Status and trends in demography of northern spotted owls, 1985-2003

Robert G. Anthony, Eric D. Forsman, Alan B. Franklin, David R. Anderson, Kenneth P. Burnham, Gary C. White, Carl J. Schwarz, James D. Nichols, James E. Hines, Gail S. Olson, Steven H. Ackers, Lawrence S. Andrews, Brian L. Biswell, Peter C. Carlson, Lowell V. Diller, Katie M. Dugger, Katherine E. Fehring, Tracy L. Fleming, Richard P. Gerhardt, Scott A. GremelR. J. Gutiérrez, Patti J. Happe, Dale R. Herter, J. Mark Higley, Robert B. Horn, Larry L. Irwin, Peter J. Loschl, Janice A. Reid, Stan G. Sovern

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126 Scopus citations


We analyzed demographic data from northern spotted owls (Strix occidentalis caurina) from 14 study areas in Washington, Oregon, and California for 1985-2003. The purpose of our analyses was to provide an assessment of the status and trends of northern spotted owl populations throughout most of their geographic range. The 14 study areas made up approximately 12% of the range of the subspecies and included federal, tribal, private, and mixed federal and private lands. The study areas also included all the major forest types that the subspecies inhabits. The analyses followed rigorous protocols that were developed a priori and were the result of extensive discussions and consensus among the authors. Our primary objectives were to estimate fecundity, apparent survival (φ), and annual rate of population change (λ) and to determine if there were any temporal trends in these population parameters. In addition to analyses of data from individual study areas, we conducted 2 meta-analyses on each demographic parameter. One meta-analysis was conducted on all 14 areas, and the other was restricted to the 8 areas that constituted the Effectiveness Monitoring Plan for northern spotted owls under the Northwest Forest Plan. The average number of years of reproductive data per study area was 14 (range = 5-19), and the average number of recapture occasions per study area was 13 (range = 4-18). Only 1 study area had <12 years of data. Our results were based on 32,054 captures and resightings of 11,432 banded individuals for estimation of survival and 10,902 instances in which we documented the number of young produced by territorial females. The number of young fledged (NYF) per territorial female was analyzed by testing a suite of a priori models that included (1) effects of age, (2) linear or quadratic time trends, (3) presence of barred owls (Strix varia) in spotted owl territories, and (4) an even-odd year effect. The NYF varied among years on most study areas with a biennial cycle of high reproduction in even-numbered years and low reproduction in odd-numbered years. These cyclic fluctuations did not occur on all study areas, and the even-odd year effect waned during the last 5 years of the study. Fecundity was highest for adults (x̄=0.372, SE=0.029), lower for 2-year-olds (x̄=0.208, SE=0.032), and very low for 1-year-olds (x̄=0.074, SE = 0.029). Fecundity was stable over time for 6 areas (Rainier, Olympic, Warm Springs, H. J. Andrews, Klamath, and Marin), declining for 6 areas (Wenatchee, Cle Elum, Oregon Coast Range, Southern Oregon Cascades, Northwest California, and Simpson), and slightly increasing for 2 areas (Tyee, Hoopa). We found little association between NYF and the proportion of northern spotted owl territories where barred owls were detected, although results were suggestive of a negative effect of barred owls on the Wenatchee and Olympic study areas. The meta-analysis on fecundity indicated substantial annual variability with no increasing or decreasing trends. Fecundity was highest in the mixed-conifer region of eastern Washington (x̄=0.560, SE=0.041) and lowest in the Douglas-fir (Pseudotsuga menziesii) region of the Oregon coast (x̄=0.306, SE=0.039). We used Cormack-Jolly-Seber open population models and Program MARK to estimate apparent survival rates of owls >1 year old. We found no differences in apparent survival rates between sexes except for 1 area (Marin), which had only 6 years of data. Estimates of apparent survival from individual study areas indicated that there were differences among age classes with adults generally having higher survival than 1- and 2-year-olds. Apparent survival rates ranged from 0.750 (SE=0.026) to 0.886 (SE=0.010) for adults, 0.626 (SE=0.073) to 0.886 (SE=0.010) for 2-year-olds, and 0.415 (SE=0.111) to 0.860 (SE=0.017) for 1-year-olds. These estimates were comparable to survival rates from previous studies on the subspecies. We found evidence for negative time trends in survival rates on 5 study areas (Wenatchee, Cle Elum, Rainier, Olympic, and Northwest California) and no trends in survival on the remaining areas. There was evidence for negative effects of barred owls on apparent survival on 3 study areas (Wenatchee, Cle Elum, and Olympic). Survival rates of adult owls on the 8 Monitoring Areas generally were high, ranging from 0.85 (SE=0.009) to 0.89 (SE=0.010), but were declining on the Cle Elum, Olympic, and Northwestern California study areas. The meta-analysis of apparent survival indicated differences among regions and changes overtime with a downward trend in the mixed-conifer and Douglas-fir regions of Washington. The meta-analysis of apparent survival also indicated that there was a negative association between fecundity and survival the following year, suggesting a cost of reproduction on survival. This effect was limited to the Douglas-fir and mixed-conifer regions of Washington and the Douglas-fir region of the Oregon Cascade Mountains. We used the reparameterized Jolly-Seber method (λRJs) to estimate annual rate of population change of territorial owls in the study areas. This estimate answers the question. Are these territorial owls being replaced in this geographically open population? Point estimates of λRJS were <1.0 for 12 of 13 study areas. The analyses provided strong evidence that populations on the Wenatchee, Cle Elum, Rainier, Olympic, Warm Springs, H. J. Andrews, Oregon Coast Ranges, and Simpson study areas were declining during the study. The mean λRJS for the 13 study areas was 0.963 (SE=0.009), suggesting that populations over all the areas were declining about 3.7% per year during the study. The mean λRJS for the 8 monitoring areas for the Northwest Forest Plan was 0.976 (SE=0.007) compared to a mean of 0.942 (SE=0.016) for the other study areas, a 2.4-versus-5.8% decline per year. This suggested that owl populations on federal lands had higher demographic rates than elsewhere; thus, the Northwest Forest Plan appeared to have a positive effect on demography of northern spotted owls. Populations were doing poorest in Washington, where apparent survival rates and populations were declining on all 4 study areas. Our estimates of λRJS were generally lower than those reported in a previous analysis (λ RJS=0.997, SE=0.003) for many of the same areas at an earlier date. The possible causes of population declines include but are not limited to habitat loss from timber harvest and fires, competition with barred owls, and weather patterns.

Original languageEnglish (US)
Pages (from-to)1-48
Number of pages48
JournalWildlife Monographs
Issue number163
StatePublished - Aug 2006


  • Annual rate of population change
  • Capture-resighting
  • Demography
  • Fecundity
  • Northern spotted owls
  • Strix occidentalis caurina
  • Survival rates


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