Modes of Pangean lake level cyclicity driven by astronomical climate pacing modulated by continental position and pCO[Formula: see text].

Orbital cyclicity is a fundamental pacemaker of Earth's climate system. The Newark-Hartford Basin (NHB) lake sediment record of eastern North America contains compelling geologic expressions of this cyclicity, reflecting variations of climatic conditions in tropical Pangea during the Late Triassic and earliest Jurassic (~233 to 199 Ma). Climate modeling enables a deeper mechanistic understanding of Earth system modulation during this unique greenhouse and supercontinent period. We link major features of the NHB record to the combined climatic effects of orbital forcing, paleogeographic changes, and atmospheric pCO[Formula: see text] variations. An ensemble of transient, orbitally driven climate simulations is assessed for nine time slices, three atmospheric pCO[Formula: see text] values, and two paleogeographic reconstructions. Climatic transitions from tropical humid to more seasonal and ultimately semiarid are associated with tectonic drift of the NHB from [Formula: see text] to [Formula: see text]. The modeled orbital modulation of the precipitation-evaporation balance is most pronounced during the 220 to 200 Ma interval, whereas it is limited by weak seasonality and increasing aridity before and after this interval. Lower pCO[Formula: see text] at around 205 Ma contributes to drier climates and could have led to the observed damping of sediment cyclicity. Eccentricity-modulated precession dominates the orbitally driven climate response in the NHB region. High obliquity further amplifies summer precipitation through the seasonal shifts in the tropical rainfall belt. Regions with other proxy records are also assessed, providing guidance toward an integrated picture of global astronomical climate forcing in the Late Triassic and ultimately of other periods in Earth history.

Variation in the sacrum of phytosaurs: New evidence from a partial skeleton of Machaeroprosopus mccauleyi.

Phytosaurs are a group of Upper Triassic semi-aquatic archosauriform reptiles. Their variable skull morphology forms the foundation of our understanding of their relationships and paleoecology, while only a few studies have focused on demonstrating the existence of postcranial variation. The numbers of vertebrae in the sacrum are thought to vary from two, the plesiomorphic condition for archosauriforms, to three, with the addition of a sacralized dorsal (i.e., dorsosacral) vertebra. In this study, we demonstrate the presence of a sacralized first caudal (i.e., caudosacral) vertebra in a sacrum belonging to Machaeroprosopus mccauleyi. We rule out taphonomic distortion or pathology as explanations for the inclusion of this element in the sacrum, suggesting instead that it occurred through modifications of the same developmental processes that likely produced dorsosacral vertebrae in phytosaurs. Additionally, we show that a dorsosacral vertebra is common in phytosaur specimens from the Chinle Formation and Dockum Group of the southwestern United States and suggest that it may be widespread among phytosaurs. The addition of sacral vertebrae potentially aided adaptation to larger body sizes or more terrestrial lifestyles in certain taxa.

A dwarf conifer tree from the Triassic of Antarctica: the first fossil evidence of suppressed growth in a favorable climate?

BACKGROUND AND AIMS: The complexity of fossil forest ecosystems is difficult to reconstruct due to the fragmentary nature of the fossil record. However, detailed morpho-anatomical studies of well-preserved individual fossils can provide key information on tree growth and ecology, including in biomes with no modern analog such as the lush forests that developed in the polar regions during past greenhouse climatic episodes. METHODS: We describe an unusual-looking stem from Middle Triassic (ca 240 Ma) deposits of Antarctica with over 100 very narrow growth-rings and conspicuous persistent vascular traces through the wood. Sections of the specimen were prepared using the cellulose acetate peel technique to determine its systematic affinities and analyse its growth. KEY RESULTS: The new fossil shows similarities with the form genus Woodworthia and with conifer stems from the Triassic of Antarctica, and is assigned to the conifers. Vascular traces are interpreted as those of small branches retained on the trunk. Growth-ring analyses reveal one of the slowest growth rates reported in the fossil record, with an average of 0.2 mm/season. While the tree was growing within the Triassic polar circle, sedimentological data and growth-ring information from other fossil trees, including from the same locality, support the presence of favorable conditions in the region. CONCLUSIONS: The specimen is interpreted as a dwarf conifer tree that grew under a generally favorable regional climate but whose growth was suppressed due to stressful local site conditions. This is the first time that a tree with suppressed growth is identified as such in the fossil record, providing new insights on the structure of polar forests under greenhouse climates and, more generally, on the complexity of tree communities in deep time.

A new pseudosuchian from the Favret Formation of Nevada reveals that archosauriforms occupied coastal regions globally during the Middle Triassic.

Recent studies suggest that both stem- and crown-group Archosauria encompassed high ecological diversity during their initial Triassic radiation. We describe a new pseudosuchian archosaur, Benggwigwishingasuchus eremicarminis gen. et sp. nov., from the Anisian (Middle Triassic) Fossil Hill Member of the Favret Formation (Nevada, USA), a pelagic setting in the eastern Panthalassan Ocean characterized by the presence of abundant ammonoids and large-bodied ichthyosaurs. Coupled with archosauriforms from the eastern and western Tethys Ocean, Benggwigwishingasuchus reveals that pseudosuchians were also components of Panthalassan ocean coastal settings, establishing that the group occupied these habitats globally during the Middle Triassic. However, Benggwigwishingasuchus, Qianosuchus, and Ticinosuchus (two other pseudosuchians known from marine sediments) are not recovered in a monophyletic group, demonstrating that a nearshore marine lifestyle occurred widely across Archosauriformes during this time. Benggwigwishingasuchus is recovered as part of an expanded Poposauroidea, including several taxa (e.g. Mandasuchus, Mambawakalae) from the Middle Triassic Manda Beds of Tanzania among its basally branching members. This implies a greater undiscovered diversity of poposauroids during the Early Triassic, and supports that the group, and pseudosuchians more broadly, diversified rapidly following the End-Permian mass extinction.

Northward dispersal of dinosaurs from Gondwana to Greenland at the mid-Norian (215-212 Ma, Late Triassic) dip in atmospheric pCO2.

The earliest dinosaurs (theropods and sauropodomorphs) are found in fossiliferous early Late Triassic strata dated to about 230 million years ago (Ma), mainly in northwestern Argentina and southern Brazil in the Southern Hemisphere temperate belt of what was Gondwana in Pangea. Sauropodomorphs, which are not known for the entire Triassic in then tropical North America, eventually appear 15 million years later in the Northern Hemisphere temperate belt of Laurasia. The Pangea supercontinent was traversable in principle by terrestrial vertebrates, so the main barrier to be surmounted for dispersal between hemispheres was likely to be climatic; in particular, the intense aridity of tropical desert belts and unstable climate in the equatorial humid belt accompanying high atmospheric pCO2 that characterized the Late Triassic. We revisited the chronostratigraphy of the dinosaur-bearing Fleming Fjord Group of central East Greenland and, with additional data, produced a correlation of a detailed magnetostratigraphy from more than 325 m of composite section from two field areas to the age-calibrated astrochronostratigraphic polarity time scale. This age model places the earliest occurrence of sauropodomorphs (Plateosaurus) in their northernmost range to ∼214 Ma. The timing is within the 215 to 212 Ma (mid-Norian) window of a major, robust dip in atmospheric pCO2 of uncertain origin but which may have resulted in sufficiently lowered climate barriers that facilitated the initial major dispersal of the herbivorous sauropodomorphs to the temperate belt of the Northern Hemisphere. Indications are that carnivorous theropods may have had dispersals that were less subject to the same climate constraints.

Hematite reconstruction of Late Triassic hydroclimate over the Colorado Plateau.

Hematite is the most abundant surficial iron oxide on Earth resulting from near-surface processes that make it important for addressing numerous geologic problems. While red beds have proved to be excellent paleomagnetic recorders, the early diagenetic origin of hematite in these units is often questioned. Here, we validate pigmentary hematite ("pigmentite") as a proxy indicator for the Late Triassic environment and its penecontemporaneous origin by analyzing spectrophotometric measurements of a 14.5-My-long red bed sequence in scientific drill core CPCP-PFNP13-1A of the Chinle Formation, Arizona. Pigmentite concentrations in the red beds track the evolving pattern of the Late Triassic monsoon and indicate a long-term rise in aridity beginning at ∼215 Ma followed by increased oscillatory climate change at ∼213 Ma. These monsoonal changes are attributed to the northward drift of the Colorado Plateau as part of Laurentia into the arid subtropics during a time of fluctuating CO2 Our results refine the record of the Late Triassic monsoon and indicate significant changes in rainfall proximal to the Adamanian-Revueltian biotic transition that thus may have contributed to apparent faunal and floral events at 216 to 213 Ma.

Evidence from South Africa for a protracted end-Permian extinction on land.

Earth's largest biotic crisis occurred during the Permo-Triassic Transition (PTT). On land, this event witnessed a turnover from synapsid- to archosauromorph-dominated assemblages and a restructuring of terrestrial ecosystems. However, understanding extinction patterns has been limited by a lack of high-precision fossil occurrence data to resolve events on submillion-year timescales. We analyzed a unique database of 588 fossil tetrapod specimens from South Africa's Karoo Basin, spanning ∼4 My, and 13 stratigraphic bin intervals averaging 300,000 y each. Using sample-standardized methods, we characterized faunal assemblage dynamics during the PTT. High regional extinction rates occurred through a protracted interval of ∼1 Ma, initially co-occurring with low origination rates. This resulted in declining diversity up to the acme of extinction near the Daptocephalus-Lystrosaurus declivis Assemblage Zone boundary. Regional origination rates increased abruptly above this boundary, co-occurring with high extinction rates to drive rapid turnover and an assemblage of short-lived species symptomatic of ecosystem instability. The "disaster taxon" Lystrosaurus shows a long-term trend of increasing abundance initiated in the latest Permian. Lystrosaurus comprised 54% of all specimens by the onset of mass extinction and 70% in the extinction aftermath. This early Lystrosaurus abundance suggests its expansion was facilitated by environmental changes rather than by ecological opportunity following the extinctions of other species as commonly assumed for disaster taxa. Our findings conservatively place the Karoo extinction interval closer in time, but not coeval with, the more rapid marine event and reveal key differences between the PTT extinctions on land and in the oceans.

Volcanically driven lacustrine ecosystem changes during the Carnian Pluvial Episode (Late Triassic).

The Late Triassic Carnian Pluvial Episode (CPE) saw a dramatic increase in global humidity and temperature that has been linked to the large-scale volcanism of the Wrangellia large igneous province. The climatic changes coincide with a major biological turnover on land that included the ascent of the dinosaurs and the origin of modern conifers. However, linking the disparate cause and effects of the CPE has yet to be achieved because of the lack of a detailed terrestrial record of these events. Here, we present a multidisciplinary record of volcanism and environmental change from an expanded Carnian lake succession of the Jiyuan Basin, North China. New U-Pb zircon dating, high-resolution chemostratigraphy, and palynological and sedimentological data reveal that terrestrial conditions in the region were in remarkable lockstep with the large-scale volcanism. Using the sedimentary mercury record as a proxy for eruptions reveals four discrete episodes during the CPE interval (ca. 234.0 to 232.4 Ma). Each eruptive phase correlated with large, negative C isotope excursions and major climatic changes to more humid conditions (marked by increased importance of hygrophytic plants), lake expansion, and eutrophication. Our results show that large igneous province eruptions can occur in multiple, discrete pulses, rather than showing a simple acme-and-decline history, and demonstrate their powerful ability to alter the global C cycle, cause climate change, and drive macroevolution, at least in the Triassic.

Arthropod and Pathogen Damage on Fossil and Modern Plants: Exploring the Origins and Evolution of Herbivory on Land.

The use of the functional feeding group-damage type system for analyzing arthropod and pathogen interactions with plants has transformed our understanding of herbivory in fossil plant assemblages by providing data, analyses, and interpretation of the local, regional, and global patterns of a 420-Myr history. The early fossil record can be used to answer major questions about the oldest evidence for herbivory, the early emergence of herbivore associations on land plants, and later expansion on seed plants. The subsequent effects of the Permian-Triassic ecological crisis on herbivore diversity, the resulting formation of biologically diverse herbivore communities on gymnosperms, and major shifts in herbivory ensuing from initial angiosperm diversification are additional issues that need to be addressed. Studies ofherbivory resulting from more recent transient spikes and longer-term climate trends provide important data that are applied to current global change and include herbivore community responses to latitude, altitude, and habitat. Ongoing paleoecological themes remaining to be addressed include the antiquity of modern interactions, differential herbivory between ferns and angiosperms, and origins of modern tropical forests. The expansion of databases that include a multitude of specimens; improvements in sampling strategies; development of new analytical methods; and, importantly, the ability to address conceptually stimulating ecological and evolutionary questions have provided new impetus in this rapidly advancing field.

Contrasting terrestrial and marine ecospace dynamics after the end-Triassic mass extinction event.

Mass extinctions have fundamentally altered the structure of the biosphere throughout Earth's history. The ecological severity of mass extinctions is well studied in marine ecosystems by categorizing marine taxa into functional groups based on 'ecospace' approaches, but the ecological response of terrestrial ecosystems to mass extinctions is less well understood due to the lack of a comparable methodology. Here, we present a new terrestrial ecospace framework that categorizes fauna into functional groups as defined by tiering, motility and feeding traits. We applied the new terrestrial and traditional marine ecospace analyses to data from the Paleobiology Database across the end-Triassic mass extinction-a time of catastrophic global warming-to compare changes between the marine and terrestrial biospheres. We found that terrestrial functional groups experienced higher extinction severity, that taxonomic and functional richness are more tightly coupled in the terrestrial, and that the terrestrial realm continued to experience high ecological dissimilarity in the wake of the extinction. Although signals of extinction severity and ecological turnover are sensitive to the quality of the terrestrial fossil record, our findings suggest greater ecological pressure from the end-Triassic mass extinction on terrestrial ecosystems than marine ecosystems, contributing to more prolonged terrestrial ecological flux.

A tiny ornithodiran archosaur from the Triassic of Madagascar and the role of miniaturization in dinosaur and pterosaur ancestry.

Early members of the dinosaur-pterosaur clade Ornithodira are very rare in the fossil record, obscuring our understanding of the origins of this important group. Here, we describe an early ornithodiran (Kongonaphon kely gen. et sp. nov.) from the Mid-to-Upper Triassic of Madagascar that represents one of the smallest nonavian ornithodirans. Although dinosaurs and gigantism are practically synonymous, an analysis of body size evolution in dinosaurs and other archosaurs in the context of this taxon and related forms demonstrates that the earliest-diverging members of the group may have been smaller than previously thought, and that a profound miniaturization event occurred near the base of the avian stem lineage. In phylogenetic analysis, Kongonaphon is recovered as a member of the Triassic ornithodiran clade Lagerpetidae, expanding the range of this group into Africa and providing data on the craniodental morphology of lagerpetids. The conical teeth of Kongonaphon exhibit pitted microwear consistent with a diet of hard-shelled insects, indicating a shift in trophic ecology to insectivory associated with diminutive body size. Small ancestral body size suggests that the extreme rarity of early ornithodirans in the fossil record owes more to taphonomic artifact than true reflection of the group's evolutionary history.

The earliest-known mammaliaform fossil from Greenland sheds light on origin of mammals.

Synapsids are unique in having developed multirooted teeth and complex occlusions. These innovations evolved in at least two lineages of mammaliamorphs (Tritylodontidae and Mammaliaformes). Triassic fossils demonstrate that close to the origins of mammals, mammaliaform precursors were "experimenting" with tooth structure and function, resulting in novel patterns of occlusion. One of the most surprising examples of such adaptations is present in the haramiyidan clade, which differed from contemporary mammaliaforms in having two rows of cusps on molariform crowns adapted to omnivorous/herbivorous feeding. However, the origin of the multicusped tooth pattern present in haramiyidans has remained enigmatic. Here we describe the earliest-known mandibular fossil of a mammaliaform with double molariform roots and a crown with two rows of cusps from the Late Triassic of Greenland. The crown morphology is intermediate between that of morganucodontans and haramiyidans and suggests the derivation of the multicusped molariforms of haramiyidans from the triconodont molar pattern seen in morganucodontids. Although it is remarkably well documented in the fossil record, the significance of tooth root division in mammaliaforms remains enigmatic. The results of our biomechanical analyses (finite element analysis [FEA]) indicate that teeth with two roots can better withstand stronger mechanical stresses like those resulting from tooth occlusion, than teeth with a single root.

Thermogenic carbon release from the Central Atlantic magmatic province caused major end-Triassic carbon cycle perturbations.

The Central Atlantic magmatic province (CAMP), the end-Triassic mass extinction (ETE), and associated major carbon cycle perturbations occurred synchronously around the Triassic-Jurassic (T-J) boundary (201 Ma). Negative carbon isotope excursions (CIEs) recorded in marine and terrestrial sediments attest to the input of isotopically light carbon, although the carbon sources remain debated. Here, we explore the effects of mantle-derived and thermogenic carbon released from the emplacement of CAMP using the long-term ocean-atmosphere-sediment carbon cycle reservoir (LOSCAR) model. We have tested a detailed emission scenario grounded by numerous complementary boundary conditions, aiming to model the full extent of the carbon cycle perturbations around the T-J boundary. These include three negative CIEs (i.e., Marshi/Precursor, Spelae/Initial, Tilmanni/Main) with sharp positive CIEs in between. We show that a total of ∼24,000 Gt C (including ∼12,000 Gt thermogenic C) replicates the proxy data. These results indicate that thermogenic carbon generated from the contact aureoles around CAMP sills represents a credible source for the negative CIEs. An extremely isotopically depleted carbon source, such as marine methane clathrates, is therefore not required. Furthermore, we also find that significant organic carbon burial, in addition to silicate weathering, is necessary to account for the positive δ13C intervals following the negative CIEs.

Molecular and isotopic evidence reveals the end-Triassic carbon isotope excursion is not from massive exogenous light carbon.

The negative organic carbon isotope excursion (CIE) associated with the end-Triassic mass extinction (ETE) is conventionally interpreted as the result of a massive flux of isotopically light carbon from exogenous sources into the atmosphere (e.g., thermogenic methane and/or methane clathrate dissociation linked to the Central Atlantic Magmatic Province [CAMP]). Instead, we demonstrate that at its type locality in the Bristol Channel Basin (UK), the CIE was caused by a marine to nonmarine transition resulting from an abrupt relative sea level drop. Our biomarker and compound-specific carbon isotopic data show that the emergence of microbial mats, influenced by an influx of fresh to brackish water, provided isotopically light carbon to both organic and inorganic carbon pools in centimeter-scale water depths, leading to the negative CIE. Thus, the iconic CIE and the disappearance of marine biota at the type locality are the result of local environmental change and do not mark either the global extinction event or input of exogenous light carbon into the atmosphere. Instead, the main extinction phase occurs slightly later in marine strata, where it is coeval with terrestrial extinctions and ocean acidification driven by CAMP-induced increases in Pco2; these effects should not be conflated with the CIE. An abrupt sea-level fall observed in the Central European basins reflects the tectonic consequences of the initial CAMP emplacement, with broad implications for all extinction events related to large igneous provinces.

CO2 -induced biochemical changes in leaf volatiles decreased fire-intensity in the run-up to the Triassic-Jurassic boundary.

The Triassic-Jurassic boundary marks the third largest mass extinction event in the Phanerozoic, characterized by a rise in CO2 -concentrations from c. 600 ppm to c. 2100-2400 ppm, coupled with a c. 3.0-4.0°C temperature rise. This is hypothesized to have induced major floral turnover, altering vegetation structure, composition and leaf morphology, which in turn are hypothesized to have driven changes in wildfire. However, the effects of elevated CO2 on fuel properties, such as chemical composition of leaves, are also important in influencing fire behaviour, but yet have not been considered. We test this by selecting three Triassic analogue species grown experimentally in different atmospheric compositions, and analyse variations in leaf chemistry, and leaf level flammability. These data were used to inform a fire behaviour model. We find that all three species tested showed a reduction in their volatile component, leading to lower flammability. Accounting for these variations in a model, our results suggest that leaf intrinsic flammability has a measurable impact on modelled fire behaviour. If scaled up to ecosystem level, periods of elevated CO2 may therefore be capable of inducing both biochemical and morphological changes in fuel properties, and thus may be capable of influencing fire behaviour.

Diphyodont tooth replacement of Brasilodon-A Late Triassic eucynodont that challenges the time of origin of mammals.

Two sets of teeth (diphyodonty) characterise extant mammals but not reptiles, as they generate many replacement sets (polyphyodonty). The transition in long-extinct species from many sets to only two has to date only been reported in Jurassic eucynodonts. Specimens of the Late Triassic brasilodontid eucynodont Brasilodon have provided anatomical and histological data from three lower jaws of different growth stages. These reveal ordered and timed replacement of deciduous by adult teeth. Therefore, this diphyodont dentition, as contemporary of the oldest known dinosaurs, shows that Brasilodon falls within a range of wide variations of typically mammalian, diphyodont dental patterns. Importantly, these three lower jaws represent distinct ontogenetic stages that reveal classic features for timed control of replacement, by the generation of only one replacement set of teeth. This data shows that the primary premolars reveal a temporal replacement pattern, importantly from directly below each tooth, by controlled regulation of tooth resorption and regeneration. The complexity of the adult prismatic enamel structure with a conspicuous intra-structural Schmelzmuster array suggests that, as in the case of extant mammals, this extinct species would have probably sustained higher metabolic rates than reptiles. Furthermore, in modern mammals, diphyodonty and prismatic enamel are inextricably linked, anatomically and physiologically, to a set of other traits including placentation, endothermy, fur, lactation and even parental care. Our analysis of the osteodental anatomy of Brasilodon pushes back the origin of diphyodonty and consequently, its related biological traits to the Norian (225.42 ± 0.37 myr), and around 25 myr after the End-Permian mass extinction event.

Femoral specializations to locomotor habits in early archosauriforms.

The evolutionary history of archosaurs and their closest relatives is characterized by a wide diversity of locomotor modes, which has even been suggested as a pivotal aspect underlying the evolutionary success of dinosaurs vs. pseudosuchians across the Triassic-Jurassic transition. This locomotor diversity (e.g., more sprawling/erect; crouched/upright; quadrupedal/bipedal) led to several morphofunctional specializations of archosauriform limb bones that have been studied qualitatively as well as quantitatively through various linear morphometric studies. However, differences in locomotor habits have never been studied across the Triassic-Jurassic transition using 3D geometric morphometrics, which can relate how morphological features vary according to biological factors such as locomotor habit and body mass. Herein, we investigate morphological variation across a dataset of 72 femora from 36 different species of archosauriforms. First, we identify femoral head rotation, distal slope of the fourth trochanter, femoral curvature, and the angle between the lateral condyle and crista tibiofibularis as the main features varying between bipedal and quadrupedal taxa, all of these traits having a stronger locomotor signal than the lesser trochanter's proximal extent. We show a significant association between locomotor mode and phylogeny, but with the locomotor signal being stronger than the phylogenetic signal. This enables us to predict locomotor modes of some of the more ambiguous early archosauriforms without relying on the relationships between hindlimb and forelimb linear bone dimensions as in prior studies. Second, we highlight that the most important morphological variation is linked to the increase of body size, which impacts the width of the epiphyses and the roundness and proximodistal position of the fourth trochanter. Furthermore, we show that bipedal and quadrupedal archosauriforms have different allometric trajectories along the morphological variation in relation to body size. Finally, we demonstrate a covariation between locomotor mode and body size, with variations in femoral bowing (anteroposterior curvature) being more distinct among robust femora than gracile ones. We also identify a decoupling in fourth trochanter variation between locomotor mode (symmetrical to semi-pendant) and body size (sharp to rounded). Our results indicate a similar level of morphological disparity linked to a clear convergence in femoral robusticity between the two clades of archosauriforms (Pseudosuchia and Avemetatarsalia), emphasizing the importance of accounting for body size when studying their evolutionary history, as well as when studying the functional morphology of appendicular features. Determining how early archosauriform skeletal features were impacted by locomotor habits and body size also enables us to discuss the potential homoplasy of some phylogenetic characters used previously in cladistic analyses as well as when bipedalism evolved in the avemetatarsalian lineage. This study illuminates how the evolution of femoral morphology in early archosauriforms was functionally constrained by locomotor habit and body size, which should aid ongoing discussions about the early evolution of dinosaurs and the nature of their evolutionary "success" over pseudosuchians.

Mapping Solar System chaos with the Geological Orrery.

The Geological Orrery is a network of geological records of orbitally paced climate designed to address the inherent limitations of solutions for planetary orbits beyond 60 million years ago due to the chaotic nature of Solar System motion. We use results from two scientific coring experiments in Early Mesozoic continental strata: the Newark Basin Coring Project and the Colorado Plateau Coring Project. We precisely and accurately resolve the secular fundamental frequencies of precession of perihelion of the inner planets and Jupiter for the Late Triassic and Early Jurassic epochs (223-199 million years ago) using the lacustrine record of orbital pacing tuned only to one frequency (1/405,000 years) as a geological interferometer. Excepting Jupiter's, these frequencies differ significantly from present values as determined using three independent techniques yielding practically the same results. Estimates for the precession of perihelion of the inner planets are robust, reflecting a zircon U-Pb-based age model and internal checks based on the overdetermined origins of the geologically measured frequencies. Furthermore, although not indicative of a correct solution, one numerical solution closely matches the Geological Orrery, with a very low probability of being due to chance. To determine the secular fundamental frequencies of the precession of the nodes of the planets and the important secular resonances with the precession of perihelion, a contemporaneous high-latitude geological archive recording obliquity pacing of climate is needed. These results form a proof of concept of the Geological Orrery and lay out an empirical framework to map the chaotic evolution of the Solar System.

The evolution of the synapsid tusk: insights from dicynodont therapsid tusk histology.

The mammalian tusk is a unique and extreme morphotype among modern vertebrate dentitions. Tusks-defined here as ever-growing incisors or canines composed of dentine-evolved independently multiple times within mammals yet have not evolved in other extant vertebrates. This suggests that there is a feature specific to mammals that facilitates the evolution of this specialized dentition. To investigate what may underpin the evolution of tusks, we histologically sampled the tusks of dicynodont therapsids: the earliest iteration of tusk evolution and the only non-mammalian synapsid clade to have acquired such a dentition. We studied the tissue composition, attachment tissues, development and replacement in 10 dicynodont taxa and show multiple developmental pathways for the adult dentitions of dicynodont tusks and tusk-like caniniforms. In a phylogenetic context, these developmental pathways reveal an evolutionary scenario for the acquisition of an ever-growing tusk-an event that occurred convergently, but only in derived members of our sample. We propose that the evolution of an ever-growing dentition, such as a tusk, is predicated on the evolution of significantly reduced tooth replacement and a permanent soft-tissue attachment. Both of these features are fixed in the dentitions of crown-group mammals, which helps to explain why tusks are restricted to this clade among extant vertebrates.

A reassessment of the enigmatic diapsid Paliguana whitei and the early history of Lepidosauromorpha.

Lepidosaurs include lizards, snakes, amphisbaenians and the tuatara, comprising a highly speciose evolutionary radiation with widely varying anatomical traits. Their stem-lineage originated by the late middle Permian 259 million years ago, but its early fossil record is poorly documented, obscuring the origins of key anatomical and functional traits of the group. Paliguana whitei, from the Early Triassic of South Africa, is an enigmatic fossil species with the potential to provide information on this. However, its anatomy and phylogenetic affinities remain highly uncertain, and have been debated since its discovery more than 100 years ago. We present microtomographic three-dimensional imaging of the cranial anatomy of P. whitei that clarifies these uncertainties, providing strong evidence for lepidosauromorph affinities based on the structure of the temporal region and the implantation of marginal dentition. Phylogenetic analysis including these new data recovers Paliguana as the earliest known stem-lepidosaur, within a long-lived group of early diverging lepidosauromorphs that persisted to at least the Middle Jurassic. Our results provide insights into cranial evolution on the lepidosaur stem-lineage, confirming that characteristics of pleurodont dental implantation evolved early on the lepidosaur stem-lineage. By contrast, key functional traits related to hearing (quadrate conch) and feeding (streptostyly) evolved later in the lepidosaur crown-group.