BREVIA

constant and provides support (6). Bends

Underwater Bipedal Locomotion propagate down single arms from base to tip
in a relatively stereotyped wavelike fashion
(7) that appears to underlie bipedal locomo-
by Octopuses in Disguise tion in these animals. This motion has been
elicited in severed arms, without direct con-
Christine L. Huffard,1* Farnis Boneka,2 Robert J. Full1 trol from the brain (7, 8), and we hypoth-
esize that it allows the octopuses described
As described in Steinbeck_s Cannery Row (this is named the Bflamboyant display[) (4). here to move bipedally with minimal neural
(1), Bthe creeping murderer, the octopusI O. aculeatus may then raise the four lateral feedback.
pretending now to be a bit of weed, now a arms and walk backward using the ventral Crypsis is the primary defense of most
rockI runs lightly on the tips of its arms.[ arms (Fig. 1D and movie S2). Relative to octopuses, yet camouflage requires cephalo-
Far from the California tide pools in this book, mantle length, O. aculeatus has longer arms pods to remain still or move only very slow-
we have observed octopuses that do indeed than O. marginatus, and this is evident in ly (9). When an octopus moves quickly, it
walk. Individuals of Octopus marginatus (from their stride. The distal 75% of the walking becomes visually conspicuous and must
Indonesia) and Octopus (Abdopus) aculeatus arm rolls under the body, sometimes complet- employ unique behaviors to evade its pred-
(from Australia) move bipedally along sand ing more than a full wheel (Fig. 1D, frame 4). ator_s search image (9). By walking, both O.
using a rolling gait. This loco- marginatus and O. aculeatus
motion differs from their normal are able to move quickly while
crawling, which usually involves A using six of their arms to remain
several arms sprawling around the 0 1 2 disguised: O. marginatus per-
body, using the suckers to push B RU LD RD LU RU LD haps as a rolling coconut and
and pull the animal along (2). O. aculeatus as a clump of algae
Underwater video allowed kin- 0 1 2 3 4 5 tiptoeing away.
ematic analyses of the strides C RD LU LD RU RD LU LD RU
(3). (A stride is defined as two D References and Notes
steps or a complete cycle of leg 1. J. Steinbeck, Cannery Row (Viking,
movements in which a leg re- New York, 1945).
2. J. A. Mather, J. Comp. Psychol. 112,
turns to its initial relative po- 306 (1998).
sition.) While crawling quickly, 3. Materials and methods are available
O. marginatus draws six arms as supporting material on Science
Online.
around its body and moves back- 4. A. Packard, G. D. Sanders, Anim. Behav.
ward on the backmost (ventral) 19, 780 (1971).
arm pair (Fig. 1A and movie S1). 5. C. L. Vaughan, J. Biomech. 36, 513
The animal is pushed back as (2003).
6. W. M. Kier, K. K. Smith, Zool. J. Linn.
each arm tip alternates rolling Soc. 83, 307 (1985).
along the oral face (sucker edge) 7. Y. Gutfreund, T. Flash, G. Fiorito, B.
of approximately its distal half. Hochner, J. Neurosci. 18, 5976 (1998).
8. G. Sumbre, Y. Gutfreund, G. Fiorito,
The phase is variable (Fig. 1B), Fig. 1. Kinematics of bipedal movements in octopuses. (A) Video frame of T. Flash, B. Hochner, Science 293,
sometimes with short periods O. marginatus walking bipedally. (B) Phase diagram of O. marginatus. L, left 1845 (2001).
during which no arm is on the arm; R, right arm; U, arm lifted from the bottom; D, arm placed on the 9. R. T. Hanlon, J. W. Forsythe, D. E.
Joneschild, Biol. J. Linn. Soc. 66, 1
bottom. On average, each arm is bottom. (C) Phase diagram of O. aculeatus. (D) Gait diagram of O. aculeatus (1999).
on the sand for more than half drawn from video (7/60-s lapse between each panel). The arrowhead in the
10. Supported by an American Malaco-
first frame indicates the direction of walking. Six arms (white) coil, raise off logical Society Student Research
the stride, which kinematically the bottom, and obscure the head and mantle (gray). The right arm (orange)
Grant to C.L.H. and by NSF Fron-
qualifies these strides as walking pushes the animal backward throughout the sequence. The left arm (blue) is tiers in Integrative Biological Research
(duty factor 0.56, SD 0.08, n 0 4 lifted from the bottom in frame 4 and replaced on the bottom in frame 5. At grant no. 0425878 to R.J.F. Video
strides, two each from two ani- least one arm is in contact with the bottom at all times. (Video courtesy of of O. marginatus is provided by
Sea Studios, Inc., Monterey, CA, USA
mals). Two individual O. mar- Sea Studios, Inc.) (cameraman, Bob Cranston).
ginatus, both 55 mm in mantle
(or body sac) length (ML), moved bipedally at The octopus rolls along the sand as if on Supporting Online Material
1.2 and 2.6 ML per s, corresponding to 0.06 alternating conveyer belts. The phase of this www.sciencemag.org/cgi/content/full/307/5717/1927/
DC1
and 0.14 m/s, respectively. These speeds are stride is also variable, but at least one of the Materials and Methods
slightly faster than the average speed cal- two arms is always on the bottom (duty fac- Movies S1 and S2
culated for the same octopuses crawling with tor: right arm, 0.69; left arm, 0.91) (Fig.
11 January 2005; accepted 14 February 2005
several arms (1.1 ML per s, SD 0.36, or 0.06 1C), kinematically qualifying this movement 10.1126/science.1109616
m/s, SD 0.02; n 0 4 strides, two each from as walking.
two animals). Bipedal locomotion has been thought to 1
Department of Integrative Biology, University of
O. aculeatus moves bipedally in a slight- require the opposition of muscle against a California, Berkeley, CA 947203140, USA. 2Depart-
ly different manner. Walking is preceded by rigid skeleton (5). Instead, transverse, lon- ment of Fisheries and Marine Science, Universitas
a cryptic display in which the octopus coils gitudinal, and oblique bands of muscle in Sam Ratulangi, Manado, North Sulawesi, Indonesia.
and raises the two front (dorsal) arms above the arms allow octopuses exceptional flex- *To whom correspondence should be addressed.
its head and generally sits on the other six ibility, while their internal volume remains E-mail: chuffard@berkeley.edu

www.sciencemag.org SCIENCE VOL 307 25 MARCH 2005 1927