MORPHOLOGY, ANATOMY AND SYSTEMATICS OF THE GENUS LATHYRUS L. (LEGUMINOSAE) IN CENTRAL ANATOLIA, TURKEY A THESIS SUBMITTED TO THE GRADUATE SCHOOL OF NATURAL AND APPLIED SCIENCES OF MIDDLE EAST TECHNICAL UNIVERSITY BY HÜSEYİN ÇİLDİR IN PARTIAL FULFILLMENT OF THE REQUIRMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPY IN BIOLOGY JULY 2011 Approval of the thesis: MORPHOLOGY, ANATOMY AND SYSTEMATICS OF THE GENUS LATHYRUS L.(LEGUMINOSAE) IN CENTRAL ANATOLIA, TURKEY submitted by HÜSEYİN ÇİLDİR in partial fulfillment of the requirements for the degree of Doctor of Philosophy of Science in Biology Department, Middle East Technical University by, Prof. Dr. Canan Özgen ____________ Dean, Graduate School of Natural and Applied Sciences Prof. Dr. Musa Doğan ____________ Head of Department, Biological Sci. Dept., METU Prof. Dr. Musa Doğan ____________ Supervisor, Biological Sci. Dept., METU Examining Committee Members: Prof. Dr. Osman Ketenoğlu ____________ Biology Dept., Ankara University Prof. Dr. Musa Doğan ____________ Biological Science Dept., METU Prof Dr. Sevil Pehlivan ____________ Biology Dept., Gazi University Assoc. Prof. Dr. Sertaç Önde ____________ Biological Science Dept., METU Assoc. Prof. Dr. Galip Akaydın ____________ Biology Dept., Hacettepe University Date: 08-07-2011 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 wok. Name, Last name: Hüseyin ÇİLDİR Signature: _____________ iii ABSRACT MORPHOLOGY, ANATOMY AND SYSTEMATICS OF THE GENUS LATHYRUS L.(LEGUMINOSAE) IN CENTRAL ANATOLIA, TURKEY Çildir, Hüseyin Ph.D., Department of Biology Supervisor: Prof. Dr. Musa Doğan July 2011, 174 pages In this study, morphology, anatomy and systematics of the Genus Lathyrus L. (Leguminosae) in Central Anatolia is presented. Comparative morphological characters and their variation in the Genus; Calyx, leaf, corolla and pollen grains micro-morphology of the species; Anatomy of the species; Ecology, endemism, phytogeography and IUCN threat categories of the species; Numerical analysis and Revision of the genus in Central Anatolia were conducted. For the first time the calyx, corolla and leaf micromorphology, and the anatomical characteristics of Lathyrus were examined. Infrageneric delimitation of the species is performed by using multivariate analysis. As a result of morphological and anatomical data, it is suggested that L. haussknechtii should be classified as different species not variety of L. brachypterus. iv The collected specimens was crosschecked with neighboring floras and the existing type specimens known from Turkey ANK, GAZİ, HUB, KNYA, CUFH, Erciyes, and Royal Botanic Garden Edinburgh (RBGE) herbaria. Morphological and micromorphological characters of pollen grains were revealed. It was determined that the pollen grains size and sculpturing were important diagnostic characters for the species. The statistical analysis was applied to compare P/E ratios of the pollen grains of studied taxa. According to this analysis, the P/E ratio is important diagnostic feature for most of the sections and the species. Sectional key and species key were prepared and updated. Expanded descriptions, GPS data, some photographs and some notes on the taxonomy of the species were given. Key words: Leguminosae, Lathyrus L., anatomy, morphology, systematics, Central Anatolia geographic region, Turkey. v ÖZ İÇ ANADOLU BÖLGESİ’NDEKİ LATHYRUS L. (LEGUMINOSAE) CİNSİNİN MORFOLOJİSİ, ANATOMİSİ VE SİSTEMATİĞİ Çildir, Hüseyin Doktora, Biyolojik Bilimler Bölümü Tez Yöneticisi: Prof. Dr. Musa Doğan July 2011, 174 sayfa Bu çalışmada, İç Anadolu bölgesinde yayılış gösteren Lathyrus L. (Leguminosae) cinsinin morfolojisi, anatomisi ve sistematiği sunulmaktadır. Karşılaştırmalı morfolojik karakterler ve bu karakterlerin cins içi varyasyonları belirlenmiş; türlerin kaliks, yaprak, korolla ve pollen mikromorfolojileri ile kök, gövde ve yaprak anatomisi özellikleri incelenmiş ; türlerin ekoloji, endemism, fitocoğrafya ve IUCN tehlike kategorileri belirlenmiş; cinsin numeric analizi ve revizyonu yapılmıştır. Bu çalışma ile Türkiye’de ilk kez Lathyrus cinsini oluşturan tülerin kaliks, korolla ve yaprak mikromorfolojisi ile anatomik özellikleri incelenmiştir. Türlerin Infrageneric sınırlandırılması çoklu varyasyon (multivariety) analiziyle yapılmıştır. Morfolojik ve anatomik veriler sonucunda L. brachypterus türünün bir varyetesi olarak sınıflandırılan L. haussknechtii’ nin ayrı bir tür olarak sınıflandırılması gerektiği önerilmiştir. vi Araziden toplanan örneklerin teşhisinde Türkiye Florası ve komşu ülkelerin floraları kullanılmıştır. Bunun yanı sıra toplanan örneklerin çapraz kontrolleri Ankara Üniversitesi herbaryumu, Gazi Üniversitesi herbaryumu, Hacettepe Üniversietsi herbaryumu, Selçuk Üniversitesi herbaryumu, Cumhuriyet üniversitesi herbaryumu, Erciyes Üniversitesi herbaryumu ve Kraliyet Botanik Bahçesi Edinburg (RBGE) herbaryumundaki mevcut örneklerle yapılmıştır. Polenlerin morfolojik ve mikromorfolojik karakterleri belirlenmesi ile polen boyutu ve yüzey yapılarının türlerin teşhisinde önemli olduğu saptanmıştır. Taksonların polenlerinin P/E oranlarında anlamlı bir farlılık olup olmadığını araştırmak için istatiksel analiz yapılmıştır. Analiz sonucunda P/E oranının türlerin teşhisinde önemli olduğu sonucuna varılmıştır. Seksiyon ve tür anahtarları hazrılanmış ve güncellenmiştir. Seksiyon ve türlerin genişletilmiş açıklamaları, GPS verileri, bazı fotoğraflar ve türlerin taksonomisi üzeirne bazı notlar verilmiştir. Anahtar kelimeler: Leguminosae, Lathyrus L., anatomi, morfoloji, sistematik, İç Anadolu Bölgesi, Türkiye. vii To my family, my wife and son. viii ACKNOWLEDGEMENT I would like to express my deepest gratitude to my supervisor Prof. Dr. Musa Doğan for his guidance, advice, criticism, encouragement and insight throughout the research. I would like to thank to members of my thesis examining committee for their suggestions and constructive critisism. I would like to thank Dr. Ferhat Celep guidances and advices. I would like to thank Dr. Ahmet Kahraman for his help about anatomical studies. I would like to thank Okan Kaan Binzat for his advice, encouragement, patience and friendship. I also would like to thank Assoc. Prof. Dr Fatma Güneş for sharing some of her specimens. I would like to thank Curators of ANK, GAZİ, HUB, KNYA, CUFH, Erciyes, RBGE for allowing me to study their Lathyrus specimens. I would like to thank my wife, and family for their patience, encouragement and support. ix TABLE OF CONTENTS ABSRACT……………………………………………………………………. iv ÖZ…………………………………………………………………………….. vi DEDICATION……………………………………………………………….. viii ACKNOWLEDGEMENTS…………………………………………………... ix TABLE OF CONTENTS……………………………………………………... x LIST OF TABLES……………………………………………………………. xii LIST OF FIGURES…………………………………………………………… xiv ABBREVIATIONS…………………………………………………………… xix CHAPTERS 1. INTRODUCTION………………………………………………………….. 1 1.1 Historical Background of the Family Leguminosae…………………… 1 1.2 The Genus Lathyrus L. in the World…………………………………... 5 1.3 The Genus Lathyrus L. in Turkey……………………………………… 9 1.4 General Description of the Study Area………………………………… 11 1.4.1 Location of the Study Area………………………………………. 11 1.4.2 Climate of the Study Area………………………………………... 13 1.4.3. Topography and Vegetation of the Study Area…………………. 14 1.5. Objectives and Scope of the study…………………………………….. 15 2. MATERIAL AND METHODS……………………………………………. 16 2.1 Collection of the Plant Materials………………………………………. 16 2.2 Morphological Studies of Vegetative and Reproductive Structures…… 17 2.3 Anatomical Method……………………………………………………. 17 2.4 Micromorphological and Statistical Method………………………….. 17 2.5. Multivariate Analysis…………………………………………………. 19 2.6 Assesment of Geographical Distribution, Endemism, Phytogeography and IUCN Threat Categories of the Species……………………………….. 19 xix 3. RESULTS………………………………………………………………….. 21 3.1 Vegetative and Reproductive Morphological Characters and Their Variation…………………………………………………………………… 21 3.1.1. Habit…………………………………………………………….. 21 3.1.2 Stem Morphology………………………………………………... 23 3.1.3. Leaf Morphology………………………………………………... 23 3.1.4 Inflorescence Morphology………………………………………. 25 3.1.5 Flowers…………………………………………………………... 27 3.1.6. Fruit……………………………………………………………... 30 3.2 Calyx, Leaf, Corolla and Pollen grains Micromorphology……………. 31 3.2.1 Calyx, Leaf and Corolla Micromorphology……………………... 31 3.2.1 Pollen grains Micromorphology…………………………………. 38 3.3. Anatomy………………………………………………………………. 73 3.3.1 Root Anatomy…………………………………………………… 75 3.2.2. Stem Anatomy…………………………………………………... 80 3.2.3. Leaf Anatomy…………………………………………………… 88 3.4. Ecology, Endemism and Phytogeography of the Genus Lathyrus …… 94 3.4.1. Habitat, Phenology and Altitude Disribution of the Species…… 94 3.4.3. Endemism, Phytogeography and IUCN Threat Categories of the Species…………………………………………………………………. 97 3.5. Numerical Analysis…………………………………………………… 105 3.6. Taxonomic Treatment………………………………………………… 110 3.6.1. The Genus Lathyrus L…………………………………………... 110 3.6.2. Sectional key……………………………………………………. 110 3.6.3. Key to Species…………………………………………………... 111 3.6.4. Species descriptions…………………………………………….. 114 4. CONCLUSION…………………………………………………………… 151 REFERENCES……………………………………………………………….. 156 CURRICULUM VITAE……………………………………………………... 174 xi LIST OF TABLES Table 1. Leguminosae subfamilies and their number of genera and species…. 2 Table 2. Characteristics of Leguminosae……………………………………..