Holt et al. (Report, 4 January 2013, p. 74) propose substantial modifications of Wallace's long-standing zoogeographic regions based on clustering of a pairwise similarity matrix of vertebrate assemblages. We worry about their compromised use of phylogenies and show that a fundamental point of their analysis-i.e., the delineation of new realms-is only weakly supported by their results and conceptually flawed.
Dividing the world into regions of similar faunistic or floristic composition and shared evolutionary history is a major aim of biogeography. The most prominent regionalization, Wallace's zoogeographic regions (1), has had tremendous influence, but its expert-based nature and lack of quantitative rigor and reproducibility have led to ongoing debates about the number and delineation of regions [reviewed in (2, 3)]. In their Report, Holt et al. (4) follow recently developed methodology and results for mammals (3) and present a quantitative global regionalization that extends to birds and amphibians. Instead of pairwise turnover of species (or genera or families), they used the number of branches in variably resolved phylogenies shared between two assemblages. The integration of global range maps and phylogenetic information to delineate biogeographical regions is promising because it may overcome problems associated with mixed results reported from studies at the species, genus, and family levels (3, 5). However, we believe that the key results of their study-i.e., an integrative delineation of the world's main biogeographical regions and the proposal of five new realms-are largely based on problematic data and methodology, as well as subjective decisions, and thus may have several conceptual flaws.First, we are concerned about the overall poor and taxonomically and geographically disparate resolution of the tree topologies invoked. These were single trees in which only 40 to 60% of species were resolved, meaning that species' contributions to the regionalization were highly nonuniform and, in some cases, very minor.Second, only counts of branches, not their actual lengths, were used for quantifying dissimilarity. This will in some cases equate species 20 million years old with those just 20,000 years old and ignores considerable differences in the ages of clades (e.g., amphibians versus birds) and regions. The authors suggest that in mammals, such differences, or even using just species' taxonomic separation instead, would not affect the existence and location of zoogeographic regions. We feel that this casts doubt, in this implementation, on the rigor and relevance of the "phylogenetic" method. New approaches are arising that account for all species in a single quantitative framework and that estimate branch lengths and uncertainty (6, 7). In the midst of such major phylogenetic advances for vertebrates, we support caution over a rush in updating Wallace's regions.Third, the authors propose the Saharo-Arabian, Sino-Japanese, and Panamanian as new "realms," even though they do not match their criterion of ...