Late Ordovician eurypterid preserves oldest euchelicerate musculature in pyrite.

Pyritization of soft tissues of invertebrates is rare in the fossil record. In New York State, it occurs in black shales of the Lorraine Group (Late Ordovician), the best-known example of which is Beecher's Trilobite Bed. Exceptional preservation at the quarry where this bed is exposed allowed detailed examination of trilobite and ostracod soft-tissue anatomy. Here, we present the first example of a eurypterid (sea scorpion) currently ascribed to Carcinosomatidae from this deposit that also preserves the first evidence for mesosomal musculature in eurypterids. This specimen demonstrates that eurypterid musculature can be preserved in pyrite and evidences the oldest example of euchelicerate muscles within the fossil record. Sulfur isotope data illustrate that pyrite rapidly replicated muscle tissue in the early burial environment, prior to the pyritization of biomineralized exoskeleton and cuticular trilobite limbs. This discovery therefore expands the limited fossil record of euchelicerate musculature, while extending the taphonomic scope for preservation of detailed internal structures, more broadly, within arthropods.

Post-Cambrian survival of the tubicolous scalidophoran Selkirkia.

Scalidophoran worms represent common infaunal components of early and middle Cambrian Burgess Shale-type fossil biotas. Early scalidophorans resemble extant priapulids based on overall morphology, but the genus Selkirkia represents the earliest record of tube dwelling for the group. Despite its ubiquitous presence in exceptional marine deposits, whether the exclusively Cambrian occurrence of Selkirkia reflects its entire evolutionary history or is affected by taphonomic biases remains unresolved. Here, we demonstrate the post-Cambrian survival of Selkirkia based on new material from the Lower Ordovician Fezouata Shale biota of Morocco. The discovery of Selkirkia in the Fezouata Shale extends the biostratigraphic range of the genus by 25 million years and its palaeobiogeographic occurrence to the high latitudes of Gondwana, strengthens the evolutionary links between Cambrian and Ordovician Burgess Shale-type biotas and increases scalidophoran diversity for the Fezouata Shale biota otherwise consisting exclusively of the palaeoscolecid Palaeoscolex? tenensis. The tube of Selkirkia underwent negligible external change for over 40 million years, indicating a high degree of morphological stasis during the Early Palaeozoic. A tubicolous mode of life is rare among extant priapulids and expressed only in Maccabeus, which forms a delicate tube from agglutinated plant debris, unlike the macroscopic secreted cuticular tube of Selkirkia.

Convergent evolution of ventral adaptations for enrolment in trilobites and extant euarthropods.

The ability to enrol for protection is an effective defensive strategy that has convergently evolved multiple times in disparate animal groups ranging from euarthropods to mammals. Enrolment is a staple habit of trilobites, and their biomineralized dorsal exoskeleton offered a versatile substrate for the evolution of interlocking devices. However, it is unknown whether trilobites also featured ventral adaptations for enrolment. Here, we report ventral exoskeletal adaptations that facilitate enrolment in exceptionally preserved trilobites from the Middle Ordovician Walcott-Rust Quarry in New York State, USA. Walcott-Rust trilobites reveal the intricate three-dimensional organization of the non-biomineralized ventral anatomy preserved as calcite casts, including the spatial relationship between the articulated sternites (i.e. ventral exoskeletal plates) and the wedge-shaped protopodites. Enrolment in trilobites is achieved by ventrally dipping the anterior margin of the sternites during trunk flexure, facilitated by the presence of flexible membranes, and with the close coupling of the wedge-shaped protopodites. Comparisons with the ventral morphology of extant glomerid millipedes and terrestrial isopods reveal similar mechanisms used for enrolment. The wedge-shaped protopodites of trilobites closely resemble the gnathobasic coxa/protopodite of extant horseshoe crabs. We propose that the trilobites' wedge-shaped protopodite simultaneously facilitated tight enrolment and gnathobasic feeding with the trunk appendages.

The Liexi fauna: a new Lagerstätte from the Lower Ordovician of South China.

The Ordovician Lagerstätten record substantial amounts of excellent preservation and soft-bodied fossils during the Great Ordovician Biodiversification Event (GOBE). However, few Lagerstätten are known from the Lower Ordovician, most of which are preserved in restricted environments and high-latitude regions. Here, we report on a new tropical Lagerstätte, Liexi fauna, which has been recently discovered from a carbonate succession within the Lower Ordovician Madaoyu Formation in western Hunan, South China. It contains a variety of soft tissues, as well as rich shelly fossils, including palaeoscolecidan worms, possible Ottoia, trilobites, echinoderms, sponges, graptolites, polychaetes, bryozoans, conodonts and other fossils. The fauna includes taxa that are not only Cambrian relics, but also taxa originated during the Ordovician, constituting a complex and complete marine ecosystem. The coexistence of the Cambrian relics and Ordovician taxa reveals the critical transition between the Cambrian and Palaeozoic Evolutionary faunas. The unusual Liexi fauna provides new evidence for understanding Ordovician macroevolution and the onset of the GOBE.

Computed tomography sheds new light on the affinities of the enigmatic euarthropod Jianshania furcatus from the early Cambrian Chengjiang biota.

BACKGROUND: The Chengjiang biota is one of the most species-rich Cambrian Konservat-Lagerstätten, and preserves a community dominated by non-biomineralized euarthropods. However, several Chengjiang euarthropods have an unfamiliar morphology, are extremely rare, or incompletely preserved. RESULTS: We employed micro-computed tomography to restudy the enigmatic euarthropod Jianshania furcatus. We reveal new morphological details, and demonstrate that the specimens assigned to this species represent two different taxa. The holotype of J. furcatus features a head shield with paired anterolateral notches, stalked lateral eyes, and an articulated tailspine with a bifurcate termination. The other specimen is formally redescribed as Xiaocaris luoi gen. et sp. nov., and is characterized by stalked eyes connected to an anterior sclerite, a subtrapezoidal head shield covering three small segments with reduced tergites, a trunk with 15 overlapping tergites with a well-developed dorsal keel, and paired tail flukes. CONCLUSIONS: The presence of antennae, biramous appendages with endopods composed of 15 articles, and multiple appendage pairs associated with the trunk tergites identify X. luoi nov. as a representative of Fuxianhuiida, an early branching group of stem-group euarthropods endemic to the early Cambrian of Southwest China. X. luoi nov. represents the fifth fuxianhuiid species described from the Chengjiang biota, and its functional morphology illuminates the ecological diversity of this important clade for understanding the early evolutionary history of euarthropods.

Soft-Bodied Fossils Are Not Simply Rotten Carcasses - Toward a Holistic Understanding of Exceptional Fossil Preservation: Exceptional Fossil Preservation Is Complex and Involves the Interplay of Numerous Biological and Geological Processes.

Exceptionally preserved fossils are the product of complex interplays of biological and geological processes including burial, autolysis and microbial decay, authigenic mineralization, diagenesis, metamorphism, and finally weathering and exhumation. Determining which tissues are preserved and how biases affect their preservation pathways is important for interpreting fossils in phylogenetic, ecological, and evolutionary frameworks. Although laboratory decay experiments reveal important aspects of fossilization, applying the results directly to the interpretation of exceptionally preserved fossils may overlook the impact of other key processes that remove or preserve morphological information. Investigations of fossils preserving non-biomineralized tissues suggest that certain structures that are decay resistant (e.g., the notochord) are rarely preserved (even where carbonaceous components survive), and decay-prone structures (e.g., nervous systems) can fossilize, albeit rarely. As we review here, decay resistance is an imperfect indicator of fossilization potential, and a suite of biological and geological processes account for the features preserved in exceptional fossils.

The appendicular morphology of Sinoburius lunaris and the evolution of the artiopodan clade Xandarellida (Euarthropoda, early Cambrian) from South China.

BACKGROUND: Artiopodan euarthropods represent common and abundant faunal components in sites with exceptional preservation during the Cambrian. The Chengjiang biota in South China contains numerous taxa that are exclusively known from this deposit, and thus offer a unique perspective on euarthropod diversity during the early Cambrian. One such endemic taxon is the non-trilobite artiopodan Sinoburius lunaris, which has been known for approximately three decades, but few details of its anatomy are well understood due to its rarity within the Chengjiang, as well as technical limitations for the study of these fossils. Furthermore, the available material does not provide clear information on the ventral organization of this animal, obscuring our understanding of phylogenetically significant details such as the appendages. RESULTS: We employed X-ray computed tomography to study the non-biomineralized morphology of Sinoburius lunaris. Due to the replacement of the delicate anatomy with pyrite typical of Chengjiang fossils, computed tomography reveals substantial details of the ventral anatomy of Sinoburius lunaris, and allow us to observe in detail the three-dimensionally preserved appendicular organization of this taxon for the first time. The dorsal exoskeleton consists of a crescent-shaped head shield with well-developed genal spines, a thorax with seven freely articulating tergites, and a fused pygidium with lateral and median spines. The head bears a pair of ventral stalked eyes that are accommodated by dorsal exoskeletal bulges, and an oval elongate ventral hypostome. The appendicular organization of the head is unique among Artiopoda. The deutocerebral antennae are reduced, consisting of only five podomeres, and bear an antennal scale on the second podomere that most likely represents an exite rather than a true ramus. The head includes four post-antennal biramous limb pairs. The first two biramous appendages are differentiated from the rest. The first appendage pair consists of a greatly reduced endopod coupled with a greatly elongated exopod with a potentially sensorial function. The second appendage pair carries a more conventionally sized endopod, but also has an enlarged exopod. The remaining biramous appendages are homonomous in their construction, but decrease in size towards the posterior end of the body. They consist of a basipodite with ridge-like crescentic endites, an endopod with seven podomeres and a terminal claw, and a lamellae-bearing exopod with a slender shaft. Contrary to previous reports, we confirm the presence of segmental mismatch in Sinoburius lunaris, expressed as diplotergites in the thorax. Maximum parsimony and Bayesian phylogenetic analyses support the monophyly of Xandarellida within Artiopoda, and illuminate the internal relationships within this enigmatic clade. Our results allow us to propose a transformation series explaining the origin of archetypical xandarellid characters, such as the evolution of eye slits in Xandarella spectaculum and Phytophilaspis pergamena as derivates from the anterolateral notches in the head shield observed in Cindarella eucalla and Luohuilinella species. In this context, Sinoburius lunaris is found to feature several derived characters within the group, such as the secondary loss of eye slits and a high degree of appendicular tagmosis. Contrary to previous findings, our analyses strongly support close affinities between Sinoburius lunaris, Xandarella spectaculum and Phytophilaspis pergamena, although the precise relationships between these taxa are sensitive to different methodologies. CONCLUSIONS: The revised morphology of Sinoburius lunaris, made possible through the use of computed tomography to resolve details of its three-dimensionally preserved appendicular anatomy, contributes towards an improved understanding of the morphology of this taxon and the evolution of Xandarellida more broadly. Our results indicate that Sinoburius lunaris possesses an unprecedented degree of appendicular tagmosis otherwise unknown within Artiopoda, with the implication that this iconic group of Palaeozoic euarthropods likely had a more complex ecology and functional morphology than previously considered. The application of computer tomographic techniques to the study of Chengjiang euarthropods holds exceptional promise for understanding the morphological diversity of these organisms, and also better reconstructing their phylogenetic relationships and evolutionary history.

Naked chancelloriids from the lower Cambrian of China show evidence for sponge-type growth.

Chancelloriids are an extinct group of spiny Cambrian animals of uncertain phylogenetic position. Despite their sponge-like body plan, their spines are unlike modern sponge spicules, but share several features with the sclerites of certain Cambrian bilaterians, notably halkieriids. However, a proposed homology of these 'coelosclerites' implies complex transitions in body plan evolution. A new species of chancelloriid, Allonnia nuda, from the lower Cambrian (Stage 3) Chengjiang Lagerstätte is distinguished by its large size and sparse spination, with modified apical sclerites surrounding an opening into the body cavity. The sclerite arrangement in A. nuda and certain other chancelloriids indicates that growth involved sclerite addition in a subapical region, thus maintaining distinct zones of body sclerites and apical sclerites. This pattern is not seen in halkieriids, but occurs in some modern calcarean sponges. With scleritome assembly consistent with a sponge affinity, and in the absence of cnidarian- or bilaterian-grade features, it is possible to interpret chancelloriids as sponges with an unusually robust outer epithelium, strict developmental control of body axis formation, distinctive spicule-like structures and, by implication, minute ostia too small to be resolved in fossils. In this light, chancelloriids may contribute to the emerging picture of high disparity among early sponges.

Water vascular system architecture in an Ordovician ophiuroid.

Understanding the water vascular system (WVS) in early fossil echinoderms is critical to elucidating the evolution of this system in extant forms. Here we present the first report of the internal morphology of the water vascular system of a stem ophiuroid. The radial canals are internal to the arm, but protected dorsally by a plate separate to the ambulacrals. The canals zig-zag with no evidence of constrictions, corresponding to sphincters, which control pairs of tube feet in extant ophiuroids. The morphology suggests that the unpaired tube feet must have operated individually, and relied on the elasticity of the radial canals, lateral valves and tube foot musculature alone for extension and retraction. This arrangement differs radically from that in extant ophiuroids, revealing a previously unknown Palaeozoic configuration.

A Silurian short-great-appendage arthropod.

A new arthropod, Enalikter aphson gen. et sp. nov., is described from the Silurian (Wenlock Series) Herefordshire Lagerstätte of the UK. It belongs to the Megacheira (=short-great-appendage group), which is recognized here, for the first time, in strata younger than mid-Cambrian age. Discovery of this new Silurian taxon allows us to identify a Devonian megacheiran representative, Bundenbachiellus giganteus from the Hunsrück Slate of Germany. The phylogenetic position of megacheirans is controversial: they have been interpreted as stem chelicerates, or stem euarthropods, but when Enalikter and Bundenbachiellus are added to the most comprehensive morphological database available, a stem euarthropod position is supported. Enalikter represents the only fully three-dimensionally preserved stem-group euarthropod, it falls in the sister clade to the crown-group euarthropods, and it provides new insights surrounding the origin and early evolution of the euarthropods. Recognition of Enalikter and Bundenbachiellus as megacheirans indicates that this major arthropod group survived for nearly 100 Myr beyond the mid-Cambrian.

A modern definition of Fossil-Lagerstätten.

Fossil-Lagerstätten are amongst the most important windows onto the paleobiology of ancient ecosystems. Inconsistencies surrounding what constitutes a Lagerstätte limits our ability to compare sites and thus their scientific potential. Here, we provide a modern and utilitarian classification scheme for Konservat-Lagerstätten, allowing for more consistent and improved scientific discourse.

The marine conservation deposits of Monte San Giorgio (Switzerland, Italy): the prototype of Triassic black shale Lagerstätten.

Marine conservation deposits ('Konservat-Lagerstätten') are characterized by their mode of fossil preservation, faunal composition and sedimentary facies. Here, we review these characteristics with respect to the famous conservation deposit of the Besano Formation (formerly Grenzbitumenzone; including the Anisian-Ladinian boundary), and the successively younger fossil-bearing units Cava inferiore, Cava superiore, Cassina beds and the Kalkschieferzone of Monte San Giorgio (Switzerland and Italy). We compare these units to a selection of important black shale-type Lagerstätten of the global Phanerozoic plus the Ediacaran in order to detect commonalities in their facies, genesis, and fossil content using principal component and hierarchical cluster analyses. Further, we put the Monte San Giorgio type Fossillagerstätten into the context of other comparable Triassic deposits worldwide based on their fossil content. The results of the principal component and cluster analyses allow a subdivision of the 45 analysed Lagerstätten into four groups, for which we suggest the use of the corresponding pioneering localities: Burgess type for the early Palaeozoic black shales, Monte San Giorgio type for the Triassic black shales, Holzmaden type for the pyrite-rich black shales and Solnhofen type for platy limestones.

Annelids from the Cambrian (Wuliuan Stage, Miaolingian) Spence Shale Lagerstätte of northern Utah, USA.

The Spence Shale Member of the Langston Formation in northern Utah and southern Idaho preserves generally non-biomineralized fossil assemblages referred to as the Spence Shale Lagerstätte. The biota of this Lagerstätte is dominated by panarthropods, both biomineralized and soft-bodied examples, but also preserves diverse infaunal organisms, including species of scalidophorans, echinoderms, lobopodians, stalked filter feeders, and various problematic taxa. To date, however, only a single annelid fossil, originally assigned to Canadia sp., has been described from the Spence Shale. This lone specimen and another recently collected specimen were analyzed in this study using scanning electron microscopy and energy dispersive X-ray spectrometry. The previous occurrence is reassigned to Burgessochaeta cf. B. setigera Walcott, 1911. The new fossil, however, is identified as a novel polychaete taxon, Shaihuludia shurikeni gen. et sp. nov., characterized by the presence of fused, bladed chaetae and a wide body. The occurrence of Burgessochaeta is the first outside the Burgess Shale and its vicinity, whereas Shaihuludia shurikeni gen. et sp. nov. adds to the diversity of annelids in the middle Cambrian and highlights the diversity of the Spence Shale Lagerstätte.

Herpetogaster collinsi from the Cambrian of China elucidates the dispersal and palaeogeographic distribution of early deuterostomes and the origin of the ambulacrarian larva.

The Cambrian Radiation represents one of the largest diversification events in Earth history. While the resulting taxonomic diversity is exceptional, relatively few of these novel species can be traced outside the boundaries of a single palaeocontinent. Many of those species with cosmopolitan distributions were likely active swimmers, presenting opportunity and means to conquer new areas, but this would not have been the case for sessile organisms. Herpetogaster is a lower to middle Cambrian (Series 2-Miaolingian, Stage 3-Wuliuan) genus of sessile, stalked, filter-feeding deuterostomes with two species, H. collinsi and H. haiyanensis, known respectively from Laurentia and Gondwana. Here, we expand the distribution of H. collinsi to Gondwana with newly discovered specimens from the Balang Formation of Hunan, China. This discovery raises questions on the origin of the genus and how sessile organisms were able to disperse over such a broad distance in the lower Cambrian. As Herpetogaster has been recovered at the base of the Ambulacrarian tree in recent phylogenies, a planktonic larval stage is suggested, which implies, that the last common ancestor of the Ambulacraria might have already had planktonic larvae or that such larvae developed multiple times within the Ambulacraria.

Before trilobite legs: Pygmaclypeatus daziensis reconsidered and the ancestral appendicular organization of Cambrian artiopods.

The Cambrian Stage 3 Chengjiang biota in South China is one of the most influential Konservat-Lagerstätten worldwide thanks to the fossilization of diverse non-biomineralizing organisms through pyritization. Despite their contributions to understanding the evolution of early animals, several Chengjiang species remain poorly known owing to their scarcity and/or incomplete preservation. Here, we use micro-computed tomography to reveal in detail the ventral appendage organization of the enigmatic non-trilobite artiopod Pygmaclypeatus daziensis-one of the rarest euarthropods in Chengjiang-and explore its functional ecology and broader evolutionary significance. Pygmaclypeatus daziensis possesses a set of uniramous antennae and 14 pairs of post-antennal biramous appendages, the latter of which show an unexpectedly high degree of heteronomy based on the localized differentiation of the protopodite, endopodite and exopodite along with the antero-posterior body axis. The small body size (less than 2 cm), the presence of delicate spinose endites and well-developed exopodites with multiple paddle-shaped lamellae on the appendages of P. daziensis indicate a nekto-benthic mode of life and a scavenging/detritus feeding strategy. Pygmaclypeatus daziensis shows that appendage heteronomy is phylogenetically widespread within Artiopoda-the megadiverse clade that includes trilobites and their relatives with non-biomineralizing exoskeletons-and suggests that a single exopodite lobe with paddle-like lamellae is ancestral for this clade. This article is part of the theme issue 'The impact of Chinese palaeontology on evolutionary research'.

Probability-based preservational variations within the early Cambrian Chengjiang biota (China).

The Chengjiang biota (Yunnan Province, China) is a treasure trove of soft-bodied animal fossils from the earliest stages of the Cambrian explosion. The mechanisms contributing to its unique preservation, known as the Burgess Shale-type preservation, are well understood. However, little is known about the preservation differences between various animal groups within this biota. This study compares tissue-occurrence data of 11 major animal groups in the Chengjiang biota using a probabilistic methodology. The fossil-based data from this study is compared to previous decay experiments. This shows that all groups are not equally preserved with some higher taxa more likely to preserve soft tissues than others. These differences in fossil preservation between taxa can be explained by the interaction of biological and environmental characteristics. A bias also results from differential taxonomic recognition, as some taxa are easily recognized from even poorly preserved fragments while other specimens are difficult to assign to higher taxa even with exquisite preservation.

The middle Cambrian Linyi Lagerstätte from the North China Craton: a new window on Cambrian evolutionary fauna.

The rapid appearance of major animal groups and complex marine communities during the Cambrian explosion is recorded in large part in Burgess Shale-type lagerstätten. However, the restricted temporal and spatial distribution of known lagerstätten continues to hinder the formation of a comprehensive perspective on Cambrian evolutionary faunas. Here we describe the Linyi Lagerstätte (ca. 504 mya), a new Cambrian Miaolingian lagerstätte from the Zhangxia Formation in Shandong Province, North China. The Linyi Lagerstätte contains a variety of well-preserved soft-bodied fossils, among which the non-trilobite arthropods, particularly the mollisoniids and radiodonts, are the most important groups. The new assemblage is remarkable for its excellent preservation of arthropod limbs, eyes and guts, as well as for its close similarity in taxonomic composition to Laurentian lagerstätten. The distinctive Linyi Lagerstätte holds great promise for providing additional insights into the morphological disparity, community structure and paleogeographic range of marine faunas during the middle Cambrian (Miaolingian).

The diverse radiodont fauna from the Marjum Formation of Utah, USA (Cambrian: Drumian).

Radiodonts have long been known from Cambrian deposits preserving non-biomineralizing organisms. In Utah, the presence of these panarthropods in the Spence and Wheeler (House Range and Drum Mountains) biotas is now well-documented. Conversely, radiodont occurrences in the Marjum Formation have remained scarce. Despite the large amount of work undertaken on its diverse fauna, only one radiodont (Peytoia) has been reported from the Marjum Biota. In this contribution we quadruple the known radiodont diversity of the Marjum fauna, with the description of the youngest members of two genera, Caryosyntrips and Pahvantia, and that of a new taxon Buccaspinea cooperi gen. et sp. nov. This new taxon can be identified from its large oral cone bearing robust hooked teeth with one, two, or three cusps, and by the unique endite morphology and organisation of its frontal appendages. Appendages of at least 12 podomeres bear six recurved plate-like endites proximal to up to four spiniform distal endites. Pahvantia hastata specimens from the Marjum Formation are particularly large, but otherwise morphologically indistinguishable from the carapace elements of this species found in the Wheeler Formation. One of the two new Caryosyntrips specimens can be confidently assigned to C. camurus. The other bears the largest spines relative to appendage length recorded for this genus, and possesses endites of variable size and unequal spacing, making its taxonomic assignment uncertain. Caryosyntrips, Pahvantia, and Peytoia are all known from the underlying Wheeler Formation, whereas isolated appendages from the Spence Shale and the Wheeler Formation, previously assigned to Hurdia, are tentatively reidentified as Buccaspinea. Notably, none of these four genera occurs in the overlying Weeks Formation, providing supporting evidence of a faunal restructuring around the Drumian-Guzhangian boundary. The description of three additional nektonic taxa from the Marjum Formation further documents the higher relative proportion of free-swimming species in this biota compared to those of the Wheeler and Weeks Lagerstätten. This could be related to a moderate deepening of the basin and/or changing regional ocean circulation at this time.

Taphonomic patterns mimic biologic structures: diagenetic Liesegang rings in Mesozoic coleoids and coprolites.

Because of physiology of coleoids, their fossils preserve soft-tissue-remains more often than other cephalopods. Sometimes, the phosphatized soft-tissues, particularly parts of the muscular mantle, display dark circular patterns. Here, we showcase that these patterns, here documented for fossil coleoids from the Jurassic of Germany and the Cretaceous of Lebanon, superficially resemble chromatophores (which enable living coleoids to alter their coloration). We examined and chemically analyzed the circular structures in these specimens, describe them, and discuss their genesis. Based on their structure and color, we visually differentiate between three types of circles. By comparison with similar structures, we suggest that these structures are not biogenic but Liesegang rings, which formed due to reaction-diffusion processes very soon after death.

Authigenic anatase within 1 billion-year-old cells.

The siliciclastic ~1 Ga-old strata of the Torridon Group, Scotland, contain some of the most exquisitely preserved three-dimensional organic-walled microfossils (OWMs) of the Precambrian. A very diverse microfossil assemblage is hosted in a dominantly phosphatic and clay mineral matrix, within the Diabaig and the Cailleach Head (CH) Formations. In this study, we report on several microfossil taxa within the CH Formation (Leiosphaeridia minutissima, Leiosphaeridia crassa, Synsphaeridium spp. and Myxococcoides spp.) that include populations of cells containing an optically transparent and highly refringent mineral, here identified using electron microscopy as anatase (TiO2 ). Most anatase crystals occur entirely within individual cells, surrounded by unbroken carbonaceous walls. Rarely, an anatase crystal may protrude outside a cell, interpreted to correspond to zones where the cell wall had broken down prior to anatase precipitation. Where an anatase crystal entombs an organic intracellular inclusion (ICI), the ICI is large and well preserved. These combined observations indicate that the intracellular anatase is an authigenic sedimentary phase, making this the first report of in situ precipitated anatase intimately associated with microfossils. The ability of anatase to preserve relatively large volumes of intracellular and cell wall organic material in these cells suggests that the crystallisation of anatase entombed cellular contents particularly quickly, soon after the death of the cell. This is consistent with the strong affinity of Ti for organic material, the low solubility of TiO2 , and reports of Ti occurring in living organisms. With the data currently available, we propose a mineralisation pathway for anatase involving Ti complexation with organic ligands within specific cells, leading to localised post-mortem anatase nucleation inside these cells as the complexes broke down. Further overgrowth of the anatase crystals was likely fuelled by very early diagenetic mobilisation of Ti that had been bound to more labile organic material nearby in the sediments.