The Origin of Birds

Technical name for birds is Aves, and

“avian” means “of or concerning birds”.
Birds have many unusual synapomorphies among
modern animals: [ Synapomorphies (shared
derived characters), representing new
specializations evolved in the most recent
common ancestor of the ingroup]

• Feathers
• Warm-blooded (also in mammals)
• Specialized lungs & air-sacs
• Hollow bones
• Toothless beaks
• Large brain
• Cervicals very different from dorsals, allowing
neck to fold into “S”-shape
• Synsacrum (sacrum fused to pelves; pelvic bones
fused together)
• Proximal caudals very mobile
• Pygostyle (distal caudals all fused together)
• Furcula - (the wishbone)
• Forelimb very long, has become wing
• Carpometacarpus (semilunate carpal block fused
to metacarpals; all metacarpals fused together)
• Three fingers, but digits all reduced so no unguals
• Backwards-pointing pubis
• Fibula reduced to proximal splint
• Astragalus & calcaneum fused to tibia
• Hinge-like ankle joint
• Tarsometatarsus (distal tarsals fused to
metatarsals; all metatarsals fused together)
• Main pedal digits II-IV
• Pedal digit I reversed, placed at bottom of
tarsometatarsus
Compare modern birds to their closest relatives, crocodilians
• Difficult to find relatives using only modern
animals (turtles have modified necks and
toothless beaks, but otherwise very
• different; bats fly and are warm-blooded,
but are clearly mammals; etc.)

• With discovery of fossils, other potential

relations: pterosaurs had big brains, “S”-
shaped neck, hinge-like foot, but wings are
VERY different.
• In 1859, Darwin published the Origin; some used
birds as a counter-example against evolution, as
there were apparently known transitional forms
between birds and other vertebrates. In 1860, a
feather (identical to modern birds' feathers) was
found in the Solnhofen Lithographic Limestone of
Bavaria, Germany: a Late Jurassic formation. The
following year, the skeleton of this feather-bearer
was found.

• This creature was named Archaeopteryx

lithographica: ancient wing of the Lithographic
limestone.
Like modern birds, it had:
• Feathered wings and a feathered tail
• Backwards-pointing pubis
• Backwards-pointing pedal digit I, located at the bottom
of the metatarsus

BUT, it was very primitive compared to other birds in

that it had:
• Individual distal caudals instead of a pygostyle
• Individual carpals and metacarpals with digits and
unguals rather than a fused carpometacarpus
• Some evidence of teeth near the damaged skull: maybe
from Archaeopteryx, maybe from something else
• Many saw Archaeopteryx as transitional between
“normal” reptiles and birds. This specimen was
bought by the British Museum (Owen's place of
work). “The London Specimen”

• A second, and much better, specimen was found in

1877: this one was bought by the Humbolt
Museum in Berlin. This specimen showed that the
skull DID have teeth! “The Berlin Specimen”

• (During the 20th Century, five more

Archaeopteryx skeletons have been found: all
from the Solnhofen Limestone).
 Archaeopteryx

the Solnhofen specimen 1960’s

the London
Specimen
1861
the Berlin
Specimen
1877
With the discovery of these primitive bird
specimens, could now better compare birds with
potential ancestors:
• Thomas Huxley (Darwin's protégé) and O. C.
Marsh suggested birds might be dinosaur
descendants (as they shared many features).
Huxley showed that bird ancestors were clearly
archosaurs, since Archaeopteryx had an antorbital
fenestra (among other features).

• During late 19th Century, the dinosaurian origin

of birds was the best option.
• During the early 20th Century, a new hypothesis
was suggested. In 1910, discovery of Euparkeria (a
very primitive archosaur about the size of a cat). Its
discover suggested it might have been “…very near
the ancestor of the Dinosaurs, Pterodacyles (sic),
Birds and Crocodiles”.

• This idea was amplified by artist and natural

historian Gerhard Heilmann. Heilmann examined
lots of Mesozoic archosaurs. He considered
coelurosaurian dinosaurs to have the most shared
features with birds, but thought that they didn't have
furculae or clavicles (and therefore didn't have
anything that could evolve into bird clavicles).
Euparkeria
• He settled instead on Euparkeria, suggesting that
it was sufficiently primitive to be a bird ancestor.
Book The Origin of Birds published in 1927 in
English (earlier Dutch version not as influential).
Became THE main hypothesis by far from the
1920s until the 1970s/1980s.

• (Heilmann also championed a particular theory of

the origin of avian flight: from quadrupedal
ground dweller to quadrupedal tree climber to
semiquadrupedal glider to bipedal flier).
• In 1970, John Ostrom was examining a
supposed pterosaur specimen in the Tyler
Museum in Haarlem, the Netherlands.
• Recognized it was really an Archaeopteryx
(had actually been found in 1855, long
before the better specimens), but also
recognized that the manus was almost
identical in every detail to that of
Deinonychus!
Ostrom documented many features found in
Archaeopteryx also found in various theropods,
most especially coelurosaurs, most especially
Deinonychus:

• Backwards pointing pubis

• Semilunate carpal block
• Three-fingered manus with digit II the longest, and
metacarpal III bowed
• Elongate forelimbs
• Functionally three-toed foot
• Reduced pedal digit I
• Etc.
• Revised the Huxley/Marsh hypothesis,
recognized that birds arose among the
coelurosaurian theropod saurischian
dinosaurs.
• Predicted that if an Archaeopteryx were
found without feather impressions, it would
be considered a coelurosaur: prediction
came true in 1970, when a specimen
(thought since 1955 to be a
Compsognathus) turned out be an
Archaeopteryx.
• Ostrom also revised an earlier idea that
flight arose from fully terrestrial bipeds
rather than going through a tree-dwelling
gliding stage; noted that birds show no sign
of using their hindlimbs in flight, whereas
almost all gliders do.

• In 1980s and 1990s cladistic work helped

establish the relationships of birds WITHIN
the dinosaurs. In mid-to-late 1990s,
discovery of other coelurosaurian dinosaurs
with feathers!!
• Under cladistics, birds are the descendants
of dinosaurs. As such, they are descendants
of the most recent common ancestor of
Megalosaurus and Iguanodon.

Therefore:

• Birds (Aves) are a type of dinosaur!

• In other words, Archaeopteryx and later
birds are

eumaniraptoran maniraptoran
coelurosaurian avetheropod tetanurine
eutheropod theropod saurischian
dinosaurian dinosauriform ornithodiran
archosaurs.
Avialae (the taxon comprised of
Archaeopteryx and more advanced birds)
share the following synapomorphies:

• Flight (probably)
• Number of caudals 25 or fewer
• Pedal digit I reversed and at bottom of
metatarsus
Many features that characterize modern birds
evolved long after Archaeopteryx. These
include:

• Loss of teeth (occurs more than once in bird

history)
• Synsacrum
• Pygostyle
• Carpometacarpus
• Tarsometatarsus
• All these features worked out by skeletal
relationships. New discovery of feathered
coelurosaurs leads to new interpretation of
these “avian” features.
• This means that feathers are NOT an
evolutionary novelty (diagnostic feature)
of birds !
• Two species of dinosaur have recently been found
in northeast China which possess feathers (Qiang
et al. 1998). Protoarchaeopteryx robusta and
Caudipteryx zoui show regiges, rectrices and
plumulaceous feather inpressions. Further, they
are not birds, lacking the more advanced
diagnostic characteristics.

• This groups Protoarchaeopteryx and Caudipteryx

with maniraptoran coelurosaurs rather than birds.
• Coelurosauria seems be characterized by
feathers, and Maniraptora by vaned
feathers.

• So, tyrannosaurids and ornithomimosaurs

were most likely feathered (or at least
descendants of feathered ancestors)!!
Since feathers are found in non-avian, non-
flying dinosaurs, feathers must have
evolved for something other than flight.
Possibilities include:

• Insulation
• Warmth for brooding eggs
• Display
• Combination of these
Flight origins:
Two traditional models are:

“Trees down” (arboreal hypothesis):

ancestors of birds were tree-dwellers, birds
went through gliding stage

“Ground up” (cursorial hypothesis):

no arboreal (tree dwelling) phase, nor
gliding stage
“Trees down” (arboreal hypothesis):
ancestors of birds were tree-dwellers, birds
went through gliding stage
“Ground up” (cursorial hypothesis):
no arboreal (tree dwelling) phase, nor
gliding stage

Given a bipedal cursorial (running) ancestor

of a flying lineage, flight must have
proceeded from the ground into the air,
assuming that the ancestor did not normally
live in trees, or if there were no trees around
A quadrupedal ancestor would have
problems evolving flight from the ground
up; it would have to be able to be bipedal in
order to use its wings at all! A cursorial
bipedal animal would have its arms free to
do with them as it pleased while running,
and its running speed would allow it to
achieve the minimum speed necessary for
liftoff.
 Why did flight evolve?
Several Main Hypotheses:
(1) To help escape from predators
(2) To help catch flying or speedy prey
(3) To help move from place to place (leaping
or gliding)
(4) To free the hindlegs for use as weapons.
(5) To gain access to new food sources or an
unoccupied niche
• Recent study on primitive modern birds
suggests wings evolved first to aid young
(and small adults) in running up sides of
trees, cliffs, etc. Later modifications
allowed true flight. If true, this behavior
was probably present in all small
maniraptorans (theropod dinosaurs), and not
just birds.
During Cretaceous, a great burst of bird
diversity, including:

• Perching feet
• Toothless beaks (convergent in different bird
groups)
• Flightless birds
• Even flightless swimming birds!

Because of the cladistic position of Aves, we now

know that Dinosauria is not extinct! (In fact, it
outnumbers mammal species 2:1).
PROTOAVIS