Gill function in an early arthropod and the widespread adoption of the countercurrent exchange mechanism.

Rising but fluctuating oxygen levels in the Early Palaeozoic provide an environmental context for the radiation of early metazoans, but little is known about how mechanistically early animals satisfied their oxygen requirements. Here we propose that the countercurrent gaseous exchange, a highly efficient respiratory mechanism, was effective in the gills of the Late Ordovician trilobite Triarthrus eatoni. In order to test this, we use computational fluid dynamics to simulate water flow around its gills and show that water velocity decreased distinctly in front of and between the swollen ends, which first encountered the oxygen-charged water, and slowed continuously at the mid-central region, forming a buffer zone with a slight increase of the water volume. In T. eatoni respiratory surface area was maximized by extending filament height and gill shaft length. In comparison with the oxygen capacity of modern fish and crustaceans, a relatively low weight specific area in T. eatoni may indicate its low oxygen uptake, possibly related to a less active life mode. Exceptionally preserved respiratory structures in the Cambrian deuterostome Haikouella are also consistent with a model of countercurrent gaseous exchange, exemplifying the wide adoption of this strategy among early animals.

The trilobite upper limb branch is a well-developed gill.

Whether the upper limb branch of Paleozoic "biramous" arthropods, including trilobites, served a respiratory function has been much debated. Here, new imaging of the trilobite Triarthrus eatoni shows that dumbbell-shaped filaments in the upper limb branch are morphologically comparable with gill structures in crustaceans that aerate the hemolymph. In Olenoides serratus, the upper limb's partial articulation to the body via an extended arthrodial membrane is morphologically comparable to the junction of the respiratory book gill of Limulus and differentiates it from the typically robust exopod junction in Chelicerata or Crustacea. Apparently limited mechanical rotation of the upper branch may have protected the respiratory structures. Partial attachment of the upper branch to the body wall may represent an intermediate state in the evolution of limb branch fusion between dorsal attachment to the body wall, as in Radiodonta, and ventral fusion to the limb base, as in extant Euarthropoda.

Early Cambrian fuxianhuiids from China reveal origin of the gnathobasic protopodite in euarthropods.

Euarthropods owe their evolutionary and ecological success to the morphological plasticity of their appendages. Although this variability is partly expressed in the specialization of the protopodite for a feeding function in the post-deutocerebral limbs, the origin of the former structure among Cambrian representatives remains uncertain. Here, we describe Alacaris mirabilis gen. et sp. nov. from the early Cambrian Xiaoshiba Lagerstätte in China, which reveals the proximal organization of fuxianhuiid appendages in exceptional detail. Proximally, the post-deutocerebral limbs possess an antero-posteriorly compressed protopodite with robust spines. The protopodite is attached to an endopod with more than a dozen podomeres, and an oval flap-shaped exopod. The gnathal edges of the protopodites form an axial food groove along the ventral side of the body, indicating a predatory/scavenging autecology. A cladistic analysis indicates that the fuxianhuiid protopodite represents the phylogenetically earliest occurrence of substantial proximal differentiation within stem-group Euarthropoda illuminating the origin of gnathobasic feeding.

Onychophoran-like musculature in a phosphatized Cambrian lobopodian.

The restricted, exclusively terrestrial distribution of modern Onychophora contrasts strikingly with the rich diversity of onychophoran-like fossils preserved in marine Cambrian Lagerstätten The transition from these early forebears to the modern onychophoran body plan is poorly constrained, in part owing to the absence of fossils preserving details of the soft anatomy. Here, we report muscle tissue in a new early Cambrian (Stage 3) lobopodian, Tritonychus phanerosarkus gen. et sp. nov., preserved in the Orsten fashion by three-dimensional replication in phosphate. This first report of Palaeozoic onychophoran musculature establishes peripheral musculature as a characteristic of the ancestral panarthropod, but documents an unexpected muscular configuration. Phylogenetic analysis reconstructs T. phanerosarkus as one of a few members of the main onychophoran lineage-which was as rare and as cryptic in the Cambrian period as it is today.

A predatory bivalved euarthropod from the Cambrian (Stage 3) Xiaoshiba Lagerstätte, South China.

Bivalved euarthropods represent a conspicuous component of exceptionally-preserved fossil biotas throughout the Lower Palaeozoic. However, most of these taxa are known from isolated valves, and thus there is a limited understanding of their morphological organization and palaeoecology in the context of early animal-dominated communities. The bivalved euarthropod Clypecaris serrata sp. nov., recovered from the Cambrian (Stage 3) Hongjingshao Formation in Kunming, southern China, is characterized by having a robust first pair of raptorial appendages that bear well-developed ventral-facing spines, paired dorsal spines on the trunk, and posteriorly oriented serrations on the anteroventral margins of both valves. The raptorial limbs of C. serrata were adapted for grasping prey employing a descending stroke for transporting it close the mouth, whereas the backwards-facing marginal serrations of the bivalved carapace may have helped to secure the food items during feeding. The new taxon offers novel insights on the morphology of the enigmatic genus Clypecaris, and indicates that the possession of paired dorsal spines is a diagnostic trait of the Family Clypecarididae within upper stem-group Euarthropoda. C. serrata evinces functional adaptations for an active predatory lifestyle within the context of Cambrian bivalved euarthropods, and contributes towards the better understanding of feeding diversity in early ecosystems.

Fuxianhuiid ventral nerve cord and early nervous system evolution in Panarthropoda.

Panarthropods are typified by disparate grades of neurological organization reflecting a complex evolutionary history. The fossil record offers a unique opportunity to reconstruct early character evolution of the nervous system via exceptional preservation in extinct representatives. Here we describe the neurological architecture of the ventral nerve cord (VNC) in the upper-stem group euarthropod Chengjiangocaris kunmingensis from the early Cambrian Xiaoshiba Lagerstätte (South China). The VNC of C. kunmingensis comprises a homonymous series of condensed ganglia that extend throughout the body, each associated with a pair of biramous limbs. Submillimetric preservation reveals numerous segmental and intersegmental nerve roots emerging from both sides of the VNC, which correspond topologically to the peripheral nerves of extant Priapulida and Onychophora. The fuxianhuiid VNC indicates that ancestral neurological features of Ecdysozoa persisted into derived members of stem-group Euarthropoda but were later lost in crown-group representatives. These findings illuminate the VNC ground pattern in Panarthropoda and suggest the independent secondary loss of cycloneuralian-like neurological characters in Tardigrada and Euarthropoda.

A superarmored lobopodian from the Cambrian of China and early disparity in the evolution of Onychophora.

We describe Collinsium ciliosum from the early Cambrian Xiaoshiba Lagerstätte in South China, an armored lobopodian with a remarkable degree of limb differentiation including a pair of antenna-like appendages, six pairs of elongate setiferous limbs for suspension feeding, and nine pairs of clawed annulated legs with an anchoring function. Collinsium belongs to a highly derived clade of lobopodians within stem group Onychophora, distinguished by a substantial dorsal armature of supernumerary and biomineralized spines (Family Luolishaniidae). As demonstrated here, luolishaniids display the highest degree of limb specialization among Paleozoic lobopodians, constitute more than one-third of the overall morphological disparity of stem group Onychophora, and are substantially more disparate than crown group representatives. Despite having higher disparity and appendage complexity than other lobopodians and extant velvet worms, the specialized mode of life embodied by luolishaniids became extinct during the Early Paleozoic. Collinsium and other superarmored lobopodians exploited a unique paleoecological niche during the Cambrian explosion.

Development and trunk segmentation of early instars of a ptychopariid trilobite from Cambrian Stage 5 of China.

Many three-dimensionally preserved exoskeletons found from the middle Cambrian (Stage 5) Gaotai Formation in Guizhou, southern China, have been assigned to the ptychopariid trilobite Gunnia sp. They represent mainly a series of early instars, exhibiting some delicate structures and morphological variation associated with their trunk segmentation and early development. Morphometric and statistical analyses indicate that the transverse joint appears to occur with the full growth of the third axial ring of the protopygidium, which increases in size much more rapidly than its corresponding protocephalon with growth. The 'one by one' sequential release of thoracic segments from a transitory pygidium does not progress exactly in accordance with the development of the pygidial axis, whose axial rings increase at a relatively faster rate, and an 'immature ring' always appears initially at the rear end of the axis. These new data set up a testable model for revealing trilobite segmentation and provide fresh insights into the development, evolution and taphonomic surroundings associated with the Cambrian trilobites.

Articulated Wiwaxia from the Cambrian Stage 3 Xiaoshiba lagerstätte.

Wiwaxia is a bizarre metazoan that has been interpreted as a primitive mollusc and as a polychaete annelid worm. Extensive material from the Burgess Shale provides a detailed picture of its morphology and ontogeny, but the fossil record outside this lagerstätte is scarce, and complete wiwaxiids are particularly rare. Here we report small articulated specimens of Wiwaxia foliosa sp. nov. from the Xiaoshiba fauna (Cambrian Stage 3, Hongjingshao Formation, Kunming, south China). Although spines are absent, the fossils' sclerites - like those of W. corrugata - are symmetrically arranged in five distinct zones. They form rows across the body, and were individually added and shed throughout growth to retain an approximately symmetrical body shape. Their development pattern suggests a molluscan affinity. The basic body plan of wiwaxiids is fundamentally conserved across two continents through Cambrian Stages 3-5 - revealing morphological stasis in the wake of the Cambrian explosion.