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.     

饲养状态真鲂(鱼弗)发声的昼夜与季节变化

本文研究了受水温和光周期等自然变化影响的饲养状态真鲂鮄发声的昼夜与季节变化,并研究了实验鱼活动(鱼类游泳次数)的日变化。声音信号的昼夜节律记录发声活动的日常水平(摄食之外的时期),但是每月变化的记录(季节性式型)则在摄食期间进行,因为摄食时声音信号增加,而日常发声活动较不频繁。实验鱼包括雌雄两性,且未达性成熟。真鲂鮄在白天发声多一些,也更活跃。声音为阵发式的,较不频繁(平均值=0.04发声/min每鱼每天)。最少的发声活动出现在晚上,凌晨和黄昏居中(声音的阵发更频繁,但是声音更少),最多的发声活动出现在白天(声音的阵发更频繁,并且含更多数目的声音)。竞争摄食时声音信号的比率不呈现季节性变化(平均值=3.98发声/min每鱼),与温度也不相关,显示出竞争摄食时声音的发出以最大比率进行。敲击声和呼噜声的某些声学特征与温度相关,特别是在较高的温度下呼噜声的节拍间隔急剧下降。敲击声和呼噜声的声音参数中的季节性变化,多数可以解释为发声肌肉和中央声音控制回路的温度效应。     

Growling for food: acoustic emissions during competitive feeding of the streaked gurnard

The streaked gurnard Trigloporus lastoviza produced only one sound type, a growl, lasting up to 3 s and consisting of repeated groups of typically one to three pulses. The foraging fish followed two different strategies. In the first, the fish circled the feeding area, grasped a food item and fled, sometimes displaying aggressively to competitors. With this foraging strategy, fish usually made sounds as they circled, grasped and fled. Fish that growled while circling were more likely to grasp a food item subsequently than were silent fish. The second feeding strategy occurred when a fish had already ingested food or failed to get any. In this case, typically fish searched for food on the substratum or approached and touched other individuals that were feeding, sometimes grabbing food that was spat out during food handling by the other fish. Although payback experiments would be needed to draw firm conclusions on the communicative function of growling during competitive feeding in the streaked gurnard, the results suggest that sound production confers advantages to individuals competing for limited food resources.     

Feeding of grey gurnard Eutrigla gurnardus (Triglidae) in the area of the Rockall Seamount

Specific features of feeding of grey gurnard Eutrigla gurnardus depending on vertical distribution and habitat conditions in the area of the Rockall Seamount in 2000 were considered. The intensity of feeding of grey gurnard from April to September decreased by several times. The main food of this species in spring are euphausiids and that in autumn are fish. Males feed slightly more intensively than females. The intensity of feeding, distribution, and behavior of grey gurnard in different seasons and at different depths near bottom and in the pelagial noticeably differ and depend on the composition of distribution and numbers of dominant food items (euphausiids, sand eel Ammodytes marinus, fish juveniles).     

Acoustic communication in an electric fish,Pollimyrus isidori (Mormyridae)

It has been known since von Frisch's work in the 1930's that mormyrid electric fishes are quite sensitive to sound. We now describe a repertoire of natural sounds produced by the mormyrid, Pollimyrus isidori, during breeding and aggression; reception of communication sounds is probably a major function for mormyrid audition. In aquaria, Pollimyrus isidori produce 'grunts', 'moans', 'growls', 'pops' and 'hoots' at various phases during nesting, courtship, and territory defense. All five sounds are produced primarily at night. Territorial males produce grunts, moans and growls during courtship. Vocalizing is stimulated by the presence of a gravid female on the male's territory and decreases with the onset of spawning. Hoots and pops are given during agonistic behavior. Grunts are bursts of acoustic pulses, stereotyped for an individual, with the potential as individual signatures. The electric organ is silent during grunts and moans and is discharged at a reduced rate during growls. The courtship and spawning of Pollimyrus isidori is described.     

Four type of sounds for one winner: vocalizations during territorial behavior in the red-mouthed goby <Emphasis Type="Italic">Gobius cruentatus</Emphasis> (Pisces Gobiidae)

During territorial encounters, the acoustic repertoire of Gobius cruentatus consists of four types of sound emissions: a tonal sound, a noisy tonal sound, a train of individual pulses, and a complex sound. The complex sound is made of two distinct elements, an initial tonal part followed by pulses. This is the largest acoustic repertoire described so far in gobiid fish during aggressive interaction. Sounds are emitted, mainly by the residents, when fish have already started the interaction but before the encounter is settled. Therefore, sounds seem to have a threatening function.     

Seasonal variation of sound production in the Lusitanian toadfish Halobatrachus didactylus

Seasonal variation of sound production, which includes boatwhistles, grunts, croaks and double croaks, was studied in the Lusitanian toadfish Halobatrachus didactylus . Boatwhistles were emitted during the mating season in contrast with the other sound types, which were emitted all year round.     

Saccular potentials of the vocal plainfin midshipman fish, <Emphasis Type="Italic">Porichthys notatus</Emphasis>

The plainfin midshipman fish, Porichthys notatus, is a vocal species of teleost fish that generates acoustic signals for intraspecific communication during social and reproductive behaviors. All adult morphs (females and males) produce single short duration grunts important for agonistic encounters, but only nesting males produce trains of grunts and growls in agonistic contexts and long duration multiharmonic advertisement calls to attract gravid females for spawning. The midshipman fish uses the saccule as the main acoustic endorgan for hearing to detect and locate vocalizing conspecifics. Here, I examined the response properties of evoked potentials from the midshipman saccule to determine the frequency response and auditory threshold sensitivity of saccular hair cells to behaviorally-relevant single tone stimuli. Saccular potentials were recorded from the rostral, medial and caudal regions of the saccule while sound was presented by an underwater speaker. Saccular potentials of the midshipman, like other teleosts, were evoked greatest at a frequency that was twice the stimulus frequency. Results indicate that midshipman saccular hair cells of non-reproductive adults had a peak frequency sensitivity that ranged from 75 (lowest frequency tested) to 145 Hz and were best suited to detect the low frequency components (≤105 Hz) of midshipman vocalizations.     

Acoustic Repertoire of Spawning Cod, Gadus morhua

We examine the vocal repertoire of spawning cod, Gadus morhua. Fish captured in northern Norway were allowed to spawn in tanks and we recorded sound during the spawning period. The cod's grunt repertoire contained more variation than previously suggested. Half of the grunts lasted more than 200ms including one grunt lasting nearly 1s and consisting of 52 pulses, which is far beyond previous reports. Moreover, increasing repetition period and increasing duration of pulses within a grunt from the first pulse towards pulses at the end of the grunt, has previously not been described. On the other hand, no knocks or series of knocks, and on average only one grunt per hour suggest that cod are less versatile vocalists than closely related gadid species, such as haddock, Melanogrammus aeglefinus.     

Vocalisation Repertoire of Female Bluefin Gurnard (Chelidonichthys kumu) in Captivity: Sound Structure,Context and Vocal Activity

Fish vocalisation is often a major component of underwater soundscapes. Therefore, interpretation of these soundscapes requires an understanding of the vocalisation characteristics of common soniferous fish species. This study of captive female bluefin gurnard, Chelidonichthys kumu, aims to formally characterise their vocalisation sounds and daily pattern of sound production. Four types of sound were produced and characterised, twice as many as previously reported in this species. These sounds fit two aural categories; grunt and growl, the mean peak frequencies for which ranged between 129 to 215 Hz. This species vocalized throughout the 24 hour period at an average rate of (18.5 ± 2.0 sounds fish^-1 h^-1) with an increase in vocalization rate at dawn and dusk. Competitive feeding did not elevate vocalisation as has been found in other gurnard species. Bluefin gurnard are common in coastal waters of New Zealand, Australia and Japan and, given their vocalization rate, are likely to be significant contributors to ambient underwater soundscape in these areas.     

Sound production, hearing and possible interception under ambient noise conditions in the topmouth minnow Pseudorasbora parva

Sounds were produced by the topmouth minnow Pseudorasbora parva , a common Eurasian cyprinid, during feeding but not during intraspecific interactions. Feeding sounds were short broadband pulses with main energies between 100 and 800 Hz. They varied in their characteristics (number of single sounds per feeding sequence, sound duration and period, and sound pressure level) depending on the food type (chironomid larvae, Tubifex worms and flake food). The loudest sounds were emitted when food was taken up at the water surface, most probably reflecting 'suctorial' feeding. Auditory sensitivities were determined between 100 and 4000 Hz utilizing the auditory evoked potentials recording technique. Under laboratory conditions and in the presence of natural ambient noise recorded in Lake Neusiedl in eastern Austria, best hearing sensitivities were between 300 and 800 Hz (57 dB re 1 μPa v . 72 dB in the presence of ambient noise). Threshold-to-noise ratios were positively correlated to the sound frequency. The correlation between sound spectra and auditory thresholds revealed that P. parva can detect conspecific sounds up to 40 cm distance under ambient noise conditions. Thus, feeding sounds could serve as an auditory cue for the presence of food during foraging.     

Vocalisations of the adult female domestic pig during a standard human approach test and their relationships with behavioural and heart rate measures

Vocal communication in the domestic pig is generally not well documented. The aim of this experiment was to categorise and ascribe the function of the vocalisations of 67 Large WhitexLandrace gilts during a standard human approach test. At testing, each group of 3-5 gilts was moved to a handling area where each individual in turn was fitted with a heart rate monitor and introduced individually to a 2.4mx2.4m test arena. After 2min familiarisation, an unfamiliar human entered the pen and stood for 3min against one wall. Behaviour and sound were recorded continuously with sound recordings transferred onto computer for analysis. Three categories of calls were initially identified: single grunts, single squeals and rapidly repeated grunts. Sixty-six gilts performed single grunts, whereas only 28 and 16 gilts performed the other two categories, respectively. Single grunts could be sub-divided into two types based on sound amplitude profile. These types differed significantly in duration. Gilts performed more short and long grunts per minute during the 3min test period than during the familiarisation period. Most short grunts observed in a subset of 15 gilts were performed with the snout close to a pen surface or the human. The rate of short grunts during the test period was negatively correlated with the time taken to make contact with the human and positively correlated with the amount of locomotor behaviour carried out, the total number of interactions with the human and the total time spent within 0.5m of the human. Most long grunts observed in a subset of 15 gilts were performed with the snout away from any surface. The rate of long grunts during the test period positively correlated with amount of locomotor behaviour and heart rate, after the effect of activity had been removed. Squeals could similarly be sub-divided into long and short types on the basis of amplitude profile. Gilts that squealed carried out more locomotor behaviour, interacted with the human more, had higher mean heart rates and lower heart rate rise when touched by the human, suggesting a higher degree of arousal. Rapidly-repeated grunts were associated with close human interaction. The results indicate that the domestic pig performs a number of distinct vocalisations during isolation. Short single grunts appear to be associated with investigatory behaviour. Long single grunts may be a form of contact call, the rate of which is related to physiological and behavioural activity. Squeals may have similar function but result from a higher level of arousal. Short, rapidly-repeated grunts appear to have either a greeting or threat function. With further research, certain pig vocalisations may be identified as providing useful additional information about an individual's welfare.     

Playbacks of food-associated calls attract chimpanzees towards known food patches in a captive setting

Food-associated calls have received much research attention due to their potential to refer to discovered food in a word-like manner. Studies have found that in many species, food-associated calls attract receivers to the food patch, suggesting these calls play roles in food sharing, cooperation and competition. Additionally, in various species, these calls play a role that has received much less attention: mediating social interactions among foragers that are already nearby or within the food patch, independently of whether they attract outside foragers. In order to increase understanding of the function of the chimpanzee (Pan troglodytes) food-associated rough grunt, we conducted captive playback studies testing whether rough grunt playbacks attract, repel or have no effect on the proximity of foragers already familiarized with the presence of food. We tested how acoustic playbacks of rough grunts (or control calls) from one of two known, identical feeding sites affected receivers’ approach and feeding behaviors. More often than expected, participants first approached the feeding site from which rough grunts, but not control calls, were broadcast. However, neither condition increased the likelihood that participants fed first from a given site. Our results support the hypothesis that rough grunts elicit an approach response in receivers, while providing no evidence that they repel. In addition, our study provides evidence that receivers may approach rough grunts even if they do not intend to feed. We discuss the information rough grunts may convey to receivers beyond information about discovered food and the potential benefits signalers may gain from this calling behavior.

     

Vocalizations and Behaviour of Pacific Humpback Dolphins Sousa chinensis

Very little is known about the acoustic repertoire of the Pacific humpback dolphin Sousa chinensis . This study, off eastern Australia, used concurrent observations of surface behaviour and acoustic recordings to gain an insight into the behavioural significance of humpback dolphin vocalizations. Humpback dolphins exhibit five different vocalization categories: broad band clicks; barks; quacks; grunts; and whistles. Broad band clicks were high in frequency (8 kHz to > 22 kHz), were directly related to foraging behaviour and may play a role in social behaviour. Barks and quacks were burst pulse sounds (frequency: 0.6 kHz to > 22 kHz, duration: 0.1–8 s) and were associated with both foraging and social behaviour. The grunt vocalization is a low frequency narrow band sound (frequency 0.5–2.6 kHz, duration 0.06–2 s) and was only heard during socializing. There were 17 different types of whistles, ranging widely in frequency (0.9–22 kHz) and vocal structure (n=329). The predominant whistle types used by the groups were type 1 (46%) and type 2 (17%). Most whistles were heard during both socializing and foraging. The number of whistles recorded in a group increased significantly as the number of mother–calf pairs increased, suggesting that whistles may be used as contact calls. Few vocalizations were heard during either travelling or milling behaviours. Broad band clicks, barks and whistle type 1 were the only vocalizations recorded during either travelling or milling.     

Spawning sounds in meagre Argyrosomus regius recorded in the Gironde estuary, France

During their spawning period (June to mid-July) in the Gironde estuary, meagre Argyrosomus regius produce two distinct sounds: regular long grunts, the most common calls, and sometimes also short grunts. It is suggested that long grunts serve the formation of spawning aggregations and short grunts announce the beginning of courtship behaviour. The meagre's long grunts include long series of 30–112 closely spaced pulses placed into call units. Each pulse produces multiple and rapidly decaying swimbladder vibrations with a dominant frequency varying between 336 and 444 Hz.     

Sound production by the river bullhead, Cottus gobio L. (Cottidae, Teleostei)

Acoustic signals of Cottus gobio consist of knocking sounds produced as single pulses (48 ms) or as trains of 4–6 pulses (230 ms). Frequencies extend up to 3 kHz, but most sound energy is concentrated between 50 and 500 Hz in both sound types. Cottus gobio is solitary, maintains territories, and defends them by threat display, seldom by biting and fighting. Threatening consists of spreading gill covers and fins, darkening, and sound production. Calling is accompanied by a nodding movement of the head, during which the pectoral girdle and the skull are moved rapidly against each other. No difference in ability of sound production was observed between sexes, but males emitted significantly more sounds than females. In the laboratory an increase in vocalization activity was observed between night and day. An increase in the number of encounters and calls was noted when temperatures were raised from 8°C to 13°C. Sound production was registered throughout the year.     

Acoustic characteristics and variations in grunt vocalizations in the oyster toadfish <Emphasis Type="Italic">Opsanus tau</Emphasis>

Acoustic communication is critical for reproductive success in the oyster toadfish Opsanus tau. While previous studies have examined the acoustic characteristics, behavioral context, geographical variation, and seasonality of advertisement boatwhistle sound production, there is limited information on the grunt or other non-advertisement vocalizations in this species. This study continuously monitored sound production in toadfish maintained in an outdoor habitat for four months to identify and characterize grunt vocalizations, compare them with boatwhistles, and test for relationships between the incidence of grunt vocalizations, sound characteristics and environmental parameters. Oyster toadfish produced grunts in response to handling, and spontaneous single (70% of all grunts), doublet (10%), and trains of grunts (20%) throughout the May to September study period. Grunt types varied in pulse structure, duration, and frequency components, and were shorter and of lower fundamental frequency than the pulse repetition rate of boatwhistles. Higher water temperatures were correlated with a greater number of grunt emissions, higher fundamental frequencies, and shorter sound durations. The number of grunts per day was also positively correlated with daylength and maximum tidal amplitude differences (previously entrained) associated with full and new moons, thus providing the first demonstration of semilunar vocalization rhythms in the oyster toadfish. These data provide new information on the acoustic repertoire and the environmental factors correlated with sound production in the toadfish, and have important implications for seasonal acoustic communication in this model vocal fish.     

Food partitioning between coexisting Atlantic salmon and brook trout in the Sainte‐Marguerite River ecosystem, Quebec

Food resource partitioning between similar‐sized, sympatric Atlantic salmon Salmo salar and brook trout Salvelinus fontinalis was examined as a possible mechanism enabling their coexistence in a stream (Allaire) of the Sainte‐Marguerite River ecosystem, Quebec, Canada. Fish stomach contents and invertebrate drift were collected concurrently during three diel cycles in August to September 1996. The food and feeding habits of an allopatric brook trout population in a nearby stream (Epinette) were studied for comparison. The diel feeding rhythms of the two coexisting fish species were similar. The composition of their diet, however, showed significant differences. Atlantic salmon predominantly (60–90%) fed on aquatic insects, mainly Ephemeroptera (35–60% of the diet). The brook trout mostly (50–80%) fed upon the allochthonous terrestrial insects (mainly adults of Coleoptera, Hymenoptera and Diptera) which comprised 5–40% of the stream drift. The allopatric brook trout fed opportunistically on the more abundant aquatic insects and terrestrial insects rarely formed 25% of its diet. The allopatric trout fed nearly twice as much as the sympatric brook trout during a day. The results suggest that the differences in feeding by brook trout in the two streams (with and without Atlantic salmon) are the result of inter‐specific interaction with Atlantic salmon and are not related to the differences in food availability between the two streams. Food resource partitioning between Atlantic salmon and brook trout may be viewed as an adaptive response resulting in a greater exploitation of available resources and coexistence.     

The basis of food selection in flounders, Platichthys flesus (L.), in the Severn Estuary

The factors influencing the selection of food by flounders, Platichthys flesus (L.), have been investigated by analysing collections made in the Severn Estuary for 1 year and the results of experiments. Flounders measuring between 6–0 and 35 cm fed heavily on the polychaete Nereis diversicolor in February and on the amphipod Gammarus salinus between February and April. Thereafter these species were replaced by the mysid Neomysis integer with the decapod Crangon vulgaris. Flounders shorter than 6.0 cm, fed mainly on Neomysis integer regardless of month. Numerous factors were involved in the choice of the food including (1) the maximum and minimum length of prey, (2) its spatial distribution in the water column, (3) its degree of concealment, (4) its motility and ability to escape predation, (5) conditioning of the flounders for certain foods, (6) the fish's swimming speed and (7) the turbidity and temperature of the water.
Although the average length of ingested prey remained unchanged as the fish grew in size, an increase in the maximum and minimum lengths was observed. The percentage of stomachs that contained no identifiable remains in flounders greater than 6–0 cm averaged 80–95 % during part of the winter and 60% in the summer and between January and March. These changes partly reflected the influence of low water temperature on the metabolic rate and availability of prey. The dry weight of the stomach contents in flounders longer than 6–0 cm was lowest in winter but high values were recorded in the spring. This latter feature was probably because the rate of feeding was greater than that of digestion. During the summer, under more normal feeding conditions, the weight of the stomach contents remained relatively low compared to other fish populations. Flounders shorter than 6–0 cm always contained more food in their stomachs on a unit weight basis than larger individuals, reflecting metabolic rate and hunting efficiency.