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Employee Stock Benefit Plans - Stock-based Compensation Expense by Financial Statement Line (Details) - USD ($)
$ in Thousands
12 Months Ended
Dec. 31, 2019
Dec. 31, 2018
Dec. 31, 2017
Share-based Compensation Arrangement by Share-based Payment Award [Line Items]      
Share-based compensation expense included in operating expenses $ 23,615 $ 28,701 $ 35,003
Research and development      
Share-based Compensation Arrangement by Share-based Payment Award [Line Items]      
Share-based compensation expense included in operating expenses 8,768 11,657 14,699
General and administrative      
Share-based Compensation Arrangement by Share-based Payment Award [Line Items]      
Share-based compensation expense included in operating expenses $ 14,847 $ 17,044 $ 20,304