Diving dinosaurs? Caveats on the use of bone compactness and pFDA for inferring lifestyle.

The lifestyle of spinosaurid dinosaurs has been a topic of lively debate ever since the unveiling of important new skeletal parts for Spinosaurus aegyptiacus in 2014 and 2020. Disparate lifestyles for this taxon have been proposed in the literature; some have argued that it was semiaquatic to varying degrees, hunting fish from the margins of water bodies, or perhaps while wading or swimming on the surface; others suggest that it was a fully aquatic underwater pursuit predator. The various proposals are based on equally disparate lines of evidence. A recent study by Fabbri and coworkers sought to resolve this matter by applying the statistical method of phylogenetic flexible discriminant analysis to femur and rib bone diameters and a bone microanatomy metric called global bone compactness. From their statistical analyses of datasets based on a wide range of extant and extinct taxa, they concluded that two spinosaurid dinosaurs (S. aegyptiacus, Baryonyx walkeri) were fully submerged "subaqueous foragers," whereas a third spinosaurid (Suchomimus tenerensis) remained a terrestrial predator. We performed a thorough reexamination of the datasets, analyses, and methodological assumptions on which those conclusions were based, which reveals substantial problems in each of these areas. In the datasets of exemplar taxa, we found unsupported categorization of taxon lifestyle, inconsistent inclusion and exclusion of taxa, and inappropriate choice of taxa and independent variables. We also explored the effects of uncontrolled sources of variation in estimates of bone compactness that arise from biological factors and measurement error. We found that the ability to draw quantitative conclusions is limited when taxa are represented by single data points with potentially large intrinsic variability. The results of our analysis of the statistical method show that it has low accuracy when applied to these datasets and that the data distributions do not meet fundamental assumptions of the method. These findings not only invalidate the conclusions of the particular analysis of Fabbri et al. but also have important implications for future quantitative uses of bone compactness and discriminant analysis in paleontology.

Spinosaurus is not an aquatic dinosaur.

A predominantly fish-eating diet was envisioned for the sail-backed theropod dinosaur Spinosaurus aegyptiacus when its elongate jaws with subconical teeth were unearthed a century ago in Egypt. Recent discovery of the high-spined tail of that skeleton, however, led to a bolder conjecture that S. aegyptiacus was the first fully aquatic dinosaur. The 'aquatic hypothesis' posits that S. aegyptiacus was a slow quadruped on land but a capable pursuit predator in coastal waters, powered by an expanded tail. We test these functional claims with skeletal and flesh models of S. aegyptiacus. We assembled a CT-based skeletal reconstruction based on the fossils, to which we added internal air and muscle to create a posable flesh model. That model shows that on land S. aegyptiacus was bipedal and in deep water was an unstable, slow-surface swimmer (<1 m/s) too buoyant to dive. Living reptiles with similar spine-supported sails over trunk and tail are used for display rather than aquatic propulsion, and nearly all extant secondary swimmers have reduced limbs and fleshy tail flukes. New fossils also show that Spinosaurus ranged far inland. Two stages are clarified in the evolution of Spinosaurus, which is best understood as a semiaquatic bipedal ambush piscivore that frequented the margins of coastal and inland waterways.

Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy "Nanotyrannus" and supports ontogenetic niche partitioning in juvenile Tyrannosaurus.

Despite its iconic status as the king of dinosaurs, Tyrannosaurus rex biology is incompletely understood. Here, we examine femur and tibia bone microstructure from two half-grown T. rex specimens, permitting the assessments of age, growth rate, and maturity necessary for investigating the early life history of this giant theropod. Osteohistology reveals these were immature individuals 13 to 15 years of age, exhibiting growth rates similar to extant birds and mammals, and that annual growth was dependent on resource abundance. Together, our results support the synonomization of "Nanotyrannus" into Tyrannosaurus and fail to support the hypothesized presence of a sympatric tyrannosaurid species of markedly smaller adult body size. Our independent data contribute to mounting evidence for a rapid shift in body size associated with ontogenetic niche partitioning late in T. rex ontogeny and suggest that this species singularly exploited mid- to large-sized theropod niches at the end of the Cretaceous.

Response to formal comment on Myhrvold (2016) submitted by Griebeler and Werner (2017).

Griebeler and Werner offer a formal comment on Myhrvold, 2016 defending the conclusions of Werner and Griebeler, 2014. Although the comment criticizes several aspects of methodology in Myhrvold, 2016, all three papers concur on a key conclusion: the metabolism of extant endotherms and ectotherms cannot be reliably classified using growth-rate allometry, because the growth rates of extant endotherms and ectotherms overlap. A key point of disagreement is that the 2014 paper concluded that despite this general case, one can nevertheless classify dinosaurs as ectotherms from their growth rate allometry. The 2014 conclusion is based on two factors: the assertion (made without any supporting arguments) that the comparison with dinosaurs must be restricted only to extant sauropsids, ignoring other vertebrate groups, and that extant sauropsid endotherm and ectotherm growth rates in a data set studied in the 2014 work do not overlap. The Griebeler and Werner formal comment presents their first arguments in support of the restriction proposition. In this response I show that this restriction is unsupported by established principles of phylogenetic comparison. In addition, I show that the data set studied in their 2014 work does show overlap, and that this is visible in one of its figures. I explain how either point effectively invalidates the conclusion of their 2014 paper. I also address the other methodological criticisms of Myhrvold 2016, and find them unsupported.

Dinosaur Metabolism and the Allometry of Maximum Growth Rate.

The allometry of maximum somatic growth rate has been used in prior studies to classify the metabolic state of both extant vertebrates and dinosaurs. The most recent such studies are reviewed, and their data is reanalyzed. The results of allometric regressions on growth rate are shown to depend on the choice of independent variable; the typical choice used in prior studies introduces a geometric shear transformation that exaggerates the statistical power of the regressions. The maximum growth rates of extant groups are found to have a great deal of overlap, including between groups with endothermic and ectothermic metabolism. Dinosaur growth rates show similar overlap, matching the rates found for mammals, reptiles and fish. The allometric scaling of growth rate with mass is found to have curvature (on a log-log scale) for many groups, contradicting the prevailing view that growth rate allometry follows a simple power law. Reanalysis shows that no correlation between growth rate and basal metabolic rate (BMR) has been demonstrated. These findings drive a conclusion that growth rate allometry studies to date cannot be used to determine dinosaur metabolism as has been previously argued.

Comment on "Evidence for mesothermy in dinosaurs".

Grady et al. (Reports, 13 June 2014, p. 1268) studied dinosaur metabolism by comparison of maximum somatic growth rate allometry with groups of known metabolism. They concluded that dinosaurs exhibited mesothermy, a metabolic rate intermediate between endothermy and ectothermy. Multiple statistical and methodological issues call into question the evidence for dinosaur mesothermy.

Semiaquatic adaptations in a giant predatory dinosaur.

We describe adaptations for a semiaquatic lifestyle in the dinosaur Spinosaurus aegyptiacus. These adaptations include retraction of the fleshy nostrils to a position near the mid-region of the skull and an elongate neck and trunk that shift the center of body mass anterior to the knee joint. Unlike terrestrial theropods, the pelvic girdle is downsized, the hindlimbs are short, and all of the limb bones are solid without an open medullary cavity, for buoyancy control in water. The short, robust femur with hypertrophied flexor attachment and the low, flat-bottomed pedal claws are consistent with aquatic foot-propelled locomotion. Surface striations and bone microstructure suggest that the dorsal "sail" may have been enveloped in skin that functioned primarily for display on land and in water.

Revisiting the estimation of dinosaur growth rates.

Previous growth-rate studies covering 14 dinosaur taxa, as represented by 31 data sets, are critically examined and reanalyzed by using improved statistical techniques. The examination reveals that some previously reported results cannot be replicated by using the methods originally reported; results from new methods are in many cases different, in both the quantitative rates and the qualitative nature of the growth, from results in the prior literature. Asymptotic growth curves, which have been hypothesized to be ubiquitous, are shown to provide best fits for only four of the 14 taxa. Possible reasons for non-asymptotic growth patterns are discussed; they include systematic errors in the age-estimation process and, more likely, a bias toward younger ages among the specimens analyzed. Analysis of the data sets finds that only three taxa include specimens that could be considered skeletally mature (i.e., having attained 90% of maximum body size predicted by asymptotic curve fits), and eleven taxa are quite immature, with the largest specimen having attained less than 62% of predicted asymptotic size. The three taxa that include skeletally mature specimens are included in the four taxa that are best fit by asymptotic curves. The totality of results presented here suggests that previous estimates of both maximum dinosaur growth rates and maximum dinosaur sizes have little statistical support. Suggestions for future research are presented.

Dinosaur census reveals abundant Tyrannosaurus and rare ontogenetic stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA.

BACKGROUND: A dinosaur census recorded during the Hell Creek Project (1999-2009) incorporates multiple lines of evidence from geography, taphohistory, stratigraphy, phylogeny and ontogeny to investigate the relative abundance of large dinosaurs preserved in the Upper Cretaceous Hell Creek Formation of northeastern Montana, USA. Overall, the dinosaur skeletal assemblages in the Hell Creek Formation (excluding lag-influenced records) consist primarily of subadult or small adult size individuals. Small juveniles and large adults are both extremely rare, whereas subadult individuals are relatively common. We propose that mature individuals of at least some dinosaur taxa either lived in a separate geographic locale analogous to younger individuals inhabiting an upland environment where sedimentation rates were relatively less, or these taxa experienced high mortality before reaching terminal size where late stage and often extreme cranial morphology is expressed. METHODOLOGY/PRINCIPAL FINDINGS: Tyrannosaurus skeletons are as abundant as Edmontosaurus, an herbivore, in the upper Hell Creek Formation and nearly twice as common in the lower third of the formation. Smaller, predatory dinosaurs (e.g., Troodon and dromaeosaurids) are primarily represented by teeth found in microvertebrate localities and their skeletons or identifiable lag specimens were conspicuously absent. This relative abundance suggests Tyrannosaurus was not a typical predator and likely benefited from much wider food choice opportunities than exclusively live prey and/or specific taxa. Tyrannosaurus adults may not have competed with Tyrannosaurus juveniles if the potential for selecting carrion increased with size during ontogeny. CONCLUSIONS/SIGNIFICANCE: Triceratops is the most common dinosaur and isolated skulls contribute to a significant portion of this census. Associated specimens of Triceratops consisting of both cranial and postcranial elements remain relatively rare. This rarity may be explained by a historical collecting bias influenced by facies and taphonomic factors. The limited discovery of postcranial elements may also depend on how extensive a fossil quarry is expanded after a skull is collected.