Oxygen as a Driver of Early Arthropod Micro-Benthos Evolution

BACKGROUND: We examine the physiological and lifestyle adaptations which facilitated the emergence of ostracods as the numerically dominant Phanerozoic bivalve arthropod micro-benthos. METHODOLOGY/PRINCIPAL FINDINGS: The PO(2) of modern normoxic seawater is 21 kPa (air-equilibrated water), a level that would cause cellular damage if found in the tissues of ostracods and much other marine fauna. The PO(2) of most aquatic breathers at the cellular level is much lower, between 1 and 3 kPa. Ostracods avoid oxygen toxicity by migrating to waters which are hypoxic, or by developing metabolisms which generate high consumption of O(2). Interrogation of the Cambrian record of bivalve arthropod micro-benthos suggests a strong control on ecosystem evolution exerted by changing seawater O(2) levels. The PO(2) of air-equilibrated Cambrian-seawater is predicted to have varied between 10 and 30 kPa. Three groups of marine shelf-dwelling bivalve arthropods adopted different responses to Cambrian seawater O(2). Bradoriida evolved cardiovascular systems that favoured colonization of oxygenated marine waters. Their biodiversity declined during intervals associated with black shale deposition and marine shelf anoxia and their diversity may also have been curtailed by elevated late Cambrian (Furongian) oxygen-levels that increased the PO(2) gradient between seawater and bradoriid tissues. Phosphatocopida responded to Cambrian anoxia differently, reaching their peak during widespread seabed dysoxia of the SPICE event. They lacked a cardiovascular system and appear to have been adapted to seawater hypoxia. As latest Cambrian marine shelf waters became well oxygenated, phosphatocopids went extinct. Changing seawater oxygen-levels and the demise of much of the seabed bradoriid micro-benthos favoured a third group of arthropod micro-benthos, the ostracods. These animals adopted lifestyles that made them tolerant of changes in seawater O(2). Ostracods became the numerically dominant arthropod micro-benthos of the Phanerozoic. CONCLUSIONS/SIGNIFICANCE: Our work has implications from an evolutionary context for understanding how oxygen-level in marine ecosystems drives behaviour

'Microdictyon' plates from the lower Cambrian Ajax Limestone of South Australia: Implications for species taxonomy and diversity

A small assemblage of isolated 'Microdictyon' plates is described from the lower Cambrian Ajax Limestone, Mt Scott Range (Flinders Ranges), South Australia. 'Microdictyon' plates are primarily known from single, isolated, perforated phosphatic sclerites; only one species ('M. sinicum') from the Chengjiang Lagerstätte is known from soft-bodied preservation of the complete organism. The isolated plates from South Australia display a wide range of morphologies potentially reflecting: 1, considerable diversification within the group at this time; 2, extensive intraspecific morphological variation; 3, different plate morphotypes along the trunk of the animal; or 4, significant ontogenetic variation in successive growth stages. The South Australian specimens are similar to several individual sclerites described from other Cambrian palaeocontinents, but appear closest to faunas described from South China. Problems associated with the taxonomy of isolated 'Microdictyon' plates are discussed, including the lack of knowledge associated with intraspecific and/or ontogenetic variability and how individual plate morphology may relate to size or relative position along the length of the complete organism

The oldest bivalved arthropods from the early Cambrian of East Gondwana: Systematics, biostratigraphy and biogeography

The oldest bradoriid fauna from Australia, occurring in the lower Cambrian Ajax and Wirrapowie limestones of the Flinders Ranges, South Australia consists of eleven taxa, including one new genus and species, 'Quadricona madonnae' gen. et sp. nov. and two new species, 'Liangshanella circumbolina' sp. nov. and 'Zepaera jagoi' sp. nov. In the Ajax Limestone, 'Liangshanella circumbolina' sp. nov. occurs c. 20 m below the FAD of the zonal trilobite 'Abadiella huoi'. This pre-trilobitic occurrence represents the oldest bivalved arthropod hitherto known from East Gondwana and suggests a lower Cambrian (Series 2, Stage 3) age for the assemblage. The recognition of distinct bradoriid assemblages associated with the 'Abadiella huoi' (Atdabanian), 'Pararaia tatei', 'P. bunyerooensis' and 'P. janeae' (all Botoman) trilobite biozones in South Australia indicates great potential for future regional biostratigraphic correlation. Quantitative biogeographic analysis including new taxonomic data from the lower Cambrian of South Australia, highlights the strong endemism displayed by early Cambrian bradoriid communities and strengthens the close faunal affinities with South China and Antarctica

Scleritome construction, biofacies, biostratigraphy and systematics of the tommotiid 'Eccentrotheca helenia' sp. nov. from the Early Cambrian of South Australia

Large collections of 'Eccentrotheca helenia' sp. nov. from the lower Cambrian Wilkawillina and Ajax limestones in the Arrowie Basin, South Australia, contain abundant low, cap-shaped and high, laterally compressed isolated sclerites in addition to partially articulated tubular specimens. The scleritome of 'Eccentrotheca helenia' sp. nov. is fully described for the first time and shown to be formed by ontogenetic fusion of sclerites into successively stacked sclerite rings, forming a larger, tubular structure. The apical termination of the tube is highly variable, but is primarily constructed by low, cap-shaped sclerites and characterised by a central aperture of variable inclination. The adapical portion of the tube is predominantly constructed by high, laterally compressed sclerites, but individual sclerite rings can contain both cap-shaped and laterally compressed sclerites along with sclerites of intermediate morphology. The apical aperture presumably housed organic structures for attachment to a hard substrate, but the scleritome also occasionally preserves small lateral perforations between fused sclerites, which may have served to stabilise the scleritome by providing additional points of anchorage. In the Arrowie Basin, 'E. helenia' is found in association with archaeocyath-microbial-spongiomorph-dominated bioherms and most likely inhabited pendant or cryptic habitats within these bioherms. 'Eccentrotheca'-like sclerites form an integral part of the scleritomes of many tommotiids which may confuse taxonomic analysis. Sclerites previously assigned to 'E.'guano, consistently occur together with sclerites of 'Kulparina rostrata' in stratigraphic intervals consistently older than strata hosting 'E. helenia'. Rare fused specimens indicate that the sclerites of 'K. rostrata' and 'E.'guano belong to the same scleritome

An encrusting kleptoparasite-host interaction from the early Cambrian

Parasite–host systems are pervasive in nature but are extremely difficult to convincingly identify in the fossil record. Here we report quantitative evidence of parasitism in the form of a unique, enduring life association between tube-dwelling organisms encrusted to densely clustered shells of a monospecific organophosphatic brachiopod assemblage from the lower Cambrian (Stage 4) of South China. Brachiopods with encrusting tubes have decreased biomass (indicating reduced fitness) compared to individuals without tubes. The encrusting tubes orient tightly in vectors matching the laminar feeding currents of the host, suggesting kleptoparasitism. With no convincing parasite–host interactions known from the Ediacaran, this widespread sessile association reveals intimate parasite–host animal systems arose in early Cambrian benthic communities and their emergence may have played a key role in driving the evolutionary and ecological innovations associated with the Cambrian radiation