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1.  Predicting bird song from space 
Evolutionary Applications  2013;6(6):865-874.
Environmentally imposed selection pressures are well known to shape animal signals. Changes in these signals can result in recognition mismatches between individuals living in different habitats, leading to reproductive divergence and speciation. For example, numerous studies have shown that differences in avian song may be a potent prezygotic isolating mechanism. Typically, however, detailed studies of environmental pressures on variation in animal behavior have been conducted only at small spatial scales. Here, we use remote-sensing data to predict animal behavior, in this case, bird song, across vast spatial scales. We use remotely sensed data to predict the song characteristics of the little greenbul (Andropadus virens), a widely distributed African passerine, found across secondary and mature rainforest habitats and the rainforest-savanna ecotone. Satellite data that captured ecosystem structure and function explained up to 66% of the variation in song characteristics. Song differences observed across habitats, including those between human-altered and mature rainforest, have the potential to lead to reproductive divergence, and highlight the impacts that both natural and anthropogenic change may have on natural populations. Our approach offers a novel means to examine the ecological correlates of animal behavior across large geographic areas with potential applications to both evolutionary and conservation biology.
doi:10.1111/eva.12072
PMCID: PMC3779089  PMID: 24062797
anthropogenic effects; avian song; behavioral ecology; random forests; remote sensing; reproductive isolation; spatial heterogeneity
2.  Trill consistency is an age-related assessment signal in banded wrens 
Older males tend to have a competitive advantage over younger males in sexual selection. Therefore, it is expected that signals used in sexual selection change with age. Although song repertoire size in songbirds is often mentioned as an age-related trait, many species, including the banded wren (Thryothorus pleurostictus), do not increase their repertoires after the first year. Here, we show that banded wrens reproduce the trill notes in their songs with less variability between them (i.e. more consistently) when they grow older. In a playback experiment, we also show that banded wrens discriminate between younger and older birds based on structural aspects of their song. In a second experiment, banded wrens also respond differentially to natural songs versus songs with artificially enhanced consistency. We argue that consistency in trill note reproduction may be achieved through practice. Sexual selection in the form of male–male competition may therefore operate on a phenotypic trait, the expression of which is enhanced by practice.
doi:10.1098/rspb.2009.0127
PMCID: PMC2677607  PMID: 19324742
sexual selection; competition; birdsong; age
3.  Syllable Type Consistency is Related to Age, Social Status, and Reproductive Success in the Tropical Mockingbird 
Animal behaviour  2009;77(3):701-706.
Many animals repeat standardized displays multiple times while attracting a mate or deterring a rival. In such contexts it is possible that the ability to perform each display or signal type in a consistent fashion is under direct selection. Studies on sexual selection on song learning in birds have focused on differences in repertoire size with less attention to the potential importance of being able to perform each song/syllable type with high consistency. We present evidence that tropical mockingbirds decrease the variation between renditions of each syllable type as they grow older (i.e., become more consistent) and that more consistent males in this species tend to have higher dominance status and reproductive success. These findings stress the importance of consistency in the performance of sexual displays and suggest that this parameter may be very relevant even in species that are selected for high vocal diversity (i.e., large repertoires). In addition to signalling dominance status and age, we hypothesize that syllable type consistency may also be an indicator of the integrity of brain function in birds analogous to the tests used for neuropsychological assessment in humans.
doi:10.1016/j.anbehav.2008.11.020
PMCID: PMC2805192  PMID: 20161558
Mimus gilvus; performance variability; song crystallization; song learning; tropical mockingbird
4.  Migration strategy and divergent sexual selection on bird song 
Migratory birds are assumed to be under stronger sexual selection pressure than sedentary populations, and the fact that their song is more complex has been taken as confirmation of this fact. However, this assumes that sexual selection pressure due to both male competition and female choice increase together. A further issue is that, in many species, songs become less complex during competitive encounters; in contrast, female choice selects for more complex song, so the two selection pressures may drive song evolution in different directions. We analysed song in two sedentary and two migratory populations of blackcaps (Sylvia atricapilla), a species in which different song parts are directed to males and females. We found that migratory populations produce longer, female-directed warbles, indicating sexual selection through female choice is the strongest in these populations. However, the part of the song directed towards males is shorter and more repetitive (as observed in individual competitive encounters between males) in non-migratory populations, indicating sedentary populations, are under stronger selection due to male competition. We show for the first time that the intensity of selection pressure from male competition and female choice varies independently between populations with different migratory behaviours. Rapid alterations in the migration patterns of species are thus likely to lead to unexpected consequences for the costs and benefits of sexual signals.
doi:10.1098/rspb.2008.1011
PMCID: PMC2664336  PMID: 18945666
blackcap; Sylvia atricapilla; song complexity; sexual selection; migration; warbler
5.  The deterrent effect of bird song in territory defense 
Behavioral Ecology  2008;20(1):200-206.
Using the responses of territory owners to playback to infer the territorial function of acoustic signals is common practice. However, difficulties with interpreting the results of such experiments have obscured our understanding of territorial signalling. For instance, a stronger response to playback is often interpreted as more aggressive, but there is no consensus as to whether this should be in response to the least or most threatening simulated intruder. Rather than following a gradual increase or decrease, the relationship between signal intensity and response strength may instead describe a peaked curve. We manipulated banded wren (Thryophilus pleurostictus) songs to simulate low-, median-, and high-performance singers and used these songs as stimuli in playback experiments. Banded wrens were less likely to approach the high-performance stimulus compared with the low- and median-performance stimuli. However, the birds that did approach the high-performance stimulus sang more than those that approached the low-performance stimulus. In addition, birds were more likely to match the songs when exposed to the median- and high-performance stimuli compared with the low-performance stimuli, and song matching predicted approach behavior. These results are in accordance with theoretical models of aggressive encounters in which low-performance opponents are challenged without further assessment. Median- and high-performance opponents, however, may require further assessment, and the latter may be perceived as too intimidating for approach.
doi:10.1093/beheco/arn135
PMCID: PMC2662740  PMID: 19337589
assessment; playback; sexual selection; song; territory defense
6.  The deterrent effect of bird song in territory defense 
Using the responses of territory owners to playback to infer the territorial function of acoustic signals is common practice. However, difficulties with interpreting the results of such experiments have obscured our understanding of territorial signaling. For instance, a stronger response to playback is often interpreted as more aggressive, but there is no consensus as to whether this should be in response to the least or most threatening simulated intruder. Rather than following a gradual increase or decrease, the relationship between signal intensity and response strength may instead describe a peaked curve. We manipulated banded wren (Thryophilus pleurostictus) songs to simulate low, median and high performance singers and used these songs as stimuli in playback experiments. Banded wrens were less likely to approach the high performance stimulus compared to the low and median performance stimuli. However, the birds that did approach the high performance stimulus sang more than those that approached the low performance stimulus. In addition, birds were more likely to match the songs when exposed to the median and high performance stimuli compared to the low performance stimuli and song matching predicted approach behavior. These results are in accordance with theoretical models of aggressive encounters in which low performance opponents are challenged without further assessment. Median and high performance opponents however, may require further assessment and the latter may be perceived as too intimidating for approach.
doi:10.1093/beheco/arn135
PMCID: PMC2662740  PMID: 19337589
playback; song; territory defense; sexual selection; assessment
7.  Habitat-dependent call divergence in the common cuckoo: is it a potential signal for assortative mating? 
The common cuckoo (Cuculus canorus) is an obligate brood parasite that mimics the eggs of its hosts. The host-specific egg pattern is thought to be inherited matrilinearly, creating female-only host-specific races. Males are thought not to be adapted to their host and they maintain the species by mating arbitrarily with respect to host specialization of females. However, recent results suggest that male cuckoos may also show host-specific adaptations and these may require assortative mating with respect to host. The calls males produce on the breeding grounds could provide a potential mechanism for assortative mating. We tested whether male cuckoo calls differ more between nearby populations that parasitize different hosts than between distant populations that parasitize the same host. We recorded the calls of geographically distant pairs of populations in Hungary, with each pair consisting of a forest population and a nearby reed bed population. Each habitat is characterized by one main host species for the common cuckoo. Our results show that calls of distant cuckoo populations from the same habitat type are more similar to each other than they are to those of nearby populations from a different habitat. These results suggest that cuckoo calls differ sufficiently to allow recognition of habitat-specific individuals.
doi:10.1098/rspb.2007.0487
PMCID: PMC2706194  PMID: 17580296
brood parasitism; cuckoo; vocalizations; mate recognition; host specific
8.  An evolutionary perspective on caching by corvids 
A principal finding in the food-caching literature is that species differences in hoarding propensity are positively correlated with species differences in degree of adaptations to caching behaviour, such as performance on spatial memory tasks and hippocampal volume. However, there are examples that do not fit this pattern. We argue that these examples can be better understood by considering the phylogenetic relatedness between species. We reconstruct the ancestral state for caching behaviour in corvids and assess when transitions in caching behaviour occurred within the corvid phylogeny. Our analysis shows that the common ancestor of all corvids was a moderate cacher. This result suggests that corvids followed a bi-directional evolutionary trajectory in which caching was secondarily lost twice and there were at least two independent transitions from moderate to specialized caching. The independent evolution of specialized cachers in the two groups must, therefore, be a case of convergent evolution. This is exemplified by the fact that specialized cachers show structurally different adaptations serving the same function to intense caching, such as different pouches to transport food. Finally, we argue that convergent evolution may have led to adaptations in memory and hippocampus that serve the same function but differ in design, and that these different adaptations may explain the examples that do not fit the pattern predicted by the adaptive specialization hypothesis.
doi:10.1098/rspb.2005.3350
PMCID: PMC1560201  PMID: 16615207
spatial memory; hippocampus; adaptive specialization; food hoarding; phylogeny; corvid
9.  Does hippocampal size correlate with the degree of caching specialization? 
A correlation between the degree of specialization for food hoarding and the volume of the hippocampal formation in passerine birds has been accepted for over a decade. The relationship was first demonstrated in family-level comparisons, and subsequently in species comparisons within two families containing a large number of hoarding species, the Corvidae and the Paridae. Recently, this approach has been criticized as invalid and excessively adaptationist. A recent test of the predicted trends with data pooled from previous studies found no evidence for such a correlation in either of these two families. This result has been interpreted as support for the critique. Here we reanalyse the original dataset and also include additional new data on several parid species. Our results show a surprising difference between continents, with North American species possessing significantly smaller hippocampi than Eurasian ones. Controlling for the continent effect makes the hoarding capacity/hippocampal formation correlation clearly significant in both families. We discuss possible reasons for the continent effect.
doi:10.1098/rspb.2004.2912
PMCID: PMC1523289  PMID: 15590591
10.  Does hippocampal size correlate with the degree of caching specialization? 
A correlation between the degree of specialization for food hoarding and the volume of the hippocampal formation in passerine birds has been accepted for over a decade. The relationship was first demonstrated in family-level comparisons, and subsequently in species comparisons within two families containing a large number of hoarding species, the Corvidae and the Paridae. Recently, this approach has been criticized as invalid and excessively adaptationist. A recent test of the predicted trends with data pooled from previous studies found no evidence for such a correlation in either of these two families. This result has been interpreted as support for the critique. Here we reanalyse the original dataset and also include additional new data on several parid species. Our results show a surprising difference between continents, with North American species possessing significantly smaller hippocampi than Eurasian ones. Controlling for the continent effect makes the hoarding capacity/hippocampal formation correlation clearly significant in both families. We discuss possible reasons for the continent effect.
doi:10.1098/rspb.2004.2912
PMCID: PMC1523289  PMID: 15590591
hippocampus; hippocampal formation; neuroecology; Paridae; Corvidae

Results 1-10 (10)