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Concept: Thrush Nightingale


Music is thought to engage its listeners by driving feelings of surprise, tension, and relief through a dynamic mixture of predictable and unpredictable patterns, a property summarized here as “expressiveness”. Birdsong shares with music the goal to attract its listeners' attention and might use similar strategies to achieve this. We here tested a thrush nightingale’s (Luscinia luscinia) rhythm, as represented by song amplitude envelope (containing information on note timing, duration, and intensity), for evidence of expressiveness. We used multifractal analysis, which is designed to detect in a signal dynamic fluctuations between predictable and unpredictable states on multiple timescales (e.g. notes, subphrases, songs). Results show that rhythm is strongly multifractal, indicating fluctuations between predictable and unpredictable patterns. Moreover, comparing original songs with re-synthesized songs that lack all subtle deviations from the “standard” note envelopes, we find that deviations in note intensity and duration significantly contributed to multifractality. This suggests that birdsong is more dynamic due to subtle note timing patterns, often similar to musical operations like accelerando or crescendo. While different sources of these dynamics are conceivable, this study shows that multi-timescale rhythm fluctuations can be detected in birdsong, paving the path to studying mechanisms and function behind such patterns.

Concepts: Sound, Music, Nightingale, Thrush, Luscinia, Thrush Nightingale


A new computational model based on an optimal power, wake-only aerodynamics method is presented to predict the interdependency of energetics and kinematics in bird and bat flight. The model is divided into offline, intermediate and online modules. In the offline module, a four-dimensional design space sweep is performed (lift, thrust, flapping amplitude and flapping frequency). In the intermediate stage, the physical characteristics of the animal are introduced (wing span, mass, wing area, aspect ratio, etc.), and a series of amplitude-frequency response surfaces are constructed for all viable flight speeds. In the online component, the amplitude-frequency response surfaces are mined for the specific flapping motions being considered. The method is applied to several biological examples including a medium sized fruit bat (Cynopterus brachyotis), and two birds: a thrush nightingale (Luscinia luscinia) and a budgerigar (Melopsittacus undulatus). For each of these animals, the power and kinematics predictions are compared with available experimental data. These examples demonstrate that this new method can reasonably predict animal flight energetics and kinematics.

Concepts: Scientific method, Futurology, Bird, Wing, Flying and gliding animals, Nightingale, Luscinia, Thrush Nightingale


Interspecific competition is assumed to play an important role in the ecological differentiation of species and speciation. However, empirical evidence for competition’s role in speciation remains surprisingly scarce. Here we studied the role of interspecific competition in the ecological differentiation and speciation of two closely related songbird species, the Common Nightingale (Luscinia megarhynchos) and the Thrush Nightingale (L. luscinia). Both species are insectivorous and ecologically very similar. They hybridize in a secondary contact zone, which is a mosaic of sites where both species co-occur (syntopy) and sites where only one species is present (allotopy). We analyzed fine-scale habitat data for both species in both syntopic and allotopic sites and looked for associations between habitat use and bill morphology, which have been previously shown to be more divergent in sympatry than in allopatry. We found that the two nightingale species differ in habitat use in allotopic sites, where L. megarhynchos occurred in drier habitats and at slightly higher elevations, but not in syntopic sites. Birds from allotopic sites also showed higher interspecific divergence in relative bill size compared to birds from syntopic sites. Finally, we found an association between bill morphology and elevation. Our results are consistent with the view that interspecific competition in nightingales has resulted in partial habitat segregation in sympatry, and that the habitat-specific food supply has in turn very likely led to bill size divergence. Such ecological divergence may enhance prezygotic as well as extrinsic postzygotic isolation and thus accelerate the completion of the speciation process. This article is protected by copyright. All rights reserved.

Concepts: Evolution, Species, Bird, Speciation, Nightingale, Thrush, Luscinia, Thrush Nightingale


1.The role of interspecific competition for generating patterns in species' distribution is hotly debated and studies taking into account processes occurring at both large and small spatial scales are almost missing. Theoretically, competition between species with overlapping niches should result in divergence of their niches in sympatry to reduce the costs of competition. Many species show a mosaic distribution within sympatric zones, with the syntopic sites occupied by both species, and allotopic sites where only one species occurs. It is unclear whether such mosaics arise as a consequence of competition-driven niche segregation or due to the decline of their abundances towards range edges driven by environmental gradients. 2.If the interspecific competition matters, we should observe (i) a shift in habitat preferences of one or both species between syntopy and allotopy, and (ii) between allopatry and allotopy. Moreover, (iii) species should show greater divergence in their habitat preferences in allotopy than in allopatry where (iv) no differences in habitat preferences may occur. Finally, (v) shifts should be generally greater in the competitively subordinate species than in the dominant species. 3.We used a unique dataset on abundance of two closely related passerine species, the Common Nightingale (Luscinia megarhynchos) and the Thrush Nightingale (Luscinia luscinia), collected across their syntopy, allotopy and allopatry. The predictions were tested within a generalized mixed-effects modelling framework. 4.After accounting for environmental gradients perpendicular to the species' contact zone, we found a strong support for all but one prediction. Habitat preferences of both species shifted markedly between syntopy and allotopy, as well as between allopatry and allotopy. Whereas the species preferred the same habitats in allopatry, their preferences became strikingly different in allotopy where the abundance of the Common Nightingale increased towards dry and warm sites with low coverage of pastures, while the abundance of the Thrush Nightingale showed exactly opposite trends. Fifth prediction was not supported. 5.Our results indicate that the competition between closely related species can result in considerable changes in habitat use across their geographic ranges accompanied with divergence in their habitat preferences in sympatry. Here the species “escape” from competition to allotopic sites covered by habitats avoided by the competitor. Therefore, we argue that the interspecific competition is an important driver of species' distribution at both large and small spatial scales. This article is protected by copyright. All rights reserved.

Concepts: Competition, Nightingale, Thrush, Luscinia, Thrush Nightingale


Species of Plasmodium (Plasmodiidae, Haemosporida) are widespread and cause malaria, which can be severe in avian hosts. Molecular markers are essential to detect and identify parasites, but still absent for many avian malaria and related haemosporidian species. Here, we provide first molecular characterization of Plasmodium matutinum, a common agent of avian malaria. This parasite was isolated from a naturally infected thrush nightingale Luscinia luscinia (Muscicapidae). Fragments of mitochondrial, apicoplast and nuclear genomes were obtained. Domestic canaries Serinus canaria were susceptible after inoculation of infected blood, and the long-lasting light parasitemia developed in two exposed birds. Clinical signs of illness were not reported. Illustrations of blood stages of P. matutinum (pLINN1) are given, and phylogenetic analysis identified the closely related avian Plasmodium species. The phylogeny based on partial cytochrome b (cyt b) sequences suggests that this parasite is most closely related to Plasmodium tejerai (cyt b lineage pSPMAG01), a common malaria parasite of American birds. Both these parasites belong to subgenus Haemamoeba, and their blood stages are similar morphologically, particularly due to marked vacuolization of the cytoplasm in growing erythrocytic meronts. Molecular data show that transmission of P. matutinum (pLINN1) occurs broadly in the Holarctic, and the parasite likely is of cosmopolitan distribution. Passeriform birds and Culex mosquitoes are common hosts. This study provides first molecular markers for detection of P. matutinum.

Concepts: Malaria, Plasmodium falciparum, Plasmodium, Apicomplexa, Nightingale, Thrush, Luscinia, Thrush Nightingale


Songbirds spend much of their time learning, producing, and listening to complex vocal sequences we call songs. Songs are learned via cultural transmission, and singing, usually by males, has a strong impact on the behavioral state of the listeners, often promoting affiliation, pair bonding, or aggression. What is it in the acoustic structure of birdsong that makes it such a potent stimulus? We suggest that birdsong potency might be driven by principles similar to those that make music so effective in inducing emotional responses in humans: a combination of rhythms and pitches -and the transitions between acoustic states-affecting emotions through creating expectations, anticipations, tension, tension release, or surprise. Here we propose a framework for investigating how birdsong, like human music, employs the above “musical” features to affect the emotions of avian listeners. First we analyze songs of thrush nightingales (Luscinia luscinia) by examining their trajectories in terms of transitions in rhythm and pitch. These transitions show gradual escalations and graceful modifications, which are comparable to some aspects of human musicality. We then explore the feasibility of stripping such putative musical features from the songs and testing how this might affect patterns of auditory responses, focusing on fMRI data in songbirds that demonstrate the feasibility of such approaches. Finally, we explore ideas for investigating whether musical features of birdsong activate avian brains and affect avian behavior in manners comparable to music’s effects on humans. In conclusion, we suggest that birdsong research would benefit from current advances in music theory by attempting to identify structures that are designed to elicit listeners' emotions and then testing for such effects experimentally. Birdsong research that takes into account the striking complexity of song structure in light of its more immediate function - to affect behavioral state in listeners - could provide a useful animal model for studying basic principles of music neuroscience in a system that is very accessible for investigation, and where developmental auditory and social experience can be tightly controlled.

Concepts: Psychology, Human, Emotion, Music, Song, Nightingale, Luscinia, Thrush Nightingale


Bird song plays an important role in the establishment and maintenance of prezygotic reproductive barriers. When two closely related species come into secondary contact, song convergence caused by acquisition of heterospecific songs into the birds' repertoires is often observed. The proximate mechanisms responsible for such mixed singing, and its effect on the speciation process, are poorly understood. We used a combination of genetic and bioacoustic analyses to test whether mixed singing observed in the secondary contact zone of two passerine birds, the Thrush Nightingale (Luscinia luscinia) and the Common Nightingale (L. megarhynchos), is caused by introgressive hybridization. We analysed song recordings of both species from allopatric and sympatric populations together with genotype data from one mitochondrial and seven nuclear loci. Semi-automated comparisons of our recordings with an extensive catalogue of Common Nightingale song types confirmed that most of the analysed sympatric Thrush Nightingale males were ‘mixed singers’ that use heterospecific song types in their repertoires. None of these ‘mixed singers’ possessed any alleles introgressed from the Common Nightingale, suggesting that they were not backcross hybrids. We also analysed songs of five individuals with intermediate phenotype, which were identified as F1 hybrids between the Thrush Nightingale female and the Common Nightingale male by genetic analysis. Songs of three of these hybrids corresponded to the paternal species (Common Nightingale) but the remaining two sung a mixed song. Our results suggest that although hybridization might increase the tendency for learning songs from both parental species, interspecific cultural transmission is the major proximate mechanism explaining the occurrence of mixed singers among the sympatric Thrush Nightingales. We also provide evidence that mixed singing does not substantially increase the rate of interspecific hybridization and discuss the possible adaptive value of this phenomenon in nightingales.

Concepts: Gene, Evolution, Bird, Speciation, Nightingale, Thrush, Luscinia, Thrush Nightingale