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Environmental change and biotic interactions both govern the evolution of the biosphere, but the relative importance of these drivers over geological time remains largely unknown. Previous work suggests that, unlike environmental parameters, diversity dynamics differ profoundly between the Palaeozoic and post-Palaeozoic eras. Here we use the fossil record to test the hypothesis that the influence of ocean chemistry and climate on the ecological success of marine calcifiers decreased throughout the Phanerozoic eon. Marine calcifiers build skeletons of calcite or aragonite, and the precipitation of these calcium carbonate polymorphs is governed by the magnesium-to-calcium ratio and temperature in abiotic systems. We developed an environmental forcing model based on secular changes of ocean chemistry and temperature and assessed how well the model predicts the proliferation of skeletal taxa with respect to calcium carbonate polymorphs. Abiotic forcing governs the ecological success of aragonitic calcifiers from the Ordovician to the Middle Jurassic, but not thereafter. This regime shift coincides with the proliferation of calcareous plankton in the mid-Mesozoic. The deposition of biomineralizing plankton on the ocean floor buffers CO2 excursions and stabilizes Earth’s biochemical cycle, and thus mitigates the evolutionary impact of environmental change on the marine biota. Controls on the ecological success of marine calcifiers changed from abiotic to biotic in the mid-Jurassic, according an environmental forcing model compared with skeletal taxa.

The extraordinary window of phosphatized and phosphatic small shelly fossils (SSF) during the early and middle Cambrian is an important testament to the radiation of biomineralizing metazoans. While SSF are well known from most Cambrian palaeocontinents during this time interval, western Laurentia has relatively few SSF faunas. Here we describe a diverse SSF fauna from the early Cambrian (Stages 3–4) Mural Formation at three localities in Alberta and British Columbia, Canada, complemented by carbon isotope measurements to aid in a potential future bio-chemostratigraphic framework. The fauna expands the recorded SSF assemblage diversity in western Laurentia and includes several brachiopods, four bradoriids, three chancelloriids, two hyoliths, a tommotiid and a helcionellid mollusc as well as echinoderm ossicles and specimens of Microdictyon, Volborthella and Hyolithellus. New taxa include the tommotiid genus Canadiella gen. nov., the new bradoriid species Hipponicharion perforata sp. nov. and Pseudobeyrichona taurata sp. nov. Compared with contemporaneous faunas from western Laurentia, the fauna is relatively diverse, particularly in taxa with originally phosphatic shells, which appear to be associated with archaeocyathid build-ups. This suggests that the generally low faunal diversity in western Laurentia may be at least partly a consequence of poor sampling of suitable archaeocyathan reef environments. In addition, the tommotiid Canadiella filigrana appears to be of biostratigraphical significance in Cambrian Stage 3 strata of western Laurentia, and the unexpected high diversity of bradoriid arthropods in the fauna also suggests that this group may prove useful for biostratigraphical resolution in the region.

Palaeozoic hypercalcified sponges were ubiquitous Ordovician—Devonian reef builders but, despite their rich fossil record, their original skeletal mineralogy and microstructure remain poorly understood. This study provides the first application of electron backscatter diffraction (EBSD) to analyse skeletal structure of Silurian and Devonian stromatoporoids. The two Silurian and two Devonian stromatoporoid taxa selected are typical of stromatoporoids in showing poor preservation. A reference sample of an exceptionally well-preserved hypercalcified chaetetid sponge from the Carboniferous Buckhorn Asphalt Quarry (a fossil lagerstatte renowned for its preservation of skeletal microstructures) contains evidence that its skeleton comprised distinct bundles of single-crystal fibres, similar to modern hypercalcifying sponges. Similar bundles of crystal fibres are proposed here as the original microstructure of stromatoporoids, and acted as precursors to the coarse fibrous calcitic overprinting recrystallisation that is orientated normal to the growth layers, seen in all stromatoporoids viewed in cross-polarised light. The studied stromatoporoids show pronounced microporosity and micro-dolomite inclusions which are circumstantial evidence of an original composition of high-Mg calcite (HMC). We propose that the evidence of fibrous structures might be linked to inclusions of hydrated amorphous calcium carbonate (ACC·H2O) in the skeleton at the time of early diagenesis, as occurs in modern calcified sponges. The possible HMC skeletal composition of Palaeozoic stromatoporoids supports earlier views that the mineral composition of hypercalcifying reef builders is linked to Phanerozoic oscillations in the ratio of Mg:Ca, expressed as aragonite-calcite seas; stromatoporoids thrived in times of calcite-seas.

Animals originated in the Neoproterozoic and ‘exploded’ into the fossil record in the Cambrian. The Cambrian also represents a high point in the animal fossil record for the preservation of soft tissues that are normally degraded. Specifically, fossils from Burgess Shale-type (BST) preservational windows give paleontologists an unparalleled view into early animal evolution. Why this time interval hosts such exceptional preservation, and why this preservational window declines in the early Paleozoic, have been long-standing questions. Anoxic conditions have been hypothesized to play a role in BST preservation, but recent geochemical investigations of these deposits have reached contradictory results with respect to the redox state of overlying bottom waters. Here, we report a multi-proxy geochemical study of the Lower Cambrian Mural Formation, Alberta, Canada. At the type section, the Mural Formation preserves rare recalcitrant organic tissues in shales that were deposited near storm wave base (a Tier 3 deposit; the worst level of soft-tissue preservation). The geochemical signature of this section shows little to no evidence of anoxic conditions, in contrast with published multi-proxy studies of more celebrated Tier 1 and 2 deposits. These data help confirm that ‘decay-limited’ BST biotas were deposited in more oxygenated conditions, and support a role for anoxic conditions in BST preservation. Finally, we discuss the role of iron reduction in BST preservation, including the formation of iron-rich clays and inducement of sealing seafloor carbonate cements. As oceans and sediment columns became more oxygenated and more sulfidic through the early Paleozoic, these geochemical changes may have helped close the BST taphonomic window.

Oscillations between the dominance of aragonite and calcite in abiotic marine CaCO3 precipitates throughout Earth history are closely coupled with the evolution of Earth’s seawater composition and represent the environmental context in which organisms evolved their ability to biomineralize. The most important factor controlling these Phanerozoic oscillations in CaCO3 polymorph composition is the ratio of Mg:Ca in seawater, which is thought to separate aragonite and calcite precipitation along a distinct temperature-controlled threshold. A sharp threshold, however, is contradicted by overlapping aragonite and calcite precipitation fields at a range of experimental conditions. Here we present experimental data that enable us to quantify the proportions of CaCO3 polymorphs as a function of Mg:Ca ratio and temperature. This allows us to convert published Mg:Ca ratio proxy data and models of the Phanerozoic Mg:Ca ratio into proportions of abiotic CaCO3 polymorphs at a given temperature, and thus provides a temperature-corrected view of aragonite-calcite sea conditions. In this revised view, abiotic calcite precipitation was inhibited during aragonite sea intervals at temperatures above 20 °C, whereas calcite sea intervals were characterized by the co-precipitation of aragonite and calcite in environments above 20 °C. This continuous prominence of aragonite precipitation in Phanerozoic warm-water environments explains the Phanerozoic increase of aragonite over calcite skeletal composition in calcifying organisms.

Understanding the influence of aragonite/calcite sea conditions on the evolution of biocalcification relies strongly on the correct interpretation of the original composition of calcareous taxa. Aragonite dissolves or inverts into calcite over geological time, and its preservation is currently unknown to predate the Pennsylvanian. Here we present direct evidence for the common occurrence of relic aragonite in Ordovician and Silurian trimerellid brachiopods, thereby extending the known range of aragonite preservation by more than 130 million years. Together with associated hypercalcifying taxa of putatively original aragonite or high-magnesium calcite composition and considerations of the temperature dependence of aragonite and calcite precipitation, our results suggest that the evolution of aragonite biomineralization might have presented an adaptive advantage in shallow marine tropical waters of calcite seas. A targeted search for Paleozoic aragonite should both resolve the original composition of consistently recrystallized taxa and enable the reassessment of the aragonite/calcite sea paradigm in a paleoenvironmental context.

Tommotiids are distinctive components of the early Cambrian small shelly fauna, almost invariably represented by isolated phosphatic sclerites derived from a multielement protective cover (scleritome). The unusual range of tommotiid sclerite morphologies and unknown construction of the scleritome have severely hampered our understanding of their phylogenetic affinities. However, recent description of rare, articulated scleritome material belonging to the tommotiid genera Eccentrotheca and Paterimitra support the hypothesis that some tommotiids fall within the stem group of the lophophorate phyla Phoronida and Brachiopoda and that at least some tommotiid sclerites are homologous precursors of the shells of organophosphatic brachiopods. Here we show that the shell microstructure of Eccentrotheca and Paterimitra share substantial similarities with paterinid brachiopods. While paterinids possess an overall brachiopod morphology, their microstructure appears more similar to Eccentrotheca and Paterimitra than to nonpaterinate lingulids. These findings strongly support the existence of a brachiopod total group that is solidly rooted within tommotiids, and identify the organophosphatic skeletal composition as plesiomorphic with calcareous shells as derived. The microstructural changes of the proposed tommotiid-brachiopod transition probably reflect an adaptation to fluctuating food and phosphorous intake that came with the switch to a sessile life style at the base of the tommotiid clade.

The tommotiid Camenella reticulosa is redescribed based on new collections of well preserved sclerites from the Arrowie Basin (Flinders Ranges), South Australia, revealing new information concerning morphology and microstructure. The acutely pyramidal mitral sclerite is described for the first time and the sellate sclerite is shown to be coiled through up to 1.5 whorls. Based on Camenella, a model is proposed by which tommotiid sclerites are composed of alternating dense phosphatic, and presumably originally organic-rich, laminae. Camenella is morphologically most similar to Lapworthella, Kennardia, and Dailyatia, and these taxa are interpreted to represent a monophyletic clade, here termed the "camenellans", within the Tommotiida. Potential reconstructions of the scleritome of Camenella are discussed and although a tubular scleritome construction was recently demonstrated for the tommotiids Eccentrotheca and Paterimitra, a bilaterally symmetrical scleritome model with the sclerites arranged symmetrically on the dorsal surface of a vagrant animal can not be ruled out.

The Early Cambrian brachiopod, Eoobolus, is one of the first representatives of the superfamily, Linguloidea, the defining characteristics of which include the classical morphology of oval shells and a pedicle that emerges from between the two valves. The material described here from the Mural Formation (Jasper National Park, Canadian Rocky Mountains) provides well-preserved muscle scars and larval shells that allow a discussion of the muscle system and the larval morphology of Eoobolus. The dorsal larval shell exhibits a morphology similar to other Cambrian linguloids, but also to paterinids, Mickwitzia muralensis, and some rhynchonelliforms. This suggests that there was a lesser degree of disparity among brachiopod larvae in the Cambrian than there is today. The muscle system of Eoobolus is similar to other linguloids, but differs from that of Recent lingulids and discinids by having one or two more pairs of oblique muscles. New data on the distribution of features characteristic of the family Eoobolidae question the validity of this family.

Early Cambrian tommotiids are problematic fossil metazoans with external organophosphatic sclerites that have been considered to be basal members of the lophophorate stem group. Tommotiids are almost exclusively known from isolated or rarely fused individual sclerites, which made previous reconstructions of the actual organism highly conjectural. However, the recent discovery of the first articulated specimens of the tommotiid Eccentrotheca revealed a tubular sclerite arrangement (scleritome) that limited the possible life habit to sessile filter feeding and thus further supported a lophophorate affinity. Here, we report the first articulated specimens of a second tommotiid taxon, Paterimitra from the Early Cambrian of the Arrowie Basin, South Australia. Articulated specimens of Paterimitra are composed of two bilaterally symmetrical sclerite types and an unresolved number of small, asymmetrical and irregular crescent-shaped sclerites that attached to the anterior margin of the symmetrical sclerites. Together, the sclerites form an open cone in which the symmetrical sclerites are joined together and form a small posterior opening near the base of the scleritome, while the irregular crescent-shaped sclerites defined a broad anterior opening. The coniform scleritome of Paterimitra is interpreted to have attached to hard substrates via a pedicle that emerged through the small posterior opening (sometimes forming a tube) and was probably a sessile filter feeder. The scleritome of Paterimitra can be derived from the tubular scleritome of Eccentrotheca by modification of basal sclerites and reduction in tube height, and probably represents a more derived member of the brachiopod stem group with the paired symmetrical sclerites possibly homologous to brachiopod valves.

New material of the enigmatic brachiopod Salanygolina obliqua Ushatinskaya from the Early Cambrian of Mongolia shows that it has a colleplax– a triangular plate – in the umbonal perforation, which is enlarged by resorption. This structure is otherwise only known from the equally enigmatic Palaeozoic orders Chileida and Dictyonellida (Rhynchonelliformea, Chileata). The colleplax in Salanygolina is here considered to be homologous with that of the chileates. Salanygolina is also provided with a ridge-like pseudodeltidium, which is another chileate feature. Other characters of Salanygolina, like the radial arrangement of adductor muscle scars and postero-medially placed internal oblique muscles are characteristic of chileates, but also found in the paterinates. In contrast, mixoperipheral dorsal valves with low rudimentary interareas are well known in paterinates, but not yet recorded from chileates. Thus, Salanygolina shows a mosaic combination of morphologic characters, known both from the paterinates and chileates, indicating that it may represent a stem group of the rhynchonelliform chileate brachiopods. The laminar phosphatic secondary shell of Salanygolina is composed of closely packed and nearly identical hexagonal prisms, oriented with their long axis normal to the laminae in a honeycomb pattern. The prism walls appear to have originally been composed of organic membranes and might represent precursors of the organic sheaths of calcite fibers that are typical of calcitic shells with a fibrous microstructure.

Phosphatised limestone-hosted and shale-hosted specimens of the obolellid Mummpikia nuda gen. et comb. nov. and two further unidentified obolellids from the Early Cambrian Mural Formation (Jasper National Park, Canadian Rocky Mountains) provide novel insights into the shell microstructure of obolellids and the nature of their pedicle. The shell was penetrated by abundant canals of a sub-μm diameter and the anterior tip of the delthyrium forms a projection into the body cavity that is penetrated by a thin canal. It is argued that both shell microstructure and posterior margin are linguliformean characters and that obolellids hold a position basal in the rhynchonelliform stem-group.

The linguliform brachiopod Eoobolus from the Early Cambrian Mural Formation (Jasper National Park, Canadian Rocky Mountains) exhibits various calcitic features in its otherwise apatitic shell. It is argued here that the decomposition of the organic matter within the shell led to a microenvironment similar to those resulting in the phosphatization of soft tissues. This diagenetic regime encouraged the initial precipitation of apatite cements followed by calcite cements. By fully coating primary structures early apatite cements separate primary structures from the later precipitation of calcite cement. Round calcareous grains, about 3 µm in size, that occur in the centre of apatite botryoids must therefore represent original components of the shell. The equivalent pits of such calcareous granules are seen in the larval shells of many Palaeozoic linguliform brachiopods. This suggests that mixed organophosphatic-calcareous shells were relatively common at that time but that they have been overlooked owing to the obliteration of original calcareous structures by traditional acid preparation methods for the extraction of phosphatic fossils. The Eoobolus shell structure is intermediate between purely organophosphatic and calcitic shells. Although one such genus is not sufficient to reconstruct the ancestral composition of the brachiopod shell, it provides a means of recognizing other transitional forms that are needed to understand fully the shift in shell mineralogy.

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.

Exceptionally preserved carbonate- and shale-hosted Mickwitzia muralensis from the Lower Cambrian Mural Formation, southern Canadian Rocky Mountains, complement one another to yield an unusually complete account of its ontogeny, ecology and phylogenetic relationships. The shell of M. muralensis is composed of dense phosphatic layers interspersed with porous organic-rich layers. At the insertion of shell-penetrating tubes, shell layers deflect inwards to produce inwardly pointing cones. The tubes are interpreted as having hosted setae that were secreted by outer-epithelial follicles. Follicular setae also occurred at the mantle margin, where they were oriented within the plane of the shell as in modern brachiopods. During ontogeny, the initial setae oriented in the plane of the shell occurred before the first shell-penetrative setae. In the juvenile and early-mature stages of shell secretion, a posterior opening was present between both valves and was used for the protrusion of an attachment structure. In the late-mature shell, this opening became fixed in the ventral valve. Based on the posterior margin and the shell microstructure, a close relationship between Mickwitzia and the paterinids is proposed with differences interpreted as heterochronic. The shell-penetrative setal apparatus of M. muralensis is distinct from that previously described of Micrina, though both types are conceivably homologous to adult and juvenile setae of modern brachiopods.

The morphological disparity of lophotrochozoan phyla makes it difficult to predict the morphology of the last common ancestor. Only fossils of stem groups can help discover the morphological transitions that occurred along the roots of these phyla. Here, we describe a tubular fossil Yuganotheca elegans gen. et sp. nov. from the Cambrian (Stage 3) Chengjiang Lagersta¨tte (Yunnan, China) that exhibits an unusual combination of phoronid, brachiopod and tommotiid (Cambrian problematica) characters, notably a pair of agglutinated valves, enclosing a horseshoe-shaped lophophore, supported by a lower bipartite tubular attachment structure with a long pedicle with coelomic space. The terminal bulb of the pedicle provided anchorage in soft sediment. The discovery has important implications for the early evolution of lophotrochozoans, suggesting rooting of brachiopods into the sessile lophotrochozoans and the origination of their bivalved bauplan preceding the biomineralization of shell valves in crown brachiopods. Origin and early evolution of major animal clades in the Cambrian Period

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Avalon-type fossils are crucial to understanding the origin of Phanerozoic ecosystems, but their usual occurrence in volcaniclastic and siliciclastic facies greatly limits their paleobiological resolution. The unique carbonate-hosted assemblage of the Khatyspyt Formation, on the Olenek uplift of north-central Siberia, promises a much enhanced anatomical and paleo-ecological view of these enigmatic organisms. Avalon-type fossils are preserved by authigenic carbonate cementation in intervals of finely laminated nodular limestones (Khatyspyt-type taphonomic window). Interbedded silicified calcareous mudstones yield diverse carbonaceous compressions, occasionally with taphonomic phantoms of Avalon-type taxa (Miaohe-type taphonomic window). Styles of moldic preservation do not appear to be taxon selective, and therefore cannot alone be responsible for the morphological distinctiveness of Ediacaran macrofossils and the profound disparity in the taxonomic composition between fossil assemblages. On the other hand, the exclusion of Avalon-type fossils from carbonaceous compressions (Miaohe-type preservational window) is a real taphonomic signal that provides an important constraint on the properties of certain Ediacaran tissues.

The morphological disparity of lophotrochozoan phyla makes it difficult to predict the morphology of the last common ancestor. Only fossils of stem groups can help discover the morphological transitions that occurred along the roots of these phyla. Here, we describe a tubular fossil Yuganotheca elegans gen. et sp. nov. from the Cambrian (Stage 3) Chengjiang Lagerstatte (Yunnan, China) that exhibits an unusual combination of phoronid, brachiopod and tommotiid (Cambrian problematica) characters, notably a pair of agglutinated valves, enclosing a horseshoe-shaped lophophore, supported by a lower bipartite tubular attachment structure with a long pedicle with coelomic space. The terminal bulb of the pedicle provided anchorage in soft sediment. The discovery has important implications for the early evolution of lophotrochozoans, suggesting rooting of brachiopods into the sessile lophotrochozoans and the origination of their bivalved bauplan preceding the biomineralization of shell valves in crown brachiopods.