Different singing styles in mated and unmated Reed Buntings Emberiza schoeniclus

Ewin described two singing styles, "rapid" and "slow", in the Reed Bunting Emberiza schoeniclus. My research has demonstrated that the prevalence of song styles is related to male mating status. Songs of mated and unmated males differed in the interval structure, number of syllables per song, number of frequency modulations per syllable and number of introductory syllables. Since both singing styles are uttered at the same intensity, it is not likely that they are parts of a continuous range and it is justified to treat them as two different song categories. After losing their mate, males sang like unpaired males and therefore a learning effect can be excluded. Both singing styles were directed to females, the slow one probably only to the mate. The use of these singing styles as a means of territory or mate defence cannot be excluded, but, if so used, it would seem to be of minor significance.     

Part-Time Mate Guarding Affects Paternity in Male Reed Buntings (Emberiza schoeniclus)

Male mate guarding by close following has been reported in many socially monogamous bird species and is generally believed to function as a paternity guard. Many aspects of the dynamics and effectiveness of this behavior are still however poorly understood. Here, we describe the temporal variation in mate guarding behavior in male reed buntings (Emberiza schoeniclus) with a particular focus on how males allocate their mating effort between mate guarding and extrapair mating in a context of intense sperm competition. In our highly synchronous study population most males have to balance the simultaneous and mutually exclusive demands of mate guarding and seeking extrapair copulations (EPCs). We found that males frequently switched between guarding their mates and performing intrusions to neighboring territories. Both activities seemed to have significant fitness payoffs, as male mate guarding effort had a positive effect on paternity, and a large fraction of extrapair fertilizations occurred during the days when the sire guarded its own female. The reed bunting is thus an example of how discontinuous or part‐time mate guarding can still be effective in securing paternity. Female reed buntings were not particularly active in initiating EPCs as they never were observed performing extraterritorial forays. We argue that the absence of female‐initiated EPCs is a prerequisite for males to trade mate guarding against seeking EPCs. Otherwise, if females circumvent male mate guarding by timing their EPCs to periods of male absence, males should guard their mates almost continuously or rely on alternative paternity guards.     

The phylogenetic affinities of the Socotra Bunting Emberiza socotrana

The monophyly of the African ‘brown bunting’ complex was corroborated by a recent molecular study. However, the little-known Socotran endemic Emberiza socotrana, which is morphologically similar to the other taxa in this complex, was not included. Here we present a hypothesis of the phylogenetic relationships of the Socotra Bunting based on one mitochondrial gene and one nuclear intron. We found the Socotra Bunting to be deeply nested within the African ‘brown bunting’ complex and, although morphologically most similar to E. capensis, it proved to be more closely related to the E. striolata/sahari and E. tahapisi/goslingi species groups. The phylogenetic uniqueness of the Socotra Bunting underpins once more the evolutionary importance of the Endemic Bird Area of Socotra, which is often considered the ‘Galápagos of the Indian Ocean’.     

Variability of response of the corn bunting,Emberiza calandra,to songs of different dialects

Experiments on the corn bunting were carried out in two localities of different dialects. In the center of the territory of each bird, we played two natural songs: either one local song and one song of the foreign dialect or two local songs coming from a neighbor and from a non-neighboring individual. The following results were obtained: (1) The behavioral responses are always more intense to the local song than to the song of a foreign dialect. Thus, the behavior of the corn bunting is similar to that of species with dialects studied up to now. (2) Eighty percent of the birds tested with the song of the foreign dialect react weakly or not at all. This peculiarity, which is specific to the corn bunting, suggests limited possibilities of generalization in this species. (3) The behavioral responses to the songs of a neighbor and of a non-neighboring individual of the same locality, are aggressive and of the same intensity. This means that the song of a neighbor, emitted from a different place than usual, is perceived by the bird as a threat to its territory.     

Weather conditions affect levels of extra-pair paternity in the reed bunting Emberiza schoeniclus

Extra-pair paternity (EPP) is common in many socially monogamous birds, but large variations in frequency of EPP are found both between and within species. Local ecological factors can affect the costs and benefits of extra-pair mating behaviour, and may therefore influence the chance that individuals engage in extra-pair copulations (EPCs). We investigated the effect of weather conditions during the peak fertile period of the female on the levels of EPP in reed buntings Emberiza schoeniclus . The reed bunting is a socially monogamous passerine, with extremely high levels of EPP (50% of offspring in 80% of broods). We found that higher daily minimum temperatures and more rainfall during the peak fertile period were associated with lower proportions of EPP. As during adverse weather conditions individuals have to invest more in self maintenance, we suggest that during long periods of rain the extra-pair mating behaviour of all individuals will be restricted, leading to lower proportions of EPP. During cold mornings, time-consuming activities such as mate guarding are likely to be more strongly affected than less time-consuming activities such as EPCs, leading to higher proportions of EPP.     

Time Course of Sensitivity of the Photoinducible Phase to Light in the Redheaded Bunting, Emberiza bruniceps

This study investigated the differential sensitivity of the photoinducible phase (Φi) to light in the redheaded bunting (Emberiza bruniceps). Using a skeleton paradigm, we assessed the rate and magnitude of testicular response as a function of the duration of an inducing light pulse and of the time of its introduction in Φi. For a period of 7 weeks, birds received at an intensity of ~100 lux, the same (6 h) entraining light stimulus with a varied inducing light pulse: 1, 2, 4 or 6 h beginning at zeitgeber time (zt) 11 (6L:5D:1L:12D, 6L:5D:2L:11D, 6L:5D:4L:9D or 6L:5D:6L:7D), or 1h pulses at zt 12 (6L:6D:1L:11D) or zt 16 (6L:10D:1L:7D). The testes were stimulated in all LD alternations, but duration- and time-dependent effects of the light pulse on the rate and magnitude of the testicular response were clearly evident. Illumination of a larger portion of Φi seems to result in higher rates of gonadal growth but there is a duration limit above which there will be no further increase of testicular response.     

Time Course of Sensitivity of the Photoinducible Phase to Light in the Redheaded Bunting,Emberiza bruniceps

This study investigated the differential sensitivity of the photoinducible phase (Φi) to light in the redheaded bunting (Emberiza bruniceps). Using a skeleton paradigm, we assessed the rate and magnitude of testicular response as a function of the duration of an inducing light pulse and of the time of its introduction in Φi. For a period of 7 weeks, birds received at an intensity of ~100 lux, the same (6 h) entraining light stimulus with a varied inducing light pulse: 1, 2, 4 or 6 h beginning at zeitgeber time (zt) 11 (6L:5D:1L:12D, 6L:5D:2L:11D, 6L:5D:4L:9D or 6L:5D:6L:7D), or 1h pulses at zt 12 (6L:6D:1L:11D) or zt 16 (6L:10D:1L:7D). The testes were stimulated in all LD alternations, but duration- and time-dependent effects of the light pulse on the rate and magnitude of the testicular response were clearly evident. Illumination of a larger portion of Φi seems to result in higher rates of gonadal growth but there is a duration limit above which there will be no further increase of testicular response.     

A territory scale analysis of habitat preferences of the declining Ortolan Bunting Emberiza hortulana

Capsule Ortolan Bunting occurrence is associated with bare ground, Lucerne, shrub cover and hedgerows/tree rows.

Aims To assess the habitat features selected by Ortolan Buntings at the territory level in semi-open landscapes, in the northern Apennines of Italy.

Methods We mapped territories in ten different plots and built a habitat selection model comparing 52 occupied cells with 52 unoccupied ones (cell size: 1?ha). We built multivariate adaptive regression splines models based on ground-measured variables.

Results The model revealed an association with intermediate Lucerne cover (50% of the cell), high shrub cover, bare ground (≥5%) and hedgerows/tree rows (≥25?m/ha). The most important driver of occurrence was bare ground (optimum at 5–20%).

Conclusion The maintenance of the mosaic and low-intensity farmed landscape, the promotion of Lucerne and the conservation/restoration of hedgerows/tree rows, may be promoted through the Rural Development Programme. The conservation of bare soil, grassland and shrubs at optimum amount at fine-scale could be the object of an agri-environment scheme targeted specifically at the Ortolan Bunting.     

Blood parasites, leucocytes and plumage brightness in the Cirl Bunting, Emberiza cirlus

1. Although a female preference for pairing with brightly plumaged males has been reported in many species, the reasons for this choice are not fully understood.
2. Parasites have been proposed as playing an important role in shaping these preferences because, by pairing with brightly coloured individuals, females can obtain parasite-free and/or more healthy mates.
3. In this paper one of the predictions of this hypothesis is tested, namely a higher health level in brightly coloured individuals, by analysing the relationships among blood parasites, leucocyte levels and plumage brightness in the Cirl Bunting, Emberiza cirlus .
4. Two species of blood parasites were detected. Whereas a lower body condition was detected in individuals infected by Leucocytozoon cambournaci, no such differences were associated with Plasmodium relictum infections. Infected individuals showed higher total leucocyte counts than non-infected individuals.
5. Colour intensity of carotenoid derived colorations was negatively correlated to the relative proportion of lymphocytes and positively correlated to the relative presence of heterophils. Furthermore, the size of yellow feathered areas was positively correlated with the absolute number of leucocytes and the relative presence of heterophils. Only some of these relationships were found for non-carotenoid derived traits.
6. These results suggest that male plumage yellow coloration in the Cirl Bunting is a reliable indicator of health status and supports the hypothesis that females obtain more parasite-free mates if they pair with brightly coloured individuals.     

The status of the Cirl Bunting Emberiza cirlus in the UK and the Channel Islands in 1998

A survey of breeding Cirl Buntings in the United Kingdom and the Channel Islands was undertaken in 1998. An estimated 453 (95% CI, 415–504) territories were recorded in 103 tetrads in Devon. This amounts to an increase of 29% in the Devon population since the last full survey in 1993, when 352 territories were recorded in 95 tetrads. In addition, nine territories were located on the island of Jersey in 1998 although none were found elsewhere. In 1993 there were 18 singing males on Jersey with a further two in both Cornwall and Somerset. The Cirl Bunting population in Devon seems to be consolidating rather than expanding as the only significant increase in numbers between 1993 and 1998 was from those tetrads occupied in both years. This paper also summarizes the results of surveys during 1989–97 and assesses the changes in the Cirl Bunting population over that period.     

Genetic variation and bill size dimorphism in a passerine bird, the reed bunting Emberiza schoeniclus

In passerine birds morphological differentiation in bill size within species is not commonly observed. Bill size is usually associated with a trophic niche, and strong differences in it may reflect the process of genetic differentiation and, possibly, speciation. We used both mitochondrial DNA (mtDNA) and nuclear microsatellites to study genetic variation between two subspecies of reed bunting, Emberiza schoeniclus schoeniclus and E.s. intermedia , along their distributional boundary in western Europe. These two subspecies are characterized by a high dimorphism in bill size and, although breeding populations of the two subspecies are found very close to each other in northern Italy, apparently no interbreeding occurs. The observed morphological pattern between the two subspecies may be maintained by geographically varying selective forces or, alternatively, may be the result of a long geographical separation followed by a secondary contact. MtDNA sequences of cytochrome b and ND5 (515 bp) showed little variation and did not discriminate between the two subspecies, indicating a divergence time of less than 500 000 years. The analysis of four microsatellite loci suggested a clear, although weak, degree of genetic differentiation in the large- and small-billed populations, as indicated by F _ST and R _ST values and genetic distances. The correlation between bill size and genetic distance between populations remained significant after accounting for the geographical distances between sampling localities. Altogether, these results indicate a very recent genetic differentiation between the two bill morphs and suggest that a strong selection for large bills in the southern part of the breeding range is probably involved in maintaining the geographical differentiation of this species.     

Variation in singing style use in the reed bunting Emberiza schoeniclus: influencing factors and possible functions

The two main functions of bird song are territory defence and mate attraction. Considerable progress has been made in understanding how species adjust the use of songs to serve these and other (presumed) functions of bird song, but the striking variety of singing behavior observable in wild birds remains enigmatic. Some species make do with simple songs and small repertoires, while others show large, complex repertoires and still others have evolved several distinct singing styles. In most species with distinct singing styles, however, the functions of singing styles are poorly understood. Two distinct singing styles (type I and II, respectively) have long been known in the reed bunting Emberiza schoeniclus, while a new third one has recently been reported to exist. We first quantitatively investigated the evidence for the existence of three singing styles. Then, we tested predictions of the mate attraction hypothesis, the mate guarding hypothesis and the territory defence hypothesis by examining the relations between singing style use with social and temporal factors. Cluster and discriminant analyses supported the existence of three (instead of two) singing styles, which could be differentiated based on four variables referring to song structure and complexity. Use of singing styles was related to male mating status (consistent with the mate attraction hypothesis), but not to female breeding stage (no support for the mate guarding hypothesis). Finally, use of singing styles differed in relation to time of day, with the dawn chorus of paired reed buntings consisting almost exclusively of songs of the recently discovered type III singing style and daytime singing primarily consisting of songs of long‐known type I (in unpaired males) or II singing styles (in paired males). Our findings suggest that one singing style (type I) primarily serves to attract a social mate, although an additional territorial function of this singing style cannot be dismissed. The function(s) of the other two singing styles, both only sung by paired males, are not related to attraction of a social mate or to the own female's fertility, but appear to be important in the context of territory defence and extra‐pair matings.     

芦鹀种群中的文化、遗传和形态进化的同时比较

我们比较了芦 (Emberizaschoeniclus)两个亚种组 ,即北部的薄喙亚种组和南部的厚喙亚种组的 10个种群中的文化、遗传和形态变异。使用了四个不同的变异标记物 ,其中两个用来测量文化分化 ,一个用来测量遗传分化 ,即微卫星等位基因的频次 ,一个用来测量种群的形态分化 ,即喙的高度。将遗传分化作为进化时间的尺度 ,我们计算了亚种组间和组内分化指标与所估计的进化率之间的相关性 ,发现只有文化定量指标和遗传分化与种群的形态分化相关 ,而两个文化分化指标之间没有关系 ,文化分化与遗传分化之间也没有关系。使用文化 -定量分化指标 ,发现亚种组间的文化进化率高于亚种内的文化进化率 ,提示鸣唱在防止杂交方面只有微弱的、也许是次要的作用。鸣唱定量特征的变异与微卫星频次相同 ,实际上在自然界中更可能是遗传决定的 ,这可以解释由于分析两个文化变异指标所得出的结果的不一致性。鸣唱的声学特性可能由于栖息地的差异或形态上的限制而发生了演变 ,而文化传播单位 (Meme)的特性可能由于学习鸣唱和文化传播而受到了影响     

Wavelength Dependency of Light-Induced Effects on Photoperiodic Clock in the Migratory Blackheaded Bunting (Emberiza melanocephala)

The effects of light wavelength on photoperiodic clock were determined in the migratory male blackheaded bunting (Emberiza melanocephala). We constructed an action spectrum for photoperiodic induction (body fattening, gain in body mass, and gonadal recrudescence) by exposing birds for 4.5 weeks to 13 h light per day (L:D = 13:11 h) of white (control), blue (450 nm), or red (640 nm) color at irradiances ranging from 0.028 to 1.4 W m^?2. The threshold light irradiance for photoinduction was about 10-fold higher for blue, compared to red and white light. Phase-dependent effects of light wavelength on the photoperiodic clock were further examined in the next two sets of skeleton photoperiods (SKPs). In the first set of SKPs, birds were exposed for four weeks to asymmetrical light periods (L:D:L:D = 6:6:1:11 h) at 0.25 ± 0.01 W m^?2; two light periods applied were of the same (450 nm: blue:blue, B:B; 640 nm, red:red, R:R) or different (blue:red, B:R or red:blue, R:B) wavelengths, or of white:white (W:W, controls). Photoperiodic induction occurred under R:R and B:R, but not under B:B and R:B light conditions; the W:W condition induced an intermediate response. The second set of SKPs used symmetrical light periods (L:D:L:D = 1:11:1:11 h), and measured effects also on the activity rhythm. Birds were first exposed to one of the four SKPs (R:R, B:B, R:B, or B:R) for three weeks, subsequently were released into dim constant light (LL_dim; ?0.01 W m^?2, the night light used in an L:D cycle) for two weeks, and then were returned to respective SKPs for another three weeks. Activity was greater in the R:R compared to B:B, and in B:R compared to R:B light condition. Zugunruhe (intense nighttime activity, indicating migratory restlessness in a caged situation) developed under the R:R and B:R, but not the B:B and R:B, light condition. Under LL_dim, all birds free-ran with a period >24 h, the Zugunruhe had a circadian period longer than the daytime activity, and the re-entrainment to SKPs was influenced by the position of light periods relative to circadian phase of the activity rhythm. Photoperiodic induction at the end of 8 weeks was found in the R:R and B:R, but not in B:B, light conditions; in the R:B condition only one bird had initiated testes. Taken together, these results suggest that in the blackheaded bunting, the circadian photoperiodic clock is differentially responsive to light wavelengths; this responsiveness is phase-dependent, and the development of Zugunruhe reflects a true circadian function. Wavelength-dependent response of the photoperiodic clock could be part of an adaptive strategy in evolution of the seasonality in reproduction and migration among photoperiodic species under wild conditions.     

Responses of a sub-oscine bird during playback: effects of different song variants and breeding period

Vermilion flycatchers (Pyrocephalus rubinus) vary their song rate and song length across the breeding season. Males sing more and longer songs after nest construction than before. Here we explored the possibility that this variation is meaningful to territorial males. Using a playback approach, we tested several males with different variations in song output (i.e. variations in song length and song rate) in different periods of the breeding season (i.e. before and after the onset of nest construction). We found that males call more in response to playbacks of long and short songs before the onset of nest construction. However, after nest construction began they responded flying more when exposed to long songs than to short songs. These results show that vermilion flycatcher discriminates between different variants of song length, and suggest that males react to long songs as if they were more threatening signals than short songs, especially after the onset of nest construction. We did not find evidence of males discriminating between a high and a low song rate. We discuss some possible implications for song function in this sub-oscine species, and compare these findings with other results in oscine species.