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(A) Dissected type I male Lusitanian toadfish ( Halobatrachus didactylus ) showing the swim bladder (SB), the gonads (G), and the accessory glands (AG). (B) Dorsal view of one lobe of the swim bladder depicting the embedded sonic muscle (SM). 

(A) Dissected type I male Lusitanian toadfish ( Halobatrachus didactylus ) showing the swim bladder (SB), the gonads (G), and the accessory glands (AG). (B) Dorsal view of one lobe of the swim bladder depicting the embedded sonic muscle (SM). 

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Animal vocalizations are good examples of signals that have been shaped by sexual selection and often contribute to resolve contests or the choice of mates. We relate the mass of the sound-producing muscles of a highly vocal fish species, the Lusitanian toadfish (Halobatrachus didactylus (Bloch and Schneider, 1801)), with the sender's physical feat...

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Context 1
... (M G ; Fig. 1A), accessory glands (in males, M AG ; Fig. 1A), and liver (M L ) mass were tallied to the nearest milligram. Both sonic muscles, which are embedded in the sides of the swim bladder (Figs. 1A, 1B), were gently cut from the swim-bladder wall with a pair of fine dissection scissors and were also weighed (M SM ) to the nearest milli- gram. ...
Context 2
... (M G ; Fig. 1A), accessory glands (in males, M AG ; Fig. 1A), and liver (M L ) mass were tallied to the nearest milligram. Both sonic muscles, which are embedded in the sides of the swim bladder (Figs. 1A, 1B), were gently cut from the swim-bladder wall with a pair of fine dissection scissors and were also weighed (M SM ) to the nearest milli- gram. We also obtained total length (TL), measured ...

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... 1), and was not observed in juvenile specimens in the present sample. The cordiform morphology cooccurs with hypertrophy of the intrinsic sonic muscles on the sides of the swim bladder, which increases the width of lobes, resulting in the heart-shaped aspect (Bass and Marchaterre, 1989a, Fig. 1;Brantley and Bass, 1994;Amorim et al., 2009;Mohr et al., 2017). ...
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Lusitanian toadfish males that provide parental care rely on acoustic signals (the boatwhistle) to attract females to their nest. We test the hypothesis that male quality, namely male size and condition that are relevant for parental success, is reflected in vocal activity and boatwhistle characteristics and thus advertised to females. We recorded 22 males over a week during the peak of the breeding season. Calling rate and calling effort (percentage of time spent calling) strongly reflected male condition (lipid content of somatic muscles) and to a smaller extent sonic muscle hypertrophy and larger gonads. Males in better condition (increased body lipid and relative higher liver mass) also contracted the sonic muscles at faster rate as shown by the shorter boatwhistle pulse periods. Amplitude modulation reflected the degree of sonic muscle hypertrophy. None of the measured male quality parameters were good predictors of boatwhistle duration and dominant frequency. Altogether this study strongly suggests that Lusitanian toadfish males advertise their quality to females primarily with boatwhistle calling rate and calling effort, which mainly reflect male condition. Because pulse period had low variability, consistent with the existence of a vocal central pattern generator, we suggest that males that sustain sonic muscles contraction at a very fast rate close to their physiological limit may be honestly advertising their quality (condition). Similarly, males that produce boatwhistles with higher amplitude modulation, a feature that seems dependent on sonic muscle hypertrophy, could be more attractive to females.
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Fishes have evolved multiple mechanisms for sound production, many 5 of which utilize sonic muscles that vibrate the swimbladder or the rubbing of bony 6 elements. Sonic muscles are among the fastest muscles in vertebrates and typically 7 drive the swimbladder to produce one sound cycle per contraction. These muscles 8 may be extrinsic, typically extending from the head to the swimbladder, or intrinsic, 9 likely a more-derived condition, in which muscles attach exclusively to the bladder 10 wall. Recently discovered in Ophidiiform fishes, slow muscles stretch the swim- 11 bladder and associated tendons, allowing sound production by rebound (cock and 12 release). In glaucosomatids, fast muscles produce a weak sound followed by a 13 louder one, again produced by rebound, which may reflect an intermediate in the 14 evolution of slow to superfast sonic muscles. Historically, the swimbladder has 15 been modeled as an underwater resonant bubble. We provide evidence for an 16 alternative hypothesis, namely that bladder sounds are driven as a forced rather than 17 a resonant response, thus accounting for broad tuning, rapid damping, and direc- 18 tionality of fish sounds. Cases of sounds that damp slowly, an indication of reso- 19 nance, are associated with tendons or bones that continue to vibrate and hence drive 20 multiple cycles of swimbladder sound. Stridulation sounds, best studied in catfishes 21 and damselfishes, are produced, respectively, as a series of quick jerks causing 22 rubbing of a ribbed process against a rough surface or rapid jaw closing mediated 23 by a specialized tendon. A cladogram of sonic fishes suggests that fish sound 24 production has arisen independently multiple times.