Bite and tooth marks on sauropod dinosaurs from the Morrison Formation.

Tooth-marked bones provide important evidence for feeding choices made by extinct carnivorous animals. In the case of the dinosaurs, most bite traces are attributed to the large and robust osteophagous tyrannosaurs, but those of other large carnivores remain underreported. Here we report on an extensive survey of the literature and some fossil collections cataloging a large number of sauropod bones (68) from the Upper Jurassic Morrison Formation of the USA that bear bite traces that can be attributed to theropods. We find that such bites on large sauropods, although less common than in tyrannosaur-dominated faunas, are known in large numbers from the Morrison Formation, and that none of the observed traces showed evidence of healing. The presence of tooth wear in non-tyrannosaur theropods further shows that they were biting into bone, but it remains difficult to assign individual bite traces to theropod taxa in the presence of multiple credible candidate biters. The widespread occurrence of bite traces without evidence of perimortem bites or healed bite traces, and of theropod tooth wear in Morrison Formation taxa suggests preferential feeding by theropods on juvenile sauropods, and likely scavenging of large-sized sauropod carcasses.

The origin of an invasive air sac system in sauropodomorph dinosaurs.

One of the most remarkable features in sauropod dinosaurs relates to their pneumatized skeletons permeated by a bird-like air sac system. Many studies described the late evolution and diversification of this trait in mid to late Mesozoic forms but few focused on the origin of the invasive respiratory diverticula in sauropodomorphs. Fortunately, it is possible to solve this thanks to the boom of new species described in the last decade as well as the broad accessibility of new technologies. Here we analyze the unaysaurid sauropodomorph Macrocollum itaquii from the Late Triassic (early Norian) of southern Brazil using micro-computed tomography. We describe the chronologically oldest and phylogenetically earliest unambiguous evidence of an invasive air sac system in a dinosaur. Surprisingly, this species presented a unique pattern of pneumatization in non-sauropod sauropodomorphs, with pneumatic foramina in posterior cervical and anterior dorsal vertebrae. This suggests that patterns of pneumatization were not cladistically consistent prior to the arrival of Jurassic eusauropods. Additionally, we describe the protocamerae tissue, a new type of pneumatic tissue with properties of both camellae and camerae. This reverts the previous hypothesis which stated that the skeletal pneumatization first evolved into camarae, and derived into delicate trabecular arrangements. This tissue is evidence of thin camellate-like tissue developing into larger chambers. Finally, Macrocollum is an example of the gradual evolution of skeletal tissues responding to the fastly specializing Respiratory System of saurischian dinosaurs.

A computed tomography-based survey of paramedullary diverticula in extant Aves.

Avian respiratory systems are comprised of rigid lungs connected to a hierarchically organized network of large, regional air sacs, and small diverticula that branch from them. Paramedullary diverticula are those that rest in contact with the spinal cord, and frequently invade the vertebral canal. Here, we review the historical study of these structures and provide the most diverse survey to date of paramedullary diverticula in Aves, consisting of observations from 29 taxa and 17 major clades. These extensions of the respiratory system are present in nearly all birds included in the study, with the exception of falconiforms, gaviiforms, podicipediforms, and piciforms. When present, they share connections most commonly with the intertransverse and supravertebral diverticula, but also sometimes with diverticula arising directly from the lungs and other small, more posterior diverticula. Additionally, we observed much greater morphological diversity of paramedullary airways than previously known. These diverticula may be present as one to four separate tubes (dorsal, lateral, or ventral to the spinal cord), or as a single large structure that partially or wholly encircles the spinal cord. Across taxa, paramedullary diverticula are largest and most frequently present in the cervical region, becoming smaller and increasingly absent moving posteriorly. Finally, we observe two osteological correlates of paramedullary diverticula (pneumatic foramina and pocked texturing inside the vertebral canal) that can be used to infer the presence of these structures in extinct taxa with similar respiratory systems.

The absence of an invasive air sac system in the earliest dinosaurs suggests multiple origins of vertebral pneumaticity.

The origin of the air sac system present in birds has been an enigma for decades. Skeletal pneumaticity related to an air sac system is present in both derived non-avian dinosaurs and pterosaurs. But the question remained open whether this was a shared trait present in the common avemetatarsalian ancestor. We analyzed three taxa from the Late Triassic of South Brazil, which are some of the oldest representatives of this clade (233.23 ± 0.73 Ma), including two sauropodomorphs and one herrerasaurid. All three taxa present shallow lateral fossae in the centra of their presacral vertebrae. Foramina are present in many of the fossae but at diminutive sizes consistent with neurovascular rather than pneumatic origin. Micro-tomography reveals a chaotic architecture of dense apneumatic bone tissue in all three taxa. The early sauropodomorphs showed more complex vascularity, which possibly served as the framework for the future camerate and camellate pneumatic structures of more derived saurischians. Finally, the evidence of the absence of postcranial skeletal pneumaticity in the oldest dinosaurs contradicts the homology hypothesis for an invasive diverticula system and suggests that this trait evolved independently at least 3 times in pterosaurs, theropods, and sauropodomorphs.

The first occurrence of an avian-style respiratory infection in a non-avian dinosaur.

Other than repaired fractures, osteoarthritis, and periosteal reaction, the vertebrate fossil record has limited evidence of non-osseous diseases. This difficulty in paleontological diagnoses stems from (1) the inability to conduct medical testing, (2) soft-tissue pathologic structures are less likely to be preserved, and (3) many osseous lesions are not diagnostically specific. However, here reported for the first time is an avian-style respiratory disorder in a non-avian dinosaur. This sauropod presents irregular bony pathologic structures stemming from the pneumatic features in the cervical vertebrae. As sauropods show well-understood osteological correlates indicating that respiratory tissues were incorporated into the post-cranial skeleton, and thus likely had an 'avian-style' form of respiration, it is most parsimonious to identify these pathologic structures as stemming from a respiratory infection. Although several extant avian infections produce comparable symptoms, the most parsimonious is airsacculitis with associated osteomyelitis. From actinobacterial to fungal in origin, airsacculitis is an extremely prevalent respiratory disorder in birds today. While we cannot pinpoint the specific infectious agent that caused the airsacculitis, this diagnosis establishes the first fossil record of this disease. Additionally, it allows us increased insight into the medical disorders of dinosaurs from a phylogenetic perspective and understanding what maladies plagued the "fearfully great lizards".

Exquisite air sac histological traces in a hyperpneumatized nanoid sauropod dinosaur from South America.

This study reports the occurrence of pneumosteum (osteohistological structure related to an avian-like air sac system) in a nanoid (5.7-m-long) saltasaurid titanosaur from Upper Cretaceous Brazil. We corroborate the hypothesis of the presence of an air sac system in titanosaurians based upon vertebral features identified through external observation and computed tomography. This is the fifth non-avian dinosaur taxon in which histological traces of air sacs have been found. We provided a detailed description of pneumatic structures from external osteology and CT scan data as a parameter for comparison with other taxa. The camellate pattern found in the vertebral centrum (ce) of this taxon and other titanosaurs shows distinct architectures. This might indicate whether cervical or lung diverticula pneumatized different elements. A cotylar internal plate of bone tissue sustains radial camellae (rad) in a condition similar to Alamosaurus and Saltasaurus. Moreover, circumferential chambers (cc) near the cotyle might be an example of convergence between diplodocoids and titanosaurs. Finally, we also register for the first time pneumatic foramina (fo) and fossae connecting camellate structures inside the neural canal in Titanosauria and the second published case in non-avian dinosaurs. The extreme pneumaticity observed in this nanoid titanosaur contrasts with previous assumptions that this feature correlates with the evolution of gigantic sizes in sauropodomorphs. This study reinforces that even small-bodied sauropod clades could present a hyperpneumatized postcranial skeleton, a character inherited from their large-bodied ancestors.

Ontogeny and the fossil record: what, if anything, is an adult dinosaur?

Identification of the ontogenetic status of an extinct organism is complex, and yet this underpins major areas of research, from taxonomy and systematics to ecology and evolution. In the case of the non-avialan dinosaurs, at least some were reproductively mature before they were skeletally mature, and a lack of consensus on how to define an 'adult' animal causes problems for even basic scientific investigations. Here we review the current methods available to determine the age of non-avialan dinosaurs, discuss the definitions of different ontogenetic stages, and summarize the implications of these disparate definitions for dinosaur palaeontology. Most critically, a growing body of evidence suggests that many dinosaurs that would be considered 'adults' in a modern-day field study are considered 'juveniles' or 'subadults' in palaeontological contexts.

Ecological correlates to cranial morphology in Leporids (Mammalia, Lagomorpha).

The mammalian order Lagomorpha has been the subject of many morphometric studies aimed at understanding the relationship between form and function as it relates to locomotion, primarily in postcranial morphology. The leporid cranial skeleton, however, may also reveal information about their ecology, particularly locomotion and vision. Here we investigate the relationship between cranial shape and the degree of facial tilt with locomotion (cursoriality, saltation, and burrowing) within crown leporids. Our results suggest that facial tilt is more pronounced in cursors and saltators compared to generalists, and that increasing facial tilt may be driven by a need for expanded visual fields. Our phylogenetically informed analyses indicate that burrowing behavior, facial tilt, and locomotor behavior do not predict cranial shape. However, we find that variables such as bullae size, size of the splenius capitus fossa, and overall rostral dimensions are important components for understanding the cranial variation in leporids.

A Ceratopsian Dinosaur from the Lower Cretaceous of Western North America, and the Biogeography of Neoceratopsia.

The fossil record for neoceratopsian (horned) dinosaurs in the Lower Cretaceous of North America primarily comprises isolated teeth and postcrania of limited taxonomic resolution, hampering previous efforts to reconstruct the early evolution of this group in North America. An associated cranium and lower jaw from the Cloverly Formation (?middle-late Albian, between 104 and 109 million years old) of southern Montana is designated as the holotype for Aquilops americanus gen. et sp. nov. Aquilops americanus is distinguished by several autapomorphies, including a strongly hooked rostral bone with a midline boss and an elongate and sharply pointed antorbital fossa. The skull in the only known specimen is comparatively small, measuring 84 mm between the tips of the rostral and jugal. The taxon is interpreted as a basal neoceratopsian closely related to Early Cretaceous Asian taxa, such as Liaoceratops and Auroraceratops. Biogeographically, A. americanus probably originated via a dispersal from Asia into North America; the exact route of this dispersal is ambiguous, although a Beringian rather than European route seems more likely in light of the absence of ceratopsians in the Early Cretaceous of Europe. Other amniote clades show similar biogeographic patterns, supporting an intercontinental migratory event between Asia and North America during the late Early Cretaceous. The temporal and geographic distribution of Upper Cretaceous neoceratopsians (leptoceratopsids and ceratopsoids) suggests at least intermittent connections between North America and Asia through the early Late Cretaceous, likely followed by an interval of isolation and finally reconnection during the latest Cretaceous.

Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus.

Skeletal pneumaticity is found in the presacral vertebrae of most sauropod dinosaurs, but pneumaticity is much less common in the vertebrae of the tail. We describe previously unrecognized pneumatic fossae in the mid-caudal vertebrae of specimens of Giraffatitan and Apatosaurus. In both taxa, the most distal pneumatic vertebrae are separated from other pneumatic vertebrae by sequences of three to seven apneumatic vertebrae. Caudal pneumaticity is not prominent in most individuals of either of these taxa, and its unpredictable development means that it may be more widespread than previously recognised within Sauropoda and elsewhere in Saurischia. The erratic patterns of caudal pneumatization in Giraffatitan and Apatosaurus, including the pneumatic hiatuses, show that pneumatic diverticula were more broadly distributed in the bodies of the living animals than are their traces in the skeleton. Together with recently published evidence of cryptic diverticula--those that leave few or no skeletal traces--in basal sauropodomorphs and in pterosaurs, this is further evidence that pneumatic diverticula were widespread in ornithodirans, both across phylogeny and throughout anatomy.

The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs.

The necks of sauropod dinosaurs were a key factor in their evolution. The habitual posture and range of motion of these necks has been controversial, and computer-aided studies have argued for an obligatory sub-horizontal pose. However, such studies are compromised by their failure to take into account the important role of intervertebral cartilage. This cartilage takes very different forms in different animals. Mammals and crocodilians have intervertebral discs, while birds have synovial joints in their necks. The form and thickness of cartilage varies significantly even among closely related taxa. We cannot yet tell whether the neck joints of sauropods more closely resembled those of birds or mammals. Inspection of CT scans showed cartilage:bone ratios of 4.5% for Sauroposeidon and about 20% and 15% for two juvenile Apatosaurus individuals. In extant animals, this ratio varied from 2.59% for the rhea to 24% for a juvenile giraffe. It is not yet possible to disentangle ontogenetic and taxonomic signals, but mammal cartilage is generally three times as thick as that of birds. Our most detailed work, on a turkey, yielded a cartilage:bone ratio of 4.56%. Articular cartilage also added 11% to the length of the turkey's zygapophyseal facets. Simple image manipulation suggests that incorporating 4.56% of neck cartilage into an intervertebral joint of a turkey raises neutral posture by 15°. If this were also true of sauropods, the true neutral pose of the neck would be much higher than has been depicted. An additional 11% of zygapophyseal facet length translates to 11% more range of motion at each joint. More precise quantitative results must await detailed modelling. In summary, including cartilage in our models of sauropod necks shows that they were longer, more elevated and more flexible than previously recognised.

Why sauropods had long necks; and why giraffes have short necks.

The necks of the sauropod dinosaurs reached 15 m in length: six times longer than that of the world record giraffe and five times longer than those of all other terrestrial animals. Several anatomical features enabled this extreme elongation, including: absolutely large body size and quadrupedal stance providing a stable platform for a long neck; a small, light head that did not orally process food; cervical vertebrae that were both numerous and individually elongate; an efficient air-sac-based respiratory system; and distinctive cervical architecture. Relevant features of sauropod cervical vertebrae include: pneumatic chambers that enabled the bone to be positioned in a mechanically efficient way within the envelope; and muscular attachments of varying importance to the neural spines, epipophyses and cervical ribs. Other long-necked tetrapods lacked important features of sauropods, preventing the evolution of longer necks: for example, giraffes have relatively small torsos and large, heavy heads, share the usual mammalian constraint of only seven cervical vertebrae, and lack an air-sac system and pneumatic bones. Among non-sauropods, their saurischian relatives the theropod dinosaurs seem to have been best placed to evolve long necks, and indeed their necks probably surpassed those of giraffes. But 150 million years of evolution did not suffice for them to exceed a relatively modest 2.5 m.

Origin of postcranial skeletal pneumaticity in dinosaurs.

The sauropodomorph Thecodontosaurus caducus and theropod Coelophysis bauri are the earliest known dinosaurs with postcranial skeletal pneumaticity. In both taxa, postcranial pneumatic features are confined to the cervical vertebrae. This distribution of pneumaticity in the skeleton is most consistent with pneumatization by diverticula of cervical air sacs similar to those of birds. Other hypotheses, including pneumatization by diverticula of the lungs, larynx and trachea, or cranial air spaces, are less well-supported.