Ecology of fish hearing

Underwater sound is directional and can convey important information about the surrounding environment or the animal emitting the sound. Therefore, sound is a major sensory channel for fishes and plays a key role in many life-history strategies. The effect of anthropogenic noise on aquatic life, which may be causing homogenisation or fragmentation of biologically important signals underwater is of growing concern. In this review we discuss the role sound plays in the ecology of fishes, basic anatomical and physiological adaptations for sound reception and production, the effects of anthropogenic noise and how fishes may be coping to changes in their environment, to put the ecology of fish hearing into the context of the modern underwater soundscape.     

The effects of anthropogenic sources of sound on fishes

There is increasing concern about the effects of pile driving and other anthropogenic (human-generated) sound on fishes. Although there is a growing body of reports examining this issue, little of the work is found in the peer-reviewed literature. This review critically examines both the peer-reviewed and 'grey' literature, with the goal of determining what is known and not known about effects on fish. A companion piece provides an analysis of the available data and applies it to estimate noise exposure criteria for pile driving and other impulsive sounds. The critical literature review concludes that very little is known about effects of pile driving and other anthropogenic sounds on fishes, and that it is not yet possible to extrapolate from one experiment to other signal parameters of the same sound, to other types of sounds, to other effects, or to other species.     

褐菖鲉的听觉阈值研究

利用听觉诱发电位记录技术研究了褐菖鲉(Sebasticus marmoratus)的听觉阈值。通过采用听觉生理系统记录和分析了8尾褐菖鲉对频率范围在100—1000 Hz的7种不同频率的声音刺激的诱发电位反应。结果表明, 褐菖鲉的听觉阈值在整体上随着频率增加而增加, 对100—300 Hz的低频声音信号敏感, 最敏感频率为150 Hz, 对应的听觉阈值为70 dB re 1 μPa。褐菖鲉的听觉敏感区间与其发声频率具有较高的匹配性, 表明其声讯交流的重要性。同时, 人为低频噪声可能对其声讯交流造成影响。     

Potential interactions between diadromous fishes of U.K. conservation importance and the electromagnetic fields and subsea noise from marine renewable energy developments

The considerable extent of construction and operation of marine renewable energy developments (MRED) within U.K. and adjacent waters will lead, among other things, to the emission of electromagnetic fields (EMF) and subsea sounds into the marine environment. Migratory fishes that respond to natural environmental cues, such as the Earth's geomagnetic field or underwater sounds, move through the same waters that the MRED occupy, thereby raising the question of whether there are any effects of MRED on migratory fishes. Diadromous species, such as the Salmonidae and Anguillidae, which undertake large-scale migrations through coastal and offshore waters, are already significantly affected by other human activities leading to national and international conservation efforts to manage any existing threats and to minimize future concerns, including the potential effect of MRED. Here, the current state of knowledge with regard to the potential for diadromous fishes of U.K. conservation importance to be affected by MRED is reviewed. The information on which to base the review was found to be limited with respect to all aspects of these fishes' migratory behaviour and activity, especially with regards to MRED deployment, making it difficult to establish cause and effect relationships. The main findings, however, were that diadromous species can use the Earth's magnetic field for orientation and direction finding during migrations. Juveniles of anadromous brown trout (sea trout) Salmo trutta and close relatives of S. trutta respond to both the Earth's magnetic field and artificial magnetic fields. Current knowledge suggests that EMFs from subsea cables may interact with migrating Anguilla sp. (and possibly other diadromous fishes) if their movement routes take them over the cables, particularly in shallow water (<20 m). The only known effect is a temporary change in swimming direction. Whether this will represent a biologically significant effect, for example delayed migration, cannot yet be determined. Diadromous fishes are likely to encounter EMFs from subsea cables either during the adult movement phases of life or their early life stages during migration within shallow, coastal waters adjacent to natal rivers. The underwater sound from MRED devices has not been fully characterized to determine its acoustic properties and propagation through the coastal waters. MRED that require pile driving during construction appear to be the most relevant to consider. In the absence of a clear understanding of their response to underwater sound, the specific effects on migratory species of conservation concern remain very difficult to determine in relation to MRED. Based on the studies reviewed, it is suggested that fishes that receive high intensity sound in close proximity to construction may be physiologically affected to some degree, whereas those at farther distances, potentially up to several km, may exhibit behaviour responses; the effect of which is unknown and will be dependent on the properties of the received sound and receptor characteristics and condition. Whether there are behavioural effects on the fishes during operation is unknown but any change to the environment and subsequent response by the fishes would need to be considered over the lifetime of the MRED. It is not yet possible to determine if effects relating to sound exposure are biologically significant. The current assumptions of limited effects are built on an incomplete understanding of how the species move around their environment and interact with natural and anthropogenic EMFs and subsea sound. A number of important knowledge gaps exist, principally whether migratory fish species on the whole respond to the EMF and the sound associated with MRED. Future research should address the principal gaps before assuming that any effect on diadromous species results in a biological effect.     

Sex-Specific Differences in Agonistic Behaviour,Sound Production and Auditory Sensitivity in the Callichthyid Armoured Catfish Megalechis thoracata

Background

Data on sex-specific differences in sound production, acoustic behaviour and hearing abilities in fishes are rare. Representatives of numerous catfish families are known to produce sounds in agonistic contexts (intraspecific aggression and interspecific disturbance situations) using their pectoral fins. The present study investigates differences in agonistic behaviour, sound production and hearing abilities in males and females of a callichthyid catfish.

Methodology/Principal Findings

Eight males and nine females of the armoured catfish Megalechis thoracata were investigated. Agonistic behaviour displayed during male-male and female-female dyadic contests and sounds emitted were recorded, sound characteristics analysed and hearing thresholds measured using the auditory evoked potential (AEP) recording technique. Male pectoral spines were on average 1.7-fold longer than those of same-sized females. Visual and acoustic threat displays differed between sexes. Males produced low-frequency harmonic barks at longer distances and thumps at close distances, whereas females emitted broad-band pulsed crackles when close to each other. Female aggressive sounds were significantly shorter than those of males (167 ms versus 219 to 240 ms) and of higher dominant frequency (562 Hz versus 132 to 403 Hz). Sound duration and sound level were positively correlated with body and pectoral spine length, but dominant frequency was inversely correlated only to spine length. Both sexes showed a similar U-shaped hearing curve with lowest thresholds between 0.2 and 1 kHz and a drop in sensitivity above 1 kHz. The main energies of sounds were located at the most sensitive frequencies.

Conclusions/Significance

Current data demonstrate that both male and female M. thoracata produce aggressive sounds, but the behavioural contexts and sound characteristics differ between sexes. Sexes do not differ in hearing, but it remains to be clarified if this is a general pattern among fish. This is the first study to describe sex-specific differences in agonistic behaviour in fishes.     

Geographical variation in sound production in the anemonefish Amphiprion akallopisos

Because of pelagic-larval dispersal, coral-reef fishes are distributed widely with minimal genetic differentiation between populations. Amphiprion akallopisos, a clownfish that uses sound production to defend its anemone territory, has a wide but disjunct distribution in the Indian Ocean. We compared sounds produced by these fishes from populations in Madagascar and Indonesia, a distance of 6500 km. Differentiation of agonistic calls into distinct types indicates a complexity not previously recorded in fishes' acoustic communication. Moreover, various acoustic parameters, including peak frequency, pulse duration, number of peaks per pulse, differed between the two populations. The geographic comparison is the first to demonstrate 'dialects' in a marine fish species, and these differences in sound parameters suggest genetic divergence between these two populations. These results highlight the possible approach for investigating the role of sounds in fish behaviour in reproductive divergence and speciation.     

Ontogenesis of agonistic vocalizations in the cichlid fish Metriaclima zebra

While acoustic communication has been described in adults of various fish species, our knowledge about the ontogeny of fish sound production is limited. In adults, sound signals are known to be involved during aggressive interactions. However, aggressive behaviour may appear early in the life of fishes due to the possible competition for food and space. If acoustic signals are used to send information to competitors, sounds are likely to play a role during interactions between juvenile fish as well. The apparition and evolution of sound production were monitored in a group of juveniles of the cichlid fish Metriaclima zebra from hatching to 4months of age. In addition, the link between vocalizations and agonistic behaviour was studied during dyadic interactions at three different ages. Sounds production appeared to be present early in the development of this fish and increased along with the number of aggressive behaviours. Recorded sounds consisted, in juveniles, in isolated pulses showing a decrease in frequency and duration as the fish grew. In adults, sounds became bursts of pulses but the transition from isolated to repetitive pulses was not observed. These results are compared to the existing literature on sound production ontogeny in fishes.     

The hearing abilities of the prawn Palaemon serratus

The mechanism of sound reception and the hearing abilities of the prawn (Palaemon serratus) have been studied using a combination of anatomical, electron microscopic and electrophysiological approaches, revealing that P. serratus is responsive to sounds ranging in frequency from 100 to 3000 Hz. It is the first time that the Auditory Brainstem Response (ABR) recording technique has been used on invertebrates, and the acquisition of hearing ability data from the present study adds valuable information to the inclusion of an entire sub-phylum of animals when assessing the potential impact of anthropogenic underwater sounds on marine organisms. Auditory evoked potentials were acquired from P. serratus, using two subcutaneous electrodes positioned in the carapace close to the supraesophageal ganglion and the statocyst (a small gravistatic organ located below the eyestalk on the peduncle of the bilateral antennules). The morphology of the statocyst receptors and the otic nerve pathways to the brain have also been studied, and reveal that P. serratus possesses an array of sensory hairs projecting from the floor of the statocyst into a mass of sand granules embedded in a gelatinous substance. It is the purpose of this work to show that the statocyst is responsive to sounds propagated through water from an air mounted transducer. The fundamental measure of the hearing ability of any organism possessing the appropriate receptor mechanism is its audiogram, which presents the lowest level of sound that the species can hear as a function of frequency. The statocyst of P. serratus is shown here to be sensitive to the motion of water particles displaced by low-frequency sounds ranging from 100 Hz up to 3000 Hz, with a hearing acuity similar to that of a generalist fish. Also, recorded neural waveforms were found to be similar in both amplitude and shape to those acquired from fish and higher vertebrates, when stimulated with low-frequency sound, and complete ablation of the electrophysiological response was achieved by removal of the statocyst.     

Agent-based models to investigate sound impact on marine animals: bridging the gap between effects on individual behaviour and population level consequences

Marine ambient sound levels have risen due to noisy human activities, such as shipping, fishing, seismic surveys and piling for windfarms. Marine mammals and fishes are two prominent taxonomic groups that are exposed to this noise pollution, which may experience detrimental effects at the population level. Acoustic effects on individual behaviour such as deterrence, disturbance, distraction and masking of biologically relevant sounds, can be translated energetically to changes in vital rates (growth, maturation, reproduction and survival) in a population consequences of acoustic disturbance (PCAD) approach. However, we typically neglect spatial variation in species distributions and noise pollution, while abiotic factors like temperature, bathymetry and currents, as well as habitat quality in terms of feeding or hiding opportunities, will also have a geographically variable impact on potential consequences. We here address the conceptual integration of agent based models (ABM) into the PCAD framework, as a suitable theoretical tool with high potential for the exploration of these spatial factors and their modifying role in noise impact assessment studies. We review five ABM case studies, including investigations into: 1) effects of movement strategy on the impact of explosions in harbour porpoise; 2) effects of disturbance sensitivity on pile driving impact on migrating cod; 3) impact of seismic survey sounds on Atlantic mackerel distribution and movement; 4) population-level impact of mitigation of harbour porpoise bycatch with pingers; and 5) population effects of alternative windfarm construction scenarios in harbour porpoise. We discuss similarities and differences among these studies in sound and species mapping approaches and we evaluate model realism and pattern validation. We believe that ABMs are a valuable tool for integrating spatial information into ecological impact studies that investigate acoustic disturbance, for any type of sound source, and for both marine mammals and fish.     

The African cichlid fish Astatotilapia burtoni uses acoustic communication for reproduction: sound production, hearing, and behavioral significance

Sexual reproduction in all animals depends on effective communication between signalers and receivers. Many fish species, especially the African cichlids, are well known for their bright coloration and the importance of visual signaling during courtship and mate choice, but little is known about what role acoustic communication plays during mating and how it contributes to sexual selection in this phenotypically diverse group of vertebrates. Here we examined acoustic communication during reproduction in the social cichlid fish, Astatotilapia burtoni. We characterized the sounds and associated behaviors produced by dominant males during courtship, tested for differences in hearing ability associated with female reproductive state and male social status, and then tested the hypothesis that female mate preference is influenced by male sound production. We show that dominant males produce intentional courtship sounds in close proximity to females, and that sounds are spectrally similar to their hearing abilities. Females were 2-5-fold more sensitive to low frequency sounds in the spectral range of male courtship sounds when they were sexually-receptive compared to during the mouthbrooding parental phase. Hearing thresholds were also negatively correlated with circulating sex-steroid levels in females but positively correlated in males, suggesting a potential role for steroids in reproductive-state auditory plasticity. Behavioral experiments showed that receptive females preferred to affiliate with males that were associated with playback of courtship sounds compared to noise controls, indicating that acoustic information is likely important for female mate choice. These data show for the first time in a Tanganyikan cichlid that acoustic communication is important during reproduction as part of a multimodal signaling repertoire, and that perception of auditory information changes depending on the animal's internal physiological state. Our results highlight the importance of examining non-visual sensory modalities as potential substrates for sexual selection contributing to the incredible phenotypic diversity of African cichlid fishes.     

Ocean acidification boosts larval fish development but reduces the window of opportunity for successful settlement

Locating appropriate settlement habitat is a crucial step in the life cycle of most benthic marine animals. In marine fish, this step involves the use of multiple senses, including audition, olfaction and vision. To date, most investigations of larval fish audition focus on the hearing thresholds to various frequencies of sounds without testing an ecological response to such sounds. Identifying responses to biologically relevant sounds at the development stage in which orientation is most relevant is fundamental. We tested for the existence of ontogenetic windows of reception to sounds that could act as orientation cues with a focus on vulnerability to alteration by human impacts. Here we show that larvae of a catadromous fish species (barramundi, Lates calcarifer) were attracted towards sounds from settlement habitat during a surprisingly short ontogenetic window of approximately 3 days. Yet, this auditory preference was reversed in larvae reared under end-of-century levels of elevated CO₂, such that larvae are repelled from cues of settlement habitat. These future conditions also reduced the swimming speeds and heightened the anxiety levels of barramundi. Unexpectedly, an acceleration of development and onset of metamorphosis caused by elevated CO₂ were not accompanied by the earlier onset of attraction towards habitat sounds. This mismatch between ontogenetic development and the timing of orientation behaviour may reduce the ability of larvae to locate habitat or lead to settlement in unsuitable habitats. The misinterpretation of key orientation cues can have implications for population replenishment, which are only exacerbated when ontogenetic development decouples from the specific behaviours required for location of settlement habitats.     

The effects of noise on the auditory sensitivity of the bluegill sunfish,Lepomis macrochirus

As concerns about the effects of underwater anthropogenic noises on the auditory function of organisms increases, it is imperative to assess if all organisms are equally affected by the same noise source. Consequently, auditory capabilities of an organism need to be evaluated and compared interspecifically. Teleost fishes provide excellent models to examine these issues due to their diversity of hearing capabilities. Broadly, fishes can be categorized as hearing specialists (broad hearing frequency range with low auditory thresholds) or hearing generalists (narrower frequency range with higher auditory thresholds). The goal of this study was to examine the immediate effects of white noise exposure (0.3-2.0 kHz, 142 dB re: 1 microPa) and recovery after exposure (1-6 days) on a hearing generalist fish, bluegill sunfish (Lepomis macrochirus). Noise exposure resulted in only a slight, but not statistically significant, elevation in auditory threshold compared to fish not exposed to noise. In combination with results from our previous studies examining effects of noise on a hearing specialist fish, the fathead minnow (Pimephales promelas), this study provides evidence supporting the hypothesis that fish's auditory thresholds can be differentially affected by noise exposure.     

Modelling larval dispersal and behaviour of coral reef fishes

Coral reef fish spend their first few weeks developing in the open ocean, where eggs and larvae appear merciless to tides and currents, before attempting to leave the pelagic zone and settle on a suitable reef. This pelagic dispersal phase is the process that determines population connectivity and allows replenishment of harvested populations across multiple coral reef habitats. Until recently this pelagic larval dispersal phase has been poorly understood and has often been referred to as the ‘black-box’ in the life-history of coral reef fishes. In this perspective article we highlight three areas where mathematical and computational approaches have been used to aid our understanding of this important ecological process. We discuss models that provide insights into the evolution of the pelagic larval phase in coral reef fish, an unresolved question which lends itself well to a modelling approach due to the difficulty in obtaining empirical data on this life history strategy. We describe how studies of fish hearing and physical sound propagation models can be used to predict the detection distance of reefs for settling larval fish, and the potential impact of anthropogenic noise. We explain how random walk models can be used to explore individual- and group-level behaviour in larval fish during the dispersal and settlement stage of their life-history. Finally, we discuss the mutual benefits that mathematical and computational approaches have brought to and gained from the field of larval behaviour and dispersal of reef fishes.     

Effects of temperature on sound production and auditory abilities in the Striped Raphael catfish Platydoras armatulus (Family Doradidae)

Background

Sound production and hearing sensitivity of ectothermic animals are affected by the ambient temperature. This is the first study investigating the influence of temperature on both sound production and on hearing abilities in a fish species, namely the neotropical Striped Raphael catfish Platydoras armatulus.

Methodology/Principal Findings

Doradid catfishes produce stridulation sounds by rubbing the pectoral spines in the shoulder girdle and drumming sounds by an elastic spring mechanism which vibrates the swimbladder. Eight fish were acclimated for at least three weeks to 22°, then to 30° and again to 22°C. Sounds were recorded in distress situations when fish were hand-held. The stridulation sounds became shorter at the higher temperature, whereas pulse number, maximum pulse period and sound pressure level did not change with temperature. The dominant frequency increased when the temperature was raised to 30°C and the minimum pulse period became longer when the temperature decreased again. The fundamental frequency of drumming sounds increased at the higher temperature. Using the auditory evoked potential (AEP) recording technique, the hearing thresholds were tested at six different frequencies from 0.1 to 4 kHz. The temporal resolution was determined by analyzing the minimum resolvable click period (0.3–5 ms). The hearing sensitivity was higher at the higher temperature and differences were more pronounced at higher frequencies. In general, latencies of AEPs in response to single clicks became shorter at the higher temperature, whereas temporal resolution in response to double-clicks did not change.

Conclusions/Significance

These data indicate that sound characteristics as well as hearing abilities are affected by temperatures in fishes. Constraints imposed on hearing sensitivity at different temperatures cannot be compensated even by longer acclimation periods. These changes in sound production and detection suggest that acoustic orientation and communication are affected by temperature changes in the neotropical catfish P. armatulus.     

Learned or innate production of acoustic signals in fishes: a test using a cyprinid

No information on the inheritance of the ability to produce sounds exists for fishes. In birds, which usually provide extensive post-hatching parental care, acoustic signals are learned in some species but are innate in others. Almost no fishes provide extensive post-hatching parental care and, consequently, the offspring have little opportunity to hear and learn sounds produced by the parents (usually the male in fishes); they may, however, be exposed to acoustic signals of conspecifics in the same habitat. We used a cyprinid, Codoma ornata, to test whether sound production is learned from the parents or whether it is innate. Fertilized eggs of this species were raised in isolation from adults. Upon maturity, these fish were tested for sound production in aggressive and reproductive contexts. Fish which had no contact with adults, and therefore no opportunity to hear the acoustic signals of their species, produced sounds that were similar to those produced by their parents, and they produced these in the same contexts. Significant differences were observed in dominant frequency for one context, with the smaller F₁ fish having signals of higher frequency than parental fish. Since no opportunity for learning existed, this provided evidence that the ability to produce sounds is innate in this minnow species.     

How Nemo finds home: the neuroecology of dispersal and of population connectivity in larvae of marine fishes

Nearly all demersal teleost marine fishes have pelagic larval stages lasting from several days to several weeks, during which time they are subject to dispersal. Fish larvae have considerable swimming abilities, and swim in an oriented manner in the sea. Thus, they can influence their dispersal and thereby, the connectivity of their populations. However, the sensory cues marine fish larvae use for orientation in the pelagic environment remain unclear. We review current understanding of these cues and how sensory abilities of larvae develop and are used to achieve orientation with particular emphasis on coral-reef fishes. The use of sound is best understood; it travels well underwater with little attenuation, and is current-independent but location-dependent, so species that primarily utilize sound for orientation will have location-dependent orientation. Larvae of many species and families can hear over a range of ~100-1000 Hz, and can distinguish among sounds. They can localize sources of sounds, but the means by which they do so is unclear. Larvae can hear during much of their pelagic larval phase, and ontogenetically, hearing sensitivity, and frequency range improve dramatically. Species differ in sensitivity to sound and in the rate of improvement in hearing during ontogeny. Due to large differences among-species within families, no significant differences in hearing sensitivity among families have been identified. Thus, distances over which larvae can detect a given sound vary among species and greatly increase ontogenetically. Olfactory cues are current-dependent and location-dependent, so species that primarily utilize olfactory cues will have location-dependent orientation, but must be able to swim upstream to locate sources of odor. Larvae can detect odors (e.g., predators, conspecifics), during most of their pelagic phase, and at least on small scales, can localize sources of odors in shallow water, although whether they can do this in pelagic environments is unknown. Little is known of the ontogeny of olfactory ability or the range over which larvae can localize sources of odors. Imprinting on an odor has been shown in one species of reef-fish. Celestial cues are current- and location-independent, so species that primarily utilize them will have location-independent orientation that can apply over broad scales. Use of sun compass or polarized light for orientation by fish larvae is implied by some behaviors, but has not been proven. Use of neither magnetic fields nor direction of waves for orientation has been shown in marine fish larvae. We highlight research priorities in this area.     

First attempt to use a remotely operated vehicle to observe soniferous fish behavior in the Gulf of Maine, Western Atlantic Ocean

Underwater sound and video observations were made at noon, sunset, and midnight in sand, gravel, and boulder habitat in the Stellwagen Bank National Marine Sanctuary, Gulf of Maine, USA in October 2001 using a remotely operated vehicle (ROV). Seventeen species of fish and squid were observed with clear habitat and time differences. Observations of feeding behavior, disturbance behavior, and both interspecific and intraspecific interactions provided numerous opportunities for potential sound production; however, sounds were recorded only during a single dive. Although high noise levels generated by the ROV and support ship may have masked some sounds, we conclude that fish sound production in the Gulf of Maine during the fall is uncommon. The recorded fish sounds are tentatively attributed to the cusk Brosme brosme. Cusk sounds consisted variously of isolated thumps, widely spaced thump trains, drumrolls, and their combinations. Frequency peaks were observed at 188, 539, and 1195 Hz. Use of a remotely operated vehicle (ROV) as a passive acoustic observation platform was problematic due to high ROV self-noise and the ROV's inability to maintain a fixed position on the bottom without thruster power. Some fishes were clearly also disturbed by ROV noise, indicating a potential ROV sampling bias. Based on our observations, we suggest that new instruments incorporating both optic and passive acoustic technologies are needed to provide better tools for in situ behavioral studies of cusk and other fishes [Current Zoology 56 (1): 90-99 2010].     

The hearing of the Atlantic Salmon, Salmo salar

The hearing of the salmon, Salmo salar L., was studied by means of a cardiac conditioning technique. Fish were trained to show a slowing of the heart, on hearing a sound, in anticipation of a mild electric shock applied later. The minimum sound level to which the fish would respond was determined for a range of pure tones, both in the sea, and in the laboratory. The fish responded only to low frequency tones (below 380 Hz), and particle motion, rather than sound pressure, proved to be the relevant stimulus. The sensitivity of the fish to sound was not affected by the level of sea noise under natural conditions but hearing is likely to be masked by ambient noise in a turbulent river. Sound measurements made in the River Dee, near Aberdeen, lead to the conclusion that salmon are unlikely to detect sounds originating in air, but that they are sensitive to substrate borne sounds. Compared with the carp and cod the hearing of the salmon is poor, and more like that of the perch and plaice.     

Ontogeny of Acoustic and Feeding Behaviour in the Grey Gurnard, Eutrigla gurnardus

Although sound production in teleost fish is often associated with territorial behaviour, little is known of fish acoustic behaviour in other agonistic contexts such as competitive feeding and how it changes during ontogeny. The grey gurnard, Eutrigla gurnardus, frequently emits knock and grunt sounds during competitive feeding and seems to adopt both contest and scramble tactics under defensible resource conditions. Here we examine, for the first time, the effect of fish size on sound production and agonistic behaviour during competitive feeding. We have made sound (alone) and video (synchronized image and sound) recordings of grey gurnards during competitive feeding interactions. Experimental fish ranged from small juveniles to large adults and were grouped in four size classes: 10–15, 15–20, 25–30 and 30–40 cm in total length. We show that, in this species, both sound production and feeding behaviour change with fish size. Sound production rate decreased in larger fish. Sound duration, pulse duration and the number of pulses increased whereas the peak frequency decreased with fish size, in both sound types (knocks and grunts). Interaction rate and the frequency of agonistic behaviour decreased with increasing fish size during competitive feeding sessions. The proportion of feeding interactions accompanied by sound production was similar in all size classes. However, the proportion of interactions accompanied by knocks (less aggressive sounds) and by grunts (more aggressive) increased and decreased with fish size, respectively. Taken together, these results suggest that smaller grey gurnards compete for food by contest tactics whereas larger specimens predominantly scramble for food, probably because body size gives an advantage in locating, capturing and handling prey. We further suggest that sounds emitted during feeding may potentially give information on the motivation and ability of the individual to compete for food resources.     

Pacific herring hearing does not include ultrasound

Recent studies have shown that some clupeid fishes, including shad and menhaden, can detect ultrasound (sound with frequencies higher than 20 kHz) and actively avoid it. However, other clupeids, including sardines and anchovies, do not detect ultrasound. The hearing abilities of herring are of particular interest because of their commercial importance, our reliance on acoustics to monitor their populations and behavioural evidence of responses to high-frequency sound by some clupeid species. We measured the hearing sensitivity of Pacific herring (Clupea pallasii) using the auditory brainstem response and found that they were unable to detect ultrasonic signals at received levels up to 185 dB re 1 microPa. Herring had hearing thresholds at lower frequencies (100-5000 Hz) that were typical of other non-ultrasound-detecting clupeids. This lower-frequency hearing sensitivity could explain the results of several earlier studies showing responses to broadband sounds.