The euarthropod Luohuilinella deletres sp. nov. is described from rare material from the Chengjiang biota, Cambrian Series 2, Stage 3, of Yunnan Province, China. Phylogenetic analysis recovers a xandarellid affinity for L. deletres, representing only the fifth described species of this clade. L. deletres possesses a head shield that is about one-fifth of the total body length and a trunk with 30 tergites, the reduced anterior-most tergite and terminal three tergites lacking pleural elongations. Anteriorly situated notches in the head shield are associated with stalked eyes, in contrast to the more posterior, enclosed eye slits present in Xandarella. Posterior to the antennae there are at least 11 pairs of biramous appendages preserved, including three pairs in the head. The morphology of the midline gut of L. deletres, in which lateral, unbranched diverticula are wider towards the front of the body, is a characteristic also found in various trilobites. The dorsoventrally flattened exoskeleton suggests a benthic or nektobenthic mode of life for L. deletres, as for other trilobitomorphs, and it likely used its well-developed anteriorly positioned eyes for searching out food, either to scavenge or to find prey.

The marine offshore shelf mudstones of the Early Cambrian Buen Formation at Sirius Passet, North Greenland, contain a rich Konservat-Lagerstätte which includes abundant well-preserved material of the bivalved arthropod Isoxys volucris new species. The new material confirms Isoxys Walcott, 1890 as a component of the earliest arthropod faunas worldwide. Isoxys species are known from the Early Cambrian of Spain, Siberia, South Australia and Southwest China and also from the Early to Middle Cambrian of Laurentian North America. Isoxys occurs in the Redlichiid, Bigotinid and Olenellid trilobite faunal realms but is restricted to within tropical/subtropical regions, attesting to possible paleolatitudinal controls on its distribution. Isoxys resembles some phyllocarid and bradoriid arthropods but without knowledge of its soft-parts the affinity of the genus remains uncertain.

The discovery of domatial dimorphism in the leperditellid ostracode Loculocavata n. gen. and the monotiopleurid Primitiella minima (Harris), and the possibility that Leperditella Ulrich exhibits domatial dimorphism, requires a revision of the concept and/or diagnosis of the Platycopina. Domatial dimorphism is manifested by a special domiciliar space for egg/brood care in females. Domatia are represented by a loculate domiciliar wall or by a more generalized space bounded anteriorly by an interior partition caused by thickening or folding (sulcation) of the shell wall. Domatial dimorphism occurs in morphologically diverse ostracodes as shown by the podocopine Mesocyprideis, kloedenellids, cytherellids, monotiopleurids, the leperditellid Loculocavata n. gen., and by its possible occurrence in Hypotetragona and Leperditella. Changes in the present concept and/or diagnosis of the Platycopina are required to account for the above. Possible classifications range from restricting the Platycopina to holosolenic ostracodes with R/L overlap and domatial dimorphism to including within the Platycopina all groups that exhibit domatial dimorphism. The position of adductor muscle attachment in Loculocavata n. gen. and Primitiella minima (Harris) suggests that L2 may be the external expression of the position of adductor muscle attachment in some ostracodes.

A fossil priapulid, Eximipriapulus globocaudatus new genus new species, is described from the Cambrian Chengjiang Lagerstätte of Yunnan, China. The exceptional preservation of the animal reveals morphological details that allow direct comparison with extant priapulids. The body is divisible into a partially eversible pharynx, a smooth collar, a scalid-bearing introvert, a neck with triangular scalids, an unsegmented trunk with annulations, and a distinctly expanded terminal region. Several specialized regions of the alimentary canal are recognized: a pharynx (lined by cuticle and bearing teeth), esophagus, midgut, hindgut, and a terminal anus. The sample includes a putative juvenile. The animal is inferred to have been an active burrower using a double-anchor strategy, practicing both deposit feeding and carnivory. Inclusion of Eximipriapulus in the most recent character matrix for cladistic analysis of fossil and Recent priapulids resolves the new genus within the priapulid crown group.

Here we report the first detailed reconstruction of the eye and visual system of a Paleozoic ostracod, namely the dimorphic primitiopsid (Palaeocopa) Primitiopsis planifrons Jones, 1887. Evidence from the cuticular lens morphology and its position on the valve suggests that P. planifrons had a naupliar eye, which is the most common optical system in crustaceans. the characters of the eye, such as the separated cuticular lens on each valve and the strongly calcified nature of the lens and its diameter, are concordant with that of some Recent podocopid ostracods in which a divided type of naupliar eye is present. Ray tracing of the reconstructed cuticular lens of the primitiopsid species demonstrates strong spherical aberration and a very long focal length (as in the eye of podocopids), which implies the presence of a spherical mirror (tapetum) below the lens. Study of the ontogenetic change of the valve thickness aids the suggestion that P. planifrons was nektobenthic. the size of its cuticular lens in relation to that of living podocopid ostracods, and the water depth at which these Recent forms occur, indicates that P. planifrons may have lived in a relatively shallow, well lit environment such as the deep subtidal zone. These interpretations are consistent with supposed paleoenvironmental conditions derived from geological evidence. Thus, analysis of the nature and relative size of the eye gives an independent test to help identify paleoenvironmental parameters of long extinct ostracods.

Systematic collecting through the upper Wenlock (upper Homerian) and lower Ludlow (Gorstian and lowermost Ludfordian) Silurian rock succession of the Long Mountain, Powys, Wales, identifies some 48 chitinozoan species that distinguish four biozones, two subzones and an interregnum. Consideration of the chitinozoan biozones together with those of the graptolites enables a local three-fold subdivision of the late Homerian lundgreni graptolite Biozone, and the distinction of lower and upper intervals for the Gorstian incipiens graptolite Biozone. The base of the Ludlow Series in the Long Mountain more or less equates to the base of the Cingulochitina acme chitinozoan Biozone, although no key chitinozoan first or last appearance datums are associated with the series boundary itself. The new graptolite–chitinozoan biozonation allows enhanced correlation between upper Wenlock and lower Ludlow sedimentary deposits of the Lower Palaeozoic Welsh depositional basin and those of the palaeo-shelf in the stratotype Wenlock and Ludlow areas of Shropshire. Chitinozoans seem affected by the phenomena that caused the late Wenlock ‘Mulde extinction’ in graptolites but, with the final disappearance of 9 species and re-appearance of 11 species following an interval of overall low diversity, they seem to have suffered less severely than their macro-zooplanktonic contemporaries.

Bradoriids are small bivalved arthropods that had global distribution for about 20 million years beginning at Cambrian Epoch 2 (c. 521 Ma). The majority of bradoriids are considered to be benthic, favouring oxygenated waters, as suggested by their anatomy, lithofacies distribution, faunal associates and provinciality. Most bradoriids were extinct by the end of the Drumian Age (middle of Cambrian Epoch 3). The post-Drumian is characterized by widespread dysoxic shelf lithofacies in southern Britain and Scandinavia and by the abundance of phosphatocopid arthropods. This interval is also associated with two bradoriid species with wide intercontinental distribution: Anabarochilina primordialis, which had a geographical range from the palaeo-tropics to high southern palaeo-latitude, and Anabarochilina australis, which extended through the palaeo-tropics from Laurentia to Gondwana. The wide environmental and geographical range of these species, coupled with a carapace anatomy that suggests an active lifestyle, is used to infer a zooplanktonic lifestyle. A possible driver of this widespread Cambrian bradoriid zooplankton was sea-level rise coupled to the periodic spread of low oxygen conditions onto continental shelves, acting in tandem with anatomical pre-adaptations for swimming. Parallels exist with the myodocope ostracod colonization of the water column during Silurian time, which may also have been influenced by extrinsic environmental controls acting on anatomical pre-adaptations for swimming. Similar biological and environmental mechanisms may have facilitated arthropod zooplankton colonizations across Phanerozoic time.

Here we record the occurrence of a new species of the Herefordshire Lagerstätte ostracod genus Pauline from the Lower Silurian (upper Telychian) of North Greenland. Pauline nivisis sp. nov. was recovered from a limestone boulder (Pentamerus Bjerge Formation) collected south of Kap Schuchert, Washington Land. It is reasonable to transpose the palaeobiology known from the Herefordshire Pauline avibella – body, limbs including swimming antennae, lateral eyes, gills and alimentary system – into the carapace of the Greenland species, which represents the oldest cylindroleberidid myodocopid and almost the oldest known myodocope, and is the first record of a Herefordshire Lagerstätte genus from outside the Welsh Borderland locality. Morphological, sedimentological and faunal evidence suggest that the Greenland species was nektobenthic. This is compatible with the notion that ostracods (specifically myodocopids) did not invade the water column until later in the Silurian, in the Wenlock and Ludlow epochs. Pauline is an Early Silurian link between ‘Baltic-British’ and North Laurentian ostracod faunas, endorsing the idea that the UK and Greenland were in close geographical proximity, near a remnant Iapetus Ocean, during late Llandovery time.

A study of type and newly collected material of the Silurian ostracod Richteria migrans (Barrande, 1872) demonstrates that it had wide distribution, occurring in at least the Czech Republic, France, Sardinia, Wales, central Asia and probably Poland. R. migrans has biostratigraphic value, as an indicator for the mid to late part of the Ludlow Series. It was almost certainly pelagic, living predominantly in probable shelf topographic lows to marginal/off-shelf environments, characteristically with cephalopod-graptolite-bivalve-dominant associates. It had at least five to six growth stages, exhibits polymorphic variation, and its morphology provides evidence to endorse the notion that ‘entomozoaceans’ are myodocopes.

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.

The Cautley Mudstone Formation and Cystoid Limestone Member of the Ashgill Formation (Windermere Supergroup; Ashgill Series), from the Cautley district of northern England, has yielded an ostracod fauna of more than 30 species. Many of these have short ranges, permitting recognition of stratigraphically successive Pusgillian–lower Cautleyan, middle–upper Cautleyan, and Rawtheyan ostracod faunas. Several species are also known from the upper Ordovician of North America (Anticosti Island), Scotland (Girvan district) and the Baltic region (Estonia, glacial erratic boulders of northern Germany), providing evidence to correlate upper Ordovician successions in these areas. The ostracods include abundant podocopes, at some horizons accounting for more than 80% of the fauna. Binodicopes are also common, but palaeocopes are rare. Assemblages are typical of a clastic dominated open marine shelf setting. Diversity at most horizons is low (c. 3–5 species), but reaches a peak of between 13–14 species in middle Cautleyan horizons. Lower diversity at Pusgillian and Rawtheyan horizons coincides with the encroachment of deeper marine-shelf facies which were probably hostile to Ordovician benthonic ostracods. Some of the ostracods (particularly Aechmina) have distributions suggesting tolerance of a range of mid- to deep shelf benthonic palaeoenvironments, but none were pelagic. During Ashgill times the Cautley district (part of palaeocontinental Avalonia) was replete with ostracod genera and species which also occur in the Baltic region (palaeocontinental Baltica; more than 90% generic similarity) and to a lesser, but nonetheless significant extent in North America and Scotland (parts of palaeocontinental Laurentia). Such trans-Tornquist Sea and Iapetus Ocean distributional patterns add to previous ostracod data that support models which show palaeogeographical proximity of Avalonia and Baltica, and Avalonia and Laurentia, by Ashgill times. The widely cited observation, that trans-Iapetus ostracod faunas remained strictly provincial until the mid-or late Silurian, cannot be sustained.

This paper evaluates the taxonomy, biostratigraphy, and palaeogeographical significance of the Cambrian bradoriid arthropods of China, the majority of which occur in the lower Cambrian of SW China. Of bradoriid faunas world-wide, Chinese occurrences yield the greatest number of specimens and a comparatively high diversity at all taxonomic levels. Nevertheless, taxonomic diversity is much less than previously supposed. Some 80 bradoriid genera and nearly 300 species have been proposed on the basis of Chinese material. By contrast, in our study, which encompasses all of the important Chinese bradoriid faunas, we recognise only 16 genera and 21 species, including those treated under open nomenclature. Interpretation of deformed specimens as discrete species and lack of application of the rules of the International Code of Zoological Nomenclature resulted in taxonomic splitting and a proliferation of names. There are an additional 12 poorly known monotypic genera of uncertain systematic status that are listed but not treated further herein. One phosphatocopid species, a group originally thought closely related to the Bradoriida, is also described.

Most Chinese bradoriid material is known from Yunnan Province; the group also occurs in Guizhou, Henan, Hubei, Hunan, Liaoning, Qinghai, Shaanxi, Sichuan, Xinjiang and Zhejiang provinces. The first bradoriids occur just below the Abadiella trilobite Biozone. They are most prolific and diverse in the Qiongzhusian Stage, constituting the most abundant animal group; the succeeding Canglangpuian Stage contains fewer individuals and species. A previously proposed bradoriid biozonal scheme lacks rigour and is of little practical value: of the five supposed biozones, two correspond to trilobite zones and three are based on taxa that herein are considered to belong to a single species.

Palaeogeographically the bradoriids occur in the Middle and especially the Western subprovinces of the Cambrian of the SW China (Yangtze) Platform. Almost all of the bradoriid genera and species are endemic to that region. The palaeogeographical links with other bradoriid faunas are mostly within the Redlichiid trilobite Realm, with areas such as N China, Australia and parts of central Asia.

Marine shelf siliciclastics of the early Cambrian Buen Formation of North Greenland have yielded the earliest abundant, well localised Bradoriida (Arthropoda) from the Laurentian part of North America: the shelf-dwelling Petrianna gen. nov. Petrianna apparently has a fairly rigid, phosphatic shell. Its assignment to the Cambriidae, a family markedly dissimilar morphologically from the phosphatocopines (the only Bradoriida hitherto regarded as having a primary phosphatic shell), implies rejection of the criterion of carapace composition in the ordinal/subordinal classification of Bradoriida.

Petrianna confirms the Bradoriida as a consistent component of the earliest arthropod faunas. Cambriids are currently known only from the early Cambrian of Greenland, Siberia and China and offer potential for correlation. Cambriid occurrences span parts of the Redlichiid, Olenellid and Bigotinid trilobite faunal realms but are known only from tropical/subtropical regions, thus indicating possible palaeolatitudinal controls on their distribution.

Cyclotron lapworthi (Groom, 1902), one of the few British Cambrian bradoriid ostracod species known from a large number of specimens, is redescribed. It has an interdorsum, thus indicating a hesslandonid affinity. The ontogeny of C. lapworthi does not demonstrate a clear distinction of its individual moult stages but does show marked changes in the development of nodes and lobes. C. lapworthi shows wide variation in carapace shape possibly indicating that its valves were thin and relatively flexible. In England and Canada C. lapworthi is restricted to the Upper Cambrian Olenus Zone. C. lapworthi may also occur coevally in the Upper Cambrian of Sweden.

The monotypic ostracode Entomozoe (pro Entomis) (Superorder Myodocopa) from the Silurian of Scotland, type genus of the Entomozoidae, is revised for the first time since being established in 1861. Based on the type and new collections of Entomozoe tuberosa (Jones, 1861), the genus shows affinity to bolbozoid myodocopes rather than to forms hitherto universally considered as ‘typical’ entomozoid myodocopes. The families Entomozoidae and Bolbozoidae are considered synonymous. In the interests of stability of nomenclature, in particular to preserve the traditionally understood usage of the taxon Entomozoidae, a case must be made to the International Commission on Zoological Nomenclature to set aside any previous designations and to establish a new type-species for Entomozoe.

Entomozoe, from the Llandovery Series, is the earliest known myodocope in the fossil record. It occurs as part of a low diversity, shallow water, nearshore assemblage. In contrast to other, but younger, Silurian bolbozoid myodocopes, it is probably a benthic form. The differences in the anterior shell morphology of Silurian bolbozoid taxa possibly reflect changes both in soft-part morphology and from benthic to pelagic lifestyles within the group.

Both bolbozoid and ‘entomozoid’ myodocopes are assigned to the Order Myodocopida rather than to the Order Halocypridida, though the taxonomic affinity of the ‘entomozoids’ is, at best, uncertain.

In this paper, on page 375, we published an Arenig palaeogeography in Figure 6 which showed the subduction zone off Laurentia as dipping northwards. This was a drafting error. The subduction zone should have dipped southward away from Laurentia. We apologise for this error and publish here a revised Figure 6 for the Arenig.

The available data from Newfoundland, the British Isles and Scandinavia suggest that by late Ordovician–early Silurian times the ocean separating Laurentia from Eastern Avalonia and Baltica had partly closed with the consumption of intervening oceanic crust. Marine seaways, however, persisted until the middle or late Silurian. Phases of crustal transtension and transpression, predominantly under a major sinistral shear couple, occurred throughout the Silurian and early Devonian until the remnant Iapetus Ocean was completely destroyed. The most appropriate Recent plate tectonic models for Silurian sedimentation between Eastern Avalonia and Laurentia are probably the deep-marine foreland basins between Timor and the northern Australian margin, or between Taiwan and mainland China.

I. ABSTRACT: Research interest in the Highland Border Complex has been pursued sporadically during the past 150 years. The results and conclusions have emphasised the problems of dealing with a lithologically disparate association which crops out in isolated, fault-bounded slivers along the line of the Highland Boundary fault. For much of the present century, the debate has centred on whether the rocks of the complex have affinities with the Dalradian Supergroup to the N, or are a discrete group. Recent fossil discoveries in a wide variety of Highland Border rocks have confirmed that many are of Ordovician age, and hence cannot have been involved in at least the early Grampian deformational events (now accurately dated as pre-Ordovician) which affect the Dalradian Supergroup. Such palaeontological discoveries form the basis for a viable biostratigraphical synthesis. On a regional scale, it is apparent that the geological history of the Highland Border rocks must be viewed in the context of plate boundary tectonism along the entire northwestern margin of Iapetus during Palaeozoic times.

II. ABSTRACT: Silicified articulate brachiopods from the Lower Ordovician (Arenig) Dounans Limestone are extremely rare but the stratigraphically diagnostic genera Archaeorthis Schuchert and Cooper, and Orthidium Hall and Clarke, have been identified. In addition, three specimens with characteristic syntrophiid morphology have been recovered. Inarticulate brachiopods are known from Stonehaven and Bofrishlie Burn near Aberfoyle, and have also been previously recorded from Arran.

III. ABSTRACT: Micropalaeontological investigation of the Highland Border Complex has produced a range of microfossils including chitinozoans, coleolids, calcispheres and other more enigmatic objects. The stratigraphical ranges of the species lie almost entirely within the Ordovician and reveal a scatter of ages for different lithologies from the Arenig through to the Caradoc or Ashgill, with a pronounced erosional break between the Llandeilo and the Caradoc.

IV. ABSTRACT: A Lower Ordovician (Arenig Series) silicified ostracode fauna from the Highland Border Dounans Limestone at Lime Craig Quarry, Aberfoyle, Central Scotland, represents the earliest record of this group of Crustacea from the British part of the early Palaeozoic ‘North American’ plate.

V. ABSTRACT: Palaeontological age determinations for a variety of Highland Border rocks are presented. The data are based on the results of recent prospecting which has demonstrated that macro- and microfossils are present in a much greater range of Highland Border lithologies than previously realised. Data from other studies are also incorporated, as are modern taxonomie re-assessments of older palaeontological discoveries, in a comprehensive survey of Highland Border biostratigraphy. These accumulated data demonstrate that all fossiliferous Highland Border rocks so far discovered are of Ordovician age, with the exception of the Lower Cambrian Leny Limestone.

VI. ABSTRACT: The Highland Border Complex consists of at least four rock assemblages: a serpentinite and possibly other ophiolitic rocks of Early or pre-Arenig age; a sequence of limestones and conglomerates of Early Arenig age; a succession of dark shales, cherts, quartz wackes, basic lavas and associated volcanogenic sediments of Llanvirn and ? earlier age; and an assemblage of limestones, breccias, conglomerates and arenites with subordinate shales of Caradoc or Ashgill age. At least three assemblages are divided by unconformities and in theirmost general aspect have similarities with coeval rocks in western Ireland.

The Highland Border Complex probably formed N of the Midland Valley arc massif in a marginal sea comparable with the Sunda shelf adjacent to Sumatra–Java. Strike-slip and thrust emplacement of the whole Complex in at least four episodes followed the probable generation of all or part of its rocks by pull-apart mechanisms.