Research Station Station de Recherche Forschungsstation

Petite Camargue Alsacienne













The Petite Camargue Alsacienne is situated in the Upper Rhine Valley, which is home of a dense population of nightingales (Luscinia megarhynchos). In 2003, we counted 194 singing males in a study area of about 18 square kilometres (see illustration below). Since 1997, we capture and ring nightingales in a study area of one square kilometre, in up to 28 territories per year. Another important basis for our studies are regular nocturnal rounds of inspection at the study site. Since only unpaired males sing regularly at night, we can determine the mating status of males by observing who sings at night and who sings only during the day. Once they are paired, male nightingales usually stop singing at night (Amrhein et a. 2002, 2004a). This suggests that the famous nocturnal song plays a role in mate attraction. In experiments using song playback before the arrival of females, we found that prospective mated males overlap more songs of the playback, i.e. seem to sing more aggressive, than do males that remain unpaired throughout the breeding cycle (Kunc et al. 2006). Females could thus gain information on the quality of males by listening to the nocturnal singing.

As in many other songbird species, male nightingales sing most during the hour before sunrise. During this dawn chorus, not only unpaired males but also paired males sing throughout the breeding season (Amrhein et al. 2004a; Kunc et al. 2005a). Dawn singing may thus be important to defend the territory against other males. This might be crucial especially if some unpaired males leave their territories in the course of the breeding cycle to become so-called floaters, and to prospect other territories before sunrise. Each year, up to 49% of the males remain unpaired (Amrhein 2004), and non-territorial males that were followed using radio-telemetry truly visited up to five occupied territories during the hour before sunrise (Amrhein et al. 2004b).

By means of radio-telemetry we could also show that territorial male nightingales intrude into the territory of their neighbour after that neighbour was challenged by a singing intruder (simulated by playback; Naguib et al. 2004). Male nightingales thus respond to territorial problems of their neighbours and can be considered members of an interactive communication network. Currently, we make paternity analyses with blood probes that we collected since 1998. First results suggest that the rate of extra-pair fertilization is not particularly high (about 7.5% of offspring; Amrhein 2004).

Study area (light grey) with the Petite Camargue Alsacienne (PCA). Dots are 194 singing nightingales and 8 additional territories in the Petite Camargue that were not monitored during the four censuses. From Amrhein & Zwygart 2004.

Publications from the nightingale research project


Roth, T. & Amrhein, V. 2010. Estimating individual survival using territory occupancy data on unmarked animals. Journal of Applied Ecology 47: 386-392. pdf

Sprau, P., Roth, T., Schmidt, R., Amrhein, V. & Naguib, M. 2010. Communication across territory boundaries: distance dependent responses in nightingales. Behavioral Ecology 21: 1011-1017. pdf

Sprau, P., Schmidt, R., Roth, T., Amrhein, V. & Naguib, M. 2010.
Effects of rapid broadband trills on responses to song overlapping in nightingales. Ethology 116: 300-308. pdf

Roth, T., Sprau, P., Schmidt, R., Naguib, M. & Amrhein, V. 2009. Sex-specific timing of mate searching and territory prospecting in the nightingale: nocturnal life of females. Proceedings of the Royal Society, Series B 276: 2045-2050. pdf. Highlighted in Nature 458: 10

Naguib, M., Schmidt, R., Sprau, P., Roth, T., Flörcke, C. & Amrhein, V. 2008. The ecology of vocal signaling: male spacing and communication distance of different song traits in nightingales. Behavioral Ecology 19: 1034-1040. pdf

Schmidt, R., Kunc, H. P., Amrhein, V. & Naguib, M. 2008. Aggressive responses to broadband trills are related to subsequent pairing success in nightingales. Behavioral Ecology 19: 635-641. pdf

Amrhein, V., Kunc, H. P., Schmidt, R. & Naguib, M. 2007. Temporal patterns of territory settlement and detectability in mated and unmated Nightingales Luscinia megarhynchos. Ibis 149: 237-244. pdf

Kunc, H. P., Amrhein, V. & Naguib, M. 2007. Vocal interactions in common nightingales (Luscinia megarhynchos): males take it easy after pairing. Behavioral Ecology and Sociobiology 61: 557-563. pdf

Schmidt, R., Amrhein, V., Kunc, H. P. & Naguib, M. 2007. The day after: effects of vocal interactions on territory defence in nightingales. Journal of Animal Ecology 76: 168-173. pdf

Kunc, H. P., Amrhein, V. & Naguib, M. 2006. Vocal interactions in nightingales, Luscinia megarhynchos: more aggressive males have higher pairing success. Animal Behaviour 72: 25-30. pdf

Schmidt, R., Kunc, H. P., Amrhein, V. & Naguib, M. 2006. Responses to interactive playback predict future pairing success in nightingales. Animal Behaviour 72: 1355-1362. pdf

Kunc, H. P., Amrhein, V. & Naguib, M. 2005a. Seasonal variation in dawn song characteristics in the common nightingale. Animal Behaviour 70: 1265-1271. pdf

Kunc, H. P., Amrhein, V. & Naguib, M. 2005b. Acoustic features of song categories and their possible implications for communication in the common nightingale (Luscinia megarhynchos). Behaviour 142: 1083-1097. pdf

Amrhein, V. & Zwygart, D. 2004. Bestand und Verpaarungsstatus von Nachtigallen Luscinia megarhynchos im elsässischen Rheintal bei Basel. Der Ornithologische Beobachter 101: 19-24. pdf

Amrhein, V., Kunc, H. P. & Naguib, M. 2004a. Seasonal patterns of singing activity vary with time of day in the Nightingale (Luscinia megarhynchos). The Auk 121: 110-117. pdf

Amrhein, V., Kunc, H. P. & Naguib, M. 2004b. Non-territorial nightingales prospect territories during the dawn chorus. Proceedings of the Royal Society of London, Series B (Suppl.) 271: S167-S169. pdf

Naguib, M., Amrhein, V. & Kunc, H. P. 2004. Effects of territorial intrusions on eavesdropping neighbors: communication networks in nightingales. Behavioral Ecology 15: 1011-1015. pdf

Amrhein, V., Korner, P. & Naguib, M. 2002. Nocturnal and diurnal singing activity in the nightingale: correlations with mating status and breeding cycle. Animal Behaviour 64: 939-944. pdf


Kugler, C. 2009. Dawn song performance predicts aggressiveness during vocal interactions in the nightingale. Master thesis, Universität Basel.

McStea, F. 2007. The effect of mating status on prospecting behavior of male Nightingales Luscinia megarhynchos in the late breeding season. Master thesis, Universität Basel.

Schmidt, R. 2007. Communication networks in nightingales: singing and territory defence in relation to mating success and settlement.
Ph.D. thesis, Universität Bielefeld. pdf

Amrhein, V. 2004. Singing activity and spatial behaviour as sexually selected traits in the Nightingale Luscinia megarhynchos. Ph.D. thesis, Universität Basel. pdf

Herhausen, C. 2004. Endoparasitism in the Nightingale Luscinia megarhynchos. Diploma thesis, Universität Bielefeld.

Kunc, H. P. 2004. Song and sexual selection in the nightingale (Luscinia megarhynchos). Ph.D. thesis, Universität Bielefeld.

Benzing, B. 2003. Spatial behaviour of mated and unmated radio-tagged nightingales (Luscinia megarhynchos) throughout the breeding season. Diploma thesis, Universität Bielefeld.

Amrhein, V. 1999. Das Revierverhalten der Nachtigall Luscinia megarhynchos. Diploma thesis, Universität Basel.