First published online March 9, 2004
Journal of Experimental Biology 207, 1345-1352 (2004)
Published by The Company of Biologists 2004
doi: 10.1242/jeb.00889
Take-off mechanics in hummingbirds (Trochilidae)
Bret W. Tobalske1,*,
Douglas L. Altshuler2 and
Donald R. Powers3
1 Department of Biology, University of Portland, 5000 North Willamette
Boulevard, Portland, OR 97203, USA
2 Bioengineering 138-78, California Institute of Technology, 1200 East
California Boulevard, Pasadena, CA 91125, USA
3 Biology Department, George Fox University, 414 N. Meridian Street,
Newberg, OR 97132, USA

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Fig. 1. Kinematic events during an autonomous take-off in a male rufous hummingbird
(Selasphorus rufus; body weight=30.7 mN). (AF) Selected frames
from 1000 Hzvideo illustrating body and wing posture; letters and timing (ms)
for each frame correspond directly to those on the graph below. Vectors
represent reaction forces produced by the hindlimbs on the perch; white scale
bars, 10 mN. This sequence lasts 150 ms, and two wingbeats were completed
prior to the end of take-off, defined as the start of the first downstroke
after end of leg thrust.
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Fig. 2. Relative timing of kinematic events and peak ground-reaction forces during
autonomous (N=6), escape (N=6) and aggressive (N=4)
take-off in the rufous hummingbird (Selasphorus rufus) and autonomous
take-off in the zebra finch (Taeniopygia guttata; N=1).
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Fig. 3. Selected kinematic and mechanical variables in the rufous hummingbird,
Selasphorus rufus, as a function of take-off treatment. (A) Peak
vertical and horizontal ground-reaction forces (N) and peak accelerations (m
s2). Vertical forces are net after subtracting
g. (B) Velocity (m s1) at the end of leg
thrust and take-off. (C) Wingbeat frequency (Hz; inverse of wingbeat duration)
during the first and second wingbeat in the take-off. N=6
hummingbirds for autonomous and escape and N=4 for aggressive; error
bars represent ± S.D.
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Fig. 4. Scaling of velocity at end of take-off in 17 bird species including, in
order of increasing body mass: rufous hummingbird (Selasphorus
rufus); zebra finch (Taeniopygia guttata); dark-eyed junco
(Junco hyemalis); house finch (Carpodacus mexicanus);
diamond dove (Geopelia cuneata); American robin (Turdus
migratorius); European starling (Sturnus vulgaris); western
scrub jay (Aphelocoma californica); northern flicker (Colaptes
auratus); ringed turtle-dove (Streptopelia risoria); European
migratory quail (Coturnix coturnix); American crow (Corvus
brachyrhynchos); rock dove (Columba livia); black oystercatcher
(Haematopus bachmani); ring-necked pheasant (Phasianus
colchicus); herring gull (Larus argentatus); wild turkey
(Meleagris gallopavo). Regression lines from reduced-major axis
regression of log-transformed data. (A) Species data, with open circle
representing rufous hummingbird; y=0.25x0.21,
r=0.68. (B) Independent contrasts. Open circle represents contrast
that includes rufous as daughter taxon; y=0.26x, r=0.65.
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© The Company of Biologists Ltd 2004