The voltage undershoot is one of the main dynamic problems of proton exchange membrane fuel cells (PEMFCs) under dynamic loadings. However, its mechanisms remain poorly understood due to its complex and coupled influencing factors. Besides, PEMFCs with a large active area are more prone to local dynamic failures for their pronounced in-plane (I-P) heterogeneity, while few studies have focused on the I-P heterogeneity of the dynamic response of large-area fuel cells. In this work, the mechanisms and the I-P heterogeneity of the voltage undershoot of a 320 cm2 PEMFC are thoroughly studied based on an in-situ monitoring method and a voltage loss decoupling method. Results show that the voltage overshoot is primarily driven by the ohmic loss (OL) overshoot and the concentration polarization (CP) overshoot. Results also prove that the OL overshoot is mainly caused by the instantaneous water shortage in the proton exchange membrane (PEM) which depends on the initial water content in the PEM. Besides, the CP overshoot is due to the instantaneous reactant shortage at the catalyst, and it is much sensitive to the change in the oxygen diffusion coefficient of the ionomer caused by the change in its water content. Meanwhile, both OL overshoot and CP overshoot demonstrate significant I-P heterogeneities under poor dynamic operating conditions, especially in cases of high load step and low inlet air humidity. Finally, the influences of operating conditions on the voltage undershoot and its I-P heterogeneity are discussed, and suggestions for improving the dynamic performance of PEMFCs are proposed.