Status and trends in demography of northern spotted owls, 1985-2003
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 (??), and annual rate of population change (??) 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 (x??=0.372, SE=0.029), lower for 2-year-olds (x??=0.208, SE=0.032), and very low for 1-year-olds (x??=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 (x??=0.560, SE=0.041) and lowest in the Douglas-fir (Pseudotsuga menziesii) region of the Oregon coast (x??=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 rate