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Lusitanian toadfish song reflects male quality.

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M. Clara P. Amorim at University of Lisbon
M. Clara P. Amorim
José Miguel Simões at Northwestern University
José Miguel Simões
Mendonça N
Mendonça N
Mendonça N
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Narcisa Maria Bandarra at Instituto Português do Mar e da Atmosfera
Narcisa Maria Bandarra
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10/12/2015 Lusitaniantoadfishsongreflectsmalequality[abstract]|Bioacousticsjournal
http://www.bioacoustics.info/article/lusitaniantoadfishsongreflectsmalequalityabstract 1/2
Authors:
Amorim,MCP,Simões,JM,Mendonça,N,Bandarra,N,Almada,VC&Fonseca,PJ
Year:2012
Volume: 21
Issue:1
Frompage:65
Topage:66
Abstract:
Duringthebreedingseason,Lusitaniantoadfishmalesbecometerritorialandemitan
advertisementcall(theboatwhistle,BW)toattractmates.Wetestthehypothesisthat
boatwhistlescanconveyinformationonindividualqualitybyrelatingcallingactivityand
signalparameterswithmalephysicalfeatures,namelymaletotallength,gonadosomatic
index,sonicmuscleindexandmalecondition,measuredbytheconditionfactorandmuscle
lipidcontent.Werecorded22malesintheTagusRiverestuary(Portugal)duringthe
breedingseason(2006and2007)foruptoeightdaysandforanaverageof35h.Single
maleswerekeptinsideclosedartificialnestsandgroupsof68individualsmaintaineda
normalchorusactivity.Acousticactivityvariedgreatlyamongsubjectmales.Allmales
producedboatwhistlesduringthestudyperiodbutcallingratevariedmarkedlyamong
them.Averagecallingratevariedfrom0.1to361.7BWh1(overallmean=39.9BWh1)and
themajorityoftimewasspentinsilence(72.4%oftherecordedhours).Boatwhistle
emissionrateandthefundamentalfrequencyofboatwhistles(equivalenttothesonic
musclecontractionfrequencyduringsoundproduction)werecorrelatedwithmale
condition(somaticandlipidcontent).Boatwhistleemissionratewasalsocorrelatedwith
sonicmuscleindex,i.e.withsonicmusclehypertrophy.Callingratevariabilityalso
decreasedwithincreasedsonicmuscleandgonadosomaticindexes.Theseresultssuggest
thatonlymaleswithhighersomaticandreproductiveconditionandwithhighersonic
musclemasswereabletosustainelevatedacousticactivityandsonicmusclecontraction
ratesduringsoundproduction.Inconclusion,boatwhistlesseemtobeabletosignalmale
qualityduringsocialinteractions.
Categories: fish
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Lusitaniantoadfishsongreflectsmalequality
[abstract]
Citation:
Bioacousticsjournal
10/12/2015 Lusitaniantoadfishsongreflectsmalequality[abstract]|Bioacousticsjournal
http://www.bioacoustics.info/article/lusitaniantoadfishsongreflectsmalequalityabstract 2/2
Amorim,MCP,Simões,JM,Mendonça,N,Bandarra,N,Almada,VC&Fonseca,PJ
(2012).Lusitaniantoadfishsongreflectsmalequality[abstract].Bioacoustics21(1):6566
... However, these frequencies are related to the pulse period (i.e., the rate of muscle contraction) and not fish size. Moreover, the pulse period has low variability in this taxa, which is consistent with their vocal central pattern generator (Bass and Baker 1990;Barimo and Fine 1998;Amorim et al. 2010). In Halobatrachus didactylus, reproductive success appears to be determined by calling rate and calling effort (i.e., the highest percentage of time spent calling). ...
... In Halobatrachus didactylus, reproductive success appears to be determined by calling rate and calling effort (i.e., the highest percentage of time spent calling). These parameters indicate male condition (Vasconcelos et al. 2012), as reflected in sonic muscle hypertrophy and larger gonads (Amorim et al. 2010). In this case, the sonic muscle mass would allow long periods of calling but not affect main frequency. ...
... In this case, the sonic muscle mass would allow long periods of calling but not affect main frequency. In other words, sounds would be related to male quality, that is to males that are likely to confer greater fitness on their offspring (Amorim and Vasconcelos 2008;Amorim et al. 2010) but not to its size. ...
Fish Sound Production: Insights
Chapter
Full-text available
  • Apr 2016
In addition to briefly reviewing sound-producing mechanisms, this chapter focuses on an under-appreciated evolutionary process, exaptation, which could aid in understanding the independent origins and high diversity of sound-producing mechanisms in fishes. Existing anatomical structures first used in non-voluntary sound production provide advantages that result in further selection and refinement of sophisticated sonic organs. Moreover, comparisons of the relationships between fish size and spectral features in multiple not phylogenetically related species highlight two acoustic patterns. In species using superfast muscles, the slope of the relationship between fish size and sound frequency is weak (1°–5°) so that emitter size is unlikely inferred from call frequency. In other species that stridulate or use bones or tendons to stimulate the swimbladder, the high slopes (25°–80°) indicate major differences in the call frequencies within a species. These signals likely convey important information (size and potential fitness of the emitter) to conspecific receivers.
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... Following Amorim et al (2010) , we used residuals of the simple linear regression of sonic muscle mass on ME (RM SM ) as a metric of sonic muscle hypertrophy. Likewise, we used the residuals of the simple linear regressions of gonads, accessory glands, and liver mass on ME (RM G , RM AG , and RM L , respectively ) to control for the influence of body size. ...
... eggs' survival as they need to be aerated for proper development and to prevent fungal infections (Ramos and Amorim, unpublished data). Therefore, in species where males provide parental care, indicators of male parental quality are expected to be important in intersexual communication and be under strong mate selection by females (Andersson 1994). Amorim et al. (2010) also reported that increased boatwhistle CR and CE strongly reflected good male condition given by the lipid content of the somatic muscles in the Lusitanian toadfish. Accordingly, we further demonstrate that these vocal parameters affect the mating success in this species and seem to inform receivers, that is, females and other competi ...
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... In courtship signaling contexts of cavity spawning fi shes, more specifi cally, females may gain a reproductive advantage by attending to reliable indicators of successful nest defense and a low likelihood of cannibalizing eggs (Manica 2004 ). Thus, informative acoustic signals in these fi shes could also include indices of male somatic condition that predict quality care of offspring (e.g., high fat reserves) (Amorim et al. 2009Amorim et al. , 2010). Sound pressure level and pulse dominant frequency are reliable indices of body size in a number of soniferous fi shes (e.g., Myrberg et al. 1993 ; Connaughton et al. 2000 ;). ...
... Condition factor (Fulton's K) was related positively to total drumming sound output in Pomatoschistus pictus and Pomatoschistus minutus (Amorim et al. 2013a ; Pedroso et al. 2013 ), and female P. pictus mated with males presenting high courtship effort, which corresponds with a high number of drumming sounds (Amorim et al. 2013a , b ). Similarly, Halobatrachus didactylus calling rate and calling effort were correlated with male size, lipid stores, and liver mass (Amorim et al. 2010 ; Vasconcelos et al. 2012) and positively predicted number of eggs in the nest (Vasconcelos et al. 2012). Sonic muscle mass is strongly related to body size and liver size in H. didactylus (Amorim et al. 2009 ), and similar relationships have been found between drumming muscles mass and body size and condition in cod, Gadus morhua (Rowe and Hutchings 2004 ). ...
Convergent Aspects of Acoustic Communication in Darters, Sculpins, and Gobies
Article
Darters (Perciformes, Percidae), sculpins (Perciformes, Cottidae), and gobioids (Gobiiformes, Gobioidei) exhibit convergent life history traits, including a benthic lifestyle and a cavity nesting spawning mode. Soniferous species within these taxa produce pulsed and/or tonal sounds with peak frequencies below 200 Hz (with some exceptions), primarily in agonistic and/or reproductive contexts. The reduced or absent swim bladders found in these taxa limit or prevent both hearing enhancement via pressure sensitivity and acoustic amplification of the contracting sonic muscles, which are associated with the skull and pectoral girdle. While such anatomies constrain communication to low frequency channels, optimization of the S/N (signal-to-noise) ratio in low frequency channels is evident for some gobies, as measured by habitat soundscape frequency windows, nest cavity sound amplification, and audiograms. Similar S/N considerations are applicable to many darter and sculpin systems. This chapter reviews the currently documented diversity of sound production in darters, sculpins, and gobioids within a phylogenetic context, examines the efficacy of signal transmission from senders to receivers (sound production mechanisms, audiograms, and masking challenges), and evaluates the potential functional significance of sound attributes in relation to territorial and reproductive behaviours.
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... Call duration and call rate may be more affected by body condition rather than body length. For example, in the Gulf toadfish (Opsanus beta), call rate is related to lipid reserves (Amorim et al. 2010). The fundamental frequency of C. carolinae tonal signals is dictated by the rate at which pulses are produced, which Fine 1978) found was determined by sonic muscle contraction rate and was independent of fish size in O. tau. ...
Sound production of the banded Sculpin, Cottus carolinae
Article
Full-text available
The use of sound in social interactions has been documented in several species of sculpin and may be used to convey information concerning species identity, intent, or fitness. Sound production may be a particularly important mode of communication in sculpin due to their nocturnal behavior and construction of nests in rock cavities where the efficacy of visual cues may be diminished. In this study, we describe acoustic signals produced during conspecific behavioral interactions in Cottus carolinae, and compare signal characteristics to the only other known sounds producers in the genus Cottus including C. gobio, C. rhenanus, C. perifretum, C. bairdii, and C. paulus. Sounds were produced during nocturnal encounters between cavity residents and intruders, and consisted of pulsatile signals that were arranged into pulse trains with remarkably long pulse intervals of approximately 1 s, as well as tonal sounds in which the periodicity was increased to approximately 60 Hz. We found no relationship between fish standard length and pulse instantaneous frequency or total pulse train duration, but did find a positive correlation between fish standard length and pulse duration, as well as position of pulse interval within the pulse train. We found that C. carolinae pulse train structure differed significantly from that of other published sculpin species, and that C. carolinae produce tonal sounds; a characteristic unique within the genus Cottus.
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... Additional stimulation by the female, or reciprocal interactions between the male and the female, for example through acoustic signals in the dark, may be necessary. Indeed, acoustic communication is an important modality in fish reproduction (for examples: [43][44][45], and it will be interesting to investigate its relevance in the reproductive behavior of cave and surface morphs of A. mexicanus. S3 Movie. ...
Breeding behavior in the blind Mexican cavefish and its river-dwelling conspecific
Article
Full-text available
Fish reproductive patterns are very diverse in terms of breeding frequency, mating system, sexual dimorphisms and selection, mate choice, spawning site choice, courtship patterns, spawning behaviors and parental care. Here we have compared the breeding behavior of the surface-dwelling and cave-dwelling morphs of the characiform A. mexicanus, with the goals of documenting the spawning behavior in this emerging model organism, its possible evolution after cave colonization, and the sensory modalities involved. Using infrared video recordings, we showed that cave and surface Astyanax spawning behavior is identical, occurs in the dark, and can be divided into 5 rapid phases repeated many times, about once per minute, during spawning sessions which last about one hour and involve one female and several males. Such features may constitute “pre-adaptive traits” which have facilitated fish survival after cave colonization, and may also explain how the two morphs can hybridize in the wild and in the laboratory. Accordingly, cross-breeding experiments involving females of one morphotype and males of the other morphotype showed the same behavior including the same five phases. However, breeding between cavefish females and surface fish males was more frequent than the reverse. Finally, cavefish female pheromonal solution was able to trigger strong behavioral responses in cavefish males–but not on surface fish males. Lastly, egg production seemed higher in surface fish females than in cavefish females. These results are discussed with regards to the sensory modalities involved in triggering reproductive behavior in the two morphs, as well as its possible ongoing evolution.
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... Toadfish may synchronize their calling with the tides, as increasing calling efforts when sound propagation was most favourable (during high tides) could improve chances of being detected by mates. Barimo and Fine (1998) observed that male oyster toadfish increased calling rates during flowing tides, and Amorim et al. (2010) noted that male Lusitanian toadfish reduced calling effort during low tides. We had no observations of how many fish were calling, and thus could not calculate individual call rates. ...
Spatial and temporal variation in toadfish ( Opsanus tau ) and cusk eel ( Ophidion marginatum ) mating choruses in eelgrass ( Zostera marina ) beds in a shallow, temperate estuary
Article
Passive acoustic recordings were made at two sites over a four-month period in eelgrass beds in a shallow estuary (Shinnecock Bay, New York, USA). Recordings were dominated by mating calls of striped cusk eels (Ophidion marginatum) at one site, and oyster toadfish (Opsanus tau) mating calls at the other. Cusk eel call characteristics (frequency and pulse period) varied significantly with time and water temperature. Fundamental frequency of toadfish calls decreased over the recording period and was not correlated with water temperature. We developed and tested automated detection algorithms to identify choruses using band-limited sound pressure levels. Distinct diel peaks in sound production were observed, with cusk eels producing morning and evening choruses, and toadfish calling mostly during daytime. Several physical and environmental variables were significantly correlated with the presence of cusk eel and toadfish choruses such as water temperature, tide state, and moon phase. The temporal variation in sound production and call characteristics differed from other studies, suggesting geographical variations in the acoustic behaviour of both species. Passive acoustic techniques can identify the location and timing of reproductive events for cryptic species that live in shallow water (<2 m) habitats, which are critical information for identification of their habitat.
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... The absence of dives containing vertical lunges during song bouts, coupled with the shallow production depths of repetitive song phrases, supports the hypothesis that feeding and singing behaviours in blue whales may be mutually exclusive states [32,33,41,79]. As song has only been recorded from male blue whales and is thought to be associated with reproduction [31,33], the repetitive production of phrases to form long song bouts in the absence of foraging could also be used as an indicator of the singing male's condition to potential mates [80][81][82]. Contrary to the consistent diving behaviour exhibited by singing blue whales, the behaviours that we observed from tagged individuals producing singular calls were much more variable, which suggests that singular A and/or B calls may have a distinct behavioural purpose from A and B sounds produced as units within phrases. Blue whales in both the Pacific and Atlantic have been shown to exhibit different behaviours when producing singular calls versus singing, with the former more frequently engaging in feeding, milling and resting, and the latter in travelling behaviours [33,83]. ...
Context-dependent variability in blue whale acoustic behaviour
Article
Full-text available
  • Aug 2018
Acoustic communication is an important aspect of reproductive, foraging and social behaviours for many marine species. Northeast Pacific blue whales (Balaenoptera musculus) produce three different call types—A, B and D calls. All may be produced as singular calls, but A and B calls also occur in phrases to form songs. To evaluate the behavioural context of singular call and phrase production in blue whales, the acoustic and dive profile data from tags deployed on individuals off southern California were assessed using generalized estimating equations. Only 22% of all deployments contained sounds attributed to the tagged animal. A larger proportion of tagged animals were female (47%) than male (13%), with 40% of unknown sex. Fifty per cent of tags deployed on males contained sounds attributed to the tagged whale, while only a few (5%) deployed on females did. Most calls were produced at shallow depths (less than 30 m). Repetitive phrasing (singing) and production of singular calls were most common during shallow, non-lunging dives, with the latter also common during surface behaviour. Higher sound production rates occurred during autumn than summer and they varied with time-of-day: singular call rates were higher at dawn and dusk, while phrase production rates were highest at dusk and night.
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... Several factors influence the acoustic behavior of toadfish species. These include individual traits such as age and physical condition (Mitchell et al. 2008;Amorim et al. 2010;Vasconcelos et al. 2015), environmental conditions such as season (Breder 1968;Fine 1978;Wall et al. 2013), lunar phase (Rice and Bass 2009;Maruska and Mensinger 2009) and the presence of predators (Remage-Healey et al. 2006), and social conditions such as the call rate of neighboring males (Fish and Offutt 1972;Remage-Healey and Bass 2005) and avoidance of call overlap (Winn 1972;Fish 1972; Thorson and Fine 2002b). Social factors are of particular importance given that hearing the calls of neighboring fish may be necessary to induce calling in an individual (Remage-Healey and Bass 2005), and reproductive success is highly dependent on calling behavior (Vasconcelos et al. 2012). ...
Acoustic communication in the Bocon toadfish (Amphichthys cryptocentrus)
Article
Full-text available
The soundscapes of many coastal habitats include vocalizations produced by species of the family Batrachoididae (toadfish and midshipman). We describe the calling and grunting behavior of male Amphichthys cryptocentrus, a tropical toadfish, and predict how these vocalizations are influenced by conspecifics. We recorded individual males, which produced broadband grunts and multi-note, harmonic “boatwhistle” calls. Grunts were either in combination with calls or stand-alone. We used a null model to test if these latter grunts were produced at random or in response to calls from conspecifics. The model supports the hypothesis that grunts were in response to calls from neighboring males, suggesting acoustic competition. Using the most conservative estimate of hearing abilities we predict that males responded to the second harmonic of neighbor’s calls (230 Hz) at amplitudes of approximately 100–125 dB re 1μPa2/Hz. We also observed that call and grunt rates increased when males were exposed to higher rates of acoustic activity from neighboring fish. Fish used grunts to respond to background calls that occurred at different amplitudes, suggesting they responded to the calls of multiple neighboring fish and not just the highest amplitude neighbor. This communication with multiple fish within hearing range suggests a communication network in which the spatial distribution of individual toadfish relative to one another will impact their vocal behavior. Thus, the density and distribution, and not just abundance, of these toadfish at a given site will influence the characteristics of the chorus and the role of this species in the local soundscape.
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... It means the size effect on the call frequency is minor and is most probably not important in the fish biology. In this case, the female choice of partner would not be related to the call frequency but to the number of calls produced per time unit or the number of pulses per call (Amorim et al., 2010Vasconcelos et al., 2012;Pedroso et al., 2013). This kind of sexual selection does not favour variation in the frequency or pulse period. ...
Consistency of spatiotemporal sound features supports the use of passive acoustics for long‐term monitoring
Article
Full-text available
  • Jul 2017
Many studies stress the usefulness of fish calls as effective indicators of distinct species occurrence. However, most of these studies have been undertaken in a given area and during restricted periods of time. There is a need to show passive acoustic monitoring is a reliable method to study vocal species over space and time. This study aims to use passive acoustic methods to follow the brown meagre Sciaena umbra at relevant temporal and spatial scales. Specimens of S. umbra were recorded in both aquarium and in the field. In situ recordings were made at two regions (Corsica and Sardinia) during four summers (2008–2012–2013–2015). Temporal and frequency parameters of the fish calls were collected by different teams and compared to test the ability to unequivocally identify the fish sound. The comparison between our data and the bibliography highlights the capability to identify S. umbra during a period of 17 years in different Mediterranean regions, clearly supporting the usefulness of acoustic monitoring to discover and protect aggregation sites of this endangered species. The sound producing mechanism in S. umbra consists of high-speed sonic muscles surrounding dorsally the posterior end of the swim bladder, which can explain the low acoustic variability that helps in the species identification. Similar mechanisms are found in other Sciaenidae, suggesting that a similar conclusion can be drawn for many other adult sciaenids that could be used as sentinel species. This study should be of high interest to policymakers and scientists because it shows passive acoustic can be confidently used in resource management.
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Concentration and Location of Metabolic Substrates in Fast Toadfish Sonic Muscle
Article
Full-text available
We measured glycogen, lipid and water concentrations in toadfish (Opsanus tau) fast sonic muscle and white body muscle, and glycogen and lipid were localized histochemically in the sonic muscle. Sonic muscle had 5.05 μg of glycogen per mg wet weight, 15.64% fat and 81% water. These values were 2.2, 6.2 and 0.18 times higher than the corresponding concentrations in white body muscle. The presence of abundant glycogen and fat provides potential fuel for anaerobic and aerobic metabolism and agrees with the designation of sonic muscle as fast oxidative glycolytic. No sexual or ontogenetic differences were present in glycogen, fat or water concentrations in sonic or body muscle, and these molecules cannot be used to explain sexual differences in sound production. Glycogen is localized in sarcoplasm under the sarcolemma, in sarcoplasmic reticulum and throughout the sarcoplasmic core. Lipids have a similar distribution, except that their distribution in the core is restricted to the periphery. These findings are in accord with ultrastructure of sonic muscle fibers.
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Crepuscular Changes in Emission Rate and Parameters of the Boatwhistle Advertisement Call of the Gulf Toadfish, Opsanus Beta
Article
Full-text available
  • Mar 2002
We quantified crepuscular variation in the emission rate and call properties of the boatwhistle advertisement call of Gulf toadfish, Opsanus beta, from a field recording of a natural population of nesting males in the Florida Keys. Their calls are more variable and complex than previously reported. A call typically starts with a grunt followed by one to five tonal boop notes (typically two or three) and lasts for over a second. The first boop is considerably longer than later ones, and intervals between boops are relatively constant until the final interval, which approximately doubles in duration. Positions of fish are fixed and calls are sufficiently variable that we could discern individual callers in field recordings. Calling rate increases after sunset when males tend to produce shorter calls with fewer notes. Analysis by number of notes per call indicates some individuals decrease the number of initial grunts and the duration of the first note, but most of the decrease results from fewer notes. To our knowledge this sort of call plasticity has not been demonstrated before in fishes. We suggest that call shortening lowers the chances of overlapping calls of other males and that the small amount of time actually spent producing sound (total on time) is an adaptation to prevent fatigue in sonic muscles adapted for speed but not endurance.
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Sound production by the Lusitanian toadfish, Halobatrachus didactylus
Article
Full-text available
Several batrachoidids have been known to produce sounds associated with courtship and agonistic interactions, and their repertoires have been studied acoustically and behaviourally. In contrast, sound production of the Lusitanian toadfish Halobatrachus didactylus, although often noted, has not been acoustically studied.This sedentary predator of Northeastern Atlantic coastal waters is usually found in sandy and muddy substrates, under rocks or crevices. Sound recordings were made in Ria Formosa, a lagoon complex in southern Portugal. The sound producing apparatus was studied in adult individuals of both sexes captured by local fishermen.It is shown that this species produces acoustic emissions similar to other batrachoidids. It produces a long, rhythmical, tonal sound, often in choruses, which is comparable to the boatwhistle or hum signals of Opsanus and Porichthys, and a complex of signals that were classified as grunts, croaks, double croaks and mixed calls (‘grunt-croak’). As in other toadfishes, H. didactylus presents sonic muscles connected to a bi-lobed swimbladder. Asynchronous contractions of the sonic muscles were detected when massaging the ventral surface of the fish.
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Do song-phrase production rate and song versatility honestly communicate male parental quality in the Gray Catbird?
Article
Sexual selection theory proposes that elaborate male secondary sexual characteristics, including complex song, may increase the attractiveness of males by honestly communicating to females their genetic quality or ability to provide material reproductive resources such as parental care. The Gray Catbird (Dumetella carolinensis) is sexually monochromatic, but males sing complex songs during the breeding season while females do not. We tested the hypotheses that song-phrase versatility and rate of song-phrase production are honest indicators of male parental effort. We predicted that both song-phrase versatility and rate of production would be positively correlated with paternal chick feeding rate. Paternal chick feeding rate was not significantly related to song-phrase versatility, but it was positively and significantly correlated with song-phrase production rate. Thus, song-phrase production rate may serve as a more reliable indicator of male parental quality than song versatility in the Gray Catbird.
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Williams’ Principle: An Explanation of Parental Care in Teleost Fishes
Chapter
  • Jan 1986
Parental care may be defined as an association between parent and offspring after fertilisation that enhances offspring survivorship. This phenomenon has attracted the attention of evolutionary biologists since Darwin; however, it was not until the recent insurgence of behavioural ecology and sociobiology (e.g. Williams 1966a; Trivers 1972; Alexander 1974; Wilson 1975) that the variety of parental care patterns in animals has attracted such rigorous study. Perhaps because we ourselves are mammals, we tend to think of parental care as being the principal occupation of females, possibly with some help from males. A survey of the vertebrates, however, reveals that mammals are merely at one end of the spectrum, with predominantly female care, and fishes are at the other end, with predominantly male care (Table 11.1; see also Chapter 10 by Turner, this volume). Within teleost fishes with external fertilisation (about 85 per cent of all teleost families), one finds that the four states of parental care, ranked in descending order of their frequencies, are: no care, male care, biparental care, and female care. This seemingly peculiar trend has attracted considerable attention from evolutionary biologists, who have proposed several hypotheses about the origins of parental care in fishes (see reviews by Maynard Smith 1977; Blumer 1979; Perrone and Zaret 1979; Baylis 1981; Gross and Shine 1981; Gross and Sargent 1985).
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Female European starling preference and choice for variation in conspecific male song
Article
  • Jan 2000
Data from several field studies support the hypothesis that female European starlings, Sturnus vulgaris, attend to variation among the songs of conspecific males when making mate-choice decisions. However, for a variety of methodological reasons, direct evidence for female preferences based on song in starlings has been lacking. This study presents a novel technique for assaying directly female preference and choice in European starlings by using the presentation of conspecific male song as an operant reinforcer in a controlled environment. Using an apparatus in which the playback of songs from different nestboxes is under the operant control of the subject, we demonstrate how the reinforcing properties of conspecific song can be used to measure female preference and choice. The results of the study suggest three conclusions. First, female starlings prefer naturally ordered conspecific male songs over reversed songs. Second, female starlings display robust preferences for longer compared with shorter male song bouts. Behaviour in the operant apparatus varied directly with male song bout length. Third, preferences based on song bout length are sex specific. Male starlings failed to respond differentially to the same stimuli for which females showed strong preferences. These results suggest that male-male variation in song bout length is important for mate choice among starlings. In addition, we detail the use of a novel behavioural assay for measuring female preferences that can be applied to similar behaviours in other species of songbirds.
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