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Biologie, Vývoj a Zoogeografie Vybraných Saproxylických Skupin

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Biologie, Vývoj a Zoogeografie Vybraných Saproxylických Skupin 1 Masarykova univerzita Přírodovědecká fakulta Katedra zoologie a ekologie Biologie, vývoj a zoogeografie vybraných saproxylických skupin orientálních druhů čeledi Stratiomyidae Diplomová práce 2007 Prof. RNDr. R. Rozkošný, Dr. Sc. Alena Bučánková 2 Biologie, vývoj a zoogeografie vybraných saproxylických skupin orientálních druhů čeledi Stratiomyidae Abstrakt Je popsána morfologie, biologie a zoogeografie larev čtyř druhů z čeledi Stratiomyidae. Dva z nich, Pegadomyia pruinosa a Craspedometopon sp. n., patří do podčeledi Pachygasterinae, další dva, Adoxomyia bistriata a Cyphomyia bicarinata , patří do podčeledi Clitellariinae. Larvy byly sbírány Dr. D. Kovacem pod kůrou padlých stromů v Malysii a Thajsku. Saproxylický způsob života larev v rámci celé čeledi je zde diskutován jako původní stav. Byly vytypovány morfologické a biologické znaky larev s možným fylogenetickým významem a jejich platnost byla vyzkoušena v kladistických programech Nona a Winclada. Zjištěný fylogenetický vztah hlavních podčeledí odpovídá v podstatě současnému systému čeledi. Překážkou detailnímu vyhodnocení jsou zatím jen nedostatečné popisy larev a jejich malá znalost, zvláště v tropických oblastech. Biology, development and zoogeography of some saproxylic Oriental species of Stratiomyidae (Diptera) Abstract The morphology, biology and zoogeography of four larvae of Stratiomyidae are described. Two of them, Pegadomyia pruinosa , Craspedometopon sp. n. belong to the subfamily Pachygasteinae, the others, Adoxomyia bistriata and Cyphomyia bicarinata are placed to the subfamily Clitellariinae. The larvae were collected under the bark of fallen trees in Malaysia and Thailand by Dr. D. Kovac. The saproxylic habitat of stratiomyid larvae is discussed in this thesis as an original state. The morphological and biological characters of possible phylogenetic significance are evaluated and their value was verified with use of Nona and Winclada programs. The results essentially agree with a recent classification of the family. The unsatisfactory descriptions of larvae and poor knowledge of them especially in tropic areas limit the detailed assesment for the time being. 3 Prohlašuji, že jsem tuto diplomovou práci vypracovala samostatně a že jsem uvedla všechny literární prameny, ze kterých jsem čerpala informace použité v této práci. Souhlasím s uložením této diplomové práce v knihovně Ústavu botaniky a zoologie PřF MU v Brně, případně v jiné knihovně MU, s jejím veřejným půjčováním a využitím pro vědecké, vzdělávací nebo jiné veřejně prospěšné účely, a to za předpokladu, že převzaté informace budou řádně citovány a nebudou využívány komerčně. V Brně dne .............. ................................ Alena Bučánková 4 PODĚKOVÁNÍ Tato práce by nevznikla bez moudrého přístupu vedoucího mé diplomové práce Prof. RNDr. R. Rozkošného, Dr.Sc., který mi, kromě v této práci popisovaných larev, fotografií vyhotovených Dr. D. Kovacem a veškeré potřebné literatury, poskytl také své rozsáhlé znalosti a bohaté zkušenosti. Děkuji Mgr . I. Malenovskému, za to, že mě naučil pracovat s programy Nona a Winclada a byl ochoten věnovat svůj drahocenný čas diskuzím o problémech vzniklých při testování vytypovaných fylogenetických znaků. Dále děkuji Mgr. P. Pařilovi a RNDr. A. Tóthové Ph. D. za vyhotovení fotografické dokumentace a svému tatínkovi za pomoc při následné úpravě fotografií. 5 Pegadomyia pruinosa : 1 – samice, 2 – samec (kreslil Dr. F. Gregor) 6 OBSAH 1. ÚVOD .....................................................................................................................................7 1.1. Současný stav poznání čeledi ..........................................................................................8 1.2. Saproxylické druhy ........................................................................................................11 1.3. Obecná morfologie čeledi ..............................................................................................13 2. MATERIÁL A METODIKA ...............................................................................................18 2.1. Popisná část ...................................................................................................................18 2.2. Fylogenetická část .........................................................................................................19 3. VÝSLEDKY .........................................................................................................................23 3.1. Studovaná vývojová stádia orientálních saproxylických druhů ....................................23 3.1.1. Pegadomyia Kertész, 1916 ....................................................................................23 3.1.2. Craspedometopon Kertész, 1909 ............................................................................28 3.1.3. Adoxomyia Kertész, 1907 .......................................................................................32 3.1.4. Cyphomyia Wiedemann, 1819 ................................................................................36 3.2. Využití znaků larev pro fylogenetické úvahy ................................................................41 3.2.1. Použité znaky ..........................................................................................................41 3.2.2. Kladistická analýza .................................................................................................46 4. DISKUZE .............................................................................................................................48 4.1. Popisná část ...................................................................................................................48 4.2. Fylogenetická část .........................................................................................................49 5. ZÁVĚR .................................................................................................................................56 6. SEZNAM LITERÁRNÍCH PRAMENŮ .............................................................................57 7. PŘÍLOHY .............................................................................................................................65 7 1. ÚVOD Bráněnkovití (Stratiomyidae) představují rozsáhlou a značně rozmanitou čeleď dvoukřídlého hmyzu, která je s výjimkou polárních oblastí a nejvyšších hor rozšířená po celém světě. Zvláště některé tropické druhy jsou nápadné svými tykadly, výrůstky na hrudi a štítku, nápadným zbarvením a velikostí. Přesto, že nejběžnější evropské druhy byly popsány téměř před 250 lety, je zvláště tropická fauna bráněnek známá zatím jen nedostatečně. Woodley (2001) zaznamenal ve svém světovém katalogu čeledi 2651 druhů, ale současný výzkum zvláště v orientální a neotropické oblasti naznačuje, že odhad 3000 druhů je zcela reálný. Jen u nepatrného počtu druhů je známa také larva. Rozmanitá morfologie dospělců je doprovázená neobyčejnou rozmanitostí morfologie, biologie a vývoje larev. Všechny larvy se liší od příbuzných čeledí (s výjimkou čeledi Xylomyidae) základním stavebním plánem originálně utvářeného ústního ústrojí a tuhým, kožovitým integumentem, který je vyztužen uhličitanem vápenatým. Larvy čtyř druhů, zde popisovaných, byly nalezeny pod kůrou mrtvých stromů v Orientální oblasti. Proto je zde také pojednáno o saproxylickém způsobu života larev bráněnek, adaptacích na tento specifický habitat a o možné evoluční historii, spojené s tímto prostředím. V rámci dvoukřídlého hmyzu je oceňován význam této čeledi pro fylogenetické úvahy. Především utváření tykadel bylo studováno již několika autory s ohledem na vývojové tendence, které jsou doloženy postupnou redukcí článků a vývojem subapikální aristy. V rámci jedné čeledi se tak objevují tendence, které se později uplatnily v plné míře u mnoha čeledí fylogeneticky mladších. Podobný význam má rozmístění konstantních štětin (tzv. chaetotaxie) na těle larev, které se stalo významným kritériem příbuznosti jednotlivých morfobiologických skupin. Při zkoumání fylogenetických vztahů u holometabolního hmyzu je důležité zahrnout, jak znaky dospělců, tak znaky larev, neboť larvy a dospělci se vyvíjejí ve zcela odlišném prostředí a jsou tudíž vystaveni jiným selekčním tlakům. Při odděleném zkoumání fylogeneze dospělců a larev je tak možné získat i zcela odlišný fylogenetický systém. Cílem diplomové práce je popis dosud neznámých larev z Orientální oblasti, pojednání o saproxylickém způsobu života a vytypování vhodných znaků larev pro fylogenetické úvahy s jejich následným otestováním v kladistických programech Nona a Winclada. 8 1.1. Současný stav poznání čeledi Čeleď Stratiomyidae, je zastoupena více než 2650 druhy ve 375 rodech (Woodley 2001). Jen u zlomku druhů je však známa také larva (příloha 7) a u ještě menšího počtu druhů je popis dostatečně podrobný, aby mohl být využit pro fylogenetickou analýzu čeledi. Z palearktické oblasti je známo 426 druhů a největší počet popsaných larev. V rámci této oblasti podrobně shrnul poznatky o 160 evropských druzích a z toho 14-ti popsaných larvách Rozkošný (1982, 1983). V této monografii je kromě popisů larev a dospělců také nejúplněji zpracován vývoj, biologie a ekologie celé čeledi. Rody palearktické oblasti včetně larev jsou stručně komentovány v práci téhož autora (1997), který podrobně pojednal i o vodních druzích střední Evropy (2000). V sérii prací Krivosheiny (1965, 1975, 1976, 1977) a Krivosheina & Rozkošný (1990) jsou dány popisy mnoha larev podčeledi Pachygastrinae převážně z ruského Dálného Východu.
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