Geodiversity,palaeodiversity or biodiversity: where is the place of palaeobiology and an understanding of taphonomy?

While some authors view biodiversity as an integral part of an over-arching geodiversity, most would see them as separate constructs. If that is the case, then the life–death interface marks the critical boundary between those that study organisms in life and the palaeontologists who try to interpret fossils in terms of their palaeobiology. The interface between life and the fossil record is characterised by the processes of taphonomy and, when fossils are collected, care must be taken to ensure that vital taphonomic information is not lost. This does not appear to be highlighted in the various fossil collecting codes and the avoidance of the loss of taphonomic information during collection should be a key feature of collecting policies and geoconservation in general.     

Taphofacies Models of Devonian in Longmenshan Area, Sichuan, China

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Was FLK North levels 1–2 a classic “living floor” of Oldowan hominins or a taphonomically complex palimpsest dominated by large carnivore feeding behavior?

From excavation at FLK North levels 1–2 in 1960–1962, Mary Leakey reported approximately 1200 Oldowan artifacts and 3300 large mammal fossils as a hominin “living floor”. Preliminary taphonomic analysis by Bunn seemed supportive, based on the presence of some cut-marked bones, the concentration of several dozen bovid individuals, and the relative abundance of limbs and mandibles over other axial elements. Recent taphonomic analysis of Leakey's entire fossil assemblage by Domínguez-Rodrigo and Barba, however, documents a minor hominin role at the site, contrasted to the dominant role of carnivores. Felids brought prey animals; hyenas scavenged from abandoned felid meals. At different times, hominins butchered several bovids and discarded artifacts at this dynamic location. Since 2006, renewed excavations at FLK North and other sites by the Olduvai Paleoanthropology and Paleoecology Project (TOPPP) have expanded artifact and fossil samples and implemented new analytical approaches to clarify taphonomic histories of the Olduvai paleolandscape. At FLK North, > 1000 new large mammal fossils from levels 1 to 2 show minimal butchery evidence amid abundant evidence of carnivore gnawing/fracture, rodent gnawing, and sediment abrasion. To help guide future excavation and analyses, we have developed several alternative working hypotheses of site formation.     

Taphonomic aspects of the Pleistocene vertebrate assemblage of Itaboraí, state of Rio de Janeiro,southeastern Brazil

Pleistocene vertebrates from Itaboraí Basin have not been taphonomically studied prior to this work, limiting the understanding of the deposition and preservation of the only Pleistocene vertebrate accumulation known for the state of Rio de Janeiro. In this work, the taphonomic signatures of the Pleistocene vertebrate assemblage of Itaboraí are identified and interpreted in order to increase the knowledge about the formation of this fossil association and the paleoecology of the region of Rio de Janeiro during the late Pleistocene. Our analysis shows that the thanatocoenosis was exposed to the biostratinomic processes during a small time span; that it is parautochthonous; and experienced short transport distances by normal fluvial streams and floods. Subsequently, the fossiliferous horizon was quickly covered by the superjacent soil. Yet, the skeletal elements were fractured and deformed during the sedimentary compaction. The differential preservation of megamammal bones is associated to the bone resistance against those destructive processes and to the specific anatomical features. Comparison between Itaboraí and other Brazilian Pleistocene vertebrate accumulations shows that the Itaboraí fossil accumulation was less affected by taphonomic processes, although it is also a time-averaged fossil concentration. Finally, some of the taphonomic features indicate an arid paleoclimate.     

Taphonomic and sedimentological aspects from PICOS II paleontological site,a quaternary pond deposit of Alagoas,Brazil

The Picos II Paleontological Site (PIIPS), located in the city of Piranhas, in Alagoas State, is a fossil deposit with a pond-like geomorphological feature, a kind of peculiar deposit for the Quaternary of Northeast Brazil. A detailed taphonomic study is presented about the PIIPS assemblage, from material collection to the laboratory analysis. The conservation state of the collected fossils in PIIPS is different from that found in other tank deposits, showing a high degree of specimen preservation. Two taxa of pleistocenic mammals have been identified: Eremotherium laurillardi and Notiomastodon platensis, and fragments of an avian bone, whose taxonomic identification was not possible. The taphonomic analysis indicates that the deposited material experienced a short transportation in high-energy event, with a short period of subaerial exposure before the final burial, where at least the final disarticulation occurred in situ, which explains the high conservation level of the material. The sedimentological analysis made in each layer of the deposit corroborates the information related to the transport of the material, which were obtained in taphonomic analysis. Regarding these aspects, PIIPS reflects nearly non-biased biocenotic patterns. The well-preserved material found in PIIPS demonstrates the great potential of deposits with pond-like geomorphological features, with sidelines that are less steep than traditional tanks, providing smoother transport and accommodation for skeletal elements, which is a distinguishing factor in preserving fossils of vertebrates.     

Systematic and taphonomic insights of fossilized feathers: A new occurrence from the Oligocene of Taubaté Basin (SE,Brazil)

Feathers are rare in the fossil record because they have a low fossilization potential. Despite their palaeobiological significance, they also provide important palaeoecological and taphonomic information. Here, we report a new occurrence of three isolated feathers from the shales of the Oligocene Tremembé Formation (Taubaté Basin, SE Brazil). Their possible taxonomic affinities and taphonomic features are also discussed. Analyses identified the specimens as representatives of two pennaceous morphotypes (i.e., a contour and a rectrice feather). Both are preserved as carbonized traces, although, due taphonomic processes, they show different degrees of preservation. Since the Tremembé Formation is responsible for the most diverse record of Cenozoic birds, and because water-adapted birds (e.g., anseriformes and phoenicopteriformes) occur in this unit, it is highly possible that these feathers belonged to these aquatic taxa. Further investigations should concentrate on geochemical and microscopic techniques in order to reveal additional taxonomic and paleoecological features currently unknown in Paleogene birds of Brazil.     

Holocene and Lateglacial summer temperature reconstruction in the Swiss Alps based on fossil assemblages of aquatic organisms: a review

The taxonomic composition of chironomid, cladoceran and diatom assemblages in small lakes in the Alpine region shows a strong relationship with summer temperature. Since fossils of all three organism groups preserve well and remain identifiable in lake sediments, summer temperature transfer-functions can be developed based on the modern distribution of these organisms and applied to fossil records to reconstruct past summer temperature variability. We provide a summary of the chironomid-, cladoceran- and diatom-based transfer functions available from the Swiss Alps and discuss the potential problem of co-variation between summer temperature and lake nutrient conditions for transfer-function development. Whereas the diatom-based summer temperature transfer function remains to be evaluated in down-core reconstructions, the cladoceran- and chironomid-based transfer functions have been used successfully to produce summer temperature records on Lateglacial and Holocene time scales that are in good agreement with other temperature reconstructions in the Alps. Major problems that can be encountered when using fossil assemblages of aquatic organisms for temperature reconstruction in the Alpine region are biases in the inferred temperatures associated with human impact on lakes and parameters other than temperature affecting the fossil assemblages. A multi-proxy approach to palaeoenvironmental reconstruction is recommended to keep a close control on past catchment and within-lake processes during the time interval of interest.     

Tube growth patterns and microbial mat-related lifestyles in the Ediacaran fossil Cloudina,Gaojiashan Lagerstätte,South China

Cloudina are important Ediacaran index fossils, as they have a consistent terminal Neoproterozoic age and a worldwide distribution. The morphology of Cloudina has been reconstructed primarily on the basis of three-dimensionally preserved Chinese and Spanish specimens, but questions remain surrounding their phylogenetic placement. Understanding Cloudina paleoecology is an important aspect for the interpretation of these enigmatic organisms. Although their paleoecological dynamics remain unresolved, a possible “mat sticker” life mode has found support from recent taphonomic investigation of Paraguayan Cloudina specimens. Here, from analysis of fossil material from the upper part of the Ediacaran Dengying Formation (Gaojiashan and Lijiagou sections, Ningqiang area, southern Shaanxi Province, South China), we document complex tube growth patterns of Cloudina in addition to a close association of these fossils with preserved microbially induced sedimentary structures (specifically matgrounds). Sedimentological and taphonomic data of Cloudina-bearing carbonate sequences at the Gaojiashan section show a relationship between Cloudina and microbial mats, and further document erectly preserved Cloudina tubes. In conjunction with observations from the Paraguay material, the Dengying fossils described here provide supportive evidence for the paleoecological interpretation of Cloudina as inhabitants of a microbially bound carbonate substrate. The apical-most part of the tube (or cone) may have served as an attachment apparatus, situated within or beneath a microbial mat. Three-dimensionally phosphatized tubes of Cloudina from the Lijiagou section reveal that single tubes often have multiple growth orientations, which is demonstrated to have resulted from the curved growth of the tube walls (both cones and funnels). As these organisms exhibited a sessile, epibenthic life mode, the observed plasticity in growth direction and orientation may have been a competition-driven adaptation to ensure wider access to available food sources and potentially greater feeding efficiency. Although Cloudina have been found to co-occur in some deposits with biostromes and bioherms (e.g., stromatolites and thrombolites), sedimentological and taphonomic features of the Ningqiang Cloudina suggest that they inhabited a stratified carbonate substrate exhibiting matground features, but where biostromes and bioherms were absent.     

There is no such thing as the ‘Ediacara Biota'

The term‘Ediacara Biota’(or many variants thereof)is commonly used to refer to certain megascopic fossils of Precambrian and early Palaeozoic age e but what does the term actually mean?What differentiates a non-Ediacaran‘Ediacaran’and an Ediacaran‘Ediacaran’from an Ediacaran non-‘Ediacaran’?Historically,the term has been used in either a geographic,stratigraphic,taphonomic,or biologic sense.More recent research and new discoveries,however,mean that the term cannot actually be defned on any of these bases,or any combination thereof.Indeed,the term is now used and understood in a manner which is internally inconsistent,and unintentionally implies that these fossils are somehow distinct from other fossil assemblages,which is simply not the case.Continued use of the term is a historical relic,which has led in part to incorrect assumptions that the‘Ediacara Biota’can be treated as a single coherent group,has obscured our understanding of the biological change over the PrecambrianeCambrian boundary,and has confused research on the early evolution of the Metazoa.In the future,the term‘Ediacaran’should be restricted to purely stratigraphic usage,regardless of affnity,geography,or taphonomy;suffcient terminology also exists where reference to specimens on a geographic,taphonomic,or biologic basis is required.It is therefore time to abandon the term‘Ediacara Biota’and to instead treat equally all of the fossils of the Ediacaran System.     

Is Darwin's ‘Abominable Mystery’ still a mystery today?

Darwin described the Cretaceous diversification and subsequent rapid rise of flowering plants (angiosperms) as an “abominable mystery”: how could they have achieved worldwide ecological dominance by early Paleogene times when the oldest angiosperm fossils are only Cretaceous in age? However, recent phylogenetic and palaeobiogeographical analyses have suggested a much earlier, perhaps Triassic origin for the stem angiosperms. We suggest that the fossil record is accurately reflecting the rapid diversification of the crown angiosperms that molecular data suggests occurred in Cretaceous–early Palaeogene times, and which coincides with similar explosive diversifications that occurred in other parts of the terrestrial biota including insects, birds and mammals. Early, stem angiosperms are poorly represented in the fossil record for a combination of taphonomic and ecological reasons. We conclude that the reported evidence shows that the Darwin's “mystery” is in fact no mystery and confirms that it merely requires an appropriate reading of the fossil record.     

Taphonomy of a Baurusuchus (Crocodyliformes,Baurusuchidae) from the Adamantina Formation (Upper Cretaceous,Bauru Basin), Brazil: Implications for preservational modes,time resolution and paleoecology

Upper Cretaceous vertebrate accumulations from the Adamantina Formation are known due to their high taxonomic diversity. On the other hand, taphonomic analyses still are rare, limiting the understanding of processes related to the biostratinomic and fossildiagenetic histories of this lithostratigraphic unit. In 2005, fossils were collected from an outcrop located at Jales municipality, state of São Paulo, southeastern Brazil. From this outcrop, a well-preserved Baurusuchus was recovered, which displays a peculiar set of taphonomic signatures. This paper identifies and interprets taphonomic features of a specimen of Baurusuchus (Crocodyliformes, Baurusuchidae; UFRJ DG 418-R) from the Adamantina Formation (Upper Cretaceous of the Bauru Basin), in Jales municipality, state of São Paulo. Brief taphonomic comparisons with other specimens previously studied (crocodiles and dinosaurs) and a lithofaciological analysis of the outcrop were undertaken in order to enhance the knowledge of the stratigraphy and paleoenvironment and improve the time resolution for the Adamantina Formation in the region of Jales. Furthermore, paleoecological data were interpreted based on the taphonomic analysis.The fossil is composed of an articulated segment of vertebral column, ribs, part of the pelvic girdle and gastralia. There is no hydraulic equivalence between both the specimen of Baurusuchus and the size of quartz grain predominant in the fossiliferous layer, suggesting death in situ or short transport as a “water carcass”. Teeth marks identified on the pubes were assigned to a small/juvenile baurusuchid crocodyliform or a theropod dinosaur. The repositioning of some elements (ribs and dorsal osteoderms) is suggestive of mummification. Desiccation marks were observed and attributed to the stage 1 of weathering. These features suggest subaerial exposure of the carcass prior to burial, however, probably after the mummification. On the other hand, the subaerial exposure was short, because the individual was not fully disarticulated. Furthermore, the degrees of articulation and preservation of the specimen nullify the hypothesis of reworking.Lithofaciological and taphonomic analyses suggest a well-drained floodplain as the burial environment and an arid or semi-arid climate in the region of Jales outcrop. In addition, the taphonomic signatures seem to indicate a time resolution about 10⁰–10⁴ years for the layer where the crocodyliforms were collected, but it seems to have, within this time limit, temporal-mixing among terrestrial crocodiles and dinosaurs collected from the same layer, suggesting time-averaging in this outcrop.     

Taphonomy and palaeoecology of the gastropod fauna from a Late Cretaceous rocky shore, Sweden

A gastropod fauna comprising 17 species, each represented by a limited number of specimens, is described from a Late Cretaceous, late early Campanian rocky shore at Ivö Klack, southern Sweden. The gastropod fauna is associated with the most diverse ancient rocky shore fauna ever found. However, the low gastropod species diversity compared to the faunas of modern rocky shores is ascribed to taphonomic factors, notably dissolution of the aragonitic shells, but the predominance of epifaunal herbivores is indicative of a guild structure similar to that found on modern rocky shores. The presence of drill holes assigned to the ichnospecies Oichnus simplex suggests the former presence of muricid gastropods which have not been found as body fossils. A single drill hole is referred to Oichnus paraboloides and was probably made by a naticid gastropod. The infaunal mode of life of naticids makes preservation of such drill holes difficult, since the majority of infaunal prey such as burrowing bivalves has aragonitic shells which are not preserved. The relatively high number of species in comparison to many other Late Cretaceous rocky shore faunas, offers an opportunity to compare gastropod guild structure at Ivö Klack with modern counterparts, even though taphonomic processes such as mechanical destruction and aragonite dissolution have played an important role in the fossil gastropod assemblage.     

Antiquity and long-term morphological stasis in a group of rove beetles (Coleoptera: Staphylinidae): Description of the oldest Octavius species from Cretaceous Burmese amber and a review of the “Euaesthetine subgroup” fossil record

The Staphylinine group of rove beetle subfamilies is a significant animal radiation, and one subordinate monophyletic clade – the ‘Euaesthetine subgroup’ – includes around 3000 species in subfamilies Euaesthetinae and Steninae and has a fossil record dating to the Early Cretaceous. Detailed morphological study of a new well-preserved Cretaceous Burmese amber fossil revealed strong evidence consistent with its taxonomic placement in the euaesthetine genus Octavius. We thus describe Octavius electrospinosus sp. nov., the first Cretaceous record of the genus and of the tribe Euaesthetini. Previously, the oldest records of Octavius and Euaesthetini were from the Eocene (Baltic amber) and discovery of O. electrospinosus sp. nov. therefore nearly doubles the minimum lineage age of Octavius, increasing it by 50 million years. We also briefly review the known Euaesthetine subgroup fossil record and tabulate summary data for all previously described fossils. All are placed in extant genera, and have visible diagnostic generic-level characters including some putative synapomorphies as judged by recent phylogenetic work. Including O. electrospinosus sp. nov., there are now four known Cretaceous species, all of which belong to either Octavius, Nordenskioldia, or Stenus. To explain the long-term morphological stasis in this group of rove beetles, we suggest that the continuous presence of mesic habitats may have buffered these lineages from strong selection for morphological change. Considering the fossils along with phylogenetic hypotheses we suggest the Euaesthetine subgroup originated in the Late Jurassic– Early Cretaceous and the Staphylinine group in the Early Jurassic. We emphasize the derived status of Cretaceous fossils in assessing possible divergence times and the significance of the pre-Cretaceous taphonomic bias for restricting more robust estimates. Further detailed morphological study of available fossils in a phylogenetic framework is badly needed to clarify the phylogenetic positions of these taxa.     

Are some fossils better than others for inferring palaeogeography?: The early Ordovician of the North Atlantic region as an example

Different kinds of fossils have different patterns of palaeogeographical distribution. This has led to differing palaeocontinental reconstructions in the early Palaeozoic. We take the early Ordovician of the North Atlantic region as a test case to investigate the influence of taxonomic group upon palaeogeographical interpretation. Using both single linkage duster analysis and parsimony methods of biogeographical analysis on fossil faunas and floras of various groups within this region it is shown that the patterns of similarity obtained, and their relationship to supposed oceanic separations, vary widely from one group to another. Some planktic fossils (graptolites, acritarchs) do not 'see' a separation between Gondwana and Baltica which is strongly supported by evidence from other groups. Trilobites and ostracodes, in particular, reveal patterns which conform well with other geological and geomagnetic evidence. At least in the Arenig-Uanvirn time period, geographical separation is seen as a more important control on faunal composition than was biofacies. Considering the 'better' fossil groups for biogeographical discrimination, different analytical methods produce similar biogeographical results.     

贵州关岭生物群海百合Traumatocrinus的再研究

在前人的研究基础上，通过对大量新发掘的贵州关岭地区晚三叠世海百合化石标本的再研究，肯定了Trauma-tocrinus的有效性。根据目前的材料，认为该属只有1个有效种，即Traumatocrinus hsui；而根据化石的埋藏学特征，认为幼年体Traumatocrinus hsui营假浮游生活，成年体是否能够适应底栖生活环境尚需进一步证实。     

浙江白垩纪恐龙化石

浙江白垩系朝川组、中戴组、金华组、赖家组和赤城山组等地层中,已发现了数以千计保存完好的蛋化石,而恐龙骨骼化石却很少而且大多很残破。其中,目前能鉴别的恐龙有暴龙类（tyrannosaurs）、伤齿龙类（troodontids）、盗蛋龙类（oviraptorids）、鸭嘴龙类（hadrosaurs）、慢龙类（segnosaurs）、泰坦巨龙类（titanosaurs）、甲龙类（ankylosaurs）和结节龙类（nodosaurs）等,分布在天台、永康、东阳、丽水、缙云、金华和江山等广大区域,说明当时浙江存在过种类繁多的恐龙动物群。根据化石埋藏和沉积特征,在白垩纪时期浙江有诸多气候炎热、四面环山的大小盆地,恐龙多沿湖泊、河流筑巢生息,季节性的暴雨导致山洪泛滥,淹没了恐龙巢穴,因泥沙的快速掩埋封闭,巢穴里的蛋形成化石后保存较好。而恐龙或逃离,或尸骨被洪水搬运散落,所以、迄今除了已发现的4具不完整的骨架化石外,绝大多数是零散的骨骼。     

Exceptionally preserved fossil assemblages through geologic time and space

Geologic deposits containing fossils with remains of non-biomineralized tissues (i.e. Konservat-Lagerstätten) provide key insights into ancient organisms and ecosystems. Such deposits are not evenly distributed through geologic time or space, suggesting that global phenomena play a key role in exceptional fossil preservation. Nonetheless, establishing the influence of global phenomena requires documenting temporal and spatial trends in occurrences of exceptionally preserved fossil assemblages. To this end, we compiled and analyzed a dataset of 694 globally distributed exceptional fossil assemblages spanning the history of complex eukaryotic life (~ 610 to 3 Ma). Our analyses demonstrate that assemblages with similar ages and depositional settings commonly occur in clusters, each signifying an ancient geographic region (up to hundreds of kilometers in scale), which repeatedly developed conditions conducive to soft tissue preservation. Using a novel hierarchical clustering approach, we show that these clusters decrease in number and shift from open marine to transitional and non-marine settings across the Cambrian-Ordovician interval. Conditions conducive to exceptional preservation declined worldwide during the early Paleozoic in response to transformations of near-surface environments that promoted degradation of tissues and curbed authigenic mineralization potential. We propose a holistic explanation relating these environmental transitions to ocean oxygenation and bioturbation, which affected virtually all taphonomic pathways, in addition to changes in seawater chemistry that disproportionately affected processes of soft tissue conservation. After these transitions, exceptional preservation rarely occurred in open marine settings, excepting times of widespread oceanic anoxia, when low oxygen levels set the stage. With these patterns, non-marine cluster count is correlated with non-marine rock quantity, and generally decreases with age. This result suggests that geologic processes, which progressively destroy terrestrial rocks over time, limit sampling of non-marine deposits on a global scale. Future efforts should aim to assess the impacts of such phenomena on evolutionary and ecological patterns in the fossil record.     

The effects of burrowing activity on archaeological sites: Ndondondwane,South Africa

Burrowing activity is a widely recognized source of site modification. Most taphonomic studies of burrowers emphasize their destructive aspects on the archaeological record. Excavations at Ndondondwane, South Africa, suggest burrowing activity is destructive in some ways, but may also preserve cultural behavior. Drawing on both direct and indirect sources of evidence, we discuss how burrowing activity by rodents, earthworms, and termites can inform about pedogenic and depositional processes at archaeological sites and both preserve and destroy evidence of intra‐settlement patterns and early African cultigens. Specifically, we demonstrate the limited effect of earthworms on site stratigraphy, how the localized activity of termites have preserved casts of early African cultigens, and how the ability of archaeologists to distinguish the devastating effects of rodent burrowing from remains of architectural features have permitted important inferences about social and ritual life in early African farming communities. © 2004 Wiley Periodicals, Inc.     

Preliminary small mammal taphonomy of FLK NW level 20 (Olduvai Gorge, Tanzania)

The Bed-I series of Olduvai Gorge (Tanzania) is a reference site in human evolution, having yielded the holotypes of Paranthropus boisei and Homo habilis, together with manufactured artefacts and abundant large and micro-fauna. Excavations in Olduvai Gorge have been recently resumed, with new aims and new results. This paper presents the results of the taphonomic analysis carried out on a fossil small-mammal assemblage recovered from FLK NW level 20, a layer overlying Tuff C, dated from 1.84 Ma. The analysis provides good evidence of a category 1 predator, most likely a barn owl, as the predator of the bone assemblage. Trampling and sediment compression might influence postdepositional breakage of the bones. This study is especially relevant since previous taphonomic analyses carried out at levels above and below this sample led to inconclusive results due to a low number of fossils (Fernández-Jalvo et al., 1998). The new sample provides new information to reconstruct the paleoenvironmental context in which early hominins inhabited.     

扬子板块西北缘下泥盆统平驿铺组双壳类遗迹化石

四川北部广元地区下泥盆统平驿铺组产出丰富的双壳类遗迹化石,但尚未开展过系统的遗迹学工作,也未进行过行为学与古生态学分析。文中针对四川广元马家剖面平驿铺组中部地层开展沉积学和系统遗迹学研究,在陆棚至近滨带下部沉积中共识别出3个属种的双壳类遗迹化石,包括Lockeia siliquaria,Protovirgularia rugosa,Ptychoplasma vagans,以及其他无脊椎动物门类遗迹化石Cruziana problematica,Dimorphichnus isp.,Lophoctenium isp.等。对双壳类遗迹化石的出现方式、形态以及保存特征的分析表明,平驿铺组中部曾出现过2种类型的双壳类,分属具分叉足(原鳃目)和楔状足的类群。原鳃目分布于过渡带和陆棚区,营沉积物摄食生活;受高能事件或沉积物掩埋影响,它们由停息状态变为“逃逸”状态的行为产生了Pro.rugosa。具楔状足的双壳类依生态类型可分为2类：在过渡带与陆棚区,漫游的、营沉积物摄食的类型产生了Pty.vagans;而在近滨下部,滤食性双壳类居群则在不同期次高能事件的间隙对沉积物进行大规模殖居,产生了L.sil...