Muscle-driven predictive physics simulations of quadrupedal locomotion in the horse.

Musculoskeletal simulations can provide insights into the underlying mechanisms that govern animal locomotion. In this study, we describe the development of a new musculoskeletal model of the horse, and to our knowledge present the first fully muscle-driven, predictive simulations of equine locomotion. Our goal was to simulate a model that captures only the gross musculoskeletal structure of a horse, without specialized morphological features. We mostly present simulations acquired using feedforward control, without state feedback ("top-down control"). Without using kinematics or motion capture data as an input, we have simulated a variety of gaits that are commonly used by horses (walk, pace, trot, tölt, and collected gallop). We also found a selection of gaits that are not normally seen in horses (half bound, extended gallop, ambling). Due to the clinical relevance of the trot, we performed a tracking simulation that included empirical joint angle deviations in the cost function. To further demonstrate the flexibility of our model, we also present a simulation acquired using spinal feedback control, where muscle control signals are wholly determined by gait kinematics. Despite simplifications to the musculature, simulated footfalls and ground reaction forces followed empirical patterns. In the tracking simulation, kinematics improved with respect to the fully predictive simulations, and muscle activations showed a reasonable correspondence to electromyographic signals, although we did not predict any anticipatory firing of muscles. When sequentially increasing the target speed, our simulations spontaneously predicted walk-to-run transitions at the empirically determined speed. However, predicted stride lengths were too short over nearly the entire speed range unless explicitly prescribed in the controller, and we also did not recover spontaneous transitions to asymmetric gaits such as galloping. Taken together, our model performed adequately when simulating individual gaits, but our simulation workflow was not able to capture all aspects of gait selection. We point out certain aspects of our workflow that may have caused this, including anatomical simplifications and the use of massless Hill-type actuators. Our model is an extensible, generalized horse model, with considerable scope for adding anatomical complexity. This project is intended as a starting point for continual development of the model and code, which we make available in extensible open-source formats.

Three-dimensional dental microwear in type-Maastrichtian mosasaur teeth (Reptilia, Squamata).

Mosasaurs (Squamata, Mosasauridae) were large aquatic reptiles from the Late Cretaceous that filled a range of ecological niches within marine ecosystems. The type-Maastrichtian strata (68-66 Ma) of the Netherlands and Belgium preserve remains of five species that seemed to have performed different ecological roles (carnivores, piscivores, durophages). However, many interpretations of mosasaur diet and niche partitioning are based on qualitative types of evidence that are difficult to test explicitly. Here, we apply three-dimensional dental microwear texture analysis (DMTA) to provide quantitative dietary constraints for type-Maastrichtian mosasaurs, and to assess levels of niche partitioning between taxa. DMTA indicates that these mosasaurs did not exhibit neatly defined diets or strict dietary partitioning. Instead, we identify three broad groups: (i) mosasaurs Carinodens belgicus and Plioplatecarpus marshi plotting in the space of modern reptiles that are predominantly piscivorous and/or consume harder invertebrate prey, (ii) Prognathodon saturator and Prognathodon sectorius overlapping with extant reptiles that consume larger amounts of softer invertebrate prey items, and (iii) Mosasaurus hoffmanni spanning a larger plot area in terms of dietary constraints. The clear divide between the aforementioned first two groups in texture-dietary space indicates that, despite our small sample sizes, this method shows the potential of DMTA to test hypotheses and provide quantitative constraints on mosasaur diets and ecological roles.

Evidence for heterothermic endothermy and reptile-like eggshell mineralization in Troodon, a non-avian maniraptoran theropod.

The dinosaur-bird transition involved several anatomical, biomechanical, and physiological modifications of the theropod bauplan. Non-avian maniraptoran theropods, such as Troodon, are key to better understand changes in thermophysiology and reproduction occurring during this transition. Here, we applied dual clumped isotope (Δ47 and Δ48) thermometry, a technique that resolves mineralization temperature and other nonthermal information recorded in carbonates, to eggshells from Troodon, modern reptiles, and modern birds. Troodon eggshells show variable temperatures, namely 42 and 29 ± 2 °C, supporting the hypothesis of an endothermic thermophysiology with a heterothermic strategy for this extinct taxon. Dual clumped isotope data also reveal physiological differences in the reproductive systems between Troodon, reptiles, and birds. Troodon and modern reptiles mineralize their eggshells indistinguishable from dual clumped isotope equilibrium, while birds precipitate eggshells characterized by a positive disequilibrium offset in Δ48. Analyses of inorganic calcites suggest that the observed disequilibrium pattern in birds is linked to an amorphous calcium carbonate (ACC) precursor, a carbonate phase known to accelerate eggshell formation in birds. Lack of disequilibrium patterns in reptile and Troodon eggshells implies these vertebrates had not acquired the fast, ACC-based eggshell calcification process characteristic of birds. Observation that Troodon retained a slow reptile-like calcification suggests that it possessed two functional ovaries and was limited in the number of eggs it could produce; thus its large clutches would have been laid by several females. Dual clumped isotope analysis of eggshells of extinct vertebrates sheds light on physiological information otherwise inaccessible in the fossil record.

The cranial anatomy and relationships of Cardiocorax mukulu (Plesiosauria: Elasmosauridae) from Bentiaba, Angola.

We report a new specimen of the plesiosaur Cardiocorax mukulu that includes the most complete plesiosaur skull from sub-Saharan Africa. The well-preserved three-dimensional nature of the skull offers rare insight into the cranial anatomy of elasmosaurid plesiosaurians. The new specimen of Cardiocorax mukulu was recovered from Bentiaba, Namibe Province in Angola, approximately three meters above the holotype. The new specimen also includes an atlas-axis complex, seventeen postaxial cervical vertebrae, partial ribs, a femur, and limb elements. It is identified as Cardiocorax mukulu based on an apomorphy shared with the holotype where the cervical neural spine is approximately as long anteroposteriorly as the centrum and exhibits a sinusoidal anterior margin. The new specimen is nearly identical to the holotype and previously referred material in all other aspects. Cardiocorax mukulu is returned in an early-branching or intermediate position in Elasmosauridae in four out of the six of our phylogenetic analyses. Cardiocorax mukulu lacks the elongated cervical vertebrae that is characteristic of the extremely long-necked elasmosaurines, and the broad skull with and a high number of maxillary teeth (28-40) which is characteristic of Aristonectinae. Currently, the most parsimonious explanation concerning elasmosaurid evolutionary relationships, is that Cardiocorax mukulu represents an older lineage of elasmosaurids in the Maastrichtian.

Correction to 'Natural Frequency Method: estimating the preferred walking speed of Tyrannosaurus rex based on tail natural frequency'.

[This corrects the article DOI: 10.1098/rsos.201441.][This corrects the article DOI: 10.1098/rsos.201441.].

Natural Frequency Method: estimating the preferred walking speed of Tyrannosaurus rex based on tail natural frequency.

Locomotor energetics are an important determinant of an animal's ecological niche. It is commonly assumed that animals minimize locomotor energy expenditure by selecting gait kinematics tuned to the natural frequencies of relevant body parts. We demonstrate that this allows estimation of the preferred step frequency and walking speed of Tyrannosaurus rex, using an approach we introduce as the Natural Frequency Method. Although the tail of bipedal dinosaurs was actively involved in walking, it was suspended passively by the caudal interspinous ligaments. These allowed for elastic energy storage, thereby reducing the metabolic cost of transport. In order for elastic energy storage to be high, step and natural frequencies would have to be matched. Using a 3D morphological reconstruction and a spring-suspended biomechanical model, we determined the tail natural frequency of T. rex (0.66 s-1, range 0.41-0.84), and the corresponding walking speed (1.28 m s-1, range 0.80-1.64), which we argue to be a good indicator of preferred walking speed (PWS). The walking speeds found here are lower than earlier estimations for large theropods, but agree quite closely with PWS of a diverse group of extant animals. The results are most sensitive to uncertainties regarding ligament moment arms, vertebral kinematics and ligament composition. However, our model formulation and method for estimation of walking speed are unaffected by assumptions regarding muscularity, and therefore offer an independent line of evidence within the field of dinosaur locomotion.

Relevance of the eastern African coastal forest for early hominin biogeography.

The influence of climate change on hominin evolution is much debated. Two issues hamper our understanding of this process: the limited hominin fossil record, and incomplete knowledge about hominin spatial occupation of Africa. Here, we analyze the presently known hominin fossil distribution pattern and explore the potential geographic distribution of hominins between ∼4.5 and ∼2.5 Ma. We focus on assessing the relevance of the Coastal Forest of Eastern Africa (CFEA) along the Indian Ocean as a core area for early hominin evolution. Based on biogeographic-phylogeographic data we propose the coastal refuge hypothesis: the CFEA provided a refugium for early hominins in periods of variable climate and strong seasonality during eccentricity maxima. From this refuge, evolved species could disperse inland (e.g. to rift basins) via vegetated humid corridors, whenever onset of stable climate periods with low seasonality during eccentricity minima allowed expansion out of the coastal enclave. We develop a conceptual model in time and space, comparing predictions with climatic and hominin fossil records. The results imply that: 1) between ∼4.5 and 3 Ma, ongoing (mostly anagenetic) hominin evolution occurred in the CFEA, punctuated by inland dispersal events at ∼4.4, 4.2, 3.8, 3.5, and 3.2 Ma; 2) before ∼3 Ma, the Afar Basin was a (sub)core area often connected to and relatively similar to the CFEA, while other inland areas were more or less marginal for early hominin habitation; 3) after ∼3 Ma, Northern Hemisphere Glaciation exerted strong influence by causing latitudinal contraction of the CFEA, leading to habitat fragmentation, isolation of hominin populations and possible cladogenetic evolution. A major challenge for the coastal refuge model is the fact that at present, no (hominin) fossils are known from the CFEA. We consider how this can be explained, and possibly overcome with targeted search efforts. Furthermore we discuss how the model can be tested, e.g. with molecular phylogeography approaches, and used to predict new hominin fossil locations. With this study, we hope to contribute a fresh perspective to the climate-evolution debate, emphasizing the role of climatic stability, length of dry season and vegetation cover to facilitate connectivity between hominin core and marginal habitats.

Synchrotron imaging of dentition provides insights into the biology of Hesperornis and Ichthyornis, the "last" toothed birds.

BACKGROUND: The dentitions of extinct organisms can provide pivotal information regarding their phylogenetic position, as well as paleobiology, diet, development, and growth. Extant birds are edentulous (toothless), but their closest relatives among stem birds, the Cretaceous Hesperornithiformes and Ichthyornithiformes, retained teeth. Despite their significant phylogenetic position immediately outside the avian crown group, the dentitions of these taxa have never been studied in detail. To obtain new insight into the biology of these 'last' toothed birds, we use cutting-edge visualisation techniques to describe their dentitions at unprecedented levels of detail, in particular propagation phase contrast x-ray synchrotron microtomography at high-resolution. RESULTS: Among other characteristics of tooth shape, growth, attachment, implantation, replacement, and dental tissue microstructures, revealed by these analyses, we find that tooth morphology and ornamentation differ greatly between the Hesperornithiformes and Ichthyornithiformes. We also highlight the first Old World, and youngest record of the major Mesozoic clade Ichthyornithiformes. Both taxa exhibit extremely thin and simple enamel. The extension rate of Hesperornis tooth dentine appears relatively high compared to non-avian dinosaurs. Root attachment is found for the first time to be fully thecodont via gomphosis in both taxa, but in Hesperornis secondary evolution led to teeth implantation in a groove, at least locally without a periodontal ligament. Dental replacement is shown to be lingual via a resorption pit in the root, in both taxa. CONCLUSIONS: Our results allow comparison with other archosaurs and also mammals, with implications regarding dental character evolution across amniotes. Some dental features of the 'last' toothed birds can be interpreted as functional adaptations related to diet and mode of predation, while others appear to be products of their peculiar phylogenetic heritage. The autapomorphic Hesperornis groove might have favoured firmer root attachment. These observations highlight complexity in the evolutionary history of tooth reduction in the avian lineage and also clarify alleged avian dental characteristics in the frame of a long-standing debate on bird origins. Finally, new hypotheses emerge that will possibly be tested by further analyses of avian teeth, for instance regarding dental replacement rates, or simplification and thinning of enamel throughout the course of early avian evolution.

Hadrosauroid Dinosaurs from the Late Cretaceous of the Sultanate of Oman.

Fragmentary post-cranial remains (femora, tibia, vertebrae) of ornithischian dinosaurs from the Late Cretaceous of the Sultanate of Oman are described and referred to hadrosauroids. The specimens come from the Al-Khod Conglomerate, of latest Campanian to Maastrichtian age, in the north-eastern part of the country. Although the fragmentary condition of the fossils precludes a precise identification, various characters, including the shape of the fourth trochanter of the femur and the morphology of its distal end, support an attribution to hadrosauroids. With the possible exception of a possible phalanx from Angola, this group of ornithopod dinosaurs, which apparently originated in Laurasia, was hitherto unreported from the Afro-Arabian plate. From a paleobiogeographical point of view, the presence of hadrosauroids in Oman in all likelihood is a result of trans-Tethys dispersal from Asia or Europe, probably by way of islands in the Tethys shown on all recent paleogeographical maps of that area. Whether hadrosauroids were widespread on the Afro-Arabian landmass in the latest Cretaceous, or where restricted to the « Oman island ¼ shown on some paleogeographical maps, remains to be determined.

Homo erectus at Trinil on Java used shells for tool production and engraving.

The manufacture of geometric engravings is generally interpreted as indicative of modern cognition and behaviour. Key questions in the debate on the origin of such behaviour are whether this innovation is restricted to Homo sapiens, and whether it has a uniquely African origin. Here we report on a fossil freshwater shell assemblage from the Hauptknochenschicht ('main bone layer') of Trinil (Java, Indonesia), the type locality of Homo erectus discovered by Eugène Dubois in 1891 (refs 2 and 3). In the Dubois collection (in the Naturalis museum, Leiden, The Netherlands) we found evidence for freshwater shellfish consumption by hominins, one unambiguous shell tool, and a shell with a geometric engraving. We dated sediment contained in the shells with (40)Ar/(39)Ar and luminescence dating methods, obtaining a maximum age of 0.54 ± 0.10 million years and a minimum age of 0.43 ± 0.05 million years. This implies that the Trinil Hauptknochenschicht is younger than previously estimated. Together, our data indicate that the engraving was made by Homo erectus, and that it is considerably older than the oldest geometric engravings described so far. Although it is at present not possible to assess the function or meaning of the engraved shell, this discovery suggests that engraving abstract patterns was in the realm of Asian Homo erectus cognition and neuromotor control.

Angolatitan adamastor, a new sauropod dinosaur and the first record from Angola.

A forelimb of a new sauropod dinosaur (Angolatitan adamastor n. gen. et sp.) from the Late Turonian of Iembe (Bengo Province) represents the first dinosaur discovery in Angola, and is one of the few occurrences of sauropod dinosaurs in sub-Saharan Africa collected with good chronological controls. The marginal marine sediments yielding the specimen are reported to be late Turonian in age and, thus it represents a non-titanosaurian sauropod in sub-Saharan Africa at a time taken to be dominated by titanosaurian forms. Moreover, Angolatitan adamastor is the only basal Somphospondyli known in the Late Cretaceous which implies in the existence of relict forms in Africa.

Angolatitan adamastor, a new sauropod dinosaur and the first record from Angola

A forelimb of a new sauropod dinosaur (Angolatitan adamastor n. gen. et sp.) from the Late Turonian of Iembe (Bengo Province) represents the first dinosaur discovery in Angola, and is one of the few occurrences of sauropod dinosaurs in sub-Saharan Africa collected with good chronological controls. The marginal marine sediments yielding the specimen are reported to be late Turonian in age and, thus it represents a non-titanosaurian sauropod in sub-Saharan Africa at a time taken to be dominated by titanosaurian forms. Moreover, Angolatitan adamastor is the only basal Somphospondyli known in the Late Cretaceous which implies in the existence of relict forms in Africa.

Um membro anterior de um novo dinossauro saurópode (Angolatitan adamastor n. gen. et sp.) do Turoniano Superior de Iembe (Bengo) representa a descoberta do primeiro dinossauro em Angola e é uma das poucas ocorrências de dinossauros saurópodes na África Subsaariana recolhidos com bons controles cronológicos. Os sedimentos marinhos marginais de onde provém o espécime estão datados do Turoniano superior e, portanto, trata-se de um saurópodes não-titanossauro na África subsaariana onde predominavam titanossauros. Além disso, Angolatitan adamastor é o único Somphospondyli basal conhecido no Cretáceo Superior, o que implica a sobrevivência de formas-relíquia na África.

First dinosaur tracks from the Arabian Peninsula.

BACKGROUND: The evolutionary history of Mesozoic terrestrial vertebrates from the Arabian Peninsula is virtually unknown. Despite vast exposures of rocky outcrops, only a handful of fossils have yet been described from the region. Here we report a multi-taxon dinosaur track assemblage near Madar village, 47 km north of Sana'a, Republic of Yemen. This represents the first dinosaur tracksite from the Arabian Peninsula, and the only multi-taxon dinosaur ichnosite in the Middle East. METHODOLOGY/FINDINGS: Measurements were taken directly from trackway impressions, following standard ichnological conventions. The presence of bipedal trackmakers is evidenced by a long series of pes imprints preserving smoothly rounded posterior margins, no evidence of a hallux, bluntly rounded digit tips and digital divarication angles characteristic of ornithopod dinosaurs. Nearby, eleven parallel quadrupedal trackways document a sauropod herd that included large and small individuals traveling together. Based on the morphology of manus impressions along with a narrow-gauged stance, the quadrupedal trackways were made by non-titanosauriform neosauropods. Additional isolated tracks and trackways of sauropod and ornithopod dinosaurs are preserved nearby. CONCLUSIONS/SIGNIFICANCE: Taken together, these discoveries present the most evocative window to date into the evolutionary history of dinosaurs of the Arabian Peninsula. Given the limited Mesozoic terrestrial record from the region, this discovery is of both temporal and geographic significance, and massive exposures of similarly-aged outcrops nearby offer great promise for future discoveries.

Europe's last Mesozoic bird.

Birds known from more than isolated skeletal elements are rare in the fossil record, especially from the European Mesozoic. This paucity has hindered interpretations of avian evolution immediately prior to, and in the aftermath of, the Cretaceous-Tertiary (K-T) extinction event. We report on a specimen of a large ornithurine bird (closely related to Ichthyornis) from the uppermost Cretaceous (Maastricht Formation) of Belgium. This is the first record of a bird from these historic strata and the only phylogenetically informative ornithurine to be recovered from the Mesozoic of Europe. Because this new specimen was collected from 40 m below the K-T boundary (approximate age of 65.8 Ma), it is also the youngest non-neornithine (= non-modern) bird known from anywhere in the world.