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Stanislav Knor Školitel

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Stanislav Knor Školitel UNIVERZITA KARLOVA V PRAZE PŘÍRODOV ĚDECKÁ FAKULTA KATEDRA ZOOLOGIE Interakce rostlin a hmyzu ve fosilním záznamu Bakalá řská práce Stanislav Knor Školitel: RNDr. Jakub Prokop, Ph.D. Praha 2008 Tímto prohlašuji, že jsem uvedenou práci vypracoval samostatn ě, s využitím uvedené citované literatury. V Praze dne 12. srpna 2008 Stanislav Knor 1 Abstrakt V sou časné dob ě je rozmanitost života na Zemi dána p ředevším druhovým bohatstvím hmyzu, cévnatých rostlin a jejich vzájemnými vztahy. Jejich po čátek spadá do doby p řed více než 400 milión ů let, kdy vznikly první suchozemské ekosystémy. Fosilní nálezy umož ňují geochronologické a taxonomické za řazení jednotlivých druh ů a jejich vývojových stádií. Přestože n ěkteré fosilní doklady jsou vzácné, existuje jich dostate čné množství k utvo ření relevantních záv ěrů. Studované fosilní nálezy související s koevolucí rostlin a hmyzu je možné roz členit následujícím zp ůsobem: poškození rostlinných tkání, koprolity, obsah trávící trubice, typy ústního ústrojí hmyzu a rozmnožovací orgány rostlin. Z t ěchto skupin pak mají pravd ěpodobn ě nejv ětší význam stopy poškození rostlinných tkání spole čně s morfologií hmyzího ústního ústrojí vytvá řející spojené funk čně potravní skupiny. Cílem bakalá řské práce je p ředstavení jednotlivých vývojových etap hmyzího ústního ústrojí, demonstrace n ěkterých p říklad ů herbivorních asociací hmyzu a rostlin, a to se zřetelem k jejich fylogenetickému postavení a stratigrafickému kontextu. Klí čová slova: Hmyz, cévnaté rostliny, fosilní asociace, herbivorie, typy ústní ústrojí, mina, hálka, poškození rostlin, koprolity, obsah trávící trubice Abstract Life biodiversity on the Earth is mainly represented by enormous richness of the vascular plants and insect species and their associations. These interactions date back more than 400 million years when the first terrestrial ecosystems appeared. Fossils enable geochronology dating and taxonomical attribution of various species and their ontogeny stages in evolution. In spite of considerable scarcity, many fossils allow to make reliable conclusions. Following types of evidence dealing with co-evolution of plant and insects are obviously distinguished: plant damages, coprolites, gut contents, insect mouthpart types and plant reproductive structures. Among them, the plant damage and corresponding functional feeding groups are of the greatest importance, although the knowledge of the mouthpart types and their evolution has also high predicative value. Aim of this thesis is review of the evolution of insect mouthpart types, showing some examples of mostly herbivore insects - vascular plants interactions including pollination and emphasis on the evolutionary aspects and their geochronology distribution. Key words: Insects, vascular plants, fossil association, herbivory, mouthpart types, mine, gall, plant damage, coprolites, gut content 2 Obsah Abstrakt 2 1. Úvod 4 2. Jednotlivé typy asociací a jejich doklady 5 2.1. Typy ústního ústrojí (Mouthpart classes) 5 2.1.1. Vývojová fáze I (spodní devon) 6 2.1.2. Vývojová fáze II (svrchní devon – svrchní karbon) 6 2.1.3. Vývojová fáze III (spodní perm – spodní trias) 7 2.1.4. Vývojová fáze IV (svrchní trias – spodní jura) 8 2.1.5. Vývojová fáze V (svrchní jura – eocén) 9 2.2. Potravn ě funk ční skupiny (Functional feeding groups) 9 2.2.1. Vn ější žír listoví (External foliage feeding) 9 2.2.2. Vnit řní žír tkání (Feeding on internal tissues) 13 2.2.2.1. Tvorba vpichu a následné sání (Piercing and sucking) 13 2.2.2.2. Vrtání (Boring) 13 2.2.2.3. Minující žír (Mining) 14 2.2.2.4. Tvorba hálek (Galling) 17 2.2.2.5. Žír semen (Seed predation) 20 2.2.3. Povrchové sání (Surface fluid feeding) 20 2.3. Koprolity (Coprolites) 22 2.4. Obsah trávící trubice (Gut content) 23 2.5 Ovipozice (Oviposition) 23 2.6. Polinace (Pollination) 24 3. Záv ěr 26 4. Pod ěkování 26 5. Literatura 27 6. P říloha 36 3 1. Úvod V sou časnosti je druhová a ekologická rozmanitost života na Zemi dána p ředevším vysokým po čtem druh ů cévnatých rostlin, hmyzu i ostatních členovc ů, a jejich vzájemnými vztahy. Studium t ěchto asociací je významným aspektem soudobého ekologického výzkumu, avšak soub ěžn ě nebyla v ěnována p řim ěř ená pozornost jejich vývoji v geologické minulosti (Labandeira, 2005). Tento nesoulad je zap řičin ěn p ředevším tím, že paleobiologické výzkumy se zam ěř ují hlavn ě na popis, systematiku a fylogenetické vztahy studovaných druh ů výše uvedených skupin, a jenom v mnohem menší mí ře se soust ředí na jejich paleoekologii. Teprve v posledních letech dochází v této problematice k zásadn ějšímu obratu a k tvorb ě řady studií, které je možno rozd ělit následujícím zp ůsobem: 1. Studium zabývající se jedním, vzácn ěji více druhy specifických asociací mezi hmyzem a hostitelskými rostlinami v rámci fosilního záznamu (Zhou a Zhang, 1989; Waggoner a Poteet, 1996). 2. Výzkum vždy jedné ze specifických, tzv. funk čně potravních skupin (functional feeding groups) a jejích ichnofosilií, jakými jsou nap říklad poz ůstatky min, hálek, vrtání nebo vn ějšího okusu na kompresních či permineralizovaných fosiliích, vyskytující se na širokém spektru rostlinných hostitel ů (Crane a Jarzembowski, 1980; Lang a kol., 1995). 3. Spíše vzácn ější typ studia, kde jeden druh nebo rod hostitelské rostliny vykazuje více typ ů asociací s r ůznými hmyzími herbivory. P říkladem m ůže být rod Psilophyton (DAWSON) HUEBER ET BANKS, z časného devonu (Trant a Gensel, 1985), pop ř. druh Psaronius chasei MORGAN, z konce svrchního karbonu (Labandeira, 1998a; Labandeira a Phillips, 1996). 4. Studie asociací cévnatých rostlin a hmyzu p říslušné lokality, zam ěř ené na jejich celkové kvalitativní i kvantitativní zhodnocení (Strauss, 1977; Scott a kol., 1985; Stephenson a Scott, 1992; Ash, 1997 resp. Labandeira a kol., 2002; Wilf a Labandeira, 1999; Wilf a kol., 2005; Beck a Labandeira, 1998). Na základ ě t ěchto studií z nejr ůzn ějších lokalit je možné vytvo řit v globálním m ěř ítku rámcový obraz paleoekologických vztah ů rostlin a hmyzu (pop ř. ostatních členovc ů) v období od nejsvrchn ějšího siluru do konce pleistocénu. Cílem této rešerše je, vzhledem k ší ři dané problematiky a omezenému rozsahu této práce, podat jen rámcový p řehled výsledk ů dosavadního výzkumu, a nastínit jeho možný vývoj a aplikace v budoucnosti. 4 2. Jednotlivé typy asociací a jejich doklady Jen nepatrnou část z popisovaných interakcí mezi rostlinami a hmyzem je možno ur čit na základ ě p římé asociace fosilního hmyzu s hostitelskou rostlinou in situ . Drtivá v ětšina výše zmín ěných asociací byla stanovena na základ ě nep římých d ůkaz ů. V zásad ě m ůžeme rozlišovat tyto doklady, a to koprolity, fosilizovaný obsah trávící trubice, poškození rostlinných tkání a morfologii fosilizovaných rostlin a hmyzu (Labandeira, 2002). U rostlin je to zejména utvá ření orgán ů spojených s reprodukcí (Friis, 1985; Crepet a Nixon, 1996) a možná i n ěkterých vegetativních struktur (Scott a Taylor, 1983; Niklas, 1986; Farrell a kol., 1991). V p řípad ě hmyzu se jedná p ředevším o utvá ření jednotlivých komponent ústního ústrojí a tvaru hlavy (Labandeira a Beal, 1990, Labandeira 1997) a v menší mí ře i kladélka (Shear a Kukalová-Peck, 1990). D ůležité jsou i povrchové struktury spojené s polina čním syndromem, jako jsou femorální korbikuly, z řetelná hrudní vestitura a t ělní kartá čky (Faegri a van der Pijl, 1980; Proctor a kol., 1996). V souvislosti s klasifikací jednotlivých typ ů asociací rozlišuje Labandeira (1997, 2002) t ři kategorie ekomorfologické disparity: (i) typy ústního ústrojí (mouthpart classes), (ii) funk čně potravní skupiny (functional feeding groups) a (iii ) potravní gildy (dietary guilds). Zjednodušen ě řečeno, první kategorie popisuje aparát, kterým hmyz přijímá potravu; druhá pak zp ůsob, jakým to činí; a t řetí, jakou potravu hmyz konzumuje. Pro pochopení vývoje vzájemných interakcí mezi rostlinami a hmyzem v geologické minulosti jsou d ůležité především první dv ě jmenované kategorie. 2.1. Typy ústního ústrojí (Mouthpart classes) Přestože k uchování tak delikátních a nepatrných struktur, jakými bývá hmyzí ústní ústrojí dochází i za výjime čně p říznivých okolností jen vzácn ě, existuje v sou časné dob ě, zejména z druhohorních uloženin (Grimaldi, 1990; Rayner a kol., 1991; Whalley a Jarzembowski, 1985), dostate čné množství dob ře zachovaných fosílií, které mimo jiné dokazují, že v ětšina sou časných typ ů ústního ústrojí má sv ůj p ůvod práv ě v druhohorách a objas ňují mnohé o evoluci, potravní strategii a mechanismech p říjmu potravy mnoha hmyzích skupin, recentních i vyhynulých. Labandeira (1997) rozlišuje celkem 34 typ ů ústního ústrojí (viz p říloha – obr. 1), p řičemž asi polovina je spjata s příjmem rostlinné potravy, k nim je pak nutno p řičíst ješt ě dva typy, též spojené s rostlinnou potravou a časov ě omezené na éru paleozoika. Podle Labandeiry 5 (1997) op ět asi polovina ústních typ ů vznikala a zanikala opakovan ě v pr ůběhu geologické minulosti. Typ ústního ústrojí vykazuje st ředn ě silnou až silnou vazbu na typ potravy. Zajímavé je, že zatímco druhová diverzita hmyzu má trvale vzestupnou tendenci, disparita ústního ústrojí se stabilizovala již n ěkdy v pr ůběhu st ředního mesozoika (Labandeira, 1997). To by vypovídalo o určitém typu ekologické saturace spíše než o plynulé expanzi potravních zdroj ů. Tento záv ěr potvrzují nezávisle na sob ě i údaje týkající se funk čně potravních skupin a potravních gild (Labandeira 2002). Pro evoluci ústního ústrojí je typický také opakovaný vznik haustelátních a obdobných struktur pro p říjem tekuté potravy u r ůzných skupin hmyzu, nap říklad u ploštic, motýl ů, dvouk řídlých a blanok řídlých, pokaždé modifikací jiných strukturálních komponent p ůvodního primitivního mandibulátního ústního ústrojí (Chaudonneret, 1990; Jervis a Vilhelmsen, 2000). Labandeira (1997) rozlišuje p ět vývojových fází v pr ůběhu geologické minulosti. První z nich spadá do spodního devonu, další do svrchního karbonu, t řetí do spodního permu, poslední dv ě pak do interval ů svrchní trias až spodní jura a svrchní jura až spodní k řída.
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