Phill Rasnick

3/27/14
Biol 433
The Emergence of Feathers in Theropods and the Evolution of Avian Flight
Introduction
The evolution of feathers was a major step in the evolution of the avian form. Feathers serve
many crucial roles for avian species including flight, insulation, protection, water repellency, display and
courtship, etc. While the development of feathers and avian flight are crucial components in the evolution
of the avian lineage, there is no conclusive evidence on the function of the most basal feather form and
the origin of avian flight. This paper discusses the likely role of basic feathers, how feathers evolved and
become more complex, as well as discussing the most likely implications for the origin of flight within
the avian lineage.
The Emergence and Evolution of Feathers
Findings of proavian theropod species from the Early Cretaceous Jehol Group of western
Liaoning, China provide much detail in the origin and evolution of feathers. The non-avian theropod
specimens found in this location demonstrate a morphologically diverse range of feathers and help prove
that feather evolution occurred prior to the origin avian species and flight. Feather structures identified
from this region and period include single filaments, compound structures composed of multiple
filaments, plumulaceous feathers, and pennaceous feathers. The feathered coelurosaurians identified also
display an evolutionary trend of increasing feather complexity and specialized feather distribution
overtime as they evolve into avian species (Xu 2006).
The most current model for feather evolution describes the earliest protofeather being composed
of a single tubular filament and them transitioning to the more complex branched feather structures seen
in modern day birds. In this model, stage I in feather development describes the tubular filament
composed of -keratin. While this model for early protofeather form is theoretical, recent discoveries of
members of the early Theropods, Therizinosauridae, show the first known examples of Stage I feathers
(Diamond et al. 2011). From there, there was the emergence of stage II feathers which are characterized
by distal branching of the filament. Stage III in feather development has been identified as the most
critical stage in heather development. It is in this stage that it is proposed that the feather follicle
developed. This stage is also marked by the development of the rachis and planar form of the feather.
Barbules which stiffen the structure feather make up stage IV in feather development while stage V is
characterized by the evolution of feather tracts (Xu 2006).
Hypotheses for the initial emergence of feathers include insulation, water repellency, courtship,
camouflage, defense, parental care and brooding, shielding nests, flight or lift, and competition among
males (Diamond et al. 2011). Recent evidence shows strong support for the first feathers being used for
display and later evolving for other adaptations including flight (Zelenitsky et al. 2012). Flight has been
discredited as the driving force for the emergence of feathers because stage I feathers would not be
capable of providing any means of lift (Diamond et al. 2011). Additionally the presence of feathers in
early flightless nonavian theropods further disproves flight as the evolutionary driving force behind
feather emergence. The most recent evidence points to display and courtship as the initial factor leading
to the appearance of feathers, with flight and other roles developing later. One reason behind this proposal
is the presence of pennibrachrium in some adult ornithomimosaurs, but not juveniles, indicating that
feathers may have arisen as a secondary sexual characteristics (Zelenitsky et al. 2012). Some researchers
also believe that while the first feathers would not have the ability to display complex colors, the clear -
keratin composition of feathers could be capable of scattering light and giving off white coloration that
could occur in various patterns. Researchers also suggest that early theropods had sufficient muscle to
shift position of their feathers enabling them to use the feathers for display purposes (Diamond et al.
2011). Later, as feathers became more complex, their functions expanded and picked up various roles
including producing lift for avian flight.
The Evolution of Avian Flight
Along with the development of feathers, there were many other morphological changes which
had to occur in avian evolution which led to the modern day birds and avian flight. As fossil evidence
concludes, feathers and bird-like wing structure arose in theropod-avian evolution prior to the evolution
of flight and the avian body form. Early fossilFossils of Archaeopteryx indicate that the avian body plan
arose from bipedal predatory theropods who had large hind limbs and tails with reduced feathered
forelimbs. In terms of skeletal morphology, both the pectoral and pelvic girdles were enlarge and
strengthened in modern birds. Forelimbs were also lengthened and lost their ability to grasp. Changes in
their hind limb structure also caused the transitioning from hip based terrestrial movement to knee based.
This transition phase was also accompanied by a reduction in size and an anterior shift in their center of
balance (Heers et al. 2012). Findings of maniraptorean fossils also demonstrate that the evolution of
flight occurred before the origin of birds as several maniraptorans could fold and move their arms in an
avian-like manner (Xu 2006).
There are several theories behind the origin of avian flight which focusses largely on whether
flight evolved in an arboreal or terrestrial environment. The arboreal theory describes that flight evolved
first in tree-dwelling theropods who used their wing structures for gliding rather than flapping. The other
theory describes the bird ancestors as cursorial theropods who developed flight off the ground. Because
extant bids flap and glide when flying, and that they spend time in both terrestrial and arboreal habitats it
is difficult to conclude which mode led the eventual evolution of modern avian flight (Heers et al. 2012,
and Dial et al. 2006).
Soft tissue analysis has demonstrated that early dromaeosaurids were better adapted to an
arboreal lifestyle than a cursorial one. It was proposed that the first dromaeosaurids had four wings with
pennaceous leg feathers also specialized for aerodynamic purposes. These early dromaeosaurs were able
to stretch out their hind limbs placing them relatively parallel to their tail, creating more surface area for
lift. During avian evolution, the hind wings would be reduced and front wings became the main structure
for generating lift. Microraptor is a species identified which represents an early stage with 4 wings and a
large tail, while Archaeopteryx shows more reduced leg feathers while still having a fairly large feathered
tail in comparison to modern day avian species (Xu 2006). Microraptor provides evidence for an arboreal
and gliding origin of flight, however, morphological evidence indicates that the species would not be
capable of adapting flapping flight (Long et al. 2003). Another issue with this hypothesis is that arboreal
gliding, without flapping, is observed in some arboreal mammals but is not observed in any extant birds
(Heers et al. 2012). It is therefore difficult to determine whether gliding was a precursor to flight or
whether this adaptation was a terminal side branch in evolution.
Evidence now shows strong support for a bipedal terrestrial origin of flight. Much of this support
is based off flightless juvenile birds and their progression towards developing flight in their first month of
life. It has been demonstrated that many current developmental transitions in extant species conceptually
parallel evolution and can give insight into the function and development of early structures. Juvenile
birds share many features with skeletons of early pre-avian theropods. These features include transitional
locomotion capacities, transitional skeletons with bird-like wings, the presence of protowings, and the
posterior center of mass due to large legs and tail and a relatively small flight apparatus (Heers et al
2012). Many juvenile, pre-flight, birds use a strategy called wing assisted incline running (WAIR) to seek
refuge from predation. This model demonstrates a possible transitioning form in the development of avian
flight from bipedal theropods. During WAIR, juvenile birds use their protowings to generate downward
force and increase traction enabling them to climb up steep surfaces prior to their fully developed flight
mechanisms. As the birds develop, the surface area of their wings increase and their feathers become
more developed leading to flight (Dial et al. 2006).
Conclusion
While the emergence of feathers was not directly involved with avian flight, their evolution
provided the avian lineage with a means of creating air foils and lift to assist in the development of flight.
Based on current evidence it is likely that the first feathers emerged for display and courtship purposes.
Fossil evidence dating back from the Cretaceous period indicate that feathers and protowings developed
earlier than flight in the avian lineage. Theropod-Avian evolution draws many parallels with juvenile to
adult morphology in extant birds which includes locomotor development, skeletal development, feather
development, and changes in size. For these reasons, the development of extant birds can provide a vast
array of knowledge in the evolution of feathers and avian flight. Based on comparisons between fossil
evidence and juvenile to adult morphology in extant birds, researchers have been able to provide strong
support for the terrestrial origin of flight, however, an arboreal origin still cannot be completely
disproven.

Works Cited
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Heers, Ashley M. and Kenneth P. Dial. "From Extant to Extinct: Locomotor Ontogeny and the Evolution
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Xu, Xing. "Feathered Dinosaurs from China and Evolution of Major Avian Characters." Integrative
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Zelenitsky, Darla K. and et al. "Feathred Non-Avian Dinosaurs from North America Provide Insight into
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