Part III The Physics of Flight - lift and propulsion.

Lift
• Flight - as a central adaptation.
- strongest selective pressure on birds.
- most prominent theme in ornithology.
- ties into their anatomy and physiology.
- birds can hover, fly upside down and Thrust Drag
backwards, and soar for days.
- human made aircraft way inferior.
- bird flight requires constant Albatross Cam
adjustments of the wing and tail.
- help from the sensory system.
Gravity
- short term energy costs - high.
- yet, most economical locomotion. - aspect ratio: wing length divided by wing width.
- costs less energy to fly 1 km than to
- wing load: body weight divided by wing area.
walk, run, or swim 1 km.

Lift and The Airfoil. 1) Bernoulis’ Principle. - lift increases with:

- airspeed and volume of air
Less Pressure deflected.
- ex. Northern Gannet with
outstretched wings facing
high winds on edge of a cliff.
Faster - lift effortlessly above edge.
- at equilibrium, will freeze Northern Gannet
A B position, motionless in the air.
- facing still air (no wind), will jump off cliff wings outstretched.
- airspeed increases.
Slower - lift and then flight.
- ex. heavy bodied birds like
camber = curvature more pressure Grebes and Loons must
of the wing. below creates Lift. “patter” across the water to
More Pressure generate lift.

- more lift. 2) Newton’s Third Law of Motion: “For every

Turkey Vulture - slots - also allow better
action there is an equal and opposite reaction.” control of air moving over
the upper surface.
- air forced from under the
- airfoil with Reaction = Lift. wing through a slot
increased expands on the upper side
angle of → reducing pressure.
attack. - creating and controlling lift.
- ex. Turkey Vulture - adjust primary
slots to control speed, lift, and
Action = Deflected Air. - Drag increases position as it searches for carrion.
with increased - alula - creates a slot at the leading
angle of attack, edge, keeps airfoil bound to the wing.
especially at low
- flips up when landing and
speeds.
taking off, with ↑ angle of attack.
- birds compensate with - commercial airplanes raise
slotting and alula. slats to create the same action.

1

• Flight - How Birds Fly
- wing level: like two half propellers
with flapping flight.
a) inner wing generates mostly lift, but
receives transferred thrust from outer.
b) outer wing generates both thrust
and lift by all 1), 2) and 3).
- control: position wings to initiate
turn and alter speed.
Willet
Forster’s Tern
Laysan Albatross b) slope soaring - ride air that is
deflected upward by a ridge or
ocean wave.
- ex. Hawks use mountain ridges,
Gulls air rising behind boats.
- mastered by long-winged
seabirds, air rising off waves.
- dynamic soaring - long narrow wings
decrease drag with a high aspect ratio.
- rise off crest of wave, sink to next wave.
- downward with fast moving upper air,
then into slow moving lower air, lose
speed and lift → rise back up and repeat.
- Albatrosses can do this for days, without flapping, non-stop.
- some believe they can dynamically soar while they sleep, like
dolphins that swim in circles while they sleep.
Propulsion Action - deflected air → action, reaction = Thrust.
3) Newton’s Second Law of
Motion: “Thrust force equals
mass (amount of air moved) Reaction
times velocity (how rapid the
wingbeat).”
- ex. canoe paddle
in the water.
- mass of water.
- times stroke of Action
the paddle. Snowy Egret
Blue Jay
- feather level: collective use
of individual primary feathers
gets transferred to the inner
wing → moves body forward.
• Kinds of Flight: Rough-legged Hawk
1) soaring flight - gliding without
flapping, no forward thrust.
- gradually lose altitude, or sink.
- sink rate increases with drag at
intermediate air speeds.
- strategy: gain altitude then sink
at most efficient rate.
a) thermal soaring - ride one thermal (column of warm rising air),
then sink to the next thermal and continue often without flapping.
- thermal air can rise at 4 m per
sec, birds can sink 1-2 m per sec.
- slots help decrease drag across
a wing with a med aspect ratio.
- ex. Broad-winged Hawks form
kettles - group of birds all rising
and gliding together.
2) flapping flight -
with forward thrust.
- change to the
horizontal → thrust.
- ex. helicopter
blades and rotor.
- each primary American Crow
works as an airfoil.
- twist during the
power downstroke.
- many tiny vertical
airfoils, thrust.
- control - from tendons and muscles attached to each primary.
- smooth integrated feather position through strokes maintained
by flexible recoil of feather vanes.
- outer wing forces transfer to inner wing forces, forward flight.
- increase speed with deeper strokes rather than faster wingbeats.
2
 - leading-edge vortices - swirls of air
• Soaring Flight coming off the leading edge that aid in lift.
• Flapping Flight - leading edge of hummingbird wing:
- proximal edge - round and thick, air
passes distally away from the body.
- distal edge - thin and tapered, air
passes distally off the tip of the wing.
- olive oil fine mist chamber revealed
vortices to aid in creating sufficient lift.
- ex. Swifts also
benefit from vortices.
- use subtle changes
in lift to turn quickly
to capture insects.
- tails - control flight position, stability, steering.
- reduces turbulence of air passing over body.
- also add lift by improving airflow over wings.

- more flapping flight: • combinations of flight:

- ex. Hummingbirds. - intermittent flight - bouts of flapping
- able to move forward, followed by non-flapping.
backward, and hover by - strategy: reduce the cost of power.
changing the direction a) flap-gliding - flap several times,
of the wingbeat. then glide. ex. Hawks and Vultures.
Red-tailed Hawk
- velocity increases with - not often flapping in open air.
tilt, top speed 26 mph. - wings spread, reduces cost at slow speeds, usually large birds.
- due to unique pectoral b) flap-bounding - flap several times, drop, then flap again.
girdle and humerus - wings folded, reduce cost at high speeds, usually small birds.
articulation. - ex. Finches and Woodpeckers.
- long primaries, short - ex. American Goldfinches sing a
secondaries - allows flight song while flapping, stop, and
figure 8 wing stroke. sing again (as they rise and fall).
- enables both a powerful downstroke and an upstroke.
c) mixed - medium sized birds.
- upstroke generates only one third that of the downstroke.
- ex. European Starlings when slow
(until recently considered to be equal).
will flap-glide, fast will flap-bound. American Goldfinch

- speed stooping - increased speeds Bar-headed Geese - flying in formation - waterbirds fly
while diving at prey, as in Falcons. in familiar “V” pattern, saves energy.
- ex. - Peregrine Falcon is the master. - high wing load, heavy body small
- angle of 30-60 degrees, begun as wings, ex. Ducks, Geese, Swans.
high as 1500 m, speeds 96-264 mph. - each flies off and behind the wing
- falconers skydive with their falcons. tip of bird in front of it.
- 144 mph, tucked wings, extended - result: cancels some air turbulence of own wing tips, less drag.
shoulders like a diamond shape. - group airstream allows those in back to glide more than leaders.
- 200 mph, elongated to streamline its Peregrine Falcon - back of the line - save up to 50% of the energy as a leader.
shape pulling wings in tight and extended its head. - landing - birds are unique in the way that they land.
- descends in a spiral pattern, to keep - birds: rotate center of mass upward to stall, land feet first.
its head as straight as possible - mammals: make contact with front limbs, rotate body downward,
(decrease drag).
then hind limbs touch
- benefit of less drag outweighs
surface.
longer descent path.
- allows birds better
- attacks birds on the wing, usually control, esp. predators.
stun or grab them. ex. Pigeons.

3
 • Wing Shape and
• Intermittent Flight Size:
• Speed Stooping - Grebes have a high
wing load.
• Formation
- Songbirds have a
low wing load.
- about 0.1-0.2 g per
Cerulean Warbler
square centimeter.
A) high aspect ratio
wings that are long
and narrow.
- ex. Albatrosses.
- high speed gliding in
high winds.
- leading edge produces more lift than the rear half.
- more lift than short broad wings of equal area and load.
- Comparative Method: Turkey Vulture ↑ aspect ratio, soar early.
- Black Vulture ↓ aspect ratio, soar later in the morning.

B) low aspect ratio wings that are short and • Bird Skeleton:
rounded for fast takeoffs and easy maneuvers. adapted for flight.
- esp. in and around vegetation. ex. Grouse. - many bones fused
- many Songbirds, escape predators and and reinforced for
catch prey. strength and to lighten
the load.
C) med aspect ratio without slots - pneumatic bones -
for fast efficient flight in open habitat. hollow inside, long
- ex. Falcons. bones have internal
cross supporting
struts.
- bones support large,
complex flight muscles.
D) med aspect ratio with slots - uncinate process - bony flaps that extend posteriorly from
for increased lift and gliding. vertical upper ribs to overlap adjacent rib → support rib cage.
- ex. Hawks. - sternum - typically keeled for flight muscle attachment.
- keel - large in powerful flyers.
- absent or reduced in flightless birds.

- coracoid - largest bone of - upstroke - supracoracoideus

the shoulder joint forming a contracts and the humerus gets
vertical column to the joint. pulled up as if by a pulley.
- scapula - long and blade- - downstroke - pectoralis
like running horizontal from contracts and the humerus gets
the shoulder joint. pulled down.
- furcula - “wishbone” serves - pectoralis - muscle that
a spring-like function. originates on the keeled sternum
- triosseal canal - space - different sections of the and inserts on the humerus.
between coracoid, scapula, muscle allow for takeoff, level - may account for as much as
and furcula through which the flight, and landing. 35% of a birds body weight.
suprcoracoideus tendon - also attaches to furcula and
passes to insert on humerus. membrane between coracoids
- supracoracoideus - muscle and the furcula.
(along with dorsal elevator muscles) used to pull the wing up. - may also attach thinly to rib
- typically smaller than the pectoralis (removed in diagram). cage in birds with shallow keels.
- when experimentally cut, cannot lift off ground, but can sustain - ex. climbers like Woodpeckers.
flight once in the air (dorsal elevator muscles are sufficient).

4
 - humerus,
• Aspect Ratio radius, and ulna
• The Bird Skeleton are homologous
to all vertebrates.
- joints allow wing
to fold in.
- as well as flexible positions during all aspects of flight.
- quill knobs - bony projections on the posterior margin of the
ulna where secondary feathers attach.
- most outer wing bones, primary feathers attach, are fused.
- radiale and ulnare form the free carpals (wrist bones).
- most other vertebrates have 10 or more carpals.
- carpometacarpus - fused first, second, and third metacarpals.
- fourth and fifth digits of the hand are lost in birds.
- first digit - phalanges 1 and 2, “thumb”, supports the alula.
- second and third digit - and associated phalanges most distal.