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Finální Verze 30.4. Tisk Odkazy OBSAH 1 ÚVOD A CÍLE PRÁCE............................................................................................ 3 2 PŘEHLED LITERATURY ....................................................................................... 4 2.1 Třída Monogenea van Beneden, 1858 .............................................................. 4 2.1.1 Základní charakteristika................................................................................ 4 2.1.2 Klasifikace .................................................................................................... 4 2.2 Ichtyofauna studovaného území ....................................................................... 7 2.3 Monogenea sladkovodních druh ů ryb povodí Nilu ........................................ 14 3 MATERIÁL A METODIKA .................................................................................. 24 3.1 Charakteristika studovaného území................................................................ 24 3.2 Metodika sb ěru dat.......................................................................................... 28 3.2.1 Vyšet řené hostitelské ryby.......................................................................... 28 3.2.2 Sb ěr a fixace monogení............................................................................... 30 3.2.3 Determinace nalezených monogeneí .......................................................... 30 4 VÝSLEDKY............................................................................................................ 33 4.1 Přehled nalezených monogeneí a jejich hostitel ů ........................................... 39 4.1.1 Annulotrema Paperna & Thurston, 1969 .................................................... 40 4.1.2 Bagrobdella Paperna, 1969......................................................................... 43 4.1.3 Bouxiella Euzet & Dossou, 1976................................................................ 49 4.1.4 Cichlidogyrus Paperna, 1960...................................................................... 51 4.1.5 Dactylogyrus Diesing, 1850 ....................................................................... 62 4.1.6 Dogielius Bychowsky, 1936....................................................................... 71 4.1.7 Enterogyrus Paperna, 1963......................................................................... 73 4.1.8 Heteronchocleidus Bychowsky, 1957 ........................................................ 77 4.1.9 Characidotrema Paperna & Thurston, 1968............................................... 78 4.1.10 Onchobdella Paperna, 1968........................................................................ 80 1 4.1.11 Protoancylodiscoides Paperna, 1969.......................................................... 83 4.1.12 Quadriacanthus Paperna, 1961................................................................... 86 4.1.13 Scutogyrus Pariselle & Euzet, 1995............................................................ 89 4.1.14 Schilbetrema Paperna & Thurston, 1968.................................................... 91 4.1.15 Synodontella Dossou & Euzet, 1993 .......................................................... 93 4.1.16 Diplectanum Diesing, 1858 ........................................................................ 96 4.1.17 Gyrodactylus Nordmann, 1832................................................................... 98 4.1.18 Heterobothrium Cerfontaine, 1895........................................................... 101 5 DISKUZE .............................................................................................................. 105 6 ZÁV ĚR.................................................................................................................. 108 2 1 ÚVOD A CÍLE PRÁCE Třída Monogenea p ředstavuje jednu z druhov ě nejbohatších skupin rybích cizopasník ů a pravd ěpodobn ě parazitických helmint ů v ůbec. Jsou to kosmopolitn ě rozší ření plošt ěnci (Platyhelminthes, Neodermata), kte ří zpravidla cizopasí na povrchu studenokrevných živo čich ů. Vyzna čují se p římým vývojovým cyklem a úzkou hostitelskou specifi čností. Úzká hostitelská specifi čnost se odráží p ředevším ve variabilit ě p řichycovacích orgán ů, což činí tuto skupinu velmi atraktivní pro studium morfologických adaptací a koevolu čních vztah ů mezi hostiteli a parazity. Řada druh ů je považována za zdroj sekundárních infekcí a je tak vnímána za d ůležité patogeny zp ůsobující úhyny ryb, zejména v akvakulturách. Tato diplomová práce se zabývá druhovou diverzitou monogeneí parazitujících na sladkovodních rybách povodí Nilu se zam ěř ením na Súdán. Dosud bylo v oblasti povodí Nilu zaznamenáno 139 druh ů t ěchto cizopasník ů, avšak v Súdánu pouze 4 druhy. Tento fakt napovídá, že Súdán dosud nebyl z hlediska t ěchto parazitických plošt ěnc ů p říliš intenzivn ě studován a práce tak nabízí originální výsledky rozši řující naše znalosti o druhové rozmanitosti monogeneí a jejich spektru hostitel ů obývajících povodí Nilu. Terénní expedice do Súdánu v roce 2010 umožnila získání rozmanitého a druhov ě bohatého materiálu monogeneí, který byl v rámci této diplomové práce morfo-metricky zpracován a taxonomicky vyhodnocen. CÍLE • Vypracovat literární rešerši týkající se druhové diverzity monogeneí parazitujících na rybách Súdánu a sestavit/aktualizovat check list (hostitel- parazit, parazit-hostitel) t ěchto cizopasník ů pro oblast povodí Nilu. • Sestavit p řehled sladkovodních druh ů ryb vyskytujících se na území Súdánu. • Ovládnout všechny pot řebné metody nezbytné pro sb ěr, fixaci a následné studium strukturální diverzity monogeneí. • Provést morfologicko-taxonomické zhodnocení monogeneí nalezených na rybách během terénního výzkumu v Súdánu (2010) a porovnat získané poznatky s dosavadním stavem poznání. 3 2 PŘEHLED LITERATURY 2.1 Třída Monogenea van Beneden, 1858 2.1.1 Základní charakteristika Monogenea pat ří mezi t ři základní skupiny nejodvozen ějších, výhradn ě parazitických plošt ěnc ů (Platyhelminthes), tj. do skupiny Neodermata (LOCKYER et al. , 2003 ). Tato třída zahrnuje p řevážn ě ektoparazitické plošt ěnce cizopasící na povrchu t ěla (k ůži, ploutvích a žábrách) mo řských, brakických a sladkovodních druh ů ryb. N ěkte ří zástupci však parazitují uvnit ř t ěla svých hostitel ů, nej čast ěji na místech, která se otevírají do vn ějšího prost ředí (nosní tká ň, trávicí a urogenitální systém) (WHITTINGTON & CHISHOLM, 2008). Všichni zástupci monogeneí jsou hermafrodité s p římým vývojovým cykle, bez ú časti mezihostitele. V ětšina z nich je vejcorodých, ale rovn ěž velká skupina monogeneí (v ětšina z čeledi Gyrodactylidae) je živorodá (BUCHMANN & BRESCIANI, 2006). Základní charakteristika, morfologie a životní cykly t ěchto cizopasník ů jsou uvedeny v literárním p řehledu bakalá řské práce (PRAVDOVÁ, 2013). V sou časné dob ě je popsáno kolem 4000 - 5000 druh ů monogeneí, což je vzhledem k bohatosti rybí fauny a vysoké specifi čnosti t ěchto cizopasník ů velmi malý po čet. Mnoho druh ů tak stále čeká na své objevení (WHITTINGTON, 1998). 2.1.2 Klasifikace Klasifikace monogeneí je dosud nejednotná. N ěkte ří auto ři rozd ělují t řídu Monogenea na podt řídu Monopisthocotylea a Polyopisthocotylea, zatímco jiní na podt řídu Polyonchoinea (18 čeledí) a Heteronchoinea (32 čeledí) (BOEGER & KRITSKY, 2001). Taxon Heteronchoinea sdružuje zástupce dvou d říve navržených podt říd, a to Oligonchoinea (30 čeledí) a Polystomatoinea (2 čeledi) (BOEGER & KRITSKY, 1993). Podt řída Polyonchoinea (= Monopisthocotylea) bývá ozna čována za nižší monogenea, zatímco podt řída Heteronchoinea (= Polyopisthocotylea) je považována za monogenea vyšší (BOEGER & KRITSKY, 1993). (OLSON & LITTLEWOOD, 2002) zkoumali fylogenezi monogeneí na základ ě srovnání sekvencí ribosomálních gen ů. Výsledná topologie fylogenetických strom ů z velké části odpovídala klasifikaci založené na morfologických znacích podle (BOEGER & 4 KRITSKY, 2001). V následujícím p řehledu jsou jednotlivé taxony řazeny dle systému uvedeného v pracích (BOEGER & KRITSKY, 1993). Podt řída: Polyonchoinea Bychowsky, 1937 (= Monopisthocotylea Odhner, 1912) Tato podt řída zahrnuje zpravidla dob ře pohyblivé ekto- a endoparazity p řevážn ě ryb, mén ě často paryb, obojživelník ů a bezobratlých (WHITTINGTON, 2005). Infikují širokou škálu mikrohabitat ů jako jsou pokožka, ploutve, žaberní a ústní dutina, nosní a oční epitel, urogenitální systém, t ělní dutina, trávicí trakt, srde ční svalovina a ob ěhový systém (WHITTINGTON & CHISHOLM, 2008). Živí se hlavn ě epiteliálními bu ňkami a pouze p říležitostn ě krví vytékající z hemoragií (BUCHMANN & BRESCIANI, 2006). Haptor se skládá z há čků a p řídatných sklerit ů (WHITTINGTON, 2005; WHITTINGTON & CHISHOLM, 2008). Marginálních há čků je obvykle p řítomno 14- 16 (v ětšinou 14). Larvy mají 2 páry o čí s čočkami. Zástupci podt řídy Polyonchoinea parazitují zejména na taxonech Agnatha, Chondrichthyes a Teleostei (BUCHMANN & BRESCIANI, 2006). Řády: Monocotylidea Lebedev, 1988 Capsalidea Lebedev, 1988 Montchadskyellidea Lebedev, 1988 Gyrodactylidea Bychowsky, 1937 Dactylogyridea Bychowsky, 1937 Podt řída Heteronchoinea Boeger & Kritsky, 2001 (= Polyopisthocotylea Odher, 1912) Tato podt řída sdružuje zástupce 2 d říve navržených podt říd Oligonchoinea a Polystomatoinea (BOEGER & KRITSKY, 1993). Infikují p řevážn ě žábry ryb a zpravidla (zvlášt ě v dosp ělosti) se nepohybují na větší vzdálenosti (WHITTINGTON & CHISHOLM, 2008). Na rozdíl od p ředešlé podt řídy se tito plošt ěnci živí p řevážn ě krví svých hostitel ů. Existují rovn ěž zástupci,
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