Společenstva Parazitických Helmintů Vybraných Skupin Endemických Ryb (Cyprinidae a Leuciscidae) Západního Mediteránu

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Společenstva Parazitických Helmintů Vybraných Skupin Endemických Ryb (Cyprinidae a Leuciscidae) Západního Mediteránu PŘÍRODOVĚDECKÁ FAKULTA Společenstva parazitických helmintů vybraných skupin endemických ryb (Cyprinidae a Leuciscidae) západního Mediteránu Diplomová práce Tereza Štolfová Vedoucí práce: Mgr. Michal Benovics, Ph.D. Ústav botaniky a zoologie Brno 2021 Bibliografický záznam Autor: Bc. Tereza Štolfová Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Společenstva parazitických helmintů vybraných skupin endemických ryb (Cyprinidae a Leuciscidae) západního Mediteránu Studijní program: Ekologická a evoluční biologie Studijní obor: Zoologie Vedoucí práce: Mgr. Michal Benovics, Ph.D. Akademický rok: 2020/2021 Počet stran: 89 + 16 Klíčová slova: parazitičtí helminti; parazitická společenstva; endemismus; hostitelská specifita; Cyprinidae; Leuciscidae; Pyrenejský poloostrov; Mediterán Bibliographic Entry Author: Bc. Tereza Štolfová Faculty of Science, Masaryk University Department of Botany and Zoology Title of Thesis: Parasitic helminth communities of selected fish taxa (Cyprinidae and Leuciscidae) in the western Mediterranean region Degree programme: Ecological and Evolutionary Biology Field of Study: Zoology Supervisor: Mgr. Michal Benovics, Ph.D. Academic Year: 2020/2021 Number of Pages: 89 + 16 Keywords: parasitic helminth; parasitic communities; endemism; host specificity; Cyprinidae; Leuciscidae; Iberian Peninsula; Mediterranean Abstrakt Sladkovodní fauna Pyrenejského poloostrova je mimořádně bohatá na endemické druhy, které patří do dvou druhově vysoce diversifikovaných čeledí – Cyprinidae a Leuciscidae. Parazitofauna těchto ryb je však stále nedostatečně prozkoumaná, a proto se v předložené diplomové práci věnuji společenstvům parazitických helmintů endemických ryb z těchto čeledí. Parazitičtí helminti byli studováni z hlediska druhového složení a diverzity společenstev. Celkem bylo identifikováno 69 taxonů parazitických helmintů z 28 druhů Cyprinidae a Leuciscidae. Největší druhová diverzita byla pozorována u řádu Monogenea, který byl zastoupen 58 druhy. U většiny druhů rodu Dactylogyrus a Gyrodactylus byl pozorován vysoký stupeň endemismu, jelikož byli nalezeni pouze na sladkovodních rybách Pyrenejského poloostrova. V rámci rodu Gyrodactylus byla provedena fylogenetická rekonstrukce, která rozdělila pyrenejské zástupce do dvou dobře podpořených skupin. První skupina zahrnovala zástupce Gyrodactylus parazitující na endemických zástupcích čeledi Leuciscidae a druhá skupina zahrnovala zástupce Gyrodactylus parazitující na endemických zástupcích čeledí Leuciscidae a Cyprinidae. Porovnání parazitofauny Pyrenejského poloostrova a severní Afriky v rámci rodu Dactylogyrus ukázalo, že i přesto, že v severní Africe bylo doposud zaznamenáno 17 druhů Dactylogyrus, tak na Pyrenejském poloostrově najdeme více druhů v poměru na hostitele. Abstract The freshwater fish fauna of Iberian Peninsula is extremely rich in endemic species, which belong to the two highly diversified families – Cyprinidae and Leuciscidae. However, the parasite fauna of these fish is still underexplored, and therefore, the proposed thesis is dealing with parasitic helminth communities of this endemic fish species belonging to these two families. The species diversity and communities composition of parasitic helminths was investigated herein. A total of 69 taxa of parasitic helminth were identified from 28 host species belonging to Cyprinidae and Leuciscidae. The highest species diversity was observed within order Monogenea, which was represented by 58 species. A high degree of endemism was observed within majority of species from genera Dactylogyrus and Gyrodactylus, as they were found only on freshwater fish of Iberian Peninsula. A phylogenetic analysis aimed to assess relationships within the genus Gyrodactylus, and dividied the endemic Iberian Gyrodactylus into two well-supported clades; the first one encompassing Gyrodactylus parasitizing endemic fish species belonging to Leuciscidae and the second one including Gyrodactylus parasitizing endemic fish species belonging to Leuciscidae and Cyprinidae. Comparison of the species diversity of the genus Dactylogyrus in the Iberian Peninsula and the North Africa within the genus Dactylogyrus showed that despite the fact that 17 species of Dactylogyrus have been recorded in the North Africa so far, there is in general higher species diversity in relation to the host diversity in the Iberian Peninsula. Poděkování Poděkování Na tomto místě bych ráda poděkovala především vedoucímu práce Mgr. Michalovi Benovicsovi, Ph.D., za jeho trpělivost, velkou vstřícnost, pomoc při zpracování dat, neustálou motivaci a všechen čas, který mi věnoval při vedení práce. Dále bych chtěla poděkovat celému parazitologickému týmu z Masarykovy univerzity v Brně za sběr parazitologického materiálu. Dále mé poděkování patří týmu z AV ČR z Českých Budějovic, především RNDr. Anně Faltýnkové, Ph.D., za pomoc při zpracování a determinaci metacerkárií a také prof. RNDr. Tomáši Scholzovi, CSc., za jeho čas a pomoc při determinaci tasemnic. V neposlední řadě bych ráda poděkovala svému příteli, rodině a přátelům za jejich podporu a trpělivost po celou dobu mého studia. Prohlášení Prohlašuji, že jsem svoji diplomovou práci vypracovala samostatně s využitím informačních zdrojů, které jsou v práci citovány. Brno 31. května 2021 …………………………………. Tereza Štolfová Obsah 1 Úvod ............................................................................................................................. 11 2 Literární přehled zkoumané problematiky ................................................................... 12 2. 1 Základní charakteristika zájmové oblasti .............................................................. 12 2. 2 Ichtyofauna Pyrenejského poloostrova ................................................................ 15 2. 2. 1 Charakteristika a distribuce Cyprinoidei ....................................................... 15 2. 2. 2 Charakteristika a distribuce ryb Pyrenejského poloostrova ......................... 16 2. 2. 3 Původ endemismu na Pyrenejském poloostrově ......................................... 19 2. 3 Paraziti kaprovitých ryb ........................................................................................ 21 2. 3. 1 Druhová diverzita parazitů ............................................................................ 21 2. 3. 2 Diverzita parazitů kaprovitých ryb Pyrenejského poloostrova ..................... 22 2. 3. 3 Hostitelská specifita parazitů ........................................................................ 31 2. 4 Cíle diplomové práce ............................................................................................ 32 3 Materiál a metodika ..................................................................................................... 33 3. 1 Materiál a vymezení studovaného území ............................................................. 33 3. 2 Parazitologická pitva a fixace parazitů ................................................................. 35 3. 3 Morfologická identifikace ..................................................................................... 35 3. 4 Úrovně hostitelské specifity ................................................................................. 36 3. 5 Analýza molekulárních dat ................................................................................... 36 3. 5. 1 Extrakce, amplifikace a sekvenování DNA .................................................... 36 3. 5. 2 Fylogenetická analýza ................................................................................... 37 3. 6 Epidemiologické parametry .................................................................................. 42 3. 7 Analýza parazitických společenstev ...................................................................... 42 4 Výsledky ........................................................................................................................ 44 4. 1 Druhová diverzita parazitických helmintů ............................................................ 44 4. 1. 1 Hostitelská specifita parazitů kaprovitých ryb Pyrenejského poloostrova ... 45 4. 2 Epidemiologie parazitů ......................................................................................... 49 4. 3 Analýza diverzity parazitických společenstev ....................................................... 57 4. 4 Fylogenetická analýza ........................................................................................... 59 5 Diskuze.......................................................................................................................... 62 6 Závěr ............................................................................................................................. 74 7 Literatura ...................................................................................................................... 76 8 Přílohy ........................................................................................................................... 90 1 Úvod Strategie parazitického života se vyvinula u mnoha odlišných skupin organismů. Vzhledem k tomu, že parazité mají většinou negativní účinky na své hostitele, je důležité poznat jejich druhovou rozmanitost a co ji ovlivňuje. Obecně je druhová rozmanitost parazitů spojena s druhovou rozmanitostí jejich hostitelů (Kamiya et al., 2014). V současné době oblast střední Evropy disponuje kvalitními informacemi ohledně druhového složení parazitů sladkovodních ryb (Moravec, 2001). Za méně prozkoumanou se považuje
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