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1.
Genetic diversity and population structure were studied in eight populations of the kestrel Falco tinnunculus to identify the genetic consequences of spatial distribution and to infer the colonization patterns of the Cape Verde archipelago. We studied genetic differentiation and gene flow among seven island populations and one mainland population using nine microsatellite loci. Within the archipelago, differentiation was strong and genetic diversity and heterozygosity were low but variable among populations. Two subspecies F. tinnunculus neglectus on the northwestern islands and F. tinnunculus alexandri on all the other islands were identified as genetically distinct units. F. t. alexandri could be further separated into two groups on eastern and southern islands. Populations are probably founded by birds originating from the mainland. Immigration is more likely to the eastern and southern populations, whereas the northwestern islands with the lowest genetic diversity and highest differentiation are likely to exhibit fewer founding events by immigrants. The number of founding events on each island may depend not only on geographical distance to neighbouring populations, but also on directional immigration due to the northeastern trade winds. This may explain differences in genetic differentiation and diversity between populations and subspecies and may enable allopatric speciation.  相似文献   

2.
Unlike other migratory hummingbirds in North America, the broad‐tailed hummingbird (Selasphorus platycercus) exhibits both long‐distance migratory behaviour in the USA and sedentary behaviour in Mexico and Guatemala. We examined the evolution of migration linked to its northward expansion using a multiperspective approach. We analysed variation in morphology, mitochondrial and nuclear DNA, estimated migration rates between migratory and sedentary populations, compared divergence times with the occurrence of Quaternary climate events and constructed species distribution models to predict where migratory and sedentary populations resided during the Last Glacial Maximum (LGM) and Last Interglacial (LIG) events. Our results are consistent with a recent northward population expansion driven by migration from southern sedentary populations. Phylogeographical analyses and population genetics methods revealed that migratory populations in the USA and sedentary populations in Mexico of the platycercus subspecies form one admixed population, and that sedentary populations from southern Mexico and Guatemala (guatemalae) undertook independent evolutionary trajectories. Species distribution modelling revealed that the species is a niche tracker and that the climate conditions associated with modern obligate migrants in the USA were not present during the LIG, which provides indirect evidence for recent migratory behaviour in broad‐tailed hummingbirds on the temporal scale of glacial cycles. The finding that platycercus hummingbirds form one genetic population and that suitable habitat for migratory populations was observed in eastern Mexico during the LIG also suggests that the conservation of overwintering sites is crucial for obligate migratory populations currently facing climate change effects.  相似文献   

3.
We investigated the genetic population structure and species status of a relatively sedentary bird that is a permanent resident of western North American forests, the blue grouse (Dendragapus obscurus). Phylogenetic analysis of complete mitochondrial control region DNA sequences resulted in the identification of three basal clades of haplotypes that were largely congruent with well-known biogeographical regions. These clades corresponded to the parapatric sooty (D. o. fuliginosus) and dusky (D. o. obscurus) subspecies groups of blue grouse plus a previously unrecognized division between northern and southern dusky grouse populations; the latter does not correspond closely to any currently recognized subspecies boundary. Approximately 66% of the total genetic variation was distributed among these three regions. Maximum likelihood estimates of gene flow between the regions were low or asymmetric; gene flow has been insufficient to prevent genetic divergence between dusky and sooty grouse. Estimates of gene flow among populations within sooty grouse were large except across the Columbia River valley. Among populations of dusky grouse, estimates of gene flow were heterogeneous and asymmetrical, reflecting large-scale fragmentation of the distribution due to landscape features and associated vegetation. Genetic, morphological and behavioural evidence suggest that sooty and dusky grouse are species-level taxa; the specific status of a third clade remains ambiguous.  相似文献   

4.
We investigated the evolutionary history of the spotted flycatcher Muscicapa striata, a long distance migratory passerine having a widespread range, using mitochondrial markers and nuclear introns. Our mitochondrial results reveal the existence of one insular lineage restricted to the western Mediterranean islands (Balearics, Corsica, Sardinia) and possibly to the Tyrrhenian coast of Italy that diverged from the mainland lineages around 1 Mya. Mitochondrial genetic distance between insular and mainland lineages is around 3.5%. Limited levels of shared nuclear alleles among insular and mainland populations further support the genetic distinctiveness of insular spotted flycatchers with respect to their mainland counterparts. Moreover, lack of mitochondrial haplotypes sharing between Balearic birds (M. s. balearica) and Corso‐Sardinian birds (M. s. tyrrhenica) suggest the absence of recent matrilineal gene flow between these two insular subspecies. Accordingly, we suggest that insular spotted flycatchers could be treated as one polytypic species (Muscicapa tyrrhenica) that differs from M. striata in morphology, migration, mitochondrial and nuclear DNA and comprises two subspecies (the nominate and M. t. balearica) that diverged recently phenotypically and in mitochondrial DNA and but still share the same nuclear alleles. This study provides an interesting case‐study illustrating the crucial role of western Mediterranean islands in the evolution of a passerine showing high dispersal capabilities. Our genetic results highlight the role of glacial refugia of these islands that allowed initial allopatric divergence of insular populations. We hypothesize that differences in migratory and breeding phenology may prevent any current gene flow between insular and mainland populations of the spotted flycatcher that temporarily share the same insular habitats during the spring migration.  相似文献   

5.
The Chiffchaff Phylloscopus collybita is represented in Sweden by two different subspecies; the northern well-established abietinus and the southern recently established collybita which has expanded its range northward during the past two decades. At present, an area approximately 500 km wide separates the two subspecies. In order to document differences between the northern and southern populations we compared morphology, vocalisation, habitat choice, and neutral genetic variation in mitochondrial (mt) DNA and at four microsatellite loci of 30 male Chiffchaffs from each subspecies. Our results show significant differences in several morphological traits and in song. Playback experiments revealed a significant difference in aggressive behaviour depending on which population-specific song that was played to the birds. Mitochondrial DNA was geographically structured with ∼90% of the birds carrying a mt haplotype matching their sample population. No allelic differences at the microsatellite loci were found between populations. Our data demonstrate a substantial differentiation between the northern and southern populations despite gene flow, clearly separating them into the subspecies abietinus and collybita .  相似文献   

6.
We used the morphological differentiation of southern, sedentary Iberian Blackcaps Sylvia atricapilla to distinguish them from migratory Blackcaps during the winter. To do so, we studied three morphometric traits (wing length, wing shape and tail length) of three migratory Iberian populations and two sedentary ones during the breeding season. We obtained a discriminant function which allowed us to differentiate the two population types (>90% correct classifications). We discuss the utility of the method for differentiating between sedentary and migratory Blackcaps in their sympatric wintering areas, as well as the implications of our results for the study and conservation of sedentary, perhaps differentiated Blackcaps in southern Iberia.  相似文献   

7.
Ma C  Yang P  Jiang F  Chapuis MP  Shali Y  Sword GA  Kang L 《Molecular ecology》2012,21(17):4344-4358
The migratory locust, Locusta migratoria, is the most widely distributed grasshopper species in the world. However, its global genetic structure and phylogeographic relationships have not been investigated. In this study, we explored the worldwide genetic structure and phylogeography of the locust populations based on the sequence information of 65 complete mitochondrial genomes and three mitochondrial genes of 263 individuals from 53 sampling sites. Although this locust can migrate over long distances, our results revealed high genetic differentiation among the geographic populations. The populations can be divided into two different lineages: the Northern lineage, which includes individuals from the temperate regions of the Eurasian continent, and the Southern lineage, which includes individuals from Africa, southern Europe, the Arabian region, India, southern China, South‐east Asia and Australia. An analysis of population genetic diversity indicated that the locust species originated from Africa. Ancestral populations likely separated into Northern and Southern lineages 895 000 years ago by vicariance events associated with Pleistocene glaciations. These two lineages evolved in allopatry and occupied their current distributions in the world via distinct southern and northern dispersal routes. Genetic differences, caused by the long‐term independent diversification of the two lineages, along with other factors, such as geographic barriers and temperature limitations, may play important roles in maintaining the present phylogeographic patterns. Our phylogeographic evidence challenged the long‐held view of multiple subspecies in the locust species and tentatively divided it into two subspecies, L. m. migratoria and L. m. migratorioides.  相似文献   

8.
Allozyme variation was investigated in 17 Japanese populations of Campanula punctata, ten from the Izu Islands and seven in the mainland (Honshu). The data indicate that there are two groups, a mainland group and an island one, and that the systematically problematic Oshima Island (northernmost Izu island) populations are closely related to those of the other islands. Nei's genetic identity values among islands and among mainland populations were 0.95 and 0.97, respectively, while the value between island and mainland populations was 0.84, suggesting that the island populations are an independent species. Total genetic variation was nearly the same among island and mainland populations. However, the apportionment of variation within and among populations was considerably different; 14% of gene diversity exists among mainland populations, while 31% of the diversity exists among island populations. Mean outcrossing rates of self-incompatible mainland and Oshima populations are 0.62–0.79, supporting xenogamy; those in self-compatible island populations are 0.37–0.57 in the northern islands, indicating a mixed mating system, and 0.16–0.25 in southern ones, indicating dominant inbreeding. Total genetic diversity in each island population decreased with distance from the mainland. Genetic and geological data suggest that the ancestors of insular populations were founded on northern islands in a relatively ancient period and that they dispersed progressively to the southern ones. Chromosome number (2n = 34) and isozyme numbers indicate gene duplications in this species, which suggests it is an ancient polyploid.  相似文献   

9.
Morphological and genetic variation is evaluated among populations of the bat, Eidolon helvum , in the islands of the Gulf of Guinea (Central Africa). The populations from the islands of Bioko, Principe, and Sao Tome do not show significant phenetic differentiation, although a trend towards a reduction of size is found in the latter two islands. The low genetic distances between populations, as well as their values of Wright's fixation indexes, suggest that gene flow has hampered differentiation on these islands. In contrast, the population from Annobon, the smallest and farthermost island, shows remarkable morphological and genetic differentiation. On the mainland, E. helvum displays unique migratory and dispersal behaviours, but migratory behaviour was not found in any of the island populations. The combination of selective forces in harsher oceanic environments and restricted gene flow among populations appears to have favoured the high degree of morphological differentiation of E. helvum on Annobon. Due to the extended length of the dry season in Annobon, an earlier achievement of sexual maturity–and consequently smaller size—may be advantageous in the absence of migration. The differentiation is more marked among females, which also suggests that selection may be linked to the reproductive pattern. The population of the island of Annobon is herein described as a new subspecies, Eidolon helvum annobonensis subsp. nov.  相似文献   

10.
Genetic variation was studied in the southern subspecies of the Asian Dolly Varden Salvelinus malma krascheninnikovi from the Kuril Islands. Thirty-six genetic loci controlling 19 enzyme systems were analyzed in 13 Dolly Varden populations from the Shumshu, Paramushir, Onekotan, Rasshua, Simushir, Urup, Iturup, and Kunashir islands. In the studied populations, the proportion of polymorphic loci was 35 to 85% and the mean heterozygosity was 0.104 to 0.173; populations from the Kunashir Island were characterized by maximum heterozygosity. In the island populations examined, significant inter-population heterogeneity of allele frequencies was found for all studied population pairs. For the total population of all islands, the inter-population diversity (GST = 0.188) was comparable to this parameter for the total population from the Kunashir Island (GST = 0.170). Genetic distances between populations did not correlate with the corresponding geographical distances, which indicates the lack of a pronounced gene exchange between the island populations. Cluster analysis and multidimensional scaling based on genetic distances did not reveal clear groups among the studied populations but indicated greater similarity within the Iturup-Simushir-Urup-Paramushir group and a greater genetic divergence of the Kunashir, Onekotan, Rasshua, and especially Shumshu populations. In the Shumshu population, allele frequencies indicate the admixture of genes of the northern Dolly Varden. The observed pattern of genetic differentiation was probably caused largely by genetic drift under the conditions of a limited gene flow because of homing (which is typical of the Dolly Varden) and the presence of isolated nonanadromous populations. The population-genetic analysis of the Dolly Varden from the Kuril Islands does not give grounds to distinguish any other isolated Dolly Varden species in this region than S. malma, which is represented by the southern form S. m. krascheninnikovi with an admixture of the northern form S. m. malma in the Shumshu Island.  相似文献   

11.
The wing shape and size morphology of populations of the medically important phlebotomine sand fly, Phlebotomus papatasi, were examined in two endemic (south of the Atlas Mountains) and nonendemic (north of the Atlas Mountains) foci of cutaneous leishmaniasis by using geometric morphometrics in Morocco. Although it is present in all of Morocco, P. papatasi is the main vector of Leishmania major in only southern part of the Atlas Mountains. There are four major mountain ranges that serve as geographical barriers for species distribution in the study area and at least four gaps were recognized among these barriers. We found statistically significant differences in wing shape morphology between southern and northern populations. Analysis clearly recognized two main groups of populations on both sides of the mountains. The graphical depiction of Principal Component Analysis (PCA) and Canonical Variates Analysis (CVA) confirmed our morphometric study suggesting that the difference in wing morphology between the populations indicates that the population of P. papatasi shows phenotypic plasticity in the study area. According to centroid size analyses, which were used as measures of wing size differences among different sites, the north population of P. papatasi had relatively larger wings than the south population.  相似文献   

12.
Hierarchical genetic structure was examined in the three geographically-defined subspecies of spotted owl (Strix occidentalis) to define relationships among subspecies and quantify variation within and among regional and local populations. Sequences (522 bp) from domains I and II of the mitochondrial control region were analyzed for 213 individuals from 30 local breeding areas. Results confirmed significant differences between northern spotted owls and the other traditional geographically defined subspecies but did not provide support for subspecific level differences between California and Mexican spotted owls. Divergence times among subspecies estimated with a 936 bp portion of the cytochrome b gene dated Northern and California/Mexican spotted owl divergence time to 115,000–125,000 years ago, whereas California/Mexican spotted owl divergence was estimated at 15,000 years ago. Nested clade analyses indicated an association between California spotted owl and Mexican spotted owl haplotypes, implying historical contact between the two groups. Results also identified a number of individuals geographically classified as northern spotted owls (S. o. caurina) that contained haplotypes identified as California spotted owls (S. o. caurina). Among all northern spotted owls sampled (n=131), 12.9% contained California spotted owl haplotypes. In the Klamath region, which is the contact zone between the two subspecies, 20.3% (n=59) of owls were classified as California spotted owls. The Klamath region is a zone of hybridization and speciation for many other taxa as well. Analyses of population structure indicated gene flow among regions within geographically defined subspecies although there was significant differentiation among northern and southern regions of Mexican spotted owls. Among all areas examined, genetic diversity was not significantly reduced except in California spotted owls where the southern region consists of one haplotype. Our results indicate a stable contact zone between northern and California spotted owls, maintaining distinct subspecific haplotypes within their traditional ranges. This supports recovery efforts based on the traditional subspecies designation for the northern spotted owl. Further, although little variation was found between California and Mexican spotted owls, we suggest they should be managed separately because of current isolation between groups.  相似文献   

13.
Migratory strategies of birds require complex orientation mechanisms, morphological adaptations, and life-history adjustments. From an evolutionary perspective, it is important to know how fast this complex combination of traits can evolve. We analyzed mitochondrial control-region DNA sequences in 241 blackcaps (Sylvia atricapilla) from 12 populations with different migratory behaviors. The sample included sedentary populations in Europe and Atlantic archipelagos and migratory populations with different distances of migration, from regional to intercontinental migrations, and different heading directions (due to a migratory divide in central Europe). There was no genetic structure between migratory and sedentary populations, or among populations from different biogeographic areas (Atlantic islands, the Iberian Peninsula, or the continent), however we found evidence of a genetic structure when comparing populations located on either side of the migratory divide. These findings support an independent evolution of highly divergent migratory strategies in blackcaps, occurring after a postglacial colonization of the continent along western and eastern routes. Accordingly, mismatch-distribution analyses suggested an expansion of blackcaps from a very small population size, and time estimates dated such an expansion during the last postglacial period. However, the populations in Gibraltar, located in a putative Mediterranean refuge, appeared to be independent of these processes, showing evidence of restricted gene flow with other populations and demonstrating insignificant historical changes in effective population size. Our results show that the interruption of gene flow between migratory and sedentary populations is not necessary for the maintenance of such a polymorphism, and that even the most divergent migratory strategies of a bird species are susceptible to evolution in response to historical environmental changes.  相似文献   

14.
The Southern Ocean contains some of the most isolated islands on Earth, and fundamental questions remain regarding their colonization and the connectivity of their coastal biotas. Here, we conduct a genetic investigation into the Cellana strigilis (limpet) complex that was originally classified based on morphological characters into six subspecies, five of which are endemic to the New Zealand (NZ) subantarctic and Chatham islands (44–52°S). Previous genetic analyses of C. strigilis from six of the seven island groups revealed two lineages with little or no within‐lineage variation. We analysed C. strigilis samples from all seven island groups using two mitochondrial (COI and 16S), one nuclear (ATPase β) and 58 loci from four randomly amplified polymorphic DNA markers (RAPDs) and confirmed the existence of two distinct lineages. The pronounced genetic structuring within each lineage and the presence of private haplotypes in individual islands are the result of little genetic connectivity and therefore very high self‐recruitment. This study supports the significance of the subantarctic islands as refugia during the last glacial maximum and adds to the knowledge of contemporary population connectivity among coastal populations of remote islands in large oceans and the distance barrier to gene flow that exists in the sea (despite its continuous medium) for most taxa.  相似文献   

15.
To investigate genetic diversity among populations of the sika deer, Cervus nippon, nucleotide sequences (705-824 bases) of the mitochondrial D-loop regions were determined in animals from 13 localities in the Japanese islands. Phylogenetic trees constructed by the sequences indicated that the Japanese sika deer is separated into two distinct lineages: the northern Japan group (the Hokkaido island and most of the Honshu mainland) and the southern Japan group (a part of the southern Honshu mainland, the Kyushu island, and small islands around the Kyushu island). All sika deer examined in this study shared four to seven units of repetitive sequences (37 to 40 bases each) within the D-loop sequences. The number of tandem repeats was different among the populations, and it was specific to each population. Six or seven repeats occurred in populations of the northern Japan group, while four or five repeats occurred in populations of the southern Japan group. Each repeat unit included several nucleotide substitutions, compared with others, and 26 types were identified from 31 animals. Sequences of the first, second, and third units in arrays were clearly different between the northern and the southern groups. Based on these D-loop data, colonization and separation of the sika deer populations in the Japanese islands were estimated to have occurred less than 0.5 million years before present. Our results provide an invaluable insight into better understanding the evolutionary history, phylogeny, taxonomy, and population genetics of the sika deer.  相似文献   

16.
The migratory locust, Locusta migratoria L., which is distributed widely in the East Hemisphere, has long been regarded as the most important agricultural pest and a model insect. Its distribution ranges from thetropic and the subtropical zone, to the temperate zone, and to the cold-temperate zone. Because of the wide geographical distribution and adaptations, many geo- graphical populations of the migratory locust dis- played apparent variation in morphology, life history, physiology, and ot…  相似文献   

17.
The random amplified polymorphic DNA (RAPD) technique was used to examine genetic divergence and interrelations of 11 geographical populations of the migratory locust in China, and the role of spatial separation in the population differentiations. AMOVA analysis of genetic variations in all the populations indicated greater within- (79.55%) than among-population variability (20.45%), and that there were significant differentiations among the populations; 11 populations were divided into four regional groups, with significantly greater variability within (82.99%) than among the groups (17.01%), and there existed apparent regional differentiations. Paired comparisons showed significantly greater variability within-than between-groups, indicating significant differentiations between populations of different regional groups. Of all the pairwise comparisons, Hainan and Tibetan groups displayed the greatest differentiation, with the difference between the two groups being seven folds of that between populations within the groups; the least differentiations were exhibited between the groups of Hainan, Xinjiang, and Inner Mongolia, with the differences between groups being only half of the differences between populations within the groups. Mantel tests of the genetic and spatial distances showed that the two matrices were significantly correlated (p<0.01), indicating that the geographical isolation played an important role in the differentiations of the geographical populations of the migratory locusts. Cluster analysis divided all populations into four major groups: Xinjiang and Inner Mongolia group, the Great Plains of North China (the Yellow River and Huai River Plains) group, Hainan group, and Tibet group. Principal component analysis (PCA) supported the division of populations based on the cluster analysis. However, analysis of individuals clustered the locusts into five populations: Xinjiang and Inner Mongolia, Hami in Xinjiang, the Great Plains of North China, Hainan, and Tibet. The locust populations in eastern China displayed apparently continous and gradient variations; as such authors consider that there were no necessity and valid reasons for further division of subspecies. The subspecific status for the main geographical populations of the migratory locusts in China was discussed.  相似文献   

18.
Hairy woodpeckers Picoides villosus are a common, year round resident with distinct plumage and morphological variation across North America. We genotyped 335 individuals at six variable microsatellite loci and analyzed 322 mtDNA control region sequences in order to examine the role of contemporary and historical barriers to gene flow. In addition we combined genetic analyses with ecological niche modelling to test if hairy woodpeckers were isolated in northern refugia (Alaska, Newfoundland and the Queen Charlotte Islands) during the last glacial maximum. Genetic analyses revealed that gene flow among North American hairy woodpecker populations is restricted, but not to the extent predicted for a sedentary species. Populations clustered into two main genetic groups, east and west of the Great Plains in the south and the Rocky Mountains in the north. Contact zones between the two main genetic groups exist in central British Columbia and Washington, but are narrow. Within each group we found additional population structure with genetic breaks between subgroups in the geographic west corresponding to breaks in forested habitat and physical barriers like open expanses of water. Population genetic patterns for hairy woodpeckers have resulted from isolation in multiple southern refugia with the current distribution of genetic groups resulting from post‐glacial expansion and subsequent reduction in gene flow. While populations in Alaska, Newfoundland and the Queen Charlotte Islands are genetically distinct from other populations, we found no evidence of these areas acting as refugia throughout the Pleistocene. Atlantic Canada populations contained unique haplotypes raising the possibility of a separate colonization from the rest of eastern Canada. The endemic subspecies on the island of Newfoundland is not genetically distinct from their closest mainland population unlike the Queen Charlotte Island subspecies.  相似文献   

19.
A migratory divide is a narrow region in which two populations showing different migratory directions meet arid presumably also mate and hybridize. Banding of willow warblers, Phylloscopus trochilus, in Europe has demonstrated a migratory divide latitudinally across central Scandinavia. In autumn, southern birds migrate southwest to tropical West Africa, whereas northern birds migrate southeast to East and South Africa. The migratory divide is associated with concordant differences in size and plumage coloration. Based on morphology, we estimate the width of the transition zone between northern and southern willow warblers to be less than 350 km. We found indication of linkage disequilibria around the migratory divide, in that measures of body size were correlated with plumage coloration within the contact zone, but uncorrelated within the populations south or north of the contact zone. The presence of linkage disequilibria and the fact that several morphological clines occur together suggest that the hybrid zone is a result of secondary contact between populations that have differentiated in allopatry. This interpretation is in accord with the knowledge of the recolonization pattern of the Scandinavian peninsula after the last glaciation; animals and plants appeared to have colonized either from the south or from the north around the northern bay of the Baltic Sea. If northern and southern willow warblers resided in allopatric populations during late Pleistocene glaciations and the hybrid zone is a result of postglacial range expansions, we would expect some degree of genetic differentiation accumulated during the period in isolation. In contrast, northern and southern willow warblers are near panmictic in the frequencies of alleles of mitochondrial DNA and at two microsatellite loci. The observed pattern, clear morphological and behavioral differentiation without genetic differentiation at neutral loci, suggests either that the differences are maintained by strong selection on the expressed genes in combination with high levels of current gene flow or, in the case of weak gene flow, that the divergence in morphology and behavior is very recent.  相似文献   

20.
In songbirds, territorial songs are key regulators of sexual selection and are learned from conspecifics. The cultural transmission of songs leads to divergence in song characteristics within populations, which can ultimately lead to speciation. Many songbirds also migrate, and individual differences in migratory behaviours can influence population genetic structure and local song differentiation. Blackcaps, Sylvia atricapilla, exhibit versatile territorial songs and show diversity in migration behaviours. They therefore comprise a good model for investigating the relationships between migratory patterns, song variation, and genetic diversity. We studied a migratory population (two groups near Paris) and a sedentary population (three groups in Corsica). All of the birds were ringed and blood sampled to investigate genetic relatedness using 17 microsatellite loci. A detailed song analysis showed that this species has a complex repertoire (> 100 syllables), which required the development of a semi‐supervised method to classify different categories of syllables and compare sequences of syllables. Our analysis showed no genetic structuring among populations: individuals belonging to the same group were not genetically closer than those from different groups. However, we found a strong wingsize difference between sedentary and migratory populations. We also showed that geographical variations in songs rely at least on both syllable and sequence content. Unexpectedly, despite a higher turnover of individuals, migratory groups share as many syllables and sequences as sedentary groups, which raises interesting issues on song learning and the maintenance of dialects in migratory birds.  相似文献   

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