Abstract
If individuals of the same population inhabit territories different in landscape structure and composition, experiencing habitat-specific demographic rates, then the landscape features become major determinants of the overall population characteristics. Few studies have tested how habitat-specific demography interacts with landscape heterogeneity to affect populations of territorial species. Here we report a 29-year study of an eagle owl (Bubo bubo) population in southern France. The aim of this study was to analyse how habitat heterogeneity could affect density and breeding performance. Mean productivity for the overall sample was 1.69±0.76 fledglings per breeding pair and, after controlling for year effect, significant differences between territories were detected for productivity. A positive correlation was found between the percentage of pairs producing 50% of the annual fledged young (an index of the distribution of fecundity among nesting territories) and the mean reproductive outputs, that is the heterogeneous structure of the population determined that most/all pairs contributed to the annual production of young during good years, but the opposite during poor years (i.e. fewer pairs produced the majority of fledglings). Mean reproductive output was positively affected by percentage of open country and diet richness. Although other factors different to territory quality could affect demography parameters (e.g. quality of breeders), our results clearly showed a significant correlation between landscape features and population productivity.
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Acknowledgments
D. Becker, M. Forero, O. Krüger, M. Mönkkönen, N. L. Rodenhouse, T. Sota and three anonymous referees made a useful critique of the first draft of the manuscript. We thank P. Roche for helping with the IDRISI program, H. Magnin, C. Horisberger and P. Horisberger and O. Maubec for the logistic help during the fieldwork. During the study, V.P. received a research grant from the Regional Park of Luberon (France) and a post-doctoral grant from the Estación Biológica de Doñana (Consejo Superior de Investigaciones Científicas, Spain).
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Appendices
Appendix 1
Territory-specific yearly data of reproductive output (i.e. number of fledglings) of the French population of eagle owls during the 29 years of the study. The years 1973 and 1974 are absent because during this period it was impossible to collect data in the field (–, missing data). Mean number of fledglings, coefficient of reproduction (CV) and percentage of contributing pairs (%) per year was only presented for the 18 years shown in Fig. 1 and during which we were able to obtain data for at least ten territories each year (see details in Materials and methods)
Territories | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | 24 | 25 | 26 | 27 | 28 | 29 | 30 | 31 | 32 | 33 | 34 | 35 | x̄ | CV | Percentage |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1971 | – | – | – | – | – | – | – | – | 2 | 3 | 2 | 2 | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – |
1972 | – | – | – | – | – | – | – | – | 3 | 2 | 2 | 1 | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – |
1975 | – | – | – | – | – | – | – | – | 2 | 2 | 2 | 2 | 2 | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – |
1976 | – | – | – | – | – | – | – | – | – | 2 | 3 | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – |
1977 | 2 | – | – | 2 | – | – | – | – | 1 | 3 | 2 | 2 | – | – | – | – | – | 1 | – | – | 1 | – | – | – | – | – | 2 | – | – | – | – | – | – | – | – | – | – | – |
1978 | – | – | – | 1 | – | – | – | – | – | 1 | 2 | 2 | 2 | 1 | – | – | – | 2 | – | – | – | 2 | – | – | – | – | 1 | – | – | – | – | 3 | 2 | – | – | 1.7 | 37.4 | 37.5 |
1979 | 2 | – | – | 2 | 2 | 1 | 2 | – | 3 | 3 | 1 | – | 1 | – | 1 | – | – | 1 | – | 1 | – | – | 1 | – | – | – | 0 | – | 1 | 2 | 2 | 2 | – | – | – | 1.5 | 50.3 | 35.3 |
1980 | 1 | 2 | 1 | 2 | 2 | – | 2 | – | 2 | 2 | 2 | 2 | 2 | 2 | 2 | – | – | 1 | 1 | 2 | 1 | 2 | – | – | 1 | – | 2 | – | 2 | 3 | – | 3 | 1 | – | – | 1.8 | 32.8 | 41.6 |
1981 | – | 1 | – | 1 | – | 2 | 1 | – | 1 | 2 | 2 | 1 | 2 | – | 1 | – | – | 2 | – | – | – | – | 1 | 1 | 2 | – | 1 | – | 2 | 2 | 1 | 3 | 2 | 1 | – | 1.5 | 39.6 | 38.1 |
1982 | 1 | 1 | – | 2 | 2 | 2 | 2 | 1 | 3 | 3 | 2 | 1 | 3 | – | – | – | – | 0 | – | 1 | 2 | 2 | – | 2 | – | – | 0 | – | 2 | 3 | 2 | 2 | – | – | – | 1.8 | 49 | 38.1 |
1983 | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | 1 | – | 2 | – | – | – | – |
1984 | 2 | 2 | 2 | 1 | 2 | 1 | 2 | – | 2 | 2 | 3 | 2 | 2 | 1 | 2 | – | – | 2 | 2 | – | 1 | – | – | 1 | 2 | – | 2 | – | 1 | 1 | – | – | – | – | – | 1.7 | 31.8 | 40.1 |
1985 | 3 | 1 | – | – | – | 2 | 2 | – | 2 | 2 | 1 | 1 | 2 | – | – | – | – | 1 | 1 | – | – | – | – | 2 | – | 1 | – | – | 2 | 0 | – | 2 | – | – | – | 1.6 | 46.6 | 40 |
1986 | 2 | – | – | – | – | 1 | 0 | – | 0 | 2 | – | 0 | 2 | – | 1 | – | – | – | 0 | – | – | – | – | – | – | – | – | – | – | – | – | 1 | – | 1 | – | 0.9 | 91.3 | 27.3 |
1987 | 1 | 0 | – | 2 | 0 | 0 | 1 | – | 2 | 2 | – | – | 3 | – | – | – | – | 2 | – | – | 1 | – | – | 2 | – | – | 1 | – | – | 2 | – | 2 | – | – | – | 1.4 | 65 | 33.3 |
1988 | 2 | 1 | – | 2 | – | – | 2 | – | 1 | 3 | 2 | – | 2 | – | – | – | – | – | 1 | – | – | – | – | 0 | – | – | – | – | – | – | 2 | 2 | – | 2 | – | 1.7 | 44.4 | 38.5 |
1989 | – | – | – | – | – | – | – | – | – | 2 | – | – | 2 | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | 2 | – | – | – | – | – | – |
1990 | 0 | – | – | – | – | – | – | – | 0 | 0 | – | 0 | 2 | – | – | – | – | 2 | 2 | – | – | 1 | – | 2 | – | 1 | 1 | – | – | 2 | 1 | 2 | – | 2 | – | 1.2 | 71.8 | 33.3 |
1991 | 2 | 1 | – | – | 2 | 2 | 2 | – | 2 | 3 | 2 | 1 | 3 | – | – | – | – | – | – | – | – | – | 2 | 2 | – | – | – | – | 2 | 3 | – | 3 | – | – | – | 2.1 | 30 | 40 |
1992 | 2 | – | – | 2 | – | – | 1 | – | 2 | 0 | 2 | – | 0 | – | – | – | – | 2 | – | – | – | – | – | – | – | – | – | – | – | – | – | 2 | – | 2 | – | 1.5 | 56.6 | 40 |
1993 | 0 | – | – | 1 | – | – | 3 | – | 2 | 2 | 2 | – | 2 | – | – | 3 | – | – | – | – | – | – | – | 1 | – | – | – | – | – | – | – | 0 | – | – | – | 1.6 | 67.1 | 40 |
1994 | 1 | 1 | – | 2 | 2 | 2 | 3 | – | 3 | 2 | 2 | – | 2 | – | 2 | 2 | – | 1 | – | – | – | – | 1 | 2 | – | – | – | – | 2 | 3 | 1 | 2 | – | – | 1 | 1.9 | 36.2 | 40 |
1995 | – | – | – | – | – | – | – | – | 2 | 3 | – | – | 2 | – | – | 3 | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – |
1996 | 2 | 1 | – | 2 | – | – | 2 | 2 | 2 | 2 | 2 | 1 | 2 | – | 1 | 2 | – | 3 | – | – | – | – | – | 2 | – | 1 | – | 2 | 3 | – | 2 | 2 | – | – | – | 1.9 | 30 | 42.1 |
1997 | 2 | – | – | – | – | – | – | – | – | 0 | – | – | – | – | – | 2 | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – | – |
1998 | 2 | – | – | 2 | – | 2 | – | 2 | 2 | 2 | 1 | – | 2 | – | – | 3 | 2 | 2 | – | – | – | – | – | – | – | – | – | 1 | 2 | 3 | – | – | – | 2 | 2 | 2 | 25.8 | 43.8 |
1999 | 1 | – | – | 2 | – | – | 2 | 2 | 2 | 2 | 0 | 2 | 2 | – | 1 | 0 | 2 | 0 | – | – | – | – | – | – | – | – | – | – | 1 | 2 | – | – | – | – | 3 | 1.5 | 59.6 | 40 |
Appendix 2
Raw data for the eight variables of landscape structure, diet and density (see Materials and methods for details) that we used to explain differences in mean reproductive output and its annual variance within the population. The results of the two forward stepwise multiple regression models that we performed using mean reproductive output and CV as dependent variables are summarised in Table 1
Territoriesa | 1 | 2 | 4 | 6 | 7 | 9 | 10 | 11 | 12 | 13 | 16 | 18 | 19 | 29 | 30 | 32 | 34 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Percentage of open country | 19.4 | 29.4 | 30.3 | 26.1 | 29.6 | 22.3 | 60.5 | 38.4 | 49.5 | 74.5 | 43.1 | 31.5 | 12.5 | 55.1 | 53.1 | 45.1 | 26.9 |
Percentage of woodland | 38.2 | 37.2 | 41.8 | 59.3 | 58.5 | 68.1 | 20.7 | 50.2 | 28.7 | 9.3 | 22.8 | 27.1 | 49.6 | 10.6 | 12.8 | 22.8 | 31.6 |
Number of ecotones | 11.5 | 14.9 | 15.1 | 14.5 | 12.3 | 13.5 | 15.4 | 14.1 | 11.2 | 11.9 | 13.5 | 13.1 | 12.8 | 13.5 | 13.5 | 13.5 | 11.9 |
Shannon diversity index | 1.7 | 1.9 | 1.9 | 1.6 | 1.6 | 1.4 | 1.9 | 1.8 | 2 | 1.8 | 2 | 2 | 1.6 | 2 | 2 | 2 | 1.8 |
NND (m) | 800 | 780 | 1200 | 670 | 1,200 | 1,000 | 760 | 760 | 2,000 | 2,550 | 900 | 3,500 | 3,600 | 1,000 | 1,500 | 1,600 | 2,500 |
Diet richness | 70 | 70 | 30 | 68 | 69 | 70 | 70 | 70 | 30 | 31 | 68 | 32 | 32 | 65 | 69 | 69 | 32 |
Diet diversity | 2.3 | 2.3 | 1.9 | 2.3 | 2.3 | 2.2 | 2.2 | 2.1 | 1.9 | 1.9 | 2.1 | 1.9 | 1.9 | 2.3 | 2.3 | 2.3 | 1.9 |
Distance (m) from the nearest patch of open country | 40.7 | 27.5 | 0.0 | 0.0 | 0.0 | 159.8 | 0.0 | 27.5 | 0.0 | 0.0 | 27.5 | 55.0 | 27.5 | 22.5 | 27.5 | 12.7 | 87.0 |
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Penteriani, V., Delgado, M.M., Gallardo, M. et al. Spatial heterogeneity and structure of bird populations: a case example with the eagle owl. Popul Ecol 46, 185–192 (2004). https://doi.org/10.1007/s10144-004-0178-8
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DOI: https://doi.org/10.1007/s10144-004-0178-8