Plant cytoplasm preservation in a baked root of Abies

Plant cytoplasm was thought impossible to be preserved in fossils. This stereotype is now undermined by increasing reports of plant cytoplasm in fossils. Previous simulations of high temperature preservation for plant cytoplasm were performed at short time scales, leaving the effectiveness and durability of such preservation an open question. Here we attempt to investigate the long time effect of high temperature on plant cytoplasm preservation in a root of Abies concolor that was baked in the wild at least 8 years ago. Light microscopy (LM), scanning electron microscopy (SEM), and transmission electron microscopy (TEM) indicate that cytoplasm and possible organelles such as mitochondria can be preserved in the baked tissue. The high frequency of wildfire in nature suggests that the potential for plant cytoplasm preservation in fossils is greater than commonly assumed. The good preservation of plant cytoplasm in the studied material after 8 years indicates that plant cytoplasm indeed can be preserved longer than previously proven.     

Molecular taphonomy of animal and plant cuticles: selective preservation and diagenesis

The nature of organic material and the environment in which it is deposited exert a major influence on the extent to which biomacromolecules are preserved in the fossil record. The role of these factors is explored with a particular focus on the cuticle of arthropods and leaves. Preservation of the original chemistry of arthropod cuticles is favoured by their thickness and degree of sclerotization, and the presence of biominerals. Decay and burial in terrestrial as opposed to marine, and anoxic rather than oxygenated conditions, likewise appear to enhance preservation. The most important factor in the long-term preservation of the chemistry of both animal and plant cuticles, however, is diagenetic alteration to an aliphatic composition. This occurs even in amber, which encapsulates the fossil, eliminating almost all external factors. Some plants contain an original decay-resistant macromolecular aliphatic component but this is not the case in arthropods. It appears that the aliphatic components of many plant as well as animal fossils may be the result of diagenetic polymerization. Selective preservation as a result of decay resistance may explain the initial survival of organic materials in sediments, but in many cases longer-term preservation relies on chemical changes. Selective preservation is only a partial explanation for the origin of kerogen.     

Experimental taphonomy of Artemia reveals the role of endogenous microbes in mediating decay and fossilization

Exceptionally preserved fossils provide major insights into the evolutionary history of life. Microbial activity is thought to play a pivotal role in both the decay of organisms and the preservation of soft tissue in the fossil record, though this has been the subject of very little experimental investigation. To remedy this, we undertook an experimental study of the decay of the brine shrimp Artemia, examining the roles of autolysis, microbial activity, oxygen diffusion and reducing conditions. Our findings indicate that endogenous gut bacteria are the main factor controlling decay. Following gut wall rupture, but prior to cuticle failure, gut-derived microbes spread into the body cavity, consuming tissues and forming biofilms capable of mediating authigenic mineralization, that pseudomorph tissues and structures such as limbs and the haemocoel. These observations explain patterns observed in exceptionally preserved fossil arthropods. For example, guts are preserved relatively frequently, while preservation of other internal anatomy is rare. They also suggest that gut-derived microbes play a key role in the preservation of internal anatomy and that differential preservation between exceptional deposits might be because of factors that control autolysis and microbial activity. The findings also suggest that the evolution of a through gut and its bacterial microflora increased the potential for exceptional fossil preservation in bilaterians, providing one explanation for the extreme rarity of internal preservation in those animals that lack a through gut.     

The role of experiments in investigating the taphonomy of exceptional preservation

In the last 20 years, much taphonomic experimentation has focused on the interpretation of exceptionally preserved fossils. Decay experiments have been used to interpret the features preserved in soft‐bodied fossils and to determine the sequence of character loss and its impact on phylogenetic position. Experiments on the impact of microbial communities on decay and mineralization have started to illuminate the processes involved in the fossilization of soft tissues, including embryos. The role of decay in promoting authigenic mineralization has been used to investigate the formation of Ediacaran macrofossils and concretions. Maturation experiments have shown how the constituents of animals and plants are transformed over time to a macromolecular material that converges on a similar stable composition. Other maturation experiments have explained how structural colours in fossils are altered from the original. A major area requiring investigation is the role of specific types of microbes in decay and their impact on sediment and pore water chemistry, as well as the environmental controls that determine their presence and level of activity. Microbial activity has received less attention than other factors in attempts to explain why the occurrence and nature of exceptional preservation varies in time and space through the fossil record.     

Cambrian anemones with preserved soft tissue from the Chengjiang biota, China

The group Cnidaria includes 'jellyfish', soft-bodied anemone and anemone-like forms and calcified corals. These diploblastic organisms have a fossil record extending back to the earliest metazoans of the Neoproterozoic; however certain cnidarians of the subclass Zoantharia, characterized by soft-bodied anemone-like forms, are absent or poorly represented in the fossil record. Despite the paucity of fossils, it is thought that calcification by soft anemone-like animals was responsible for producing the skeleton that allowed the preservation of the first corals. We report discovery of an abundant assemblage of in situ soft-bodied polyps with tissues. They are preserved in exquisite detail and come from the well-known Lower Cambrian Chengjiang biota of Yunnan, China. The soft-bodied polyps display a simple anatomy that is comparable to some extant anemones of the order Actinaria. The new fossils are assigned to Archisaccophyllia kunmingensis n. gen. et n. sp. Their simple and conservative form suggests that these fossils may represent some kind of ancestral rootstock. The preserved life assemblage provides a unique snapshot of Lower Cambrian anemone life and provides clues for relationships with extant actiniarians as well as calcified corals.     

THE FIDELITY OF THE FOSSIL RECORD: THE IMPROBABILITY OF PRESERVATION

Abstract:  The fidelity of the fossil record reflects how accurately it preserves the history of life. Since Darwin's time any mismatch between our theories and the fossil record has been attributed to the imperfections of the record. For over a century scarcity of gradual evolutionary trends was explained in this way until the punctuated equilibrium model was proposed. A null hypothesis that all morphological patterns in the fossil record are unbiased random walks can be rejected because it predicts far more apparent trends than exist. Current best estimates suggest that trends occur in at most 5% of characters. When an organism dies either it becomes fossilized or it doesn't. To be confident a species has not been preserved the probability against preservation must be significantly larger than the total number of individuals of that species that ever existed. For skeletized species preservation was the norm not the exception. Nevertheless, fossils must then avoid subsequent destruction and be discovered to be useful.     

Remarkably preserved annelid worms from the La Meseta Formation (Eocene), Seymour Island, Antarctica

Abstract: Worm tubes, which exhibit the replaced tube-lining membrane, have been collected from the lowermost Lower Eocene Acantilados Allomember of the La Meseta Formation, Seymour Island, Antarctica. The discovery represents the oldest examples of preservation of the tube-lining membrane of tube-dwelling Polychaeta in the fossil record. A new genus and species, Caprascolex antarcticus , are described. The specimens are preserved as thin coatings of amorphous iron oxide on the inner surface of the moulds interpreted to be the replaced tube-lining membrane. Examination of the rarely described tube-lining mucosoid membrane in extant polychaetes shows that the fossils are nearly identical in morphology and scale to extant forms. These fossils record in remarkable detail the morphology of the tube-lining membrane, which appears to be composed of growth bands formed as the worm constructed the tube. The tube-lining is believed to have been originally preserved as pyrite, with subsequent oxidation to iron oxide. The tube-lining membrane of worm tubes possibly is known from only one other fossil occurrence.     

The shape of pterosaur evolution: evidence from the fossil record

Abstract Although pterosaurs are a well‐known lineage of Mesozoic flying reptiles, their fossil record and evolutionary dynamics have never been adequately quantified. On the basis of a comprehensive data set of fossil occurrences correlated with taxon‐specific limb measurements, we show that the geological ages of pterosaur specimens closely approximate hypothesized patterns of phylogenetic divergence. Although the fossil record has expanded greatly in recent years, collectorship still approximates a sigmoid curve over time as many more specimens (and thus taxa) still remain undiscovered, yet our data suggest that the pterosaur fossil record is unbiased by sites of exceptional preservation (lagerstätte). This is because as new species are discovered the number of known formations and sites yielding pterosaur fossils has also increased – this would not be expected if the bulk of the record came from just a few exceptional faunas. Pterosaur morphological diversification is, however, strongly age biased: rarefaction analysis shows that peaks of diversity occur in the Late Jurassic and Early Cretaceous correlated with periods of increased limb disparity. In this respect, pterosaurs appear unique amongst flying vertebrates in that their disparity seems to have peaked relatively late in clade history. Comparative analyses also show that there is little evidence that the evolutionary diversification of pterosaurs was in any way constrained by the appearance and radiation of birds.     

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.     

Buoyancy mechanisms limit preservation of coleoid cephalopod soft tissues in Mesozoic Lagerstätten

Coleoid cephalopods are characterized by internalization of their shell, and are divided into the ten‐armed Decabrachia (squids and cuttlefish) and the eight‐armed Vampyropoda (octopuses and vampire squid). They have a rich fossil record predominantly of the limited biomineralized skeletal elements they possess: arm hooks, statoliths, mouthparts (the buccal mass) and internal shell (gladius or pen), although exquisitely preserved soft tissue coleoids are known from several Lagerstätten worldwide. Recent studies have shown that although morphological similarities between extant decabrachian gladii and fossil examples exist, no known examples of fossil decabrachians are currently known. However, molecular clock data and phylogenetic bracketing suggest that they should be present in Lagerstätten that are rich in vampyropod soft tissue fossils (i.e. Hâkel and Hâdjoula Lagerstätten, Cretaceous, Lebanon). We propose that a hitherto unknown taphonomic bias pertaining to the differing methods of buoyancy control within coleoid groups limits preservation potential. Both negatively and neutrally buoyant decabrachians use chemical buoyancy control (ammonia) whereas vampyropods do not. In the event of rapid burial in an environment conducive to exceptional preservation, ammonia dramatically decreases the ability of the decabrachian carcass to generate the required pH for authigenic calcium phosphate replacement, limiting its preservation potential. Moreover, the greater surface area and comparatively fragile dermis further decrease the potential for fossilization. This taphonomic bias may have contributed to the lack of preserved labile soft‐tissues in other cephalopods groups such as the ammonoids.     

Exceptional fossil preservation and the cambrian explosion

Exceptionally preserved, non-biomineralizing fossils contributeimportantly to resolving details of the Cambrian explosion,but little to its overall patterns. Six distinct "types" ofexceptional preservation are identified for the terminal Proterozoic-Cambrianinterval, each of which is dependent on particular taphonomiccircumstances, typically restricted both in space and time.Taphonomic pathways yielding exceptional preservation were particularlyvariable through the Proterozoic-Cambrian transition, at leastin part a consequence of contemporaneous evolutionary innovations.Combined with the reasonably continuous record of "Doushantuo-typepreservation," and the fundamentally more robust records ofshelly fossils, phytoplankton cysts and trace fossils, thesetaphonomic perturbations contribute to the documentation ofmajor evolutionary and biogeochemical shifts through the terminalProterozoic and early Cambrian. Appreciation of the relationship between taphonomic pathwayand fossil expression serves as a useful tool for interpretingexceptionally preserved, often problematic, early Cambrian fossils.In shale facies, for example, flattened non-biomineralizingstructures typically represent the remains of degradation-resistantacellular and extracellular "tissues" such as chaetae and cuticles,whereas three-dimensional preservation represents labile cellulartissues with a propensity for attracting and precipitating earlydiagenetic minerals. Such distinction helps to identify theacuticular integument of hyolithids, the chaetae-like natureof Wiwaxia sclerites, the chaetognath-like integument of Amiskwia,the midgut glands of various Burgess Shale arthropods, and themisidentification of deposit-feeding arthropods in the Chengjiangbiota. By the same reasoning, putative lobopods in the SiriusPasset biota and putative deuterostomes in the Chengiang biotaare better interpreted as arthropods.     

Animal behavior frozen in time: gregarious behavior of Early Jurassic lobsters within an ammonoid body chamber

Direct animal behavior can be inferred from the fossil record only in exceptional circumstances. The exceptional mode of preservation of ammonoid shells in the Posidonia Shale (Lower Jurassic, lower Toarcian) of Dotternhausen in southern Germany, with only the organic periostracum preserved, provides an excellent opportunity to observe the contents of the ammonoid body chamber because this periostracum is translucent. Here, we report upon three delicate lobsters preserved within a compressed ammonoid specimen of Harpoceras falciferum. We attempt to explain this gregarious behavior. The three lobsters were studied using standard microscopy under low angle light. The lobsters belong to the extinct family of the Eryonidae; further identification was not possible. The organic material of the three small lobsters is preserved more than halfway into the ammonoid body chamber. The lobsters are closely spaced and are positioned with their tails oriented toward each other. The specimens are interpreted to represent corpses rather than molts. The lobsters probably sought shelter in preparation for molting or against predators such as fish that were present in Dotternhausen. Alternatively, the soft tissue of the ammonoid may have been a source of food that attracted the lobsters, or it may have served as a long-term residency for the lobsters (inquilinism). The lobsters represent the oldest known example of gregariousness amongst lobsters and decapods in the fossil record. Gregarious behavior in lobsters, also known for extant lobsters, thus developed earlier in earth's history than previously known. Moreover, this is one of the oldest known examples of decapod crustaceans preserved within cephalopod shells.     

The Stomatal Ontogeny and Structure of the Liassic Pteridosperm Sagenopteris (Caytoniales) from Hungary

Stomatal ontogeny is often inferred but rarely documented for extinct fossil plants because it requires observations from young leaves that are rarely preserved as fossils. The discovery of several very young leaves of the Jurassic plant Sagenopteris (Caytoniales) in the Mecsek Mountains (southern Hungary) in a good state of preservation provides the opportunity for studying the stomatal ontogenesis of this genus. The specimens show perigenous anomocytic stomata. This feature confirms the evolutionarily high position of Sagenopteris among fossil gymnosperms and supports the opinion that the ancestors of angiosperms and some groups of pteridosperms might be closely related. Such clear examples of stomatal development have not previously been documented for fossil material.     

Miocene Ethiopian amber: A new source of fossil cryptogams

Amber is renowned for the exceptional preservation state of its inclusions, allowing detailed morphological analysis and providing relevant environmental, palaeoecological, geographical, and geological information. Amber deposits are predominantly known from North America, Europe, and Asia, and are considered to be rare on the continents that formed Gondwana. The recent discovery of fossiliferous amber deposits in Ethiopia, therefore, provides an inimitable opportunity to close gaps in the fossil record of African terrestrial biota and to study organisms which are otherwise rare in the fossil record. Here we show that diverse cryptogams are preserved in highest fidelity in Miocene Ethiopian amber. We describe gametophyte fragments of four liverworts: Thysananthus aethiopicus sp. nov. (Porellales, Lejeuneaceae), Lejeunea abyssinicoides sp. nov. (Porellales, Lejeuneaceae), Frullania shewanensis sp. nov. (Porellales, Frullaniaceae), and Frullania palaeoafricana sp. nov. (Porellales, Frullaniaceae). Furthermore, we describe a pleurocarpous moss of the extant genus Isopterygium (Hypnales, Pylaisiadelphaceae) and a lichen representing the order Lecanorales. These new specimens represent the first amber fossils of liverworts, mosses, and lichens from the African continent and render Ethiopian amber as one of the few worldwide amber deposits preserving bryophytes (mosses and liverworts) or lichens. Fossil species of Thysananthus were recorded in Eocene Baltic and Oligocene Bitterfeld as well as Miocene Dominican and probably also Miocene Mexican ambers. Fossils that can unequivocally be assigned to Lejeunea have only been found in Dominican amber so far. Neotropical ambers contain only one taxon of Frullania to date, while the genus is most diverse in Baltic, Bitterfeld, and Rovno ambers, formed in temperate regions. The new fossils support a tropical to subtropical origin of Ethiopian amber. The new African liverwort fossils are included in an updated list of leafy liverworts described from worldwide Cenozoic ambers to date.     

Influence of microbial biofilms on the preservation of primary soft tissue in fossil and extant archosaurs

Background

Mineralized and permineralized bone is the most common form of fossilization in the vertebrate record. Preservation of gross soft tissues is extremely rare, but recent studies have suggested that primary soft tissues and biomolecules are more commonly preserved within preserved bones than had been presumed. Some of these claims have been challenged, with presentation of evidence suggesting that some of the structures are microbial artifacts, not primary soft tissues. The identification of biomolecules in fossil vertebrate extracts from a specimen of Brachylophosaurus canadensis has shown the interpretation of preserved organic remains as microbial biofilm to be highly unlikely. These discussions also propose a variety of potential mechanisms that would permit the preservation of soft-tissues in vertebrate fossils over geologic time.

Methodology/Principal Findings

This study experimentally examines the role of microbial biofilms in soft-tissue preservation in vertebrate fossils by quantitatively establishing the growth and morphology of biofilms on extant archosaur bone. These results are microscopically and morphologically compared with soft-tissue extracts from vertebrate fossils from the Hell Creek Formation of southeastern Montana (Latest Maastrichtian) in order to investigate the potential role of microbial biofilms on the preservation of fossil bone and bound organic matter in a variety of taphonomic settings. Based on these analyses, we highlight a mechanism whereby this bound organic matter may be preserved.

Conclusions/Significance

Results of the study indicate that the crystallization of microbial biofilms on decomposing organic matter within vertebrate bone in early taphonomic stages may contribute to the preservation of primary soft tissues deeper in the bone structure.     

The fossil record of the Peronosporomycetes (Oomycota)

Evidence of fossil Peronosporomycetes has been slow to accumulate. In this review various fossils historically assigmed to the Peronosporomycets are dicussed briefly and an explanation is provided as to why the fossil record of this grouop has remained inconsistent. In recent year there has been several new reports of fossil peronosporomycetes based on structurally preserved oogonium-antheridium complexes from Derovonian and Carboniferous rocks that demonstrate the existence of these organisms as fossils and refute the long-standing assumption that they are too delicate to be preserved. Among these are serral tyoes characterized by oogonial surface members of the group. To date at last three groups of fossil vascular plants (i.e. lycophytes, ferns and seed ferns) are known to host peronosporomycetes aas endophytes; however only one form has been identified as a parasite.     

Vibrational spectroscopy of fossils

Over the last two decades, there has an been increasing interest in applying vibrational spectroscopy in palaeontological research. For example, this chemical analytical technique has been used to elucidate the chemical composition of a wide variety of fossils, including Archaean putative microfossils, stromatolites, chitinozoans, acritarchs, fossil algae, fossil plant cuticles, putative fossil arthropods, conodonts, scolecodonts and dinosaur bones. The insights provided by these data have been equally far ranging: to taxonomically identify a fossil, to determine biogenicity of a putative fossil, to identify preserved biologically synthesized compounds and to elucidate the preservational mechanisms of fossil material. Vibrational spectroscopy has clearly been a useful tool for investigating various palaeontological problems. However, it is also a tool that has been misapplied and misinterpreted, and thus, this review is dedicated to providing a palaeontologist who is new to vibrational spectroscopy with a basic understanding of these techniques, and the types of chemical information that can be obtained. Two example applications of these techniques are discussed in detail, one looking into fossil palynomorph taxonomy and other into the enigmatic Burgess Shale‐type preservation.     

Unique preservation of siliceous dinoflagellate motile cells from the Oligocene fossil Lagerstätte of Sieblos,Germany

The Triassic to Recent fossil record of the dinoflagellates is represented overwhelmingly by geologically resistant, organic-walled, non-motile resting cysts; such cysts are formed following the sexual phase in the life cycle. Very few confirmed records exist of the motile stage being preserved in the fossil record. This paper reports the occurrence of two very unusual dinoflagellate taphofacies, one developed in bituminous shales and the other in micrites, from the Oligocene fossil Lagerstätte at Sieblos, Hesse, Germany. A new dinoflagellate taxon, Sieblososphaera martini sp. nov. has been identified through analysis of dissociated skeletal elements in the bituminous shales and external moulds and casts in the micrites. The unique preservation of these fossils confirms them not only as primary biogenically silicified motile thecate cells, but also indicates that there was a much greater range of tabulation present within the subfamily Lithoperidiniaceae than has hitherto been recognized.     

Names for trace fossils: a uniform approach

The taxonomic treatment of trace fossils needs a uniform approach, independent of the ethologic groups concerned. To this aim, trace fossils are rigorously defined with regard to biological taxa and physical sedimentary structures. Potential ichnotaxobases are evaluated, with morphology resulting as the most important criterion. For trace fossils related to bioerosion and herbivory, substrate plays a key role, as well as composition for coprolites. Size, producer, age, facies and preservation are rejected as ichnotaxobases. Separate names for undertracks and other poorly preserved material should gradually be replaced by ichnotaxa based on well-preserved specimens. Recent traces may be identified using established trace fossil taxa but new names can only be based on fossil material, even if the distinction between recent and fossil may frequently remain arbitrary. It is stressed that ichnotaxa must not be incorporated into biological taxa in systematics. Composite trace fossil structures (complex structures made by the combined activity of two or more species) have no ichnotaxonomic standing but compound traces (complex structures made by one individual tracemaker) may be named separately under certain provisions. The following emendations are proposed to the International Code of Zoological Nomenclature: The term 'work of an animal' should be deleted from the code, and ichnotaxa should be based solely on trace fossils as defined herein.     

陕西宁强胡家坝地区晚埃迪卡拉世高家山生物群岩相变化与化石保存关系

我国上扬子陕西宁强-勉县境内产出埃迪卡拉纪晚期高家山生物群,根据大量野外追踪、实地观察与室内岩相分析,对碎屑岩相黄铁矿化软躯体化石的出现与岩相横向变化之间的关系以及相应的埋藏相进行初步探讨,认为胡家坝露头带为相对远离风暴源的沉积序列,最利于化石的保存;高家山露头带则属于相对靠近风暴源的沉积序列,化石保存潜力不如胡家坝露头带的沉积序列。