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Marc Naguib

    Marc Naguib

    ABSTRACT Birdsong is among the most well-studied biological models and attracts attention from a broad range of biological and non-biological disciplines. Across species, there is striking variety in the complexity of song and its... more
    ABSTRACT Birdsong is among the most well-studied biological models and attracts attention from a broad range of biological and non-biological disciplines. Across species, there is striking variety in the complexity of song and its delivery. In some species, song can be a simple repetition of few basic notes, but in others there can be hundreds of different song patterns arranged in complex manners. Within this structural diversity, song functions as a general advertisement signal to attract mates and mark a territory against same-sex competitors. Individual differences in singing often provide very specific information about origin, motivation, and phenotypic and genetic differences among individuals. Like human speech, birdsong develops under the influence of vocal learning during an early sensitive period, and during a period of motor practice it advances into a highly complex signal. Such parallels have established birdsong as the animal model system for vocal learning for comparison with human speech.
    Animal communication often occurs in communication networks in which multiple signalers and receivers are within signaling range of each other. In such networks, individuals can obtain information on the quality and motivation of... more
    Animal communication often occurs in communication networks in which multiple signalers and receivers are within signaling range of each other. In such networks, individuals can obtain information on the quality and motivation of territorial neighbors by eavesdropping on their signaling interactions. In songbirds, extracting information from interactions involving neighbors is thought to be an important factor in the evolution of strategies of territory defense. In a playback experiment with radio-tagged nightingales Luscinia megarhynchos we here demonstrate that territorial males use their familiar neighbors’ performance in a vocal interaction with an unfamiliar intruder as a standard for their own response. Males were attracted by a vocal interaction between their neighbor and a simulated stranger and intruded into the neighbor’s territory. The more intensely the neighbor had interacted with playback, the earlier the intrusions were made, indicating that males eavesdropped on the ...
    ... er al., 1992), Heuglin's robin chats (Cossypha heuglini; Todt et al., 198l), plain 272 DIETMAR TODT AND MARC NAGUIB titmice (Parus inornutus; Dixon, 1969), and calls of red-winged blackbirds (Beletsky et al., 1986), song sparrows... more
    ... er al., 1992), Heuglin's robin chats (Cossypha heuglini; Todt et al., 198l), plain 272 DIETMAR TODT AND MARC NAGUIB titmice (Parus inornutus; Dixon, 1969), and calls of red-winged blackbirds (Beletsky et al., 1986), song sparrows (Kramer and Lemon, 1983; Kramer et aL ...
    Recently, Lévy walks have been put forward as a new paradigm for animal search and many cases have been made for its presence in nature. However, it remains debated whether Lévy walks are an inherent behavioural strategy or emerge from... more
    Recently, Lévy walks have been put forward as a new paradigm for animal search and many cases have been made for its presence in nature. However, it remains debated whether Lévy walks are an inherent behavioural strategy or emerge from the animal reacting to its habitat. Here, we demonstrate signatures of Lévy behaviour in the search movement of mud snails (Hydrobia ulvae) based on a novel, direct assessment of movement properties in an experimental set-up using different food distributions. Our experimental data uncovered clusters of small movement steps alternating with long moves independent of food encounter and landscape complexity. Moreover, size distributions of these clusters followed truncated power laws. These two findings are characteristic signatures of mechanisms underlying inherent Lévy-like movement. Thus, our study provides clear experimental evidence that such multi-scale movement is an inherent behaviour rather than resulting from the animal interacting with its en...
    For many animals, long-range signalling is essential to maintain contact with conspecifics. In territorial species, individuals often have to balance signalling towards unfamiliar potential competitors (to solely broadcast territory... more
    For many animals, long-range signalling is essential to maintain contact with conspecifics. In territorial species, individuals often have to balance signalling towards unfamiliar potential competitors (to solely broadcast territory ownership) with signalling towards familiar immediate neighbours (to also maintain so-called "dear enemy" relations). Hence, to understand how signals evolve due to these multilevel relationships, it is important to understand how general signal traits vary in relation to the overall social environment. For many territorial songbirds dawn is a key signalling period, with several neighbouring individuals singing simultaneously without immediate conflict. In this study we tested whether sharing a territory boundary, rather than spatial proximity, is related to similarity in dawn song traits between territorial great tits (Parus major) in a wild personality-typed population. We collected a large dataset of automatized dawn song recordings from 72 ...
    Research Interests:
    Most studies on sexual selection focus on male characteristics such as male song in songbirds. Yet female vocalizations in songbirds are growing in interest among behavioural and evolutionary biologists because these vocalizations can... more
    Most studies on sexual selection focus on male characteristics such as male song in songbirds. Yet female vocalizations in songbirds are growing in interest among behavioural and evolutionary biologists because these vocalizations can reveal the female's preferences for male traits and may affect male display. This study was designed to test whether male song performance influences the different female signals in the domestic canary ( Serinus canaria ). Female canaries were exposed to three types of song performance, differing in the repetition rate of sexy syllables. This experiment demonstrates that female birds are engaged in multimodal communication during sexual interaction. The results support the copulation solicitation hypothesis for female-specific trills: these trills were positively correlated and had a similar pattern to the copulation solicitation displays; responses were higher to the songs with higher performance and responses decreased with the repetition of the ...
    In most animals, communication plays a central role in a variety of contexts. In this chapter, we synthesize studies on vocal communication and spatial behavior in nightingales, Luscinia megarhynchos, with other research on songbirds to... more
    In most animals, communication plays a central role in a variety of contexts. In this chapter, we synthesize studies on vocal communication and spatial behavior in nightingales, Luscinia megarhynchos, with other research on songbirds to emphasize the need to integrate studies on communication with spatial and movement data to be able to understand communication in a dynamic social and communication
    Many animals use long-range signals to compete over mates and resources. Optimal transmission can be achieved by choosing efficient signals, or by choosing adequate signalling perches and song posts. High signalling perches benefit sound... more
    Many animals use long-range signals to compete over mates and resources. Optimal transmission can be achieved by choosing efficient signals, or by choosing adequate signalling perches and song posts. High signalling perches benefit sound transmission and reception, but may be more risky due to exposure to airborne predators. Perch height could thus reflect male quality, with individuals signalling at higher perches appearing as more threatening to rivals. Using playbacks on nightingales (Luscinia megarhynchos), we simulated rivals singing at the same height as residents, or singing three metres higher. Surprisingly, residents increased song output stronger, and, varying with future pairing success, overlapped more songs of the playback when rivals were singing at the same height than when they were singing higher. Other than expected, rivals singing at the same height may thus be experienced as more threatening than rivals singing at higher perches. Our study provides new evidence t...
    Any signal must get from a sender to a receiver if information is to be transmitted. In the case of bird song, the acoustic properties of the habitat may hinder this being achieved. However, birds as senders and receivers have evolved... more
    Any signal must get from a sender to a receiver if information is to be transmitted. In the case of bird song, the acoustic properties of the habitat may hinder this being achieved. However, birds as senders and receivers have evolved numerous adaptations to overcome the problem of getting the message across. In this chapter, we explore habitat-dependent patterns of
    SUMMARY Female birds incorporate in the yolks of their eggs significant concentrations of a number of different androgens. Yolk androgen has been shown to positively affect several fitness components at the embryo, nestling and juvenile... more
    SUMMARY Female birds incorporate in the yolks of their eggs significant concentrations of a number of different androgens. Yolk androgen has been shown to positively affect several fitness components at the embryo, nestling and juvenile stages. Previous experiments have shown that females lay eggs with higher androgen concentrations when they are paired with highly ornamented males. This pattern suggests that yolk androgens are costly to females. In this study, we experimentally manipulated adult female condition in zebra finches Taeniopygia guttata by modifying the level of developmental stress they suffered as nestlings. This was achieved by cross-fostering nestlings to broods of varying brood size. Subsequently, we measured the yolk testosterone contents of the female offspring that resulted from the experimental manipulation. As predicted, females deposited decreasing concentrations of testosterone with increasing brood sizes experienced as nestlings: testosterone concentration ...
    Environmental factors play a key role in the expression of phenotypic traits and life-history decisions, specifically when they act during early development. In birds, brood size is a main environmental factor affecting development.... more
    Environmental factors play a key role in the expression of phenotypic traits and life-history decisions, specifically when they act during early development. In birds, brood size is a main environmental factor affecting development. Experimental manipulation of brood sizes can result in reduced offspring condition, indicating that developmental deficits in enlarged broods have consequences within the affected generation. Yet, it is unclear whether stress during early development can have fitness consequences projecting into the next generation. To study such trans-generational fitness effects, we bred female zebra finches, Taeniopygia guttata , whose mothers had been raised in different experimental brood sizes. We found that adult females were increasingly smaller with increasing experimental brood size in which their mother had been raised. Furthermore, reproductive success at hatching and fledging covaried negatively with the experimental brood size in which their mothers were ra...
    Formal models have shown that diel variation in female mate searching is likely to have profound influence on daily signalling routines of males. In studies on acoustic communication, the temporal patterns of the receivers' signal... more
    Formal models have shown that diel variation in female mate searching is likely to have profound influence on daily signalling routines of males. In studies on acoustic communication, the temporal patterns of the receivers' signal evaluation should thus be taken into account when investigating the functions of signalling. In bird species in which diel patterns of signalling differ between males singing to defend a territory or to attract a mate, the diel patterns of mate and territory prospecting are suggested to depend on the sex of the prospector. We simulated newly arriving female nightingales ( Luscinia megarhynchos ) by translocating radio-tagged females to our study site. The mate-searching females prospected the area mostly at night, visiting several singing males. The timing of female prospecting corresponded to the period of the night when the singing activity of unpaired males was higher than that of paired males. In contrast to females, territory searching males have ...
    In animal communication, elaborate signals have been shown to be under sexual selection and often to reliably indicate a signaler's quality, condition, or motivation. For instance, the performance of physically challenging signals... more
    In animal communication, elaborate signals have been shown to be under sexual selection and often to reliably indicate a signaler's quality, condition, or motivation. For instance, the performance of physically challenging signals such as trills – i.e. rapidly repeated elements of broad frequency bandwidth – is considered to reflect signaler quality. Nightingales Luscinia megarhynchos are renowned for their outstanding song repertoire sizes, and most songs include a variety of complex trills. In the present study, we examined whether performance of trills can reliably reflect male quality. We show that vocal performance of trills predicts the age of a male. Older males sang trills that were closer to the performance limit than did younger males. Moreover, males with narrower beaks sang more consistent trills than did males with wider beaks. Vocal performance of trills, however, did not significantly predict other measures of biometric quality such as body size or body condition ...
    To investigate aspects of neighbor recognition in a migratory bird with large and complex song repertoires we conducted playback experiments in the field with male territorial Thrush Nightingales Luscinia luscinia as subjects. Males sing... more
    To investigate aspects of neighbor recognition in a migratory bird with large and complex song repertoires we conducted playback experiments in the field with male territorial Thrush Nightingales Luscinia luscinia as subjects. Males sing with immediate variety and neighbors can share a considerable part of their repertoire of song components and song types. Subjects responded significantly more to their neighbors' song broadcast from the unshared (opposite) territorial boundary than to playback of the same song broadcast from the shared territorial boundary. Thus, this study did not detect constraints on neighbor recognition imposed for instance by large and complex song repertoires, high singing versatility, or sharing of song patterns. Because Thrush Nightingales sing shared patterns in different ways, either by constructing different song types out of the shared pool of elements or by singing shared song types in different sequential order, they could attend to song components and their syntactical arrangement, full songs, or their sequential delivery for neighbor recognition. Which mechanism they use might depend on the similarity of the song and singing style of a singer compared to other familiar conspecifics.
    In many bird species the sex ratio of adults is male‐biased, which is likely to have consequences for the ecology as well as for the conservation of a species. For example, when some males remain unpaired in a population, there should be... more
    In many bird species the sex ratio of adults is male‐biased, which is likely to have consequences for the ecology as well as for the conservation of a species. For example, when some males remain unpaired in a population, there should be strong selection on behavioural traits that enhance pairing success. A surplus of males is also likely to have important implications for the interpretation of breeding bird survey data. In our study population of Nightingales Luscinia megarhynchos, about half of the males stayed unpaired, suggesting that the number of males encountered singing was greater than the number of breeding pairs. Furthermore, the detectability (the probability of encountering a male singing) of mated males was only two‐thirds that of unmated males when censused in the morning or late in the breeding season. The relative detectability was more similar early in the season and during the twilight periods before sunrise and after sunset. Males that arrived earlier on the bree...
    ABSTRACT Variation of signals is a widespread feature in animal communication. Signals usually vary among signallers and in some cases vary across geographic areas. Individual variation provides the basis for individual recognition and... more
    ABSTRACT Variation of signals is a widespread feature in animal communication. Signals usually vary among signallers and in some cases vary across geographic areas. Individual variation provides the basis for individual recognition and thus has important implications for social interactions. Microgeographic variation can reveal insights into patterns of dispersal and into timing and mechanism of acquisition or development of behavioral traits. In addition, it may reflect adaptations of signals to the transmission characteristics of local habitat types. In song birds these kinds of variation have been documented in particular for male song. Here, we examined individual signature cues, site-specific variation, and variation among habitat types in tonal contact calls of 26 chiffchaffs, Phylloscopus collybita canarensis, recorded on two Canary islands. Multiparametric analyses of calls and subsequent discriminant function analyses revealed clear individual differences as well as microgeographic variation in call structure. Call structures differed not only between islands but also among the different locations on Tenerife, indicating that individuals on the same island do not share the same call, as suggested earlier. Calls recorded in different types of habitat, however, did not differ in structure as predicted by the ‘acoustic adaptation hypothesis’. The findings indicate that individual recognition may be possible on the basis of calls and they contribute to the understanding of development of behavioural trails in relation to life history patterns such as the timing and pattern of dispersal.
    In animal communication, sexually selected signals have been shown to often signal individual attributes such as motivation or quality. Birdsong is among the best studied signalling systems, and song traits vary substantially among... more
    In animal communication, sexually selected signals have been shown to often signal individual attributes such as motivation or quality. Birdsong is among the best studied signalling systems, and song traits vary substantially among individuals. The question remains if variation in signalling also reflects more general and consistent individual characteristics. Such consistent individual differences in behaviour that are relatively stable over time and contexts are referred to as personality or behavioural syndromes. Here, we studied the relation between singing and explorative behaviour, a well‐studied personality trait, using great tits (Parus major) under standardized aviary conditions. The results show that singing activity measured as the number of songs sung in spring prior to breeding correlated with male but not with female explorative behaviour. In contrast, song repertoire was not related to explorative behaviour but varied over the day. The link between explorative and sin...
    In communication, animals often use complex signals with different traits carrying different information. In the song of some songbirds, both trills and song overlapping signal arousal or the readiness to escalate a contest in male‐male... more
    In communication, animals often use complex signals with different traits carrying different information. In the song of some songbirds, both trills and song overlapping signal arousal or the readiness to escalate a contest in male‐male interactions, yet they also differ inherently from each other. Song overlapping is restricted to interactions and has a clear directive function as the songs are timed specifically to the songs of a counterpart. Trills, however, can be used without opponents actively singing and do not have such a directional character unless when combined with directed traits. This difference raises the question whether trills can enhance the agonistic function of song overlapping when being used simultaneously. Here, we exposed male nightingales (Luscinia megarhynchos) prior to pairing to overlapping playback treatments differing in the presence or absence of rapid broadband trills. Males responded differently to the two playback treatments suggesting that song ove...
    Environmental conditions during early development play a crucial role in shaping an organism's phenotype. To test how social group size affects stress response and behavioral characteristics, we used great tits (Parus major) from... more
    Environmental conditions during early development play a crucial role in shaping an organism's phenotype. To test how social group size affects stress response and behavioral characteristics, we used great tits (Parus major) from selection lines for exploratory behavior, a proxy for an avian personality trait, and birds from the wild in a brood size manipulation experiment. Nestlings were tested for stress response using an established stress test and after independence subjects were tested for exploratory behavior. Nestlings from small broods showed a stronger stress response than nestlings from normal‐sized broods. Exploratory behavior was not affected by brood size but associated with sex ratio in the nest. Birds from female‐biased broods became faster explorers than those from male‐biased broods. The results demonstrate that early social conditions can affect physiological stress responses in nestlings and that behavioral personality traits measured after fledging can be aff...
    Songbirds are an important model system for the study of the neurological bases of song learning, but variation in song learning accuracy and adult song complexity remains poorly understood. Current models of sexual selection predict that... more
    Songbirds are an important model system for the study of the neurological bases of song learning, but variation in song learning accuracy and adult song complexity remains poorly understood. Current models of sexual selection predict that signals such as song must be costly to develop or maintain to constitute honest indicators of male quality. It has been proposed that reductions of nestling condition during song development might limit the expression of song learning. Adult song could thus act as an indicator of early stress as only males that enjoy good condition during development could learn accurately and sing long songs or large repertoires. We tested this hypothesis in the zebra finch by modifying early condition through cross‐fostering chicks to small, medium, and large broods. Song learning was very accurate and was found to reflect very closely tutor song characteristics and to depend on the number of males in the tutoring group. Although the brood size manipulation stron...
    The nutritional and social conditions that individuals experience during early development can have profound effects on their morphology, physiology, behaviour and life history. Experimental increases in brood size in birds can result in... more
    The nutritional and social conditions that individuals experience during early development can have profound effects on their morphology, physiology, behaviour and life history. Experimental increases in brood size in birds can result in reduced offspring condition and survival, indicating that developmental deficits in enlarged broods have negative fitness consequences within the affected generation. To study long-term effects (i.e. transgenerational effects of developmental stress), we conducted a two-step breeding experiment in which we manipulated early developmental conditions in zebra finches Taeniopygia guttata . We raised zebra finches by manipulating brood sizes and controlled for maternal and genetic effects by cross-fostering. In a previous study, we showed that offspring condition and body size decreased with increasing brood size. Here we show that this effect was carried over to the next generation. Body size in nestlings and at nutritional independence was affected by...
    Offspring signalling can serve to communicate need to the parents and thus influence parental readiness to provide care. Offspring stimuli that affect parental care have been investigated extensively. Yet much less is known about the... more
    Offspring signalling can serve to communicate need to the parents and thus influence parental readiness to provide care. Offspring stimuli that affect parental care have been investigated extensively. Yet much less is known about the mechanisms leading to a decline in maternal motivation when conflicts of provisioning current and future offspring may arise. Here we tested responses by pregnant and non-pregnant female guinea pigs (Cavia aperea f. porcellus) to playback of pup calls during their period of lactation for dependent offspring. Most concurrently pregnant and lactating females did not respond to pup calls, whereas non-pregnant lactating females responded strongly. Our findings expand on previous studies by showing that female behavioural responsiveness to pup stimuli is strongly reduced by concurrent pregnancy and lactation. These instantaneous measurements of female responsiveness to young show more directly than standard measures like nursing performance or time to weaning how female motivation to care for current offspring is diminished by simultaneous gestation.

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