Taphonomic versus ecologic controls on taxonomic relative abundance patterns in tempestites

Evaluation of taphonomic sample bias in tempestites is an important prerequisite for paleoecologic analysis in storm-dominated shelf settings. Data from tempestites in the Upper Cambrian Bison Creek Formation of southern Alberta indicate that size, sorting greatly influences trilobite relative abundance patterns. Shifts in relative abundances of taxa both within and between beds are matched by changes in trilobite size. That is, collections fall into two intergrading types of assemblage which are dominated by taxa which reach large and small adult sizes, respectively. These shifts in dominance are interpreted as taphonomic variability within a single trilobite biofacies. Biofacies differentiation is expressed as abrupt changes in relative abundance patterns that are independent of trilobite size. Qualitative general models for size-sorting in tempestites are developed. Size-sorting in graded tempestites produces vertical shifts in relative abundances of taxa that mimic those expected in short-term autogenic ecologic succession. Size-sorting between beds in a single section could lead to an overestimation of the number of biofacies present. Size-sorting between proximal and distal portions of a depth gradient might generate abundance patterns that mimic those expected for 'depth-related communities', even in the complete absence of any underlying ecologic differentiation. The models represent taphonomic null hypotheses which must be rejected prior to making paleoecologic inferences in storm-dominated shelf environments. 0 Taphonomy, tempestites, Trilobita, Upper Cambrian, paleoecology.     

湘西中寒武世多节类三叶虫新属种

描记湘西江南斜坡带中寒武统花桥组的多节类三叶虫４新属７新种：Ｃｈａｎｇｑｉｎｇｉａｌａｅｖｉｓｓｐ．ｎｏｖ，Ｆｉｓｓａｎｏｍｏｃａｒｅｌｌａｐａｉｂｉｅｎｓｉｓｇｅｎ．ｅｔｓｐ．Ｎｏｖ．，Ｈｕａｙｕａｎｉａｓｕｂｃａｌｖａｇｅｎ．ｅｔｓｐ．ｎｏｖ，Ｌｕａｓｐｉｓｄｅｃｏｒｏｓａｇｅｎ．ｅｔｓｐ．Ｎｏｖ．。Ｐａｒａｎｏｍｏｃａｒｅｌｌａｆｏｒｔｉｓｓｐ．Ｎｏｖ．，Ｗａｎｇｃｕｎｉａｗａｎｇｃｕｎｅｎｓｉｓｇｅｎ．ｅｔｓｐ．ｎｏｖ和Ｚｈｕｊｉａｈｕｎａｎｅｎｓｉｓｓｐ．Ｎｏｖ．。Ｆｉｓｓａｎｏｍｏｃａｒｅｌｌａｇｅｎ．ｎｏｖ．属Ａｎｏｍｏｃａｒｅｌｌｉｎｄａｅ科；Ｗａｎｇｃｕｎｉａｓｅｎ．ｎｏｖ．和Ｈｕａｙｕａｎｉａｓｅｎ．ｎｏｖ．分别存疑地归入Ｐａｐｙｒａｓｐｉｄａｅ科和Ａｎｏｍｏｃａｒｅｌｌｉｄａｅ科；Ｌｕａｓｐｉｓｇｅｎ．ｎｏｖ．的科级分类位置未定。所有新属种与华北和东北南部中寒武世动物群关系较密切，是斜坡相混生动物群的重要分子。     

Trait variability and stress: canalization, developmental stability and the need for a broad approach

Trait variability (particularly fluctuating asymmetry) may provide a general measure of environmental stress applicable across taxa but consistent empirical support is lacking. Historically, stress effects were considered to act independently on trait canalization, developmental noise and trait size. However, in trait comparisons these processes are often assumed to be associated. Here we reconsider this issue and implications for detecting stress effects using trait variability. Published studies that consider multiple environments report little association between the effects of environmental variation on trait canalization and on developmental noise measured as fluctuating asymmetry, sug-gesting that environmental effects often act independently on these processes. To further test the usefulness of trait variability as an indicator of stress, comparisons across environ-ments should take a broad approach and report on several measures of trait variability, rather than focusing on only one index of fluctuating asymmetry as is commonly done.     

A statistical/computer-graphic technique for assessing variation in tectonically deformed fossils and its application to Cambrian trilobites from Kashmir

Hughes, N. C. & Jell, P. A. 1992 07 15: A stdlislical/compuler-graphic technique for assessing variation in tectonically deformed fossils and its application to Cambrian trilobites from Kashmir.
A combined approach using statistics and computer graphics can help resolve patterns of morphological variation in tectonically deformed fossils. Bivariate analyses and Principal Components Analysis can be used to identify a generalized strain vector in populations of deformed fossils, the identification of which permits discrimination of biological and tectonically induced variation. Results can be used to determine the number of taxa and growth relationships within the population. Statistically resolved taxa can be compared with species described from other areas using computer-aided shape restorations. Application of these techniques to a sample of trilobite cranidia from the Cambrian of Kashmir demonstrates that the variation of characters used to diagnose several genera and species are ontogeny- or deformation-controlled. Seven previously described species of Saukia, Prosaukia, Hundwarella and Anomocare are best considered as a single species, Hundwarella personara . These results suggest a Middle Cambrian age for the fauna, which has affinities with other faunas described from India and northern China. Hundwarella personara shows a pattern of developmental flexibility similar to that seen in other Cambrian trilobites. Morphometrics, computer-aided restoration, deformed fossils, allometry, trilobites, Cambrian, Kashmir, developmental flexibility .     

Trilobite body patterning and the evolution of arthropod tagmosis

Preservation permitting patterns of developmental evolution can be reconstructed within long extinct clades, and the rich fossil record of trilobite ontogeny and phylogeny provides an unparalleled opportunity for doing so. Furthermore, knowledge of Hox gene expression patterns among living arthropods permit inferences about possible Hox gene deployment in trilobites. The trilobite anteroposterior body plan is consistent with recent suggestions that basal euarthropods had a relatively low degree of tagmosis among cephalic limbs, possibly related to overlapping expression domains of cephalic Hox genes. Trilobite trunk segments appeared sequentially at a subterminal generative zone, and were exchanged between regions of fused and freely articulating segments during growth. Homonomous trunk segment shape and gradual size transition were apparently phylogenetically basal conditions and suggest a single trunk tagma. Several derived clades independently evolved functionally distinct tagmata within the trunk, apparently exchanging flexible segment numbers for greater regionally autonomy. The trilobite trunk chronicles how different aspects of arthropod segmentation coevolved as the degree of tagmosis increased.     

THE ROLE OF HETEROCHRONY IN THE EVOLUTION OF CAMBRIAN TRILOBITES

1. Renewed interest in the role of changes to developmental regulation in organisms has highlighted the importance of heterochrony in the evolution of the Metazoa.
2. Beecher's interpretation of the evolution of the Trilobita as having been principally by peramorphosis is examined, as is the view of later workers, principally Stubblefield and Hupé, that paedomorphosis was a dominant factor in trilobite evolution.
3. Both peramorphosis and paedomorphosis are considered to have been important in trilobite evolution.
4. The role of paedomorphosis in the evolution of major morphological novelties is critically examined. Its importance in changes to the structure of the glabella is discussed and new terms proposed to describe the ontogenetic and phylogenetic state of the glabella.
5. The highly variable nature of early Cambrian trilobites, in particular the large degree of ontogenetic change, is considered, along with possible poor developmental control of the growth and moulting hormonal systems, to have been significant in providing a high degree of intrapopulational morphological variability. Selection of these heterochronic variants was responsible for the rapid diversification of the Trilobita during the Cambrian.     

Head segmentation of trilobites

Although trilobites have provided research subjects for more than two centuries, their head segmentation has remained unresolved. Four glabellar furrows (SO and S1–S3) marking the segmental boundaries are generally present in the cephalic axis, but there are trilobites with one more pair of furrows, the so‐called S4, in the cephalic axis, causing confusion in understanding trilobite head segmentation. Recent advances in developmental biology and palaeontology have shed light on the arthropod head problem, and thus, trilobite head segmentation can be reviewed in the light of this knowledge. Based on the information from the anatomy of exceptionally preserved trilobites and artiopodans closely related to trilobites, it is inferred that trilobite head contains five segments: the anteriormost ocular segment potentially associated with the hypostome, the antennal segment and the following three segments with walking legs. When present, the S4 furrows are situated where the eye ridges meet the cephalic axis of trilobites, indicating that the furrows are incised ‘within’ the anteriormost segment in trilobites with an anteriorly enlarged frontal lobe. Trilobites of the Order Redlichiida, the most primitive stock, show variable conditions in the frontal glabellar conditions, while in other more derived groups, the condition is rather constant. The frontal glabellar condition, therefore, could provide a clue to elucidate the unresolved Cambrian trilobite phylogeny and the Cambrian roots of the post‐Cambrian trilobites.     

湘西晚寒武世多节类三叶虫新种及Erixanium属首次发现

文中描记 3个多节类三叶虫新种 :Hardyoidesdamaensissp .nov .,Meringaspisdamaensissp .nov .,andRhyssometopus (Rostrifinis)nitidussp .nov .,均产自湘西凤凰花桥组中部。同时描记首次在江南斜坡带中发现的Erixanium一属的保留命名种Erixaniumcf.E .sentum。Erixanium是全球性分布的多节类三叶虫 。     

Modelling enrolment in Cambrian trilobites

Trilobites were capable of enroling in different ways based on the flexible articulation of thoracic segments and associated interlocking devices; the type of enrolment (spiral or sphaeroidal) is thought to have largely depended on the coaptative devices that each trilobite used to enclose the body. Based on X‐ray microtomography scans of complete enrolled specimens from the Cambrian, we created three‐dimensional (3D) computer models to assess the kinematics needed to achieve both enrolment types. We demonstrate that closely related trilobites with little morphological variation (Bailiaspis?, Conocoryphe and Parabailiella) developed different enrolment types as a result of small variations in the number of thoracic segments and the angle between adjacent segments. Moreover, our models indicate that sphaeroidal enrolment, which is associated with a smaller number of thoracic segments, enabled faster encapsulation. This supports the hypothesis that there was a trend in the evolution of trilobites towards reduction in the number of thoracic segments in phylogenetically derived taxa in order to enhance the efficiency of enrolment.     

The Phylogeny and Systematics of Blind Cambrian Ptychoparioid Trilobites

The paraphyletic trilobite suborder Ptychopariina includes a large proportion of Cambrian trilobite diversity and is probably ancestral to most groups of post-Cambrian trilobites. Resolution of the phylogenetic relationships within the group is therefore crucial to a better understanding of the initial radiation of trilobites. The recognition of approaches that can successfully resolve the relationships of ptychoparioid taxa is an important first step towards this aim. Cladistic analysis was used to determine relationships within the Cambrian ptychoparioid trilobite family Conocoryphidae, and to test claims that the family is polyphyletic. Ninety-seven characters were coded for 40 conocoryphid species and nine non-conocoryphids. The results indicate that the family consists of four distantly related clades. Three are recognized here as distinct families, including an extensively revised Conocoryphidae, and the families Holocephalidae and Atopidae. The fourth clade is referred to the subfamily Acontheinae (Corynexochida) as the new Tribe Hartshillini. Analysis of the disparity of these four clades shows that they are significantly less morphologically variable than the original polyphyletic taxon, demonstrating the possible effects of taxonomic error on macroevolutionary studies of morphological disparity.     

Ontogeny of the Furongian (late Cambrian) trilobite Proceratopyge cf. P. Lata Whitehouse from northern Victoria Land,Antarctica, and the evolution of metamorphosis in trilobites

There were multiple origins of metamorphosis‐undergoing protaspides in trilobite evolution: within the superfamilies Remopleuridioidea, Trinucleoidea, and within the Order Asaphida. Recent studies have revealed that the protaspides of the Cambrian representatives of the Remopleuridioidea and the Trinucleoidea did not undergo metamorphosis. However, ontogeny of the Cambrian members of the Order Asaphida has remained unknown. This study documents the ontogeny of the Furongian asaphoidean ceratopygid trilobite, Proceratopyge cf. P. lata Whitehouse, from northern Victoria Land, Antarctica. Two stages for the protaspid phase, five developmental stages for the post‐protaspid cranidia, and ten stages for the post‐protaspid pygidia have been identified. Interestingly, the protaspis directly developed into a meraspis without metamorphosis. A new cladistic analysis resulted in a single most parsimonious tree, according to which the presence of the bulbous commutavi protaspis turns out to be a synapomorphy for Asaphidae + Cyclopygoidea, not a synapomorphy for the Order Asaphida as previously suggested. In addition, it is inferred that there was convergent evolution of indirectly‐developing commutavi protaspides during the Furongian and Early Ordovician. Metamorphosis‐entailing planktonic larvae evolved in many different metazoan lineages near the Cambrian–Ordovician transition, due to the escalating ecological pressure of the Great Ordovician Biodiversification Event. Since the bulbous commutavi protaspid morphology is thought to be an adaptation for a planktonic life mode, the convergent evolution of the indirect development in the three trilobite lineages at this period might have been a result of adaptation to the early phase of the Great Ordovician Biodiversification Event.     

Phylogenetic support for the monophyly of proetide trilobites

The monophyly of the order Proetida, the only trilobite group to survive the end‐Devonian mass extinction, has been regularly questioned since its erection almost three decades ago. Through analysis of a novel phylogenetic data set comprising 114 characters coded for 55 taxa, including both traditional members of the Proetida along with a number of other trilobite groups, the monophyly of proetide trilobites is rigorously tested for the first time. Proetida is shown to be monophyletic, united by the initial compound eye formation in early protaspids occurring at the lateral margin rather than the anterior margin, and the form of the protaspid glabella being tapering with a pre‐glabellar field. A number of adult characters, including the possession of a quadrate or shield‐shaped hypostome with angular posterior margins, the hypostome median body being divided by a deep groove that entirely traverses the median body, the presence of an enlarged thoracic spine on the sixth tergite and a tergite count of between 7 and 10, also define the basal node. Hystricurid and dimeropygoid trilobites are shown to resolve at the base of the group, while the remaining proetide taxa are divided between large proetoid and aulacopleuroid clades. Some taxa previously allied with Aulacopleuroidea, such as rorringtoniids and scharyiids, are retrieved as basal members of the Proetoidea.     

Ontogeny of the redlichiid trilobite Eoredlichia intermediata from the Chengjiang Lagerstätte,lower Cambrian,southwest China

A detailed investigation of the morphology and ontogeny of the redlichioid trilobite Eoredlichia intermediata (Bulletin of the Geological Society of China, 3–4, 1940, 333) from the lower Cambrian Yu'anshan Member of the Heilinpu Formation, in Kunming, Yunnan Province, southwest China, is presented. The new material comprises a relatively complete ontogenetic series ranging from the early meraspid to the holaspid period, which reveals more details on morphological variation such as the appearance of bacculae in some holaspid specimens, contraction and disappearance of fixigenal spines, and macropleural spines of the first and second thoracic segments, which are all documented for the first time and can also be used as developmental markers defining ontogenetic phases. Two distinct morphotypes, possibly an expression of intraspecific variation or sexual dimorphism, are distinguished by the morphology of pleural spines of the second thoracic segment in meraspid degree 14 and holaspides. The trunk segmentation schedule of E. intermediata is also discussed and conforms to the protarthrous developmental mode. The distinction of the thoracic region into two portions can be observed during late meraspid and early holaspid periods, which might be considered as a reference for a better understanding on the relationship of tagmosis and growth segmentation among the Cambrian redlichiid trilobites.     

The evolutionary history of crustacean segmentation: a fossil-based perspective

The evolution of segmentation in Crustacea, that is, the formation of sclerotized and jointed body somites and arrangement of somites into tagmata, is viewed in light of historical traits and functional constraints. The set of Early to Late Cambrian 'Orsten' arthropods have informed our current views of crustacean evolution considerably. These three-dimensionally preserved fossils document ancient morphologies, as opposed to purely hypothetical models and, because of the unusual preservation of larval stages, provide us with unparalleled insight into the morphogenesis of body somites and their structural equipment. The variety of evolutionary levels represented in the 'Orsten' including lobopodians, tardigrades, and pentastomids also allows phylogenetic interpretations far beyond the Crustacea. The 'Orsten' evidence and data from representatives of the Lower Cambrian Chengjiang biota in southwestern China, including phylogenetically earlier forms, form the major source of our morphology-based review of structural and functional developments that led toward the Crustacea. The principal strategy of arthropods is the simultaneous development of head somites, as expressed in a basal "head larva," and a successive addition of postcephalic somites from a preterminal budding zone with progressive maturation of metameric structures. This can be recognized in the developmental patterns of extant and fossil representatives of several euarthropod taxa, particularly crustaceans, trilobites, and chelicerates (at least basally). The development of these taxa points to an early somite-poor and free-living hatching stage. Embryonic development to a late stage within an egg, as occurring in recent onychophorans and certain in-group euarthropods, is regarded as achieved several times convergently.     

Trilobite larvae and larval ecology

Silicified trilobite faunas yield a well preserved and abundant record of early development in many taxa. Although their morphologies are well known, protaspid larvae are poorly understood in terms of trilobite life history and developmental biology. Two distinct morphologic types are recognized among the great diversity of protaspides studied; these are termed adult‐like and nonadult‐tike according to gross similarity to later developmental stages. Transitions between nonadult‐like and adult‐like morphologies are abrupt, occurring between successsive instars, and, thereby, constitute metamorphoses.

By analogy with developmental patterns among modern marine arthropod taxa, metamorphoses during early trilobite ontogeny correspond to radical modifications in life mode and ecology. Adult‐like trilobite protaspides possess a dorso‐ventrally flattened tergum which displays a coarsely featured prosopon and surface‐parallel spines; these are interpreted as benthic larvae. In contrast, nonadult‐like protaspid larvae display an ovoid to spheroidal shape with spine pairs that project at right angles to one another; these are considered to have been planktonic. Protaspid morphologies are compared and discussed in light of these inferred modes of life.     

Mansuyia Sun,and Tsinania Walcott,from the Furongian of North China and the evolution of the trilobite family Tsinaniidae

The Furongian trilobite family Tsinaniidae is characterized by a highly effaced surface and forms an important constituent of the Furongian trilobite faunas of east Gondwana. However, the origin of the characteristic morphology of this family has remained unclear. Only recently has the tsinaniid trilobite Lonchopygella megaspina been suggested to represent an intermediate stage in the evolutionary transition to other tsinaniids on the basis of the trunk segmentation. Here, we report successive occurrences of four species of the kaolishaniid genus Mansuyia and a tsinaniid trilobite Tsinania canens from the Furongian (late Cambrian) Chaomidian Formation in Shandong Province, China. A cladistic analysis including these taxa reveals that the four species of Mansuyia constitute stem‐group taxa to the family Tsinaniidae, rendering Mansuyia and the Kaolishaniidae paraphyletic. The youngest species of Mansuyia, M. taianfuensis, turns out to be the immediate sister taxon of the Tsinaniidae, displaying a closely similar morphology to the tsinaniid trilobite, Shergoldia laevigata. The generic and familial boundary therefore situated between M. taianfuensis and S. laevigata.     

Canalization of Gene Expression and Domain Shifts in the Drosophila Blastoderm by Dynamical Attractors

The variation in the expression patterns of the gap genes in the blastoderm of the fruit fly Drosophila melanogaster reduces over time as a result of cross regulation between these genes, a fact that we have demonstrated in an accompanying article in PLoS Biology (see Manu et al., doi:10.1371/journal.pbio.1000049). This biologically essential process is an example of the phenomenon known as canalization. It has been suggested that the developmental trajectory of a wild-type organism is inherently stable, and that canalization is a manifestation of this property. Although the role of gap genes in the canalization process was established by correctly predicting the response of the system to particular perturbations, the stability of the developmental trajectory remains to be investigated. For many years, it has been speculated that stability against perturbations during development can be described by dynamical systems having attracting sets that drive reductions of volume in phase space. In this paper, we show that both the reduction in variability of gap gene expression as well as shifts in the position of posterior gap gene domains are the result of the actions of attractors in the gap gene dynamical system. Two biologically distinct dynamical regions exist in the early embryo, separated by a bifurcation at 53% egg length. In the anterior region, reduction in variation occurs because of stability induced by point attractors, while in the posterior, the stability of the developmental trajectory arises from a one-dimensional attracting manifold. This manifold also controls a previously characterized anterior shift of posterior region gap domains. Our analysis shows that the complex phenomena of canalization and pattern formation in the Drosophila blastoderm can be understood in terms of the qualitative features of the dynamical system. The result confirms the idea that attractors are important for developmental stability and shows a richer variety of dynamical attractors in developmental systems than has been previously recognized.     

Cambrian biostratigraphy of the Bowers back-arc basin,Northern Victoria Land,Antarctica — A review

The Bowers Mountains in Northern Victoria Land contain the richest Cambrian Series 3 (Miaolingian, middle Cambrian) and Cambrian Series 4 (Furongian, late Cambrian) fossiliferous successions in Antarctica. Almost all the fossils are found within the Bowers Supergroup, which outcrops within the Bowers Terrane, a fault-bounded northwest-southeast oriented strip in Northern Victoria Land. The fossils provide the main age control on the history and evolution of the Bowers volcanic arc and back-arc basin. The great bulk of the fossils occur within the Spurs Formation. The fossil assemblages are dominated by agnostoids and polymerid trilobites with most ranging in age from Drumian to Paibian, although one fauna is of Jiangshanian age. Over 40 agnostoid taxa and over 100 polymerid trilobite taxa have been recorded from the rocks of the Bowers Supergroup. The youngest fauna occurs within the adjacent Robertson Bay Terrane, where a limited fauna of polymerid trilobites and conodonts from within a limestone olistolith have a very late Cambrian or early Ordovician age. Faunal affinities are mainly with Australia, New Zealand, North and South China and the Himalaya with lesser ties to Iran, Kazakhstan, Siberia and Laurentia.