Primitive birds flying

Primitive birds (basalmost paraves) flapped their wings and flew using the same set of muscles as their pterosaur ancestors.

http://press.princeton.edu/witton/sa...pterosaurs.pdf At one stage, it was thought that the flight muscles of pterosaurs were very [modern] birdlike, with the arm lifted by a muscle, m. supracoracoideus, anchoring on the sternum rather than the shoulders. In birds, this muscle arcs over the glenoid to attach on the dorsal surface of the humerus, elevating the wing with a pulley-like system (e.g., Kripp 1943; Padian 1983a; Wellnhofer 1991a). Detailed reconstruction of the proximal arm musculature of pterosaurs shows that this is not the case, however, and that the [pterosaur] arm was more likely lifted by large muscles anchored on the scapula and back, and lowered by those attached to the sternum and coracoid (fig. 5.8; Bennett 2003a). Unlike [modern] birds, where two vastly expanded muscles are mainly used to power flight, it appears that pterosaurs used several muscle groups to form their flapping strokes.

https://repository.si.edu/bitstream/...=1&isAllowed=y

Furthermore, the supracoracoideus muscle, and hence an ossified sternum, is not necessary to effect the recovery stroke of the wing. Thus the main evidence for Archaeopteryx having been a terrestrial, cursorial predator is invalidated. There is nothing in the structure of the pectoral girdle of Archaeopteryx that would preclude its having been a powered flier.

Concerning the lack of asymmetric feathers in flying basalmost paraves, the following seems relevant: https://gwawinapterus.wordpress.com/...-introduction/ The basal deinonychosaur Anchiornis [a primitive bird] might offer a possible explanation. It had symmetrical feathers, but they were arranged in an unique away; in species with asymmetrical feathers, the most distally attached wing feathers are the longest ones. In Anchiornis, the longest are anchored near the wrist, making the center of the wing the broadest area. This is not an unusual profile among flightless maniraptors – oviraptors like Caudipteryx have this sort of arrangement as well. Anchiornis, however, differs in that the feathers at the front (as in, anchored more distally) of the longest feather decrease rapidly in size as they are closer to the end of the supporting digit; this results in a rounded, yet slightly pointy wing shape. It is possible that this arrangement could had been an early adaptation to the demands of powered flight, before true asymmetrical feathers evolved. If so, it is possible that Anchiornis did engage in powered flight, or even a method of escape akin to rudimentary WAIR. So far, no tests have been conducted to examine the aerodynamic capacities of it’s wings. In Anchiornis, the entire wing forms the airfoil. http://www.sciencedirect.com/science/article/pii/S0960982212011943 Primitive bird: In modern birds (Neornithes), the wing is composed of a layer of long, asymmetrical flight feathers overlain by short covert feathers [1-3]. It has generally been assumed that wing feathers in the Jurassic bird Archaeopteryx [4-9] and Cretaceous feathered dinosaurs [10, 11] had the same arrangement. Here, we redescribe the wings of the archaic bird Archaeopteryx lithographica [3-5] and the dinosaur Anchiornis huxleyi [12, 13] and show that their wings differ from those of Neornithes in being composed of multiple layers of feathers. In Archaeopteryx, primaries are overlapped by long dorsal and ventral coverts. Anchiornis has a similar configuration but is more primitive in having short, slender, symmetrical remiges. Archaeopteryx and Anchiornis therefore appear to represent early experiments in the evolution of the wing. This primitive configuration has important functional implications: although the slender feather shafts of Archaeopteryx [14] and Anchiornis [12] make individual feathers weak, layering of the wing feathers may have produced a strong airfoil. Furthermore, the layered arrangement may have prevented the feathers from forming a slotted tip or separating to reduce drag on the upstroke. The wings of early birds therefore may have lacked the range of functions seen in Neornithes, limiting their flight ability. Longrich NR, Vinther J, Meng Q, Li Q, Russell AP. https://en.wikipedia.org/wiki/Bird_flight Elliptical wings are short and rounded, having a low aspect ratio, allowing for tight maneuvering in confined spaces such as might be found in dense vegetation. As such they are common in forest raptors (such as Accipiter hawks), and many passerines, particularly non-migratory ones (migratory species have longer wings). They are also common in species that use a rapid take off to evade predators, such as pheasants and partridges.

Altogether we have a picture of a flying, feathered, 4 winged, arboreal, primitive bird with a long bony tail, that flew like a pterosaur. With elliptical wings and symmetric feathers

At times people have argued that a flying pterosaur would not devolve into a gliding primitive bird. But this is misguided, because the evidence indicates that flying pterosaurs evolved into flying primitive birds, not into gliding primitive birds.

Here are the aspects related to flight capability:

• muscles used 
• keeled or not keeled sternum 
• flight feathers (asymmetric or not) 
• feathered hindlimbs

See here for more details:
http://pterosaurnet.blogspot.ca/2010/05/keeled-breastbone.html
http://pterosaurnet.blogspot.ca/2015/07/primitive-birds-flying.html
http://pterosaurnet.blogspot.ca/2014/09/flight-without-supracoracoideus.html
http://pterosaurnet.blogspot.ca/2013/02/flight-stroke.html


http://www.nature.com/articles/ncomms14576 (2017)

A stiffer feathered postpatagium in Anchiornis may have compensated for its aerodynamically inferior arm feathers to some degree