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Cervus Elaphus) Females in North-Eastern Poland

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Cervus Elaphus) Females in North-Eastern Poland University of Warsaw Faculty of Biology Tomasz Borowik Factors affecting fertility of red deer (Cervus elaphus) females in north-eastern Poland Doctoral dissertation in biological sciences Discipline: biology Thesis conducted at the Mammal Research Institute Polish Academy of Sciences, Białowieża Dissertation conducted under supervision of: Prof. Dr habil. Bogumiła Jędrzejewska Mammal Research Institute, Polish Academy Sciences Warsaw – Białowieża, July 2014 Uniwersytet Warszawski Wydział Biologii Tomasz Borowik Czynniki kształtujące płodność samic jelenia (Cervus elaphus) w północno-wschodniej Polsce Rozprawa doktorska w zakresie nauk biologicznych w dyscyplinie biologii Praca wykonana w Instytucie Biologii Ssaków Polskiej Akademii Nauk w Białowieży Praca wykonana pod kierunkiem Prof. dr hab. Bogumiły Jędrzejewskiej Instytut Biologii Ssaków Polskiej Akademii Nauk Warszawa – Białowieża, lipiec 2014 Oświadczenie kierującego pracą Oświadczam, że niniejsza praca została przygotowana pod moim kierunkiem i stwierdzam, że spełnia ona warunki do przedstawienia jej w postępowaniu o nadanie stopnia doktora nauk biologicznych w zakresie nauk biologicznych w dyscyplinie biologii. Data Podpis kierującego pracą Oświadczenie autora pracy Świadom odpowiedzialności prawnej oświadczam, że niniejsza rozprawa doktorska została napisana przeze mnie samodzielnie i nie zawiera treści uzyskanych w sposób niezgodny z obowiązującymi przepisami. Oświadczam również, że przedstawiona praca nie była wcześniej przedmiotem procedur związanych z uzyskaniem stopnia doktora w innej jednostce. Oświadczam ponadto, że niniejsza wersja pracy jest identyczna z załączoną wersją elektroniczną. Data Podpis autora pracy Streszczenie Celem badań, przeprowadzonych w Puszczach Augustowskiej, Knyszyńskiej i Białowieskiej (północno-wschodnia Polska) w latach 2006-2011, było określenie parametrów rozrodczych samic jelenia szlachetnego (Cervus elaphus) oraz czynników kształtujących ich płodność. Parametry rozrodcze określono w oparciu o analizę narządów rozrodczych samic. Łącznie we wszystkich objętych badaniami kompleksach leśnych zebrano 541 prób. W porównaniu z innymi europejskimi populacjami, jelenie w badanych kompleksach leśnych występowały w niskich zagęszczeniach, łanie wykazywały duży potencjał rozrodczy, który był wyrażony wysokim poziomem płodności (85%), wczesnym przystępowaniem do rozrodu (1,5-roczne samice) oraz brakiem spadku płodności z wiekiem. W skali biogeograficznej wykazano, iż poziom płodności łań był ujemnie skorelowany z zagęszczeniem populacji. W północno-wschodniej Polsce prawdopodobieństwo płodności samic wzrastało z ich wiekiem i kondycją. Czynniki środowiskowe (niskie temperatury i obfite opady latem) mogły mieć pośredni, negatywny wpływ na płodność poprzez kondycję samic. Wśród płodów wykazano większy udział płodów męskich (1:0.68). Zależne od wieku różnice w kondycji fizycznej samic wykazywały tendencję, w której łanie w lepszej kondycji fizycznej miały większy udział płodów męskich, co potwierdzałoby hipotezę Triversa Willarda. Poziom śmiertelności cieląt w pierwszych 6 miesiącach życia był wysoki (41%) i wzrastał wraz z wiekiem samic. Oczekiwana długość życia nowo narodzonych samic wynosiła 4,3 roku. Wykazano zbliżony, niezależny od wieku, poziom śmiertelności samic. Młode samice (1-3-letnie) miały największy wkład w rozrodczość populacji (46%). W latach badań rozrodczość bilansowała ubytki powodowane przez śmiertelność, dlatego przewiduje się stabilizację badanej populacji jeleni. Uzyskane wyniki mogą mieć praktyczne zastosowanie w zarządzaniu populacjami jelenia i planowaniu wielkości odstrzałów łowieckich. Słowa kluczowe jeleń szlachetny, Cervus elaphus, płodność, rozrodczość, samice, zagęszczenie populacji, kondycja, proporcja płci, GIS, meta-analiza Tytuł pracy w języku angielskim Factors affecting fertility of red deer (Cervus elaphus) females in north-eastern Poland Acknowledgements First and foremost, I would like to express my special appreciation and thanks to my supervisor, Professor Bogumiła Jędrzejewska for her invaluable help and unceasing support and advice starting from the study planning, fund raising, project realization to writing the thesis. I am indebted to Professor Jędrzejewska for her patience, encouragement and knowledge sharing. I would like to thank all foresters and hunters representing the State Forests (National Forest Holding) and the Polish Hunting Association for their great contribution in the sample collection process and drive censuses. I am deeply grateful to Dr. Piotr Wawrzyniak (Regional Directorate of State Forests in Białystok) for his remarkable engagement and helping hand in the coordination of sampling and for providing many expert suggestions, which improved my study. Furthermore, a thank you to all the staff from Mammal Research Institute who always kindly helped me in solving any problems I came across, and who created a friendly working atmosphere. My special thanks go to Dr. Karol Zub and Jakub Bubnicki for their help in statistical and GIS analyses. Finally, I want to thank my family for their continuous love and support. The project was financed by the Ministry of Science and Higher Education (grant number N N304 094034) and the budget of the Mammal Research Institute PAS. Table of contents ABSTRACT ........................................................................................................................ 1 ABSTRAKT ........................................................................................................................ 3 1. INTRODUCTION ........................................................................................................... 5 2. AIMS OF THE STUDY .................................................................................................. 11 3. METHODS AND MATERIAL ...................................................................................... 12 3.1. Study area ........................................................................................................... 12 3.2. Characteristics of the red deer population .......................................................... 17 3.3. Data collection .................................................................................................... 19 3.3.1. Habitat productivity assessment and weather data acquisition ............. 19 3.3.2. Estimation of red deer population density ............................................. 19 3.3.3. Collection of red deer samples ............................................................... 21 3.3.4. Determination of the body condition and the age of hinds .................... 23 3.3.5. Ascertainment of fertility and sex of foetuses ......................................... 24 3.4. Statistical analysis .............................................................................................. 25 4. RESULTS ........................................................................................................................ 29 4.1. Habitat productivity and red deer population density ......................................... 29 4.2. Body condition of hinds ...................................................................................... 31 4.3. Pregnancy and lactation rates in relation to female age, body mass and marrow fat .................................................................................................... 38 4.4. Factors explaining the variability in red deer female fertility ............................ 43 4.5. Sex ratio of red deer foetuses .............................................................................. 50 4.6. Life tables of the deer population ....................................................................... 51 5. DISCUSSION ................................................................................................................. 58 5.1. Recapitulation of the results ............................................................................... 58 5.2. Reproductive parameters of red deer females in NE Poland compared to other European populations ............................................................................ 60 5.3. Density-dependence in red deer fertility rates .................................................... 63 5.4. Other factors affecting the probability of female fertility ................................... 65 5.5. Body condition of mothers and the sex ratio of foetuses .................................... 69 5.6. Red deer population dynamics – reproductive input under the existing mortality level ..................................................................................................... 70 6. CONCLUSIONS ............................................................................................................. 73 7. REFERENCES ................................................................................................................ 75 APPENDIX 1. Red deer population density in the studied forests ..................................... 84 APPENDIX 2. Body condition indices in relation to the age of red deer females – results of statistical testing ..................................................................... 85 APPENDIX 3. Reproduction parameters in relation to the age of hinds – results of statistical testing ..................................................................... 88 APPENDIX 4. Factors affecting the red deer female fertility – confidence sets of models (GLMs) .......................................................
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