First published online March 2, 2007
Journal of Experimental Biology 210, 993-1005 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.001990
Clicking caterpillars: acoustic aposematism in Antheraea polyphemus and other Bombycoidea
Sarah G. Brown1,
George H. Boettner2 and
Jayne E. Yack1,*
1 Department of Biology, Carleton University, Ottawa, Ontario, K1S 5B6,
Canada
2 Plant Soil and Insect Sciences, University of Massachusetts, Amherst, MA
01003, USA

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Fig. 2. Sound-producing structures of Antheraea polyphemus larvae. (A)
Close-up of the head region. (B) Scanning electron micrograph of the
mouthparts. The left and right mandibles (arrows) lie below the labrum. (C,D)
Scanning electron micrographs of the left and right mandibles, respectively.
Scale bars, 250 µm.
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Fig. 3. Oscillograms of Antheraea polyphemus sounds recorded from fifth
instar larvae. (A) Sounds recorded from an individual feeding on Quercus
rubra leaves at 10 cm. (B) A click train, showing the typical click
pattern of a larva after being pinched with forceps at 10 cm. (C) Three clicks
from the train in B (denoted by black circles) with an expanded time scale
showing the multiple components of clicks. The most typical pattern, a
double-component click, is shown on the left. (D) Click spectra from five
different individuals. The bandwidth varies, but in all clicks most energy is
between 8 and 18 kHz.
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Fig. 4. (A) Oscillograms of late instar Antheraea polyphemus sounds
obtained during one-, two- and five-pinch trials. Arrows indicate the times
when each larva was attacked. (B) Number of clicks in 60 s produced by late
instar larvae following the first attack in one- (20.6±35.5), two-
(25.1±22.5) and five-pinch (54.3±46.1) trials (means ±
s.d.). The total amount of signaling is positively correlated with the number
of attacks (*P<0.001).
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Fig. 5. (A) A representative oscillogram of sounds produced by a fifth instar
Antheraea polyphemus larva when pinched five consecutive times with
forceps. The larva first regurgitated (R) after the second pinch was
administered. (B) Defensive regurgitation in a fifth instar larva in response
to a simulated predator attack. (C) The percentage of regurgitating late
instar larvae in one-, two- and five-pinch trials. (D) The number of pinches
required to elicit regurgitation in late instar larvae. Larvae most commonly
regurgitated after the second pinch.
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Fig. 6. (A) The escalation in behavioural responses of late instar Antheraea
polyphemus larvae to increasing levels of disturbance. CR,
neither clicking nor regurgitation; C+R, clicking only; CR+,
regurgitation only; C+R+, clicking and regurgitation. C+R+ increased between
one-, two- and five-pinch trials, whereas C+R decreased between one-,
two- and five-pinch trials. (B) The temporal relationship between clicking and
regurgitation. In almost all five-pinch trials, the acoustic signaling
occurred before regurgitation. *Significant difference
(P<0.001).
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Fig. 7. The amount of time taken for ants to carry untreated mealworm segments,
segments covered with distilled water, and segments covered in regurgitant,
into one of their foraging holes. Note that two trials in which mealworms were
coated in regurgitant were not included for analysis, because the mealworms
were never accepted by ants. Values are means ± s.d. *Significant
difference (P<0.001).
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Fig. 8. Mean consumption (± s.d.) of mouse chow coated in distilled water
(control diet) versus mouse chow coated in regurgitant (experimental
diet). (N=10). *Significant difference (P<0.005).
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Fig. 9. Fifth instar (A) Actias luna and (B) Manduca sexta
larvae. Scale bars, 1 cm. (C,D) Oscillograms of sounds produced by A.
luna and M. sexta, respectively, showing the typical click
patterns of larvae after being pinched with forceps. (E,F) Clicks from the
trains in C and D, respectively, with an expanded time scale showing that
clicks generally have two components.
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© The Company of Biologists Ltd 2007