Scale effect and bimodality in the frequency distribution of species occupancy

Community patterns in species-by-site matrices provide valuable clues for inferring ecological processes at work. One such pattern is the occupancy frequency distribution (OFD) depicting the frequency distribution of row sums (i.e., occupancy) with a quarter OFDs of bimodal forms. Another pattern that also reflects the structure of row sums is the ranked species occupancy curve (RSOC), and has been shown to imply a 50% of bimodality in OFDs. The use of RSOCs has been advocated in literature over the OFD based on two conclusions from a 6-model inference using only 24 matrices: (i) RSOCs have two general forms, with half representing bimodal OFDs; (ii) there are no effects of spatial and study scales on RSOCs of different forms. Using a much more representative dataset of 289 matrices, I cast doubt on these two conclusions. A missing but dominant RSOC model (the truncated power law) is added. The number of species and the nestedness of the community differ significantly among matrices of different RSOC forms; however, the number of sites and the taxa in the studies do not differ among RSOC or OFD forms. The quarter OFDs of bimodal forms is reassured, with the least frequent occupancy consistent with Raunkiaer's law of frequency. Importantly, a RSOC is mathematically transferrable to an OFD, with the derivative of the occupancy ranking curve being equal to the negative reciprocal of the occupancy frequency. Based on the type of the community (null versus interactive) and site environment (homogenous versus heterogeneous), four scenarios are needed to identify pre-inferring assemblage mechanisms. The results highlight the need for shifting research from the emphasis of marginal sums to the analysis of matrix structure for an in-depth understanding of the community assemblage patterns and mechanisms.