Abstract
Biparental incubation is frequent among shorebirds and is expected when the survival prospects of offspring increase relative to uniparental incubation. To understand why this occurs, it is important to identify the factors that constrain uniparental incubation. It is assumed that birds choose nesting sites that provide an appropriate microclimate for incubation. Many shorebirds nest in sites with no or little cover, where ambient temperatures at ground level might be >50°C during very hot days. Shorebirds nest in exposed sites because predation risk on incubating adults is higher in covered sites. In hot environments, incubating shorebirds might experience heat stress in exposed sites, and this may compromise nesting success if adults are unable to attend their nests continuously, limiting the possibilities of uniparental incubation and thus the expression of a sexual conflict over incubation. The operative temperatures of Kentish plovers (Charadrius alexandrinus) were recorded in exposed and covered sites, and the thermal behaviour and incubating tactics of pair members were studied in a hot environment. During the hottest part of the day, there was a difference of 10–15°C in the operative temperatures of plovers between covered and exposed sites. Plovers in covered sites did not exhibit any thermoregulatory behaviour indicative of thermal stress, probably because the thermal range encountered by them in such places during most of the daytime was close to the thermo-neutral zone. The frequency with which plovers in exposed sites exhibited thermoregulatory behaviour was related to ambient temperature. Under very hot conditions, incubating birds were probably unable to maintain homeostasis for long periods and pair members resorted to shortening incubation bouts. Female Kentish plovers mainly incubate in the daytime and males during the night. However, the probability of diurnal incubation by males increased with ambient temperature in exposed nests, but not in covered ones. In fact, the frequency of participation in diurnal incubation by males was greater in exposed than in covered sites, suggesting that the participation of males in diurnal incubation may be related to the inability of females to stay at the nest during long periods when the ambient temperature is high. Even after resorting to shortened incubation bouts, the plovers may be unable to attend their nests continuously during heat waves, and the nests may be deserted. The propensity of plovers to desert their nests was affected by proximity to water, with nests located close to water being deserted less frequently. It seems likely that susceptibility to thermal stress changed in relation to proximity to water because in sites close to water it was possible to belly-soak, which would allow a more continuous nest attendance. Therefore, despite the adoption of behavioural solutions to face heavy heat loads, nesting success was vulnerable to these solutions because heat stress during extended periods may constrain parental nest attendance, and this may limit the opportunities for sexual conflicts over incubation.
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Acknowledgements
Consejería de Medio Ambiente (Junta de Andalucía) gave permission to conduct this research at Reserva Natural Laguna de Fuente de Piedra, as well as providing lodging and many other facilities. Manuel Rendón-Martos, director of the Nature Reserve, facilitated our work during all phases of the project. In the field, we were assisted by A. Arroyo, J.L. Arroyo, R. Camarena, J.M. Ramírez, J. Rubio, M. Siquier and M. Vázquez. G. Nehls assembled the taxidermic mounts, and F. Chiara gave us technical assistance with these mounts. Our thanks also go to F. Álvarez, I. Quintero, T. Székely, J. Wright and a referee for many helpful comments on the manuscript. Financial support was provided by Plan Andaluz de Investigación (research group RNM 0105) and Dirección General de Investigación Científica y Técnica (grants PB92-0115 and PB95-0110).
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Amat, J.A., Masero, J.A. How Kentish plovers, Charadrius alexandrinus, cope with heat stress during incubation. Behav Ecol Sociobiol 56, 26–33 (2004). https://doi.org/10.1007/s00265-004-0758-9
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DOI: https://doi.org/10.1007/s00265-004-0758-9