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.     

Song learning in montane white-crowned sparrows: from whom and when

White-crowned sparrow, Zonotrichia leucophrys, adults and nestlings were colour-banded and songs of mature progeny recorded to determine whether songs were learned from fathers, age peers or adults other than fathers. Four out of 17 sons and daughters sang the same song types as their fathers, indicating that transmission of song from fathers to their young (vertical transmission) is not the norm. Since juveniles do not sing full (‘crystallized’) songs until they are older, transmission from juvenile to juvenile (horizontal) is unlikely on the breeding grounds but may occur if siblings migrate together and hear each other's song on the wintering grounds. The usual mode of transmission is via other adults (oblique). Birds learn one or two songs prior to or during migration, and upon settling on the breeding area the following year may utter only one theme, that which matches the neighbours' songs during matched countersinging bouts. A bird's song dialect is thus not necessarily an indicator of its birthplace. One wild and one laboratory-reared bird sang songs of a Lincoln sparrow, Melospiza lincolnii, and a strawberry finch, Amandava amandava, respectively. These data and data from field playback studies lend support to the notion that interspecific terrioriality may lead to acquisition of alien song for use in matched countersinging during territorial encounters with allospecific neighbours.     

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

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

Energy metabolism, testosterone and corticosterone in white-crowned sparrows

The influence of the steroid hormones testosterone and corticosterone on energy metabolism and activity of birds is largely enigmatic. We measured resting metabolic rate during night and day in 12 long-term castrated and 12 intact male white-crowned sparrows (Zonotrichia leucophrys gambelii) under short-day (8:16 SD), long-day (20:4 LD), LD+testosterone implant and LD−testosterone implant conditions. Each male was sequentially measured under all four conditions. Photostimulation increased testosterone, resting metabolic rate, food intake, hopping activity and body mass in castrates and intact males. Surprisingly, testosterone levels and metabolic rates did not differ between intact and castrated males. Testosterone implantation increased activity and food intake, but decreased body mass and resting metabolic rate in both groups. Removing testosterone implants reversed the effects on resting metabolic rate, activity and food intake. Corticosterone levels, measured immediately at the end of metabolism measurements, showed birds were not stressed. Corticosterone had no apparent relationship with resting metabolic rate and there was no interaction between corticosterone and testosterone. Overall, positive changes in testosterone levels resulted in a decrease of resting metabolic rate. We speculate that testosterone increases activity, and birds compensate for increased activity metabolism by reducing resting metabolic rate. Accepted: 18 July 1999     

Aspects of risk-aversion in foraging white-crowned sparrows

Two series of aviary experiments with white-crowned sparrows (Zonotrichia leucophrys) are reported. The first series attempts to mimic foraging choices between microhabitats. The second series examines foraging choices made within a single artificial patch, a design intended to resemble dietary selection. In all experiments subjects chose between a constant food reward and a variable reward, the mean of which equalled the constant reward. Each experiment allowed the bird to feed at an overall rate greater than that minimally required for daily energy balance. In both series of experiments, the birds usually preferred the constant reward. Results of the first series do not differ significantly from previously reported preferences of dark-eyed juncos (Junco hyemalis), birds weighing 10 g less than the white-crowned sparrows.     

Yelkouan shearwater nest-cavity selection and breeding success

We describe, for the first time, nest-cavities selected by the Mediterranean endemic yelkouan shearwater Puffinus yelkouan on French islands, comparing it with the sympatric Cory's shearwater Calonectris diomedea. By monitoring 179 suitable cavities during four years, we show that yelkouan shearwaters select deeper cavities and with more winding tunnels than those selected by Cory's shearwaters or unoccupied cavities. Logistic regression modelling indicates that cavities with nest-like characteristics are not limited. We show that breeding success is enhanced in deeper cavities, with winding tunnels and higher block covers. We do not find any apparent competition between the two shearwater species.     

Effects of breeding success, mate fidelity and senescence on breeding dispersal of male and female blue-footed boobies

1. Understanding the effects of individual and population factors on variation in breeding dispersal (the movement of individuals between successive breeding sites) is key to identifying the strategies behind breeders' movements. Dispersal is often influenced by multiple factors and these can be confounded with each other. We used 13 years of data on the locations, mates, breeding success and ages of individuals to tease apart the factors influencing breeding dispersal in a colonially breeding long-lived seabird, the blue-footed booby Sula nebouxii. 2. Breeding dispersal varied among and within years. Males dispersed further in years of higher population density, and late breeding males and females dispersed further than early breeders. This temporal variation related to changes in competition for territory was taken into account in all tests of individual factors influencing breeding dispersal. 3. Individuals that retained their mates from the previous year dispersed shorter distances than those that changed their mates. 4. The effect of previous breeding success depended on mate fidelity. Unsuccessful breeding induced greater dispersal in birds that changed their mates but not in birds that retained their mates, indicating that breeders who change mates may take their own previous breeding experience into account during habitat selection. Faithful individuals may have to stay close to their previous sites to encounter their mates. 5. Male divorcees dispersed over shorter distances than their former mates, possibly because males contribute more than females to establishing territories. 6. Dispersal of males and females declined with increasing age over the first 10-11 years of life, then increased in old age, possibly due to senescent decay in the ability to compete for mates and territories.     

Re-expression of songs deleted during vocal development in white-crowned sparrows, Zonotrichia leucophrys

White-crowned sparrows learn and produce multiple song types as juveniles, but most individuals stop singing all except one by the end of the first singing season. This single song type is generally maintained throughout adulthood. We demonstrate that, at the start of the second and subsequent singing seasons, this species can recall songs that had been deleted during the first singing season. The re-expression of song occurred in both the oriantha and the gambelii subspecies. Although all our males recrystallized the original song in the second year, our results indicate a mechanism for seasonal song change without new song memorization. The traditional dichotomy of closed-ended versus open-ended learning is inadequate for birds that learn early in life but can change their song output seasonally. We suggest that species can exhibit a closed sensitive period for song memorization and first production, with the ability to recall deleted songs later in life. This type of learning, selective attrition followed by subsequent re-expression, may be used by some species currently considered open-ended learners. Copyright 2000 The Association for the Study of Animal Behaviour.     

Signals of status in wintering white-crowned sparrows, Zonotrichia leucophrys gambelii

The possibility of plumage status signalling within the social systems of wintering birds has been a controversial issue. Our results are the first to demonstrate conclusively the reality of such signalling. Data from eight groups of captive white-crowned sparrows (Zonotrichia leucophrys gambelii), each with 8 to 11 different individuals, show that immature and adult females with crowns painted to resemble more brightly coloured, dominant adult males consistently win encounters with control birds of their own age and sex. These experiments demonstrate that signals that correlate with age (adult versus immature) and sex (adult male versus adult female) are used by the birds as reliable indicators of relative dominance position. Our demonstration of status signalling draws attention to the need to explain how such a system can be evolutionarily stable and we discuss some suitable models.     

Optimization of selection contribution and mate allocations in monoecious tree breeding populations

Background

The combination of optimized contribution dynamic selection and various mating schemes was investigated over seven generations for a typical tree breeding scenario. The allocation of mates was optimized using a simulated annealing algorithm for various object functions including random mating (RM), positive assortative mating (PAM) and minimization of pair-wise coancestry between mates (MCM) all combined with minimization of variance in family size and coancestry. The present study considered two levels of heritability (0.05 and 0.25), two restrictions on relatedness (group coancestry; 1 and 2%) and two maximum permissible numbers of crosses in each generation (100 and 400). The infinitesimal genetic model was used to simulate the genetic architecture of the trait that was the subject of selection. A framework of the long term genetic contribution of ancestors was used to examine the impacts of the mating schemes on population parameters.

Results

MCM schemes produced on average, an increased rate of genetic gain in the breeding population, although the difference between schemes was small but significant after seven generations (up to 7.1% more than obtained with RM). In addition, MCM reduced the level of inbreeding by as much as 37% compared with RM, although the rate of inbreeding was similar after three generations of selection. PAM schemes yielded levels of genetic gain similar to those produced by RM, but the increase in the level of inbreeding was substantial (up to 43%).

Conclusion

The main reason why MCM schemes yielded higher genetic gains was the improvement in managing the long term genetic contribution of founders in the population; this was achieved by connecting unrelated families. In addition, the accumulation of inbreeding was reduced by MCM schemes since the variance in long term genetic contributions of founders was smaller than in the other schemes. Consequently, by combining an MCM scheme with an algorithm that optimizes contributions of the selected individuals, a higher long term response is obtained while reducing the risk within the breeding program.     

Seasonal changes in song nuclei and song behavior in Gambel's white-crowned sparrows

In males of several songbird species, the morphology of forebrain nuclei that control song changes seasonally. The only seasonally breeding songbird in which seasonal changes in the structure of song control nuclei have been reported not to occur is the nonmigratory Nuttall's subspecies of white-crowned sparrow. In the present study, we manipulated photoperiod and plasma testosterone concentrations in captive male white-crowned sparrows of the migratory Gambel's subspeices. Males exposed to photoperiods and plasma testosterone concentrations typical of those experienced by wild breeding males had larger song control nuclei than males held on a winter photoperiod. We also found seasonal change in stereotypy of spectral and temporal parameters of song in wild Gambel's white-crowned sparrows. We hypothesize that seasonal changes in song control nuclei may correlate with seasonal changes in song stereotypy. © 1995 John Wiley & Sons, Inc.     

Genetic analysis of song dialect populations in Puget Sound white-crowned sparrows

The relationship between cultural variation and biological variationamong natural populations has been the subject of both theoreticaland empirical study. Zonotrichia leucophrys pugetensis is oneof three subspecies of white-crowned sparrow known to form geographicalsong dialects. We investigated whether these dialects correspondto genetic differences among Z. l. pugetensis populations. Wecompared allele frequencies at four microsatellite loci in malesfrom 11 sites spanning six dialects over the subspecies' rangein Oregon and Washington. Cluster analysis and genotype assignmenttests indicated no tendency for sample sites within dialectareas to be genetically more similar than are sites from differentdialect areas. AMOVA tests revealed high within-site variationand low but significant cross-site and cross-dialect-area variation.Finally, genetic distance between sites was not correlated withdialect differences when the effect of geographic distance wascontrolled statistically. We compare our finding of low geneticdifferentiation among Z. l. pugetensis dialect populations toresults of previous studies on Z. l. nuttalli and Z. l. oriantha.Because genetic structuring appears weaker than cultural (songdialect) structure in this species, we discuss the behavioralmechanisms underlying dialect maintenance in the presence ofapparent gene flow.     

Habitat selection and breeding success in Yellow-legged Gulls Larus cachinnans

The habitat selection and breeding performance of Yellow-legged Gulls Larus cachinnans were studied in the Medes Islands colony, northeastern Spain, during 1995 and 1996. Of the three main habitats on the islands (shrubs, grass and bare areas), gulls first occupied those with the highest percentage of tall vegetation. Gulls tended to select nest sites with 20–75% cover despite great differences in the cover in the habitats and territories, suggesting that the presence of a suitable nest site may play a major role in the choice of breeding habitat. Nest-site tenacity did not influence the preferences of gulls at any level since the same pattern of choice was observed in an area subjected to annual culls (i.e. where most of the breeding pairs were culled annually and replaced by naive birds). In spite of great differences in the physical characteristics of the habitats, little difference was found in breeding performance of the gulls between habitats. Gulls nesting in the least preferred habitat (i.e. mainly bare) had smaller clutches than those nesting in the other two habitats, possibly as a result of their later seasonal laying. Despite the similar breeding success in different habitats, gulls did not seem to distribute according to the ideal free model reported for Herring Gulls Larus argentatus since the density in the preferred habitat (i.e. shrubs) was never higher than in the other two. We suggest that the habitat selection by Yellow-legged Gulls within the colony could follow an ideal despotic distribution.     

Resource availability, breeding site selection, and reproductive success of red-winged blackbirds

Red-winged blackbirds are polygynous and show strong breeding site preferences, but it is unclear which environmental factors regulate their reproductive success and are ultimately responsible for shaping their patterns of habitat selection and their mating system. We evaluated the effect of variation in insect emergence rates on the reproductive success of male and female redwings nesting on replicate ponds. The number of male and female redwings that settled on a pond varied two- to three-fold among ponds, but was not related to insect emergence rates. Insect emergence rates had a positive effect on the number of nestlings successfully fledged by females, the number of nestlings fledged from male territories, and on the mass of nestlings at fledging. Typha stem density also varied widely among ponds, and was positively related to male and female settling density and mass of nestlings at fledging, but not to the number of nestlings fledged by females or males. We conclude that alternative breeding sites differ in their ability to support redwing reproduction, and that the availability of emerging odonates is an important environmental factor influencing the reproductive success of both male and female red-winged blackbirds. Received: 31 March 1997 / Accepted: 3 July 1997     

From song dialects to speciation in white‐crowned sparrows

The behavioural signals used in mate selection are a key component in the evolution of premating isolating barriers and, subsequently, the formation of new species. The importance of mating signals has a long tradition of study in songbirds, where many species differ in their song characteristics. In oscine songbirds, individual birds usually learn their songs from a tutor. Mistakes during learning can help generate geographic dialects, akin to those within human language groups. In songbirds, dialect differences can often be substantial and there is an intuitive connection between the evolution of song amongst populations at a small scale, and the more substantive song differences between bird species and presumably used in species recognition. However, studies investigating the concordance between putative genetic and behavioural boundaries have generated mixed results. In many cases, this is possibly a function of the poor resolving power of the genetic markers employed. In this issue of Molecular Ecology, Lipshutz et al. ( 2017 ) combine genomic markers with a robust behavioural assay to address the importance of song variation amongst white‐crowned sparrow (Zonotrichia leucophrys) subspecies.     

Pike predation affects breeding success and habitat selection of ducks

1. Fish and ducks often belong to the same local food web, and several studies indicate that there is a general negative effect of fish on breeding ducks. This pattern has so far been addressed mainly within the framework of competition for common invertebrate prey, while predation by large fish as a force behind settlement and abundance patterns in ducks remains largely unknown. This is the first study to address the effect of fish predation on breeding ducks, isolated from that of competition, and the first experiment to explore the ability of ducks to identify and avoid lakes with high risk of fish predation. 2. We used a before–after control–impact design and 11 naturally fishless lakes. Waterfowl on the lakes were surveyed during the breeding season of 2005. Large adult pike (Esox lucius) were added to two lakes in early spring 2008, and waterfowl surveys were repeated on all 11 lakes. 3. Pike introduction did not affect the number of pairs on lakes during the nesting season in any of three focal duck species (mallard Anas platyrhynchos, teal Anas crecca, and goldeneye Bucephala clangula). During the brood‐rearing season, however, there was a decrease in duck days in teal and goldeneye in lakes with pike, with similar trends observed in mallard. The number of goldeneye ducklings was also significantly lower in lakes with pike. We were unable to determine whether the response was attributable to direct pike predation or to broods leaving experimental lakes, but in either case, our study demonstrates high fitness costs for ducks breeding on lakes with pike. 4. The apparent inability of nesting ducks to detect pike and the clear fitness implications may influence the annual recruitment of ducks on a larger scale as pike are both common and widespread. Vegetation complexity and food abundance are likely to be of overriding importance when breeding ducks are choosing a nesting site. As pike have a strong influence on breeding birds, relying on vegetation and cues of food abundance, while ignoring indicators of predation risk from fish, could lead to lakes with pike acting as an ecological trap.