A new small duckbilled dinosaur (Hadrosauridae: Lambeosaurinae) from Morocco and dinosaur diversity in the late Maastrichtian of North Africa.

In the Late Cretaceous, northern and southern hemispheres evolved distinct dinosaurian faunas. Titanosaurians and abelisaurids dominated the Gondwanan continents; hadrosaurids, ceratopsians and tyrannosaurs dominated North America and Asia. Recently, a lambeosaurine hadrosaurid, Ajnabia odysseus, was reported from the late Maastrichtian phosphates of the Oulad Abdoun Basin Morocco, suggesting dispersal between Laurasia and Gondwana. Here we report new fossils from the phosphates of Morocco showing lambeosaurines achieved high diversity in the late Maastrichtian of North Africa. A skull represents a new dwarf lambeosaurine, Minqaria bata. Minqaria resembles Ajnabia odysseus in size, but differs in the ventrally positioned jugal facet and sinusoidal toothrow. The animal is small, ~ 3.5 m long, but the fused braincase shows it was mature. A humerus and a femur belong to larger hadrosaurids, ~ 6 m long, implying at least three species coexisted. The diversity of hadrosaurids in Europe and Africa suggests a dispersal-driven radiation, with lambeosaurines diversifying to take advantage of low ornithischian diversity. African lambeosaurines are small compared to North American and Asia hadrosaurids however, perhaps due to competition with titanosaurians. Hadrosaurids are unknown from eastern Africa, suggesting Moroccan hadrosaurids may be part of a distinct insular fauna, and represent an island radiation.

A giant tyrannosaur from the Campanian-Maastrichtian of southern North America and the evolution of tyrannosaurid gigantism.

Tyrannosaurid dinosaurs dominated as predators in the Late Cretaceous of Laurasia, culminating in the evolution of the giant Tyrannosaurus rex, both the last and largest tyrannosaurid. Where and when Tyrannosaurini (T. rex and kin) originated remains unclear. Competing hypotheses place tyrannosaurin origins in Asia, or western North America (Laramidia). We report a new tyrannosaurin, Tyrannosaurus mcraeensis, from the Campanian-Maastrichtian Hall Lake Formation of New Mexico, based on a fossil previously referred to T. rex. T. mcraeensis predates T. rex by ~ 6-7 million years, yet rivaled it in size. Phylogenetic analysis recovers T. mcraeensis as sister to T. rex and suggests Tyrannosaurini originated in southern Laramidia. Evolution of giant tyrannosaurs in southern North America, alongside giant ceratopsians, hadrosaurs, and titanosaurs suggests large-bodied dinosaurs evolved at low latitudes in North America.

Evolution and dispersal of snakes across the Cretaceous-Paleogene mass extinction.

Mass extinctions have repeatedly shaped global biodiversity. The Cretaceous-Paleogene (K-Pg) mass extinction caused the demise of numerous vertebrate groups, and its aftermath saw the rapid diversification of surviving mammals, birds, frogs, and teleost fishes. However, the effects of the K-Pg extinction on the evolution of snakes-a major clade of predators comprising over 3,700 living species-remains poorly understood. Here, we combine an extensive molecular dataset with phylogenetically and stratigraphically constrained fossil calibrations to infer an evolutionary timescale for Serpentes. We reveal a potential diversification among crown snakes associated with the K-Pg mass extinction, led by the successful colonisation of Asia by the major extant clade Afrophidia. Vertebral morphometrics suggest increasing morphological specialisation among marine snakes through the Paleogene. The dispersal patterns of snakes following the K-Pg underscore the importance of this mass extinction event in shaping Earth's extant vertebrate faunas.

Cretaceous dinosaur bone contains recent organic material and provides an environment conducive to microbial communities.

Fossils were thought to lack original organic molecules, but chemical analyses show that some can survive. Dinosaur bone has been proposed to preserve collagen, osteocytes, and blood vessels. However, proteins and labile lipids are diagenetically unstable, and bone is a porous open system, allowing microbial/molecular flux. These 'soft tissues' have been reinterpreted as biofilms. Organic preservation versus contamination of dinosaur bone was examined by freshly excavating, with aseptic protocols, fossils and sedimentary matrix, and chemically/biologically analyzing them. Fossil 'soft tissues' differed from collagen chemically and structurally; while degradation would be expected, the patterns observed did not support this. 16S rRNA amplicon sequencing revealed that dinosaur bone hosted an abundant microbial community different from lesser abundant communities of surrounding sediment. Subsurface dinosaur bone is a relatively fertile habitat, attracting microbes that likely utilize inorganic nutrients and complicate identification of original organic material. There exists potential post-burial taphonomic roles for subsurface microorganisms.

The chances of establishing a real-world Jurassic Park are slim. During the fossilization process, biological tissues degrade over millions of years, with some types of molecules breaking down faster than others. However, traces of biological material have been found inside some fossils. While some researchers believe these could be the remains of ancient proteins, blood vessels, and cells, traditionally thought to be among the least stable components of bone, others think that they have more recent sources. One hypothesis is that they are in fact biofilms formed by bacteria. To investigate the source of the biological material in fossil bone, Saitta et al. performed a range of analyses on the fossilized bones of a horned dinosaur called Centrosaurus. The bones were carefully excavated in a manner to reduce contamination, and the sediment the bones had been embedded in was also tested for comparison. Saitta et al. found no evidence of ancient dinosaur proteins. However, the fossils contained more organic carbon, DNA, and certain amino acids than the sediment surrounding them. Most of these appeared to have a very recent source. Sequencing the genetic material revealed that the fossil had become a habitat for an unusual community of microbes that is not found in the surrounding sediment or above ground. These buried microbes may have evolved unique ways to thrive inside fossils. Future work could investigate how these unusual organisms live and whether the communities vary in different parts of the world.

Correction: Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary.

[This corrects the article DOI: 10.1371/journal.pbio.2001663.].

Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary.

Pterosaurs were the first vertebrates to evolve powered flight and the largest animals to ever take wing. The pterosaurs persisted for over 150 million years before disappearing at the end of the Cretaceous, but the patterns of and processes driving their extinction remain unclear. Only a single family, Azhdarchidae, is definitively known from the late Maastrichtian, suggesting a gradual decline in diversity in the Late Cretaceous, with the Cretaceous-Paleogene (K-Pg) extinction eliminating a few late-surviving species. However, this apparent pattern may simply reflect poor sampling of fossils. Here, we describe a diverse pterosaur assemblage from the late Maastrichtian of Morocco that includes not only Azhdarchidae but the youngest known Pteranodontidae and Nyctosauridae. With 3 families and at least 7 species present, the assemblage represents the most diverse known Late Cretaceous pterosaur assemblage and dramatically increases the diversity of Maastrichtian pterosaurs. At least 3 families-Pteranodontidae, Nyctosauridae, and Azhdarchidae-persisted into the late Maastrichtian. Late Maastrichtian pterosaurs show increased niche occupation relative to earlier, Santonian-Campanian faunas and successfully outcompeted birds at large sizes. These patterns suggest an abrupt mass extinction of pterosaurs at the K-Pg boundary.

Maaqwi cascadensis: A large, marine diving bird (Avialae: Ornithurae) from the Upper Cretaceous of British Columbia, Canada.

Mesozoic bird fossils from the Pacific Coast of North America are rare, but small numbers are known from the Late Cretaceous aged sediments of Hornby Island, British Columbia. Most are unassociated fragments that offer little information, but additional preparation of a large coracoid has revealed more details of its structure, as well as three associated wing bones. Phylogenetic analysis suggests that Maaqwi cascadensis, gen. et sp. nov. represents a derived crown or near-crown member of Ornithurae, and specifically suggests affinities with Vegaviidae. M. cascadensis is characterized by large size, and regressions based on dimensions of the coracoid suggest a large bird, with an estimated body mass of approximately 1.5 kilograms. The bones are robust, with thick walls, suggesting that M. cascadensis was a bird adapted for diving, similar to modern loons and grebes. The wings are short, while the coracoid is unusually short and broad, similar to modern loons. Along with the Ichthyornithes and Hesperornithes, M. cascadensis and Vegaviidae appear to represent a third clade of bird that evolved to exploit marine habitats in the Late Cretaceous, one specialized for foot-propelled diving and rapid cruising flight over water.

The first oviraptorosaur (Dinosauria: Theropoda) bonebed: evidence of gregarious behaviour in a maniraptoran theropod.

A monodominant bonebed of Avimimus from the Nemegt Formation of Mongolia is the first oviraptorosaur bonebed described and the only recorded maniraptoran bonebed from the Late Cretaceous. Cranial elements recovered from the bonebed provide insights on the anatomy of the facial region, which was formerly unknown in Avimimus. Both adult and subadult material was recovered from the bonebed, but small juveniles are underrepresented. The taphonomic and sedimentological evidence suggests that the Avimimus bonebed represents a perimortem gregarious assemblage. The near absence of juveniles in the bonebed may be evidence of a transient age-segregated herd or 'flock', but the behaviour responsible for this assemblage is unclear. Regardless, the Avimimus bonebed is the first evidence of gregarious behaviour in oviraptorosaurs, and highlights a potential trend of increasing gregariousness in dinosaurs towards the end of the Mesozoic.

EVOLUTION. A four-legged snake from the Early Cretaceous of Gondwana.

Snakes are a remarkably diverse and successful group today, but their evolutionary origins are obscure. The discovery of snakes with two legs has shed light on the transition from lizards to snakes, but no snake has been described with four limbs, and the ecology of early snakes is poorly known. We describe a four-limbed snake from the Early Cretaceous (Aptian) Crato Formation of Brazil. The snake has a serpentiform body plan with an elongate trunk, short tail, and large ventral scales suggesting characteristic serpentine locomotion, yet retains small prehensile limbs. Skull and body proportions as well as reduced neural spines indicate fossorial adaptation, suggesting that snakes evolved from burrowing rather than marine ancestors. Hooked teeth, an intramandibular joint, a flexible spine capable of constricting prey, and the presence of vertebrate remains in the guts indicate that this species preyed on vertebrates and that snakes made the transition to carnivory early in their history. The structure of the limbs suggests that they were adapted for grasping, either to seize prey or as claspers during mating. Together with a diverse fauna of basal snakes from the Cretaceous of South America, Africa, and India, this snake suggests that crown Serpentes originated in Gondwana.

Biogeography of worm lizards (Amphisbaenia) driven by end-Cretaceous mass extinction.

Worm lizards (Amphisbaenia) are burrowing squamates that live as subterranean predators. Their underground existence should limit dispersal, yet they are widespread throughout the Americas, Europe and Africa. This pattern was traditionally explained by continental drift, but molecular clocks suggest a Cenozoic diversification, long after the break-up of Pangaea, implying dispersal. Here, we describe primitive amphisbaenians from the North American Palaeocene, including the oldest known amphisbaenian, and provide new and older molecular divergence estimates for the clade, showing that worm lizards originated in North America, then radiated and dispersed in the Palaeogene following the Cretaceous-Palaeogene (K-Pg) extinction. This scenario implies at least three trans-oceanic dispersals: from North America to Europe, from North America to Africa and from Africa to South America. Amphisbaenians provide a striking case study in biogeography, suggesting that the role of continental drift in biogeography may be overstated. Instead, these patterns support Darwin and Wallace's hypothesis that the geographical ranges of modern clades result from dispersal, including oceanic rafting. Mass extinctions may facilitate dispersal events by eliminating competitors and predators that would otherwise hinder establishment of dispersing populations, removing biotic barriers to dispersal.

A suspension-feeding anomalocarid from the Early Cambrian.

Large, actively swimming suspension feeders evolved several times in Earth's history, arising independently from groups as diverse as sharks, rays and stem teleost fishes, and in mysticete whales. However, animals occupying this niche have not been identified from the early Palaeozoic era. Anomalocarids, a group of stem arthropods that were the largest nektonic animals of the Cambrian and Ordovician periods, are generally thought to have been apex predators. Here we describe new material from Tamisiocaris borealis, an anomalocarid from the Early Cambrian (Series 2) Sirius Passet Fauna of North Greenland, and propose that its frontal appendage is specialized for suspension feeding. The appendage bears long, slender and equally spaced ventral spines furnished with dense rows of long and fine auxiliary spines. This suggests that T. borealis was a microphagous suspension feeder, using its appendages for sweep-net capture of food items down to 0.5 mm, within the size range of mesozooplankton such as copepods. Our observations demonstrate that large, nektonic suspension feeders first evolved during the Cambrian explosion, as part of an adaptive radiation of anomalocarids. The presence of nektonic suspension feeders in the Early Cambrian, together with evidence for a diverse pelagic community containing phytoplankton and mesozooplankton, indicate the existence of a complex pelagic ecosystem supported by high primary productivity and nutrient flux. Cambrian pelagic ecosystems seem to have been more modern than previously believed.

Distributions of cranial pathologies provide evidence for head-butting in dome-headed dinosaurs (Pachycephalosauridae).

Pachycephalosaurids are small, herbivorous dinosaurs with domed skulls formed by massive thickening of the cranial roof. The function of the dome has been a focus of debate: the dome has variously been interpreted as the product of sexual selection, as an adaptation for species recognition, or as a weapon employed in intraspecific combat, where it was used in butting matches as in extant ungulates. This last hypothesis is supported by the recent identification of cranial pathologies in pachycephalosaurids, which appear to represent infections resulting from trauma. However, the frequency and distribution of pathologies have not been studied in a systematic fashion. Here, we show that pachycephalosaurids are characterized by a remarkably high incidence of cranial injury, where 22% of specimens have lesions on the dome. Frequency of injury shows no significant difference between different genera, but flat-headed morphs (here interpreted as juveniles or females) lack lesions. Mapping of injuries onto a digitial pachycephalosaurid skull shows that although lesions are distributed across the dome, they cluster near the apex, which is consistent with the hypothesis that the dome functioned for intraspecific butting matches.

Mass extinction of lizards and snakes at the Cretaceous-Paleogene boundary.

The Cretaceous-Paleogene (K-Pg) boundary is marked by a major mass extinction, yet this event is thought to have had little effect on the diversity of lizards and snakes (Squamata). A revision of fossil squamates from the Maastrichtian and Paleocene of North America shows that lizards and snakes suffered a devastating mass extinction coinciding with the Chicxulub asteroid impact. Species-level extinction was 83%, and the K-Pg event resulted in the elimination of many lizard groups and a dramatic decrease in morphological disparity. Survival was associated with small body size and perhaps large geographic range. The recovery was prolonged; diversity did not approach Cretaceous levels until 10 My after the extinction, and resulted in a dramatic change in faunal composition. The squamate fossil record shows that the end-Cretaceous mass extinction was far more severe than previously believed, and underscores the role played by mass extinctions in driving diversification.

Primitive wing feather arrangement in Archaeopteryx lithographica and Anchiornis huxleyi.

In modern birds (Neornithes), the wing is composed of a layer of long, asymmetrical flight feathers overlain by short covert feathers. It has generally been assumed that wing feathers in the Jurassic bird Archaeopteryx and Cretaceous feathered dinosaurs had the same arrangement. Here, we redescribe the wings of the archaic bird Archaeopteryx lithographica and the dinosaur Anchiornis huxleyi and show that their wings differ from those of Neornithes in being composed of multiple layers of feathers. In Archaeopteryx, primaries are overlapped by long dorsal and ventral coverts. Anchiornis has a similar configuration but is more primitive in having short, slender, symmetrical remiges. Archaeopteryx and Anchiornis therefore appear to represent early experiments in the evolution of the wing. This primitive configuration has important functional implications: although the slender feather shafts of Archaeopteryx and Anchiornis make individual feathers weak, layering of the wing feathers may have produced a strong airfoil. Furthermore, the layered arrangement may have prevented the feathers from forming a slotted tip or separating to reduce drag on the upstroke. The wings of early birds therefore may have lacked the range of functions seen in Neornithes, limiting their flight ability.

A transitional snake from the Late Cretaceous period of North America.

Snakes are the most diverse group of lizards, but their origins and early evolution remain poorly understood owing to a lack of transitional forms. Several major issues remain outstanding, such as whether snakes originated in a marine or terrestrial environment and how their unique feeding mechanism evolved. The Cretaceous Coniophis precedens was among the first Mesozoic snakes discovered, but until now only an isolated vertebra has been described and it has therefore been overlooked in discussions of snake evolution. Here we report on previously undescribed material from this ancient snake, including the maxilla, dentary and additional vertebrae. Coniophis is not an anilioid as previously thought a revised phylogenetic analysis of Ophidia shows that it instead represents the most primitive known snake. Accordingly, its morphology and ecology are critical to understanding snake evolution. Coniophis occurs in a continental floodplain environment, consistent with a terrestrial rather than a marine origin; furthermore, its small size and reduced neural spines indicate fossorial habits, suggesting that snakes evolved from burrowing lizards. The skull is intermediate between that of lizards and snakes. Hooked teeth and an intramandibular joint indicate that Coniophis fed on relatively large, soft-bodied prey. However, the maxilla is firmly united with the skull, indicating an akinetic rostrum. Coniophis therefore represents a transitional snake, combining a snake-like body and a lizard-like head. Subsequent to the evolution of a serpentine body and carnivory, snakes evolved a highly specialized, kinetic skull, which was followed by a major adaptive radiation in the Early Cretaceous period. This pattern suggests that the kinetic skull was a key innovation that permitted the diversification of snakes.

Torosaurus is not Triceratops: ontogeny in chasmosaurine ceratopsids as a case study in dinosaur taxonomy.

BACKGROUND: In horned dinosaurs, taxonomy is complicated by the fact that the cranial ornament that distinguishes species changes with age. Based on this observation, it has been proposed that the genera Triceratops and Torosaurus are in fact synonymous, with specimens identified as Torosaurus representing the adult form of Triceratops. The hypothesis of synonymy makes three testable predictions: 1) the species in question should have similar geographic and stratigraphic distributions, 2) specimens assigned to Torosaurus should be more mature than those assigned to Triceratops, and 3) intermediates should exist that combine features of Triceratops and Torosaurus. The first condition appears to be met, but it remains unclear whether the other predictions are borne out by the fossil evidence. METHODOLOGY/PRINCIPAL FINDINGS: We assessed the relative maturity of Torosaurus and Triceratops specimens by coding skulls for characters that vary with maturity, and then using a clustering analysis to arrange them into a growth series. We found that a well-defined sequence of changes exists in horned dinosaurs: development of cranial ornament occurs in juveniles, followed by fusion of the skull roof in subadults, and finally, the epoccipitals, epijugals, and rostral fuse to the skull in adults. Using this scheme, we identified mature and immature individuals of both Torosaurus and Triceratops. Furthermore, we describe the ventral depressions on the frill of Triceratops, and show that they differ in shape and position from the parietal fenestrae of Torosaurus. Thus, we conclude that these structures are not intermediates between the solid frill of Triceratops and the fenestrated frill of Torosaurus. CONCLUSIONS/SIGNIFICANCE: Torosaurus is a distinct genus of horned dinosaur, not the adult of Triceratops. Our method provides a framework for assessing the hypothesis of synonymy through ontogeny in the fossil record.

Mass extinction of birds at the Cretaceous-Paleogene (K-Pg) boundary.

The effect of the Cretaceous-Paleogene (K-Pg) (formerly Cretaceous-Tertiary, K-T) mass extinction on avian evolution is debated, primarily because of the poor fossil record of Late Cretaceous birds. In particular, it remains unclear whether archaic birds became extinct gradually over the course of the Cretaceous or whether they remained diverse up to the end of the Cretaceous and perished in the K-Pg mass extinction. Here, we describe a diverse avifauna from the latest Maastrichtian of western North America, which provides definitive evidence for the persistence of a range of archaic birds to within 300,000 y of the K-Pg boundary. A total of 17 species are identified, including 7 species of archaic bird, representing Enantiornithes, Ichthyornithes, Hesperornithes, and an Apsaravis-like bird. None of these groups are known to survive into the Paleogene, and their persistence into the latest Maastrichtian therefore provides strong evidence for a mass extinction of archaic birds coinciding with the Chicxulub asteroid impact. Most of the birds described here represent advanced ornithurines, showing that a major radiation of Ornithurae preceded the end of the Cretaceous, but none can be definitively referred to the Neornithes. This avifauna is the most diverse known from the Late Cretaceous, and although size disparity is lower than in modern birds, the assemblage includes both smaller forms and some of the largest volant birds known from the Mesozoic, emphasizing the degree to which avian diversification had proceeded by the end of the age of dinosaurs.

Cranial ontogeny in Stegoceras validum (Dinosauria: Pachycephalosauria): a quantitative model of pachycephalosaur dome growth and variation.

Historically, studies of pachycephalosaurs have recognized plesiomorphically flat-headed taxa and apomorphically domed taxa. More recently, it has been suggested that the expression of the frontoparietal dome is ontogenetic and derived from a flat-headed juvenile morphology. However, strong evidence to support this hypothesis has been lacking. Here we test this hypothesis in a large, stratigraphically constrained sample of specimens assigned to Stegoceras validum, the best known pachycephalosaur, using multiple independent lines of evidence including conserved morphology of ornamentation, landmark-based allometric analyses of frontoparietal shape, and cranial bone histology. New specimens show that the diagnostic ornamentation of the parietosquamosal bar is conserved throughout the size range of the sample, which links flat-headed specimens to domed S. validum. High-resolution CT scans of three frontoparietals reveal that vascularity decreases with size and document a pattern that is consistent with previously proposed histological changes during growth. Furthermore, aspects of dome shape and size are strongly correlated and indicative of ontogenetic growth. These results are complementary and strongly support the hypothesis that the sample represents a growth series of a single taxon. Cranial dome growth is positively allometric, proceeds from a flat-headed to a domed state, and confirms the synonymy of Ornatotholus browni as a juvenile Stegoceras. This dataset serves as the first detailed model of growth and variation in a pachycephalosaur. Flat-headed juveniles possess three characters (externally open cranial sutures, tuberculate dorsal surface texture, and open supratemporal fenestrae) that are reduced or eliminated during ontogeny. These characters also occur in putative flat-headed taxa, suggesting that they may also represent juveniles of domed taxa. However, open cranial sutures and supratemporal fenestrae are plesiomorphic within Ornithischia, and thus should be expected in the adult stage of a primitive pachycephalosaur. Additional lines of evidence will be needed to resolve the taxonomic validity of flat-headed pachycephalosaur taxa.

The bizarre wing of the Jamaican flightless ibis Xenicibis xympithecus: a unique vertebrate adaptation.

Birds have frequently evolved to exploit insular environments by becoming adapted to a terrestrial lifestyle and losing the ability to fly, usually via reducing the wings and pectoral girdle. The enigmatic flightless ibis Xenicibis xympithecus (Threskiornithidae) from the Quaternary of Jamaica provides a rare example of flight loss in ibises. We report on previously undescribed fossils of Xenicibis, and show that the wing differed radically from that of all other birds, flightless or volant. The metacarpus is elongate, grotesquely inflated and has extremely thick walls; phalanges are short and block-like; the radius is distally expanded; and the humerus is elongate. The furcula, coracoid and sternum are all well developed. We propose that the elongate forelimb and massive hand functioned in combat as a jointed club or flail. This hypothesis is supported by the morphology of the carpometacarpus, by features permitting rapid extension of the wing and by the presence of fractures in wing bones. Although other birds use the wings as weapons, none resemble Xenicibis, which represents a unique and extraordinary morphological solution to this functional problem. Xenicibis strikingly illustrates how similar selective pressures, acting on a similar starting point, can result in novel outcomes.

Spatial niche partitioning in dinosaurs from the latest cretaceous (Maastrichtian) of North America.

We examine patterns of occurrence of associated dinosaur specimens (n = 343) from the North American Upper Cretaceous Hell Creek Formation and equivalent beds, by comparing their relative abundance in sandstone and mudstone. Ceratopsians preferentially occur in mudstone, whereas hadrosaurs and the small ornithopod Thescelosaurus show a strong association with sandstone. By contrast, the giant carnivore Tyrannosaurus rex shows no preferred association with either lithology. These lithologies are used as an indicator of environment of deposition, with sandstone generally representing river environments, and finer grained sediments typically representing floodplain environments. Given these patterns of occurrence, we argue that spatial niche partitioning helped reduce competition for resources between the herbivorous dinosaurs. Within coastal lowlands ceratopsians preferred habitats farther away from rivers, whereas hadrosaurs and Thescelosaurus preferred habitats in close proximity to rivers, and T. rex, the ecosystem's sole large carnivore, inhabited both palaeoenvironments. Spatial partitioning of the environment helps explain how several species of large herbivorous dinosaurs coexisted. This study emphasizes that different lithologies can preserve dramatically dissimilar vertebrate assemblages, even when deposited in close proximity and within a narrow window of time. The lithology in which fossils are preserved should be recorded as these data can provide unique insights into the palaeoecology of the animals they preserve.