Habitat preference of endangered Eastern Iberian Reed Buntings Emberiza schoeniclus witherbyi

Capsule Careful management is required to maintain or create habitats with the correct proportions of rush with reeds that are preferred by this subspecies.

Aims To determine the habitat requirements of Eastern Iberian (Western Iberian Reed Buntings were not included in the study) Reed Buntings Emberiza schoeniclus witherbyi, a subspecies endemic to a few marshes in France, Spain and Morocco, with a population of 254–360 breeding pairs.

Methods Twenty-five wetlands in Spain were surveyed and presence/absence of Iberian Reed Buntings was deteremined together with data on 41 environmental variables. The relationships between the birds and habitat data were investigated using glm.

Results Proportion of rush with reeds, proportion of reed–bulrush patches and perimeter length of rush–reed patches, were the variables that primarily explained the present distribution of Eastern Iberian Reed Buntings. Other vegetation classes traditionally associated with the presence of the Eastern Iberian Reed Buntings, such as sedges, were not found to be important in the selected models.

Conclusion Conservation planning and management guidelines for this subspecies should focus on encouraging these specific-vegetation parameters. Flood control and grazing are proposed as the best practices that will favour these vegetation variables and avoid simplification of the plant community.     

The breeding ecology of Reed Buntings Emberiza schoeniclus in farmland and wetland habitats in lowland England

We studied the nesting and feeding ecology of Reed Buntings Emberiza schoeniclus breeding on farmland and wetland habitats along the Trent Valley in Nottinghamshire, England. Rank and emergent vegetation accounted for most nests and most foraging by provisioning adults. Caterpillars and spiders accounted for 70% of chick invertebrate prey and all broods were fed cereal grain or other vegetable matter. Variation in the abundance of key invertebrate prey across habitats accounted for the foraging preferences of adult buntings. Depredation was the main cause of nest failure, and survival of nests at the egg stage was positively related to the extent of nest concealment. A measure of total brood biomass was positively related to the abundance of key invertebrate prey within 100 m of nests. Rank and emergent vegetation provided Reed Buntings with greater nest concealment and a richer source of invertebrate prey than agricultural habitats such as set-aside, cereals and oilseed rape. The provision of rank and emergent vegetation on farmland is likely to increase the nesting opportunities and productivity of Reed Buntings in agricultural landscapes.     

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     

ASYMMETRIC RESPONSE PATTERNS TO SUBSPECIES‐SPECIFIC SONG DIFFERENCES IN ALLOPATRY AND PARAPATRY IN THE GRAY‐BREASTED WOOD‐WREN

Song divergence between closely related taxa may play a critical role in the evolutionary processes of speciation and hybridization. We explored song variation between two Ecuadorian subspecies of the gray‐breasted wood‐wren (Henicorhina leucophrys) and tested the impact of song divergence on response behaviors. Songs were significantly different between the two subspecies, even between two parapatric populations 10 km apart. Playback experiments revealed an asymmetric response pattern to these divergent subspecies specific songs; one subspecies responded more to songs of its own subspecies than to the other subspecies’ songs, whereas the second responded equally strongly to songs of both subspecies. While song parameters revealed a mixed pattern of divergence between allopatric and parapatric populations, the majority of spectral characteristics showed increased divergence in parapatry, suggestive of character displacement. This increased song divergence in parapatry appeared to affect behavioral responses to playback as discriminating responses were most prominent in parapatry and against parapatric songs. The clear behavioral impact of subspecies‐specific song differences supports a potential role for song as an acoustic barrier to gene flow. The asymmetric nature of the responses suggests that song divergence could affect the direction of gene flow and the position of the subspecies‐specific transition.     

BEHAVIOURAL RESPONSE TO SONG PLAYBACK BY MALE AND FEMALE WHITE-CROWNED SPARROWS OF TWO SUBSPECIES

ABSTRACT

Nuttall's white-crowned sparrow Zonotrichia leucophrys nuttalli and the Puget Sound white-crowned sparrow Z. i. pugetensis intergrade through a zone of hybridization in northern California. The songs of the two differ in the sequence of syllable types and in the phonology of the complex syllables. We did a playback experiment to males of both subspecies in the field and to females of both subspecies in the laboratory to determine their response to the two subspecies songs. By making alterations of features of the songs, we also attempted to identify characteristics that account for subspecific recognition. Males of both subspecies were more responsive to their own subspecies song than to that of the other subspecies. Substitution of Puget Sound complex syllables into the Nuttall's song caused a reduction in response by male Nuttall's, but alteration of a Nuttall's song to a Puget Sound sequence of syllables had no effect. Females of both subspecies gave more sexual displays when played a song of their own subspecies than when played that of the other subspecies. Female Nuttall's gave an intermediate response when played Nuttall's songs with Puget Sound complex syllables or when played Nuttall's songs altered to a Puget Sound sequence. Puget Sound females did not discriminate between their own subspecies song and either of the altered stimulus songs separately. They did respond less to a song that combined both alterations than to their own subspecies song.     

Seed food preferences of granivorous farmland passerines

Capsule Large buntings prefer cereal grains whilst sparrows also take oily seeds.

Aims To determine seed food preferences of Tree Sparrow Passer montanus and Corn Bunting Emberiza calandra.

Methods Forty seed choice experiments were conducted at two sites over two winters. In each experiment, two seed types were provided and the number of visits made by birds to each type was recorded over a set period. At one site, Tree Sparrows were colour-ringed, allowing choices made by individual birds to be recorded. Data were also collected for House Sparrow Passer domesticus, Yellowhammer Emberiza citrinella and Reed Bunting E. schoeniclus.

Results All five species fed intensively on cereal grain, and wheat and oats were consistently preferred to barley. Sparrows and Reed Buntings also took maize, which was avoided by Corn Bunting and Yellowhammer. Tree Sparrow exhibited a broad diet, selecting cereal grain and oily seeds including sunflower and oilseed rape, but rye-grass seed was almost completely avoided.

Conclusion Cereal grain should be a key component of over-winter provision of seed for farmland passerines, especially when targeted at Corn Bunting and Yellowhammer. Oily seeds such as brassicas and sunflower will benefit species with more generalist diets, including Tree Sparrows.     

Nest predation in Reed Buntings Emberiza schoeniclus: an experimental study

Predation is a major cause of nest failure in many bird species. High levels of nest loss may be a consequence of habitat fragmentation, leading to increased amounts of edge habitat. Yet the evidence for generally high nest predation rates along edges in fragmented landscapes is ambiguous. Using real nests of Reed Buntings Emberiza schoeniclus in which artificial Reed Bunting and real Japanese Quail Coturnix japonica eggs were placed, we experimentally tested for edge effects on nest predation in highly fragmented reed Phragmites sp. habitats in the Swiss lowlands. We also examined seasonal patterns of predation and the impacts of nest visits by observers. We found evidence for an edge effect at the water-sided reed edge, with nests located closer to the water being more likely to be predated than those further away. Predation probability increased from early to late season, suggesting that nest predation may be density dependent. Probability of nest predation was only weakly influenced by whether or not a nest was visited. Our results suggest that the intensive reed management currently applied in Swiss nature reserves may result in unnaturally high levels of nest losses in the Reed Bunting, because reed bands are not wide enough to allow nest placement at a safe distance from reed edges.     

Territorial reactions of male Yellowhammers (Emberiza citrinella) toward a specific song structure

Male songbirds assess their rivals by listening to their songs and, in some species, are especially sensitive to certain parts of the song. Yellowhammers (Emberiza citrinella) sing songs with different structures consisting of an initial phrase of stereotyped and repeated notes in a trill, and a terminal phrase, present or not, which consists of low or high frequency whistles that we called elements. We analyzed 1,055 songs from 25 different males and found that four different structures could be distinguished in a French Yellowhammer population. We found that the frequency ranges of the elements were more constant across individuals than the ranges of the notes, and among them, the low frequency element was nearly twice as conserved as the high frequency element. Then, in the wild, we tested the territorial responses of males to the four different structures and found that the full song structure, which is composed of the initial phrase followed by a high frequency element and a low frequency element, induced stronger territory defence responses from the males than any of the other structures. As only the structure containing the initial phrase and the low frequency element also induced a strong response, we propose that the main element responsible for the enhanced reaction of the males is the low frequency element, which is consistent with the fact that it is more conserved than the other constituents.     

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.     

Signal value of alternating and overlapping singing in the Yellowhammer<Emphasis Type="Italic"> Emberiza citrinella</Emphasis>

Interactive playback experiments were used to study the signal value of alternating and overlapping singing in the Yellowhammer Emberiza citrinella. We carried out interactive playback experiments in which 13 males were subjects of two treatments that differed in temporal pattern of playback song delivery (alternating vs overlapping). We measured 12 parameters of males' response, belonging to three categories: song output, call output and mobility (distance from loudspeaker and flights). The results do not confirm the hypothesis that the overlapping pattern is a signal of stronger threat, as compared to the alternating pattern. Overlapping and alternating playbacks generally elicited a similar response, characteristic for highly aroused males. The only significant difference found concerned latency of approach to the loudspeaker during the playback stage. When playback alternated with songs of males, the birds gained higher scores on the axis of that response measure. Such a result was linked to differences in how the alternating and overlapping playbacks affect detection and localisation of simulated rivals, rather than to the signal's threat value itself. The strong response of Yellowhammer males is consistent with the high intrusion rate and possibility of extra-pair copulation in this species, which probably simplifies close-range communication towards more aggressive and unequivocal repelling of rivals irrespective of the temporal patterning of song.     

The geographical distribution of populations of the large-billed subspecies of reed bunting matches that of its main winter food

We investigated the effects of resource distribution on the population structure and distribution of a polytypic bird species. We compared the presence of insect remains (mainly dormant larvae) in the winter diet of two reed bunting subspecies, small billed E. s. schoeniclus and large billed E. s. intermedia and studied the distribution of this resource within reed ( Phragmites sp.) stems in seven north Italian localities where the two subspecies breed (three schoeniclus and four intermedia populations). We also tested if the distributions of winter insect resources and breeding populations of the large bill subspecies overlapped. The distribution of the larvae in reed stems matched closely the distribution of large billed breeding populations. The winter diets of the two subspecies were significantly different in terms of frequency of insect remains. These results, when compared to theoretical models of para-patric distribution, suggest that the two subspecies may be subject to ecological (vicariant) selection maintaining their reproductive isolation. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 21–26.     

What exactly is ‘local song’ in a population of ortolan buntings with a common dialect?

Previous studies have shown that ortolan buntings (Emberiza hortulana) exhibit apparent dialect variation. Neighbouring males typically share the same final phrase of a simple two-part song. Consequently, the final phrase was considered to be the dialect cue important for discriminating between males from a local population and strangers. Recently, it was shown that in an isolated and fragmented population of the ortolan buntings in Norway there was no local dialect in the above-mentioned sense. Norwegian males often had song types with different final phrases in their repertoire, and had larger repertoires and a lower level of song type sharing than in other populations. It was experimentally revealed that only local songs (L) evoked a strong response in Norwegian males, while these birds did not respond strongly to foreign (F) or hybrid songs composed of local and foreign initial and final phrases in any composition (i.e. both FL and LF songs). These results suggest that, in the Norwegian population, the final phrase of the song is not a sufficient cue for local song dialect recognition. This paper is a further study in which we tested the response of the ortolan bunting males to L, F, FL and LF songs in a typical continuous population (in Poland) in which males share the same single final phrase, i.e. have a common dialect. We found that ortolan bunting males in Poland responded with similar strength to L, FL and LF songs. The majority of males did not respond as strongly to playback of only F songs. Our results suggest that a common final phrase for a population should not be treated as the only signal of ‘locality’. Our results show clear asymmetry in response to hybrid songs with non-local phrases in different populations.     

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.     

A survey for Yersinia pseudotuberculosis in migratory birds in coastal Japan

Yersinia pseudotuberculosis was isolated from three specimens of two species of birds, the black-faced bunting (Emberiza spodocephala) and pied wagtail (Motacilla alba), of 528 specimens of birds examined from coastal regions in Japan. The two isolated strains of Y. pseudotuberculosis were identified as serovar 4b and serovar 3. This is the first isolation of Y. pseudotuberculosis from birds in Japan. Yersinia enterocolitica was isolated from three specimens of the pied wagtail, one specimen of the reed bunting (Emberiza schoeniclus) and one specimen of the rustic bunting (Emberiza rustica). Yersinia frederiksenii was isolated from two specimens of the gray-rumped sandpiper (Heteroscelus brevipes). Yersinia intermedia was isolated from one specimen of the pied wagtail.     

Behavioural response to song and genetic divergence in two subspecies of white‐crowned sparrows (Zonotrichia leucophrys)

Divergence in sexual signals may drive reproductive isolation between lineages, but behavioural barriers can weaken in contact zones. Here, we investigate the role of song as a behavioural and genetic barrier in a contact zone between two subspecies of white‐crowned sparrows (Zonotrichia leucophrys). We employed a reduced genomic data set to assess population structure and infer the history underlying divergence, gene flow and hybridization. We also measured divergence in song and tested behavioural responses to song using playback experiments within and outside the contact zone. We found that the subspecies form distinct genetic clusters, and demographic inference supported a model of secondary contact. Song phenotype, particularly length of the first note (a whistle), was a significant predictor of genetic subspecies identity and genetic distance along the hybrid zone, suggesting a close link between song and genetic divergence in this system. Individuals from both parental and admixed localities responded significantly more strongly to their own song than to the other subspecies song, supporting song as a behavioural barrier. Putative parental and admixed individuals were not significantly different in their strength of discrimination between own and other songs; however, individuals from admixed localities tended to discriminate less strongly, and this difference in discrimination strength was explained by song dissimilarity as well as genetic distance. Therefore, we find that song acts as a reproductive isolating mechanism that is potentially weakening in a contact zone between the subspecies. Our findings also support the hypothesis that intraspecific song variation can reduce gene flow between populations.     

Neighbour-stranger song discrimination in territorial ortolan bunting Emberiza hortulana males

Neighbour-stranger discrimination has been demonstrated in many species, but the mechanisms employed in discrimination vary. We tested whether an oscine bird with small repertoire size, the ortolan bunting Emberiza hortulana , discriminated between songs of neighbours and strangers. We performed playback experiments to measure response of males to a repeated single example of a single song type derived from a repertoire of a neighbour or stranger. Thirteen males were tested twice each, and in both cases songs were broadcast from the territory boundary shared by the subject male and the neighbour. Subjects responded more aggressively to songs of strangers than neighbours, i.e. they approached the loudspeaker faster and came closer and did more flights during the playback of stranger song. We found no significant differences in vocal response between treatments. We conclude that ortolan bunting can discriminate between songs of neighbours and strangers. This study provides experimental evidence for ortolan buntings in neighbour-stranger discrimination. It also demonstrates that a single example of song is enough to discriminate between neighbours and strangers. We discuss which song characteristics are the possible acoustic basis for discrimination in the studied species.     

Individuelles Erkennen des Gesangs durch die Weibchen und Gesangsaktivit?t der M?nnchen bei der Rohrammer (Emberiza schoeniclus)

Zusammenfassung Die vorliegende Untersuchung ist der zweite experimenelle Nachweis, daß Singvogelweibchen im Freiland die Gesänge der eigenen Männchen erkennen können. Es ist zu erwarten, daß es sich um ein bei vielen Vogelarten verbreitetes Phänomen handelt. Wie gezeigt werden konnte, ergeben sich daraus weitreichende Folgen für Erklärungen zur Funktion des Gesangs und des Paarungsverhaltens.

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Individual recognition of song by the female and song activity of the male in the Reed Bunting (Emberiza schoeniclus)

Playback of tape recorded songs in the field showed that females of the Reed Bunting approached only songs of their mates. This is the second experimental evidence for songbirds that females are able to recognize the songs of individual males. This ability of the females is seen as a possible cause for increase of song activity after the start of breeding. Females should choose the male next to their nest to get his help for rearing their young. Therefore, the males should stay near the nests and sing to ensure that they are recognized and chosen during the next fertile period. Alternative hypotheses are discussed.

     

Morphological sexing of passerines: not valid over larger geographical scales

Sex determination of birds is important for many ecological studies but is often difficult in species with monomorphic plumage. Morphology often provides a possibility for sex determination, but the characters need to be verified. We tested whether five passerine species can be sexed according to standard morphological measurements applying a forward logistic regression with sex determined by molecular analysis as the dependent variable. Furthermore, we tested whether the results can be used on a larger geographic scale by applying morphological sexing methods gained by similar studies from other regions to our data set. Of the five species of this study only Garden Warblers Sylvia borin could not be sexed morphologically. In the Robin Erithacus rubecula, 87.2% of all individuals were sexed correctly. For Reed Warblers Acrocephalus scirpaceus, Willow Warblers Phylloscopus trochilus and Reed Buntings Emberiza schoeniclus, the respective values were 77.6, 89.4 and 86.4%. When the logistic regression functions from similar studies on Robins and Reed Buntings in Denmark and Scotland were applied to the birds from south-western Germany, they performed less well compared to the original dataset of these studies and compared to the logistic regression function of our own study. The same was the case for Willow Warblers when a wing length criterion used in Great Britain was applied to the birds of our study. These discrepancies may have several explanations: (1) the models are optimised for the dataset from which they were extracted, (2) inter-ringer variation in measurements, (3) the use of different age cohorts, (4) different morphology due to different habitat availability around the study site, or, most likely, (5) different morphology due to different migratory behaviour. We recommend that morphological sex differentiation methods similar to this study (1) be only used population specific, (2) only with one age cohort and (3) to adjust the extracted equations from time to time.