Fossilization Of Soft Tissues Research Articles

Soft-tissue fossilization illuminates the stepwise evolution of the ray-finned fish brain

A complex brain is central to the success of backboned animals. However, direct evidence bearing on vertebrate brain evolution comes almost exclusively from extant species, leaving substantial knowledge gaps. Although rare, soft-tissue preservation in fossils can yield unique insights on patterns of neuroanatomical evolution. Paleontological evidence from an exceptionally preserved Pennsylvanian (∼318 Ma) actinopterygian, Coccocephalus, calls into question prior interpretations of ancestral actinopterygian brain conditions. However, the ordering and timing of major evolutionary innovations, such as an everted telencephalon, modified meningeal tissues, and hypothalamic inferior lobes, remain unclear. Here, we report two distinct actinopterygian morphotypes from the latest Carboniferous-earliest Permian (∼299 Ma) of Brazil that show extensive soft-tissue preservation of brains, cranial nerves, eyes, and potential cardiovascular tissues. These fossils corroborate inferences drawn from ✝Coccocephalus, while adding new information about neuroanatomical evolution. Skeletal features indicate that one of these Brazilian morphotypes is more closely related to living actinopterygians than the other, which is also reflected in soft-tissue features. Significantly, the more crownward morphotype shows a key neuroanatomical feature of extant actinopterygians-an everted telencephalon-that is absent in the other morphotype and ✝Coccocephalus. All preserved Paleozoic actinopterygian brains show broad similarities, including an invaginated cerebellum, hypothalamus inferior lobes, and a small forebrain. In each case, preserved brains are substantially smaller than the enclosing cranial chamber. The neuroanatomical similarities shared by this grade of Permo-Carboniferous actinopterygians reflect probable primitive conditions for actinopterygians, providing a revised model for interpreting brain evolution in a major branch of the vertebrate tree of life.

Preservation of Soft-Bodied Organisms in Early Cambrian Carbonates

Abstract —Preservation of soft-bodied organisms as casts is common in the Ediacaran fossil record and extremely rare in Cambrian rocks. Among the factors, limiting fossilization of soft tissues, there is bioturbation–sediment disturbance by burrowing organisms. It is the emergence of burrowing metazoans and associated sediment bioturbation that is thought to be one of the major causes for the disappearance of Ediacaran soft-bodied organisms from the fossil record around ~540 Ma. Here, we study an assemblage of fossils preserved as casts in the Fortunian carbonates of the Olenek Uplift (northeastern Siberian Platform) in association with a typical Fortunian ichnoassemblage. The overall morphology and preservation of the fossils reveal that they comprise microbially induced sedimentary structures and soft-bodied holdfasts. The latter are vaguely reminiscent of some Ediacaran holdfasts, though it is unlikely that they include their phylogenetic descendants. Three-dimensional preservation of the studied fossils was caused by authigenic crystallization of calcite and its further early-diagenetic dolomitization. Our study confirms the critical importance of specific environmental conditions ensuring preservation of soft-bodied organisms as three-dimensional molds and casts. This unique interplay of environmental factors became rare in the Cambrian Period, which was caused by intensification and expansion of bioturbation in marine basins.

Sediment-encased pressure–temperature maturation experiments elucidate the impact of diagenesis on melanin-based fossil color and its paleobiological implications

AbstractMelanin pigments are central to colors and patterns in modern vertebrate integuments, which inform upon ecological and behavioral strategies like crypsis, aposematism, and sociosexual selection. Over the last decade, melanin has emerged as a valuable tool for predicting color in exceptionally preserved fossil feathers and subsequent testing of paleobiological hypotheses in long-extinct dinosaurs and birds. Yet much remains to be learned about melanin stability, diagenetic alterations to melanin chemistry, and their implications for “paleocolor reconstruction.” Pressure–temperature maturation experiments with modern feathers offer a way to examine these topics but have mostly been conducted in closed-system capsules or open-system aluminum foil. Both methods have operational limitations and do not consider the filtering effect of porous sediment matrices on thermally labile chemical groups versus stable ones during natural fossilization. We use sediment-encased maturation to resolve this issue and demonstrate replication of organic preservation of melanin highly comparable to compression fossils. Our experiments, coupled with time-of-flight secondary ion mass spectrometry, show predictable volatilization of N/S-bearing molecules and increased melanin cross-linking with elevated temperatures. We also suggest that eumelanin is more stable compared with pheomelanin at higher temperatures, explaining why eumelanic colors (black, dark brown, iridescent) are better preserved in fossils than pheomelanic ones (reddish brown). Furthermore, we propose that proteins preferentially undergo hydrolysis more so than forming N-heterocycles in selectively open systems analogous to natural matrices. Thus, we conclude that melanin pigments and not diagenetically altered protein remnants are the key players in promoting fossilization of soft tissues like feathers.

Preservation of Membrana Testacea in titanosaurid dinosaur eggshells from the Upper Cretaceous Deccan volcano-sedimentary strata of Central India

AbstractThe calcitic eggshell units of amniote eggs are underlain by a soft organic layer, the Membrana Testacea (MT), which has a mesh-like texture consisting of fibrils of organic material. Because of its soft anatomy, the MT is rarely preserved in fossils and only a few fossil reports of MT are known so far. Here we report the preservation of a mineralized MT layer in titanosaur eggshells recovered from a marlstone facies interbedded with the Deccan lava flows exposed near Piplanarayanwar village of Chhindwara District, Madhya Pradesh state in Central India. The MT layer is mesh-like, resembling protein membranes of extant reptiles and the MT reported in titanosaurid eggshells of the Upper Cretaceous Anacleto Formation at Auca Mahuevo, Argentina. The presence of tendrils and fibres of calcite in the MT layer testifies to the fact that the calcium layer represents the original fibrous MT. It also supports the view that fossilization of soft tissues like MT is possible because of the inferred anaerobic conditions that prevailed during the deposition of Piplanarayanwar intertrappean sediments in lacustrine or paludal bodies in a coastal-plain setting.

The complex role of microbial metabolic activity in fossilization.

Bacteria play an important role in the fossilization of soft tissues; their metabolic activities drive the destruction of the tissues and also strongly influence mineralization. Some environmental conditions, such as anoxia, cold temperatures, and high salinity, are considered widely to promote fossilization by modulating bacterial activity. However, bacteria are extremely diverse, and have developed metabolic adaptations to a wide range of stressful conditions. Therefore, the influence of the environment on bacterial activity, and of their metabolic activity on fossilization, is complex. A number of examples illustrate that simple, general assumptions about the role of bacteria in soft tissue fossilization cannot explain all preservational pathways: (i) experimental results show that soft tissues of cnidaria decay less in oxic than anoxic conditions, and in the fossil record are found more commonly in fossil sites deposited under oxic conditions rather than anoxic environments; (ii) siderite concretions, which often entomb soft tissue fossils, precipitate due to a complex mixture of sulfate- and iron reduction by some bacterial species, running counter to original theories that iron reduction is the primary driver of siderite concretion growth; (iii) arthropod brains, now widely accepted to be preserved in many Cambrian fossil sites, are one of the first structures to decay in taphonomic experiments, indicating that their fossilization processes are complex and influenced by bacterial activity. In order to expand our understanding of the complex process of bacterially driven soft tissue fossilization, more research needs to be done, on fossils themselves and in taphonomic experiments, to determine how the complex variation in microbial metabolic activity influences decay and mineralization.

A high-resolution timescale for the Miocene Shanwang diatomaceous shale lagerstätte (China): development of Wavelet Scale Series Analysis for cyclostratigraphy

The Miocene aged Shanwang Formation from the Shanwang National Geopark in China represents a succession of lacustrine diatomaceous shales containing an abundant and diverse biota with lagerstatte fossilization of soft tissues. To date, the Shanwang Formation has not been investigated for cyclostratigraphy nor has it been dated with high precision methods. Now we use thorium data as a paleoenvironmental and paleoclimatic proxy to conduct a detailed cyclostratigraphic analysis. A new and simple cyclostratigraphic method, Wavelet Scale Series Analysis (WSSA) is developed to recognize Milankovitch cycles. A total of three short eccentricity and fifteen precession cycles are identified; obliquity cycles are not apparent. In the sedimentary succession, the corresponding precession and short eccentricity cycles are 1.17 m and 4.98 m thick respectively, with this verified by Correlation Coefficient (COCO) analysis and Multitaper-Method (MTM) spectral analysis. We estimate the studied interval was deposited over a duration of 0.3 Myr with a depositional rate of c. 5.7 cm/kyr. Paleomagnetic and radio isotope dating data shows that the diatomaceous shale was deposited during Chron C5En, which places it at approximately 18.5 Ma during the Burdigalian stage of the Early Miocene, rather than in the Middle Miocene as previously thought. The Shanwang lagerstatte biota therefore predates the Middle Miocene Climate Optimum (MMCO) and did not form within it. The geological time scale with a high resolution of 20 kyr was set accordingly.

Mineral composition of host sediments influences the fossilization of soft tissues

Cambrian Lagerstätten host rocks are frequently composed of kaolinite and chlorite in varying amounts; accordingly, our goal was to study the preservation potential of crustaceans in these two clays. We conducted long-term experiments (12–18 months, the longest duration of actualistic taphonomy experiments from published literature) on the decay of Artemia salina in these clay sediments. The degree of preservation, transformed mineralogical composition of the sediments, and the elemental composition of the nauplial remains were examined. We demonstrate that the kaolinite and chlorite sediment enhanced the preservation (in the kaolinite the effect was considerably higher than in the chlorite) compared with the sediment-free control. pH inside the sediments dropped to 6.5–7.1 and was even lower (<4) around the buried carcasses, facilitating the dissolution of clays. This phenomenon was confirmed by mineralogical analyses of the experimental sediments, which showed mineralogical signatures of such dissolution and new mineral phases. According to the variations in the dissolved minerals in the sediments, different cations entered the buried remains as was shown by the multiple energy dispersive X-ray analyses. An increased level of Mg was detected in the carcasses buried in chlorite, whereas Al and Si concentrations were higher in the kaolinite; in both cases, Ca rapidly entered the decaying tissues from marine water. Bacteria underwent similar mineralization as the macroremains and apparently had no direct effect on the mineralization. The results confirmed an important role of dissolved Al ions in preservation of soft-bodied organisms in clay-dominated sediments and explained wide variation in chemical composition of their fossils.

Rocks with Soft-Tissue Fossils Share a Mineral Fingerprint

Discovering new resting places of these rare and information-rich fossils will be critical to understanding the largest expansion of life in Earth’s history, according to researchers.

Fossilization of soft tissues

Fossilization of soft tissues

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

AbstractColeoid 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.

The role of experiments in investigating the taphonomy of exceptional preservation

AbstractIn 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.

SEDIMENT PERMEABILITY AND THE PRESERVATION OF SOFT-TISSUES IN CONCRETIONS: AN EXPERIMENTAL STUDY

Sediment permeability is hypothesized to affect soft tissue fossilization within concretions through its effects on organic decay and concretion growth. The role of permeability was tested in a series of experiments in which cod tissue was decayed in glass beads of varying permeabilities. Decay was measured using infrared gas analysis (IRGA), and mineral precipitation within the beads (a proxy for concretion growth) was measured using micro CT scanning. The interactions of the three variables—sediment permeability, decay, and mineralization—were assessed with MANOVA and with linear regressions of decay and precipitation per unit decay on permeability. These two linear regressions were combined into a more general, nonlinear expression of the relationship between permeability and total mineral precipitation. The results show that sediment permeability has two competing effects on precipitation, the strength of each varying dynamically depending on permeability. Low permeability environments inhibit decay, thus enhancing fossilization but inhibiting overall precipitation because a build-up of decay products is necessary to promote mineral formation (the “decay effect”). However, low permeability environments can also increase precipitation per unit decay by inhibiting the diffusion of decay products away from the carcass, allowing for a faster build-up of decay products (the “mineralization effect”). At low permeabilities, the decay effect dominates (decay controlled), and precipitation is positively correlated with permeability. At higher permeabilities, the mineralization effect dominates (permeability controlled), and precipitation is negatively correlated with permeability. The experiments show that fossilization within concretions is promoted by decay inhibition (at low permeabilities) and rapid concretion growth (at intermediate permeabilities). Thus, the effects of permeability on fossilization are complex, and influence the mechanism of fossilization.

A unifying model for Neoproterozoic-Palaeozoic exceptional fossil preservation through pyritization and carbonaceous compression.

Soft-tissue fossils capture exquisite biological detail and provide our clearest views onto the rise of animals across the Ediacaran-Cambrian transition. The processes contributing to fossilization of soft tissues, however, have long been a subject of debate. The Ediacaran Gaojiashan biota displays soft-tissue preservational styles ranging from pervasive pyritization to carbonaceous compression, and thus provides an excellent opportunity to dissect the relationships between these taphonomic pathways. Here geochemical analyses of the Gaojiashan fossil Conotubus hemiannulatus show that pyrite precipitation was fuelled by the degradation of labile tissues through bacterial sulfate reduction (BSR). Pyritization initiated with nucleation on recalcitrant tube walls, proceeded centripetally, decelerated with exhaustion of labile tissues and possibly continued beneath the BSR zone. We propose that pyritization and kerogenization are regulated principally by placement and duration of the decaying organism in different microbial zones of the sediment column, which hinge on post-burial sedimentation rate and/or microbial zone thickness.

Bias and sensitivity in the placement of fossil taxa resulting from interpretations of missing data.

The utility of fossils in evolutionary contexts is dependent on their accurate placement in phylogenetic frameworks, yet intrinsic and widespread missing data make this problematic. The complex taphonomic processes occurring during fossilization can make it difficult to distinguish absence from non-preservation, especially in the case of exceptionally preserved soft-tissue fossils: is a particular morphological character (e.g., appendage, tentacle, or nerve) missing from a fossil because it was never there (phylogenetic absence), or just happened to not be preserved (taphonomic loss)? Missing data have not been tested in the context of interpretation of non-present anatomy nor in the context of directional shifts and biases in affinity. Here, complete taxa, both simulated and empirical, are subjected to data loss through the replacement of present entries (1s) with either missing (?s) or absent (0s) entries. Both cause taxa to drift down trees, from their original position, toward the root. Absolute thresholds at which downshift is significant are extremely low for introduced absences (two entries replaced, 6% of present characters). The opposite threshold in empirical fossil taxa is also found to be low; two absent entries replaced with presences causes fossil taxa to drift up trees. As such, only a few instances of non-preserved characters interpreted as absences will cause fossil organisms to be erroneously interpreted as more primitive than they were in life. This observed sensitivity to coding non-present morphology presents a problem for all evolutionary studies that attempt to use fossils to reconstruct rates of evolution or unlock sequences of morphological change. Stem-ward slippage, whereby fossilization processes cause organisms to appear artificially primitive, appears to be a ubiquitous and problematic phenomenon inherent to missing data, even when no decay biases exist. Absent characters therefore require explicit justification and taphonomic frameworks to support their interpretation.

The Role of Biology in the Fossilization of Embryos and Other Soft-Bodied Organisms: Microbial Biofilms and Lagerstätten

Soft-tissue fossils are among the most striking and informative remains of extinct organisms. Although relatively rare, they are diverse, ranging from single microbial cells to nuclei and chromosomes; algae; metazoan embryos and larvae; flowers; complete, small, soft-bodied metazoans, metazoan tissues; integumentary structures such as melanosomes; skin texture, vertebrate feathers and hair, insect wings with color patterns, and sometimes even the entire bodies of large animals. The susceptibility of newly dead soft tissues to physical destruction, consumption, and microbial decay makes their preservation unlikely under most taphonomic conditions. In addition, their vulnerability to rapid autolysis, bioturbation, and destructive physical processes requires that rapid biological events must occur as the critical first steps of fossilization. An understanding of the processes by which biological remains enter the fossil record is important in inferring what non-microbial and microbial processes were operative in Lagerstätten. Paleontologists have recognized that microbial biofilms often accompany soft-tissue fossils, and have suggested that bacteria play an active role in soft tissue fossilization, but that role must be determined experimentally with living bacteria and dead tissue.Marine embryos and marine bacteria are used to investigate the processes that mediate early steps in soft-tissue preservation because they offer simple systems for laboratory investigation of the roles of autolysis-blocking environments, microbial interactions, biofilm formation, and authigenic mineralization in taphonomy. Understanding microbially mediated preservation of embryos may supply new insights into a more general biology of fossilization.

Concretions as Agents of Soft-Tissue Preservation: A Review

Carbonate concretions may preserve exceptional soft-tissue fossils. Organismal decay in sediment can produce HCO3−faster than it diffuses away, creating a local microenvironment of high alkalinity around the decaying organism that promotes carbonate precipitation. Infilling of sediment pore-space around the decaying organism decreases permeability and inhibits decay, thus increasing preservation potential within the concretion and promoting soft-tissue preservation. Different patterns of concretion growth may promote different mechanisms of soft-tissue preservation. Other factors such as shifting salinity and clay mineral chemistry, which increase preservation potential in the depositional environment, also promote soft tissue preservation within concretions. The rate of concretion nucleation and growth affects preservation; the faster concretions nucleate and grow, the better the preservation. Concretionary preservation is biased both by concretion-promoting environments and organism size effects on concretion formation.

Exceptional soft tissue fossilization of a Pleistocene vulture (Gyps fulvus): new evidence for emplacement temperatures of pyroclastic flow deposits

Volcanic sediments are often unsuitable for the fossilization of both hard and soft organic tissues, however, in some circumstances, they can provide unusual conditions for the preservation of remains. Here we report an exceptional case of soft tissue fossilization of a Late Pleistocene Eurasian griffon vulture (Gyps fulvus) in the pyroclastic sequence of the Alban Hills volcanic region (SE Rome, Italy). CT analyses have revealed an exceptional natural cast of the complete head and neck that preserve extraordinary detail including the fossilized everted tongue, beak, feather insertions and the first record of the nictitating membrane of the eye. This fossilization (superior in detail even to the victims of the AD 79 Plinian eruption of Vesuvius) reveals no evidence of burning and requires re-evaluation of the thermal constraints in operation for the preservation of organic materials within pyroclastic sediments. The analysis of the external morphological features has provided key information regarding the taphonomic processes in operation, the emplacement temperatures of distal pyroclastic flow deposits and the relationships between organic materials and low temperature phreatomagmatic flows. This sheds light not only on the extremely rare situation of fossilization in volcanic contexts but also provides a new perspective on taphonomic studies of highly detailed casts of fossil vertebrates.

Inorganic and biogenic As-sulfide precipitation at seafloor hydrothermal fields

Highlights: • As-sulfides are minor minerals at volcanic arc and back-arc hydrothermal settings. • There are two modes of As-sulfide precipitation: inorganic and biogenic. • The most stable As-sulfide at low-T seafloor hydrothermal conditions is orpiment. • Biogenic type of orpiment represents completely mineralized fungal hyphae. • Fungal orpiment precipitation is a mechanism of bio-sequestration of hydrothermal As. We investigated As-sulfides (orpiment, As2S3 and realgar, As4S4) from four seafloor hydrothermal fields: three from back-arc and one from mid-ocean ridge settings. Our studies suggest two different modes of As-sulfide precipitation at the seafloor hydrothermal sites: inorganic and biogenic. The Eh–pH modeling shows that the most stable As-sulfide at low-temperature (T < 100 °C) seafloor hydrothermal conditions is orpiment, whereas realgar appears as a stable As-sulfide phase at T = 25 °C and under a narrow range of Eh–pH values. Inorganically precipitated realgar forms euhedral crystals whereas the orpiment is mostly colloform. The biogenic type of orpiment (no realgar of this type was found) represents completely mineralized fungal hyphae. The proposed scenario of biogenic orpiment formation assumes that crack-dwelling fungal filaments in the sub-seafloor were killed by an As-rich hydrothermal fluid moving along the same crack network. The fungal organic matter served as a geochemical trap for hydrothermal As which reacted with S (leached from the basement or reduced from seawater sulfate) and mineralized the fungal hyphae as As2S3. This process of fungal orpiment precipitation is a mechanism of bio-sequestration of hydrothermal As and another mode of soft-tissue fossilization not described so far.

Contingent interactions among biofilm‐forming bacteria determine preservation or decay in the first steps toward fossilization of marine embryos

Fossils of soft tissues provide important records of early animals and embryos, and there is substantial evidence for a role for microbes in soft tissue fossilization. We are investigating the initial events in interactions of bacteria with freshly dead tissue, using marine embryos as a model system. We previously found that microbial invasion can stabilize embryo tissue that would otherwise disintegrate in hours or days by generating a bacterial pseudomorph, a three dimensional biofilm that both replaces the tissue and replicates its morphology. In this study, we sampled seawater at different times and places near Sydney, Australia, and determined the range and frequency of different taphonomic outcomes. Although destruction was most common, bacteria in 35% of seawater samples yielded morphology‐preserving biofilms. We could replicate the taphonomic pathways seen with seawater bacterial communities using single cultured strains of marine gammaproteobacteria. Each given species reproducibly generated a consistent taphonomic outcome and we identified species that yielded each of the distinct pathways produced by seawater bacterial communities. Once formed,bacterial pseudomorphs are stable for over a year and resist attack by other bacteria and destruction by proteases and other lytic enzymes. Competition studies showed that the initial action of a pseudomorphing strain can be blocked by a strain that destroys tissues. Thus embryo preservation in nature may depend on contingent interactions among bacterial species that determine if pseudomorphing occurs.We used Artemia nauplius larvae to show that bacterial biofilm replacement of tissue is not restricted to embryos, but is relevant for preservation of small multicellular organisms. We present a model for bacterial self‐assembly of large‐scale three‐dimensional tissue pseudomorphs, based on smallscaleinteractions among individual bacterial cells to form local biofilms at structural boundaries within the tissue. Localbiofilms then conjoin to generate the pseudomorph.

Exceptional Soft-Tissue Preservation in Boring Ctenostome Bryozoans and Associated “Fungal” Borings from the Early Devonian of Podolia, Ukraine

Colonies of boring ctenostome bryozoans and microborings of “fungi” that occur in the Early Devonian (Lochkovian, ∼416 Ma) of Podolia, western Ukraine, have soft-tissue preserved by phosphatization. These comprise exceptional three-dimensional body walls of feeding zooids with probable parietal muscles inserted on the cystid wall, and setigerous collars twisted within the vestibulum. The presence of collars in this Early Devonian ctenostomes proves the existence of this feature for more than 416 Ma of ctenostome evolution. Phosphatized remains of the zooid walls are interpreted as relicts of the originally chitinous cystid walls. This is the first record of soft-tissue fossilization in a boring bryozoan. The presence of cavities (specialized heterozooids), empty or filled with laminated calcium phosphate, is also documented in bryozoans for the first time. These cavities are interpreted as “store-rooms” in which the bryozoans accumulated nutrients. The new taxon, Podoliaporu doroshevi gen. et sp. nov. is ...