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Parzitofauna Plazů Evropy JIHOČESKÁ UNIVERZITA V ČESKÝCH BUDĚJOVICÍCH ZEMĚDĚLSKÁ FAKULTA Studijní program: B4106 Zemědělská specializace Studijní obor: Biologie a ochrana zájmových organismů Katedra: Katedra biologických disciplín Vedoucí katedry: doc. RNDr. Ing. Josef Rajchard, Ph.D. DIPLOMOVÁ PRÁCE Parzitofauna plazů Evropy Vedoucí diplomové práce: Mgr. Miloslav Jirků, Ph.D. Konzultant diplomové práce: Mgr. Michal Berec, Ph.D. Autor: Bc. Jakub Žídek České Budějovice, 2014 Prohlašuji, že v souladu s § 47b zákona č. 111/1998 Sb. v platném znění souhlasím se zveřejněním své diplomové práce, a to - v nezkrácené podobě - v úpravě vzniklé vypuštěním vyznačených částí archivovaných zemědělskou fakultou - elektronickou cestou ve veřejně přístupné části databáze STAG provozované Jihočeskou univerzitou v Českých Budějovicích na jejích internetových stránkách, a to se zachováním mého autorského práva k odevzdanému textu této kvalifikační práce. Souhlasím dále s tím, aby toutéž elektronickou cestou byly v souladu s uvedeným ustanovením zákona č. 111/1998 Sb. zveřejněny posudky školitele a oponentů práce i záznam o průběhu a výsledku obhajoby kvalifikační práce. Rovněž souhlasím s porovnáním textu mé kvalifikační práce s databází kvalifikačních prací Theses.cz provozovanou Národním registrem vysokoškolských kvalifikačních prací a systémem na odhalování plagiátů. Datum: Podpis studenta: Souhrn: Předložená diplomová práce předkládá historicky nejucelenější přehled parazitů plazů žijících v Evropě, včetně základních taxonomických informací, známého geografického rozšíření, hostitelského spektra a nejdůležitějších referencí. Data potřebná k vytvoření přehledu byla získána z věděckých článků. V předložené práci jsou zaznamenání paraziti všech evropských plazů v celém jejich areálu rozšíření. Na víc byl vytvořen ucelený seznam evropské herpetofauny, které byly doposud nekompletní, včetně ohrožení a endemických druhů. Klíčová slova: review, plazi, paraziti, Evropa Abstract: This thesis presents the most comprehensive historical overwiev of reptile parasites living in Europe, including basic taxonomic information, the known geographic range, host range and important references. Data needed to create a host-parasite list was obtained from scientific articles. In the present work are recorded parasites of the Eropean reptiles throughout their entire range. Moreover was created a comprehensive list of the European herpetofauna that were still incomplete, including threat status and endemic species. Key words: review, reptiles, parasites, Europe Poděkování: Tímto bych chtěl poděkovat především mému školiteli Miloslavu Jirků, Ph.D, za vedení mé práce. Dále bych chtěl poděkovat své rodině a přátelům, kteří mi byli oporou nejen při psaní této práce, ale po celou dobu mého studia. Obsah 1. Úvod...................................................................................................................... 8 2. Cíle ........................................................................................................................ 9 3. Literární přehled ................................................................................................. 10 3.1 Charakteristika herpetofauny Evropy ................................................................. 10 3.1.1 Ohrožení................................................................................................... 11 3.1.2 Endemismus ............................................................................................. 13 3.2 Novodobý výzkum parazitů evropských plazů: od 70. let po současnost .......... 14 3.3 Charakteristika jednotlivých skupin parazitů plazů Evropy ............................... 19 3.3.1 Mikrosporidie (Microsporidia) ................................................................ 19 3.3.2 Výtrusovci (Apicomplexa) ...................................................................... 20 3.3.3 Hemogregariny (Haemogregarinidae ...................................................... 20 3.3.4 Kokcidie (Eimeriorina) ............................................................................ 21 3.3.5 Bičíkatá protista ....................................................................................... 22 3.3.6 Rybomorky (Myxozoa)............................................................................ 22 3.3.7 Pijavky (Hirudinae).................................................................................. 23 3.3.8 Roztoči (Acari) ........................................................................................ 23 3.3.9 Jednorodí (Monogenea) ........................................................................... 24 3.3.10 Motolice (Trematoda) .............................................................................. 25 3.3.11 Hlístice (Nematoda) ................................................................................. 25 3.3.12 Tasemnice (Cestoda)................................................................................ 27 3.3.13 Vrtejši (Acanthocephala) ......................................................................... 28 4. Metodika ............................................................................................................. 29 4.1 Specifikace cílové oblasti ................................................................................... 29 4.2 Vymezení spektra cílových druhů plazů ............................................................. 29 4.3 Specifikace parazitů ............................................................................................ 29 4.4 Výběr a zpracování literárních dat ...................................................................... 30 5. Výsledky ............................................................................................................. 32 6. Diskuse................................................................................................................ 42 7. Závěr ................................................................................................................... 47 8. Citace .................................................................................................................. 48 9. Přilohy ................................................................................................................. 60 1. Úvod Parazitismus je jednou z nejvíce rozšířených životních strategií organismů a hraje důležitou roli v evoluci. Zhruba před třiceti lety prošla parazitologie významnými změnami. Od dříve převládajícího spíše deskriptivního přístupu, který vycházel ze zoologických základů a byl velmi důležitý pro poznání širokého spektra skupin i jednotlivých druhů parazitů, došlo k přechodu na ekologické, fyziologické, imunologické, biochemické a v posledních 15-20 letech fylogenetické pojetí parazitologie. I přes bezpočet novodobých objevů v rámci parazitologie, zůstávají paraziti i nadále závažným problémem, mnohým se navíc vlivem lidské činnosti a globálních změn zvětšuje areál rozšíření a vzrůstá tak nebezpečí jejich šíření do nových oblastí a naivních hostitelů. Na druhou stranu vliv těchto faktorů vede také k úbytku řady druhů živočichů, kteří jsou jejich hostiteli. Sami paraziti, zejména ti hostitelsky specifičtí, tedy úzce vázaní na určitý taxon (např. plazího) hostitele se tak ocitají v pozici ohrožených, neboť bez svých ubývajících hostitelů nejsou schopni existovat. V neposlední řadě paraziti plazů představují zřejmě nejopomíjenější část globální parazitofauny obratlovců a představují tak její nejméně probádanou oblast zasluhující zvláštní pozornost. Přes relativně velké množství relevantních prací dodnes neexistuje kompletní seznam parazitofauny evropských plazů, který by ulehčil determinaci jednotlivých druhů, pomohl pochopit jejich ekologii a umožnil poodhalit podstatu složení a struktury jejich společenstev. Právě z těchto důvodů bylo v této práci stanoveno za cíl sestavení co možná nejkompletnějšího seznamu parazitů plazů vyskytujících se v Evropě, spolu s přehledem známého geografického rozšíření a spektra hostitelů jednotlivých druhů a základní vyhodnocení shromážděných dat. 8 2. Cíle Cílem této práce je: Vytvoření kompletního seznamu parazitů plazů Evropy, jejich geografického rozšíření a hostitelského spektra. Vytvoření kompletního seznamu plazů Evropy a jejich parazitů. Analýza získaných dat - charakteristika společenstev parazitů jednotlivých vyšších taxonů hostitelů. Analýza společenstev parazitů z hlediska jejich diverzity, životních strategií a biologie, hostitelského spektra a biogeografie. Zpracování části získaných dat formou rukopisu vědeckého článku. 9 3. Literární přehled 3.1 Charakteristika herpetofauny Evropy U plazů je jasný gradient růstu druhové diverzity směrem od severu k jihu s největší druhovou diverzitou v jižních oblastech, konkrétně na Iberském a Balkánském, příp. Apeninském poloostrově a v oblasti Kavkazu. Glaciální refugia, jako jsou například právě Iberský, Apeninský a Balkánský poloostrov, jsou důležitými centry druhové rozmanitosti, stejně jako Středomořské ostrovy. Nejvíce druhů plazů, tak nalezneme ve Španělsku (65 druhů), Řecku (55), Itálii (50), Francii (38) a Bulharsku (33) (Cox and Temple 2009). Celkové počty evropských druhů udává každý autor odlišné. Například Sillero et al. (2014) udávají 145 druhů plazů, zatímco Cox and Temple (2009) jich udávají 151. Právě tito autoři se nejvíce liší ve výběru druhů, které do Evropy zahrnují. Sillero et al. (2014) do Evropy nezahrnují druhy žijící na Kanárských ostrovech (např. ještěrky a gekony rodu Gallotia, resp. Tarentola), ostrovech Středozemního
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