Spatial Congruence Analysis (SCAN): A method for detecting biogeographical patterns based on species range congruences

Species with congruent geographical distributions, potentially caused by common historical and ecological spatial processes, constitute biogeographical units called chorotypes. Nevertheless, the degree of spatial range congruence characterizing these groups of species is rarely used as an explicit parameter. Methods conceived for the identification of patterns of shared ranges often suffer from scale bias associated with the use of grids, or the incapacity to describe the full complexity of patterns, from core areas of high spatial congruence, to long gradients of range distributions expanding from these core areas. Here, we propose a simple analytical method, Spatial Congruence Analysis (SCAN), which identifies chorotypes by mapping direct and indirect spatial relationships among species. Assessments are made under a referential value of congruence as an explicit numerical parameter. A one-layered network connects species (vertices) using pairwise spatial congruence estimates (edges). This network is then analyzed for each species, separately, by an algorithm which searches for spatial relationships to the reference species. The method was applied to two datasets: a simulated gradient of ranges and real distributions of birds. The simulated dataset showed that SCAN can describe gradients of distribution with a high level of detail. The bird dataset showed that only a small portion of range overlaps is biogeographically meaningful, and that there is a large variation in types of patterns that can be found with real distributions. Species analyzed separately may converge on similar or identical groups, may be nested in larger chorotypes, or may even generate overlapped patterns with no species in common. Chorotypes can vary from simple ones, composed by few highly congruent species, to complex, with numerous alternative component species and spatial configurations, which offer insights about possible processes driving these patterns in distinct degrees of spatial congruence. Metrics such as congruence, depth, richness, and ratio between common and total areas can be used to describe chorotypes in detail, allowing comparisons between patterns across regions and taxa.