20080706155500!Basilosaurus 135F~Dinosaurs-in-River-Posters 1 asteroid-impact-by-david-hardy cladogramDino pterosaur-color-sm I10-77-Ediacara Systematics and taxonomy systematics, paleontology ® history of evolutionary changes systematics = study of relationships between organisms taxonomy = theory and practice of classification category: class, order, family, species, … taxon: Mammalia, Primates, Hominidae, Homo sapiens, … 1. Before Linnaeus honeybee = Apis pubescens, thorace subgriseo, abdomine fusco, pedibus posticis glabris utrinque margine ciliatis [Bee with soft short hairs, gray chest, dark brown abdomen, legs with no hair, and small sacs with hair-like outgrowths along the edge] Acaciae quodammodo accedens, Myrobalano chebulo Veslingii similis arbor Americana spinosa, foliis ceratoniae in pediculo geminatis, siliqua bivalvi compressa corniculata seu cochlearum vel arietinorum cornuum in modum incurvata, sive Unguis cati [A spiny American tree, in some way resembling Acacia, similar to Vesling´s Myrobalanus chebulae, with Ceratonia leaves in pairs on the pedicle, a silique with two valves, which is compressed, and horn-shaped or curved like the horns of snailshells or ram´s horns, or like a cat´s claws] European bison = buffle, urus, bubalus, catoblepas, theur, the bubalus of Belon, Scottish bison … Aristotle: bonasus ® the same? Carolus Linnaeus 2. Carolus Linnaeus: 1735 Systema Naturae binomial nomenclature: genus + species hierarchical classification: kingdom, class, order, genus, species, (variety/subspecies) 3. Darwin: cladogenesis (branching) and anagenesis (change without branching) a system should reflect a real phylogeny ® but HOW? Evolutionary systematics before 1950: common ancestor + adaptive divergence discussions if adaptive or neutral traits better subjective and unclear criteria of choosing and weighing of traits Þ taxonomy in crisis (Þ the word „taxonomy“ itself replaced by „systematics“) controversy between splitters and lumpers E. Mayr Numerical taxonomy (phenetics) 1957: Charles Michener, Robert Sokal, P.H.A. Sneath taxonomy should be based on a total similarity rather than on a small number of „important“ traits Þ as many traits as possible numerical methods: morphological and genetic distances, ordination (PCA, DFA, CVA, MDS, ...), cluster analysis (UPGMA) phenograms problems: homoplasy (= convergence, parallelism, reversion) shared primitive (ancestral) traits unequal rate of evolution http://1.bp.blogspot.com/-FTv4KO-M7Ng/UUdZeqGAakI/AAAAAAAACAw/ZX9OXu3fGZk/s1600/spreadlabels1.png phenogram distance/ similarity Výsledek obrázku pro hoplitis image Hoplitis Phylogenetic systematics (cladistics) 1950, 1966: Willi Hennig: Phylogenetic Systematics only genealogies, not adaptive divergence strict monophyly monophyletic group = clade monophyly paraphyly polyphyly W. Hennig IV.jpg “Reptilia” “Pongidae” characters: plesiomorphic (= ancestral, „primitive“) symplesiomorphic (= shared ancestral) apomorphic (= derived) synapomorphic (= shared derived) autapomorphic (= unique derived) clades defined only by synapomorphies symplesiomorphy autapomorphy synapomorphy mutation mutation clade.jpg crown group stem group Definition of clades and classification of extinct taxa: origin of an apomorphy arbitrary dimension principle of parsimony: Occam´s razor (William of Ockham, 14th century) cladograms http://pics.livejournal.com/earth_wizard/pic/0007r2b0 PhyloCode (International Code of Phylogenetic Nomenclature) till now somewhat controversial and impractical problems: homoplasy, rapid evolution Rivet: the simplest structure Þ we suppose that it is the most similar to the common ancestor of all modern types of fasteners We can define 7 derived character states (ie. those nonexistent in rivets): 1) notched heads, 2) rounded heads, 3) hex heads, 4) threaded shafts, 5) tapered shafts, 6) pointed tips, and 7) thick diameter. Cladistics and phenetics exemplified by the „evolution“ of fasteners Character states of all 4 types are listen in the following table where „0“ = plesiomorphic („rivet-like“) state „1“ = apomorphic (derived) state Capture d’écran 2013-11-22 à 11.47.47.png Cladistics and phenetics exemplified by the „evolution“ of fasteners If we use the phenetic approach we count the total number of shared states (both ancestral and derived). For example rivet vs. nail: 6 similarities, 1 difference Cladistics and phenetics exemplified by the „evolution“ of fasteners Capture d’écran 2013-11-22 à 11.51.19.png If we use the cladistic approach we take into account only shared derived states. For example. screw vs. bolt: 2 synapomorphies Cladistics and phenetics exemplified by the „evolution“ of fasteners Capture d’écran 2013-11-22 à 11.51.19.png Capture d’écran 2013-11-22 à 11.51.29.png •Cladistics and phenetics exemplified by the „evolution“ of fasteners Evolutionary systematics – a response phylogenetic relationships + degree of divergence Þ combination of phenetic and cladistic approach reflection of both clades and grades An evolutionary grade is a group of similar species that has given rise to another group that differs markedly from the ancestral condition, and is thus not considered part of the ancestral group. Þ the ancestral group is then paraphyletic eg. reptiles (without birds), fishes in a traditional sense E. Mayr HISTORY OF LIFE ON EARTH http://thumbnails-visually.netdna-ssl.com/the-great-tree-of-life_505ba0c07cda2.gif Precambrian Hadean Archean Proterozoic Phanerozoic eon Výsledek obrázku pro geological time Precambrian Hadean Archean Proterozoic Phanerozoic eon Ediacaran (Vendian) fauna ~635-542 M File:Leicestershire UK locator map 2010.svg Charnwood, Leicestershire ~ 560 M https://aroundtheredmap.files.wordpress.com/2015/04/nfld_fullmap.gif http://upload.wikimedia.org/wikipedia/commons/thumb/9/9a/Newfoundland_and_Labrador_in_Canada.svg/20 00px-Newfoundland_and_Labrador_in_Canada.svg.png Mistaken Point, Newfounland ~ 565 M http://chigaku.ed.gifu-u.ac.jp/chigakuhp/html/news_chikyuh/060725/pic01.jpg https://www.dfat.gov.au/aib/images/australia-topographic-map-960.jpg http://palaeo.gly.bris.ac.uk/palaeofiles/lagerstatten/Ediacara/map.gif Ediacara Hills, Australia Dickinsonia ~ 580 M Spriggina Charnia http://museumvictoria.com.au/pages/17084/ImageGallery/pic-41264c-40.jpg http://firstlifeseries.com/wp-content/gallery/stills/firstlife_6.19_Spriggina_CGI.jpg http://upload.wikimedia.org/wikipedia/commons/5/5f/Bomakellia_kelleri.JPG https://www.sciencemag.org/content/319/5870/1660/F2.large.jpg Funisia: sex? Spriggina Paleozoic Mesozoic Cenozoic Cambrian explosion ~ 542-520 M Phanerozoic era Cambrian explosion Burgess Burgess Shale ~ 542-520 M burgess continent Canadian Rockies, Yoho National Park Wallcott Charles Doolittle Walcott (1909) conway_morris 421319a-i1 Simon Conway Morris 225px-Charles_Walcott 6a00e553df64898834011168520f8e970c-800wi 9780192862020 Yohoia Anomalocaris Marella Marella Anomalocaris nathorsti A. canadensis burgess_shale Yohoia Halucigenia Pikaia2 Pikaia gracilens Hallucigenia Aysheaia2 Leachoilia Nectocaris Odaraia Opabinia2 Wiwaxia Aysheaia Opabinia Wiwaxia Nectocaris Leachoilia Odaraia slide0006_image035 Hallucigenia Halucigenia EP_pikaia Pikaia gracilens (Chordata) http://jto.s3.amazonaws.com/wp-content/uploads/2015/06/f-hallucigenia-b-20150626.jpg http://jto.s3.amazonaws.com/wp-content/uploads/2015/06/f-hallucigenia-a-20150626.jpg http://i.dailymail.co.uk/i/pix/2015/06/25/11/29F5D5F000000578-0-Hallucigenia_sparsa_lived_in_the_oc eans_around_505_million_years-a-15_1435227606620.jpg http://fc02.deviantart.net/fs71/i/2013/049/1/d/aysheaia_by_paleopatrick-d5uj8yg.jpg http://burgess-shale.rom.on.ca/images/zoomify/aysheaia-usnm-83942a.jpg Aysheaia Transition from sea to land? http://fc02.deviantart.net/fs71/i/2013/049/1/d/aysheaia_by_paleopatrick-d5uj8yg.jpg http://burgess-shale.rom.on.ca/images/zoomify/aysheaia-usnm-83942a.jpg Onychophora diversity and disparity: interpretation of Burgess fossils Stephen Jay Gould vs. Simon Conway Morris diversity = number of species disparity = number of „Bauplans“ traditional Gould Conway Morris http://scilib.narod.ru/Biology/Dennett/DDI/images/fig_10_12.png http://home.cogeco.ca/~parksidescienceq/SBI%203U/evolution%201/gould_stephen_jay.gif http://www.nature.com/nature/journal/v421/n6921/images/421319a-i1.0.jpg biodiversita Phanerozoic increasing diversity Jack J. Sepkoski (1981): logistic model Biodiversity2 Michael J. Benton (1997): curve for terrestrial organisms differs exponential model sepkoski_and_ronnie modern fauna Cambrian fauna Paleozoic fauna Biodiversita3 Biodiversita4 If we take into account incompleteness of the fossil record ® no trend? biodiversita Extinction: background extinctions mass extinctions ® „Big Five“ greatest: end of the Permian Ordovician Devonian Permian Triassic K/T Paleozoic Cambrian: single supercontinent Rodinia (Proterozoic) ® Gondwana, Laurentia, Baltica, Angara (Siberia), Avalonia … Ordovician: increase of diversity (marine organisms) the end: 1st mass extinction Silurian: gnathostomes first terrestrial o. (plants, scorpions) Laurentia+Baltica = Laurasia Devon: radiace ryb, první žraloci, lalokoploutví, obojživelníci na konci 2. masová extinkce Výsledek obrázku pro amphibian evolution tiktaalik Ichthyostega 365 Ma Devon: radiace ryb, první žraloci, lalokoploutví, obojživelníci na konci 2. masová extinkce Acanthostega Ichthyostega Karbon: přesličky, hmyz, první plazi Archaeothyris (Synapsida) Perm: Pangea Therapsida (® savci) na konci 3. masová extinkce Edaphosaurus (Pelycosauria) Mesozoic Triassic: butterflies, dipterans radiation of reptiles (tortoises, ichthyosars, plesiosaurs, pterosaurs) the end: dinosars, mammals, 4th extinction ichthyosaurs pterosaurs cynodont (Cynognathus) primitive mammal (Castorocauda) Therapsida synapsid Pelycosauria (Palaeohatteria) plesiosaurs https://ad8d0c4b-a-62cb3a1a-s-sites.googlegroups.com/site/palaeocritti/by-group/biarmosuchia/Biarmo suchus_NT.jpg?attachauth=ANoY7crfJHTDoL0B20prGrFWaRTRXv2QeHcBbqqa4JSu_qMfYFr5qLWmqeTjtJUi7d2QMa0jCR Asa_m-3UIArR_TkhL7T0O9B-wfvtG8meuKYvAGMaNHcNbEFSLut5_tyJ3tNFNQtPUEV0ksgsdyqC5L3YBNfIDLQJbL4QRnMFTZV 62lw7bBFSf4PZUE_dIEmIFrNFQiBd4KBpbN6OrCIbYBuE6AuF5f-0Xs-cP-sPtgyzQt2Bvrf09BJij_0jeUNi7-GiX0i9fp&att redirects=0 Sphenacodon: Lower Permian (270 M ) – mandible from several bones, reptile-like articulation, no eardrum Biarmosuchia: Upper Permian – one of the first therapsids, articulation already more mammal-like, knit upper jaw, hind legs more upright Biarmosuchus Evolution of mammals Procynosuchus: end of Permian – primitive cynodont Thrinaxodon: Lower Triassic – more derived cynodont, eardrum in the lower jaw http://upload.wikimedia.org/wikipedia/commons/9/97/Procynosuchus_BW.jpg http://palaeos.com/vertebrates/cynodontia/images/thrinaxodon1.jpg http://upload.wikimedia.org/wikipedia/commons/d/d4/Morganucodon.jpg Probainognathus: Middle Triassic (~ 235 M) – 2 joints: mammalian+reptilian Diarbrognathus: Lower Jurassic (~ 209 M) – advanced cynodont, although still 2 joints, but the reptilian one used almost entirely for hearing Morganucodon: Lower Jurassic (~ 220 M) – still a residue of the reptilian joint http://fc06.deviantart.net/fs70/i/2009/354/d/d/Probainognathus_by_Theropsida.jpg Hadrocodium: Lower Jurassic – transition of the middle ear ossicles from the jaw to the cranium http://24.media.tumblr.com/tumblr_mduve8jcgm1qc2w0eo1_500.jpg Jurassic: bone fishes bird evolution Saurischia Ornithischia Allosaurus Stegosaurus dinosaurs Mesozoic 104Saurischia Brachiosaurus theropod dinosaurs tyrannosaurs (Cretaceous) Maniraptora Archaeopteryx lithographica birds Microraptor gui Mesozoic t-rex_vs_gigantosaurus T. rex Gigantosaurus ap_dinobirds Výsledek obrázku pro microraptor gui image Jurassic: bone fishes bird evolution Cretaceous: angiosperms modern sharks and rays, mosasaurs, first snakes, birds mammals: divergenc of marsupials and placentals mosasaurs Hesperornis the end: 5th extinction, 66 M ® the cause?? Mesozoic Ichthyornis_BW Ichthyornis Extinkce na K/T* (K/Pg**) boundary: 1980 Louis Alvarez et al.: catastrophic hypothesis – asteroid 10 km in diameter 109 ´ more than Hiroshima pic L. Alvarez Dinosauri.jpg *) Cretaceous/Tertiary **) Cretaceous /Paleogene Ukázalo se, že extinkce savců alespoň v Severní Americe byla mnohem vážnější (93 % druhů, 86 % rodů) a zotavení rychlejší než se původně myslelo. Během 300 ky lokální diverzita na K úrovni, morfologická disparita dokonce vyšší; díky vysoké míře endemismu regionální diverzita dokonce 2X vyšší. Důvodem to, že extinkce se týkala především druhů s menším rozšířením, proto bias – ten se může týkat i dalších extinkcí. Extinction on the K/T (K/Pg) boundary: iridium on the boundary cca. 100-fold increase of the amount of iridium K/T boundary Chicxulub crater (Mexico) cenotes syncline tektites from the K/T map of gravitational field shock crystal Problems of the impact theory: extinctions not that sudden for most animals, occurring before the impact species have been disappearing in several phases from more thermophilic to less thermophilic the impact by ca. 300 ky older than the extinction (but it may have trigger tsunamies and earthquakes Þ mixing of layers) locality El Penon (Mexico): same species above and below the „impact“ layer) Alternative hypothesis: gradual cooling caused by giant volcano erruptions on the Deccan Plateau in India basalt layer 1200-1800 m thick, 100 000 km2 Þ during 1 MY ® min. 1,5 mil. km3 of basalts the origin of the plateau at the turn of the Cretaceous and Tertiary Recent findings: According to new dating the Deccan event appeared before the impact – problém: inaccurate dating of the Indian event More precise dating: the Chicxulub crater corresponds with the extinction ~100 ky before the impact cooling by 6–8 °C, probably as a consequence of the Deccan catastrophe – the impact then as the „coup of grace“ Cyanobacteria as a result of the greenhouse effect? Some theories: two consecutive impacts Animations: eg. https://www.youtube.com/watch?v=bU1QPtOZQZU Paleozoic Mesozoic Cenozoic Phanerozoic era eon: epoch Paleogene Neogene Paleocene Eocene Oligocen Miocene Plio- Pleisto- period Paleontological vs. molecular data When have animal phyla and mammalian/bird orders emerged? Cambrian explosion? molecular data (Wray et al. 1996): Protostomia-Deuterostomia ~ 1200 M Chordata-Echinodermata ~ 1000 M „phylogenetic fuse“? Cambrian explosion? recent molecular estimates closer to the Cambrian explosion: Metazoa ~ 650 M (Peterson et al. 2004) Protostomia-Deuterostomia ~ 582 M (Aris-Brosou and Yang 2003) Paleontological vs. molecular data When have animal phyla and mammalian/bird orders emerged? Cambrian explosion? fauna of Chengjiang (Chína) ~ 525 M Yunnanozoon lividum Myllokunmingia http://cambrian-cafe.up.n.seesaa.net/cambrian-cafe/image/Yunnanozoon_lividum_02.jpg?d=a1 http://www.geocities.co.jp/NatureLand/5218/yunnanozu-n.jpg http://upload.wikimedia.org/wikipedia/commons/7/78/Haikouella_lanceolata.jpg http://bio.sunyorange.edu/updated2/THINKING_EVOLUTION/prehistoric/early_ans/transitional/transition al/image102.jpg Haikouella lanceolata jawlss fish Cambrian explosion? fauna of Chengjiang (China) ~ 525 M Doushantuo formation (S China), 590–560 M: many species early embryonic stages? Haikouichthys ercaicunensis 525 M http://images.sciencedaily.com/2004/11/041104005307.jpg http://upload.wikimedia.org/wikipedia/commons/9/9f/Haikouichthys_3d.png cetacean evolution: mesonychids ® moving to water ® cetaceans Andrewsarchus mongolicus recent groups of mammals and birds and the K/T boundary Mesonyx Spinosaurus Gigantosaurus Allosaurus Tyrannosaurus Andrewsarchus polar bear Paleontological vs. molecular data When have animal phyla and mammalian/bird orders emerged? cetacean evolution mesonychids ~ 56 M Pakicetus 56-34 M Ambulocetus 50-49 M Rodhocetus 47 M Basilosaurus 40-34 M Dorudon 41-33 M Cetotherium 15 M Hazardni hrac General principles diversity: stock market analogy extinction: foot soldier model lifetime of lineages: gambler´s bankruptcy model D. Raup J. J. Sepkoski random walk David Raup, Jack Sepkoski: periodicity? (26 M)