Denver W. Fowler

Predatory Functional Morphology in Raptors: Interdigital Variation in Talon Size Is Related to Prey Restraint and Immobilisation Technique

Despite the ubiquity of raptors in terrestrial ecosystems, many aspects of their predatory behaviour remain poorly understood. Surprisingly little is known about the morphology of raptor talons and how they are employed during feeding behaviour. Talon size variation among digits can be used to distinguish families of raptors and is related to different techniques of prey restraint and immobilisation. The hypertrophied talons on digits (D) I and II in Accipitridae have evolved primarily to restrain large struggling prey while they are immobilised by dismemberment. Falconidae have only modest talons on each digit and only slightly enlarged D-I and II. For immobilisation, Falconini rely more strongly on strike impact and breaking the necks of their prey, having evolved a ‘tooth’ on the beak to aid in doing so. Pandionidae have enlarged, highly recurved talons on each digit, an adaptation for piscivory, convergently seen to a lesser extent in fishing eagles. Strigiformes bear enlarged talons with comparatively low curvature on each digit, part of a suite of adaptations to increase constriction efficiency by maximising grip strength, indicative of specialisation on small prey. Restraint and immobilisation strategy change as prey increase in size. Small prey are restrained by containment within the foot and immobilised by constriction and beak attacks. Large prey are restrained by pinning under the bodyweight of the raptor, maintaining grip with the talons, and immobilised by dismemberment (Accipitridae), or severing the spinal cord (Falconini). Within all raptors, physical attributes of the feet trade off against each other to attain great strength, but it is the variable means by which this is achieved that distinguishes them ecologically. Our findings show that interdigital talon morphology varies consistently among raptor families, and that this is directly correlative with variation in their typical prey capture and restraint strategy.

Reanalysis of “Raptorex kriegsteini”: A Juvenile Tyrannosaurid Dinosaur from Mongolia

The carnivorous Tyrannosauridae are among the most iconic dinosaurs: typified by large body size, tiny forelimbs, and massive robust skulls with laterally thickened teeth. The recently described small-bodied tyrannosaurid Raptorex kreigsteini is exceptional as its discovery proposes that many of the distinctive anatomical traits of derived tyrannosaurids were acquired in the Early Cretaceous, before the evolution of large body size. This inference depends on two core interpretations: that the holotype (LH PV18) derives from the Lower Cretaceous of China, and that despite its small size, it is a subadult or young adult. Here we show that the published data is equivocal regarding stratigraphic position and that ontogenetic reanalysis shows there is no reason to conclude that LH PV18 has reached this level of maturity. The probable juvenile status of LH PV18 makes its use as a holotype unreliable, since diagnostic features of Raptorex may be symptomatic of its immature status, rather than its actual phylogenetic position. These findings are consistent with the original sale description of LH PV18 as a juvenile Tarbosaurus from the Upper Cretaceous of Mongolia. Consequently, we suggest that there is currently no evidence to support the conclusion that tyrannosaurid skeletal design first evolved in the Early Cretaceous at small body size.

The Predatory Ecology of Deinonychus and the Origin of Flapping in Birds

Most non-avian theropod dinosaurs are characterized by fearsome serrated teeth and sharp recurved claws. Interpretation of theropod predatory ecology is typically based on functional morphological analysis of these and other physical features. The notorious hypertrophied ‘killing claw’ on pedal digit (D) II of the maniraptoran theropod Deinonychus (Paraves: Dromaeosauridae) is hypothesized to have been a predatory adaptation for slashing or climbing, leading to the suggestion that Deinonychus and other dromaeosaurids were cursorial predators specialized for actively attacking and killing prey several times larger than themselves. However, this hypothesis is problematic as extant animals that possess similarly hypertrophied claws do not use them to slash or climb up prey. Here we offer an alternative interpretation: that the hypertrophied D-II claw of dromaeosaurids was functionally analogous to the enlarged talon also found on D-II of extant Accipitridae (hawks and eagles; one family of the birds commonly known as “raptors”). Here, the talon is used to maintain grip on prey of subequal body size to the predator, while the victim is pinned down by the body weight of the raptor and dismembered by the beak. The foot of Deinonychus exhibits morphology consistent with a grasping function, supportive of the prey immobilisation behavior model. Opposite morphological trends within Deinonychosauria (Dromaeosauridae + Troodontidae) are indicative of ecological separation. Placed in context of avian evolution, the grasping foot of Deinonychus and other terrestrial predatory paravians is hypothesized to have been an exaptation for the grasping foot of arboreal perching birds. Here we also describe “stability flapping”, a novel behaviour executed for positioning and stability during the initial stages of prey immobilisation, which may have been pivotal to the evolution of the flapping stroke. These findings overhaul our perception of predatory dinosaurs and highlight the role of exaptation in the evolution of novel structures and behaviours.

Evolutionary trends in Triceratops from the Hell Creek Formation, Montana

Significance The deciphering of evolutionary trends in nonavian dinosaurs can be impeded by a combination of small sample sizes, low stratigraphic resolution, and lack of ontogenetic (developmental) details for many taxa. Analysis of a large sample (n > 50) of the famous horned dinosaur Triceratops from the Hell Creek Formation of Montana incorporates new stratigraphic and ontogenetic findings to permit the investigation of evolution within this genus. Our research indicates that the two currently recognized species of Triceratops (T. horridus and T. prorsus) are stratigraphically separated and that the evolution of this genus likely incorporated anagenetic (transformational) change. These findings impact interpretations of dinosaur diversity at the end of the Cretaceous and illuminate potential modes of evolution in the Dinosauria. The placement of over 50 skulls of the well-known horned dinosaur Triceratops within a stratigraphic framework for the Upper Cretaceous Hell Creek Formation (HCF) of Montana reveals the evolutionary transformation of this genus. Specimens referable to the two recognized morphospecies of Triceratops, T. horridus and T. prorsus, are stratigraphically separated within the HCF with the T. prorsus morphology recovered in the upper third of the formation and T. horridus found lower in the formation. Hypotheses that these morphospecies represent sexual or ontogenetic variation within a single species are thus untenable. Stratigraphic placement of specimens appears to reveal ancestor–descendant relationships. Transitional morphologies are found in the middle unit of the formation, a finding that is consistent with the evolution of Triceratops being characterized by anagenesis, the transformation of a lineage over time. Variation among specimens from this critical stratigraphic zone may indicate a branching event in the Triceratops lineage. Purely cladogenetic interpretations of the HCF dataset imply greater diversity within the formation. These findings underscore the critical role of stratigraphic data in deciphering evolutionary patterns in the Dinosauria.

Ontogenetic influence on neural spine bifurcation in Diplodocoidea (Dinosauria: Sauropoda): a critical phylogenetic character.

Within Diplodocoidea (Dinosauria: Sauropoda), phylogenetic position of the three subclades Rebbachisauridae, Dicraeosauridae, and Diplodocidae is strongly influenced by a relatively small number of characters. Neural spine bifurcation, especially within the cervical vertebrae, is considered to be a derived character, with taxa that lack this feature regarded as relatively basal. Our analysis of dorsal and cervical vertebrae from small‐sized diplodocoids (representing at least 18 individuals) reveals that neural spine bifurcation is less well developed or absent in smaller specimens. New preparation of the roughly 200‐cm long diplodocid juvenile Sauriermuseum Aathal 0009 reveals simple nonbifurcated cervical neural spines, strongly reminiscent of more basal sauropods such as Omeisaurus. An identical pattern of ontogenetically linked bifurcation has also been observed in several specimens of the basal macronarian Camarasaurus, suggesting that this is characteristic of several clades of Sauropoda. We suggest that neural spine bifurcation performs a biomechanical function related to horizontal positioning of the neck that may become significant only at the onset of a larger body size, hence, its apparent absence or weaker development in smaller specimens. These results have significant implications for the taxonomy and phylogenetic position of taxa described from specimens of small body size. On the basis of shallow bifurcation of its cervical and dorsal neural spines, the small diplodocid Suuwassea is more parsimoniously interpreted as an immature specimen of an already recognized diplodocid taxon. Our findings emphasize the view that nonmature dinosaurs often exhibit morphologies more similar to their ancestral state and may therefore occupy a more basal position in phylogenetic analyses than would mature specimens of the same species. In light of this, we stress the need for phylogenetic reanalysis of sauropod clades where vital characters may be ontogenetically variable, particularly when data is derived from small individuals. J. Morphol., 2012.

The First Giant Titanosaurian Sauropod from the Upper Cretaceous of North America

Argentinosaurus (Cenomanian, Argentina) is generally accepted as being the largest dinosaur so far discovered and is one of several giant titanosaurian sauropods known from the Upper Cretaceous of South America and Asia, but surprisingly not from North America. Here we present the first evidence of giant titanosaurian sauropods from the Upper Cretaceous of North America: two enormous vertebrae and a partial femur, from the Naashoibito Member of the Ojo Alamo Formation, New Mexico, and referred to Alamosaurus sanjuanensis. One of the new vertebrae, a posterior cervical, is comparable in size to a posterior cervical described for Puertasaurus: an Argentinosaurus-sized titanosaurian from the Maastrichtian of Argentina. This makes A. sanjuanensis the largest dinosaur from North America, and among the largest in the world. These findings indicate that A. sanjuanensis is diagnosed based on immature remains, which may have implications for cladistic analyses.

Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America

Interbasinal stratigraphic correlation provides the foundation for all consequent continental-scale geological and paleontological analyses. Correlation requires synthesis of lithostratigraphic, biostratigraphic and geochronologic data, and must be periodically updated to accord with advances in dating techniques, changing standards for radiometric dates, new stratigraphic concepts, hypotheses, fossil specimens, and field data. Outdated or incorrect correlation exposes geological and paleontological analyses to potential error. The current work presents a high-resolution stratigraphic chart for terrestrial Late Cretaceous units of North America, combining published chronostratigraphic, lithostratigraphic, and biostratigraphic data. 40Ar / 39Ar radiometric dates are newly recalibrated to both current standard and decay constant pairings. Revisions to the stratigraphic placement of most units are slight, but important changes are made to the proposed correlations of the Aguja and Javelina formations, Texas, and recalibration corrections in particular affect the relative age positions of the Belly River Group, Alberta; Judith River Formation, Montana; Kaiparowits Formation, Utah; and Fruitland and Kirtland formations, New Mexico. The stratigraphic ranges of selected clades of dinosaur species are plotted on the chronostratigraphic framework, with some clades comprising short-duration species that do not overlap stratigraphically with preceding or succeeding forms. This is the expected pattern that is produced by an anagenetic mode of evolution, suggesting that true branching (speciation) events were rare and may have geographic significance. The recent hypothesis of intracontinental latitudinal provinciality of dinosaurs is shown to be affected by previous stratigraphic miscorrelation. Rapid stepwise acquisition of display characters in many dinosaur clades, in particular chasmosaurine ceratopsids, suggests that they may be useful for high resolution biostratigraphy.

Direct U-Pb dating of Cretaceous and Paleocene dinosaur bones, San Juan Basin, New Mexico: COMMENT

Based on U-Pb dating of two dinosaur bones from the San Juan Basin of New Mexico (United States), [Fassett et al. (2011)][1] claim to provide the first successful direct dating of fossil bones and to establish the presence of Paleocene dinosaurs. Fassett et al. ignore previously published work that

A new multi-faceted framework for deciphering Diplodocid ontogeny

Determining maturity in sauropod dinosaurs histologically is problematic as rapid growth leads to remodeling of Lines of Arrested Growth (LAGs). Although a complimentary system has been devised utilizing several factors including relative amounts of remodeling (Histologic Ontogenetic Stage [HOS]), most assessments of sauropod maturity are based on morphologic indicators. To better assess skeletal maturity and morphologic change through ontogeny, we examined cranial and postcranial material from over 20 diplodocid individuals (Apatosaurus and Diplodocus) from the Upper Jurassic Morrison Formation. Here we describe consistent combinations of morphologic and histologic features that can be used to ascertain maturity. Small diplodocids (femoral lengths ≤120 cm) display nonto weakly bifurcated cervical and dorsal neural spines, acamerate to camerate centra, two to six preserved LAGs in dorsal ribs, and a maximum femoral designation of HOS 7. Larger individuals (femoral length ~125 cm) have more developed internal pneumatic structures, greater neural spine bifurcation, preserve up to eight LAGs, and a femoral designation of HOS 9. In contrast, skeletally mature sauropods (femoral lengths >150 cm) have complex pneumatic structures, extended neural spine bifurcation (also within anterior caudals), and a femoral HOS between 11-13. Further, all of the preserved small diplodocid skulls exhibit a postparietal foramen (previously suggested to be an apomorphy of Dicraeosauridae), which is absent in large skulls (where preserved), suggesting that it is an ontogenetic character. These findings support the hypothesis of significant ontogenetic morphological change in diplodocid sauropods and suggest caution when describing new taxa on the basis of small-bodied holotypes. D. Cary Woodruff. Royal Ontario Museum; University of Toronto, Toronto, ON, Canada; and Great Plains Dinosaur Museum and Field Station, Malta, MT, United States of America. sauropod4@gmail.com Denver W. Fowler. Dickinson Museum Center, Dickinson, ND, United States of America. df9465@hotmail.com John R. Horner. Burke Museum of Natural History and Culture, University of Washington, Seattle, WA, United States of America. johnrhorner@mac.com Woodruff, D. Cary, Fowler, Denver W., and Horner, John R. 2017. A new multi-faceted framework for deciphering diplodocid ontogeny. Palaeontologia Electronica 20.3.43A: 1-53 palaeo-electronica.org/content/2017/1918-deciphering-diplodocid-growth Copyright: September 2017 Palaeontology Association. This is an open access article distributed under the terms of AttributionNonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0), which permits users to copy and redistribute the material in any medium or format, provided it is not used for commercial purposes and the original author and source are credited, with indications if any changes are made. creativecommons.org/licenses/by-nc-sa/4.0/ WOODRUFF, FOWLER, & HORNER: DECIPHERING DIPLODOCID GROWTH