Praca Doktorska Wykonana W Katedrze Ekologii I Zoologii Kręgowców Instytutu Ekologii I Ochrony Środowiska

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Praca Doktorska Wykonana W Katedrze Ekologii I Zoologii Kręgowców Instytutu Ekologii I Ochrony Środowiska Stacjonarne Studia Doktoranckie Ekologii i Ochrony Środowiska Maciej Jażdżewski Porównanie strategii żerowania kozy Cobitis taenia complex (Linneusz, 1758) i kozy bałtyckiej Sabanejewia baltica (Witkowski, 1994) (Pisces, Cobitidae) Comparison of the foraging strategies of spined loach Cobitis taenia complex (Linnaeus, 1758) and northern golden loach Sabanejewia baltica (Witkowski, 1994) (Pisces, Cobitidae) Praca doktorska wykonana w Katedrze Ekologii i Zoologii Kręgowców Instytutu Ekologii i Ochrony Środowiska Promotor: prof. dr hab. Mirosław Przybylski Promotor pomocniczy: dr Eliza Szczerkowska-Majchrzak Łódź, 2020 PODZIĘKOWANIA Pragnę serdecznie podziękować mojemu Promotorowi, prof. dr. hab. Mirosławowi Przybylskiemu za wskazanie tematu badań, opiekę naukową na każdym etapie powstawania rozprawy doktorskiej, za pomoc i wyrozumiałość. Mojej Promotor Pomocniczej, dr Elizie Szczerkowskiej-Majchrzak dziękuję za wielki trud włożony w oznaczenie diety kóz. Pani dr hab. Krystynie Żuwale z Zakładu Anatomii Porównawczej Uniwersytetu Jagiellońskiego dziękuję za nieocenioną pomoc okazaną przy badaniu narządów węchu. Kierownikowi Katedry Ekologii i Zoologii Kręgowców, prof. dr. hab. Andrzejowi Krukowi dziękuję za wsparcie oraz życzliwość. Mojej drogiej Koleżance, dr Lidii Marszał oraz serdecznym Kolegom: dr. Bartoszowi Janicowi, dr. Dariuszowi Pietraszewskiemu i mgr. Szymonowi Tybulczukowi składam wyrazy wdzięczności za duchowe wsparcie oraz za pomoc w pracach terenowych i przy przygotowaniu maszynopisu. Wszystkim Koleżankom i Kolegom z Katedry Ekologii i Zoologii Kręgowców pragnę podziękować za gotowość niesienia pomocy oraz przyjazną atmosferę, którą mnie otoczyli. Mojej kochanej Rodzinie i Bliskim dziękuję za bezwarunkowe wspieranie mnie w trudnych momentach oraz za nieskończoną cierpliwość. 2 Mojej najdroższej Żonie Eli oraz ukochanym Dzieciom, Jasiowi i Antosiowi 3 Spis treści WPROWADZENIE ..................................................................................................................6 I. Porównanie diety i aktywności żerowania kozy Cobitis taenia complex (Linneusz, 1758) i kozy bałtyckiej Sabanejewia baltica (Witkowski, 1994) ........................................................ 10 I. 1. WSTĘP ........................................................................................................................ 10 I. 2. TEREN BADAŃ ......................................................................................................... 16 I. 3. MATERIAŁ I METODY ............................................................................................. 17 I. 3.1. Aktywność żerowania ............................................................................................... 18 I. 3.2. Ogólna analiza diety .................................................................................................. 19 I. 4. WYNIKI ...................................................................................................................... 25 I. 4.1. Aktywność żerowania ............................................................................................... 25 I. 4.2. Ogólna analiza diety .................................................................................................. 27 I. 4.3. Różnice w diecie między klasami wielkości kóz w porach roku ................................. 30 I. 4.4. Preferencje siedliskowe względem wyróżnionych kategorii pokarmowych ................ 32 I. 4.5. Analiza zawartości detrytusu w przewodach pokarmowych ryb ................................. 33 I. 4.6. Szerokość i nakładanie się niszy pokarmowej ............................................................ 33 I. 5. DYSKUSJA ................................................................................................................. 37 I. 5.1. Aktywność żerowania ............................................................................................... 37 I. 5.2. Ogólna analiza diety .................................................................................................. 44 I. 5.3. Różnice w diecie między klasami wielkości kóz w porach roku ................................. 49 I. 5.4. Preferencje siedliskowe kóz w oparciu o wyróżnione kategorie pokarmowe .............. 50 I. 5.5. Analiza zawartości detrytusu w przewodach pokarmowych ryb ................................. 51 I. 5.6. Szerokość i nakładanie się niszy pokarmowej ............................................................ 53 I. 6. LITERATURA ............................................................................................................ 56 I. 7. TABELE...................................................................................................................... 68 I. 8. RYCINY...................................................................................................................... 74 Apendyks 1 ......................................................................................................................... 84 II. Budowa narządów węchu kozy Cobitis taenia complex (Linneusz, 1758) i kozy bałtyckiej Sabanejewia baltica (Witkowski, 1994) ................................................................................. 86 II. 1. WSTĘP ...................................................................................................................... 86 II. 2. MATERIAŁ I METODY ............................................................................................ 92 II. 3. WYNIKI..................................................................................................................... 94 II. 4. DYSKUSJA ............................................................................................................... 96 II. 5. LITERATURA ......................................................................................................... 105 4 II. 6. TABELE .................................................................................................................. 110 II. 7. RYCINY .................................................................................................................. 110 PODSUMOWANIE .............................................................................................................. 113 PIŚMIENNICTWO .............................................................................................................. 115 STRESZCZENIE .................................................................................................................. 119 SUMMARY.......................................................................................................................... 122 5 WPROWADZENIE Dobór naturalny sprzyja zwierzętom, które wydajnie żerują, maksymalizując stosunek korzyści do kosztów zdobywania pokarmu. Korzyści obejmują pozyskane składniki odżywcze o znaczeniu energetycznym, budulcowym i regulatorowym, podczas gdy do kosztów należy zużycie energii, czas poświęcony na zdobycie pokarmu oraz narażenie na oddziaływanie drapieżników lub pasożytów (Helfman i inni 2009). Współistniejące gatunki ewoluują pod względem efektywności wykorzystania poszczególnych zasobów, zmniejszając tym samym skutki konkurencji międzygatunkowej. Efektem konkurencji może być zróżnicowanie niszy w obrębie populacji, ale zwiększona różnorodność może wynikać również z plastyczności behawioralnej, bez konieczności powszechnie zakładanych ewolucyjnych zmian o charakterze morfologicznym i anatomicznym (Svanbäck i Bolnick 2007). Występowanie dwóch sympatrycznych gatunków może prowadzić do różnych rodzajów koegzystencji (Ross 1986, Hartney 1989). Gatunki ryb mogą współistnieć w tym samym siedlisku dzięki drobnym różnicom w wykorzystaniu zasobów na poziomie mikrosiedliska lub w wyniku przemieszczenia jednego z nich do siedliska alternatywnego (Larson 1980, Werner i inni 1983). Należy się spodziewać, że problem współwystępowania będzie szczególnie interesujący w przypadku gatunków o bardzo zbliżonej morfologii i biologii. W polskiej ichtiofaunie do takich gatunków należą dwie ryby bentoniczne: koza Cobitis taenia (Linneusz, 1758) i koza bałtycka Sabanejewia baltica (Witkowski, 1994) z rodziny kozowatych (Cobitidae). Ryby należące do rodzaju Cobitis są szeroko rozpowszechnione w Eurazji i północnej Afryce (Bǎnǎrescu 1990, Perdices i Doadrio 2001, Kottelat i Freyhof 2007). Stanowią szczególny przedmiot zainteresowań naukowców z powodu hybrydyzacji 6 zachodzącej pomiędzy nimi, prowadzącej do powstawania poliploidów wskutek dodania genomu innego gatunku. Badania chromosomów wykazały, że gatunki z rodzaju Cobitis składają się zarówno z populacji diploidalnych (jednogatunkowych), jak i hybryd diploidalno-poliploidalnych, w których osobniki jednego gatunku współwystępują z poliploidalnymi hybrydami. Ich różnorodność biologiczna jest przedmiotem intensywnych badań, które oprócz analizy morfologicznej (Vasil’eva i Vasil’ev 1998, Jeleń i inni 2008, Kotusz 2008) i molekularnej (Janko i inni 2005, 2007), obejmują również zmienność chromosomalną (Boroń 1992, 1999, 2001, 2003, Ráb i inni 2000, Boroń i inni 2006). Na występujące w Polsce ryby z rodzaju Cobitis składają się diploidalne osobniki C. taenia oraz tri- i tetraploidalne osobniki hybrydowe powstałe między gatunkami należącymi do: C. tanaitica, C. elongatoides oraz C. taenia (Boroń 2003, Boroń i inni 2006). Rozróżnienie osobników diploidalnych od poliploidalnych możliwe jest w zasadzie tylko przy pomocy badań cytogenetycznych (Kotusz 2000, Bohlen i inni 2005). Kozy występujące na stanowisku badań (w Mysiakowcu na rzece
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