Eyespot patterns have evolved in many prey species. These patterns were traditionally explained by the eye mimicry hypothesis, which proposes that eyespots resembling vertebrate eyes function as predator avoidance. However, it is possible that eyespots are not the mimicry of eyes: according to the conspicuousness hypothesis, eyespots are just one form of vivid aposematic signals where only conspicuousness matters. To test these hypotheses and explore factors influencing predators’ responses, we conducted a meta-analysis with 33 empirical papers focusing on bird responses to lepidopterans having conspicuous patterns (eyespots and non-eyespots). Supporting the latter hypothesis, the results showed no clear difference in predator avoidance efficacy between eyespots and non-eyespots. When comparing geometric pattern characteristics, bigger pattern sizes and smaller numbers of patterns were more effective in preventing avian predation. This finding indicates that paired concentric patterns have weaker deterring effects than single ones. Taken together, our study supports the conspicuousness hypothesis more than the eye mimicry hypothesis. Due to the number and species coverage of published studies so far, the generalisability of our conclusion may be limited. The findings highlight that pattern conspicuousness is key to eliciting avian avoidance responses, shedding a different light on this classic example of signal evolution.
PLOS ONE 18 (10) e0292074 - e0292074 2023/10/25 [Refereed]
Conspicuous facial features, such as blushing in primates, can communicate social/emotional/physiological states in animals. However, the role of bare facial features is less well studied in birds than in humans or primates. We investigate the Java sparrow, which is characterised by conspicuous rings of swollen and blushed bare skin around the eye. Eye rings show no clear sex difference, although the swelling is associated with breeding. Java sparrows are socially monogamous, with mutual courtships and long-term pair-bonding. Therefore, it is plausible that eye rings function in within-pair communication. Specifically, do eye rings reflect psychophysiological conditions after pair formation? We assessed variations in ring thickness in pair-bonded birds and compared them with single birds and pairs of non-bonded individuals. Over the 12-week experimental period, pair-bonded males and females had an increased ring thickness, unlike the controls. We suggest eye rings convey breeding motivations or serve as fertility signals. This would be of great importance for ensuring reproductive synchrony in tropical birds like the Java sparrow. Our results contribute to understanding the evolution of facial ornamentation in birds, which was often overlooked in the past studies.
Royal Society Open Science 10 (10) 2023/10/18 [Refereed]
The diverse characteristics of animal signal designs can be explained by the sensory bias hypothesis, which suggests that natural selection shapes sensory bias and preferences associated with signal design. Traditionally, this hypothesis has focused on female sensory biases and male sexual traits. However, considering shared sensory systems between males and females in non-sexual contexts, existing sensory bias possibly contributes to the evolution of shared social and sexual traits. Our previous studies on the family Estrildidae supported this idea. An evolutionary relationship probably existed between diet and white dot plumage, and a species of estrildid finches showed a visual preference for white dot patterns. To investigate this further, we examined hunger-related visual preferences using phylogenetic comparative methods and behavioural experiments. Specifically, we compared the gazing responses of 12 species of estrildids to monochromatic printed white dot and stripe patterns, considering their phylogenetic relationships. The results support our idea that the common estrildid ancestor had a hunger-related visual preference for white dot patterns. Subject species generally preferred white dots to stripes. Furthermore, males and females showed a similar preference towards dots. Our findings provide insights into the role of sensory bias in the evolution of mutual ornamentation.
Abstract
Many animals have polka dot patterns on their body surface, some of which are known to have signalling functions; however, their evolutionary origins remain unclear. Dot patterns can trigger a fear response (trypophobia) in humans and are known to function as aposematic signals in non-human animals, suggesting that dots may deserve attention for biological reasons. Interestingly in many birds, plumage dot patterns serve for social/sexual signalling. To understand their evolution, we have focused on the sensory bias hypothesis, which predicts the role of pre-existing sensory preference driven by natural selection in shaping signal design. Our previous phylogenetic comparative study supported the hypothesis and showed that diet-driven visual preference promoted the evolution of plumage patterns, as there was an evolutionary correlation between termite-eating (white roundish gregarious prey) and the presence of plumage dot patterns in species of the family Estrildidae. This suggests that these species possess an intrinsic preference for dots. To test this, we compared the responses of an Estrildid species with dot plumage pattern (star finch Neochmia ruficauda) towards simultaneously presented monochrome-printed white dot vs white stripe patterns under both food-deprived and -supplied conditions. Overall, star finches preferred dots to stripes. They showed foraging-like behaviours almost only toward dots when hungry and gazed at dots frequently even when food was available, suggesting both hunger-related and hunger-neutral dot preferences. These results are rather surprising, given how strongly the subjects were attracted to abstract dot patterns without organic structure, but provided good support for the sensory bias hypothesis.
The division of cognitive processing between the two hemispheres of the brain causes lateralized eye use in various behavioral contexts. Generally, visual lateralization is shared among vertebrates to a greater extent, with little interspecific variation. However, previous studies on the visual lateralization in mating birds have shown surprising heterogeneity. Therefore, this systematic review paper summarized and analyzed them using phylogenetic comparative methods. The review aimed to elucidate why some species used their left eye and others their right to fixate on individuals of the opposite sex, such as mating partners or prospective mates. It was found that passerine and non-passerine species showed opposite eye use for mating, which could have stemmed from the difference in altricial vs. precocial development. However, due to the limited availability of species data, it was impossible to determine whether the passerine group or altricial development was the primary factor. Additionally, unclear visual lateralization was found when studies looked at lek mating species and males who performed courtship. These findings are discussed from both evolutionary and behavioral perspectives. Possible directions for future research have been suggested.
Koen de Reus, Masayo Soma, Marianna Anichini, Marco Gamba, Marianne de Heer Kloots, Miriam Lense, Julia Hyland Bruno, Laurel Trainor, Andrea Ravignani
Philosophical Transactions of the Royal Society B: Biological Sciences 376 (1835) 20200337 - 20200337 0962-8436 2021/08/23 [Refereed]
This review paper discusses
rhythmic
interactions and distinguishes them from
non-rhythmic
interactions. We report on communicative behaviours in social and sexual contexts, as found in dyads of humans, non-human primates, non-primate mammals, birds, anurans and insects. We discuss observed instances of rhythm in dyadic interactions, identify knowledge gaps and propose suggestions for future research. We find that most studies on rhythmicity in interactive signals mainly focus on one modality (acoustic or visual) and we suggest more work should be performed on multimodal signals. Although the social functions of interactive rhythms have been fairly well described, developmental research on rhythms used to regulate social interactions is still lacking. Future work should also focus on identifying the exact timing mechanisms involved. Rhythmic signalling behaviours are widespread and critical in regulating social interactions across taxa, but many questions remain unexplored. A multidisciplinary, comparative cross-species approach may help provide answers.
This article is part of the theme issue ‘Synchrony and rhythm interaction: from the brain to behavioural ecology’.
Rebecca N. Lewis, Masayo Soma, Selvino R. de Kort, R. Tucker Gilman
Frontiers in Psychology 12 654198 - 654198 2021/06/04 [Refereed][Invited]
Social learning of vocalizations is integral to song inheritance in oscine passerines. However, other factors, such as genetic inheritance and the developmental environment, can also influence song phenotype. The relative contributions of these factors can have a strong influence on song evolution and may affect important evolutionary processes such as speciation. However, relative contributions are well-described only for a few species and are likely to vary with taxonomy. Using archived song data, we examined patterns of song inheritance in a domestic population of Java sparrows (Lonchura oryzivora), some of which had been cross-fostered. Six-hundred and seventy-six songs from 73 birds were segmented and classified into notes and note subtypes (N = 22,972), for which a range of acoustic features were measured. Overall, we found strong evidence for cultural inheritance of song structure and of the acoustic characteristics of notes; sons’ song syntax and note composition were similar to that of their social fathers and were not influenced by genetic relatedness. For vocal consistency of note subtypes, a measure of vocal performance, there was no apparent evidence of social or genetic inheritance, but both age and developmental environment influenced consistency. These findings suggest that high learning fidelity of song material, i.e., song structure and note characteristics, could allow novel variants to be preserved and accumulate over generations, with implications for evolution and conservation. However, differences in vocal performance do not show strong links to cultural inheritance, instead potentially serving as condition dependent signals.
The duets of birds have intrigued biologists for a long time, yet much remains unknown about the evolution of these striking collective displays. This is partly because previous studies on duet evolution have been biased to songbirds and neglected other bird groups. In songbirds, the absence of migration has been found to predict the occurrence of duetting, indirectlysupporting the idea that duet communication is linked with pair bonding. Here, we used phylogenetic comparative analyses in a sedentary clade of non-songbirds, the barbets (Capitonidae), to reveal new correlates of duet evolution. We found (i) that duets evolved several times independently in different barbet lineages and (ii) that duetting evolved in association with group living (i.e. the presence of helpers or non-breeding adults during the breeding period), but not with sexual monochromatism or habitat type. Our findings are consistent with a duet function in mate guarding and dominance against subordinate group members as well as joint territory defence. Altogether, the results highlight the importance of the social environment for the evolution of collective signalling.
Color patterns, such as bars or dots that cover the body surface of animals are generally thought to play roles in signaling and camouflage. In birds, however, the macroscopic aspects of plumage coloration are less well understood, as past studies typically described plumage colorations by using spectrophotometric analyses. To provide insight into the evolution of plumage patterns as sexual signals, we characterized interspecific and intersexual variations in the plumage patterns of estrildid finches and tested their associations with other courtship signals and life-history traits using a comparative phylogenetic approach. Our results support the idea that plumage patterns in estrildids are favored by sexual selection because large-sized conspicuous plumage patterns are possessed by species with an elaborate courtship dance. These plumage patterns may also play roles in social signaling because patterns are more conspicuous in species with intraspecific brood parasitism. We predict that pattern traits can be favored by mate choice or intrasexual competition when they can serve as honest indicators of individual condition. As our results are consistent between the sexes, we suggest that the same selective force is acting on the evolution of plumage patterns in males and females in parallel. Finally, we also found a trade-off between large size and vivid color patterns, suggesting that too conspicuous patterns are costly, presumably because of the risk of catching the eyes of potential predators. Therefore, plumage patterns are also shaped by natural selection.
Sexual traits (e.g. visual ornaments, acoustic signals, courtship behaviour) are often displayed together as multimodal signals. Some hypotheses predict joint evolution of different sexual signals (e.g. to increase the efficiency of communication) or that different signals trade off with each other (e.g. due to limited resources). Alternatively, multiple signals may evolve independently for different functions, or to communicate different information (multiple message hypothesis). We evaluated these hypotheses with a comparative study in the family Estrildidae, one of the largest songbird radiations, and one that includes many model species for research in sexual selection and communication. We found little evidence for either joint evolution or trade-offs between song and colour ornamentation. Some negative correlations between dance repertoire and song traits may suggest a functional compromise, but generally courtship dance also evolved independently from other signals. Instead of correlated evolution, we found that song, dance and colour are each related to different socio-ecological traits. Song complexity evolved together with ecological generalism, song performance with investment in reproduction, dance with commonness and habitat type, whereas colour ornamentation was shown previously to correlate mostly with gregariousness. We conclude that multimodal signals evolve in response to various socio-ecological traits, suggesting the accumulation of distinct signalling functions.
PLOS ONE 12 (3) e0172655 1932-6203 2017/03 [Refereed][Not invited]
Mutual interactions between sexes have multiple signalling functions. Duet singing in songbirds is related to mutual mate guarding, joint resource defence, and signalling commitment. Coordinated visual displays of mating pairs are thought to perform similar functions, but are less well understood. The current study evaluated mutual interactions in an Estrildid species to explore the relative importance of duet dancing and male singing in mating success of pairs in a first encounter. When Java sparrows (Lonchura oryzivora) court prospective mates, only males sing. However, both males and females perform courtship dances, often in a duet-like manner. These dances are typically terminated by female copulation solicitation displays (CSDs). In the current study, we observed higher mating success when courtship dances were mutually exchanged, and when males sang. However, the sex initiating the courtship did not affect mating success. Most females produced CSDs after duet dancing but before hearing the entire song, indicating that duet dancing played a crucial role in mating. This finding highlights an unexplored aspect of duetting behaviour in the process of mutual mate choice. These results conflict with the majority of past songbird research, which has interpreted songs as primary behavioural sexual signals.
BIOACOUSTICS-THE INTERNATIONAL JOURNAL OF ANIMAL SOUND AND ITS RECORDING 26 (2) 161 - 168 0952-4622 2017 [Refereed][Not invited]
Vocalizations have been elucidated in previous songbird studies, whereas less attention has been paid to non-vocal sounds.In the bluecapped cordon-bleu (Uraeginthus cyanocephalus), both sexes perform courtship displays that are accompanied by singing and distinct body movements (i.e.dance).Our previous study revealed that their courtship bobbing includes multiple rapid steps.This behaviour is quite similar to human tap dancing, because it can function as both visual and acoustic signals.To examine the acoustic signal value of such steps, we tested if their high-speed step movements produce non-vocal sounds that have amplitudes similar to vocal sounds. We found that step behaviour affected step sound amplitude. Additionally, the dancing step sounds were substantially louder than feet movement sounds in a non-courtship context, and the amplitude range overlapped with that of song notes.These results support the idea that in addition to song cordon-bleus produce acoustic signals with their feet.
Song is a notable sexual signal of birds, and serves as an honest indicator of male quality. Condition dependence of birdsong has been well examined from the viewpoint of the developmental stress hypothesis, which posits that complex songs assure fitness because learned acoustic features of songs are especially susceptible to early-life stress that young birds experience in song learning periods. The effect of early stress on song phenotypes should be crucial, especially in age-limited song learners which sing stereotyped songs throughout life. However, little attention has been paid to non-learned song features that can change plastically even in adulthood of age-limited song-learners. Although it has been shown that food availability affects song rate in wild songbirds, there is limited evidence of the link between favorable nutritional conditions and song phenotypes other than song rate. Under the prediction that singing behavior reflects an individual's recent life history, we kept adult Bengalese finch males under high-nutrition or normal diet for a short term, and examined changes in body mass and songs. We found that birds on a high-nutrition diet showed higher song output (e.g. song rate and length) compared with those of the control group, while changes in body mass were moderate. In addition, note repertoire became more consistent and temporal structures got faster in both nutrition and control groups, which indicates that songs were subject to other factors than nutrition. Considering that female estrildid finches, including Bengalese and zebra finches, show a preference toward complex songs as well as longer songs and higher song rate, it is plausible that different aspects of singing behavior signal different male qualities, and provide multifaceted clues to females that choose mates.
According to classical sexual selection theory, complex multimodal courtship displays have evolved in males through female choice. While it is well-known that socially monogamous songbird males sing to attract females, we report here the first example of a multimodal dance display that is not a uniquely male trait in these birds. In the blue-capped cordon-bleu (Uraeginthus cyanocephalus), a socially monogamous songbird, both sexes perform courtship displays that are characterised by singing and simultaneous visual displays. By recording these displays with a high-speed video camera, we discovered that in addition to bobbing, their visual courtship display includes quite rapid step-dancing, which is assumed to produce vibrations and/or presumably non-vocal sounds. Dance performances did not differ between sexes but varied among individuals. Both male and female cordon-bleus intensified their dance performances when their mate was on the same perch. The multimodal (acoustic, visual, tactile) and multicomponent (vocal and non-vocal sounds) courtship display observed was a combination of several motor behaviours (singing, bobbing, stepping). The fact that both sexes of this socially monogamous songbird perform such a complex courtship display is a novel finding and suggests that the evolution of multimodal courtship display as an intersexual communication should be considered.
PLOS ONE 10 (5) 1932-6203 2015/05 [Refereed][Not invited]
Music and dance are two remarkable human characteristics that are closely related. Communication through integrated vocal and motional signals is also common in the courtship displays of birds. The contribution of songbird studies to our understanding of vocal learning has already shed some light on the cognitive underpinnings of musical ability. Moreover, recent pioneering research has begun to show how animals can synchronize their behaviors with external stimuli, like metronome beats. However, few studies have applied such perspectives to unraveling how animals can integrate multimodal communicative signals that have natural functions. Additionally, studies have rarely asked how well these behaviors are learned. With this in mind, here we cast a spotlight on an unusual animal behavior: nonvocal sound production associated with singing in the Java sparrow (Lonchura oryzivora), a songbird. We show that male Java sparrows coordinate their bill-click sounds with the syntax of their song-note sequences, similar to percussionists. Analysis showed that they produced clicks frequently toward the beginning of songs and before/after specific song notes. We also show that bill-clicking patterns are similar between social fathers and their sons, suggesting that these behaviors might be learned from models or linked to learning-based vocalizations. Individuals untutored by conspecifics also exhibited stereotypical bill-clicking patterns in relation to song-note sequence, indicating that while the production of bill clicking itself is intrinsic, its syncopation appears to develop with songs. This paints an intriguing picture in which non-vocal sounds are integrated with vocal courtship signals in a songbird, a model that we expect will contribute to the further understanding of multimodal communication.
Birdsong is a sexual signal that serves as an indicator of male quality. There is already abundant evidence that song elaboration reflects early life-history because early developmental stress affects neural development of song control systems, and leaves irreversible adverse effects on song phenotypes. Especially in closed-ended vocal learners, song features crystallized early in life are less subject to changes in adulthood. This is why less attention has been paid to lifelong song changes in closed-ended learners. However, in the eyes of female birds that gain benefits from choosing mates based on male songs, not only past but also current conditions encoded in songs would be meaningful, given that even crystallized songs in closed-ended learners would not be identical in the long term. In this study, we examine within-individual song changes in the Java sparrow Lonchura oryzivora, with the aim of shedding light on the relationship between song and long-term life history. Specifically, we compared song length, tempo, and song complexity measures between the point just after song crystallization and around 1 yr later, and also compared those traits between fathers and sons to clarify the effect of vocal learning. While it is not surprising that song complexity did not differ depending on age or between fathers and sons, we found that song length and tempo increased with age. Follow-up analyses have revealed that frequency bandwidth and peak frequency of song notes also elevated with age. Our results show that song performance related to motor skills can be improved even after song crystallization. We also suggest that song performance in closed-ended vocal learners gives a reliable clue for mate choice by reflecting male quality with aging.
Bird songs have evolved under sexual selection pressure. Songs include multiple features that are subject to female preference, but recent comparative research has indicated evolutionary tradeoffs between song performance and complexity in some species. Trill, a repetition of the same sound, is a performance-related song trait; higher trill performance can be achieved at the cost of song complexity at the among-species or population level. The aim of this study was to examine whether such tradeoffs also account for within-species variation in Java sparrow songs, which include both multiple trill types and non-trill parts. We found a great individual variation in trill proportion, trill performance, and song complexity. A positive association between trill performance and body size suggested that trills can serve as an indicator of male quality. However, contrary to the tradeoffs predicted by previous studies based on other passerine species, trill performance and song complexity, i.e., note repertoire, were positively correlated: males in better condition can sing songs with larger note repertoires and higher trill performance, which may explain how trills and non-trill notes are both maintained and have co-evolved by sexual selection in Java sparrow songs. (C) 2013 Elsevier B.V. All rights reserved.
Postmating sexual selection plays an important part in the evolution of secondary sexual characters. Based on differential allocation theory that predicts biased reproductive investment of females depending on the attractiveness of mates, a number of previous studies have shown that egg production is related in various ways to ornamental sexual traits of males, but evidence for behavioural sexual traits is less abundant. In this study we examine such maternal effects in relation to birdsong. Because the Bengalese finch is a monomorphic songbird, courtship song serves a key role in mate choice. To take into account individual female differences in egg production performance, we sequentially paired naive, captive, female Bengalese finches to two different males, and investigated if their reproductive investment (clutch size, egg mass and hatchling sex ratio) was related to the song traits of their mates. We found that clutch size and egg mass were highly repeatable within individual females while sex ratio was not. Despite the inflexibility of egg mass within each female, egg mass increased when females were mated to males with longer songs. In addition, we found a non-significant weak tendency toward male-biased sex ratio in relation to longer song duration of mates. Our findings suggest that females mated to better mates adjusted their reproductive investment by producing heavier eggs and possibly offspring of the more costly sex.
Birdsong is an important sexually selected trait, and its acoustic features are socially transmitted in the process of song learning. Though a great deal of research has been conducted to shed light on the mechanisms and functions of song learning, we still do not have clear answers as to why for some species, including Estrildid finches and humans, the ability of "social" vocal learning has evolved. Therefore, this paper addresses social factors responsible for song learning, especially focusing on tutor choice in Estrildid finches, with the aim of elucidating what is already clear about how the social environment shapes songs and what is needed in future studies.
Estrildine finches are important model species in experimental studies on female mating preferences, but research has focused on only two species from this family, namely, the Zebra Finch Taeniopygia guttata and the Bengalese Finch Lonchura striata var. domestica, and we know comparatively little about other closely related species. Therefore, we investigated sexual dimorphism and female choice in the Java Sparrow Padda oryzivora, which also belongs to the family Estrildidae, to open up the potential for comparative research and to better understand sexual selection in this family. First, we took measurements of eight morphological traits of six male and six female Java Sparrows: natural wing length, maximum wing length, tarsus length, tail length, exposed culmen length, bill width, bill depth and body mass. We quantitatively confirmed that Java Sparrows show sexual dimorphism in bill depth, with males having deeper bills than females. Second, we examined individual variation in male courtship songs by analyzing their acoustic and syntactical structure. After collecting data on male morphological and song-related traits, we conducted two-way choice tests to ask which kinds of male traits predicted which males were preferred by female Java Sparrows. In the choice tests, we put a cage with a female in between two cages each containing one male and recorded the position of the female every 30 seconds by point sampling. We performed a stepwise regression analysis to assess the relationship between the time females spent in front of each male and male morphological and song-related traits. The results indicated that females base their preference upon large body size, which is likely to act as a good indicator of male quality. However, no preference was observed for song-related traits or sexually dimorphic bill depth. Perhaps the sexual dimorphism in bill size is instead the evolutionary outcome of male-male competition.
The theory of sexual selection predicts a relationship between male sexual traits and reproductive success. This prediction has been tested extensively using the complexity of birdsong as a model for trait elaboration. However, contradictory results have emerged. Some studies have demonstrated that males with large repertoires enjoy a reproductive advantage, whereas other studies have failed to support this prediction. To make general inferences from this mixed evidence, we quantitatively reviewed the relevant literature using a meta-analytic approach. The mean effect size for the song/mating success association was significant, but the effects were generally weak, affected by publication bias, confounded by uncontrolled variables, and differing across the traits examined. Effect sizes were heterogeneous across studies due to species-specific effects, differences in mating systems, and song phenotypes. The degree of association between song complexity and reproductive success was independent of the strength of sexual selection, as assessed by the degree of polygyny and extrapair paternity. Our results highlight the importance of considering various biological factors to understand the role of repertoires in mediating mating success in different species.
Birdsong is an important sexually selected trait, and its acoustic features are socially transmitted in the process of song learning. The preference of some female songbirds for song complexity as measured by repertoire size has driven the evolution of many song traits and song-learning programs in males. If juvenile birds adopt the attractiveness of model birds by copying their repertoires, tutor choice in the early song-learning period should be crucial for future reproductive success. Thus, we hypothesized that the song quality of tutors influences the tutor choice of chicks in the Bengalese finch, Lonchura striata var. domestica. Each subject chick was reared by a foster pair in an individual breeding cage. At around the fledging period, we introduced another unrelated adult male (subtutor) into each cage to simulate the natural social environment in which chicks have opportunities to hear the songs of nonfather males after fledging. Subject chicks learned from both tutors when the total note repertoire size of the two models was small; they tended to learn from the subtutor when the father had a smaller note repertoire size. The acquired note repertoire size was not affected by whether subjects learned songs from the foster father or not. Song learning from the subtutor contributed to a larger note repertoire. Observed patterns in tutor choice, in particular song learning from a subtutor, could be an adaptive strategy that helps chicks compensate for exposure to smaller repertoire size in models. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
Birdsong differs from other sexual traits in that the acquisition process involves learning. Especially in close-ended learning species like the Bengalese finch, conditions experienced during the critical song-learning period can have a profound influence on song quality. Therefore, to understand song evolution from a life-history perspective, we investigated early ontogenetic effects on song quality. In particular, we focused on maternal effects and sibling competition. In asynchronously hatching bird species, the age hierarchy among nestlings affects physical development due to competition for food; mothers may influence this competition by adjusting their investment in each egg according to its sequence in the laying order. To independently assess these effects, chicks of the Bengalese finch were cross-fostered so that the age hierarchies formed in fostered broods were independent of the laying order. Our results indicate that song quality partially reflects early ontogenetic conditions, whereas song duration and note-type repertoire were independent of either laying order or age hierarchy. The syntactical complexity of note order declined over the laying sequence. This finding suggests that the song learning ability is influenced by within-clutch variation in maternal investment toward eggs. Considering that song syntactical complexity is subject to female preference in the Bengalese finch, it is likely that maternal resource allocation strategies play a role in song evolution.
Recent studies of the vocal mechanisms of songbirds have shown that there are physical effects on sound production. Interspecific and phylogenetic comparisons have shown that body mass and bill morphology are two major factors affecting vocalizations such as songs. We analyzed the distance calls of female Bengalese finches Lonchurastriata var. domestica, the non-vocal learning sex, to assess the potential physical effects on the acoustic structure of vocalizations. By experimentally controlling rearing condition using cross-fostering we could examine the effects of the developmental environment and genetic background. None of the tested factors affected the peak frequency of the distance calls, but we found that larger-billed birds tended to produce shorter bout calls with higher trill rates. These results suggest that the divergence of bill morphology can affect acoustic features at the within-population level. We also found that the birds reared in the same foster brood and siblings from the same genetic parents tended to produce calls with similar trill rates. This implies that the trill rate is under the influence of developmental and genetic factors.
Maternal effects, such as investment in eggs, have profound effects on offspring fitness. Mothers are expected to skew their investment depending on the laying order and sex when unequal sibling competition occurs within a brood because of sex-specific vulnerability and age hierarchy caused by asynchronous hatching. The Bengalese finch hatches asynchronously and shows a moderate reversed sexual size dimorphism. However, contrary to commonly accepted assumptions of size-dependent vulnerability, the smaller sex (male) is more vulnerable to developmental stress caused by sibling competition. We investigated whether maternal investment would be biased by the position in laying order and the sex of eggs, and also explored the possible differences in growth patterns depending on sex, laying order, and age hierarchy by observing chicks fostered to experimentally manipulated broods where brood composition was controlled and age hierarchy was more enhanced than in natural breeding conditions. We found that overall patterns of maternal investment favored the disadvantageous sectors of sibling competition, i.e., eggs of later laying order and sons over those of early laying order and daughters. We also examined the effect of laying order on adult body size and sex differences in growth patterns. When reared in the subordinate age hierarchy, females could compensate for the deficit of decreased growth rate by taking longer to mature, whereas males could not. We suggest that this sex-specific growth pattern could be the cause of sex differences in vulnerability to early developmental stress.
Birdsong is an acoustic ornament. According to indicator models, a trait must be costly to act as an honest signal, but the potential costs of elaborate songs are still poorly understood. The developmental stress hypothesis Suggests that learned song characteristics Could be an honest indicator of early developmental conditions because the brain structures associated with learning songs are susceptible to early developmental stress, which could thus affect song development. Unlike previous studies of developmental stress that examined the effect of a stress hormone or restricted nutrition, we observed Bengalese finches under semi-natural breeding conditions in captivity to investigate the relationship between early rearing conditions (e.g., brood size and sex ratio) and the subsequent variation in body size and song among individuals. Our results suggest that the early rearing environment directly affects body size and song complexity, whereas song output is determined mainly by' body size. These results support the developmental stress hypothesis. Moreover, Our findings are the first to show that developmental condition affects not only the number of note types but also the syntactical complexity of the song.
Social foraging is a far more complex phenomenon than individual foraging because of the many social interactions and communications that affect individual behaviour. Social dominance and social learning markedly influence the foraging efficiency of individuals in an unfamiliar environment. This study investigated how these two factors affect the costs and benefits of social foraging. We presented a novel feeding environment to captive budgerigars, Melopsittacus undulatus, and then compared the latency to feeding and number of pecks at the food for social and individual conditions, and for high-, middle-, and low-ranking birds. When in the social condition, birds started foraging faster and pecked more food than in the individual condition. Presumably, this lowered neophobia and enhanced foraging efficiency in the social condition was caused by social facilitation. Low-ranking birds had less resource accessibility in the social condition, probably because they were constrained by the existence of higher-ranking birds when it came to accessing the feeder. Nevertheless, the food intake of low-ranking birds almost equaled that of high- or middle-ranking birds in the social condition. In summary, high status is definitely an advantage, while low status adds some costs to individuals. Nevertheless, low-ranking birds compensate for this through enhanced foraging. It was clear that social foraging provides a great advantage to foragers of each rank, because of social facilitation.
Recruitment calls are reported as one mechanism used by some birds and mammals for attracting conspecifics once an individual has found a food source, hence leading to the formation of foraging flocks/groups. Jungle Crows Corvus macrorhynchos sometimes give "kakaka" calls while foraging in flocks. We conducted field observations and playback experiments to examine whether these calls serve as recruitment calls leading to flock formation. Crows gave "kakaka" calls while foraging, and the size of foraging flocks was positively correlated with the number of calls given. Broadcasting "kakaka" calls attracted more Jungle Crows. There was also a positive correlation between pecking rates and the size of foraging flocks. The results indicate that "kakaka" calls are effective recruitment calls and that Jungle Crows forage more efficiently in flocks.
Japan Society for the Promotion of Science:Grants-in-Aid for Scientific Research
Date (from‐to) : 2016/04 -2020/03
Author : Soma Masayo
Estrildid finches are known for great interspecific diversity in the degree of elaboration in courtship dance, song and plumage coloration and also for the considerable sex differences in these traits within species. The present series of studies examined the function and evolution of such diverse sexual signals. Ethological studies revealed that song and dance have different functions even when they are expressed simultaneously. Song serves as a condition dependent signal giving a clue for mate choice, while dance plays an important role in pair bonding. Phylogenetic comparative studies also showed that song and dance evolved independently. However, plumage pattern conspicuousness and courtship dance complexity are correlated among species, meaning that static and dynamic aspects of visual sexual signals are interlinked with each other.
Japan Society for the Promotion of Science:Grants-in-Aid for Scientific Research Grant-in-Aid for Young Scientists (A)
Date (from‐to) : 2011/04 -2015/03
Author : SOMA Masayo
Estrildid finches have multiple sexual traits, including courtship display that is composed of song and dance and ornamental plumage colors. This study shed light on song and dance, and asked why multiple sexual signals evolved and how they function. Estrildid finches showed a capacity to coordinate multimodal signals, such as singing, dancing, and bill sounds. In addition, young finches of a Estrildid species show practicing behavior of dance very early in their life, which might contribute to master fine coordination between singing and dancing. As dances are often displayed in parallel with songs, they are likely to evolve non-independently of songs. However, by using phylogenetic comparative approaches I found that dance and song evolved independently. The mechanisms and functions of the two behaviors are at least partially independent.