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Diptera) S Novými Poznatky O Identifikaci, Ekologii a Rozšíření Středoevropských Druhů

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Diptera) S Novými Poznatky O Identifikaci, Ekologii a Rozšíření Středoevropských Druhů MASARYKOVA UNIVERZITA Přírodovědecká fakulta Olga KOMZÁKOVÁ Studium fylogeneze evropských rodů čeledi Anthomyiidae (Diptera) s novými poznatky o identifikaci, ekologii a rozšíření středoevropských druhů Dizertační práce Školitel: Prof. RNDr. Rudolf Rozkošný, DrSc. Brno, 2012 Bibliografická identifikace Jméno a příjmení autora: Olga Komzáková Název disertační práce: Studium fylogeneze evropských rodů čeledi Anthomyiidae (Diptera) s novými poznatky o identifikaci, ekologii a rozšíření středoevropských druhů Název disertační práce anglicky: A study on the phylogeny of the European genera of Anthomyiidae (Diptera) with new data on identification, ecology and the distribution of the Central European species Studijní progarm: Biologie Studujní obor (směr), 15001V - Zoologie kombinace oborů: Školitel: Prof. RNDr. Rudolf Rozkošný, DrSc Rok obhajoby: 2012 Klíčová slova: Anthomyiidae, fylogenetická analýza, kladogram, molekulární analýza, fylogram, ovipositor, Botanophila, synekologické analýzy, checklist, rozšíření. Klíčová slova anglicky: Anthomyiidae, phylogenetic analysis, cladogram, molecular analysis, phylogram, ovipositor, Botanophila, synecological analysis, check list, distribution. © Olga Komzáková, Masarykova univerzita, 2012 Poděkování Na tomto místě bych ráda poděkovala zejména svému školiteli prof. Rudolfu Rozkošnému za vzorné odborné vedení, obrovskou trpělivost a ochotu. Velký dík patří i Dr. Verneru Michelsenovi, který mi byl nápomocen při determinacích obtížných druhů a při studiu entomologických sbírek v Zoologickém muzeu v Kodani. Také mi zapůjčil svou sbírku preparátů samičích kladélek mnoha evropských druhů, která se stala podkladem pro návrh determinačního klíče na samice rodu Botanophila. Při studiu tohoto rodu mi cennými radami pomohl také D.M. Ackland (Oxford). Dále jsem velice zavázána Dr. Andrei Tóthové za analýzu DNA květilek a Dr. Igoru Malenovskému a Dr. Marii Zhai za pomoc s dalšími analýzami. Prof. Miroslavu Bartákovi děkuji za plodnou spolupráci při psaní článků o květilkách a hlavně za pravidelný přísun bohatého sbírkového materiálu k determinaci. Dr. Jindřich Roháček (Opava) mi umožnil zpracování rozsáhlého materiálu z projektu „Poľana“. Dr. Františku Gregorovi (Brno) děkuji za svolení k reprodukci dvou jeho originálních ilustrací květilek. Závěrem chci samozřejmě poděkovat zejména svému manželovi, sestře a rodičům za psychickou podporu a trpělivost a rodičům manžela za čas věnovaný mé malé dceři, který mi umožnil připravit se na státní zkoušku a sepsat tuto práci. Studijní pobyt v Kodani byl financován z programu SYNTHESIS, projekt DK-TAF 0655 a DK-TAF 3147. Abstrakt Studium fylogeneze evropských rodů čeledi Anthomyiidae (Diptera) s novými poznatky o identifikaci, ekologii a rozšíření středoevropských druhů Olga Komzáková Čeleď květilkovití (Anthomyiidae) patří mezi početné a hospodářsky významné zástupce dvoukřídlého hmyzu (Diptera, Brachycera). Studium čeledi bylo v minulosti často opomíjeno z důvodu nesnadné druhové identifikace a absence určovacích klíčů na samice. První část předložené dizertační práce je věnována fylogenetickým vztahům mezi evropskými rody čeledi Anthomyiidae, které byly dosud studovány pouze klasickými metodami na základě morfologických znaků. V rámci morfologické analýzy je v dizertaci prověřeno 32 evropských rodů z celkového počtu 39, u každého je vyhodnoceno 39 znaků. Výsledkem analýzy morfologických znaků je konsenzuální fylogenetický strom, který v podstatě potvrdil úvahy o příbuznosti rodů v monografiích W. Henniga a G.C.D. Griffithse. Analýza molekulárních znaků byla provedena na sekvencích dvou mitochondriálních genů 12S, 16S a jednoho jaderného genu 28S, opět u vybraných druhů zastupujících 32 evropských rodů. Fylogramy sestavené pouze na základě získaných molekulárních znaků mnoho informací nepřinesly, protože podpořily navržené vztahy pouze mezi několika rody. Výsledkem kombinovaného vyhodnocení morfologických znaků a znaků zjištěných molekulární analýzou byly bayesiánskou metodou (BI) a metodou maximální parsimonie (MP) vytvořeny dva fylogramy. Větší množství informací poskytl fylogram vyhodnocený metodou MP. Monofyletické skupiny byly potvrzeny u skupin sdružených kolem rodů, Botanophila, Delia (včetně skupiny Hydrophoria), Pegomya a Pegoplata. Nejasné zůstává postavení rodů Acyglossa, Mycophaga a Paregle, stejně jako postavení několika dalších rodů, které vytvořily jen příbuzné dvojice bez dalších vztahů. Další studium příbuzenských vztahů mezi rody bude zřejmě vyžadovat analýzy většího počtu genů a vyjasnění vztahů k příbuzné čeledi Scathophagidae. Dále byl navržen determinační klíč na samice středoevropských druhů rodu Botanophila s využitím morfologických znaků na jejich ovipositorech a předložen návrh skupin středoevropských druhů tohoto rodu. Výsledky studie byly srovnány s návrhem příbuzenských skupin navržených D.M. Acklandem podle morfologických znaků na samčích termináliích. Znaky vyšetřených samičích ovipositorů v rámci středoevropských druhů potvrdily Acklandovo rozčlenění rodu na skupiny druhů brunneilinea, dissecta, fugax, gnava, seneciella, striolata a varicolor. U skupiny cordifrons je navrženo její rozdělení a druhy helviana a verticella byly povýšeny na samostatné skupiny. Na základě determinace sbírkového materiálu pocházejícího z šesti šumavských rašelinišť byly provedeny analýzy druhového složení s ohledem na dominanci, konstanci, sezónní výskyt a rozšíření druhů v rámci těchto lokalit. Studovaná společenstva květilek jsou porovnávána na základě počtu druhů, indexů druhové bohatosti, diverzity a equitability, koeficientů podobnosti a analýzou hlavních komponent (PCA). Nejpočetnějšími druhy byly Pegoplata aestiva, Delia platura, Hylemya nigrimana (eudominantní), Delia cardui a Botanophila fugax (dominantní), Delia cardui a Botanophila fugax se navíc s druhy Delia platura, Heterostylodes nominabilis a Hylemya vagans vyskytovaly na všech sledovaných lokalitách (eukonstantní). Výsledky dále ukázaly, že nejvyššího počtu druhů, jedinců a zároveň indexu druhové bohatosti bylo dosaženo při sběru pomocí žlutých misek. V rámci předložené dizertace byl také doplněn a aktualizován druhový seznam květilek České republiky a Slovenska. Počet druhů známých z České republiky tak vzrostl o 36 druhů a ze Slovenska o 12 druhů. Druhový seznam květilek České republiky je srovnán se seznamy sousedních států; čímž byly vymezeny druhy, které lze očekávat i na našem území. Klíčová slova: Anthomyiidae, fylogenetická analýza, kladogram, molekulární analýza, fylogram, ovipositor, Botanophila, synekologické analýzy, checklist, rozšíření. Abstract A study on the phylogeny of European genera of Anthomyiidae (Diptera) with new data on identification, ecology and distribution of Central European species Olga Komzáková Anthomyiidae is a numerous and economically important family of Diptera (Brachycera). In the past a study of this family was often neglected owing to difficult identification of species and absence of keys to females in many cases. The first part of the submitted dissertation is devoted to relationships among European anthomyiid genera that have been studied almost exclusively by classical methods based on morphological characters. In the framework of the morphological analysis, 33 European genera (from 39 genera in total) are examined and the character state is defined for each of the evaluated 39 characters. A consensual phylogenetic tree, as a result of morphological analysis, essentially confirmed considerations on generic relationships included in the W. Henning’s and G.C.D. Griffiths’ monographs. The analysis of molecular characters is based on sequences of two mitochondrial genes 12S, 16S and a nuclear gene 28S of 32 species representing European genera. The phylograms, formed base on molecular data only, provided little information about relationships among genera. Two phylograms resulted from the combined evaluation of morphological and molecular characters constructed by Bayesian (BI) and maximal parsimony (MP) methods. MP phylogram provided a larger amount of information. Monophyletic groups were accumulated around genera Botanophila, Delia (including Hydrophoria), Pegomya and Pegoplata. The phylogenetic position of Acyglossa, Mycophaga a Paregle is still unclear, as well as the position of several further genera forming only couples without additional relations. A further study of relationships among anthomyiid genera will apparently need an evaluation of further genes and elucidation of relations between Anthomyiidae and Scathophagidae as sister groups within Calyptratae. Female postabdominal sclerites of 33 Central European species of Botanophila were examined in detail and evaluated. Based on differences of these structural characters, the first identification key to females of this large genus can be compiled. Relationships proved among the examined females indicated distinct species groups. Results of this evaluation were compared with species groups proposed by D.M. Ackland based on structures of the male terminalia. Species groups based on female ovipositors confirmed the seven Acklands´species groups (brunneilinea, dissecta, fugax, gnava, seneciella, striolata and varicolor). The cordifrons group should be divided into two groups and helviana and verticella are each treated as a separate species group. The species composition of the family Anthomyiidae in six peat-bogs in the Šumava Mts is analysed with regards to dominance, constancy, seasonal occurrence and type of distribution of each species. Anthomyiid communities in all sites are
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