Characterizing the Change of Annual Cone Production in Longleaf Pine Forests

Sporadic cone (or seed) production challenges longleaf pine forest restoration. Characterizing annual cone production change from long-term monitored data provides functional information on the foundational species of this imperiled ecosystem. In this study, permutation entropy (PE) and phase change were used to analyze longleaf pine annual cone production based on cone counts from four sites (Escambia Experimental Forest, Blackwater River State Forest, The Jones Center at Ichauway, and Sandhills State Forest). PE is an analytical tool to measure the complexity of a dynamic system while phase change characterizes the stage of self-organization. Results indicate that PE at each site was close to 1.0 (largely random changes in annual cone production) and generally increased with time. The positive association between the permutation entropy of cone production and average air temperature at different times was significant at two sites (Blackwater River State Forest and The Jones Center at Ichauway). The frequency distribution of phases (e.g., Poor (P), Fair (F), Good (G), Bumper (B)) in cone production followed negative power laws. Phase P could transition to any stage, but more than 50% remained in P across sites. For phase G, it would revert to P more than 50% of the time. Phase B would shift to P, except at Sandhills State Forest. The average lasting time of phase P was approximately 3.7 years. The overall relationship between the interval time of phase B and cone production was not statistically significant. Similarly, the overall relationship between cone production in phase B and the phase change times between consecutive B phases was not statistically significant. These results provide information on the ecological complexity of cone (or seed) production. Our methods can be helpful for estimating the occurrence of bumper cone (or seed) production, the lasting period between phase changes, and providing a tool for predicting natural regeneration potential over time for longleaf pine and other tree species (e.g., masting species).