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Individual quality is often signaled by phenotypic flags, such as bright plumage patches in birds. Extended phenotype signals can similarly show quality, but in these cases the signals are external to the individual, often taking the form of objects scavenged from the environment. Through multiple manipulative experiments, we showed that objects used for nest decoration by a territorial raptor, the black kite (Milvus migrans), act as reliable threats to conspecifics, revealing the viability, territory quality, and conflict dominance of the signaler. Our results suggest that animal-built structures may serve as signaling devices much more frequently than currently recognized.

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1. The deployment of electronic devices on animals is rapidly expanding and producing leapfrog advances in ecological knowledge. Even though their effects on the ecology and behaviour of the marked subjects are potentially important, <10% of the studies are accompanied by an evaluation of impact, and comprehensive, long-term assessments have been few. Therefore, there is an urgent need to test for impacts, especially for tags that are heavy and deployed for long time periods, such as satellite transmitters. 2. We marked 110 individuals of a medium-sized, migratory raptor, the black kite Milvus migrans, with GPS satellite tags, representing about 4% of the body mass and attached as backpacks through a Teflon harness. Tagged individuals were compared to control animals of similar sex, age and breeding status for a large number of behavioural, condition-related and ecological traits. 3. Despite a sample size two-to threefold greater than most previous assessments that reported significant impacts, there was no detectable difference between tagged and control individuals in key vital rates such as survival probability, longevity, recruitment, age of first breeding, reproductive performance and timing of breeding. 4. Tagged and untagged kites showed similar social dominance during fights over food and a similar capability to provision nestlings, which prevented carry-over effects on the stress levels and condition of their offspring. 5. Synthesis and applications. Radio-marking studies are growing exponentially in the current 'movement ecology era', and impact assessments will be ever more important. In principle, tags of up to 4% mass-load can be deployed without apparent harm on some avian soaring species, but impacts should be properly evaluated on a case-by-case basis. Resilient species for which impacts seem weak could be used as early warning systems for trials of new devices: if impacts are observed, they are likely to be even greater on more vulnerable species. Finally, individual fatalities caused by marking should be taken into serious account, but comprehensively evaluated in the light of broader population-level impacts. Future initiatives to minimize tagging impacts could include more stringent licensing criteria enforcing attendance at training courses or incorporation of impact evaluations into study designs, increased availability of training courses for tagging, and enhanced sharing of information through blogs, workshops or specialized journal sections.

a b s t r a c tPreserving large predators is important but challenging because these species are typically wide-ranging, select multiple habitats at different scales and often present spatial or habitat separation between the breeder and floater sectors of a population. In addition, most of our knowledge on raptor floaters' habitat requirements comes from large solitary species, whose floaters often occupy temporary settlement areas spatially separate from breeding locations. Here, we examine space and habitat use by a loosely colonial, wetland-dependent raptor, the Black kite ( Milvus migrans ), in a population where floaters co-exist with territory holders, enabling a direct comparison of their habitat preferences. The study was conducted in Doñana National Park (South-Western Spain), a seasonally drying marshland currently surrounded by intensive agriculture and rice-fields. Intensive radio-tracking revealed that breeders and floaters selected and avoided the same habitats despite a radical, four-to-eight fold difference in their homerange dimensions: all kites over-selected open habitats suitable for their aerial foraging modes and avoided woodland and farmland. These results suggest a continuum of raptor population structures ranging from solitary species whose floaters select different habitats than breeders and are concentrated in spatially separate settlement areas, to colonial and semi-social species whose floaters fully coexist with breeders with shared habitat preferences. Both extremes of this continuum will pose challenges for conservation management. In solitary species, special conservation efforts may be required to identify and manage temporary settlement areas, while in gregarious species, the larger ranges of floaters may expose them to different threats than breeders, whose occurrence and consequences may be subtle to identify..

The annual cycle of most animals is structured into discrete stages, such as breeding, migration and dispersal. While there is growing appreciation of the importance of different stages of an organism’s annual cycle for its fitness and population dynamics, almost nothing is known about if and how such seasonal effects can change through a species lifespan. Here, we take advantage of the opportunity offered by a long-term satellite/GPS-tracking study and a reliable method of remote death-detection to show that certain stages of both the annual and life cycle of a migratory long-lived raptor, the Black kite Milvus migrans, may represent sensitive bottlenecks for survival. In particular, migratory journeys caused bursts of concentrated-mortality throughout life, but the relative importance of stage-specific survival changed with age. On the other hand, the balance between short-stages of high mortality and long-stages of low mortality made population-growth similarly dependent on all portions of the annual cycle. Our results illustrate how the population dynamics of migratory organisms can be inextricably linked to ecological pressures balanced over multiple stages of the annual cycle and thus multiple areas of the globe, suggesting the frequent need for challenging conservation strategies targeting all portions of a species year-round range.

No environment is truly constant in time. As a result, animals have evolved multiple adjustments to cope with such fluctuations. However, the allocation of effort to costly activities that imply long-term commitments, such as breeding, may be extremely challenging when future resources change constantly and unpredictably, a context that has received little investigation. To fill this gap, we studied the breeding response by a wetland-dependent raptor, the black kite Milvus migrans, to within and between-years fluctuations in resource availability (inundation levels). The breeding performance of the population was decomposed into reproductive components expressed in a sequence of successive tasks along the breeding cycle (e.g. timing of laying, clutch size, hatching success, brood reduction). Variation in each component was related to resource levels observed at different key dates of the season in order to test whether and when population-level reproduction was adjusted to available resources. Along a 22-year time-series, inundation levels fluctuated unpredictably within and among years, and mostly affected the later components of kites' reproduction, such as hatching success and the incidence of brood reduction, which were the main determinants of the population yearly breeding output. Results were consistent with multiple adjustments to cope with uncertainty. As the season progressed and resources became easier to assess, a bet-hedging waiting strategy based on a conservatively small, invariant and asynchronous clutch gave way to real-time resource-tracking mechanisms mediated by progressive adjustments to current prey availability, so that population-level breeding rates were determined and tuned to resources rather late in the season. Such adjustments were the likely outcome of the interaction between parental tactics and environmental constraints. Behavioural flexibility, such as dietary opportunism, probably promoted further resistance to resource oscillations. Given that all ecosystems show some degree of unpredictability, resource-tracking adjustments, such as the ones depicted here, are likely to be commonplace in most communities.

Large, long-lived species with slow life histories and protracted pre-breeding stages are particularly susceptible to declines and extinction, often for unknown causes. Here, we show how demographic modeling of a medium-sized raptor, the Red Kite Milvus milvus, can aid to refocus conservation research and attention on the most likely mechanisms driving its decline. Red Kites' survival and reproduction increased through three sequential stages for 1-2, 3-6, and 7-30 yr of age, mainly corresponding to individuals that are dispersing, attempting to gain a territory, and breeding. As typical of long-lived species, elasticities were highest for adult (≥7 yr old) survival, but this was high, with little scope for improvement. Instead, the declines were driven by an extremely low survival of pre-adults in their first years of life, which weakened the whole demographic system by nullifying the offspring contribution of adults and curtailing their replacement by recruits. For example, 27 pairs were necessary to generate a single prime age adult. Simulation of management scenarios suggested that the decline could be halted most parsimoniously by increasing pre-adult survival to the mean levels recorded for other areas, while only the synergistic, simultaneous improvement of breeding success, adult and pre-adult survival could generate a recovery. We propose three actions to attain such goals through selective supplementary feeding of both breeding and non-breeding individuals, and through mortality improvement by GPS remote-sensing devices employed as surveillance monitoring tools. Our results show how improving demographic models by using real, local vital rates rather than "best guess" vital rates can dramatically improve model realism by refocusing attention on the actual stages and mortality causes in need of manipulation, thus building precious time and resources for conservation management. These results also highlight the frequent key role of pre-adult survival for the management of long-lived species, coherent with the idea of demographic systems as integrated chains only as strong as their weakest link.