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Geografski Obrasci Genetičke I Morfološke Varijabilnosti Predstavnika Grupe Silene Saxifraga (Caryophyllaceae)

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Geografski Obrasci Genetičke I Morfološke Varijabilnosti Predstavnika Grupe Silene Saxifraga (Caryophyllaceae) UNIVERZITET U BEOGRADU BIOLOŠKI FAKULTET Sanja Z. Đurović GEOGRAFSKI OBRASCI GENETIČKE I MORFOLOŠKE VARIJABILNOSTI PREDSTAVNIKA GRUPE SILENE SAXIFRAGA (CARYOPHYLLACEAE) doktorska disertacija Beograd, 2017 UNIVERSITY OF BELGRADE FACULTY OF BIOLOGY Sanja Z. Đurović GEOGRAPHICAL PATTERNS OF GENETIC AND MORPHOLOGICAL VARIABILITY OF THE TAXA BELONGING TO SILENE SAXIFRAGA GROUP (CARYOPHYLLACEAE) Doctoral Dissertation Belgrade, 2017 Mentori: dr Marjan Niketić naučni i muzejski savetnik, Prirodnjački muzej u Beogradu dr Gordana Tomović vanredni profesor, Univerzitet u Beogradu - Biološki fakultet Članovi komisije: dr Vladimir Stevanović redovni profesor u penziji, Univerzitet u Beogradu - Biološki fakultet, redovni član SANU dr Maja Lazarević docent, Univerzitet u Beogradu - Biološki fakultet dr Božo Frajman vanredni profesor, Univerzitet u Insbruku - Biološki fakultet Datum odbrane: _______________________ Izjave zahvalnosti Ova doktorska disertacija je realizovana u okviru projekta Ministarstva prosvete, nauke i tehnološkog razvoja Republike Srbije br. 173030 pod nazivom „Biodiverzitet biljnog sveta Srbije i Balkanskog poluostrva – procena, održivo korišćenje i zaštita“, na Katedri za ekologiju i geografiju biljaka Biološkog fakulteta Univerziteta u Beogradu. Molekularne analize su sprovedene u Institutu za Botaniku u Insbruku, u Austriji, a horološka istraživanja su u velikoj meri pomognuta SYNTHESYS stipendijama (AT-TAF-3503 i CZ-TAF-4439). Mnogi ljudi su me zadužili svojim savetima i pomoći tokom pripremanja ove teze i neću imenovati ni približno sve njih. Posebno se zahvaljujem mentorima dr Gordani Tomović i dr Marjanu Niketiću na ukazanoj šansi i poverenju, bez čijih ideja, znanja i iskustva ne bi bilo ove disertacije, ali i na velikoj pomoći prilikom terenskih istraživanja i na brojnim uspomenama sa zajedničkih putovanja, kao i na svim smernicama tokom izrade ove teze. Svojim učešćem u komisiji su mi veliku čast ukazali dr Vladimir Stevanović, dr Maja Lazarević i dr Božo Frajman, i zahvalna sam im na korisnim savetima, posebno tokom završnih faza izrade teze. Božo Frajman je ujedno i jedna od najzaslužnijih osoba zahvaljujući kojima su urađene molekularne analize i hvala mu na pruženoj prilici i uvek pozitivnom stavu tokom naše saradnje. Pored njega, mnogo su mi pomogli i dr Peter Schönswetter, Marianne Magauer i Daniela Pirkebner. Kustosi i botaničari koji su mi izašli u susret tokom poseta herbarskim zbirkama, a kojima sam veoma zahvalna, su: dr Snežana Vukojičić (BEOU), dr Zoltan Barina (BP), Otakar Šida (PR), Patrik Mráz (PRC), Dimitar Dimitrov (SO), Kiril Stoyanov (SOA), Chavdar Gusev i Svetlana Bancheva (SOM), dr Ernst Vitek (W), Walter Till (WU), dr Ivana Rešetnik i dr Antun Alegro (ZA, ZAHO). Snežana Vukojičić je bila mnogo više od jednog kustosa tokom posete zbirci. Njen vedri duh i veliko znanje su bili pokretačka snaga u mnogim trenucima. Neizmerno hvala na svim diskusijama i divnom društvu na terenskim istraživanjima. Zahvalna sam i brojnim profesorima i kolegama, koji su izvrsni učitelji, pomoć i inspiracija. Posebno hvala dr Dmitru Lakušiću, dr Jasmini Šinžar-Sekulić i dr Ljiljani Tomović, koji su nesebično podelili svoja znanja, koja su mi mnogo značila u izradi ove teze. Veoma sam zahvalna svojoj porodici i prijateljima, koji su ulepšali izradu ove teze svojom podrškom. Geografski obrasci genetičke i morfološke varijabilnosti predstavnika grupe Silene saxifraga (Caryophyllaceae) Sažetak Uzimajući u obzir prethodne klasifikacije i nejasne odnose između taksona u okviru ili dovođenih u vezu sa grupom Silene saxifraga, korišćene su nuklearne sekvence ITS, hloroplastne sekvence rps16 i polimorfizmi dužine umnoženih fragmenata (AFLP-ovi), za utvrđivanje filogenetičkih odnosa između ispitivanih taksona. Filogenetička analiza uzoraka 112 populacija, iz čitavog areala grupe, koji predstavljaju 33 taksona, je pokazala da su S. cephallenia, S. congesta, S. linoides, S. multicaulis, S. schwarzenbergeri i S. waldsteinii jasno izdiferencirane od ostalih analiziranih taksona, koji su formirali redefinisanu grupu S. saxifraga. Filogenetički odnosi u okviru ove grupe su slabo razjašnjeni i nisu u potpunosti u skladu sa aktuelnom taksonomijom. Na filogenetičkom stablu se uglavnom grupišu geografski bliske populacije, a ne one koje pripadaju istom taksonu, verovatno zato što je grupa nastala u skorije vreme, u pleistocenu, nakon čega je došlo do brze radijacije i ekspanzije areala. Redefinisanje taksona u okviru grupe S. saxifraga izvršeno je na osnovu multivarijantne analize morfometrijskih podataka i razlika u kvalitativnim karakterima. Analizirani su uzorci 122 populacije koje predstavljaju 22 taksona, uključujući S. linoides, S. multicaulis i S. waldsteinii. Izdvojeno je deset taksona u rangu vrste i osam podvrsta u okviru vrste S. saxifraga. Opisana je jedna nova vrsta (S. ancrae) i jedna nova podvrsta (S. saxifraga subsp. lurensis). Šest taksona je ušlo u novu kombinaciju, a od njih je pet dobilo i novi status. Ključ za identifikaciju taksona u okviru revidirane grupe S. saxifraga prate dopune originalnih opisa, kao i horološki podaci i karakteristike staništa. Centar genetičkog i taksonomskog diverziteta grupe se nalazi u južnim delovima Balkanskog poluostrva. Ključne reči: AFLP; Balkansko poluostrvo; diverzifikacija; filogeografija; horologija; ITS; morfometrija; Silene sect. Saxifragoideae; S. saxifraga grupa, taksonomija Naučna oblast: Biologija Uža naučna oblast: Ekologija, biogeografija i zaštita životne sredine UDK broj: [581.961+575.22]:582.661.51(497)(043.3) Geographical patterns of genetic and morphological variability of the taxa belonging to Silene saxifraga group (Caryophyllaceae) Abstract Considering previous classifications and unresolved relationships among taxa belonging to or hypothesized to be closely related to Silene saxifraga group, we used sequences of the nuclear ribosomal internal transcribed spacer (ITS), the plastid rps16 intron and amplified fragment length polymorphisms (AFLPs) to elucidate phylogenetic relationships, covering the section’s entire geographical range. Phylogenetic analyses of 112 populations belonging to 33 taxa showed that S. cephallenia, S. congesta, S. linoides, S. multicaulis, S. schwarzenbergeri and S. waldsteinii are clearly differentiated from all other taxa, which are included in the newly circumscribed S. saxifraga group. Unresolved phylogenetic relationships governed by geographic proximity rather than by taxonomic identity within S. saxifraga group are probably the result of recent rapid radiation and range expansion and/or local adaptation to divergent environments. Delimitation among taxa belonging to S. saxifraga group is discussed on the basis of multivariate analysis of morphometric data and differences in qualitative characters. Samples from 122 populations belonging to 22 taxa, including S. linoides, S. multicaulis and S. waldsteinii were analysed. Ten taxa are treated as species, with eight subspecies within S. saxifraga. One new species (S. ancrae) and one new subspecies (S. saxifraga subsp. lurensis) are desribed, whereas six taxa are given as new combination and five out of them as new status. Diagnostic key followed by addition to the original description of taxa belonging to revised S. saxifraga group is given, together with chorological data and habitat features. The southern part of the Balkan Peninsula is the centre of genetic and taxonomic diversity of the group. Key words: AFLP; Balkan Peninsula; chorology; diversification; ITS; phylogeography; morphometrics; Silene sect. Saxifragoideae; S. saxifraga group, taxonomy Scientific field: Biology Scientific subfield: Ecology, biogeography and environmental protection UDC number: [581.961+575.22]:582.661.51(497)(043.3) SADRŽAJ 1. UVOD ........................................................................................................................... 1 1.1. Istorijat klasifikacije tribusa Sileneae i roda Silene na osnovu morfologije i filogenetička istraživanja. ............................................................................................... 1 1.2. Filogeografska istraživanja južnoevropskih taksona. ................................................. 10 1.3. Karakteristike istraživanih taksona, koji su tokom istorije dovedeni u vezu sa grupom S. saxifraga. ..................................................................................................... 11 1.4. Pregled klasifikacije vrsta koje su tokom istorije dovedene u vezu sa grupom S. saxifraga. ....................................................................................................................... 45 2. CILJEVI RADA ......................................................................................................... 53 3. MATERIJAL I METODE .......................................................................................... 54 3.1. Molekularne analize ...................................................................................................... 54 3.1.1. Biljni materijal............................................................................................... 54 3.1.2. Rad u laboratoriji (ekstrakcija DNK, ITS, rps16, AFLP). ............................ 55 3.1.3. Analiza sekvenci. .......................................................................................... 57 3.1.4. Analiza polimorfizma dužine umnoženih fragmenata. ................................. 57 3.2. Morfometrijske analize ................................................................................................
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