The origin of echinoid shell beds in siliciclastic shelf environments: three examples from the Miocene of Sardinia,Italy

Three mass accumulations of sea urchins from the Miocene of Sardinia show a number of taphonomic features which set them apart from previously described echinoid assemblages from the Cenozoic in which they represent: (1) monotypic assemblages; (2) include very well‐preserved remains of either regular or spatangoid echinoids; and (3) originate in deeper water environments. These accumulations are compared using a detailed sedimentological and taphonomic analysis including preservational fabrics, taphonomic signatures, size frequency distributions, density of occurrences and preferred orientations. The possible role of gregarious behaviour contributing to mass occurrences and the specific sedimentary events leading to the excellent preservation are discussed. The interpreted depositional environment of all three deposits is that of a storm‐dominated, siliciclastic shelf environment. A phymosomatid assemblage represents rapid burial through obrution of a highly dense, freshly dead community. A Brissopsis‐dominated spatangoid assemblage represents a mixed accumulation of parautochthonous and transported skeletons. The third assemblage consisting of regular echinoid spines and rare tests represents a composite tempestite. Differences in the depositional environments are related to their position along onshore–offshore gradient with the first two beds originated in a deeper setting than that of the spine accumulation. This study shows that the preservation of assemblages containing complete regular echinoids and spatangoids is higher in deeper water settings than in shallow water environments.     

陕西西乡寒武纪幸运期宽川铺生物群网格状微体化石埋藏学与生物属性研究*

磷酸盐化保存是软躯体化石特异埋藏的一个重要途径, 而微生物在软躯体磷酸盐化过程中可能发挥了重要作用。前人通过埋藏学实验发现, 微生物会在动物胚胎等软躯体组织内部快速滋生, 充填生物体内部空间, 以微生物假形的方式复制了生物体的原始形态。但化石的磷酸盐化过程是否与埋藏学实验模拟的过程一致, 目前仍有争议。本次研究在寒武纪早期宽川铺生物群中发现了一类网格状微体化石。此类化石的保存状态可以根据其中丝状微生物滋生的程度分为三种类型, 它们展示了生物从死亡到微生物侵入、滋生, 最后被磷酸盐化的全过程。这些标本显示, 微生物假形在生物软组织磷酸盐化过程中扮演了重要角色, 但并不是化石磷酸盐化的必由之路, 尤其是当生物体具有矿化硬骨骼或者几丁质软骨骼等抗腐性较强的结构时。此类标本多以不完整保存的残片为主, 正反两面结构一致, 具有典型列状排列的近圆形与哑铃形网孔。由于化石结构简单, 生物学性状较少, 因此它们的亲缘关系尚不明确, 是一类需要继续研究的疑难化石。     

The tenth skeletal specimen of Archaeopteryx

We describe the tenth skeletal specimen of the Upper Jurassic Archaeopterygidae. The almost complete and well-preserved skeleton is assigned to Archaeopteryx siemensii Dames, 1897 and provides significant new information on the osteology of the Archaeopterygidae. As is evident from the new specimen, the palatine of Archaeopteryx was tetraradiate as in non-avian theropods, and not triradiate as in other avians. Also with respect to the position of the ectopterygoid, the data obtained from the new specimen lead to a revision of a previous reconstruction of the palate of Archaeopteryx . The morphology of the coracoid and that of the proximal tarsals is, for the first time, clearly visible in the new specimen. The new specimen demonstrates the presence of a hyperextendible second toe in Archaeopteryx . This feature is otherwise known only from the basal avian Rahonavis and deinonychosaurs (Dromaeosauridae and Troodontidae), and its presence in Archaeopteryx provides additional evidence for a close relationship between deinonychosaurs and avians. The new specimen also shows that the first toe of Archaeopteryx was not fully reversed but spread medially, supporting previous assumptions that Archaeopteryx was only facultatively arboreal. Finally, we comment on the taxonomic composition of the Archaeopterygidae and conclude that Archaeopteryx bavarica Wellnhofer, 1993 is likely to be a junior synonym of A. siemensii , and Wellnhoferia grandis Elżanowski, 2001 a junior synonym of A. lithographica von Meyer, 1861.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 97–116.     

Experimental analysis of soft‐tissue fossilization: opening the black box

Taphonomic experiments provide important insights into fossils that preserve the remains of decay‐prone soft tissues, tissues that are usually degraded and lost prior to fossilization. These fossils are among the most scientifically valuable evidence of ancient life on Earth, giving us a view into the past that is much less biased and incomplete than the picture provided by skeletal remains alone. Although the value of taphonomic experiments is beyond doubt, a lack of clarity regarding their purpose and limitations, and ambiguity in the use of terminology, are hampering progress. Here we distinguish between processes that promote information retention and those that promote information loss, in order to clarify the distinction between fossilization and preservation. Recognizing distinct processes of decay, mineralization and maturation, the sequence in which they act, and the potential for interactions, has important consequences for analysis of fossils, and for the design of taphonomic experiments. The purpose of well‐designed taphonomic experiments is generally to understand decay, maturation and preservation individually, thus limiting the number of variables involved. Much work remains to be done, but these methodologically reductionist foundations will allow researchers to build towards more complex taphonomic experiments and a more holistic understanding and analysis of the interactions between decay, maturation and preservation in the fossilization of non‐biomineralized remains. Our focus must remain on the key issue of understanding what exceptionally preserved fossils reveal about the history of biodiversity and evolution, rather than on debating the scope and value of an experimental approach.     

甘肃灵台上新世哺乳动物化石埋藏学

灵台任家坡动物群年龄组成以老年、幼年个体为主,显示动物正常死亡条件下的骨骼富集积累过程。由于动物死亡与掩埋之间时限的长短不同导致个体间风化痕迹、关节分解、破损程度不同。化石具有明显的定向性及分选性,经过季节性水流短距离搬运和较弱的分选作用。化石的埋藏特点与泛滥平原骨骼富集有较多的相似性,但沉积物组成差别很大,暗示中国红粘土物源的特殊性。     

Taphonomic field experiments and the role of the Shelf and Slope Experimental Taphonomy Initiative

Fossilization of organism remains is the result of biological, physical, and chemical processes at work in the environment of deposition. The pathways taken by the remains of organisms through death, decay, burial, and diagenesis (known as taphonomic pathways) affect the fossil record of life in important ways, from complete removal and recycling of remains to preservation of hard parts and sometimes to exceptional preservation of soft tissues. Our understanding of taphonomic processes informs our interpretations of the fossil record and helps in the reconstruction of ancient environments. Research into taphonomic pathways in modern environments provides important insight into the fossilization process. In 1993, a group of paleontologists formed the Shelf and Slope Experimental Taphonomy Initiative (SSETI) and began a long-term comparative study of taphofacies and differential decay of taxonomic groups by placing experimental arrays containing bivalves, gastropods, decapod crustaceans, sea urchins, and wood species at multiple marine shelf and slope sites in the Bahamas and Gulf of Mexico. The goal of SSETI is to observe taphonomic processes over a period of decades to better understand taphonomic pathways for these various groups of taxa in a wide variety of depositional environments beyond the shallow near shore region. This special volume brings together eight studies that are the result of the SSETI program. The focus of this contribution is to first review the body of published work concerned with the fossilization process through the experimental manipulation of the remains of organisms in modern marine environments. We also present the methods of the SSETI project and its contributions to our understanding of taphonomic processes in relation to carbonate and wood recycling and preservation.     

Estimation of adult skeletal age‐at‐death using the Sugeno fuzzy integral

Age‐at‐death estimation of an individual skeleton is important to forensic and biological anthropologists for identification and demographic analysis, but it has been shown that the current aging methods are often unreliable because of skeletal variation and taphonomic factors. Multifactorial methods have been shown to produce better results when determining age‐at‐death than single indicator methods. However, multifactorial methods are difficult to apply to single or poorly preserved skeletons, and they rarely provide the investigator with information about the reliability of the estimate. The goal of this research is to examine the validity of the Sugeno fuzzy integral as a multifactorial method for modeling age‐at‐death of an individual skeleton. This approach is novel because it produces an informed decision of age‐at‐death utilizing multiple age indicators while also taking into consideration the accuracies of the methods and the condition of the bone being examined. Additionally, the Sugeno fuzzy integral does not require the use of a population and it qualitatively produces easily interpreted graphical results. Examples are presented applying three commonly used aging methods on a known‐age skeletal sample from the Terry Anatomical Collection. This method produces results that are more accurate and with smaller intervals than single indicator methods. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Taphonomy of coral reefs: a review

This review summarises the major factors that affect the post-mortem history of skeletons in a coral reef environment. Skeletal material is traced from life, through death, breakdown, transport, burial and diagenesis to its final fossil form. The fact that most reef sediments are of skeletal composition poses problems of concentration or dilution of individual grain types in taphonomic analysis of reefs. Rates of supply of grains vary, not only with organism abundance and skeletal growth rates, but also with rates of physical and biological breakdown to transportable sediment. Physical and organic processes affect sedimentary structures and textures by mixing or segregating skeletal grains, though biogenic processes normally dominate in the protected setting of reef lagoons. The soft and hard substrates associated with reefs present different media for calcium carbonate accumulation and post-depositional disturbance, for example, loose sediments suffer bioturbation and rocks surfaces suffer bioerosion. The wide range of durability of skeletons and their susceptibility to diagenesis contribute further to the complexities of the preservation of coral reefs.     

Proportions and variability in the Indian lorises: Bone weight observations onLoris tardigradus lydekkerianus andNycticebus coucang

Thirty uniformly prepared, disarticulated complete skeletons of adultLoris tardigradus lydekkerianus andNycticebus coucang have been weighed for preliminary study of the weight relations between different major skeletal parts. In these forms, combined relative weight of the precaudal vertebrae, ribs and sternum surpasses the averages for the rest of the skeletal parts. The weight of the skull equals one-fifth of the total skeletal weight inLoris andNycticebus. The weight of the skeletons of the limbs (including the shoulder girdle and the hip bones) contributes less than half of the total skeletal weight in lorises. The bones of the upper limbs are lighter than those of the lower limbs in the 2 series examined.Loris andNycticebus are distinguished by having the proportionately heaviest axial skeleton and skull and the lowest relative weights of the bones of the 4 limbs. The combined relative skeletal weight of the upper and lower limbs ranges between 45.34 and 50.01 inLoris andNycticebus. The relative weight of the skeletons of the hands and feet are almost similar to the corresponding weights in the 2 series.As far as the weights of the different skeletal parts are concerned, there is no relative asymmetry in the 2 series ofLoris andNycticebus examined.     

Precambrian animal life: Taphonomy of phosphatized metazoan embryos from southwest China

Phosphatized fossils from the Neoproterozoic Doushantuo Formation have provided valuable insight into the early evolution of metazoans, but the preservation of these spectacular fossils is not yet fully understood. This research begins to address this issue by performing a detailed specimen-based taphonomic analysis of the Doushantuo Formation phosphatized metazoan embryos. A total of 206 embryos in 65 thin sections from the Weng'an Phosphorite Member of the Doushantuo Formation were examined and their levels of pre-phosphatization decay estimated. The data produced from this examination reveal a strong taphonomic bias toward earlier (2-cell and 4-cell) cleavage stages, which tend to be well-preserved, and away from later (8-cell and 16-cell) cleavage stages, which tend to exhibit evidence for slight to intense levels of organic decay. In addition, the natural abundances of these embryos tend to decrease with advancement in cleavage stage, and no evidence of more advanced (beyond 16-cell) cleavage stages or eventual adult forms were found in this study. One possible explanation for this taphonomic bias toward early cleavage stages is that later cleavage stages and adult forms were more physically delicate, allowing them to be more easily damaged during burial and reworking, allowing for more rapid decay. The spectacular preservation of these embryos was probably aided by their likely internal enrichment in phosphate-rich yolk, which would have caused their internal dissolved phosphate levels to reach critical levels with only miniscule organic decay, thereby hastening phosphatization. If internal sources of phosphate did indeed play a role in the phosphatization of these embryos, it may explain their prolific abundance in these rocks compared to other phosphatized fossils as well as indicating that metazoans lacking such internal phosphate sources were likely much more difficult to preserve. The phosphatic fossils of the Doushantuo Formation, therefore, provide an indispensable, yet restricted, window into Neoproterozoic life and metazoan origins.     

Dental health and lifestyle in four early mediaeval juvenile populations: Comparisons between urban and rural individuals, and between coastal and inland settlements

Dental stress markers such as enamel hypoplasia and caries are suitable indicators of population health and lifestyle, although they must be recorded and interpreted carefully. To date, they have been predominantly studied in adult samples, whereas juvenile remains are also affected by these lesions. In this study, dental enamel hypoplasia and caries were both evaluated on 613 non-adult individuals from four early mediaeval Moravian and Frankish skeletal series, who had experienced contrasting environments and lifestyles. The aim of this study is to assess the relationship between these biological traits and living conditions, and how this is manifested in the juvenile dental remains.Significant differences between populations were found in stress markers, dental lesions and the way these were manifested. Exposure to stressful conditions varies between urban and rural populations and is related to age groups. Although the children under investigation seem to have had different diets, it is difficult to distinguish the biological contribution (different enamel susceptibility) from the lifestyle contribution (different food, environment) in the formation of caries. Moreover, such studies must be interpreted carefully due to the possibility of intra- and inter-observer errors and the subjectivity of the scoring techniques.Nevertheless, this study also demonstrates that results of an investigation of juvenile skeletal remains can be as informative as a study of adults and that juvenile skeletons can be included in large bioarchaeological population studies.     

Methods for estimating missing human skeletal element osteometric dimensions employed in the revised fully technique for estimating stature

One of the greatest limitations to the application of the revised Fully anatomical stature estimation method is the inability to measure some of the skeletal elements required in its calculation. These element dimensions cannot be obtained due to taphonomic factors, incomplete excavation, or disease processes, and result in missing data. This study examines methods of imputing these missing dimensions using observable Fully measurements from the skeleton and the accuracy of incorporating these missing element estimations into anatomical stature reconstruction. These are further assessed against stature estimations obtained from mathematical regression formulae for the lower limb bones (femur and tibia). Two thousand seven hundred and seventeen North and South American indigenous skeletons were measured, and subsets of these with observable Fully dimensions were used to simulate missing elements and create estimation methods and equations. Comparisons were made directly between anatomically reconstructed statures and mathematically derived statures, as well as with anatomically derived statures with imputed missing dimensions. These analyses demonstrate that, while mathematical stature estimations are more accurate, anatomical statures incorporating missing dimensions are not appreciably less accurate and are more precise. The anatomical stature estimation method using imputed missing dimensions is supported. Missing element estimation, however, is limited to the vertebral column (only when lumbar vertebrae are present) and to talocalcaneal height (only when femora and tibiae are present). Crania, entire vertebral columns, and femoral or tibial lengths cannot be reliably estimated. Further discussion of the applicability of these methods is discussed.     

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.     

Ancestral patterns in bird limb development: A new look at Hampé's experiment

The “Archaeopteryx limb” of experimentally treated bird embryos has become a standard quotation in the growing literature on developmental factors in evolution. It has not only been claimed that an early manipulation of the chick limb produces a series of atavistic skeletal features, but the experiment is also frequently interpreted within a genome-centered concept of atavism. The present study provides a morphological, quantitative, and comparative analysis of the skeletal and muscular reactions to the classic barrier insertion experiment of Hampé. The main result of this operation, traditionally seen as a “full length fibula”, is shown to be a relative effect due to tibia shortening, while all the other ancestral skeletal features, which are usually pointed out as being provoked by the elongated fibula, do not appear. The experimentally generated fibula/tibia length ratio and distance, however, mimic the pattern in developing reptilian limbs and are seen to induce secondary effects in the muscular system that are reminiscent of archosaur reptiles. Similar muscle patterns are found as interspecific variations in several bird species. The revised view of the skeletal changes and the additional data on muscular effects allow for a renewed interpretation of the experiment, shifting the emphasis from atavisms to the role of heterochrony, developmental integration, and epigenetic constraint in the evolutionary modification of organismic structures.     

Preservational modes of some ichthyosaur soft tissues (Reptilia,Ichthyopterygia) from the Jurassic Posidonia Shale of Germany

Konservat-Lagerstätten, such as the Toarcian (Early Jurassic) Posidonia Shale of southwestern Germany, are renowned for their spectacular fossils. Ichthyosaur skeletons recovered from this formation are frequently associated with soft tissues; however, the preserved material ranges from three-dimensional, predominantly phosphatized structures to dark films of mainly organic matter. We examined soft-tissue residues obtained from two ichthyosaur specimens using an integrated ultrastructural and geochemical approach. Our analyses revealed that the superficially-looking ‘films’ in fact comprise sections of densely aggregated melanosome (pigment) organelles sandwiched between phosphatized layers containing fibrous microstructures. We interpret this distinct layering as representing condensed and incompletely degraded integument from both sides of the animal. When compared against previously documented ichthyosaur fossils, it becomes readily apparent that a range of preservational modes exists between presumed ‘phosphatic’ and ‘carbonized’ soft-tissue remains. Some specimens show high structural fidelity (e.g. distinct integumentary layering), while others, including the fossils examined in this study, retain few original anatomical details. This diversity of soft-tissue preservational modes among Posidonia Shale ichthyosaurs offers a unique opportunity to examine different biostratinomic, taphonomic and diagenetic variables that potentially could affect the process of fossilization. It is likely that soft-tissue preservation in the Posidonia Shale was regulated by a multitude of factors, including decay efficiency and speed of phosphatic mineral nucleation; these in turn were governed by a seafloor with sustained microbial mat activity fuelled by high organic matter input and seasonally fluctuating oxygen levels.     

Biomineralization in bryozoans: present,past and future

Many animal phyla have the physiological ability to produce biomineralized skeletons with functional roles that have been shaped by natural selection for more than 500 million years. Among these are bryozoans, a moderately diverse phylum of aquatic invertebrates with a rich fossil record and importance today as bioconstructors in some shallow‐water marine habitats. Biomineralizational patterns and, especially, processes are poorly understood in bryozoans but are conventionally believed to be similar to those of the related lophotrochozoan phyla Brachiopoda and Mollusca. However, bryozoan skeletons are more intricate than those of these two phyla. Calcareous skeletons have been acquired independently in two bryozoan clades – Stenolaemata in the Ordovician and Cheilostomata in the Jurassic – providing an evolutionary replicate. This review aims to highlight the importance of biomineralization in bryozoans and focuses on their skeletal ultrastructures, mineralogy and chemistry, the roles of organic components, the evolutionary history of bimineralization in bryozoans with respect to changes in seawater chemistry, and the impact of contemporary global changes, especially ocean acidification, on bryozoan skeletons. Bryozoan skeletons are constructed from three different wall types (exterior, interior and compound) differing in the presence/absence and location of organic cuticular layers. Skeletal ultrastructures can be classified into wall‐parallel (i.e. laminated) and wall‐perpendicular (i.e. prismatic) fabrics, the latter apparently found in only one of the two biomineralizing clades (Cheilostomata), which is also the only clade to biomineralize aragonite. A plethora of ultrastructural fabrics can be recognized and most occur in combination with other fabrics to constitute a fabric suite. The proportion of aragonitic and bimineralic bryozoans, as well as the Mg content of bryozoan skeletons, show a latitudinal increase into the warmer waters of the tropics. Responses of bryozoan mineralogy and skeletal thickness to oscillations between calcite and aragonite seas through geological time are equivocal. Field and laboratory studies of living bryozoans have shown that predicted future changes in pH (ocean acidification) combined with global warming are likely to have detrimental effects on calcification, growth rate and production of polymorphic zooids for defence and reproduction, although some species exhibit reasonable levels of resilience. Some key questions about bryozoan biomineralization that need to be addressed are identified.     

On the nature and causes of luminescent lines and bands in coral skeletons: II. Contribution of skeletal crystals

Slices cut from skeletons of massive Porites display two types of luminescence when illuminated by ultra-violet (UV) light: (1) faint luminescent banding associated with the annual skeletal density banding pattern and (2) narrow lines of strong luminescence associated with monsoonal runoff of fresh water from nearby land. Barnes and Taylor [Barnes, D.J. Taylor, R.B. 2001a. On the nature and causes of luminescent lines and bands in coral skeletons. Coral Reefs 19, 221-230] showed how larger skeletal holes could give rise to increased luminescence—thus accounting for the link between skeletal density banding and faint luminescent banding. Work described here tests the notion that strongly luminescent lines are also regions of lower skeletal density. Experiments involving real and artificial coral skeletons indicated that likely changes in hole size in real skeletons cannot account for the amount of luminescence associated with luminescent lines. Larger particles (< 50 μm) of powdered skeleton from cut from luminescent lines were more luminescent than similar particles cut from adjacent less luminescent skeleton. However, very small particles (< 3 μm) from the two regions of skeleton showed no difference in luminescence. Since skeletal crystals would have been largely destroyed by powdering skeleton to very small particle sizes, most of the luminescence of strongly luminescent lines is probably associated with changed crystal size and packing, with changed crystal chemistry, or with a combination of these possibilities.     

Likelihood reinstates Archaeopteryx as a primitive bird

The widespread view that Archaeopteryx was a primitive (basal) bird has been recently challenged by a comprehensive phylogenetic analysis that placed Archaeopteryx with deinonychosaurian theropods. The new phylogeny suggested that typical bird flight (powered by the front limbs only) either evolved at least twice, or was lost/modified in some deinonychosaurs. However, this parsimony-based result was acknowledged to be weakly supported. Maximum-likelihood and related Bayesian methods applied to the same dataset yield a different and more orthodox result: Archaeopteryx is restored as a basal bird with bootstrap frequency of 73 per cent and posterior probability of 1. These results are consistent with a single origin of typical (forelimb-powered) bird flight. The Archaeopteryx-deinonychosaur clade retrieved by parsimony is supported by more characters (which are on average more homoplasious), whereas the Archaeopteryx-bird clade retrieved by likelihood-based methods is supported by fewer characters (but on average less homoplasious). Both positions for Archaeopteryx remain plausible, highlighting the hazy boundary between birds and advanced theropods. These results also suggest that likelihood-based methods (in addition to parsimony) can be useful in morphological phylogenetics.     

Atlas of vertebrate decay: a visual and taphonomic guide to fossil interpretation

Like many other important evolutionary transitions, our knowledge of the origin of vertebrates is limited to windows of exceptional preservation of soft‐bodied fossils. Unfortunately, these fossils are rare and have been subjected to complex taphonomic filters including decay, collapse and distortion. To maximize our ability to utilize these crucial fossils to reconstruct the timing and sequence of evolutionary events, we are in the need of a robust taphonomic framework with in which to interpret them. Here, we report the results of a series of experiments designed to examine patterns of transformation and loss during decay of important anatomical characters of chordates and primitive vertebrates (ammocoete, adult lamprey, hagfish, juvenile chondrichthyans and a non‐vertebrate chordate, Branchiostoma). Complex and repeated patterns of transformation during decay are identified and figured for informative character complexes including eyes, feeding apparatus, skull and brain, muscles, branchial apparatus, axial structures, viscera, heart and fins. The resulting data regarding character decay and relative loss serve as a guide to recognition and interpretation of the anatomy of non‐biomineralized fossil vertebrates. The methods and techniques outlined are eminently applicable to other soft‐bodied groups and present a new way to interpret the exceptionally preserved fossil record.