Cambrian Fossil Record Research Articles

Origin and Early Evolution of Echinoderms

Echinoderms are a major group (phylum) of invertebrate animals with a rich fossil record stretching back to the Cambrian period, approximately 518 million years ago. While all modern species are characterized by pentaradial (i.e., fivefold) symmetry, Cambrian echinoderms also include taxa with different types of symmetry (e.g., bilateral symmetry). These distinct forms were present from very early in the phylum's history, demonstrating that the initial diversification of echinoderm body plans was extremely rapid. The phylogenetic relationships of Cambrian echinoderms have long been debated, hindering efforts to reconstruct the evolution of the phylum, but recent analyses have consistently recovered bilaterally symmetrical forms as the earliest-diverging echinoderms. This reveals the sequence of character acquisition in echinoderm evolution, indicating that radial symmetry is a derived character of the group, which evolved after the acquisition of a mineralized skeleton. Cambrian echinoderms were adapted to diverse modes of life, with ecology an important factor shaping their early evolution. However, the reasons why echinoderms evolved their unique pentaradial body plan remain unclear. ▪The Cambrian fossil record provides valuable insights into the origin and early evolution of echinoderms over half a billion years ago.▪Cambrian echinoderms were morphologically diverse, with several extinct groups exhibiting character combinations that distinguish them from living species.▪Phylogenetic analyses of bilateral, asymmetrical, triradial, and pentaradial fossils have allowed us to decipher the assembly of the modern echinoderm body plan.▪Echinoderms became ecologically diverse early in their history, with varied modes of feeding, locomotion, and attachment.

Shadows of the past: An Ediacara-style taphonomic window in advent of Cambrian bioturbators

One of the hypotheses, explaining a sudden vanishing of the Ediacaran biota at the Ediacaran-Cambrian transition, correlates this phenomenon with a disappearance of specific taphonomical factors providing fossilization of soft-bodied organisms (‘Cheshire Cat’ hypothesis). The terminal Ediacaran-Lower Cambrian fossil record does reveal a progressive development of bioturbation and associated decrease of microbial mats, which played a crucial role in the late Precambrian ecosystems. But the tempo of the evolution of organism-sediment interactions significantly varied between different zones of marine basins. Hence, if the ‘Cheshire Cat’ hypothesis is correct, the sedimentary environments barely (if any) colonized by the burrowing eumetazoans might provide the unique interplay of conditions for habitat and Ediacara-style preservation of soft-bodied organisms even in the early Cambrian. Herein we present the new carbon isotope data calibrating the Terreneuvian record in the Ura Uplift of South Siberia and also report fossilized soft-bodied organisms morphologically-close to the Ediacaran holdfasts preserved in association with microbially-induced sedimentary structures in the Cambrian Stage 2 carbonate-siliciclastic Nokhtuisk Formation. Although the late Terreneuvian age, this marginal marine succession is poorly bioturbated and hosts the soft-bodied fossils in the intervals undisturbed by the burrowing organisms. Our study demonstrates the possibility of Ediacara-style preservation of soft-bodied organisms in specific habitat and taphonomic conditions until being replaced by the Cambrian endobenthos.

Morphometric analysis ofSkiagia-plexus acritarchs from the early Cambrian of North Greenland: toward a meaningful evaluation of phenotypic plasticity

AbstractThe Cambrian evolutionary radiations are marked by spectacular biotic turnovers and the establishment of increasingly tiered food chains. At the base of these food chains are primary producers, which in the Cambrian fossil record are chiefly represented among organic-walled microfossils. The majority of these microfossil remains have traditionally been attributed to an informal category of incertae sedis called “acritarchs,” based entirely on form taxonomy. Acritarch form taxa have been intensely used for biostratigraphy and in large-scale studies of phytoplankton diversity. However, both prospects have been challenged by cases of taxonomic inconsistencies and oversplitting arising from the large phenotypic plasticity seen among these microfossils. The acritarch form genusSkiagiastands as an ideal case study to explore these taxonomic challenges, because it encompasses a number of form species widely used in lower Cambrian biostratigraphy. Moreover, subtle morphological differences amongSkiagiaspecies were suggested to underlie key evolutionary innovations toward complex reproductive strategies. Here we apply a multivariate morphometric approach to investigate the morphological variation ofSkiagia-plexus acritarchs using an assemblage sourced from the Buen Formation (Cambrian Series 2, Stages 3–4) of North Greenland. Our analysis showed that the species-level classification ofSkiagiadiscretizes a continuous spectrum of morphologies. While these findings bring important taxonomic and biostratigraphic hurdles to light, the unequal frequency distribution of life cycle stages amongSkiagiaspecies suggests that certain elements of phytoplankton paleobiology are nonetheless captured bySkiagiaform taxonomy. These results demonstrate the value of using morphometric tools to explore acritarch phenotypic plasticity and its potential ontogenetic and paleoecological drivers in Cambrian ecosystems.

Nonmineralized triradial conulariids from the lowermost Cambrian Stage 2 of the Olenek Uplift, Siberian Platform

AbstractIn the early Cambrian fossil record, triradial symmetry is typical for anabaritids and occurs among carinachitids. The former are an extinct group of minute benthic cnidarians covered with a calcareous tubular exoskeleton. The origin of the anabaritids is poorly understood, but previously reported triradial pyramid-shaped steinkerns and molds of the oldest conulariids, Vendoconularia, from the upper Ediacaran of the White Sea region suggested the anabaritids were closely related to conulariids. However, triradial symmetry could originate independently in different lineages in the late Ediacaran and early Cambrian. Herein we describe a new taxon, Ilankirus kessyusensis new genus new species, from the base of the Cambrian Stage 2 of the Olenek Uplift (Siberian Platform). These fossils occur as ornamented steinkerns in the shape of trilateral pyramids and lack any relics of a mineralized exoskeleton. Abundant plastic deformations and fractures of the casts suggest the organism was weakly if at all mineralized. The steinkerns are encrusted with a thin patina of iron-rich chlorite (chamosite) formed because of a multistage diagenetic replacement of authigenic glauconite (glauconite–berthierine–chamosite) under reducing conditions of oxygen-depauperate pore- and seawater. Both lacking two major autapomorphies of the crown-group conulariids (mineralized periderm and quadrate cross section of the oral region of the periderm), the late Ediacaran triradial Vendoconularia and Terreneuvian Ilankirus represent stem-group conulariids.UUID: http://zoobank.org/a2ce04fa-36a5-485a-806c-f13f4749fc7f

The 'biomineralization toolkit' and the origin of animal skeletons.

Biomineralized skeletons are widespread in animals, and their origins can be traced to the latest Ediacaran or early Cambrian fossil record, in virtually all animal groups. The origin of animal skeletons is inextricably linked with the diversification of animal body plans and the dramatic changes in ecology and geosphere-biosphere interactions across the Ediacaran-Cambrian transition. This apparent independent acquisition of skeletons across diverse animal clades has been proposed to have been driven by co-option of a conserved ancestral genetic toolkit in different lineages at the same time. This 'biomineralization toolkit' hypothesis makes predictions of the early evolution of the skeleton, predictions tested herein through a critical review of the evidence from both the fossil record and development of skeletons in extant organisms. Furthermore, the distribution of skeletons is here plotted against a time-calibrated animal phylogeny, and the nature of the deep ancestors of biomineralizing animals interpolated using ancestral state reconstruction. All these lines of evidence point towards multiple instances of the evolution of biomineralization through the co-option of an inherited organic skeleton and genetic toolkit followed by the stepwise acquisition of more complex skeletal tissues under tighter biological control. This not only supports the 'biomineralization toolkit' hypothesis but also provides a model for describing the evolution of complex biological systems across the Ediacaran-Cambrian transition.

Bradoriids (Arthropoda) and the Cambrian Diversification

Bradoriids, among the earliest arthropods to appear in the fossil record, are extinct, ostracod-like bivalved forms that ranged from the early Cambrian to the Middle Ordovician. Bradoriids are notable for having appeared in the Cambrian fossil record before the earliest trilobites, and considering their rapid ascent to high genus-level diversity, provide key data for our understanding of the evolutionary dynamics of the Cambrian Explosion. This paper presents a broad review of bradoriid paleobiology. It is hypothesized here that an allele of Antennapedia determines whether bradoriid shields are preplete, amplete, or postplete. The preplete configuration of the shields of Cambroarchilocus tigris gen. nov. sp. nov. suggests that shield rowing motion may have propelled the animal backwards. Arcuate scars attributed here to a microdurophagous predator (Arcuoichnus pierci nov. ichnogen. nov. ichnosp.) occur on the paratype of Cambroarchilocus tigris gen. nov. sp. nov.

Trilobite evolutionary rates constrain the duration of the Cambrian explosion

Trilobites are often considered exemplary for understanding the Cambrian explosion of animal life, due to their unsurpassed diversity and abundance. These biomineralized arthropods appear abruptly in the fossil record with an established diversity, phylogenetic disparity, and provincialism at the beginning of Cambrian Series 2 (∼521 Ma), suggesting a protracted but cryptic earlier history that possibly extends into the Precambrian. However, recent analyses indicate elevated rates of phenotypic and genomic evolution for arthropods during the early Cambrian, thereby shortening the phylogenetic fuse. Furthermore, comparatively little research has been devoted to understanding the duration of the Cambrian explosion, after which normal Phanerozoic evolutionary rates were established. We test these hypotheses by applying Bayesian tip-dating methods to a comprehensive dataset of Cambrian trilobites. We show that trilobites have a Cambrian origin, as supported by the trace fossil record and molecular clocks. Surprisingly, they exhibit constant evolutionary rates across the entire Cambrian, for all aspects of the preserved phenotype: discrete, meristic, and continuous morphological traits. Our data therefore provide robust, quantitative evidence that by the time the typical Cambrian fossil record begins (∼521 Ma), the Cambrian explosion had already largely concluded. This suggests that a modern-style marine biosphere had rapidly emerged during the latest Ediacaran and earliest Cambrian (∼20 million years), followed by broad-scale evolutionary stasis throughout the remainder of the Cambrian.

Northwest Africa’s Ediacaran to early Cambrian fossil record, its oldest metazoans and age constraints for the basal Taroudant Group (Morocco)

The Anti-Atlas area in southern Morocco preserves one of the most complete latest Ediacaran to Cambrian shallow-marine carbonate records in the world and serves as a standard for global chemostratigraphic δ13Ccarb curves. Contrary to the excellent chemostratigraphic coverage, neither metazoan fossils (only stromatolites and calcified algae) nor radiometric age constraints were hitherto available for the basal part of this carbonate record, the Tifnout Member of the Adoudou Formation of the Taroudant Group. We report an Ediacara-type fauna from shallow-marine sandstones from the basal Tabia Member and microfossils from dolostones from both the Tabia and the Tifnout members of the Adoudou Formation. The former are discoidal fossils and share many characteristics with the form genus Aspidella and exhibit a peculiar style of preservation, pointing towards an early diagenetic two-stage (carbonate followed by silica) cementation history, leading to good preservation of these supposedly once soft-bodied organisms. The dolostone-hosted microfossils are preserved as sparite-filled molds enclosed by a thin opaque envelope. Despite their small size, they resemble late Ediacaran calcifying metazoans such as Namacalathus and tubular small shelly fossils. Together, these fossils constitute the oldest known direct evidence for presumably metazoan life from Northwest Africa. We also report new time constraints for the Adoudou Formation and fossils enclosed therein based on combined LA-ICP-MS and high-precision CA-ID-TIMS U-Pb zircon geochronology of detrital and volcanic zircons. Together with new chemostratigraphic (δ13Ccarb) and literature data, these new age constraints suggest a late Ediacaran age (time span between ca. 561 and 542 Ma) for the discoidal Ediacara-type fossils from the basal Tabia Member, and a latest Ediacaran to earliest Cambrian (time span between ca. 544 and 540 Ma) and an early Cambrian age (time span between ca. 528 and 523 Ma) for the two microfossil faunas from the Tifnout Member.

Ichnological evidence for meiofaunal bilaterians from the terminal Ediacaran and earliest Cambrian of Brazil.

The evolutionary events during the Ediacaran-Cambrian transition (~541 Myr ago) are unparalleled in Earth history. The fossil record suggests that most extant animal phyla appeared in a geologically brief interval, with the oldest unequivocal bilaterian body fossils found in the Early Cambrian. Molecular clocks and biomarkers provide independent estimates for the timing of animal origins, and both suggest a cryptic Neoproterozoic history for Metazoa that extends considerably beyond the Cambrian fossil record. We report an assemblage of ichnofossils from Ediacaran-Cambrian siltstones in Brazil, alongside U-Pb radioisotopic dates that constrain the age of the oldest specimens to 555-542 Myr. X-ray microtomography reveals three-dimensionally preserved traces ranging from 50 to 600 μm in diameter, indicative of small-bodied, meiofaunal tracemakers. Burrow morphologies suggest they were created by a nematoid-like organism that used undulating locomotion to move through the sediment. This assemblage demonstrates animal-sediment interactions in the latest Ediacaran period, and provides the oldest known fossil evidence for meiofaunal bilaterians. Our discovery highlights meiofaunal ichnofossils as a hitherto unexplored window for tracking animal evolution in deep time, and reveals that both meiofaunal and macrofaunal bilaterians began to explore infaunal niches during the late Ediacaran.

Current Understanding of Ecdysozoa and its Internal Phylogenetic Relationships.

Twenty years after its proposal, the monophyly of molting protostomes-Ecdysozoa-is a well-corroborated hypothesis, but the interrelationships of its major subclades are more ambiguous than is commonly appreciated. Morphological and molecular support for arthropods, onychophorans and tardigrades as a clade (Panarthropoda) continues to be challenged by a grouping of tardigrades with Nematoida in some molecular analyses, although onychophorans are consistently recovered as the sister group of arthropods. The status of Cycloneuralia and Scalidophora, each proposed by morphologists in the 1990s and widely employed in textbooks, is in flux: Cycloneuralia is typically non-monophyletic in molecular analyses, and Scalidophora is either contradicted or incompletely tested because of limited genomic and transcriptomic data for Loricifera, Kinorhyncha, and Priapulida. However, novel genomic data across Ecdysozoa should soon be available to tackle these difficult phylogenetic questions. The Cambrian fossil record indicates crown-group members of various ecdysozoan phyla as well as stem-group taxa that assist with reconstructing the most recent common ancestor of panarthropods and cycloneuralians.

Meiofaunal deuterostomes from the basal Cambrian of Shaanxi (China).

Deuterostomes include the group we belong to (vertebrates) as well as an array of disparate forms that include echinoderms, hemichordates and more problematic groups such as vetulicolians and vetulocystids. The Cambrian fossil record is well-populated with representative examples, but possible intermediates are controversial and the nature of the original deuterostome remains idealized. Here we report millimetric fossils, Saccorhytus coronarius nov. gen., nov. sp., from an Orsten-like Lagerstätte from the earliest Cambrian period of South China, which stratigraphically are amongst the earliest of deuterostomes. The bag-like body bears a prominent mouth and associated folds, and behind them up to four conical openings on either side of the body as well as possible sensory structures. An anus may have been absent, and correspondingly the lateral openings probably served to expel water and waste material. This new form has similarities to both the vetulicolians and vetulocystids and collectively these findings suggest that a key step in deuterostome evolution was the development of lateral openings that subsequently were co-opted as pharyngeal gills. Depending on its exact phylogenetic position, the meiofaunal habit of Saccorhytus may help to explain the major gap between divergence times seen in the fossil record and estimates based on molecular clocks.

Preservation and phylogeny of Cambrian ecdysozoans tested by experimental decay of Priapulus.

The exceptionally preserved Cambrian fossil record provides unique insight into the early evolutionary history of animals. Understanding of the mechanisms of exceptional soft tissue preservation frames all interpretations of the fauna and its evolutionary significance. This is especially true for recent interpretations of preserved nervous tissues in fossil ecdysozoans. However, models of soft tissue preservation lack empirical support from actualistic studies. Here experimental decay of the priapulid Priapulus reveal consistent bias towards rapid loss of internal non-cuticular anatomy compared with recalcitrant cuticular anatomy. This is consistent with models of Burgess Shale-type preservation and indicates that internal tissues are unlikely to be preserved with fidelity if organically preserved. This pattern, along with extreme body margin distortion, is consistent with onychophoran decay, and is therefore resolved as general for early ecdysozoans. Application of these patterns to phylogenetic data finds scalidophoran taxa to be very sensitive to taphonomically informed character coding, but not panarthropodan taxa. Priapulid decay also have unexpected relevance for interpretation of myomeres in fossil chordates. The decay data presented serve not only as a test of models of preservation but also a framework with which to interpret ecdysozoan fossil anatomies, and the subsequent evolutionary inferences drawn from them.

Arthropod eyes: The early Cambrian fossil record and divergent evolution of visual systems

Four types of eyes serve the visual neuropils of extant arthropods: compound retinas composed of adjacent facets; a visual surface populated by spaced eyelets; a smooth transparent cuticle providing inwardly directed lens cylinders; and single-lens eyes. The first type is a characteristic of pancrustaceans, the eyes of which comprise lenses arranged as hexagonal or rectilinear arrays, each lens crowning 8–9 photoreceptor neurons. Except for Scutigeromorpha, the second type typifies Myriapoda whose relatively large eyelets surmount numerous photoreceptive rhabdoms stacked together as tiers. Scutigeromorph eyes are facetted, each lens crowning some dozen photoreceptor neurons of a modified apposition-type eye. Extant chelicerate eyes are single-lensed except in xiphosurans, whose lateral eyes comprise a cuticle with a smooth outer surface and an inner one providing regular arrays of lens cylinders. This account discusses whether these disparate eye types speak for or against divergence from one ancestral eye type. Previous considerations of eye evolution, focusing on the eyes of trilobites and on facet proliferation in xiphosurans and myriapods, have proposed that the mode of development of eyes in those taxa is distinct from that of pancrustaceans and is the plesiomorphic condition from which facetted eyes have evolved. But the recent discovery of enormous regularly facetted compound eyes belonging to early Cambrian radiodontans suggests that high-resolution facetted eyes with superior optics may be the ground pattern organization for arthropods, predating the evolution of arthrodization and jointed post-protocerebral appendages. Here we provide evidence that compound eye organization in stem-group euarthropods of the Cambrian can be understood in terms of eye morphologies diverging from this ancestral radiodontan-type ground pattern. We show that in certain Cambrian groups apposition eyes relate to fixed or mobile eyestalks, whereas other groups reveal concomitant evolution of sessile eyes equipped with optics typical of extant xiphosurans. Observations of fossil material, including that of trilobites and eurypterids, support the proposition that the ancestral compound eye was the apposition type. Cambrian arthropods include possible precursors of mandibulate eyes. The latter are the modified compound eyes, now sessile, and their underlying optic lobes exemplified by scutigeromorph chilopods, and the mobile stalked compound eyes and more elaborate optic lobes typifying Pancrustacea. Radical divergence from an ancestral apposition type is demonstrated by the evolution of chelicerate eyes, from doublet sessile-eyed stem-group taxa to special apposition eyes of xiphosurans, the compound eyes of eurypterids, and single-lens eyes of arachnids. Different eye types are discussed with respect to possible modes of life of the extinct species that possessed them, comparing these to extant counterparts and the types of visual centers the eyes might have served.

Homology of Head Sclerites in Burgess Shale Euarthropods

The Cambrian fossil record of euarthropods (extant arachnids, myriapods, crustaceans, hexapods) has played a major role in understanding the origins of these successful animals and indicates that early ancestors underwent an evolutionary transition from soft-bodied taxa (lobopodians) to more familiar sclerotized forms with jointed appendages [1-3]. Recent advances in paleoneurology and developmental biology show that this major transformation is reflected by substantial changes in the head region of early euarthropods, as informed by the segmental affinity of the cephalic appendages [1, 4-6]. However, data on the implications of this reorganization for non-appendicular exoskeletal structures are lacking, given the difficulty of inferring the precise segmental affinities of these features. Here, I report neurological remains associated with the stalked eyes and "anterior sclerite" in the (middle Cambrian) Burgess Shale euarthropods Helmetia expansa and Odaraia alata and provide evidence that these features are associated with nerve traces originating from the anterior brain region, the protocerebrum. The position of the protocerebral ganglia in exceptionally preserved Cambrian euarthropods indicates the homology of the anterior sclerite in extinct groups (e.g., fuxianhuiids, bivalved forms, artiopodans [7, 8]) and allows new comparisons with the dorsal cephalic plate of radiodontans, large nektonic predators whose anterior segmental organization bears fundamental similarities to that of Paleozoic lobopodians [1, 6, 9, 10]. These observations allow reconstruction of the segmental architecture of the head region in the earliest sclerotized euarthropods and demonstrate the deep homology between exoskeletal features in an evolutionary continuum of taxa with distinct types of body organization.

Intelligent Design

Darwin’s Doubt claims to review research on the earliest forms of metazoan life. Part One presents a history of discovery and interpretation of the Cambrian animals through both paleontology and genetics; Part Two, research on protein evolution, evo-devo, and epigenetics; Part Three, ideas of “self-organization.” The final four chapters purport to demonstrate how Intelligent Design provides the ultimate explanation of all these phenomena.Meyer, Director of the Center for Science and Culture at the Discovery Institute, emphasizes that our understanding of the Cambrian explosion remains incomplete. He recapitulates the arguments from his earlier book, Signature in the Cell, and from his Discovery Institute colleagues Michael Behe, William Dembski, Jonathan Wells, and Douglas Axe, who all maintain that there have never been a sufficient number of cell divisions, nor a sufficient number of years in the history of the universe for unguided processes to bring about animal forms. Darwin’s Doubt thoroughly reviews the arguments of the Intelligent Design movement and, thus, could save a person the trouble of reading all their previous publications.Reporting on the discovery that developmental gene regulatory networks are highly conserved and not tolerant of mutations, Meyer asserts that this must have always been the case, thereby preventing the development of varied animal body plans: “The system of gene regulation that controls animal-body-plan development is exquisitely integrated, so that significant alterations in these gene regulatory networks inevitably damage or destroy the developing animal. But given this, how could a new animal body plan, and the new dGRNs necessary to produce it, ever evolve gradually via mutation and selection from a preexisting body plan and set of dGRN’s?” (p. 269). The idea that in early metazoan populations such networks, and the interplay among them, might have evolved spectacular diversity before they became so highly integrated is dismissed, even though this idea is central to current research.Meyer moves to the crux of his argument on page 337: “Neither neo-Darwinian nor a host of more recent proposals [he includes a list] have succeeded in explaining the origin of the novel animal forms that arose in the Cambrian period. Yet all these evolutionary theories have two things in common: they rely on strictly material processes, and they also have failed to identify a cause capable of generating the information necessary to produce new forms of life…. Is it possible that intelligent design – the purposeful action of a conscious and rational agent – might have played a role in the Cambrian explosion?”The reliance on “strictly material processes” that Meyer finds limiting is the basis of modern (since the 17th century) science. His concern for “generating the information necessary” reflects the Intelligent Design movement’s premise that anything really complex requires a blueprint, despite the fact that modern biology has shown that this is not true: DNA is not a blueprint and organisms are not constructed like machines.In addressing how a designer actually manipulated atoms and molecules to create brand-new life forms, all Meyer claims is that “intelligent agents can act suddenly or discretely in accord with their powers of rational choice or volition, even if they do not always do so…. If body plans arose as the result of an intelligent agent actualizing an immaterial plan or idea, then an extensive series of material precursors to the first animals need not exist in the fossil record…. Mental plans or concepts need not leave a material trace” (p. 375). To quote Nick Matzke, formerly of the National Center for Science Education, this explains the appearance of the first animals and their body plans by saying, in effect, “POOF!”The author and his institute claim that Darwin’s Doubt is an important contribution to modern biological science. There are major clues that it isn’t. HarperCollins chose to publish this under their HarperOne imprint, and HarperOne describes itself thus: The most important books across the full spectrum of religion, spirituality, and personal growth…. A more important clue is found in the last chapter, entitled What’s at Stake. In science, what would be at stake would be a better understanding of the Cambrian fossil record and the processes of life and evolution. But Meyer instead tells us: “Modern life suspends many of us, so we feel, high over a chasm of despair. It provokes feelings of dizzying anxiety – in a word, vertigo. The evidence of a purposeful design behind life, on the other hand, offers the prospect of significance, wholeness, and hope.” A heavy burden for a trilobite to bear!**for the confusion this book causes about the evidence and the science of studying the evidence. It rates four frogs as a review of the thinking of the Intelligent Design movement.

The Cambrian explosion: the construction of animal biodiversity

In 1909 Charles Walcott discovered the remarkable soft-bodied animal fauna of the Middle Cambrian age Burgess Shale in British Columbia. These fossils have become the iconic representation of the astonishing expansion of animal diversity during the Cambrian, including the first appearance in the fossil record of essentially all living phyla, known as the “Cambrian explosion.” Although the Burgess Shale biota is the public face of the Cambrian explosion, even appearing on a 1995 cover of Time, it is only a small part of the story (and somewhat late in the tale). Most readers will be familiar with the Cambrian explosion through Stephen Jay Gould’s 1989 book Wonderful Life (1989) or perhaps Martin Brasier’s more recent Darwin’s Lost World (2009). Both of these books were written to reach a general audience. What has been lacking to date is a more technical book, aimed at students and professionals, which summarizes and introduces the tremendous amount of research that this episode of earth history has received in the last 25 years. The Cambrian Explosion: The Construction of Animal Biodiversity by Douglas H Erwin and James W Valentine more than fills this need. It will become required reading for anyone who wants to understand the multiple lines of evidence, from geology, geochemistry, paleontology, genetics, phylogenetics, comparative morphology, and ecology, that have to be integrated to understand one of the most important episodes in the history of life. The authors, both paleontologists, are deeply respected authorities on evolution and the Cambrian fossil record, with a welldeserved reputation for innovative approaches. This book necessarily covers a tremendous amount of ground; the forty-two pages of references demonstrate the depth and breadth of scholarship that the authors summarize. The first section covers the geological context of the explosion, stretching back to the Snowball Earth episodes of the Cryogenian. This includes summaries of the current status of the absolute and relative time scales for this period and what geochemical proxies reveal about ocean chemistry, including oxygenation states and changes in the carbon cycle. The latter is unfortunately the weakest part of the book; there is very poor integration between figures (which come from external sources) and text; the figure captions and labels are inadequate. The book hits its stride in the next section. It begins with a brief review of phylogenetics and of living metazoan morphology and relationships, a subject Valentine explored in more depth in his comprehensive earlier book, On the Origin of Phyla (2004). This is followed

Deuterostomic Development in the Protostome Priapulus caudatus

The fate of the blastopore during development in the bilaterian ancestor is currently not well understood. In deuterostomes, the blastopore forms the anus, but its fate in protostome groups is variable. This variability, combined with an absence of information from key taxa, hampers the reconstruction of the ancestral developmental mode of the Protostomia and the Bilateria. The blastopore fate of the bilaterian ancestor plays a crucial role in understanding the transition from radial to bilateral symmetric organisms. Priapulids have a conservative morphology, an abundant Cambrian fossil record, and a phylogenetic position that make them a key group in understanding protostome evolution. Here, we characterize gastrulation and the embryonic expression of genes involved in bilaterian foregut and hindgut patterning in Priapulus caudatus. We show that the blastopore gives rise to the anus at the vegetal pole and that the hindgut markers brachyury and caudal are expressed in the blastopore and anus, whereas the foregut markers foxA and goosecoid are expressed in the mouth in the animal hemisphere. Thereby, gastrulation in the conservatively evolving protostome P. caudatus follows strictly a deuterostomic pattern. These results are more compatible with a deuterostomic rather than protostomic (blastopore forms the mouth) or amphistomic (mouth and anus are formed simultaneously) mode of development in the last common bilaterian ancestor.

The advent of hard-part structural support among the Ediacara biota: Ediacaran harbinger of a Cambrian mode of body construction

The apparent lack of taxonomic continuity between the Precambrian and Cambrian fossil records has led to controversial and conflicting interpretations about the Ediacara biota and their place in the evolution of metazoan life on this planet. This has been further complicated by the absence of similar modes of construction between these faunas and the rarity of Precambrian skeletonized fossils. We describe a new Ediacaran organism that represents the oldest multielement organism with structural support through either biomineralization or chitin. Coronacollina acula gen. et sp. nov. from the Ediacara Member (Rawnsley Quartzite) was constructed from a framework of rigid and brittle elements that disarticulated after death. It reveals a constructional mode not recognized previously among members of this assemblage, but one that was prevalent among Cambrian organisms. Coronacollina consists of a truncated cone associated with spicules, up to 37 cm in length, diverging radially from the cone. This constructional morphology is similar to the Cambrian Choia , a low conical demosponge with a corona of long spicules, providing a long-predicted constructional link between the Ediacara biota and the Cambrian fossil record.

The role of docosahexaenoic and the marine food web as determinants of evolution and hominid brain development: The challenge for human sustainability

Life originated on this planet about 3 billion years ago. For the first 2.5 billion years of life there was ample opportunity for DNA modification. Yet there is no evidence of significant change in life forms during that time. It was not until about 600 million years ago, when the oxygen tension rose to a point where air-breathing life forms became thermodynamically possible, that a major change can be abruptly seen in the fossil record. The sudden appearance of the 32 phyla in the Cambrian fossil record was also associated with the appearance of intracellular detail not seen in previous life forms. That detail was provided by cell membranes made with lipids (membrane fats) as structural essentials. Lipids thus played a major, as yet unrecognised, role as determinants in evolution. The compartmentalisation of intracellular, specialist functions as in the nucleus, mitochondria, reticulo-endothelial system and plasma membrane led to cellular specialisation and then speciation. Thus, not only oxygen but also the marine lipids were drivers in the Cambrian explosion. Docosahexaenoic acid (DHA) (all-cis-docosa-4,7,10,13,16,19-hexaenoic acid, C22:6ω3 or C22:6, n-3, DHA) is a major feature of marine lipids. It requires six oxygen atoms to insert its six double bonds, so it would not have been abundant before oxidative metabolism became plentiful. DHA provided the membrane backbone for the emergence of new photoreceptors that converted photons into electricity, laying the foundation for the evolution of other signalling systems, the nervous system and the brain. Hence, the ω3 DHA from the marine food web must have played a critical role in human evolution. There is also clear evidence from molecular biology that DHA is a determinant of neuronal migration, neurogenesis and the expression of several genes involved in brain growth and function. That same process was essential to the ultimate cerebral expansion in human evolution. There is now incontrovertible support of this hypothesis from fossil evidence of human evolution taking advantage of the marine food web. Lipids are still modifying the present evolutionary phase of our species; their signature is evident in the changing panorama of non-communicable diseases. The most worrying change in disease pattern is the sharp rise in brain disorders, which, in the European Union, has overtaken the cost of all other burdens of ill health at €386 billion for the 25 member states at 2004 prices. In 2007, the UK cost was estimated at £77 billion and confirmed in 2010 at £105 billion - greater than heart disease and cancer combined. The rise in mental ill health is now being globalised. The solution to the rising vascular disorders in the last century and now brain disorders in this century lies in a radical reappraisal of the food system, which last century was focussed on protein and calories, with little attention paid to the requirements of the brain - the very organ that was the determinant of human evolution. With the marine fish catch having plateaued 20 years ago and its sustainability now under threat, a critical aspect of this revision is the development of marine agriculture from estuarine, coastal and oceanic resources. Such action is likely to play a key role in future health and intelligence.

First record of a bivalved larval shell in Early Cambrian tommotiids and its phylogenetic significance

Abstract: Brachiopods are marine Lophotrochozoa whose soft parts are enclosed in a bivalved shell. Although brachiopods are represented by a rich record from the Early Cambrian to the present, the origin of their bivalved body plan remains controversial. The Early Cambrian organophosphatic tommotiids Micrina and Paterimitra from Australia have been proposed as stem brachiopods. Here, we describe their earliest ontogeny, indicating that tommotiids possessed bivalved planktotrophic larvae. The curious combinations of characters in Micrina and Paterimitra indicate that they may belong to the stems of the Linguliformea and Rhynchonelliformea, respectively. The bivalved shell of adult living brachiopods may represent a plesiomorphic character retained from planktic tommotiid larvae; the crown group body plan of the Brachiopoda may have evolved through the paedomorphic retention of a bivalved larval state.