Milhiser is a native of Belle Mead, New Jersey and he graduated from Montgomery High School in Skillman, New Jersey.
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In 2014 Milhiser will appear in the film Camp Takota with Youtube personalities, actresses and comediennes Grace Helbig, Hannah Hart and Mamrie Hart.
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He has been a regular performer at the Upright Citizens Brigade Theater since 2005, where he was a member of the sketch comedy group Serious Lunch, who have been featured on Late Night with Jimmy Fallon and Attack of the Show.
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Mooney, along with John Milhiser, Michael Patrick O'Brien, Noël Wells, Brooks Wheelan, and his comedy partner and friend, Beck Bennett, joined the thirty-ninth season of Saturday Night Live to fill the void left by the recent departures of Fred Armisen, Bill Hader, and Jason Sudeikis.
Serious Lunch members John Milhiser and Bob Officer appeared on Late Night with Jimmy Fallon Thursday, March 18 (s.1 : ep.211) to set the world record for touching noses.