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Masarykova Univerzita Přírodovědecká Fakulta Ústav Botaniky a Zoologie Masarykova univerzita Přírodovědecká fakulta Ústav botaniky a zoologie Srovnání taxocenóz a vertikální stratifikace letové aktivity kůrovců (Coleoptera: Curculionidae: Scolytinae) v tvrdém luhu a horských jedlobučinách Diplomová práce Brno 2011 Autor: Jiří Procházka Vedoucí DP: Dipl. Biol. Jiří Schlaghamerský, Ph.D. 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 převzaté informace budou řádně citovány a nebudou vyuţívány komerčně. V Brně 10. 5. 2011 Poděkování Na tomto místě bych chtěl poděkovat dr. Jiřímu Schlaghamerskému za vedení mé práce, dr. M. Kníţkovi za pomoc s determinací materiálu, D. Hauckovi za pomoc při sběru dat, P. Průdkovi za pomoc s tříděním vzorků a P. Šebkovi za cenné rady, týkající se statistického hodnocení dat. Také bych rád poděkoval rodině, bez jejíţ podpory by tato práce nikdy nevznikla. Výzkum byl proveden v rámci projektu financovaného grantem GAAV KJB 600960705 a výzkumným záměrem MSM 0021622416. ABSTRAKT Společenstva kůrovců v oblasti soutoku Moravy a Dyje a v Beskydech byla studována prostřednictvím odchytu do nárazových pastí, uspořádaných do vertikálních transektů ( 0,5 aţ 21 m nad zemí). V oblasti Soutoku bylo uloveno 1428 jedinců, patřících 30 druhům, z nichţ čtyři byly v daném území nově zjištěné, šest druhů je zařazeno v červeném seznamu ohroţených druhů České republiky do kategorie zranitelný a tři druhy jsou v ČR nepůvodní. V Beskydech bylo zachyceno 6698 jedinců 27 druhů, z nichţ výskyt tří druhů byl v této oblasti dosud neznámý, jeden je klasifikován v červeném seznamu ohroţených druhů České republiky jako zranitelný a jeden je v ČR nepůvodní. Kůrovci tedy byli mnohem hojnější v Beskydech, avšak společenstvo Soutoku se vyznačovalo větší druhovou diverzitou. Byla vyhodnocena fenologie 16 nejhojnějších druhů. Pozorován byl monovoltinní i bivoltinní vývojový cyklus i sesterské generace. Nejvíce jedinců bylo zaznamenáno v 21 metrech, kdeţto druhově nejbohatší se ukázala výška 7 metrů nad zemí. Výšková preference jednotlivých druhů se však výrazně lišila, pravděpodobně kvůli různým potravním nárokům. Dále bylo potvrzeno, ţe většina kůrovců nepreferuje osvětlenější stanoviště. ABSTRACT Comparison of assemblages and vertical stratification of flight activity of bark beetles (Coleoptera: Curculionidae: Scolytinae) in hard-wood floodplain forests and montane fir-beech forests. Assemblages of bark and ambrosia beetles were assessed in the floodplain close to the confluence of the Morava and Dyje Rivers and in the Beskydy mountains (Czech Republic) by using flight interception traps arranged into vertical transects (0,5 to 21 metres above ground level). 1428 specimens belonging to 30 species were collected in the area of Confluence. Four of them are new for this area, six species are included in the Red list for Czech Republic as vulnerable and three species are considered as non-indigenous in the Czech Republic. 6698 specimens of 27 species were collected in the Beskydy mountains. Three of them are new for this area, one species is included in the Red list for Czech Republic as vulnerable and one species is considered as non-indigenous in the Czech republic. Scolytids were more abundant in the Beskydy mountains, while the assemblage was more diverse at the Confluence. Phenology was assessed for 16 of the most abundant species. Univoltine and bivoltine life cycles and sister generations were observed. Most of the specimens were recorded 21 m above ground, while most of the species were collected at 7 m. Flight activity of single species differed, probably due to food preferences. It was also confirmed that most of the bark and ambrosia beetles do not prefer insolated habitats. OBSAH 1. ÚVOD……………………………………………………………......…………….… 7 2. MATERIÁL A METODIKA…………………………………….…...….……….. 13 2.1. Charakteristika oblasti soutoku Moravy a Dyje………………………….… 13 2.2. Charakteristika lokalit v oblasti Soutoku………………....………..…...…... 15 2.3. Charakteristika Chráněné krajinné oblasti Beskydy……………..……...… 17 2.4. Charakteristika beskydských lokalit………………………………...…..….. 19 2.5. Metodika sběru dat……………………………………………………........… 22 2.6. Analýza dat…………………………………………………………..……...… 25 3. VÝSLEDKY…………………………………………………………………….…. 30 3.1. Zhodnocení účinnosti odchytu do nárazových pastí…………………...….... 30 3.2. Obecné charakteristiky společenstev kůrovců…...…………………..……... 32 3.3. Srovnání společenstev kůrovců na Soutoku a v Beskydech………..…....…. 33 3.4. Srovnání jednotlivých lokalit na Soutoku…………………...…………….... 40 3.5. Srovnání jednotlivých lokalit v Beskydech…………………………...…...… 42 3.6. Srovnání transektů na jednotlivých lokalitách na Soutoku a v Beskydech………………………………………………………….………..…. 44 3.7. Fenologie nejhojnějších druhů Soutoku…………………….…………..……45 3.8. Fenologie nejhojnějších druhů Beskyd………………………………...……. 50 3.9. Porovnání vertikálního rozmístění letové aktivity kůrovců na Soutoku a v Beskydech………………………………………………………………..... 56 3.10. Vertikální stratifikace letové aktivity nejhojnějších druhů Soutoku…….. 60 3.11. Vertikální stratifikace letové aktivity nejhojnějších druhů Beskyd…....... 66 3.12. Analýza hlavních komponent (PCA)…………………………………..…... 71 3.13. Testování vztahu kůrovců ke světelnosti……………………….……..….... 73 4. DISKUZE…………………………………………………………………...……... 74 4.1. Použití nárazových pastí………………………………………………..……. 74 4.2. Kůrovci v jihomoravských luzích…………………...………………...……... 74 4.3. Kůrovci v Beskydech…………………………………………….……..…….. 75 4.4. Srovnání taxocenóz kůrovců Soutoku a Beskyd……………………..……... 76 4.5. Srovnání výskytu kůrovců na jednotlivých lokalitách na Soutoku a v Beskydech……………………………….………………….…….…….... 78 4.6. Výskyt kůrovců v průběhu sezóny na Soutoku……………………...….…... 79 4.7. Výskyt kůrovců v průběhu sezóny v Beskydech…………………...…….…. 80 4.8. Vertikální distribuce kůrovců v porostu……………………………...…….. 80 4.9. Vztah kůrovců ke světelnosti……………………………….………..…......... 81 5. ZÁVĚR………………………………………………………………….…..……... 82 6. LITERATURA………………………………...……………………………..……. 83 7. PŘÍLOHY………………………………………………………………………….. 92 7 1. ÚVOD Rozrůznění nik je zajímavý způsob, jak se vyhnout kompetici. Jednou z moţností je časový posun, kdy si organismy nekonkurují kvůli asynchronnímu vývojovému cyklu. Např.kudlanka Tenodera sinensis se v oblasti společného výskytu s kudlankou náboţnou (Mantis religiosa) líhne o 2–3 týdny dříve, takţe její nymfy jsou vţdy větší, čímţ se úspěšně vyhýbají mezidruhové kompetici (Hurd & Eisenberg 1990). Další moţností, jak se vyhnout případnému konkurentovi je vertikální stratifikace, coţ znamená, ţe soutěţící druhy preferují různou výšku. Tento jev byl zaznamenán např. v tropickém deštném lese v Ekvádoru u sympatrického komplexu babočkovitých motýlů z podčeledi Ithomiinae. Výška, ve které motýli nejčastěji létali, se mezi druhy výrazně lišila a zároveň pozitivně korelovala s výškou, kde rostla hostitelská rostlina (Baccaloni 1997). Podobně dva druhy opylovačů v tropickém deštném lese obsadily dolní část koruny a jiné dva druhy opylovaly pouze květy v horní části (Perry 1984). Pokud ulovíme hmyz v určité výšce, nemusí to nutně znamenat, ţe se tam hledá potravu či se rozmnoţuje, ale můţeme takto zaznamenat i disperzi či migraci, ať uţ v preferované a nebo náhodné výšce nad zemí (Ulyshen & Hanula 2007). Typickým příkladem jsou švábi (Blattodea), u nichţ bylo zjištěno, ţe mobilnější druhy létají nejvýše, naopak špatní letci spíše při zemi. Zároveň ale samci vţdy létají výš neţ samice, aby tak lépe zachytili jejich feromony (Schal 1982). Aktivita ţivočichů v různých výškách porostu je do značné míry ovlivněna strukturou porostu. Podle Parkera (1995) se nejníţe nachází vrstva lesní hrabanky, nad níţ je bylinné patro, tvořené bylinnou vegetací a semenáčky dřevin. Následuje podrost, který je tvořen zejména křovinami a odrůstajícími stromy, na nějţ navazují niţší koruny stromů, které tvoří přechod mezi podrostem a plně či z větší části osluněnou hlavní úrovní korun. Horní část hlavní úrovně, plně exponovaná vnějším vlivům, se nazývá vnější koruna, nad níţ výjimečně vystupují pouze koruny nejvyšších stromů, neboli nadúroveň. Jennings et al. (1999) rozdělili koruny stromů podle osvětlení na dominantní patro, které vystupuje nad hlavní korunové patro a které je plně exponované slunečnímu záření shora i z boku, kodominantní patro, coţ je hlavní korunové patro, plně osvětlené shora, střední patro, které se nachází pod hlavním korunovým patrem, ale stále je alespoň částečně osvětleno, a spodní korunové patro, ke kterému se skrz předchozí patra dostává jen minimum světla. Pokud se zaměříme na hmyzí faunu v korunách stromů opadavého temperátního lesa, zjistíme, ţe převaţují síťokřídlí (Neuroptera) a ploštice (Heteroptera), významný podíl 8 mají i brouci (Coleoptera), přestoţe jde jen o několik málo druhů, např. kůrovců (Scolytinae) nebo drabčíků (Staphylinidae), jak zjistili např. Müller et al. (2007) a Gruppe et al. (2008). I kdyţ je mezi brouky spousta dobrých letců, zdá se, ţe většina druhů preferuje spíše podrost oproti koruně (Müller et al. 2007, Ulyshen & Hanula 2007 a Vodka et al. 2008). Je škoda, ţe tolik prací, zaměřených na prostorovou distribuci hmyzu, se omezuje pouze na vyšší taxonomické skupiny, jelikoţ aktivita konkrétních druhů v rámci skupiny se můţe velmi lišit (Ulyshen & Hanula 2007). I kdyţ se zdá, ţe většina druhů kůrovců je rovnoměrně distribuována napříč porostními patry s mírnou preferencí střední výšky (Wermelinger
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