title: Historical biogeography of Reticulitermes termites (Isoptera: Rhinotermitidae) inferred from analyses of mitochondrial and nuclear loci.
abstract: Termites of the genus Reticulitermes are ecologically and economically important wood-feeding social insects that are widespread in the Holarctic region. Despite their importance, no study has yet attempted to reconstruct a global time-scaled phylogeny of Reticulitermes termites. In this study, we sequenced mitochondrial (2096bp) and nuclear (829bp) loci from 61 Reticulitermes specimens, collected across the genus' entire range, and one specimen of Coptotermes formosanus, which served as an outgroup. Bayesian and Maximum likelihood analyses conducted on the mitochondrial and nuclear sequences support the existence of four main lineages that span four global geographical regions: North America (NA lineage), western Europe (WE lineage), a region including eastern Europe and western Asia (EA+WA lineage), and eastern Asia (EA lineage). The mitochondrial data allowed us to clarify the phylogenetic relationships among these lineages. They were also used to infer a chronogram that was time scaled based on age estimates for termite fossils (including the oldest Reticulitermes fossils, which date back to the late Eocene-early Oligocene). Our results support the hypothesis that the extant Reticulitermes lineage first differentiated in North America. The first divergence event in the ancestral lineage of Reticulitermes occurred in the early Miocene and separated the Nearctic lineages (i.e., the NA lineages) from the Palearctic lineages (i.e., WE, EE+WA, and EA lineages). Our analyses revealed that the main lineages of Reticulitermes diversified because of vicariance and migration events, which were probably induced by major paleogeographic and paleoclimatic changes that occurred during the Cenozoic era. This is the first global and comprehensive phylogenetic study of Reticulitermes termites, and it provides a crucial foundation for studying the evolution of phenotypic and life-history traits in Reticulitermes. For instance, the phylogeny we obtained suggested that 'asexual queen succession', a unique reproductive system, independently evolved at least three times during the diversification of the genus.
authors.affiliation: Institut de Recherche sur la Biologie de l'Insecte, UMR 7261, Université François Rabelais de Tours, Parc Grandmont, 37200 Tours, France. Electronic address: franck.dedeine@univ-tours.fr.
authors.forename: Franck
authors.initials: F
authors.lastname: Dedeine
authors.affiliation: Institut de Recherche sur la Biologie de l'Insecte, UMR 7261, Université François Rabelais de Tours, Parc Grandmont, 37200 Tours, France. Electronic address: simon.dupont@univ-tours.fr.
authors.forename: Simon
authors.initials: S
authors.lastname: Dupont
authors.affiliation: Institut de Recherche sur la Biologie de l'Insecte, UMR 7261, Université François Rabelais de Tours, Parc Grandmont, 37200 Tours, France. Electronic address: sylv.guyot@orange.fr.
authors.forename: Sylvain
authors.initials: S
authors.lastname: Guyot
authors.affiliation: Laboratory of Insect Ecology, Graduate School of Agriculture, Kyoto University, Oiwakecho, Kitashirakawa, Sakyo-ku, Kyoto 606-8502, Japan. Electronic address: kenjijpn@kais.kyoto-u.ac.jp.
authors.forename: Kenji
authors.initials: K
authors.lastname: Matsuura
authors.affiliation: Department of Entomology, Rutgers University, New Brunswick, NJ 08901, USA. Electronic address: cwang@aesop.rutgers.edu.
authors.forename: Changlu
authors.initials: C
authors.lastname: Wang
authors.affiliation: Department of Plant Protection, College of Agriculture, University of Shahid-Chamran, Ahwaz, Iran. Electronic address: habibpour_b@scu.ac.ir.
authors.forename: Behzad
authors.initials: B
authors.lastname: Habibpour
authors.affiliation: Institut de Recherche sur la Biologie de l'Insecte, UMR 7261, Université François Rabelais de Tours, Parc Grandmont, 37200 Tours, France. Electronic address: bagneres@univ-tours.fr.
authors.forename: Anne-Geneviève
authors.initials: AG
authors.lastname: Bagnères
authors.affiliation: Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Università di Bologna, via Selmi 3, 40126 Bologna, Italy. Electronic address: barbara.mantovani@unibo.it.
authors.forename: Barbara
authors.initials: B
authors.lastname: Mantovani
authors.affiliation: Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Università di Bologna, via Selmi 3, 40126 Bologna, Italy. Electronic address: andrea.luchetti@unibo.it.
authors.forename: Andrea
authors.initials: A
authors.lastname: Luchetti
mesh_terms: D000818:Animals; D005075:Biological Evolution; D002467:Cell Nucleus; D004272:DNA, Mitochondrial; D017344:Genes, Insect; D049810:Genetic Speciation; D020049:Isoptera / Q000145:classification* / Q000235:genetics; D010802:Phylogeny; D058974:Phylogeography
chemical_list: D004272:DNA, Mitochondrial
keywords: Asexual queen succession; Continental vicariance; Divergence times; Reproductive system; Social insects; Time-scaled phylogeny