A multi-level signal receiver includes a data sampler having (M−1) sense amplifiers therein, which are configured to compare a multi-level signal having one of M voltage levels with (M−1) reference voltages, to thereby generate (M−1) comparison signals. The data sampler is further configured to generate a target data signal including N bits, where M is an integer greater than two and N is an integer greater than one. An equalization controller is provided, which is configured to train the (M−1) sense amplifiers by: (i) adjusting at least one of (M−1) voltage intervals during a first training mode, and (ii) adjusting levels of the (M−1) reference voltages during a second training mode, based on equalized values of the (M−1) comparison signals, where each of the (M−1) voltage intervals represents a difference between two adjacent voltage levels from among the M voltage levels.