title: Into the Andes: multiple independent colonizations drive montane diversity in the Neotropical clearwing butterflies Godyridina.
abstract: Understanding why species richness peaks along the Andes is a fundamental question in the study of Neotropical biodiversity. Several biogeographic and diversification scenarios have been proposed in the literature, but there is confusion about the processes underlying each scenario, and assessing their relative contribution is not straightforward. Here, we propose to refine these scenarios into a framework which evaluates four evolutionary mechanisms: higher speciation rate in the Andes, lower extinction rates in the Andes, older colonization times and higher colonization rates of the Andes from adjacent areas. We apply this framework to a species-rich subtribe of Neotropical butterflies whose diversity peaks in the Andes, the Godyridina (Nymphalidae: Ithomiini). We generated a time-calibrated phylogeny of the Godyridina and fitted time-dependent diversification models. Using trait-dependent diversification models and ancestral state reconstruction methods we then compared different biogeographic scenarios. We found strong evidence that the rates of colonization into the Andes were higher than the other way round. Those colonizations and the subsequent local diversification at equal rates in the Andes and in non-Andean regions mechanically increased the species richness of Andean regions compared to that of non-Andean regions ('species-attractor' hypothesis). We also found support for increasing speciation rates associated with Andean lineages. Our work highlights the importance of the Andean slopes in repeatedly attracting non-Andean lineages, most likely as a result of the diversity of habitats and/or host plants. Applying this analytical framework to other clades will bring important insights into the evolutionary mechanisms underlying the most species-rich biodiversity hotspot on the planet.
authors.affiliation: Institut de Systématique, Évolution, Biodiversité, ISYEB - UMR 7205 - CNRS MNHN UPMC EPHE, Muséum national d'Histoire naturelle, Sorbonne Universités, 57 rue Cuvier CP50, F-75005, Paris, France.|Department of Biology, University of Lund, 223 62, Lund, Sweden.
authors.forename: Nicolas
authors.initials: N
authors.lastname: Chazot
authors.affiliation: McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, FL, 32611, USA.
authors.forename: Keith R
authors.initials: KR
authors.lastname: Willmott
authors.affiliation: CNRS, UMR 5554 Institut des Sciences de l'Evolution (Université de Montpellier), Place Eugène Bataillon, 34095, Montpellier, France.|Department of Biological Sciences, University of Alberta, T6G 2E9, Edmonton, AB, Canada.
authors.forename: Fabien L
authors.initials: FL
authors.lastname: Condamine
authors.affiliation: Institut de Systématique, Évolution, Biodiversité, ISYEB - UMR 7205 - CNRS MNHN UPMC EPHE, Muséum national d'Histoire naturelle, Sorbonne Universités, 57 rue Cuvier CP50, F-75005, Paris, France.
authors.forename: Donna Lisa
authors.initials: DL
authors.lastname: De-Silva
authors.affiliation: Departamento de Zoologia and Museu de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, Campinas, São Paulo, Brazil.
authors.forename: André V L
authors.initials: AV
authors.lastname: Freitas
authors.affiliation: Museo de Historia Natural, Universidad Nacional de San Marcos, Lima, Peru.
authors.forename: Gerardo
authors.initials: G
authors.lastname: Lamas
authors.affiliation: IBENS, Ecole Normale Supérieure, UMR 8197 CNRS, Paris, France.
authors.forename: Hélène
authors.initials: H
authors.lastname: Morlon
authors.affiliation: Grupo de Investigación de Sanidad Vegetal, Universidad Católica de Oriente, Rionegro, Antioquia, Colombia.
authors.forename: Carlos E
authors.initials: CE
authors.lastname: Giraldo
authors.affiliation: Department of Zoology, University of Cambridge, Cambridge, UK.
authors.forename: Chris D
authors.identifier: http://orcid.org/0000-0002-7809-062X
authors.initials: CD
authors.lastname: Jiggins
authors.affiliation: Centre d'Ecologie Fonctionnelle et Evolutive, CEFE, UMR 5175 CNRS - EPHE - Université de Montpellier - Université Paul Valéry Montpellier, 34293, Montpellier 5, France.
authors.forename: Mathieu
authors.initials: M
authors.lastname: Joron
authors.affiliation: Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA, 02138, USA.
authors.forename: James
authors.initials: J
authors.lastname: Mallet
authors.affiliation: Universidad Nacional de Colombia, sede Medellín, Medellín, Colombia.
authors.forename: Sandra
authors.initials: S
authors.lastname: Uribe
authors.affiliation: Institut de Systématique, Évolution, Biodiversité, ISYEB - UMR 7205 - CNRS MNHN UPMC EPHE, Muséum national d'Histoire naturelle, Sorbonne Universités, 57 rue Cuvier CP50, F-75005, Paris, France.
authors.forename: Marianne
authors.initials: M
authors.lastname: Elias
mesh_terms: D000818:Animals; D044822:Biodiversity; D002080:Butterflies / Q000235:genetics*; D017753:Ecosystem; D049810:Genetic Speciation; D010802:Phylogeny; D013020:South America
keywords: Andes; Godyridina; Ithomiini; Lepidoptera; Neotropics; biogeography; trait-dependent diversification