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1.1 První Chobotnatci 5 1.2 Plesielephantiformes 5 1.3 Elephantiformes 6 1.3.1 Mammutida 6 1.3.2 Elephantida 7 1.3.3 Elephantoidea 7 2

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1.1 První Chobotnatci 5 1.2 Plesielephantiformes 5 1.3 Elephantiformes 6 1.3.1 Mammutida 6 1.3.2 Elephantida 7 1.3.3 Elephantoidea 7 2 MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV GEOLOGICKÝCH VĚD Jakub Březina Rešerše k bakalářské práci Využití mikrostruktur klů neogenních chobotnatců na příkladu rodu Zygolophodon Vedoucí práce: doc. Mgr. Martin Ivanov, Dr. Brno 2012 OBSAH 1. Současný pohled na evoluci chobotnatců 3 1.1 První chobotnatci 5 1.2 Plesielephantiformes 5 1.3 Elephantiformes 6 1.3.1 Mammutida 6 1.3.2 Elephantida 7 1.3.3 Elephantoidea 7 2. Kly chobotnatců a jejich mikrostruktura 9 2.1 Přírůstky v klech chobotnatců 11 2.1.1 Využití přírůstků v klech chobotnatců 11 2.2 Schregerův vzor 12 2.2.1 Stavba Schregerova vzoru 12 2.2.2 Využití Schregerova vzoru 12 2.3 Dentinové kanálky 15 3 Sedimenty s nálezy savců v okolí Mikulova 16 3.1 Baden 17 3.2 Pannon a Pont 18 1. Současný pohled na evoluci chobotnatců Současná systematika chobotnatců není kompletně odvozena od jejich fylogeneze, rekonstruované pomocí kladistických metod. Diskutované skupiny tak mnohdy nepředstavují monofyletické skupiny. Přestože jsou taxonomické kategorie matoucí (např. Laurin 2005), jsem do jisté míry nucen je používat. Některým skupinám úrovně stále přiřazeny nebyly a zde této skutečnosti není přisuzován žádný význam. V této rešerši jsem se zaměřil hlavně na poznatky, které následovaly po vydání knihy; The Proboscidea: Evolution and Paleoecology of Elephants and Their Relatives, od Shoshaniho a Tassyho (1996). Chobotnatci jsou součástí skupiny Tethytheria společně s anthracobunidy, sirénami a desmostylidy (Shoshani 1998; Shoshani & Tassy 1996; 2005; Gheerbrant & Tassy 2009). Základní klasifikace sestává ze dvou skupin. Ze skupiny Plesielephantiformes, do které patří čeledě Numidotheriidae, Barytheriidae a Deinotheridae a ze skupiny Elephantiformes, do které patří čeledě Palaeomastodontidae, Phiomiidae, Mammutida, Gomphotheriidae, tetralofodontní gomfotéria, Stegodontidae a Elephantidae (Shoshani & Marchant 2001; Shoshani & Tassy 2005; Gheerbrant & Tassy 2009). Čeleď Moeritheriidae byla vyjmuta z Plesielephantiformes, kvůli jejich bunodoncii a zařazena byla samostatně v rámci Proboscidea (Shoshani a Tassy 2005). Ze znaků používaných k fylogenetickým rekonstrukcím byl v minulosti dáván neobvykle přehnaný důraz na znaky na molárech (Shoshani & Tassy 1996). V současnosti je kladen důraz také na morfometrii klů, maxily a mandibuly, mandibulárního kanálu s přidruženými otvory (Ferretti & Debruyne 2011) a jazylkového aparátu (obr. 1) zvaného hyoideum (Shoshani & Marchant 2001). Obr. 1 - Fylogeneze chobotnatců na základě jazylkového aparátu (Shoshani & Marchant 2001; 2005, upraveno). Znaky na jazylkovém aparátu jsou vhodné pro fylogenetické analýzy, jelikož je svou funkcí u chobotnatců zcela unikátní a větších rozměrů, které dovolují zachování této většinou drobné kůstky. Je spojen s polykáním a dýcháním (je připojen na jazyk), pomáhá v komunikaci (pomocí něj chobotnatci vydávají zvuky) a s přidruženým svalstvem pomáhá ukládat vodu do pharyngeálního vaku v období stresu (Shoshani & Marchan 2001; Shoshani & Tassy 2005). Podle Shoshaniho a Tassyho (2005) se jazylkový aparát vyvinul v blízkosti hranice oligocén/miocén asi před 25 miliony let. | 3 Přestože, kromě paleontologického materiálu, lze u chobotnatců studovat DNA žijících slonů a zmrzlých mršin mamutů a mamutidů ze Sibiře a Aljašky, v jejich fylogenezi je stále mnoho nejasností. Shoshani a Tassy (1996) a Shoshani (1998) rozlišují hlavní evolučními trendy u chobotnatců jako; 1) zvětšování tělesných rozměrů (největší až kolem 4,5 m v kohoutku), s tím spojené; 2) prodlužování končetin a vývoj krátkého a širokého chodidla (graviportální a gravigrádní adaptace); 3) zvětšování lebky do mimořádných rozměrů; 4) zkracování krku (začalo u oligocenních chobotnatců); 5) zvětšování mozku; 6) prodlužování mandibuly a později jako paralelismus také sekundární zkrácení v oblasti symfýzy spojené se ztrátou mandibulárních klů; 7) vývoj chobotu, který se vyvíjel spolu s protažením a pozdějším zkrácením mandibuly a patrně se také vyvíjel paralelně mezi elefantidy, mamutidy a gomfotérii; 8) horizontální výměa molárů a premolárů; 9) postupná redukce v počtu zubů; 10) hypertrofie (nadměrný růst) druhých řezáků do formy klů, které byly rovné, stáčely se dolů nebo nahoru a jsou tak největšími známými zuby; 11) rozšíření a specializace molárů: zvyšování počtu centrálních hrbolů, kuželů a počtu příčných hřebenů, nebo lamel (od dvou hřebenů u nejprimitivnějších a až po 30 hřebenů u nejpokročilejších chobotnatců), přičemž je v tomto směru paralelismus patrný mezi skupinami Elephantinae a Stegodonty. Evoluce chobotnatců se význačuje třemi radiacemi (Shoshani & Tassy 1996; Shoshani 1998; Gheerbrant & Tassy 2009). První je paleocén-oligocénní, kdy se objevily listožravé (folivorní) druhy s vertikální výměnou zubů a nízkými korunkami (brachyodontní), jsou to plesielefantiformové, meritéria a první elefantiformové. Druhá je miocenní a je pro ni typická radiace gomfotérií sensu lato a stegodontů. Živili se převážně trávou, všichni měli horizontální výměnu zubů, horní tři moláry měly sedm lamel a byly brachyodontní nebo hypsodontní (vysoké korunky). Třetí radiace trvá od miocénu do současnosti (7 milionů let) a jde o radiaci moderní čeledi Elephantidae. Většina se živila a živí trávou a všichni mají horizontální výměnu zubů (až do 30 lamel) a vysoké korunky (hypsodontní). Nejvyšší taxonomická diverzita chobotnatců proběhla v miocénu (obr. 2). Graviportální postoj chobotnatců zřejmě vznikl během oligocénu a hrál jistě klíčovou roli v jejich rozšíření během miocénu (Delmer 2009). Chobotnatci se vyskytovali v různých biotopech; od pouští po hory a rozšířili se na všechny kontinenty vyjma Austrálie a Antarktidy. Obr. 2 - Fylogeneze chobotnatců (Gherbrant & Tassy 2009, upraveno). | 4 1.1 První chobotnatci Za nejstarší chobotnatce jsou považovány druhy Phosphatherium escuilliei a Daouitherium rebouli, které pocházejí ze svrchnopaleocenní (stupeň thanet) lokality Ouled Abdoun v Maroku (Gheerbrant et al. 1996; 1998; Gheerbrant et al. 2001; 2002). Mají mnoho společných znaků s Numidotherium koholense z časně eocenní lokality El Kohol v Alžírsku. Šlo o listožravá (folivorní) zvířata a je pro ně charakteristický striktně lofodontní chrup. Velikost rodu Phosphatherium v kohoutku byla pravděpodobně menší než 1 m. Obr. 3 - Rekonstrukce bazálního chobotnatce druhu Phosphatherium a Daouitherium Phosphatherium escuilliei (Gheerbrant & Tassy 2009). neukazují žádné z nejnápadnějších znaků chobotnatců, jako chobot, kly, horizontální výměnu molárů a graviportální chůzi (obr. 3). Přítomny jsou však méně nápadné byť zásadní synapomorfie na lebce a zubech (Gheerbrant & Tassy 2009). P. escuilliei je známo jen z kraniálního materiálu. Chybí rostrální část, která by mohla pomoci s homologií horních klů chobotnatců (Delmer 2009) a nález postkrania by mohl osvětlit předpoklad, zda Tethytheria měla semiakvatické předky (Gheerbrant & Tassy 2009). P. escuilliei a D. rebouli dokazují raný vznik chobotnatců, jejich Africký původ, ale ukazují také časnou radiaci moderních skupin savců explozivního charakterem ve spojení s vymíráním na hranici křída/paleocén (Gheerbrant et al. 1996; Gheerbrant 1998; Gheerbrant & Tassy 2009). Primitivní chobotnatci měli lofodontní chrup. Bunolofodontní moláry byly vyvinuty jen u skupin Moeritheriidae a Elephantiformes. Nález druhu Arcanotherium savagei (Delmer 2009) ze spodnooligocénní lokality Dor El Talha v Lybii svými unikátními znaky, kterými se liší od rodů Barytherium i Numidotherium, představuje jakýsi morfologický mezičlánek mezi striktně lofodontními eocenními chobotnatci a bunolofodontními. Stejně tak Moeritherium chehbeurameuri ze středno až svrchno-eocéní lokality Bir El Ater v Alžírii (Gheerbrant & Tassy 2008). Údaje Delmera (2009) o mandibulární symfýze A. savagei, podporují hypotézu homologie dolních řezáků časných chobotnatců a stále rostoucích klů u elephantiformů. Nový nález barytherida druhu Omanitherium dhofarens (Seiffert et al. 2012) ze spodního oligocénu Ománu ukazuje svojí denticí na morfologický přechod mezi rody Arcanotherium a Barytherium. Tabuce et al. (2007) se zabýval fylogenetickými vztahy primitivních chobotnatců pomocí mikrostruktur skloviny (obr. 9). 1.2 Plesielephantiformes Patří sem všichni bazální chobotnatci kromě meritérií (Numidotheriidae, Barytheriidae a Deinotheriidae). Skupina Deinotheriidae je kromě značných rozměrů charakteristická stále rostoucím párem mandibulárních klů, absencí klů maxilárních a reverzí k lofodontnímu chrupu, čímž se výrazně liší od ostatních skupin chobotnatců. Deinotéria mají původ v Africe. Předek deinotérií je patrně rod Chilgatherium z pozdního eocénu Etiopie (Sanders et al. 2004). Poslední deinotéria Deinotherium bossi přetrvala v Africe do doby před 1,5 miliony let (Shoshani & Tassy 1996; Gheerbrant & Tassy 2009). Původ deinotérií je třeba hledat u skupin Numidotheriidae, Barytheriidae a Moeritheriidae se kterými sdílejí některé | 5 morfologické znaky (Shoshani & Tassy 1996). Nový nález barytherida druhu Omanitherium dhofarens ze spodního oligocénu Ománu ukazuje svojí prodlouženou symfýzou a spodními řezáky na možný původ deinotérií u skupiny Barytheriidae (Seiffert et al. 2012). 1.3 Elephantiformes Nejstaršími elefantiformy jsou rody Palaeomastodon (lofodontní) a Phiomia (bunodontní). Za nejprimitivnější je považován druh Palaeomastodon beadnelli. Není však jisté který plesielefantiform je jejich předkem, protože morfologické rozdíly mezi prvními zástupci Elephantiformes a archaickými chobotnatci jsou dost nápadné. Do skupiny Elephantimorpha spadají skupiny Mammutida, Elephantida do které
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