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Anatomy, Palynology and Floral Diversity of the Genus Lamium L

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Anatomy, Palynology and Floral Diversity of the Genus Lamium L ANATOMY, PALYNOLOGY AND FLORAL DIVERSITY OF THE GENUS LAMIUM L. (LAMIACEAE) IN TURKEY A THESIS SUBMITTED TO THE GRADUATE SCHOOL OF NATURAL AND APPLIED SCIENCES OF MIDDLE EAST TECHNICAL UNIVERSITY BY ZEYNEP ATALAY IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN BIOLOGY JANUARY 2016 ii Approval of the thesis: ANATOMY, PALYNOLOGY AND FLORAL DIVERSITY OF THE GENUS LAMIUM L. (LAMIACEAE) IN TURKEY submitted by ZEYNEP ATALAY in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the Department of Biological Sciences, Middle East Technical University by, Prof. Dr. M. Gülbin Düral Ünver ___________________ Dean, Graduate School of Natural and Applied Sciences Prof. Dr. Orhan Adalı ___________________ Head of Department, Biology, METU Prof. Dr. Musa Doğan ___________________ Supervisor, Dept. of Biology, METU Examining Committee Members: Prof. Dr. Zeki Kaya ___________________ Dept. of Biological Sciences, METU Prof. Dr. Musa Doğan ___________________ Dept. of Biological Sciences, METU Prof. Dr. Osman Ketenoğlu ___________________ Dept. of Biology, Ankara University Assoc. Prof. Ferhat Celep ___________________ Dept. of Biology, Gazi University Assoc. Prof. Nursen Coruh ___________________ Dept. of Chemistry, METU_ D Date.______ Date: __________________ iii I hereby declare that all information in this document has been obtained and presented in accordance with academic rules and ethical conduct. I also declare that, as required by these rules and conduct, I have fully cited and referenced all material and results that are not original to this work. Name, Last Name : Zeynep ATALAY Signature :______.________ iv ABSTRACT ANATOMY, PALYNOLOGY AND FLORAL DIVERSITY OF THE GENUS LAMIUM L. (LAMIACEAE) IN TURKEY Atalay, Zeynep Ph. D., Department of Biological Sciences Supervisor: Prof. Dr. Musa Doğan January 2016, 208 pages Lamium L. is the type genus of subfamily Lamioideae and the family Lamiaceae. Due to the absence of recognizable morphological characters, circumscription of the genus has undergone many changes through time. Turkey is the main biodiversity center of the genus with 36 taxa, 18 of which are endemic. Even though many genera in the subfamiliy Lamioideae have been subjected to various anatomical and micromorphological studies, a comprehensive investigation of the genus Lamium is still lacking. In the first part of this study, the anatomical (root, stem, leaf and petiole) and micromorphological (trichome and pollen morphology) characteristics of the Lamium species in Turkey are presented and their taxonomic implications are discussed. Together with gross morphology, such anatomical and micromorphological characters are proved to be taxonomically informative in the delimitation of certain infrageneric taxa. v In the second part of this study, pollination biology and floral diversity of bee pollinated Lamium species; including Lamium villosifolium and L. album subsp. crinitum, which grow sympatrically in Isik Mountain, Kizilcahamam, were investigated. Two Lamium species are clearly adapted to pollination by Apiodea. Field and laboratory investigations revealed that co-occuring Lamium species are isolated from each other on the basis of morphological, mechanical and altitudinal aspects. The diversity of floral constructions increases the mechanical isolation by attracting different pollinators and by loading pollen on different parts of pollinator`s bodies. Change in the plant population density along the altitudinal gradient clearly triggered the pollinator assemblage variation. Keywords: Anatomy, Palynology, Micromorphology, Lamium, Pollination, Systematics vi ÖZ TÜRKİYE’DE BULUNAN LAMIUM L. (LAMIACEAE) CİNSİNİN ANATOMİ, POLEN VE ÇİÇEK ÇEŞİTLİLİGİ ÜZERİNE ÇALIŞMALAR Atalay, Zeynep Doktora, Biyoloji Bölümü Tez Yöneticisi: Prof. Dr. Musa Doğan Ocak 2016, 208 sayfa Lamium L. cinsi Lamiaceae familyasının ve Lamioideae alt familyasının tip cinsidir. Ayırt edici karakterlerin azlığı sebebi ile, cinsin sınırları zaman içinde pek çok kez değişikliğe uğramıştır. Türkiye`de, 16 tanesi endemik olmak üzere, 36 takson yayılış göstermektedir ve ülkemiz cinsin biyoçeşitlilik merkezidir. Altfamilya Lamioideae içindeki pek çok cins üzerinde anatomik ve mikromorfolojik çalışmalar yapılmış olsa da, Lamium cinsi üzerinde bu tip çalışmalar oldukça azdır. Bu çalışmanın ilk kısmında, Türkiye`deki Lamium türlerinin anatomik (kök, gövde, yaprak ve petiol anatomisi) ve mikromorfolojik (tüy ve polen morfolojisi) özellikleri verilmiş, bu karakterlerin taksonomik önemi tartışılmıştır. Bütüncül morfolojik özellikler ile beraber, anatomik ve mikromorfolojik karakterlerin bazı tür ve tür altı kategorilerde taksonomik öneminin olduğu belirlenmiştir. Bu çalışmanın ikinci kısmında, Kızılcahamam Işık Dağı alanında yayılış gösteren iki Lamium; vii L. villosifolium ve L. album subsp. crinitum türlerinin tozlaşma biyolojisi ve çiçek çeşitliliği üzerinde çalışmalar yapılmıştır. Bu iki Lamium türü, belirgin bir şekilde arılarla tozlaşmaya adapte olmuştur. Arazi ve laboratuvar calışmaları ışığında iki Lamium türü birbirlerinden morfolojik, mekanik ve yükseklik özellikleri açısından ayrılmaktadır. Çiçek morfolojilerindeki farklılık, mekanik izolasyonu arttırmakta, polenlerin tozlaştırıcı vücütlarının farklı bölgelerinde taşınmasını sağlamaktadır. Yüksekliğe bağlı olarak değişim gösteren bitki popülasyon yoğunluğu, tozlaştırıcı birliğinde farklılıklara sebep olmaktadır. Anahtar kelimeler: Anatomi, Palinoloji, Mikromorfoloji, Lamium, Polinasyon, Sistematik viii To my parents ix ACKNOWLEDGEMENTS I would like to express my deepest appreciation to my supervisor Prof. Dr. Musa Doğan for his guidance, encouragement, constructive criticism and insight throughout this research. I would like to thank my thesis examining comittee Prof. Dr. Zeki Kaya, Prof. Dr. Osman Ketenoğlu, Assoc. Prof. Dr. Ferhat Celep, and Assoc. Prof. Dr. Nursen Coruh for their suggestions and constructive criticism. I would like to thank Assoc. Prof. Dr. Ferhat Celep for his help and insights in my studies. I would like to thank Assist. Prof. Dr. Fatih Dikmen for identification and measurement of the bees and Assoc. Prof. Dr. Hakan Mete Doğan for providing distribution maps. I would like to thank the Scientific and Technical Research Council of Turkey (TUBITAK/112T-113) and the Scientific Research Projects of Middle East Technical University (BAP/07-02-2014-007-299) for their financial assistance. I would like to thank my lab-mates Deniz Tiambeng, Jelena Erdal and Tural Cavadzade for their help and suggestions. My dear friends; Tuğce, Isil, Begum, Eren, Cansu and Selen. The fun times we had together gave me strength through this tough period. My dearest friends from METU; Gozde, Semra and Mustafa, you are always in my heart. My dear friend Aybars Demiralp, thank you for sharing your beautiful and peaceful thoughts with me. The final credit goes to my family members, especially my mother and father who support me endlessly. x TABLE OF CONTENTS ABSTRACT…………………………………………………………………………V ÖZ……………………………………………………………………….…………VII TABLE OF CONTENTS…………………………………………..….……………XI LIST OF TABLES……………………………………………………..…………..XV LIST OF FIGURES……………………………………………….…..………….XVII ABBREVIATIONS………………………………………………..……………..XXII CHAPTERS 1. INTRODUCTION………………………………………………….……………..1 1.1 Brief History of the Lamiaceae……………………………..………..…..1 1.2 General Characteristics of Lamium species…………………….…….......4 1.3 Taxonomic History of the genus Lamium..................................................6 1.4 Anatomical and Micromorphological Studies on the Lamiaceae…….....12 1.4.1 Anatomy…………………………………………….…….…..12 1.4.2 Palynology…………………………………………….…..…..14 1.4.3 Trichome Morphology………………………………...……....15 1.5 Phytogeographic Distribution and Endemism Ratio of Lamium Species in Turkey……………………….……………………………………………....15 1.6 Historical Background of Pollination Biology………………………….16 1.7 Why Study Pollination Biology…………………………………...…….20 xi 1.8 Pollination Syndromes………………………………………...………...22 1.9 Evolutionary Patterns in the Lamiaceae……………………….………..25 1.10 Objectives of the Study………………………………………….…….28 2. MATERIAL AND METHODS……………………………………………….....31 2.1 Plant Material………………………………………………..………….31 2.2 Anatomical Investigations…………………………………...………….31 2.3 Palynological Investigations …………………………………...……….32 2.4 Trichome Micromorphology Investigations……………….……………33 2.5 Distribution and Ecological Data of the Lamium species………….……33 2.6 Pollination Biology Investigations…………………………….…..……33 2.6.1 Lamium species…………………………………………..……34 2.6.2 Study Sites……………………………………………….……35 2.6.3 Morphometric Measurements…………………………………35 2.6.4 Pollinator Observations…………………………………….....37 2.6.5 Statistics……………………………………………….………38 3. RESULTS AND DISCUSSION….…………………………………….……..….39 3.1 Final Taxonomic Treatment Results……………………………….……39 3.2 Ecology, Endemism and Phytogeography of the Lamium Species……..44 3.2.1 Endemism and Phytogeography……………………………....52 3.2.2 Phenology and Altitudinal Range of the Lamium Species…....54 3.2.3 Geographical Distribution of the Lamium Species………....…59 3.3 Anatomical Features of the Genus Lamium……………….……………64 3.3.1 Root Anatomy………………………………………….……..64 xii 3.3.2 Stem Anatomy………………………………………….……..66 3.3.3 Leaf Anatomy…………………………………………….…...68 3.3.4 Petiole Anatomy…………………………..……………….….70 3.3.5 Systematic Implications of Vegetative Anatomy……………..72 3.4 Trichome Micromorphology…………………………………...……….91 3.4.1 Nonglandular Trichomes..........................................................91 3.4.2 Glandular
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