This contribution examines the Modern Synthesis in the light of its historical underpinnings, attempts at expansion and treatment of macroevolutionary theory. Particular emphasis is given to the need to better understand the patterns and processes operating on species and higher-level biological entities within a hierarchical framework, as they are often not congruent with lower-level processes. We then focus on the oft-overlooked importance of extrinsic abiotic drivers (e.g. tectonic change, sea-level and climatic fluctuations, as well as factors resulting in mass extinctions) in directing evolution. We propose that the fundamental niche is a species-level property useful in testing for the effects of extrinsic abiotic drivers on macroevolutionary patterns and potential instances of species-level selection. Integration of a diverse array of fields requires both the addition of new information and a standardized theoretical framework for understanding how the effects of change at one level reverberate to other levels. These are some of the remaining ingredients necessary for building a truly synthetic theory of evolution.

The heteromorph ammonite Pravitoceras sigmoidale from the Upper Cretaceous Seidan Formation (Izumi Group) in south-west Japan is frequently encrusted by sessile anomiid bivalves. Fossils of P. sigmoidale with anomiids are often concentrated at the top of or just above turbidite sandstones. Projecting retroversal hooks and apertures of P. sigmoidale are usually intact, and some individuals are associated with jaw apparatuses near apertures. Anomiids are found on both sides and ventral peripheries of P. sigmoidale conchs, attached predominantly to body chambers. These modes of occurrence suggest that the encrustation by anomiids occurred not on post-mortem floating or sunken carcasses but on live conchs and that these organisms were rapidly buried by turbidity current deposits shortly after death. Attachment to both flanks and ventral peripheries of the retroversal hooks may indicate that at least adult individuals of P. sigmoidale did not lie on the sea floor and did not drag their body chambers. It is suggested that fully mature individuals of this ammonite species lived for a long period of time after having formed the retroversal hook because a few generations of anomiids have colonized a single body chamber. Such colonization by anomiids is also observed on Didymoceras awajiense, which is considered to be the closely related ancestral species of P. sigmoidale. This anomiid–heteromorph ammonite commensal relationship might continue to persist in descendants during the course of evolution of these heteromorph ammonites.

Organised mineralised structures observed in large inoceramids (valves on a metre scale) from the Late Albian, Toolebuc Formation, Australia (Inoceramus sutherlandi McCoy, 1865), and the Santonian, Niobrara Formation, USA (Platyceramus sp.), were investigated using variable pressure scanning electron microscope (SEM) with energy-dispersive X-ray spectroscopy (EDX), X-ray microcomputed tomography (micro-CT) and X-ray diffraction (XRD) analyses. These indicate that the structures comprised a phosphate framework of aligned tubes and shallow troughs overlain perpendicularly by evenly spaced structures. In the Toolebuc Inoceramus, these are U-shaped cross-structures, whilst in the Niobrara Platyceramus, they comprise bundled fibre elements. Comparison with modern bivalves indicates that the observed phosphatised structures represent soft-tissue preservation of the gills, as suggested in earlier publications. The tubes and troughs are remnants of a filamental support framework comprising ordinary and primary filaments, whilst the U-shaped cross-structures (I. sutherlandi) and fibrous bands (Platyceramus) represent preserved longitudinal gill musculature. Internal perforate and strand-like fabric observed on the internal surface of some Platyceramus tubular structures suggests that the framework comprised collagen. The presence of primary and ordinary filaments in numerous unusually large plicae, in at least two lamellae, indicates that the gill structures were heterorhabdic. Each plica has at least 40 ordinary filaments, an exceptional number when compared with the maximum of 20 present in modern heterorhabdic gills. The absence of incontrovertible interfilament junctions makes it difficult to say whether inoceramids were filibranch, pseudolamellibranch or eulamellibranch. However, structures that are best attributed to intraplical junctions between filaments suggest the Inoceramidae had gills akin to those of pseudolamellibranch bivalves, although their unusually large number of filaments per plica is more reminiscent of homorhabdic eulamellibranch gills. The general form of the gill is similar to that described in some other pteriomorphs, most specifically Pteria. However, it has more complex junctions and interconnections, although these are not as intricate or pervasive as those observed in the pseudolamellibranch Ostreidae. The connections and well-developed filament framework allowed the gill to reach its unusually large size, supporting the large size of these inoceramid species. The unusually large size of the gill and its components indicate that the organism fed on the larger suspended organic particles in the water column. It would also have been capable of processing large volumes of water quickly, leading to greater potential for food accumulation and with likely implications for respiratory efficiency. This may help explain the common association of inoceramids with oxygen-deficient palaeoenvironments, particularly as the general structure of the inoceramid gill is very different to that observed in the commonest extant chemosymbiotic bivalves.

Calamitalean stems that are preserved in growth position in Carboniferous sedimentary deposits have been described many times in the literature as either pith casts or stem casts. Many of these in situ stems show branching, which gives some information on their patterns of growth. Their manner of preservation is discussed in the light of a new stand of well-preserved in situ stem and branch pith casts of Calamites discovered in mid to upper Duckmantian sandstone at Brymbo in the Wrexham Coalfield of North Wales. Analysis of a brown mineralized layer surrounding the casts and below the black compression remains of the stem tissues has shown the presence of goethite, muscovite, quartz and kaolin. Deposition of these minerals around the inside of the central stem cavity would have provided rigidity and sufficient support, while the pith cavity filled with sediments. The outer tissues would then have been compressed to form a thin coal layer around the mineralized infill of the pith cavity. Cross sections of stems were found clustered together in relatively small areas, and kernel density map and nearest neighbour analysis suggest that each small patch of these pith casts represents an individual plant spread by rhizomatous growth. Stems found in ironstone nodules are external casts of leafy stems preserved by the deposition of siderite on their surfaces. A length of rhizome found at Brymbo was similarly preserved as a cast in ironstone.

The earliest record of a seed with a pappus-like, parachute seed dispersal mechanism, Edenia villisperma gen. et sp. nov., is described from the Upper Triassic of the eastern United States. The seed is small and roughly triangular. Clusters of long hairs emerge from a whorl of at least five circular scars just below the proximal end. This morphology indicates that the hair clusters represent modified lateral structures similar to the pappus of several eudicot angiosperm groups, but probably representing a case of convergent evolution of a similar structure in a gymnosperm. The seeds are usually found isolated, but one specimen indicates that they were born tightly packed together on an axis. A few earlier records exist of dispersal hairs, but this is the first clearly indicating a pappus-like structure. Although the exact affinities of Edenia are uncertain, this seed demonstrates that plants with highly advanced wind dispersal mechanisms occurred at least 55 million years earlier than previously thought.

An adult skeleton of the pistosauroid sauropterygian Yunguisaurus liae reveals a number of morphological features not observed in the holotype, such as the complete morphology of the skull roof, stapes, atlas and axis, ventral view of the postcranium, and nearly complete limbs and tail. Size and morphological differences between the two specimens are mostly regarded as ontogenetic variation, and newly added data did not affect the phylogenetic relationships with other pistosauroids significantly. The number of mesopodia (11 carpals and 8 tarsals) exceeds that known in any other Triassic marine reptiles and does not serve as a precursor of the plesiosaurian pattern with fewer mesopodia of different topology; it demonstrates variability of the limb morphology among the Triassic pistosauroids. The pectoral girdles of Corosaurus, Augustasaurus and Yunguisaurus may indicate early stages of the adaptation towards the plesiosaurian style of paraxial limb movements with ventroposterior power stroke.

Elevated concentrations of an organic compound, 24-isopropylcholestane, found in the Precambrian Huqf Supergroup of Oman may provide the oldest known sponge ‘fossil’. This evidence is of critical importance for a properly balanced understanding of the origin of animals. Several different pelagophyte (Class Pelagophyceae part of the Stramenopiles within the Chromaveolata) algae are also capable of producing these exact compounds, and may similarly have done so in deep time. Modern marine algae are also reported to produce structural isomers that are compositionally identical to the sponge marker; they do this in copious quantities. Further, 24-isopropylcholestane can be produced by diagenetic alteration of compounds produced in large quantities by algae. It is also possible that contamination by petroleum derived lubricating oil used when coring while extracting these compounds from subsurface layers, has affected the data. All extinct organisms that may have produced this compounds are unavailable for analysis by the modern organic chemist and cannot be eliminated from the list of possible producers of the sponge marker. There are also significant uncertainties regarding the dating of the strata from which these ancient compounds are found. Although the compounds are widely reported as c. 751 Ma, they are younger than 645 Ma. It seems more likely that these compounds represent algal biochemical evolution at a time when algal burial occurred in great quantity with well known coeval algal fossils but no sponge fossils. The macroalgal biomass may have declined during the agronomic revolution at the base of the Cambrian Period owing to processing by metazoans, accounting for the comparative scarcity of these sponge markers in Phanerozoic sediments, after which time sponge spicules and body fossils become evident.

The end-Triassic mass extinction (c. 201.6 Ma) was one of the five largest mass-extinction events in the history of animal life. It was also associated with a dramatic, long-lasting change in sedimentation style along the margins of the Tethys Ocean, from generally organic-matter-poor sediments during the Triassic to generally organic-matter-rich black shales during the Jurassic. New core material from Germany provides biomarker evidence of persistent photic-zone euxinia during the Hettangian, the onset of which is associated with a series of both negative and positive carbon isotope excursions. Combined inorganic and organic geochemical and micropalaeontological analyses reveal strong similarities between the Hettangian and the better-known Toarcian anoxic event. These events appear to be the most clearly expressed events within a series of anoxic episodes that also include poorly studied black shale intervals during the Sinemurian and Pliensbachian. Both the Hettangian and Toarcian events are marked by important changes in phytoplankton assemblages from chromophyte- to chlorophyte-dominated assemblages within the European Epicontinental Seaway. Phytoplankton changes occurred in association with the establishment of photic-zone euxinia, driven by a general increase in salinity stratification and warming of surface waters. For both events, the causes of large negative carbon isotope excursions remain incompletely understood; evidence exists for both variation in the δ¹³C of atmospheric CO₂ and variation in the sources of organic carbon. Regardless of the causes of δ¹³C variability, long-term ocean anoxia during the Early Jurassic can be attributed to greenhouse warming and increased nutrient delivery to the oceans triggered by flood basalt volcanism.

Two species of Anomalocaris co-occur in the Emu Bay Shale (Cambrian Series 2, Stage 4) at Big Gully, Kangaroo Island. Frontal appendages of Anomalocaris briggsi Nedin, 1995, are more common than those of Anomalocaris cf. canadensis Whiteaves, 1892, at a quarry inland of the wave-cut platform site from which these species were originally described. An oral cone has the three large, node-bearing plates recently documented for Anomalocaris canadensis, confirming that Anomalocaris lacks a tetraradial ‘Peytoia’ oral cone and strengthening the case for the identity of the Australian specimens as Anomalocaris. Disarticulated anomalocaridid body flaps are more numerous in the Emu Bay Shale than in other localities, and they preserve anatomical details not recognized elsewhere. Transverse lines on the anterior part of the flaps, interpreted as strengthening rays or veins in previous descriptions of anomalocaridids, are associated with internal structures consisting of a series of well-bounded, striated blocks or bars. Their structure is consistent with a structural function imparting strength to the body flaps. Setal structures consisting of a series of lanceolate blades are similar to those of other anomalocaridids and are found in isolation or associated with body flaps. A single specimen also preserves putative gut diverticula. The morphology of the appendages, oral cone, gut diverticula and compound eyes of Anomalocaris, along with its large size, suggests that it was an active predator, and specimens of coprolites containing trilobite fragments and trilobites with prominent injuries have been cited as evidence of anomalocaridid predation on trilobites. Based on frontal appendage morphology, Anomalocaris briggsi is inferred to have been a predator of soft-bodied animals exclusively and only Anomalocaris cf. canadensis may have been capable of durophagous predation on trilobites, although predation (including possible cannibalism) by Redlichia could also explain the coprolites and damage to trilobite exoskeletons found in the Emu Bay Shale.

Protoheliolites is an early heliolitine coral characterized by closely spaced corallites separated in places by sparse coenenchyme. Growth characteristics in the type species, P. norvegicus, are revealed by detailed analysis based on serial peels and thin sections of coralla from the uppermost Katian of north-western Estonia. Colonies of this species had a strong ability to recover from damage and partial mortality, resulting in various forms of rejuvenation, regeneration, fusion and reorganization of corallites; in some cases, this involved relatively large areas of undifferentiated soft parts. The shells of commensal cornulitids became enclosed in host coralla during colony growth. Coralla of P. norvegicus exhibit distinctive growth cycles due to responses to seasonal changes. The production of new corallites by coenenchymal increase usually occurred in low-density bands, in which corallites generally display round to subrounded transverse outlines. In high-density bands, the corallites became crenulated, their wall thickness increased, septal development was more pronounced, and the amount of coenenchyme increased. In addition to these cyclomorphic changes, there were significant astogenetic changes during growth. Compared with the early stage of colony development, distinctive characteristics in the late astogenetic stage include a decrease in the growth rate of the colony, better coordination among corallites, maximum development of corallite crenulations and septa in high-density bands, more numerous coenenchymal tubules and a greater proportion of corallum area occupied by coenenchyme. In general, the role of polyps in determining morphological characteristics of individual corallites, such as tabularium area, corallite crenulations and wall thickness, was subordinate to the astogeny of the colony. Growth characteristics including colony-wide coordination of polyp behaviour and subjugation of individuals to restore the colony following damage suggest a strong astogenetic control and high level of colony integration. Protoheliolites probably arose from a heliolitine genus rather than from a nonheliolitine group as some authors have proposed.

New silicified topotypic material of the Upper Ordovician Streptis undifera (Schmidt, 1858) from the stratotype of the Porkuni Regional Stage in Estonia provides important data on triplesiid morphology and ontogeny, which has substantial implications for our understanding of the affinity of this group of brachiopods. In particular, the new material shows that the early ontogeny of Streptis includes evidence for a cicatrix attachment and colleplax-like structure in the ventral valve. It is likely also that a lecithotrophic feeding habit evolved in triplesiidines sometime in the Ordovician, which sets them apart from all other known strophomenates. A neotype is selected among the specimens of Streptis undifera from the type locality in Porkuni. In Baltoscandia, Streptis undifera appears first in Norway in late Katian below the strata with the pentamerid brachiopod Holorhynchus. In Estonia, Streptis undifera is an index species of the early Hirnantian and occurs in association with stromatoporoid-coral reefs and related inter-reef deposits, which overly the strata with Holorhynchus. In Baltoscandia, the distribution areas of Streptis and Holorhynchus are more or less identical in spite of some differences in age.

Despite an impressive radiation of more than 30 species in the wake of the end-Permian mass extinction, the taxonomic study of Saurichthys has suffered from a lack of universally diagnostic features and a lack of tested quantitative schemes that can be applied to analyse interspecific morphological differences. In this study, we provide an initial quantitative framework for morphological evolution in Saurichthys by focusing on a single bone, the opercle and exploring patterns of interspecific variability in shape using outline-based geometric morphometrics and linear measurements. For the six species examined, comprising 155 specimens and representatives from the Early, Middle and Late Triassic, our results indicate that interspecific shape differences largely reflect an anterior–posterior dimension decrease (= craniocaudal direction) as the dorso-ventral dimension remains similar. In contrast, intraspecific variability in shape is subtle and spread across the outline of the bone, such that counter-acting dimension differences (increase/decrease) were found to occur along a single margin at oblique axes in several species. Our quantitative scheme, which is widely applicable to other groups, provides a useful description of the broad modes of opercle shape change that may help as a starting framework from which to develop character states for opercle morphology in future study.

The morphological variation of stromatoporoids, which are solitary organisms, is partitioned into its presumably genetic and environmental components. Potentially heritable, environmentally mediated and residual components of morphological variability were estimated in a test set containing Devonian stromatoporoids of the genus Gerronostromaria from southern Poland using analysis of variance. The taxonomic importance of traditional morphometric features is limited, because they are dominated by the intra-skeletal component of variance. Conventional metrics were therefore replaced by stereological and textural quantities. Both stereological and textural features are dominated by the inter-skeletal and inter-locality components of variation and thus may be valuable in taxonomic and environmental studies of stromatoporoids. Statistical analyses of these characters (principal component analysis and cluster analysis) were performed. Of 13 characters considered most useful in taxonomic studies, only five have been used previously in conventional species definitions.

In trilobites as in many others extinct organisms, our understanding of the mechanisms of evolution is based on morphological and ontogenetic features. Data from ontogenetic development are essential to provide an insight into the origins of evolutionary changes. In phacopid trilobites, detailed studies of ontogenetic series have been achieved using quantitative methods. A comparison of ontogenetic trajectories of closely related species has been conducted to understand how the both morphological disparity and developmental patterning of different ontogenetic stages were structured and to determine how the environmental or developmental constraints influenced morphological changes in trilobites. Study of such morphological disparity in developmental evolution requires knowledge of morphological features and timing and rate of development among closely related species. At a macroevolutionary scale, such quantitative studies should allow us (1) to determine the impact of individual ontogenies on morphological diversification and (2) to identify the evolutionary patterns in Phacopidae during 100 Ma of existence.

A sea urchin placed on the sea floor near an active brine seep was recovered after 13 years with detailed soft-tissue preservation. Growth of an amorphous calcium carbonate solid with small amounts of the mineral bassanite occurred on the spines and test. The solid also exhibits striations at both the macro- and microscopic scales that preserve the muscle texture of the sea urchin. Such soft-tissue replacement and mineralization could lead to exquisite fossilization. Soft-tissue mineralization has been previously replicated in controlled laboratory conditions; however, this is the first report of the lithologic replication of soft tissues in an open marine experiment. Examples of extraordinary fossil preservation, or Lagersätten, give a distinct snapshot of the past and have led to a greater understanding of the history of life. Soft-tissue lithification occurs in special circumstances in which local chemical conditions (often mediated by decay or bacterial factors) promote early diagenetic mineralization, the first steps of which are observed in this instance. The preservation of articulated skeletons, especially within echinoderms, is normally attributed to rapid burial, but that may not be necessary given that this urchin was at or very near the sediment–water interface for 13 years.

Upper Serpukhovian to lower Bashkirian conodonts studied from the lower and middle part of the Barcaliente Formation type section (NW Spain) are not abundant, but the sedimentary record seems unusually well represented. The first occurrence of Declinognathodus bernesgae occurs more than 140 m below the first occurrence of D. inaequalis, the taxon of the D. noduliferus species group appearing in the bed of the Global Standard Stratotype-section and Point (GSSP) for the Mid-Carboniferous Boundary (Arrow Canyon, Nevada). P1 elements transitional between D. bernesgae and D. inaequalis are described and considered relevant to the global correlation of the Mid-Carboniferous Boundary. In addition, D. praenoduliferus, D. cf. noduliferus, D. lateralis, Idiognathoides asiaticus, I. corrugatus and Neognathodus sp. are described and illustrated from the Barcaliente Formation. A phylogeny is proposed for the early species of these genera on the basis of the morphological changes of the P1 element, where the rostral parapet and area, the surface roughness and the length and depth of the adcarinal troughs are diagnostic characters at the specific level, while caudal ornamentation is a secondary taxonomic character.

The first dicynodont reported for the Permian of South America is described in detail here. The specimen consists of a partial skull and associated lower jaws, collected from the Serra do Cadeado locality, Rio do Rasto Formation, Brazil. The specimen was assigned to the genus Endothiodon Owen in a preliminarily description published in the 1970s. This identification implied a direct correlation with some of the well-established biozones of the Beaufort Group, South Africa, where this taxon is known to occur. Although badly weathered, the Brazilian material preserves several features that allow us to confirm its assignment to Endothiodon, such as the pineal foramen situated on a boss, the bulbous swellings of the dentaries, a boss situated on the ventral margin of the jugal, the extensive number of dentary teeth and the upturned pointed beak of the lower jaw. However, this set of characters is not enough to establish a specific taxonomic assignment for the specimen, although it seems to be more closely related to E. bathystoma, E. uniseries and E. whaitsi than to E. mahalanobisi. The biostratigraphical correlations proposed for the tetrapod faunas of the Rio do Rasto Formation highlight the similarities between these associations and others from the Mid and Late Permian of South Africa and Eastern Europe. However, until now, these correlations are tentative because only a few faunal elements have been recovered from Brazil, and most of them lack stratigraphic data on the levels from where they were collected. Due to the presence of Endothiodon, at least part of the Rio do Rasto Formation can be correlated with deposits in India, Malawi, Mozambique, South Africa, Tanzania, Zambia and Zimbabwe.

Phalarodon atavus from the Germanic Muschelkalk Basin was previously represented only by cranial elements. Here we report a nearly complete and articulated specimen of P. atavus from the Middle Triassic Luoping Biota, Yunnan, South China. This is the first specimen of P. atavus from outside the Germanic Basin. This discovery demonstrates a peri-Tethyan distribution of P. atavus. The new specimen is also the first one preserving the postcranial anatomy of this species, providing the opportunity to evaluate its sustained swimming ability. Inferences made on its functional morphology suggest that this species was probably adapted for active foraging. Tooth crown morphology suggests that P. atavus may have preferred externally soft prey.

Pinnatiramosus qianensis Geng, 1986, is a plant with a complex, extensive pinnate branching system and pitted tracheids, collected from marine Lower Silurian (Llandovery; c. 430 Ma) rocks in Guizhou Province, China. It challenges long-held theories on the origin and early evolution of vascular plants in the Silurian and Devonian. However, there is a hypothesis that the fossils were not syngenetic with the entombing rock, but were the rooting systems of much younger plants, probably of Permian age. New sections and collections of P. qianensis have been subjected to detailed analyses, which indicate that P. qianensis belongs to an early Permian (c. 285 Ma) rooting system growing down into lower Silurian rocks.

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.

Biomineralized tissues are chemically altered after death, and this diagenetic alteration can obscure original biological chemical features or provide new chemical information about the depositional environment. To use the chemistry of fossil biominerals to reconstruct biological, environmental or taphonomic information, a solid appreciation of biomineralization, mineral diagenesis and biomineral–water interaction is needed. Here, I summarize the key recent developments in the fields of biomineralization and post-mortem trace element exchange that have significant implications for our understanding of the diagenetic behaviour of biominerals and the ways in which biomineral chemistry can be used in palaeontological and taphonomic research.

Ecosystems today are under growing pressure, with human domination at many scales. It is difficult, however, to gauge what has changed or been lost – and why – in the absence of data from periods before human activities. Actualistic taphonomic studies, originally motivated to understand preservational controls on deep-time fossil records, are now providing insights into modern death assemblages as historical archives of present-day ecosystems, turning taphonomy on its head. This article reviews the past 20 years of work on the temporal resolution and ability of time-averaged skeletal assemblages to capture ecological information faithfully, focusing primarily on molluscs from soft-sediment seafloors. Two promising arenas for ‘applied taphonomy’ are then highlighted: (1) using live-dead mismatch – that is, observed discordance in the diversity, species composition, and distribution of living animals and co-occurring skeletal remains – to recognize recent anthropogenic change, and (2) using time-averaged death assemblages as windows into regional diversity and long-term baselines, as a supplement or substitute for conventional live-collected data. Meta-analysis and modelling find that, in unaltered habitats, live-dead differences in community-level attributes can be generated largely or entirely by time-averaging of natural spatial and temporal variability in living assemblages, on time frames consistent with the range of shell ages observed in death assemblages. Time-averaging coarsens the temporal and spatial resolution of biological information in predictable ways; by comparison, taphonomic bias of information arising from differential preservation, production and transport of shells is surprisingly modest. Several challenges remain for basic taphonomic research, such as empirical and analytical methods of refining the temporal resolution of death assemblages; assessing the fate of resolution and fidelity with progressive burial; and expanding our understanding of the dynamics of skeletal accumulation in other groups and settings. Rather than shunning human-impacted areas as inappropriate analogues of the deep past, we should capitalize on them to explore the fundamental controls on skeletal accumulation and to develop robust protocols for bringing time-averaged death assemblages into the toolkits of conservation biology and environmental management.

Siliceous hot spring deposits form at Earth's surface above terrestrial hydrothermal systems, which create low-sulphidation epithermal mineral deposits deeper in the crust. Eruption of hot spring waters and precipitation of opal-A create sinter apron complexes and areas of geothermally influenced wetland. These provide habitat for higher plants that may be preserved in situ, by encrustation of their surfaces and permineralization of tissues, creating T⁰ plant assemblages. In this study, we review the fossil record of hot spring floras from subfossil examples forming in active hot spring areas, via fossil examples from the Cenozoic, Mesozoic and Palaeozoic to the oldest known hot spring flora, the Lower Devonian Rhynie chert. We demonstrate that the well-known megabias towards wetland plant preservation extends to hot spring floras. We highlight that the record of hot spring floras is dominated by plants preserved in situ by permineralization on geothermally influenced wetlands. Angiosperms (members of the Cyperaceae and Restionaceae) dominate Cenozoic floras. Equisetum and gleicheniaceous ferns colonized Mesozoic (Jurassic) geothermal wetlands and sphenophytes and herbaceous lycophytes late Palaeozoic examples. Evidence of the partitioning of wetland hydrophytic and dryland mesophytic communities, a feature of active geothermal areas, is provided by well-preserved and well-exposed fossil sinter apron complexes, which record flooding of dryland environments by thermal waters and decline of local forest ecosystems. Such observations from the fossil record back-up hypotheses based on active hot springs and vegetation that suggest removal of taphonomic filtering in hot spring environments is accompanied by an increase in ecological and ecophysiological filtering. To this end we also demonstrate that in the hot spring environment, the wetland bias extends beyond broad ecology. We show that ecosystems preserved from the Cenozoic to the Mesozoic provide clear evidence that the dominant plants preserved in situ by hot spring activity are also halophytic, tolerant of high pH and high concentrations of heavy metals. By extension, we hypothesize that this is also the case in Palaeozoic hot spring settings and extended to the early land plant flora of the Rhynie chert.

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.