Song overproduction, selective attrition and song dialects in the white-crowned sparrow

Male Puget Sound white-crowned sparrows, Zonotrichia leucophrys pugetensis, form large vocal dialects along the Pacific northwest coast of North America. Most adult males sing a single song dialect throughout their lives. To determine how dialects are maintained in this species, I studied two populations at the Columbia River mouth in a contact zone between two dialects. Singers of each of the two dialects clustered together in space, forming local song 'neighbourhoods'. Two hypotheses for the maintenance of dialects were tested. The late acquisition hypothesis predicts that yearling or adult male immigrants memorize their song from the territory neighbours they settle next to. The selective attrition hypothesis predicts that males memorize a variety of dialects early in life, overproduce song dialects upon arrival, and later selectively retain the one prelearned dialect that best matches what their neighbours sing. In 1997 and 1998, 35-40% of new territory occupants sang two dialects upon arrival in April, and then over the course of days to several weeks, discarded one dialect from their repertoire. Fourteen of 16 (88%) kept as their adult song the dialect that matched the dialect sung by the majority of their neighbours. No male added a new dialect to his repertoire after arrival, nor did males alter their retained dialect to more closely resemble their neighbours' songs. New arrivals that overproduced dialects upon arrival were significantly more likely to match their neighbours' dialect than males that did not overproduce upon arrival. In a playback experiment, males in the overproduction stage engaged in matched countersinging to the dialect played to them. These observations and the experiment support the selective attrition hypothesis: males visit and memorize a variety of dialects, probably in their hatching-year summer, overproduce dialects upon arrival the next spring, and then selectively retain the one dialect that matches the local song culture. Vocal plasticity late in life is not the result of a late sensitive phase for song memorization, but rather results from behavioural selection operating on a pre-existing repertoire of song dialects. Copyright 2000 The Association for the Study of Animal Behaviour.     

Nonlocal male mountain white-crowned sparrows have lower paternity and higher parasite loads than males singing local dialect

Bird song often varies geographically, and when this geographicvariation has distinct boundaries, the shared song types arereferred to as song dialects. We investigated the role of songdialect in male mating success in a wild breeding populationof mountain white-crowned sparrows (Zonotrichia leucophrysoriantha). In 2 of 3 years, males singing unusual songs ("nonlocal"males) had lower total fertilization success (measured by microsatellitepaternity analysis) than did males singing the local dialect ("local" males). Similarly, females produced disproportionately more young with local than with nonlocal males. However, dialectwas not a significant predictor of male mating success whencontrolling for other factors that might affect paternity.Instead, the low mating success of nonlocal males was apparentlydue to an interaction between song dialect and parasite load.Nonlocal males were more severely infected by bloodborne Haemoproteusthan were local males, although they did not differ in anyother measured aspect of quality. Immigrant birds may be immunologically disadvantaged, possibly due to a lack of previous experiencewith the local parasite fauna, resulting in low mating success.     

Species recognition in Darwin's finches (Geospiza, Gould). III. Male responses to playback of different song types,dialects and heterospecific songs

The songs of the six different species of Darwin's ground finches (Geospiza) on the Galápagos Islands are difficult to distinguish unambiguously because of high levels of intraspecific variation and interspecific similarity in some cases. We recorded the responses of males on five islands to playback of (a) the two main conspecific song types, A and B, (b) local conspecific and heterospecific song, and (c) local and foreign dialects. Males reacted equally strongly to different conspecific song types (A and B), but responded significantly more strongly to local conspecific song than to either heterospecific song or foreign dialect. These results are inconsistent with earlier suggestions that song types subdivide Geospiza populations and that Geospiza song lacks species-distinctness because of loss-of-contrast or character convergence. The apparent paradox of low song specificity and well-developed acoustic discrimination is discussed in the light of other data showing that close-range species recognition also depends on visual cues.     

Cultural and genetic evolution in mountain white-crowned sparrows: song dialects are associated with population structure

Bird song often varies geographically within a species; when this geographic variation has distinct boundaries, the shared song types are referred to as song dialects. How dialects are produced and their adaptive significance are longstanding problems in biology, with implications for the role of culture in the evolution and ecology of diverse organisms, including humans. Here we test the hypothesis that song dialect, a culturally transmitted trait, is related to the population genetic structure of mountain white-crowned sparrows (Zonotrichia leucophrys oriantha). To address this, we compared microsatellite allele frequencies from 18 sample sites representing eight dialect regions in the Sierra Nevada. Pairwise genetic distances were not significantly correlated with geographic distances either within or between dialects, nor did dialect groups form distinct genetic groups according to neighbor-joining or UPGMA analysis, and most variation in allele frequencies occurred among individuals rather than at higher levels. However, most of the remaining variation was attributable to differences among, rather than within, dialect regions, and this among-dialect component of variance was statistically significant. Moreover, when controlling for the effect of geographic distance, song dissimilarity and genetic distance between site pairs were significantly correlated. Thus, song dialects appear to be associated with reductions in, but not strict barriers to, gene flow among dialect regions.     

Experimental evidence that distinct song phrases in the Grey‐cheeked Fulvetta Alcippe morrisonia permit species and local dialect recognition

Birds with small song repertoires have a limited number of song types which may serve multiple functions. The Grey‐cheeked Fulvetta Alcippe morrisonia is a non‐migratory species of lowland forests in Taiwan. Its song consists of two distinct phrases: a whistled phrase and a harmonic one. Each individual usually sings only one type of whistled phrase, and the geographical patterns of songs can be distinguished by the motif of this phrase. We proposed a dual adaptation hypothesis for the functions of these two phrases. Playback experiments including six sound stimuli (familiar whistled phrases, familiar harmonic phrases familiar complete songs, foreign whistled phrases, foreign harmonic phrases and foreign complete songs) were conducted at 12 sites in the Shoushan Nature Park of Kaohsiung, Taiwan. Grey‐cheeked Fulvettas came closer to the playback speaker and increased counter‐singing more in response to the familiar whistled phrase (local dialect) than to a foreign whistled phrase (distant dialect). However, birds responded equally to the playback of harmonic stimuli from familiar and foreign sources. We suggest that in this species, whistled phrases are used for local recognition and harmonic phrases are used for species recognition in short‐range communication.     

SONG INHERITANCE AND MATING IN A SONGBIRD WITH LOCAL DIALECTS

ABSTRACT

In this paper we document the pattern of geographic variation in song of the Corn Bunting in a marked population in Sussex. Song variation is best described as a system of local dialects with three song types in each dialect. We examine the inheritance of dialects from father to son; sons sing the same dialect as their nearest neighbour, rather than inheriting the dialect of the father. Therefore songs seem to be learned after dispersal. We also compare the dialects of mates and fathers of females: our results suggest that females do not rely on dialects when pairing. These results are discussed in the context of the current controversy surrounding other species with dialects and hypotheses relating dialects to the genetic structure of populations.     

What exactly is ‘local song’ in a population of ortolan buntings with a common dialect?

Previous studies have shown that ortolan buntings (Emberiza hortulana) exhibit apparent dialect variation. Neighbouring males typically share the same final phrase of a simple two-part song. Consequently, the final phrase was considered to be the dialect cue important for discriminating between males from a local population and strangers. Recently, it was shown that in an isolated and fragmented population of the ortolan buntings in Norway there was no local dialect in the above-mentioned sense. Norwegian males often had song types with different final phrases in their repertoire, and had larger repertoires and a lower level of song type sharing than in other populations. It was experimentally revealed that only local songs (L) evoked a strong response in Norwegian males, while these birds did not respond strongly to foreign (F) or hybrid songs composed of local and foreign initial and final phrases in any composition (i.e. both FL and LF songs). These results suggest that, in the Norwegian population, the final phrase of the song is not a sufficient cue for local song dialect recognition. This paper is a further study in which we tested the response of the ortolan bunting males to L, F, FL and LF songs in a typical continuous population (in Poland) in which males share the same single final phrase, i.e. have a common dialect. We found that ortolan bunting males in Poland responded with similar strength to L, FL and LF songs. The majority of males did not respond as strongly to playback of only F songs. Our results suggest that a common final phrase for a population should not be treated as the only signal of ‘locality’. Our results show clear asymmetry in response to hybrid songs with non-local phrases in different populations.     

Effects of early song experience on song preferences and song control and auditory brain regions in female house finches (Carpodacus mexicanus)

We examined the effects of song tutoring on adult song preferences, volume of song-control brain regions, and activity of auditory brain regions in female house finches (Carpodacus mexicanus). Hand-reared females were tutored with local songs, foreign songs, or no song. We then examined adult song preferences, determined the Nissl-defined volume of the song-control nuclei HVc, Area X, and RA, and compared the number of cells immunoreactive for Zenk protein in the auditory regions NCM and cmHV, following playback of songs heard early in life (Tutor/Playback Match) versus not heard (Tutor/Playback Nonmatch). All hand-reared birds exhibited preferences for locally recorded song over foreign or heterospecific song. We found no difference in the volume of song-control nuclei among the three groups. As well, we found no difference in the number of Zenk immunoreactive cells in NCM and cmHV between females in the Tutor/Playback Match group and females in the Tutor/Playback Nonmatch group. Isolate-reared birds showed greater Zenk immunoreactivity following song playback than either tutored group. Thus, early auditory experience may not play a role in adult geographic song preferences, suggesting that genetic factors can lead to preferences for songs of local dialects. Song tutoring did not influence the size of song-control regions nor Zenk induction levels following song playback, suggesting that early experience with particular songs does not influence Zenk expression. However, overall greater activation in isolate females in auditory areas suggests that exposure to song early in life may increase the selectivity of Zenk activation to song playback in auditory areas.     

Randomization Analyses: Mimicry, Geographic Variation and Cultural Evolution of Song in Brood-Parasitic Straw-Tailed Whydahs, Vidua fischeri

Although bird song has been an important model for investigating questions of behavior development, cultural evolution and population differentiation, the quantitative methods of analysis have been problematic. Here we develop and apply quantitative randomization methods to test hypotheses about these processes in a natural population of birds. Songs of the African brood-parasitic straw-tailed whydahs ( Vidua fischeri ) and songs of their host species, the purple grenadier ( Granatina ianthinogaster ), were compared in audiospectrograms for similarity to test the following hypotheses: Whydahs mimic the songs of their host species, they have local song dialects, neighboring males match their song themes, local males match the songs of local hosts, remote populations have different songs according to their geographic distance, and songs undergo cultural evolution over time across generations. Randomization analyses were completed using (1) Mantel matrix statistics and (2) tree-based measures employing Sankoff optimization of Manhattan matrices and approximate randomizations. Our results provide evidence for song mimicry, local song dialects, matching song themes between neighboring males, song matching of local whydah mimics and grenadier song models, correspondence of song differences and geographic distance, and cultural continuity with change in song traditions within a local population. These randomization methods may be useful in other studies of animal communication, and they are sufficiently general for use both with distance matrices derived either from naturalistic impressions of song similarity as in our example or from acoustic measurements.     

Variability of response of the corn bunting,Emberiza calandra,to songs of different dialects

Experiments on the corn bunting were carried out in two localities of different dialects. In the center of the territory of each bird, we played two natural songs: either one local song and one song of the foreign dialect or two local songs coming from a neighbor and from a non-neighboring individual. The following results were obtained: (1) The behavioral responses are always more intense to the local song than to the song of a foreign dialect. Thus, the behavior of the corn bunting is similar to that of species with dialects studied up to now. (2) Eighty percent of the birds tested with the song of the foreign dialect react weakly or not at all. This peculiarity, which is specific to the corn bunting, suggests limited possibilities of generalization in this species. (3) The behavioral responses to the songs of a neighbor and of a non-neighboring individual of the same locality, are aggressive and of the same intensity. This means that the song of a neighbor, emitted from a different place than usual, is perceived by the bird as a threat to its territory.     

Non‐Salient Geographic Variation in Birdsong in a Species That Learns by Improvisation

Geographic variation in birdsong and differential responses of territorial males to local and non‐local song variants have been documented in a number of songbird species in which males learn their songs through imitation. Here, we investigated geographic song variation and responses to local and non‐local song in the grasshopper sparrow (Ammodramus savannarum), a species in which males develop song by improvisation rather than imitation, as a first step toward understanding how the extent and salience of geographic song variation is related to the mode of song development. To describe the geographic variation in song, we compared songs from populations in eastern Maryland and central Ohio, USA, using multiple acoustic analysis techniques. We then conducted a playback experiment in Maryland using local and non‐local (Ohio) songs to test how territorial males responded to this geographic variation. We found acoustic differences between songs from the two sites. However, males responded similarly to playback of these songs, suggesting that this geographic variation is not behaviorally salient in a territorial context. Together with previous studies, our results suggest that across species, geographic song variation and the extent to which this variation functions in communication may be correlated with the accuracy with which song models are imitated during song development.     

High levels of gene flow among song dialect populations of the Puget Sound white‐crowned sparrow

Populations within a species can show geographic variation in behavioral traits that affect mating decisions or limit dispersal. This may lead to restricted gene flow, resulting in a correlation between behavioral variation and genetic differentiation. Populations of a songbird that differ in a learned behavioral trait, their song dialects, may also differ genetically. If song dialects function as mating barriers, evolutionary processes such as genetic drift should lead to divergence in allele frequencies among dialect populations. The Puget Sound white‐crowned sparrow (Zonotrichia leucophrys pugetensis) is an excellent study system with a well‐defined series of song dialects along the Pacific Northwest coast. A previous study found low genetic differentiation based on four microsatellite loci; however, available loci and analyses techniques have since dramatically improved and allow us to reassess gene flow in this species. We also add extra samples to fill in gaps and add a new level of analysis of geographic variation. Based on acoustic similarities, we group six song dialects into two geographically larger “northern” and “southern” song themes. One southern dialect is acoustically more similar to dialects in the north, which makes the genetic profile of birds singing this dialect particularly interesting. Traditional F‐statistics, analysis of molecular variance as well as Bayesian techniques confirmed the earlier result that geographic variation in song does not correlate with the neutral genetic structure of the sampled dialect populations. The song themes also did not differ genetically, and the origin of the extralimital northern‐theme dialect cannot be determined. We compare this result to findings in several other species and discuss how the timing of learning and dispersal allow vocalizations to vary independently of patterns of genetic divergence.     

Detailed large‐scale mapping of geographical variation of Yellowhammer Emberiza citrinella song dialects in a citizen science project

Song dialects, as a special case of geographical variation in vocalization, are useful tools in the study of a number of topics ranging from cultural evolution to the emergence of reproductive barriers, and thus continue to be the focus of many bird‐song studies. The Yellowhammer Emberiza citrinella is a common Palaearctic bird with a long breeding season and song period, whose distinctive song exhibits clearly distinguishable dialects differing in the composition of the final phrase. The species is therefore particularly suitable for large‐scale studies of dialect distribution involving not only experts but also members of the public. Here we demonstrate that citizen science can indeed allow mapping of songbird dialect distribution on a national scale in unprecedented detail. During the project Dialects of Czech Yellowhammers, which collected almost 4000 recordings over 6 years (2011–2016), we obtained information on dialect distribution for almost 90% of the territory of Czechia. We detected most of the common dialect types known elsewhere in Europe, and these were distributed in a fragmented fashion, with several unusual and rare dialects also detected in small areas. However, the highly scattered distribution of birds using some final phrases traditionally assigned to distinct dialects suggests that classification of Yellowhammer dialects warrants a thorough re‐assessment based on quantitative data. Successful involvement of the public in this and other ongoing citizen science projects has contributed to the establishment of the Yellowhammer as a model species for dialect research. The dataset will serve as a foundation for future studies investigating processes responsible for the origin and maintenance of avian dialects, and may be particularly useful for evaluating their spatial and temporal stability.     

The Response of Corn Buntings to Playback of Dialects

The response to playback of three different dialects by 10 territorial male corn buntings was investigated; the dialects played were own dialect, the dialect of a neighbouring population (NEAR) and a dialect from 40 km away (FAR). A significant effect of dialects was found for two measures of response, latency and singing rate. Response strength varies in the pattern OWN > NEAR > FAR for these measures. This pattern of response would not maintain the observed homogeneity of dialect populations. The results are similar to those obtained by experiments on other dialect species.     

A geographical perspective on the relationship between Impatiens spur lengths and bill lengths of sunbirds in Afrotropical mountains

Trait matching—a correlation between the morphology of plants and their pollinators—has been frequently observed in pollination interactions. Different intensities of natural selection in individual regions should cause such correlations to be observable across different local assemblages. In this study, we focused on matching between spur lengths of the genus Impatiens and bill lengths of sunbirds in tropical Africa. For 25 mountain and island locations, we compiled information about the composition and traits of local Impatiens and sunbird assemblages. We found that assemblage mean and maximum values of bill lengths were positively correlated with mean and maximum spur lengths across locations. Moreover, our results suggest that the positive correlations hold only for forest sunbird assemblages sharing the same habitat with Impatiens species. We further show that long‐billed sunbirds seem to locally match the morphology of multiple Impatiens plant species, not vice versa. Our observation implies that trait matching significantly contributes to structuring of Impatiens–sunbird pollination systems. We suggest that special habitat preferences together with spatial isolation of mountain environment might play a role in this case.     

Song perception among incipient species as a mechanism for reproductive isolation

Many functions in behavioural processes of small passerines are regulated via vocal messages. Song plays an important role in the development of reproductive barriers and thus playback experiments can often be used for investigating the potential for reproductive isolation through behavioural mechanisms. Moltoni's warbler Sylvia (cantillans) moltonii is characterized by diagnostic vocalizations and a peculiar pattern of distribution, being parapatric and partly sympatric with the nominate Sylvia c. cantillans. With this work, we test whether these two closely related taxa react equally to their own song and to the song of the other taxon, shedding light on whether they perceive each other's songs as coming from the same species. We carried out 184 playback experiments within the mainland range of the two forms. We judged the response of the bird on a scale of scores. Each taxon responded more strongly to playback when faced with the song of its own taxon. This held true when applied only to males or females. Additionally, birds tested for both songs showed a stronger response to the song of their own taxon. The distributional context (sympatry vs. allopatry) did not affect bird response. Results indicate that a certain degree of reproductive isolation between the two taxa (because of diverged mate recognition systems) already exists; consistently with genetic data and with the peculiar pattern of distribution; this suggests that the two taxa have reached species status.     

Geographic variation of reproductive traits and competition for pollinators in a bird‐pollinated plant

Geographic variation in the reproductive traits of animal‐pollinated plants can be shaped by spatially variable selection imposed by differences in the local pollination environment. We investigated this process in Babiana ringens (Iridaceae), an enigmatic species from the Western Cape region of South Africa. B. ringens has evolved a specialized perch facilitating cross‐pollination by sunbirds and displays striking geographic variation in perch size and floral traits. Here, we investigate whether this variation can be explained by geographic differences in the pollinator communities. We measured floral and inflorescence traits, and abiotic variables (N, P, C, and rainfall) and made observations of sunbirds in populations spanning the range of B. ringens. In each population, we recorded sunbird species identity and measured visitation rates, interfloral pollen transfer, and whether the seed set of flowers was pollen limited. To evaluate whether competition from co‐occurring sunbird‐pollinated species might reduce visitation, we quantified nectar rewards in B. ringens and of other co‐flowering bird‐pollinated species in local communities in which populations occurred. Variation in abiotic variables was not associated with geographical variation of traits in B. ringens. Malachite sunbirds were the dominant visitor (97% of visits) and populations with larger‐sized traits exhibited higher visitation rates, more between‐flower pollen transfer and set more seed. No sunbirds were observed in four populations, all with smaller‐sized traits. Sunbird visitation to B. ringens was not associated with local sunbird activity in communities, but sunbird visitation was negatively associated with the amount of B. ringens sugar relative to the availability of alternative nectar sources. Our study provides evidence that B. ringens populations with larger floral traits are visited more frequently by sunbirds, and we propose that visitation rates to B. ringens may be influenced, in part, by competition with other sunbird‐pollinated species.     

Song dialects in the yellowhammer Emberiza citrinella : bioacoustic variation between and within dialects

The yellowhammer Emberiza citrinella is a common European bird that sings in dialects that for decades have been distinguished by the existence of one single element (called a “specific”). In this study we looked into other possibilities for dialect discrimination, measuring 24 different variables. For the first time, multivariate statistics were used to discriminate dialect in yellowhammer song. Two similar dialects (XlB and XsB) that are not clearly defined in the literature were studied. Statistics incorporated (1) all variables, (2) no variables of “specific” elements, and (3) no variables under the influence of these “specific” variables. Multivariate statistics support dialect discrimination by ear and confirmed that only one element in yellowhammer song characterises dialect. In addition, we looked for local differences within two dialects and found that one local observation area showed a higher separation than the other sites (Meck1). However, as yet there is insufficient evidence for the existence of a new subdialect. The experiments comply with the current laws of the study area.     

杭州市区白头鹎鸣声的微地理差异

2002年5—8月,对中国东部浙江省杭州市区的白头鹎鸣声的微地理差异进行研究。在约60 km2的 研究区内,选择8个调查点(4个城区,4个丘陵山地),录制了80个雄性白头鹎的511个鸣声样本,并随机选 取每一调查点的20个鸣声样本进行分析。其结果显示,白头鹎在杭州市区至少有8种微地理鸣声方言,每个微 地理鸣声方言都有一典型鸣句;它们在听觉上、波形结构、音节组成、音节频谱特征等方面均不相同。有的一 路之隔的相邻微地理鸣声方言之间存在明显差异;部分区域出现鸣声混合现象;有的个体还具有“多语”功 能。白头鹎鸣声产生微地理差异的原因可能与鸟类个体的扩散和城市中鸟类栖息地的人为改变有关。