hydrothermal plumes: plumes of buoyant fuid were trapped by water column stratifcation because their buoyancy with respect to the environment reversed, fuids became heavier than their surroundings and gravitational forces brought them to a halt, spreading out laterally from originating vents and resulting in the lateral dispersion of efuents and sulphide particle settling. Subsequently, polymetallic exhalites were sealed by carbonate crusts displaying three generations of ikaite-to-aragonite palisade crystals, now recrystallized to calcite and subsidiary vaterite. Th of fuid inclusions in early calcite crystals, ranging from 65 to 78 ºC, provide minimum entrapment temperatures for carbonate precipitation and early recrystallization. δ13Ccarb (‒1.1 to+ 1.6‰) and δ18Ocarb (‒7.6 to ‒6.5‰) values are higher than those preserved in contemporaneous glendonite concretions (‒8.5 to ‒4.7‰ and ‒12.4 to ‒9.1‰, respectively) embedded in kerogenous shales, the latter related to thermal degradation of organic matter. Hydrothermal discharges graded from highly reduced, acidic, metalliferous, and hot (~ 150 ºC) to slightly alkaline, calcium-rich and warm (< 100 ºC), controlling the precipitation of authigenic carbonates.
Semantic segmentation has been proposed as a tool to accelerate the processing of natural history collection images. However, developing a flexible and resilient segmentation network requires an approach for adaptation which allows processing different datasets with minimal training and validation. This paper presents a cross-validation approach designed to determine whether a semantic segmentation network possesses the flexibility required for application across different collections and institutions. Consequently, the specific objectives of cross-validating the semantic segmentation network are to (a) evaluate the effectiveness of the network for segmenting image sets derived from collections different from the one in which the network was initially trained on; and (b) test the adaptability of the segmentation network for use in other types of collections. The resilience to data variations from different institutions and the portability of the network across different types of collections are required to confirm its general applicability. The proposed validation method is tested on the Natural History Museum semantic segmentation network, designed to process entomological microscope slides. The proposed semantic segmentation network is evaluated through a series of cross-validation experiments designed to test using data from two types of collections: microscope slides (from three institutions) and herbarium sheets (from seven institutions). The main contribution of this work is the method, software and ground truth sets created for this cross-validation as they can be reused in testing similar segmentation proposals in the context of digitization of natural history collections. The cross-validation of segmentation methods should be a required step in the integration of such methods into image processing workflows for natural history collections.
The faunistic knowledge of the Diptera of Morocco recorded from 1787 to 2021 is summarized and updated in this first catalogue of Moroccan Diptera species. A total of 3057 species, classified into 948 genera and 93 families (21 Nematocera and 72 Brachycera), are listed. Taxa (superfamily, family, genus and species) have been updated according to current interpretations, based on reviews in the literature, the expertise of authors and contributors, and recently conducted fieldwork. Data to compile this catalogue were primarily gathered from the literature. In total, 1225 references were consulted and some information was also obtained from online databases. Each family was reviewed and the checklist updated by the respective taxon expert(s), including the number of species that can be expected for that family in Morocco. For each valid species, synonyms known to have been used for published records from Morocco are listed under the currently accepted name. Where available, distribution within Morocco is also included. One new combination is proposed: Assuania melanoleuca (Séguy, 1941), comb. nov. (Chloropidae)
Although phosphatized bryozoans have been described recently from the early Cambrian, the first unequivocal bryozoan fossils with hard skeletons are known from the Ordovician. Recent discoveries of bryozoans in the early Ordovician (Tremadocian) of South China have greatly expanded our understanding of the diversification of these colonial lophophorates. In particular, the Fenhsiang Formation of Late Tremadocian age (Migneintian) in Hubei Province is proving to be particularly rich in bryozoans. Here we record 24 species, including several yet to be formally described, belonging to 18 genera and four palaeostomate suborders (Esthonioporata, Cystoporata, Trepostomata, and Cryptostomata). Bryozoan diversity in the Fenhsiang Formation matches levels more typical of younger faunas of Middle Ordovician age. The presence of diverse and morphologically disparate taxa close to the base of the Ordovician suggests rapid diversification following the first appearance of bryozoans with calcified skeletons, and/or the existence of as yet unknown biomineralized bryozoans in the Cambrian.
The Endeavour voyage was the first expedition from Britain to carry professional botanists and natural history artists. The main purpose was to observe the Transit of Venus. However, Cook had secret orders to explore and map the elusive southern continent. Joseph Banks, a wealthy naturalist, was consumed with excitement at the prospect of undiscovered plants, animals, and people. He arranged for a party of naturalists to accompany the expedition. Banks employed three artists: Parkinson; Buchan; and Spöring. Daniel Solander was his scientific partner. Contemporary diaries reveal how these men, from different backgrounds and nations, worked together to compile 3,000 specimens and over 1,300 drawings. Banks intended to publish the drawings quickly, but publication was delayed. When Solander died, the publication lost momentum and Banks was preoccupied by other projects. Banks’ Florilegium was finally published over 200 years later, a lasting legacy of the talents, dedication, and bravery of the artists.
South African marine biogeography has received a great deal of attention in the past. However, offshore biogeographic breaks are based on abiotic variables or specialist opinion rather than data-driven approaches. Here we investigate biogeographic breaks based on the distribution of offshore marine benthic molluscs collected during the Natal Museum Dredging Programme (1981-1997) along the coast of South Africa, between the South East Atlantic and South West Indian Oceans. Spatially constrained (depth, latitude and longitude) cluster analyses were applied to three resemblance measures; the Jaccard coefficient, popular in biogeographic studies, and gamma+ and theta+ coefficients, where taxonomic relatedness is considered when comparing sites. We used these resemblances to 1) find the most consistent biogeographic breaks and 2) compare the results of different measures to better understand the patterns and possible processes involved in the biogeographical classification. Although there were differences in the overall results from the three resemblance measures, they produced comparable findings at depths shallower than 80 m. Our data-driven approach indicated that the biogeographic break between the warm-temperate and subtropical assemblages was 140 km north of the current inshore break. The break between the subtropical and tropical mollusc assemblages agreed more closely with the present Ecoregions. Compared to Jaccard, where relatedness is not considered, gamma+ and theta+ results revealed a more substantial depth effect, and samples collected in the same depth range were more closely related and shared a more recent evolutionary history than geographically close samples from different depths. Furthermore, assemblages found deeper than ~80 m demonstrated greater diversity in terms of relatedness, an essential consideration in marine spatial planning and consequent designation of protected areas. This study reveals the importance of including in-situ data and interspecies relationships in future marine spatial planning to ensure that these evolutionary rich animal communities are considered adequately.
Indirect development with an intermediate larva exists in all major animal lineages, and thus larvae are central to most scenarios for animal evolution. Yet how larvae evolved remains disputed. Here we show that changes in the timing of trunk formation underpin the diversification of larvae and bilaterian life cycles. Combining chromosome-scale genome sequencing with transcriptomic and epigenomic profiling in the slow-evolving oweniid Owenia fusiformis, we found that different genes and genomic regulatory elements control the development of its feeding larva and adult stage. First, O. fusiformis embryos develop into an enlarged anterior domain that forms larval tissues and the adult head, as posterior growth and trunk patterning is deferred to pre-metamorphic stages. These traits also occur in the so-called "head larvae" of other bilaterians, with whom O. fusiformis larva shows extensive transcriptomic similarities. Conversely, animals with non-feeding larvae and gradual metamorphoses, such as the annelid Capitella teleta, start trunk differentiation during embryogenesis, like direct developers. Together, our findings suggest that the ancestral temporal decoupling of head and trunk formation, as retained in extant "head larvae", allowed larval evolution in Bilateria, questioning prevailing scenarios that propose either co-option or innovation of gene regulatory programmes to explain larva and adult origins.
Present-day people from England and Wales harbour more ancestry derived from Early European Farmers (EEF) than people of the Early Bronze Age¹. To understand this, we generated genome-wide data from 793 individuals, increasing data from the Middle to Late Bronze and Iron Age in Britain by 12-fold, and Western and Central Europe by 3.5-fold. Between 1000 and 875 bc, EEF ancestry increased in southern Britain (England and Wales) but not northern Britain (Scotland) due to incorporation of migrants who arrived at this time and over previous centuries, and who were genetically most similar to ancient individuals from France. These migrants contributed about half the ancestry of Iron Age people of England and Wales, thereby creating a plausible vector for the spread of early Celtic languages into Britain. These patterns are part of a broader trend of EEF ancestry becoming more similar across central and western Europe in the Middle to Late Bronze Age, coincident with archaeological evidence of intensified cultural exchange2–6. There was comparatively less gene flow from continental Europe during the Iron Age, and Britain’s independent genetic trajectory is also reflected in the rise of the allele conferring lactase persistence to ~50% by this time compared to ~7% in central Europe where it rose rapidly in frequency only a millennium later. This suggests that dairy products were used in qualitatively different ways in Britain and in central Europe over this period.
The rhipidoglossan radula, consisting of numerous teeth in each transverse row, is characteristic of phylogenetically distant groups of gastropods, including Vetigastropoda, Neritimorpha, and "lower" Heterobranchia. Previous studies have revealed the main patterns in the formation of the rhipidoglossan radula of vetigastropods, the main feature of which is the division of the formation zone into two horns, where marginal teeth are formed by a multilayered epithelium (odontoblasts). This work is devoted to the study of the formation of the rhipidoglossan radula of Nerita litterata using light and electron microscopy. The data obtained show that, despite the different external morphology of the radular sac of neritids and vetigastropods, the radular sac of N. litterata, like that of vetigastropods, is divided into two parts, in which the marginal teeth are similarly formed by odontoblasts located in more than one layer. It seems probable that this complex, three-dimensional structure of the formation zone is associated with a broad radula with numerous elongate marginal teeth and could be characteristic for other gastropods with this type of radula. Additional supporting rods located along the odontoblasts and consisting of vacuolated cells were first discovered in Nerita.
You can download this paper for free until September 29, 2021, at: https://authors.elsevier.com/c/1dYtGACeY7xhD
Evidence of Late Triassic large tetrapods from the UK is rare. Here, we describe a track-bearing surface located on the shoreline near Penarth, south Wales, United Kingdom. The total exposed surface is c. 50 m long and c. 2 m wide, and is split into northern and southern sections by a small fault. We interpret these impressions as tracks, rather than abiogenic sedimentary structures, because of the possession of marked displacement rims and their relationship to each other with regularly spaced impressions forming putative trackways. The impressions are large (up to c. 50 cm in length), but poorly preserved, and retain little information about track-maker anatomy. We discuss alternative, plausible, abiotic mechanisms that might have been responsible for the formation of these features, but reject them in favour of these impressions being tetrapod tracks. We propose that the site is an additional occurrence of the ichnotaxon Eosauropus , representing a sauropodomorph trackmaker, thereby adding a useful new datum to their sparse Late Triassic record in the UK. We also used historical photogrammetry to digitally map the extent of site erosion during 2009–2020. More than 1 m of the surface exposure has been lost over this 11-year period, and the few tracks present in both models show significant smoothing, breakage and loss of detail. These tracks are an important datapoint for Late Triassic palaeontology in the UK, even if they cannot be confidently assigned to a specific trackmaker. The documented loss of the bedding surface highlights the transient and vulnerable nature of our fossil resources, particularly in coastal settings, and the need to gather data as quickly and effectively as possible.
Biological invasions are common among freshwater molluscs, with the North American planorbid gastropod Gyraulus parvus being reported from Europe (Germany) by the 1970s. It has since spread across Central and Western Europe, mostly living in artificial and highly modified habitats. However, considerable conchological and anatomical similarity exists between it and the native European G. laevis. Using four other European and one North American Gyraulus species as outgroups, separate phylogenetic analyses using mitochondrial and nuclear DNA sequences show that G. parvus and G. laevis are in fact part of the same species-level clade, with the former having nomenclatural priority. However, the structure within the mitochondrial tree suggests a North American origin of the invasive populations. It also makes it possible to track down the distribution of both races. Although native and non-native races in Europe tend to possess some differences in conchology and ecology, the degree of overlap makes it impossible to accurately distinguish between them without the DNA barcode data. Our results change the outlook on the conservation of the rare native race. While interspecific competition among snail species is rare, invasion on an intraspecific level may represent a serious threat for native populations.
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.
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