The two phases of the Cambrian Explosion

Abstract The dynamics of how metazoan phyla appeared and evolved – known as the Cambrian Explosion – remains elusive. We present a quantitative analysis of the temporal distribution (based on occurrence data of fossil species sampled in each time interval) of lophotrochozoan skeletal species (n = 430) from the terminal Ediacaran to Cambrian Stage 5 (~545 – ~505 Million years ago (Ma)) of the Siberian Platform, Russia. We use morphological traits to distinguish between stem and crown groups. Possible skeletal stem group lophophorates, brachiopods, and molluscs (n = 354) appear in the terminal Ediacaran (~542 Ma) and diversify during the early Cambrian Terreneuvian and again in Stage 2, but were devastated during the early Cambrian Stage 4 Sinsk extinction event (~513 Ma) never to recover previous diversity. Inferred crown group brachiopod and mollusc species (n = 76) do not appear until the Fortunian, ~537 Ma, radiate in the early Cambrian Stage 3 (~522 Ma), and with minimal loss of diversity at the Sinsk Event, continued to diversify into the Ordovician. The Sinsk Event also removed other probable stem groups, such as archaeocyath sponges. Notably, this diversification starts before, and extends across the Ediacaran/Cambrian boundary and the Basal Cambrian Carbon Isotope Excursion (BACE) interval (~541 to ~540 Ma), ascribed to a possible global perturbation of the carbon cycle. We therefore propose two phases of the Cambrian Explosion separated by the Sinsk extinction event, the first dominated by stem groups of phyla from the late Ediacaran, ~542 Ma, to early Cambrian stage 4, ~513 Ma, and the second marked by radiating bilaterian crown group species of phyla from ~513 Ma and extending to the Ordovician Radiation

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

Evolutionary origins of animal skeletal biomineralization

The evolutionary history of biomineralization in animals is crucial to our understanding of modern mineralized tissues. Traditional methods of unravelling this history have aimed to derive a theory of the development of biomineralization through evolution by the comparison of mineralized systems in model organisms. This has led to the recognition of the ‘biomineralization toolkit’ and raised the question of the homology of mineralized tissues versus convergent or parallel evolution. The ‘new animal phylogeny’ reveals that many of the groups known to biomineralize sit among close relatives that do not, and it favours an interpretation of convergent or parallel evolution for biomineralization in animals. In addition, the fossil record of the earliest mineralized skeletons presents a rapid proliferation of biomineralization across a range of animal phyla with fossil representatives of many modern biomineralizing phyla. A synthesis of molecular, developmental, phylogenetic and fossil evidence demonstrates the convergent or parallel evolution of biomineralization in animals at the phylum level. The fossil record of the Cambrian explosion not only provides vital evidence for the evolution of animal mineralized tissues but also suggests a mechanism for its rapid and synchronous convergent origin

Cutting the first ‘teeth’: a new approach to functional analysis of conodont elements

The morphological disparity of conodont elements rivals the dentition of all other vertebrates, yet relatively little is known about their functional diversity. Nevertheless, conodonts are an invaluable resource for testing the generality of functional principles derived from vertebrate teeth, and for exploring convergence in a range of food-processing structures. In a few derived conodont taxa, occlusal patterns have been used to derive functional models. However, conodont elements commonly and primitively exhibit comparatively simple coniform morphologies, functional analysis of which has not progressed much beyond speculation based on analogy. We have generated high-resolution tomographic data for each morphotype of the coniform conodont Panderodus acostatus. Using virtual cross sections, it has been possible to characterize changes in physical properties associated with individual element morphology. Subtle changes in cross-sectional profile have profound implications for the functional performance of individual elements and the apparatus as a whole. This study has implications beyond the ecology of a single conodont taxon. It provides a basis for reinterpreting coniform conodont taxonomy (which is based heavily on cross-sectional profiles), in terms of functional performance and ecology, shedding new light on the conodont fossil record. This technique can also be applied to more derived conodont morphologies, as well as analogous dentitions in other vertebrates and invertebrates

Role OSN v prevenci a řešení mezinárodních konfliktů: Případová studie bývalé Jugoslávie a Sierry Leone

The aim of the Masters Thesis is to analyze the role of the United Nations in conflict prevention and resolution. The theoretical framework lists a set of tools that should be followed in order to achieve a success in UN peacekeeping operations. Based on the list of the tools the Masters Thesis continues with two case studies. The case study of Former Yugoslavia and its peacekeeping operation UNPROFOR is considered to be one of the biggest failures of the UN peacekeeping. In contrary, the case study of Sierra Leoneans peacekeeping operation UNAMSIL then represents a successful UN peacekeeping operation. The Masters Thesis is concluded by a comparative analysis of these two case studies which evaluates the development and the implementation of these tools in each of the case studies. The analysis brings the answer to the research question which asks why some UN peacekeeping operations are successful whereas others completely fail

Functional adaptation underpinned the evolutionary assembly of the earliest vertebrate skeleton

Conodonts are the first vertebrates to bear a mineralized skeleton, restricted to an array of tooth‐like feeding elements. The functional implications for the development of tooth‐like elements differentiated into two tissues is tested using 2D finite element modeling, mapping the patterns of stress and strain that elements with differing material properties exhibited during function. Addition of a stiff crown does not change the patterns of stress, rather it reduces the deformation of the element under the same force regime, and distributes stress more evenly across the element. The euconodont crown, like vertebrate dental enamel, serves to stiffen the element and protect the underlying dentine. Stiffness of the crown may be a contributing factor to the subsequent diversity of euconodont form, and logically function, by allowing a greater range of feeding strategies to be employed. The euconodont crown also serves as an analogue to enamel and enameloid, demonstrating that enamel‐like tissues have evolved multiple times in independent vertebrate lineages, likely as a response to similar selective pressures. Conodonts can, therefore, serve as an independent test on hypotheses of the effect of ecology on the development of the vertebrate skeleton

A new tannuolinid problematic from the lower Cambrian of the Sukharikha River in northern Siberia

A new species of tannuolinid, Tannuolina pavlovi, is reported from the basal Krasnoporog Formation cropping out along the Sukharikha River at the northwestern margin of the Siberian Platform. The new material expands the geographic range of tannuolinids onto the Siberian Platform and extends their stratigraphic range into the basal Tommotian Stage of the traditional Lower Cambrian. The oldest appearance of tannuolinids in the fossil record is shown herein to have occurred as early as the oldest appearance of the linguliformean brachiopods, of which tannuolinids have been suggested to be a stem group. The sellate sclerites of T. pavlovi sp. nov. are different from those of the other known species of Tannuolina in the consistent absence of a pronounced sella and duplicature on the respective sides of sclerites, as well as in having regularly distributed large setal pores on the lateral edges. The mitral sclerites of T. pavlovi sp. nov. do not clearly show asymmetry of left and right forms, manifested in other species by the presence of the carina on the interior surface. Large pores aligned along the lateral margins of sellate sclerites regularly coalesce below the wall surface from an apically and an aperturally directed basal branch, forming a nearly right angle