This paper outlines my research path over three decades while providing a review on the role of fish sounds in mate choice and reproduction. It also intends to provide advice to young scientists and point toward future avenues in this field of research. An overview of studies on different fish model species shows that male mating acoustic signals can inform females and male competitors about their size (dominant frequency, amplitude, and sound pulse rate modulation), body condition (calling activity and sound pulse rate), and readiness to mate (calling rate, number of pulses in a sound). At least in species with parental care, such as toadfishes, gobies, and pomacentrids, calling activity seems to be the main driver of reproductive success. Playback experiments ran on a restricted number of species consistently revealed that females prefer vocal to silent males and select for higher calling rates. This personal synthesis concludes with the suggestion to increase knowledge on fish mating signals, especially considering the emerging use of fish sounds to monitor aquatic environments due to increasing threats, like noise pollution.

As an undergraduate biology student at the University of Lisbon, I got an Erasmus grant to England to study foraging behaviour in fish. At the time, I had already decided to pursue a research career in Animal Behaviour. There, while reading the book, “The Behaviour of Teleost Fishes” (Pitcher, 1992), I came across Anthony Hawkins's chapter on underwater sound and fish behaviour (Hawkins, 1992). I found the idea of fish producing sounds to communicate during social interactions so fascinating that I wrote to Anthony Hawkins showing my interest in carrying out a PhD with him at the Marine Laboratory in Aberdeen. Soon after, I received a positive and warm reply letter and started my PhD later that year, focussing on the acoustic communication in triglids. That chapter, and Anthony Hawkins himself, have thus shaped my early research path.

During my PhD, I studied several highly soniferous triglid species. Captive triglids grunted and growled while being fed, providing the opportunity to investigate the role of acoustic signals during competitive feeding (Amorim and Hawkins, 2000; Amorim , 2004). I described their sounds and associated behaviour in detail, and investigated the ontogeny and the temporal patterns of sound production (seasonal and daily), among other aspects (Amorim, 1997, 2005; Amorim and Hawkins, 2000, 2005; Amorim , 2004). It was an exciting opportunity to work with one of the founders of fish bioacoustics and an enthusiast to this day.

Early researchers in fish bioacoustics provided very interesting insights on the communication function of acoustic signals (Hawkins, 1992; Crawford , 1997; Lugli , 1996; Bass and McKibben, 2003; Ladich, 2004), and researchers, like Friedrich Ladich (University of Vienna, Austria), Marco Lugli (University of Parma, Italy), John Crawford (University of Pennsylvania, USA), or Andrew Bass (Cornell University, USA), heavily influenced my early steps as I maintained a keen interest to pursue research in bioacoustics from a behavioural ecology perspective.

My research on fish acoustic communication continued with a post-doctoral (post-doc) fellowship at ISPA-Instituto Universitário (Portugal). During my first post-doc, I carried out research with the Mozambique tilapia, Oreochromis mossambicus (Amorim , 2003; Amorim and Almada, 2005) and stayed 6 months in Michael Fine's laboratory (Virginia Commonwealth University, USA) investigating the metabolic costs of sound production in the oyster toadfish, Opsanus tau (Amorim , 2002). While establishing my research in Portugal, and until now, Michael Fine has been my unseen mentor, providing valuable guidance, including numerous and extremely constructive reviews of my papers. During my second post-doc, still affiliated with ISPA, I founded the Fish Bioacoustics Lab (https://www.fishbioacoustics.pt/) with Paulo Fonseca (University of Lisbon, Portugal) and have since carried out most research either at the Faculty of Sciences of the University of Lisbon or in situ.

During this early phase of my career, I also realised the multiple challenges associated with studying fish acoustic communication. Triglids, such as the grey (Eutrigla gurnardus) and the streaked (Chelidonichthys lastoviza) gurnards, are coastal-water species found at down to over 150 m depth, imposing limitations to study them in situ. Capturing these fishes for laboratory experiments often requires decompression, since gases in the swimbladder expand when hauled to the surface. Observing the full span of their natural behaviour in laboratory conditions proved to be difficult. For example, triglids did not show reproductive behaviour in captivity—a common context for sound production (Hawkins, 1992). Importantly, I was often faced with the inability to identify the sound producer in a group of interacting fish (Amorim , 2004). Also, fish did not respond to acoustic playbacks (Amorim, 1997), hindering the collection of direct evidence of the function of acoustic signals. These difficulties are not inherent to this fish family but reflect the challenges scientists face to study acoustic communication in fishes (Ladich, 2004), including the mere identification of sound-producing fish species (Looby , 2022; Rice , 2022; Parsons , 2022). In particular, acoustic playback experiments, that have been widely used to show the behavioural significance of sounds in terrestrial animals (McGregor, 1992), have often failed to elicit a response in fishes and, when successful, typically require the presence of an associated visual stimulus (Ladich, 2004; Wackermannova , 2017).

After my PhD, I intended to define relevant questions while choosing adequate model species to pursue investigation in fish bioacoustics. It seemed to me that, although acoustic communication in fishes is more conspicuous during the reproductive season, there was still a considerable lack of knowledge regarding the function of fish mating acoustic signals. Several questions remained largely unanswered:

  • What is the information transmitted in male courtship acoustic signals? Are sounds advertising male quality?

  • Do females use acoustic information during mate choice?

  • If so, does acoustic signalling influence both male and female reproductive success?

  • How are acoustic communication and underlying fitness traits affected in an ever-changing world, for example, by increasing man-made sound or climate change?

  • And finally, which fish models are more suited to address these challenging themes?

Important requisites to choose adequate model species include easiness to distinguish males from females based on external morphology or nuptial colouration. Also, fish should show clear and easy-to-measure behavioural patterns, reproduce well in captive conditions or, if in the field, in places where you can observe them easily. In good model species, the sound emitter should also be readily identified, typically by sound production being associated with clear visual displays. Ideally, these species should also respond to acoustic or video playback. Finally, model species should be easy to obtain or access. Below, I provide a brief description of the fish taxa that helped me answer some of the above questions, highlighting some advantages, but also some disadvantages, as model species to study the role of fish mating acoustic signals (Table S1).

Cichlids make widespread use of acoustic communication during courtship (Ripley and Lobel, 2004; Amorim, 2006). Species, like the Mozambique tilapia, a maternal mouthbrooder, are easily bred in the laboratory (Oliveira and Almada, 1998). When adult fish are placed in a tank, a dominance structure appears within few hours. After 1–5 days, dominant males establish territories, actively dig and defend a pit on the substrate, and start courting nearby females (Amorim , 2003). Males are not only sexually dimorphic (larger bodies and jaws) but exhibit a distinct black colouration from females or subordinate males that depict a silver dull colour (Oliveira and Almada, 1998). They also show conspicuous visual behaviours during territorial defence and courtship that are easily recognisable by the observer (Oliveira and Almada, 1998; Amorim , 2003). Courtship and concurrent sound production is readily elicited when males are allowed to interact with females, for example, after removing a partition that was previously separating them. This can be very helpful, as acoustic recordings require pumps, thermostats, and filters to be switched off to allow a good signal- to-noise ratio (SNR), thus imposing a limited duration to recording sessions. In addition, the Mozambique tilapia is resilient to laboratory conditions. Other cichlids can be used as models to study sound production and behaviour (Ripley and Lobel, 2004; Amorim , 2008; Maruska , 2012). Particularly, cichlids from the great African lakes are exciting model species because of the potential involvement of acoustic communication in species recognition and mate choice, which is hypothesised to have influenced their rapid speciation (Amorim , 2008).

Gobies are ecologically relevant species that are renowned models in bioacoustics and behavioural ecology (Myrberg and Lugli, 2006; Amorim , 2013a; Pedroso , 2013). They mate easily in captivity and, like cichlids, exhibit conspicuous, stereotyped, easy-to-measure behaviours. During the breeding season, males defend nests and attract mates by making visual and acoustic signals from the nest. Females enter the nest and lay eggs on the nest's ceiling, after which they leave all parental care to the male (Amorim , 2013a). Gobies develop nuptial colouration that allows for discrimination between males and females (Amorim and Neves, 2007). In the painted gobies (Pomatoschistus pictus), it is also possible to distinguish different males by the pattern of black dots in the dorsal fin. Sounds can be recorded in relatively short sessions as painted gobies readily show courtship behaviour when males are provided access to females. Because most of the sounds produced by males are within the nest, it is easy to obtain good SNR recordings by placing the hydrophone in a chimney fitted to the nest (Amorim , 2013a).

The Lusitanian toadfish (Halobatrachus didactylus) is a highly soniferous species that produces sounds audible to conspecifics more than 10 m away, even in very shallow waters (2–3 m depth) (Alves , 2016). This contrasts with cichlids and gobies that can only be detected up to three body lengths away (Simões , 2008; Amorim , 2018). Mate attraction relies on boatwhistle production from within the nest (Vasconcelos , 2012) and courtship does not involve visual displays (Brantley and Bass, 1994). This species is an excellent model to test the role of acoustic signalling in mating outcome in the field, as males readily occupy artificial concrete nests and use them to mate (Vasconcelos , 2012; Amorim , 2016). After mating, they stay in the nest, tending the eggs and clinging larvae, while they continue to call to attract more females. The strategic deployment of artificial nests in the lower intertidal area makes them accessible at low spring tides, making it easier to check for occupancy and reproductive success (number of eggs). Importantly, it also allows for recording males individually by placing a hydrophone next to each nest. Unfortunately, Lusitanian toadfish are not good models to study mating behaviour in captivity (personal observation).

The meagre (Argyrosomus regius) and other sciaenids are also notably soniferous and excellent subjects for field studies and can also be studied in the laboratory (Pereira , 2020; Vieira , 2022). Being demersal broadcast spawners (i.e., with no parental care) that form large breeding aggregations, they provide the opportunity to study acoustic communication in a species with a contrasting life strategy to the previous model species. Also, as they are one of the largest, most valuable sciaenids in the world, learning about the relation between calling activity and spawning opens the possibility to use acoustic information for conservation and fisheries management or for aquaculture production methods (Bolgan , 2020). However, like triglids, they do not perform obvious displays during sound production, hindering the identification of the sound emitter in captivity (personal observation).

In species with conventional sex roles, in which males invest in courtship and females in mate choice, females may increase their fitness by choosing a mate that provides direct benefits, such as territories or parental care, that will enhance offspring development and survival (Andersson, 1994). Alternately, females could gain indirect benefits, such as “good genes,” that will increase the genetic fitness for the offspring (Andersson, 1994). While the role of acoustic signals in mate choice is well established in mammals, birds, frogs, and insects (Wilkins , 2013), it remains underappreciated in fish. To date, there is a paucity of data on whether females use male acoustic signals to assess indirect (male quality) or direct benefits, namely parental abilities, to make mate choices in fish. In this context, a first step in my research has been to investigate which information is conveyed by fish mating sounds and then study its use in mating decisions. Possible approaches to test the function of fish sounds in mate choice are association studies, muting experiments, and playback experiments. The sections below are organised by topics, rather than by a chronological order, which can be inferred by publication dates.

Male size, condition, and other features, such as motivation, are important features to secure better territories and other resources and for successful parenting. I hypothesised that the ability to convey this information to females through acoustic signalling is likely widespread in soniferous fishes and investigated this topic with the above models (see Table I for a summary of main findings). As the sound-producing apparatus and associated muscles usually scale with body size, it is expected that larger animals will produce lower frequency and louder sounds (Connaughton , 2000; Fine and Parmentier, 2015; Parmentier and Fine, 2016; Sprague , 2022). Also, body condition likely limits acoustic performance, namely sustained muscle contraction rate (and hence, sustained pulse rate) and calling rate, as they can be energetically demanding (Mitchel , 2008).

TABLE I.

Information content of fish mating signals relevant for mate choice. The table summarises the main findings of acoustic features that are positively (↗), negatively (↘), or not associated (—) to male traits, such as male size, condition (K, COND, lipid content), and motivation. ↗— denotes an initial positive association followed by a plateau. Note that the table is not intended to be exhaustive and mainly summarises examples provided in the text. Common and scientific names are given except for lake Malawi cichlids (genus Maylandia) for which only scientific names are provided. SPL, sound pressure level (dB); PF, peak frequency (Hz); f0–f10, frequency harmonics; PRM, pulse rate modulation within a sound; PR, pulse rate; SD, sound duration (ms); NP, number of pulses in a sound; CR, call rate; CE, calling effort (percentage time spent calling); K, Fulton's condition factor; COND, residuals between male weight and length.

Male trait
Family Species Body size K COND Fat content Motivation Reference
Batrachoididae  Lusitanian toadfish, Halobatrachus didactylus  All acoustic features —  All acoustic features —  SD↗  Mean CR ↗
CE ↗
PR ↗ 
CR ↗  Amorim (2010); Jordão (2012)  
Plainfin midshipman, Porichthys notatus  SPL ↗  f0-f10↗—  f0-f10↗—      Balebail and Sisneros (2022)  
Gobiidae  Painted goby, Pomatoschistus pictus  PF ↘
SPL ↗
PR↘ 
All acoustic features —    Max CR ↗
Mean CR ↗ 
NP↗  Amorim (2013a); Petrisinec (2016)  
Sand goby, P. minutus  PF —
SPL ↗
PR↘ 
All acoustic features —    Max CR ↗
Mean CR ↗ 
  Pedroso (2013)  
African cichlids  Mozambique tilapia, Oreochromis mossambicus  PF ↘  PF↗        Amorim (2003)  
Maylandia “zebra gold”  PF —  PF—        Amorim (2008)  
M. zebra  PF ↘  PF—        Amorim (2008)  
M. callainos  PF ↘  PF—        Amorim (2008)  
M. fainzilberi  PF —  PF—        Amorim (2008)  
M. emmiltos  PF —  PF—        Amorim (2008)  
Pomacentridae  Bicolor damselfish, Stegastes partitus  PF ↘          Myrberg (1993)  
Male trait
Family Species Body size K COND Fat content Motivation Reference
Batrachoididae  Lusitanian toadfish, Halobatrachus didactylus  All acoustic features —  All acoustic features —  SD↗  Mean CR ↗
CE ↗
PR ↗ 
CR ↗  Amorim (2010); Jordão (2012)  
Plainfin midshipman, Porichthys notatus  SPL ↗  f0-f10↗—  f0-f10↗—      Balebail and Sisneros (2022)  
Gobiidae  Painted goby, Pomatoschistus pictus  PF ↘
SPL ↗
PR↘ 
All acoustic features —    Max CR ↗
Mean CR ↗ 
NP↗  Amorim (2013a); Petrisinec (2016)  
Sand goby, P. minutus  PF —
SPL ↗
PR↘ 
All acoustic features —    Max CR ↗
Mean CR ↗ 
  Pedroso (2013)  
African cichlids  Mozambique tilapia, Oreochromis mossambicus  PF ↘  PF↗        Amorim (2003)  
Maylandia “zebra gold”  PF —  PF—        Amorim (2008)  
M. zebra  PF ↘  PF—        Amorim (2008)  
M. callainos  PF ↘  PF—        Amorim (2008)  
M. fainzilberi  PF —  PF—        Amorim (2008)  
M. emmiltos  PF —  PF—        Amorim (2008)  
Pomacentridae  Bicolor damselfish, Stegastes partitus  PF ↘          Myrberg (1993)  

1. Male size

Figure 1 depicts the relation between male size and sound dominant frequency (the frequency with most energy) in model species that I have studied: Batrachoididae–Lusitanian toadfish; Gobiidae–painted goby and sand goby, Pomatoschistus minutus; African cichlids–Mozambique tilapia and lake Malawi cichlids, Maylandia “zebra gold,” Maylandia zebra, Maylandia callainos, Maylandia fainzilberi, and Maylandia emmiltos (Amorim , 2003, 2008, 2010, 2013a; Pedroso , 2013). Results from the bicolor damselfish, Stegastes partitus (Myrberg , 1993), are included as a reference for a known frequency-size dependence [Fig. 1(A)]. In Lusitanian toadfish, in the sand goby, and in the lake Malawi cichlids Maylandia “zebra gold,” M. fainzilberi, and M. emmiltos, no relation was found (Pearson correlation, P > 0.05). In the remaining species, the correlation between size and sound frequency was significant but only the bicolor damselfish presented a steep slope, and only bicolor damselfish (R2 = 0.87), Mozambique tilapia (R2 = 0.88), and the lake Malawi cichlid M. zebra (R2 = 0.60) presented a R2 higher than 0.5. Figure 1(B) highlights the lack of relationship between both the fundamental (muscle contraction rate) and the dominant frequency of mating boatwhistles with male size in the Lusitanian toadfish. It also highlights the invariance in muscle contraction rate (determined by the firing rate of central pattern generators) with male length, which contrasts with a more variable dominant frequency that can be represented by the fundamental frequency or the first or second harmonics (Amorim and Vasconcelos, 2008).

FIG. 1.

(Color online) Relation between fish size and peak (or dominant) frequency of mating signals in species with (A) slow and (B) fast pulse rate sounds. Slow pulse rate sounds include mating acoustic signals from gobies (Pomatoschistus pictus, P. minutus), African cichlids (Oreochomis mossambicus, Maylandia “zebra gold,” M. zebra, M. callainos, M. fainzilberi, M. emmiltos), pomacentrids (Stegastes partitus), while advertisement boatwhistles made by batrachoidids (Halobatrachus didactylus) have a fast pulse rate (Amorim , 2003; Amorim , 2008; Amorim , 2010; Amorim , 2013a; Pedroso , 2013; Myrberg , 1993). PF, peak frequency; PF2, second peak frequency around 450 Hz; F0, fundamental frequency.

FIG. 1.

(Color online) Relation between fish size and peak (or dominant) frequency of mating signals in species with (A) slow and (B) fast pulse rate sounds. Slow pulse rate sounds include mating acoustic signals from gobies (Pomatoschistus pictus, P. minutus), African cichlids (Oreochomis mossambicus, Maylandia “zebra gold,” M. zebra, M. callainos, M. fainzilberi, M. emmiltos), pomacentrids (Stegastes partitus), while advertisement boatwhistles made by batrachoidids (Halobatrachus didactylus) have a fast pulse rate (Amorim , 2003; Amorim , 2008; Amorim , 2010; Amorim , 2013a; Pedroso , 2013; Myrberg , 1993). PF, peak frequency; PF2, second peak frequency around 450 Hz; F0, fundamental frequency.

Close modal

This lack of relationship between mating sound spectral properties and male size is expected in the Lusitanian toadfish (Amorim , 2010). In species that produce sounds by contracting sonic muscles at a high rate (50–300 Hz), the contraction rate dictates the sound fundamental frequency (and overall spectral features). Fish size has little effect on this feature, as it mainly depends on muscle twitch parameters (Parmentier and Fine, 2016; Parmentier , 2019). However, fishes with a slow pulse rate [i.e., with pulsed sounds; all species in Fig. 1(A)], may show an inverse steep relationship between dominant frequency and fish size (Parmentier and Fine, 2016). The strength of this relationship is likely related to the variations in scaling of the sonic muscles with fish growth (larger fish produce longer twitches and thus, lower frequency sounds), and with the lack of a sharp frequency tuning (Fine and Parmentier, 2015). Most fish sounds have a wide, rather than a tuned, frequency spectrum, which makes this spectral parameter less reliable and less objective to measure. Considering that some fish are capable of resolving differences in frequency by approximately 10% (Fay, 1988), in species like the bicolor damselfish, the Mozambique tilapia, and the lake Malawi cichlid (M. zebra), that present a tight relation (higher R2 and steep slopes) with male size in adults, dominant frequency could be a reliable indicator of size.

Sound amplitude can also be a predictor of body size, though measured less often because many fish make sounds while moving (Amorim , 2015; Parmentier , 2021). We measured sound pressure level (SPL) (dB) in both painted and sand gobies, in males making mating drums while stationary in the nest (Amorim , 2013a; Pedroso , 2013). In both species, SPL was a significant predictor of body size: painted goby (R2 = 0.42); sand goby (R2 = 0.36) (Fig. 2). In both species, a variation of 1 cm in standard length (SL) corresponds to an expected change of 24 dB in sound amplitude (Amorim , 2013a; Pedroso , 2013). For comparison, larger plainfin midshipman, Porichthys notatus (Batrochoididae), also produce mating hums from their nest with higher amplitude (R2 = 0.60), but a variation of 1 cm in male SL only increases on average 1 dB in SPL (Balebail and Sisneros, 2022). These differences suggest contrasting scaling effects in the sound production systems of gobies and batrachoidids: pectoral-girdle–based and swimbladder-based mechanisms (Cohen and Winn, 1967; Parmentier , 2017). Interestingly, some fishes seem to be able to adjust sound amplitude. The blacktail shiner, Cyprinella venusta (Cyprinidae) (Holt and Johnston, 2014) and two batrachoidids—the oyster toadfish (Luczkovich , 2016) and the plainfin midshipman (Brown , 2021)—have been shown to increase the amplitude of their sounds in the presence of noise (known as Lombard effect). This could be achieved, for example, by changing swimbladder volume, consistent with plainfin midshipman males inflating the swimbladder when humming to entice females to their nests, likely to enhance sound amplitude and attractiveness (Bass , 2015). As sound amplitude attenuates rapidly, this parameter should be more relevant for mate choice when a female is already in close proximity to the male, though louder sounds will still increase the chances of mate detection and attraction.

FIG. 2.

(Color online) Relation between fish size and sound pressure level (SPL) of mating signals in the painted goby Pomatoschistus pictus and in the sand goby P. minutus (Amorim , 2013a; Pedroso , 2013).

FIG. 2.

(Color online) Relation between fish size and sound pressure level (SPL) of mating signals in the painted goby Pomatoschistus pictus and in the sand goby P. minutus (Amorim , 2013a; Pedroso , 2013).

Close modal

Fish size can be further inferred from mating sounds by pulse rate modulation (PRM), i.e., the decrease in pulse emission rate or increase in pulse period throughout a sound, named fatigue resistance in Amorim (2013a). We only investigated this parameter in gobies. In the painted goby, PRM was the best predictor of male size after SPL, accounting for 45% of the variability in male SL (Amorim , 2013a). PRM was not related to male condition in the painted goby, indicating it mainly depends on body scaling and is therefore a good indicator of male size. The increase in pulse period in a mating drum is however unlikely to be resolved by the goby auditory system (Amorim , 2018). Interestingly, no such relation was found in the sand goby, despite their higher drum pulse rate (Pedroso , 2013), suggesting interspecific differences in physiological constraints of sound production. Other sound features may vary with fish size. For example, we found that the number of pulses of mating drums increases with male length in the painted goby but this feature also changes with male condition (Amorim , 2013a) and is highly dependent on motivation.

2. Male condition

Male condition is a phenotypic quality important in mate choice. A number of studies have related mating sound features with male condition, such as the Fulton's condition factor (K, ratio between body weight and cubed standard length) or the residuals of the regression between male weight and length (COND) (e.g., Amorim and Neves, 2007; Amorim , 2010). We found a significant positive correlation between dominant frequency and male K in the Mozambique tilapia (R2 = 45.7%) (and not with any other measured sound feature; Amorim , 2003) but not in lake Malawi cichlids (Amorim , 2008). Likewise, we did not find a correlation between any acoustic parameter and male COND in the Lusitanian toadfish (Amorim , 2010). In the painted and the sand gobies, regression models showed that male K was not predicted by any acoustic trait (Amorim , 2010, 2013a; Pedroso , 2013). Interestingly, a recent study has shown that harmonic frequencies (f0–f10) of the tonal advertisement hum of the plainfin midshipman are predictors of body condition (K and COND), with males with higher body condition producing hums with higher harmonic frequencies (Balebail and Sisneros, 2022). The above take into consideration nonlethal proxies of fish nutritional state that assume that fish in a better condition are heavier for a given length. However, these metrics may be poorly correlated with physiological condition and may not describe adequately fat content (Sutton , 2000).

We further investigated the relationship between sound production and male fat content in gobies and the Lusitanian toadfish (Fig. 3). Although male K was not correlated with any acoustic feature in the painted and sand gobies, acoustic activity (maximum drumming rate and mean drumming rate considering only the vocally active minutes, respectively) was a good predictor of male body fat in both species (Amorim , 2013a; Pedroso , 2013) [Fig. 3(A)]. In the Lusitanian toadfish, male lipid content was also strongly related to mean calling rate and calling effort (percentage time spent calling) (Amorim , 2010) [Fig. 3(B)]. Contrasting results were found for the plainfin midshipman where body condition (K and COND), or size, did not present a relationship with calling activity, perhaps because its mating sound is much longer than in other batrachoidids or gobies (on the order of minutes to hours instead of milliseconds or seconds), thus hindering direct mate assessment by females (Balebail and Sisneros, 2022).

FIG. 3.

(Color online) Relation between body lipid content and (A) calling rate in the gobies Pomatoschistus pictus and P. minutus and with (B) calling effort and (C) pulse rate in the Lusitanian toadfish Halobatrachus didactylus. Calling activity data concerns maximum drumming rate per minute for the painted goby, active drumming rate (mean drumming rate in the minutes when there was calling activity) for the sand goby, and calling effort (percentage of time spent calling at the hour level) for the Lusitanian toadfish. Recording sessions were 20 min for painted and sand gobies, but up to a fortnight for Lusitanian toadfish. Lipid content is expressed relative to 100 g of fresh tissue (Amorim , 2010; Amorim , 2013a; Pedroso , 2013).

FIG. 3.

(Color online) Relation between body lipid content and (A) calling rate in the gobies Pomatoschistus pictus and P. minutus and with (B) calling effort and (C) pulse rate in the Lusitanian toadfish Halobatrachus didactylus. Calling activity data concerns maximum drumming rate per minute for the painted goby, active drumming rate (mean drumming rate in the minutes when there was calling activity) for the sand goby, and calling effort (percentage of time spent calling at the hour level) for the Lusitanian toadfish. Recording sessions were 20 min for painted and sand gobies, but up to a fortnight for Lusitanian toadfish. Lipid content is expressed relative to 100 g of fresh tissue (Amorim , 2010; Amorim , 2013a; Pedroso , 2013).

Close modal

Because sonic muscle contraction is determined by the firing rate of a central pattern generator in the hindbrain (Bass , 2015), finding that boatwhistle pulse period or pulse rate (the inverse of pulse period) presents low intra- and interindividual variability in the Lusitanian toadfish did not come as a surprise (Amorim and Vasconcelos, 2008; Amorim , 2010; Amorim , 2011). However, an exciting result from field recordings of Lusitanian toadfish males was that male fat content was a predictor of faster pulse rate, i.e., shorter pulse periods [Fig. 3(C)] (Amorim , 2010). Consistent with these findings, Balebail and Sisneros (2022) found that that the fundamental frequency (hence, the pulse rate) of the plainfin midshipman hum increased asymptoticly with male COND up to a threshold and plateaued at higher body condition. Together, these studies suggest that males from both batrachoidid species may be advertising their neuromuscular performance to females by sustaining sonic muscle contraction close to their physiological limit. These shifts in fundamental frequency of the mating call translate into higher frequency harmonics that could be better detected and localised by females in shallow waters where low frequencies do not propagate far (Alves , 2016). Pushing the limits of neuromuscular performance in courtship displays used in mate choice is widespread in vertebrates (Schwark , 2022). In songbirds, for example, males that approach the limit of vocal performance constraints in terms of syllable repetition rate and frequency bandwidth are preferred by females (Podos, 2001; Goller, 2022).

3. Other messages

Sound production can also signal motivation, such as readiness to mate. Motivation can be signalled by increasing calling rate or increasing the number of pulses in a sound. For example, in Lusitanian toadfish and other batrachoidids, the calling rate of nesting males is influenced by the acoustic behaviour of neighbours (Winn, 1967; Fish, 1972; Jordão , 2012; Vieira , 2021a). We found that when engaged in sustained calling, Lusitanian toadfish males tend to match their neighbours' calling rate, seemingly to avoid calling at a rate higher than necessary (Jordão , 2012), while also avoiding call overlap (Vieira , 2021a). This could be a strategy to maximise mate attraction without incurring unnecessary costs. Indeed, calling rate is known to increase in the presence of a prospective mate in batrachoidids (Gray and Winn, 1961).

We also investigated changes of calling behaviour in the painted goby in relation to different stages of mating (Petrisinec, 2016): (1) when both male and female were still outside the nest; (2) when the male was inside the nest but the female was still outside; (3) when both male and female were inside the nest; (4) similar to the previous stage, but the female was rolling in the nest, likely spawning. In this study sounds were not detected in stage 1. Painted goby males produced most thumps in stage 2, when the female was still outside the nest, and mainly drummed when the female was inside the nest [Fig. 4(A)]. This suggests that thumps serve in mate attraction, whereas drums are courtship signals, which are typically emitted in close range and contain features relevant for mate assessment. Notably, drums emitted during more advanced courtship (stages 3 and 4) had a higher number of pulses than the drums produced when the female was outside the nest (stage 2) [Fig. 4(B)]. This suggests that the number of pulses in a drum signals male readiness to mate and is a feature likely relevant to mate choice.

FIG. 4.

(Color online) Relation between (A) number of thumps and drums and (B) number of pulses in a drum observed in different courtship stages in the painted goby Pomatoschistus pictus. In stage 2, the male is inside the nest and the female is still outside; in stage 3, both male and female are inside the nest; in stage 4, the couple is inside but the female is rolling in the nest, likely spawning. Data concern 17 males recorded in 20 min male–female interactions (Petrisinec, 2016).

FIG. 4.

(Color online) Relation between (A) number of thumps and drums and (B) number of pulses in a drum observed in different courtship stages in the painted goby Pomatoschistus pictus. In stage 2, the male is inside the nest and the female is still outside; in stage 3, both male and female are inside the nest; in stage 4, the couple is inside but the female is rolling in the nest, likely spawning. Data concern 17 males recorded in 20 min male–female interactions (Petrisinec, 2016).

Close modal

The link of acoustic communication with other aspects of male quality, such as disease resistance, an individual's developmental trajectory, sperm quality, ability to find food, or escape predators, has not been studied thus far.

If the above acoustic features (sound frequency, SPL, number of pulses in a sound, pulse rate, PRM, and calling rate) are used by females in mate choice, then they should relate to fish reproductive success. Table II summarises the main findings of the studies that addressed the role of acoustic communication in fish reproductive success.

TABLE II.

Association of acoustic activity and acoustic features of mating signals with reproductive success. Note that although acoustic activity has been linked to spawning in other fishes, such as sciaenids and epinephelids, the relationship between acoustic features or calling rate and spawning success remains to be tested in these groups. CR, call rate; CE, calling effort (percentage time spent calling); SD, sound duration (ms); NP, number of pulses in a sound.

Family Species Mating success Clutch size % of males that mated Setup Observations Reference
Batrachoididae  Lusitanian toadfish, Halobatrachus didactylus  Mean CR
Max CR 
Max CR
CE 
25  Field  Mesh around nests restraining passage to males and large females  Vasconcelos (2012)  
  COND  84.8  Field  Open nests; easy entrance  Amorim (2016)  
  Max CR
Mean CR
CE 
30  Field  Mesh around nests restraining passage to males and large females; opening slightly larger than in Vasconcelos (2012); data for control fish in muting experiments  Amorim (2016)  
Gobiidae  Painted goby, Pomatoschistus pictus  Mean CR
CE
NP (and SD) 
  52.6  Laboratory  Single-choice experiment  Amorim (2013a)  
CR    69  Laboratory  Single-choice experiment.
Data for control fish in added sound exposure experiment 
De Jong (2018a); De Jong (2018b)  
Two spotted goby,
P. flavescens 
CR  CR  48.7  Laboratory  Single-choice experiment.
Data for two temperatures (16 °C and 20 °C) 
Albouy (2023)  
Pomacentridae  Hawaiian domino damselfish, Dascyllus albisella    CR    Field  Reproduction success was tallied as no. of egg clutches per cycle  Oliver and Lobel (2013)  
Gadidae  Pollack, Pollachius pollachius    CR    Laboratory  Reproduction success was tallied as total egg and fertilized daily egg production  Wilson (2014)  
Atlantic cod Gadus morhua    CR    Laboratory  Reproduction success was tallied as volume of eggs  Rowe and Hutchings (2006)  
Family Species Mating success Clutch size % of males that mated Setup Observations Reference
Batrachoididae  Lusitanian toadfish, Halobatrachus didactylus  Mean CR
Max CR 
Max CR
CE 
25  Field  Mesh around nests restraining passage to males and large females  Vasconcelos (2012)  
  COND  84.8  Field  Open nests; easy entrance  Amorim (2016)  
  Max CR
Mean CR
CE 
30  Field  Mesh around nests restraining passage to males and large females; opening slightly larger than in Vasconcelos (2012); data for control fish in muting experiments  Amorim (2016)  
Gobiidae  Painted goby, Pomatoschistus pictus  Mean CR
CE
NP (and SD) 
  52.6  Laboratory  Single-choice experiment  Amorim (2013a)  
CR    69  Laboratory  Single-choice experiment.
Data for control fish in added sound exposure experiment 
De Jong (2018a); De Jong (2018b)  
Two spotted goby,
P. flavescens 
CR  CR  48.7  Laboratory  Single-choice experiment.
Data for two temperatures (16 °C and 20 °C) 
Albouy (2023)  
Pomacentridae  Hawaiian domino damselfish, Dascyllus albisella    CR    Field  Reproduction success was tallied as no. of egg clutches per cycle  Oliver and Lobel (2013)  
Gadidae  Pollack, Pollachius pollachius    CR    Laboratory  Reproduction success was tallied as total egg and fertilized daily egg production  Wilson (2014)  
Atlantic cod Gadus morhua    CR    Laboratory  Reproduction success was tallied as volume of eggs  Rowe and Hutchings (2006)  

1. The Lusitanian toadfish

We carried out in situ the first study addressing the relationship between male acoustic signalling and reproductive success using the Lusitanian toadfish as a model species (Vasconcelos , 2012). Males that spontaneously occupied artificial concrete shelters in the lower intertidal within the Tagus estuary (Portugal) were acoustically monitored individually for up to 2 weeks (N = 56). During recordings, the entrance of the nests was covered with a plastic mesh with an opening that prevented the large nest-holder males from exiting but allowed smaller females, sneaker males, and prey items to enter. After this period, the nests were inspected during low spring tides for the presence and number of eggs, a proxy for reproductive success. Males in nests with egg clutches presented a significantly higher calling activity [mean and maximum calling rate per hour and mean calling effort; Fig. 5(A)] and lower boatwhistle dominant frequency. From these features, only the maximum calling rate and calling effort were good predictors of the number of eggs found in the nest, explaining 52% and 6% of clutch size variability, respectively.

FIG. 5.

(Color online) Comparison of calling activity between males with and without clutches in the nests of (A) Lusitanian toadfish Halobatrachus didactylus, (B) painted goby Pomatoschistus pictus, (C) two spotted goby P. flavescens. Calling effort is the percentage of time spent calling at the hour level. Drum rate is the mean number of sounds per min. Recording sessions were 20 min for the painted goby, 3 × 30 min for the two spotted goby, and up to a fortnight for the Lusitanian toadfish (Vasconcelos , 2012; Amorim , 2013a; Albouy , 2023).

FIG. 5.

(Color online) Comparison of calling activity between males with and without clutches in the nests of (A) Lusitanian toadfish Halobatrachus didactylus, (B) painted goby Pomatoschistus pictus, (C) two spotted goby P. flavescens. Calling effort is the percentage of time spent calling at the hour level. Drum rate is the mean number of sounds per min. Recording sessions were 20 min for the painted goby, 3 × 30 min for the two spotted goby, and up to a fortnight for the Lusitanian toadfish (Vasconcelos , 2012; Amorim , 2013a; Albouy , 2023).

Close modal

We further tested whether acoustic activity and several measures of male quality, including circulating steroid levels, related to male reproduction success. This was done in a follow-up experiment with a similar, but more natural setup, where males (N = 33) could move freely in the breeding area (Amorim , 2016). This experiment showed with paternity analysis that the degree of cuckoldry was low and that clutch size was a reliable measure of reproductive success. In contrast to the earlier study, clutch size was not related to male acoustic activity. Instead, it was positively related to male condition (COND) and negatively with testosterone, which tends to decrease throughout the breeding season (Modesto , 2003). The contrast in the relevance of calling activity to reproduction outcome with the previous findings (Vasconcelos , 2012) could relate to an erroneous attribution of vocal activity to subject males in the open-nest experiment due to nest takeovers. An alternative explanation could be linked with the contrasting percentage of males that succeeded in mating in the two experimental setups: 25% in the restrained nests (Vasconcelos , 2012) vs 84.8% in the open nests (Amorim , 2016). This difference suggests that females do not enter the restrained nests so easily or do not find them attractive. In addition, the small nest openings in the early study (Vasconcelos , 2012) possibly also influenced mate choice decisions by increasing the costs of sampling for males if going through the opening was difficult.

To confirm the role of acoustic activity in the mating success of the Lusitanian toadfish, we made an additional muting field experiment with three treatments: muted (with the swimbladder deflated by cutting a small portion of its wall), sham operated and unmanipulated (Amorim , 2016). Here, we restrained males in the nests, as in Vasconcelos (2012), but left a slightly larger opening. In this, as in the previous experiments, silent males failed to mate, but 30% of the vocal males obtained clutches, comparable to the success rate observed in Vasconcelos (2012). As in the study by Vasconcelos (2012), maximum calling rate was a predictor of the number of eggs in the nest. Taken together, this set of field experiments suggests that calling is required for mating in the Lusitanian toadfish as silent males failed to mate. High calling activity, especially maximum calling rate (signalling male condition), appears to be key for reproductive success although additional factors seem to be also at play. These could include mate sampling costs (easiness to access different nests), or non-auditory sensory cues used while inside the nest, such as mechanosensory or chemical.

2. Gobies

Unlike batrachoidids, gobies have nuptial colours and elaborate courtship visual displays, suggesting that visual communication should be relevant in mate choice and that acoustic signalling could play, perhaps, a minor role. We investigated this question in the painted goby in a single-choice experiment where ripe females interacted with a single male but could still decide whether to mate or not, i.e., delay spawning (Amorim , 2013a). As in Lusitanian toadfish, silent painted goby males failed to mate successfully, suggesting that in this species, acoustic communication is also required for mating. From the soniferous males, approximately half (10 out of 19) succeeded in obtaining clutches in their nest. Successful males showed a significantly higher calling activity [higher mean drumming rate and calling effort; Fig. 5(B)], made longer sounds with longer pulse periods, and had a higher fat body content than unsuccessful males. Surprisingly, neither visual courtship rate nor male size differed between these two groups of males.

A subsequent study supported the relevance of acoustic courtship in mating in this species. While studying the effect of added noise on multimodal communication in the painted goby in similar one-choice experiments, we found that the likelihood of successful spawning was predominantly correlated with male acoustic courtship in the control treatment (i.e., no added noise) (de Jong , 2018a). However, when exposed to masking noise, visual courtship becomes more relevant in determining spawning events, suggesting visual behaviour as a backup sensory channel when acoustic communication is compromised.

We recently investigated the role of acoustic communication in the reproductive success of the two spotted goby Pomatoschistus flavescens and found congruent results (Albouy , 2023). This species uses both visual and acoustic signals and emits similar sound types to the painted goby (de Jong , 2018b). In the two spotted goby, drumming rate was the only predictor of spawning success and was higher in males that obtained clutches [Fig. 5(C)].

Although the link between acoustic signalling and reproductive outcome was not tested in the sand goby, Pedroso (2013) also observed that silent males did not mate successfully.

Together, these studies point to a high reliance on acoustic activity for mate choice, in both the Lusitanian toadfish and gobies, which seems to be informative of male fat reserves. Male condition is a particularly relevant trait for female mate choice in fish species with paternal care, as it is linked with paternal abilities and associated higher egg-hatching success (e.g., Knapp and Kovach, 1991; Lindström , 2006; Sisneros , 2009). Whether acoustic signalling advertises male paternal abilities remains untested in fish. Although it is thought that acoustic communication is important for reproductive success in fish (Parmentier and Fine, 2016), few other studies have also shown this link (see below).

3. Other model fish species

In a field work, Oliver and Lobel (2013) observed several reproductive cycles in the Hawaiian domino damselfish, Dascyllus albisella and demonstrated that male reproductive success, measured as the number of egg clutches per cycle, was correlated with visual and acoustic (concurrent) courtship rate. Although these authors did not measure fat reserves in the studied males (N = 26), a previous study linked courtship rate with fat reserves in this species (Knapp and Kovach, 1991) suggesting that females are selecting condition dependent traits that advertise paternal care abilities.

In Gadidae, Wilson (2014) showed that in the pollack, Pollachius pollachius, grunt sound production was significantly correlated with both total egg and fertilized daily egg production. Rowe and Hutchings (2006) also found that grunt production was significantly associated with the volume of eggs produced in the Atlantic cod Gadus morhua. Grunt production was associated with both agonistic and courtship behaviour in cod and agonistic interactions often preceded mounting attempts, indicating that intrasexual competition is a strong component of the mating system (Rowe and Hutchings, 2006). In a follow-up study carried out in a large mesocosm, Rowe and Hutchings (2008) showed that reproduction in Atlantic cod is largely skewed and that a significant correlate of male mating success (number of ventral mounts initiated by a male) was the mass of the drumming muscles, supporting the relationship between sound production and mating success in this species.

Acoustic activity, such as chorusing, has been linked to spawning in sciaenids (e.g., field: Mok and Gilmore, 1983; Luczkovich , 1999; captivity: Montie , 2016; Montie , 2017; Vieira , 2019; Bolgan , 2020) and in groupers (epinephelids) (field: Koenig , 2017, Rowell , 2019) through concurrent appearance of eggs and larvae in the water with recordings of fish calls. However, the relationship between acoustic features, male quality, mate choice, and spawning success (offspring number) remains to be tested in these groups. Sciaenids and epinephelids are large fishes, spawn in large aggregations, often in the dark, and are broadcast spawners releasing milt and eggs in the water column, characteristics that constrain both field and laboratory studies. One such example is the meagre, that can attain up to 2 m and breed in large aggregations at dusk, in murky estuarine waters (Vieira , 2022). While it is possible to breed this species in captivity, it is difficult to keep them in large aggregations or provide space to allow their full behavioural repertoire (Pereira , 2020).

Early playback studies of conspecific courtship calls have shown an attraction function (Gerald, 1971; Lugli , 1996; Myrberg , 1986; McKibben and Bass, 1998), preference for soniferous over silent males (Tavolga, 1956; Myrberg and Stadler, 2002), or a facilitation of courtship (Myrberg and Spires, 1972). Playback of courtship sounds influence affiliation preference in African cichlids when acoustic and visual signals are decoupled in time, suggesting that information on the presence and breeding motivation of a nearby conspecific male may influence female mate preference before visual contact (Verzijden , 2010; Maruska , 2012).

Perhaps the first evidence of the use of acoustic signals in mate choice was provided by the seminal field experiment in which Myrberg (1986) showed that females of the bicolor damselfish were preferentially attracted to played-back courtship chirps of lower dominant frequency—a characteristic of larger males (Table III). In another landmark study, McKibben and Bass (1998) demonstrated that the plainfin midshipman females show a robust phonotactic response to the playback of mating hums, differentiating between hum-like signals differing in fundamental frequency, intensity, duration, amplitude modulation, and fine temporal pattern. These authors showed that gravid females preferred the more intense of two signals differing by 3 dB (McKibben and Bass, 1998), an acoustic feature that is linked with male size (Balebail and Sisneros, 2022). As larger males father more offspring (Sisneros , 2009), it is likely that midshipman females are using male acoustic signals for mate choice (Table III).

TABLE III.

Role of fish acoustic signals in mate choice. The table summarises the main findings but is not intended to be exhaustive. Note that only one study was carried out in the field (Myrberg , 1986). PBK, playback experiment; CR, call rate; PF, peak frequency.

Family Species PBK test Visual stimuli Outcome Observations Reference
Gobiidae  Painted goby, Pomatoschistus pictus  Sound vs control  Absent  No preference  Mating sounds were played back at a medium CR; control was silence or white noise  Amorim (2013b);
Amorim (2013b)  
Sound vs control  Present  Preference for males associated with sound  Males and female separated by a clear partition 
High CR vs low CR  Present  No preference  Males and female separated by a clear partition  Vicente (2013); unpublished data 
High CR vs low CR  Present  No preference  Females has access to male compartmentbut male is in a jar near the nest 
High CR vs low CR  Present  Preference for males associated with high CR  Females has full access to the territorial male 
Pomacentridae  Bicolor damselfish, Stegastes partitus  High PF vs low PF  Absent  Preference for males with low PF, indicative of their larger size  The experiment used only the sounds of two males; field experiment  Myrberg (1986)  
Batrachoididae  Plainfin midshipman, Porichthys notatus  Sounds differing by 3 dB  Absent  Preference for louder signals  Two-choice tests for several stimuli pairs differing by 3 dB; other tests are reported in this study  McKibben and Bass (1998)  
Family Species PBK test Visual stimuli Outcome Observations Reference
Gobiidae  Painted goby, Pomatoschistus pictus  Sound vs control  Absent  No preference  Mating sounds were played back at a medium CR; control was silence or white noise  Amorim (2013b);
Amorim (2013b)  
Sound vs control  Present  Preference for males associated with sound  Males and female separated by a clear partition 
High CR vs low CR  Present  No preference  Males and female separated by a clear partition  Vicente (2013); unpublished data 
High CR vs low CR  Present  No preference  Females has access to male compartmentbut male is in a jar near the nest 
High CR vs low CR  Present  Preference for males associated with high CR  Females has full access to the territorial male 
Pomacentridae  Bicolor damselfish, Stegastes partitus  High PF vs low PF  Absent  Preference for males with low PF, indicative of their larger size  The experiment used only the sounds of two males; field experiment  Myrberg (1986)  
Batrachoididae  Plainfin midshipman, Porichthys notatus  Sounds differing by 3 dB  Absent  Preference for louder signals  Two-choice tests for several stimuli pairs differing by 3 dB; other tests are reported in this study  McKibben and Bass (1998)  

While the above studies have pointed towards a significant role of mating signals, much was left unanswered. We have carried out a suite of dichotomous-choice playback studies in the painted goby to investigate the use of acoustic signals in mate choice (Table III). We showed that females do not prefer conspecific sound over a control (silence or white noise) (Amorim , 2013b). However, when sound was combined with visual access of a territorial male, females preferred to affiliate with the male associated with the courtship sound playback. These results support the relevance of acoustic courtship but only when associated with visual cues. However, the question whether painted goby females use calling rate to evaluate male condition needed further investigation. We carried out three playback experiments (following Amorim , 2013b), in which painted goby females were allowed to choose between two matched size males associated with the playback of either a high or a low drumming rate (8 vs 2 drums min−1) (Vicente, 2013; unpublished data). In experiment 1, transparent partitions separated the female from the male (as in Amorim , 2013b). In experiment 2, the partition was removed, allowing the female access to the male compartment, but they could not interact freely as the male and his nest were inside a jar. In the third experiment, the partition was removed, and fish could interact freely. Affiliation time did not differ between high and low drumming rate males in experiments 1 and 2, but in experiment 3, females preferred the males associated with the high drumming rate. This set of experiments suggests that drumming rate is important in mate choice in this species but it needs to be associated with the full suite of courtship stimuli; visual signals, plus likely chemosensory, mechanosensory, and tactile cues that are exchanged mostly inside the nest. The findings warrant for future studies addressing the relative role of different sensory modalities in mate choice, especially in late courtship stages near spawning.

The above examples with painted gobies clearly showed that acoustic courtship is essential for mating, but mating decisions require other sensory components. Even in batrachoidids, which are mostly reliant on acoustic communication, there seems to be other factors at play. For example, just changing the size of the male's nest opening altered the association between male calling activity and reproductive success in the Lusitanian toadfish. It thus seems that acoustic signals are a part of a more complex multimodal system that still needs to be unveiled.

Multimodal communication is classically viewed to provide back-up messages or convey multiple meanings (Halfwerk , 2019). For example, acoustic signals associated with visual displays could both inform on a male's intention to mate (if one sensory channel would be compromised, the message could still be conveyed through the other channel), or convey different information at the same time, such as species identity and intention to mate. However, the sensory integration of multimodal signals by the receiver can give rise to a multimodal percept involving unique, smaller, or larger responses when compared to unimodal stimuli (Halfwerk , 2019). This means that receivers can gain additional information that is only available in the combined signal components. In addition, sensory integration of multimodal mating systems is affected by the ecological and social scenario with the weight of different sensory modalities changing according to context (Amorim , 2019). In the future. we need to take into consideration factors such as habitat's transmission properties, distance between signallers and receivers, predation risk, physiological state of the sender and the receiver, and number of male competitors and of sneaker males. Advances in this area will require fish models that respond to manipulated signals, such as acoustic playbacks (McGregor, 1992) or robotic fish (Bierbach , 2018).

We still know little about the link of acoustic communication with mate choice and reproductive success and, as reviewed above, this has only been investigated in a few polygamic fish species with paternal care. However, fishes have an astounding diversity in reproductive models and mating systems (DeWoody and Avise, 2001). Strategies can range from monogamy, sequential polygamy, obligate trio spawning to group spawning, and involve different parental care strategies (internal brooding in pouches, such as in seahorses, maternal or paternal mouthbrooding, such as in cichlids, and mono- or biparental care) or no parental care. Such a range of mating and parental care systems may have given rise to different acoustic communication strategies. From the studies described above, acoustic indicators of male size and mainly male condition (such as calling rate) seem to play a role in reproductive success in fishes with paternal care. Yet, do these acoustic features also drive reproductive success in species without paternal care?

For example, some broadcaster spawners, such as sciaenids and epinephelids (and many others) also rely heavily on acoustic communication for mating as they form large and loud chorus aggregations. Is the dependence on acoustic signalling in broadcast spawners similar to the one found in nesting fishes? Do they use sounds for mate choice or does sound only serve to form and maintain spawning aggregations? One could argue that individual male sounds have a limited role in mate choice as they are likely masked by the loud chorus in spawning aggregations (Lobel , 2010). Instead, could chorusing function to modulate reproductive hormones to coordinate male and female mating behaviour and synchronise milt and egg release? A recent study has shown that in the blacktail shiners, a freshwater fish where males defend territories for reproduction, male sounds contribute to the onset of spawning behaviours in female (Crovo , 2022), suggesting mating sounds could prime and coordinate mating behaviour in fishes. Thus, it is plausible that chorusing in large fish aggregations could serve a similar purpose—a hypothesis that deserves investigation.

Several authors have pleaded for the need to create a global library of fish sounds and increase the inventory of soniferous species (Rountree , 2020; Parsons , 2022; Looby , 2023). The argument is that the documentation of sounds and their sources is needed to have an adequate appreciation of aquatic soundscapes. Fish sounds provide significant knowledge about fish acoustic communities, biodiversity, and habitat health (e.g., Carriço , 2020) and the spatiotemporal patterns of spawning events (Luczcovitch , 1999; Rowell , 2019). They are also key to characterizing responses to environmental drivers, such as temperature, salinity, lunar phase, tide, and time of sunset (e.g., Vieira , 2022), climate change and extreme weather events (Gordon , 2018; Boyd , 2021), and anthropogenic noise (e.g., Vieira , 2021b), or map the spreading of invasive species (Amorim , 2023). While even unidentified fish sounds can provide information on local communities and allow biodiversity assessment, known sound sources are needed to assess them individually or to understand their contribution to the overall soundscape (Carriço , 2020; Mooney , 2020). Passive acoustic monitoring (PAM) of known fish sounds could thus contribute to more effective conservation and fisheries management (Parsons , 2022). Increasing the documentation of mating sounds could bring the information provided by PAM one step forward as it refines the assessment of spatiotemporal patterns of fish spawning patterns, population state, and the disruption of reproduction due to anthropogenic activities and global changes. It could also facilitate monitoring the performance of marine protected areas (La Manna , 2021).

The increase in noise pollution underwater resulting from human activities has been the focus of a growing number of studies, showing it can have detrimental effects on fitness (Hawkins and Popper, 2018; de Jong , 2020). For example, in situ experiments with the Lusitanian toadfish showed that boat noise playback depressed male vocal activity and impacted reproductive success by decreasing the likelihood of mating, clutch size, and larvae development (Amorim , 2022; Faria , 2022). Vieira (2021b) also showed that sound from passing boats caused significant masking (assessed through auditory evoked potentials) and depressed calling activity in the meagre in the wild. While it is argued that pelagic fish can escape sound sources (unlike nest-holder toadfishes), they may still decide to stay in disturbed areas following the “best-of-a-bad-job” strategy if they lack alternatives or if nearby places are not suitable to breed; this seems to be the case in the brown meagre, Sciaena umbra, that prefers to spawn in the noisiest areas of the Venice tidal inlets (Picciulin , 2021). Although the effect of man-made sound on fishes has received some attention (Popper and Hawkins, 2019; de Jong , 2020; Amorim , 2022), there is a great need for more studies, especially carried out in situ, addressing the impact of man-made underwater sound pollution on soniferous fishes (and fishes in general) and how it impacts mate choice and mating decisions.

Another major concern is the increase in ocean temperature, projected to increase up to 4 °C until the end of the century (Intergovernmental Panel on Climate Change, 2021). Fish peripheral and central mechanisms of acoustic signal production and detection are affected by temperature, as they are ectotherms. For example, when there is a direct relation between sonic muscle contraction and sound pulses, pulse repetition rate (and fundamental frequency) is positively correlated with temperature (Ladich, 2018). This is the case of the painted goby in which the pulse repetition rate of mating drums increases 2.9 Hz on average per °C (Vicente , 2015). Other features, like sound dominant frequency or calling rate, may also vary with temperature in different species (Ladich, 2018). Higher water temperatures have also been found to increase hearing sensitivity, especially in higher frequencies, but these temperature-dependent changes have mainly focussed on fishes with hearing specializations (i.e., possessing accessory hearing structures that enhance hearing, both in terms of sensitivity and detectable hearing range) (Ladich, 2018; Schliwa and Ladich, 2021). We still know very little on the features used for mating decisions in fish and even less whether temperature induces parallel changes in male sound features and in female preference (known as temperature coupling). So far, only one study suggests that there may be temperature coupling in acoustic communication in fishes (Mckibben and Bass, 1998). The possible deleterious effects of temperature uncoupling on mating success needs to be addressed. Importantly, the combined effects of noise pollution, global warming, and other stressors, warrants investigation.

See supplementary material for a description of features that facilitate studying the function of mating acoustic signals in fish species (Table S1).

This research journey would not have been meaningful and rewarding without all the people who have contributed to the body of research that is presented here. Many thanks to all the students, collaborators, and partners in science who have been a part of this research. The author also thankful to Yorgos Stratoudakis for reviewing an earlier draft of this manuscript and to two anonymous reviewers who provided constructive comments that greatly improved the paper.

The authors have no conflicts to disclose.

The data that support the findings of this study are available from the corresponding author upon reasonable request.

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