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Concept: Paleontology

554

We review the salient evidence consistent with or predicted by the Hoyle-Wickramasinghe (H- W) thesis of Cometary (Cosmic) Biology. Much of this physical and biological evidence is multifactorial. One particular focus are the recent studies which date the emergence of the complex retroviruses of vertebrate lines at or just before the Cambrian Explosion of ∼500 Ma. Such viruses are known to be plausibly associated with major evolutionary genomic processes. We believe this coincidence is not fortuitous but is consistent with a key prediction of H-W theory whereby major extinction-diversification evolutionary boundaries coincide with virus-bearing cometary-bolide bombardment events. A second focus is the remarkable evolution of intelligent complexity (Cephalopods) culminating in the emergence of the Octopus. A third focus concerns the micro-organism fossil evidence contained within meteorites as well as the detection in the upper atmosphere of apparent incoming life-bearing particles from space. In our view the totality of the multifactorial data and critical analyses assembled by Fred Hoyle, Chandra Wickramasinghe and their many colleagues since the 1960s leads to a very plausible conclusion - life may have been seeded here on Earth by life-bearing comets as soon as conditions on Earth allowed it to flourish (about or just before 4.1 Billion years ago); and living organisms such as space-resistant and space-hardy bacteria, viruses, more complex eukaryotic cells, fertilised ova and seeds have been continuously delivered ever since to Earth so being one important driver of further terrestrial evolution which has resulted in considerable genetic diversity and which has led to the emergence of mankind.

Concepts: DNA, Gene, Genetics, Bacteria, Evolution, Organism, Species, Paleontology

427

New ages for flowstone, sediments and fossil bones from the Dinaledi Chamber are presented. We combined optically stimulated luminescence dating of sediments with U-Th and palaeomagnetic analyses of flowstones to establish that all sediments containing Homo naledi fossils can be allocated to a single stratigraphic entity (sub-unit 3b), interpreted to be deposited between 236 ka and 414 ka. This result has been confirmed independently by dating three H. naledi teeth with combined U-series and electron spin resonance (US-ESR) dating. Two dating scenarios for the fossils were tested by varying the assumed levels of (222)Rn loss in the encasing sediments: a maximum age scenario provides an average age for the two least altered fossil teeth of 253 +82/-70 ka, whilst a minimum age scenario yields an average age of 200 +70/-61 ka. We consider the maximum age scenario to more closely reflect conditions in the cave, and therefore, the true age of the fossils. By combining the US-ESR maximum age estimate obtained from the teeth, with the U-Th age for the oldest flowstone overlying Homo naledi fossils, we have constrained the depositional age of Homo naledi to a period between 236 ka and 335 ka. These age results demonstrate that a morphologically primitive hominin, Homo naledi, survived into the later parts of the Pleistocene in Africa, and indicate a much younger age for the Homo naledi fossils than have previously been hypothesized based on their morphology.

Concepts: Electron, Spin, Sediment, Sedimentary rock, Geology, Fossil, Paleontology, Geologic time scale

359

The fossil record of ceratopsid dinosaurs between the occurrence of their proximate sister taxa in the Turonian and the beginning of their well-documented radiation from the late Campanian of North America onwards (approximately 90 and 77 Ma) is poor, with only seven taxa described from this early period in their evolution. We describe a new taxon of a highly adorned basal centrosaurine, Wendiceratops pinhornensis gen. et sp. nov., from the lower part of the Oldman Formation (middle Campanian, approximately 78-79 Ma), Alberta, Canada. Over 200 bones derived from virtually all parts of the skeleton, including multiple well-preserved specimens of the diagnostic parietosquamosal frill, were collected from a medium-density monodominant bonebed, making the new taxon one of the best-represented early ceratopsids. The new taxon is apomorphic in having epiparietals at loci 2 and 3 developed as broad-based, pachyostotic processes that are strongly procurved anterodorsally to overhang the posterior and lateral parietal rami, and an ischium with a broad, rectangular distal terminus. Although the morphology of the nasal is incompletely known, Wendiceratops is inferred to have a large, upright nasal horn located close to the orbits, which represents the oldest occurrence of this feature in Ceratopsia. Given the phylogenetic position of the new taxon within Centrosaurinae, a enlarged nasal horn is hypothesized to have arisen independently at least twice in ceratopsid evolution.

Concepts: Phylogenetics, Cladistics, Fossil, Paleontology, Dinosaur, Taxon, Ceratopsidae, Albertaceratops

281

Large plant-eating dinosaurs are usually presumed to have been strictly herbivorous, because their derived teeth and jaws were capable of processing fibrous plant foods. This inferred feeding behavior offers a generalized view of dinosaur food habits, but rare direct fossil evidence of diet provides more nuanced insights into feeding behavior. Here we describe fossilized feces (coprolites) that demonstrate recurring consumption of crustaceans and rotted wood by large Late Cretaceous dinosaurs. These multi-liter coprolites from the Kaiparowits Formation are primarily composed of comminuted conifer wood tissues that were fungally degraded before ingestion. Thick fragments of laminar crustacean cuticle are scattered within the coprolite contents and suggest that the dinosaurian defecators consumed sizeable crustaceans that sheltered in rotting logs. The diet of decayed wood and crustaceans offered a substantial supply of plant polysaccharides, with added dividends of animal protein and calcium. Nevertheless, it is unlikely that the fossilized fecal residues depict year-round feeding habits. It is more reasonable to infer that these coprolites reflected seasonal dietary shifts-possibly related to the dinosaurs' oviparous breeding activities. This surprising fossil evidence challenges conventional notions of herbivorous dinosaur diets and reveals a degree of dietary flexibility that is consistent with that of extant herbivorous birds.

Concepts: Nutrition, Extinction, Cretaceous, Fossil, Paleontology, Dinosaur, Carnivore, Late Cretaceous

261

Countershading was one of the first proposed mechanisms of camouflage [1, 2]. A dark dorsum and light ventrum counteract the gradient created by illumination from above, obliterating cues to 3D shape [3-6]. Because the optimal countershading varies strongly with light environment [7-9], pigmentation patterns give clues to an animal’s habitat. Indeed, comparative evidence from ungulates [9] shows that interspecific variation in countershading matches predictions: in open habitats, where direct overhead sunshine dominates, a sharp dark-light color transition high up the body is evident; in closed habitats (e.g., under forest canopy), diffuse illumination dominates and a smoother dorsoventral gradation is found. We can apply this approach to extinct animals in which the preservation of fossil melanin allows reconstruction of coloration [10-15]. Here we present a study of an exceptionally well-preserved specimen of Psittacosaurus sp. from the Chinese Jehol biota [16, 17]. This Psittacosaurus was countershaded [16] with a light underbelly and tail, whereas the chest was more pigmented. Other patterns resemble disruptive camouflage, whereas the chin and jugal bosses on the face appear dark. We projected the color patterns onto an anatomically accurate life-size model in order to assess their function experimentally. The patterns are compared to the predicted optimal countershading from the measured radiance patterns generated on an identical uniform gray model in direct versus diffuse illumination. These studies suggest that Psittacosaurus sp. inhabited a closed habitat such as a forest with a relatively dense canopy. VIDEO ABSTRACT.

Concepts: Habitat, Light, Sunlight, Color, Lighting, Pigment, Paleontology, Dinosaur

251

Defecation in communal latrines is a common behaviour of extant mammals widely distributed among megaherbivores. This behaviour has key social functions with important biological and ecological implications. Herbivore communal latrines are only documented among mammals and their fossil record is exceptionally restricted to the late Cenozoic. Here we report the discovery of several massive coprolite associations in the Middle-Late Triassic of the Chañares Formation, Argentina, which represent fossil communal latrines based on a high areal density, small areal extension and taphonomic attributes. Several lines of evidence (size, morphology, abundance and coprofabrics) and their association with kannemeyeriiform dicynodonts indicate that these large synapsids produced the communal latrines and had a gregarious behaviour comparable to that of extant megaherbivores. This is the first evidence of megaherbivore communal latrines in non-mammal vertebrates, indicating that this mammal-type behaviour was present in distant relatives of mammals, and predates its previous oldest record by 220 Mya.

Concepts: Ecology, Synapsid, Fossil, Paleontology, Dinosaur, Triassic, Mesozoic, Permian

235

Sauropod dinosaurs are a group of herbivorous dinosaurs which exceeded all other terrestrial vertebrates in mean and maximal body size. Sauropod dinosaurs were also the most successful and long-lived herbivorous tetrapod clade, but no abiological factors such as global environmental parameters conducive to their gigantism can be identified. These facts justify major efforts by evolutionary biologists and paleontologists to understand sauropods as living animals and to explain their evolutionary success and uniquely gigantic body size. Contributions to this research program have come from many fields and can be synthesized into a biological evolutionary cascade model of sauropod dinosaur gigantism (sauropod gigantism ECM). This review focuses on the sauropod gigantism ECM, providing an updated version based on the contributions to the PLoS ONE sauropod gigantism collection and on other very recent published evidence. The model consist of five separate evolutionary cascades (“Reproduction”, “Feeding”, “Head and neck”, “Avian-style lung”, and “Metabolism”). Each cascade starts with observed or inferred basal traits that either may be plesiomorphic or derived at the level of Sauropoda. Each trait confers hypothetical selective advantages which permit the evolution of the next trait. Feedback loops in the ECM consist of selective advantages originating from traits higher in the cascades but affecting lower traits. All cascades end in the trait “Very high body mass”. Each cascade is linked to at least one other cascade. Important plesiomorphic traits of sauropod dinosaurs that entered the model were ovipary as well as no mastication of food. Important evolutionary innovations (derived traits) were an avian-style respiratory system and an elevated basal metabolic rate. Comparison with other tetrapod lineages identifies factors limiting body size.

Concepts: Evolution, Metabolism, Bird, Paleontology, Sauropoda, Dinosaur, Sauropodomorpha, Titanosaur

224

The highly derived morphology and astounding diversity of snakes has long inspired debate regarding the ecological and evolutionary origin of both the snake total-group (Pan-Serpentes) and crown snakes (Serpentes). Although speculation abounds on the ecology, behavior, and provenance of the earliest snakes, a rigorous, clade-wide analysis of snake origins has yet to be attempted, in part due to a dearth of adequate paleontological data on early stem snakes. Here, we present the first comprehensive analytical reconstruction of the ancestor of crown snakes and the ancestor of the snake total-group, as inferred using multiple methods of ancestral state reconstruction. We use a combined-data approach that includes new information from the fossil record on extinct crown snakes, new data on the anatomy of the stem snakes Najash rionegrina, Dinilysia patagonica, and Coniophis precedens, and a deeper understanding of the distribution of phenotypic apomorphies among the major clades of fossil and Recent snakes. Additionally, we infer time-calibrated phylogenies using both new ‘tip-dating’ and traditional node-based approaches, providing new insights on temporal patterns in the early evolutionary history of snakes.

Concepts: Evolution, Biology, Phylogenetics, Fossil, Paleontology, Snake, Dinosaur, Najash

215

The use of limbs for foraging is documented in both marine and terrestrial tetrapods. These behaviors were once believed to be less likely in marine tetrapods due to the physical constraints of body plans adapted to locomotion in a fluid environment. Despite these obstacles, ten distinct types of limb-use while foraging have been previously reported in nine marine tetrapod families. Here, we expand the types of limb-use documented in marine turtles and put it in context with the diversity of marine tetrapods currently known to use limbs for foraging. Additionally, we suggest that such behaviors could have occurred in ancestral turtles, and thus, possibly extend the evolutionary timeline of limb-use behavior in marine tetrapods back approximately 70 million years. Through direct observationin situand crowd-sourcing, we document the range of behaviors across habitats and prey types, suggesting its widespread occurrence. We argue the presence of these behaviors among marine tetrapods may be limited by limb mobility and evolutionary history, rather than foraging ecology or social learning. These behaviors may also be remnant of ancestral forelimb-use that have been maintained due to a semi-aquatic life history.

Concepts: Evolution, Ecology, Evolutionary biology, Behavior, Tetrapod, Paleontology, Evolutionary history of life, Ichthyostega

215

Fossorialism is a beneficial adaptation for brooding, predator avoidance and protection from extreme climate. The abundance of fossilised burrow casts from the Early Triassic of southern Africa is viewed as a behavioural response by many tetrapods to the harsh conditions following the Permo-Triassic mass-extinction event. However, scarcity of vertebrate remains associated with these burrows leaves many ecological questions unanswered. Synchrotron scanning of a lithified burrow cast from the Early Triassic of the Karoo unveiled a unique mixed-species association: an injured temnospondyl amphibian (Broomistega) that sheltered in a burrow occupied by an aestivating therapsid (Thrinaxodon). The discovery of this rare rhinesuchid represents the first occurrence in the fossil record of a temnospondyl in a burrow. The amphibian skeleton shows signs of a crushing trauma with partially healed fractures on several consecutive ribs. The presence of a relatively large intruder in what is interpreted to be a Thrinaxodon burrow implies that the therapsid tolerated the amphibian’s presence. Among possible explanations for such unlikely cohabitation, Thrinaxodon aestivation is most plausible, an interpretation supported by the numerous Thrinaxodon specimens fossilised in curled-up postures. Recent advances in synchrotron imaging have enabled visualization of the contents of burrow casts, thus providing a novel tool to elucidate not only anatomy but also ecology and biology of ancient tetrapods.

Concepts: Evolution, Tetrapod, Synapsid, Paleontology, Frog, Dinosaur, Triassic