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[dinosaur] Evolution of avian flight + more papers (1 old one)

Ben Creisler

Some updates to earlier posts and some recent non-dino papers:

This is a ref that I sent to the DML back in December 2015. It was apparently blocked, which I missed. It was just brought to my attention that the paper had not appeared on the DML.   I'll try again with the official version that came out in February of this year...

Alexander O. Averianov (2015)
Frontal bones of non-avian theropod dinosaurs from the Upper Cretaceous (Santonian – ? Campanian) Bostobe Formation of the northeastern Aral Sea region, Kazakhstan.
Canadian Journal of Earth Sciences 53(2): 168-175 
doi: 10.1139/cjes-2015-0099

Three theropod taxa have been identified based on isolated frontal bones from the Santonian–?Campanian Bostobe Formation at the Shakh-Shakh locality, in the northeastern Aral Sea region of Kazakhstan. A frontal referable to Ornithomimosauria indet. is distinct in having a slightly pronounced cerebral dome. A frontal of an unidentified therizinosaur resembles the element in Erlikosaurus (Cenomanian—Turonian of Mongolia) in most features, but it differs in having an anterolateral prong along the orbital rim, a less-developed facet for contact with the prefrontal, and a larger depression for the cerebral hemisphere. A frontal identifiable as Troodontidae indet. differs from that of Troodon (Campanian—Maastrichtian of North America) and resembles that of the Mongolian Campanian Gobivenator in having a larger supratemporal fossa and the anterior process of the parietals wedged between the frontals. This specimen is the first reliable record for Troodontidae from the Bostobe Formation.


Also, the official English language version of a paper I posted from the original Russian source (with a rough translation)

A. O. Averianov , M. S. Arkhangelsky & S. M. Merkulov (2016)
An azhdarchid humerus (Pterosauria, Azhdarchidae) from the Upper Cretaceous of Saratov Region.
Paleontological Journal  50(4): 414-417
DOI: 10.1134/S0031030116040031

A proximal humerus fragment referred to as Azhdarchidae indet. from the Rybushka Formation (Upper Cretaceous, Lower Campanian) of the Beloe Ozero locality in Saratov Region is described. The proximal articular surface is not saddle-shaped, has a weakly convex profile in the frontal section. The most posteriorly projecting part of the proximal articular surface is displaced ventrally. A large pneumatic foramen is located on the anterior surface ventral to the base of deltopectoral crest and close to the proximal articular surface. The humeral head is slightly declined from the diaphysis and only slightly overhangs the diaphysis posteriorly. This proximal humerus fragment possibly belongs to Volgadraco bogolubovi Averianov, Arkhangelsky et Pervushov, 2008, described from the Rybushka Formation of the Shirokii Karamysh 2 locality in Saratov Region.

News story:



From the new volume of Philosophical Transactions of the Royal Society B on flight:

Moving in a moving medium: new perspectives on flight



Bret W. Tobalske (2016)
Evolution of avian flight: muscles and constraints on performance
Philosophical Transactions of the Royal Society B: 371 20150383
DOI: 10.1098/rstb.2015.0383

Competing hypotheses about evolutionary origins of flight are the ‘fundamental wing-stroke’ and ‘directed aerial descent’ hypotheses. Support for the fundamental wing-stroke hypothesis is that extant birds use flapping of their wings to climb even before they are able to fly; there are no reported examples of incrementally increasing use of wing movements in gliding transitioning to flapping. An open question is whether locomotor styles must evolve initially for efficiency or if they might instead arrive due to efficacy. The proximal muscles of the avian wing output work and power for flight, and new research is exploring functions of the distal muscles in relation to dynamic changes in wing shape. It will be useful to test the relative contributions of the muscles of the forearm compared with inertial and aerodynamic loading of the wing upon dynamic morphing. Body size has dramatic effects upon flight performance. New research has revealed that mass-specific muscle power declines with increasing body mass among species. This explains the constraints associated with being large. Hummingbirds are the only species that can sustain hovering. Their ability to generate force, work and power appears to be limited by time for activation and deactivation within their wingbeats of high frequency. Most small birds use flap-bounding flight, and this flight style may offer an energetic advantage over continuous flapping during fast flight or during flight into a headwind. The use of flap-bounding during slow flight remains enigmatic. Flap-bounding birds do not appear to be constrained to use their primary flight muscles in a fixed manner. To improve understanding of the functional significance of flap-bounding, the energetic costs and the relative use of alternative styles by a given species in nature merit study.


Free pdf:

Emily L. C. Shepard, Andrew N. Ross & Steven J. Portugal (2016)
Introduction: Moving in a moving medium: new perspectives on flight.
Philosophical Transactions of the Royal Society B: 371 20150382; 
DOI: 10.1098/rstb.2015.0382

One of the defining features of the aerial environment is its variability; air is almost never still. This has profound consequences for flying animals, affecting their flight stability, speed selection, energy expenditure and choice of flight path. All these factors have important implications for the ecology of flying animals, and the ecosystems they interact with, as well as providing bio-inspiration for the development of unmanned aerial vehicles. In this introduction, we touch on the factors that drive the variability in airflows, the scales of variability and the degree to which given airflows may be predictable. We then summarize how papers in this volume advance our understanding of the sensory, biomechanical, physiological and behavioural responses of animals to air flows. Overall, this provides insight into how flying animals can be so successful in this most fickle of environments.


Patrick J. Butler (2016)
The physiological basis of bird flight
Philosophical Transactions of the Royal Society B: 371 20150384
Published 15 August 2016.
DOI: 10.1098/rstb.2015.0384

Flapping flight is energetically more costly than running, although it is less costly to fly a given body mass a given distance per unit time than it is for a similar mass to run the same distance per unit time. This is mainly because birds can fly faster than they can run. Oxygen transfer and transport are enhanced in migrating birds compared with those in non-migrators: at the gas-exchange regions of the lungs the effective area is greater and the diffusion distance smaller. Also, migrating birds have larger hearts and haemoglobin concentrations in the blood, and capillary density in the flight muscles tends to be higher. Species like bar-headed geese migrate at high altitudes, where the availability of oxygen is reduced and the energy cost of flapping flight increased compared with those at sea level. Physiological adaptations to these conditions include haemoglobin with a higher affinity for oxygen than that in lowland birds, a greater effective ventilation of the gas-exchange surface of the lungs and a greater capillary-to-muscle fibre ratio. Migrating birds use fatty acids as their source of energy, so they have to be transported at a sufficient rate to meet the high demand. Since fatty acids are insoluble in water, birds maintain high concentrations of fatty acid–binding proteins to transport fatty acids across the cell membrane and within the cytoplasm. The concentrations of these proteins, together with that of a key enzyme in the β-oxidation of fatty acids, increase before migration.


Also, from another source;

Tarah N. Sullivan, Andreï Pissarenko, Steven A. Herrera, David Kisailus, Vlado A. Lubarda&  Marc A. Meyers (2016)
A lightweight, biological structure with tailored stiffness: The feather vane.
Acta Biomaterialia 41: 27-39

The flying feathers of birds are keratinous appendages designed for maximum performance with a minimum weight penalty. Thus, their design contains ingenious combinations of components that optimize lift, stiffness, aerodynamics, and damage resistance. This design involves two main parts: a central shaft that prescribes stiffness and lateral vanes which allows for the capture of air. Within the feather vane, barbs branch from the shaft and barbules branch from barbs, forming a flat surface which ensures lift. Microhooks at the end of barbules hold barbs tightly together, providing the close-knit, unified structure of the feather vane and enabling a repair of the structure through the reattachment of un-hooked junctions. Both the shaft and barbs are lightweight biological structures constructed of keratin using the common motif of a solid shell and cellular interior. The cellular core increases the resistance to buckling with little added weight. Here we analyze the detailed structure of the feather barb and, for the first time, explain its flexural stiffness in terms of the mechanics of asymmetric foam-filled beams subjected to bending. The results are correlated and validated with finite element modeling. We compare the flexure of single barbs as well as arrays of barbs and find that the interlocking adherence of barbs to one another enables a more robust structure due to minimized barb rotation during deflection. Thus, the flexure behavior of the feather vane can be tailored by the adhesive hooking between barbs, creating a system that mitigates damage. A simplified three-dimensional physical model for this interlocking mechanism is constructed by additive manufacturing. The exceptional architecture of the feather vane will motivate the design of bioinspired structures with tailored and unique properties ranging from adhesives to aerospace materials.

Statement of Significance

Despite its importance to bird flight, literature characterizing the feather vane is extremely limited. The feather vane is composed of barbs that branch from the main shaft (rachis) and barbules that branch from barbs. In this study, the flexural behavior of the feather barb and the role of barbule connections in reinforcing the feather vane are quantitatively investigated for the first time, both experimentally and theoretically. Through the performed experiments, structure-function relationships within the feather vane are uncovered. Additionally, in the proposed model the sophisticated structure of the barbs and the interlocking mechanism of the feather vane are simplified to understand these processes in order to engineer new lightweight structures and adhesives.


Some non-dino papers:

Nicole Klein, Rainer R. Schoch & Günter Schweigert (2016)
A juvenile eurysternid turtle (Testudines: Eurysternidae) from the upper Kimmeridgian (Upper Jurassic) of Nusplingen (SW Germany).
Geobios (advance online publication)

A juvenile turtle from the upper Kimmeridgian (Upper Jurassic) of Nusplingen is identified as an eurysternid turtle. It differs in plastral morphology from a juvenile eurysternid turtle from the latest Kimmerdigian of Kelheim described in the 19th century, which represents a comparably early developmental stage. Both juveniles have primordial ribs not yet transformed into costals and lack all other carapacial elements whereas the plastral elements are well developed. The new specimen from Nusplingen has a more robust plastron type when compared to the very gracile, bow- or arc-shaped plastron type of the formerly described juvenile. Both plastron types are also represented by yet undescribed additional juvenile, medium-sized and/or larger eurysternid specimens. The juvenile specimens thus likely document the presence of two morphologically very similar eurysternid taxa in the Upper Jurassic of southern Germany. Both plastron types are different from those described for Idiochelys and Solnhofia but may resemble plastron morphology of Eurysternum, which is, however, only incompletely known.


Aodhán Ó Gogáin, Howard J. Falcon-Lang, David K. Carpenter, Randall F. Miller, Michael J. Benton, Peir K. Pufahl, Marcello Ruta, Thomas G. Davies, Steven J. Hinds & Matthew R. Stimson  (2016)
Fish and tetrapod communities across a marine to brackish salinity gradient in the Pennsylvanian (early Moscovian) Minto Formation of New Brunswick, Canada, and their palaeoecological and palaeogeographical implications.
Palaeontology (advance online publication)
DOI: 10.1111/pala.12249

Euryhaline adaptations in Pennsylvanian vertebrates allowed them to inhabit the marine to freshwater spectrum. This is illustrated by new assemblages of fish and tetrapods from the early Moscovian Minto Formation of New Brunswick, Canada. Fish include chondrichthyans (xenacanthids and the enigmatic Ageleodus), acanthodians (gyracanthids and acanthodiforms), sarcopterygians (rhizodontids, megalichthyids and dipnoans), and actinopterygians (eurynotiforms). Tetrapods include small- to medium-sized, and largely aquatic, stem tetrapods (colosteids) and anthracosaurs (embolomeres). A key finding is that the parautochthonous fossil assemblages are preserved across a salinity gradient, with diversity (measured by the Simpson Index) declining from open marine environments, through brackish embayments, and reaching a nadir in tidal estuaries. Chondrichthyans dominate the entire salinity spectrum (65% of fossils), a distribution that demonstrates a euryhaline mode of life, and one large predatory chondrichthyan, Orthacanthus, may have practised filial cannibalism in coastal nurseries because its heteropolar coprolites contain juvenile xenacanthid teeth. In contrast, other fish communities were more common in open marine settings while tetrapods were more common in coastal brackish waters. While all these faunas were also likely to have been euryhaline, their osmoregulation was, perhaps, less versatile. The demonstration of widespread euryhalinity among fish and aquatic tetrapods explains why Pennsylvanian faunas generally show a cosmopolitan biogeography because taxa were able to disperse via seaways. It also resolves the paradox of enriched strontium isotopic signatures observed in these faunas because organisms would have been, at times, exposed to continental water bodies as well. Therefore, our new findings contribute to the long-running debate about the ecology of Pennsylvanian fishes and tetrapods.


Stephen E. Grasby, Benoit Beauchamp  and Jochen Knies (2016)
Early Triassic productivity crises delayed recovery from world's worst mass extinction.
Geology (advance online publication)

The recovery of life after the latest Permian extinction was protracted over Early Triassic time. Detailed geochemistry of marine sections along northwest Pangea indicates that upwelling ceased at the extinction event. Nitrogen stable isotope data suggest that this was associated with progressive increase in nutrient stress throughout the Early Triassic, coincident with a significant decrease in organic carbon content despite pervasive anoxic to euxinic conditions. We argue that the Early Triassic hothouse both reduced marine productivity and deepened the nutricline, reducing the overall rate of nutrient delivery to the photic zone, creating an Early Triassic nutrient gap. When oceans finally cooled by Middle Triassic time, renewed nutrient upwelling and onset of organic-rich shale deposition occurred across northwest Pangea, marking the final return of global marine productivity.


for World Lizard Day....

Yannick Bucklitsch, Wolfgang Böhme & André Koch (2016)

Scale Morphology and Micro-Structure of Monitor Lizards (Squamata: Varanidae: Varanus spp.) and their Allies: Implications for Systematics, Ecology, and Conservation.

Zootaxa 4153(1): 1-192




We analysed scale morphology and micro-structure from five different body regions using scanning electron microscopy (SEM) across all nine recognized subgenera of the monitor lizard genus Varanus including 41 different species investigated. As far as we are aware, this qualitative visual technique was applied by us for the first time to most monitor lizard species and probably also to the primary outgroup and sister species Lanthanotus borneensis. A comprehensive list of 20 scalation characters each with up to seven corresponding character states was established and defined for the five body regions sampled. For the phylogenetic approach, parsimony analyses of the resulting morphological data matrix as well as Bremer and bootstrap support calculations were performed with the software TNT. Our results demonstrate that a variety of micro-ornamentations (i.e., ultra- or micro-dermatoglyphics) as seen in various squamate groups is hardly present in monitor lizards. In several species from six out of nine subgenera, however, we found a honeycomb-shaped micro-structure of foveate polygons. Two further samples of Euprepiosaurus Fitzinger, 1843 exhibit each another unique microscopic structure on the scale surface. Notably, the majority of species showing the honeycombed ultra-structure inhabit arid habitats in Australia, Africa and the Middle East. Therefore, it can be inferred that this microscopic scalation feature, which has also been identified in other desert dwelling lizard species, is taxonomically and ecologically correlated with a xeric habitat type in varanids, too. In addition, the systematic affiliation of V. spinulosus, an endemic monitor lizard species from the Solomon Islands with an extraordinary scale shape, is discussed in the light of current hypotheses about its phylogenetic position within the Varanidae. Due to its unique scalation characteristics, in combination with other morphological evidence, a new monotypic subgenus, Solomonsaurus subgen. nov., is erected for this enigmatic monitor lizard species. Furthermore, we propose a taxonomic splitting of the morphologically and ecologically heterogeneous subgenus Euprepiosaurus comprising the Pacific or mangrove and the tree monitor lizards, respectively, again based on the SEM data. Thus, for the members of the highly arboreal V. prasinus species group erection of a new subgenus, Hapturosaurus subgen. nov., is justified based on the autapomorphic scale shape in concert with further morphological, phylogenetic and ecological evidence. In addition, V. reisingeri originally described as a distinct species is considered conspecific with the wide-spread V. prasinus due to joint synapormorphic features in the ventral scale micro-structure. Consequently, V. prasinus is (again) rendered polytypic with the taxon reisingeri being assigned subspecies status here.


In conclusion, the established scalation characters allow discrimination of single species even among closely-related Varanus species, such as the members of the V. indicus species group. Together with a recently published identification key for Southeast Asian monitor lizards based on macroscopic phenotypic characters (Koch et al. 2013), the SEM-pictures of the present study may serve as additional references for the microscopic identification of CITES-relevant monitor lizard skins and products, respectively.