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Genetics and Evolution of Marine Vertebrates (Without Dispersive Larval Stages)

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Genetics and Evolution of Marine Vertebrates (Without Dispersive Larval Stages) GENETICS AND EVOLUTION OF MARINE VERTEBRATES (WITHOUT DISPERSIVE LARVAL STAGES) Ana Veríssimo – GEOA The Marine Environment nmh.ac.uk (Natural History Museum) The Horizontal Dimension The Vertical Dimension Epipelagic 200 m Ø decreasing light levels Mesopelagic Ø increasing pressure (1 atm per 10 m) 1000 m Ø decreasing temperatures (down to ~ 4⁰C) Bathypelagic 4000 m Abyssopelagic earthguide.ucsd.edu Stable environmental conditions at depths > 1000 m. Marine Biogeographic Regions Briggs & Bowen 2012 J. Biogeogr. Species Distribution at Depth Priede 2017. Marine Organisms Without Larval Dispersal ¨ All cartilaginous fishes (chimaeras, sharks, rays) ¨ Many bony fishes (direct development) ¨ All sea turtles ¨ All marine mammals Vertebrate Tree of Life Chondrichthyes Skates Sharks Chimaeras Li et al. 2012 Mol Phylo Evol Chondrichthyes Rays (Batoidea) Rapid radiation of: skates vs. sharks (~400 Myr) Sharks (Galeoidea) Drivers of diversification not clear yet… Li et al. 2012 Mol Phylo Evol Sharks (Selachii) Phylogenetic relationships among and within major shark lineages not yet resolved… Sharks (Selachii) Eukaryota; Metazoa; Vertebrata; Chondrichthyes 321 Carcharhinidae 44 Hemigaleidae 42 Triakidae-1 40 35 Triakidae-2 Leptochariidae 22 Pseudotriakidae 21 25 Recent studies suggest that Proscylliidae Carcharhiniformes 16 Scyliorhinidae-1 most families appeared before the Scyliorhinidae-2 Alopiidae 39 end of the Cretaceous, Megachasmidae 13 33 Odontaspididae 31 Pseudocarchariidae 29 Lamnidae 36 Cetorhinidae Lamniformes with major radiations between 20 32 Carchariidae 7 Mitsukurinidae Jurassic and Cretaceous. Ginglymostomatidae 45 38 Stegosomatidae Hemiscylliidae 5 19 Brachaeluridae 27 14 Orectolobidae Orectolobiformes Parascylliidae Heterodontidae Somniosidae Heterodontiformes Heterodontiformes Ancient divergences, oceanic habits Oxynotidae 28 3 Dalatiidae and large variation in divergence 23 Etmopteridae Centrophoridae Squaliformes timings among studies are challenging 12 Squalidae Squatinidae 9 18 Echinorhinidae Squalimorphii in reconstructingGaleomorphii the biogegraphic Squatiniformes Squatiniformes Pristiophoridae 4 Echinorhiniformes Notorynchidae history of this group. 43 15 Hexanchidae Pristiophoriformes Chlamydoselachidae (continued on next page) Hexanchiformes O S D C P Tr J K Pg PALEOZOIC MESOZOIC CZ 400 300 200 100 0 Million years ago Fig. 2 Continues HHedges.indbedges.indb 332121 11/28/2009/28/2009 11:28:04:28:04 PPMM Rays (Batoidea) 322 THE TIMETREE OF LIFE (continued on previous page) Myliobatidae 34 Gymnuridae The same studies suggest that Dasyatidae-1 2 most batoid families also Dasyatidae-2 30 Urolophidae appeared before the end of Hexatrygonidae the Cretaceous. Plesiobatidae Myliobatiformes Potamotrygonidae 11 Urotrygonidae Rhinidae 41 10 Rhynchobatidae 24 Rhiniformes 1 Pristidae Rhinobatidae Rhynchobatiformes Pristiformes 8 Platyrhinidae Torpedinidae Rhinobatiformes 26 Platyrhiniformes Narcinidae-2 6 Narcinidae-1 Rajidae Torpediniformes Rhinochimaeridae Rajiformes 37 17 Chimaeridae Callorhinchidae Holocephali Batoidea Chimaeriformes O S D C P Tr J K Pg PALEOZOIC MESOZOIC CZ 400 300 200 100 0 Million years ago Fig. 2 A timetree of cartilaginous fi shes (Chondrichthyes). for paraphyletic and/or polyphyletic groups are as follows: Divergence times are from Table 1. Galeomorphii, Triakidae-1 (Mustelus), Triakidae-2 (Triakis), Scyliorhinidae-1 Squalimorphii, and Batoidea comprise the Subclass (Pentanchinae), Scyliorhinidae-2 (Scyliorhininae), Dasyatidae-1 Elasmobranchii. Abbreviations: C (Carboniferous), CZ (Cenozoic), (Dasyatis), Dasyatidae-2 (Himantura), Narcinidae-1 (Narcininae), D (Devonian), J (Jurassic), K (Cretaceous), O (Ordovician), and Narcinidae-2 (Narkinae). P (Permian), Pg (Paleogene), S (Silurian), and Tr (Triassic). Codes Division of living cartilaginous " shes into Elasmo- compressed sawsharks (Pristiophoriformes) and angel branchii and Holocephali is strongly supported by sharks (Squatiniformes) (15). ! ese in turn group morphological analyses, as is uniting these groups to with the Orders Squaliformes, Hexanchiformes, and form Chondrichthyes (6, 9, 11). Within Holocephali, it Echinorhiniformes in the Hypnosqualean clade. ! e is believed that Rhinochimaeridae and Chimaeridae Orders Lamniformes, Carcharhiniformes, Orectolob- form a group to the exclusion of Callorhinchidae (12). iformes, and Heterodontiformes are grouped as Galea (3, ! e relationships of the more species-rich elasmo- 4). Minor modi" cations to Shirai’s original 1992 hypoth- branchs are more contentious. Early studies suggested esis of elasmobranch interrelationships were made by de a basal split between sharks and rays (13, 14). In 1992, Carvalho in 1996 (4). ! is hypothesis remains the con- Shirai published an extensive and in# uential analysis sensus from morphological data. of morphological variation among sharks and rays Because the monophyly of Chondrichthyes and recip- in which he proposed a “Hypnosqualean hypothesis” rocal monophyly of Elasmobranchii and Holocephali wherein the batoids fall together with the dorsoventrally have not been controversial and are supported by HHedges.indbedges.indb 332222 11/28/2009/28/2009 11:28:05:28:05 PPMM Rays (Batoidea) Skates (Rajoidei) Thornbacks (Platirhinoidei) Electric rays (Torpedinoidei) Guitarfishes I Guitarfishes II Panrays (Zanobatoidei) Stingrays (Myliobatoidei) Aschliman 2011 Rays (Batoidea) Evolution of Batoids Long internal branches (old lineages) with recent species radiation Concordant across major lineages Radiations after major mass extinction event (K/T transition) into vacated niche space Convergence of body features across different lineages Mass extinction Aschliman 2011 Drivers of Elasmobranch Diversity ¨ Specific life history traits ¤ Habit: benthic, benthopelagic, pelagic ¤ Habitat: coastal/shelf waters, oceanic, deepwater ¤ Behavior: female phylopatry to discrete nursery areas Habit: Benthic vs. Pelagic Clades with exclusive but contiguous geographic ranges, and strong genetic differentiation among clades. Habit: Benthic vs. Pelagic A single Indo-Pacific clade, although with some evidence of further geographic structure. Habitat: Oceanic vs. Coastal No genetic differentiation detected across the Atlantic Ocean. mtDNA CR Habitat: Oceanic vs. Coastal Significant overall genetic differentiation, with two main genetic stocks. Habitat: Deepwater No genetic structure detected along the eastern Atlantic, or between the eastern Atlantic & Australia. Strong divergence of Mediterranean population, with different life history traits. Female Phylopatry to Nursery Areas Sampling of adult females and small juveniles, using 17 nuclear microsatellites for parental reconstruction. Most adult females gave birth at their natal island, and also returned to that same nursery every breeding cycle. Female Phylopatry to Nursery Areas Female Phylopatry to Nursery Areas Individual site fidelity Population site fidelity Things to have in mind … ¨ Know your species! ¤ How is dispersal/gene flow accomplished? ¤ What may be potential barriers/limitations to gene flow? ¤ What life history traits may influence the distribution of individuals and their genes?.
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