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Due to inadequate preservation, pterobranchs are often difficult to identify in the fossil record, and a better understanding of preservational modes and diagenetic and metamorphic effects is needed for their recognition. Pterobranch hemichordates are common in Cambrian Stage 5 and younger sedimentary rocks, but are frequently overlooked. Often, pterobranch hemichordate colonies have been considered to be algal remains or hydroids. Re-examination of Cambrian Burgess Shale algae reveals that the genera Yuknessia and Dalyia can be recognized as putative early representatives of pterobranch hemichordates. Distinct fusellar construction of the individual zooidal tubes and branching of the creeping proximal part of the colonies are found in the morphologically similar rhabdopleurid pterobranch genus Sphenoecium. The erect tubes of Sphenoecium do not branch and can reach a length of several centimetres. The development of the fusellar construction in this taxon shows a highly irregular development of the suture patterns, but a fairly consistent height of the individual fuselli. The taxon is widely distributed in the Cambrian Series 3, but has regularly been identified as a hydroid or an alga. Sphenoecium wheelerensis from the Cambrian Wheeler Shale of Utah is described as new.

Characterized by atypical frontalmost appendages, leanchoiliids are early arthropods whose phylogenetic placement has been much debated. Morphological interpretations have differed, some of which concern critical characters such as the number of eyes and head appendages, but methodological approaches also have diverged. Here, we describe a new leanchoiliid, Yawunik kootenayi gen. et sp. nov., based on 42 specimens from the newly discovered Marble Canyon locality of the Burgess Shale (Kootenay National Park, British Columbia; middle Cambrian). This new morphotype demonstrates the presence of a four-segmented head in leanchoiliids, along with two small antero-median eyes in addition to lateral eyes. Yawunik is characterized by a 12-segmented trunk and a carinate, lanceolate telson adorned with minute spines. The ‘great appendages’ of the animal bear teeth on their two distal rami, which would have enhanced their ability to grasp prey. Attitudes of specimens, resulting from burial at multiple aspects of bedding, suggest the ‘great appendages’ were flexible and capable of antero-posterior rotation. We also discuss the nature of intersegmental tissues and filaments present within the ‘great appendages’. Our phylogenetic analyses extend the monophyly of leanchoiliids to include Haikoucaris and Yohoia in a new clade, the Cheiromorpha nom. nov. (within Heptopodomera nom. nov.). Other nodes are poorly resolved unless implied weights are used, and in this case, the topology is critically sensitive to the coding prerogative of inapplicable states (NAs). Both the traditional ‘Arachnomorpha’ hypothesis (NAs as additional states) and the more recently favoured ‘Artiopoda + Crustacea’ (NAs as uncertainties) were obtained using the same data set and outgroup. This result stresses, first, the historical importance of polarization over data content in scenarios of early arthropod evolution, and second, a pressing need to investigate the impacts of coding inapplicables, especially given the inflating effect of implied weights.

The soft-bodied Cambrian organism Wiwaxia poses a taxonomic conundrum. Its imbricated dorsal scleritome suggests a relationship with the polychaete annelid worms, whereas its mouthparts and naked ventral surface invite comparison with the molluscan radula and foot. 476 new and existing specimens from the 505-Myr-old Burgess Shale cast fresh light on Wiwaxia's sclerites and scleritome. My observations illuminate the diversity within the genus and demonstrate that Wiwaxia did not undergo discrete moult stages; rather, its scleritome developed gradually, with piecewise addition and replacement of individually secreted sclerites. I recognize a digestive tract and creeping foot in Wiwaxia, solidifying its relationship with the contemporary Odontogriphus. Similarities between the scleritomes of Wiwaxia, halkieriids, Polyplacophora and Aplacophora hint that the taxa are related. A molluscan affinity is robustly established, and Wiwaxia provides a good fossil proxy for the ancestral aculiferan � and perhaps molluscan � body plan.

We redescribe the morphology of Yohoia tenuis (Chelicerata sensu lato) from the Cambrian Burgess Shale Lagerstatte. The morphology of the most anterior, promi- nent, so-called great appendage changes throughout ontog- eny. While its principal morphology remains unaltered, the length ratios of certain parts of the great appendage change significantly. Furthermore, it possesses a special jack-knifing mechanism, i.e. an elbow joint: the articulation between the distal one of the two peduncle elements and the most proxi- mal of the four spine-bearing claw elements. This morphol- ogy might have enabled the animal to hunt like a modern spearer-type mantis shrimp, an analogy enhanced by the sim- ilarly large and protruding eyes. For comparison, details of specimens of selected other great-appendage arthropods from the Lower Cambrian Chengjiang Lagerstatte have been inves- tigated using fluorescence microscopy. This revealed that the morphology of the great appendage of Y. tenuis is much like that of the Chengjiang species Fortiforceps foliosa and Jianfen- gia multisegmentalis. The morphology of the great appendage of the latter is even more similar to the morphology devel- oped in early developmental stages of Y. tenuis, while the morphology of the great appendage of F. foliosa is more sim- ilar to that of later developmental stages of Y. tenuis. The arrangement of the elbow joint supports the view that the great appendage evolved into the chelicera of Chelicerata sensu stricto, as similar joints are found in various ingroup taxa such as Xiphosura, Opiliones or Palpigradi. With this, it also supports the interpretation of the great appendage to be homologous with the first appendage of other arthropods.

Abundant material from a new quarry excavated in the lower Cambrian Emu Bay Shale (Kangaroo Island, South Australia) and, particularly, the preservation of soft-bodied features previously unknown from this Burgess Shale-type locality, permit the revision of two bivalved arthropod taxa described in the late 1970s, Isoxys communis and Tuzoia australis. The collections have also produced fossils belonging to two new species: Isoxys glaessneri and Tuzoia sp. Among the soft parts preserved in these taxa are stalked eyes, digestive structures and cephalic and trunk appendages, rivalling in quality and quantity those described from better-known Lagerstatten, notably the lower Cambrian Chengjiang fauna of China and the middle Cambrian Burgess Shale of Canada.

The first lobopodian known from the Ordovician is described from the Soom Shale Lagerstatte, South Africa. The organism shows features homologous to Palaeozoic marine lobopodians described from the Middle Cambrian Burgess Shale, the Lower Cambrian Chengjiang biota, the Lower Cambrian Sirius Passet Lagerstatte and the Lower Cambrian of the Baltic. The discovery provides a link between marine Cambrian lobopodians and younger forms from the Silurian and Carboniferous. The new fossil preserves an annulated trunk, lobopods with clear annulations, and curved claws. It represents a rare record of a benthic organism from the Soom Shale, and demonstrates intermittent water oxygenation during the deposition of the unit.

Current models for the exceptional preservation of Burgess Shale fossils have focused on either the HF-extractable carbonaceous compressions or the mineral films identified by elemental mapping. BSEM, EDX and microprobe analysis of two-dimensionally preserved Marpolia, Wiwaxia and Burgessia identifies the presence of both carbonaceous and aluminosilicate films for most features, irrespective of original lability. In the light of the deep burial and greenschist facies metamorphism documented for the Burgess Shale, the aluminosilicate films are identified as products of late-stage volatilization and coincident mineralization of pre-existing compression fossils, whereas the three-dimensionally preserved gut-caecal system of Burgessia is interpreted as an aluminosilicate replacement of a pre-existing carbonate phase. The case for late diagenetic emplacement of aluminosilicate minerals is supported by the extensive aluminosilicification of trilobite shell and (originally) calcareous veinlets in the Burgess Shale, as well as documentation of other secondarily aluminosilicified compression fossils. By distinguishing late diagenetic alteration from the early diagenetic processes responsible for exceptional preservation, it is possible to reconcile the range of preservational modes currently expressed in the Burgess Shale.

All possible combinations of six tiering positions in relation to the substratum/water interface, six motility levels and six feeding strategies define a complete theoretical ecospace of 216 potential modes of life for marine animals. The number of modes of life actually utilized specifies realized ecospace. Owing to constraints of effectiveness and efficiency the modern marine fauna utilizes only about half the potential number of modes of life, two-thirds of which (62 of 92) are utilized by animals with readily preserved, mineralized hard parts. Realized ecospace has increased markedly since the early evolution of animal ecosystems. The Ediacaran fauna utilized at most 12 modes of life, with just two practised by skeletal organisms. A total of 30 modes of life are recorded in the Early and Middle Cambrian, 19 of which were utilized by skeletal organisms. The other 11 are documented from soft-bodied animals preserved in the Chengjiang and Burgess Shale Konservat-Lagerstatten. The number of modes of life utilized by skeletal organisms increased by more than 50 per cent during the Ordovician radiation to a Late Ordovician total of 30. Between the Late Ordovician and the Recent the number of utilized modes of life has doubled again. The autecological and taxonomic diversity histories of the marine metazoa appear to be broadly parallel, and future studies of theoretical ecospace utilization should provide more detailed tests of pattern and process in the ecological history of the metazoa.

A new bivalved arthropod is described from the Lower Cambrian (?Upper Atdabanian) Buen Formation of North Greenland. Pauloterminus spinodorsalis gen. et sp. nov. possesses a bivalved carapace that covers the head, which has a single pair of antennae, and anteriormost thorax. No mouthparts are visible. The five-segmented abdomen was limbless and terminated in a telson plus a pair of large, lobate uropods. A suite of at least six biramous thoracic limbs are present: the short endopods are made up of small, serial podomeres, while the exopods are lobate and may have functioned as gills as well as in swimming. Partially infilled guts are occasionally visible, suggesting that this animal may have been a sediment feeder. It is compared to other Cambrian bivalved arthropods, especially the waptiids Chuandianella ovata from the Lower Cambrian Chengjiang fauna (China) and Waptia fieldensis from the Middle Cambrian Burgess Shale (British Columbia). Of these three animals, the Greenland and Chinese taxa appear to be the most closely related. P. spinodorsalis possesses many typical arthropod features, but it also demonstrates more primitive characters that are more reminiscent of the lobopodians.

A problematic organism, Bowengriphus perphlegis gen. et sp. nov., is described based on two specimens from the Late Permian Rangal Coal Measures of eastern central Queensland. It displays a double-looped feeding apparatus bearing small conical elements, considered homologous with that of the supposed lophophorate Odontogriphus omalus Conway Morris, 1976, from the Middle Cambrian Burgess Shale of British Columbia, Canada. Odontogriphids are thus interpreted as a group that survived through the Palaeozoic and made the transition from marine to freshwater environments. Recent proposals that odontogriphids are stem-group chordates are not well established.

More than 300 specimens of the previously rare arthropod Alalcomenaeus cambricus Simonetta have been collected from a new Burgess Shale locality in the Glossopleura Zone on Mount Stephen, British Columbia. This new material provides much more complete information on its morphology. The cephalon was covered by a shield. A pair of pedunculate eyes and three median eyes were followed by a large anterior appendage, the ‘great appendage’, bearing three long flagella. The two posterior head appendages, like those of the trunk, were biramous. They consisted of a segmented, inner branch, and a flap-like outer branch, fringed with long filaments. The trunk consisted of 11 somites, each protected by a tergite and bearing a pair of biramous limbs. The telson was paddle-like and fringed posteriorly with wide flat spines. Alalcomenaeus was probably a predator, moving mainly by swimming. It is now known to be one of the more abundant, widely distributed and longest ranging of Burgess Shale arthropod genera. Its affinities lie with the Arachnomorpha.

The Sirius Passet fauna from the Lower Cambrian of North Greenland yields a diverse suite of poorly sclerotized arthropods. A new nektaspid from the fauna described here differs from typical naraoiids in having six thoracic segments and in being isopygous. Soft parts are poorly known, but posterior limbs are preserved in some specimens. Although specimens are invariably highly flattened, details of the axial articulation suggest that the animal could enrol. The widths of the articulating half rings can be used to reconstruct both the degree of rotation at each articulation during enrolment and the height at which each point along the half ring is above the fulcrum, and this in turn may be used to make a full three-dimensional reconstruction of the exoskeleton. This method of retrodeformation is in principle applicable to the reconstruction of other taxa, especially trilobites, found in shales.

The stem-group priapulid Ottoia Walcott, 1911, is the most abundant worm in the mid-Cambrian Burgess Shale, but has not been unambiguously demonstrated elsewhere. High-resolution electron and optical microscopy of macroscopic Burgess Shale specimens reveals the detailed anatomy of its robust hooks, spines and pharyngeal teeth, establishing the presence of two species: Ottoia prolifica Walcott, 1911, and Ottoia tricuspida sp. nov. Direct comparison of these sclerotized elements with a suite of shale-hosted mid-to-late Cambrian microfossils extends the range of ottoiid priapulids throughout the middle to upper Cambrian strata of the Western Canada Sedimentary Basin. Ottoiid priapulids represented an important component of Cambrian ecosystems: they occur in a range of lithologies and thrived in shallow water as well as in the deep-water setting of the Burgess Shale. A wider survey of Burgess Shale macrofossils reveals specific characters that diagnose priapulid sclerites more generally, establishing the affinity of a wide range of Small Carbonaceous Fossils and demonstrating the prominent role of priapulids in Cambrian seas.