Imitation and Invention in Song Learning in Nightingales (Luscinia megarhynchos B., Turdidae)

Song learning in oscine birds is often defined solely as a process of song imitation; nonetheless, not all songs produced by laboratory‐tutored birds are imitations of the model songs. If song learning were strictly a process of imitation, these non‐imitated songs (inventions) would be expected to contain no learned attributes. To determine whether species‐typical song attributes can be learned in the absence of imitation, we compared the imitations and inventions of laboratory‐tutored nightingales (Luscinia megarhynchos B.) with the songs of wild nightingales and the songs of laboratory‐reared, untutored nightingales. The species‐typical song attribute measured was stereotypy. We quantified stereotypy by four variables: (1) percentage of notes shared between two renditions of the same song type (2) difference in repetition rates of the same trill in two renditions of the same song type (3) acoustic similarity of the same note in two renditions of the same song type, and (4) acoustic similarity of the same note repeated within a trill. Wild songs and imitated songs were significantly more stereotypical than the songs of untutored birds for all measures. For the percentage of notes shared (1), and the acoustic similarity of notes in two renditions of the same song type (3), invented songs did not differ from the songs of untutored birds, suggesting that imitation is necessary for the acquisition of these song characteristics. However, invented songs were significantly more stereotypical than the songs of untutored birds for measures of stereotypy related to trills (2 and 4), and neither imitated nor invented songs differed significantly from the songs of wild birds in terms of trill rate stereotypy (2). Thus, it appears that the process of learning to produce trills may differ from the process of learning non‐repetitive song components: increased stereotypy in trills occurs even when the trills themselves are not copied from song models. Strict imitation does not fully account for the acquisition of some learned song attributes.     

Conditions Affecting Song Acquisition in Nightingales (Luscinia megarhynchos L.)

Hand raised nightingales were alternatively confronted with different series of songs which permitted labeling of particular learning situations and allowed detection of specific consequences of diverse learning conditions. We found that visual contact with a tutor affected both quality and quantity of acquired patterns. Without visual contact the birds acquired only song sections consisting of repeated vocal units', which proved to be relevant for species recognition. With visual contact the birds learnt every presented song type completely (→ song type sharing between tutor and learner). Furthermore, the birds developed additional song types individually: by distinct parameter variation, or by recombination of particular tutor song units (Fig. 1; Table 1). Functional aspects of this type of song acquisition and development are discussed.     

Spring arrival patterns of the Cuckoo Cuculus canorus,Nightingale Luscinia megarhynchos and Spotted Flycatcher Musciapa striata in Britain

Data on the arrival dates of Cuckoo, Nightingale and Spotted Flycatcher published in the Royal Meteorological Society phenological reports are presented. From these records, contour maps showing the pattern of progression through Britain were produced for each species. Of the three species, the Cuckoo arrived earliest and progressed the slowest from south to north; it entered Britain by the southeast. The Nightingale arrived later than the Cuckoo and progressed along a southeast to northwest axis. The Spotted Flycatcher was the last to arrive, but progressed with the greatest speed from south to north. Arrival dates of all three species were influenced by the temperature in southern Europe in the month preceding arrival in Britain and by the difference between these temperatures in southern Europe and those a month later in France or Britain. The diminishing population levels of Nightingale may have had a masking effect on its arrival dates. Although mean arrival dates for the three species were strongly influenced by temperature, long-term trend effects should be interpreted with caution as other factors also influence mean arrival dates.     

Status of the Nightingale Luscinia megarhynchos in Britain at the end of the 20th Century with particular reference to climate change

Capsule The population level may be unchanged but the range has contracted.

Aims To establish the current status of the Nightingale in Britain and explore causes for any changes.

Methods Over 3000 sites where Nightingales were known to have occurred since 1980 were surveyed by volunteers between mid-April and early June 1999. A selection of 135 random tetrads were also surveyed to gauge the efficiency of the volunteer survey in locating Nightingales.

Results The survey located 4565 singing male Nightingales while the random tetrad surveys suggest that c. 32% of birds occur away from known sites, increasing the estimate for the British Nightingale population to 6700 males (95% confidence limits 5600–9350) in 1999. A higher proportion of Nightingales was found in scrub (46.7%) than in 1976 (28.4%), suggesting a recent shift in habitat use.

Conclusion There is little evidence of a change in the size of the British Nightingale population, probably because earlier surveys underestimated numbers. The range has contracted markedly over the last few decades and numbers outside the core areas in southeast England are now low. Changes in habitat quality and increasing deer populations have caused decreases on a local scale. Changes in climate on the breeding grounds and general changes in climate or habitat suitability on the African winter quarters are likely to be important in influencing the distribution within England. Models of the effects of future climate change on Nightingale distribution in Britain predict that numbers and range should increase over the next few decades.     

Memorization and reproduction of songs in nightingales (Luscinia megarhynchos): evidence for package formation

Summary To examine the properties of avian song memorization, handraised nightingales (Lusdnia megarhynchos) were exposed to three strings composed of 21 or 12 different song types (Fig. 1) which they heard only during the tutoring programs. The analysis of the birds' singing performed several months later revealed a kind of song memorization and production which, hitherto, had not been studied in song birds and is called package formation.Each package was a temporally consistent group of acquired song types which could be traced back to a coherent succession of, usually, three to five (max. seven) model song types. Model song types which a bird failed to acquire were located outside those sequences of model song types that had been acquired and later formed the packages. In contrast to the order of the tutored strings, packages were sung in a variable order. In parallel, neither the frequency of occurrence of the packages nor the performance length was related to the size of the package.We infer from these results that package formation reflects basic properties of the nightingales' song-type memories, which — through segmentation of serially presented learning stimuli — may facilitate the acquisition of large signal repertoires.     

First-Year Common Nightingales (Luscinia megarhynchos) Have Smaller Song-Type Repertoire Sizes Than Older Males

Based on the assumptions that birdsong indicates male quality and that quality is related to age, one might expect older birds to signal their age. That is, in addition to actual body condition, at least some song features should vary with age, presumably towards more complexity. We investigated this issue by comparing repertoire sizes of free‐ranging common nightingale males in their first breeding season with those of older males. Nightingales are a good model species as they are open‐ended learners, where song acquisition is not confined to an early sensitive period of learning. Moreover, nightingales develop an extraordinarily large song‐type repertoire (approx. 180 different song types per male), and differences in repertoire size among males are pronounced. We analysed repertoire characteristics of the nocturnal song of nine nightingales in their first breeding season and compared them with the songs of nine older males. The repertoire size of older males was on average 53% larger than that of yearlings. When analysing two song categories of nightingales, whistle and non‐whistle songs separately, we found similar results. Our findings show marked differences in repertoire size between age categories, suggesting that this song feature may reflect a male's age. We discuss those mechanisms that may constrain the development of larger repertoires in first‐year males. Whether repertoire sizes are crucial for female mate choice or in vocal interactions among conspecific males remains open to further investigations.     

Seasonal changes in the nocturnal singing routines of Common Nightingales Luscinia megarhynchos

Male Common Nightingales Luscinia megarhynchos famously sing at night. There are two distinct types of nocturnal singing routine (the dusk-to-dawn pattern of variation in song rate): (1) dusk and dawn choruses, with little or no song during the middle of the night; (2) a rapid increase in song rate after dusk, reaching a broad peak in the middle of the night, declining towards dawn, and followed by a dawn chorus. Males sing different nocturnal singing routines at different times in the breeding season. Earlier in the breeding season, most males sing Type 2 routines. Later in the breeding season, most males sing Type 1 routines, as do birds on the wintering grounds. At least some individuals may also sing Type 1 routines during the first few days after their arrival on their breeding territories, before the arrival of females. The main function of nocturnal song appears to be mate attraction. The patterns of variation in song rate over the course of the night are qualitatively similar to those predicted by stochastic dynamic programming (SDP) models of daily singing and foraging routines, for birds that do not forage at night, in circumstances when birds can pair at night (Type 2 routines), and when they cannot (Type 1 routines). The observed seasonal changes in the types of routine sung are also consistent with the predictions of these models.     

The predictive value of trill performance in a large repertoire songbird,the nightingale Luscinia megarhynchos

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 of the males. The findings suggest that receivers could benefit from the predictive value of physically demanding song traits in assessing age as an important quality component of potential mates or rivals. Particularly in species with high singing versatility, signaler assessment based on readily assessable structures may be adaptive, as this will allow receivers to quickly gather relevant information about the singer without attending to the full song repertoire.     

Simple Mechanisms Can Explain Social Learning in Domestic Dogs (Canis familiaris)

Recent studies have suggested that domestic dogs (Canis familiaris) engage in highly complex forms of social learning. Here, we critically assess the potential mechanisms underlying social learning in dogs using two problem‐solving tasks. In a classical detour task, the test dogs benefited from observing a demonstrator walking around a fence to obtain a reward. However, even inexperienced dogs did not show a preference for passing the fence at the same end as the demonstrator. Furthermore, dogs did not need to observe a complete demonstration by a human demonstrator to pass the task. Instead, they were just as successful in solving the problem after seeing a partial demonstration by an object passing by at the end of the fence. In contrast to earlier findings, our results suggest that stimulus enhancement (or affordance learning) might be a powerful social learning mechanism used by dogs to solve such detour problems. In the second task, we examined whether naïve dogs copy actions to solve an instrumental problem. After controlling for stimulus enhancement and other forms of social influence (e.g. social facilitation and observational conditioning), we found that dogs’ problem solving was not influenced by witnessing a skilful demonstrator (either an unknown human, a conspecific or the dog’s owner). Together, these results add to evidence suggesting that social learning may often be explained by relatively simple (but powerful) mechanisms.     

Song Sharing in Neighboring and Non-Neighboring Thrush Nightingales (Luscinia luscinia) and its Implications for Communication

In many passerines, males have repertoires of different songs of which some songs are often shared with other males. Sharing of song repertoires among males can provide insights into the context in which songs were acquired and on the role of song repertoires in inter- and intrasexual communication. Here we studied repertoire sharing in male territorial thrush nightingales ( Luscinia luscinia ). We compared male vocal repertoires of the basic song components, full songs, and the sequencing of songs in a bout. The results show that males differed significantly in the size of their song repertoires but not in the size of the repertoire of basic song components. Moreover, they shared almost all (80%) the repertoire of song components but only 30% of their song types. Neighboring males shared significantly more song types than did non-neighboring males but interestingly they did not share more basic song components than non-neighboring males. These results show that the repertoire of basic song components is under much less sexual selection than the size of song repertoires. Sharing of song repertoires among neighbors presumably results from repertoire conversion over time and from males returning to their territories in the following season. Repertoire sharing then can be an indicator of territory tenure and thus it can be important in repelling rivals and in female choice.     

Song Learning in the Anna Hummingbird (Calypte anna)

Songs of wild male Anna hummingbirds (Calypte anna) consist of syllables grouped into phrases. Nearest neighbors tend to share similar syllable types, rhythms and syntax. Songs from different localities contain different syllable types, syntax and repetition indices. A male raised by hand in isolation produced a song consisting of highly variable syllable types of a wide frequency range. The song was simple in structure, and syllables were not grouped into phrases. Three males raised by hand as a group sang songs containing two stereotyped syllable types sung in alternating sequence and without phrase structure. These three males shared syllable types and syntax. The data from our study indicate that despite its relatively simple syrinx the Anna hummingbird learns syllable types, frequency, rhythm and syntax (as do oscines with their more complex syringes) during the song development process.     

Nightingales Luscinia megarhynchos in scrub habitats in the southern fens of East Anglia, England: associations with soil type and vegetation structure

The southeast corner of the East Anglian Fens supports a large concentration of Nightingale Luscinia megarhynchos territories. A total of 382 territories were located in extensive surveys in 1999 and 2000, probably representing over 5% of the English population. Transect counts revealed that the Fenland population is restricted in distribution and is associated with localized thickets of scrub. The highest densities are found on humus‐rich soils, suggesting that soil type, in addition to habitat availability, may have a strong influence on the Nightingale's distribution in this region. This paper provides the first assessment of habitat requirements of the Nightingale in scrub, which now forms a principal habitat for the species in England. Detailed studies of the attributes of over 100 Nightingale territories revealed subtle differences in the vegetation structure of these thickets when compared with paired, unoccupied, but apparently similar thickets. The Nightingale territories tended to have a higher proportion of bare ground or short vegetation in the field layer under the canopy, whereas paired sites were more likely to have low field layer vegetation beneath the canopy. The bare ground within the thickets is a feature of shading beneath very dense foliage cover. Within Nightingale territories, low field layer volume and shrub twig volume at the thicket edges was higher than in unoccupied thickets. The differences detected in vegetation structure suggest that a dense and continuous canopy forming a shell over bare ground but with dense low foliage at thicket edges provides the ideal vegetation structure for Nightingales in scrub habitats. Our study suggests that Nightingales occupy scrub of a very specific structure, and specific stage in vegetation succession. This structure probably provides an optimal combination of foraging habitat, microclimate and cover from predators. It is suggested that humus‐rich soils may be preferred because they may support a particularly rich source of invertebrate food, but this remains to be tested empirically.     

斑胸草雀的鸣声学习

鸣声是鸟类重要的生物学特征和分类特征。由于生理结构和神经控制方面的差异,不同种群和同一种群的不同个体都有自己特有的鸣声特征,正是利用这些特征,鸟类才能在集群、取食、迁徙、喂雏、御敌和求偶等行为中进行通讯。作为一种能够利用语音信息进行交流的高等脊椎动物,斑胸草雀的鸣声特别是鸣啭能力获得,需要有一个适宜的学习对象,并在一段特定的时间内,经历复杂的学习过程才能实现。     

Molecular and phenotypic divergence in the bluethroat (Luscinia svecica) subspecies complex

Subspecies complexes may provide valuable insights into the early stages of the speciation process. The bluethroat (Luscinia svecica) consists of many morphologically distinct subspecies that differ most strikingly in the ornamental colour pattern of the male throat. We investigated the genetic and phenotypic differentiation in this subspecies complex, using (i) microsatellite genotyping (11 loci) of a sample of 364 individuals from bluethroat populations in Europe and Asia, and (ii) spectrometric and morphological measurements of a sample of 131 museum skin specimens. Population genetic analyses, based on microsatellite allele frequency variation, revealed a slight but significant overall population differentiation (F(ST) = 0.042). There was a well-differentiated southern group of subspecies with white or no throat spots and a less-differentiated northern group of chestnut-spotted populations. Phylogenetic analyses indicated that the southern all-blue and white-spotted forms are ancestral to the chestnut-spotted subspecies. In addition to the qualitative variation in throat plumage pattern already described in the literature, we found significant quantitative variation among subspecies in hue, chroma and brightness of the ultraviolet (UV)/blue throat coloration, and this variation seemed to be unrelated to the phylogenetic distance between subspecies.     

Power Dissipation in the Cochlea Can Enhance Frequency Selectivity

The cochlear cavity is filled with viscous fluids, and it is partitioned by a viscoelastic structure called the organ of Corti complex. Acoustic energy propagates toward the apex of the cochlea through vibrations of the organ of Corti complex. The dimensions of the vibrating structures range from a few hundred (e.g., the basilar membrane) to a few micrometers (e.g., the stereocilia bundle). Vibrations of microstructures in viscous fluid are subjected to energy dissipation. Because the viscous dissipation is considered to be detrimental to the function of hearing—sound amplification and frequency tuning—the cochlea uses cellular actuators to overcome the dissipation. Compared to extensive investigations on the cellular actuators, the dissipating mechanisms have not been given appropriate attention, and there is little consensus on damping models. For example, many theoretical studies use an inviscid fluid approximation and lump the viscous effect to viscous damping components. Others neglect viscous dissipation in the organ of Corti but consider fluid viscosity. We have developed a computational model of the cochlea that incorporates viscous fluid dynamics, organ of Corti microstructural mechanics, and electrophysiology of the outer hair cells. The model is validated by comparing with existing measurements, such as the viscoelastic response of the tectorial membrane, and the cochlear input impedance. Using the model, we investigated how dissipation components in the cochlea affect its function. We found that the majority of acoustic energy dissipation of the cochlea occurs within the organ of Corti complex, not in the scalar fluids. Our model suggests that an appropriate dissipation can enhance the tuning quality by reducing the spread of energy provided by the outer hair cells’ somatic motility.     

Recent evolutionary history of the bluethroat (Luscinia svecica) across Eurasia

We analysed mitochondrial DNA (mtDNA) sequences from 154 bluethroats (Luscinia svecica) sampled at 21 sites throughout much of their Eurasian range. A previously reported, single base-pair mtDNA difference between L. s. svecica and L. s. namnetum was inconsistent upon expanded geographical sampling. A significant FST value (0.29) and an isolation-by-distance effect show the existence of geographical differentiation. Phylogenetic analysis of haplotypes revealed northern and southern groups, although lineage sorting is incomplete. There was no geographical structure to the haplotype tree within groups, and currently recognized subspecies were not supported. A minimum evolution tree based on pairwise mtDNA genetic distances among average samples showed the same two broadly distributed northern and southern groups. These groups abut in the centre of the latitudinal range, and were possibly isolated by forest that developed and spread westward over the last 15 000 years. Pairwise FST values averaged 0.16 in the southern group, 0.04 in the northern group, and 0.42 between groups. Mismatch distributions suggested population growth in each group, with that in the south being more recent. In the northern group, the geographical pattern in tau suggested northward and eastward expansion. Analysis of nucleotide diversity suggested westward expansion in the southern group. The northern group had higher nucleotide diversity than the southern group, consistent with a larger current population size in the north. Given the significant FST, incompletely sorted haplotype tree, and broadly patterned minimum evolution tree, L. svevica appears to represent a species at an intermediate stage of differentiation between panmixia and reciprocal monophyly.     

Broad Selectivity for Courtship Song in the Cricket Gryllus bimaculatus

Various characteristics of a long‐distance acoustic signal have been shown to vary to different degrees. It has been suggested that female preferences based on stable song parameters are stabilising or weakly directional, and preferences based on variable parameters are strongly directional. We tested this hypothesis based on a short‐distance signal (courtship song) produced by the field cricket, Gryllus bimaculatus. We studied the degree of variability of different courtship song parameters and the behavioural importance of several parameters using synthesised song models in playback experiments. We found that most of the courtship song elements of G. bimaculatus were quite variable (coefficient of variation, CV, in the range of 20–53%). The most variable parameter of the courtship song was the relative amplitude of two elements: high‐amplitude ticks and low‐amplitude pulses. Because songs containing only ticks (of rare occurrence) appeared to be more effective than songs with both ticks and pulses (of frequent occurrence), we consider female preferences to be directional. Alteration of less variable traits, such as the carrier frequency and duration of ticks (CV = 20–25%), had a different effect on female responsiveness. The synthesised songs with different carrier frequencies of ticks were as attractive to females as the positive control (courtship of muted males accompanied by playback of the recorded song). Altering the duration of ticks had a crucial effect on the female response rate, decreasing female responsiveness to the level observed in the negative control (courtship of muted males). Thus, we did not find a strong relationship between the variability of individual song parameters and their potential importance in song recognition and the evaluation of male quality. The partial inconsistency of our results with the data of other authors may be due to different patterns of past and current selection on long‐distance and short‐distance acoustic signals.     

Age-Related Variation in Mate-Guarding Intensity in the Bluethroat (Luscinia s. svecica)

Female extra‐pair copulations (EPCs) have selected for male paternity guarding strategies in many bird species. In the bluethroat, Luscinia s. svecica, males guard their mates closely during the last 2 d before the start of egg laying, but there is great individual variation in the intensity of mate guarding. Here we show that some of this variation is related to male age. Old males guarded their mates with much lower intensity and sang more than young males, although the latter difference was not statistically significant. Controlling for male age, male and female coloration and size were not significantly related to the intensity of mate guarding. We have previously shown that young and old males had a similar paternity loss in their own broods. On the other hand, old males were far more successful than young males in achieving extra‐pair fertilizations. These patterns suggest that young and old males have different trade‐offs between preventing paternity loss in own nest and gaining paternity in others, because male skills in obtaining EPCs improve with experience and/or because of female preferences for old males as copulation partners. There were no significant relationships between paternity and male mate‐guarding behaviour during the fertile period, indicating that mate guarding is not a very effective paternity‐assurance strategy in the bluethroat.