First published online February 1, 2008
Journal of Experimental Biology 211, 502-509 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.008474
Development of vocalization, auditory sensitivity and acoustic communication in the Lusitanian toadfish Halobatrachus didactylus
Raquel O. Vasconcelos1,2,3,* and
Friedrich Ladich2
1 Departamento de Biologia Animal e Centro de Biologia Ambiental, Faculdade de
Ciências da Universidade de Lisboa. Bloco C2 Campo Grande, 1749-0161
Lisbon, Portugal
2 Unidade de Investigação em Eco-Etologia, I.S.P.A. Rua Jardim do
Tabaco 34, 1149-041 Lisbon, Portugal
3 Department of Behavioural Biology, University of Vienna, Althanstrasse 14,
1090 Vienna, Austria

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Fig. 1. Oscillogram of a single grunt of a juvenile H. didactylus showing
temporal sound characteristics analyzed (PP, pulse period).
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Fig. 2. Sonogram and oscillogram of a grunt call produced by a representative
H. didactylus of (A) group G2 (6.1 cm SL) and (B) group G5
(28.5 cm SL), showing two single grunts (A) and a part of a grunt
train (B). Note the shorter grunt duration and lower dominant frequency in B.
Sampling frequency 6 kHz, filter bandwidth 15 Hz, 70% overlap,
Blackman–Harris window.
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Fig. 3. Log–log plot of mean number of pulses against standard length
(SL, in cm). Regression equation: log number of
pulses=–0.72xlog(SL)+1.47. N=44,
P<0.001, r2=0.579.
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Fig. 4. Correlation between mean pulse period and standard length (SL).
Regression equation: pulse period=0.12xSL+6.08. N=44,
P<0.001, r2=0.480.
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Fig. 5. Correlation between mean sound pressure level and log standard length
(SL, in cm). Regression equation: sound pressure
level=34.70xlog(SL)+92.56. N=38, P<0.001,
r2=0.903.
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Fig. 6. Cepstrum-smoothed sound power spectra (mean) of the Lusitanian toadfish
grunt call from groups G1 (3.8 cm SL, N=1); G2
(5.4–6.6 cm SL, N=5); G3 (8.0–10.2 cm
SL, N=9); G4 (12.4–15.3 cm SL, N=6);
and G5 (20.2–31.8 cm SL, N=8). Sampling frequency 6
kHz, filter bandwidth 1 Hz, 75% overlap, Blackman–Harris window.
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Fig. 7. Auditory thresholds of juveniles from the five different size groups. G1,
2.8–3.8 cm SL (N=6); G2, 5.4–6.6 cm SL
(N=6); G3, 8.0–10.2 cm SL (N=7); G4,
12.4–15.3 cm SL (N=6); and G5, 20.2–31.8 cm
SL (N=9). Values are mean ± s.e.m. Asterisks indicate
highly statistically significant differences between groups (one-way ANOVA);
**P<0.001.
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Fig. 8. Relation between auditory sensitivities and cepstrum-smoothed sound power
spectra of grunt calls in the five size groups calculated for a distance of 10
cm. For size ranges see Figs 6
and 7.
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© The Company of Biologists Ltd 2008