Winter GPS tagging reveals home ranges during the breeding season for a boreal-nesting migrant songbird, the Golden-crowned Sparrow.

Determining space use for species is fundamental to understanding their ecology, and tracking animals can reveal insights into their spatial ecology on home ranges and territories. Recent technological advances have led to GPS-tracking devices light enough for birds as small as ~30 g, creating novel opportunities to remotely monitor fine-scale movements and space use for these smaller species. We tested whether miniaturized GPS tags can allow us to understand space use of migratory birds away from their capture sites and sought to understand both pre-breeding space use as well as territory and habitat use on the breeding grounds. We used GPS tags to characterize home ranges on the breeding grounds for a migratory songbird with limited available breeding information, the Golden-crowned Sparrow (Zonotrichia atricapilla). Using GPS points from 23 individuals across 26 tags (three birds tagged twice), we found home ranges in Alaska and British Columbia were on average 44.1 ha (95% kernel density estimate). In addition, estimates of territory sizes based on field observations (mean 2.1 ha, 95% minimum convex polygon [MCP]) were three times smaller than 95% MCPs created using GPS tags (mean 6.5 ha). Home ranges included a variety of land cover classes, with shrubland particularly dominant (64-100% of home range cover for all but one bird). Three birds tracked twice returned to the same breeding area each year, supporting high breeding site fidelity for this species. We found reverse spring migration for five birds that flew up to 154 km past breeding destinations before returning. GPS-tracking technology allowed for critical ecological insights into this migratory species that breeds in very remote locations.

Modeling how population size drives the evolution of birdsong, a functional cultural trait.

Oscine songbirds have been an important study system for social learning, particularly because their learned songs provide an analog for human languages and music. Here, we propose a different analogy: from an evolutionary perspective, could birds' songs change over time more like arrowheads than arias? Small improvements to a bird's song can lead to large fitness differences for its singer, which could make songs more analogous to human tools than languages. We modify a model of human tool evolution to accommodate cultural evolution of birdsong: each song learner chooses the most skilled available tutor to emulate, and each is more likely to produce an inferior copy than a superior one. Similar to human tool evolution, our model suggests that larger populations of birds could foster improvements in song over time, even when learners restrict their pool of tutors to a subset of individuals in their social network. We also demonstrate that song elements could be simplified instead of lost after population bottlenecks if lower quality traits are easier to imitate than higher quality ones. We show that these processes could plausibly generate empirically observed patterns of song evolution for some song traits, and we make predictions about the types of song elements most likely to be lost when populations shrink. More broadly, we aim to connect the modeling approaches used in human and nonhuman systems, moving toward a cohesive theoretical framework that accounts for both cognitive and demographic processes.

To accept or reject heterospecific mates: behavioural decisions underlying premating isolation.

Premating isolation in animals involves decision-making processes that affect whether individuals accept or reject heterospecific mates. An integrative understanding of the behavioural processes underlying heterospecific acceptance can clarify the conditions under which premating isolation evolves. As an illustration, we review how Reeve's (Reeve 1989 Am. Nat.133, 407-435. (doi:10.1086/284926)) acceptance threshold model can help make sense of patterns of premating isolation in nature. This model derives a threshold trait value for acceptance for rejection of recipients of an action (e.g. mating) based on the fitness consequences of these decisions. We show that the maintenance of partial reproductive isolation can be an outcome of optimal acceptance thresholds, even in the face of reinforcement. We also use this model to clarify how the composition of multispecies communities can shape premating isolation. The acceptance threshold model can also be viewed as the behavioural underpinning of reproductive character displacement and cascading reinforcement. Finally, we highlight potential limitations of the acceptance threshold model with respect to investigating the role of sexual selection in speciation, and we propose that integration of behavioural models in speciation research will help us gain a full picture of the mechanisms underlying premating isolation. This article is part of the theme issue 'Signal detection theory in recognition systems: from evolving models to experimental tests'.

Pervasive reinforcement and the role of sexual selection in biological speciation.

Sexual selection has been widely implicated as a driver of speciation. However, allopatric forms are often defined as species based on divergence in sexually selected traits and it is unclear how much such trait differences affect reproductive isolation upon secondary contact, the defining feature of biological species. We show that in birds, divergence in song and plumage in allopatry corresponds poorly with whether species mate assortatively in hybrid zones and argue that this is because many other factors besides trait divergence affect propensity to hybridize, including rarity of conspecific mates and choice based on territory rather than male traits. We then present a general model for the establishment of sympatry that assumes a period of differentiation in allopatry followed by secondary contact and often hybridization, with hybridization subsequently reduced by reinforcement of mate preferences. We suggest that reinforcement commonly operates by a narrowing of a "window of recognition" for traits that are different between the species, rather than evolution of the traits themselves. Our arguments imply that it is important to study postmating as well as premating reproductive isolation in limiting sympatry and suggest that studies of reinforcement should focus on evolution of female preferences for diagnostic traits, rather than evolution of traits per se.