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Koexistence a Rozdělení Niky U Pavouků Rodu Philodromus

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Koexistence a Rozdělení Niky U Pavouků Rodu Philodromus Masarykova univerzita Přírodovědecká fakulta Ústav botaniky a zoologie Koexistence a rozdělení niky u pavouků rodu Philodromus Diplomová práce Autor: Radek Michalko Brno 2012 Vedoucí DP: doc. Mgr. Stano Pekár Ph.D. 1 Souhlasím s uloţením této diplomové 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 8.1.2012 ………………………………… Podpis 2 PODĚKOVÁNÍ Zejména bych chtěl poděkovat vedoucímu mé diplomové práce panu docentu Stanu Pekárovi, ţe mi umoţnil pracovat na tomto tématu, za trpělivé vedení a uţitečné rady. Dále bych chtěl velice poděkovat mým rodičům, bez jejichţ osobní a finanční podpory by tato práce nevznikla. Rovněţ bych chtěl poděkovat Lence Sentenské, Evě Líznarové, Pavlovi Šebkovi a Stanovi Korenkovi za podporu a cenné rady všeho druhu. 3 ABSTRAKT Koexistence a rozdělení niky pavouků rodu Philodromus V této diplomové práci byl zkoumán mechanismus umoţňující koexistenci mezi Philodromus albidus, P. aureolus a P. cespitum. Studie probíhala na území významného krajinného prvku U Kříţe v Brně Starém Lískovci. Studované území se skládá ze třech typů biotopů: listnatý les, křoviny a monokultura švestek. Pavouci byli získáváni pomocí sklepávání. U zkoumaných druhů byly porovnávány různé dimenze niky. Prostorová nika byla zkoumána na základě mikro- aţ makrobiotopových preferencí. Trofická nika byla zkoumána na základě velikosti a typu přirozené kořisti a pomocí laboratorních experimentů potravních preferencí. Časová nika byla zkoumána na základě fenologie jednotlivých druhů. Studované druhy se lišily v prostorové a trofické nice. Přestoţe se všechny tři druhy vyskytovaly ve všech třech typech biotopů, tak kaţdý druh preferoval jiný. P. albidus a P. aureolus se vyskytovali na vegetaci výše nez P. cespitum. P. albidus lovil menší kořist neţ ostatní dva druhy. Na koexistenci se podílelo jednak rozdělení niky podle zdrojů a jednak prostorový efekt zásoby. 4 ABSTRACT Coexistence and niche partitioning in spiders of the genus Philodromus This thesis investigates the possible mechanisms of coexistence among three foliage spider species: Philodromus albidus, P. aureolus and P. cespitum. The study took place in an abandoned orchard in Brno. The study area consists of three habitats: deciduous forest, scrub and plum tree monoculture. Spiders were collected by beating shrubs and trees. The micro – macrohabitat preferences were investigated in order to find differences in the spatial niche. The differences in the trophic niches were studied by analyzing the actual and potential prey types and sizes. The natural prey analyses were supplemented with the laboratory experiments on prey preferences. The differences in the temporal niche were studied by comparing their phenologies. The studied species differed in their spatial and trophic niches. All species occurred in all habitats, but each species preferred different one. P. albidus and P. aureolus also occurred higher on the vegetation than P. cespitum. P. albidus utilized smaller prey than the other two species. The coexistence is mediated by two mechanisms: the resources partitioning and spatial storage effect. 5 OBSAH 1. ÚVOD……………………………………………………………………………………………………………….6 1.1. Ekologická nika a její evoluce……………………………………………………………………...6 1.2. Koexistence druhů…………………………………………………………………………………...9 1.2.1. Mechanismy nezávislé na fluktuaci podmínek: Diferenciace nik podle zdrojů…….10 1.2.2. Mechanismy závislé na fluktuaci podmínek prostředí………………………………..11 1.2.3. Neutrální teorie biodiverzity a biogeografie……………………………………………14 1.3. Koexistence mezi pavouky……………………………………………………………………….15 1.3.1. Rozdělení trofické niky…………………………………………………………………...16 1.3.2. Rozdělení prostorové niky……………………………………………………………….18 1.3.3. Rozdělení časové niky…………………………………………………………………...19 1.4. Cíle práce......................................................................................................................20 2. MATERIÁL A METODIKA……………………………………………………………………………………...21 2.1. Studované druhy……………………………………………………………………………………21 2.2. Oblast výzkumu……………………………………………………………………………………..21 2.3. Prostorová nika……………………………………………………………………………………..24 2.3.1. Biotopové preference…………………………………………………………………….24 2.3.2. Meso-Mikrobiotop………………………………………………………………………...25 2.4. Trofická nika…………………………………………………………………………………………26 2.4.1. Přirozená a potencionální kořist………………………………………………………...26 2.4.2. Potravní preference………………………………………………………………………28 2.5. Časová nika………………………………………………………………………………………….29 3. VÝSLEDKY……………………………………………………………………………………………………….30 3.1. Prostorová nika……………………………………………………………………………………..30 3.1.1. Biotopopové preference………………………………………………………………….30 3.1.2. Meso-mikrobiotop………………………………………………………………………...36 3.2. Trofická nika…………………………………………………………………………………………39 3.2.1. Přirozená a potencionální kořist………………………………………………………...39 3.2.2. Potravní preference………………………………………………………………………45 3.3. Časová nika………………………………………………………………………………………….49 4. DISKUZE………………………………………………………………………………………………………….51 4.1. Rozdělení niky u Philodromus spp………………………………………………………………51 4.2. Mechanismus koexistence Philodromus spp………………………………………………....58 5. LITERATURA…………………………………………………………………………………………………….62 6 1. ÚVOD 1.1. Ekologická nika a její evoluce Pojem nika poprvé pouţil GRINNEL (1917), v jehoţ podání nika znamená místo, které určitý organismus v přírodě obývá tedy v podstatě nároky, které vyţaduje pro svůj výskyt. Další koncept niky poskytl ELTON (1927), který ji naopak chápal jakoţto funkci, kterou určitý druh v přírodě plní (např. opylovač). První oficiální koncept ekologické niky formuloval HUTCHINSON (1957). Ekologická nika je podle něj celkový soubor poţadavků na zdroje a tolerancí k podmínkám (biotickým i abiotickým), které jedinci určitého druhu potřebují k ţivotu respektive k udrţení ţivotaschopné rostoucí populace. Nika je mnohorozměrný prostor tzn., ţe kaţdý organismus má různé tolerance k mnoha různým podmínkám (teplota, vlhkost, zastínění aj.) a zároveň vyuţívá mnoho různých zdrojů (prostor, potrava atd.). Jednotlivé podmínky a zdroje nutné pro přeţití daného druhu v prostředí vytváří osy niky. CHASE & LEIBOLD (2003) definují niku jako spojení podmínek prostředí, které umoţňují druhu uspokojit jeho minimální poţadavky tak, aby natalita byla rovna nebo vyšší neţ mortalita, a vlivu tohoto druhu na tyto podmínky. Tzn. nejen poţadavky druhu pro jeho existenci v daném prostředí, ale zároveň i vliv tohoto druhu na dané prostředí. Např. kdyţ určitý druh konzumuje určité zdroje, tak zároveň sniţuje jejich koncentraci. Nika má základní dva aspekty: pozici a šířku niky. Pozici niky určuje optimum: hodnota podmínek a zdrojů, za kterých má daná populace největší fitness. Šířka niky je rozsah podmínek a zdrojů, při kterém je natalita alespoň rovna mortalitě. S šířkou niky souvisí pojmy fundamentální a realizovaná nika. Fundamentální nika popisuje celkový potenciál druhu, je determinována geneticky a fyziologicky, v prostředí bez nepřátel (kompetitorů, predátorů, parazitů a patogenů). Realizovaná nika je podoblast fundamentální niky, kterou daný druh skutečně obývá v prostředí, kde dochází k interakcím s ostatními organismy. Podle šířky niky rozlišujeme stenoekní (s úzkou nikou) a euryekní (s širokou nikou) druhy. Stenoekní druhy jsou schopny tolerovat nebo vyuţívat pouze malý rozsah podmínek respektive zdrojů, naopak euryekní druhy jsou sto tolerovat nebo vyuţívat široké spektrum podmínek respektive zdrojů (HUTCHINSON 1957). Kaţdý druh ţije ve společenstvu mnoha jiných druhů, které si niku nejen vzájemně ovlivňují, ale zároveň i částečně utváří. Niky jednotlivých druhů ţijících ve společenstvu se buď vůbec nepřekrývají, nebo můţe docházet k reciprokému překryvu, kdy se fundamentální niky překrývají na různě velké šířce, ale kaţdá z nich obsahuje exkluzivní region korespondující s podmínkami prostředí, ve kterých jiné druhy nedokáţou perzistovat (např. členové jedné gildy). Zvláštním případem překryvu nik je zahrnutí niky, kdy fundamentální nika jednoho druhu je kompletně vnořena v nice druhého druhu (např. generalista x specialista) (COLWELL & FUENTES 1975). Dříve se mělo za to, ţe ekologická nika druhu je neměnná a je to jakási vlastnost druhu. Ve skutečnosti ekologická nika druhu vykazuje různou dynamiku v prostoru a čase (PEARMAN ET AL. 2008, CAVENDER-BARES ET AL. 2009). K dynamice niky dochází pravděpodobně u všech druhů, ale na různé časové a prostorové škále a za různých situací. Dynamiku a evoluci niky lze zkoumat jednak porovnáváním různých populací jednoho druhu nebo porovnáváním niky příbuzných druhů a sledovat, zda došlo k výraznější diverzifikaci od niky jejich společného předka (EMERSON & GILLESPIE 2008, 7 PEARMAN ET AL. 2008, POLECHOVÁ & STORCH 2008, CAVENDER-BARES ET AL. 2009, PETERSON 2011). Posun niky znamená jakákoliv změna (šířka, poloha, tvar) buď fundamentální nebo realizované niky nebo obojí. Ke změně niky můţe docházet na různých časových škálách (ekologické a evoluční) jako důsledek biologických interakcí (kompetice, predace, potrava, hostitel), nebo jako důsledek adaptace k místním abiotickým podmínkám (při změně klimatických podmínek nebo osidlování nových území) či jejich vzájemné interakci. K posunu můţe dojít na jakékoliv z jejich dimenzí a zároveň určitá dimenze můţe být labilnější neţ jiná. Někdy se nika můţe měnit čistě z důvodu fenotypové plasticity, jindy můţe být změna fixována i geneticky
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