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Festuca Ovina and F MASARYKOVA UNIVERZITA V BRNĚ Přírodovědecká fakulta DISERTAČNÍ PRÁCE Petr ŠMARDA Brno, 2005 MASARYKOVA UNIVERZITA V BRNĚ Přírodovědecká fakulta Petr ŠMARDA BIOSYSTEMATICKÁ STUDIE STŘEDOEVROPSKÝCH KOSTŘAV FESTUCA SER. PSAMMOPHILAE PAWLUS. Disertační práce Školitel: doc. RNDr. Petr Bureš Ph.D. Brno, 2005 Bibliografická identifikace Jméno a příjmení autora: Mgr. Petr Šmarda Název disertační práce: Biosystematická studie středoevropských kostřav Festuca ser. Psammophilae Pawlus. Název disertační práce anglicky: Biosystematical study of Central European fescues Festuca ser. Psammophilae Pawlus. Studijní program: botanika Studijní obor (směr), kombinace oborů: Školitel: doc. RNDr. Petr Bureš Ph.D. Rok obhajoby: 2005 Klíčová slova v češtině: Festuca L., úzkolisté kostřavy, taxonomie, systematika, evoluce, karyologie, morfometrie, AFLP, obsah DNA, variabilita velikosti genomu, průtoková cytometrie, nomenklatura, rozšíření, střední Evropa, suché trávníky, skalní stepi, váté písky Klíčová slova v angličtině: Festuca L., fine-leaved fescues, taxonomy, systematics, evolution, karyology, multivariate morfometrics, AFLP, DNA content, genome size variability, flow cytometry, nomenclatory, distribution, Central Europe, dry grasslands, rocky steppes, wind-blown sands © Petr Šmarda, Masarykova univerzita v Brně, 2005 Poděkování: Za vedení práce a řadu cenných rad a námětů bych rád poděkoval bývalému i současnému školiteli doc. RNDr. Marii Dvořákové CSc. a doc. RNDr. Petru Burešovi Ph.D., jakož i mnoha dalším členům katedry. Svým spolužákům Tomáši Vymyslickému, Honzovi Rolečkovi, Katce Kočí, Kláře Helánové, Zuzce Rozbrojové, Katce Krátké, Janě Božkové a dalším vděčím za mnohé sběry kostřav a krásné chvíle na společných exkurzích. Výsledky předkládané práce byly finančně podpořeny z grantů MSM 143100010, FRVŠ 555-G4/2004, GAČR 206/03/0228, GAČR 206/98/1545, GAČR 524/05/H536 a MSM 0021622416. Abstrakt Festuca ser. Psammophilae Pawlus (Poeae, Poaceae) je kritickou skupinou středoevropských úzkolistých kostřav sekce Festuca. Jednotlivé taxony jsou si morfologicky velmi podobné a taxonomické postavení některých taxonů není zcela jasné. Klasickými karyologickými metodami nebo cytometricky byl zjištěn stupeň ploidie u několika set rostlin zahrnujících všechny sledované druhy a materiál z velké části jejich areálu. Na základě výsledků mnohorozměrných analýz morfologických znaků, studia AFLP polymorfizmu a studia typového materiálu bylo ve střední Evropě rozlišeno 6 druhů: F. pallens Host (reliktní skalní biotopy, diploid); F. csikhegyensis Simonk. (bazické skalní biotopy, tetraploid); F. psammophila (Čelak) Fritsch (kyselé písky) se dvěma poddruhy F. p. subsp. psammophila a F. p. subsp. dominii (Krajina) P. Šmarda nom. prov. (dolní Pomoraví, stř. a vých. Polsko); F. vaginata Willd. (bazické písky panonské oblasti); F. polesica Zapał. (přímořské a vnitro- zemské písečné duny); F. pseudovaginata Penksza (bazické písky panonské oblasti). K jednotlivým druhům jsou doplněny mapky rozšíření, determinační klíč, informace o typovém materiálu, exsikátových sbírkách, synonymice a hybridech. Diskutována je možná evoluce okruhu s ancestrální pozicí F. pallens a izolovaným postavením F. polesica. Při studiu byla objevena možnost cytometrického stanovení stupně ploidie ze suchého herbářového materiálu a snižování pozorovaného obsahu suchého materiálu oproti živým vzorkům. U F. pallens a několika dalších druhů byla zjištěna na vnitrodruhová variabilita v obsahu DNA. U F. pallens s.l. se tato variability projevuje na intrapopulační i lokálni úrovni a v rámci celého areálu se obsah DNA v rámci diploidů i tetraploidů liší až o 16,6 %. Obsah DNA obou ploidních je geograficky provázaný (P<0,001) a sleduje podobný geografický trend s rostlinami s větším genomem v jihovýchodní části areálu (P<0,01). U diploidů větší obsah DNA koresponduje s reliktním charakterem stanoviště (P<0,001) a typem vegetace na konci posledního glaciálu (P<0,05). Obsah DNA u diploidů se liší v jednotlivých fytogeo- grafických oblastech (P<0,001) a u tetraploidů mezi jednotlivými geografickými typy (P<0,001). Monoploidní obsah DNA je u tetraploidů nižší než u diploidů (P<0,001). Dissertation Abstract Festuca ser. Psammophilae Pawlus (Poeae, Poaceae) is the critical groups of Central European fine leaved fescues of sect. Festuca. Particular taxa are very close morphologically and taxonomic status of some taxa is still not clearly resolved. For several hundreds of plants, including all taxa studied and material from major parts of their distribution areas, ploidy level was estimated by classical karyological methods and flow cytometry. Based on the multivariate morphological analysis, AFLP analysis, and the study of type specimens, following six species were recognized: F. pallens Host (relict rocky habitats, diploid); F. csikhegyensis Simonk. (basic rocky habitats, tetraploid); F. psammophila (Čelak) Fritsch (acidic sands) with two subspecies F. p. subsp. psammophila and F. p. subsp. dominii (Krajina) P. Šmarda nom. prov. (SW Moravia, C and E Poland); F. vaginata Willd. (basic sands mainly in Pannonian region); F. polesica Zapał. (seaside and inland sand dunes); F. pseudovaginata Penksza (basic sands in Pannonian lowland). Determination key, distribution maps, informations about the type specimens, exsiccata collections, synonyms, and hybrids are provided. Evolutionary relationships with the assumed putative taxon F. pallens and the rather isolated position of F. polesica are discussed.. Within the study, flow cytometry was revealed to enable determination of ploidy level from dry herbarium material. Observed DNA content of herbarium specimens decrease in comparison with the fresh plants. Intraspecific variability of DNA content was observed in F. pallens and several further species. In F. pallens s.l., DNA content variability appears both on intrapopulation and landscape scales. Within the whole distribution area, at maximum 1.166 fold difference were observed both within the diploids and tetraploids. DNA content in both ploidy levels is spatially related (P<0.001) and reflects the same geographical gradient with plants with larger genomes growing in South-eastern part of the distribution area. In diploids, larger DNA content correlates with relict habitats (P<0.01) and character of palaeo-vegetation 20 000 BP (P<0.02). DNA content in diploids vary among phytogeographical regions (P<0.001), and among patricular tetraploid types (P<0.001). Relative monoploid DNA content of tetraploids is smaller than diploids (P<0.001). Obsah 1. Rod Festuca L. a současné názory na jeho evoluci a členění ............................................ 2 1.1 Popis........................................................................................................................... 2 1.2 Postavení v rámci čeledi Poaceae Barnhart............................................................... 3 1.3 Infragenerické členění ................................................................................................ 4 1.4 Evoluce rodu .............................................................................................................. 7 2. Festuca ser. Psammophilae ve střední Evropě................................................................. 10 2.1 Úvod......................................................................................................................... 10 2.2 Konspekt středoevropských druhů........................................................................... 10 2.3 Určovací klíč ............................................................................................................ 16 2.4 Evoluce a systematika okruhu.................................................................................. 17 3. Konspekt disertační práce ................................................................................................ 20 3.1 Cíle disertační práce................................................................................................. 20 3.2 Použité metody......................................................................................................... 20 3.3 Přehled publikovaných a k publikaci připravených prací ........................................ 20 3.4 Hlavní výsledky disertační práce ............................................................................. 22 3.5 Náměty k dalšímu studiu a rozpracovaná témata..................................................... 22 4. Seznam použité literatury................................................................................................. 25 (1) Šmarda P., Müller J., Vrána J. et Kočí K. (2005): Ploidy level variability of some Central European fescues (Festuca subg. Festuca L., Poaceae). – Biologia (Bratislava) 60: 25–36. (2) Šmarda P. et Kočí K. (2003): Chromosome number variability in Central European members of Festuca ovina and F. pallens groups (sect. Festuca). – Folia Geobot. 38: 65–95. (3) Grulich V., Krahulec F., Šmarda P. et Kočí K. (2002): Festuca. p. 828–834. In: Kubát K. [ed.], Klíč ke květeně České republiky. – Academia, Praha. (4) Šmarda P. (submitted): DNA ploidy levels of Romanian fescues (Festuca L., Poaceae), measured in living plants and herbarium specimens. – Folia Geobot. (5) Šmarda P., Šmerda J., Knoll A. et Bureš P. (in prep.): Revision of Central European taxa of Festuca ser. Psammophilae Pawlus – morphometrical, karyological and AFLP analysis. (6) Šmarda P. et Bureš P. (submitted): Intraspecific genome size variability in Festuca pallens on different geographical scales
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