Mate choice by female white-crowned sparrows in a mixed-dialect population

Summary It has been argued that the song dialects of white-crowned sparrows, Zonotrichia leucophrys, may function to restrict gene flow between dialect populations, thus promoting adaptation to local environmental conditions. One proposed mechanism whereby this may occur is if a female mates preferentially with males singing her natal dialect, typically the song type of her father. Previous field and laboratory studies of the role played by song differences in mate choice by female white-crowned sparrows have produced equivocal results. These investigations were conducted in populations in which a single song dialect was predominant. We examined mate choice in a population in which two song dialects are equally common. We found that songs of mates of individual females in successive years were no more likely to be of the same dialect than expected by chance. In addition, individual females did not mate preferentially with males whose songs matched those of their fathers. We conclude that, in this population, females choose mates on the basis of characteristics other than song type.     

Song dialects do not restrict gene flow in an urban population of the orange-tufted sunbird, <Emphasis Type="Italic">Nectarinia osea</Emphasis>

Geographic variation in vocalizations is widespread in passerine birds, but its origins and maintenance remain unclear. In this study, we test the hypothesis that song dialect, a culturally transmitted trait, is related to the population genetic structure of the orange-tufted sunbird, Nectarinia osea. To address this, we compared mitochondrial DNA (mtDNA) sequence variation together with allele frequencies at five microsatellite loci from an urban population of sunbirds exhibiting two distinct song dialects on a microgeographic scale. Our findings reveal no association between dialect membership and genetic composition. All genetic measures, from both mitochondrial and nuclear DNA, indicate high levels of gene flow between both dialect populations. The low F _ST values obtained from mtDNA and microsatellite analysis imply that the variation among dialects does not account for more than 2%, at best, of the overall genetic variation found in the entire population. These measures fall well within the range of similar measures obtained in other studies of species exhibiting vocal dialects, most of which fail to detect any dialect-based genetic differentiation. The persistence of dialects in the orange-tufted sunbird may thus best be explained by dispersal of individuals across dialect boundaries and possibly from surrounding areas, followed by postdispersal vocal matching. Because genetic structuring appears weaker than cultural structure in this species, we discuss the behavioral mechanisms underlying dialect maintenance in the presence of apparent gene flow.     

Song overproduction and selective attrition lead to song sharing in the field sparrow (Spizella pusilla)

Summary Male field sparrows settling on particular territories for the first time often sing two or more song types. Vocal experience with neighboring males then leads in most cases to the deletion from the male's repertoires of all but one song type. In 2 years, settling males retained the song type that most closely resembled that of a neighbor. Focal observations in 1 year revealed that the most actively singing neighbor was the one the new birds resembled. Songs were not selected on the basis of their absolute acoustic characteristics, but rather by their resemblance to neighbors' songs. After the attrition process is complete, males retain their song for the rest of their lives. Evidence supports the hypothesis that the actual acquisition of a song need not take place at the time and place as males occupy territories. Rather, males may acquire an abundance of song models earlier in life and then select songs from this repertoire that most closely resemble those of their neighbors. The development of mature repertoires and of local dialects is the outcome of one process in this species.     

Population structure of humpback whales in the western and central South Pacific Ocean as determined by vocal exchange among populations

For cetaceans, population structure is traditionally determined by molecular genetics or photographically identified individuals. Acoustic data, however, has provided information on movement and population structure with less effort and cost than traditional methods in an array of taxa. Male humpback whales (Megaptera novaeangliae) produce a continually evolving vocal sexual display, or song, that is similar among all males in a population. The rapid cultural transmission (the transfer of information or behavior between conspecifics through social learning) of different versions of this display between distinct but interconnected populations in the western and central South Pacific region presents a unique way to investigate population structure based on the movement dynamics of a song (acoustic) display. Using 11 years of data, we investigated an acoustically based population structure for the region by comparing stereotyped song sequences among populations and years. We used the Levenshtein distance technique to group previously defined populations into (vocally based) clusters based on the overall similarity of their song display in space and time. We identified the following distinct vocal clusters: western cluster, 1 population off eastern Australia; central cluster, populations around New Caledonia, Tonga, and American Samoa; and eastern region, either a single cluster or 2 clusters, one around the Cook Islands and the other off French Polynesia. These results are consistent with the hypothesis that each breeding aggregation represents a distinct population (each occupied a single, terminal node) in a metapopulation, similar to the current understanding of population structure based on genetic and photo‐identification studies. However, the central vocal cluster had higher levels of song‐sharing among populations than the other clusters, indicating that levels of vocal connectivity varied within the region. Our results demonstrate the utility and value of using culturally transmitted vocal patterns as a way of defining connectivity to infer population structure. We suggest vocal patterns be incorporated by the International Whaling Commission in conjunction with traditional methods in the assessment of structure.     

Characterizing the spatial structure of songbird cultures

Recent advances have shown that human-driven habitat transformations can affect the cultural attributes of animal populations in addition to their genetic integrity and dynamics. Here I propose using the song of oscine birds for identifying the cultural spatial structure of bird populations and highlighting critical thresholds associated with habitat fragmentation. I studied song variation over a wide geographical scale in a small and endangered passerine, the Dupont's Lark Chersophilus duponti, focusing on (1) cultural population structure, to determine a statistical representation of spatial variation in song and identify cultural units, and (2) the minimum patch size needed for an individual to develop a stable repertoire. I found that overall song diversity depends on variation among populations (beta-cultural diversity). Abrupt thresholds occurred in the relationships between individual song dissimilarity and geographic distance, and between individual song diversity and patch area. Spatial autocorrelation analysis showed that populations located as little as 5 km apart may have independently evolved their song traditions. Song diversity stabilized in patches as small as 100 ha supporting as few as 8-20 males. Song repertoires of smaller patches were significantly poorer. Almost one-quarter of the study populations inhabited patches <100 ha, and their cultural traditions appear to have eroded. The analysis of spatial patterns in birdsong may be a useful tool for detecting subpopulations prone to extinction.     

Effects of Population Size on Singing Behavior of a Rare Duetting Songbird

Although the genetic and ecological effects of population declines in endangered species have been well studied, little is known of the social consequences. Changes in signaling behavior may result in disrupted communication and affect both reproductive and conflict‐resolution activities. The North Island Kōkako (Callaeas wilsoni) is an endangered, duetting (i.e., alternating, coordinated singing by mated pairs) songbird endemic to New Zealand temperate rain forests. Scattered populations (approximately 1500 individuals in 13 surviving and 11 translocated populations) in isolated conservation areas of different sizes have been rescued from extirpation and are currently recovering. We examined key song attributes of the Kōkako to assess whether population size or growth rate are related to song complexity, the reduction of which may compromise effective communication. We analyzed song repertoire size and phrase‐type sharing (i.e., Jaccard index of similarity), vocal performance (singing rates, song switching rates, and diversity of phrase types), and song syntactical characteristics (i.e., unpredictability in sequences of phrase types) in surviving and translocated populations (populations of approximately 19–250 territorial individuals). Population size was positively correlated with a population's song repertoire, song diversity, and switching of song phrase types and negatively correlated with shared phrase types and variation in syntactical structure of songs. Population growth rate correlated positively with pair repertoire size, population repertoire size, and singing rates during song bouts. As for solo‐singing species in fragmented landscapes, songs in the fragmented populations of Kōkako appear to be undergoing microevolution as occurs in island colonization events. Our results suggest that vocal changes in small populations could affect population establishment and growth, particularly in multiple‐source translocations. We believe measurement of vocal behavior could be used as a supplement to periodic population censuses to allow more frequent monitoring of population size. Efectos de la Conducta de Canto sobre el Tamaño Poblacional de una Ave Canora Rara     

Genes and song: genetic and social connections in fragmented habitat in a woodland bird with limited dispersal

Understanding the processes leading to population declines in fragmented landscapes is essential for successful conservation management. However, isolating the influence of disparate processes, and dispersal in particular, is challenging. The Grey Shrike-thrush, Colluricincla harmonica, is a sedentary woodland-dependent songbird, with learned vocalizations whose incidence in suitable habitat patches falls disproportionally with decline in tree cover in the landscape. Although it has been suggested that gaps in tree cover might act as barriers to its dispersal, the species remains in many remnants of native vegetation in agricultural landscapes, suggesting that it may have responded to habitat removal and fragmentation by maintaining or even increasing dispersal distances. We quantified population connectivity of the Grey Shrike-thrush in a system fragmented over more than 120 years using genetic (microsatellites) and acoustic (song types) data. First, we tested for population genetic and acoustic structure at regional and local scales in search of barriers to dispersal or gene flow and signals of local spatial structuring indicative of restricted dispersal or localized acoustic similarity. Then we tested for effects of habitat loss and fragmentation on genetic and acoustic connectivity by fitting alternative models of mobility (isolation-by-distance [the null model] and reduced and increased movement models) across treeless vs. treed areas. Birds within -5 km of each other had more similar genotypes and song types than those farther away, suggesting that dispersal and song matching are limited in the region. Despite restricted dispersal detected for females (but not males), populations appeared to be connected by gene flow and displayed some cultural (acoustic) connectivity across the region. Fragmentation did not appear to impact greatly the dispersal of the Grey Shrike-thrush: none of the mobility models fit the genetic distances of males, whereas for females, an isolation-by-distance model could not be rejected in favor of the models of reduced or increased movement through treeless gaps. However, dissimilarities of the song types were more consistent with the model of reduced cultural connectivity through treeless areas, suggesting that fragmentation impedes song type sharing in the Grey Shrike-thrush. Our paper demonstrates that habitat fragmentation hinders important population processes in an Australian woodland bird even though its dispersal is not detectably impacted.     

Male's return rate, rather than territory fidelity and breeding dispersal, explains geographic variation in song sharing in two populations of an oscine passerine (Oreothlypis celata)

Males of some oscine passerines learn and share songs of neighboring males. This process can lead to the formation of song pattern neighborhoods or microhabitat song dialects. The degree to which song sharing occurs between populations and the spatial scale over which neighboring males share songs, however, can vary widely, and interpopulation comparisons have suggested that song sharing is more common in residents than in migrants. Here, we examine two populations of the orange-crowned warbler (Oreothlypis celata) to quantify patterns of song sharing at the northern (long-distance migrant) and southern (short-distance migrant) edges of the breeding distribution and to test if return rate, territory fidelity, and breeding dispersal explain the patterns found in the two populations. The southern population (O. celata sordida breeding on Santa Catalina Island, California; 33°N) had a higher annual return rate to their territories and exhibited higher song sharing in neighborhoods than their counterparts (O. celata celata breeding in Fairbanks, Alaska; 64°N). Year-to-year patterns of territory fidelity and breeding dispersal distances were similar between populations. Our results suggest that if migratory distance generally covaries with the proportion of returning males, this could explain different levels of song sharing between the short- and long-distance migrants.     

Conservation Genetics in the Management of Desert Fishes

Abstract: The status and security of fishes in North American deserts have steadily declined in this century due to man's activities in this naturally fragile region. We address genetic aspects of the population structure of desert fishes as applicable to conservation and recovery programs by developing two zoogeographic models of isolation and gene flow. In the Death Valley model populations are isolated, with no chance of natural gene flow among them. Genetic diversity within populations tends to be low, but genetic divergence among populations within a species is high. In the Stream Hierarchy model, a complicated hierarchical genetic structure exists and is a function of geographic proximity and connectivity of habitats. Within-habitat genetic diversity tends to be higher, and among-habitat differentiation lower, than in the Death Valley model. These two systems must be recognized as distinct and managed differently. We also suggest three areas of experimentation needed to better understand and manage genetic stocks of desert fishes: relationships between heterozygosity and fitness, experimental mixing of similar stocks to examine effects of increased heterozygosity, and analysis of the relative roles of genetic adaptation and phenotypic plasticity in local differentiation.     

Culturally correct song dialects are correlated with male age and female song preferences in wild populations of brown-headed cowbirds

Variation in vocal behavior among local populations of songbirds may be significant to females in mate choice. In a study of dialect populations of brown-headed cowbirds, estradiol-implanted females from two dialects held the pre-copulatory lordosis posture longer in response to playback of the local flight whistle song than to foreign whistle types from adjacent and more distant dialects. Females were held in captivity for a relatively brief period prior to testing and received no tutoring so discrimination was based solely on experience in the wild. This is the first study to show evidence of discrimination by female cowbirds based on flight whistle type. Evidence is presented from one of the study dialects that the majority of yearling male cowbirds are vocally distinct from resident adults in having either foreign or incomplete local whistles. Although these yearlings are fully mature sexually, they have little or no mating success. The correlation between female whistle preference and male mating success suggests that the ability to give the correct local whistle type may be a characteristic used by females to assess age and quality of a potential mate. A learned female preference for the predominant local song type may be a factor in the stability of these dialects by making it adaptive for males to conform to the local dialect.     

Song of the humpback whale — Population comparisons

Summary Humpback whale songs recorded on tropical calving grounds exhibit different dialects depending on the oceanic basin. Songs sampled simultaneously from two populations in the North Pacific (Hawaii and Mexico) were essentially identical. These North Pacific songs were clearly different from the song type shared by two populations in the North Atlantic (Cape Verde Islands and West Indies). Songs from the Southern Hemisphere (Tonga) represent a third distinct dialect. Our evidence shows that, despite annual change in song organization, significant differences in humpback song occur between isolated ocean basins, while only subtle differences exist within an oceanic population (Hawaii and Pacific Mexico).     

Song sharing correlates with lifetime social pairing success but not territory tenure in the Puget Sound white-crowned sparrow

Imitative song learning in birds often results in the formation of vocal dialects or local song neighborhoods in which males in close proximity share very similar songs. If song sharing confers a selective advantage due to intra-sexual competition or inter-sexual mate choice, song sharing should enhance the singer’s territory tenure or lifetime pairing success. We tested this in a migratory population of the Puget Sound white-crowned sparrow, Zonotrichia leucophrys pugetensis, in which some territory neighbors share whole songs. Song sharing with territory neighbors in the first year on territory did not correlate with territory tenure but correlated positively with social pairing success throughout a male’s lifetime when length of territory tenure was controlled. Thus, song sharing in this migratory population does not appear to confer a benefit in male–male interactions, at least as measured by the length of territory tenure. However, males that share songs do benefit from higher lifetime pairing success. Combined with the data available for other species, it appears that intra- and inter-sexual selection may have stronger effects on song sharing in sedentary and migratory populations, respectively.     

Song sharing in two populations of song sparrows (Melospiza melodia)

Sharing song types with immediate neighbors is widespread in birds with song repertoires, and sharing songs may confer a selective advantage in some cases. Levels of song sharing vary between different geographical populations of several bird species, and ecological differences often correlate with differences in singing behavior; in particular, males in migratory subspecies often share fewer songs than males in resident subspecies. The song sparrow (Melospiza melodia) appears to fit this pattern: resident song sparrows in western North America generally share 20–40% of their repertoire (of about eight songs) with each neighbor, while migratory subspecies from eastern North America often share 10% or less. We compared song sharing in two populations within a single subspecies of song sparrow (M. m. morphna) in Washington State. These populations, separated by only 120 km, nonetheless differ in migratory tendencies and several other ecological and life history variables. We recorded complete song repertoires from 11 male song sparrows in a high-elevation, migrating population at Gold Creek in west-central Washington, and compared them to two samples (n = 15 and n = 36) from a coastal, resident population at Discovery Park, Seattle, Washington. Despite major differences in habitat, population density, and migratory tendencies, song sharing among Gold Creek males was as high as that among Discovery Park males. In both populations, sharing was highest between immediate neighbors, and declined with distance. We conclude that at the within-subspecies level, neither migration nor population density affect song sharing in song sparrows, a song repertoire species. Received: 26 November 1998 / Received in revised form: 1 May 1999 / Accepted: 29 May 1999     

Bearded seal males perceive geographic variation in their trills

Geographic variation in animal vocalisations, ranging from micro- to macro-variations, has now been widely documented. These dialects can impair communication between separated groups or populations and thus may play a role in speciation processes. Although the existence of geographical variation has been already shown in some pinnipeds species, the extent to which individuals perceive it is poorly understood. Here, we studied this question in the bearded seal Erignatus barbatus, a species found across different arctic regions. As in other phocids, bearded seal males emit sophisticated acoustic displays while defending an aquatic territory during the breeding season. First, we used playbacks to demonstrate that the trill has a function in territorial defence. Second, we used synthetic trills from two distinct populations (Nunavut and Greenland), to show that bearded seal males perceived the geographic variation in the trills. Males from Greenland responded more strongly to trills from local males than those from a distant area (Nunavut). This study provides the first experimental evidence that phocid males can perceive geographic variation in their vocalisations. Acoustic analyses combined with playback experiments can help to identify distinct populations in bearded seals. This may then provide information on both strength and scale of breeding site fidelity in this and potentially other species.     

Experiments on the origin of dialects in the short-toed tree creeper (Certhia brachydactyla)

Summary Wild short-toed tree creepers in southern Germany were exposed to tape recordings of seven song types from other dialects or song institutions and sixteen song types from thirteen handreared Middle European birds. One song type from southern Germany was used as a control. The males reacted strongly to two slightly different Spaish dialects and not at all or only feebly to one song institution from Cyprus and four from Morocco. The song types of the hand-reared birds had less effect than controls. The songs of two birds from southern Germany hand-raised together were quite similar to those from Cyprus. We conclude that Cyprus and North Africa on the outskirts of the species distribution were colonized by short-toed tree creepers who had had no opportunity to learn their songs from adult conspecifics.     

The song of the humpback whale Megaptera novaeangliae in the West Indies

Songs of the humpback whale Megaptera novaeangliae were recorded and analyzed from Grand Turks in the Bahamas to Venezuela. The design features of the song are as follows. The basic song evolves through a series of different sounds in a fixed order. The song is produced only in the winter tropical calving grounds, just before the whales arrive on the banks. Redundancy is high in that syllables, motifs, phrases and the entire song are repeated. Low, intermediate, and high-frequency sounds are scattered throughout the song. One sound is associated with blowing. The song appears to be partially different each year and there are some differences within a year between banks which may indicate that dialects are present. It is suggested that songs from other populations are quite different. The apparent yearly changes do not occur at one point in time. Only single individuals produce the song and they are hypothesized to be young, sexually mature males. The implications of these various design features are discussed.     

Sympatry and allopatry of song in North American tanagers

Summary The possibility that the hypotheses of character convergence and character displacement might explain interspecific variation in song features among North American tanagers is examined. Scarlet and summer tanagers (Piranga olivacea and P. rubra) are the most sympatric among the four species and have the largest differences in song characteristics. However, songs of scarlet tanagers do not differ significantly between sympatry and allopatry. Songs of summer tanagers more resemble those of scarlet tanagers in allopatry, but this geographical variation has been explained by variation in habitat characteristics. Both species countersing in sympatry, and react aggressively to each other's songs in sympatry as well as in allopatry, probably due to similarities in the song. The limited data from the narrow sympatric zone between western and hepatic tanagers (P. ludoviciana and P. flava) indicate that their songs differ to a great extent. Therefore, the results do not support the hypothesis of character convergence. Character displacement might explain partially interspecific differences in song features between scarlet and summer tanagers but cannot be tested further.     

Local adaptation and sexual selection: a reciprocal transfer experiment with the grasshopper Chorthippus biguttulus

Immigration into locally adapted populations has been suggested, among other potential causes, to maintain genetic variance in fitness necessary for good-genes models. Using a reciprocal transplant experiment we examined whether females prefer native to transferred males in the grasshopper Chorthippus biguttulus. On average, native and transferred males did not differ in their attractiveness, measured as female response rate to playbacks of male acoustic courtship signals. In line with this result, we found no significant effect of transfer on body size, condition, fluctuating asymmetry or song traits. However, the reciprocal transplant experiment showed that environmental conditions did influence body condition and maximum loudness of the calling song, but that the genetic origin of male grasshoppers had no significant effect on any of the analysed traits.Communicated by L. Simmons     

Song type variants and aggressive context

We asked whether switching among song type variants functions as a graded signal of aggression in song sparrows (Melospiza melodia). Song type variants are different versions of a given song type and are defined by much smaller acoustic differences than those defining separate song types. The principal evidence that switching between song types functions as an aggressive signal, in song sparrows and other species, is that song type switching frequencies increase during aggressive contexts, for example during simulated territorial intrusions. We tested whether variant switching frequencies change in the same way, using playback of conspecific song to simulate intrusions. Male song sparrows showed a significant increase in variant switching frequencies during intrusion relative to before intrusion. After intrusions, variant switching frequencies returned to pre-intrusion levels. These results are consistent with the hypothesis that variant switching is a graded aggressive signal, but other interpretations are possible, for example, that variant switching is a signal of submission or represents simple production error. Received: 20 January 2000 / Revised: 15 July 2000 / Accepted: 30 July 2000     

Securing the Demographic and Genetic Future of Tuatara through Assisted Colonization

Abstract: Climate change poses a particular threat to species with fragmented distributions and little or no capacity to migrate. Assisted colonization, moving species into regions where they have not previously occurred, aims to establish populations where they are expected to survive as climatic envelopes shift. However, adaptation to the source environment may affect whether species successfully establish in new regions. Assisted colonization has spurred debate among conservation biologists and ecologists over whether the potential benefits to the threatened species outweigh the potential disruption to recipient communities. In our opinion, the debate has been distracted by controversial examples, rather than cases where assisted colonization may be a viable strategy. We present a strategic plan for the assisted migration of tuatara (Sphenodon punctatus), an endemic New Zealand reptile. The plan includes use of extant populations as reference points for comparisons with assisted‐colonization populations with respect to demography, phenotypic plasticity, and phenology; optimization of genetic variation; research to fill knowledge gaps; consideration of host and recipient communities; and inclusion of stakeholders in the planning stage. When strategically planned and monitored, assisted colonization could meet conservation and research goals and ultimately result in the establishment of long‐term sustainable populations capable of persisting during rapid changes in climate.