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Filogenetički Odnosi I Genska Raznolikost Roda Squalius (Cypriniformes, Actinopteri) U Jadranskom Slijevu

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Filogenetički Odnosi I Genska Raznolikost Roda Squalius (Cypriniformes, Actinopteri) U Jadranskom Slijevu PRIRODOSLOVNO-MATEMATIČKI FAKULTET BIOLOŠKI ODSJEK Kristina Čavlović FILOGENETIČKI ODNOSI I GENSKA RAZNOLIKOST RODA SQUALIUS (CYPRINIFORMES, ACTINOPTERI) U JADRANSKOM SLIJEVU DOKTORSKI RAD Mentor: prof. dr. sc. Davor Zanella Zagreb, 2019. Faculty of Science Department of Biology Kristina Čavlović PHYLOGENETIC RELATIONSHIPS AND GENETIC DIVERSITY OF THE GENUS SQUALIUS (CYPRINIFORMES, ACTINOPTERI) IN ADRIATIC SEA BASIN DOCTORAL DISSERTATION Supervisor: prof. dr. sc. Davor Zanella Zagreb, 2019 „Ovaj je doktorski rad izraĊen na Zoologijskom zavodu, pod vodstvom prof. dr. sc. Davora Zanelle, u sklopu Sveuĉilišnog poslijediplomskog doktorskog studija Biologije pri Biološkom odsjeku Prirodoslovno-matematiĉkog fakulteta Sveuĉilišta u Zagrebu“. INFORMACIJE O MENTORU ZANELLA, Davor (Zagreb, 19. travnja 1973.) Redoviti profesor, 235863 U Zagrebu je završio osnovnu i srednju školu. Maturirao je 1992. g. na V. gimnaziji, a iste je godine upisao studij biologije, smjer profesor biologije na PMF-u u Zagrebu. Diplomirao je 1997. g. Od 2000. g. zaposlen je u Zoologijskom zavodu kao znanstveni novak. Godine 2003. obranio je magistarski rad, a 2007. i doktorsku disertaciju. Na mjesto docenta izabran je 2009., 2013. g. u izvanrednog profesora, a 2019. u redovitog profesora. Usavršavao se iz ekologije slatkih voda (odreĊivanje kvalitete vode pomoću biotiĉkog indeksa) i populacija riba rijeĉnih tokova i jezera u Laboratoriju za ihtiologiju i plankton Sveuĉilišta u Trstu, boravio je u Tromsu, Norveška na struĉnom usavršavanju iz podruĉja ihtiologije u sklopu projekta "Management of Freshwater Fisheries of Bordering Rivers" u NIVA (Norwegian Institute for Water Research). Do 2009. g. vodi praktikume na preddiplomskom studiju iz kolegija Kralježnjaci, kao i terensku nastavu iz zoologije. Od 2009. g. nositelj je ili sunositelj kolegija Vertebrata, Upravljanje prirodnim populacijama, Ekologija i zaštita prirode, Zoologija 3 (Kralježnjaci), Ihtiologija, Akvakultura i ribarstvo slatkih voda i Akvakultura za studente preddiplomskog i diplomskog studija razliĉitih smjerova. Sudjeluje u terenskoj nastavi iz navedenih kolegija. Voditelj je i 12 diplomskih radova te dva magistarska i 15 završnih radova. Njegova znanstvena djelatnost obuhvaća istraživanja raznolikosti i staništa slatkovodnih riba Hrvatske, posebno endemskih vrsta (inventarizaciju i utvrĊivanje stupnja ugroženosti te bioloških i ekoloških znaĉajki pojedinih vrsta). Drugo podruĉje njegova znanstvenog rada usmjereno je na prouĉavanje prirodnih populacija te njihovom upravljanju. Objavio je kao autor ili koautor 51 znanstveni rad (41 u ĉasopisima indeksiranim u CC, 10 u ĉasopisima indeksiranim u drugim sekundarnim i tercijarnim bazama), Koautor tri knjige te jednog poglavlja u knjizi. Sudjelovao na meĊunarodnim skupovima s 46 priopćenja te domaćim skupovima s 12 priopćenja. Sudjelovao je na brojnim meĊunarodnim i domaćim skupovima sa 60-tak priopćenja. Aktivno je sudjelovao u izvedbi triju znanstvena projekta. Bio je i suradnik na dva meĊunarodna projekta Jedan je od osnivaĉa, tajnik i blagajnik te ĉlan Upravnog odbora Hrvatskog ihtiološkog društva od 1999. godine. Od 2005. do 2012. g. je tajnik, blagajnik i ĉlan Upravnog odbora Hrvatskog ekološkog društva. ZAHVALE Stvaranje ove doktorske disertacije ne bi bilo moguće bez svih izvanrednih ljudi koji su bili uz mene tijekom dobrih i loših trenutaka. Najveća hvala prof. dr. sc. Davoru Zanelli koji mi je omogućio izradu rada pod svojim mentorstvom, pružajući mi veliko znanstveno iskustvo. Hvala na svim korisnim savjetima i mirnoći koja mi je pomogla da ostanem fokusirana na sam znanstveni rad, te bezrezervnom željom za pomoć u svakom trenutku. Zahvaljujem se na velikoj strpljivosti u svim trenucima neuspjeha i nepredvidivih dogaĊaja tijekom cijelog puta. Ogromno hvala doc. dr. sc. Ivani Buj na odvojenom vremenu koje mi je posvetila, te na svom prenešenom znanju i pomoći oko laboratorijskih metoda i raĉunalnih programa, bez ĉega bih u više sluĉajeva bila potpuno izgubljena. Isto tako, želim se zahvaliti i na periodu osamostaljenja, bez ĉega ne bih nauĉila samostalno znanstveno razmišljati. Isto tako, htjela bih se zahvaliti svim djelatnicima Laboratorija za vertebrata Zoologijskog zavoda, koji su mi omogućili vrlo ugodan boravak te znanstvenu podršku. Osobito bih se zahvalila dr. sc. Zoranu Marĉiću za slike i svu pomoć oko terenskih pitanja. Veliko hvala svim kolegama sa Zavoda za humanu reprodukciju KBC-a Rijeka, bez ĉijeg razumijevanja ne bi bili mogući svi moji izostanci zbog potreba istraživanja i pisanja ove disertacije. Veliko hvala prof. dr. sc. Nedi Smiljan Severinski na zapoĉetom putu, a posebno hvala kolegicama Lindi, Tamari i Vanji na svim pretrpljenim „fazama“, razumijevanju, dizanju morala i savjetima koji su doprinijeli znanstvenom sazrijevanju. Posebna zahvala ide dragim prijateljima i kolegama, Miljenku i Petri, bez kojih se ne bih upustila u ovo putovanje. Vaše vjerovanje i stalno podupiranje davalo mi je vjetar u leĊa. Jelka, hvala ti na smještaju i psihološkoj podršci, te neiscrpnom izvoru smijeha. Hvala Tini, Ivani, Ani, Jeleni i Korani i svim ostalim prijateljima koji su bili zakinuti za druženja kroz svo ovo vrijeme, ali uvijek s razumijevanjem. Posebno veliko hvala mojoj obitelji, mami i tati, jer bez vašeg truda i odricanja ostvarenje ovog sna ne bi bilo moguće. Hvala na bezuvjetnoj ljubavi i podršci koju nikad nisam izgubila, a niti morala tražiti. Na kraju, Ivane, hvala ti što si nam krojio život dok sam se ja borila s raĉunalnim simulacijama. Hvala ti na konstantnoj potpori, ljubavi i motivaciji, a posebice na zdravom i ukusnom jelovniku. Još jedno hvala - Mucanu i Mucki. Uz njih je svako ostajanje doma uz znanstveni rad bilo puno lakše. Sveuĉilište u Zagrebu Doktorski rad Prirodoslovno-matematiĉki fakultet Biološki odsjek FILOGENETIČKI ODNOSI I GENSKA RAZNOLIKOST RODA SQUALIUS (CYPRINIFORMES, ACTINOPTERI) U JADRANSKOM SLIJEVU KRISTINA ĈAVLOVIĆ Zoologijski zavod Rod Squalius (Leuciscidae, Cypriniformes) živi u slatkovodnim vodotocima Europe i Azije, s velikom raznolikosti u mediteranskoj regiji. Cilj istraživanja bio je razriješiti taksonomske nedoumice klenova rijeka jadranskog slijeva u Hrvatskoj i BiH, te odrediti rasprostranjenosti pojedinih vrsta, povezanost populacija i razinu genske raznolikosti, te ugroženost pojedinih populacija u svrhu djelotvornije zaštite. Filogenetiĉka rekonstrukcija napravljena na mitohondrijskim i jezgrinim molekularnim biljezima (cyt b, rhod, rag1 i S7) pokazala je dvije nezavisne, genetiĉki udaljene evolucijske grupe: mediteransku (S. svallize, S. microlepis, S. tenellus, S. illyricus i S. zrmanjae) i S. cephalus grupu (S. squalus), zahvaljujući davnom vremenu divergencije. Na raznolikost navedenih vrsta najviše je utjecala alopatrijska specijacija, ĉemu je doprinijela i prirodna hibridizacija. Ovim istraživanjem nije potvrĊeno postojanje istarske vrste S. janae, te se predlaže da se taj naziv koristi kao mlaĊi sinonim vrste S. squalus. Manja zabilježena unutarvrsna genska raznolikost u dalmatinskoj regiji objašnjava se povećanom osjetljivosti klenova na antropogene utjecaje. (197 stranica, 48 slika, 47 tablica, 301 literaturni navod, jezik izvornika: hrvatski jezik) Kljuĉne rijeĉi: filogenetička rekonstrukcija, raznolikost vrsta, taksonomija, mitohondrijski i jezgrini molekularni biljezi, genska udaljenost, dijagnostička mjesta, migracije. Mentor: Davor Zanella, prof. dr. sc. Ocjenjivaĉi: Ivana Buj, doc. dr. sc., Marko Ćaleta, izv. prof. dr. sc.; Ana Galov, izv. prof. dr. sc. University of Zagreb Doctoral Thesis Faculty of Science Department of Biology PHYLOGENETIC RELATIONSHIPS AND GENETIC DIVERSITY OF THE GENUS SQUALIUS (CYPRINIFORMES, ACTINOPTERI) IN ADRIATIC SEA BASIN KRISTINA ĈAVLOVIĆ Division of Zoology Background: Genus Squalius (Leuciscidae, Cypriniformes) inhabits european and asian freshwater bodies, with particularly high diversity in Mediterranean area. The taxonomic status of many populations is still matter of debate. This paper constitute first attempt at deciphering mitochondrial and nuclear relationships of Adriatic Squalius species. Aim: The aim of this investigation was to resolve taxonomic uncertainties of the Adriatic chubs from Croatia and Bosnia and Herzegovina and to determine distribution of species, population's connection, status and genetic diversity for future conservation purposes. Also, to test biogeographical hypotheses that could have determined the current distribution of Adraitic Squalius species. Methods: In order to reveal phylogeographic and population genetic structure of the genus Squalius in the Adriatic watershed, samples from 19 localities in Croatia and Bosnia and Herezegovina were obtained. The isolation of DNA, gel electrophoresis, PCR and sequencing of DNA from frozen fin tissue. Selective neutrality of mutations and recombinations was tested by statistical tests D* and F*, Tajima test and recombination test. Haplotype variants of nuclear genes in heterozygous individuals were resolved using a Bayesian statistical method with 1000 iterations. Phylogenetic resconstruction was based on mitochondrial and nuclear genetic markers (cytochrome b, rhodopsin, recombination activating gene 1 and first intron of ribosomal protein S7). The complete mitochondrial cytochrome b (total of 1140 bp) from 166 sequences belonging to the Squalius species was obtain from Adriatic rivers, Europe and Asia. For the nuclear phylogenetic analysis, a subsets of 158 rhod sequences (782 pb), 136 rag1 sequences (1443 pb) and 47 S7 sequences (726 pb) was selected. Phylogenetic
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