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Brachiopods are among the first animal phyla to emerge from the Cambrian Explosion, rapidly diversifying to all major palaeocontinental blocks within 20 million years. The group underwent another steep rise in diversity during the Ordovician, and their relative abundance and diversity made them one of the most successful invertebrate groups during the entire Palaeozoic. During this time, brachiopods lived in a range of environments and represented a significant component of marine ecosystems, yet information regarding their modes of life and ecology is somewhat limited. Recent studies, primarily from the Chengjiang and Burgess Shale Lagerstätten, have revealed that by the Middle Cambrian (Series 3, Stage 5) brachiopods from across the phylum had already developed a range of ecological strategies and life modes. Cambrian brachiopods occupied distinct trophic niches on soft and hard substrates and exhibited at least five types of lifestyles: pedicle attachment, pedicle anchoring, cemented, free‐lying and semi‐infaunal. Comparisons with Ordovician benthic assemblages show that despite the explosion of brachiopod taxa witnessed in the Ordovician, with the exception of the appearance of burrowing brachiopods, life strategies of brachiopods remained largely the same. Indicating that the majority of life modes observed in brachiopods had rapidly evolved and was already in place prior to the Great Ordovician Biodiversity Event., Funding via a Marie Curie COFUND Research Fellowship (Durham University) is acknowledged (Topper); Harper thanks the Leverhulme Trust for a research fellowship and the Wenner‐Gren Foundation for support.

The Sirius Passet Lagerstatte (SP), Peary Land, North Greenland, occurs in black slates deposited at or just below storm wave base. It represents the earliest Cambrian microbial mat community with exceptional preservation, predating the Burgess Shale by 10 million years. Trilobites from the SP are preserved as complete, three-dimensional, concave hyporelief external moulds and convex epirelief casts. External moulds are shown to consist of a thin veneer of authigenic silica. The casts are formed from silicified cyanobacterial mat material. Silicification in both cases occurred shortly after death within benthic cyanobacterial mats. Pore waters were alkali, silica-saturated, high in ferric iron but low in oxygen and sulphate. Excess silica was likely derived from remobilized biogenic silica. The remarkable siliceous death mask preservation opens a new window on the environment and location of the Cambrian Explosion. This window closed with the appearance of abundant mat grazers later as the Cambrian Explosion intensified.

Schoenemann, B. & Clarkson, E.N.K. 2012: The eyes of Leanchoilia. Lethaia, Vol. 45, pp. 524–531. Leanchoilia is one of the commonest non-trilobite arthropod taxa of the Lower and Middle Cambrian. It has recently been regarded to be part of the ‘short great appendage arthropods’ clade and is normally considered a predator, both on account of its raptorial appendages and serial midgut digestive glands. The presence or absence of eyes has been one of the most controversial aspects of its morphology, despite pedunculate eyes having been described in the original descriptions of L. superlata (Walcott 1912) and L. illecebrosa Hou 1987. Other authors considered all species of this genus to be blind. There are two pairs of pits on the head, which have been interpreted as eyes attached near the front of the headshield. Pedunculate eyes have been found in four of about fifty specimens of L. illecebrosa from the Chengjiang and Haikou areas. They are delicate and slender, which explains their rare preservation. Distally they are structured like a bunch of grapes, with about 100 spherical units, presumably lenses, each with a diameter of about 55 μm indicating, by calculation, adaptation to dim light. They probably could detect movement, but could not form images. Furthermore, L. illecebrosa possesses two, probably ventral, median eyes, autapomorphies of euarthropods. Thus, like many modern Euarthropoda, L. illecebrosa has two kinds of eyes – two stalked ventral eyes and two pairs of probably ventral (ocellar) median eyes. Its short stalked eyes, orientated downwards and covered by the headshield, suggest that species of Leanchoilia were benthic animals, feeding on benthic or infaunal animals. □Arthropoda, Burgess Shale, Cambrian, Chengjiang, Compound eye, Lagerstatte, Leanchoilia.

Garson, D.E., Gaines, R.R., Droser, M.L., Liddell, W.D. & Sappenfield, A. 2011: Dynamicpalaeoredox and exceptional preservation in the Cambrian Spence Shale of Utah. Le-thaia, DOI:10.1111⁄j.1502-3931.2011.00266.xBurgess Shale-type faunas provide a unique glimpse into the diversification of metazoanlife during the Cambrian. Although anoxia has long been thought to be a pre-requisitefor this particular type of soft-bodied preservation, the palaeoenvironmental conditionsthat regulated extraordinary preservation have not been fully constrained. In particular,the necessity of bottom water anoxia, long considered a pre-requisite, has been the sub-ject of recent debate. In this study, we apply a micro-stratigraphical, ichnologicalapproachtodeterminebottom wateroxygen conditionsunder,which BurgessShale-typebiotas were preserved in the Middle Cambrian Spence Shale of Utah. Mudstones of theSpence Shale are characterizedby fine scale (mm-cm)alternation between laminated andbioturbated intervals, suggesting high-frequency fluctuations in bottom water oxygena-tion. Whilst background oxygen levels were not high enough to support continuousinfaunal activity, brief intervals of improved bottom water oxygen conditions punctuatethe succession. A diverse skeletonized benthic fauna, including various polymerid trilo-bites, hyolithids, brachiopods and ctenocystoids suggests that complex dysoxic benthiccommunity was established during times when bottom water oxygen conditions werepermissive. Burgess Shale-type preservation within the Spence Shale is largely confinedto non-bioturbated horizons, suggesting that benthic anoxia prevailed in intervals, wherethese fossils were preserved. However, some soft-bodied fossils are found within weaklyto moderately bioturbated intervals (Ichnofabric Index 2 and 3). This suggests that Bur-gess Shale-type preservation is strongly favoured by bottom water anoxia, but may notrequire it in all cases. h Anoxia, Burgess Shale, Burgess Shale type-preservation, LangstonFormation, Spence Shale Member,Utah.Daniel E. Garson [daniel.garson@gmail.com], Department of Earth Sciences, University ofCalifornia, Riverside, CA 92521, USA; Robert R. Gaines [Robert.Gaines@pomona.edu],Geology Department, Pomona College, 185 E. Sixth St., Claremont, CA 91711, USA; MaryL. Droser [mary.droser@ucr.edu], Department of Earth Sciences, University of California,Riverside, CA 92521, USA; W. David Liddel [dave.liddell@usu.edu], Department ofGeology, Utah State University, Logan, UT 84322-4505, USA; Aaron Sappenfield[aaron.sappenfield@email.ucr.edu], Department of Earth Sciences, University of California,Riverside, CA 92521, USA; manuscript received on 14 September 2010; manuscript acceptedon 04 February 2011.

Opabinia regalis has long been regarded as a curious animal, with its five eyes, its long flexible anterior process, and gill lamellae carried on the outside of overlapping lateral lobes. More recently, Opabinia has been reconstructed with lobopod limbs lying adaxial but separate from the lateral lobes. This version of Opabinia represented a lobopod–arthropod transition and prompted a hypothesis for the origin of the biramous limb that involved uniting the lobopod limb with a lateral lobe. New evidence of elemental maps is consistent with previous interpretations of the triangular structures in Opabinia as lateral extensions of the gut; there is no convincing evidence for the presence of lobopod limbs. Re-examination of critical specimens reveals that the gill lamellae are not on the outside of the lateral lobes. The limbs of Opabinia resemble the phyllopodous exopod of arthropods; the posterior margin is fringed with blades. Opabinia remains on the stem of euarthropods but not as a part of a paraphyletic Lobopodia. The Lobopodia is a clade of Cambrian armoured lobopods and onychophorans. A new hypothesis for the origin of the arthropod biramous limb from an exopod like that in Opabinia is presented, which involves an endite-bearing phyllopodous limb as an intermediate stage.

Echmatocrinus from the Middle Cambrian Burgess Shale of British Columbia was originally described as the earliest crinoid(?) known from the fossil record. Recently, Conway Morris and Ausich & Babcock have questioned whether Echmatocrinus is in fact an echinoderm, comparing it instead to cnidarians with a polyp-like body and pinnate tentacles, and other authors are beginning to use this reinterpretation. We studied the well-preserved holotype of Echmatocrinus brachiatus, two paratypes, and 18 new specimens recovered from different levels in the Burgess Shale sequence at three localities. All are preserved as pyrite films in dark shale with relatively little relief, suggesting a lightly skeletized body. Complete specimens have a long, slightly tapering, large-plated attachment stalk, a conical cup or calyx with numerous small to medium-sized irregular plates, and 7–10 short arms with heavier plating and (in the holotype) soft appendages alternating from opposite sides of several arms. Several morphologic features indicate that Echmatocrinus is an echinoderm and has crinoid affinities: (1) Sutured plates, shown by darker depressed sutures, slightly raised plate centers, and oriented plate ornament, cover all major parts of the body; (2) reticulate surface ornament in the pyrite film on the plates of all specimens matches the ornament in the Burgess Shale edrioasteroid Walcottidiscus, an undoubted echinoderm, but not the pyritized surfaces of other metazoans in the fauna; (3) this distinctive ornament may represent the surface expression of microporous stereom; (4) possible ligament or muscle pads are present between the arm ossicles to fold and unfurl the more heavily plated arms. Within the echinoderms, only crinoids commonly have a calyx attached by a stalk or stem to the substrate and bear erect, moveable, uniserial arms for feeding. Although Echmatocrinus shows some resemblance to octocorals in overall body shape as an attached suspension feeder, almost all the details are different, indicating that Echmatocrinus is most likely unrelated to this group. All complete specimens of Echmatocrinus are attached to hard substrates, either another fossil or skeletal debris. The new specimens indicate that Echmatocrinus was twice as common (about 0.02%) in the Burgess Shale fauna as previously recorded and represents one of the earliest attached, medium-level, skeletized, suspension feeders or microcarnivores in the fossil record.

The lower Rabbitkettle Formation of northwestern Canada is a monofacial Upper Cambrian unit of variably calcareous, argillaceous siltstone and fine-grained sandstone with rare bioclastic grainstone, deposited on a gentle slope below fair-weather wave base with no discernible fluctuation in water depth. The trilobite fauna is a mixture of pandemic agnostoids and Laurentian polymeroids, including protaspides and meraspides, and individuals are disarticulated, non-abraded and mostly oriented convex-upward. Bioclasts are interpreted as in situ elements affected only by weak bottom currents and storm-induced turbulence. A major proportion of the larger (≥5 mm across) polymeroid cranidia and pygidia in the lower part (Marjuman) of the formation are broken; large thoracic segments are often broken at the axial furrow and some broken free cheeks occur, but essentially no broken agnostoids or hypostomes were observed. Trilobites are not broken in upper beds (Steptoean), above the base of the Glyptagnostus retculatus Zone. Physical breakage cannot be dismissed entirely, but most damage is interpreted to be due to size-selective predation, possibly through lethal blows similar to those delivered by some extant stomatopod crustaceans. A possible culprit may be an animal akin to Yohoia, known from the Middle Cambrian Burgess Shale. The distribution of attacked trilobites serves as a proxy for the presence and disappearance of soft-bodied carnivores. In the Rabbitkettle Formation, it suggests that Burgess Shale-type animals may have persisted into the Late Cambrian but suffered extinction at the Marjuman-Steptoean ‘biomere’ event when most trilobite species vanished.

Opabinia regalis Walcott is an enigmatic fossil from the Middle Cambrian Burgess Shale of uncertain affinities. Recent suggestions place it in a clade with Anomalocaris Whiteaves from the Burgess Shale and Kerygmachela Budd from the Greenlandic Sirius Passet Fauna; these taxa have been interpreted as ‘lobopods’. Consideration of available Opabinia specimens demonstrates that reflective extensions from the axial region, previously thought to be either gut diverticula or musculature, can be accommodated in neither the trunk nor the lateral lobes that arise from it. They must therefore be external structures independent of the lateral lobes. On the basis of their sub-triangular appearance, size and taphonomy, they are considered here to represent lobopod limbs. Some evidence for the existence of terminal claws is also presented. The question of whether Kerygmachela, Opabinia and Anomalocaris constitute a monophyletic or paraphyletic grouping is considered. While they share several characters, most of these are plesiomorphies. Further, Opabinia and Anomalocaris share several arthropod-like characters not possessed by Kerygmachela. It is concluded that these three taxa probably form a paraphyletic grouping at the base of the arthropods. Retention of lobopod-like characters within the group provides important documentation of the lobopod-arthropod transition. A proper understanding of Opabinia and its close relatives, which may include the tardigrades, opens the way for a reconstruction of the arthropod stem-group. This in turn allows the construction of a speculative but satisfying scenario for the evolution of major arthropod features, including the origin of the biramous limb, tergites and arthropod segmentation. ‘Arthropodization’ may thus be seen not to be a single event but a series of adaptive innovations.OPABINIA, ANOMALOCARIS, KERYGMACHELA, Burgess Shale, problematica, Lobopodia, Arthropoda.

The exoskeleton in the Middle Cambrian Burgess Shale arthropod Tegopelte has previously been described as divided into tergites. Evidence from its sister taxon Saperion and from a Naraoia specimen from the Lower Cambrian Chengjiang fauna enables a reinter

Burgess-Shale-type preservation is defined as a taphonomic pathway involving the exceptional organic preservation of non-mineralizing organisms in fully marine siliciclastic sediments. In the Phanerozoic it occurs widely in Lower and Middle Cambrian sequences but subsequently disappears as a significant taphonomic mode. The hypothesis that this distribution derives solely from a secular increase in the depth of bioturbation is falsified: low bioturbation indices do not prevent the rapid enzymatic degradation of organic structure, nor do they account for the conspicuous absence of comparable preservation during the Vendian. An earlier, Late Riphean (ca. 750–850 Ma), interval of enhanced organic-walled fossil preservation suggests a long-term recurrence in Burgess-Shale-type taphonomy that is independent of metazoan activity. A model based on the potentially powerful anti-enzymatic and/or stabilizing effects of clay minerals on organic molecules is proposed to account for Burgess-Shale-type preservation. Long-term changes in average clay mineralogies and the ocean chemistry that determines their interaction with organic molecules are likely to have induced the pronounced secular distribution of these fossil biotas, while regional variations in tectonism, weathering, etc., explain their non-uniform geographic distribution; the close correlation between exceptional, organic-walled fossil preservation and volcano-genic sedimentation in Tertiary lake deposits provides a compelling analogue. Recognition of a temporal control on Burgess-Shale-type preservation constrains the evolutionary scenarios that can be drawn from such biotas; significantly, neither the initial rate of appearance, nor the ultimate fate of Burgess-Shale-type taxa can be directly assessed. □Taphonomy, exceptional preservation, organic preservation, fossil Lagerstatten, Burgess Shale, clay mineralogy, clay-organic interactions, secular change, Cambrian, Proterozoic.

Despite many papers devoted to it, the morphology of the Burgess Shale animal Opabinia regalis continues to excite controversy. In particular, the trunk region remains incompletely understood, leading to several recent attempts to interpret the fossil in radically different ways. New material of Opabinia from the Royal Ontario Museum and the Smithsonian collection, together with the recent description of comparative material of the Burgess Shale anomalocaridid Hurdia, help clarify details of its morphology, in particular with regards to the lateral lobes and setal blades. A recent reconstruction of the trunk lobes is rejected, and further evidence for the presence of trunk limbs is presented. Despite disagreements over its morphology, the phylogenetic placement of Opabinia is now relatively uncontroversial, although various derived aspects of its morphology complicate placing it precisely. □Biramous limb, Burgess Shale, exceptional preservation, Opabinia regalis. © 2011 The Authors, Lethaia © 2011 The Lethaia Foundation.

Aronson, R. B. 1992 07 15: Decline of the Burgess Shale fauna: ecologic or taphonomic restriction? An important alternative has not been tested in the debate over the origin, significance, and decline of Burgess Shale taxa: their observed stratigraphic distribution could be an artifact of increasing bioturbation. Based on information currently available, comparisons of the observed and expected stratigraphic distributions of Burgess Shale-type faunas falsify bioturbation as the exclusive cause. Ecological factors and/or taphonomic controls other than bioturbation were also responsible for the post-Cambrian decline of Burgess Shale-type faunas. Bioturbation, Burgess Shale fauna, soft-bodied animals, taphonomy.

Two Cambrian lepidote metazoans known from different respective types of preservation have been compared in order to elucidate their biology and affinities. The widely distributed Lower Cambrian Halkieria is represented by isolated hollow sclerites, probably of originally calcareous composition. The Middle Cambrian Wiwaxia is known from the Burgess Shale as isolated sclerites (scales and spines) and as more or less complete individuals. Although Halkieria sclerites were mineralized and those of Wiwaxia were probably not, there are fundamental structural and morphological similarities between the two. Both bad an imbricating scaly and spiny armour consisting of hollow sclerites with a longitudinally fibrous structure. The sclerites did not grow, but were probably moulted during the course of ontogenetic growth. Halkieria and Wiwaxia are regarded as closely related. Both are referred to the Order Sachitida He 1980. The sclerite armour of Halkieria is reconstructed on the template provided by Wiwaxia. The interpretation of sachitid sclerites as protective armour is an alternative to the interpretation by Jell (1981, Alcheringa 5)that sachitid sclerites were respiratory organs in an animal of probable annelid affinities. Sachitids are interpreted as sluggish, benthic deposit feeders that do not belong to any recognized phylum.

Hyoliths are operculate calcareous shells found in Palaeozoic rocks. Runnegar et al. (1975) suggested that they be referred to a new phylum (Hyolitha) but Marek & Yochelson (1976) and Dzik (1978) preferred to regard them as an extinct class of the Mollusca. Since the hyolith cone is not easily homologized with the monoplacophoran shell, the exoskeletons of the shelled Mollusca and the Hyolitha appear to have developed independently. Reconstructions of the anatomy of hyoliths indicate that it is unlikely that both groups shared a common molluscan ancestor. Therefore, hyoliths are probably not molluscs. Previous reconstructions of articulated hyolithids have suggested that left and right appendages (helens) curved dorsally. Crushed articulated specimens from the Burgess Shale indicate that this conclusion is incorrect; hyoltthid helens seem to have curved ventrally when the animals were alive.

Neither collapse, due to decay in a soft-bodied organism, nor compaction, due to overburden pressure, normally lead to significant lateral expansion in flattened fossils except in the case of some with rigidly mineralized skeletons. The fossils are thus analogous to a variety of two-dimensional views of a three-dimensional object. This realization provides a foundation for drawing and testing a reconstruction using either computer or manual graphic restoration methods. A complementary approach based on the photography of simple models, which is particularly useful where a complex three-dimensional morphology is under study, is described and illustrated by two examples, the Middle Cambrian arthropod Odaraia and the Upper Ordovician graptolite Direllograptus.

The Phyllopod Bed of the Burgess Shale, in which Walcott found the famous soft bodied fossils, consists of thin graded beds of calcareous siltstone and mud-stone, which are probably turbidites. The Burgess Shale was deposited on a reef front submarine fan, and the preservation of the fossils is probably due to rapid burial.

Conway Morris, S., Whittington, H. B., Briggs, D. E. G., Hughes, C. P. & Bruton, D. L. 1982: Atlas of the Burgess Shale..