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Název Práce: Plžů MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE ŽIVOTNÍ CYKLUS A POPULAČNÍ DYNAMIKA SUCHOZEMSKÝCH PLŽŮ Bakalářská práce Denisa Pavézková Vedoucí práce: doc. RNDr. Michal Horsák, Ph.D. Brno 2014 Bibliografický záznam Autor: Denisa Pavézková Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Životní cyklus a populační dynamika suchozemských Název práce: plžů Studijní program: Ekologická a evoluční biologie Studijní obor: Zoologie Vedoucí práce: doc. RNDr. Michal Horsák, Ph.D. Akademický rok: 2013/2014 Počet stran: 60+4 Klíčová slova: Suchozemští plži; životní cyklus; reprodukce; životní strategie; r/K-strategie; délka života; věková struktu- ra; Vertigo Bibliographic Entry Author Denisa Pavézková Faculty of Science, Masaryk University Department of botany and zoology Life cycle and population dynamics of terrestrial Titleof Thesis: gastropods Degreeprogramme: Ecology and evolutionary biology Fieldof Study: Zoology Supervisor: doc. RNDr. Michal Horsák, Ph.D. AcademicYear: 2013/2014 NumberofPages: 60+4 Keywords: Terrestrial gastropods; life cycle; reproduction; life history strategies; r/K-strategy; life span; age structu- re; Vertigo Abstrakt Suchozemští plţi jsou skupina hermafroditních ţivočichů obývající širokou škálu stanovišť. Podle prostředí, ve kterém se vyskytují, uplatňují rozdílné ţivotní strategie. Tato práce shrnuje základní poznatky o ţivotním cyklu suchozemských plţů, samostat- ně pak dosavadní informace o počtu a velikosti vajíček, délce reprodukční sezóny, vej- corodosti nebo vejcoţivorodosti, semelparii nebo iteroparii, délce ţivotního cyklu a délce ţivota u suchozemských plţů, vyskytujících se i na území České republiky. Praktická část je zaměřena na stanovení věkové struktury u rodu vrkoč (Vertigo). Díky korozním znakům na ulitě je moţné určit stáří jedince s přesností na roky. Jedinci stu- dovaných druhů (Vertigo alpestris, V. angustior, V. antivertigo, V. substriata) se doţí- vají aţ tří let, nejčastěji byli zastoupeni jedinci ve věku 6 aţ 18 měsíců. Práce rozšiřuje dosavadní poznatky o délce ţivota těchto plţů, na základě populací z území České a Slovenské republiky. Abstract Terrestrial gastropods are a group of hermaphroditic animals inhabiting wide range of habitats. According to the environment which they live in, they apply different life history strategies. This thesis summarizes a basic knowledge about the life cycle of ter- restrial gastropods. Furthermore, it presents information about the species of the terres- trial gastropods which are known from the Czech Republic, number and size of their eggs, length of reproduction season, oviparity or ovoviviparity, semelparity or iteropar- ity, length of the life cycle and life span. The second part is focused on determining the age structure of four Vertigo species. Thanks to the corrosion signs at the shell, it is pos- sible to determine the age of individuals with the accuracy of years. The individuals of the studied species (Vertigo angustior, V. alpestris, V. substriata, V. antivertigo) lived up to three years. Mostly frequent individuals were from the age of six to the eighteen months. This thesis adds to the growing body of literature about the life span of the small gastropods based on the populations from the Czech and the Slovak Republic. Poděkování Na prvním místě bych ráda poděkovala vedoucímu práce doc. RNDr. Michalovi Horsákovi, PhD. za rady, opravy, čas, který mi věnoval, ochotu a hlavně za trpěli- vost. Dále pak Honzovi za pomoc s formátováním textu, za podporu a povzbuzování a všem, co si na mě při psaní vzpomněli nebo mi jakkoliv pomohli. Děkuji. Čestné prohlášení Souhlasím s uloţením této bakalářské práce v knihovně Ústavu botaniky a zoologie PřF MU v Brně, případně v jiné knihovně MU, s jejím veřejným půjčováním a vyuţitím pro vědecké, vzdělávací nebo jiné veřejně prospěšné účely, a to za předpokladu, ţe pře- vzaté informace budou řádně citovány a nebudou vyuţívány komerčně. Prohlašuji, ţe jsem svoji bakalářskou práci vypracovala samostatně s vyuţitím in- formačních zdrojů, které jsou v práci citovány. Brno 28. dubna 2014 ……………………………… Jméno Příjmení Obsah 1 ÚVOD 9 2 ŽIVOTNÍ CYKLUS 11 2.1 Samooplození a páření ...................................................................................... 11 2.2 Páření ................................................................................................................ 13 2.2.1 Opakované páření ..................................................................................... 15 2.2.2 Šípy lásky ................................................................................................. 16 2.3 Kladení vajíček ................................................................................................. 17 2.3.1 Počet a velikost vajíček ............................................................................ 19 2.4 Oviparie a zadrţování vajíček .......................................................................... 21 2.5 R a K-strategie .................................................................................................. 23 2.5.1 Semelparie a iteroparie ............................................................................. 24 2.6 Délka ţivota ...................................................................................................... 26 2.7 Dosavadní poznatky o ţivotních cyklech středoevropských suchozemských plţů ................................................................................................................... 29 2.7.1 Diskuze ..................................................................................................... 30 3 STANOVENÍ VĚKOVÉ STRUKTURY RODU VERTIGO 32 3.1 Metodika a materiál .......................................................................................... 32 3.1.1 Materiál .................................................................................................... 32 3.1.2 Odběr a zpracování vzorků ...................................................................... 33 3.2 Výsledky ........................................................................................................... 36 3.3 Diskuze ............................................................................................................. 49 4 ZÁVĚR 52 5 LITERATURA 53 6 PŘÍLOHY 61 8 1 ÚVOD Ţivotní cyklus je proces opakující se generace po generaci, kterým prochází všechny ţivé organismy. Lze jej definovat jako sérii postupných změn, kterými organismus pro- chází od narození po vznik jeho potomků (BELL & KOUFOPANOU 1991). Populační dynamika vypovídá o velikostních a věkových změnách celé populace. Působením vnitřních a vnějších vlivů dochází k uplatňování takových strategií, které zajistí repro- dukci organismu, a zároveň co nejvyšší moţnou plodnost, při ohledu na ţivotní projevy jedince. Proto je moţné i mezi plţi pozorovat K stratégy, kteří zvyšují svoji reprodukční zdatnost opakovanou reprodukcí během delšího ţivota. A r stratégy, u kterých je vlivem nestabilního vnějšího prostředí nejlepší strategií, vloţit většinu energie do jedné masové reprodukce (semelparie), ovšem na úkor budoucího přeţití (PIANKA 1970). Dalšími strategiemi, které jsou mezi suchozemskými plţi uplatňovány i podle vnějších podmí- nek, jsou oviparie a zadrţování vajíček. Při vejcorodosti jsou kladena vajíčka ihned po jejich vytvoření, naopak při zadrţování vajíček můţe v extrémním případě dojít k jejich vylíhnutí uvnitř rodiče, nebo neprodleně po nakladení, coţ se označuje jako vejcoţivo- rodost (TOMPA 1979a). Měkkýši tvoří jednu z nejpočetnějších skupin ţivočichů s 117 358 druhy (ZHANG 2011). Většina suchozemských plţů patří do skupiny Stylommatophora, která pojímá 1 400 rodů z 60 čeledí. V České republice je známo 221 druhů plţů, z toho 171 sucho- zemských (HORSÁK et al. 2013). Plicnatí plţi jsou hermafroditi, produkující samčí i samičí gamety. Během ţivota u nich můţe docházet k časově oddělenému dozrávání samčích a samičích gamet. Při rozmnoţování většinou kopulují s jedinci stejného druhu, s nimiţ si vyměňují spermie a tím i genetický materiál. Jejich reprodukci ovlivňují nejen vnitřní faktory, jako napří- klad produkce hormonů, ale také podmínky vnějšího prostředí, kterými mohou být tep- lota, nadmořská výška, vlhkost a populační hustota. Díky podmínkám, ve kterých se vyskytují a zdrojům, které mají k dispozici, vznikla vysoká variabilita ţivotních strate- gií, které zvyšují jejich úspěšnost při rozmnoţování, růstu a přeţívání. Ţivotní cyklus plţů je dobře prostudované téma, většinou však na přehledu několika vybraných druhů. Mezi nejčastěji popisované suchozemské druhy, které se vyskytují i na území České republiky, patří plamatka lesní (Arianta arbustorum). První část rešeršní práce je zamě- 9 řena na shrnutí dostupných obecných informací o ţivotním cyklu suchozemských plţů. Zahrnuje informace o hermafroditismu a s ním spojené samooplození a páření, známé skutečnosti o výběru partnera, ţivorodosti a vejcoţivorodosti, velikosti a počtu vajíček, reprodukčních strategiích semelparii a iteroparii a délce ţivota. Druhá část je věnována srovnávacím poznatkům o rozmnoţování druhů, které se vyskytují i v naší fauně, a případně i jejich populační dynamice. Vzhledem k tomu, ţe konkrétní délka ţivota není známa u mnoha druhů plţů, je na základě nedávno publikované studie o biologii několika druhů skupiny Vertiginidae (MYZYK 2011) postavena praktická část bakalářské práce, jejím cílem je popsat délku ţivota čtyř vybraných druhů rodu Vertigo. Hlavním určujícím faktorem pro stanovení stáří jedince,
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