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Stock-Based Compensation - Expense (Details) - USD ($)
$ in Thousands
3 Months Ended 9 Months Ended
Sep. 30, 2017
Sep. 30, 2016
Sep. 30, 2017
Sep. 30, 2016
Stock-based compensation expense        
Total stock-based compensation expense $ 2,739 $ 3,796 $ 9,728 $ 11,709
Research and development        
Stock-based compensation expense        
Total stock-based compensation expense 1,546 2,042 5,310 5,866
General and administrative        
Stock-based compensation expense        
Total stock-based compensation expense $ 1,193 $ 1,754 $ 4,418 $ 5,843