First published online March 16, 2007
Journal of Experimental Biology 210, 1148-1160 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.002493
Scaling of the axial morphology and gap-bridging ability of the brown tree snake, Boiga irregularis
Bruce C. Jayne1,* and
Michael A. Riley2
1 Department of Biological Sciences, University of Cincinnati, PO Box
210006, Cincinnati, OH 45221-0006, USA
2 Department of Psychology, University of Cincinnati, PO Box 210376,
Cincinnati, OH 45221-0376, USA

View larger version (16K):
[in this window]
[in a new window]
|
Fig. 1. The smallest (SVL=43 cm, mass=9.9 g) and second largest
(SVL=184 cm, mass=1137 g) brown tree snakes used to determine the
scaling of anatomy and gap-bridging performance. The smaller snake is near the
size of a hatchling.
|
|

View larger version (5K):
[in this window]
[in a new window]
|
Fig. 2. Methods used to quantify the posture of the snake one frame before bridging
a gap. The image is a tracing of a right lateral view of a video of a snake
with SVL=57 cm crossing a 30 cm gap. The pegs in the large horizontal
cylinder are inclined approximately 45° relative to the horizontal plane,
and the tips of those in the foreground were oriented towards the camera. The
origin of the coordinate system used for kinematic analysis was the top center
end of the perch upon which the snake was initially placed. The longitudinal
positions were measured or calculated for the most posterior location of the
snake (Xposterior), mid-way between the end of the initial
perch and Xposterior (Xmid-perch) and
the most anterior location of the snake that was underneath part of the
initial perch or one of its pegs (third right peg from the edge in this
example) so that upward forces could be applied (Xanchor).
The x and y coordinates were digitized for the lowest
location of the mid-dorsal line and center of mass (COM) for the portion of
the body suspended in the gap between the two perches.
|
|

View larger version (20K):
[in this window]
[in a new window]
|
Fig. 3. Schematic right lateral view of the major epaxial muscles of the brown tree
snake at mid-body. SSP-SP, semispinalis-spinalis muscle; M, multifidus muscle;
LD, longissimus dorsi muscle; IL iliocostalis muscle. The colored areas
indicate contractile tissue, the tendons are white, and the skeletal
structures are gray. Vertebrae are numbered beginning at the anterior
attachment site of a single segment of the SSP-SP, and five vertebrae
(37) are not shown. The thin horizontal line represents the shelf
formed by the pre- and postzygapophyses.
|
|

View larger version (91K):
[in this window]
[in a new window]
|
Fig. 4. Cross-sections at mid-body of the smallest (SVL=43 cm) and largest
(SVL=188 cm) snakes. On the large section the major epaxial muscles
for which cross-sectional area were determined are tinted blue
(semispinalis-spinalis), yellow (multifidus), red (longissimus dorsi) and
green (iliocostalis).
|
|

View larger version (32K):
[in this window]
[in a new window]
|
Fig. 5. Tracings from videotapes showing lateral (left) and dorsal (right) views of
different gap-bridging behaviors. (A) Crawling behavior of a snake with
SVL=136 cm crossing a 67 cm gap. (B) Lunging and tail wrapping
behaviors of a snake with SVL=106 cm crossing a 52 cm gap. The most
anterior anchor points in A and B are the fifth right peg and the third left
peg from the edge of the initial perch, respectively. The white marks indicate
intervals approximately one-eighth SVL along the body of the snake.
The times before contacting the destination perch are to the left of each
figure. The portion of the snake spanning the gap was nearly straight during
crawling, whereas early in the lunge the body had several lateral bends that
were later straightened. Videos of these behaviors can be viewed at
http://bioweb.ad.uc.edu/faculty/jayne/videos.htm.
|
|

View larger version (6K):
[in this window]
[in a new window]
|
Fig. 6. The scaling of maximal gap distance crossed versus snake size. The
solid line is the least-squares regression for log10-transformed
data, and the broken line indicates a 1:1 ratio of snake SVL and gap
distance. Each symbol represents one snake. Circles indicate the crawling
behavior (see Fig. 5A);
triangles, lunging behavior; inverted triangles, tail-wrapping behavior.
Squares indicate a lunge while the tail was wrapped around the initial perch
(Fig. 5B). A solid symbol
indicates that the snake had a conspicuous anchor point on the initial perch
against which upward forces could be applied
(Fig. 2), whereas a hollow
symbol indicates the lack of such a point. See
Table 1 for regression
statistics.
|
|

View larger version (7K):
[in this window]
[in a new window]
|
Fig. 7. The kinematics of a snake using the crawling behavior to cross a gap. (A)
Forward (B) vertical and (C) lateral displacement of the snout of the snake
versus time. The time intervals between successive points are all 100
ms. The snake touched the perch on the far side of the gap at 0 s. After
moving forward with a nearly constant speed (4.5 to 2.5 s), the
snake gradually slowed before reaching the destination perch. Gap distance, 38
cm; snake SVL, 65 cm.
|
|

View larger version (10K):
[in this window]
[in a new window]
|
Fig. 8. Variation in the paths traveled by the snout of snakes while crawling
across a gap. (A,C,E) Lateral views and (B,D,F) overhead views. (A,B) Snake
SVL=65 cm, gap distance=38 cm; (C,D) SVL=54 cm, gap
distance=30 cm; (E,F) SVL=87 cm, gap distance=38 cm. The time
intervals between successive points are all 100 ms. The end and top center of
the initial perch are at the origin of the x, y and z
axes.
|
|

View larger version (16K):
[in this window]
[in a new window]
|
Fig. 9. The kinematics and the path traveled by a snake using the lunging behavior
to cross a gap. (A) Forward and (B) lateral displacement versus time,
and (C) a lateral view of the path traveled by the snout of the snake
(SVL=106 cm) while crossing a 52 cm gap. The times between the last
thirteen successive points are 33 ms; all other time intervals between
successive points are 200 ms. The snake touched the perch on the far side of
the gap at 0 s. Different colored symbols indicate four attempts to bridge the
gap, for which only the last was successful. The rapid forward and vertical
velocities (slope) within the last 0.5 s indicate when the snake lunged. The
end and top center of the initial perch are at the origin of the x, y
and z axes.
|
|
© The Company of Biologists Ltd 2007