title: A dated molecular phylogeny of manta and devil rays (Mobulidae) based on mitogenome and nuclear sequences.
abstract: Manta and devil rays are an iconic group of globally distributed pelagic filter feeders, yet their evolutionary history remains enigmatic. We employed next generation sequencing of mitogenomes for nine of the 11 recognized species and two outgroups; as well as additional Sanger sequencing of two mitochondrial and two nuclear genes in an extended taxon sampling set. Analysis of the mitogenome coding regions in a Maximum Likelihood and Bayesian framework provided a well-resolved phylogeny. The deepest divergences distinguished three clades with high support, one containing Manta birostris, Manta alfredi, Mobula tarapacana, Mobula japanica and Mobula mobular; one containing Mobula kuhlii, Mobula eregoodootenkee and Mobula thurstoni; and one containing Mobula munkiana, Mobula hypostoma and Mobula rochebrunei. Mobula remains paraphyletic with the inclusion of Manta, a result that is in agreement with previous studies based on molecular and morphological data. A fossil-calibrated Bayesian random local clock analysis suggests that mobulids diverged from Rhinoptera around 30 Mya. Subsequent divergences are characterized by long internodes followed by short bursts of speciation extending from an initial episode of divergence in the Early and Middle Miocene (19-17 Mya) to a second episode during the Pliocene and Pleistocene (3.6 Mya - recent). Estimates of divergence dates overlap significantly with periods of global warming, during which upwelling intensity - and related high primary productivity in upwelling regions - decreased markedly. These periods are hypothesized to have led to fragmentation and isolation of feeding regions leading to possible regional extinctions, as well as the promotion of allopatric speciation. The closely shared evolutionary history of mobulids in combination with ongoing threats from fisheries and climate change effects on upwelling and food supply, reinforces the case for greater protection of this charismatic family of pelagic filter feeders.
authors.affiliation: Department of Marine Benthic Ecology and Evolution, Centre for Ecological and Evolutionary Studies, University of Groningen, Nijenborgh 7, 9747 AG Groningen, The Netherlands; Department of Ecology & Evolutionary Biology, University of California Santa Cruz, 100 Shaffer Road, Santa Cruz, CA 95060, USA; Faculty of Biosciences and Aquaculture, Universitetet i Nordland, Universitetsalleen 11, 8049 Bodø, Norway. Electronic address: m.poortvliet@rug.nl.
authors.forename: Marloes
authors.initials: M
authors.lastname: Poortvliet
authors.affiliation: Department of Marine Benthic Ecology and Evolution, Centre for Ecological and Evolutionary Studies, University of Groningen, Nijenborgh 7, 9747 AG Groningen, The Netherlands. Electronic address: J.L.Olsen@rug.nl.
authors.forename: Jeanine L
authors.initials: JL
authors.lastname: Olsen
authors.affiliation: Department of Ecology & Evolutionary Biology, University of California Santa Cruz, 100 Shaffer Road, Santa Cruz, CA 95060, USA. Electronic address: dcroll@ucsc.edu.
authors.forename: Donald A
authors.initials: DA
authors.lastname: Croll
authors.affiliation: Department of Ecology & Evolutionary Biology, University of California Santa Cruz, 100 Shaffer Road, Santa Cruz, CA 95060, USA. Electronic address: bernardi@ucsc.edu.
authors.forename: Giacomo
authors.initials: G
authors.lastname: Bernardi
authors.affiliation: Department of Ecology & Evolutionary Biology, University of California Santa Cruz, 100 Shaffer Road, Santa Cruz, CA 95060, USA. Electronic address: newton@ucsc.edu.
authors.forename: Kelly
authors.initials: K
authors.lastname: Newton
authors.affiliation: Faculty of Biosciences and Aquaculture, Universitetet i Nordland, Universitetsalleen 11, 8049 Bodø, Norway. Electronic address: Spyros.Kollias@uin.no.
authors.forename: Spyros
authors.initials: S
authors.lastname: Kollias
authors.affiliation: Monterey Bay Aquarium, 886 Cannery Row, Monterey, CA 93940, USA. Electronic address: JOSullivan@mbayaq.org.
authors.forename: John
authors.initials: J
authors.lastname: O'Sullivan
authors.affiliation: Department of Biology and Environmental Science, Linnaeus University, SE 39182 Kalmar, Sweden; The Manta Trust, Catemwood House, Corscombe, Dorchester, Dorset DT2 0NT, United Kingdom. Electronic address: daniel@mantatrust.org.
authors.forename: Daniel
authors.initials: D
authors.lastname: Fernando
authors.affiliation: The Manta Trust, Catemwood House, Corscombe, Dorchester, Dorset DT2 0NT, United Kingdom. Electronic address: guy@mantatrust.org.
authors.forename: Guy
authors.initials: G
authors.lastname: Stevens
authors.affiliation: Centro Interdisciplinario de Ciencias Marinas, Instituto Politécnico Nacional, Av. IPN s/n, La Paz, Baja California Sur 23096, Mexico. Electronic address: galvan.felipe@gmail.com.
authors.forename: Felipe
authors.initials: F
authors.lastname: Galván Magaña
authors.affiliation: Muséum national d'Histoire naturelle, Département Systématique et Evolution, CP 51, rue Buffon, 75231 Paris cedex 05, France. Electronic address: seret@mnhn.fr.
authors.forename: Bernard
authors.initials: B
authors.lastname: Seret
authors.affiliation: KwaZulu-Natal Sharks Board, 1A Herrwood Drive, Umhlanga Rocks 4320, South Africa; Biomedical Resource Unit, University of KwaZulu-Natal, Westville Campus, University Road, Westville 3600, South Africa. Electronic address: wintner@shark.co.za.
authors.forename: Sabine
authors.initials: S
authors.lastname: Wintner
authors.affiliation: Faculty of Biosciences and Aquaculture, Universitetet i Nordland, Universitetsalleen 11, 8049 Bodø, Norway. Electronic address: Galice.Guillaume.Hoarau@uin.no.
authors.forename: Galice
authors.initials: G
authors.lastname: Hoarau
mesh_terms: D000818:Animals; D001499:Bayes Theorem; D005075:Biological Evolution; D002467:Cell Nucleus / Q000235:genetics; D005580:Fossils; D054629:Genome, Mitochondrial; D059014:High-Throughput Nucleotide Sequencing; D016013:Likelihood Functions; D008957:Models, Genetic; D010802:Phylogeny; D016415:Sequence Alignment; D017422:Sequence Analysis, DNA; D012860:Skates, Fish / Q000145:classification*
keywords: Divergence times; Manta; Mitogenome; Mobula; Molecular clock; Phylogenetics