Eleven (44%) of the 25 randomly selected candidates with outlier phenotypes (Arhgap30, Arrdc5, ClusterXN1, Cpgi81, Gsdme, Hpf1, Josd1, Mkrn2, Scaf11, Slamf9, Smg9) had no prior links to osteoarthritis or skeletal biology based on association with human disease and structured literature searching, whereas only two (20%) of the 10 prioritized candidates (Scaf11, Smg9) had no prior association (Fig. 1, Supplementary Data 7). Here, JOSD1 is linked to osteoarthritis.