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While at the academy, Mills initially played as a midfielder, but his playing position changed with the arrival of George Prost who was impressed with the way Mills "got on the ball and tried to dictate play" and switched him into a central–defence role, a position he plays today.
However, Mills remained out of the first team for almost half of the season until he finally made his Southampton debut, where he made his first start, in a 3–0 loss against Watford on 26 December 2005.
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Born in Swindon, Mills started his youth career with the academy of Swindon Town before switching to the Southampton Academy in 1999.
He turned professional in 2002 and had loan stints with Coventry City and Bournemouth during his time at the Saints. However, his playing time was limited due to injuries.
At the start of the 2006–07 season, Mills suffered an ankle injury, but recovered and made his first appearance of the season, in a 4–0 win over Wigan Athletic on 21 October 2006.
In order to gain first team football Mills expressed a desire to join Doncaster Rovers after winning promotion at Wembley, and was expected to sign in the close season.