Complete phylogeny of Micrathena spiders suggests multiple dispersal events among Neotropical rainforests, islands, and landmasses, and indicates Andean orogeny promotes speciation

Abstract

The Neotropical region is the most diverse on the planet, largely due to its mosaic of tropical rainforests. Multiple tectonic and climatic processes have been hypothesized to play a role in generating this diversity; these include Andean orogeny, closure of the Panama Isthmus, the hypothesized GAARlandia land bridge, and putative historical connections among currently isolated forests. Micrathena spiny spiders (Araneidae) include [~]120 species distributed mostly in forests from Mexico to Argentina, including the Antilles. Here, we use it as a model to study the biogeographic history of Neotropical rainforests by estimating a complete, dated phylogeny using morphological data for all species and molecular data for a subsample of 79 species. This resulted in a phylogeny that is mostly robust and supports most of the previously recognized species groups, although with uncertainty in the phylogenetic position of some species, especially those lacking sequences. The genus began diversifying about 25 million years ago. We use an event-based approach and biogeographic stochastic mapping to estimate ancestral distributions and the timing and direction of dispersal events, and to identify areas where diversity was generated, while accounting for phylogenetic uncertainty. Andean cloud forests generated the majority of species through in-situ speciation, but the Amazon was the major source of species for adjacent areas through dispersal; on the other extreme, the Dry Diagonal received species from other areas but generated very little diversity. Species exchange between Central and South America was intense, with [~]23 dispersal events beginning at least 20 million years ago, indicating that Micrathena dispersed between these continents before closure of the Isthmus of Panama. We also inferred [~]4 over-water dispersal events from Central and North America to the Antilles, which happened in the last 20 million years, and thus much after the proposed age of the GAARlandia land bridge. We identified an important species exchange route between the Amazon and the Atlantic forests. Sampling all species of the genus was fundamental to some of the conclusions above, especially in identifying the Andes as the area that generated the majority of species. This study highlights the importance of a solid and complete taxonomic sampling in biogeographic studies.