ࡱ> cebh{TV7 AbjbjUU +7|7|Ml,,,,,,,@R R R 8  ,@(N ^4"VVViiiU"x'$) ,|',ieiii',,VVe(i,V,Vi=,,eV @CP@ R Ue${(0(],+,e@@,,,,Prof. dr. sc. uro Huber 27. o~ujka 2001. Zavod za biologiju 16. travnja 2001. Veterinarski fakultet 06. svibnja 2001. Heinzelova 55 10000 Zagreb Tel. 2390-141, Fax. 244-1390 E-mail: huber@vef.hr Stan: Slavka Kolara 45b 10410 Velika Gorica (Tel. 6218-345) Za "LOVSTVO" (za Hrvatski lova ki savez) 4. LOVNO GOSPODARENJE 4. 1. Bioloaki temelji uzgoja i zaatite divlja i (zooekologija) 4. 1. 1. Populacija vrste i njeno podru je 4. 1. 1. 1. Brojnost populacije i zakonitosti koje ju reguliraju Pod populacijom podrazumijevamo skupinu jedinki iste vrste koje istodobno ~ive na odreenom prostoru i do odreene su mjere reproduktivno izolirane od susjednih populacija te vrste. Tako neko jezero ima svoju populaciju odreenih vrsta riba, neki grad svoju populaciju golubova ili neki planinski lanac svoju populaciju medvjeda. Susjedne se populacije mogu razlikovati u u estalosti pojedinih obilje~ja odnosno gena koji ta obilje~ja uvjetuju. Populacije meu kojima postoji ja a i dugotrajnija izolacija u pravilu se viae meusobno razlikuju. Gledano kroz geoloaka razdoblja takova izolacija mo~e dovesti i do nastajanja odvojenih vrsta tijekom mnogo tisua godina. Svaka populacija ima sposobnost i tendenciju rasta, jer bioloaki mo~e proizvesti viae potomaka nego ato je jedinki u roditeljskoj generaciji i viae nego ih ugiba od starosti. Sposobnost proizvodnje "viaka" potomaka jedna je od bitnih poluga evolucijskog izbora odnosno prirodne selekcije. Samo uz postojanje "viaka" mogu je odabir "bolje prilagoenih" koji kroz veu reprodukciju tijekom niza generacija dovode do evolucijskog pomaka odnosno do ustaljivanja ponekih novih osobina i mogueg nastanka nove vrste. Istodobno je sposobnost populacije da raste ograni ena uvjetima okoliaa. Ekoloako je pravilo da se otpor okoliaa na rast populacije poveava sa porastom broja jedinki u populaciji. To zna i da po etna populacija u povoljnim uvjetima okoliaa brzo raste, ali se taj rast usporava sa pribli~avanjem noseeg kapaciteta staniata, te se dalje zaustavlja oko te vrijednosti (Slika 1). Slika 1.Predvidljivi rast neke populacije na odgovarajuem novom staniatu tijekom vremena. Postoji faza blagog ubrzavanja rasta, prekofaze eksponencijalno brzog rasta i postupnog usporavanja (sigmoidna krivulja). Rast se zaustavlja pri dosizanju kapaciteta staniata (K), ovdje previenom na 1900 jedinki.  EMBED Excel.Chart.8 \s  4. 1. 1. 2. Sastav populacije Sastav populacije specifi an je za pojedine vrste ovisno o njenom evolucijskom razvoju i bioloakim osobitostima. Na primjeru vrsta gdje su spolovi odvojeni i gdje novoroeni potomci predstavljaju samostalne jedinke, sastav se pojedine populacije mo~e prikazati kao "dobna" ili "uzrastna" piramida. Istodobno se mo~e prikazati i udio jedinki pojedinog spola (Slika 2a, b, c). Dok je omjer spolova, barem pri roenju, za veinu vrsta ravnomjeran, udio po dobnim skupinama normalno je najvei u najmlaih pripadnika populacije i progresivno se smanjuje s njihovim starenjem. Brzina gubljenja jedinki sa dobi ovisna je o generacijskom intervalu i fizioloakom trajanju ~ivota pojedine vrste. To mo~e biti mjereno brojem dana ili tjedana za mnoge beskraljeanjake, odnosno brojem mjeseci i godina za razne kraljeanjake. `iroka baza svake takove piramide ozna ava veliki reprodukcijski uspjeh i potencijal rasta. Baza piramide koja je u~a od udjela starijih dobnih kategorija upozorava na zastoj u reprodukciji i vodit e k izumiranju populacije ako se ograni avajui inioci na vrijeme ne promijene. Slika 2a. Normalni hipotetski model dobne i spolne strukture populacije jelena  EMBED Excel.Chart.8 \s  Slika 2b. Model populacije jelena sa uspjenom reprodukcijom ali bez viih dobnih razreda  EMBED Excel.Chart.8 \s  Slika 2c. Populacija jelena sa slabom reprodukcijom i loim izgledima za opstanak  EMBED Excel.Chart.8 \s  4. 1. 1. 3. Prirast populacije i njena kolebanja Prema standardnom modelu prikazanom na Slici 1 nova populacija u odgovarajuem staniatu raste dok ne dosegne njegov nosei kapacitet. Slijedi najdulje vrijeme postojanja neke populacije, a to je kada je ona u razini kapaciteta staniata. Pogreano je vjerovati da se ona u tom razdoblju stalno u istom broju. Stalna su kolebanja iznad i ispod stvarnog kapaciteta, tako da populacije mnogih vrsta gotovo nikada i nisu u stvarnoj ravnote~i sa svojim staniatem. Kolebanja ili oscilacije mogu biti u pravilnim vremenskim razmacima pa ih zovemo periodi kima, ili su nepravilna, odnosno neperiodi ka (Slika 3). Periodi ka kolebanja: Mnoge vrste pokazuje velike sezonske promjene brojnosti jedinki, a jedan od primjera su populacije planktona u nekom jezeru ljeti u odnosu na zimu. Klasi ni ekoloaki primjer viaegodianje periodi ke oscilacije su leminzi (Lemmus sp.) u tundri. Oni ~ive u razmjernoj sigurnosti svojih podzemnih hodnika i hrane se podzemnim i nadzemnim dijelovima biljaka. To im omoguuje nesmetan rast populacije koja svake tree do etvrte godine premaai kapacitet staniata. U nestaaici hrane i u dodatnom stresu zbog velike brojnosti vlastite vrste leminzi napuataju svoja skloniata i kreu u masovne migracije. U tim uvjetima predstavljaju obilje hrane za lokalne predatore, posebno za polarne lisice i snje~ne sove, koji se posljedi no tome izuzetno uspjeano razmno~avaju. Populacija leminga u istoj sezoni padne na mali postotak po etne (Slika 3 - Periodi ke oscilacije). Tada se vegetacija tundre mo~e oporaviti, ali istovremeno predatori viae ne nalaze hrane za svoje poveane populacije. Snje~ne sove tada esto lete daleko na jug u potrazi za hranom. Veina tih sova se nikada ne vrati u tundru, a njihovo periodi ko pojavljivanje svake 3 do 4 u nekim gradovima u Sjedinjenim Ameri kim Dr~avama prepoznato je kao odraz drasti nog pada populacije leminga u dalekoj tundri. Ciklus populacije bijelog zeca i kanadskog risa traje 9 do 10 godina i drugi je klasi an primjer u ekologiji populacija. Opisan je u poglavlju odnosa predatora i plijena (4. 1. 2. 1. 2. 2.). Sli ne cikluse pokazuju i neki kukci ije se li inke hrane aumskim liaem kao i migratorni postinjski skakavci (Locusta migratoria). Neperiodi ka kolebanja: Porasti i padovi veli ina nekih populacija, a koji se ne ponavljaju u pravilnim vremenskim razmacima, obi no slijede nagle promjene u staniatu koje su mogli izazvati po~ari, poplave, erupcije vulkana, ili zahvati ovjeka. Slika 3. Pojednostavljeni prikaz rasta populacije do kapaciteta staniata (K), te razdoblja periodi kih (u ciklusu od 5 godina) i neperiodi kih kolebanja brojnosti jedinki sa kona nim izumiranjem populacije  EMBED Excel.Chart.8 \s  Ope je pravilo za sve vrste oscilacija populacija da su one ja e i da imaju trajnije posljedice u jednoli nijim staniatima. Primjer takovog staniata je tundra gdje razmjerno malen broj vrsta ~ivi sa velikim brojem jedinki. ovjek takoer ima sklonost proizvoditi jednoli na staniata gdje pokuaava uzgajati po~eljne vrste u monokulturi; bilo da su to farme sa velikim brojem domaih ~ivotinja, polja sa poljoprivrednim kulturama, planta~ne aume ili ograena uzgajaliata divlja i. U svim tim uvjetima mo~e neki iznenadni vanjski utjecaj dovesti do propasti takovih populacija: novi virusni ili bakterijski uzro nik bolesti, novi nametnik ili kukac koji se hrani tim vrstama, kao i nagla klimatska promjena. Nasuprot tome ekosustav bogat razli itim vrstama, poput tropske vla~ne aume, ima mnogostruke mehanizme koji djeluju im neka od lokalnih populacija po ne odstupati od ravnote~e oko kapaciteta staniata. Zato su oscilacije tamo bla~e i kratkotrajnijeg u inka. Mo~e se zaklju iti da, iako svaka populacija te~i poveanju svoje brojnosti, da hiperpopulacija nije niti jednoj u interesu. Zato su se razvili brojni mehanizmi samoregulacije, od kojih neki djeluju brzo i ne dopuataju znatnija kolebanja, a drugi se aktiviraju tek kod veih odstupanja pa su kolebanja dramati nija. 4. 1. 2. Okolina i njen upliv na populaciju 4. 1. 2. 1. }ivi imbenici okoline i meusobna ovisnost 4. 1. 2. 1. 1. Populacija vrste kao imbenik okoline Svaka vrsta ima svoju odreenu ekoloaku niau koja definira njenu ulogu u ekosustavu, odnosno njene odnose sa cjelokupnim ~ivim i ne~ivim okoliaem. Putem svoje niae pojedina vrsta izravno utje e na brojna meudjelovanja u prehrambenom lancu, te neizravno i na sve procese u sustavu uklju ujui i njegove ne~ive inioce. Za upravljanje vrstom i njenim staniatem po~eljno je poznavati ekoloaku niau te vrste. U praksi se naj eae mnogi elementi niae neke vrste vidi tek kad ta vrsta nestane. Sli an mo~e biti i u inak unoaenja nove vrste u neko staniata gdje ona, ili ne mo~e nai svoju niau pa propadne, ili ue u niau jedne ili viae autohtonih vrsta te njih potisne. Sve te situacije upozoravaju da svaka vrsta ima u inak na sveukupne odnose u staniatu i da to treba poatovati u svim gospodarskim zahvatima. 4. 1. 2. 1. 2. Populacija drugih vrsta kao imbenik okoline Izmeu pripadnika razli itih vrsta razvili su se vrlo razli iti odnosi, koje nazivamo interspecijskim odnosima. Meusobni u inak dviju vrsta mo~e imati negativni predznak (-), pozitivni predznak (+), ili biti neutralan (0). Sve mogue kombinacije tih predznaka daju ukupno 9 skupina razli itih interspecijskih odnosa: Vrsta A Vrsta B Odnos 0 : 0 Neutralizam (Za svaku od vrsta je prisue druge bez utjecaja. Rijetko ili teako je dokazati da je upravo tako.) - : - Kompeticija - aktivna (Obje vrste trpe i meusobno se aktivno bore za prostor i hranu; npr. toksi nim tvarima.) - : - Kompeticija - pasivna (Obje vrste trpe, jer si meusobno iscrpljuju izvore hrane, vode i drugih potreba) - : 0 Amenzalizam (Vrsti A smeta prisue vrste B dok na nju vrsta A nema utjecaja.) + : - Prazitizam (Vrsta A je nametnik koji ima koristi od svog nosioca B, a on zbog toga trpi i odnos traje dulje.) + : - Predacija (Vrsta A je grabe~ljivac koji u kraem odnosu nanosi tetu jedinki vrste B koja predstavlja plijen.) + : 0 Komenzalizam (Vrsta A je komenzal koji ima koristi od vrste B, a za koju je odnos neutralan) + : + Protokooperacija (Obje vrste imaju korist ali odnos nije obligatan.) + : + Simbioza (Obje vrste imaju korist i odnos je obligatan, odnosno niti jedna od vrsta ne mo~e ~ivjeti samostalno.) 4. 1. 2. 1. 2. 1. Odnosi meu konkurentskim vrstama Od ovih nabrojanih moguih odnosa izmeu razli itih vrsta, odnosi "kompeticije" su jedni od naj eaih. Primjeri gdje se konkurentske vrste aktivno meusobno bore rjei su u odnosu na u estalost pasivne konkurencije. Tako se jedna vrsta trave u ameri koj preriji zove prstenasta trava, jer izlu uje toksi nu tvar koja sprje ava rast drugih vrsta trava u krugu oko nje. Drugi primjer su razli ite vrste nametni kih helminata (crva) koji takoer pomou otrovnih izlu evina atite svoj prostor u probavnom traktu nosioca od drugih vrsta nametnika. U prirodi je naj eaa pasivna kompeticija. Tako su sve ~ivotinje koje se hrane travom i dijele isti paanjak u tom odnosu. To vrijedi kako za razli ite vrste divljih travojeda, tako i odnose meu vrstama domaih ~ivotinja ili za konkurenciju pri paai domaih i divljih ~ivotinja. Sli no je i sa ograni enim koli inama vode na primjeru suane sezone u afri koj savani. Iako se te ~ivotinje rijetko meusobno tjeraju sa zaliha zajedni kih potreba, sve ato potroai jedna vrsta izravno nedostaje drugoj, te bi svakoj vrsti odgovaralo da one druge nema. Opisani negativni u inci ipak su ovdje promatrani prvenstveno na razini jedinke. Dakle ono ato odreenoj jedinci neke vrste predstavlja nesumnjivi minus, ne mora uvijek biti i minus za cijelu vrstu. Konkurentski odnosi zaoatravaju selekcijski pritisak, te osiguravaju da e se uspjeano razmno~avati i svoje gene prenositi u potomstvo samo najbolje prilagoene jedinke. 4. 1. 2. 1. 2. 2. Odnosi izmeu predatora i plijena Za razliku od parazita i njegova nosioca, predator (grabe~ljivac) je u pravilu vei ili je podjednak svom plijenu (~rtvi), kratko ga koristi, a to za ~rtvu gotovo uvijek zna i i smrt. Evolucijski odnosi su predatoru omoguili razvoj mnogih tjelesnih obilje~ja za uspjeanije nala~enje, hvatanja i savladavanje plijena: dobra osjetila, brzinu tr anja, letenja ili plivanja, sna~ne eljusti, zube, kljun, pane, lovke, otrove... Istovremeno je i plijen razvijao sposobnosti izbjegavanja postajanja ~rtvom: prikrivanjem u boji okoline (mimikrija), skrivanjem u predatoru nedostupna skloniata, brzinom bijega, oklopima, bodljama, otrovima... U tome treba posebno promatrati ulogu ovjeka kao predatora koji se pojavio i tehnoloaki se razvio neusporedivo br~e nego ato su evolucijski procesi mogli razviti odgovarajue obrambene mehanizme u vrsta plijena. Stoga ovjek nema ekoloaku ulogu usporedivu onoj ostalih predatora. On mora svjesno prilagoavati svoje predatorske moi kako ne bi bitno naruaio ekoloake odnose meu vrstama svog plijena, a koje uklju uju i prirodne vrhunske predatore. Joa izra~enije nego kod odnosa konkurencije, negativan utjecaj (-) predatora na plijen treba promatrati strogo na razini jedinke. Svaka populacija plijena ima dugoro no koristi od svojih neposrednih predatora: putem prirodne selekcije najprilagoenijih i regulacijom veli ine populacije prilagoene kapacitetu staniata. Kad predatori izgube mogunost takovog djelovanja, regulaciju veli ine plijena preuzimaju ograni ene koli ine hrane u staniatu, te razne nametni ke, bakterijske ili virusne bolesti. U nekih vrsta to ograni enje mogu biti socijalni odnosi ili pak nedostatak mjesta za skrivanje, odmaranje ili odgoj potomstva. Zajedni kom evolucijom sa svojim predatorima plijen je postupno prilagoavao svoje tjelesne, psihi ke i reprodukcijske sposobnosti upravo tako da opstane u danim uvjetima. Da je njegova evolucija tekla bez prisua odreenog predatora zasigurno bi rezultat te evolucije bile ~ivotinje bitno razli itih obilje~ja. Primjere za to imamo u vrstama koje su se razvile na raznim oto nim skupinama bez predatora, te su posljedi no vrlo trome, bez mogunosti leta ili obrane. Sjetimo se istrijebljene ptice Dodo sa otoka Mauricijusa ili mnogih vrsta ptica sa Havajskog oto ja. Veli ine populacija u ponekim sustavima plijena i predatora mijenjaju se u ciklusima, odnosno pokazuju periodi ka kolebanja. Poznati i to no dokumetirani takav primjer su populacije bijelog zeca i kanadskog risa (Slika 4). Zahvaljujui sretnoj okolnosti, da su inovnici Haudson Bay kompanije pri dugogodianjem otkupljivanju krzna od trapera za svako preuzeto krzno zabilje~ili vrstu i datum, danas raspola~emo nizom od 90 godina podataka gdje brojevi u pojedinim godinama dosi~u i 150 tisua jedinki. Veliki brojevi omoguuju statisti ku obradu sa razlikovanjem trendova koji su posljedica slu ajnosti od onih koji imaju zna ajnu zakonitost. Tako ove krivulje pokazuju da obje vrste imaju razdoblja velikih populacija, kao i razdoblja svojih minimuma. Redoviti razmak izmeu dva vrha ili dva dna krivulje je 9 ili 10 godina. Takoer se vidi da uzlazne i silazne krivulje zeca prate iste takove krivulje risa sa zaostatkomod jedne do dvije godine. Naravno brojevi uhvaenih i oguljenih risova bili su u pravilu oko 50% manji nego brojevi istih u ze eva. Na tom primjeru objavljen je vei broj klasi nih znanstvenih ekoloakih populacijskih studija. Ozbiljne analize su dokazale da kretanje broja ze eva izravno utje e na veli inu populacije risa, ali ne i obratno; stvarni broj risova ima malen utjecaj na kolebanje populacije zeca. Tako godine rasta broja ze eva omoguuju bolju prehranu, a time i uspjeaniju reprodukciju risova, i to putem ranijeg spolnog sazrijevanja ~enki, njihovog veeg postotka oploenosti, veeg broja roene mladun adi i, ato je posebno va~no, njihovog velikog postotka pre~ivljavanja. Ovo posljednje je mogue jer dobro hranjena majka proizvodi viae mlijeka, a kad mladun ad po inje jesti meso ona ih ne mora zbog odlaska u lov predugo ostavljati same i nezaatiene, te se redovito vraa i sa uhvaenim plijenom. Takova situacija omoguava rast populacije risa i za preko 50% godianje. Nasuprot tome, kada je smjer krivulje populacije zeca u opadanju, a risova joa uvijek ima mnogo, u jednom trenutku risovi viae nemaju dovoljno svoje glavne hrane. Odrasli risovi to uglavnom pre~ivljavaju ali se tada dogaa suprotan trend u reprodukciji. Malo ili nimalo mladun adi pre~ivljava do spolne zrelosti, te i populacija risa naglo dobiva negativni trend. Dok je zec risu glavni plijen, ris je samo jedan od vie predatora na zeca. Ipak svi ti predatori zajedno imaju manji utjecaj na populaciju zeca od drugih mehanizama kontrole njegove populacije. Kad se populacija zeca pribli~i vrhuncu njima ponestaje hrane i skloniata, te postaju ranjivi za razne bolesti. Uzro nici svih bolesti mnogo se lakae i br~e aire meu svojim nositeljima kada je njihova populacija guaa i kontakt eai. Neka novija istra~ivanja pokazuju da ak i mijene sun eve aktivnosti utje u na ~ivotne uvjete, a time i na veli inu populacije zeca. Slika 4. Ciklus bijelog zeca i kanadskog risa na osnovi broja otkupljenih krzna u Kanadi.  Postoje i primjeri kako predatori kontroliraju veli inu populacije svog plijena. To se najbolje vidi ako se u nekom podru ju uklone svi predatori. Na zaravni Kaibab u Arizoni (SAD) ljudi su po etku 20. stoljea, to nije 1907. istrijebili sve velike zvijeri: vuka, pumu i kojota (Slika 5). Akcije nije bila poduzeta zbog ekoloakog pokusa nego zbog tra~enja farmera da se zaatiti njihova stoka. U narednih 11 godina populacija jelena je sada po etnih 4000 narasla 10 puta, a do 1925. i na 100000 jedinki. Uskoro je staniate bilo iscrpljeno prekomjernom paaom i bratenjem, a za jakih zima 1925. i 1926. od gladi je uginulo preko 60000 jelena. Preostala populacija jelena vie nikada nije dosegla svoj vrhunac ali se stabilizirala na broju viem nego to bi on bio da su prisutni predatori. Slika 5. Odnos populacije predatora i jelena, te samoregulacija prenamno~ene populacije jelena nakon iscrpljivanja staniata.  4. 1. 2. 1. 3. Raslinje kao osnova za opstanak divlja i inioci ne~ive prirode poput tla i klimatskog re~ima odreuju kakav e se razviti vegetacijski pokrov na nekom podru ju. Biljke tog vegetacijskog pokrova su jedini organizmi na Zemlji koji mogu u zna ajnijim koli inama od anorganske materije proizvoditi organsku, i to putem sun eve energije u procesu fotosinteze. Organska tvar nastala fotosintezom su ugljikohidrati koji u sebi nose vezanu energiju poteklu od svjetlosti sa Sunca. }ivotinje kao svoju hranu mogu rabiti isklju ivo organsku tvar, pa tako sva hrana o kojoj ovise potje e od biljaka. ak i ako je ta ~ivotinja meso~der, ona neizravno dobiva energiju svojedobno uhvaenu fotosintezom. Naravno da koli ina biljne mase u nekom staniatu ograni ava broj odnosno masu ~ivotinja koje se mogu tim biljem hraniti. Biljna masa je, kao i svi ~ivi organizmi, obnovljivi prirodni izvor. Njena proizvodnja je evolucijski prilagoena potroanji od strane ~ivotinja. U uravnote~enom ekoloakom sustavu su proizvodnja i potroanja biljne mase podjednaki. Do poremetnji te ravnote~e mo~e doi zbog zastoja u proizvodnji biljne mase ili zbog porasta potroanje povezanog sa rastom populacije biljo~dernih ~ivotinja. Klimatske i druge nepogode poput suaa, dugih zima, kasnih mrazeva, ili po~ara i poplava mogu bitno smanjiti veli inu biljne produkcije u nekom podru ju kroz odreeno vrijeme. Porast populacije biljo~dera, bez obzira na razlog tom porastu, mo~e pak bitno oatetiti biljni pokriva  nekog podru ja. Primjer porasta populacije jelena zbog eliminacije predatora i posljedi no iscrpljivanje biljne osnove staniata praeno pomorom tih jelena, opisan je u prethodnom poglavlju (Slika 5). Slika X. Prora un veli ine populacije prema udjelu obilje~enih jedinki pri ponovljenom hvatanju. Rezultat treba korigirati za postotak promjene sastava populacije u meuvremenu: roene i uginule, te imigrirale i emigrirale ~ivotinje. Slika X. Promjene u sastavu populacije prepelica tijekom tri godine PAGE  PAGE 2 8  1 2 3 4 !!!!!!!P!(,(22P6R666666@@K2K~þ{>*OJQJmHsH jw(Ujb> UV6OJQJ]mHsH jUj9> CJOJQJUV jDUj> UV jUj[> CJOJQJUVj> UV jUCJOJQJmHsH5CJOJQJ\mHsHOJQJmHsH/\ &@z8prt0 (*$dha$dhd'@ 5 6 !!!!P!0&^&'>01224P666>@@@XAdh`dhXAAHHHK2KK"LLMMMpNN"OOP=P{PPPP1QrRtRR pdh^p`dh`dhRWf[J^L^^&`6gpĆƆȆʆ~Ռ֌׌،ٌ"ĝdh`dh~ĝ'(./0239:;<=@A0JmHnHu0J j0JUmHsHjkOJQJUmHsHOJQJmHsHj0OJQJUmHsHĝƝ'012=>?@A&`#$dh ) 0 00&P . 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