The osteology of the Late Triassic reptile Scleromochlus taylori from µCT data.

Scleromochlus taylori is one of the most enigmatic members of the herpetofauna from the Lossiemouth Sandstone Formation (Upper Triassic) of Elgin (Moray, Scotland). For many years it was thought to be closely related to pterosaurs and dinosaurs, but the anatomy of this animal is difficult to interpret because of the notoriously poor preservation of the six available specimens, which comprise void space in the sandstone after the bones were destroyed by diagenesis. Historically, these fossils have been studied using physical molds, which provide only incomplete, and potentially distorted, information. Due to these uncertainties, interpretations of the anatomy, phylogenetic relationships, and paleobiology of Scleromochlus taylori have remained contentious. Here, we use microcomputed tomographic (µCT) techniques to redescribe and illustrate the osteology of Scleromochlus in detail, building upon a short redescription of keystone features of the anatomy that we recently published. We digitally visualize, describe, and figure previously inaccessible-and thus unaltered-portions of its skeleton, as well as providing new observations on the exposed parts of each specimen. This work reveals many novel features of the skull, mandible, trunk, tail, girdles, forelimb, and hindlimb (particularly of the manus, femur, and pes), demonstrating that historic molding techniques failed, in some cases, to accurately capture the anatomy of Scleromochlus. Our review sheds light on some of the most controversial aspects of Scleromochlus morphology showing that this taxon retains plesiomorphic features of Avemetatarsalia in the postcranial skeleton, alongside a suite of synapomorphies diagnostic of pterosauromorphs (the broad clade of pterosaurs and taxa more closely related to them than dinosaurs), particularly one subgroup, the lagerpetids. Consistent with recent work, our updated phylogenetic analyses (Maximum Parsimony and Bayesian Inference) demonstrate that Scleromochlus taylori is an avemetatarsalian archosaur that is recovered firmly in an early diverging position within Pterosauromorpha, as a member of Lagerpetidae, thus shedding important information on the origin of pterosaurs, the first group of vertebrates to evolve powered flight.

The virtual brain endocast of Trogosus (Mammalia, Tillodontia) and its relevance in understanding the extinction of archaic placental mammals.

After successfully diversifying during the Paleocene, the descendants of the first wave of mammals that survived the end-Cretaceous mass extinction waned throughout the Eocene. Competition with modern crown clades and intense climate fluctuations may have been part of the factors leading to the extinction of these archaic groups. Why these taxa went extinct has rarely been studied from the perspective of the nervous system. Here, we describe the first virtual endocasts for the archaic order Tillodontia. Three species from the middle Eocene of North America were analyzed: Trogosus hillsii, Trogosus grangeri, and Trogosus castoridens. We made morphological comparisons with the plaster endocast of another tillodont, Tillodon fodiens, as well as groups potentially related to Tillodontia: Pantodonta, Arctocyonidae, and Cimolesta. Trogosus shows very little inter-specific variation with the only potential difference being related to the fusion of the optic canal and sphenorbital fissure. Many ancestral features are displayed by Trogosus, including an exposed midbrain, small neocortex, orbitotemporal canal ventral to rhinal fissure, and a broad circular fissure. Potential characteristics that could unite Tillodontia with Pantodonta, and Arctocyonidae are the posterior position of cranial nerve V3 exit in relation to the cerebrum and the low degree of development of the subarcuate fossa. The presence of large olfactory bulbs and a relatively small neocortex are consistent with a terrestrial lifestyle. A relatively small neocortex may have put Trogosus at risk when competing with artiodactyls for potentially similar resources and avoiding predation from archaic carnivorans, both of which are known to have had larger relative brain and neocortex sizes in the Eocene. These factors may have possibly exacerbated the extinction of Tillodontia, which showed highly specialized morphologies despite the increase in climate fluctuations throughout the Eocene, before disappearing during the middle Eocene.

Evidence for a novel cranial thermoregulatory pathway in thalattosuchian crocodylomorphs.

Thalattosuchian crocodylomorphs were a diverse clade that lived from the Early Jurassic to the Early Cretaceous. The subclade Metriorhynchoidea underwent a remarkable transition, evolving from semi-aquatic ambush predators into fully aquatic forms living in the open oceans. Thalattosuchians share a peculiar palatal morphology with semi-aquatic and aquatic fossil cetaceans: paired anteroposteriorly aligned grooves along the palatal surface of the bony secondary palate. In extant cetaceans, these grooves are continuous with the greater palatine artery foramina, arteries that supply their oral thermoregulatory structures. Herein, we investigate the origins of thalattosuchian palatal grooves by examining CT scans of six thalattosuchian species (one teleosauroid, two early-diverging metriorhynchoids and three metriorhynchids), and CT scans of eleven extant crocodylian species. All thalattosuchians had paired osseous canals, enclosed by the palatines, that connect the nasal cavity to the oral cavity. These osseous canals open into the oral cavity via foramina at the posterior terminus of the palatal grooves. Extant crocodylians lack both the external grooves and the internal canals. We posit that in thalattosuchians these novel palatal canals transmitted hypertrophied medial nasal vessels (artery and vein), creating a novel heat exchange pathway connecting the palatal vascular plexus to the endocranial region. Given the general hypertrophy of thalattosuchian cephalic vasculature, and their increased blood flow and volume, thalattosuchians would have required a more extensive suite of thermoregulatory pathways to maintain stable temperatures for their neurosensory tissues.

Paleontology: All dinosaurs big and small.

Two new studies illuminate how some dinosaurs were able to grow to enormous sizes, while others shrank to become today's birds.

Shifts in food webs and niche stability shaped survivorship and extinction at the end-Cretaceous.

It has long been debated why groups such as non-avian dinosaurs became extinct whereas mammals and other lineages survived the Cretaceous/Paleogene mass extinction 66 million years ago. We used Markov networks, ecological niche partitioning, and Earth System models to reconstruct North American food webs and simulate ecospace occupancy before and after the extinction event. We find a shift in latest Cretaceous dinosaur faunas, as medium-sized species counterbalanced a loss of megaherbivores, but dinosaur niches were otherwise stable and static, potentially contributing to their demise. Smaller vertebrates, including mammals, followed a consistent trajectory of increasing trophic impact and relaxation of niche limits beginning in the latest Cretaceous and continuing after the mass extinction. Mammals did not simply proliferate after the extinction event; rather, their earlier ecological diversification might have helped them survive.

The ecological diversification and evolution of Teleosauroidea (Crocodylomorpha, Thalattosuchia), with insights into their mandibular biomechanics.

Throughout the Jurassic, a plethora of marine reptiles dominated ocean waters, including ichthyosaurs, plesiosaurs and thalattosuchian crocodylomorphs. These Jurassic ecosystems were characterized by high niche partitioning and spatial variation in dietary ecology. However, while the ecological diversity of many marine reptile lineages is well known, the overall ecological diversification of Teleosauroidea (one of the two major groups within thalattosuchian crocodylomorphs) has never been explored. Teleosauroids were previously deemed to have a morphologically conservative body plan; however, they were in actuality morphofunctionally more diverse than previously thought. Here we investigate the ecology and feeding specializations of teleosauroids, using morphological and functional cranio-dental characteristics. We assembled the most comprehensive dataset to date of teleosauroid taxa (approximately 20 species) and ran a series of principal component analyses (PC) to categorize them into various feeding ecomorphotypes based on 17 dental characteristics (38 specimens) and 16 functionally significant mandibular characters (18 specimens). The results were examined in conjunction with a comprehensive thalattosuchian phylogeny (153 taxa and 502 characters) to evaluate macroevolutionary patterns and significant ecological shifts. Machimosaurids display a well-developed ecological shift from: (1) slender, pointed tooth apices and an elongate gracile mandible; to (2) more robust, pointed teeth with a slightly deeper mandible; and finally, (3) rounded teeth and a deep-set, shortened mandible with enlarged musculature. Overall, there is limited mandibular functional variability in teleosaurids and machimosaurids, despite differing cranial morphologies and habitat preferences in certain taxa. This suggests a narrow feeding ecological divide between teleosaurids and machimosaurids. Resource partitioning was primarily related to snout and skull length as well as habitat; only twice did teleosauroids manage to make a major evolutionary leap to feed distinctly differently, with only the derived machimosaurines successfully radiating into new feeding ecologies.

Scleromochlus and the early evolution of Pterosauromorpha.

Pterosaurs, the first vertebrates to evolve powered flight, were key components of Mesozoic terrestrial ecosystems from their sudden appearance in the Late Triassic until their demise at the end of the Cretaceous1-6. However, the origin and early evolution of pterosaurs are poorly understood owing to a substantial stratigraphic and morphological gap between these reptiles and their closest relatives6, Lagerpetidae7. Scleromochlus taylori, a tiny reptile from the early Late Triassic of Scotland discovered over a century ago, was hypothesized to be a key taxon closely related to pterosaurs8, but its poor preservation has limited previous studies and resulted in controversy over its phylogenetic position, with some even doubting its identification as an archosaur9. Here we use microcomputed tomographic scans to provide the first accurate whole-skeletal reconstruction and a revised diagnosis of Scleromochlus, revealing new anatomical details that conclusively identify it as a close pterosaur relative1 within Pterosauromorpha (the lagerpetid + pterosaur clade). Scleromochlus is anatomically more similar to lagerpetids than to pterosaurs and retains numerous features that were probably present in very early diverging members of Avemetatarsalia (bird-line archosaurs). These results support the hypothesis that the first flying reptiles evolved from tiny, probably facultatively bipedal, cursorial ancestors1.

The origin of placental mammal life histories.

After the end-Cretaceous extinction, placental mammals quickly diversified1, occupied key ecological niches2,3 and increased in size4,5, but this last was not true of other therians6. The uniquely extended gestation of placental young7 may have factored into their success and size increase8, but reproduction style in early placentals remains unknown. Here we present the earliest record of a placental life history using palaeohistology and geochemistry, in a 62 million-year-old pantodont, the clade including the first mammals to achieve truly large body sizes. We extend the application of dental trace element mapping9,10 by 60 million years, identifying chemical markers of birth and weaning, and calibrate these to a daily record of growth in the dentition. A long gestation (approximately 7 months), rapid dental development and short suckling interval (approximately 30-75 days) show that Pantolambda bathmodon was highly precocial, unlike non-placental mammals and known Mesozoic precursors. These results demonstrate that P. bathmodon reproduced like a placental and lived at a fast pace for its body size. Assuming that P. bathmodon reflects close placental relatives, our findings suggest that the ability to produce well-developed, precocial young was established early in placental evolution, and that larger neonate sizes were a possible mechanism for rapid size increase in early placentals.

'Ear stones' in crocodylians: a cross-species comparative and ontogenetic survey of otolith structures.

The vestibular system of the inner ear is a crucial sensory organ, involved in the sensation of balance and equilibrium. It consists of three semicircular canals that sense angular rotations of the head and the vestibule that detects linear acceleration and gravity. The vestibule often contains structures, known as the otoliths or 'ear stones'. Otoliths are present in many vertebrates and are particularly well known from the fossil record of fish, but surprisingly have not been described in detail in most tetrapods, living or extinct. Here, we present for the first time a survey of the otoliths of a broad sample of extant crocodylian species, based on computed tomography scans. We find that otoliths are present in numerous crocodylian species of different growth stages, and they continue to increase in size during ontogeny, with positive allometry compared to skull length. Our results confirm that otoliths are a common component of the crocodylian vestibular system, and suggest they play an important role in sensory detection. Otoliths are likely common, but overlooked, constituents of the inner ear in tetrapods, and a broader study of their size, shape and distribution promises insight into sensory abilities.

Brawn before brains in placental mammals after the end-Cretaceous extinction.

Mammals are the most encephalized vertebrates, with the largest brains relative to body size. Placental mammals have particularly enlarged brains, with expanded neocortices for sensory integration, the origins of which are unclear. We used computed tomography scans of newly discovered Paleocene fossils to show that contrary to the convention that mammal brains have steadily enlarged over time, early placentals initially decreased their relative brain sizes because body mass increased at a faster rate. Later in the Eocene, multiple crown lineages independently acquired highly encephalized brains through marked growth in sensory regions. We argue that the placental radiation initially emphasized increases in body size as extinction survivors filled vacant niches. Brains eventually became larger as ecosystems saturated and competition intensified.

An exquisitely preserved in-ovo theropod dinosaur embryo sheds light on avian-like prehatching postures.

Despite the discovery of many dinosaur eggs and nests over the past 100 years, articulated in-ovo embryos are remarkably rare. Here we report an exceptionally preserved, articulated oviraptorid embryo inside an elongatoolithid egg, from the Late Cretaceous Hekou Formation of southern China. The head lies ventral to the body, with the feet on either side, and the back curled along the blunt pole of the egg, in a posture previously unrecognized in a non-avian dinosaur, but reminiscent of a late-stage modern bird embryo. Comparison to other late-stage oviraptorid embryos suggests that prehatch oviraptorids developed avian-like postures late in incubation, which in modern birds are related to coordinated embryonic movements associated with tucking - a behavior controlled by the central nervous system, critical for hatching success. We propose that such pre-hatching behavior, previously considered unique to birds, may have originated among non-avian theropods, which can be further investigated with additional discoveries of embryo fossils.

A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs.

Pterosaurs were the first vertebrates to evolve flight1,2 and include the largest flying animals in Earth history.3,4 While some of the last-surviving species were the size of airplanes, pterosaurs were long thought to be restricted to small body sizes (wingspans ca. <1.8-1.6 m) from their Triassic origins through the Jurassic, before increasing in size when derived long-skulled and short-tailed pterodactyloids lived alongside a diversity of birds in the Cretaceous.5 We report a new spectacularly preserved three-dimensional skeleton from the Middle Jurassic of Scotland, which we assign to a new genus and species: Dearc sgiathanach gen. et sp. nov. Its wingspan is estimated at >2.5 m, and bone histology shows it was a juvenile-subadult still actively growing when it died, making it the largest known Jurassic pterosaur represented by a well-preserved skeleton. A review of fragmentary specimens from the Middle Jurassic of England demonstrates that a diversity of pterosaurs was capable of reaching larger sizes at this time but have hitherto been concealed by a poor fossil record. Phylogenetic analysis places D. sgiathanach in a clade of basal long-tailed non-monofenestratan pterosaurs, in a subclade of larger-bodied species (Angustinaripterini) with elongate skulls convergent in some aspects with pterodactyloids.6 Far from a static prologue to the Cretaceous, the Middle Jurassic was a key interval in pterosaur evolution, in which some non-pterodactyloids diversified and experimented with larger sizes, concurrent with or perhaps earlier than the origin of birds. VIDEO ABSTRACT.

Rostral neurovasculature indicates sensory trade-offs in Mesozoic pelagic crocodylomorphs.

Metriorhynchoid thalattosuchians were a marine clade of Mesozoic crocodylomorphs that evolved from semi-aquatic, "gharial"-like species into the obligately pelagic subclade Metriorhynchidae. To explore whether the sensory and physiological demands of underwater life necessitates a shift in rostral anatomy, both in neurology and vasculature, we investigate the trigeminal innervation and potential somatosensory abilities of metriorhynchoids by digitally segmenting the rostral neurovascular canals in CT scans of 10 extant and extinct crocodyliforms. The dataset includes the terrestrial, basal crocodyliform Protosuchus haughtoni, two semi-aquatic basal metriorhynchoids, four pelagic metriorhynchids and three extant, semi-aquatic crocodylians. In the crocodylian and basal metriorhynchoid taxa, we find three main neurovascular channels running parallel to one another posteroanteriorly down the length of the snout, whereas in metriorhynchids there are two, and in P. haughtoni only one. Crocodylians appear to be unique in their extensive trigeminal innervation, which is used to supply the integumentary sensory organs (ISOs) involved with their facial somatosensory abilities. Crocodylians have a far higher number of foramina on the maxillary bones than either metriorhynchoids or P. haughtoni, suggesting that the fossil taxa lacked the somatosensory abilities seen in extant species. We posit that the lack of ISO osteological correlates in metriorhynchoids is due to their basal position in Crocodyliformes, rather than a pelagic adaptation. This is reinforced by the hypothesis that extant crocodyliforms, and possibly some neosuchian clades, underwent a long "nocturnal bottleneck"-hinting that their complex network of ISOs evolved in Neosuchia, as a sensory trade-off to compensate for poorer eyesight.

Paranasal sinus system and upper respiratory tract evolution in Mesozoic pelagic crocodylomorphs.

Thalattosuchians were a predominately marine clade of Mesozoic crocodylomorphs, including semi-aquatic teleosauroid and obligately pelagic metriorhynchid subclades. Recent advances in our understanding of thalattosuchian endocranial anatomy have revealed new details of the evolutionary transition from terrestrial to marine to pelagic taxa. Paranasal sinuses, however, have received little attention. Herein, we investigate the evolution of the paranasal sinus system and part of the upper respiratory system (nasopharyngeal ducts) in Thalattosuchia, by reconstructing the nasal and paranasal anatomy in CT scans of seven thalattosuchian skulls: one teleosauroid, two basal metriorhynchoids and four metriorhynchids. Our outgroups were: three extant crocodylian species (including adult and subadult skulls) and the basal crocodyliform Protosuchus. We found thalattosuchians exhibit exceptionally reduced paranasal sinus systems, solely comprising the antorbital sinus, as has been previously proposed. The semi-aquatic basal thalattosuchians Palgiopthalmosuchus gracilirostris and Pelagosaurus typus both have an antorbital sinus partially located medial to a reduced external antorbital fenestra and broadly communicating with the dorsal alveolar canal. In pelagic metriorhynchids, the antorbital cavity is more extensive than in basal taxa and possibly had an active function associated with a hypothesized accessory suborbital diverticulum, but our reconstructions are insufficient to confirm or reject the presence of such a diverticulum. The nasopharyngeal ducts of metriorhynchids are dorsoventrally enlarged, possibly enabling stronger ventilation. The sequence of acquisition of craniofacial adaptations show a mosaic pattern and appears to predate many skeletal adaptations, suggesting these changes occurred early in the thalattosuchian marine transition.

Ontogenetic variation in the crocodylian vestibular system.

Crocodylians today live in tropical to subtropical environments, occupying mostly shallow waters. Their body size changes drastically during ontogeny, as do their skull dimensions and bite forces, which are associated with changes in prey preferences. Endocranial neurosensory structures have also shown to change ontogenetically, but less is known about the vestibular system of the inner ear. Here we use 30 high-resolution computed tomography (CT) scans and three-dimensional geometric morphometrics to investigate the size and shape changes of crocodylian endosseous labyrinths throughout ontogeny, across four stages (hatchling, juvenile, subadult and adult). We find two major patterns of ontogenetic change. First, the labyrinth increases in size during ontogeny, with negative allometry in relation to skull size. Second, labyrinth shape changes significantly, with hatchlings having shorter semicircular canal radii, with thicker diameters and an overall dorsoventrally shorter labyrinth than those of more mature individuals. We argue that the modification of the labyrinth during crocodylian ontogeny is related to constraints imposed by skull growth, due to fundamental changes in the crocodylian braincase during ontogeny (e.g. verticalisation of the basicranium), rather than changes in locomotion, diet, or other biological functions or behaviours.

Spinosaurus.

Steve Brusatte introduces Spinosaurus, the largest known predatory dinosaur.

Petrosal Anatomy of the Paleocene Eutherian Mammal Deltatherium fundaminis (Cope, 1881).

We describe the tympanic anatomy of the petrosal of Deltatherium fundaminis, an enigmatic Paleocene mammal based on cranial specimens recovered from New Mexico, U.S.A. Although the ear region of Deltatherium has previously been described, there has not been a comprehensive, well-illustrated contribution using current anatomical terminology. The dental and cranial anatomy of Deltatherium is a chimera, with morphological similarities to both 'condylarth' and 'cimolestan' taxa. As such, the phylogenetic relationships of this taxon have remained elusive since its discovery, and it has variably been associated with Arctocyonidae, Pantodonta and Tillodontia. The petrosal of Deltatherium is anteriorly bordered by an open space comprising a contiguous carotid opening and pyriform fenestra. The promontorium features both a small rostral tympanic process and small epitympanic wing but lacks well-marked sulci. A large ventral facing external aperture of the canaliculus cochleae is present and bordered posteriorly by a well-developed caudal tympanic process. The hiatus Fallopii opens on the ventral surface of the petrosal. The tegmen tympani is mediolaterally broad and anteriorly expanded, and its anterior margin is perforated by a foramen for the ramus superior of the stapedial artery. The tympanohyal is small but approximates the caudal tympanic process to nearly enclose the stylomastoid notch. The mastoid is widely exposed on the basicranium and bears an enlarged mastoid process, separate from the paraoccipital process. These new observations provide novel anatomical data corroborating previous hypotheses regarding the plesiomorphic eutherian condition but also reveal subtle differences among Paleocene eutherians that have the potential to help inform the phylogeny of Deltatherium. Supplementary Information: The online version contains supplementary material available at 10.1007/s10914-021-09568-3.

Pterosaurs.

150 million years ago, an animal no bigger than a herring gull soared above shallow lagoons in what is now Bavaria. It had obscenely long fingers on each hand, which anchored thin wings of skin, and long toothy jaws, perfectly suited to snatching squids and fishes from the warm waters. Our animal was probably doing just that before its untimely death. For one reason or another, the creature crashed into the water and sank to its anoxic depths, where it was eventually buried by layers of lime. The animal fossilised with sprawled arms and jaws agape - as if in shock and disbelief.