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Lamiaceae) in East and Southeast Anatolia, Turkey MORPHOLOGY, ANATOMY AND SYSTEMATICS OF THE GENUS SALVIA L. (LAMIACEAE) IN EAST AND SOUTHEAST ANATOLIA, TURKEY A THESIS SUBMITTED TO THE GRADUATE SCHOOL OF NATURAL AND APPLIED SCIENCES OF MIDDLE EAST TECHNICAL UNIVERSITY BY AHMET KAHRAMAN IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN BIOLOGY JULY 2011 Approval of the thesis MORPHOLOGY, ANATOMY AND SYSTEMATICS OF THE GENUS SALVIA L. (LAMIACEAE) IN EAST AND SOUTHEAST ANATOLIA, TURKEY submitted by AHMET KAHRAMAN in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Department of Biological Sciences, Middle East Technical University by, Prof. Dr. Canan Özgen ___________________ Dean, Graduate School of Natural and Applied Sciences Prof. Dr. Musa Doğan ___________________ Head of Department, Biology, METU Prof. Dr. Musa Doğan ___________________ Supervisor, Dept. of Biology, METU Examining Committee Members: Prof. Dr. N. Münevver Pınar ___________________ Dept. of Biology, Ankara University Prof. Dr. Musa Doğan ___________________ Dept. of Biology, METU Assoc. Prof. Dr. Sertaç Önde ___________________ Dept. of Biological Sciences, METU Assoc. Prof. Dr. C. Can BĠLGĠN ___________________ Dept. of Biology, METU Assist. Prof. Dr. H. Nurhan Büyükkartal ___________________ Dept. of Biology, Ankara University Date: 11.07.2011 ii 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: AHMET KAHRAMAN Signature : iii ABSTRACT MORPHOLOGY, ANATOMY AND SYSTEMATICS OF THE GENUS SALVIA L. (LAMIACEAE) IN EAST AND SOUTHEAST ANATOLIA, TURKEY Kahraman, Ahmet Ph.D., Department of Biological Sciences Supervisor: Prof. Dr. Musa Doğan July 2011, 336 pages This study aims to revise the genus Salvia L. in East and Southeast Anatolia in Turkey on the basis of macromorphological, anatomical, palynological, mericarp micro-morphological, ecological and numerical analysis. Towards achieving this goal, about 2500 specimens of Salvia were collected and examined during extensive field studies between July 2005 and June 2009. The materials collected by other researchers either from Turkey or abroad were also investigated. The results of the taxonomic revision show that the study area includes 59 taxa, 24 (40.7%) of which are endemic and the remaining 35 (59.3%) are non-endemic. Salvia siiirtica is described as new to science. S. macrosiphon is described as a new record for Turkey and S. cerino-pruinosa and S. pseudeuphratica are re-evaluated as valid species as well as S. ballsiana is rediscovered. Macromorphological characters that were considered to have taxonomically diagnostic value are investigated and their possible variations are discussed. Habit, iv stem, leaf, inflorescence, bract, calyx, corolla and stamen properties are compared at infrageneric and species level. The first comprehensive evaluation of the systematic value of anatomy, palynology and mericarp micromorphology of Salvia are presented. Some characters, such as row numbers of ray cells in the root, type of the leaf blade structure, shape of cross section of the petiole, shape and number of median vascular bundles, pollen size, size of mericarps and diameter of abscission scars, are useful for infrageneric delimitation. Variation in some anatomical characters, such as number of cell layers of collenchyma and cortex, number of cell layers of palisade parenchyma, size of petiole, number of lateral vascular bundles, exine ornamentation type of pollen, presence of large lumina in the middle of the primary lumen, the muri thickness, mericarp shape, mericarp length/width ratio and surface ornamentation type, can be used for separating species within the sections. Based on the updated geographical and ecological distribution, conservation status of the taxa is reassessed at regional, national and global levels. At the regional scale, the distribution is CR for 6 taxa EN for 9 taxa, VU for 5 taxa, NT for 10 taxa and LC for 29 taxa. At the national scale, the distribution is CR for 5 taxa, EN for 5 taxa, VU for 7 taxa, NT for 10 taxa and LC for 32 taxa. At the global scale, the distribution is CR for 5 taxa, EN for 3 taxa, VU for 6 taxa, NT for 10 taxa NT and LC for 35 taxa. The main threats in the study area are overgrazing, constructions, land clearing, fire, urbanization and tourism. The infrageneric delimitation is performed using multivariate analysis. Identification keys to sections and species are given. Synonymy, updated descriptions, phenology, distribution and habitats in Turkey, general distribution outside Turkey, distribution maps, phytogeography, specimen citations, some notes on taxonomy and photographs showing general appearance of the taxa are also provided. Keywords: Lamiaceae, Salvia, systematics, morphology, anatomy, palynology, East and Southeast Anatolia, Turkey. v ÖZ TÜRKĠYE’NĠN DOĞU VE GÜNEY DOĞU ANADOLU BÖLGELERĠ’NDE BULUNAN SALVIA L. (LAMIACEAE) CĠNSĠNĠN MORFOLOJĠSĠ, ANATOMĠSĠ VE SĠSTEMATĠĞĠ Kahraman, Ahmet Doktora, Biyolojik Bilimler Bölümü Tez Yöneticisi: Prof. Dr. Musa Doğan Temmuz 2011, 336 sayfa Bu çalışma makromorfolojik, anatomik, palinolojik, merikarp mikromorfolijisi, ekolojik ve numerik analizler kullanılarak Türkiye’nin Doğu ve Güneydoğu bölgerinde bulunan Salvia cinsinin revizyonunu gerçekleştirmeyi amaçlar. Bunun için, Salvia taksonlarının yaklaşık 2500 örneği Temmuz 2005 ve Haziran 2009 arasındaki yoğun arazi çalışmaları boyunca toplanmış ve incelenmiştir. Ayrıca, yerli ve yabancı araştırmacılar tarafından toplanmış materyaller de incelenmiştir. Taksonomik revizyonun sonuçları çalışma alanında 59 taksonun bulunduğunu, bunların 24 (40.7%)’ünün endemik geriye kalan 35 (59.3%) taksonun ise endemik olmadığını gösterir. Salvia siirtica bilim dünyası için yeni bir tür olarak tanımlanmıştır. S. macrosiphon Türkiye için yeni kayıt olarak bulunmuş, S. cerino- pruinosa ve S. pseudeuphratica yeniden tür olarak değerlendirilmiş ayrıca S. ballsiana yeniden keşfedilmiştir. vi Taksonomik olarak ayırt edici değeri olduğu düşünülen makromorfolojik karakterler araştırılmış ve bunların olası varyasyonları tartışılmıştır. Habit, gövde, yaprak, çiçek durumu, brakte, çanak yaprak, taç yaprak ve stamen özellikleri cins içi ve tür seviyesinde karşılaştırılmıştır. Salvia’nın anatomisi, palinolojisi ve merikarp mikromorfolojisinin sistematik önemi üzerine ilk kapsamlı değerlendirme sunulmuştur. Bazı karakterler cins içi sınıflandırmada faydalı bulunmuştur örneğin; kökteki ışın hücrelerinin sıra sayısı, yaprak ayası tipi, yaprak sapının enine kesitinin şekli, ortadaki iletim demetlerinin şekli ve sayısı, polen büyüklüğü, merikarpın büyüklüğü ve hilumun çapı. Bazı anatomik karekterlerdeki varyasyon türlerin ayırt edilmesinde kullanılabilir; korteks ve kollenkima hüclerinin tabaka sayısı, palizat parenkima hücrelerinin tabaka sayısı, yaprak sapının büyüklüğü, yandaki iletim demetlerinin sayısı, polen eksin ornamentasyonunun tipi, primer luminanın ortasında geniş sekonder luminanın varlığı, merikarp şekli, uzunluk/genişlik oranı ve yüzey ornamentasyon tipi gibi. Güncellenen coğrafik ve ekolojik dağılımlar dikkate alınarak, taksonların koruma statüleri bölgesel, ulusal ve uluslararası ölçekte tekrar değerlendirilmiştir. Bölgesel ölçekte 6 takson CR, 9 takson EN, 5 takson VU, 10 takson NT ve 29 takson LC; ulusal ölçekte 5 takson CR, 5 takson EN, 7 takson VU, 10 takson NT ve 32 takson LC; uluslararası ölçekte 5 takson CR, 3 takson EN, 6 takson VU, 10 takson NT ve 35 takson LC kategorisindedir. Çalışma alanındaki en önemli tehtitler; aşırı otlatma, inşaat çalışmaları, arazi açma, yangınlar, kentleşme ve turizmdir. Cins içi sınıflandırma çoklu varyasyon analizi yardımı ile yapılmıştır. Seksiyon ve tür anahtarları verilmiştir. Ayrıca, taksonların sinonimi, tip örnekleri, güncellenmiş betimleri, fenolojisi, Türkiye’deki dağılımları ve habitatları, Türkiye dışındaki dağılımları, dağılım haritaları, fitocoğrafyası, incelenen örnekler, taksonomisi üzerine bazı notlar ve genel görünümlerini gösteren fotoğraflar verilmiştir. Anahtar Kelimeler: Lamiaceae, Salvia, sistematik, morfoloji, anatomi, palinoloji, Doğu ve Güneydoğu Anadolu, Türkiye. vii To my wife and family viii ACKNOWLEDGMENTS I would like to express my deepest gratitude to my supervisor Prof. Dr. Musa Doğan for his valuable guidance, advice, criticism, encouragement and insight throughout the research. I would like to thank to the examining committee members of my thesis Prof. Dr. N. Münevver Pınar, Assoc. Prof. Dr. C. Can Bilgin, Assoc. Prof. Dr. Sertaç Önde and Assist. Prof. Dr. H. Nurhan Büyükkartal for their valuable suggestions and constructive criticsm to the thesis. I am also very grateful to Prof. Dr. Sevil Pehlivan, Assoc. Prof. Dr. Esra Martin, Assoc. Prof. Dr. Galip Akaydın, Dr. Greg G. Guerin, Assist. Prof. Dr. Hülya Özler, Assist. Prof. Dr. Hatice Çölgeçen, Assist. Prof. Dr. Hakan Mete Doğan and Assist. Prof. Dr. Birol Başer for their help and collaboration. I wish to thank my wife Hilal Kahraman and parents for their patience, encouragement and endless moral support. I give my special thanks to my father Süleyman Kahraman for his valuable assistance and support in field studies. I am grateful for the help of the members of the Scanning Electron Microscope (SEM) Laboratory of Department of Metallurgical and Materials Engineering in METU. I also thank to
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