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Korelacioni Odnosi Dnevnih Leptira Srbije I Vaskularnih Biljaka… Jakšić, P. Korelacioni odnosi dnevnih leptira Srbije i vaskularnih biljaka… th Niš (Serbia ), September 01-03, 2007 9 Symposium on Flora of Southeastern Serbia and Proceeding, 225-230 Neighbouring Regions ! Korelacioni odnosi dnevnih leptira Srbije i vaskularnih biljaka kao njihovih ovipozijskih i nutritivnih biljaka Predrag Jakšić Odsek za biologiju sa ekologijom PMF-a Univerziteta u Nišu Abstract: Jakšić, P.: Correlation relationshipbetween butterflies of Serbia and vascular plants as their oviposition and nutritive plants. Proceeding of the 9th Symposium of flora of Southeastern Serbia and Neighbouring Regions, Nis, 2007. The adaptive plasticity in relation to environmental conditions, which is manifested by character of correlation relationship between vascular plants as ovipositional and nutritive plants, and butterfly caterpillars, has been investigated. In this paper were specially analyzed polyphagian butterflies- if a nutritive plant of their caterpillars belongs to the same or to the different life forms, and what are ecological implications of this relations. The pattern was established by this analysis. Key words: living form, vascular plants, butterflies, Serbia ! ! Uvod sledeće životne forme biljaka: fanerofite (p), ! nanofanerofite (np), fanerofitske lijane (pl), Tokom evolucije organske vrste pokazuju drvenaste hamefite (dc), zeljaste hamefite (zc), adaptivnu plastičnost na uslove spoljne sredine, hemikriptofite (h), geofite (g), terofite (t), manifestovanu posebnim načinom života. terofite/hamefite (th), akvatične helo-hidrofite (a). Adaptivni karakter je rezultat reakcije organizma na Najveći broj gusenica dnevnih leptira Srbije hrani uticaje spoljne sredine. Pošto se spoljna sredina se biljkama koje su hemikriptofite (h). Prema neprekidno menja to je i reakcija organizma T i s c h l e r o v o j (1949) klasifikaciji životnih kontinuirana, to je ekološka osnova evolucionog formi životinja gusenice najvećeg broja vrsta naših procesa. Adaptivni karakter jedne vrste se najbolje dnevnih leptira su herbikoli, podgrupa epifitobionti, uočava analizom njenih morfofizioloških i što znači da žive na biljkama. Gusenice nekih vrsta ekofizioloških odlika. S t a n k o v i ć (1961) je dnevnih leptira (Maculinea sp.) su u prvoj polovini životnu formu definisao na sledeći način: «Skup života herbikoli, podgrupa endofitobionti, što znači adaptivnih odlika jedne organske vrste, saglašenih sa da žive u biljkama (u cvasti) a u drugoj polovini konkretnom sredinom u kojoj se ona susreće, čini larvenog života su mirmekofili, što znači da žive u životnu formu te vrste.» Životnom formom se mravinjacima hraneći se larvama mrava. Za adultne odražavaju neposredni odnosi vrste prema određenom razvojne stadijume dnevnih leptira bi se moglo reći kompleksu životnih uslova pod kojima živi. da su aerobionti koji se hrane cvetnim nektarom. Klasifikaciju životnih formi vaskularnih Veza između vaskularnih biljaka i dnevnih biljaka dao je R a u n k i a e r (1934), a detaljnu leptira je višestruka. Poznato je da ženke leptira klasifikaciju po ovom principu kod nas su dali imaju strogo definisan odnos prema biljkama kao Kojić i saradnici (1997). Klasifikaciju životnih hrani njihovih gusenica. U tom kontekstu formi životinja dao je T i s c h l e r (1949) a kod nas razlikujemo ovipozicijsku biljku – na kojoj ženka su sličnu klasifikaciju dali Matvejev i Dimovski polaže jaja, i nutritivnu biljku – kojom se hrane (1964). Prema tim klasifikacijama razlikujemo gusenice. Ovipozicijska biljka je po pravilu 225 9th Symposium on Flora of Southeastern Serbia and Neighbouring Regions Proceeding istovremeno i nutritivna biljka, dok neke nutritivne Materijali i metode biljke ne moraju biti i ovipozicijske. Ovipozicija je genetski fiksirana radnja. Zbog ovoga se areal jedne U radu smo primenili metodu nepotpune vrste uvek prostire unutar areala ovipozicijske indukcije (M i l a n k o v i J a k š i č , 2006). biljke. Pored ovoga, leptiri su često i oprašivači Objedinili smo i sistematizovali sve do sada pojedinih vaskularnih biljaka, hrane se nektarom i poznate podatke o ovipozicijskim i nutritivnim mogu pokazivati specifičnost u izboru biljne vrste vrstama vaskularnih biljaka Srbije, i šire. Iz takve kao mesta za odmor (resting plants). Gusenice se baze podataka nasumice smo odabrali 36 vrsta mogu hraniti samo ovipozicijskom biljkom, tada dnevnih leptira čije su gusenice monofagne i 13 govorimo o monofagnim vrstama, ili mogu vrsta dnevnih leptira čije su gusenice polifagne. uzimati i neke druge vrste, često i iz različitih Potom smo analizirali životne forme vaskularnih rodova, kada govorimo o polifagnim vrstama. biljaka prema Raunkiaeru odabranih vrsta dnevnih Filogenetska veza između vaskularnih biljaka i leptira. Smisao analize je bio da pronadjemo tzv. dnevnih leptira, iskazana ovipozicijom i nutritivnim «zavisne» i «nezavisne» varijable i da na bazi toga odnosima, ima svoje biohemijsko-fiziološko formulišemo ekološke i biogeografske reperkusije. utemeljenje. Ono se ogleda najpre u setu specifičnih enzima digestivnog trakta gusenica, Rezultati i diskusija prilagođenih nutritivnim biljkama. Ali, mnogo suptilnija veza se uočava analizom mehanizma Rezultate komparativne analize prikazani su odbrane gusenice, zasnovanom na specifičnim na Tab. 1 - Tab. 4. jedinjenjima dobijenim iz nutritivne biljke. Dobro Najzad, ako analiziramo polifagne vrste je proučen primer cijanoglukozida, preuzetih iz dnevnih leptira onda uočavamo određene Sedum vrsta, kojima se hrane gusenice vrste pravilnosti, to je ilustrovano na primeru vrsta Parnassius apollo. Nishida i Rothschild Ochlodes venata i Plebeius idas (Tab. 3). (1995) su pokazali da se visoka koncentracija ovih Istovetnu analizu smo uradili za 13 nasumice jedinjenja nalazi ne samo u telu leptira, već i u odabranih vrsta dnevnih leptira, čije gusenice u njegovim krilima (aposematička obojenost), pa čak ishrani koriste oko 70 vrsta vaskularnih biljaka. i u jajima koja ženka polaže. W a h l b e r g (2001) Rezultate te analize u sumarnoj formi prikazujemo je utvrdio da gusenice 65 vrsta Melitaeina na tabeli 4. (Lepidoptera: Nymphalidae) zasnivaju svoju odbranu na iridoidima, jedinjenjima koja se javljaju Zaključci kao sekundarni metaboliti kod biljaka iz 16 različitih familija. Autor je zaključio da je Analizom table 1 uočavamo specifičnu prisustvo iridoida glavni faktor u izboru nutritivne adaptaciju pojedinih vrsta vaskularnih biljaka biljke, bez obzira na njenu taksonomsku pripadnost. prema ekološkim faktorima. Tako je, na pr., Rumex B r a b y i T r u e m a n (2006) su dokazali acetosa submezofita (prema faktoru vlage u postojanje veze između asocijacije vaskularnih zemljištu), neutrofilna (kiselost zemljišta), biljaka i gusenica kao osnove evolucionog mezotrofna (sadržaj azota u zemljištu), mehanizma adaptivne radijacije kod dnevnih leptira polusciofita-heliofita (svetlost) i mezotermna iz familije Pieridae. I u ovom slučaju autori su (toplota) vrsta biljke. Na osnovu ovakve slike nije koren veze našli u biohemijsko-fiziološkoj osnovi - moguće uočiti specifični odgovor vrste dnevnog glukozinolatu kao aktivnoj supstanci koju moraju leptira prema ovipozicijskoj i nutritivnoj biljci imati prehrambene biljke. gusenica. Ali, ako taj odnos posmatramo kroz Ako je međuodnos vaskularnih biljaka i životne forme vaskularnih biljaka (tabla 2) onda dnevnih leptira filogenetski utemeljen na bazi već uočavamo izvesne pravilnosti. biohemijsko-fizioloških mehanizama, i ako je U ishrani gusenica dnevnih leptira zastupljene iskazan njihovom adaptivnom radijacijom, onda su vaskularne biljke koje pripadaju svim poznatim verujemo da tu relaciju možemo ustanoviti i na životnim formama. Od 36 analiziranih vrsta ekološkom nivou. Izvesno je da su za takvu analizu dnevnih leptira sa table 1, čije su gusenice pogodnije monofagne vrste dnevnih leptira. monofagne, 25 vrsta dnevnih leptira, što čini 72%, Međutim, u ovom radu smo posebno analizirali je vezano za vaskularne biljke koje su polifagne vrste dnevnih leptira – da li te nutritivne hemikriptofite (h). Sa druge strane, nijedna od vrste biljaka pripadaju istoj životnoj formi ili analiziranih vrsta nije vezana za životnu formu različitim životnim formama, i kakve su fanerofitnih lijana (pl) ili terofita/hamefita (th). I biogeografske implikacije takvih odnosa. kod 13 analiziranih polifagnih vrsta vidimo da je 226 Jakšić, P. Korelacioni odnosi dnevnih leptira Srbije i vaskularnih biljaka… najveći broj njih vezan za biljke iz grupe 3). Bilo bi značajno da se ovakva analiza uradi za hemikriptofita (h). Najzad, ako analiziramo sve vrste dnevnih leptira Srbije. polifagne vrste dnevnih leptira, one čije se gusenice Polifagne vrste dnevnih leptira možemo podeliti u hrane ne sa jednom već sa većim brojem dve grupe: specifično zavisne i nespecifično vaskularnih biljaka, onda vidimo punu pravilnost. zavisne vrste. Specifično zavisne vrste dnevnih Dve nasumice odabrane vrste dnevnih leptira – leptira imaju gusenice koje se hrane većim brojem Pyrgus malvae (L.) i Plebeius idas (L.) pokazuju vrsta vaskularnih biljaka, često i iz različitih diferenciranost prema vaskularnim biljkama rodova, ali koje uvek pripadaju istoj životnoj formi. određene životne forme. Vrsta P. malvae (L.) Toj grupi pripada prvih pet vrsta navedenih u tabeli preferira hemikriptofite (h) pa je i njeno prisustvo 4. Nespecifično zavisne vrste dnevnih leptira imaju na datom staništu indikator te životne forme. gusenice koje se hrane većim brojem vrsta Suprotno tome, vrsta P. idas (L.) nije usko vaskularnih biljaka, često i iz različitih rodova
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