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Diverzitet Aerofitskih Cijanobakterija I Algi U Biofilmu Odabranih Pećina U Srbiji

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Diverzitet Aerofitskih Cijanobakterija I Algi U Biofilmu Odabranih Pećina U Srbiji UNIVERZITET U BEOGRADU BIOLOŠKI FAKULTET Slađana S. Popović DIVERZITET AEROFITSKIH CIJANOBAKTERIJA I ALGI U BIOFILMU ODABRANIH PEĆINA U SRBIJI doktorska disertacija Beograd, 2018 UNIVERSITY OF BELGRADE FACULTY OF BIOLOGY Slađana S. Popović DIVERSITY OF AEROPHYTIC CYANOBACTERIA AND ALGAE IN BIOFILM FROM SELECTED CAVES IN SERBIA Doctoral Dissertation Belgrade, 2018 Mentor: dr Gordana Subakov Simić, vanredni profesor, Univerzitet u Beogradu, Biološki fakultet Članovi komisije: dr Jelena Krizmanić, vanredni profesor, Univerzitet u Beogradu, Biološki fakultet dr Olivera Krunić, redovni profesor, Univerzitet u Beogradu, Rudarsko-geološki fakultet Datum odbrane:___________________________ Doktorska disertacija realizovana je na Katedri za algologiju, mikologiju i lihenologiju Instituta za botaniku i botaničke bašte „Jevremovac“, Biološkog fakulteta Univerziteta u Beogradu. Koristim priliku da se zahvalim svim kolegama i koleginicama sa katedre na pomoći i lepim trenucima. Najveću zahvalnost dugujem svojoj izuzetnoj mentorki prof. dr Gordani Subakov Simić na velikoj podršci tokom izrade doktorske disertacije, ohrabrenju, usmeravanju, podsticanju istraţivačkog duha, korisnim savetima, ukazanom poverenju i volji i pomoći u svakom trenutku. TakoĎe se zahvaljujem dr Jeleni Krizmanić na maksimalnoj posvećenosti tokom identifikacije silikatnih algi i sugestijama koje su pozitivno uticale na kvalitet disertacije. Zahvalnost takoĎe dugujem Danijeli Vidaković i Olgi Jakovljević. Prof dr Oliveri Krunić se zahvaljujem na korisnim savetima, velikoj pomoći i volji da uvek izaĎe u susret, kao i njenom saradniku Vladimiru Šarabi. Prof dr Milici Ljaljević Grbić i njenim saradnicima Milošu Stuparu, Nikoli Unkoviću i Ţeljku Savkoviću se zahvaljujem na kreativnim idejama i aktivnom učešću u odreĎenim delovima studije, čija je stručnost znatno doprinela kvalitetu publikacija. Prof dr Aleksandri Korać se zahvaljujem na neizrecivoj upornosti prilikom rada na analizi nove vrste za nauku, cijanobakterije Nephrococcus serbicus, kao i njenim saradnicima Igoru Goliću, Maji Bogdanović i Aniti Lazarević. Prof dr Kristini Šarić dugujem zahvalnost za pokazano intresovanje i korisne informacije vezane za uzorke stena, kao i strpljivom radu na uzorcima. TakoĎe se zahvaljujem dr Milki Vidović, direktorki Centra za ekologiju i tehnoekonomiku NU IHTM na podršci tokom izrade i pisanja doktorske disertacije kao i svim kolegama iz IHTM. Posebnu zahvalnost dugujem Ani Pantelić i Ţeljki Milovanović. Veliku zahvalnost dugujem Prof. RNDr. Jiří Komáreku iz Češke Republike na bezrezervnoj pomoći prilikom opisa nove vrste cijanobakterija, ali i na pomoći prilikom identifikacije problematičnih taksona. Zahvalnost dugujem takoĎe i DCC timu (Jeffrey Johansen, Jan Kaštovský i Tomáš Hauer) koji su se maksimalno potrudili da nam problematiku identifikacije cijanobakterija iznesu na najbolji način. TakoĎe se zahvaljujem profesorima Jan Lepšu i Petr Šmilaueru čiji je Canoco statistički program otvorio mnoga nova vrata u istraţivanjima. Srdačno se zahvaljujem svim kolegama i prijateljima koji su izašli u susret prilikom terenskih istraţivanja pećina na teritoriji Srbije, posebno JP “Srbijašume” Beograd i ŠG “Timočke šume”, kao i upravljačima svih pećina, ali i Vojislavu Simiću, Darku Draguloviću, Bojanu Gavriloviću na pomoći oko uzorkovanja, kao i baki iz Visoke bez koje neverovatna Boţanina pećina ne bi bila otkrivena. Dragim Savetama i Molerima, Jeleni Jovanović, Dragani Predojević, Ivani Trbojević, Ani Blagojević, Ani Andjelković (koja je imala volje da pročita celu doktorsku disertaciju), Miljani Jakovljević, Dragani Antonijević, svim ostalim dragim prijateljima, Sneţi, Jeleni, Isidori, Antoniji, Ivani, Nataši Vama Zago i tebi teta Rado, dugujem ogromnu zahvalnost na podršci, pomoći, ohrabrenju i što ste uvek bili uz mene. Disertaciju posvećujem svojim roditeljima, mami Gordani i tati Stevu, kao i sestri Dragani koji su me uvek podrţavali i bez čije pomoći i podstreha koji me je naveo da obiĎem obliţnje pećine, doktorat ne bi krenuo u ovom smeru. Hvala im na beskrajnoj podršci, razumevanju, ljubavi i veri koju su imali u moj uspeh. The cave you fear to enter holds the treasure you seek Joseph Campbell Diverzitet aerofitskih cijanobakterija i algi u biofilmu odabranih pećina u Srbiji Sažetak Predmet ove doktorste disertacije je analiza zajednice aerofitskih cijanobakterija i algi iz biofilma sa stenovitog supstrata ulaza petnaest odabranih pećina iz Srbije kao i nekoliko tačaka u blizini veštačkog osvetljenja. Akcenat istraživanja bio je na fototrofnim mikroorganizmima iz biofilma, dok su kao dodatak endolitski predstavnici identifikovani iz nekoliko uzoraka stene. Analizirana je i morfologija, stepen razvijenosti biofilma, kao i udeo sadržaja vode, organske i neorganske materije u biofilmu. U svim pećinama, mereni su ekološki parametri i određene su primarna produkcija (preko koncentracije hlorofila a) i petrografska analiza stenovitog supstrata. Na odabranim uzorcima rađena je kvantitativna i sezonska analiza cijanobakterija i algi. Statističke analize su poslužile kao alat za sagledavanje odnosa zabeleženih fototrofnih mikroorganizama i različitih ekoloških i opisnih parametara. Zabeleženo je prisustvo ukupno 290 taksona cijanobakterija i algi koje su svrstane u 4 razdela: Cyanobacteria (134 taksona, od kojih 82 taksona pripada kokoidnim formama), Bacillariophyta (129 taksona), Chlorophyta (26 taksona) i Xanthophyta (1 takson). Najveći broj taksona Cyanobacteria zabeležen je u rodovima Gloeocapsa, Chroococcus, Gloeothece, Leptolyngbya i Scytonema. U razdelu Bacillariophyta se izdvajaju rodovi Luticola i Humidophila, a u razdelu Chlorophyta rod Cosmarium. Iz biofilma Božanine pećine opisana je nova vrsta cijanobakterije za nauku, Nephrococcus serbicus S. Popović, G. Subakov Simić i J. Komárek, sp. nov. pri čemu je sam rod po prvi put zabeležen na teritoriji Evrope. Cyanobacteria po broju taksona dominiraju na svim lokalitetima, osim Rćanske i Bogovinske pećine. Najveći broj pripadnika Bacillariophyta i Chlorophyta zabeležen je u Rćanskoj (usled prisustva vode na biofilmu) i pećini Samar (zbog sezonskog uzorkovanja). Upravo se značaj sezonskog uzorkovanja biofilma u pećinama Samar i Jezava pre svega ogleda kroz znatno veći diverzitet aerofitskih cijanobakterija i algi u odnosu na druge pećine. Predstavnici Cyanobacteria dominirali su u pećinama čiji je supstrat rekristalisali i organogeni krečnjak, a Bacillariophyta na klastičnom i mikritskom krečnjaku. U dobro razvjenim, želatinoznim biofilmovima dominirale su cijanobakterije. U tankom sloju biofilma na stenovitom supstratu dominirale su zelene alge ukoliko su biofilmovi bili suvi, odnosno silikatne alge ukoliko su bili vlažni. Vrednosti ekoloških parametara na svakoj tački uzorkovanja dovedeni su u vezu sa lokacijom pećine, morfologijom i veličinom pećinskog ulaza, godišnjim i dnevnim variranjima meteoroloških činilaca. Vrednosti temperature i relativne vlažnosti vazduha bile su prilično ujednačene na nivou pojedinačne pećine, dok su vrednosti intenziteta svetla pokazale varijacije i zavisnost u odnosu na blizinu tačke uzorkovanja ulazu, kao i njene ekspoziciju. Koncentracija hlorofila a, kao i sadržaj vode, organske i neorganske materije bili su korelisani sa dobro razvijenim biofilmom na ulazima pećina i biofilmom na horizontalnoj površini. Sezonska analiza biofilma u odabranim pećinama pokazala je da odnos vrednosti parametara u biofilmu zavisi od sezone, što se najviše odrazilo na sadržaj vode. Vrednosti hlorofila a bile su više u hladnijim mesecima. Najveća vrednost biomase je u Degurićkoj pećini izračunata na mestu gde je visok sadržaj vode, kao i viša koncentracija hlorofila a, a u pećini Vernjikica na mestu gde je zabeležen i visok sadržaj organske materije. Ključne reči: pećine, biofilm, fototrofni mikroorganizmi, aerofitske cijanobakterije, aerofitske alge, Nephrococcus serbicus , biomasa, stenoviti supstrat, krečnjak Naučna oblast: Biologija Uža naučna oblast: Algologija UDK broj: 581.91/.92:581.96:[582.232+582.26][551.435.84+552.54](497.11)(043.3) Diversity of aerophytic cyanobacteria and algae in biofilm from selected caves in Serbia Abstract The aim of this PhD thesis is the analysis of the aerophytic cyanobacterial and algal community from biofilm developed on stone substrate at the entrance of fifteen selected caves from Serbia and several sites near the artificial light in caves. The focus of the study was on phototrophic microorganisms from the biofilm, while endolithic representatives were additionaly identified from several stone samples. Morphology, degree of biofilm development, as well as the content of water, organic and inorganic matter in biofilm were also observed. At all sampling sites, ecological parameters, primary production (through concentration of chlorophyll a), as well as petrographic analysis of rocky substrates were measured and determined. Quantitative and seasonal analyzes of cyanobacteria and algae were assessed in selected samples. Statistical analyzes have served as a tool for examining the relationship between recorded phototrophic microorganisms and various ecological and descriptive parameters. A total of 290 cyanobacterial and algal taxa were recorded and classified into 4 divisions: Cyanobacteria (134 taxa, of which 82 taxa belong to the coccoid forms), Bacillariophyta (129 taxa), Chlorophyta (26 taxa) and Xanthophyta (1 taxon). The highest number of cyanobacterial taxa was documented within the following genera: Gloeocapsa, Chroococcus, Gloeothece, Leptolyngbya and Scytonema.
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