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Druhová Diverzita a Evoluce Monogeneí Parazitujících Kaprovité Ryby Vybraných Oblastí Afriky

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Druhová Diverzita a Evoluce Monogeneí Parazitujících Kaprovité Ryby Vybraných Oblastí Afriky PŘÍRODOVĚDECKÁ FAKULTA Diverzita a evoluce monogeneí parazitujících kaprovité ryby vybraných oblastí Afriky DIPLOMOVÁ PRÁCE VOJTĚCH DLAPKA Vedoucí práce: prof. RNDr. Andrea Vetešníková Šimková, Ph.D. Ústav botaniky a zoologie Brno 2020 Bibliografický záznam Autor: Bc. Vojtěch Dlapka Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Diverzita a evoluce monogeneí parazitujících kaprovité ryby vybraných oblastí Afriky Studijní program: Ekologická a evoluční biologie Studijní obor: Zoologie Vedoucí práce: prof. RNDr. Andrea Vetešníková Šimková, Ph.D. Konzultant práce: Mgr. Eva Řehulková, Ph.D. Akademický rok: 2019/2020 Počet stran: 72 + 9 Klíčová slova: Monogenea, Afrika, Cyprinoidei, diverzita, fylogeneze, hostitelská specifita Bibliographic entry Author: Bc. Vojtěch Dlapka Faculty of Science, Masaryk University Department of Botany and Zoology Title of thesis: Diversity and evolution of monogeneans parasitizing cyprinid fish in the selected areas of Africa Degree Program: Ecological and Evolutionary Biology Field of study: Zoology Supervisor: prof. RNDr. Andrea Vetešníková Šimková, Ph.D. Consultant: Mgr. Eva Řehulková, Ph.D. Academic Year: 2019/2020 Number of Pages: 72 + 9 Keywords: Monogenea, Africa, Cyprinoidei, diversity, phylogeny, host specifity Abstrakt Diplomová práce je tematicky věnována druhové diverzitě a evoluci parazitů taxonu Monogenea u vybraných druhů kaprovitých ryb z různých geografických oblastí Afriky – Senegal (západní Afrika), Keňa (východní Afrika), Súdán (severovýchodní Afrika), Maroko (severní Afrika) a Kamerun (rovníková Afrika). V oblasti Afriky je v současné době známo kolem 500 druhů Cyprinoidei, tento taxon je druhý druhově nejbohatší. Převážná většina druhů patří do čeledi Cyprinidae, avšak znalosti diverzity jejich parazitofauny a evoluční vztahy mezi parazity afrických Cyprinoidei jsou pořád velice limitovány. Teoretická část této diplomové práce je věnována (1) druhovému složení kaprovitých ryb na africkém kontinentu, historické biogeografii této skupiny a (2) diverzitě a evoluci parazitů taxonu Monogenea u kaprovitých ryb Afriky. Praktická část je zaměřena na morfologickou i molekulární determinaci monogeneí kaprovitých ryb vybraných států Afriky, analýzu diverzity parazitických společenstev, základní epidemiologické ukazatele a fylogenetickou rekonstrukci parazitů taxonu Monogenea nalezených na Cyprinoidei Afriky. Abstract The diploma thesis is dealing with the species diversity and evolution of parasites of Monogenea of selected species of Cyprinoidei in different geographical regions of Africa – Senegal (West Africa), Kenya (East), Sudan (Northeastern Africa), Morocco (North Africa) and Cameroon (equatorial Africa). Around 500 Cyprinoidei species are currently known in Africa, where this taxon is the second most species-rich concerning the freshwater fishes. The vast majority of species belong to the Cyprinidae family, but knowledge of the diversity of their parasitofauna and the evolutionary relationships between parasites of African Cyprinoidei are still very limited. The theoretical part of this thesis is focused on (1) the species composition of Cyprinoidei on the African continent, the historical biogeopraphy of this freshwater fish group and (2) diversity and evolution of monogenean parasites of Cyprinoidei in Africa. The practical part of this thesis is focused on the morphological and molecular determination of cyprinoid Monogenea, the analysis of diversity of parasitic communities, basic epidemiological indicators and phylogenetic reconstruction of monogenean parasites of Cyprinoidei in Africa. Poděkování Na tomto místě bych chtěl poděkovat zejména vedoucí mojí diplomové práce prof. RNDr. Andree Vetešníkové Šimkové, Ph.D. za trpělivost, odborné vedení a cenné rady a připomínky, mojí konzultantce Mgr. Evě Řehulkové, Ph.D. a paní RNDr. Boženě Koubkové, Ph.D. za pomoc při determinaci parazitických skupin. Dále mé poděkování patří Mgr. Márii Seifertové, Ph.D. a Kristýně Koukalové za velkou pomoc při zpracování molekulárních dat a Mgr. Michalu Benovicsovi za pomoc se zpracováním fylogenetických rekonstrukcí. V neposlední řadě chci poděkovat rodině, přítelkyni a přátelům za podporu a pomoc v průběhu celého mého studia. Čestné prohlášení Prohlašuji, že jsem svoji diplomovou práci vypracoval samostatně pod vedením vedoucí práce s využitím informačních zdrojů, které jsou v práci citovány. V Brně dne: ……………………. Vojtěch Dlapka Obsah 1. Úvod a cíle práce .............................................................................................................. 11 2. Ichtyofauna Afriky ............................................................................................................ 12 2.1. Charakteristika a distribuce kaprovitých ryb Afrického kontinentu .......................... 12 2.2. Klasifikace a distribuce Cyprinoidei .......................................................................... 13 2.3. Biogeografie africké ichtyofauny a významné taxony Cyprinoidei .......................... 15 3. Paraziti kaprovitých ryb .................................................................................................... 18 3.1. Obecná charakteristika taxonu Monogenea ............................................................... 18 3.2. Hostitelská specifita ................................................................................................... 19 3.3. Diverzita parazitů kaprovitých ryb Afriky ................................................................. 20 3.4. Diverzita monogeneí kaprovitých ryb Afriky ............................................................ 20 4. Materiál a metodika .......................................................................................................... 23 4.1. Oblasti výzkumu, odlov ryb a přehled studovaných druhů taxonu Cyprinoidei........ 23 4.2. Parazitologická pitva ryb a fixace parazitů ................................................................ 25 4.3. Morfologická determinace parazitů ........................................................................... 26 4.4. Analýza molekulárních dat ........................................................................................ 27 4.4.1. Extrakce, amplifikace a sekvenování DNA ....................................................... 27 4.4.2. Fylogenetická analýza ........................................................................................ 32 4.5. Epidemiologické parametry ....................................................................................... 33 4.6. Analýza diverzity parazitických společenstev ........................................................... 33 5. Výsledky ........................................................................................................................... 36 5.1. Druhy Cyprinoidei parazitované zástupci třídy Monogenea ..................................... 36 5.2. Druhová diverzita parazitů třídy Monogenea ............................................................ 37 5.3. Epidemiologické parametry ....................................................................................... 41 5.4. Analýza diverzity parazitických společenstev ........................................................... 48 5.5. Fylogenetická analýza ................................................................................................ 51 6. Diskuze ............................................................................................................................. 59 7. Závěr ................................................................................................................................. 63 8. Literatura ........................................................................................................................... 64 Obrazová příloha ...................................................................................................................... 73 1. Úvod a cíle práce Monogenea jsou považována za jednu z druhově nejpočetnějších a nejrozšířenějších skupin rybích parazitů. Zástupci této třídy zpravidla patří mezi ektoparazity sladkovodních i mořských ryb, napadající zejména žábra, ploutve a povrch těla svých hostitelů. Monogenea se vyznačují velice úzkou hostitelskou specifitou (tj. jsou vázána na jednoho hostitele nebo úzké spektrum často příbuzných hostitelů), vysokou druhovou diverzitou a morfologickou adaptací přichycovacího orgánu, tzv. opisthaptoru, která je spojena právě s hostitelskou specifitou a specifitou na úrovni mikrohabitatu (selekce preferovaných pozic na žábrech ryb). Vzhledem k úzké hostitelské specifitě jsou Monogenea často využívána v evolučně ekologických studiích jako modelová skupina, a to pro studium parazito-hostitelských vztahů, koevoluce, morfologických adaptací parazitů a ekologie společenstev. Taxon Cyprinoidei je všeobecně považován za jednu z největších a nejdiverzifikovanějších skupin sladkovodních obratlovců a nejinak je tomu i na africkém kontinentu, kde jsou Cyprinoidei zastoupeni především čeledí Cyprinidae. Lze proto předpokládat, že vysoká diverzita hostitelů je spojená s vysokou diverzitou jejich hostitelsky specifických parazitů třídy Monogenea. V současné době je na afrických sladkovodních rybách známo přes 480 druhů monogeneí, avšak znalosti diverzity afrických monogeneí i evolučních vztahů mezi zástupci této výjimečné skupiny parazitů jsou stále limitovány a zasluhují další vědeckou pozornost. Cílem diplomové práce bylo studium diverzity a evoluce parazitů taxonu Monogenea u ryb taxonu
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