Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria.

Pterosaurs were the first vertebrates to evolve powered flight1 and comprised one of the main evolutionary radiations in terrestrial ecosystems of the Mesozoic era (approximately 252-66 million years ago), but their origin has remained an unresolved enigma in palaeontology since the nineteenth century2-4. These flying reptiles have been hypothesized to be the close relatives of a wide variety of reptilian clades, including dinosaur relatives2-8, and there is still a major morphological gap between those forms and the oldest, unambiguous pterosaurs from the Upper Triassic series. Here, using recent discoveries of well-preserved cranial remains, microcomputed tomography scans of fragile skull bones (jaws, skull roofs and braincases) and reliably associated postcrania, we demonstrate that lagerpetids-a group of cursorial, non-volant dinosaur precursors-are the sister group of pterosaurs, sharing numerous synapomorphies across the entire skeleton. This finding substantially shortens the temporal and morphological gap between the oldest pterosaurs and their closest relatives and simultaneously strengthens the evidence that pterosaurs belong to the avian line of archosaurs. Neuroanatomical features related to the enhanced sensory abilities of pterosaurs9 are already present in lagerpetids, which indicates that these features evolved before flight. Our evidence illuminates the first steps of the assembly of the pterosaur body plan, whose conquest of aerial space represents a remarkable morphofunctional innovation in vertebrate evolution.

Empirical evidence for stability of the 405-kiloyear Jupiter-Venus eccentricity cycle over hundreds of millions of years.

The Newark-Hartford astrochronostratigraphic polarity timescale (APTS) was developed using a theoretically constant 405-kiloyear eccentricity cycle linked to gravitational interactions with Jupiter-Venus as a tuning target and provides a major timing calibration for about 30 million years of Late Triassic and earliest Jurassic time. While the 405-ky cycle is both unimodal and the most metronomic of the major orbital cycles thought to pace Earth's climate in numerical solutions, there has been little empirical confirmation of that behavior, especially back before the limits of orbital solutions at about 50 million years before present. Moreover, the APTS is anchored only at its younger end by U-Pb zircon dates at 201.6 million years before present and could even be missing a number of 405-ky cycles. To test the validity of the dangling APTS and orbital periodicities, we recovered a diagnostic magnetic polarity sequence in the volcaniclastic-bearing Chinle Formation in a scientific drill core from Petrified Forest National Park (Arizona) that provides an unambiguous correlation to the APTS. New high precision U-Pb detrital zircon dates from the core are indistinguishable from ages predicted by the APTS back to 215 million years before present. The agreement shows that the APTS is continuous and supports a stable 405-kiloyear cycle well beyond theoretical solutions. The validated Newark-Hartford APTS can be used as a robust framework to help differentiate provinciality from global temporal patterns in the ecological rise of early dinosaurs in the Late Triassic, amongst other problems.

The precise temporal calibration of dinosaur origins.

Dinosaurs have been major components of ecosystems for over 200 million years. Although different macroevolutionary scenarios exist to explain the Triassic origin and subsequent rise to dominance of dinosaurs and their closest relatives (dinosauromorphs), all lack critical support from a precise biostratigraphically independent temporal framework. The absence of robust geochronologic age control for comparing alternative scenarios makes it impossible to determine if observed faunal differences vary across time, space, or a combination of both. To better constrain the origin of dinosaurs, we produced radioisotopic ages for the Argentinian Chañares Formation, which preserves a quintessential assemblage of dinosaurian precursors (early dinosauromorphs) just before the first dinosaurs. Our new high-precision chemical abrasion thermal ionization mass spectrometry (CA-TIMS) U-Pb zircon ages reveal that the assemblage is early Carnian (early Late Triassic), 5- to 10-Ma younger than previously thought. Combined with other geochronologic data from the same basin, we constrain the rate of dinosaur origins, demonstrating their relatively rapid origin in a less than 5-Ma interval, thus halving the temporal gap between assemblages containing only dinosaur precursors and those with early dinosaurs. After their origin, dinosaurs only gradually dominated mid- to high-latitude terrestrial ecosystems millions of years later, closer to the Triassic-Jurassic boundary.

Extreme ecosystem instability suppressed tropical dinosaur dominance for 30 million years.

A major unresolved aspect of the rise of dinosaurs is why early dinosaurs and their relatives were rare and species-poor at low paleolatitudes throughout the Late Triassic Period, a pattern persisting 30 million years after their origin and 10-15 million years after they became abundant and speciose at higher latitudes. New palynological, wildfire, organic carbon isotope, and atmospheric pCO2 data from early dinosaur-bearing strata of low paleolatitudes in western North America show that large, high-frequency, tightly correlated variations in δ(13)Corg and palynomorph ecotypes occurred within a context of elevated and increasing pCO2 and pervasive wildfires. Whereas pseudosuchian archosaur-dominated communities were able to persist in these same regions under rapidly fluctuating extreme climatic conditions until the end-Triassic, large-bodied, fast-growing tachymetabolic dinosaurian herbivores requiring greater resources were unable to adapt to unstable high CO2 environmental conditions of the Late Triassic.

The Fossil Calibration Database-A New Resource for Divergence Dating.

Fossils provide the principal basis for temporal calibrations, which are critical to the accuracy of divergence dating analyses. Translating fossil data into minimum and maximum bounds for calibrations is the most important-often least appreciated-step of divergence dating. Properly justified calibrations require the synthesis of phylogenetic, paleontological, and geological evidence and can be difficult for nonspecialists to formulate. The dynamic nature of the fossil record (e.g., new discoveries, taxonomic revisions, updates of global or local stratigraphy) requires that calibration data be updated continually lest they become obsolete. Here, we announce the Fossil Calibration Database (http://fossilcalibrations.org), a new open-access resource providing vetted fossil calibrations to the scientific community. Calibrations accessioned into this database are based on individual fossil specimens and follow best practices for phylogenetic justification and geochronological constraint. The associated Fossil Calibration Series, a calibration-themed publication series at Palaeontologia Electronica, will serve as a key pipeline for peer-reviewed calibrations to enter the database.

Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira.

The early evolutionary history of Ornithodira (avian-line archosaurs) has hitherto been documented by incomplete (Lagerpeton) or unusually specialized forms (pterosaurs and Silesaurus). Recently, a variety of Silesaurus-like taxa have been reported from the Triassic period of both Gondwana and Laurasia, but their relationships to each other and to dinosaurs remain a subject of debate. Here we report on a new avian-line archosaur from the early Middle Triassic (Anisian) of Tanzania. Phylogenetic analysis places Asilisaurus kongwe gen. et sp. nov. as an avian-line archosaur and a member of the Silesauridae, which is here considered the sister taxon to Dinosauria. Silesaurids were diverse and had a wide distribution by the Late Triassic, with a novel ornithodiran bauplan including leaf-shaped teeth, a beak-like lower jaw, long, gracile limbs, and a quadrupedal stance. Our analysis suggests that the dentition and diet of silesaurids, ornithischians and sauropodomorphs evolved independently from a plesiomorphic carnivorous form. As the oldest avian-line archosaur, Asilisaurus demonstrates the antiquity of both Ornithodira and the dinosaurian lineage. The initial diversification of Archosauria, previously documented by crocodilian-line archosaurs in the Anisian, can now be shown to include a contemporaneous avian-line radiation. The unparalleled taxonomic diversity of the Manda archosaur assemblage indicates that archosaur diversification was well underway by the Middle Triassic or earlier.

A palaeoequatorial ornithischian and new constraints on early dinosaur diversification.

Current characterizations of early dinosaur evolution are incomplete: existing palaeobiological and phylogenetic scenarios are based on a fossil record dominated by saurischians and the implications of the early ornithischian record are often overlooked. Moreover, the timings of deep phylogenetic divergences within Dinosauria are poorly constrained owing to the absence of a rigorous chronostratigraphical framework for key Late Triassic-Early Jurassic localities. A new dinosaur from the earliest Jurassic of the Venezuelan Andes is the first basal ornithischian recovered from terrestrial deposits directly associated with a precise radioisotopic date and the first-named dinosaur from northern South America. It expands the early palaeogeographical range of Ornithischia to palaeoequatorial regions, an area sometimes thought to be devoid of early dinosaur taxa, and offers insights into early dinosaur growth rates, the evolution of sociality and the rapid tempo of the global dinosaur radiation following the end-Triassic mass extinction, helping to underscore the importance of the ornithischian record in broad-scale discussions of early dinosaur history.

The femora of Drepanosauromorpha (Reptilia: Diapsida): Implications for the functional evolution of the thigh of Sauropsida.

The femora of diapsids have undergone morphological changes related to shifts in postural and locomotor modes, such as the transition from plesiomorphic amniote and diapsid taxa to the apomorphic conditions related to a more erect posture within Archosauriformes. One remarkable clade of Triassic diapsids is the chameleon-like Drepanosauromorpha. This group is known from numerous articulated but heavily compressed skeletons that have the potential to further inform early reptile femoral evolution. For the first time, we describe the three-dimensional osteology of the femora of Drepanosauromorpha, based on undistorted fossils from the Upper Triassic Chinle Formation and Dockum Group of North America. We identify apomorphies and a combination of character states that link these femora to those in crushed specimens of drepanosauromorphs and compare our sample with a range of amniote taxa. Several characteristics of drepanosauromorph femora, including a hemispherical proximal articular surface, prominent asymmetry in the proximodistal length of the tibial condyles, and a deep intercondylar sulcus, are plesiomorphies shared with early diapsids. The femora contrast with those of most diapsids in lacking a crest-like, distally tapering internal trochanter. They bear a ventrolaterally positioned tuberosity on the femoral shaft, resembling the fourth trochanter in Archosauriformes. The reduction of an internal trochanter parallels independent reductions in therapsids and archosauriforms. The presence of a ventrolaterally positioned trochanter is also similar to that of chameleonid squamates. Collectively, these features demonstrate a unique femoral morphology for drepanosauromorphs, and suggest an increased capacity for femoral adduction and protraction relative to most other Permo-Triassic diapsids.

Delayed recovery of non-marine tetrapods after the end-Permian mass extinction tracks global carbon cycle.

During the end-Permian mass extinction, marine ecosystems suffered a major drop in diversity, which was maintained throughout the Early Triassic until delayed recovery during the Middle Triassic. This depressed diversity in the Early Triassic correlates with multiple major perturbations to the global carbon cycle, interpreted as either intrinsic ecosystem or external palaeoenvironmental effects. In contrast, the terrestrial record of extinction and recovery is less clear; the effects and magnitude of the end-Permian extinction on non-marine vertebrates are particularly controversial. We use specimen-level data from southern Africa and Russia to investigate the palaeodiversity dynamics of non-marine tetrapods across the Permo-Triassic boundary by analysing sample-standardized generic richness, evenness and relative abundance. In addition, we investigate the potential effects of sampling, geological and taxonomic biases on these data. Our analyses demonstrate that non-marine tetrapods were severely affected by the end-Permian mass extinction, and that these assemblages did not begin to recover until the Middle Triassic. These data are congruent with those from land plants and marine invertebrates. Furthermore, they are consistent with the idea that unstable low-diversity post-extinction ecosystems were subject to boom-bust cycles, reflected in multiple Early Triassic perturbations of the carbon cycle.

Reevaluation of the cranial osteology and phylogenetic position of the early crocodyliform Eopneumatosuchus colberti, with an emphasis on its endocranial anatomy.

Eopneumatosuchus colberti Crompton and Smith, 1980, known from a single partial skull, is an enigmatic crocodylomorph from the Lower Jurassic Kayenta Formation. In spite of its unique morphology, an exceptionally pneumatic braincase, and presence during a critical time period of crocodylomorph evolution, relatively little is known about this taxon. Here, we redescribe the external cranial morphology of E. colberti, present novel information on its endocranial anatomy, evaluate its phylogenetic position among early crocodylomorphs, and seek to better characterize its ecology. Our examination clarifies key aspects of cranial suture paths and braincase anatomy. Comparisons with related taxa (e.g., Protosuchus haughtoni) demonstrate that extreme pneumaticity of the braincase may be more widespread in protosuchids than previously appreciated. Computed tomography scans reveal an endocranial morphology that resembles that of other early crocodylomorphs, in particular the noncrocodyliform crocodylomorph Almadasuchus figarii. There are, however, key differences in olfactory bulb and cerebral hemisphere morphology, which demonstrate the endocranium of crocodylomorphs is not as conserved as previously hypothesized. Our phylogenetic analysis recovers E. colberti as a close relative of Protosuchus richardsoni and Edentosuchus tienshanensis, contrasting with previous hypotheses of a sister group relationship with Thalattosuchia. Previous work suggested the inner ear has some similarities to semi-aquatic crocodyliforms, but the phylogenetic placement of E. colberti among protosuchids with a terrestrial postcranial skeletal morphology complicates paleoecological interpretation.

Osteology and relationships of Revueltosaurus callenderi (Archosauria: Suchia) from the Upper Triassic (Norian) Chinle Formation of Petrified Forest National Park, Arizona, United States.

Once known solely from dental material and thought to represent an early ornithischian dinosaur, the early-diverging pseudosuchian Revueltosaurus callenderi is described from a minimum of 12 skeletons from a monodominant bonebed in the upper part of the Chinle Formation of Arizona. This material includes nearly the entire skeleton and possesses a combination of plesiomorphic and derived character states that help clarify ingroup relationships within Pseudosuchia. A phylogenetic analysis recovers R. callenderi in a clade with Aetosauria and Acaenasuchus geoffreyi that is named Aetosauriformes. Key autapomorphies of R. callenderi include a skull that is longer than the femur, a complete carapace of dermal armor including paramedian and lateral rows, as well as ventral osteoderms, and a tail end sheathed in bone. Histology of the femur and associated osteoderms demonstrate that R. callenderi was slow growing and that the individuals from the bonebed were not young juveniles but had not ceased growing. A review of other material assigned to Revueltosaurus concludes that the genus cannot be adequately diagnosed based on the type materials of the three assigned species and that only R. callenderi can be confidently referred to Revueltosaurus.

Grouping behavior in a Triassic marine apex predator.

Marine tetrapods occupy important roles in modern marine ecosystems and often gather in large aggregations driven by patchy prey distribution,1,2 social or reproductive behaviors,3,4 or oceanographic factors.5 Here, we show that similar grouping behaviors evolved in an early marine tetrapod lineage, documented by dozens of specimens of the giant ichthyosaur Shonisaurus in the Luning Formation in West Union Canyon, Nevada, USA.6,7 A concentration of at least seven skeletons closely preserved on a single bedding plane received the bulk of previous attention. However, many more specimens are preserved across ∼106 square meters and ∼200 stratigraphic meters of outcrop representing an estimated >105-6 years. Unlike other marine-tetrapod-rich deposits, this assemblage is essentially monotaxic; other vertebrate fossils are exceptionally scarce. Large individuals are disproportionately abundant, with the exception of multiple neonatal or embryonic specimens, indicating an unusual demographic composition apparently lacking intermediate-sized juveniles or subadults. Combined with geological evidence, our data suggest that dense aggregations of Shonisaurus inhabited this moderately deep, low-diversity, tropical marine environment for millennia during the latest Carnian Stage of the Late Triassic Period (237-227 Ma). Thus, philopatric grouping behavior in marine tetrapods, potentially linked to reproductive activity, has an antiquity of at least 230 million years.

Histological analysis of ankylothecodonty in Silesauridae (Archosauria: Dinosauriformes) and its implications for the evolution of dinosaur tooth attachment.

Dinosaurs possess a form of tooth attachment wherein an unmineralized periodontal ligament suspends each tooth within a socket, similar to the condition in mammals and crocodylians. However, little information is known about tooth attachment and implantation in their close relatives, the silesaurids. We conducted a histological survey of several silesaurid taxa to determine the nature of tooth attachment in this phylogenetically and paleoecologically important group of archosaurs. Our histological data demonstrate that these early dinosauriforms do not exhibit the crocodilian/dinosaur condition of a permanent gomphosis, nor the rapid ankylosis that is plesiomorphic for amniotes. Instead, all sampled silesaurids exhibit delayed ankylosis, a condition in which teeth pass through a prolonged stage where the teeth are suspended in sockets by a periodontal ligament, followed by eventual mineralization and fusion of the tooth to the jaws. This suggests that tooth attachment in crocodylians and dinosaurs represent the further retention of an early ontogenetic stage compared to silesaurids, a paedomorphic trend that is mirrored in the evolution of synapsid tooth attachment. It also suggests that the dinosaur and crocodylian gomphosis was convergently acquired via heterochrony or, less likely, that the silesaurid condition represents a reversal to a plesiomorphic state. Moreover, if Silesauridae is nested within Ornithischia, a permanent gomphosis could be convergent between the two main dinosaur lineages, Ornithischia and Saurischia. These results demonstrate that dental characters in early archosaur phylogenies must be chosen and defined carefully, taking into account the relative duration of the different phases of dental ontogeny.

The limits of convergence: the roles of phylogeny and dietary ecology in shaping non-avian amniote crania.

Cranial morphology is remarkably varied in living amniotes and the diversity of shapes is thought to correspond with feeding ecology, a relationship repeatedly demonstrated at smaller phylogenetic scales, but one that remains untested across amniote phylogeny. Using a combination of morphometric methods, we investigate the links between phylogenetic relationships, diet and skull shape in an expansive dataset of extant toothed amniotes: mammals, lepidosaurs and crocodylians. We find that both phylogeny and dietary ecology have statistically significant effects on cranial shape. The three major clades largely partition morphospace with limited overlap. Dietary generalists often occupy clade-specific central regions of morphospace. Some parallel changes in cranial shape occur in clades with distinct evolutionary histories but similar diets. However, members of a given clade often present distinct cranial shape solutions for a given diet, and the vast majority of species retain the unique aspects of their ancestral skull plan, underscoring the limits of morphological convergence due to ecology in amniotes. These data demonstrate that certain cranial shapes may provide functional advantages suited to particular dietary ecologies, but accounting for both phylogenetic history and ecology can provide a more nuanced approach to inferring the ecology and functional morphology of cryptic or extinct amniotes.

Osteology of the Late Triassic Bipedal Archosaur Poposaurus gracilis (Archosauria: Pseudosuchia) from Western North America.

Poposaurus gracilis is a bipedal pseudosuchian archosaur that has been poorly understood since the discovery of the holotype fragmentary partial postcranial skeleton in 1915. Poposaurus. gracilis is a member of Poposauroidea, an unusually morphologically divergent clade of pseudosuchians containing taxa that are bipedal, quadrupedal, toothed, edentulous, and some individuals with elongated thoracic neural spines (i.e., sails). In 2003, a well preserved, fully articulated, and nearly complete postcranial skeleton of P. gracilis was discovered with some fragmentary cranial elements from the Upper Triassic Chinle Formation of Grand Staircase-Escalante National Monument of southern Utah, USA. The aim of this work is to describe the osteology of this specimen in detail and compare P. gracilis to other closely related pseudosuchian archosaurs. The open neurocentral sutures throughout the majority of the vertebral column, the small size of this individual, and the presence of seven evenly spaced cyclic growth marks in the histologically sectioned femur indicate that this specimen was a skeletally immature juvenile, or subadult when it died. The pes of P. gracilis contains multiple skeletal adaptations and osteological correlates for soft tissue structures that support a hypothesis of digitigrady for this taxon. When coupled with the numerous postcranial characters associated with cursoriality, and the many anatomical traits convergent with theropod dinosaurs, this animal likely occupied a similar ecological niche with contemporaneous theropods during the Late Triassic Period. Anat Rec, 303:874-917, 2020. © 2019 American Association for Anatomy.

Repeated Evolution of Herbivorous Crocodyliforms during the Age of Dinosaurs.

Extinct crocodyliforms from the age of dinosaurs (Mesozoic Era) display an impressive range of skeletal morphologies, suggesting a diversity of ecological roles not found in living representatives [1-6]. In particular, unusual dental morphologies develop repeatedly through the evolutionary history of this group [2, 4-9]. Recent descriptions of fossil crocodyliforms and their unusual teeth provide the inferential basis for a wide range of feeding ecologies. However, tests of these hypotheses are hindered by the lack of directly comparable dental morphologies in living reptiles and mammals, thereby preventing an accurate ecosystem reconstruction. Here, we demonstrate, using a combination of the orientation patch count rotated method and discrete morphological features, that Mesozoic crocodyliforms exploited a much greater range of feeding ecologies than their extant relatives, including likely omnivores and herbivores. These results also indicate that crocodyliforms independently developed high-complexity dentitions a minimum of three times. Some taxa possess teeth that surpass the complexities of living herbivorous lizards and rival those of omnivorous and herbivorous mammals. This study indicates that herbivorous crocodyliforms were more common than previously thought and were present throughout the Mesozoic and on most continents. The occurrence of multiple origins of complex dentitions throughout Crocodyliformes indicates that herbivory was a beneficial dietary strategy and not a unique occurrence. Many of these crocodyliforms lived alongside omnivorous or herbivorous synapsids, illustrating an ecological partition that is not observed today.

Integrating gross morphology and bone histology to assess skeletal maturity in early dinosauromorphs: new insights from Dromomeron (Archosauria: Dinosauromorpha).

Understanding growth patterns is central to properly interpreting paleobiological signals in tetrapods, but assessing skeletal maturity in some extinct clades may be difficult when growth patterns are poorly constrained by a lack of ontogenetic series. To overcome this difficulty in assessing the maturity of extinct archosaurian reptiles-crocodylians, birds and their extinct relatives-many studies employ bone histology to observe indicators of the developmental stage reached by a given individual. However, the relationship between gross morphological and histological indicators of maturity has not been examined in most archosaurian groups. In this study, we examined the gross morphology of a hypothesized growth series of Dromomeron romeri femora (96.6-144.4 mm long), the first series of a non-dinosauriform dinosauromorph available for such a study. We also histologically sampled several individuals in this growth series. Previous studies reported that D. romeri lacks well-developed rugose muscle scars that appear during ontogeny in closely related dinosauromorph taxa, so integrating gross morphology and histological signal is needed to determine reliable maturity indicators for early bird-line archosaurs. We found that, although there are small, linear scars indicating muscle attachment sites across the femur, the only rugose muscle scar that appears during ontogeny is the attachment of the M. caudofemoralis longus, and only in the largest-sampled individual. This individual is also the only femur with histological indicators that asymptotic size had been reached, although smaller individuals possess some signal of decreasing growth rates (e.g., decreasing vascular density). The overall femoral bone histology of D. romeri is similar to that of other early bird-line archosaurs (e.g., woven-bone tissue, moderately to well-vascularized, longitudinal vascular canals). All these data indicate that the lack of well-developed femoral scars is autapomorphic for this species, not simply an indication of skeletal immaturity. We found no evidence of the high intraspecific variation present in early dinosaurs and other dinosauriforms, but a limited sample size of other early bird-line archosaur growth series make this tentative. The evolutionary history and phylogenetic signal of gross morphological features must be considered when assessing maturity in extinct archosaurs and their close relatives, and in some groups corroboration with bone histology or with better-known morphological characters is necessary.

Caveats on the use of fossil calibrations for molecular dating: a comment on Near et al.

We reassess a study on a fossil-calibrated molecular clock that provides a new method for evaluating the accuracy of calibration points. We address several pitfalls that molecular systematists should be aware of when calculating rates of molecular evolution based on fossil calibrations. These caveats involve the substantiation and accurate use of geologic dates, the inappropriate use of fixed calibration points, and the explicit and objective phylogenetic placement of fossil taxa. Paleontological data, like molecular data, should be treated with the utmost rigor.

Including autapomorphies is important for paleontological tip-dating with clocklike data, but not with non-clock data.

Tip-dating, where fossils are included as dated terminal taxa in Bayesian dating inference, is an increasingly popular method. Data for these studies often come from morphological character matrices originally developed for non-dated, and usually parsimony, analyses. In parsimony, only shared derived characters (synapomorphies) provide grouping information, so many character matrices have an ascertainment bias: they omit autapomorphies (unique derived character states), which are considered uninformative. There has been no study of the effect of this ascertainment bias in tip-dating, but autapomorphies can be informative in model-based inference. We expected that excluding autapomorphies would shorten the morphological branchlengths of terminal branches, and thus bias downwards the time branchlengths inferred in tip-dating. We tested for this effect using a matrix for Carboniferous-Permian eureptiles where all autapomorphies had been deliberately coded. Surprisingly, date estimates are virtually unchanged when autapomorphies are excluded, although we find large changes in morphological rate estimates and small effects on topological and dating confidence. We hypothesized that the puzzling lack of effect on dating was caused by the non-clock nature of the eureptile data. We confirm this explanation by simulating strict clock and non-clock datasets, showing that autapomorphy exclusion biases dating only for the clocklike case. A theoretical solution to ascertainment bias is computing the ascertainment bias correction (Mkparsinf), but we explore this correction in detail, and show that it is computationally impractical for typical datasets with many character states and taxa. Therefore we recommend that palaeontologists collect autapomorphies whenever possible when assembling character matrices.