Development and

Evolutionary
Origin of Feathers
Richard O. Prum
Received 9 June 1999; Accepted 3 September 1999

Reported by:
Kyra Mari Dominique E. Aldaba
Sushmita B. Ramos

Definition of Terms

Feathers

Rachis central shaft of the feather

Ramus main shaft of the barb
Barb primary branches of the rachis
Barbules secondary branches of the feather
Pennulum series of long distal cells

Synapomorphy -An apomorphy (derived or

branched or pinnate epidermal derivative

composed of a matrix of intracellular keratin

specialised character) shared by two or more groups which

originated in their last common ancestor

Apomorphy- A derived or specialised character

Definition of Terms
Pennaceous - contour-feather
Plumulaceous Downy feather
Nonavian theropod dinosaurs - dinosaurs from the

Cretaceous had feathers, nests, laid eggs and roosted like birds

Coelurisaurian dinosaurs

theropod dinosaurs

closely related to birds

Pleisiomorphy - An ancestral or primitive character

Pterylae - An area on the skin of a bird from which

feathers grow

Placode -

platelike structure, especially a thickened

plate of ectoderm in the early embryo, from which a sense
organ develops

Introduction

Evolutionary origin of feathers is controversial

and poorly understood because the lack of
ancestral feather morphologies or structural
antecedents.
First fossil record with feathers, Archeopteryx,
had them in the modern form.
Discoveries of the novel epidermal structures in
the coelurisaurian dinosaurs, Sinosaupteryx and
Beipiaosaurus, and the descirption of fully
pennaceous feathers of identified nonavian
theropod dinosaurs, Protoarcheopteryx and
Caudipteryx had lead to focused on the current
theories of feather origin.

Introduction

Current theories of feather origins lack

consensus about the morphology of the earliest
feather. Usual hypotheses of these theories
focused on structural variation of feather and
their taxonomic distribution among extant birds.
Other theories focused about the adaptive and
functional explanations of the evolution of
feathers from primitive reptilian scale.

FEATHER
MORPHOLOGY

Development of Feather Follicle

Epidermal
placode

Ramogenic
zone

Feather
follicle

Dermal
condensation

Dermal
papilla

Differenti
ation
inside
the
Feather
Follicle

Developmental
model of Feather

Feather Follicle Transition

Stage I
- undifferentiated tubular collar
yielded the first feather

Stage II
- differentiated barb ridges resulted to
unbranched barb and a basal calamus

Stage III
- Involves two development novelties
* Stage IIIa
- evolution of the helical displacement
of barb ridges resulting to the origin
of rachis.
* Stage IIIb
- origin of peripheral barbule plates
Stage IV
- origin of the proximal and distal
barbules

Stage V
* Stage Va
- lateral displacement of the new
barb
* Stage Vb
- division and lateral
displacement of the new barb
loci

DISCUSSION

The cylindrical organization of the

follicle is the defining developmental
and morphological characteristic of
feathers
Follicles and feathers are diversified
through a series of derived novelties in
the developmental mechanisms within
the follicle.
These subsequent novelty mechanisms
gave rise to the rest of feather
diversity.

Discussion

The model polarizes most of the

developmental novelties required to
evolve the entire structural diversity of
feathers.
Open pennaceous feather originated
before closed pennaceous feather.
Afterfeather evolved from main feather.
Exceptions of the highly derived
contour feathers are the flightless
Australo-Papauan ratites, Dromaius
and Casuaria.

Discussion

A few novelties are not completely

polarized by the model.
Morphology found in the model exists
among the feathers of extant birds and
are known products of avian feather
follicles.
Additional intermediate could have
occurred between some of the stages
of the proposed model.

Discussion

Polarities of the developmental

novelties in the model are congruent
with von Baers rule the hypothesis
that stages that occur earlier in
development are phylogenetically
more broadly distributed and
historically plesiomorphic.

Discussion

The structure of the invaginated follicle

creates a unique cylindrical sandwich
of epidermal and dermal tissue layers
that permits (a) continuous interaction
of epidermal and dermal; (b)
indeterminate growth of the epidermis;
and (c) continuous nourishment of the
epidermis by the dermis without
continued growth in the volume of the
dermal pulp.

Discussion

Brushs phylogram of feather evolution

is congruent with the proposed model.
Many additional topics in feather
evolution remain to be studied.

Functional Hypotheses of
feather evolution

Flight
- feathers have been hypothesized to have evolved through
natural selection on primitive scales for an aerodynamic function
- But this aerodynamic hypothesis is not plausible due to the ff.
reasons:
-presence of the follicle structure is the only similarity bet. scales
and follicle.
-bipinnate structure of feathers could not have evolved from a
closed pennaceous feather.
- selection for an aerodynamic function could only have taken
place after the evolution of the closed pennaceous, bipinnate
feather
- aerodynamis function probably only gave rise to the
asymmetrical vane and robust rachis found in the rectrices and
remiges of most flighted bird

Thermal Insulation
- first filamentous, nonpinnate feathers could
certainly have been plumulaceous and
provided thermal insulation

Heat Shielding
- feathers evolved from elongate, crudely
pennaceous scales as a shield from excessive
solar radiation.
- but the protofeathers proposed appears to
be intermediate bet. scales and feathers.
Thus, impossible to grow in any plausible
developmental mechanism.

Water Repellancy

- it is plausible that the first filamentous, nonpinnate

feathers have functioned for water repellency
Communication and crypsis(camouflaged) - the
simplest possible control could have yielded feathers
with horizontal stripes or longitudinal gradients in
color.

Defense
- modern feathers do not provide significant physical
defense to birds
- pointed keratinaceous structures on the integument
could have providedprotection to the body

Integumental structures of
theropod dinosaurs

The integumental structure found in the

Sinosauropteryx and Beipiaosaurus could be
homologous to avian feathers but there were
many questions about its morphology.
The model provides functionally neutral criteria
to evaluate the homology between avian
feathers and other fossil integumental structures.
Additional examination is required to establish
more information about Sinosauropteryx and
Beipiaosaurus.

Sinosauropteryx and
Beipiaosaurus

Figure 8. The elongated, single filament feathers

ofBeipiaosaurus. The yellow arrows point to feathers on
the head and neck (right) and tail (above)
(From: Xu et al. 2009).
Figure 7. Restoration of a Sinosauropteryx
(Sinosauropteryx prima) with its body covered
with feathers that were likely important for
thermoregulation
(From: Chuong et al. 2001, and based on Chen
et al. 1998).

Integumental structures of theropod

dinosaurs

The pennaceous integumental structures

described from the forelimbs and tails of
Protarcheopteryx and Caudipteryx are
homologous with avian feathers but do
not function directly for flight.
If additional phylogenetic analyses
confirms that Protarcheopteryx and
Caudipteryx are primarily flightless, then
it will confirm that closed pennaceous
feathers preceeded the asymmetrical
flight feathers.

Molecular mechanisms of feather

development

The research has confirmed the general paradigm in

molecular developmental biology that a common,
plesiomorphic set of genes plays an important role in
pattern specification and morphogenesis of structures in a
diversity of metazoans. Thus, the broadly distributed Hox
genes, Wnt-7a, Sonic hedgehog, N-CAM, L-CAM, BMP2, and
TGF are now known to be involved in specifying the
pattern of feathers placodes within ptyerylae and
morphogenesis within the feather placodes and follicles.
As more is learned about the molecular basis of feather
development, it will be important to establish a similar
distinction of between the plesiomorphic mechanisms that
are shared with other epidermal appendages and the
derived developmental novelties that are unique to
feathers.