title: The comparative phylogeography of fruit bats of the tribe Scotonycterini (Chiroptera, Pteropodidae) reveals cryptic species diversity related to African Pleistocene forest refugia.
abstract: The hypothesis of Pleistocene forest refugia was tested using comparative phylogeography of Scotonycterini, a fruit bat tribe endemic to Africa containing four species: Scotonycteris zenkeri, Casinycteris argynnis, C. campomaanensis, and C. ophiodon. Patterns of genetic structure were assessed using 105 Scotonycterini (including material from three holotypes) collected at 37 localities, and DNA sequences from the mitochondrial cytochrome b gene (1140 nt) and 12 nuclear introns (9641 nt). Phylogenetic trees and molecular dating were inferred by Bayesian methods. Multilocus analyses were performed using supermatrix, SuperTRI, and *BEAST approaches. Mitochondrial analyses reveal strong phylogeographical structure in Scotonycteris, with four divergent haplogroups (4.9-8.7%), from Upper Guinea, Cameroon, western Equatorial Africa, and eastern Democratic Republic of the Congo (DRC). In C. argynnis, we identify two mtDNA haplogroups corresponding to western and eastern Equatorial Africa (1.4-2.1%). In C. ophiodon, the mtDNA haplotypes from Cameroon and Ivory Coast differ by only 1.3%. Nuclear analyses confirm the validity of the recently described C. campomaanensis and indicate that western and eastern populations of C. argynnis are not fully isolated. All mtDNA clusters detected in Scotonycteris are found to be monophyletic based on the nuclear dataset, except in eastern DRC. In the nuclear tree, the clade from western Equatorial Africa is closely related to individuals from eastern DRC, whereas in the mitochondrial tree it appears to be the sister-group of the Cameroon clade. Migrate-n analyses support gene flow from western Equatorial Africa to eastern DRC. Molecular dating indicates that Pleistocene forest refugia have played an important role in shaping the evolution of Scotonycterini, with two phases of allopatric speciation at approximately 2.7 and 1.6 Mya, resulting from isolation in three main forest areas corresponding to Upper Guinea, Cameroon, and Equatorial Africa. Two cryptic species and two subspecies are described herein in the genus Scotonycteris. Female philopatry and male biased dispersal are supported for the smallest taxa, i.e., the three species of Scotonycteris and C. argynnis. The Congo, Ntem, and Sanaga rivers are identified as biogeographic barriers to the dispersal of Scotonycteris during interglacial periods. A greater capacity for long-distance dispersal is inferred for the largest species, C. ophiodon.
authors: Sorbonne Universités, Université Paris-6 (UPMC), UFR927, 75005 Paris, France; Muséum national d'Histoire naturelle, ISYEB UMR 7205, CP51, 75005 Paris, France; Muséum national d'Histoire naturelle, UMS 2700, 75005 Paris, France. Electronic address: hassanin@mnhn.fr.
authors: Alexandre
authors: A
authors: Hassanin
authors: Muséum national d'Histoire naturelle, ISYEB UMR 7205, CP51, 75005 Paris, France; Muséum national d'Histoire naturelle, UMS 2700, 75005 Paris, France.
authors: Souraya
authors: S
authors: Khouider
authors: Université de Kisangani, Faculté des Sciences, BP 2012, Kisangani, Democratic Republic of the Congo.
authors: Guy-Crispin
authors: GC
authors: Gembu
authors: Field Museum of Natural History, 1400, South Lake Shore Drive, IL 60605, Chicago, USA.
authors: Steven M
authors: SM
authors: Goodman
authors: Université Felix-Houphouet-Boigny, UFR Biosciences, 22, BP 582 Abidjan 22, République de Côte d'Ivoire, Côte d'Ivoire.
authors: Blaise
authors: B
authors: Kadjo
authors: Muséum national d'Histoire naturelle, ISYEB UMR 7205, CP51, 75005 Paris, France; Muséum national d'Histoire naturelle, UMS 2700, 75005 Paris, France.
authors: Nicolas
authors: N
authors: Nesi
authors: Centre international de recherches médicales de Franceville, BP 769, Franceville, Gabon.
authors: Xavier
authors: X
authors: Pourrut
authors: Institut Pasteur de Bangui, BP 923, Bangui, Central African Republic.
authors: Emmanuel
authors: E
authors: Nakouné
authors: Muséum national d'Histoire naturelle, ISYEB UMR 7205, CP51, 75005 Paris, France; Muséum national d'Histoire naturelle, UMS 2700, 75005 Paris, France.
authors: Céline
authors: C
authors: Bonillo
mesh_terms: D000349:Africa; D000818:Animals; D001483:Base Sequence; D001499:Bayes Theorem; D002685:Chiroptera / Q000145:classification* / Q000235:genetics; D045303:Cytochromes b / Q000235:genetics; D004272:DNA, Mitochondrial / Q000235:genetics*; D005260:Female; D065928:Forests; D051456:Gene Flow; D006239:Haplotypes; D007438:Introns / Q000235:genetics; D008297:Male; D010802:Phylogeny; D058974:Phylogeography; D013045:Species Specificity
chemical_list: D004272:DNA, Mitochondrial; D045303:Cytochromes b
keywords: African mammals; Ancient DNA; Casinycteris; New species; Phylogeny; Quaternary; Rainforests; Refuge theory; Scotonycteris; Taxonomy