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The Eocene Geiseltal Konservat-Lagerstätte (Germany) is famous for reports of three dimensionally preserved soft tissues with sub-cellular detail. The proposed mode of preservation, direct replication in silica, is not known in other fossils and has not been verified using modern approaches. Here, we investigated the taphonomy of the Geiseltal anurans using diverse microbeam imaging and chemical analytical techniques. Our analyses confirm the preservation of soft tissues in all body regions but fail to yield evidence for silicified soft tissues. Instead, the anuran soft tissues are preserved as two layers that differ in microstructure and composition. Layer 1 comprises sulfur-rich carbonaceous microbodies interpreted as melanosomes. Layer 2 comprises the mid-dermal Eberth–Katschenko layer, preserved in calcium phosphate. In addition, patches of original aragonite crystals define the former position of the endolymphatic sac. The primary modes of soft tissue preservation are therefore sulfurization of melanosomes and phosphatization of more labile soft tissues, i.e., skin. This is consistent with the taphonomy of vertebrates in many other Konservat-Lagerstätten. These findings emphasize an emerging model for pervasive preservation of vertebrate soft tissues via melanosome films, particularly in stagnation-type deposits, with phosphatization of more labile tissues where tissue biochemistry is favorable. [ABSTRACT FROM AUTHOR]

Melanin pigments play a critical role in physiological processes and shaping animal behaviour. Fossil melanin is a unique resource for understanding the functional evolution of melanin but the impact of fossilisation on molecular signatures for eumelanin and, especially, phaeomelanin is not fully understood. Here we present a model for the chemical taphonomy of fossil eumelanin and phaeomelanin based on thermal maturation experiments using feathers from extant birds. Our results reveal which molecular signatures are authentic signals for thermally matured eumelanin and phaeomelanin, which signatures are artefacts derived from the maturation of non-melanin molecules, and how these chemical data are impacted by sample preparation. Our model correctly predicts the molecular composition of eumelanins in diverse vertebrate fossils from the Miocene and Cretaceous and, critically, identifies direct molecular evidence for phaeomelanin in these fossils. This taphonomic framework adds to the geochemical toolbox that underpins reconstructions of melanin evolution and of melanin-based coloration in fossil vertebrates. Reconstructing original fossil colour provides insights into the behaviour of ancient animals but is challenging because phaeomelanin pigments have a poor fossil record. Here, the authors present experimental data that predict the composition of fossil melanins and support the molecular preservation of phaeomelanin in 10 million year old frogs. [ABSTRACT FROM AUTHOR]

Many fossil insects show monochromatic colour patterns that may provide valuable insights into ancient insect behaviour and ecology. Whether these patterns reflect original pigmentary coloration is, however, unknown, and their formation mechanism has not been investigated. Here, we performed thermal maturation experiments on extant beetles with melanin-based colour patterns. Scanning electron microscopy reveals that melanin-rich cuticle is more resistant to degradation than melanin-poor cuticle: with progressive maturation, melanin-poor cuticle regions experience preferential thinning and loss, yet melanin-rich cuticle remains. Comparative analysis of fossil insects with monotonal colour patterns confirms that the variations in tone correspond to variations in preserved cuticle thickness. These preserved colour patterns can thus be plausibly explained as melanin-based patterning. Recognition of melanin-based colour patterns in fossil insects opens new avenues for interpreting the evolution of insect coloration and behaviour through deep time. [ABSTRACT FROM AUTHOR]

The Geiseltal biota is an Eocene lacustrine Konservat‐Lagerstätte in central Germany. Despite its rich fauna and flora (over 50 000 fossil vertebrates, insects and other invertebrates, plants and trace fossils) the taphonomy of the biota, and of the anurans in particular, is poorly understood. We analysed the skeletal taphonomy of 168 anurans, scoring each specimen for orientation, completeness and articulation. Most are partial skeletons, truncated during preparation. This process introduces measurable artefacts into the taphonomic data, which requires further study. We identified recurring taphonomic states, including disarticulation or loss of abdominal bones, loss of one or more limbs, extensive disarticulation, and extensive loss of elements across the body. Relatively rare states include: loss of distal limb elements only, scattered bones, and isolated, articulated limbs. These data inform a new taphonomic model for the Geiseltal anurans. Comparative analysis of patterns in completeness and articulation in these specimens and anurans from other Cenozoic Lagerstätten reveals repeated patterns in preservation in lacustrine‐hosted settings, such as proximal to distal trends in completeness, preferential loss of small peripheral bones, disarticulation of abdominal bones, and reorientation of major body regions. Taphonomic controls in lacustrine systems are surface water temperature, lake depth, vegetation, exposure to bottom currents, transport mechanisms, scavenging, decay regime, bone size and location in the skeleton; all ultimately controlled by palaeoclimate, lake physiography and hydrology, and anuran anatomy. A universal approach to quantitative analysis of skeletal taphonomy metrics will enable comparative testing of the relationship between preservation and palaeoenvironment for diverse fossil vertebrates. [ABSTRACT FROM AUTHOR]

A key feature of the pigment melanin is its high binding affinity for trace metal ions. In modern vertebrates trace metals associated with melanosomes, melanin‐rich organelles, can show tissue‐specific and taxon‐specific distribution patterns. Such signals preserve in fossil melanosomes, informing on the anatomy and phylogenetic affinities of fossil vertebrates. Fossil and modern melanosomes, however, often differ in trace metal chemistry; in particular, melanosomes from fossil vertebrate eyes are depleted in Zn and enriched in Cu relative to their extant counterparts. Whether these chemical differences are biological or taphonomic in origin is unknown, limiting our ability to use melanosome trace metal chemistry to test palaeobiological hypotheses. Here, we use maturation experiments on eye melanosomes from extant vertebrates and synchrotron rapid scan‐x‐ray fluorescence analysis to show that thermal maturation can dramatically alter melanosome trace element chemistry. In particular, maturation of melanosomes in Cu‐rich solutions results in significant depletion of Zn, probably due to low pH and competition effects with Cu. These results confirm fossil melanosome chemistry is susceptible to alteration due to variations in local chemical conditions during diagenesis. Maturation experiments can provide essential data on melanosome chemical taphonomy required for accurate interpretations of preserved chemical signatures in fossils. [ABSTRACT FROM AUTHOR]

Recent discoveries of nonintegumentary melanosomes in extant and fossil amphibians offer potential insights into the physiological functions of melanin not directly related to color production, but the phylogenetic distribution and evolutionary history of these internal melanosomes has not been characterized systematically. Here, we present a holistic method to discriminate among melanized tissues by analyzing the anatomical distribution, morphology, and chemistry of melanosomes in various tissues in a phylogenetically broad sample of extant and fossil vertebrates. Our results show that internal melanosomes in all extant vertebrates analyzed have tissue-specific geometries and elemental signatures. Similar distinct populations of preservedmelanosomes in phylogenetically diverse vertebrate fossils often map onto specific anatomical features. This approach also reveals the presence of various melanosome-rich internal tissues in fossils, providing a mechanism for the interpretation of the internal anatomy of ancient vertebrates. Collectively, these data indicate that vertebratemelanins share fundamental physiological roles in homeostasis via the scavenging and sequestering of metals and suggest that intimate links between melanin and metal metabolism in vertebrates have deep evolutionary origins. [ABSTRACT FROM AUTHOR]

Abstract: 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Colouration is an important multifunctional attribute of modern animals, but its evolutionary history is poorly resolved, in part because of our limited ability to recognize and interpret fossil evidence of colour. Recent studies on structural and pigmentary colours in fossil insects and feathers have illuminated important aspects of the anatomy, taphonomy, evolution and function of colour in these fossils. An understanding of the taphonomic factors that control the preservation of colour is key to assessing the fidelity with which original colours are preserved and can constrain interpretations of the visual appearance of fossil insects and theropods. Various analytical approaches can identify anatomical and chemical evidence of colour in fossils; experimental taphonomic studies inform on how colour alters during diagenesis. Preservation of colour is controlled by a suite of factors, the most important of which relate to the diagenetic history of the host sediment, that is, maximum burial temperatures and fluid flow, and subsurface weathering. Future studies focussing on key morphological and chemical aspects of colour preservation relating to cuticular pigments in insects and keratinous structures and nonmelanin pigments in feathers, for example, will resolve outstanding questions regarding the taphonomy of colour and will enhance our ability to infer original colouration and its functions in fossil insects and theropods. [ABSTRACT FROM AUTHOR]

McNamara, M.E., Orr, P.J., Manzocchi, T., Alcalá, L., Anadón, P. & Peñalver, E. 2011: Biological controls upon the physical taphonomy of exceptionally preserved salamanders from the Miocene of Rubielos de Mora, northeast Spain. Lethaia, Vol. 45, pp. 210-226. The middle Miocene Rubielos de Mora Konservat-Lagerstätte of northeast Spain is hosted within profundal, finely laminated, lacustrine mudstones. The diverse biota includes abundant salamanders. Most individuals died during separate episodes and sank rapidly postmortem. Specimens are typically preserved in dorso-ventral aspect, the most hydrodynamically stable orientation. The near-cylindrical morphology of the body, however, allowed some carcasses to settle in or subsequently re-orientate into, lateral orientations. Loss of skeletal elements (i.e. reduced completeness) reflects their location within the body and followed a distal to proximal trend. Two stages are identified: initial loss of a small number of phalanges, followed by loss of more proximal limb bones plus additional phalanges. Disarticulation is more complex: it occurred via several mechanisms (notably, abdominal rupture and re-orientation of part of the body and limbs during decay) and shows no consistent pattern among specimens. The physical taphonomy of the salamanders is controlled predominantly by intrinsic biological factors, i.e. the geometry of the body and of individual skeletal elements, the orientation, inherent strength and location of specific joints and the extent to which soft tissues, particularly the skin, persist during decay. These biological factors probably control patterns of physical taphonomy of other fossil tetrapods with a similar skeletal configuration. □ Articulation, completeness, Konservat-Lagerstätten, orientation, quantitative taphonomy, salamanders. [ABSTRACT FROM AUTHOR]

McNamara, M.E., Orr, P.J., Kearns, S.L., Alcalá, L., Anadón, P. & Peñalver-Mollá, E. 2010: Exceptionally preserved tadpoles from the Miocene of Libros, Spain: ecomorphological reconstruction and the impact of ontogeny upon taphonomy. Lethaia, Vol. 43, pp. 290–306. The Libros exceptional biota from the Upper Miocene of NE Spain includes abundant frog tadpoles ( Rana pueyoi) preserved in finely laminated lacustrine mudstones. The tadpoles exhibit a depressed body, short tail, low tail fins, dorso-laterally directed eyes and jaw sheaths; these features identify the Libros tadpoles as members of the benthic lentic ecomorphological guild. This, the first ecomorphological reconstruction of a fossil tadpole, supports phylogenetic evidence that this ecology is a conserved ranid feature. The soft-tissue features of the Libros tadpoles are characterized by several modes of preservation. The space occupied previously by the brain is defined by calcium carbonate, the nerve cord is defined by calcium phosphate, and jaw sheaths and bone marrow are preserved as organic remains. Gut contents (and coprolites adjacent to specimens) comprise ingested fine-grained sedimentary detritus and epiphyton. The body outline and the eyespots, nares, abdominal cavity, notochord, caudal myotomes and fins are defined by a carbonaceous bacterial biofilm. A similar biofilm in adult specimens of R. pueyoi from Libros defines only the body outline, not any internal anatomical features. In the adult frogs, but not in the tadpoles, calcium phosphate and calcium sulphate precipitated in association with integumentary tissues. These differences in the mode of preservation between the adult frogs and tadpoles reflect ontogenetic factors. □ Anuran, ecology, soft-tissue, tadpoles, taphonomy. [ABSTRACT FROM AUTHOR]

Bone marrow in ca. 10 Ma frogs and salamanders from the Miocene of Libros, Spain, represents the first fossilized example of this extremely decay-prone tissue. The bone marrow, preserved in three dimensions as an organic residue, retains the original texture and red and yellow color of hematopoietic and fatty marrow, respectively; moldic osteoclasts and vascular structures are also present. We attribute exceptional preservation of the fossilized bone marrow to cryptic preservation: the bones of the amphibians formed protective microenvironments, and inhibited microbial infiltration. Specimens in which bone marrow is preserved vary in their completeness and articulation and in the extent to which the body outline is preserved as a thin film of organically preserved bacteria. Cryptic preservation of these labile tissues is thus to a large extent independent of, and cannot be predicted by, the taphonomic history of the remainder of the specimen. [ABSTRACT FROM AUTHOR]

Abstract The Middle–Upper Jurassic Yanliao Lagerstätte contains numerous exceptionally preserved fossils of aquatic and land organisms, including insects, salamanders, dinosaurs, pterosaurs and mammaliaforms. Despite extensive study of the diversity and evolutionary implications of the biota, the palaeoenvironmental setting and taphonomy of the fossils remain poorly understood. We reconstruct both the palaeoenvironment of the Daohugou area (one of the most famous Yanliao fossil areas), and the biostratinomy of the fossils. We use high-resolution stratigraphic data from field investigation and excavations to document in detail the stratigraphic succession, lithofacies, facies associations, and biostratinomic features of the Lagerstätte. Our results show that frequent volcanic eruptions generated an extensive volcaniclastic apron and lake(s) in the studied area. The frequent alternation of thin lacustrine deposits and thick volcaniclastic apron deposits indicates either that the studied area was located in the marginal regions of a single lake, where the frequent influx of volcaniclastic apron material caused substantial fluctuations in lake area and thus the frequent lateral alternation of the two facies, or that many short-lived lakes developed on the volcaniclastic apron. Most terrestrial insects were preserved in the laminated, normally graded siltstone, claystone and tuff facies that form many thin intervals with deposits of graded sandstone, siltstone and tuff in between. Within each interval the terrestrial insects occur in many laminae associated with abundant aquatic organisms, but are particularly abundant in some laminae that directly underlie tuff of fallout origin. Most of these terrestrial insects are interpreted to have been killed in the area adjacent to the studied palaeolake(s) during volcanic eruptions. Their carcasses were transported by influxes of fresh volcaniclastic material, primarily meteoric runoff and possibly minor distal pyroclastic flow into the palaeolake(s), and were buried in palaeolake deposits prior to extended decay probably due to a combination of rapid vertical settling, ash fall and water turbulence. Highlights • Frequent volcanic eruptions generated extensive volcaniclastic apron and lake(s). • Mass mortality of land animals occurred during volcanic eruptions. • Terrestrial animal remains were transported into lake(s) and rapidly buried. • Yanliao ecosystem was frequently disturbed by active volcanism. [ABSTRACT FROM AUTHOR]

Few fossil vertebrate skeletons are complete and fully articulated. Various taphonomic processes reduce the skeletal fidelity of decaying carcasses, the effects of most of which are reasonably well understood. Some fossil vertebrates, however, exhibit patterns of disarticulation and loss of completeness that are difficult to explain. Such skeletons are one of two variants. They are incomplete, often markedly so, but the preserved parts are highly articulated. Alternatively, they are complete, or nearly so, but articulation varies markedly between parts of the body. A characteristic feature is the absence of skeletal elements that, on the basis of their larger size and/or greater density, would be predicted to be present. Here we erect a model, termed “stick ‘n’ peel”, that explains how these distinctive patterns originate. The model emphasizes the role of decay products, especially fluids released from the carcass while resting on the sediment surface. These fluids permeate the sediment below and around the carcass. As a result, skeletal elements on the downward facing side of the carcass become adhered to the sediment surface, and are less likely to be remobilized as a result of current activity than others. The pattern of articulation and, especially, completeness is thus not what would be predicted on the basis of the size, shape and density of the skeletal elements. The effects of stick ‘n’ peel are difficult to predict a priori. Stick ‘n’ peel has been identified in vertebrate fossils in lacustrine and marine settings and is likely to be a common feature of the taphonomic history of many vertebrate assemblages. Specimens becoming adhered to the substrate may also explain the preservation in situ of the multi-element skeletons of invertebrates such as echinoderms, and integumentary structures such as hair and feathers in exceptionally preserved fossils. [ABSTRACT FROM AUTHOR]