Following the 1989 'Exxon Valdez' oil spill in Prince William Sound, Alaska, we studied the status of recovery of harlequin duck Histrionicus histrionicus populations during 1995 to 1998. We evaluated potential constraints on full recovery, including (1) exposure to residual oil; (2) food limita- tion; and (3) intrinsic demographic limitations on population growth rates. In this paper, we synthe- size the findings from our work and incorporate information from other harlequin duck research and monitoring programs to provide a comprehensive evaluation of the response of this species to the 'Exxon Valdez' spill. We conclude that harlequin duck populations had not fully recovered by 1998. Furthermore, adverse effects continued as many as 9 yr after the oil spill, in contrast to the conven- tional paradigm that oil spill effects on bird populations are short-lived. These conclusions are based on the findings that (1) elevated cytochrome P450 (CYP1A) induction on oiled areas indicated con- tinued exposure to oil in 1998; (2) adult female winter survival was lower on oiled than unoiled areas during 1995 to 1998; (3) fall population surveys by the Alaska Department of Fish and Game indi- cated numerical declines in oiled areas during 1995 to 1997; and (4) densities on oiled areas in 1996 and 1997 were lower than expected using models that accounted for effects of habitat attributes. Based on hypothesized links between oil contamination and demography, we suggest that harlequin duck population recovery was constrained primarily by continued oil exposure. Full population recovery will also be delayed by the time necessary for intrinsic population growth to allow return to pre-spill numbers following cessation of residual oil spill effects. Although not all wildlife species were affected by the 'Exxon Valdez' oil spill, and some others may have recovered quickly from any effects, harlequin duck life history characteristics and benthic, nearshore feeding habits make them susceptible to both initial and long-term oil spill effects.
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