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Jepice Hotovo S Opravou MASARYKOVA UNIVERZITA PŘÍRODOV ĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE Parthenogeneze jako rozmnožovací strategie u jepic (Ephemeroptera) Bakalá řská práce Jan Šupina Vedoucí práce: doc. RNDr. Sv ětlana Zahrádková, Ph.D. BRNO 2012 Bibliografický záznam Autor: Jan Šupina Přírodov ědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Parthenogeneze jako rozmnožovací strategie u jepic (Ephemeroptera) Studijní program: Bakalá řský studijní program Studijní obor: Systematická biologie a ekologie Vedoucí práce: doc. RNDr. Sv ětlana Zahrádková, Ph.D. Akademický rok: 2011/2012 Po čet stran: 51 Klí čová slova: nepohlavní rozmnožování, chov, embryonální vývoj, geografická parthenogeneze Bibliographic Entry Author: Jan Šupina Faculty of Science, Masaryk Univeristy Department of Botany and Zoology Title of thesis: Parthenogenesis as reproductive stategy of mayflies (Ephemeroptera) Degree programme: Bachelor's degree programme Field of study: Systematic Biology and Ecology Supervisor: doc. RNDr. Sv ětlana Zahrádková, Ph.D. Academic Year: 2011/2012 Number of Pages: 51 Keywords: asexual reproduction, rearing, embryonic development, geographic parthenogenesis Abstrakt V práci se zabývám jepicemi (Ephemeroptera), které se rozmnožují nepohlavn ě pomocí parthenogeneze (tychoparthenogeneze a obligátní parthenogeneze). Sou částí práce je literární rešerše, v ěnovaná shrnutí informací o tomto jevu, zejména pro druhy jepic uvád ěných z České republiky. Druhá část práce je zam ěř ena na metody studia partenogeneze a shrnuje publikované zkušenosti v této oblasti. Tato práce se dále zabývá publikovanými poznatky z laboratorního chovu jepic, a také poznatky mého experimentu-chovu druhu Baetis rhodani . Seznam jepic druh ů po celém sv ětě s výskytem partenogeneze je uveden vp říloze. Abstract In the present thesis I deal with mayflies (Ephemeroptera), which reproduce asexually by parthenogenesis (both tychoparthenogenesis and obligate parthenogenesis). The thesis includes a literature review summarizing available information about this phenomenon with focus on mayfly species reported from the Czech Republic. The second part of the thesis focuses on methods of study of parthenogenesis and summarises the published experience in this field. This work further deal with the published results from laboratory rearings of mayflies and presents results of my own experiment-the rearing of Baetis rhodani . A world list of mayfly species with documented occurrence of parthenogenesis is given in the attachment. Pod ěkování Na tomto míst ě bych cht ěl pod ěkovat vedoucí práce doc. RNDr. Sv ětlan ě Zahrádkové, Ph.D. za vedení, pomoc a trp ělivost p ři zpracovávání bakalá řské práce, Jan ě Zahrádkové za překlad st ěžejního díla profesora Charlese Degrange z francouštiny, a také Mgr. Markovi Poláškovi za pomoc p ři sb ěru materiálu a při řešení n ěkterých technických problém ů. Prohlášení Prohlašuji, že jsem svoji bakalá řskou práci vypracoval samostatn ě s využitím informa čních zdroj ů, které jsou v práci citovány. Souhlasím s uložením této bakalá řské práce v knihovn ě Ústavu botaniky a zoologie P řF MU v Brn ě, p řípadn ě v jiné knihovn ě MU, s jejím ve řejným půjčováním a využitím pro v ědecké, vzd ělávací nebo jiné ve řejn ě prosp ěšné ú čely, a to za předpokladu, že p řevzaté informace budou řádn ě citovány a nebudou využívány komer čně. Brno 7. kv ětna 2012 ............................ Jan Šupina Obsah 1. ÚVOD.....................................................................................................................................9 1. 1. Charakteristika řádu Ephemeroptera ..............................................................................9 1. 2. Zp ůsoby rozmnožování ................................................................................................10 1. 3. Parthenogeneze jako zp ůsob rozmnožování.................................................................10 2. VÝSKYT PARTHENOGENEZE U JEPIC.........................................................................13 2. 1. Ameletidae....................................................................................................................14 2. 2. Caenidae .......................................................................................................................15 2. 3. Baetidae ........................................................................................................................16 2. 4. Ephemerellidae .............................................................................................................20 2. 5. Ephemeridae .................................................................................................................22 2. 6. Heptageniidae ...............................................................................................................23 2. 7. Leptophlebiidae ............................................................................................................27 2. 8. Palingeniidae ................................................................................................................27 2. 9. Polymitarcyidae............................................................................................................28 2. 10. Siphlonuridae..............................................................................................................29 3. METODY STUDIA PARTHENOGENEZE U JEPIC ........................................................31 3. 1. Sestavení a testování metodiky chovu jepic v laboratorních podmínkách...................31 3. 1. 1. Získávání vají ček a jejich chov ............................................................................31 3. 1. 2. Chov larev ............................................................................................................32 3. 1. 3. Chov subimág.......................................................................................................33 3. 1. 4. Chovná za řízení....................................................................................................33 3. 1. 5. Vlastní chov..........................................................................................................35 4. MOŽNOSTI A CÍLE STUDIA PARTHENOGENEZE U JEPIC .......................................38 5. ZÁV ĚR.................................................................................................................................39 6. P ŘÍLOHA.............................................................................................................................40 7. SEZNAM POUŽITÉ LITERATURY..................................................................................45 8 1. ÚVOD 1. 1. Charakteristika řádu Ephemeroptera Řád jepice, latinsky Ephemeroptera , pat ří mezi nejstarší řády recentního pterygotního hmyzu. Tento hmyz s prom ěnou nedokonalou obývá stojaté i tekoucí vody. Ve světě je zatím známo 42 čeledí s více než 3000 druhy (BARBER-JAMES et al. 2008), z toho zhruba 350 druh ů se vyskytuje v Evrop ě a 670 v Severní Americe (BRITTAIN & SARTORI 2003). ZAHRÁDKOVÁ et al. (2009) uvádí, že v naší faun ě, tedy na území České republiky, je známo 107 druh ů jepic, pat řící do 30 rod ů a 16 čeledí. Tento řád obsahuje malé až st ředn ě velké druhy, jejichž velikost m ůže kolísat mezi 3–35 mm. T ělo dosp ělc ů je jemné, štíhlé a bez chloupk ů. Hlava je malá se dv ěma krátkými tykadly. Hru ď je ve srovnání s hlavou velká a zavalitá. Z hrudi vyr ůstají 2 páry blanitých k řídel, přičemž zadní k řídla jsou mnohem menší a s postupující specializací se siln ě zmenšují (Baetis ), n ěkdy až v nepatrné proužky ( Centroptilum ), nebo mizí úpln ě ( Cloeon , Caenis ). Jsou jemná, pr ůhledná, lesklá, bezbarvá, často lehce nažloutlá nebo nahn ědlá, n ěkdy hn ědě skvrnitá (LANDA 1969). Zade ček je tvo řen 10 články, kon čící dv ěma dlouhými tenkými št ěty, mezi nimiž bývá často dlouhé koncové vlákno-pašt ět. Nohy mohou být u n ěkterých druh ů redukované. Same čci mají první pár nohou delší než sami čky, které jim slouží k přichycení sami čky p ři rozmnožování. Tvary t ěla larev se mohou zna čně lišit, podle jejich přizp ůsobení se k r ůzným p řírodním podmínkám. LANDA (1969) popisuje stavbu t ěla larev, které se skládá ze z řeteln ě odd ělené hlavy, mohutné hrudi a z deseti článkového zade čku, na jehož posledním článku jsou dva št ěty a st řední pašt ět. Siln ě chitinizovaná hlava larev je na rozdíl od dosp ělc ů dob ře vyvinutá, je prognátní nebo ortognátní a široká jako hru ď. Hlava nese velké složené o či a dv ě jednoduchá o čka. Složené o či jsou vypouklé, oválné, s vnit řním okrajem často zarovnaným. Narozdíl od dosp ělc ů, u nichž je ústní ústrojí zakrn ělé (nedochází k přijímání potravy), je u larev tohoto řádu ústní ústrojí pln ě vyvinuté a p řizp ůsobené nap říklad k seškrabování nárostu rostlin na kamenech, nebo k pojídání detritu na dn ě. Larvy jepic dýchají vzdušnicovými (tracheálními) žábrami. Vzdušnice vybíhají vždy ze zadního kraje článku zade čku a žábry jsou tvo řeny párem lupínk ů anebo ke říčků vláken r ůzného tvaru a velikosti. Nejv ětší bývají žábry 2. a 3. pár ů, poslední pár bývá již velmi malý. Žaberní plátky však mohou plnit i jinou funkcí než dýchání, nap říklad u rodu Epeorus vytváří velkou přísavku, která umož ňuje larvám udržet se v prudce tekoucí vod ě. 9 BRITTAIN & SARTORI (2003) hovo ří o jepicích jako o ojedin ělém řádu v hmyzí říši, protože mají 2 ok řídlená stádia, stádium subimaga (polodosp ělce) a stádium imaga (dosp ělce). Subimago má všechny hlavní znaky dosp ělce. Celé
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