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First published online September 19, 2006
Journal of Experimental Biology 209, 3812-3827 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02443

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The locomotor kinematics of Asian and African elephants: changes with speed and size

John R. Hutchinson^1,*, Delf Schwerda², Daniel J. Famini³, Robert H. I. Dale⁴, Martin S. Fischer² and Rodger Kram⁵

¹ Structure and Motion Laboratory, Department of Veterinary Basic Sciences, The Royal Veterinary College, University of London, Hatfield, Hertfordshire, AL9 7TA, UK
² Institut fuer Spezielle Zoologie und Evolutionsbiologie, mit Phyletischem Museum, Jena, 07743, Germany
³ Department of Integrative Biology, University of California, Berkeley, CA 94720-3140, USA
⁴ Department of Psychology, Butler University, Indianapolis, IN 46208, USA
⁵ Department of Integrative Physiology, University of Colorado, Boulder, CO 80309-0354, USA

View larger version (127K): [in a new window]   Fig. 1. Mid-stance phase of the right hindlimb, shown in right lateral view for representative slower and faster locomotion of an adult (h=1.93 m) African elephant (A, 1.7 m s-1; B, 4.1 m s-1) and a subadult (h=1.41 m) Asian (C, 1.2 m s-1; D, 5.4 m s-1) elephant. Limb phases were: (A) Plf=0.15, Prh=0.51, Prf=0.66; (B) Pf=0.23, Prh=0.52, Prf=0.73; (C) Plf=0.18, Prh=0.48, Prf=0.65; (D) Plf=0.22, Prh=0.52, Prf=0.71 (compare with Fig. 3). Markers are explained in the text.

View larger version (10K): [in a new window]   Fig. 2. Typical elephant foot-on (A) and foot-off (B) events. Right hind foot of an African elephant shown in lateral view during slow walking (1.2 m s-1; tst=1.5 s).

View larger version (36K): [in a new window]   Fig. 3. Relative limb phase (P; with left hind foot contact defined as 0.0; others as fraction of a stride following that contact) plotted against dimensionless speed for African (filled symbols) and Asian (open symbols) elephants. Circles, left front (Plf); triangles, right hind (Prh); squares, right front (Prf) feet.

View larger version (18K): [in a new window]   Fig. 4. Duty factor (A) and fore-minus hindlimb duty factor difference (ßdiff; B) plotted against dimensionless speed for African (blue; filled triangles) and Asian (red; open triangles) elephants.

View larger version (15K): [in a new window]   Fig. 5. Stance (blue triangles) and swing (green squares) phase durations (A) and relative stride lengths (triangles) and frequencies (squares) (B) plotted against dimensionless speed for African (filled symbols) and Asian (open symbols) elephants.

View larger version (20K): [in a new window]   Fig. 6. Relationships of body mass for each individual African (blue triangles) and Asian (red squares) elephant with: near-maximal forward velocity (A), minimum observed duty factor (B), maximum observed stride length (triangles) and frequency (squares) (C; symbols are dark for African, light for Asian), and hip height for all elephants combined (D). Hip height (h) least-squares power curve fit shown: h=0.23M 0.26b; N=60 [excluding two elephants (from Christian et al., 1999)], R2=0.73; upper/lower 95% confidence intervals 0.26/0.33. This result does not change markedly if the baby elephant outlier is excluded (h=0.26M 0.24b; R2=0.69), or if RMA (Model II) regression is used (slope=0.29).

View larger version (34K): [in a new window]   Fig. 7. Relationships of stride length (blue triangles) and stride frequency (green squares) to forward velocity for African (filled symbols) and Asian (open symbols) elephants. Circled data points indicate data for the smallest elephant (African; 116 kg).

View larger version (16K): [in a new window]   Fig. 8. Vertical displacements (from the lowest point shown) of the right (in lateral view) shoulder (red triangles) and hip (blue circles) joints for a representative Asian elephant (h=1.41 m) moving at 1.2 m s-1 (in A; 3 strides) and 5.4 m s-1 (in B; <2.5 strides), plotted against time. Large symbols indicate motion during the stance phase; small symbols are motion during the swing phase of the respective limb.

View larger version (28K): [in a new window]   Fig. 9. Duty factor minus right forelimb relative limb phase (Prf) (see Discussion) required to have an aerial phase for the entire body, plotted against dimensionless speed for African (filled symbols) and Asian (open symbols) elephants. A value of 0 or less would require a whole-body aerial phase.