Prostorna Distribucija I Ekološke Varijacije Staništa HipogeiNih Makromiceta ( Mycota ) U Srbiji

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Prostorna Distribucija I Ekološke Varijacije Staništa Hipogei�Nih Makromiceta ( Mycota ) U Srbiji UniverzitetuBeogradu Biološkifakultet BorisN.Ivanevi PROSTORNADISTRIBUCIJAIEKOLOŠKE VARIJACIJESTANIŠTAHIPOGEINIH MAKROMICETA(MYCOTA)USRBIJI doktorskadisertacija Beograd 2016 UniversityofBelgrade FacultyofBiology BorisN.Ivanevi SPATIALDISTRIBUTIONANDHABITAT VARIABILITYOFHYPOGAEOUS MACROMYCETES(MYCOTA)INSERBIA DoctoralDissertation Belgrade 2016 MENTORI dr Dmitar Lakuši dr Mitko Karadelev redovni profesor redovni profesor Univerzitet u Beogradu, Univerzitet “Sv. Kiril i Biološki fakultet Metodije”u Skoplju, Prirodno-matematiki fakultet 7LANOVI KOMISIJE dr Dmitar Lakuši redovni profesor Univerzitet u Beogradu, Biološki fakultet dr Mitko Karadelev redovni profesor Univerzitet “Sv. Kiril i Metodije”u Skoplju Prirodno-matematiki fakultet dr Jasmina Šinžar-Sekuli vanredni profesor Univerzitet u Beogradu, Biološki fakultet DATUM ODBRANE: Autor ilustracije Golden Celtic Tree Of Life sa naslovne strane je Serge Averbukh. (Slika je neznatno modifikovana) Drvo je arhetipski simbol u ljudskoj kulturi koji se manifestuje u mnogo oblika, od mitskog svetog drveta, zapisa i Igdrasila, preko keltskog drveta života do metafore evolucije arlsa Darvina. Da bi drvo opstalo i napredovalo, neophodne su gljive sa kojima je u simbiozi. Želim da uputim svoju zahvalnost kolegama i prijateljima koji su pomogli da ova disertacija dobije svoj definitivni oblik. Pre svega, dr Miroljubu Milenkovi6u na poverenju i želji da podeli svoje jedinstveno i dragoceno znanje o podzemnim gljivama Srbije, i poveri vrednu zbirku koja je rezultat rada i istraživanja tokom dvadeset pet godina. Takoe i za nesebi8nu pomo pri sreivanju i tuma8enju podataka, gde je njegovo iskustvo bilo nezamenljivo. Mentorima Prof. Dmitru Lakušiu i Prof. Mitku Karadelevu dugujem posebnu zahvalnost za podstrek i konstruktivan pristup, kao i za kljune sugestije. Dr David Minter i Prof. Maria awrynowicz su mi dali odline globalne uvide u problematiku prouavanja i posebno ouvanja podzemnih gljiva. Za obradu, organizaciju i rukovanje podacima imao sam neprocenjivu pomo Dušana Gvozdenovia, a za specifi8ne procene diverziteta Prof. Jasmine Šinžar-Sekuli. Tekst je složen zahvaljuju6i pomoi Predraga Ilia, potrebne prevode ljubazno je uradila Vojislava Tasi, a originalni akvarel Bora Mili6evi6. Najveu zahvalnost upuujem porodici za bezrezervnu podršku. Prostorna distribucija i ekološke varijacije staništa hipogeinih makromiceta ( Mycota ) u Srbiji R EZIME Jedan broj gljiva iz razliitih taksonomskih grupa procesom konvergentne evolucije razvio je zajedniku ekološku strategiju formiranja makroskopskih podzemnih sporokarpa. Zbog te osobine nazvane su hipogeine ili podzemne gljive. One imaju znaajnu ulogu u šumskim ekosistemima kao fungalni partneri u simbiotskom odnosu sa drveem, a ovaj odnos je obostrano od esencijalne koristi za biljku i gljivu. Takoe su i važan izvor hrane za šumske životinje. U ovoj grupi najpoznatiji su tartufi, cenjene i skupe gljive koje se koriste u ljudskoj ishrani i imaju ekonomski znaaj, ali pored njih i brojne druge vrste imaju veliku i važnu ulogu u prirodi. Iako najstariji podaci o hipogeinim gljivama na podruju Srbije potiu još iz 18. veka, do danas je vrlo malo poznato o njima i postoji samo petnaestak objavljenih naunih radova i nekoliko kongresnih saopštenja, uglavnom s kraja 20. i poetka 21. veka. U njima su prvenstveno obraivani tartufi, dok o drugim grupama gotovo da nema nikakvih podataka i one su predstavljale veliku nepoznatu komponentu biodiverziteta Srbije. Jedan od razloga za mali stepen istraženosti hipogeinih gljiva, ne samo u Srbiji ve i globalno, je problematika njihovog prouavanja i pronalaženja u prirodi zbog kriptinog, podzemnog razvoja sporokarpa na osnovu kojih je prvenstveno mogue detektovati i locirati individue gljiva. Poev od 1991. godine zapoeta su istraživanja hipogeinih gljiva uz pomo obuenih pasa, inicirana od strane dr Miroljuba Milenkovia u kojima je uestvovalo više saradnika. Na taj nain formirana je zbirka koja je 2012. godine smeštena u fungarijum Prirodnjakog muzeja u Beogradu, i od tada je nastavljeno sa njenim prinavljanjem. Najvei broj nalaza i taksona koji su u njoj deponovani nisu do sada objavljeni. Glavni cilj ove disertacije je da se rasvetli i analizira diverzitet i položaj hipogeinih gljiva u ekosistemima Srbije, da se napravi sveobuhvatan uvid i analiza podataka, objedine znanja i na taj nain doprinese i unapredi fundamentalno poznavanje i razumevanje distribucije i ekologije hipogeinih gljiva u Srbiji i na evropskom kontinentu. Tokom rada nainjen je ukupan pregled istraživanja hipogeinih makromiceta na teritoriji Srbije sa istoriografskim podacima, kritiki su analizirana prethodna istraživanja i uraena je taksonomska revizija postojeih podataka. Kao glavni resurs korišen je materijal iz navedene zbirke koja je temeljno obraena, a svi nalazi su dopunjeni podacima o tanim geografskim lokacijama (georeferencirani su) i ekološkim uslovima na staništima gde su hipogeine gljive zabeležene u Srbiji. Na taj nain je izraen kompletan registar do sada poznatih i otkrivenih vrsta na podruju Srbije, a podaci o distribuciji tih vrsta su analizirani u cilju utvrivanja korelacija sa tipovima staništa i drugim ekološkim faktorima. Prisustvo veine konstatovanih vrsta je sada po prvi put publikovano za teritoriju Srbije, a jedan od taksona iz zbirke je nov za nauku. Zabeleženo je ukupno 68 taksona, što ini oko 40% poznatih evropskih vrsta, tako da je konstatovano daleko vee bogatstvo nego što je do sada bilo poznato, i popunjena je praznina na mapi rasprostranjenja hipogeinih gljiva u Evropi. Za sve vrste su prikazani opsezi variranja ekoloških parametara, optimalne vrednosti i ekološke valence, stepeni vezanosti, preferencije, specijalizacije i variranja staništa, prostorna i vremenska distribucija, areal karte, vrednosti ukupnog i procene potencijalnog diverziteta, kao i zavisnost od bioklimatskih inilaca. Razmotreni su faktori ugrožavanja i procenjene mere ouvanja, izdvojena su potencijalno najvrednija podruja za zaštitu sa najveom proporcijom komplementarnog diverziteta. Uoene su specifinosti i razlike hipogeine fungije Srbije i drugih evropskih regiona. Osim toga izneta su zapažanja o posebnostima metodologije i nainima za efikasno sprovoenje terenskih istraživanja podzemnih gljiva. Na ovaj nain su sumirana sva postojea znanja o hipogeinim gljivama u Srbiji. Izneti podaci i rezultati analiza osim naunog znaaja imaju temeljnu vrednost za aktivnosti procene i upravljanja prirodnim resursima, posebno u domenu zaštite i o8uvanja. K LJUNE REI: Biodiverzitet, Ekologija gljiva, fungija Srbije, hipogeine gljive N AUNA OBLAST: Ekologija U ŽA NAUNA OBLAST: Ekologija gljiva UDK BROJ: 581.52:582.28(043.3) Spatial distribution and habitat variability of hypogaeous macromycetes ( Mycota ) in Serbia APSTRACT Hypogaeous fungi have evolved a common ecological strategy of forming macroscopic underground sporocarps as a result of convergent evolution. This polyphyletic group includes a number of species from different taxonomic units. They play an important role in forest ecosystems as fungal partners in essential mutualistic symbiotic associations with trees where both organisms derive benefit. Furthermore, they are an important source of food for forest animals. The truffles, the prized and expensive fungi of culinary and economic importance, are the best known of this group, however, there are other numerous species that serve a large and significant role in nature. Although the oldest data on hypogaeous fungi from the territory of Serbia date back to the 18th century, very little is known about them to this day and there are only about a dozen published original scientific articles and several conference papers, mostly dating from the late 20th and the early 21st centuries. These articles and papers primarily addressed the truffles, and offered hardly any data about other groups, which presented a large unknown component of Serbia’s biodiversity. A reason that hypogaeous fungi had not been researched to a greater degree, either in Serbia or worldwide, lies in the difficulties associated with studying and detecting these fungi in nature, due to the cryptic, underground development of sporocarps, whose presence basically allows for the detection and location of the fungal individuals. The year 1991 saw a commencement of investigation of hypogaeous fungi with the help of trained dogs, initiated by Dr Miroljub Milenkovi involving a number of associates. Their efforts led to the formation of a collection which was housed in the Fungarium of the Natural History Museum in Belgrade in 2012. The collection has grown since then. Most findings and taxa deposited in the Fungarium had not been published until now. The main goal of this doctoral dissertation was to research, analyse and shed light on the diversity and status of hypogaeous fungi in Serbia’s ecosystems, provide a comprehensive survey and analysis of data, consolidate data from multiple sources and contribute to and advance the fundamental knowledge and understanding of distribution and ecology of hypogaeous fungi in Serbia and Europe. In the course of the research, an exhaustive survey of studies of hypogaeous macromycetes in Serbia’s territory was produced, complete with historiographic data, a critical analysis of earlier studies and a taxonomic revision of the existing data. The material from the above-mentioned collection served as the main resource. It was thoroughly examined and all findings were supplemented with georeferences and data on ecological
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