Female, but not male, tropical sparrows respond more strongly to the local song dialect: implications for population divergence

In addition to the observed high diversity of species in the tropics, divergence among populations of the same species exists over short geographic distances in both phenotypic traits and neutral genetic markers. Divergence among populations suggests great potential for the evolution of reproductive isolation and eventual speciation. In birds, song can evolve quickly through cultural transmission and result in regional dialects, which can be a critical component of reproductive isolation through variation in female preference. We examined female and male behavioral responses to local and nonlocal dialects in two allopatric populations of rufous-collared sparrows (Zonotrichia capensis) in the Andes Mountains of Ecuador. Here we show that female sparrows prefer their natal song dialect to the dialect of an allopatric population that is just 25 km away and separated by an unsuitable higher-elevation habitat (pass of 4,200 m), thus providing evidence of prezygotic reproductive isolation among populations. Males showed similar territorial responses to all conspecific dialects with no consistent difference with respect to distance, making male territoriality uninformative for estimating reproductive isolation. This study provides novel evidence for culturally based prezygotic isolation over very short distances in a tropical bird.     

Social dialect and men's voice pitch influence women's mate preferences

Low male voice pitch may communicate potential benefits for offspring in the form of heritable health and/or dominance, whereas access to resources may be indicated by correlates of socioeconomic status, such as sociolinguistic features. Here, we examine if voice pitch and social dialect influence women's perceptions of men's socioeconomic status and attractiveness. In Study 1, women perceived lower pitched male voices as higher in socioeconomic status than higher pitched male voices. In Study 2, women independently perceived lower pitched voices and higher status sociolinguistic dialects as higher in socioeconomic status and attractiveness. We also found a significant interaction wherein women preferred lower pitched men's voices more often when dialects were lower in sociolinguistic status than when they were higher in sociolinguistic status. Women also perceived lower pitched voices as higher in socioeconomic status more often when dialects were higher in sociolinguistic status than when lower in sociolinguistic status. Finally, women's own self-rated socioeconomic status was positively related to their preferences for voices with higher status sociolinguistic dialects, but not to their preferences for voice pitch. Hence, women's preferences for traits associated with potentially biologically heritable benefits, such as low voice pitch, are moderated by the presence of traits associated with resource accrual, such as social dialect markers. However, women's preferences for language markers of resource accrual may be functionally independent from preferences for potential biological indicators of heritable benefits, such as voice pitch.     

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.     

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.     

Urban noise and the cultural evolution of bird songs

In urban environments, anthropogenic noise can interfere with animal communication. Here we study the influence of urban noise on the cultural evolution of bird songs. We studied three adjacent dialects of white-crowned sparrow songs over a 30-year time span. Urban noise, which is louder at low frequencies, increased during our study period and therefore should have created a selection pressure for songs with higher frequencies. We found that the minimum frequency of songs increased both within and between dialects during the 30-year time span. For example, the dialect with the highest minimum frequency is in the process of replacing another dialect that has lower frequency songs. Songs with the highest minimum frequency were favoured in this environment and should have the most effective transmission properties. We suggest that one mechanism that influences how dialects, and cultural traits in general, are selected and transmitted from one generation to the next is the dialect''s ability to be effectively communicated in the local environment.     

Pair Duets in the Yellow-Naped Amazon (Psittaciformes: Amazona auropalliata): Responses to Playbacks of Different Dialects

Yellow-naped amazons, Amazona auropalliata , have regional dialects in which several functional classes of vocalization, including contact calls and pair duets, change their acoustic structure at the same geographic boundaries. Here we examine the responses of 11 pairs of yellow-naped amazons to playbacks of duets from other pairs nesting near the same roost, other roosts within the same dialect, and roosts in foreign dialect areas. Overall, pairs responded more strongly to duets from their own dialect than to those of the foreign dialect. Pairs responded to both treatments from their own dialect (local same dialect and distant same dialect) with movement towards the broadcasting loudspeaker and more rarely with squeals, a vocalization typically observed only in the context of aggressive chases. These aggressive responses were never observed during playbacks of the foreign dialect treatment or congeneric controls. There were no differences among treatments in the incidence of contact calls or pair duets. A similar pattern of stronger aggressive responses to local than to foreign dialects has been found in a wide range of oscine songbirds. The results of the present experiment suggest that a general function may underlie this behavioral response both in oscines and in other bird taxa with vocal learning.     

Dialect Discrimination by Male Orange-Tufted Sunbirds (Nectarinia osea): Reactions to Own vs. Neighbor Dialects

Male orange-tufted sunbirds ( Nectarinia osea ) exhibit distinct song dialects throughout Israel. Recently, two distinct local dialects with a sharp boundary were discovered in a small (1.5 km²) urban neighborhood densely inhabited by 63 territorial sunbird pairs. We conducted playback experiments to determine song dialect discrimination capability by sunbird males in this neighborhood. Males of both dialects responded significantly more strongly to playback of their own dialect than to that of the adjacent dialect. In spite of the extreme proximity between the two dialect areas, we found no effect of distance to the neighboring dialect on the intensity of any the behavioral responses. We suggest that due to the complex acoustic properties of this urban neighborhood, sunbirds are extremely limited in the number of neighboring males they can assess to establish what the local song is. A stronger response to one's own dialect is therefore expected, and we discuss how local dialects could be maintained via this mechanism regardless of the very small distances between territories and dialect populations.     

Quantifying cultural macro-evolution: a case study of the hinoeuma fertility drop

Understanding patterns and underlying processes of human cultural diversity has been a major challenge in evolutionary anthropology. Recent developments in the study of cultural macro-evolution have illuminated various novel aspects of cultural phenomena at the population level. However, limitations in data availability have constrained previous analyses to use simplest models ignoring factors that potentially affect cultural evolutionary dynamics. Here, we focus on two such factors: accumulated effects of cultural transmission between populations over time and variation in social influence among populations. As a test case, we analyze data on the hinoeuma fertility drop, the Japanese nation-wide drastic decline in the number of births caused by a culturally-transmitted superstition recurring every sixty years, to show that these factors do play significant roles. Specifically, our results suggest that transmission of the superstition in a short timescale has tended to occur among neighboring populations, while transmission in a long timescale is likely to have occurred between populations culturally close to each other, with the cultural closeness being measured by similarity in dialects. The results also indicate a special role played by a population occupying a center in a language–distance network (the cultural center) in the spread of the superstition.     

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.     

Birds of a feather speak together: Understanding the different dialects of animals can help to decipher their communication

As humans, many animal species that communicate via vocalization show a wide range of accents and dialects driven by environmental and social factors.

All human languages come with dialects not just as a result of geographical distance between different tribes or groups but also as a part of their local culture and heritage. “Dialect or the speech of the people is capable of expressing whatever the people are,” said Sterling K Brown, an American actor. A Southerner in the USA, a Sicilian in Italy or a Bavarian in Germany would be easily recognizable as such, but also often stresses his or her dialect to signal where they come from geographically and culturally.

… many animal species from multiple taxa have evolved forms of vocal communication that are subject to geographic, genetic, environmental, behavioural and social variations.

Dialect and accents are not unique to humans: Many animal species from multiple taxa have evolved forms of vocal communication that are subject to geographic, genetic, environmental, behavioral, and social variations. One more or less universal factor behind the variety of animal dialects is that some convergence or conservation of vocalization can confer selective or competitive advantages.One area of confusion lies in the use of terms such as dialect and accent, which have more clearly defined meanings in the context of human languages. Here, accents refer to the way the same words are spoken, varying in acoustic frequency or intonation but not in enunciation or form. Accents are thus a subset of dialects, which also bring variations in vocabulary, grammatical structure and idiom within a common underlying language.     

Performance of a Receptive Language Test among Young Children in Madagascar

Language tests developed and validated in one country may lose their desired properties when translated for use in another, possibly resulting in misleading estimates of ability. Using Item Response Theory (IRT) methodology, we assess the performance of a test of receptive vocabulary, the U.S.-validated Peabody Picture Vocabulary Test-Third Edition (PPVT-III), when translated, adapted, and administered to children 3 to 10 years of age in Madagascar (N = 1372), in the local language (Malagasy). Though Malagasy is considered a single language, there are numerous dialects spoken in Madagascar. Our findings were that test scores were positively correlated with age and indicators of socio-economic status. However, over half (57/96) of items evidenced unexpected response variation and/or bias by local dialect spoken. We also encountered measurement error and reduced differentiation among person abilities when we used the publishers’ recommended stopping rules, largely because we lost the original item ordering by difficulty when we translated test items into Malagasy. Our results suggest that bias and testing inefficiency introduced from the translation of the PPVT can be significantly reduced with the use of methods based on IRT at both the pre-testing and analysis stages. We explore and discuss implications for cross-cultural comparisons of internationally recognized tests, such as the PPVT.     

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.     

Dialect use in large assemblies: a study in European starling Sturnus vulgaris roosts

Dialects may signal social or population identity and increase tolerance within communities. We hypothesized that in European starling Sturnus vulgaris communal roosts, birds coming from the same breeding area, i.e. dialectal zone, might tend to stay together within the roost. Recordings were performed in the colonies, revealed in earlier studies, multiple dialects and small sectors where birds shared the same variants at the different levels. We also performed recordings in different locations within night roosts. The dialects recorded in the roosts were the same as those recorded at nest sites during the day and they were not distributed randomly within roosts: birds from the same geographical diurnal origin would gather and stay together, either because they arrived together or were attracted to their dialect. Although our results have to be confirmed by the study of identifiable individuals, we propose original lines of thought on roost structuring and on the role of song dialects.     

Winter dialects in the bronzed cowbird and their relationship to breeding-season dialects

Studies of geographical variation in animal signals generally focus on breeding-season behaviour but, in many species, signalling persists throughout the year. In passerine birds, patterns of variation in the nonbreeding season might provide opportunities for vocal learning that have been neglected by a historic focus on breeding-season behaviour. This study provides the first example of dialect variation outside of the breeding season. Quantitative analysis of acoustic similarity showed discrete differences between the songs of bronzed cowbirds, Molothrus aeneus, in four winter flocks. Most songs produced by members of a given flock were classified as belonging to the same dialect. Songs from one of the four winter dialects were indistinguishable from songs recorded in the breeding season in the same region. Depending on migratory patterns, dialects in one season may be a consequence of dialects in the other season, or the two seasonal patterns may be the result of independent social or evolutionary forces. Because the nonbreeding season is an important period of vocal learning in some bird species, winter dialects might limit the range of signals available for individuals to learn to produce. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Population genetic structure and vocal dialects in an amazon parrot

The relationship between cultural and genetic evolution was examined in the yellow-naped amazon Amazona auropalliata. This species has previously been shown to have regional dialects defined by large shifts in the acoustic structure of its learned contact call. Mitochondrial DNA sequence variation from a 680 base pair segment of the first domain of the control region was assayed in 41 samples collected from two neighbouring dialects in Costa Rica. The relationship of genetic variation to vocal variation was examined using haplotype analysis, genetic distance analysis, a maximum-likelihood estimator of migration rates and phylogenetic reconstructions. All analyses indicated a high degree of gene flow and, thus, individual dispersal across dialect boundaries. Calls sampled from sound libraries suggested that temporally stable contact call dialects occur throughout the range of the yellow-naped amazon, while the presence of similar dialects in the sister species Amazona ochrocephala suggests that the propensity to form dialects is ancestral in this clade. These results indicate that genes and culture are not closely associated in the yellow-naped amazon. Rather, they suggest that regional diversity in vocalizations is maintained by selective pressures that promote social learning and allow individual repertoires to conform to local call types.     

Genetic population structure in an equatorial sparrow: roles for culture and geography

Female preference for local cultural traits has been proposed as a barrier to breeding among animal populations. As such, several studies have found correlations between male bird song dialects and population genetics over relatively large distances. To investigate whether female choice for local dialects could act as a barrier to breeding between nearby and contiguous populations, we tested whether variation in male song dialects explains genetic structure among eight populations of rufous‐collared sparrows (Zonotrichia capensis) in Ecuador. Our study sites lay along a transect, and adjacent study sites were separated by approximately 25 km, an order of magnitude less than previously examined for this and most other species. This transect crossed an Andean ridge and through the Quijos River Valley, both of which may be barriers to gene flow. Using a variance partitioning approach, we show that song dialect is important in explaining population genetics, independent of the geographic variables: distance, the river valley and the Andean Ridge. This result is consistent with the hypothesis that song acts as a barrier to breeding among populations in close proximity. In addition, songs of contiguous populations differed by the same degree or more than between two populations previously shown to exhibit female preference for local dialect, suggesting that birds from these populations would also breed preferentially with locals. As expected, all geographic variables (distance, the river valley and the Andean Ridge) also predicted population genetic structure. Our results have important implications for the understanding whether, and at what spatial scale, culture can affect population divergence.     

Dialects of an invasive songbird are preserved in its invaded but not native source range

Biological invasions are not only events with substantial environmental and socioeconomic impacts but are also interesting natural experiments, allowing the study of phenomena such as the cultural evolution of bird song following introduction. We took an excellent opportunity to compare the distribution of dialects of the yellowhammer Emberiza citrinella, a small Eurasian passerine, in its native source region (Great Britain) and invaded range (New Zealand) more than hundred years after relocation. Recent field recordings (including those provided by volunteers within a citizen science project) were complemented by those from archives, each assigned to appropriate dialect by visual inspection of a sonogram, and the resulting spatial patterns of dialect distribution were interpreted using historical data on the yellowhammer invasion. The two countries differ markedly in the composition and distribution of dialects. New Zealand populations sing a greater number of different dialects, seven in total, five of which were not detected in the current British population, but have been reported by previous studies from the continental Europe. Two identified localities of capture (Brighton, Sussex, UK) and release (Dunedin, Otago, NZ) differ even more strikingly, having no dialects in common. The largely sedentary nature of yellowhammers allows for two mutually exclusive explanations for European dialects being detected in New Zealand but not in Great Britain: 1) the corresponding song types have emerged de novo in New Zealand, through convergent cultural evolution; 2) the dialects have disappeared from Great Britain, while being preserved in New Zealand. Indirect evidence from the widespread occurrence of these dialects in continental Europe and the reported stability of yellowhammer song, supports the latter explanation. We suggest that the yellowhammer dialect system is an avian equivalent of a phenomenon already noted in human languages, in which ancient words or structures are retained in expatriate communities.     

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.     

Do surname differences mirror dialect variation?

Our focus in this paper is the analysis of surnames, which have been proven to be reliable genetic markers because in patrilineal systems they are transmitted along generations virtually unchanged, similarly to a genetic locus on the Y chromosome. We compare the distribution of surnames to the distribution of dialect pronunciations, which are clearly culturally transmitted. Because surnames, at the time of their introduction, were words subject to the same linguistic processes that otherwise result in dialect differences, one might expect their geographic distribution to be correlated with dialect pronunciation differences. In this paper we concentrate on the Netherlands, an area of only 40,000 km2, where two official languages are spoken, Dutch and Frisian. We analyze 19,910 different surnames, sampled in 226 locations, and 125 different words, whose pronunciation was recorded in 252 sites. We find that, once the collinear effects of geography on both surname and cultural transmission are taken into account, there is no statistically significant association between the two, suggesting that surnames cannot be taken as a proxy for dialect variation, even though they can be safely used as a proxy for Y-chromosome genetic variation. We find the results historically and geographically insightful, hopefully leading to a deeper understanding of the role that local migrations and cultural diffusion play in surname and dialect diversity.