Conclusions
The present study demonstrated the strong parameter dependency of temporal beta-diversity patterns on the four fundamental parameters of the neutral model. The results of the present study caution that macroecological studies of temporal beta diversity are challenging without considering these four parameters in any case. Neutral dynamics is a starting point for discussing the real drivers of temporal beta-diversity patterns. In simulation studies, relaxing the zero-sum and neutrality assumptions would provide more realistic patterns comparable to empirical patterns (Jabot, 2010; Jabot & Chave, 2011; Munoz & Huneman, 2016; Takeuchi et al., 2022). Moving forward, two future study directions of temporal beta diversity, more generally biodiversity changes, are recommeded. One is toward the linking of biodiversity changes with the modern co-existence theory and other studies focusing on species co-existence, which would pave the way for a new field of study. For example, under variable climate conditions, different types of storage effects, which is a kind of mechanism facilitating species co-existence (Chesson & Warner, 1981; Chesson, 2000), would affect the apparent compositional changes; the persistence of long-lived individuals could contribute to decreasing the change in the short term, but the existence of buried seeds could contribute to less variability in community composition in the long term. The other is toward an explicit consideration of the spatial connections among the temporal beta-diversity patterns. Macroecological studies of temporal beta diversity often consider different sites as independent, but in reality, close distance sites are often connected. For example, animals move across sites and plants provide seeds to neighbouring sites (Howe & Smallwood, 1982; Bowler & Benton, 2005). For a deeper mechanistic understanding of temporal beta diversity, it is important to identify the direct effects through connections from simple synchronic responses to similar climate changes due to site closeness. Many future tasks remain to be accomplished in the study of temporal beta diversity, which is important in both basic and applied research.