Michael Pittman

A monodactyl nonavian dinosaur and the complex evolution of the alvarezsauroid hand

Digital reduction is a striking evolutionary phenomenon that is clearly exemplified in theropod dinosaurs by the functionally didactyl manus of tyrannosaurids, the flight-adapted manus of birds (Aves), and the tridactyl but digit II-dominated manus of alvarezsauroids. The enlargement of manual digit II in alvarezsauroids and the concurrent reduction of the lateral digits have been interpreted as adaptations for digging, although no detailed biomechanical analysis of hand function has so far been carried out for this group. In the derived alvarezsauroid clade Parvicursorinae, the lateral digits are so small as to be presumably vestigial. Here we report a new alvarezsauroid, Linhenykus monodactylus gen. et sp. nov., based on a specimen from the Upper Cretaceous Wulansuhai Formation of Inner Mongolia, China. Cladistic analysis identifies Linhenykus as the most basal parvicursorine, and digit II of the manus retains a slender morphology and other primitive features. However, Linhenykus is also highly apomorphic in exhibiting the most extreme reduction of the lateral manual digits seen in any alvarezsauroid. Phalanges are retained only on the most medial digit (digit II), making Linhenykus the only known functionally monodactyl nonavian dinosaur. Other parvicursorines are more primitive in retaining a tridactyl manus but more derived in that digit II is highly robust and shows other apomorphic features in both of its phalanges. The unexpected combination of features seen in the hand of Linhenykus points to a complex mosaic pattern of manual evolution in alvarezsauroids, with loss of the presumably vestigial outer digits being decoupled from change in the form of digit II.

3D Camouflage in an Ornithischian Dinosaur

Summary 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, 4, 5, 6]. Because the optimal countershading varies strongly with light environment [7, 8, 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, 11, 12, 13, 14, 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

Osteology of the Late Cretaceous alvarezsauroid Linhenykus monodactylus from China and comments on alvarezsauroid biogeography

The alvarezsauroid theropod Linhenykus monodactylus from the Upper Cretaceous of Inner Mongolia, China is the first known monodactyl non-avian dinosaur, providing important information on the complex patterns of manual evolution seen in alvarezsauroids. Here we provide a detailed description of the osteology of this taxon. Linhenykus shows a number of features that are transitional between parvicursorine and non-parvicursorine alvarezsauroids, but detailed comparisons also reveal that some characters had a more complex distribution. We also use event-based tree-fitting to perform a quantitative analysis of alvarezsauroid biogeography incorporating several recently discovered taxa. The results suggest that there is no statistical support for previous biogeographic hypotheses that favour pure vicariance or pure dispersal scenarios as explanations for the distributions of alvarezsauroids across South America, North America and Asia. Instead, statistically significant biogeographic reconstructions suggest a dominant role for sympatric (or “within area”) events, combined with a mix of vicariance, dispersal and regional extinction. At present the alvarezsauroid data set is too small to completely resolve the biogeographic history of this group: future studies will need to create larger data sets that encompass additional clades.

Laser-Stimulated Fluorescence in Paleontology

Fluorescence using ultraviolet (UV) light has seen increased use as a tool in paleontology over the last decade. Laser-stimulated fluorescence (LSF) is a next generation technique that is emerging as a way to fluoresce paleontological specimens that remain dark under typical UV. A laser’s ability to concentrate very high flux rates both at the macroscopic and microscopic levels results in specimens fluorescing in ways a standard UV bulb cannot induce. Presented here are five paleontological case histories that illustrate the technique across a broad range of specimens and scales. Novel uses such as back-lighting opaque specimens to reveal detail and detection of specimens completely obscured by matrix are highlighted in these examples. The recent cost reductions in medium-power short wavelength lasers and use of standard photographic filters has now made this technique widely accessible to researchers. This technology has the potential to automate multiple aspects of paleontology, including preparation and sorting of microfossils. This represents a highly cost-effective way to address paleontologys preparatory bottleneck.

Mosaic evolution in an asymmetrically feathered troodontid dinosaur with transitional features

Asymmetrical feathers have been associated with flight capability but are also found in species that do not fly, and their appearance was a major event in feather evolution. Among non-avialan theropods, they are only known in microraptorine dromaeosaurids. Here we report a new troodontid, Jianianhualong tengi gen. et sp. nov., from the Lower Cretaceous Jehol Group of China, that has anatomical features that are transitional between long-armed basal troodontids and derived short-armed ones, shedding new light on troodontid character evolution. It indicates that troodontid feathering is similar to Archaeopteryx in having large arm and leg feathers as well as frond-like tail feathering, confirming that these feathering characteristics were widely present among basal paravians. Most significantly, the taxon has the earliest known asymmetrical troodontid feathers, suggesting that feather asymmetry was ancestral to Paraves. This taxon also displays a mosaic distribution of characters like Sinusonasus, another troodontid with transitional anatomical features.

Shake a Tail Feather: The Evolution of the Theropod Tail into a Stiff Aerodynamic Surface

Theropod dinosaurs show striking morphological and functional tail variation; e.g., a long, robust, basal theropod tail used for counterbalance, or a short, modern avian tail used as an aerodynamic surface. We used a quantitative morphological and functional analysis to reconstruct intervertebral joint stiffness in the tail along the theropod lineage to extant birds. This provides new details of the tail’s morphological transformation, and for the first time quantitatively evaluates its biomechanical consequences. We observe that both dorsoventral and lateral joint stiffness decreased along the non-avian theropod lineage (between nodes Theropoda and Paraves). Our results show how the tail structure of non-avian theropods was mechanically appropriate for holding itself up against gravity and maintaining passive balance. However, as dorsoventral and lateral joint stiffness decreased, the tail may have become more effective for dynamically maintaining balance. This supports our hypothesis of a reduction of dorsoventral and lateral joint stiffness in shorter tails. Along the avian theropod lineage (Avialae to crown group birds), dorsoventral and lateral joint stiffness increased overall, which appears to contradict our null expectation. We infer that this departure in joint stiffness is specific to the tail’s aerodynamic role and the functional constraints imposed by it. Increased dorsoventral and lateral joint stiffness may have facilitated a gradually improved capacity to lift, depress, and swing the tail. The associated morphological changes should have resulted in a tail capable of producing larger muscular forces to utilise larger lift forces in flight. Improved joint mobility in neornithine birds potentially permitted an increase in the range of lift force vector orientations, which might have improved flight proficiency and manoeuvrability. The tail morphology of modern birds with tail fanning capabilities originated in early ornithuromorph birds. Hence, these capabilities should have been present in the early Cretaceous, with incipient tail-fanning capacity in the earliest pygostylian birds.

Basal paravian functional anatomy illuminated by high-detail body outline

Body shape is a fundamental expression of organismal biology, but its quantitative reconstruction in fossil vertebrates is rare. Due to the absence of fossilized soft tissue evidence, the functional consequences of basal paravian body shape and its implications for the origins of avians and flight are not yet fully understood. Here we reconstruct the quantitative body outline of a fossil paravian Anchiornis based on high-definition images of soft tissues revealed by laser-stimulated fluorescence. This body outline confirms patagia-bearing arms, drumstick-shaped legs and a slender tail, features that were probably widespread among paravians. Finely preserved details also reveal similarities in propatagial and footpad form between basal paravians and modern birds, extending their record to the Late Jurassic. The body outline and soft tissue details suggest significant functional decoupling between the legs and tail in at least some basal paravians. The number of seemingly modern propatagial traits hint that feathering was a significant factor in how basal paravians utilized arm, leg and tail function for aerodynamic benefit.

New Material of Zhenyuanopterus (Pterosauria) from the Early Cretaceous Yixian Formation of Western Liaoning

A new partial specimen of Zhenyuanopterus from the Lower Cretaceous Yixian Formation of Beipiao city, Liaoning Province is described. This specimen is slightly larger than half the size of the holotype. Although it is not complete, it provides new information on Zhenyuanopterus both in taxonomy and ontogeny: the total number of the caudal vertebrae is 15; the forelimb is more robust than the hindlimb; the growth rate of the humerus and femur is constant, and their ratio is about 1. Compared with the holotype, the scapula and coracoid grow faster than the humerus and the femur. The scapula and coracoid may slow their growth rate after a certain ontogenetic stage. The pectoral girdle, robust forelimb and weak hindlimb of the new material (XHPM1088) together with data from the holotype of Zhenyuanopterus indicate that this taxon spent less time on land than in the sky or other ecological niches such as forests and cliffs.

Osteology of a New Late Cretaceous Troodontid Specimen from Ukhaa Tolgod, Ömnögovi Aimag, Mongolia

ABSTRACT A new troodontid dinosaur, Almas ukhaa, from the Late Cretaceous deposits of the Djadokhta Formation at Ukhaa Tolgod, Mongolia, is described here. The holotype specimen (IGM 100/1323) comprises an almost complete and articulated cranium and partial articulated postcranial skeleton. This specimen has a small body size and a short snout as in basal paravians, but it exhibits a number of derived troodontid features that differentiate Almas ukhaa from the Early Cretaceous troodontids reported from China and unite this new taxon with other Late Cretaceous troodontids. Relative to other troodontids, Almas ukhaa is autapomorphic in the presence of a posteriorly curved pterygoid flange, absence of a lateral groove on the anterior part of the dentary, presence of a distinct spikelike process on the ischium, and elongate chevrons. The eggshell associated with IGM 100/1323 can be assigned to Prismatoolithidae indet. based on the smooth surface, eggshell thickness, and microstructural characteristics, and also preserves attributes similar to Protoceratopsidovum minimum. A unique relationship between Byronosaurus and the perinate troodontids IGM 100/972 and IGM 100/974 is no longer supported based on the new observations of Almas ukhaa and Gobivenator.

A specimen of Paralycoptera Chang & Chou 1977 (Teleostei: Osteoglossoidei) from Hong Kong (China) with a potential Late Jurassic age that extends the temporal and geographical range of the genus

We describe a Mesozoic fish Paralycoptera sp. (Teleostei: Osteoglossoidei), on the basis of a postcranial skeleton collected from the volcaniclastic mudstones of the Lai Chi Chong Formation of Hong Kong, China. The new finding—representing the city’s first Mesozoic fish—extends the geographical distribution of Paralycoptera from eastern mainland China into Hong Kong, demonstrating a wider distribution than previously appreciated for this genus. A radiometric age for the Lai Chi Chong Formation of 146.6 ± 0.2 Ma implies a temporal range expansion for Paralycoptera of approximately 40 million years back from the Early Cretaceous (∼110 Ma). However, spores found in the Formation suggest an Early Cretaceous age that is consistent with the existing age assignment to Paralycoptera. We argue that the proposed temporal range extension is genuine because it is based on recent high precision radiometric age data, but given the discrepancies with the biostratigraphic ages further investigation is needed to confirm this. This study provides an important step towards revealing Hong Kong’s Mesozoic vertebrate fauna and understanding its relationship to well-studied mainland Chinese ones.