Temporal patterns of territory settlement and detectability in mated and unmated Nightingales Luscinia megarhynchos

In many bird species the sex ratio of adults is male-biased, which is likely to have consequences for the ecology as well as for the conservation of a species. For example, when some males remain unpaired in a population, there should be strong selection on behavioural traits that enhance pairing success. A surplus of males is also likely to have important implications for the interpretation of breeding bird survey data. In our study population of Nightingales Luscinia megarhynchos, about half of the males stayed unpaired, suggesting that the number of males encountered singing was greater than the number of breeding pairs. Furthermore, the detectability (the probability of encountering a male singing) of mated males was only two-thirds that of unmated males when censused in the morning or late in the breeding season. The relative detectability was more similar early in the season and during the twilight periods before sunrise and after sunset. Males that arrived earlier on the breeding grounds were more successful in attracting a mate than males arriving later. Some of the unmated males deserted their territories and prospected areas up to 4000 m distant, whereas others settled on the study site only late in the season and may actually have changed territories. We suggest that adult sex ratios and the time of the census should be taken into account when interpreting the results of breeding bird surveys.

Acoustic features of song categories and their possible implications for communication in the common nightingale (Luscinia megarhynchos)

In many passerine species, males sing more than one distinct song type. Commonly, songs are assigned to different song types or song categories based on phonological and syntactical dissimilarities. However, temporal aspects, such as song length and song rate, also need to be considered to understand the possible functions of different songs. Common nightingales (Luscinia megarhynchos) have large vocal repertoires of different song types but their songs additionally can be grouped into two distinct categories (particular groups of song types): whistle songs and nonwhistle songs. Whistle songs are hypothesised to be important to attract migrating females. We studied temporal properties of whistle songs and nonwhistle songs and examined the relationship between those song parameters and song output parameters, such as song rate and song length. To investigate how song parameters vary among males, we calculated the coefficients of variation for different song traits. We found that the variation in the proportion of whistle songs was significantly higher among males than variation in other song parameters. Furthermore, the proportion of whistle songs was negatively correlated with other sona output patterns. These findings suggest that the production of whistle songs might be constrained and/or that whistle songs and their succeeding pauses may act as a functional unit in communication.

Seasonal patterns of singing activity vary with time of day in the nightingale (Luscinia megarhynchos)

Seasonal patterns of singing activity of male birds have been thoroughly studied, but little is known about how those patterns vary with time of day. Here, we censused mated and unmated male Nightingales (Luscinia megarhynchos) at four different hours of the day throughout the breeding cycle. In unmated males, singing activity increased until the young hatched in their neighborhood, and the seasonal variation was similar at each of the four hours of the day. In mated males, however, the seasonal patterns of singing activity differed between hours of the day. In morning (about the hour of egg-laying) and during the dusk chorus, the singing activity of mated males was strongly influenced by the females' reproductive state: singing activity was low before egg-laying and during incubation, but high during the egg-laying period. In the dawn chorus, however, singing activity showed a similar seasonal pattern in mated and unmated males and was high until late stages of the breeding cycle. Our results suggest that the social context influences singing behavior to a varying degree across the season, and that this variation also depends on time of day. The hour of data collection thus is an important but often neglected factor when seasonal changes of singing activity are studied.

De Luscinia quae est apud veteres.

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云南新平哀牢山夜间捕获鸟类的多样性

2007-09-05-2007-11-05和2008-09-03-2008-11-07期间,在云南省新平县哀牢山金山丫口用灯光诱捕对秋季夜间迁徙鸟类组成进行了调查.共捕获鸟类129种,7943只,分属于14日23科.雀形目鸟类为网捕主要鸟类,占总网捕数量的58.9%.其中红尾伯劳(Lanius cristatus)、小杜鹃(Cuculus poliocephalus)和红喉歌鸲(Luscinia calliope)3种是该地夜间网捕优势种鸟类,三者数量皆超过网捕岛类总数的5%.每天夜间20:00-22:00和次日凌晨04:00-4)6:00是捕获鸟类数最的高峰时段.网捕期间,早期Shannon-Wiener指数和Pielou指数最低,优势指数C最高;中期Shannon-Wiener指数和Pielou指数最高,优势度指数C最低.2008与2007年相比,优势指数C和G-F指数较低.但Shannon-Wiener指数和Pielou指数较高.云南省不同"打雀山"仪问捕获鸟类的种类和数量存在差异.地理和气候可能是造成这些差异的主要因素.

哀牢山金山丫口夜间网捕鸟类多样性及其影响因素

2007 年9 月5 日至11 月5 日和2008 年9 月3 日至11 月7 日期间，主要利用灯光诱捕，对云南省新平县哀牢山金山丫口（23°57′N, 101°30′E）夜间迁徙鸟类多样性进行调查。同时记录网捕鸟类的时序和几种主要鸟类的迁徙时序。分析各种因子（地理、温度、湿度、雾天、风向、风力以及月相周期）对夜间网捕鸟类数量和种类的影响。 2 年期间共网捕鸟类7943 只，129 种，分别属于14 目23 科。雀形目鸟类占总网捕数量的58.9%。网捕数量超过1%的科共10 个科。网捕数量占总网捕数量1%以上的共14 种。其中红尾伯劳Lanius cristatus、小杜鹃Cuculus poliocephalus、红喉歌鸲Luscinia calliope 三者的数量超过网捕鸟类总数的5%。所有网捕鸟类中，东洋型鸟类占最多共54 种，占41.85%。 2 年期间网捕鸟类多样性指数H′为2.60989，均匀度指数J 为0.29063，优势度指数C 为 0.22503，G-F 指数为0.71366。2007 年的优势指数C 和G-F 指数较2008 年高，但2008 年的 Shannon-Wiener 指数和Pielou 指数较2007 年高。红尾伯劳是造成两年多样性测度指数差异的主要原因。云南省不同“打雀山”夜间捕获鸟类的种类和数量存在差异。地理和气候可能是造成这些差异的主要因素。 2 年工作期间将观察时间平均分为早期（9 月3 日至9 月24 日）、中期（9 月25 日至10 月16 日）、晚期（10 月17 日至11 月8 日）三个时期。早期网捕鸟类数量最多5263 只，多样性指数最低，优势度指数最高；中期记录的种类最多86 种，多样性指数和均匀度指数中期最高。造成这种差异的主要原因是早期红尾伯劳的数量尤其突出。单独分析2007 年和2008 年各时期，除了2007 年晚期种类最多67 种以外，其余变化都与2 年总体一致。每个时期都有部分鸟类数量达到高峰，而也有部分鸟类数量处在低谷期。飞经哀牢山金山丫口的秋季夜间迁徙鸟类开始迁徙的时间大致在8 月中旬，结束在11 月下旬。每种鸟类的迁徙持续时间不一致，其中迁徙持续时间超过30 天的鸟有36 种。同种鸟类不同年份迁徙时间不一致。夜间每个时段捕获数量和种类有差异，在夜间20:00-22:00 和凌晨04:00-06:00 捕获数量和种类较多。24:00-02:00 网捕鸟类数量和种类最少。调查结果显示不是所有鸟类都是整夜迁徙，夜间开始迁徙较早的鸟类清晨也停止迁徙较早，夜间开始迁徙较晚的鸟类清晨停止迁徙较晚。除红喉歌鸲以外的4 种鸟，它们的迁徙时间是从9 月前至11 月后。红尾伯劳、小杜鹃以及大杜鹃的迁徙高峰期是9 月上旬至9 月中旬。厚嘴苇莺Acrocephalus aedon 的迁徙高峰期为 9 月中旬至10 月下旬。红喉歌鸲的迁徙时间是9 月下旬至11 月上旬后，迁徙高峰期为整个 10 月。5 个优势种都是整夜迁徙，不过每种鸟的迁徙高峰时段和低谷时段不一致。红尾伯劳、小杜鹃、大杜鹃在20:00-22:00 捕获数量最多；红喉歌鸲、厚嘴苇莺在凌晨04:00-06:00 捕获数量最多。红尾伯劳、红喉歌鸲、大杜鹃在24:00-02:00 捕获数量最少；厚嘴苇莺在22:00-24:00 捕获数量最少；小杜鹃在凌晨02:00-04:00 捕获数量最少。对5 个优势种在各迁徙时期的体重进行分析，采用双样本T 检验（Independent-Samples T Test）表明：红喉歌鸲和小杜鹃早中晚期的体重差异极为显著（P<0.01）。运用单因素方差分析（One-Way ANONA）表明：小杜鹃早中晚期的体重变化极其显著（P<0.01）。进一步采用双样本T 检验（Independent-Samples T Test）：小杜鹃晚期体重与早期和中期体重差异都极其著（P<0.01），早期与中期的体量没有显著差异（P>0.01）。其余3 个优势种各时期体量差异不显著（P>0.01）。影响夜间网捕鸟类种类和数量的因素较多。地理特点是影响夜间网捕鸟类种类的主要因素。气候条件是影响夜间网捕鸟类种类和数量的重要因素。风向和风力对夜间网捕数量和种类都有极其显著的影响（P<0.01），其中风向对其影响比风力大，西南风时夜间网捕鸟类数量和种类较多。雾天对夜间网捕数量和种类影响极其显著（P<0.01），大雾天气夜间网捕鸟类数量和种类较多。月相周期对夜间网捕数量及其种类的影响极其显著 (P<0.01)，新月和残月期间夜间网捕鸟类数量和种类较多。温湿度对夜间网捕鸟类数量和种类影响不显著(P>0.05)。除此之外，光也是捕获夜间鸟类的必要条件。

Responses to interactive playback predict future pairing success in nightingales

Birdsong is a sexually selected trait that serves in territory defence and mate choice. Individual song traits can be affected by the body condition of the male and thus may reflect his quality. Such relations between male quality and general singing performance raise the question whether differences in male quality also affect response strategies used in dyadic interactions. To address this question, we studied the relation between pairing success of male common nightingales, Luscinia megarhynchos, and their responses to rivals posing different levels of threat. Using interactive playback, we exposed males prior to mating to either aggressively or moderately singing rivals (by song overlapping and song alternating, respectively). Males that remained unpaired throughout the season (bachelors) interrupted their singing significantly more often after being overlapped than after alternating playback, whereas subsequently mated males kept the number of singing interruptions more constant across playback treatment. This suggests that subsequently paired males are less discriminative than are bachelors when challenged by rivals varying in aggressiveness. Regardless of playback treatment, males that later became paired responded significantly more strongly than did bachelor males. Thus, an increase in singing after a vocal interaction prior to mating predicted future mating success. (c) 2006 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Seasonal variation in dawn song characteristics in the common nightingale

Many male temperate zone passerines show a marked peak of singing activity before sunrise. The two main functions of this so-called dawn chorus are mate attraction and territory defence. We examined how seasonal patterns of different dawn song characteristics were related to mating status and to the breeding cycle of females in the common nightingale, Luscinia megarhynchos. We investigated two measures of song output: song rate and percentage performance time. We also calculated the proportion of 'whistle songs', a song category that is thought to be important in female choice. We predicted that if the main function of dawn singing in nightingales is to attract a social mate, then mated males should change their dawn singing behaviour after pair formation. In contrast, if dawn singing is mainly used in territory defence, we expected no difference in song traits between mated and unmated males throughout the season. We found that song rate and the proportion of whistle songs were low at the beginning of the season and did not predict future mating status. After arrival of females, all measures of dawn song performance remained largely constant throughout the breeding season, and we did not find significant differences in the seasonal variation between mated and unmated males. These findings are consistent with the hypothesis that song at dawn is important to defend a territory throughout the breeding season. (c) 2005 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Effects of territorial intrusions on eavesdropping neighbors: communication networks in nightingales

Animal communication often occurs in communication networks in which multiple signalers and receivers are within signaling range of each other. In such networks, individuals can obtain information on the quality and motivation of territorial neighbors by eavesdropping on their signaling interactions. In songbirds, extracting information from interactions involving neighbors is thought to be an important factor in the evolution of strategies of territory defense. In a playback experiment with radio-tagged nightingales Luscinia megarhynchos we here demonstrate that territorial males use their familiar neighbors' performance in a vocal interaction with an unfamiliar intruder as a standard for their own response. Males were attracted by a vocal interaction between their neighbor and a simulated stranger and intruded into the neighbor's territory. The more intensely the neighbor had interacted with playback, the earlier the intrusions were made, indicating that males eavesdropped on the vocal contest involving a neighbor. However, males never intruded when we had simulated by a second playback that the intruder had retreated and sang outside the neighbor's territory. These results suggest that territorial males use their neighbors' singing behavior as an early warning system when territorial integrity is threatened. Simultaneous responses by neighboring males towards unfamiliar rivals are likely to be beneficial to the individuals in maintaining territorial integrity.

Aggressive responses to broadband trills are related to subsequent pairing success in nightingales

In sexually selected signals, distinct components often have specific signal value in mate choice or male-male competition. In songbirds, structural song traits such as trills, that is, a series of repetitive notes, can be important in female choice. However, little is known about their signal value in male-male interactions. Here, we investigated the hypothesis that males assess the competitive abilities of rivals based on the use and performance of rapid broadband trills produced within songs. Using a 2-speaker playback experiment, we exposed territorial male nightingales, Luscinia megarhynchos, that differed in their subsequent pairing success, to a simulated vocal interaction between 2 unfamiliar rivals. The singing of the 2 simulated rivals differed in the number of songs containing rapid broadband trills. Subjects responded significantly more strongly to the loudspeaker that broadcast songs containing such trills than to the loudspeaker that broadcast exclusively songs without such trills. Moreover, responses also depended on the fine structure of trills. Males that became paired later in the season significantly increased their response intensity with increasing trill performance, whereas males that remained unpaired responded in the opposite way and decreased their response intensity with increasing trill performance. These results indicate that rapid broadband trills are a signal of aggression and that the nature of the response in vocal interactions reflects aspects of the challenged male's fitness. © The Author 2008. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology. All rights reserved.


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