Computational biomechanical analyses demonstrate similar shell-crushing abilities in modern and ancient arthropods
The biology of the American horseshoe crab, Limulus polyphemus, is well documented-including its dietary habits, particularly the ability to crush shell with gnathobasic walking appendages-but virtually nothing is known about the feeding biomechanics of this iconic arthropod. Limulus polyphemus is also considered the archetypal functional analogue of various extinct groups with serial gnathobasic appendages, including eurypterids, trilobites and other early arthropods, especially Sidneyia inexpectans from the mid-Cambrian (508 Myr) Burgess Shale of Canada. Exceptionally preserved specimens of S. inexpectans show evidence suggestive of durophagous (shell-crushing) tendencies-including thick gnathobasic spine cuticle and shelly gut contents-but the masticatory capabilities of this fossil species have yet to be compared with modern durophagous arthropods. Here, we use advanced computational techniques, specifically a unique application of 3D finite-element analysis (FEA), to model the feeding mechanics of L. polyphemus and S. inexpectans: the first such analyses of a modern horseshoe crab and a fossil arthropod. Results show that mechanical performance of the feeding appendages in both arthropods is remarkably similar, suggesting that S. inexpectans had similar shell-crushing capabilities to L. polyphemus This biomechanical solution to processing shelly food therefore has a history extending over 500 Myr, arising soon after the first shell-bearing animals. Arrival of durophagous predators during the early phase of animal evolution undoubtedly fuelled the Cambrian 'arms race' that involved a rapid increase in diversity, disparity and abundance of biomineralized prey species.
Cambrian of the Stansbury and Arrowie Basins
Cambrian sedimentation in southeastern South Australia was the culmination of deposition in the Adelaide Fold Belt, a meridional belt of deep subsidence and correspondingly thick sediment accumulation initiated in the early Cryogenian (Fig. 7). Sedimentation took place on a basement of Archaean to early Mesoproterozoic Gawler Craton to the west, and Palaeo- to Mesoproterozoic Curnamona Province to the east (Gravestock & Gatehouse 1995, Belperio et al. 1998, Preiss 2000, 2002).
Complex axial growth patterns in an early Cambrian trilobite from South Australia
The exceptional fossil record of trilobites provides our best window on developmental processes in early euarthropods, but data on growth dynamics are limited. Here, we analyse post-embryonic axial growth in the Cambrian trilobite Estaingia bilobata from the Emu Bay Shale, South Australia. Using threshold models, we show that abrupt changes in growth trajectories of different body sections occurred in two phases, closely associated with the anamorphic/epimorphic and meraspid/holaspid transitions. These changes are similar to the progression to sexual maturity seen in certain extant euarthropods and suggest that the onset of maturity coincided with the commencement of the holaspid period. We also conduct hypothesis testing to reveal the likely controls of observed axial growth gradients and suggest that size may better explain growth patterns than moult stage. The two phases of allometric change in E. bilobata, as well as probable differing growth regulation in the earliest post-embryonic stages, suggest that observed body segmentation patterns in this trilobite were the result of a complex series of changing growth controls that characterized different ontogenetic intervals. This indicates that trilobite development is more complex than previously thought, even in early members of the clade.
Disparate compound eyes of Cambrian radiodonts reveal their developmental growth mode and diverse visual ecology
Radiodonts are nektonic stem-group euarthropods that played various trophic roles in Paleozoic marine ecosystems, but information on their vision is limited. Optical details exist only in one species from the Cambrian Emu Bay Shale of Australia, here assigned to Anomalocaris aff. canadensis We identify another type of radiodont compound eye from this deposit, belonging to 'Anomalocaris' briggsi. This ≤4-cm sessile eye has >13,000 lenses and a dorsally oriented acute zone. In both taxa, lenses were added marginally and increased in size and number throughout development, as in many crown-group euarthropods. Both species' eyes conform to their inferred lifestyles: The macrophagous predator A. aff. canadensis has acute stalked eyes (>24,000 lenses each) adapted for hunting in well-lit waters, whereas the suspension-feeding 'A.' briggsi could detect plankton in dim down-welling light. Radiodont eyes further demonstrate the group's anatomical and ecological diversity and reinforce the crucial role of vision in early animal ecosystems.
The Early Cambrian trilobite family Emuellidae Pocock, 1970: Systematic position and revision of Australian species
The family Emuellidae Pocock, 1970 was established for Emuella Pocock, 1970 and Balcoracania Pocock, 1970 from the Lower Cambrian of South Australia. Based on their peculiar trunk tagmosis, emuellids have been interpreted as the sister group of all other trilobites with dorsal facial sutures, and classified as high as the ordinal level. Cladistic analysis with a range of exemplar taxa of the Olenellina and Redlichiina instead resolves the emuellids within the Redlichiina, with tagmosis into a prothorax and opisthothorax ("telosoma") nonhomologous in olenellines and emuellids. A taxonomic revision of Australian species identifies Balcoracania flindersi as a junior subjective synonym of B. dailyi, whereas the two named species of Emuella are considered to be distinct. Balcoracania dailyi possesses up to 103 thoracic segments, the maximum number recorded in any trilobite
Shelly fossils from the lower Cambrian White Point Conglomerate, Kangaroo Island, South Australia
The lower Cambrian (Series 2) White Point Conglomerate (WPC) on Kangaroo Island, South Australia contains exotic clasts representing a diverse array of lithologies, including metamorphics, chert, sandstone, and abundant carbonates, notably archaeocyath-rich bioclastic limestone. Acetic acid digestion of the WPC bioclastic limestone clasts reveals a diverse shelly fauna. This assemblage includes abundant organophosphatic brachiopods such as Cordatia erinae Brock and Claybourn gen. et sp. nov., Curdus pararaensis, Eodicellomus elkaniformiis, Eohadrotreta sp. cf. E. zhenbaensis, Eoobolus sp., Kyrshabaktella davidii, and Schizopholis yorkensis. Additional shelly taxa include the solenopleurid trilobite Trachoparia? sp., the tommotiids Dailyatia odyssei, Dailyatia decobruta Betts sp. nov., Kelanella sp., and Lapworthella fasciculata, spines of the bradoriid arthropod Mongolitubulus squamifer, and several problematica, such as Stoibostrombus crenulatus and a variety of tubular forms. The upper age limit for the WPC is constrained by biostratigraphic data from the overlying Marsden Sandstone and Emu Bay Shale, which are no younger than the Pararaia janeae Trilobite Zone (Cambrian Series 2, Stage 4). The shelly fossil assemblage from the WPC limestone clasts indicates an upper Dailyatia odyssei Zone (= Pararaia tatei to lower P. janeae trilobite zones), equivalent to the Atdabanian-early Botoman of the Siberian scheme. This contrasts with the previously suggested late Botoman age for the limestone clasts, based on the diverse archaeocyath assemblage. The minor age difference between the WPC and its fossiliferous limestone clasts suggests relatively rapid reworking of biohermal buildups during tectonically-active phases of deposition in the Stansbury Basin.
Revision of 'Discomesites' and 'Estaingia' (Trilobita) from the Lower Cambrian Cymbric Vale Formation, Western New South Wales: Taxonomic, Biostratigraphic and Biogeographic Implications
The taxonomy of Discomesites and Estaingia from the Lower Cambrian Cymbric Vale Formation of western New South Wales is revised. Discomesites is regarded as a valid subgenus of Pagetides. Pagetides (Discomesites) fragum is considered a senior subjective synonym of P. (D.) lunatulus. Pagetides (Discomesites) spinosus from the Shackleton Limestone in the Holyoake Range, Transantarctic Mountains, is considered to be a junior subjective synonym of P. (D.) fragum. Estaingia cerastes from the Cymbric Vale Formation is considered to be synonymous with Hsuaspis cf. H. bilobata from the Shackleton Limestone. The Cymbric Vale Formation trilobite fauna is of late Early Cambrian (late Botoman) age, equivalent to the Pararaia janeae Zone of South Australia, based on correlation of the Syringocnema favus archaeocyathan fauna. Absolute ages of recently dated tuffs from the Cymbric Vale and Billy Creek Formations are questioned, based on new information regarding the stratigraphic position of the Cymbric Vale Formation tuff in relation to archaeocyathan and trilobite biostratigraphy. The co-occurrence of Pagetides (Discomesites) fragum and Estaingia cerastes in the upper part of the Cymbric Vale Formation and in the Shackleton Limestone represents the first species-level correlation between the Lower Cambrian of Australia and Antarctica using trilobites. The distribution of these trilobite species, in association with the Syringocnema favus archaeocyathan fauna, provides supporting evidence that Australia and Antarctica were connected by a continuous carbonate-detrital shelf during the late Early Cambrian (mid-late Botoman),allowing faunal exchange between these regions.
Calcitic shells in the aragonite sea of the earliest Cambrian
The initial acquisition of calcium carbonate polymorphs (aragonite and calcite) at the onset of skeletal biomineralization by disparate metazoans across the Ediacaran-Cambrian transition is thought to be directly influenced by Earth's seawater chemistry. It has been presumed that animal clades that first acquired mineralized skeletons during the so-called "aragonite sea" of the latest Ediacaran and earliest Cambrian (Terreneuvian) possessed aragonite or high-Mg calcite skeletons, while clades that arose in the subsequent "calcite sea" of Cambrian Series 2 acquired low-Mg calcite skeletons. Here, contrary to previous expectations, we document shells of one of the earliest helcionelloid molluscs from the basal Cambrian of southwestern Mongolia that are composed entirely of low-Mg calcite and formed during the Terreneuvian aragonite sea. The extraordinarily well-preserved Postacanthella shells have a simple prismatic microstructure identical to that of their modern low-Mg calcite molluscan relatives. High-resolution scanning electron microscope observations show that calcitic crystallites were originally encased within an intra- and interprismatic organic matrix scaffold preserved by aggregates of apatite during early diagenesis. This indicates that not all molluscan taxa during the early Cambrian produced aragonitic shells, weakening the direct link between carbonate skeletal mineralogy and ambient seawater chemistry during the early evolution of the phylum. Rather, our study suggests that skeletal mineralogy in Postacanthella was biologically controlled, possibly exerted by the associated prismatic organic matrix. The presence of calcite or aragonite mineralogy in different early Cambrian molluscan taxa indicates that the construction of calcium carbonate polymorphs at the time when skeletons first emerged may have been species dependent.
The Emu Bay Shale Konservat-Lagersattte: a view of Cambrian life from East Gondwana
Recent fossil discoveries from the lower Cambrian Emu Bay Shale (EBS) on Kangaroo Island, South Australia, have provided critical insights into the tempo of the Cambrian explosion of animals, such as the origin and seemingly rapid evolution of arthropod compound eyes, as well as extending the geographical ranges of several groups to the East Gondwanan margin, supporting close faunal affinities with South China. The EBS also holds great potential for broadening knowledge on taphonomic pathways involved in the exceptional preservation of fossils in Cambrian Konservat-Lagerstätten. EBS fossils display a range of taphonomic modes for a variety of soft tissues, especially phosphatization and pyritization, in some cases recording a level of anatomical detail that is absent from most Cambrian Konservat-Lagerstätten.
A review of the Cambrian biostratigraphy of South Australia
Cambrian rocks in South Australia occur in the Stansbury, Arrowie, eastern Officer and Warburton Basins. The succession in the Stansbury and Arrowie Basins can be divided into three sequence sets (supersequences), C1, C2 and C3. Sequence set C1 can be divided into five third-order sequences: C1.0, C1.1A, C1.1B, C1.2 and C1.3. Trilobites from the Stansbury and Arrowie Basins are restricted largely to the lower part of the succession. Four trilobite zones are recognized: 'Abadiella huoi' (latest Atdabanian–earliest Botoman), 'Pararaia tatei', 'Pararaia bunyerooensis' and 'Pararaia janeae' Zones (all Botoman). Trilobites higher in the succession are known from only a few horizons and in part correlate with the upper Lower Cambrian Lungwangmiaoan Stage of China, equivalent to the top Toyonian. 'Pagetia sp.' has been reported in the Coobowie Formation of the Stansbury Basin, thus suggesting an early Middle Cambrian age. The Cambrian faunas of the Warburton Basin range in age from early Middle Cambrian (Late Templetonian) to very Late Cambrian, although the richest faunal assemblages are late Middle Cambrian ('Ptychagnostus punctuosus to Goniagnostus nathorsti' Zones). Conodonts, including 'Cordylodus proavus', occur in a Datsonian fauna. The Arrowie Basin contains the most complete and best studied archaeocyath succession in the Australia–Antarctica region. The 'Warriootacyathus wilkawillensis', 'Spirillicyathus tenuis' and 'Jugalicyathus tardus' Zones from the lower Wilkawillina Limestone (Arrowie Basin) and equivalents are correlated with the Atdabanian. Botoman archaeocyathids occur higher in the Wilkawillina Limestone. The youngest (Toyonian) archaeocyath fauna in Australia occurs in the Wirrealpa Limestone (Arrowie Basin). Brachiopods and molluscs of the Arrowie and Stansbury Basins can be divided into four biostratigraphic assemblages. Several informal Early Cambrian SSF biostratigraphic assemblages are recognized. Probable tabulate-like corals occur in the Botoman Moorowie Formation. Seven informal acritarch assemblages occur in the Early Cambrian of the Stansbury and Arrowie Basins. Trace fossils may mark the Precambrian–Cambrian boundary. Only two of several tuffaceous horizons from the Stansbury and Arrowie Basins have been dated (i) a date of 522.0 ± 2.1 Ma from the Heatherdale Shale of the Stansbury Basin, about 400m above latest Atdabanian archaeocyathids and (ii) a date of 522.0 ± 1.8 Ma from the lower part of the Billy Creek Formation in the Arrowie Basin. Neither date is regarded as reliable.