The Early Palaeozoic was a critical interval in the evolution of marine life on our planet. Through a window of some 120 million years, the Cambrian Explosion, Great Ordovician Biodiversification Event, End Ordovician Extinction and the subsequent Silurian Recovery established a steep trajectory of increasing marine biodiversity that started in the Late Proterozoic and continued into the Devonian. Biogeography is a key property of virtually all organisms; their distributional ranges, mapped out on a mosaic of changing palaeogeography, have played important roles in modulating the diversity and evolution of marine life. This Memoir first introduces the content, some of the concepts involved in describing and interpreting palaeobiogeography, and the changing Early Palaeozoic geography is illustrated through a series of time slices. The subsequent 26 chapters, compiled by some 130 authors from over 20 countries, describe and analyse distributional and in many cases diversity data for all the major biotic groups plotted on current palaeogeographic maps. Nearly a quarter of a century after the publication of the ‘Green Book’ (Geological Society, London, Memoir 12, edited by McKerrow and Scotese), improved stratigraphic and taxonomic data together with more accurate, digitized palaeogeographic maps, have confirmed the central role of palaeobiogeography in understanding the evolution of Early Palaeozoic ecosystems and their biotas.

Abstract New palaeogeographical reconstructions are presented at 10 myr intervals from the Lower Cambrian at 540 Ma to the Lower Devonian at 400 Ma, showing continental crustal fragments and oceans (not lands and seas), with appropriate kinematic continuity between successive maps. The maps were chiefly generated by revised and selected palaeomagnetic data and revised Apparent Polar Wandering paths linked to present-day polygons from the main continents. These have been reinforced by analysis of the distributions of some fossils and sediments. Gondwana was the dominating supercontinent from its final assembly in the Latest Neoproterozoic at about 550 Ma until the Carboniferous, and covered much of the Southern Hemisphere. The Northern Hemisphere was largely occupied by the vast Panthalassic Ocean. The relative positions of the major continents and the latitudes and rotation histories of Gondwana, Baltica, Siberia and Laurentia (Laurussia from the mid-Silurian) are now well known. Although Laurentia was oriented in a similar direction to the present, Siberia was inverted throughout the Lower Palaeozoic, and Baltica too was initially inverted, but rotated through 120° between the Late Cambrian and Late Ordovician before collision with Laurentia in the mid-Silurian Caledonide Orogeny. Through reconstructions of the Caledonide and some other orogenies, the progressive history of the Iapetus Ocean between Laurentia and Baltica/Gondwana is well constrained. Less major continents whose positions are also well known include Avalonia (initially peri-Gondwanan but migrating in the Early Ordovician to join Baltica by the end of the Ordovician), Sibumasu (now considered an integral part of Gondwana) and Mongolia (adjacent to Siberia). A large number of other terranes are reviewed and plotted on the reconstructions with varying degrees of certainty. However, significant continents with less well constrained or controversial positions are South China, North China (Sinokorea), Annamia (Indochina) and Arctic Alaska–Chukotka. The European areas of France, Iberia and southern Italy, previously considered by some as a separate Armorican Terrane Assemblage, remained parts of core Gondwana until the opening of the Palaeotethys Ocean near the end of the Silurian, but it is uncertain whether Perunica (Bohemia) was one of that group or whether it left Gondwana during the Middle Ordovician.

Abstract Palaeogeographers, geographers and structural geologists use different well-defined terms to designate continental and tectonic units, whereas biogeographers, palaeobiogeographers and palaeontologists use a wide range of subjective terminologies to describe biogeographical and palaeobiogeographical units. The absence of clear definitions and of rules or guidelines for palaeobiogeographical nomenclature has resulted in frequent misunderstandings and general confusion, in particular when applied to ancient time periods, such as the Palaeozoic. Palaeogeographical and palaeobiogeographical terminology used in Palaeozoic geology and palaeontology is reviewed, and recent attempts to standardize palaeobiogeographical nomenclature summarized. We make a number of proposals for future use of terms to avoid confusion and misunderstandings.

Abstract The Cambrian geographical and temporal distributions of many clades remain poorly understood, despite their importance for elucidating the palaeobiogeographical context of the Cambrian radiation. New species and genus level occurrence databases were developed to analyse temporal and geographical distributional patterns in taxa belonging to 14 clades from over 60 globally distributed early and Middle Cambrian Burgess Shale-type lagerstätten. Analyses demonstrate that clades with confirmed Precambrian origins were, on average, more widespread and temporally persistent than clades with first fossil occurrences in the Cambrian. Despite their dominance in diversity, arthropods were less widely distributed and temporally persistent than many other groups. Finally, a significant correlation between geographical range and temporal persistence is demonstrated, supporting the hypothesis that Cambrian taxa with wider geographical ranges were less likely to go extinct than those with narrower ranges. Supplementary material: Species and genus data tables are available at: http://www.geolsoc.org.uk/SUP18665

Abstract We examined the palaeogeographical distribution of selected Cambrian trace fossils. Astropolichnus hispanicus , Climactichnites , Syringomorpha nilssoni and early examples of Paleodictyon all have a restricted palaeogeographical distribution, probably representing that of their producers. A cosmopolitan distribution is seen in Didymaulichnus miettensis and in early examples of Rusophycus and Dactyloidites . Oldhamia shows a wide distribution in Lower Cambrian deep-sea sediments although that of Oldhamia geniculata is restricted.

Abstract The Archaeocyatha is a group of Cambrian fossils successively considered as cnidarians or sponges or as an independent phylum convergent with many groups lacking clear affinities. Comparisons with Recent calcified sponges discovered in submarine caves have demonstrated that the sponge model is consistent with their structural organization. Thus their systematic position is now agreed as a class within the phylum Porifera, permitting realistic investigations of their comparative physiology and life strategies. Archaeocyatha is an important part of research programmes on the Cambrian System, initiated by different commissions of the IUGS since 1970. Archaeocyathan biozones are available in some key regions. Faunal and palaeocommunities distribution, especially of the reefs they helped build in epeiric seas, and migration pathways constrain Cambrian palaeogeographical reconstructions. A database, using recent compilations of the group, is now online. This free access data source offers specialists a tool, easy to use not only as an identification key but also to establish faunal, geographical and stratigraphical distributions of archaeocyathan genera, and a rapid first step towards Cambrian palaeogeographical reconstructions.

Abstract Stromatoporoid sponges first appeared during the late Mid-Ordovician (mid–late Darriwilian) accompanying an important ‘Chazy’ reef-building episode. Representatives of the order Labechiida appeared first, initially splitting into two sister groups: those from North China (nine genera) and those from Laurentia (four genera). Two genera were common to the two regions, but others in North China and Siberia were endemic. This initial provincialism was not maintained into the Late Ordovician (Sandbian) as labechiids attained a wider dispersal, covering Laurentia with peripheral terranes in Northwest Scotland and Chukchi Peninsula, cratonic Siberia, the Urals (eastern margins of Baltica), and East Gondwanan blocks of Tarim North China, marginal Tasmania and the peri-Gondwanan New South Wales island-arc terrane. Only a few endemics remained present in the Sandbian – three in Laurentia, and one in Tarim. In the Katian, maximum diversification of labechiids occurred (19 genera, including a genus possibly transitional to first actinostromatids). Also four genera of the Clathrodictyida first appeared, and overall distribution continued to increase. The Hirnantian marked a dramatic decline in the global distribution (only Anticosti Island, Manitoba, Norway and Estonia) and marked diversity loss of labechiids. Altogether c. 70% of stromatoporoid species disappeared in response to end-Ordovician global cooling events. Llandovery stromatoporoids were widespread in Laurentia, Baltica and Siberia, with clathrodictyids as dominant and labechiids accessory (the latter group more common in Siberia and China). Gradual diversification and expansion of stromatoporoids followed in the late Llandovery, with appearances of the orders Actinostromatida and Stromatoporida. The stromatoporoids became most widespread and most diversified during the Wenlock, with clathrodictyids maintaining their leading position, and the appearances of the earliest Stromatoporellida and Syringostromatida. Rapid spread of new phylogenetic stocks indicates that widespread pandemism prevailed among Wenlock stromatoporoids. The Ludlow was characterized by final closure of the Iapetus Ocean, accompanying uplift of palaeocontinental regions, and declining prominence of stromatoporoid-bearing shallow carbonate shelves. However, clathrodictyids and actinostromatids remained dominant, whereas the incoming of Amphiporida in several regions (Somerset Island, Baltic area, West Ukraine, West and East Urals, Novaya Zemlya, Tien Shan, Japan and New South Wales) and restricted distribution of some other taxa suggest a certain provincialism developed. With regional regression and stratigraphic hiatuses, the stromatoporoids (mostly hangovers from the Ludlow) became less common in the Pridoli, and apparently even entirely absent from Gondwana and Siberia.

Abstract The Cambrian, Ordovician and Silurian distributions of non-stromatoporoid sponges are reviewed. The earliest Cambrian faunas contain mostly hexactinellids, with protomonaxonids dominating middle Cambrian assemblages. There are no obvious palaeobiogeographical patterns, with many genera being found widely. Vauxiids, lithistids and heteractinids are apparently confined to low latitudes, but this may be due to a poor fossil record. Most known Ordovician faunas are from low latitudes, although some high-latitude faunas are known, which contain reticulosan hexactinellids and protomonaxonids. There is some division of faunas within Laurentia, into eastern and western provinces, with the western assemblage extending across low northern latitudes during the Late Ordovician. During the Silurian Period, sponge diversity was very low during the Llandovery Epoch, probably partly owing to lack of habitat for taxa restricted to carbonate facies, and also because of sampling bias. There was a dramatic increase in diversity through the Silurian Period, mostly owing to an apparent diversification in the demosponges; however, there are many ghost lineages, indicating that their fossil record is poorly known. Non-lithistid sponges are very poorly known, with few recorded outside Euramerica. The currently available data for Early Palaeozoic sponges are too incomplete to allow any reliable palaeobiogeographical inferences. Supplementary material: the compilation of Silurian sponge occurrences is available at: http://www.geolsoc.org.uk/SUP18666 .

Abstract During the Late Ordovician, Cincinnatian, the epicontinental seas and continental margin of Laurentia provided habitats that were suitable for corals. Biogeographical differentiation occurred within this equatorially placed continent, when corals were introduced to areas that had fundamentally different environments. There were four biogeographical divisions, characterized by distinctive faunas that included some endemic taxa: the Red River–Stony Mountain Province, Richmond Province, Edgewood Province and the less well understood, informal ‘Continental Margin’ Area. In each division, the potential for diversification and the capacity for diversity were determined by factors such as the duration and size of the division, the amount of immigration, the extent of evolution and biogeographical differentiation, faunal responses to changes in sea-level and climate, and the complexity of the ecological structure. The development of multiple biogeographical divisions, each contributing to overall diversity, enhanced the ‘Great Ordovician Biodiversification Event’. During the latest Ordovician mass extinction, there was a reduction of diversity and loss of biogeographical divisions within Laurentia. The divisions were terminated when their characteristic taxa disappeared, in response to major environmental changes associated with glaciation in Gondwana and subsequent global warming.

Abstract The biogeographical patterns shown by Ordovician linguliform and craniiform brachiopods are greatly influenced by their dominance in low-diversity associations in marginal environments. This is particularly evident in the Early Ordovician, when linguliform-dominated dysaerobic assemblages are widely distributed along the deep shelves of Gondwana, the Kazakhstanian terranes and in Baltica. By the Darriwilian, micromorphic linguliforms are characteristic components of the pantropical climatic-controlled faunas of Laurentia, Cuyania and Kazakhstanian terranes, which – in spite of separation by extensive oceans – retain a distinct similarity. Analysis of craniiform biogeographical distribution is impeded significantly by the poor state of craniide taxonomy and lack of reliable data from most regions. However, in general their biogeographical dispersion is similar to other groups of the Palaeozoic Evolutionary Fauna. Unlike the linguliforms, which are important members of the Cambrian Evolutionary Fauna, there is no convincing Cambrian craniiform record; they may have evolved and dispersed from Gondwana and associated microcontinents and island arcs. The earliest well-established record is from the late Tremadocian of temperate to high-latitude peri-Gondwana. During most of the Ordovician, they have a peri-Iapetus distribution. They are very rare or absent in tropical Gondwana, South China and Kazakhstanian terranes and are not yet documented from Siberia. The trimerellides probably evolved in tropical peri-Gondwanan island arc settings. Their dispersion and major features of biogeography mirror those of atrypides. Gold Open Access: This article is published under the terms of the http://creativecommons.org/licenses/by/3.0/ .

Abstract The phylogeographical evolution and the consequent changing distribution and diversity of rhynchonelliform brachiopods through the Ordovician are linked to the dynamic palaeogeography of the period. The Early Ordovician (Tremadocian and Floian) is characterized by globally low-diversity faunas with local biodiversity epicentres, notably on the South China Palaeoplate; low-latitude porambonitoid-dominated faunas with early plectambonitoid and clitambonitoid representatives, as well as high-latitude assemblages mostly dominated by orthoids, can be recognized, but many taxa are rooted in Late Cambrian stocks. The Early Ordovician displays a steady increase in rhynchonelliformean biodiversity, which was mostly driven by the increasing success of the Porambonitoidea and Orthoidea, but the billingsellids and early plectambonitoids also contributed to this expansion. During the Early to Mid Ordovician (Dapingian–Darriwilian), marine life experienced an unprecedented hike in diversity at the species, genus and family levels that firmly installed the suspension-feeding benthos as the main component of the Palaeozoic fauna. However, this may have occurred in response to an early Darriwilian annihilation of existing clades, some of which had been most successful during the Early Ordovician. New clades rapidly expanded. The continents were widely dispersed together with a large number of microcontinents and volcanic arcs related to intense magmatic and tectonic activity. Climates were warm and sea-levels were high. Pivotal to the entire diversification is the role of gamma (inter-provincial) diversity and by implication the spread of the continents and frequency of island arcs and microcontinents. The phylogeographical analysis demonstrates that this new palaeogeographical configuration was particularly well explored and utilized by the strophomenides, especially the Plectambonitoidea, which radiated rapidly during this interval. The porambonitoids, on the other hand, were still in recovery following the early Darriwilian extinctions. Orthides remained dominant, particularly at high latitudes. Biodiversity epicentres were located on most of the larger palaeoplates, as well as within the Iapetus Ocean. Provincial patterns were disrupted during the Sandbian and early Katian with the migration of many elements of the benthos into deeper-water regimes, enjoying a more cosmopolitan distribution. Later Katian faunas exhibit a partition between carbonate and clastic environments. During the latest Katian, biogeographical patterns were disrupted by polewards migrations of warm-water taxa in response to the changing climate; possibly as a consequence of low-latitude cradles being developed in, for instance, carbonate reef settings. Many clades were well established with especially the strophomenides beginning to outnumber the previously successful orthides, although this process had already begun, regionally, in the mid to late Darriwilian. At the same time, atrypoid and pentameroid clades also began to radiate in low-latitude faunas, anticipating their dominance in Silurian faunas. The Hirnantian was marked by severe extinctions particularly across orthide-strophomenide clades within the context of few, but well-defined, climatically controlled provincial belts. Supplementary material: The individual localities and a reference list for the data sources are provided at: http://www.geolsoc.org.uk/SUP18667

Abstract The palaeogeographical distributions of Early Palaeozoic bryozoan faunas are reviewed. Previous studies are examined and new databases have been assembled of the stratigraphical and geographical distribution of Ordovician and Silurian taxa. Analysis was carried out using cluster analysis based on Jaccard’s coefficient and paired group method, as well as principal coordinate analysis based on Jaccard’s coefficient, to examine the relationships between different localities. Bryozoan faunas increased in diversity throughout the Ordovician peaking with 133 genera during the Katian. In the earliest Ordovician provincialism is difficult to determine, but by the Darriwilian five distinct provinces developed, decreasing to four in the Sandbian. There was a decrease in provinciality throughout the Katian as faunas became less endemic, caused by the reduction of geographical barriers. Following the extinction of many genera at the end of the Ordovician, early Silurian faunas contain remnant taxa. Subsequently fenestrates began to dominate faunas. During the Llandovery bryozoans began to show distinct provincialism, but this declined during the Wenlock, only to re-emerge during the Ludlow. Late Silurian (Pridoli) faunas are sparse but nevertheless show possible division into two provinces.

Abstract The distribution of all known Cambrian echinoderm taxa, encompassing both articulated specimens and taxonomically diagnostic isolated ossicles, is documented for the first time. The database described by 2011 comprises 188 species recorded from 65 formations from around the world. Formations that have yielded articulated echinoderms are unequally distributed in space and time. Only Laurentia and West Gondwana provide reasonably complete records at the resolution of Stage. The review of the biogeographical distributions of the eight major echinoderm clades shows that faunas from Laurentia and Northeast Gondwana (China and Korea) are distinct from those of West Gondwana and Southeast Gondwana (Australia); other regions are too poorly sampled to make firm palaeobiogeographical statements. Analysis of alpha diversity (species per formation) shows that diversity rose initially to Cambrian Stage 5, declined into Guzhangian and Paibian before returning to Stage 5 levels by the end of the Cambrian. This pattern is replicated in Laurentia and West Gondwana. We show that taxonomically diagnostic ossicles found in isolation typically occur significantly earlier than the first articulated specimens of the same taxa and provide important information on the first occurrence and palaeobiogeographical distribution of key taxa, and of the phylum as a whole. Supplementary material: Articulated Cambrian echinoderms and Isolated plates of Cambrian echinoderms are provided at: http://www.geolsoc.org.uk/SUP18668

Abstract The palaeobiogeographical distribution of the six major clades of Ordovician echinoderms (asterozoans, blastozoans, crinoids, echinozoans, edrioasteroids and stylophorans) is analysed based on a comprehensive and up-to-date database compiling 3701 occurrences (1938 species recorded from 331 localities) of both complete specimens and isolated ossicles. Although historically biased towards a limited number of regions (Europe, North America, Russia), the resulting dataset makes it possible to identify six main palaeobiogeographical provinces for Ordovician echinoderms: Laurentia, Baltica, West Gondwana, East Gondwana, Avalonia and Siberia. At a global scale, the high endemicity of echinoderms during the Early to Middle Ordovician coincides with the time of maximum dispersal of continental masses. Late Ordovician faunas tend to become more cosmopolitan, possibly as a consequence of changing palaeogeography and/or relatively higher sea-levels in the Sandbian–Katian interval. Regional biodiversity patterns of Ordovician echinoderms confirm that their major diversification during the Ordovician is not a single, universal evolutionary event, but rather results from the complex addition of contrasted local evolutionary trends.

Abstract The biogeographical distribution of Ordovician and Silurian gastropods, monoplacophorans and mimospirids has been analysed on a generic level. The dataset contains 334 genera and 2769 species, yielding 1231 records of genera with 2274 occurrences worldwide. There is a bias towards eastern Laurentia, Baltica and Perunica records. Some 53.1% of the records are Ordovician. The study demonstrates that these molluscs are well suited to being used to improve understanding of Ordovician and Silurian biogeographical provinciality. Specific points are that: a Lower Ordovician assemblage is evident in Laurentia; the fauna of the Argentinean Precordillera is Laurentian until the Darriwilian, when taxa are shared with North China; Late Silurian gastropods from the Alexander terrane (SE Alaska) are unknown in Laurentia, but support a rift origin of this terrane from NE Siberia; Perunica, Ibero-Armorica and Morocco cluster together throughout the Ordovician but Perunica and Morocco are closer; Darriwilian–Sandbian deep-water Bohemian taxa occur in Baltica; a Laurentian–Baltica proximity is unsupported until the Silurian; Siberia clusters with North China and eastern Laurentia during the Tremadocian–Darriwilian; during the Gorstian–Pridoli Siberia clusters with the Farewell and Alexander terranes; North China may have been close to Laurentia and the Argentinean margin of Gondwana; and the affinity of Tarim taxa is problematic.

Abstract Bivalves first appeared in the Early Cambrian and were virtually cosmopolitan. These very small and insignificant molluscs were probably surface crawlers on the microbial mat floors of the Cambrian sea. They evolved little further in the Mid Cambrian and by the Late Cambrian had apparently disappeared from the fossil record. Their re-appearance in the Early Ordovician coincided with a major diversification in which all the principal bivalve clades evolved, but the class was confined to Gondwana; their habitat was now principally infaunal in siliciclastic sediments. In the Mid Ordovician a few forms reached Baltica and the eastern Laurentian margin, but it was the Late Ordovician before bivalves once again became cosmopolitan. This geographical dispersal allowed bivalves to colonize the low-latitude carbonate platforms and led to the development of diverse epifaunal faunas, although most remained as infaunal forms in siliciclastic sediments. The end-Ordovician regression occasioned by the Hirnantian glaciation caused major extinction of those epifaunal forms restricted to the carbonate platforms. The Silurian faunas were cosmopolitan and the major evolutionary event was caused by the appearance of a Gondwanan cephalopod limestone facies that provided sites for epibyssate praecardiidinid bivalves (=Nepiomorphia) that evolved rapidly and were able to withstand short periods of anoxia.

Abstract The geographical distribution of rostroconch taxa of the orders Ribeirioida and Conocardiida, mostly across the equatorial continents and oceans, is compiled for six presumed diversity acmes in the early Early Ordovician ( c. 485 Ma), the late Early–early Middle Ordovician ( c. 475 Ma), the Late Ordovician ( c. 455 Ma), the Early Silurian ( c. 435 Ma), the late Early–early Late Silurian ( c. 425 Ma) and the early Early Devonian ( c. 415 Ma), based on our present, uneven knowledge. Rostroconchs show distribution patterns which enable a provisional separation of biogeographical provinces at least from the Silurian onward, comparable with Late Palaeozoic rostroconch distributional patterns. The distributions of tabulate corals, trilobites and bivalves appear roughly comparable, but not the nektic and planktic groups. A restriction to low latitudes (‘tropical’ realms) is clear for members of Ribeiria and Eopteria in the Early Ordovician, similar to proven patterns for hippocardiid rostroconchs from the Late Silurian onward until the Middle Permian. Preliminary rostroconch provinces or subprovinces, respectively, are currently discernable in the Silurian and Devonian for northwestern Laurentia, south-central Laurentia/Baltica and the north-central margin of Gondwana (i.e. Perunica, Bohemia). Rostroconch distributional data for SE Asia, China, Kazakhstan, Siberia and Australia are sparse and require further studies.

Abstract Eunicidan polychaetes formed a significant part of Early Palaeozoic marine invertebrate communities, as shown by the abundance and diversity of scolecodonts (polychaete jaws) in the fossil record. In this study we summarize the early radiation and biodiversity trends and discuss the palaeobiogeography of these fossils. The oldest (latest Cambrian–Early Ordovician) representatives had primitive, usually symmetrical, placognath/ctenognath type jaw apparatuses. The first more advanced taxa, possessing labidognath-type jaw apparatuses or placognath apparatuses with compound maxillae, are first recorded in the Middle Ordovician. The most significant increase in generic diversity occurred in the Darriwilian, when many common taxa appeared and diversified. The Ordovician and Silurian scolecodont occurrences allow some palaeobiogeographical units and distribution patterns to be explored and outlined. The most robust data presently at hand derive from successions in Baltica and Laurentia. That information, together with new records from other palaeocontinents, reveals a wide distribution for the most frequent and species-rich genera and families, similar to the biogeographical patterns of extant polychaetes. Like many other benthic and pelagic fossil groups, scolecodont-bearing polychaetes show an increased cosmopolitan character in the Silurian as compared with the Ordovician. Species-level endemism appears to be relatively common, inferring a potential for scolecodonts as biogeographical tools in the future.