Patrick J. Orr

Ordovician faunas of Burgess Shale type

The renowned soft-bodied faunas of the Cambrian period, which include the Burgess Shale, disappear from the fossil record in the late Middle Cambrian, after which the Palaeozoic fauna dominates. The disappearance of faunas of Burgess Shale type curtails the stratigraphic record of a number of iconic Cambrian taxa. One possible explanation for this loss is a major extinction, but more probably it reflects the absence of preservation of similar soft-bodied faunas in later periods. Here we report the discovery of numerous diverse soft-bodied assemblages in the Lower and Upper Fezouata Formations (Lower Ordovician) of Morocco, which include a range of remarkable stem-group morphologies normally considered characteristic of the Cambrian. It is clear that biotas of Burgess Shale type persisted after the Cambrian and are preserved where suitable facies occur. The Fezouata biota provides a link between the Burgess Shale communities and the early stages of the Great Ordovician Biodiversification Event.

Fossilized melanosomes and the colour of Cretaceous dinosaurs and birds

Spectacular fossils from the Early Cretaceous Jehol Group of northeastern China have greatly expanded our knowledge of the diversity and palaeobiology of dinosaurs and early birds, and contributed to our understanding of the origin of birds, of flight, and of feathers. Pennaceous (vaned) feathers and integumentary filaments are preserved in birds and non-avian theropod dinosaurs, but little is known of their microstructure. Here we report that melanosomes (colour-bearing organelles) are not only preserved in the pennaceous feathers of early birds, but also in an identical manner in integumentary filaments of non-avian dinosaurs, thus refuting recent claims that the filaments are partially decayed dermal collagen fibres. Examples of both eumelanosomes and phaeomelanosomes have been identified, and they are often preserved in life position within the structure of partially degraded feathers and filaments. Furthermore, the data here provide empirical evidence for reconstructing the colours and colour patterning of these extinct birds and theropod dinosaurs: for example, the dark-coloured stripes on the tail of the theropod dinosaur Sinosauropteryx can reasonably be inferred to have exhibited chestnut to reddish-brown tones.

The arthropod Offacolus kingi (Chelicerata) from the Silurian of Herefordshire, England: computer based morphological reconstructions and phylogenetic affinities

The small, non–biomineralized, three–dimensionally preserved arthropod Offacolus kingi Orr et al. from the Wenlock Series (Silurian) of Herefordshire, England, is re–evaluated, and the new family Offacolidae erected. This new study is based on specimens which have been serially ground, reconstructed by computer and rendered in the round as coloured models. Offacolus possesses a prosomal appendage array similar to that of Limulus, but also bears robust and setose exopods on appendages II–V which are unlike those found in any other arthropods. Opisthosomal appendages are similar in number and morphology to the book–gills of Limulus. Cladistic analysis places Offacolus basally within the Chelicerata, as a sister taxon to the eurypterids and extant chelicerates, but more derived than the Devonian Weinbergina.

Post-Cambrian closure of the deep-water slope-basin taphonomic window

Exceptional faunas (Konservat-Lagerstatten that preserve traces of volatile nonmineralized tissues) are statistically overabundant in the Cambrian Period; almost all examples preserved in continental-slope and shelf-basin environments are of this age. The hypothesis that an increase in the amount and complexity of bioturbation was an important agent in the elimination of this deep-water slope-basin taphonomic window is supported. Post-Cambrian ichnofaunal assemblages contain a higher proportion of pascichnia and agrichnia, ethologies produced by a mobile infauna. They also illustrate the lateral partitioning of organisms into different environmental niches; both opportunistic and equilibrium infaunas occur in low-oxygen environments in which the preservation of nonbiomineralized tissues was favored. Direct consumption of carcasses by bioturbating organisms was less important than changes to sediment properties as a result of bioturbation, notably enhanced microbial degradation of reactive organic matter, increased permeability, and the disruption of geochemical gradients necessary for mineral authigenesis.

Three‐dimensional preservation of a non‐biomineralized arthropod in concretions in Silurian volcaniclastic rocks from Herefordshire, England

Three‐dimensional preservation of a non‐biomineralized arthropod occurs in carbonate concretions in a volcaniclastic deposit from the Wenlock Series of Herefordshire, England. Specimens are preserved in calcite that co‐precipitated with framboids and polyhedra of pyrite. The texture of the calcite indicates that it is a void infill. It forms a cast of the external surface of the arthropod, having precipitated after decay of even the most recalcitrant biological tissues. Incorporation of the fossils into concretions ensured their long term preservation but was not, at least in most examples, responsible for preventing potential collapse and occlusion of voids in the interval between the decay of tissues and the precipitation of calcite. The precipitation and/or accumulation of clay minerals adjacent to specimens during decay was important in this process, as were possibly the geotechnical properties of the ash itself. Limited dolomitization of the calcite around the edges of the fossils and in the matrix of the concretion occurred at a later stage.

Soft-tissue preservation in Miocene frogs from Libros, Spain: Insights into the genesis of decay microenvironments

Abstract The Late Miocene Libros biota is a lacustrine-hosted, Konservat-Lagerstätte from Libros, near Teruel in northeast Spain. Adult frogs are characterized by the preservation of their soft tissues, some in histological detail. The soft tissues of the body outline are preserved as a layered structure, which comprises a central carbonaceous bacterial biofilm enveloped by the phosphatized remains of the mid-dermal Eberth-Katschenko layer, external to which is a second, thinner, carbonaceous bacterial biofilm. Bacterial autolithification is restricted to limited phosphatization of the cell margins of bacteria adjacent to phosphatized dermis. Phosphatization occurred during the late stages of decay; phosphate was sourced primarily from the dermis itself. Other tissues and organs are also defined in authigenic minerals: nervous tissue (aragonite), the stomach (calcium phosphate), and collagen fibers of the dermal stratum compactum (calcium sulphate); bone marrow is organically preserved. The disparate modes of soft-tissue preservation within individual specimens reflects development of several highly localized, chemically distinct microenvironments within the frog carcasses during decay. These microenvironments correspond to individual organs and tissues, were established at different times during decay, and varied in their duration. The preservation of soft tissues via multiple taphonomic pathways was controlled ultimately by anatomical and physiological factors.

The remarkable fossils from the Early Cretaceous Jehol Biota of China and how they have changed our knowledge of Mesozoic life

Palaeontologists and others have been repeatedly amazed by reports of spectacularly well-preserved fossils from China, and one of the key sources has been the Jehol Biota of Liaoning, Hebei and Inner Mongolia in NE China. The Jehol Biota consists of three main horizons, the Dabeigou, Yixian and Jiufotang formations, spanning the late Hauterivian to early Aptian (131-120 Ma) of the Early Cretaceous and, collectively, these have produced thousands of essentially complete specimens of plants, insects, aquatic invertebrates, fishes, frogs, salamanders, turtles, lizards, choristoderes, pterosaurs, dinosaurs, birds and mammals. Most of the specimens show some aspect of exceptional preservation, ranging from clear impressions of the body outlines to traces of soft tissues (liver, teleost air sac, eye spots) and external body coverings (scales, feathers, hair). The claim was made that these discoveries have revolutionized our understanding of evolution through this critical part of the Cretaceous Terrestrial Revolution. Key insights have come from the numerous specimens of dinosaurs with feathers, but numerical study shows that only the finds of birds and mammals have substantially changed our views about global diversity and patterns of evolution through the Early Cretaceous.

High-fidelity organic preservation of bone marrow in ca. 10 Ma amphibians

Bone marrow in ca. 10 Ma frogs and salamanders from the Miocene of Libros, Spain, represents the first fossilized example of this extremely decay-prone tissue. The bone marrow, preserved in three dimensions as an organic residue, retains the original texture and red and yellow color of hematopoietic and fatty marrow, respectively; moldic osteoclasts and vascular structures are also present. We attribute exceptional preservation of the fossilized bone marrow to cryptic preservation: the bones of the amphibians formed protective microenvironments, and inhibited microbial infiltration. Specimens in which bone marrow is preserved vary in their completeness and articulation and in the extent to which the body outline is preserved as a thin film of organically preserved bacteria. Cryptic preservation of these labile tissues is thus to a large extent independent of, and cannot be predicted by, the taphonomic history of the remainder of the specimen.

Experimental maturation of feathers: implications for reconstructions of fossil feather colour

Fossil feathers often preserve evidence of melanosomes—micrometre-scale melanin-bearing organelles that have been used to infer original colours and patterns of the plumage of dinosaurs. Such reconstructions acknowledge that evidence from other colour-producing mechanisms is presently elusive and assume that melanosome geometry is not altered during fossilization. Here, we provide the first test of this assumption, using high pressure–high temperature autoclave experiments on modern feathers to simulate the effects of burial on feather colour. Our experiments show that melanosomes are retained despite loss of visual evidence of colour and complete degradation of other colour-producing structures (e.g. quasi-ordered arrays in barbs and the keratin cortex in barbules). Significantly, however, melanosome geometry and spatial distribution are altered by the effects of pressure and temperature. These results demonstrate that reconstructions of original plumage coloration in fossils where preserved features of melanosomes are affected by diagenesis should be treated with caution. Reconstructions of fossil feather colour require assessment of the extent of preservation of various colour-producing mechanisms, and, critically, the extent of alteration of melanosome geometry.

The original colours of fossil beetles

Structural colours, the most intense, reflective and pure colours in nature, are generated when light is scattered by complex nanostructures. Metallic structural colours are widespread among modern insects and can be preserved in their fossil counterparts, but it is unclear whether the colours have been altered during fossilization, and whether the absence of colours is always real. To resolve these issues, we investigated fossil beetles from five Cenozoic biotas. Metallic colours in these specimens are generated by an epicuticular multi-layer reflector; the fidelity of its preservation correlates with that of other key cuticular ultrastructures. Where these other ultrastructures are well preserved in non-metallic fossil specimens, we can infer that the original cuticle lacked a multi-layer reflector; its absence in the fossil is not a preservational artefact. Reconstructions of the original colours of the fossils based on the structure of the multi-layer reflector show that the preserved colours are offset systematically to longer wavelengths; this probably reflects alteration of the refractive index of the epicuticle during fossilization. These findings will allow the former presence, and original hue, of metallic structural colours to be identified in diverse fossil insects, thus providing critical evidence of the evolution of structural colour in this group.