97
Male Cottus typically excavate nest cavities beneath rocks and guard the developing
eggs after spawning (Morris 1954 ; Downhower and Brown 1980 ; Goto 1993 ).
Cottus sounds have been primarily observed in agonistic contexts as a mechanism
to deter intruders (Ladich 1989 ; Colleye et al. 2013 ), though it is unclear whether
the prevalence of agonistic sound observations refl ects context-specifi c signaling
rates or rather the diffi culty of observing spawning in the laboratory (but see Kierl
and Johnston 2010 ). Marine sculpins exhibit a diversity of spawning modes, but
soniferous Myoxocephalus scorpius males are known to guard crevice-laid eggs
until hatching (Ennis 1970 ). The presence of paternal care in the soniferous
Myoxocephalus octodecemspinous and Myoxocephalus aenaeus is uncertain. Male
Myoxocephalus jaok and M. brandti guard eggs (Panchenko 2001a , b ), but their
acoustic behaviour has not been tested.
The gobiid pattern of reproduction is quite stereotyped in its basic scheme across
species: the male selects and occupies a nest cavity below a hard object, often excavating or at least adjusting this cavity, and defends it by patrolling the area surrounding the nest (Tavolga 1956 ; Torricelli et al. 1985 ). The hard object varies greatly
within and between species, but commonly includes lamellibranch or gastropod
valves, stones, reedstand, or artifi cial objects. Once a ripe female is in a male’s
visual fi eld, the male swims towards her, attempting to lead the female into the nest
cavity with stereotyped swimming movements (“approaching-leading behaviours”).
Sounds are typically emitted throughout this courtship sequence, which occurs primarily outside the nest. Once in the nest and ready to spawn, the female inverts her
position to release eggs onto the nest ceiling. At this stage the male may emit the
so-called pre-spawning sounds before inverting to release sperm. Upside-down
behaviours of both partners, male sound emission and patrol of the nest entrance are
alternated throughout the duration of spawning (Lugli et al. 1997 ; Lugli and
Torricelli 1999 ; Malavasi et al. 2008 ).
Gobiid sounds are mainly documented in agonistic and/or reproductive contexts.
In eight species the vocal behaviour was shown to be associated with both contexts
(Table 2 ): Padogobius bonelli (formerly P. martensii ), Gobius paganellus , Gobius
niger , Zosterisessor ophiocephalus , Pomatoschistus canestrinii , Pomatoschistus
pictus , Bathygobius fuscus , Gobiosoma bosci . For some of the remaining species,
the occurrence of sounds in only one context could result from a lack of deeper
investigation rather than the real absence of acoustic signaling in a given context;
reproductive behaviour is diffi cult to achieve in captivity in some species (e.g. mudskippers). Despite the high prevalence of sound production among gobies, detailed
descriptions of sound production and associated behaviours are available for only a
few species: P. bonelli (Lugli et al. 1995 , 1997 ), Pomatoschistus minutus (Lindström
and Lugli 2000 ), P. canestrinii (Malavasi et al. 2009 ), and P. pictus (Amorim and
Neves 2007 ). Within the reproductive context, these species’ sounds are associated
with pre-spawning and/or courtship phases.
Convergent Aspects of Acoustic Communication in Darters, Sculpins, and Gobies
Male Cottus typically excavate nest cavities beneath rocks and guard the developing
eggs after spawning (Morris 1954 ; Downhower and Brown 1980 ; Goto 1993 ).
Cottus sounds have been primarily observed in agonistic contexts as a mechanism
to deter intruders (Ladich 1989 ; Colleye et al. 2013 ), though it is unclear whether
the prevalence of agonistic sound observations refl ects context-specifi c signaling
rates or rather the diffi culty of observing spawning in the laboratory (but see Kierl
and Johnston 2010 ). Marine sculpins exhibit a diversity of spawning modes, but
soniferous Myoxocephalus scorpius males are known to guard crevice-laid eggs
until hatching (Ennis 1970 ). The presence of paternal care in the soniferous
Myoxocephalus octodecemspinous and Myoxocephalus aenaeus is uncertain. Male
Myoxocephalus jaok and M. brandti guard eggs (Panchenko 2001a , b ), but their
acoustic behaviour has not been tested.
The gobiid pattern of reproduction is quite stereotyped in its basic scheme across
species: the male selects and occupies a nest cavity below a hard object, often excavating or at least adjusting this cavity, and defends it by patrolling the area surrounding the nest (Tavolga 1956 ; Torricelli et al. 1985 ). The hard object varies greatly
within and between species, but commonly includes lamellibranch or gastropod
valves, stones, reedstand, or artifi cial objects. Once a ripe female is in a male’s
visual fi eld, the male swims towards her, attempting to lead the female into the nest
cavity with stereotyped swimming movements (“approaching-leading behaviours”).
Sounds are typically emitted throughout this courtship sequence, which occurs primarily outside the nest. Once in the nest and ready to spawn, the female inverts her
position to release eggs onto the nest ceiling. At this stage the male may emit the
so-called pre-spawning sounds before inverting to release sperm. Upside-down
behaviours of both partners, male sound emission and patrol of the nest entrance are
alternated throughout the duration of spawning (Lugli et al. 1997 ; Lugli and
Torricelli 1999 ; Malavasi et al. 2008 ).
Gobiid sounds are mainly documented in agonistic and/or reproductive contexts.
In eight species the vocal behaviour was shown to be associated with both contexts
(Table 2 ): Padogobius bonelli (formerly P. martensii ), Gobius paganellus , Gobius
niger , Zosterisessor ophiocephalus , Pomatoschistus canestrinii , Pomatoschistus
pictus , Bathygobius fuscus , Gobiosoma bosci . For some of the remaining species,
the occurrence of sounds in only one context could result from a lack of deeper
investigation rather than the real absence of acoustic signaling in a given context;
reproductive behaviour is diffi cult to achieve in captivity in some species (e.g. mudskippers). Despite the high prevalence of sound production among gobies, detailed
descriptions of sound production and associated behaviours are available for only a
few species: P. bonelli (Lugli et al. 1995 , 1997 ), Pomatoschistus minutus (Lindström
and Lugli 2000 ), P. canestrinii (Malavasi et al. 2009 ), and P. pictus (Amorim and
Neves 2007 ). Within the reproductive context, these species’ sounds are associated
with pre-spawning and/or courtship phases.
Convergent Aspects of Acoustic Communication in Darters, Sculpins, and Gobies