Phylogeny and taxonomic revisions of Piper (Piperaceae) in Taiwan

Taiwan, located at the northernmost extent of Huxley's Line, marks the furthest point along this biogeographic boundary where exchanges of flora and fauna occur. The genus Piper, a globally distributed and taxonomically complex group of both economic and evolutionary importance, has a convoluted taxonomic history in this transitional zone that requires comprehensive study. Here, we integrate morphological, molecular, and distributional data to revise the taxonomy of Taiwanese Piper, resolve their evolutionary relationships, and infer their biogeographic history. Our revision recognizes 11 species in Taiwan, including seven native taxa, of which three are endemic. The present analysis establishes the first comprehensive phylogenetic framework for Taiwanese Piper and reveals that native species arose from lineages on both sides of Wallace's Line. Collectively, these findings provide the first evidence of bidirectional expansion within a genus in Taiwan and position Piper as an exemplary system for examining phylogeographic dynamics in transitional biogeographic zones.