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

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

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 :

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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.

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

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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 my lab-mates Ferhat Celep, Safi Bagherpour, Hüseyin Çildir, Okan Binzat and Evren Cabi.

I would like to thank the curators of the herbaria, AEF, ANK, B, BM, CBB, E, FUH, G, GAZI, HUB, ISTE, ISTF, K, KNYA, LE, MO, VANF and W for allowing us to ix

study their Salvia collections, and finally the Scientific and Technical Research Council of Turkey (TUBĠTAK-TBAG 104 T 450) for their financial support for the research.

This study was supported by Faculty Develeopment Program (ÖYP) of Middle East Technical University financed by DPT.

TABLE OF CONTENTS

ABSTRACT……………………………………………………………………….…iv ÖZ…………………………………………………………………………...... ……..vi ACKNOWLEDGMENT……………………………………………………...... …....ix TABLE OF CONTENTS…………………………………………………...... ……...xi LIST OF TABLES……………………………………………………...…………..xiv LIST OF FIGURES…………………………………………………..….…………xvi LIST OF ABBREVIATIONS………………………………………..…..……...... xxv CHAPTERS 1. INTRODUCTION……………………………………………………………...….1 1.1. Historical Background of the Family Lamiaceae…………………………...... 1 1.2. Historical Background of the Genus Salvia L………………………..………7 1.3. General Description of the Study Area………………..……...………..……14 1.3.1. Location……………………………….…..………………………….14 1.3.2. Topographic Characteristics………………………………..……...…17 1.3.3. Climatic Characteristics……………………………...……………….18 1.3.4. Soil Characteristics……………………….………..…………………19 1.3.5. Objectives of the study…………………………...…………..………20 2. MATERIALS AND METHODS……………………………………...…...…….21 2.1. Field Surveys and Plant Material Collection…………………..…………….21 2.2. Methods used in Systematics………………..……………………….……...21 2.3. Methods used in Macromorphological Studies of Vegetative and Reproductive Structures……………………………………………………..22 2.4. Methods used in Anatomical Studies………………………..……………...23 2.5. Methods used in Palynological Studies……………………….………….....23 2.6. Methods used in Mericarp Micromorphological Studies………………...... 24 2.7. Assessment of the Threat Categories………………...……………………..24 2.8. Ecological Analysis……………………..…………………………..….…..25 2.9. Numerical Taxonomic Methods…………………..…………..……………25 xi

3. RESULTS AND DISCUSSION…..………..…………………………...……….27 3.1. Gross Morphology………..……………..…………………………………..27 3.1.1. Life Form and Growth Habit…………….………………..………….27 3.1.2. Stem……………………………………………….………………….27 3.1.3. Leaves…………...……………………………………………………29 3.1.4. Inflorescence…………...………………………………………….….31 3.1.5. Bracts……………………………...……...…………………………..32 3.1.6. Calyx………………………...………………………………………..33 3.1.7. Corolla……………………..………………...…………………..……33 3.1.8. Stamens………………...……………………………………………..35 3.1.9. Style…………………………………..……………………………….36 3.2. Anatomy of the Vegetative Organs………………………...……………....37 3.2.1. Root Anatomy……………………….…...………………………….37 3.2.2. Stem Anatomy………………………….…..…………….…………45 3.2.3. Leaf Blade Anatomy………………...…………..………………..…51 3.2.4. Petiole Anatomy…………………………..….…………………..…62 3.2.5. Infrageneric Anatomical Descriptions of Salvia…………………….73 3.2.6. Systematic Implications of Anatomical Characteristics……….……82 3.3. Pollen Micromorphology……………...…………………………………...84 3.3.1. Size and Shape…………………………...……………….…………84 3.3.2. Apertures…………………………….………..………………….…85 3.3.3. Exine Sculpturing……………………….………..………………....88 3.3.4. Infrageneric Pollen Descriptions of Salvia……………….……..…100 3.3.5. Systematic Implications of Pollen Characteristics…………………102 3.4. Mericarp Micromorphology…..…………..……………………………....115 3.4.1. Size, Shape and Colour………………………………………...…..115 3.4.2. Abscission Scar Diameter and Shape………………….……..….....116 3.4.3. Surface Sculpturing…………………………………………...……116 3.4.4. Infrageneric Mericarp Descriptions of Salvia…………………...…124 3.4.5. Systematic Implications of Mericarp Characteristics……...……….125 3.5. Ecology and Distribution……………..…………………………………..138 3.5.1. Distribution, Endemism and Phytogeography……………..……... 138 xii

3.5.2. Habitat, Phenology and Altitudinal Range………………...…..…..144 3.5.3. Soil Properties…………………………...…………………………151 3.6. Assessment of Threat Categories of the Salvia Taxa……………………..156 3.7. Numerical taxonomy of Salvia……………….…………..……..…………166 3.8. Systematics….…………….…………..………………………….……….171 4. CONCLUSION………………..………………………………………………..310 REFERENCES………………..……………………………..…………………….315 CURRICULUM VITAE………………………...…………………………..……..331

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LIST OF TABLES

TABLES Table 1. Genera of the subtribe Salvinaeae sensu Harley et al. (2004) and their previous taxonomic positions. …………...……………...……….…………………..5 Table 2. Subgenera and sections of the genus Salvia classified by Bentham (1832- 1836)………………………………………………………..……..………….………9 Table 3. The number of endemic plant species growing in different phytogeographical regions of Turkey (Ekim et al., 2000)…………………………..15 Table 4. Anatomical characteristics of roots in the Salvia taxa studied. All sizes are in µm. Numbers refer to minimum-maximum (mean)…………………………..….42 Table 5. Anatomical characteristics of stems in the taxa studied of Salvia. All sizes are in µm. Numbers refer to minimum-maximum (mean). SD = Stem diameter, SC = Stem contour, COL = Thickness of collenchyma, CHL = Thickness of the chlorenchyma, COR = Thickness of the cortex, PI = pith region, VBC = Thickness of vascular bundles at the corners, VBB = Thickness of vascular bundles between the corners……………………………….…………………………………..…….…….48 Table 6. Anatomical characteristics of leaf blades in the taxa studied of Salvia. All sizes are in µm. Numbers refer to minimum-maximum (mean). L: Length, W: Width…………………………………………………………….....………………..56 Table 7. Anatomical characteristics of leaf blades in the taxa studied of Salvia. All sizes are in µm. Numbers refer to minimum-maximum (mean)…………………….60 Table 8. Anatomical characteristics of petioles in the taxa studied of Salvia. All sizes are in µm. Numbers refer to minimum-maximum (mean)……………………….…67 Table 9. Pollen characteristics in the taxa studied of Salvia based on LM. All sizes are in µm. Numbers refer to minimum-maximum (mean±standard deviation). Os = Oblate-spheroidal, Ps = Prolate-spheroidal, So = Suboblate, Sp = Subprolate. -, absent, +, rarely present, ++, densely present……………………………………...108 Table 10. Pollen characteristics in the taxa studied of Salvia based on SEM. All sizes are in µm. -, absent, +, present………………………………………………..……112 xiv

Table 11. Quantitative mericarp characteristics of mericarps in the taxa studied of Salvia. All sizes are in mm except for length/with ratio. Numbers refer to minimum- maximum (mean)…………………………………………….………………….…132 Table 12. Qualitative mericarp characteristics in the taxa studied of Salvia. S = Near spherical, Pr = Prolate-spheroidal. -, absent, +, rarely present, ++, densely present…………………………………………….………………….…………….135 Table 13. The taxa distributed in the study area and their distribution data. Details of the taxa known only within the area, endemism, phtogeographic regions, phenology and altitudinal range of the taxa are also provided. *: taxa known only within the study area in Turkey. Euro-Sib.: Euro-Siberian element, Ir.-Tur.: Irano-Turanian element, Medit.: Mediterranean element, Unk. or Multi.: Unknown or Multiregional. The taxa are listed in alphabetical order…………………………………………...140 Table 14. Physical and chemical properties of soil samples taken from the natural habitat of the taxa……………………………………………………………...... …153 Table 15. The recent IUCN threat categories with criteria and an estimated proportion (%) of the global population of the taxa distributed in the research area. Column 1: Taxa, Column 2: Threat categories according to Turkish Red Data Book (Ekim et al., 2000), Column 3: Recommended threat categories for regional scale, Column 4: Recommended threat categories for national scale, Column 5: Recommended threat categories for global scale, Column 6: An estimate of the proportion (%) of the global population occurring within the region, Column 7: IUCN Red List Criteria (2001). The taxa are listed in alphabetical order. +: the taxa endemic, *: taxa known only within the study area in Turkey, euro-sib.: euro-siberian element, ir.-tur.: irano-turanian element, medit.: mediterranean element, unk. or multi.: unknown or multiregional. The taxa are listed in alphabetical order………161 Table 16. The main threats to the taxa in the study area. +: the taxa endemic, *: taxa known only within the study area in Turkey. The taxa are listed in alphabetical order………………………………………………………………………………..164 Table 17. List of macromorphological, anatomical, pollen and mericarp micromorphological characters scored for the numerical taxonomic analysis….…166

LIST OF FIGURES

FIGURES Figure 1. Worldwide distribution of species of the genus Salvia. Numbers refer to approximate number of species and circles show present centers of diversity (Walker et al., 2004)……………………..………………………………..………...…………8 Figure 2. Representations of the stamen types defined per clade suggested by the molecular results (Walker & Sytsma, 2007)………..………………………...……..12 Figure 3. Main phytogeographical regions in Turkey………..………………..……16 Figure 4. Location of the research area………………………………...... …...……17 Figure 5. Location of stations visited within the study area between 2005 and 2009……………………………………………………………………………….....22 Figure 6. Stem types in Salvia………………..…………………………..…………28 Figure 7. A. S. indicia, a perennial herbaceous species growing up to 1.5 m high; B. S. hedgeana, a dwarf perennial suffruticose species barely reaching 17 cm high...... 29 Figure 8. Leaf characteristics in Salvia…………………………...…………..……..30 Figure 9. The number of flowers per verticillaster in inflorescences in Salvia…...... 32 Figure 10. Variation in bract, calyx and corolla morphology in Salvia……………..34 Figure 11. Representations of the stamen types in Salvia species in study area (Hedge 1982a). c: connective, db: dolabriform sterile tissue; f: filament…....……...………36 Figure 12. Cross section of the root of S. atropatana. ca: cambium, cor: cortex, pe: periderm, ph: phloem sc: sclerenchyma, x: xylem, xr: xylem rays………...... ….38 Figure 13. Cross section of the root of S. hedgeana………….……………….…….39 Figure 14. Cross section of the root of S. pachystachys…….………………..……..39 Figure 15. Cross section of the root of S. viridis…………………………….…..….40 Figure 16. Cross section of the root of S. ceratophylla……………………….…….40 Figure 17. Cross section of the root of S. dicroantha………………………..…...…41 Figure 18. Cross section of the root of S. verticillata subsp. amasiaca……………..41 Figure 19. Cross section of the stem of S. divaricata. chl: chlorenchyma, col: collenchyma, cor: cortex, ep: epidermis, ph: phloem, pi: pith…………………..…..46 xvi

Figure 20. Cross section of the stem of S. caespitosa……………………...………..46 Figure 21. Cross section of the stem of S. cerino-pruinosa……………………..…..46 Figure 22. Cross section of the stem of S. virgata………………………...………...47 Figure 23. Cross section of the stem of S. blepharochleana…………….…...……...47 Figure 24. Cross section of the stem of S. pachystachys………………………...…..47 Figure 25. Cross section of the leaf blade of S. ballsiana. cu: cuticle, hy: hypodermis, le: lower epidermis, mid: midrib, mv: median vascular bundle, pp: palisade parenchyma, ue: upper epidermis……………..……………………………….…....53 Figure 26. Cross section of the leaf blade of S. indica. ab. abaxial side, ad: adaxial side, le: lower epidermis, mid: midrib, mv: median vascular bundle, pp: palisade parenchyma, sp: spongy parenchyma, ue: upper epidermis, vb: vascular bundle…….……………………………………………………………...…..………53 Figure 27. Cross section of the leaf blade of S. verticillata subsp. verticillata. eg: eglandular hairs, hyp: hypodermis, le: lower epidermis, lv. lateral vascular bundle, mid: midrib, mv: median vascular bundle, pp: palisade parenchyma, sp: spongy parenchyma, ue: upper epidermis……………………..………....………………….54 Figure 28. Cross section of the leaf blade of S. macrochlamys. gh: glandular hairs, hyp: hypodermis, le: lower epidermis, lv. lateral vascular bundle, mv: median vascular bundle, pp: palisade parenchyma, ue: upper epidermis……………………………………………………………..……………...54 Figure 29. Cross section of the leaf blade of S. aethiopi…………..…………..……55 Figure 30. Cross section of the leaf blade of S. microstegia…………..…………….55 Figure 31. Cross section of the petiole of S. pinnata. ab: abaxial side, ad: adaxial side, col: collenchymas, lv: lateral vascular bundle, mv: median vascular bundle……………………………………………………………………………….63 Figure 32. Cross section of the petiole of S. pachystachys…………………….……64 Figure 33. Cross section of the petiole of S. viridis……………………………..…..64 Figure 34. Cross section of the petiole of S. glutinosa……………………………...65 Figure 35. Cross section of the petiole of S. syriaca………………………..……....65 Figure 36. Cross section of the petiole of S. verticillata subsp. verticillata……...…66 Figure 37. Cross section of the petiole of S. cerino-pruinosa………………………66

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Figure 38. LM micrographs of pollen grains in the Salvia taxa examined: a-b. S. anatolica, c-d. S. ballsiana, e-f. S. bracteata, g-h. S. caespitosa, i-j. S. divaricata, k-l. S. hedgeana, m-n. S. huberi, o-p. S. kurdica (a, c, e, g, i, k, m, o: Equatorial view; b, d, f, h, j, l, p: Polar view). Scale bar = 20 µm……………………………………….86 Figure 39. LM micrographs of pollen grains in the Salvia taxa examined: a-b. S. macrochlamys, c-d. S. pachystachys, e-f. S. tomentosa, g-h. S. trichoclada, i-j. S. euphratica var. euphratica, k-l. S. kronenburgii, m-n. S. atropatana, o-p. S. brachyantha (a, c, e, g, i, k, m, o: Equatorial view; b, d, f, h, j, l, p: Polar view). Scale bar = 20 µm………………………………………………………………………….87 Figure 40. LM micrographs of pollen grains in the Salvia taxa examined: a-b. S. hypargeia, c-d. S. indica, e-f. S. palaestina, g-h. S. glutinosa, i-j. S. dicroantha, k-l. S. staminea, m-n. S. russellii, o-p. S. verticillata subsp. verticillata (a, c, e, g, i, k, m, o: Equatorial view; b, d, f, h, j, l, p: Polar view). Scale bar = 20 µm…………….....88 Figure 41. SEM micrographs showing pollen morphology of S. huberi (Type 1a)...89 Figure 42. SEM micrographs showing pollen morphology of S. rosifolia (Type 1a)…………………………………………………………………………………...90 Figure 43. SEM micrographs showing pollen morphology of S. absconditiflora (Type 1a) ………………………………..……………………………….…………90 Figure 44. SEM micrographs showing pollen morphology of S. multicaulis (Type 1a)…………………………………………………………………………………...90 Figure 45. SEM micrographs showing pollen morphology of S. syriaca (Type 1a)…………………………………………………………………………………...91 Figure 46. SEM micrographs showing pollen morphology of S. virgata (Type 1a) ………………………………………………………………………………………91 Figure 47. SEM micrographs showing pollen morphology of S. verticillata subsp. verticillata (Type 1a)….…………..………………..……………………………….91 Figure 48. SEM micrographs showing pollen morphology of S. bracteata (Type 1b)………………………………………………………………………….………..92 Figure 49. SEM micrographs showing pollen morphology of S. macrochlamys (Type 1b) ………………………………..…………..……………..………………………92 Figure 50. SEM micrographs showing pollen morphology of S. trichoclada (Type 1b).. ………………………………..………………..………………………………93 xviii

Figure 51. SEM micrographs showing pollen morphology of S. hypargeia (Type 1b) ……………………………………………………………………………………….93 Figure 52. SEM micrographs showing pollen morphology of S. ballsiana (Type 2)…………………………………………………………………………………….94 Figure 53. SEM micrographs showing pollen morphology of S. blepharochleana (Type 3a)… ………………………………..……………………..………………....95 Figure 54. SEM micrographs showing pollen morphology of S. aethiopis (Type 3a)…………………………………………………………………………………...95 Figure 55. SEM micrographs showing pollen morphology of S. eriophora (Type 3a) ……………………………………………………………………………………….95 Figure 56. SEM micrographs showing pollen morphology of S. glutinosa (Type 3a)…………………………………………………………………………………...96 Figure 57. SEM micrographs showing pollen morphology of S. pachystachys (Type 3b).. ………………………………..………………………………..….….….....….97 Figure 58. SEM micrographs showing pollen morphology of S. euphratica var. euphratica (Type 3b)… ………………………………..…………………..…....….97 Figure 59. SEM micrographs showing pollen morphology of S. viridis (Type 3b)……………………………………………………………………………..….....97 Figure 60. SEM micrographs showing pollen morphology of S. cilicica (Type 3b)…………………………………………………………………………………...98 Figure 61. SEM micrographs showing pollen morphology of S. cyanescens (Type 3b)… ………………………………..…………………………..……………...…...98 Figure 62. SEM micrographs showing pollen morphology of S. indica (Type 3b)..……………………………………………………………………………….....98 Figure 63. SEM micrographs showing pollen morphology of S. limbata (Type 3b)……………………………………………………………………………….…..99 Figure 64. SEM micrographs showing pollen morphology of S. palaestina (Type 3b)….. …………………………….……………...………………………...…….. ..99 Figure 65. SEM micrographs showing pollen morphology of S. dicroantha (Type 3b)… ………………………………..………………….…..……………………….99 Figure 66. SEM micrographs showing pollen morphology of S. nemorosa (Type 3b)… ………………………………..………………………………...………...…100 xix

Figure 67. SEM micrographs showing pollen morphology of S. staminea (Type 3b)… ………………………………..…………………………………….…….…100 Figure 68. SEM micrographs showing mericarp morphology of S. ballsiana (Type 1)… ………………………………..……………………………………..…….….117 Figure 69. SEM micrographs showing mericarp morphology of S. caespitosa (Type 1)… ………………………………..……………………………..…………….….118 Figure 70. SEM micrographs showing mericarp morphology of S. macrochlamys (Type 1)…………… ………………………………..…………………………...... 118 Figure 71. SEM micrographs showing mericarp morphology of S. pilifera (Type 1) ……………………………………………………………………………………...118 Figure 72. SEM micrographs showing mericarp morphology of S. euphratica var. leiocalycina (Type 1)… ………………………………..……………..…..……….119 Figure 73. SEM micrographs showing mericarp morphology of S. aethiopis (Type 1)…………………………………..……………………………..……...……..…..119 Figure 74. SEM micrographs showing mericarp morphology of S. rosifolia (Type 1)… ………………………………..…………………………………....…………119 Figure 75. SEM micrographs showing mericarp morphology of S. kronenburgii (Type 1)… ………………………………..……………………………..…………120 Figure 76. SEM micrographs showing mericarp morphology of S. huberi (Type 1)…………………………………………………………………………….……..120 Figure 77. SEM micrographs showing mericarp morphology of S. russellii (Type 1)………………………………………………………………….………………..120 Figure 78. SEM micrographs showing mericarp morphology of S. pinnata (Type 2)……………………………………………………………….…………………..121 Figure 79. SEM micrographs showing mericarp morphology of S. atropatana (Type 2)………………………………………..………………………………………….121 Figure 80. SEM micrographs showing mericarp morphology of S. indica (Type 2)…………………………………………………………………..……………….122 Figure 81. SEM micrographs showing mericarp morphology of S.recognita (Type 3)...… ………………………………..…………………………………………….122 Figure 82. SEM micrographs showing mericarp morphology of S. euphratica var. euphratica (Type 3)……… ………………………………..…..………………….123 xx

Figure 83. SEM micrographs showing mericarp morphology of S. ceratophylla (Type 4)… ………………………………..………………………………..…...... 123 Figure 84. The richest three regions in terms of the endemic Salvia taxa in the study area shaded. These regions are indicated by circles. : East Anatolia, : Southeast Anatolia…………………………………..……………………………..139 Figure 85. The number of the Salvia taxa studied within each square. Endemics/Non- endemics……………………………..…………………………….………………139 Figure 86. S. hydrangea growing on rocky volcanic slopes in Ağrı Mountain……145 Figure 87. S. longipedicellata growing in fields………………………………..….146 Figure 88. S. euphratica growing on marly banks………………………………....146 Figure 89. S. caespitosa growing on steppe……………………….…..…..…….....147 Figure 90. S. divaricata growing on limestone slopes…………………...………...147 Figure 91. S. poculata growing in subalpine meadows……………..……………..148 Figure 92. S. multicaulis growing on calcareous rocky slopes……………..……...148 Figure 93. S. aethiopis growing along roadsides…………………..…………..…..149 Figure 94. S. glutinosa growing on moist places in Picea orientalis forest…….....149 Figure 95. S. ballsiana growing on moist places in Quercus scrub…………...…...150 Figure 96. Distribution of the Salvia taxa studied according to the 2001 IUCN Red List categories at regional, national and global scales, phytogeographic elements, and geographic regions…………………………………………....……………………158 Figure 97. Phenogram showing the seven section of Salvia by means of UPGMA algorithm and Gower General Similarity. (1) Sect. Hemisphace, (2) Sect. Drymosphace, (3) Sect. Plethiosphace, (4) Sect. Aethiopis, (5) Sect. Horminum, (6) Sect. Hymenosphace, (7) Sect. Salvia…………………………………………...…170 Figure 98. General appearance of S. divaricata in its natural habitat……...………181 Figure 99. General appearance of S. tomentosa in its natural habitat………...……183 Figure 100. Photograph of a herbarium sheet of S. kurdica…………..…..…………185 Figure 101. General appearance of S. macrochlamys in its natural habitat……..…187 Figure 102. Distribution of S. divaricata, S. tomentosa, S. kurdica and S. macrochlamys in the study area…………………..…………..………………..…..188 Figure 103. General appearance of S. recognita in its natural habitat……...…...…189 Figure 104. General appearance of S. pilifera in its natural habitat………….....….191 xxi

Figure 105. General appearance of S. pinnata in its natural habitat………...…..…193 Figure 106. Distribution of S. recognita, S. pilifera and S. pinnata in the study area…………………………………………………………………………………193 Figure 107. General appearance of S. bracteata in its natural habitat…...……...…195 Figure 108. General appearance of S. trichoclada in its natural habitat………...... 198 Figure 109. General appearance of S. anatolica in its natural habitat………..……200 Figure 110. Distribution of S. bracteata, S. trichoclada and S. anatolica in the study area…………………...……………………..……………………………...………200 Figure 111. General appearance of S. rosifolia in its natural habitat……………....202 Figure 112. General appearance of S. huberi in its natural habitat……………...…204 Figure 113. General appearance of S. caespitosa in its natural habitat……………206 Figure 114. General appearance of S. pachystachys in its natural habitat…………208 Figure 115. General appearance of S. hedgeana in its natural habitat………...…...210 Figure 116. Distribution of S. rosifolia, S. huberi, S. caespitosa, S. pachystachys and S. hedgeana in the study area………………...…………..………………………...211 Figure 117. General appearance of S. suffruticosa in its natural habitat……….…..214 Figure 118. General appearance of S. ballsiana in its natural habitat…………...... 217 Figure 119. Distribution of S. suffruticosa and S. ballsiana in the study area……..217 Figure 120. General appearance of S. blepharochleana in its natural habitat…...... 219 Figure 121. General appearance of S. hydrangea in its natural habitat……………222 Figure 122. General appearance of S. euphratica var. euphratica in its natural habitat.... …………………………………….……………..……………………....225 Figure 123. General appearance of S. euphratica var. euphratica in its natural habitat………………………………..……………………………….………….…225 Figure 124. General appearance of S. pseudeuphratica in its natural habitat…..….227 Figure 125. General appearance of S. cerino-pruinosa in its natural habitat. …….229 Figure 126. General appearance of S. kronenburgii in its natural habitat……..…..231 Figure 127. Distribution of S. blepharochleana, S. hydrangea, S. euphratica var. euphratica, S. euphratica var. leiocalycina, S. pseudeuphratica, S. cerino-pruinosa and S. kronenburgii in the study area……………………………………..………..232 Figure 128. General appearance of S. multicaulis in its natural habitat…………....234 Figure 129. General appearance of S. absconditiflora in its natural habitat...……..236 xxii

Figure 130. Distribution of S. multicaulis and S. absconditiflora in the study area…………………………………………………………………………………237 Figure 131. General appearance of S. viridis in its natural habitat……………..….239 Figure 132. Distribution of S. viridis in the study area……….………..………..…239 Figure 133. General appearance of S. syriaca in its natural habitat……………….242 Figure 134. General appearance of S. hypargeia in its natural habitat…………….244 Figure 135. General appearance of S. montbretii in its natural habitat……………246 Figure 136. Distribution of S. syriaca, S. hypargeia and S. montbretii in the study area……….………….…………………………..………………………………....246 Figure 137. General appearance of S. spinosa in its natural habitat…………….…248 Figure 138. General appearance of S. macrosiphon in its natural habitat………....250 Figure 139. General appearance of S. palaestina in its natural habitat………..…..253 Figure 140. Distribution of S. spinosa, S. macrosiphon and S. palaestina in the study area…..…………………..………..………………………………..……………....253 Figure 141. General appearance of S. eriophora in its natural habitat………….....255 Figure 142. General appearance of S. brachyantha in its natural habitat………….257 Figure 143. Distribution of S. eriophora and S. brachyantha in the study area…...258 Figure 144. General appearance of S. sclarea in its natural habitat…………...…..260 Figure 145. General appearance of S. aethiopis in its natural habitat…………..…262 Figure 146. Distribution of S. sclarea and S. aethiopis in the study area………….262 Figure 147. General appearance of S. ceratophylla in its natural habitat……….…264 Figure 148. General appearance of S. longipedicellata in its natural habitat…...... 266 Figure 149. Distribution of S. ceratophylla and S. longipedicellata in the study area ……………………………………………………………………………………..267 Figure 150. General appearance of S. atropatana in its natural habitat…...……....269 Figure 151. General appearance of S. microstegia in its natural habitat………...... 271 Figure 152. General appearance of S. xanthocheila in its natural habitat………....273 Figure 153. Distribution of S. atropatana, S. microstegia and S. xanthocheila in the study area…………………..…………..………………………………..…………274 Figure 154. General appearance of S. frigida in its natural habitat………...…...…276 Figure 155. General appearance of S. poculata in its natural habitat……..…….....278 Figure 156. General appearance of S. odontochlamys in its natural habitat…….....280 xxiii

Figure 157. Distribution of S. frigida, S. poculata and S. odontochlamys in the study area……………………...………..………………………………..……………….280 Figure 158. General appearance of S. candidissima subsp. candiddissima in its natural habitat………...……………..………………………………..……………282 Figure 159. General appearance of S. cyanescens in its natural habitat……..….....284 Figure 160. General appearance of S. cilicica in its natural habitat………...……..286 Figure 161. Distribution of S. candidissima subsp. candidissima, S. cyanescens and S. cilicica in the study area…………………………….…………………………..286 Figure 162. General appearance of S. limbata in its natural habitat………..……...288 Figure 163. General appearance of S. indica in its natural habitat……………...... 290 Figure 164. General appearance of S. siirtica in its natural habitat…………….…292 Figure 165. Distribution of S. limbata, S. indica and S. siirtica in the study area....293 Figure 166. General appearance of S. glutinosa in its natural habitat………..…....294 Figure 167. Distribution of S. glutinosa in the study area………………..………..295 Figure 168. General appearance of S. staminea in its natural habitat…………...... 297 Figure 169. General appearance of S. virgata in its natural habitat………...……..299 Figure 170. General appearance of S. nemorosa in its natural habitat……...……..301 Figure 171. General appearance of S. dicroantha in its natural habitat………...…303 Figure 172. Distribution of S. staminea, S. virgata, S. nemorosa and S. dicroantha in the study area……………………………………………………………………...303 Figure 173. General appearance of S. verticillata subsp. verticillata in its natural habitat………………...…………………………………………………………....305 Figure 174. General appearance of S. verticillata subsp. amasiaca in its natural habitat………………………………………………………………..……………307 Figure 175. General appearance of S. russellii in its natural habitat…………..….309 Figure 176. Distribution of S. verticillata subsp. amasiaca, S. verticillata subsp. verticillata and S. russellii in the study area……………………..……………..…309

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LIST OF ABBREVIATIONS c., ca……………………..circa (about) cm………………………..centimeter D………………………....P.H. Davis Da………………………..Mountain E………………………….East Euro-Sib…………………Euro-Siberian E. Karabacak…………….Ersin Karabacak F. Celep………………….Ferhat Celep holo……………………....Holotype Ir.-Tur……………………Irano-Turanian IUCN…………………….the International Union for Conservation Nature iso………………………..isotype Medit…………………….Mediterranean m…………………………meter mm……………………….millimeter N…………………………North S………………………….South S. Bagherpour……………Safi Bagherpour sect……………………….section subsp……………………..subspecies Syn……………………….Synonym W…………………………West var………………………...variety

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CHAPTER 1

INTRODUCTION

1.1. Historical Background of the Family Lamiaceae

The family name “Labiatae”, first having been used in the Genera Plantarum of de Jussieu (1789) originates from the Latin word “labium”, referring to bilipped corolla of zygomorphic flowers, a characteristic feature of the family. As an alternative to Labiatae, the name “Lamiaceae” was proposed by the English botanist John Lindley in 1836 and later on approved by the International Code of Botanical Nomenclature in 1935. Though this is still considered an acceptable alternative name, most botanists now use the name Lamiaceae in referring to this family.

The family Lamiaceae is contained within the order Lamiales along with more than 23000 species in at least 23 families (Schäferhoff et al., 2010). It is one of the largest and most distinctive angiosperm families with almost 7200 species belonging to 236 genera, including many well-known herbs, subshrubs, shrubs and trees of horticultural, economic and medicinal significance (Harley et al., 2004). In Turkey, Lamiaceae is the third largest family with 45 genera and 574 species, 256 of which are endemic. The rate of endemism is about 44.5% in this family (Davis, 1965-1985; Davis et al., 1988; Güner et al., 2000).

In Lamiaceae species, stems are often quadrangular and erect to prostrate, usually with iridoids and phenolic glycosides. Leaves are often opposite, occasionally whorled, very rarely alternate, simple, sometimes lobed or dissected, or pinnately or palmately compound, entire to serrate, and petiolate or sessile. Inflorescences have an intermediate main axis and determinate (cymosely branched) lateral axes, often congested into pseudowhorls, terminal or axillary. Bracts are usually present, persistent or deciduous, and bracteolate or not. Flowers are bisexual and often bilateral. Calyces are 4-5 (-9),

persistent, connate, actinomorphic (=radial) to zygomorphic (=bilateral), usually 2- lipped, tubular, bell-shaped or wheel-shaped, and occasionally enlarged in fruit. Corollas are often 5, connate, actinomorphic to more often slightly to strongly zygomorphic, and often 2-lipped (rarely 1-lipped). Corolla tube is short to elongate, rarely spurred, often with annulus of hairs or appendaged within. Stamens are attached within corolla tube, and are 4 or 2 by abortion. Filaments are short of often elongate and are usually exserted from corollas (Judd et al., 2002; Harley et al., 2004).

Lamiaceae is characterized by aromatic plants which have been widely used since ancient times in many parts of Asia and the Mediterranean region. It is known for medicinal, culinary and ornamental herbs such as basil (Ocimum), rosemary (Rosmarinus), thyme (Thymus), mint (Mentha), sage (Salvia), savory (Satureja), balm (Melissa), catmint (Nepeta) and oregano (Origanum). Most species are frequently cultivated owing to not only their aromatic qualities but also their ease of cultivation (Wagstaff et al., 1998; van Jaarsveld, 2002). Many other aromatic species are also used in perfumes or toilet preparations. Another significant attribute of many Lamiaceae species is as a rich source of nectar for many honeys. They play a significant role in the production of many Mediterranean honeys (Harley et al., 2004).

Although Lamiaceae has an almost cosmopolitan distribution, with the exception of the coldest polar regions, many species of the family are especially well found in tropical and temperate areas such as the Mediterranean region and tropical upland savannas (Dorman et al., 2004). A review of the biogeography of the family was performed by Hedge (1992), who recognized six main regions of Lamiaceae diversity: (1) Mediterranean and Southwest Central Asia, (2) Africa south of Sahel and Madagascar, (3) China, (4) Australia, (5) South America and (6) North America and Mexico. Indomalesian region is also a major diversity center of subfamilies Symphorematoideae and Viticoideae (Harley et al., 2004).

The showy flowers of Lamiaceae are pollinated by bees, wasps, butterflies, moths, flies, beetles and birds. The upper lip of the two-lipped corolla protects the stamens and

stigma, whereas the lower lip provides a landing platform. The pollinator is dusted with pollen on its back or head as it probes for nectar. However, the stamens of Ocimum and its relatives lie close to the lower lip and deposit pollen on the underside of the pollinator (Judd et al., 2002). One of the best known and most elaborate types of pollination mechanism within Lamiaceae is that of Salvia (Harley et al., 2004). The flowers possess only two stamens, normally with a single fertile theca and a very elongate connective which articulates with the often shorter filament. The lower theca forms a sterile appendage that usually acts as a lever, tapping the pollen directly onto the back of the pollinator. There is rather variation on this theme within the genus, which includes both insect and bird pollinators. In some species, such as S. verticillata, there is no articulation between the filament and connective in that the posterior lip of the corolla is basally narrowed and readily folds back exposing stamens. In a number of genera there has been a shift towards bird pollination, especially in such groups as the New World Salvia subgenus Calosphace, many of whose red and large flowered species pollinated hummingbirds (Harley et al., 2004). The nutlets of many species of Lamiaceae are only shaken from calyx by the action of wind or disturbance of the plant. They may be also eaten by birds or dispersed by water (Judd et al., 2002).

The earliest taxonomic treatments of Lamiaceae were Bentham‟s monograph (1832- 1836) and his later modifications (Bentham, 1848; Bentham, 1876). Bentham (1876) subdivided the family into eight tribes, which are Ocimeae (=Ocimoideae), Mentheae (=Satureineae), Salvieae (=Monardeae), Nepeteae, Lamieae (=Stachydeae), Prasieae, Prostanthereae and Ajugeae (=Ajugoideae). Bentham‟s (1876) classification was followed by that of Briquet (1895-1897) with various modifications. He increased the rank of some of Bentham‟s tribes and subtribes to the subfamilial level and reclassified a few genera (Cantino & Sanders, 1986; Cantino, 1992a). More recent versions of the classification were made by many researchers (Junell, 1934; Erdtman, 1945; Melchior, 1964; Wunderlich, 1967; El-Gazzar & Watson, 1968, 1970; Sanders & Cantino, 1984; Cantino & Sanders, 1986; Cantino, 1992a, b; Cantino et al., 1992; Thorne, 1992; Wagstaff et al., 1995, 1997; Harley et al., 2004).

An alternative classification of Lamiaceae was proposed by Erdtman (1945) based on palynological properties, such as the number of apertures and nuclei. He subdivided the family into the two subfamilies: Lamioideae and Nepetoideae. Whereas the subfamily Lamiodieae has mostly tricolpate pollen shed in the two-celled stage, albuminous seeds, spatulate embryos and the presence of iridoid glycosides, the subfamily Nepetoideae is often characterized by hexacolpate (rarely 8-12 colpate) pollen shed in the three-celled stage, exalbuminous seeds with an investing embryo, the absence of iridoid glycosides. Additionally, most species of the Nepetoideae are strongly aromatic with terpenoids and rosmarinic acid present, and possess highly unsaturated seed oils and usually mucilaginous pericarps (Erdtman, 1945; Martin, 1946; Wunderlich, 1967; Kooiman, 1972; Zoz & Litvinenko, 1979; Cantino & Sanders, 1986; Ryding, 1992; Harley et al., 2004). Erdtman's subfamilial classification is highly agree with Bentham's (1876) tribal classification, four of Bentham's tribes constituting subfamily Lamioideae, and the other four constituting subfamily Nepetoideae (Cantino & Sanders, 1986). Wunderlich's (1967) extensive pollen survey also provided strong support for Erdtman's classification through the addition of many new genera to the palynological data base. Based on morphology, but affected by molecular phylogenetic analyses (Cantino, 1992a, b; Wagstaff, 1992), Cantino et al. (1992) proposed a new classification of Lamiaceae that substantially differed from earlier taxonomic treatments (Bentham, 1876; Briquet, 1895- 1897), however was very similar to a classification proposed earlier by Junell (1934). Their classification was supported by Thorne (1992).

In the most recent classification of the Lamiaceae (Harley et al. 2004), it is divided into the seven subfamilies: Ajugoideae Kostel., Lamioideae Harley, Nepetoideae (Dumort.) Luerss., Prostantheroideae Luerss., Scutellarioideae (Dumort.) Caruel, Symphorematoideae Briq. and Viticoideae Briq. Among these subfamilies Nepetoideae is one of the most distinctive groups, which is consistently monophyletic in various molecular and morphological analyses (Cantino & Sanders, 1986; Cantino, 1992a, b; Cantino et al., 1992; Wagstaff et al., 1995, 1998; Wagstaff & Olmstead, 1997). Although broadening taxonomic coverage and increasing amounts of molecular data have begun to improve resolution of phylogenetic relationships among tribes, subtribes

and genera within the subfamily, they are poorly understood since complex and possibly homoplasious morphological characters make taxa difficult to distinguish. The large subfamily Nepetoideae is divided into the three tribes: Elsholtzieae, Mentheae and Ocimeae. Mentheanae is the largest and economically most significant tribe of the Nepetoideae, consisting almost 65 genera (Harley et al., 2004). Mentheae is divided into the three subtribes: Salviinae, Menthinae and Nepetinae (Harley et al. 2004). The subtribe Salviinae is represented by eight genera, which have been classified differently by several authors (Table 1; Harley et al., 2004). Based on the current molecular data, the Salviinae is recognized as a well supported monophyletic subtribe within the Mentheae, however its largest genus Salvia seems to be polyphyletic (Walker et al., 2004 ; Walker & Sytsma, 2007).

Table 1. Genera of the subtribe Salvinaeae sensu Harley et al. (2004) and their previous taxonomic positions. NI = Not indicated. The number in parentheses shows the total number of species in each genus. Harley et al. (2004) Wunderlich Briquet Bentham (1967) (1897) (1876) Chaunostmata Donn. Sm. (1) NI Incertae sedis NI Dorystaechas Boiss. & Heldr. (1) Meriandreae Meriandreae Monardeae Lepechiana Willd. (40) Lepechinieae Lepechinieae Satureineae Meriandra Benth. (2) Meriandreae Meriandreae Monardeae Perovskia Kar. (7) Meriandreae Meriandreae Monardeae Rosmarinus L. (3) Ajugoideae Ajugoideae Monardeae Salvia (1000) Salviaeae Salviaeae Monardeae Zhumeria Rech. f. & Wendelbo (1) NI NI NI

It is widely accepted that Lamiaceae evolved from Verbenaceae (Cronquist, 1981). The two families share many characters, such as opposite leaves, zygomorphic flowers and bicarpellate gynoecium. The Lamiaceae is generally separated from Verbenaceae on the basis of its gynoecium and fruit structure. The Lamiaceae often possesses genera with a

deeply four-lobed ovary with a gynobasic style while most Verbenaceae have an unlobed ovary with a terminal style. However, taxa with an intermediate morphology exist in both families, and the boundary between the two families is somewhat arbitrary (Cronquist 1981). Cantino (1992a) observed that the Lamiaceae as circumscribed by Bentham (1876) and Briquet (1895-1897) was polyphyletic, with at least four lineages of Lamiaceae arisen independently within a paraphyletic Verbenaceae.

Pollen micromorphological studies for Lamiaceae has been carried out by many researchers (Erdtman, 1945; Emboden, 1964; Henderson et al., 1968; Rudal, 1980; Basset & Munro, 1986; Cantino et al., 1992a, b; Demissew & Harley, 1992; Harley, 1992; Harley et al., 1992, 2004; Abu-Asab & Cantino, 1992, 1993, 1994; Celenk et al., 2008a, b; Salmaki et al., 2008a; Moon et al., 2008a, b; Bagherpour et al., 2009; Bagherpour et al., 2010; Hassan et al., 2009; Kahraman et al., 2009b, c, 2010b, c, d; Kahraman & Doğan, 2010; Özler et al., 2011). Some of these studies have been proved to be useful in systematic of Lamiaceae and its certain genera (Erdtman, 1945; Emboden, 1964; Henderson et al., 1968; Cantino et al., 1992; Harley et al., 1992, Abu- Asab & Cantino, 1994, Moon et al., 2008a, b).

Mericarp (nutlet) micromorphology provide useful systematic characters at various taxonomic levels in Lamiaceae, depending on the characters chosen and the variation present (Isley, 1947; Wojciechowska, 1961, 1966; Husain et al., 1990; Rejdali, 1990; Demissew & Harley, 1992; Marin et al., 1994, 1996; Oran, 1996; Turner & Delprete, 1996; Zhou et al., 1997; Guerin, 2005; Moon & Hong, 2006; Kaya & Dirmenci, 2008; Salmaki et al., 2008b; Bagherpour et al., 2010; Kahraman & Doğan, 2010; Kahraman et al., 2010b, c; Büyükkartal et al., 2011; Kahraman et al., 2011a, b). Husain et al. (1990) investigated the mericarp micromorphology in the Lamiaceae tribe Saturejeae. In the tribe Saturejeae, sculpturing patterns were the most useful characters. Mericarp shape and the nature of abscission scars were invariable in the tribe Saturejeae, but in the tribe Westringieae these characters varied significantly. Marin et al. (1994) characterized mericarps of Teucrium by or absence and type of oil glands on the mericarp surface and concluded that these characters can be used as taxonomic markers within the family at

genus, section and species levels. Guerin (2005) found that mericarp shape, the nature of abscission scars, indumentum, surface sculpturing, exocarp cell shape and sculpturing were useful characters at infrageneric classifications of Hemigenia and Microcorys.

1.2. Historical Background of the Genus Salvia L.

The genus name “Salvia” is derived from the Latin “salvare”, meaning “to heal or save”, and “salvus”, meaning uninjured or whole, and refers to medicinal properties of some species of Salvia (Clebsch, 2003). The derivation of the common English name is through the Old French form “sauge”, then to the old English “sawge”, and eventually to the modern English “sage” (Dweck, 2000). The genus is known to the Egyptians as “anusi”, to the Greeks as “elelisphakon” and to Spanish and Moroccan Arab herbalists as “salima” or “asphacus” (Rivera et al., 1994).

In Turkey, Salvia L. is known as “adaçayı” and is frequently consumed as teas (Baytop, 1999). Many Salvia species are used in folk medicines throughout the world, possessing antioxidant, antidiabetic, antimicrobial, antitumor, antiplasmodial and anti-inflammatory properties (Ulubelen, 2003; Kamatou et al., 2008; Şenol et al., 2010). Some Salvia species have been also recorded to be used against memory loss in Europe folk medicine (Perry et al., 1996). Additionally, a number of Salvia species are also used in food, cosmetics, perfumery and the pharmaceutical industry (Chalchat et al., 1998; Baylac & Racine, 2003). In addition to medicinal uses, they are also grown in parks and gardens as ornamental plants (Marin et al., 1996).

Salvia is the largest genus of the family Lamiaceae and comprises almost one quarter of the family, composed of approximately 1000 species occurring worldwide. The genus is distributed extensively in tropical and temperate areas of the Old and New World, with three distinct regions of population: Central and South America (ca. 500 spp.), Western Asia (ca. 200 spp.) and Eastern Asia (ca. 100 spp.) (Walker & Sytsma, 2007) (Figure 1).

Figure 1. Worldwide distribution of species of the genus Salvia. Numbers refer to approximate number of species and circles show present centers of diversity (Walker et al., 2004).

The oldest findings of pollen of the genus Salvia show to originate from the lower Miocene in Mexico (Graham, 1999) and from the upper Miocene in Alaska (Müller, 1981). Therefore, this indicates an age of Salvia about 25 million years. In accordance with Himmelbaur & Stibal (1933-1935), East Asia is the occurrence center of the genus on the basis of floral morphology. It is assumed at the present time that Salvia originated in the Old World, probably from an ancient Mediterranean species (Dieringer et al., 1991; Baikova 1999). Salvia species migrated during the Miocene and Pliocene via tropical mountains to South America and South Africa (Baikova, 1999).

The first monographic treatment of Salvia was made by Etlinger (1777) who recognized 48 species. Bentham (1832-1836, 1876) gave a full monographic account of the genus and subdivided it into four subgenera, twelve sections (Table 2) and many series on the basis of differences in leaf, calyx, corolla and stamen morphology. The section Eusphace created by Bentham was later designated as the type section of Salvia and changed to the section Salvia (Hedge, 1972). Bentham‟s system remains until now the most widely accepted treatment of Salvia, even though over 500 new species have been

discovered. Bentham‟s (1876) classification system was modified by a number of researchers (Briquet, 1897; Ferald, 1900; Stibal, 1934, 1935; Epling, 1938, 1939; Pobedimova 1954; Hruby, 1962; El-Gazzar & Watson, 1968; Hedge 1972). However, Hedge (1974, 1982a, b) avoided presenting Bentham‟s subgeneric groupings.

Fernald (1900) revised the Mexican and Central American species of Salvia, following Bentham‟s classification of the genus. Epling (1939), who provided the most recent revision of the subgenus Calosphace, recognized 468 species in 91 sections, generally raising Bentham's subsections and series to sections. These sections were characterized mainly by floral characters and geography. Epling‟s classification of Calosphace still remains the best available as it is the most comprehensive, covering twice as many species as Bentham‟s treatments (Santos, 2004). More recently, Pobedimova (1954) proposed seven subgenera, eight sections, and many series for the genus in Flora U.S.S.R. El-Gazzar & Watson (1968), in a numerical revision of morphological characters of Salvia, found the traditional subgenera and sections and concluded that the genus can be separated into a New World group and an Old World group.

Table 2. Subgenera and sections of the genus Salvia classified by Bentham (1832-1836). Subgenera Sections Calosphace Calosphace Leonia Echinosphace Pycnospahce Heterosphace Notiosphace Hemisphace Salvia Eusphace (synonym of Salvia) Hymenosphace Drymosphace Sclarea Horminum Aethiopis Plethiosphace

Salvia is separated from the other 72 genera in the tribe Mentheae by the presence of two aborted posterior stamens and a markedly elongated connective tissue separating thecae of the two expressed stamens, which may form a lever mechanism of pollination (Huck, 1992; Claβen-Bockhoff et al., 2003; Reith et al., 2007). Since the dorsal pollination mechanism in Salvia was first described by Sprengel (1793), the importance of the lever mechanism to reproductive biology has been investigated by many researchers in Salvia (Hruby, 1934; Claβen-Bockhoff et al., 2003, 2004a, b; Reith et al., 2007; Walker & Sytsma, 2007). The evolution of the unique lever mechanism and stamen connective has occurred parallel in the Old World and the New World (Himmelbaur & Stibal, 1932-1934; Claβen-Bockhoff et al., 2004a; Walker & Sytsma, 2007).

It was long assumed that an unusual pollination and stamen structure of Salvia had evolved only once, and that thus Salvia was monophyletic, meaning that all taxa of the genus evolved from one ancestor. However, recent molecular phylogenetic analyses in Salvia and related genera have revealed that Salvia is polyphyletic, with three major lineages and five other genera intercalated within it and the staminal lever mechanism evolved three times independently, each time with a distinct morphology (Walker et al., 2004; Walker & Sytsma, 2007). In accordance with Walker & Sytsma (2007), “Salvia clade” contains all examined members of Salvia and the genera Dorystaechas, Meriandra, Perovskia, Rosmarinus and Zhumeria, and is characterized morphologically by the abortion of the two adaxial stamens. The genera Lepechinia and Melissa are closely related, and together with the Salvia clade constitute a monophyletic group within the Mentheae. The Salvia clade comprises three distinct lineages, namely Salvia clade I, Salvia clade II and Salvia clade III. Therefore, Salvia is not monophyletic. The Salvia clade I sensu Walker & Sytsma (2007), which is a sister to Rosmarinus and Perovskia, comprises European, Central African, Southern African and West Asian Salvia species. The Salvia clade II, which is sister to Meriandra and Dorystaechas, contains New World species composed of subgenus Calosphace and section Audibertia. The Salvia clade III, which is sister to Zhumeria, is composed of West Asian, Central Asian, East Asian, Mediterranean and African species. According to the literature

(Epling, 1938, 1939; Hedge, 1974, 1982a, b; Walker & Sytsma, 2007), the Salvia clades I, II and III are represented by about 250, 520 and 105 species, respectively.

Based on the staminal morphology of Salvia, nine distinct stamen types are identified in species within Salvia clades I, II and III (Walker & Sytsma, 2007). Two different stamen types (stamen types A and B) are observed in about 23 species within the Salvia clade I. In the stamen type A, the two posterior thecae are expressed and not fused (Figure 2; e.g. S. aucheri var. aucheri and S. hydrangea known from Turkey). In the stamen type B, the two posterior thecae are not expressed, and the distal posterior ends of the adjacent connectives are fused into a complex structure blocking access to nectar (Figure 2; e.g. S. aethiopis, S. candidissima, S. sclarea and S. verbascifolia = synonym of S. microstegia known from Turkey). The A and B stamen types characterized by Hedge (1982a) and (Walker & Sytsma, 2007) are identical.

Rosmarinus possesses a significantly elongated connective tissue in its two stamens, and a total abortion of the posterior branch of the connective and the posterior theca (stamen type C; Figure 2). Perovskia has a slightly elongate connective tissue in its two expressed stamens (stamen type D; Figure 2).

Five stamen types (stamen types E to I) are recognized in 49 species within the Salvia clade II. The subgenus Calosphace, which is composed of nearly 500 species found throughout the New World, is characterized by E to G types of stamens. In the stamen type G, both posterior thecae are expressed, and not fused to one another (Figure 2; S. axillaris in the monotypic section Axillares). In the stamen type F, both posterior thecae are aborted, and the adjacent posterior thecae are not or only little fused (Figure 2; e.g. species in sections Blakea, Hastatae and Standleyana). In the stamen type E, both posterior thecae are aborted and adjacent posterior connective branches are fused (Figure 2; e.g. the remaining species of the subgenus Calosphace). The section Audibertia, which is confined to the California Floristic Province and adjacent deserts (Walker et al., 2004), is characterized by H and I types of stamens. The stamen type H has a completely aborted posterior theca and connective arm (Figure 2; e.g. S. dorrii, S. leucophylla and S.

melilifera) while the stamen type I has a reduced posterior theca (Figure 2; e.g. S. californica and S. greatai).

Meriandra has slightly elongate connectives (stamen type J, Figure 2) whereas Dorystaechas possesses slightly elongate and swollen connectives (stamen type K, Figure 2).

Figure 2. Representations of the stamen types defined per clade suggested by the molecular results (Walker & Sytsma, 2007). 12

Two stamen types (stamen types M to N) are recognized in 11 species within the Salvia clade III. In the stamen type M, both posterior thecae are expressed and not fused to one another (Figure 2; e.g. S. aristata known from Turkey, S. aegyptiaca and S. tetradonta). In the stamen type N, both posterior thecae are aborted, or expressed and produce little or no pollen (Figure 2; e.g. S. glutinosa known from Turkey and S. roborowskii).

Zhumeria has the thecae of the two fertile stamens somewhat distinguished, though without a distinct connective (stamen type L, Figure 2).

The first attempt at an infrageneric classification of Salvia was that of Bentham (1833), which listed 22 species from Turkey and divided them into seven sections: Eusphace Benth., Hymenosphace Benth., Horminum Benth., Aethiopis Benth., Drymosphace Benth., Plethiosphace Benth. and Hemisphace Benth. The section Eusphace was later changed to the section Salvia (Hedge, 1972). This was followed by the 1879 arrangement of Boissier, which recognized 75 species of Salvia in Turkey. Leaf, calyx, corolla and stamen characteristics are the most important means for distinguishing the sections (Bentham, 1833; Boissier, 1879; Hedge, 1972; Kahraman et al., 2010b).

Hedge (1982a), in his comprehensive treatment of Salvia in Turkey, recognized 86 species with two subspecies, two varieties and one naturally occurring hybrid. He grouped the species by stamen characters and other morphological similarities. Since the publication of Flora of Turkey and the East Aegean Islands, seven new species, namely S. nydeggeri (Huber-Morath, 1982), S. aytachii (Vural & Adıgüzel, 1996), S. hedgeana (Dönmez, 2001), S. anatolica (Hamzaoğlu et al., 2005), S. marashica (İlçim et al., 2009), S. ekimiana (Celep & Doğan, 2009) and S. siirtica (Kahraman et al., 2011b), two species re-evaluated as valid species, namely S. cerino-pruinosa and S. pseudeuphratica (Kahraman et al., 2010b), three new records, namely S. macrosiphon (Kahraman et al., 2009a), S. viscosa (Celep et al., 2009a) and S. aristata (Behçet & Avlamaz, 2009) and two new varieties, namely S. sericeo-tomentosa var. hatayica (Celep et al., 2009b) and S. cadmica var. bozkiriensis (Celep et al., 2011a) have been described from Turkey. S. aucheri Benth. var. canescens Boiss. & Heldr. has been also raised to subspecies rank (Celep et al., 2011b). The number of species now reaches 98 with three subspecies and 13

three varieties and the rate of endemism is 53.1%, showing that Turkey is a major centre of diversity for the genus in Asia. Distribution of the genus in neighbouring countries is as follows: 75 species in the former USSR (Pobedimova, 1954), 70 in Flora Iranica (Hedge, 1982b), 36 in Europe (Hedge, 1972), and 21 in Flora Palaestina (Zohary, 1966).

1.3. General Description of the Study Area

1.3.1. Location

Turkey is located between 36o-42o north latitude and 26o-45o east longitude and in the northern hemisphere, near the junction of European, Asian and African continents (Figure 3 & 4). It is geographically placed in the temperate zone with climate types in different regions and surrounded by eight countries along its land borders. In the west it borders on Greece and Bulgaria, in the east on Georgia, Armenia, Azerbaijan and Iran, and in the south on Iraq and Syria. Turkey has a land area of 774815 km2, 97% of which lies in Asia (the Anatolian Plateau) and the remaining 3% in Europe (Thrace). Turkey is divided in seven geographic regions: the Marmara Region, the Black Sea Region, the Mediterranean Region, the East Anatolian Region (the study area), the Southeast Anatolian Region (the study area), the Central Anatolian Region and the Aegean Region, each of which has different climatic and ecological properties. Approximate 85% of the land is at an elevation of at least 450 m and the average altitude of the country is 1332 m (Turkish e-Government Portal, 2010; Wikipedia, 2011).

Turkey has nearly 12000 natural vascular plant taxa and almost 30% of them are endemic. Endemic plants are composed of 18.6% of total plant diversity in Spain, 14.9% in Greece, 2.9% in France and 0.1% in Poland (Türe & Böcük, 2008). Thus, Turkey is one of the richest centers in the world for plant diversity and endemism owing to its great diversity in climate, geomorphology, topography and soil. The region differs from most countries in Europe and the Middle East with its rich plant diversity and ranks in 9th order in terms of biodiversity richness among continental countries (Avcı, 2005; 14

Davis, 1971; Ekim et al., 2000). It is the meeting ground of three different phytogeographical regions called Euro-Siberian, Mediterranean and Irano-Turanian (Figure 3; Davis, 1971, Türe et al., 2005). Each phytogeographical region varies in terms of floristic composition and vegetation structure. The Euro-Siberian phytogeographical region includes the Euxine province. The Mediterranean phytogeographical region includes the Mediterranean provinces of the West Anatolian Region, Amanous and Taurus Mountains. The Irano-Turanian phytogeographical region is represented by the Central, East and Southeast Anatolian geographical divisions of Turkey. A major part of this region is covered by grass, some of which are used for grazing in summer and some harvested for winter fodder. The region has the richest plant diversity in Turkey and is dominated by a typical steppe vegetation (Öztürk et al., 2008). These look like the steppes of Iran and Central Asia (Breeckle, 1986; Akhani, 2006). The Irano-Turanian and Mediterranean phytogeographical regions have more endemic taxa than the Euro-Siberian phytogeographical region in Turkey (Table 3). High representation of Irano-Turanian elements is because of conversion of Anatolian steppes in the Irano-Turanian region into farm fields (Türe & Böcük, 2008). The endemism ratio of the Black Sea (Euro-Siberian) region is about 5%. This is due to the presence of more stable environmental factors in comparison to the Central, Eastern and South Anatolian regions (Türe & Tokur, 2000; Türe et al., 2004). The west and east parts of the Anatolian Diagonal are under the effect of Irano-Turanian phytogeographical region, however they differ in terms of floristic features (Erik & Tarıkahya, 2004; Kaya & Aksakal, 2005). Many of the species forming the flora of Turkey are found along this diagonal.

Table 3. The number of endemic plant species growing in different phytogeographical regions of Turkey (Ekim et al., 2000). PhyTogeographical regions The number of endemic species Irano-Turanian 1220 Mediterranean 1050 Euro-Siberian 300

Figure 3. Main phytogeographical regions in Turkey.

The study area is situated in the East and Southeast Anatolian geographic regions of Turkey and covers an area of 224612 km2. According to Davis‟ grid square system (1965), the area is located within the A8-A9, B6-B10 and C6-C10 grid squares (Figure 4). It falls within the Irano-Turanian, Mediterranean and Euro-Siberian phytogeographical regions. East Anatolia ranks in 2th order with 380 species after the Mediterranean region in terms of the number of endemic plants among geographic regions in Turkey (Ekim et al., 2000). Southeast Anatolia ranks in the last order with 35 species in terms of the number of endemic plants among geographic regions in Turkey (Ekim et al., 2000).

Figure 4. Location of the research area.

1.3.2. Topographic Characteristics

East Anatolia, which is known as the Anti-Taurus, is the largest (165436 km2) and highest (5137 m) region in Turkey. The region is rough topographically, plains and plateaus being very high and vast around Ağrı, Hakkari, Erzurum and Van where high altitude mountains are gathered together (Öztürk et al., 2008). The montane (1200-1700 m), oreal (1700-2200 m), subalpine (2200-2700 m) and alpine (2700-3200 m) zones, as well as in high altitudes subnival and nival zones are recognized (Hamzaoğlu, 2006). There are numerous inactive quaternary volcanoes in the region, including Ağrı Mountain (5137 m), at the highest point in Turkey, and Süphan (4548 m) Tendürek (3680 m) and Nemrut Mountains (2935 m). Basic igneous rocks of various kinds cover much of East Anatolia (e.g. between Malatya and Kangal and between Elazığ and Maden), however hard limestone is present in Munzur Mountains. Cilo Mountain (4168 m) is largely dolomitic (Davis, 1965).

There are long and generally narrow valleys abruptly formed between mountains. The eastern Anatolia is the richest in terms of the number of lakes and contains Turkey's

largest lake, Lake Van (3713 km2), and the lakes of Erçek, Çıldır and Hazar. The headwaters of three major rivers arise in the region: the east-flowing Aras, the south- flowing Fırat (Euphrates) and the south-flowing Dicle (Tigris).

Southeast Anatolia is notable for the uniformity of its landscape even though the eastern part of the region is comparatively more uneven than its western part. The region is composed of rolling hills and a broad plains surface extending into Syria. In the region, elevations fall gradually, from about 800 m in the north to about 400 m near the Syrian frontier. Karacadağ (1957 m) is the highest point of the region. The west part of Karacadağ is covered by Eocene and Miocene limesotenes and resembles a desert with its unique character (Kaya et al., 2009). Mardin stands near the western rim of a limestone escarpment. Towards the south of Mardin, the vast expanse of the Syrian Desert lying below is seen (Davis, 1965).

1.3.3. Climatic Characteristics

East Anatolia is characterized by a continental type of climate with hot and extremely dry summers and very cold winters with frequently heavy snowfall. The coldest areas of Turkey are found in the region. On some high mountains (e.g. Ağrı and Cilo Mountains) harsh climatic conditions persist all the year round and even small glaciers are generated. Annual mean temperature is -7 to -10°C in winter and 17 to 19°C in summer. The hottest month is August, with a mean maximum temperature ranging varying between 24 and 28°C. Mean minimum temperature in the coolest month is between -16° and -17°C. Extremely cold temperatures of -45.6°C have been recorded around Ağrı while around Malatya and Elazığ summer temperatures are about 42°C. Rains are common in the spring and autumn. Summer rains are more extreme than those in winter, especially in the northeast around Erzurum and Kars provinces. Highest rainfall is in Tunceli (1003.8 mm) and lowest in Erzincan (359.6 mm). Mean annual precipitation varies between 430 and 790 mm (Öztürk et al., 2008).

Southeast Anatolia is under the influence of the steppe climate with very hot summers and severe aridity, with evaporation, reaching up to 1000-2000 mm (DIE, 2000). Winters are hardly frost, however severe dry winds dominate the summer season. Annual mean temperature is between 15 and 16°C, in winter between 3 and 4°C, in summer between 30° and 35°C. Average daily maximum temperature is about 30oC and the minimum temperature is between 1.5 and 6oC. Annual mean precipitation ranges from 400 to 700 mm, however the Harran plain gets only 330 mm of annual average rainfall, most of which falls in the winter (Öztürk et al., 2008).

1.3.4. Soil Characteristics

Main soil types found in the study area are alluvial, colluvial, chestnut, brown, regosols, basaltic, organic, and arid soils (Öztürk et al., 2004, 2006). According to soil properties of samples obtained from different basins, pH varies between 7.5 and 7.9, and the ranges + + 2+ 2+ – – of electrical conductivity Na , K , Ca , Mg , Cl and SO4 values are 2.4-20.50 (meq/100 g), 0.01-0.86 (meq/100 g), 1.5–31.97 (meq/100 g), 10.82-84.75 (meq/100 g), 0.58-29.41 (meq/100 g), and 1.68-21.55 (meq/100 g), respectively (Öztürk et al., 2006). The soils of the Harran plain are alluvial, dark-red or reddish brown clay, humid and calcareous, with a pH varying between 7.3 and 7.6 and total salt level ranges from 0.075 to 1.450% (Dinç and Şenol, 1988). The Aras valley between Kağızman and the Armenian border is full of impressive salt outlets, some pouring directly out of the mountains and thus resembling snow patches from the distance. In the south and south east of Sivas and around Gürün large areas of gypseous hills are found with a very special flora.

1.4. Objectives of the study

The objectives of this study were as follows: (i) to carry out a taxonomic revision of Salvia in East and Southeast Anatolia based on extensive field studies and numerous recent collections, (ii) to describe anatomical features of vegetative organs (e.g. roots, stems, leaf blades and petioles) of the taxa studied and to assess the variation in these characteristics in respect to their potential systematic value, (iii) to present a detailed account of the pollen and mericarp micromorphology of the taxa, to evaluate the utility of the pollen and mericarp data for classification at infraspecific and sectional level within the genus and provide keys for the identification of the taxa, (iv) to construct an infrageneric grouping in the genus with the use of numerical taxonomic methods, (v) to determine the conservation status of the taxa according to IUCN threat categories and criteria, (vi) to present descriptions, illustrations as well as data on distribution (including maps), phytogeography, phenology and ecology for each taxon, (vii) to provide a key for their identification on the basis of vegetative and floral characters of the taxa studied.

CHAPTER 2

MATERIALS AND METHODS

2.1. Field Surveys and Plant Material Collection

As a part of the taxonomic revision of Turkish Salvia, field surveys were carried out and fresh plant materials were collected from about 600 different populations during field trips in most parts of East and Southeast Anatolia between July 2005 and June 2009. Location of stations visited within the study area is shown in Figure 5. The specimens collected were pressed and dried according to standard techniques for further laboratory analysis (Davis & Heywood, 1973), and then they were deposited in Plant Systematics Laboratory, Department of Biological Sciences, Middle East Technical University (METU), Ankara. The study also included many herbarium specimens kept at the following local and international herbaria (for abbreviations, http://sciweb.nybg.og/science2/IndexHerbariorum.asp): AEF, ANK, B, BM, CBB, E, FUH, G, GAZI, HUB, ISTE, ISTF, K, KNYA, LE, MO, VANF and W.

During the extensive field studies, the type localities and other known localities of the Salvia taxa, and also a number of other sites where they can potentially distribute were visited. In the field, the data on distribution and habitat, GPS coordinates, population sizes, the number of mature individuals, phenological and ecological features and threat factors were recorded (Davis & Heywood, 1973).

2.2. Methods used in Systematics

The specimens were all cross-checked by using the relevant literature, such as Flora Orientalis (Boissier, 1879), Flora of Syria, Palestine and Sinai (Post, 1933), Flora of the USSR (Pobedimova, 1954), Flora Europaea (Hedge, 1972), Flora of Turkey and the

East Aegean Islands (Hedge, 1982a), Flora Iranica (Hedge, 1982b) and Nouvelle Flore du Liban et de la Syrie (Mouterde, 1983). The plant authorities were specified according to Authors of Plant Names (Brummitt & Powell, 1992).

An identification key for determination of the species studied is provided. Detailed descriptions of each species are followed by data on phenology, distribution, habitat and phytogeography and photographs. Distribution maps of the taxa are given according to Davis‟ grid square system (1965). Based on the recent taxonomic revision, herbarium materials and local floristic investigations, distribution maps were prepared.

Figure 5. Location of stations visited within the study area between 2005 and 2009.

2.3. Methods used in Macromorphological Studies of Vegetative and Reproductive Structures

Morphological studies were carried out on living as well as herbarium materials. Leaf, stem, inflorescence, calyx, corolla and stamen characters of each taxon were examined and measured so as to determine their morphological characteristics using a Leica S8AP0 stereomicroscope. Minimum and maximum ranges of measured characters are provided. 22

2.4. Methods used in Anatomical Studies

Anatomical investigations were performed for mature specimens fixed in 70% alcohol or herbarium specimens. Roots, stems, leaf blades and petioles were separated from specimens and then dissected with a razor blade. All samples were dehydrated through a graded ethanol series, transferred into xylene, embedded in paraffin wax and then sectioned at 5-20 µm thickness with a Leica RM2125RT rotary microtome. The sections were transferred onto slides and stained in solutions of 1% safranin and fast green (Johansen, 1944) with some modifications relating to staining time and amount of additions to the stains, and then permanently mounted on slides using Canada Balsam and Entellan. At least five permanent slides were prepared and examined, and photographs were taken with a Leica DM1000 light microscope.

2.5. Methods used in Palynological Studies

For light microscopy (LM), pollen grains were first treated with 70% alcohol to remove oily substances and then embedded in glycerine jelly stained with basic fuchsin following the method of Wodehouse (1935). Polar axis (P), equatorial axis (E), colpus length (Clg), colpus width (Clt), exine thickness (Ex), intine thickness (In) and apocolpium diameter (Ap) were measured from at least 30 fully developed grains per sample under a Leica DMLB2 microscope (1000x). Results are provided as minimum, maximum and mean±standard deviations.

For scanning electron microscopy (SEM), pollen grains were transferred directly to stubs with double-sided adhesive tape and micrographs were obtained using JEOL-6060 SEM at Gazi University and JEOL JSM-6400 SEM at Middle East Technical University. The terminology follows mainly that of Henderson et al. (1968), Faegri and Iversen (1989), Abu-Asab & Cantino (1994) and Punt et al. (2007).

2.6. Methods used in Mericarp Micromorphological Studies

Mature mericarps of Salvia taxa studied were collected during the fruiting period. Two to four samples were examined for each taxon after a number of specimens had been compared under a Leica S8AP0 binocular stereoscopic microscope for similarity. For the mericarp length, width, length/width ratio and abscission scar diameter, at least 20 samples were measured for each taxon. Results are provided as minimum, maximum and mean.

For SEM, selected mature mericarps were mounted on aluminum stubs using double- sided adhesive tape, coated with gold with a Hummle VII sputter coater, viewed with JEOL JSM-6400 and JEOL JSM-6490LV SEM and micrographed. After all micrographs were examined, micrographs representing each mericarp sculpturing type were selected. The micrographs were used to describe surface sculpturing of the mericarps. The descriptive terminology of Ryding (1995), Stearn (2004) and Salmaki et al. (2008b) for mericarp characters was generally followed.

2.7. Assessment of the Threat Categories

Based on the current surveys, conservation status of Salvia taxa collected from different populations is assessed at regional, national and global levels according to version 3.1 of the IUCN Red List categories and criteria (IUCN, 2001) and with reference to Gärdenfors (2001) and Gärdenfors et al. (2001). The proportion (%) of the global population of the taxa within the area was also provided. Details of the threats are determined for each taxa and comments are made in accordance with Broughton & McAdam (2002).

Definitions of IUCN threat categories adopted are as follows: (1) Critically Endangered (CR): A taxon is Critically Endangered when it has an extremely high risk of extinction in the wild, (2) Endangered (EN): A taxon is Endangered when it has a very high risk of extinction in the wild, (3) Vulnerable (VU): A taxon is Vulnerable when it has a high

risk of extinction in the wild in the medium term future, (4) Near Threatened (NT): A taxon is Near Threatened when it does not qualify Critically Endangered, Endangered or Vulnerable, whereas it is close to qualifying for or is likely to qualify for a threatened category in the near future, and (5) Least Concern (LC): A taxon is Least Concern when it does not qualify for Critically Endangered, Endangered or Vulnerable, but widespread and abundant taxa are included in this category.

2.8. Ecological Analysis

Ecological properties of the taxa studied were determined on the basis of soil samples taken around their growth environment at 0-30 cm in depth. Texture, organic matter, total salt, pH, CaCO3, K, N and P analyses of all soil samples were made using standard techniques (Bayraklı, 1987) at the Soil, Fertilizer and Water Resources Central Research Institute, Ankara. Soil structure findings were evaluated according to Kaçar (1972).

2.9. Numerical Taxonomic Methods

For morphometric analysis, the specimens stemming from own collections were selected as OTUs (Operational Taxonomic Units, the specimens). The basic units of comparison in numerical phenetics are termed as OTUs (Ward, 1993). The selection was undertaken according to well-preserved and dried, and mature specimens. In the field, we also noted some morphological characters, such as colour and length of calyx and corolla, which are hard to infer from the dried specimens. 60 diagnostic vegetative, reproductive, anatomical, palynological and mericarp characters were chosen and scored for each OTU. The raw data matrix is composed of binary (absent: 0, present: 1), quantitative and qualitative multistate variables.

For the multivariate analysis, a similarity matrix was created first using Gower‟s (1971) general coefficient similarity index (Sneath & Sokal, 1973), which allows the use of binary, alternative, qualitative, quantitative and semi-quantitative characters as well as

taking into account missing values, advantages not found together in other available similarity coefficients (St-Laurent et al., 2000). This similarity matrix was then clustered with the help of UPGMA (Unweighted Pair Group Method with Arithmetic Means) and the results are shown in the phenogram. UPGMA is the most frequently used method (Romesburg, 1984) and also appears to produce the best results (Radford, 1986) in terms of following criteria: accurate reflection of the similarity matrix, symmetrical hierarchical structure, and congruence with classification derived by traditional methods (Ward, 1993). The characters used in the analysis were assumed to be as important as each other and were unweighted. The MVSP software version 3.2 (a multivariate statistics package for IBM PC and compatibles) program package was used for the analysis (Kovach, 1999).

CHAPTER 3

RESULTS AND DISCUSSION

3.1. Gross Morphology

3.1.1. Life Form and Growth Habit

Salvia species are herbaceous often with a woody rootstock (e.g. S. blepharochleana, S. brachyantha, S. divaricata, S. eriophopra, S. frigida, S. macrosiphon, S. microstegia, S. multicaulis, S. russellii), shrubby or clearly woody at base (e.g. S. caespitosa, S. euphratica, S. huberi, S. hydrangea, S. pachystachys, S. rosifolia). Most species are perennials, however, the only one exception is S. viridis, which appears to be an annual. Several herbaceous plants (e.g. S. aethiopis, S. ceratophylla, S. sclarea) are also biennials or short-lived perennials. In general, the rooting system of mature shrubby or woody plants is composed of a deep and strong taproot with numerous lateral roots accounting for the majority of the underground biomass.

3.1.2. Stem

The stems are procumbent (e.g. S. caespitosa, S. hedgeana, S. macrochlamys, S. pinnata), ascending-erect (e.g. S. blepharochleana, S. bracteata, S. dicroantha, S. huberi, S. pilifera, S. rosifolia, S. suffruticosa, S. verticillata) or erect (e.g. S. aethiopis, S. ballsiana, S. divaricata, S. glutinosa, S. limbata, S. palaestina, S. recognita, S. sclarea, S. staminea, S. virgata) (Figure 6).

Herbaceous species are mostly erect while shrubby species are mainly ascending to erect. Whereas herbaceous species (e.g. S. indica, S. recognita) grow up to 180 cm tall (Figure 7A), dwarf shrubby species (e.g. S. caespitosa, S hedgeana, S. pachstachys) can be from 5 to 40 cm (Figure 6B). 27

The stems are solitary (e.g. S. frigida, S. longipedicellata, S. macrosiphon, S. spinosa), several (e.g. S. atropatana, S. eriophora, S. microstegia) or many (e.g. S. macrochlamys, S. russellii, S. suffruticosa, S. verticillata). Branched stems are found almost all species, but a few species, such as S. huberi, S. hypargeia, S. montbretii and S. rosifolia, have often unbranched stems.

Figure 6. Stem types in Salvia. A. S. macrochlamys, with procumbent stems; B. S. suffruticosa, with ascending-erect stems; C. S. ballsiana, with erect stems; D. S. virgata, with erect stems.

Figure 7. A. S. indica, a perennial herbaceous species growing up to 1.5 m high; B. S. hedgeana, a dwarf perennial suffruticose species barely reaching 17 cm high.

3.1.3. Leaves

The lamina of fully developed leaves is always petiolate, but in S. hypargeia (Figure 8A) and S. montbretti petioles are also indistinct. S. ceratophylla has also subabsent petioles (Figure 8C). The leaves are mostly thick-textured and occasionally membranous or submembranous leaves as in S. bracteata, S. pilifera (Figure 8B), S. pinnata and S. rosifolia.

The leaves are simple, pinnatisect, trifoliate, pinnatified or lyrate in shape. The simple leaves are observed in most Salvia species (e.g. S. cilicica, S. euphratica, S. hypargeia, S. indica, S. microstegia, S. russellii, S. tomentosa, S. viridis, S. virgata) (Figure 8A, E, F). S. blepharochleana, S. bracteata, S. huberi, S. hydrangea, S. pilifera, S. pinnata, S. rosifolia, S. suffruticosa, S. trichoclada have always pinnatisect leaves (Figure 8B). S. hedgeana is characterized by its trifoliate leaves. Pinnatified leaves are found only in S. ceratophylla (Figure 8C). Lyrate leaves are sometimes found in S. palaestina (Figure 8D), S. verticillata and S. russellii. 29

Figure 8. Leaf characteristics in Salvia. A. S. hypargeia, with simple, linear to linear- oblong leaves, subentire margins and indistinct petioles; B. S. pilifera, with pinnatisect, oblong-ovate and submembranous leaves; C. S. ceratophylla, with pinnatified leaves and subsessile petioles; D. S. palaestina, with lyrate, oblong-ovate leaves and erose margins; E. S. microstegia, with simple, oblong-ovate leaves and serrate margins; F. S. cilicica, with simple, orbicular leaves.

The leaves vary in their outline from linear to orbicular. Many species have oblong to ovate leaves (e.g. S. macrochlamys, S. microstegia, S. palaestina, S. pilifera S. poculata, S. sclarea, S. verticillata, S. virgata) (Figure 8B, D, E). Linear or linear-oblong leaves are found in S. atropatana, S. eriophora, S. hypargeia (Figure 8A), S. montbretii and S. rusellii. The leaves of S. brachyantha and S. russellii vary from linear-oblong to ovate. Suborbicular or orbicular leaves are found in S. cilicica (Figure 8F), S. cyanescens and S. kurdica.

The leaves vary considerably in margins both within and between species of Salvia such as serrate, serrulate, erose, crenate, crenulate, subentire and entire (Figure 8). The most common margin type is crenulate as in S. eriophora, S. euphratica, S. multicaulis, S. viridis and S. dicroantha.

3.1.4. Inflorescence

The inflorescence of Salvia is variously arranged into oppositely reduced cymes, with one to many-flowered interrupted whorls (verticillasters). It matures acropetally and is branched or unbranched. For instance, the inflorescence of S. divaricata is an elongate and widely branched paniculate, but S. hedgeana and S. pachystachys have unbranched inflorescences.

The number of flowers per verticillaster is of taxonomic significance. Some species, such as S. kurdica and S. macrochlamys (Figure 9A), appear always to have one or two flowers per verticillaster, whereas in most species, the number of flowers reaches 10 or more. The number of flowers per verticillaster of S. verticillata can be up to 40 (Figure 9B). The density of the raceme is also important as a character. In some species, such as S. macrochlamys, verticillasters are approximating, while in majority, they are clearly distant.

Figure 9. The number of flowers per verticillaster in inflorescences in Salvia. A. S. macrochlamys, with two flowers in each verticillaster; B. S. verticillata, with flowers up to 40 flowers in each verticillaster.

3.1.5. Bracts

Bracts are present in all species except for S. divaricata. Bract size, shape and colour have little diagnostic value at taxonomic level since they are sometimes quite variable within the same species. Often the bracts can be nearly as long as or shorter than calyces or nearly as long as calyces. S. macrochlamys, S. palaestina and S. sclarea possess very large bracts apparently exceeding calyces. The bracts range in color from green, gray, violoet, pink to purple, depending on the species. In shape they are usually ovate, ovate- oblong or oblong, seldom obovate or ovate-elliptic and have short or long, acuminate apices. The largest bracts are produced at the base of the inflorescence, where they are sometimes transitional with foliage leaves. The bracts are deciduous or persistent. Bracteoles are present in many species, however absent in several species, such as S. ballsiana, S. cerino-pruinosa, S. macrosiphon, S. montbretii and S. spinosa (Figure 10).

3.1.6. Calyx

The calyces of all species are persistent, and previous year‟s calyces may be found on dried inflorescences adjacent to flowering inflorescences. The calyx characters, such as 32

size, shape and texture, can be used as the basis for infrageneric classification and to distinguish species. In shape the calyx is campanulate (e.g. S. caespitosa, S. huberi, S. pachystachys, S. rosifolia), infundibular (e.g. S. blepharochleana, S. pilifera) or tubular (e.g. S. palaestina, S. spinosa, S. syriaca, S. tomentosa, S. viridis), and bilabiate (Figure 10).

The calyx shows little or considerable growth or change in shape after anthesis. The fruiting calyces are membranous only in sect. Hymenosphace, but they are is thick- textured in the other sections (Figure 10).

The upper lip of the calyx is tri-lobed or obsoletely so or subentire and the lower is two- lobed. The apices of the calyx lobe are sometimes rigid and spine-tipped or not (Figure 10).

3.1.7. Corolla

The corollas of all species are tubular and bilabiate. They vary in colour from white to pink, blue, violet and yellow. Because of their size variation, full developed flowers can provide reliable morphological characters by which to classify and identify species. In some species, such as S. blepharochleana and S. kronenburgii, the corollas can be up to 50 mm long. Some species, such as S. russelli and S. verticillata, have small corollas barely reaching 15 cm long (Figure 10).

Length of upper lip of the corolla relative to the lower lip (labellum) and size of the tube vary significantly among species. The upper lip of the corolla is straight (e.g. S. absconditiflora, S. macrochlamys, S. multicaulis, S. russellii, S. suffruticosa) or falcate (hook-shaped) (e.g. S. glutinosa, S. hypargeia, S. montbretii, S. palaestina, S. spinosa, S. virgata). The tube is straight or curved, ventricose or not, invaginated and with a small internal scale (squamula) or not, annulate (hairy ring) or not (Figure 10).

Figure 10. Variation in bract, calyx and corolla morphology in Salvia. A. S. macrochlamys; B. S. caespitosa; C. S. euphratica var. leiocalycina; D. S.kronenburgii; E. S. palaestina; F. S. indica; G. S. ceratophylla; H. S. glutinosa; I. S. staminea; J. S. russellii. 34

3.1.8. Stamens

Staminal morphology has been commonly figured in infrageneric taxonomy of Salvia (e.g. Bentham, 1848; Hildebrand, 1865; Epling, 1939; El-Gazzar et al., 1968; Claβen- Bockhoff et al., 2003, 2004a, b; Hedge, 1972, 1974, 1982a, b; Reith et al., 2007; Walker & Sytsma, 2007). El-Gazzar et al. (1968) identified four major stamen morphologies in Salvia. These were defined largely by the relative lengths of the two portions of the connective and whether the portion posterior to the filament is sterile. Walker & Sytsma (2007) recognized fifteen distinct stamen types (Figure 2; stamen types A to O) in various lineages of Salvia and the related genera.

Based on the staminal morphology of Salvia, three distinct stamen types (Figure 11; stamen types A, B and C) are identified in species of Salvia in Turkey (Hedge (1982a). In the stamen type A, upper theca is larger than the poorly developed lower theca which bears at least some pollen and staminal connectives are shorter than or nearly equal to filaments. This stamen type is found in species belonging to sec. Salvia and Hymenosphace. In the stamen type B (the most common type), staminal connectives are prominently longer than filaments, the lower theca is reduced to a usually dolabriform plate and sterile, and stamens are articulating. Species of sect. Aethiopis, Drymosphace, Horminum and Plethiosphace possess this stamen type. In the stamen type C, staminal connectives are longer than filaments, the lower theca is subulate and sterile, and stamens are not articulating. This stamen type is found in members of sect. Hemisphace. Stamens are included within upper lip of corolla in all species except for S. staminea, with long exserted stamens beyond upper lip of the corolla (Figure 10I). The stamen properties can be utilized in the taxonomy of the genus at the infrageneric level.

Figure 11. Representations of the stamen types in Salvia species in study area (Hedge 1982a). A: Stamen type A, B: Stamen type B, C: Stamen type C. c: connective, db: dolabriform sterile tissue; f: filament.

3.1.9. Style

The style is variable in length, glabrous, is slightly or long exserted from corolla lips and divided in two parts at apex.

3.2. Anatomy of the Vegetative Organs

Anatomy, or the internal form and structure of plant organs, is a significant source of data used in plant classification. The utility of anatomy and its implication in the systematics of various genera of Lamiaceae are well known (e.g. Bokhari & Hedge, 1971; Bokhari & hedge, 1976; Metcalfe & Chalk, 1950; Kahraman et al. 2010; Kahraman et al., 2009c). However, a thorough anatomical study of the genus Salvia is lacking to date. There is rathet limited published information on anatomy of the vegetative organs of species of Salvia (Özdemir & Şenel, 1999; Kaya et al., 2007; Bagherpour et al., 2010; Kahraman & Doğan, 2010; Kahraman et al., 2009b, c, 2010b, c, d) and much of what is available comprising assorted data for individual species.

The first comprehensive evaluation of the systematic value of root, stem, leaf blade and petiole anatomy of Salvia is presented here. The main anatomical characters of the taxa studied are summarized in Tables 4-8. LM micrographs of transverse sections of roots, stems, leaf blades and petioles are illustrated in Figures 12-37.

3.2.1. Root Anatomy (Figures 12-18, Table 4)

The outermost surface of the roots is coated by a thin or thick periderm of which cells are dark colored, crushed, break up or sometimes fall out. Beneath the periderm, the cortex is relatively large, with multi-layered layers of large parenchymatous cells which are heterogeneous in shape and size. There are several or many groups of sclerenchymatous fibers above the phloem. Phloem and xylem elements can be distinguished in the vascular tissue. The vascular cambium, originating between the xylem and the phloem, is sometimes inconspicuous. Among the taxa examined, the thickness of the phloem varies between 50 µm and 200 µm while the thickness of the xylem varies between 300 µm to 3500 µm. The xylem consists of vessel members and tracheids. The vessel members are oval, round or polygonal and their diameter ranges from 10 µm to 130 µm. The tracheids range from 4 µm to 25 µm in diameter. The xylem ray cells are composed of 1 to 40 radial rows of square to rectangular parenchymatous

cells. In the taxa of the sections Salvia and Hymenosphace the maximum number of rays is 6 (7 only in S. pseudeuphratica), but in the taxa of the others the maximum is 8 or more. They range from 8 µm to 90 µm in length and from 3 µm to 30 µm width. In the roots of some taxa, there is no pith as the xylem fills all the central portion of the root. Some taxa have a small central pith comprising polygonal, round or oval parenchymatous cells since the xylem occupies much of the pith. These cells vary between 15 µm and 150 µm.

Figure 12. Cross section of the root of S. atropatana. ca: cambium, cor: cortex, pe: periderm, ph: phloem sc: sclerenchyma, x: xylem, xr: xylem rays.

Figure 13. Cross section of the root of S. hedgeana.

Figure 14. Cross section of the root of S. pachystachys.

Figure 15. Cross section of the root of S. viridis.

Figure 16. Cross section of the root of S. ceratophylla.

Figure 17. Cross section of the root of S. dicroantha.

Figure 18. Cross section of the root of S. verticillata subsp. amasiaca. 41

Table 4. Anatomical characteristics of roots in the Salvia taxa studied. All sizes are in µm. Numbers refer to minimum-maximum (mean). Taxa and sections Ray characteristics Size of phloem Size of xylem Diameter of Diameter of Diameter of pith tissue tissue vessels tracheids cells Row Length Width number Sect. Salvia S. anatolica 1-3 12-25 (18) 6-10 (8) 140-180 (165) 1050-1250 (1125) 20-45 (30) 6-14 (9) 35-90 (60) S. ballsiana 1-3 (-4) 15-40 (25) 8-20 (15) 50-80 (70) 400-600 (480) 20-110 (50) 8-18 (12) 25-90 (55) S. bracteata 1-3 15-35 (25) 8-18 (13) 50-70 (60) 1200-1400 (1300) 30-90 (60) 7-17 (12) 30-100 (60) S. caespitosa 1-2 13-26 (20) 4-10 (7) - 1400-1600 (1500) 15-60 (40) 6-12 (8) - S. divaricata 1-2 (-3) 12-28 (19) 6-15 (12) 70-120 (95) 500-650 (560) 30-55 (40) 7-20 (12) 50-120 (85) S. hedgeana 1-2 10-20 (15) 4-8 (6) 140-200 (175) 1500-2000 (1750) 15-85 (40) 5-10 (8) - S. huberi 1-2 10-25 (18) 4-9 (7) 50-70 (60) 900-1100 (1000) 15-80 (45) 5-17 (11) - S. macrochlamys 1-4 (-6) 9-40 (25) 3-16 (9) - - 20-90 (50) 8-20 (15) -

42 S. pachystachys 1-2 10-25 (15) 4-11 (8) - 2500-3000 (2750) 20-70 (45) 5-15 (10) -

S. pilifera 1-4 10-30 (20) 5-17 (11) 70-100 (80) 680-800 (750) 20-70 (40) 4-10 (7) 30-120 (75) S. pinnata 1-6 15-30 (22) 6-13 (10) 70-120 (90) 900-1150 (1050) 15-60 (35) 6-15 (10) 30-110 (70) S. recognita 1-5 15-35 (25) 8-20 (14) 100-150 (120) 1600-2200 (1900) 25-120 (60) 5-15 (10) 30-140 (85) S. rosifolia 1-2 13-30 (22) 5-11 (8) 100-130 (110) 800-1150 (1000) 20-95 (50) 4-11 (7) S. suffruticosa 1-2 12-30 (20) 6-15 (10) 60-90 (70) 400-580 (500) 20-100 (50) 6-12 (9) 25-95 (60) S. tomentosa 1-2 10-22 (15) 6-12 (9) 100-160 (130) 850-1020 (950) 15-70 (40) 5-17 (10) 30-120 (70) S. trichocloda 1-3 12-23 (17) 7-20 (12) 50-80 (65) 1200-1350 (1275) 20-60 (45) 5-13 (10) 30-75 (50) Sect. Hymenosphace S. absconditiflora 1-3 (-4) 10-20 (15) 4-10 (6) 50-80 (65) 400-500 (450) 12-40 (25) 4-10 (7) 25-80 (50) S. blepharochleana 1-2 10-19 (15) 4-11 (7) - - 25-80 (50) 4-10 (7) - S. cerino-pruinosa 1-2 10-20 (15) 4-10 (6) - - 20-65 (40) 6-13 (9) 20-80 (50) S. euphratica var. 1-3 (-4) 8-20 (15) 8-18 (14) 50-70 (60) 800-1000 (900) 20-80 (50) 5-15 (10) 20-100 (60) euphratica S. euphratica var. 1-3 (-4) 10-25 (17) 5-15 (10) 60-80 (70) 950-1200 (1050) 20-85 (50) 5-15 (10) 20-95 (55) leiocalycina S. hydrangea 1-2 8-20 (14) 5-10 (8) 65-100 (80) 550-700 (625) 20-80 (45) 20-70 (45) S. kronenburgii 1-2 10-35 (22) 5-12 (8) 95-130 (110) 825-930 (880) 18-85 (50) 5-15 (10) 20-85 (55)

Table 4 (continued) S. multicaulis 1-3 9-20 (15) 4-10 (6) 60-80 (70) 300-400 (350) 10-35 (20) 3-9 (6) 20-90 (50) S. pseudeuphratica 1-6 (-7) 15-25 (20) 5-14 (10) - 900-1000 (950) 20-100 (50) 5-15 (10) - Sect. Horminum S. viridis 1-20 12-35 (22) 5-15 (10) 50-70 (60) 1200-1450 (1350) 15-85 (50) 5-20 (11) - Sect. Aethiopis S. aethiopis 1-8 20-80 (45) 8-18 (13) - - 15-80 (45) 5-20 (12) -

S. atropatana 1-12 15-75 (40) 7-20 (13) 120-140 (110) 1700-1900 (1800) 20-90 (50) 5-20 (12) - S. brachyantha 1-8 20-60 (40) 8-20 (14) - 800-900 (8509 15-55 (35) 5-18 (11) - S. candidissima subsp. 1-15 15-35 (25) 8-12 (10) - 3000-3500 (3250) 25-130 (75) 5-20 (12) - candidissima S. ceratophylla 2-40 15-50 (30) 8-20 (14) - - 20 60 (40) 6-20 (13) - S. cilicica 3-20 20-60 (40) 10-20 (15) - 900-1300 (1100) 25-75 (50) 5-17 (12) 25-85 (55) S. cyanescens 1-10 15-40 (25) 8-15 (12) 80-100 (90) - 20-80 (50) 5-15 (10) - S. eriophora 1-10 20-50 (35) 7-18 (13) - - 20-70 (45) 5-20 (13) -

4 S. frigida 1-8 15-33 (25) 9-18 (13) 75-85 (80) 500-550 (525) 15-40 (25) 5-12 (8) 15-55 (35)

S. hypargeia 1-10 15-30 (50) 7-15 (11) 90-100 (95) - 15-50 (35) 5-12 (98) - S. limbata 2-8 15-40 (25) 8-20 (14) - 1150-1350 (1250) 20-70 (50) 5-25 (15) 25-130 S. longipedicellata 1-9 25-70 (50) 9-23 (16) - 950-1200 (1100) 30-130 (70) 5-20 (12) - S. montbretii (1-) 3-15 20-50 (35) 12-25 (18) - 1800-2400 20-90 (50) 6-15 (10) - (21000) S. palaestina 1-8 (-10) 15-70 (45) 8-30 (18) - - 20-65 (45) 5-25 (15) - S. poculata 1-10 20-50 (35) 8-15 (11) - - 15-65 (40) 5-20 (12) - S. sclarea 1-10 20-90 (50) 8-20 (14) - - 40-110 (75) 5-20 (12) - S. siirtica 1-12 20-50 (35) 10-18 (14) - - 20-60 (40) 5-15 (10) - S. xanthocheila 1-11 20-60 (40) 8-16 (12) - - 20-60 (45) 5-15 (10) - Sect. Drymosphace S. glutinosa 1-30 15-35 (25) 7-20 (13) - 1300-1600 (1450) 20-80 5-20 (12) - Sect. Plethiosphace S. dicroantha 1-12 12-45 (30) 5-18 (12) - 700-900 (800) 15-60 (35) 5-24 (15) 20-70 (45) S. nemorosa 1-8 13-55 (33) 7-17 (12) - 750-1050 (900) 15-60 (35) 5-17 (12) 20- 65 (40) S. staminea (1-) 3-8 20-40 (30) 9-20 (15) - - 20-70 (45) 5-20 (12) -

Table 4 (continued) S. virgata 1-8 15-50 (30) 9-17 (13) - - 20-55 (35) 5-15 (10) 20-60 (40) Sect. Hemisphace S. russellii 1-10 12-50 (30) 10-20 (15) 70-90 (080) 1300-1900 (1600) 25-75 (50) 5-20 (12) 25-100 (60) S. verticillata subsp. (1-) 3-15 20-70 (45) 15-30 (23) 95-120 (110) 1400-1750 (1575) 15-50 (35) 8-23 (15) 30-150 (80) amasiaca S. verticillata subsp. 1-7 15-50 (35) 10-24 (16) - - 15-45 (32) 8-20 (14) - verticillata

3.2.2. Stem Anatomy (Figure 19-24, Table 5)

In cross section, the stems are between 1200 µm and 7000 µm in diameter and mostly quadrangular to more or less circular or six-sided in one species S. hedgeana. The epidermis consists of a single layer of almost square, rectangular or oval, compactly arranged cells. The upper and lower walls of the epidermis are thicker than the lateral walls. The upper surface of the epidermis is covered by a thin or thick cuticule. On the epidermis, there are peltate glandular, capitate glandular or non-glandular trichomes. Under the epidermis, 1-9 layers (20-250 µm thick) of collenchymatous cells are found at the corners. The collenchyma tissue is narrow in some species, such as S. divaricata, S. caespitosa, S. hedgeana, S. huberi, S. pachystachys and S. rosifolia. However, some taxa, such as S. candidissima subsp. candididssima, S. eriophora and S. syriaca, has a wide collenchyma tissue. There are 1-4 layers of chlorenchymatous cells situated between the corners. In S. anatolica, S. caespitosa, S. rosifolia and S. tomentosa have only one layer of chlorenchymatous cells. The cortex contains of 1-15 layers (15-650 µm thick) of irregularly oval, round or polyhedral parenchymatous cells, with or without intercellular spaces. Some species, such as S. ballsiana, S. divaricata, S. huberi, S. pachystachys, S. rosifolia, S. absconditiflora, S. viridis, S. atropatana, S. ceratophylla, S. hypargeia and S. russellii, has a narrow cortex. In some taxa, such as S. euphratica var. leiocalycina, S. candidissima subsp. candidissima, S. montbretii, S. syriaca, have a very large cortex. Vascular cambium forms xylem inward and phloem outward. Several or many scattered groups of sclerenchymatous fibers are concentrated outside the phloem consisting of several layers of irregular cells. The vascular bundles at the corners (150- 900 µm) are larger than those between the corners (60-800 µm). The pith is 600 to 6000 µm and comprises thin walled polygonal or circular parenchymatous cells, usually with abundant intercellular spaces.

Figure 19. Cross section of the stem of S. divaricata. chl: chlorenchyma, col: collenchyma, cor: cortex, ep: epidermis, ph: phloem, pi: pith.

Figure 20. Cross section of the stem of S. caespitosa.

Figure 21. Cross section of the stem of S. cerino-pruinosa.

Figure 22. Cross section of the stem of S. virgata.

Figure 23. Cross section of the stem of S. blepharochleana.

Figure 24. Cross section of the stem of S. pachystachys.

Table 5. Anatomical characteristics of stems in the taxa studied of Salvia. All sizes are in µm. Numbers refer to minimum-maximum (mean). SD = Stem diameter, SC = Stem contour, COL = Thickness of collenchyma, CH = Thickness of the chlorenchyma, COR = Thickness of the cortex, PI = pith region, VBC = Thickness of vascular bundles at the corners, VBB = Thickness of vascular bundles between the corners. Taxa and sections SD SC COL CH COR VBC VBB PI Sect. Salvia S. anatolica 1900-2300 (2100) Quadrangular 110-140 (120) 1 200-310 (250) 300-350 (325) 250-320 (300) 800-950 (850) S. ballsiana 3000-3100 (3150) Quadrangular to 80-100 (90) 2-4 15-80 (50) 230-280 (250) 70-110 (90) 2600-2700 (2650) more or less circular S. bracteata 1500-1900 (1700) Quadrangular 70-90 (80) 1-2 80-150 (120) 200-350 (250) 100-200 (150) 800-1200 (1000) S. caespitosa 1500-1900 (1750) Quadrangular 40-60 (50) 1 100-180 (150) 170-200 (185) 100-180 (160) 1100-1200 (1150) S. divaricata 2000-3000 (2500) Quadrangular 55-75 (65) 1-2 35-80 (55) 300-500 (400) 200-350 (290) 1500-2500 (2000) S. hedgeana 1700-3000 (2250) Hexangular / 30-60 (50) - 120-300 (225) 150-300 (230) 100-250 (200) 750-1200 (1000) Quadrangular S. huberi 1850-2000 (1950) Quadrangular 30-50 (40) 1-2 30-100 (50) 300-350 (330) 210-280 (250) 1200-1300 (1250) S. macrochlamys 4000-5000 (4500) Quadrangular 60-80 (70) 1-2 70-250 (150) 750-900 (825) 600-800 (700) 2800-3000 (2800)

8 S. pachystachys 1400-1600 (1500) Quadrangular 30-60 (45) 1-2 40-80 (50) 190-220 (210) 150-190 (170) 850-950 (900)

S. pilifera 3500-4500 (4000) Quadrangular 100-140 (120) 2 100-200 (150) 350-400 (380) 200-250 (220) 2500-3000 (2750) S. pinnata 2800-3100 (3000) Quadrangular 140-170 (150) 1-2 110-170 (140) 420-500 (350) 250-350 (300) 1850-1950 (1900) S. recognita 2250-2800 (2500) Quadrangular 70-100 (85) 2-3 35-90 (65) 300-350 (325) 180-240 (210) 1600-1800 (1700) S. rosifolia 1900-2070 (2000) Quadrangular 30-50 (40) 1 30-65 (50) 300-350 (330) 250-300 (280) 1250-1400 (1320) S. suffruticosa 2500-2900 (2700) Quadrangular 120-160 (150) 2-3 170-350 (250) 350-430 (390) 200-280 (230) 1500-1600 (1550) S. tomentosa 1950-2150 (2050) Quadrangular 80-120 (100) 1 - 270-290 (280) 250-275 (265) 1000-1100 (1050) S. trichocloda 1700-2500 (2000) Quadrangular 90-150 (120 1-2 100-180 (140) 250-400 (275) 150-250 (200) 1000-1350 (1250) Sect. Hymenosphace S. absconditiflora 1700-2000 (1850) Quadrangular 90-120 (105) 1-2 30-70 (50) 200-250 (220) 120-170 (140) 1200-1500 (1350) S. blepharochleana 3000-3300 (3150) Quadrangular to 65-90 (75) 3-4 190-290 (250) 180-230 (205) 100-175 (130) 2050-2475 (2250) more or less circular S. cerino-pruinosa 2600-2900 (2750) Quadrangular to 130-150 (140) 2-4 150-250 (200) 320-340 (330) 280-315 (300) 1300-1525 (1450) more or less circular

Table 5 (continued) S. euphratica var. 2300-2700 (2500) Quadrangular 90-120 (105) 1-2 170-300 (240) 250-325 (290) 150-210 (180) 1600-1800 (1700) euphratica S. euphratica var. 2150-2600 (2400) Quadrangular 80-105 (95) - 280-500 (370) 310-330 (320) 310-330 (320) 600-650 (625) leiocalycina S. hydrangea ------S. kronenburgii 2000-3000 (2500) Quadrangular 100-120 (110) 1-2 140-260 (200) 250-350 (300) 150-250 (200) 1400-2150 (1800) S. multicaulis 1600-1900 (1750) Quadrangular 65-85 (75) 1-2 70-170 (110) 180-210 (200) 100-170 (130) 1100-1350 (1250) S. pseudeuphratica 1200-1500 (1350) Quadrangular 50-70 (60) 1-2 50-160 (100) 150-200 (180) 90-130 (110) 700-800 (750) Sect. Horminum S. viridis 2300-2500 (2400) Quadrangular 20-60 (40) 2-3 15-100 (60) 350-400 (375) 100-200 (150) 1750-1850 (1800) Sect. Aethiopis S. aethiopis 5000-7000 (6000) Quadrangular 110-150 (130) 2-3 100-300 (200) 400-670 (550) 180-300 (250) 4000-6000 (5000) S. atropatana 3500-3700 (3600) Quadrangular 80-100 (90) 3 50-100 (70) 350-500 (425) 130-220 (170) 2800-3000 (2900) S. brachyantha 2850-3200 (3000) Quadrangular 140-170 (155) 1-2 110-280 (200) 300-400 (350) 120-250 (180) 1850-2200 (2000) S. candidissima 4000-6500 (5250) Quadrangular 130-185 (160) 2-3 280-400 (330) 350-500 (440) 110-300 (210) 3000-5300 (4500) subsp. candidissima

4 S. ceratophylla 2700-4500 (3750) Quadrangular 65-100 (80) 2-3 55-90 (75) 300-520 (420) 90-250 (160) 2200-4000 (3250)

S. cilicica 2500-3500 (3000) Quadrangular - 2-3 100-220 (160) 350-470 (410) 100-200 (150) 1850-2900 (2400) S. cyanescens 3000-3500 (3250) Quadrangular 80-120 (100) - 100-200 (150) 400-500 (450) 150-300 (225) 2300-2850 (2600) S. eriophora 3300-3600 (3450) Quadrangular 170-200 (185) - 150-320 (240) 400-530 (470) 180-400 (280) 2200-2500 (2350) S. frigida 2300-2650 (2500) Quadrangular 90-110 (100) 1-2 75-150 (105) 250-300 (275) 100-200 (150) 1650-1950 (1800) S. hypargeia 2200-2650 (2400) Quadrangular 70-90 (80) 2-3 30-110 (60) 250-300 (275) 120-230 (170) 1750-2000 (1850) S. microstegia 2900-3300 (3100) Quadrangular 115-135 (125) 2-3 90-220 (150) 300-450 (375) 90-200 (145) 2050-2350 (2200) S. montbretii 2700-3200 (2900) Quadrangular 70-100 (85) 2-3 220-400 (310) 325-450 (380) 150-300 (220) 1250-1750 (1500) S. sclarea 3000-3500 (3250) Quadrangular 80-120 (100) 2-3 100-220 (190) 400-450 (425) 150-300 (230) 1800-2250 (2000) S. siirtica 2000-2600 (2250) Quadrangular 60-120 (90) 1-3 60-150 (110) 250-400 (225) 100-210 (160) 1300-2050 (1650) S. syriaca 3000-4500 (3750) Quadrangular 160-250 1-2 250-650 (425) 300-850 (650) 250-700 (350) 1000-1900 (1500) S. xanthocheila 3500-4000 (3750) Quadrangular - 1-2 4-8 300-600 (500) 300-600 (500) 2100-2700 (2400) 110-250 (180) Sect. Drymosphace S. glutinosa 3000-3500 (3250) Quadrangular 120-150 (135) 1-3 50-200 (120) 370-430 (400) 100-200 (150) - Sect. Plethiosphace

Table 5 (continued) S. dicroantha 1650-1950 (1800) Quadrangular 75-100 (90) 2-3 40-120 (80) 200-240 (220) 90-180 (140) 1200-1300 (1250) S. nemorosa 2300-2500 (2400) Quadrangular 120-140 (130) 3 70-240 (150) 300-350 (325) 100-220 (160) 1450-1650 (1550) S. staminea 3000-3500 (3250) Quadrangular 80-160 2-4 125-350 (240) 350-450 (400) 150-375 (250) 1700-2200 (2000) S. virgata 2600-3000 (2800) Quadrangular 90-120 1-5 20-150 (85) 225-275 (250) 60-160 (110) 1900-2000 (1950) Sect. Hemisphace S. russellii 1500-1750 (1650) Quadrangular 70-100 (85) 2-4 25-85 (55) 200-275 (240) 100-180 (140) 1000-1100 (1050) S. verticillata subsp. 3500-4000 (3750) Quadrangular 150-190 (170) 2-3 120-275 (200) 650-800 (725) 350-550 (450) 2700-3000 (2850) amasiaca S. verticillata subsp. 3700-4700 (4100) Quadrangular 150-180 (165) 2-3 120-260 (190) 600-700 (650) 300-480 (380) 2600-3300 (2900) verticillata

3.2.3. Leaf Blade Anatomy (Figures 25-30, Tables 6-7)

The leaf blades (75-350 µm thick) have an epidermis composed of one layer of oval, squarish or rectangular cells and covered by a thin or thick cuticle (up to 18 µm). The blades of S. anatolica, S. ballsiana, S. pilifera, S. caespitosa, S. pachystachys, S. huberi, S. rosifolia, S. blepharochleana, S. hydrangea and S. syriaca have a thicker cuticle than 5 µm, however, the other taxa have a thinner cuticle. Peltate glandular, capitate glandular or non-glandular trichomes are present on both sides of the blade.

The leaf blades are amphistomatic or hypostomatic, with diacytic stomata. The blades of all taxa are amphistomatic except the leaf blades of S. glutinosa which are hypostomatic. In the amphistomatic taxa, stomata are more frequent on the abaxial surface of the blades. The upper epidermal cells are larger than the lower epidermal cells (e.g. S. pilifera, S. tometosa, S. multicaulis, S. viridis, S.aethiopis, S. hypargeia, S. virgata, S. russellii) or nearly equal to them (S. bracteata, S. macrochlamys, S. blepharochleana, S. ceratophylla, S. palaestina, S. glutinosa). Among the studied taxa, the upper epidermal cells are 8-60 µm in length and 10-100 µm in width, whereas the lower epidermal cells are 4-45 µm in length and 7-80 µm in width. The upper and lower walls of the epidermal cells were thicker than the lateral walls. Most taxa of the section Savia and sect. Hymenosphace present hypodermis with one layer of large, polyhedral-isodiametric or rectangular cells under both epidermis. In the taxa of the section Hemisphace, there is a monolayer hypodermis often becoming palisade-like under the upper epidermis. All taxa of the sections Horminum, Aethiopis, Drymosphace and Plethiosphace do not present hypodermis.

The mesophyll is isobilateral (e.g. S. bracteata, S. caespitosa, S. pachystachys, S. euphratica var. euphratica, S. euphratica var. leiocalycina, S. syriaca, S. virgata) or dorsiventral (e.g. S. pilifera, S. pinnata, S. absconditiflora, S. aethiopis, S. frigida, S. indica, S. sclarea, S. russelii, S. verticillata subsp. amasiaca, S. verticillata subsp. verticillata). In the isobilateral leaves, the mesophyll is undifferentiated into palisade and spongy tissues and two to six layers of the palisade parenchyma is found on both

sides of the blade. In the dorsiventral leaves, the mesophyll is differentiated into palisade and spongy tissues and consists of one to four layers of the palisade parenchyma followed by one to three layers of the spongy parenchyma. The palisade parenchyma is 10-90 µm in length and 4-23 µm in width.

The abaxial side of the midrib is mostly flat to slightly convex, strongly convex in S. ballsiana or slightly concave in S. aethiopis, while it is convex on the adaxial side. The width of the midrib ranges from 170 µm in S. caespitosa to 2200 µm in S. aethiopis and S. siirtica. The length of the midrib varies between 220 µm in S. caespitosa and 2150 µm in S. palaestina. The midrib has an epidermis with one layer of rectangular or oval cells. Collenchymatous cells have one to two layers below the abaxial epidermis and one to three below the adaxial epidermis.

In all taxa of sect. Salvia, Hymenosphace, Drymosphace, Hemisphace there is a large central vascular bundle, but S. viridis in sect. Horminum, most taxa of sect. Aethiopis and Plethiosphace have two to seven central vascular bundles. The central vascular bundle(s) is/are open U-shaped sometimes with incurved ends in all taxa except S. montbretii which are open crescent shaped. Some species, such as S. candidissima subsp. candidissima, S. cilicica, S. cyanescens, S. limbata, S. siirtica and S. virgata, have several small inverse bundles on the adaxial side. In many taxa there are no small accessory vascular bundles in the midrib. S. ballsiana, S. macrochlamys, S. pinnata in sect. Salvia, S. viridis in sect. Horminum, S. nemorosa, S. staminea, S. verticillata subsp. verticillata in sect. Hemisipahce and most species of sect. Aethiopis contain one or two small accessory vascular bundles in each side of the midrib.

Figure 25. Cross section of the leaf blade of S. ballsiana. cu: cuticle, hy: hypodermis, le: lower epidermis, mid: midrib, mv: median vascular bundle, pp: palisade parenchyma, ue: upper epidermis.

Figure 26. Cross section of the leaf blade of S. indica. ab. abaxial side, ad: adaxial side, le: lower epidermis, mid: midrib, mv: median vascular bundle, pp: palisade parenchyma, sp: spongy parenchyma, ue: upper epidermis, vb: vascular bundle.

Figure 27. Cross section of the leaf blade of S. verticillata subsp. verticillata. eg: eglandular hairs, hyp: hypodermis, le: lower epidermis, lv. lateral vascular bundle, mid: midrib, mv: median vascular bundle, pp: palisade parenchyma, sp: spongy parenchyma, ue: upper epidermis.

Figure 28. Cross section of the leaf blade of S. macrochlamys. gh: glandular hairs, hyp: hypodermis, le: lower epidermis, lv. lateral vascular bundle, mv: median vascular bundle, pp: palisade parenchyma, ue: upper epidermis.

Figure 29. Cross section of the leaf blade of S. aethiopis.

Figure 30. Cross section of the leaf blade of S. microstegia.

Table 6. Anatomical characteristics of leaf blades in the taxa studied of Salvia. All sizes are in µm. Numbers refer to minimum- maximum (mean). L: Length, W: Width. Taxa and sections Leaf Leaf type Size of upper Size of lower Comparison of Layers of Size of palisade Hypodermis thickness epidermal cells epidermal cells upper and palisade parenchyma (L x W) (L x W) lower parenchyma (L x W) epidermal cells Sect. Salvia S. anatolica 200-240 İsobilateral, 15-30 (21) x 20- 15-25 (20) x 20- Nearly equal 4-5 15-30 (22) x 9-15 Present (270) amphistomatic 60 (40) 50 (30) S. ballsiana 210-260 İsobilateral, 15-36 (23) x 20- 14-30 (19) x 17- Nearly equal 3-5 20-36 (27) x 7-15 Present (240) amphistomatic 55 (38) 60 (30) (11) S. bracteata 120-160 İsobilateral, 8-18 (13) x 15-45 8-15 (12) x 15-40 Nearly equal 2-3 20-45 (35) x 10-15 Present (140) amphistomatic (30) (27) (12) S. caespitosa 250-300 İsobilateral, 17-35 (25) x 25- 15-35 (25) x 18- Nearly equal 5-6 17-35 (25) x 9-16 Present (280) amphistomatic 60 (40) 45 (30) (13) S. divaricata 190-250 Dorsiventral, 15-35 (25) x 20- 10-20 (15) x 15- Larger upper 2-3 40-60 (48) x 8-15 Absent

56 (215) amphistomatic 55 (35) 35 (22) epidermal cells (11)

S. hedgeana 250-300 İsobilateral, 15-45 (30) x 20- 15-45 (30) x 15- Nearly equal 4-5 18-40 (25) x 9-16 Present (280) amphistomatic 65 (40) 55 (35) (12) S. huberi 280-350 İsobilateral, 15-30 (25) x 20- 15-25 (20) x 15- Nearly equal 4-6 15-35 (25) x 7-13 Present (320) amphistomatic 50 (35) 40 (27) (10) S. macrochlamys 210-250 İsobilateral l, 15-23 (20) x 15- 10-20 (16) x 15- Nearly equal 4-5 18-35 (25) x 7-13 Present (230) amphistomatic 40 (25) 40 (25) (10) S. pachystachys 200-250 İsobilateral, 15-25 (20) x 20- 15-27 (20) x 17- Nearly equal 4-5 15-35 (25) x 8-14 Present (230) amphistomatic 55 (40) 50 (35) (11) S. pilifera 200-250 Dorsiventral, 15-30 (20) x 15- 10-15 (13) x 10- Larger upper 3-4 30-60 (50) x 7-15 Absent (225) amphistomatic 45 (30) 30 (18) epidermal cells (11) S. pinnata 180-200 Dorsiventral, 10-32 (20) x 15- 8-20 (15) x 10-22 Larger upper 2-3 40-90 (70) x 11-17 Absent (190) amphistomatic 37 (25) (17) epidermal cells (14) S. recognita 200-250 Dorsiventral, 15-27 (21) x 25- 12-27 (19) x 20- Larger upper 2-3 20-50 (40) x 9-15 Absent (230) amphistomatic 65 (45) 45 (30) epidermal cells (12) S. rosifolia 235-285 İsobilateral, 14-27 (20) x 20- 10-25 (18) x 20- Nearly equal 4-5 20-45 (35) x 8-13 Present (260) amphistomatic 50 (35) 50 (30) (11)

Table 6 (continued) S. suffruticosa 240-300 İsobilateral, 10-32 (25) x 18- 10-25 (17) x 15- Nearly equal 3-4 25-60 (40) x 9-17 Present (265) amphistomatic 55 (35) 55 (30) (12) S. tomentosa 100-160 Dorsiventral, 10-20 (15) x 15- 8-15 (11) x 12-30 Larger upper 1-2 20-35 (30) x 6-14 Absent (125) amphistomatic 45 (30) (21) epidermal cells (10) S. trichocloda 200-260 İsobilateral, 10-25 (20) x 20- 10-20 (15) x 20- Nearly equal 3-4 20-40 (30) x 10-15 Present (240) amphistomatic 60 (40) 50 (35) (12) Sect. Hymenosphace S. absconditiflora 180-230 Dorsiventral, 14-25 (20) x 15- 10-20 (15) x 12- Larger upper 2-3 30-50 (40) x 7-12 Absent (200) amphistomatic 50 (35) 30 (20) epidermal cells (10) S. blepharochleana 230-270 İsobilateral, 10-20 (15) x 15- 9-18 (14) x 15-45 Nearly equal 3-4 25-40 (35) x 8-15 Present (250) amphistomatic 45 (30) (30) (12) S. cerino-pruinosa 210-250 İsobilateral, 10-20 (15) x 15- 8-20 (14) x 13-40 Nearly equal 3-4 25-50 (38) x 8-15 Present (240) amphistomatic 45 (30) (25) (12) S. euphratica var. 140-175 İsobilateral, 9-20 (15) x 15-45 5-18 (10) x 13-37 Larger upper 2-3 17-30 (25) x 5-10 Present euphratica (155) amphistomatic (30) (22) epidermal cells (8)

57 S. euphratica var. 95-140 İsobilateral, 11-25 (18( x 20- 9-20 (14) x 15-30 Larger upper 2-3 15-35 (25) x 5-10 Present

leiocalycina (120) amphistomatic 50 (35) (23) epidermal cells (7) S. hydrangea 200-250 İsobilateral, 10-22 (17) x 15- 10-20 (15) x 13- Nearly equal 4-5 18-40 (30) x 6-12 Present (225) amphistomatic 35 (25) 35 (25) (9) S. kronenburgii 180-230 İsobilateral, 8-18 (13) x 15-35 6-12 (9) x 10-20 Larger upper 2-3 20-40 (30) x 5-10 Present (200) amphistomatic (25) (15) epidermal cells (8) S. multicaulis 150-210 Dorsiventral, 15-30 (22) x 20- 10-21(15) x 15- Larger upper 2 30-55 (40) x 10-15 Absent (180) amphistomatic 70 (45) 30 (25) epidermal cells (13) S. pseudeuphratica 95-140 İsobilateral, 6-10 (8) x 10-30 4-8 (6) x 10-28 Nearly equal 2-3 15-25 (20) x 4-9 Present (115) amphistomatic (20) (18) (7) Sect. Horminum S. viridis 230-270 Dorsiventral, 20-60 (40) x 20- 9-35 (20) x 15-50 Larger upper 2 30-75 (50) x 7-16 Absent (250) amphistomatic 100 (60) (30) epidermal cells (11) Sect. Aethiopis S. aethiopis 130-165 Dorsiventral, 20-45 (28) x 25- 7-15 (10) x 15-45 Larger upper 2-3 20-30 (25) x 8-12 Absent (145) amphistomatic 75 (50) (30) epidermal cells (10) S. atropatana 125-160 Dorsiventral, 15-30 (23) x 25- 10-20 (15) x 15- Larger upper 2-3 20-40 (30) x 6-12 Absent (140) amphistomatic 50 (38) 35 (25) epidermal cells (10)

Table 6 (continued) S. brachyantha 120-150 Dorsiventral, 10-25 (17) x 25- 10-18 (14) x 18- Larger upper 2-3 20-40 (30) x 7-13 Absent (135) amphistomatic 55 (40) 40 (28) epidermal cells (10) S. candidissima 195-260 Dorsiventral, 15-25 (20) x 35- 8-13 (10) x 20-30 Larger upper 2-3 30-60 (45) x 9-15 Absent subsp. candidissima (225) amphistomatic 60 (45) (25) epidermal cells (12) S. ceratophylla 135-200 Dorsiventral, - - Nearly equal 2-3 20-45 (32) x 10-15 Absent (165) amphistomatic (12) S. cilicica 150-220 Dorsiventral, 10-20 (15) x 20- 7-12 (9) x 18-30 Larger upper 2 20-50 (35) x 8-15 Absent (180) amphistomatic 50 (35) (24) epidermal cells (11) S. cyanescens 75-120 Dorsiventral, 7-15 (11) x 15-30 5-8 (6) x 12-20 Larger upper 2-3 20-40 (30) x 6-10 Absent (95) amphistomatic (23) (16) epidermal cells (8) S. eriophora 170-210 Dorsiventral, 15-25 (20) x 20- 10-20 (15) x 15- Larger upper 2-3 20-30 x 8-12 (10) Absent (190) amphistomatic 50 (35) 35 (25) epidermal cells S. frigida 100-140 Dorsiventral, 9-23 (16) x 20-60 7-13 (10) x 10-30 Larger upper 2-3 15-25 (20) x 7-12 Absent (120) amphistomatic (40) (20) epidermal cells (10) S. hypargeia 85-110 Dorsiventral, 10-18 (15) x 20- 5-8 (7) x 10-25 Larger upper 1-2 20-40 (30) x 6-8 Absent (100) amphistomatic 45 (34) (13) epidermal cells (7)

58 S. indica 265-285 Dorsiventral, 26-50 (38) x 50- 18-30 (25) x 20- Larger upper 2-3 45-80 (60) x 8- 23 Absent

(275) amphistomatic 100 (75) 60 (40) epidermal cells (15) S. limbata 185-260 İsobilateral, 15-35 (25) x 25- 7-25 (15) x 10-30 Larger upper 4-5 25-60 (42) x 10-16 Absent (220) amphistomatic 65 (45) (20) epidermal cells (13) S. longipedicellata 75-100 Dorsiventral, 15-20 (18) x 20- 8-15 (11) x 10-20 Larger upper 2 15-25 (20) x 6-8 Absent (90) amphistomatic 35 (28) (15) epidermal cells (7) S. microstegia 105-150 Dorsiventral, 10-20 (15) x 20- 8-12 (10) x 15-25 Larger upper 2-3 15-30 (22) x 6-10 Absent (125) amphistomatic 45 (32) (20) epidermal cells (8) S. montbretii 125-150 Dorsiventral, 10-20 (15) x 20- 8-16 (12) x 15-35 Larger upper 1-2 25-50 (39) x 8-15 Absent (135) amphistomatic 60 (40) (25) epidermal cells (12) S. palaestina 210-260 İsobilateral, 15-35 (25) x 20- 15-35 (25) x 20- Nearly equal 3-4 20-70 (45) x 10-20 Absent (235) amphistomatic 85 (50) 80 (50) S. poculata 115-145 Dorsiventral, 14-20 (17) x 20- 10-20 (15) x 15- Nearly equal 2-3 10-30 (20) x 8-12 Absent (130) amphistomatic 40 (30) 40 (26) (10) S. sclarea 100-130 Dorsiventral, 10-25 (17) x 15- 7-15 (10) x 15-30 Larger upper 2-3 20-30 (25) x 6-10 Absent (115) amphistomatic 50 (30) (21) epidermal cells

Table 6 (continued) S. siirtica 80-110 Dorsiventral, 15-30 (22) x 17- 10-25 (17) x 15- Larger upper 1 20-30 (25) x 8-12 Absent (95) amphistomatic 60 (35) 53 (33) epidermal cells S. syriaca 175-220 İsobilateral, 15-35 (25) x 22- 7-20 (11) x 10-20 Larger upper 4-5 - Absent (200) amphistomatic 45 (34) (13) epidermal cells S. xanthocheila 125-180 Dorsiventral, 10-30 (20) x 18- 6-14 (9) x 10-20 Larger upper 2-3 23-45 (35) x 6-12 Absent (150) amphistomatic 55 (35) (15) epidermal cells (10) Sect. Drymosphace S. glutinosa 165-240 Dorsiventral, 15-35 (25) x 15- 15-32 (23) x 15- Nearly equal 2-3 22-35 (30) x 8-12 Absent (200) hypospomatic 40 (27) 35 (25) (10) Sect. Plethiosphace S. dicroantha 130-160 İsobilateral, 15-40 (27) x 25- 8-20 (15) x 10-35 Larger upper 3-4 14-32 (25) x 6-13 Absent (145) amphistomatic 60 (43) (20) epidermal cells (10) S. nemorosa 170-220 İsobilateral, 20-50 (35) x 20- 11-25 (17) x 13- Larger upper 4-5 20-55 (35) x 7-15 Absent (195) amphistomatic 53 (36) 25 (19) epidermal cells (11) S. staminea 130-215 İsobilateral, 17-50 (33) x 30- 12-30 (21) x 20- Larger upper 4-5 20-35 (27) x 7-11 Absent

59 (170) amphistomatic 65 (47) 50 (36) epidermal cells (9)

S. virgata 110-160 İsobilateral, 12-45 (28) x 12- 9-25 (16) x 13-40 Larger upper 2-4 20-60 (40) x 8-20 Absent (135) amphistomatic 50 (30) (25) epidermal cells (15) Sect. Hemisphace S. russellii 125-205 Dorsiventral, 10-30 (20) x 15- 6-10 (8) x 10-30 Larger upper 1-2 20-45 (35) x 6-9 Present (165) amphistomatic 65 (40) (20) epidermal cells (7) S. verticillata subsp. 180-235 Dorsiventral, 10-25 (18) x 20- 8-15 (11) x 15-30 Larger upper 2 20-50 (35) x 8-13 Present amasiaca (205) amphistomatic 50 (35) (25) epidermal cells (11) S. verticillata subsp. 155-245 Dorsiventral, 10-27 (19) x 20- 8-16 (12) x 15-30 Larger upper 2 18-35 (26) x 8-12 Present verticillata (200) amphistomatic 45 (33) (25) epidermal cells (10)

Table 7. Anatomical characteristics of leaf blades in the taxa studied of Salvia. All sizes are in µm. Numbers refer to minimum-maximum (mean). Taxa and sections Shape of the Width of midrib Length of midrib Number Lateral adaxial side of of median bundles in the midrib bundles each wing Sect. Salvia S. anatolica Flat to slightly 600-700 (650) 600-650 (625) 1 0 convex S. ballsiana Strongly convex 1000-1300 (1200) 1150-1550 (1400) 1 1 S. bracteata Flat to slightly 450-550 (500) 400-450 (425) 1 0 convex S. caespitosa Flat 170-210 (190) 220-260 (240) 1 0 S. divaricata Slightly convex 1000-1250 (1100) 850-1050 (950) 1 1 S. hedgeana Flat 400-450 (425) 430-450 (440) 1 0 S. huberi Flat 250-300 (280) 280-320 (300) 1 0 S. macrochlamys Slightly convex 1800-2000 (1900) 1400-1520 (1450) 1 1-2 S. pachystachys Flat 240-280 (260) 240-290 (270) 1 0 S. pilifera Slightly convex 750-850 (800) 700-800 (770) 1 0 S. pinnata Flat 650-700 (680) 500-550 (525) 1 1 S. recognita Flat 1050-1150 (1100) 750-870 (800) 1 0 S. rosifolia Flat 300-340 (320) 350-380 (360) 1 0 S. suffruticosa Flat 600-620 (610) 570-590 (580) 1 0 S. tomentosa Flat 900-920 (910) 690-715 (700) 1 0 S. trichocloda Flat to slightly 550-600 (580) 500-550 (530) 1 0 convex Sect. Hymenosphace S. absconditiflora Flat 500-800 (650) 500-700 (600) 1 0 S. blepharochleana Flat to slightly 550-650 (600) 550-650 (600) 1 0 convex S. cerino-pruinosa Flat ? ? 1 0 S. euphratica var. Flat 550-800 (675) 525-800 (650) 1 0 euphratica S. euphratica var. Flat - - 1 0 leiocalycina S. hydrangea Flat 400-450 (425) 380-420 (400) 1 0 S. kronenburgii Flat 700-950 (850) 750-900 (830) 1 0 S. multicaulis Flat 600-700 (650) 450-550 (500) 1 0 S. pseudeuphratica Flat 400-450 (425) 380-420 (400) 1 0 Sect. Horminum S. viridis Flat 1500-1700 (1600) 550-600 (580) 2-3 2 Sect. Aethiopis S. aethiopis Flat to slightly 2000-2200 (2100) 1700-1800 (1750) 3 1 concave S. atropatana Slightly convex 1480-1560 (1525) 1440-1500 (1470) 2 (-3) 1 S. brachyantha Slightly convex 1150-1250 (1200) 1050-1100 (1080) 2-3 1 S. candidissima Slightly convex 1600-2000 (1800) 1700-1950 (1800) 1 0 subsp. candidissima S. ceratophylla Convex 1000-1200 (1100) 1400-1550 (1450) 1 1 S. cilicica Slightly convex 1050-1150 (1100) 1000-1000 (1050) 1 0 S. cyanescens Slightly convex 750-800 (775) 800-850 (825) 1 0 S. eriophora Slightly convex 1250-1350 (1300) 1150-1250 (1200) 2 1 S. frigida Flat 930-970 (950) 730-770 (750) 2 1 60

Table 7 (continued) S. hypargeia Slightly convex 950-1050 (1000) 1050-1250 (1150) 1 1 S. indica Slightly convex 1650-1850 (1750) 1340-1480 (1400) 2-3 1 S. limbata Slightly convex 1650-1750 (1700) 1640-1730 (1685) 4-6 1 S. longipedicellata Slightly convex 1100-1200 (1150) 1100-1200 (1150) 1 1 S. microstegia Slightly convex 1450-1500 (1480) 1050-1100 (1070) 4 1 S. montbretii Flat 1250-1300 (1275) 1080-1120 (1100) 1 1 S. palaestina Slightly convex 1900-2000 (1950) 2050-2150 (2100) 1 1 S. poculata Flat 900-1000 (950) 900-1000 (950) 1 1 S. sclarea Flat 1750-1800 (1775) 1510-1550 (1530) 1 1 S. siirtica Flat 2100-2200 (2150) 1700-1770 (1735) 4-6 1 S. syriaca Slightly convex 1300-1400 (1350) 1100-1160 (1130) 6 0 S. xanthocheila Slightly convex 1800-1900 (1850) 1450-1550 (1500) 3 1 Sect. Drymosphace S. glutinosa Slightly convex 1250-1350 (1300) 1450-1550 (1500) 1 0 Sect. Plethiosphace S. dicroantha Flat 1200-1250 (1225) 850-900 (975) 3 0 S. nemorosa Flat 1450-1500 (1470) 1220-1260 (1240) 3-4 1 S. staminea Slightly convex 1650-1700 (1675) 1700-1750 (1725) 1 1 S. virgata Slightly convex 1325-1750 (1500) 1175-1425 (1300) 4-7 0 Sect. Hemisphace S. russellii Flat 850-900 (875) 650-700 (675) 1 0 S. verticillata subsp. Flat to slightly 1500-1600 (1550) 1300-1400 (1350) 1 1 amasiaca convex S. verticillata subsp. Flat to slightly - - 1 0 verticillata convex

3.2.4. Petiole Anatomy (Figures 31-37, Table 8)

In cross section, the petioles can be categorized into seven types: open U-shaped (e.g. S. huberi, S. rosifolia, S. hydrangea, S. dicroantha), U-shaped with obtuse or erect margins (e.g. S. suffruticosa, S. tomentosa, S. viridis, S. glutinosa, S. sclarea, S. nemorosa), open V-shaped (e.g. S. anatolica, S. divaricata, S. belpharochleana, S. euphratica var. euphratica, S. euphratica var. leiocalycina), D-shaped with more or less procumbent margins (e.g. S. aethiopis, S. candidissima subsp. candidissima, S. frigida, S. hypargeia, S. syriaca, S. virgata, S. russellii, S. verticillata subsp. amasiaca, S. verticillata subsp. verticillata), obtuse triangular (e.g. S. bracteata, S. trichoclada), triangular (e.g. S. ballsiana, S. macrochlamys, S. absconditiflora, S. multicaulis) or open crescent shaped (e.g. S. caespitosa, S. pachystachys). The width of the petioles ranges from 1000 µm in S. blepharochleana to 2200 µm in S. aethiopis. The length of the midrib varies between 550 µm in S. pachystachys and 2150 µm in S. aethiopis.

The epidermal cells of the petioles are oval, rectangular and quadrangular. The upper and lower cell walls of the epidermis are thicker than the lateral walls. The upper surface is covered with a thin or thick cuticle. Peltate glandular, capitate glandular or non- glandular trichomes are found on the adaxial and abaxial sides. Collenchymatous cells can be observed into two different types: angular or annular. The annualar collenchymatous cells have one to two layers below the abaxial epidermis and one to four below the adaxial epidermis. The angular collenchymatous cells are situated under the epidermis in the corners of the petioles and have one to eight layers. Ground tissue is composed of parenchymatous cells which are heterogeneous in size.

The median vascular bundle is continuous with one large bundle or is divided into two to eight separate bundles. The shape of the central vascular bundle(s) is/are open U-shaped occasionally with incurved ends (e.g. S. ballsiana, S. pilifera, S. suffruticosa, S. tomentosa, S. aethiopis, S. brachyantha), U-shaped (e.g. S. divaricata, S. rosifolia), V- shaped (e.g. S. glutinosa) or open crescent shaped (e.g. S. anatolica, S. cerino-pruinosa, S. euphratica var. euphratica, S. euphratica var. leiocalycina). Some species, such as S.

candidissima subsp. candidissima, S. cilicica, S. cyanescens, S. limbata, S. palaestina, S. siirtica, S.syriaca and S. virgata, have also discontinuous central vascular bundles with incurved ends and one to several inverse bundles on the adaxial side. There are one to five small accessory vascular bundles in each side. In some species (e.g. S. aethiopis, S. brachyantha, S. eriophora, S. longipedicellata, S. microstegia, S. siirtica, S. nemorosa, S. verticillata) sclerenchyma tissue is found on both the phloem and the xylem, while in some species (e.g. taxa of sect. Hymenosphace and Salvia except S. ballsiana) sclerenchyma tissue is present only above the phloem. However, the petioles of S. ballsiana have vascular bundles without sclerenchymatous fibers.

Figure 31. Cross section of the petiole of S. pinnata. ab: abaxial side, ad: adaxial side, col: collenchymas, lv: lateral vascular bundle, mv: median vascular bundle.

Figure 32. Cross section of the petiole of S. pachystachys.

Figure 33. Cross section of the petiole of S. viridis.

Figure 34. Cross section of the petiole of S. glutinosa.

Figure 35. Cross section of the petiole of S. syriaca.

Figure 36. Cross section of the petiole of S. verticillata subsp. verticillata.

Figure 37. Cross section of the petiole of S. cerino-pruinosa.

Table 8. Anatomical characteristics of petioles in the taxa studied of Salvia. All sizes are in µm. Numbers refer to minimum-maximum (mean). Taxa and sections Petiole shape Width of Length of Number of Shape of median Numbers of Layers of angular Shape of the petiole petiole median bundles shape lateral bundles in collenchyma adaxial side of the bundles each wing petiole Sect. Salvia S. anatolica Open V- 2000-2600 1000-1400 1 Open crescent shaped 3 4-5 Concave shaped (2350) (1200) S. ballsiana Triangular 2000-2200 1500-1670 1 Open U-shaped with 2-3 4-5 Slightly convex (2100) (1600) incurved ends S. bracteata Obtuse 2300-2500 1000-1200 1 Open U-shaped 2 7-8 Flat triangular (2400) (1100) S. caespitosa Open crescent 1800-2200 650-900 1 Open U-shaped 1 (-2) 2-3 Concave shaped (2000) (800) S. divaricata Open V- 2100-2500 1300-1400 1 U-shaped 1 5-6 Concave

67 shaped (2300) (1350)

S. hedgeana Open V- 1750-1800 900-950 1 Open U-shaped 3 6-7 Concave shaped (1780) (930) S. huberi Open U- 1450-1500 800-885 1 Open U-shaped 2-3 3-4 Concave shaped (1480) (850) S. macrochlamys Triangular 2400-2500 700-740 1 Open crescent shaped 1-2 5-6 Flat to slightly (2400) (720) convex S. pachystachys Open crescent 1450-1600 550-650 1 Open U-shaped 1 (-2) 1-2 Concave shaped (1500) (600) S. pilifera Triangular 1800-1900 1400-1460 1 Open U-shaped with 2 6-7 Flat (1850) (1430) incurved ends S. pinnata Triangular 1300-1600 860-1040 1 Open U-shaped 1-2 5-6 Flat (1450) (950) S. rosifolia Open U- 1550-1700 1200-1300 1 U-shaped 2-3 5-7 Concave shaped (1625) (1250) S. suffruticosa U-shaped 1350-1900 1000-1425 1 Open U-shaped with 2-3 3-4 Concave with obtuse (1650) (1200) incurved ends margins

Table 8. (continued) S. tomentosa U-shaped 1085-1200 1000-1050 1 Open U-shaped with 1 5 Concave with obtuse (1200) (1025) incurved ends margins S. trichocloda Obtuse 2400-2700 1200-1250 1 Open U-shaped with 2-3 7-8 Flat triangular (2600) (1220) incurved ends Sect. Hymenosphace S. absconditiflora Triangular 1500-1750 870-1000 1 Open U-shaped 1 (-2) 5-7 Flat (1600) (950) S. blepharochleana Open V- 1000-1575 800-925 1 Open U-shaped 1 (-2) 2-3 Concave shaped (1550) (870) S. cerino-pruinosa Open V- 1200-2000 800-1400 3-4 Open crescent shaped 1 6-8 Concave shaped (1650) (1150) S. euphratica var. Open V- 1400-1950 1000-1330 3 Open crescent shaped 1 (-2) 3-5 Concave euphratica shaped (1700) S. euphratica var. Open V- 1400-1500 800-1000 3 Open crescent shaped 1 (-2) 3-4 Concave

68 leiocalycina shaped (1450) (900)

S. hydrangea Open U- 1250-1450 1040-1060 1 Open U-shaped 1 (-2) 2-4 Concave shaped (1350) (1050) S. kronenburgii Triangular 1600-2200 950-1500 1 Open U-shaped 1 3-4 Nealy flat (1900) (1250) S. multicaulis Triangular 1200-1400 600-680 1 Open U-shaped 1 (-2) 4-5 Flat (11300) (650) S. pseudeuphratica Open U- 1050-1500 800-1000 1 Open U-shaped 1-2 3-4 Concave shaped (1300) (900) Sect. Horminum S. viridis U-shaped 2300-2500 1800-1925 2-3 Open U-shaped 2-4 1-2 Concave with erect (2400) (1850) margins Sect. Aethiopis S. aethiopis D-shaped 6000-7000 5000-6000 5-8 Open U-shaped with 3 6-7 Flat with (6500) (5500) incurved ends procumbent margins 68

Table 8. (continued) S. atropatana D-shaped 1600-2000 1200-1500 3 Open U-shaped 2-3 3-4 Flat to slightly with (1800) (1350) convex procumbent margins S. brachyantha D-shaped 1800-2500 1300-1650 3-6 Open U-shaped with 2 3-4 Flat to slightly with (2150) (1500) incurved ends convex procumbent margins S. candidissima D-shaped 2800-3200 2700-3100 4-6 Discontinuous arc of 3 4-5 Flat subsp. candidissima with (3000) (2900) abaxially bundles with procumbent incurved ends and margins several inverse bundles on the adaxial side S. ceratophylla D-shaped 3000-3200 2500-2700 6 Open U-shaped 3 - Flat

69 with (3100) (2600)

procumbent margins S. cilicica D-shaped 1900-2000 1600-1700 3 Discontinuous arc of 2 4-5 Slightly convex with (1950) (1650) abaxially vascular with procumbent incurved ends and margins several inverse bundles on the adaxial side S. cyanescens D-shaped 2950-3150 2900-3100 2 Discontinuous arc of 2-4 4 Flat with (3050) (3000) abaxially vascular with procumbent incurved ends and margins several inverse bundles on the adaxial side S. eriophora D-shaped - - 2-3 Open U-shaped 2-3 - Flat with procumbent margins

Table 8. (continued) S. frigida D-shaped 1480-1550 1040-1060 4 Open U-shaped 1-2 - Flat with (1510) (1050) procumbent margins S. hypargeia D-shaped 1800-1850 1250-1300 2-3 Open U-shaped 2 - Slightly convex with (1825) (1275) procumbent margins S. limbata D-shaped 2800-2900 2400-2500 4-8 Discontinuous arc of 4 - Concave with (2850) (2450) abaxially bundles with procumbent incurved ends and one margins inverse bundles on the adaxial side S. longipedicellata D-shaped 2800-3000 2250-2400 3 Open U-shaped 2 - Slightly concave with (2900) (2325) 70 procumbent margins S. microstegia D-shaped 1900-1950 1300-1330 3 Open U-shaped 2 3-4 Flat with (1925) (1315) procumbent margins S. palaestina D-shaped 2900-3000 2350-2400 4-6 Discontinuous arc of 2-3 4-5 Flat with (2950) (2370) abaxially bundles with procumbent incurved ends and one margins inverse bundles on the adaxial side S. poculata D-shaped 2400-2900 1650-1750 4 Open U-shaped 3 4-5 Flat with (2700) (1700) procumbent margins

Table 8. (continued) S. sclarea U-shaped 3000-3400 2500-2800 3 or more Discontinuous arc of 4-5 4-5 Concave with erect (3200) (2650) abaxially bundles with margins incurved ends and several inverse bundles on the adaxial side S. siirtica U-shaped 3000-3500 2850-3350 8 Discontinuous arc of 3-4 - Concave with erect (3250) (3100) abaxially bundles with margins incurved ends and several inverse bundles on the adaxial side (kenarlarda da var) S. syriaca D-shaped 2000-2600 1300-2000 3-4 Discontinuous arc of 1-2 3-4 Flat with (2300) (1650) abaxially bundles with procumbent incurved ends and one margins inverse bundles on the 71 adaxial side S. xanthocheila D-shaped - - 3 Open U-shaped 2 4-5 Flat to slightly with concave procumbent margins Sect. Drymosphace S. glutinosa U-shaped 3000-3100 2650-2750 1 Open V-shaped with 3-4 4-5 Concave with obtuse (3050) (2700) incurved ends margins Sect. Plethiosphace S. dicroantha Open U- 1500-1500 950-1000 3 Open U-shaped 3 2-3 Concave shaped (1450) (975) S. nemorosa U-shaped 2000-2500 1800-1950 3-4 Open U-shaped 3-4 4-6 Concave with erect (2250) (1875) margins S. staminea D-shaped 2600-2700 2250-2300 1 Open U-shaped 2-3 2-3 Concave (2650) (2275)

Table 8. (continued) S. virgata D-shaped 2050-2300 1650-1750 6-7 Discontinuous arc of 1-2 4-5 Flat (2200) (1700) abaxially bundles with incurved ends and two inverse bundles on the adaxial side Sect. Hemisphace S. russellii D-shaped 1900-2000 1500-1550 1 Open U-shaped 1 - Slightly concave with (1950) (1525) procumbent margins S. verticillata subsp. D-shaped 2250-3000 1550-2050 2-5 Open U-shaped 2 4-5 Slightly concave amasiaca with (2600) (1800) procumbent margins S. verticillata subsp. D-shaped 2600-3200 1600-2050 4-5 Open U-shaped 2 5-6 Slightly concave

72 verticillata with (2900) (1850)

procumbent margins

3.2.5. Infrageneric Anatomical Descriptions of Salvia

Sect. Salvia Root anatomy: The thickness of the phloem ranges from 50 µm to 200 µm. The thickness of the xylem varies between 400 µm to 3000 µm. The vessel members range from 15 µm to 110 µm in diameter. The tracheids vary between 4 µm to 20 µm in diameter. The xylem ray cells are composed of 1 to 6 radial rows. They range from 9 µm to 40 µm in length and from 3 µm to 20 µm width. The pith cells vary between 25 µm and 140 µm (Table 4).

Stem anatomy: The stems are between 1400 µm and 5000 µm in diameter and mostly quadrangular to more or less circular or six-sided. Under the epidermis, 1-7 layers (40- 160 µm thick) of collenchymatous cells are found at the corners. There are 1-4 layers of chlorenchymatous cells situated between the corners. The cortex contains of 1-8-layered (15-350 µm thick). The vascular bundles at the corners (150-900 µm) are larger than those between the corners (70-800 µm). The pith is 750 to 3000 µm (Table 5).

Leaf blade anatomy: The leaf blades are amphistomatic with 100-350 µm thick. The upper epidermal cells are mostly equal to lower epidermal cells. The upper epidermal cells are 8-45 µm in length and 15-65 µm in width, whereas the lower epidermal cells are 8-45 µm in length and 12-60 µm in width. Hypodermis is present under both epidermis or absent. The mesophyll is mostly isobilateral. In the isobilateral leaves, two to six layers of the palisade parenchyma is found on both sides of the blade. In the dorsiventral leaves, the mesophyll consists of two to four layers of the palisade parenchyma followed by one to three layers of the spongy parenchyma. The palisade parenchyma is 15-90 µm in length and 6-17 µm in width. The abaxial side of the midrib is mostly flat to slightly convex or strongly convex, while it is convex on the adaxial side. The width of the midrib ranges from 170 µm to 2000 µm. The length of the midrib varies between 220 µm and 1520 µm. Collenchymatous cells have one layer below the abaxial epidermis and one to two below the adaxial epidermis. There is one large central vascular bundle. The central vascular bundle is open U-shaped sometimes sometimes

with incurved ends. There are no inverse bundles on the adaxial side. Accessory vascular bundles are mainly absent (Table 6-7).

Petiole anatomy: The petioles are open U-shaped, U-shaped with obtuse margins, open V-shaped, obtuse triangular, triangular and open crescent shaped. The width of the petioles ranges from 1085 µm to 1200 µm. The length of the midrib varies between 550 µm and 2600 µm. The annualar collenchymatous cells have one layer below the abaxial epidermis and one to three below the adaxial epidermis. The angular collenchymatous cells have two to eight layers. The median vascular bundle is continuous with one large bundle. The shape of the central vascular bundle is open U-shaped sometimes with incurved ends, U-shaped and open crescent shaped. Inverse bundles on the adaxial side are absent. There are one to four small accessory vascular bundles in each side (Table 8).

Sect. Hymenosphace Root anatomy: The thickness of the phloem ranges from 50 µm to 130 µm. The thickness of the xylem varies between 300 µm to 1200 µm. The vessel members range from 10 µm to 100 µm in diameter. The tracheids vary between 3 µm to 15 µm in diameter. The xylem ray cells are composed of 1 to 7 radial rows. They range from 8 µm to 25 µm in length and from 4 µm to 15 µm width. The pith cells vary between 20 µm and 100 µm (Table 4).

Stem anatomy: The stems are between 1200 µm and 3300 µm in diameter and quadrangular to more or less circular. Under the epidermis, 3-7 layers (50-150 µm thick) of collenchymatous cells are found at the corners. There are 1-4 layers of chlorenchymatous cells situated between the corners. The cortex contains of 2-9-layered (30-500 µm thick). The vascular bundles at the corners (150-350 µm) are larger than those between the corners (90-315 µm). The pith is 600 to 2450 µm (Table 5). Leaf blade anatomy: The leaf blades are amphistomatic with 95-270 µm thick. The upper epidermal cells are mostly equal to lower epidermal cells. The upper epidermal cells are 6-30 µm in length and 10-70 µm in width, whereas the lower epidermal cells are 4-20 µm in length and 10-45 µm in width. Hypodermis is present under both 74

epidermis or absent. The mesophyll is mostly isobilateral. In the isobilateral leaves, two to five layers of the palisade parenchyma is found on both sides of the blade. In the dorsiventral leaves, the mesophyll consists of two to three layers of the palisade parenchyma followed by one to two layers of the spongy parenchyma. The palisade parenchyma is 15-55 µm in length and 4-15 µm in width. The abaxial side of the midrib is mostly flat or rarely flat to slightly convex, while it is convex on the adaxial side. The width of the midrib ranges from 4000 µm to 950 µm. The length of the midrib varies between 380 µm and 900 µm. Collenchymatous cells have one layer below the abaxial and adaxial epidermis. There is one large central vascular bundle. The central vascular bundle is open U-shaped. There are no inverse bundles on the adaxial side. Accessory vascular bundles are absent (Table 6-7).

Petiole anatomy: The petioles are open U-shaped, open V-shaped and triangular. The width of the petioles ranges from 1000 µm to 2000 µm. The length of the midrib varies between 600 µm and 1330 µm. The annualar collenchymatous cells have one layer below the abaxial epidermis and one to two below the adaxial epidermis. The angular collenchymatous cells have two to eight layers. The median vascular bundle is continuous with one large bundle except S. cerino-pruinosa and S. euphratica which have three or four median bundles. The shape of the central vascular bundle is open U- shaped and open crescent shaped. Inverse bundles on the adaxial side are absent. There are one to two small accessory vascular bundles in each side (Table 8).

Sect. Horminum Root anatomy: The thickness of the phloem ranges from 50 µm to 70 µm. The thickness of the xylem varies between 1200 µm to 1450 µm. The vessel members range from 15 µm to 85 µm in diameter. The tracheids vary between 5 µm to 20 µm in diameter. The xylem rays are up to 20 radial rows. They range from 12 µm to 35 µm in length and from 5 µm to 15 µm width. There is no pith as the xylem fills all the central portion of the root (Table 4).

Stem anatomy: The stems are between 2300 µm and 2500 µm in diameter and quadrangular. Under the epidermis, 1-4 layers (20-60 µm thick) of collenchymatous cells are found at the corners. There are 2-3 layers of chlorenchymatous cells situated between the corners. The cortex contains of 1-4-layered (15-100 µm thick). The vascular bundles at the corners (350-400 µm) are larger than those between the corners (100-200 µm). The pith is 1750 to 1850 µm (Table 5).

Leaf blade anatomy: The leaf blades are amphistomatic with 230-270 µm thick. The upper epidermal cells are larger than the lower epidermal cells. The upper epidermal cells are 20-60 µm in length and 20-100 µm in width, whereas the lower epidermal cells are 9-35 µm in length and 15-50 µm in width. Hypodermis is absent. The mesophyll is dorsiventral with two layers of the palisade parenchyma followed by two to four layers of the spongy parenchyma. The palisade parenchyma is 30-75 µm in length and 7-16 µm in width. The abaxial side of the midrib is flat, while it is convex on the adaxial side. The width of the midrib ranges from 1500 µm to 1700 µm. The width of the midrib varies between 550 µm and 600 µm. Collenchymatous cells have one layer below the abaxial and adaxial epidermis. There are two to three large central vascular bundles. The central vascular bundle is open U-shaped. There are no inverse bundles on the adaxial side. Two small accessory vascular bundles are present in each side of the midrib (Table 6-7).

Petiole anatomy: The petioles are U-shaped with erect margins. The width of the petioles ranges from 2300 µm to 2500 µm. The length of the midrib varies between 1800 µm and 1920 µm. The annualar collenchymatous cells have one to two layers below the abaxial epidermis and one to two below the adaxial epidermis. The angular collenchymatous cells have one to two layers. The median vascular bundle is discontinuous with two to three large bundles. The shape of the central vascular bundle is open U-shaped. Inverse bundles on the adaxial side are absent. There are two to four small accessory vascular bundles in each side (Table 8).

Sect. Aethiopis Root anatomy: The thickness of the phloem ranges from 75 µm to 140 µm. The thickness of the xylem varies between 500 µm to 3500 µm. The vessel members range from 15 µm to 130 µm in diameter. The tracheids vary between 5 µm to 25 µm in diameter. The xylem rays are up to 40 radial rows. They range from 15 µm to 90 µm in length and from 7 µm to 25 µm width. The pith cells vary between 15 µm and 130 µm (Table 4).

Stem anatomy: The stems are between 2000 µm and 7000 µm in diameter and quadrangular. Under the epidermis, 3-9 layers (60-250 µm thick) of collenchymatous cells are found at the corners. There are 1-3 layers of chlorenchymatous cells situated between the corners. The cortex contains of 1-15-layered (30-650 µm thick). The vascular bundles at the corners (250-850 µm) are larger than those between the corners (90-700 µm). The pith is 1750 to 1850 µm (Table 5).

Leaf blade anatomy: The leaf blades are amphistomatic with 75-285 µm thick. The upper epidermal cells are mostly larger than the lower epidermal cells. The upper epidermal cells are 7-50 µm in length and 15-100 µm in width, whereas the lower epidermal cells are 5-35 µm in length and 10-60 µm in width. Hypodermis is absent. The mesophyll is mostly dorsiventral. In the isobilateral leaves, three to five layers of the palisade parenchyma is found on both sides of the blade. In the dorsiventral leaves, the mesophyll consists of one to three layers of the palisade parenchyma followed by one to four layers of the spongy parenchyma. The palisade parenchyma is 10-80 µm in length and 6-23 µm in width. The abaxial side of the midrib is mostly flat to slightly convex or rarely slightly concave, while it is convex on the adaxial side. The width of the midrib ranges from 900 µm to 2200 µm. The width of the midrib varies between 730 µm and 2150 µm. Collenchymatous cells have one to two layers below the abaxial epidermis and one to three below the adaxial epidermis. There are one to six large central vascular bundles. The central vascular bundle is open U-shaped sometimes with incurved ends or open crescent shaped. There are several small inverse bundles on the

adaxial side in some species. One small accessory vascular bundle is present in each side of the midrib (Table 6-7).

Petiole anatomy: The petioles are mostly D-shaped with procumbent margins or U- shaped. The width of the petioles ranges from 1480 µm to 7000 µm. The length of the midrib varies between 1040 µm and 6000 µm. The annualar collenchymatous cells have one to two layers below the abaxial epidermis and one to four below the adaxial epidermis. The angular collenchymatous cells have three to seven layers. The median vascular bundle is discontinuous with two to eight large bundles. The shape of the central vascular bundle is open U-shaped occasionally with incurved ends. Inverse bundles on the adaxial side are sometimes present. There are one to five small accessory vascular bundles in each side (Table 8).

Sect. Drymosphace Root anatomy: The thickness of the xylem varies between 1300 µm to 1600 µm. The vessel members range from 20 µm to 80 µm in diameter. The tracheids vary between 5 µm to 20 µm in diameter. The xylem rays are up to 30 radial rows. They range from 15 µm to 35 µm in length and from 7 µm to 20 µm width. There is no pith as the xylem fills all the central portion of the root (Table 4).

Stem anatomy: The stems are between 3000 µm and 3500 µm in diameter and quadrangular. Under the epidermis, 7-8 layers (120-150 µm thick) of collenchymatous cells are found at the corners. There are 1-3 layers of chlorenchymatous cells situated between the corners. The cortex contains of 3-6-layered (50-200 µm thick). The vascular bundles at the corners (370-430 µm) are larger than those between the corners (100-200 µm) (Table 5).

Leaf blade anatomy: The leaf blades are hypostomatic with 165-240 µm thick. The upper epidermal cells are equal to the lower epidermal cells. The upper epidermal cells are 15-35 µm in length and 15-40 µm in width, whereas the lower epidermal cells are 15-32 µm in length and 15-35 µm in width. Hypodermis is absent. The mesophyll is 78

dorsiventral with two to three layers of the palisade parenchyma followed by three to four layers of the spongy parenchyma. The palisade parenchyma is 22-35 µm in length and 8-12 µm in width. The abaxial side of the midrib is slightly convex, while it is convex on the adaxial side. The width of the midrib ranges from 1250 µm to 1350 µm. The width of the midrib varies between 1450 µm and 1550 µm. Collenchymatous cells have one layer below the abaxial epidermis and one to two below the adaxial epidermis. There is one large central vascular bundle. The central vascular bundle is open U-shaped with incurved ends. There are several small inverse bundles on the adaxial side in some species. Accessory vascular bundles are absent (Table 6-7).

Petiole anatomy: The petioles are U-shaped with obtuse margins. The width of the petioles ranges from 3000 µm to 3100 µm. The length of the midrib varies between 2650 µm and 2750 µm. The annualar collenchymatous cells have two layers below the abaxial epidermis and two to three below the adaxial epidermis. The angular collenchymatous cells have four to five layers. The median vascular bundle is continuous with one large bundle. The shape of the central vascular bundle is open V-shaped with incurved ends. Inverse bundles on the adaxial side are absent. There are three to four small accessory vascular bundles in each side (Table 8).

Sect. Plethiosphace Root anatomy: The thickness of the xylem varies between 700 µm to 1050 µm. The vessel members range from 15 µm to 70 µm in diameter. The tracheids vary between 5 µm to 24 µm in diameter. The xylem rays are up to 12 radial rows. They range from 12 µm to 55 µm in length and from 5 µm to 20 µm width. The pith cells vary between 20 µm and 70 µm (Table 4).

Stem anatomy: The stems are between 1650 µm and 3500 µm in diameter and quadrangular. Under the epidermis, 5-8 layers (75-160 µm thick) of collenchymatous cells are found at the corners. There are 1-5 layers of chlorenchymatous cells situated between the corners. The cortex contains of 2-7-layered (20-350 µm thick). The vascular

bundles at the corners (200-450 µm) are larger than those between the corners (60- 375µm). The pith is 1200 to 2200 µm (Table 5).

Leaf blade anatomy: The leaf blades are hypostomatic with 110-220 µm thick. The upper epidermal cells are larger than the lower epidermal cells. The upper epidermal cells are 12-50 µm in length and 12-60 µm in width, whereas the lower epidermal cells are 8-30 µm in length and 10-50 µm in width. Hypodermis is absent. The mesophyll is isobilateral with two to five layers of the palisade. The palisade parenchyma is 14-60 µm in length and 6-20 µm in width. The abaxial side of the midrib is flat or slightly convex, while it is convex on the adaxial side. The width of the midrib ranges from 1200 µm to 1650 µm. The width of the midrib varies between 850 µm and 1700 µm. Collenchymatous cells have one layer below the abaxial epidermis and one to three below the adaxial epidermis. There are one to seven large central vascular bundles. The central vascular bundle is open U-shaped. There are several small inverse bundles on the adaxial side in S. virgata. Accessory vascular bundles are absent or present (Table 6-7).

Petiole anatomy: The petioles are open U-shaped with erect margins, U-shaped and D- shaped with procumbent margins. The width of the petioles ranges from 1500 µm to 2700 µm. The length of the midrib varies between 950 µm and 2300 µm. The annualar collenchymatous cells have one layer below the abaxial epidermis and one to three below the adaxial epidermis. The angular collenchymatous cells have two to six layers. One to seven median vascular bundles are present. The shape of the central vascular bundle is open U-shaped. Inverse bundles on the adaxial side are present only in S. virgata. There are one to four small accessory vascular bundles in each side (Table 8).

Sect. Hemisphace Root anatomy: The thickness of the phloem ranges from 70 µm to 120 µm. The thickness of the xylem varies between 1300 µm to 1900 µm. The vessel members range from 15 µm to 75 µm in diameter. The tracheids vary between 5 µm to 23 µm in diameter. The xylem rays are up to 15 radial rows. They range from 12 µm to 70 µm in

length and from 10 µm to 30 µm width. The pith cells vary between 25 µm and 150 µm (Table 4).

Stem anatomy: The stems are between 1500 µm and 4700 µm in diameter and quadrangular. Under the epidermis, 5-8 layers (70-190 µm thick) of collenchymatous cells are found at the corners. There are 2-4 layers of chlorenchymatous cells situated between the corners. The cortex contains of 1-10-layered (25-275 µm thick). The vascular bundles at the corners (200-800 µm) are larger than those between the corners (100-550 µm). The pith is 1000 to 3300 µm (Table 5).

Leaf blade anatomy: The leaf blades are hypostomatic with 125-245 µm thick. The upper epidermal cells are larger than the lower epidermal cells. The upper epidermal cells are 10-30 µm in length and 15-65 µm in width, whereas the lower epidermal cells are 6-16 µm in length and 10-30 µm in width. Hypodermis is present under the upper epidermis. The mesophyll is dorsiventral with one to two layers of the palisade parenchyma followed by two to three layers of the spongy parenchyma. The palisade parenchyma is 18-50 µm in length and 6-13 µm in width. The abaxial side of the midrib is flat to slightly convex, while it is convex on the adaxial side. The width of the midrib ranges from 850 µm to 1600 µm. The width of the midrib varies between 650 µm and 1400 µm. Collenchymatous cells have one layer below the abaxial epidermis and two below the adaxial epidermis. There is one large central vascular bundle. The central vascular bundle is open U-shaped with incurved ends. There are no small inverse bundles on the adaxial side. Accessory vascular bundles are absent or present (Table 6- 7).

Petiole anatomy: The petioles are D-shaped with procumbent margins. The width of the petioles ranges from 1900 µm to 3200 µm. The length of the midrib varies between1500 µm and 2050 µm. The annualar collenchymatous cells have one layer below the abaxial epidermis and one to three below the adaxial epidermis. The angular collenchymatous cells have four to six layers. One to five median vascular bundles are present. The shape

of the central vascular bundle is open U-shaped. Inverse bundles on the adaxial side are absent. There are one to two small accessory vascular bundles in each side (Table 8).

3.2.6. Systematic Implications of Anatomical Characteristics

Root anatomy of the family Lamiaceae (Metcalfe & Chalk, 1950) is characterized by xylem rays of roots composed of 2–12 or more rowed cells. The present study shows they are composed of 1-40 radial rows of square to rectangular parenchymatous cells. While the taxa of the sections Salvia and Hymenosphace have 1-6 (-7) rows of xylem ray cells, in the taxa of the others the maximum row number of xylem ray cells is 8 or more.

According to Metcalfe and Chalk (1950), stems of Lamiaceae species were quadrangular and had collenchyma cells covering a broad area at the corners and sclerenchyma cells surrounding the vascular tissue. The results show that the examined taxa of Salvia have mainly quadrangular to more or less circular or six-sided stems, 1-9 layers of collenchymatous cells, 1-4 layers of chlorenchymatous cells and several or many scattered groups of sclerenchymatous fibers outside the phloem.

In Lamiaceae, the vascular system of the leaf blade and petiole has been received special attention to whether or not they are useful in separating different genera in previous studies (e.g. Bokhari & Hedge, 1971; Metcalfe & Chalk 1950). The present result reveal that the shape and number of the central vascular bundle has diagnostic significance in identifying some sections and species.

The most significant diagnostic characters, particularly helpful for infrageneric delimitation, are row numbers and length of ray cells in the root, presence of hypodermis, presence of stomata on the upper and lower side of the leaf, type of structure of the leaf, shape of cross section, shape and number of median vascular bundles in the petiole. Variation in some anatomical characters, such as number of cell layers of collenchyma and cortex, size of vascular bundles in the stem, number of cell 82

layers of palisade parenchyma, number of median and lateral vascular bundles in the midrib, size of midrib, size of petiole, number of lateral vascular bundles and number of cell layer of collenchyma in the petiole, are useful for separating species within the sections.

A diagnostic key to sections of Salvia using anatomical characters are provided below:

1. Leaves hypostomatic and the median vascular bundle in the petiole open-V shaped…...……… ………………………………..…………………………………………5. sect. Drymosphace 1. Leaves ampipostomatic and the median vascular bundle in the petiole not open-V shaped…..2 2. Hypodermis present only under the upper epidermis of the leaf………7. sect. Hemisphace 2. Hypodermis present under the upper and lower epidermis of the leaf or not………………3 3. Petiole U-shaped with 2-3 median vascular bundles…………………3. sect. Horminum 3. Petiole U-shaped with 3-8 median vascular bundles or other shapes…………...……….4 4. Leaves rarely isobilateral………………….…………………………4. sect. Aethiopis 4. Leaves always or mostly isobilateral………………………...…………………..……5 5. The maximum row number of pith rays in the roots more than 8-12……………….. ……………………..…………………………………………6. sect. Plethiosphace 5. The maximum row number of pith rays in the roots up to 7………...……………… …………………………………………….…1-2. sect. Salvia and Hymenosphace

3.3. Pollen Micromorphology

The pollen micromorphology of some genera of Labiatae has been studied intensively by a number of authors (e.g. Erdtman, 1945; Cantino, 1992a, b; Harley et al., 1992, 2004; Abu-Asab & Cantino, 1992, 1993, 1994; Salmaki et al., 2008a; Moon et al., 2008a, b), but there are only few investigations dealing with pollen micromorphology of the genus Salvia (Emboden, 1964; Henderson et al., 1968; Hassan et al., 2009; Bagherpour et al., 2009; Bagherpour et al., 2010; Hassan et al., 2009; Kahraman & Doğan, 2010; Kahraman et al., 2009b, c, 2010b, c, d; Özler et al., 2011). Henderson et al. (1968) presented brief descriptions of pollen morphology of 59 Salvia taxa, some of which grow in Turkey, using mainly light microscopic observations. Özler et al. (2011) examined the surface sculpturing of pollen of 30 taxa belonging to five sections of the genus in Turkey by means of LM and SEM and defined three exine sculpturing types: reticulate-perforate, reticulate-granulate and bireticulate.

The major pollen characteristics for the taxa studied are given in Tables 9 and 10. Representative LM and SEM micrographs of pollen grains are illustrated in Figures 38- 67. The pollen morphological variation within the genus is described below for the following pollen properties: size and shape of pollen grains, number, position and morphology of apertures and exine sculpturing.

3.3.1. Size and Shape

The pollen grains are dispersed as monads and their size varies from small to large: polar axis (P) = 23.04-65.28 µm, equatorial axis (E) = 26.88-66.24 µm (Table 9). Of the taxa investigated, the smallest pollen grains, on average, are observed in S. verticillata subsp. verticillata.

The shape of the pollen grains in equatorial view ranges from suboblate to subprolate while their shape in polar view is more or less circular (Table 9; Figures 38-52, 54-67). Often grain shapes are variable among the taxa and suboblate to oblate-spheroidal forms

are common even within the same taxa. The pollen grains are radially symmetric and isopolar in all taxa studied.

3.3.2. Apertures

The pollen grains are mostly hexacolpate, sometimes intermixed with heptacolpate grains in S. kurdica (Figure 38o-p), S. candidissima subsp. candidissima and S. siirtica, and octacolpate grains in S. recognita.

The colpus length (Clg) ranges from 20.16 µm, in S. brachyantha, S. verticillata subsp. verticillata, to 60.48 µm, in S. blepharochleana. The colpus length is strongly correlated with length of polar axis. The colpus width (Clt) varies from 2.64 µm, in S. poculata, to 12.48 µm, in S. ballsiana, S. pachystachys. Colpi are distributed symmetrically, elongated, usually shallow and narrowing towards the poles, and their ends are acute. Colpus membranes are granulate (e.g. S. tomentosa, S. glutinosa, S. dicroantha), granulate-scabrate (e.g. S. ballsiana, S. pachystachys, S. viridis, S. nemorosa, S. verticillata subsp. verticillata, S. verticillata subsp. amasiaca) or granulate-scabrate- gemmate (e.g. S. bracteata) (Figures 47, 48, 52, 56, 57, 59, 65, 66). In polar view, two of the mesocolpia are larger than the other four. The apocolpium diameter (Ap) varies from 3.36 µm, in S. absconditiflora, S. verticillata subsp. amasiaca, to 13.96 µm, in S. macrosiphon (Table 9).

Figure 39. LM micrographs of pollen grains in the Salvia taxa examined: a-b. S. macrochlamys, c-d. S. pachystachys, e-f. S. tomentosa, g-h. S. trichoclada, i-j. S. euphratica var. euphratica, k-l. S. kronenburgii, m-n. S. atropatana, o-p. S. brachyantha (a, c, e, g, i, k, m, o: Equatorial view; b, d, f, h, j, l, p: Polar view). Scale bar = 20 µm.

Figure 40. LM micrographs of pollen grains in the Salvia taxa examined: a-b. S. hypargeia, c-d. S. indica, e-f. S. palaestina, g-h. S. glutinosa, i-j. S. dicroantha, k-l. S. staminea, m-n. S. russellii, o-p. S. verticillata subsp. verticillata (a, c, e, g, i, k, m, o: Equatorial view; b, d, f, h, j, l, p: Polar view). Scale bar = 20 µm.

3.3.3. Exine Sculpturing

The range of the exine thickness (Ex) is between 0.72-2.16 µm whereas the range of the intine thickness (In) is 0.24-1.20 µm (Table 10). Among the taxa examined, the thinnest exine, on average, is observed in S. caespitosa and the thickest exine, on average, is observed in S. candidissima subsp. candidissima. The thinnest intine, on average, is

observed in S. blepharochleana and S. euphratica var. euphratica and the thickest exine, on average, is observed in S. pachystachys.

In examinations of the exine sculpturing with SEM, three basic types of surface structures can be distinguished: reticulate-perforate, reticulate-granulate and bireticulate (the most frequent type), respectively. Based on the detailed configuration of the exine sculpturing both reticulate-perforate and bireticulate patterns can be subdivided into two subtypes.

Type 1: Reticulate-perforate

The reticulate-perforate sculpturing pattern occurs in 26 taxa and can be divided into two subtypes according to the number of perforations.

The number of perforation is < 20 in S. hedgeana, S. huberi, S. pinnata, S. rosifolia, S. absconditiflora, S. hydrangea, S. multicaulis, S. pseudeuphratica, S. brachyantha, S. montbretii, S. syriaca, S. virgata, S. russellii, S. verticillata subsp. verticillata and S. verticillata subsp. amasiaca (Type 1a, Figures 41-47). S. verticillata subsp. verticillata and S. verticillata subsp. amasiaca also possess holes on their colpus membrane ornamentation (Figure 47). Moreover, thickness of the primary and secondary muri is rather similar in S. russellii.

Figure 41. SEM micrographs showing pollen morphology of S. huberi (Type 1a). 89

Figure 42. SEM micrographs showing pollen morphology of S. rosifolia (Type 1a).

Figure 43. SEM micrographs showing pollen morphology of S. absconditiflora (Type 1a).

Figure 44. SEM micrographs showing pollen morphology of S. multicaulis (Type 1a).

Figure 45. SEM micrographs showing pollen morphology of S. syriaca (Type 1a).

Figure 46. SEM micrographs showing pollen morphology of S. virgata (Type 1a).

Figure 47. SEM micrographs showing pollen morphology of S. verticillata subsp. verticillata (Type 1a).

The number of perforation is ≥ 20 in S. bracteata, S. caespitosa, S. divaricata, S. kurdica, S. macrochlamys, S. pilifera, S. tomentosa, S. trichloclada, S. cerino-pruinosa, S. frigida and S. hypargeia (Type 1b; Figures 48-51). Furthermore, S. bracteata and S. trichoclada have undulate muri surfaces (Figures 48-50).

Figure 48. SEM micrographs showing pollen morphology of S. bracteata (Type 1b).

Figure 49. SEM micrographs showing pollen morphology of S. macrochlamys (Type 1b).

Figure 50. SEM micrographs showing pollen morphology of S. trichoclada (Type 1b).

Figure 51. SEM micrographs showing pollen morphology of S. hypargeia (Type 1b).

Type 2: Reticulate-granulate

The reticulate-granulate sculpturing pattern is recognized only in S. ballsiana (Figure 52). Lumina vary from 0.45 µm to 1.70 µm in diameter and more or less rounded. Muri thickness ranges between 0.20 µm and 0.60 µm (Table 10).

Figure 52. SEM micrographs showing pollen morphology of S. ballsiana (Type 2).

Type 3: Bireticulate

The bireticulate sculpturing pattern is the most common (32) among the taxa examined. Primary lumina are continuous and vary in shape from irregular ellipses to polygons. Secondary lumina are ≥ 1 µm in length. The size and number of the secondary lumina decrease towards the poles and apertures. Two subtypes can be defined based on the number of secondary lumina per primary lumen.

The number of secondary lumina is < 10 per primary lumen in S. suffruticosa, S. blepharochleana, S. aethiopis, S. eriophora, S. microstegia, S. odontochlamys and S. glutinosa (Type 3a; Figures 53-56). In S. eriophora, S. microstegia and S. odontochlamys, the secondary reticulum is also composed of one or several large secondary lumina in the middle of the primary lumen. Additionally, S. aethiopis (Figure 54), S. eriophora, S. microstegia and S. glutinosa (Figure 56), have the primary muri with more than twice as thick as the secondary muri.

Figure 53. SEM micrographs showing pollen morphology of S. blepharochleana (Type 3a).

Figure 54. SEM micrographs showing pollen morphology of S. aethiopis (Type 3a).

Figure 55. SEM micrographs showing pollen morphology of S. eriophora (Type 3a).

Figure 56. SEM micrographs showing pollen morphology of S. glutinosa (Type 3a).

The number of secondary lumina is ≥ 10 per primary lumen in S. anatolica, S. pachystachys, S. recognita, S. euphratica var. euphratica, S. euphratica var. leiocalycina, S. kronenburgii, S. viridis, S. atropatana, S. candidissima subsp. candidissima, S. ceratophylla, S. cilicica, S. cyanescens, S. indica, S. limbata, S. longipedicellata, S. macrosiphon, S. palaestina, S. poculata, S. sclarea, S. siirtica, S. spinosa, S. xanthocheila, S. dicroantha, S. nemorosa and S. staminea (Type 3b, Figures 57-67). Moreover, S. viridis, S. atropatana, S. candidissima subsp. candidissima, S. ceratophylla, S. cilicica, S. cyanescens, S. indica, S. limbata, S. longipedicellata, S. macrosiphon, S. palaestina, S. poculata, S. sclarea, S. siirtica, S. spinosa, S. xanthocheila, S. staminea have one or several large secondary lumina in the middle of the primary lumen (Figures 59-64, 67). In S. atropatana, S. candidissima subsp. candidissima, S. ceratophylla, S. cilicica, S. cyanescens, S. indica, S. limbata, S. macrosiphon, S. palaestina, S. sclarea, S. siirtica, S. spinosa, S. xanthocheila and S. staminea, the primary muri are also twice thicker than the secondary muri (Figures 60- 64, 67).

Figure 57. SEM micrographs showing pollen morphology of S. pachystachys (Type 3b).

Figure 58. SEM micrographs showing pollen morphology of S. euphratica var. leiocalycina (Type 3b).

Figure 59. SEM micrographs showing pollen morphology of S. viridis (Type 3b).

Figure 60. SEM micrographs showing pollen morphology of S. cilicica (Type 3b).

Figure 61. SEM micrographs showing pollen morphology of S. cyanescens (Type 3b).

Figure 62. SEM micrographs showing pollen morphology of S. indica (Type 3b).

Figure 63. SEM micrographs showing pollen morphology of S. limbata (Type 3b).

Figure 64. SEM micrographs showing pollen morphology of S. palaestina (Type 3b).

Figure 65. SEM micrographs showing pollen morphology of S. dicroantha (Type 3b).

Figure 66. SEM micrographs showing pollen morphology of S. nemorosa (Type 3b).

Figure 67. SEM micrographs showing pollen morphology of S. staminea (Type 3b).

3.3.4. Infrageneric Pollen Descriptions of Salvia

Sect. Salvia: Pollen grains are hexacolpate to octacolpate, suboblate to subprolate, with a polar axis diameter of 32.64-64.80 µm and an equatorial axis diameter of 35.52-66.24 µm. The colpus length is 26.88-60.37 µm. The colpus width is 2.88-12.48 µm. The exine thickness is 0.72-1.92 µm. The intine thickness is 0.48-1.20 µm. The apocolpium diameter is 4.80-13.75 µm (Table 9). Reticulate-perforate, reticulate-granulate and bireticulate sculpturing patterns are observed (Table 10).

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Sect. Hymenosphace: Pollen grains are hexacolpate, suboblate to subprolate, with a polar axis diameter of 31.68-65.28 µm and an equatorial axis diameter of 34.56-66.24 µm. The colpus length is 25.92-60.48 µm. The colpus width is 4.56-10.75 µm. The exine thickness is 0.96-1.92 µm. The intine thickness is 0.24-1.20 µm. The apocolpium diameter is 3.36-11.86 µm (Table 9). The section shows two types of sculpturing: reticulate-perforate and bireticulate (Table 10).

Sect. Horminum: Pollen grains of S. viridis belonging to this section are hexacolpate, mostly suboblate to rarely prolate-spheroidal, with a polar axis diameter of 31.68-37.44 µm and an equatorial axis diameter of 37.44-47.04 µm. The colpus length is 26.88-34.56 µm. The colpus width is 6.72-9.12 µm. The exine thickness is 0.96-1.44 µm. The intine thickness is 0.50-0.96 µm. The apocolpium diameter is 6.72-8.64 µm (Table 9). This section has the bireticulate sculpturing pattern with one or several central lumina in the primary lumen (Table 10).

Sect. Aethiopis: Pollen grains of are hexacolpate to heptacolpate, suboblate to prolate- spheroidal, with a polar axis diameter of 24.96-62.40 µm and an equatorial axis diameter of 30.72-67.20 µm. The colpus length is 20.16-54.72 µm. The colpus width is 2.64- 12.00 µm. The exine thickness is 1.17-2.16 µm. The intine thickness is 0.24-0.96 µm. The apocolpium diameter is 4.56-13.96 µm (Table 9). The section show reticulate- perforate and bireticulate sculpturing patterns. Most of the taxa in the section possess one or several large lumina in the middle of the primary lumen and the primary muri with more than twice as thick as the secondary muri (Table 10).

Sect. Drymosphace: Pollen grains of S. glutinosa belonging to this section are hexacolpate, mostly suboblate to rarely prolate-spheroidal, with a polar axis diameter of 39.36-45.12 µm and an equatorial axis diameter of 48.00-53.76 µm. The colpus length is 34.56-40.32 µm. The colpus width is 5.76-8.64 µm. The exine thickness is 1.44-1.92 µm. The intine thickness is 0.48-0.96 µm. The apocolpium diameter is 6.24-8.64 µm (Table 9). The species show the bireticulate type of sculpturing (Table 10).

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Sect. Plethiosphace: Pollen grains are hexacolpate, suboblate to prolate-spheroidal, with a polar axis diameter of 26.88-47.04 µm and an equatorial axis diameter of 29.76-54.72 µm. The colpus length is 21.12-42.24 µm. The colpus width is 3.84-10.56 µm. The exine thickness is 1.20-1.92 µm. The intine thickness is 0.48-0.96 µm. The apocolpium diameter is 3.84-8.64 µm (Table 9). The section shows reticulate-perforate and bireticulate sculpturing patterns (Table 10).

Sect. Hemisphace: Pollen grains are hexacolpate, suboblate to oblate-spheroidal, with a polar axis diameter of 23.04-37.44 µm and an equatorial axis diameter of 26.88-41.28 µm. The colpus length is 20.16-32.64 µm. The colpus width is 3.35-8.64 µm. The exine thickness is 0.83-1.68 µm. The intine thickness is 0.48-0.72 µm. The apocolpium diameter is 3.36-6.96 µm (Table 9). The section shows the reticulate-perforate type of sculpturing (Table 10).

3.3.5. Systematic Implications of Pollen Characteristics

There is variation among the studied taxa in some pollen characters such as pollen size and shape, exine sculpturing, the number of perforations, the number and size of secondary lumina, the presence of one or several central large lumina in the primary lumen, the shape of primary lumen and the ratio of the muri thickness. The observed variation appears to have systematic significance particularly at species level, including morphologically related species. Nevertheless, these characters are of restricted value for infrageneric delimitation of Salvia. For example, the section Hemisphace can be distinguished from the other sections (P x E = 30.69-59.58 x 34.43-61.55 µm, on average) by its smaller pollen grains (P x E = 26.75-30.27 x 30.66-34.27 µm, on average). S. verticillata subsp. verticillata and S. verticillata subsp. amasiaca in this section are also characterized by holes on colpus membrane sculpturing. Furthermore, S. viridis in sect. Horminum, S. staminea in sect. Plethiosphace and most of the species in sect. Aethiopis differ from the members of the other sections in having one or several large lumina in the middle of the primary lumen. As well as, S. ballsiana in sect. Salvia is clearly distinct from the other taxa examined by a reticulate-granulate sculpturing.

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A diagnostic key are given using pollen characters. From pollen micromorphological point of view, it is not possible to find any significant difference among some species. Therefore, in the diagnostic key presented below, these species are represented in one branch of the key.

1. Average pollen size (P x E) 26.75-30.27 x 30.66-34.27 µm…………..……………………2 2. Small holes on colpus membrane sculpturing present……………….…………………… ………………...……………………S. verticillata subsp. amasiaca & subsp. verticillata 2. Small holes on colpus membrane sculpturing absent………...…………………S. russellii 1. Average pollen size (P x E) 30.69-59.58 x 34.43-61.55 µm………..………………………3 3. Type of exine sculpturing reticulate-granulate………………..………………S. ballsiana 3. Type of exine sculpturing reticulate-perforate or bireticulate………….…………………4 4. Type of exine sculpturing reticulate-perforate………………..………………………5 5. Perforation number less than 20………………………...…………………………6 6. Muri surface undulate………………………………………………S. montbretii 6. Muri surface smooth……………….……..……………………………………7 7. Suboblate pollen grains present…………….…………………………...……8 8. Apocolpium diameter 8.64-11.52 µm………....………..…………S. rosifolia 8. Apocolpium diameter 3.36-7.20 µm………………….……….……………9 9. Intine thickness 0.24-0.48 µm………………..…………S. absconditiflora 9. Intine thickness 0.48-0.96 µm………………………………………….10 10. Thickness of the secondary muri 25-45 µm………..………S. hedgeana 10. Thickness of the secondary muri 11-20 µm………..……S. brachyantha 7. Suboblate pollen grains absent………………………………………………11 11. Prolate-spheroidal pollen grains absent……………………………S. virgata 11. Prolate-spheroidal pollen grains present………………………………….12 12. Apocolpium diameter 8.64-11.52 µm…………………………S. pinnata 12. Apocolpium diameter 4.56-8.50 µm………………….………………13 13. Exine thickness 0.96-1.44 µm……………………………………14 14. Oblate-spheroidal pollen grains dense……...…S. pseudeuphratica

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14. Oblate-spheroidal pollen grains rare……….……………S. huberi 13. Exine thickness 1.44-1.92 µm……………………………………15 15. Thickness of the secondary muri 0.14-0.20 µm…………S.syriaca 15. Thickness of the secondary muri 0.20-0.36 µm………………..16 16. Oblate-spheroidal pollen grains dense………..…S. hydrangea 16. Oblate-spheroidal pollen grains rare……………S. multicaulis 5. Perforation number 20 or more…………………………………………………..17 17. Muri surface undulate…………………………………..……………………18 18. Apocolpium diameter 5.52-7.22 µm……………………..………S. bracteata 18. Apocolpium diameter 9.60-12.48 µm………………………….S. trichoclada 17. Muri surface smooth…………………………………………………………19 19. Subprolate pollen grains present……………………………S. macrochlamys 19. Subprolate pollen grains absent…………………………………………...20 20. Prolate-spheroidal pollen grains present………………………………21 21. Apocolpium diameter 10.73-13.75 µm……………………S. kurdica 21. Apocolpium diameter 5.52-9.82 µm……….………..……………22 22. Exine thickness 0.72-1.20 µm…………………...…S. caespitosa 22. Exine thickness 1.44-1.92 µm…………………...……………23 23. Intıne thickness 0.48-0.72 µm…………………...S. hypargeia 23. Intıne thickness 0.72-0.92 µm…………………...S. divaricata 20. Prolate-spheroidal pollen grains absent……………………………….24 24. Apocolpium diameter 8.86-11.86 µm…………………………….25 25. Intine thickness 0.72-1.20 µm…………………………S. pilifera 25. Intıne thickness 0.50-0.72 µm…………………S. cerino-pruinosa 24. Apocolpium diameter 4.80-8.64 µm………………………………26 26. Suboblate pollen grains dense…………………………S. frigida 26. Suboblate pollen grains rare………………..………S. tomentosa 4. Type of exine sculpturing bireticulate………………………………………………27 27. Secondary lumen number less than 10…………………………………………28 28. Large secondary lumina in middle of the primary lumen present……………29

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29. The primary muri less than twice as thick as the secondary muri……..…… .………………………………………………………S. odontochlamys 29. The primary muri more than twice as thick as the secondary muri………30 30. Colpus width 7.20-9.84 µm……………………………S. microstegia 30. Colpus width 4.80-6.72 µm……………………………..S. eriophora 28. Large secondary lumina in the midle of the primary lumen absent……….…31 31. The primary muri less than twice as thick as the secondary muri………..32 32. Polar axis 49.92-61.44 µm, intine thickness 0.24-0.48 µm……………. …………………………………………………S. blepharochleana 32. Polar axis 34.56-48.96 µm, intine thickness 0.64-0.72 µm……………. ………………………………………………………S. suffruticosa 31. The primary muri more than twice as thick as the secondary muri………33 33. Thickness of the secondary muri 0.20-0.30 µm……………S. glutinosa 33. Thickness of the secondary muri 0.10-0.20 µm……………S. aethiopis 27. Secondary lumen number 10 or more………………………………………….34 34. Large secondary lumina in middle of the primary lumen absent…………….35 35. Suboblate pollen grains present………………………………………..36 36. Apocolpium diameter 9.60-10.56 µm…………………S. pachystachys 36. Apocolpium diameter 4.80-7.68 µm……………………………….37 37. Prolate-spheroidal pollen grains absent………………..S. anatolica 37. Prolate-spheroidal pollen grains present…….………..S. dicroantha 35. Suboblate pollen grains absent…………………………………………38 38. Apocolpium diameter 9.60-11.52 µm…………………….S. recognita 38. Apocolpium diameter 4.80-9.60 µm……………………………….39 39. Polar axis 26.88-38.40 µm………….………………...S. nemorosa 39. Polar axis 38.40-65.28 µm……………………………………..40 40. Subprolate pollen grains present…S. euphratica var. euphratica 40. Subprolate pollen grains absent……………………………… …………….S. euphratica var. leiocalycina & S. kronenburgii 34. Large secondary lumina in middle of the primary lumen present……………41 41. The primary muri less than twice as thick as the secondary muri………..42

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42. Thickness of the primary muri 0.30-0.40 µm, exine thickness 1.44-1.92 µm………………………………………………S. longipedicellata 42. Thickness of the primary muri 0.20-0.30 µm, exine thickness 0.96-1.44 µm………………………….…………………………….S. viridis 41. The primary muri more than twice as thick as the secondary muri………43 43. Suboblate pollen grains absent…………………………………….44 44. Thickness of the primary muri 0.08-0.15 µm…………S. palaestina 44. Thickness of the primary muri 0.16-0.25 µm…………..S. poculata 43. Suboblate pollen grains present……………………………………45 45. Apocolpium diameter 9.89-13.86 µm...S. macrosiphon & S. spinosa 45. Apocolpium diameter 3.84-9.81 µm…………………………...46 46. Prolate-spheroidal pollen grains present……………………47 47. Apocolpium diameter 6.73-9.81 µm…………….S. siirtica 47. Apocolpium diameter 3.84-6.73 µm……………………48 48. Suboblate pollen grains rare……………S. xanthocheila 48. Suboblate polen grains dense………………S. staminea 46. Prolate-spheroidal pollen grains absent…………………….49 49. Oblate-spheroidal pollen grains absent…………...S. indica 49. Oblate-spherodal pollen grains present…………………50 50. Prolate-spheroidal pollen grains dense………S. limbata 50. Prolate-spheroidal pollen grains rare…….S. atropatana, .…S. cilicica, S. cyanescens, S. sclarea, S. ceratophylla, ………………….S. candidissima subsp. candidissima

Cantino et al. (1992) revised the classification of certain genera in the Lamiaceae on the basis of palynological features and placed Salvia within the subfamily Nepetoideae since it had hexacolpate pollen grains. The present study revelas that the studied taxa mainly possess hexacolpate pollen grains, sometimes intermixed with heptacolpate or octacolpate grains in several species, such as S. kurdica, S. candidissima subsp. candidissima, S. siirtica and S. recognita.

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Henderson et al. (1968) using only LM sorted pollen grain types of Salvia and its allies into somewhat arbitrary groups. They divided them into nine types, respectively Salvia type, Salvia leucophylla type, Salvia apiana type, Salvia japonica type, Salvia nipponica type, Salvia hirtella type, Horminum type, Dorystoechas type and Salviastrum type. The Salvia type was characterized to have prolate to spheroidal pollen grains (30–49 µm) and unequal mesocolpia four of which were smaller and the other two larger. The exine thickness was also determined 1.5 µm. The ektexine was much thicker than the endexine; muri were simplibaculate. This Salvia type of pollen grains was further divided into four subtypes including species with scattered perforations in the tectum of each lumen (their type 1a), with one or two bacula occasionally, or none in each reticular loculus (their type 1b), with only marginal perforations in the membrane of each reticular loculus (their type 1c) and with each major reticular loculus with secondary reticulum (their type 1d). Included in this analysis were fifteen taxa investigated also in the present study. Henderson et al. (1968) placed S. caespitosa, S. hydrangea, S. macrochlamys, S. montbretii, S. pilifera, S. rosifolia, S. suffruticosa, S. tomentosa (syn. S. grandiflora), S. trichloclada and S. verticillata in their Salvia type 1a and S. brachyantha, S. glutinosa, S. sclarea, S. virgata and S. viridis (syn. S. horminum) in their Salvia type 1b. Conforming with their findings the present work also shows that S. brachyantha, S. caespitosa, S. hydrangea, S. macrochlamys, S. montbretii, S. pilifera, S. rosifolia, S. tomentosa, S. trichloclada, S. verticillata and S. virgata have perforations in each lumen. However, the closer examination gave evidence that S. glutinosa, S. sclarea, S. suffruticosa and S. viridis have a bireticulate sculpturing pattern. This discrepancy can be easily explained by the lower magnification used by Henderson et al. (1968) that was insufficient showing details of exine sculpturing pattern.

Henderson et al. (1968) reported that in pollen grains of a few Salvia species the two larger mesocolpia differed in the exine structure from the four smaller mesocolpia. Similarly, heteromorphic exine ornamentation was recorded in Agastache scrophulariifolia by Moon et al. (2008b) and in Nepeta sibthorpii subsp. tumeniana by Celenk et al. (2008a). However, we have not observed this phenomenon in the taxa studied here.

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Table 9. Pollen characteristics in the taxa studied of Salvia based on LM. All sizes are in µm. Numbers refer to minimum-maximum (mean±standard deviation). Os = Oblate-spheroidal, Ps = Prolate-spheroidal, So = Suboblate, Sp = Subprolate. -, absent, +, rarely present, ++, densely present.

Taxa and sections Polar axis (P) Equatorial Shape Colpus length Colpus width Exine (Ex) Intine (In) Apocolpium axis (E) (Clg) (Clt) (Ap) So Os Ps Sp Sect. Salvia S. anatolica 38.40-44.16 45.12-52.80 ++ + - - 33.60-39.36 6.72-9.12 1.20-1.68 0.48-0.96 4.80-6.72 (41.61±1.37) (49.40±2.05) (37.27±1.49) (7.89±0.64) (1.53±0.17) (0.70±0.17) (5.54±0.57) S. ballsiana 44.16-56.64 51.84-63.36 + ++ - - 37.44-49.92 6.72-12.48 0.72-1.44 0.48-0.96 10.56-13.44 (51.90±3.63) (57.51±3.42) (44.41±3.53) (9.55±1.57) (1.19±0.22) (0.77±0.14) (11.71±0.92) S. bracteata 38.40-52.32 34.72-48.00 - + ++ - 32.64-43.20 5.76-7.58 0.96-1.44 0.48-0.72 5.52-7.20 (48.38±2.55) (43.35±2.19) (38.62±2.09) (6.63±0.53) (1.23±0.17) (0.60±1.11) (6.13±0.48) S. caespitosa 39.36-56.64 42.24-58.56 ++ ++ + - 28.80-51.84 6.72-10.56 0.72-1.20 0.48-0.96 6.72-9.50 (45.05±4.29) (51.28±3.69) (39.82±5.17) (9.26±0.90) (0.95±0.15) (0.72±0.16) (8.21±0.91) 108 S. divaricata 45-75-55.00 49.20-55.63 - ++ + - 39.65-49.36 6.00-9.51 1.47-1.90 0.72-0.92 6.89-9.82 (49.78±3.59) (52.75±2.29) (42.68±3.50) (7.89±1.16) (1.75±0.21) (0.78±0.08) (7.60±1.02)

S. hedgeana 34.56-48.00 41.28-52.80 ++ ++ + - 31.68-42.24 5.76-7.68 1.44-1.92 0.48-0.72 5.76-7.20 (39.49±2.61) (46.57±2.30) (35.43±1.31) (6.62±0.43) (1.75±0.16) (0.56±0.11) (6.53±0.46) S. huberi 39.36-58.56 38.40-52.80 - + ++ - 33.60-46.08 5.76-8.64 0.96-1.44 0.48-0.72 4.80-6.72 (47.57±3.71) (43.50±3.68) (38.50±3.48) (6.64±0.77) (1.27±0:15) (0.59±0.11) (5.88±0.63) S. kurdica 53.28-64.80 51.58-64.13 - + ++ - 48.07-60.37 3.84-8.31 1.16-1.70 0.53-0.75 10.73-13.75 (59.58±3.38) (57.14±3.51) (54.81±3.46) (5.93±1.26) (1.35±0.18) (0.65±0.07) (12.22±1.23) S. macrochlamys 39.36-53.76 35.52-46.08 - + + ++ 29.76-48.00 6.72-9.12 1.68-1.92 0.72-0.96 5.76-7.68 (48.44±2.69) (41.59±2.54) (43.21±2.73) (7.95±0.69) (1.84±0.11) (0.87±0.10) (6.83±0.64) S. pachystachys 48.00-57.60 52.80-66.24 ++ + + - 41.28-50.88 8.64-12.48 0.96-1.44 0.72-1.20 9.60-10.56 (52.18±2.35) (61.55±3.18) (45.12±2.56) (10.68±1.01) (1.21±0.11) (0.91±0.18) (10.01±0.41) S. pilifera 35.52-52.80 40.32-50.88 + ++ - - 30.72-45.12 2.88-5.76 0.96-1.44 0.72-1.20 9.60-11.52 (44.85±3.66) (45.63±3.32) (38.46±3.10) (4.39±1.04) (1.13±0.17) (0.88±0.18) (10.83±0.80) S. pinnata 38.40-53.76 41.28-48.96 - + ++ - 32.64-46.08 3.84-7.68 0.96-1.44 0.65-0.96 8.64-11.52 (46.38±3.73) (45.12±2.19) (39.32±3.52) (5.77±1.06) (1.11±0.16) (0.74±0.14) (10.00±0.84) S. recognita 40.32-47.04 43.20-49.92 - ++ + - 28.80-39.36 6.72-9.60 0.72-1.20 0.48-0.96 9.60-11.52 (43.40±2.11) (47.48±1.58) (35.91±2.42) (8.41±0.85) (0.96±0.12) (0.74±0.13) (10.65±0.65) S. rosifolia 32.64-44.16 39.36-50.88 ++ + - - 26.88-38.40 3.84-9.60 0.72-1.20 0.48-1.20 8.64-11.52 (39.99±2.65) (47.37±3.03) (33.39±3.09) (7.00±1.01) (0.97±0.15) (0.72±0.15) (10.33±0.92)

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Table 9 (continued) S. suffruticosa 34.56-48.96 37.44-49.92 - ++ + - 28.80-43.20 3.84-6.72 0.72-1.20 0.64-0.72 7.20-9.60 (41.94±4.00) (45.02±2.87) (35.01±4.09) (5.26±0.81) (0.99±0.14) (0.68±0.11) (8.75±0.68) S. tomentosa 38.40-43.20 45.12-48.96 + ++ - - 33.60-39.36 7.68-9.60 1.44-1.92 0.48-0.96 5.76-8.64 (41.21±1.41) (47.19±1.15) (37.14±1.57) (8.92±0.64) (1.70±0.17) (0.76±0.17) (7.52±0.90) S. trichocloda 38.40-54.72 37.44-60.00 - ++ + - 35.52-47.04 3.84-10.56 0.96-1.44 0.48-0.96 9.60-12.48 (47.48±4.29) (47.48±4.29) (41.63±3.49) (7.38±1.62) (1.14±0.16) (0.69±0.12) (10.82±0.99) Sect. Hymenosphace S. absconditiflora 31.68-47.04 38.40-48.96 + ++ - - 25.92-41.28 6.48-8.64 1.20-1.92 0.24-0.48 3.36-6.72 (39.52±3.65) (44.58±2.70) (34.11±3.73) (7.55±0.70) (1.65±0.22) (0.41±0.11) (5.25±1.02) S. blepharochleana 49.92-61.44 45.12-65.28 + + ++ - 48.00-60.48 5.28-8.16 0.96-1.44 0.24-0.48 4.80-7.58 (56.96±2.80 (52.96±4.95) (52.93±3.20) (6.51±0.84) (1.17±0.18) (0.32±0.12) (6.39±0.94) S. cerino-pruinosa 39.75-55.12 47.62-58.32 + ++ - - 32.50-43.54 4.84-10.75 1.20-1.52 0.50-0.72 8.86-11.86 (46.18±4.31) (52.26±3.52) (39.14±5.72) (7.37±1.84) (1.30±0.12) (0.63±0.11) (9.23±1.86) S. euphratica var. 38.40-51.84 34.56-52.80 - - ++ + 34.56-46.08 4.80-6.72 1.20-1.68 0.24-0.48 5.76-7.44 euphratica (46.11±2.81) (42.75±4.21) (40.13±3.17) (5.28±0.66) (1.42±0.17) (0.32±0.12) (6.37±0.51) S. euphratica var. 40.32-51.84 42.24-61.44 - ++ + - 33.60-46.08 6.72-10.08 1.44-1.92 0.48-0.96 7.20-9.60

109 leiocalycina (46.43±3.59) (49.95±4.59) (40.03±3.83) (8.66±0.87) (1.62±0.21) (0.74±0.20) (8.38±0.85) S. hydrangea 32.64-43.20 36.48-49.92 - ++ + - 27.84-38.40 4.80-7.44 1.44-1.68 0.48-0.72 4.56-5.76 (40.42±2.20) (44.38±2.99) (35.10±1.93) (5.55±0.75) (1.57±0.12) (0.62±0.12) (5.30±0.44) S. kronenburgii 42.24-65.28 45.12-66.24 - ++ + - 36.48-59.52 6.24-9.60 1.20-1.92 0.24-0.72 5.28-7.44 (49.50±5.79) (52.13±4.91) (45.07±6.13) (7.62±0.83) (1.48±0.25) (0.53±0.19) (6.43±0.59) S. multicaulis 36.48-48.96 38.40-48.00 - + ++ - 32.64-41.28 4.56-6.72 1.44-1.92 0.48-1.20 4.80-6.96 (42.21±3.08) (41.54±2.98) (36.03±2.38) (5.64±0.56) (1.62±0.18) (0.63±0.25) (5.71±0.71) S. pseudeuphratica 39.26-59.31 44.59-65.28 - ++ + - 32.05-53.66 5.21-9.22 1.00-1.41 0.42-0.58 6.36-8.50 (47.84±7.26) (50.31±7.52) (42.40±7.50) (7.62±1.28) (1.17±0.15) (0.49±0.05) (7.54±1.09) Sect. Horminum S. viridis 31.68-37.44 37.44-47.04 ++ + + 26.88-34.56 6.72-9.12 0.96-1.44 0.50-0.96 6.72-8.64 (34.08±1.63) (42.08±2.30) (29.66±2.17) (7.72±0.67) (1.20±0.20) (0.71±0.19) (7.37±0.59) Sect. Aethiopis S. aethiopis 24.96-33.60 35.52-43.20 ++ + - - 22.08-28.80 6.24-9.60 1.20-1.68 0.48-0.96 4.80-6.72 (30.69±2.44) (39.20±2.20) (26.14±2.01) (7.65±0.92) (1.50±0.17) (0.62±0.18) (5.68±0.46) S. atropatana 38.40-48.00 46.08-58.56 ++ + - - 30.72-40.32 7.44-11.52 1.20-1.92 0.48-0.96 4.80-8.16 (42.50±2.61) (49.22±2.63) (36.83±2.66) (9.29±1.12) (1.55±0.20) (0.70±0.20) (6.77±0.92) S. brachyantha 24.96-39.36 32.64-46.08 ++ + - - 20.16-35.52 5.76-8.64 1.20-1.68 0.48-0.96 4.32-6.00 (30.78±3.81) (38.59±3.73) (25.02±3.30) (7.08±0.75) (1.51±0.17) (0.66±0.19) (4.98±0.57) S. candidissima subsp. 36.48-62.40 46.08-67.20 ++ + - - 32.64-54.72 8.64-12.00 1.68-2.16 0.48-0.96 6.48-8.64 candidissima (46.66±6.12) (59.68±4.59) (40.70±5.42) (10.21±1.01) (1.91±0.15) (0.78±0.17) (7.64±0.74)

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Table 9 (continued) S. ceratophylla 31.68-43.20 44.16-52.80 ++ + - - 24.96-36.48 7.68-10.56 1.44-1.92 0.48-0.96 5.52-8.40 (38.37±2.88) (47.46± 2.49) (32.51±3.02) (9.19±0.79) (1.60±0.20) (0.69±0.19) (6.89±0.81) S. cilicica 34.56-43.20 43.20-51.84 ++ + - - 29.76-38.40 6.72-10.56 1.44-1.92 0.48-0.96 5.28-8.16 (39.07±2.02) (47.74±2.23) (33.70±2.05) (8.43±0.99) (1.64±0.20) (0.73±0.20) (7.02±0.88) S. cyanescens 37.44-56.64 46.08-62.40 ++ + - - 31.68-48.96 6.72-9.60 1.20-1.68 0.48-0.72 5.04-6.96 (46.53±5.63) (54.30±5.20) (40.54±5.21) (8.07±0.84) (1.39±0.19) (0.65±0.11) (6.03±0.57) S. eriophora 26.88-37.44 32.64-42.24 ++ + - - 22.08-31.68 4.80-6.72 1.20-1.68 0.48-0.72 4.56-6.00 (32.58±2.75) (38.50±2.60) (27.07±2.01) (5.92±0.57) (1.40±0.19) (0.62±0.12) (5.42±0.51) S. frigida 30.72-44.16 37.44-46.08 ++ + - - 24.00-37.44 7.20-9.60 1.20-1.68 0.48-0.72 4.80-7.68 (37.25±3.11) (42.46±2.62) (31.30±3.69) (8.24±0.70) (1.43±0.18) (0.63±0.12) (5.81±0.84) S. hypargeia 38.40-48.00 42.24-52.80 ++ + + - 32.64-42.24 6.72-9.60 1.44-1.92 0.48-0.72 5.52-7.68 (42.37±3.30) (48.77±2.69) (37.18±2.99) (7.61±0.75) (1.70±0.18) (0.54±0.11) (6.70±0.72) S. indica 36.48-48.00 45.12-54.72 ++ - - - 28.80-41.28 7.20-9.60 1.44-1.92 0.48-0.96 6.72-9.36 (42.66±3.54) (49.09±2.48) (36.48±2.88) (8.54±0.78) (1.70±0.19) (0.71±0.18) (8.34±0.69) S. limbata 35.52-54.72 43.20-65.28 ++ ++ - - 29.76-47.04 7.68-10.56 1.20-1.68 0.48-0.72 5.76-6.72 (43.62±4.87) (52.70±4.88) (37.09±4.20) (9.20±0.89) (1.42±0.21) (0.57±0.12) (6.24±0.38)

110 S. longipedicellata 32.64-48.00 40.32-52.80 ++ + + - 25.92-39.36 7.20-10.56 1.44-1.92 0.48-0.96 4.56-6.96 (38.94±4.52) (46.02±3.04) (31.14±3.58) (8.51±0.75) (1.66±0.21) (0.66±0.19) (6.04±0.67)

S. macrosiphon 45.23-53.52 46.46-55.73 + ++ + - 39.52-46.93 5.25-7.80 1.33-1.70 0.50-0.84 9.89-13.96 (49.32±3.75) (52.12±3.45) (42.65±3.60) (6.30±0.88) (1.49±0.10) (0.65±0.12) (11.96±1.24) S. microstegia 32.64-50.88 39.36-60.48 ++ + - - 26.88-43.20 7.20-9.84 1.20-1.68 0.24-0.72 4.80-6.48 (41.34±6.11) (47.74±5.49) (34.66±5.28) (8.62±0.89) (1.42±0.219 (0.53±0.18) (5.44±0.43) S. montbretii 34.56-45.12 43.20-53.76 ++ + + - 28.80-40.32 7.68-11.04 1.20-1.68 0.24-0.72 6.24-9.36 (39.74±2.97) (47.71±2.81) (33.95±3.09) (8.89±0.97) (1.50±0.17) (0.50±0.19) (8.05±0.86) S. odontochlamys 38.22-46.61 44.51-47.66 - ++ - - 33.51-41.46 5.20-6.98 1.17-1.52 0.50-0.86 6.44-8.29 (42.65±3.31) (46.44±1.35) (37.38±2.99) (6.42±0.71) (1.33±0.16) (0.67±0.14) (7.17±0.71) S. palaestina 31.68-42.24 33.60-46.08 - ++ ++ - 25.92-36.48 4.08-7.68 1.20-1.68 0.24-0.72 5.52-6.72 (36.22±3.21) (39.68±3.51) (31.17±3.16) (5.78±1.18) (1.46±0.21) (0.57±0.16) (6.14±0.36) S. poculata 33.27-48.09 35.99-46.57 - + ++ - 24.94-42.57 2.64-6.77 1.25-1.82 0.63-0.86 6.07-9.11 (42.37±5.41) (43.18±3.09) (35.91±6.45) (4.11±1.32) (1.50±0.21) (0.79±0.08) (7.44±0.93) S. sclarea 38.40-47.04 47.04-59.52 ++ + - - 33.60-41.28 7.20-10.56 1.20-1.68 0.24-0.72 6.00-8.64 (42.34±2.55) (52.99±3.41) (37.22±2.48) (8.79±0.78) (1.42±0.19) (0.54±0.17) (7.42±0.84) S. siirtica 39.64-54.21 45.15-57.29 ++ ++ + - 32.51-47.71 6.51-10.45 1.35-1.90 0.52-0.86 6.73-9.81 (46.27±2.73) (50.67±2.98) (39.94±2.80) (8.62±0.88) (1.61±0.11) (0.70±0.12) (8.09±0.78) S. spinosa 38.40-51.84 47.04-58.56 ++ + + - 30.72-48.00 7.68-10.56 1.44-1.92 0.48-0.96 9.91-10.78 (42.66±3.42) (51.17±2.73) (37.28±5.39) (8.85±0.77) (1.74±0.19) (0.77±0.18) (10.32±0.34) S. syriaca 29.76-40.32 30.72-44.16 - + ++ - 24.00-35.52 6.00-8.64 1.44-1.92 0.48-0.96 4.80-7.68 (35.52±3.07) (39.10±3.17) (29.57±3.16) (7.29±0.76) (1.62±0.21) (0.64±0.18) (5.97±0.74) 110

Table 9 (continued) S. xanthocheila 38.40-51.84 39.36-53.76 + ++ + - 32.64-46.08 6.72-11.52 1.44-1.92 0.24-0.72 4.80-6.72 (43.07±3.35) (46.88±3.04) (37.12±3.30) (8.75±1.37) (1.58±0.17) (0.50±0.18) (6.00±0.50) Sect. Drymosphace S. glutinosa 39.36-45.12 48.00-53.76 ++ + + - 34.56-40.32 5.76-8.64 1.44-1.92 0.48-0.96 6.24-8.64 (42.37±1.85) (51.46±1.83) (37.47±1.66) (7.38±0.83) (1.76±0.20) (0.77±0.19) (7.55±0.79) Sect. Plethiosphace S. dicroantha 31.68-46.08 38.40-48.96 ++ ++ + - 26.88-42.24 6.00-8.64 1.20-1.68 0.48-0.96 5.76-7.68 (36.54±4.29) (41.86±2.59) (32.26±4.22) (7.28±0.85) (1.46±0.19) (0.77±0.19) (6.72±0.66) S. nemorosa 26.88-38.40 29.76-38.40 - ++ - - 21.12-32.64 3.84-8.64 1.20-1.92 0.48-0.96 4.80-6.72 (30.98±3.06) (34.43±2.35) (25.76±3.10) (6.11±1.19) (1.66±0.28) (0.73±0.17) (5.63±0.69) S. staminea 33.60-45.12 40.32-54.72 ++ + + - 28.80-40.32 7.44-10.56 1.44-1.92 0.48-0.96 3.84-5.28 (37.79±3.09) (43.90±3.48) (32.38±3.09) (8.98±0.92) (1.71±0.21) (0.76±0.18) (4.52±0.49) S. virgata 34.56-47.04 39.36-50.88 - ++ - - 29.76-42.24 6.48-8.16 1.20-1.92 0.48-0.96 5.76-8.64 (39.49±2.69) (44.35±2.93) (34.85±2.55) (7.23±0.57) (1.57±0.23) (0.76±0.17) (7.58±0.96) Sect. Hemisphace S. russellii 27.08-29.90 29.03-31.87 - ++ - - 22.75-25.58 3.35-4.90 0.83-1.17 0.48-0.71 5.02-6.96 (28.42±0.87) (30.72±1.00) (23.95±0.99) (3.67±0.80) (1.03±0.11) (0.59±0.08) (6.04±0.79)

111 S. verticillata subsp. 24.96-37.44 30.72-41.28 ++ ++ - - 21.12-32.64 5.76-8.64 0.96-1.44 0.48-0.72 3.84-6.72

amasiaca (30.27±2.64) (34.27±2.49) (24.77±2.83) (7.70±0.78) (1.28±0.17) (0.58±0.12) (5.14±0.90) S. verticillata subsp. 23.04-29.76 26.88-34.56 ++ ++ - - 20.16-25.92 4.80-8.16 0.96-1.68 0.48-0.72 3.36-4.80 verticillata (26.75±1.76) (30.66±2.05) (22.79±1.61) (6.54±1.00) (1.30±0.28) (0.63±0.12) (3.89±0.38)

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Table 10. Pollen characteristics in the taxa studied of Salvia based on SEM. All sizes are in µm. -, absent, +, present. Taxa and sections Diameter of Diameter of Thickness Thickness of Presence of large Presence of holes on Sculpturing type primary secondary of primary secondary secondary lumina in colpus membrane lumina lumina muri muri the middle of the sculpturing primary lumen Sect. Salvia S. anatolica 1.00-3.57 0.32-1.55 0.30-0.70 0.15-0.40 - - Type 3b S. ballsiana 0.45-1.70 - 0.20-0.60 - - - Type 2 S. bracteata 0.74-2.98 0.20-0.81 0.21-0.55 0.15-0.35 - - Type 1b S. caespitosa 0.90-3.43 0.22-0.97 0.25-0.41 0.15-0.29 - - Type 1b S. divaricata 0.75-3.23 0.10-0.30 0.25-0.40 0.13-0.24 - - Type 1b S. hedgeana 0.85-2.90 0.40-0.90 0.30-0.60 0.25-0.45 - - Type 1a S. huberi 0.75-2.96 0.29-0.81 0.26-0.45 0.13-0.30 - - Type 1a S. kurdica 0.95-4.32 0.23-0.79 0.23-0.42 0.14-0.25 - - Type 1b S. macrochlamys 1.00-5.22 0.10-0.61 0.30-0.50 0.17-0.35 - - Type 1b S. pachystachys 1.12-3.88 0.33-1.30 0.30-0.48 0.16-0.30 - - Type 3b 112 S. pilifera 0.70-2.59 0.16-0.60 0.19-0.45 0.14-0.31 - - Type 1b

S. pinnata 0.65-3.15 0.15-0.58 0.20-0.31 0.12-0.29 - - Type 1a S. recognita 0.97-3.76 0.30-1.40 0.30-0.75 0.16-0.43 - - Type 3b S. rosifolia 0.80-2.33 0.12-0.64 0.24-0.44 0.23-0.34 - - Type 1a S. suffruticosa 0.52-.2.16 0.20-1.10 0.28-0.54 0.11-0.29 - - Type 3a S. tomentosa 0.87-3.13 0.16-0.65 0.22-0.41 0.15-0.26 - - Type 1b S. trichoclada 0.64-2.52 0.16-0.75 0.19-0.42 0.12-0.34 - - Type 1b Sect. Hymenosphace S. absconditiflora 0.43-2.48 0.16-0.52 0.30-0.54 0.15-0.39 - - Type 1a S. blepharochleana 0.75-2.30 0.35-1.24 0.35-0.58 0.20-0.40 - - Type 3a S. cerino-pruinosa 0.64-3.42 0.20-0.90 0.17-0.40 0.12-0.35 - - Type 1b S. euphratica var. 0.60-3.27 0.23-1.13 0.23-0.60 0.17-0.38 - - Type 3b euphratica S. euphratica var. 0.94-4.07 0.25-1.31 0.25-0.75 0.23-0.45 - - Type 3b leiocalycina S. hydrangea 0.61-3.17 0.22-0.83 0.28-0.44 0.20-0.36 - - Type 1a S. kronenburgii 1.07-3.58 0.31-1.19 0.22-0.35 0.16-0.31 - - Type 3b S. multicaulis 0.50-2.01 0.12-0.49 0.25-0.40 0.20-0.30 - - Type 1a

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Table 10 (continued) S. pseudeuphratica 0.50-1.71 0.15-0.49 0.20-0.40 0.18-0.34 - - Type 1a Sect. Horminum S. viridis 0.99-3.05 0.20-1.55 0.20-0.30 0.11-0.20 + - Type 3b Sect. Aethiopis S. aethiopis 0.53-1.66 0.20-1.13 0.32-0.45 0.10-0.20 - - Type 3a S. atropatana 0.75-2.70 0.15-1.05 0.43-0.70 0.10-0.20 + - Type 3b S. brachyantha 0.50-1.75 0.15-0.98 0.35-0.62 0.11-0.20 + Type 1a S. candidissima 0.85-3.49 0.21-1.72 0.32-0.53 0.12-0.22 + - Type 3b subsp. candidissima S. ceratophylla 0.93-3.42 0.40-1.60 0.40-0.72 0.10-0.20 + - Type 3b S. cilicica 0.61-3.59 0.20-1.86 0.35-0.60 0.10-0.20 + - Type 3b S. cyanescens 1.00-3.70 0.15-1.39 0.35-0.50 0.11-0.21 + - Type 3b S. eriophora 0.50-1.98 0.15-1.35 0.25-0.50 0.09-0.20 + - Type 3a S. frigida 0.61-3.54 0.16-0.79 0.25-0.45 0.15-0.25 - - Type 1b S. hypargeia 0.85-4.33 0.14-0.71 0.28-0.39 0.17-0.29 - - Type 1b 113 S. indica 0.95-4.03 0.30-1.55 0.35-0.53 0.09-0.21 + - Type 3b

S. limbata 1.09-4.07 0.30-1.73 0.30-0.40 0.11-0.18 + - Type 3b S. longipedicellata 0.91-2.71 0.15-1.17 0.30-0.40 0.17-0.25 + - Type 3b S. macrosiphon 1.29-3.70 0.25-1.20 0.35-0.50 0.15-0.24 + - Type 3b S. microstegia 0.70-2.85 0.28-1.46 0.30-0.50 0.10-0.17 + - Type 3a S. montbretii 0.76-2.93 0.20-0.80 0.30-0.40 0.18-0.31 - - Type 1a S. odontochlamys 0.80-2.49 0.25-1.10 0.30-0.40 0.16-0.23 + - Type 3a S. palaestina 1.66-4.32 0.25-2.05 0.35-0.50 0.08-0.15 + - Type 3b S. poculata 0.67-2.94 0.20-1.64 0.35-0.45 0.16-0.25 + - Type 3b S. sclarea 1.00-4.20 0.18-1.30 0.32-0.42 0.10-0.20 + - Type 3b S. siirtica 1.90-4.96 0.40-2.37 0.30-0.55 0.08-0.20 + - Type 3b S. spinosa 1.33-3.88 0.25-1.43 0.36-0.54 0.14-0.26 + - Type 3b S. syriaca 0.50-2.71 0.22-0.63 0.25-0.37 0.14-0.20 - - Type 1a S. xanthocheila 0.80-4.58 0.30-1.96 0.43-0.65 0.10-0.18 + - Type 3b Sect. Drymosphace S. glutinosa 0.60-2.06 0.25-1.00 0.42-0.60 0.20-0.30 - - Type 3a Sect. Plethiosphace S. dicroantha 0.66-2.67 0.30-1.02 0.25-0.38 0.16-0.24 - - Type 3b 113

Table 10 (continued) S. nemorosa 0.70-3.65 0.23-1.20 0.29-0.41 0.15-0.27 - - Type 3b S. staminea 0.67-2.70 0.15-1.80 0.46-0.70 0.10-0.19 + - Type 3b S. virgata 0.63-2.78 0.17-0.67 0.30-0.41 0.20-0.30 - - Type 1a Sect. Hemisphace S. russelii 0.40-2.05 0.15-0.73 0.20-0.30 0.18-0.30 - - Type 1a S. verticillata subsp. 0.50-2.56 0.11-0.56 0.20-0.25 0.12-0.22 - + Type 1a amasiaca S. verticillata subsp. 0.53-2.61 0.15-0.66 0.20-0.28 0.11-0.23 - + Type 1a verticillata

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3.4. Mericarp Micromorphology

Though the mericarp (nutlet) micromorphology are investigated by many researchers (e.g. Husain et al., 1990; Rejdali, 1990; Marin et al., 1994; Turner & Delprete, 1996; Guerin, 2005; Moon & Hong, 2006; Salmaki et al., 2008b), data on the mericarp micromorphology of representatives of Salvia are rather limited and mostly restricted to taxonomic treatments (Marin et al., 1996; Oran, 1996; Özkan et al., 2009; Bagherpour et al., 2010; Kahraman & Doğan, 2010; Kahraman et al., 2010b, c; Büyükkartal et al., 2011; Kahraman et al., 2011a, b). Marin et al. (1996) and Oran (1996) found that gross morphology of mericarps and surface sculpturing in some species of Salvia were variable and taxonomically useful. Kahraman et al. (2011a) investigated Salvia sect. Hymenosphace and pointed out significant variation in mericarp shape and size, the nature of the abscission scar, shape of transverse sections, surface sculpturing and exocarp cell shape was sufficient to distinguish taxa at species level, including morphologically similar species.

The results obtained from micromorphological investigations are described below and illustrated in Figures 68-83. The main mericarp characteristics for the taxa examined, such as mericarp size, shape and colour, diameter of abscission scar and nutlet surface sculpturing type, are provided in Tables 11 and 12.

3.4.1. Size, Shape and Colour

Mericarps are bilaterally symmetric based on the position of the abscission (=attachment) scar. They range from 1.52 mm (S. nemorosa) to 5.38 mm in length (S. euphratica var. euphratica) and from 1.02 mm (S. verticillata subsp. verticillata) to 4.70 mm in width (S. cerino-pruinosa). They range in length to width ratio from 1.01 (S. bracteata and S. kurdica) to 1.88 (S. russellii) (Table 11).

Shape of mericarps are near spherical (e.g. S. kurdica, S. pseudeuphratica, S. indica) or prolate-spheroidal (the most common shape; e.g. S. viridis, S. longipedicellata, S.

115

nemorosa, S. verticillata subsp. verticillata, S. verticillata subsp. amasiaca). Shape of transverse section of mericarps are rounded (e.g. S. pinnata, S. pseudeuphratica) or rounded-trigonous (e.g. S. rosifolia, S. palaestina, S. virgata). When mericarps were not fullymatured, their colour was greenish at first, and then turned to dark or light brown (e.g. S. bracteata, S. absconditiflora, S. frigida, S. staminea, S. russellii) or black (e.g. S. divaricata, S. pilifera, S. euphratica var. euphratica, S. euphratica var. leiocalycina) (Table 12).

3.4.2. Abscission Scar Diameter and Shape

Abscission scars are located either at the center of the basal end or slightly shifted towards the ventral side. Diameter of the abscission scars vary between 0.15 mm (S. verticillata subsp. verticillata) and 1.28 mm (S. kronenburgii) (Table 11). Their shape is rounded (e.g. S. caespitosa, S. muticaulis, S. dicroantha), U-shaped (e.g. S. atropatana, S. hypargeia, S. syriaca), V-shaped (e.g. S. viridis, S. frigida, S. glutinosa) or squared (e.g. S. nemorosa) (Table 12).

3.4.3. Surface Sculpturing

Mericarp surface presents colliculate (small hill-like eminences, spaced, covering throughout the mericarp surface), reticulate (a reticulum or net and interspaces characterized with raised walls), verrucate (irregular projections or knobs) and rugose sculpturing patterns (wrinkled, the irregular elevations making up the wrinkles and running mostly in one direction), mostly as a result of the sculpturing of the exocarp cell walls with the pattern determined by whether the periclinal walls were concave or convex and whether the anticlinal walls were raised or sunken.

Type 1: Colliculate

The colliculate sculpturing pattern is present 37 taxa and can be divided into three subtypes based on shape of exocarp cells. 116

Pentangular or hexangular exocarp cells are present in S. anatolica, S. ballsiana, S. bracteata, S. caespitosa, S. hedgeana, S. macrochlamys, S. pachystachys, S. pilifera, S. tomentosa, S. trichoclada, S. blepharochleana, S. euphratica var. leiocalycina, S. hydrangea, S. aethiopis, S. brachyantha, S. candidissima subsp. candidissima, S. cyanescens, S. frigida, S. longipedicellata, S. macrosiphon, S. spinosa, S. syriaca and S. staminea (Figures 68-73).

Rounded exocarp cells are observed in S. divaricata, S. rosifolia, S. suffruticosa, S. absconditiflora and S. kronenburgii (Figures 74, 75).

Irregular shaped exocarp cells are recognized in S. huberi, S. cilicica, S. hypargeia, S. limbata, S. odontochlamys, S. palaestina, S. poculata, S. sclarea and S. russellii (Figures 76, 77).

Figure 68. SEM micrographs showing mericarp morphology of S. ballsiana (Type 1).

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Figure 69. SEM micrographs showing mericarp morphology of S. caespitosa (Type 1).

Figure 70. SEM micrographs showing mericarp morphology of S. macrochlamys (Type 1).

Figure 71. SEM micrographs showing mericarp morphology of S. pilifera (Type 1).

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Figure 72. SEM micrographs showing mericarp morphology of S. euphratica var. leiocalycina (Type 1).

Figure 73. SEM micrographs showing mericarp morphology of S. aethiopis (Type 1).

Figure 74. SEM micrographs showing mericarp morphology of S. rosifolia (Type 1).

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Figure 75. SEM micrographs showing mericarp morphology of S. kronenburgii (Type 1).

Figure 76. SEM micrographs showing mericarp morphology of S. huberi (Type 1).

Figure 77. SEM micrographs showing mericarp morphology of S. russellii (Type 1).

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Type 2: Reticulate

The reticulate sculpturing pattern is observed in 10 species as in S. kurdica, S. pinnata, S. pseudeuphratica, S. viridis, S. atropatana, S. indica, S. microstegia, S. montbretii, S. siirtica and S. glutinosa (Figures 78-80)

Figure 78. SEM micrographs showing mericarp morphology of S. pinnata (Type 2).

Figure 79. SEM micrographs showing mericarp morphology of S. atropatana (Type 2).

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Figure 80. SEM micrographs showing mericarp morphology of S. indica (Type 2).

Type 3: Verrucate

The verrucate sculpturing pattern is observed in 8 taxa as in S. recognita, S. cerino- pruinosa, S. euphratica var. euphratica, S. multicaulis, S. xanthocheila, S. nemorosa, S. verticillata subsp. amasiaca and S. verticillata subsp. verticillata (Figures 81, 82).

Figure 81. SEM micrographs showing mericarp morphology of S.recognita (Type 3).

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Figure 82. SEM micrographs showing mericarp morphology of S. euphratica var. euphratica (Type 3).

Type 4: Rugose

The rugose sculpturing pattern is present in 3 species as in S. ceratophylla, S. dicroantha and S. virgata (Figure 83).

Figure 83. SEM micrographs showing mericarp morphology of S. ceratophylla (Type 4).

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3.4.4. Infrageneric Mericarp Descriptions of Salvia

Sect. Salvia: Mericarps range from 2.46 to 5.26 mm in length and from 1.83 to 4.50 mm in width. Lenthg to width ratio varies between 1.01 and 1.60. Shape of the mericarps is mostly prolate-spheroidal and shape of their transverse sections is rounded-trigonous in all almost species except for S. kurdica and S. pinnata. The colour is mostly black or rarely brown. Diameter of abscission scars ranges from 0.41 to 1.25 mm and their shape is rounded. Regarding sculpturing pattern of the mericap surface, three types can be distinguished: colliculate, reticulate and verrucate (Tables 11, 12).

Sect. Hymenosphace: Mericarps range from 2.50 to 5.38 mm in length and from 2.04 to 4.70 mm in width. Lenthg to width ratio varies between 1.02 and 1.40. Shape of the mericarps is mostly prolate-spheroidal and shape of their transverse sections is rounded to rounded-trigonous. The colour is mostly black or rarely brown. Diameter of abscission scars ranges from 0.70 to 1.28 mm and their shape is rounded. Regarding sculpturing pattern of the mericap surface, three types can be distinguished: colliculate, reticulate and verrucate (Tables 11, 12).

Sect. Horminum: Mericarps range from 2.85 to 3.34 mm in length and from 1.55 to 1.77 mm in width. Lenthg to width ratio varies between 1.70 and 2.04. Shape of the mericarps is prolate-spheroidal and shape of their transverse sections is trigonous. The colour is brown. Diameter of abscission scars ranges from 0.27 to 0.36 mm and their shape is V-shaped. The mericap surface shows the reticulate sculpturing pattern (Tables 11, 12).

Sect. Aethiopis: Mericarps range from 2.17 to 4.50 mm in length and from 1.59 to 3.50 mm in width. Lenthg to width ratio varies between 1.02 and 1.61. Shape of the mericarps is mostly prolate-spheroidal and shape of their transverse sections is rounded- trigonous. The colour is brown in all almost species except for S. indica. Diameter of abscission scars ranges from 0.26 to 0.68 mm and their shape is rounded, U-shaped or

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V-shaped. Regarding sculpturing pattern of the mericap surface, four types can be distinguished: colliculate, reticulate, verrucate and rugose (Figure 11, 12).

Sect. Drymosphace: Mericarps range from 2.47 to 3.04 mm in length and from 1.77 to 2.02 mm in width. Lenthg to width ratio varies between 1.36 and 1.46. Shape of the mericarps is prolate-spheroidal and shape of their transverse sections is rounded- trigonous. The colour is brown. Diameter of abscission scars ranges from 0.29 to 0.38 mm and their shape is V-shaped. The mericap surface shows the reticulate sculpturing pattern (Tables 11, 12).

Sect. Plethiosphace: Mericarps range from 1.52 to 2.41 mm in length and from 1.03 to 1.85 mm in width. Lenthg to width ratio varies between 1.07 and 1.61. Shape of the mericarps is mostly prolate-spheroidal and shape of their transverse sections is rounded- trigonous. The colour is black or brown. Diameter of abscission scars ranges from 0.26 to 0.40 mm and their shape is rounded, U-shaped or squared. Regarding sculpturing pattern of the mericap surface, three types can be distinguished: colliculate, verrucate and (Tables 11, 12).

Sect. Hemisphace: Mericarps range from 1.61 to 2.45 mm in length and from 1.02 to 1.97 mm in width. Lenthg to width ratio varies between 1.19 and 1.88. Shape of the mericarps is prolate-spheroidal and shape of their transverse sections is rounded- trigonous. The colour is black or brown. Diameter of abscission scars ranges from 0.15 to 0.35 mm and their shape is U-shaped or V-shaped. Regarding sculpturing pattern of the mericap surface, two types can be distinguished: colliculate and verrucate (Tables 11, 12).

3.4.4. Systematic Implications of Mericarp Characteristics

In the taxa investigated, some mericarp characters, such as mericarp size and shape, mericarp length/width ratio, surface sculpturing, abscission scar diameter and shape and exocarp cell shape, vary significantly. These characters are more useful in separation of 125

species in the sections, including morphologically close species. For instance, the morphologically similar endemic species S. cerino-pruinosa, S. euphratica var. euphratica, S. euphratica var. leiocalycina, S. kronenburgii and S. pseudeuphratica can be separated from each other by the mericarp size, sculpturing type, exocarp cell shape, mericarp length/width ratio and shape of transverse sections. The verrucate sculpturing type is found S. cerino-pruinosa and S. euphratica var. euphratica, the colliculate sculpturing type in S. euphratica var. leiocalycina and S. kronenburgii, and the reticulate sculpturing type in S. pseudeuphratica. S. pseudeuphratica (3.22-3.68 x 3.04-3.53 mm) has smaller mericarps than the others (4.00-5.38 x 3.10-4.70 mm). S. euphratica var. leiocalycina has pentangular-hexangular exocarp cells while S. kronenburgii has rounded exocarp cells.

Mericarp size and abscission scar diameter are mainly useful at the infrageneric level. For example, sect. Hymenosphace (0.70-1.28 mm) can be clearly distinguished in its large abscission scars from all the other sections (0.15-0.65 mm) except for sect. Salvia. Moreover, sect. Plethiosphace (1.52-2.41 x 1.03-1.85 mm) and sect. Hemisphace (1.61- 2.45 x 1.02-1.97 mm) differs from sect. Hymenosphace (2.50-5.38 x 2.04-4.70 mm) and sect. Salvia (2.46-5.26 x 1.83-4.50 mm) by their smaller mericarps.

A diagnostic key are provided using mericarp characters. Mericarp micromorphology does not provide any support in separating some species. Thus, in the diagnostic key presented below, these species are represented in one branch of the key.

1. Largest diameter of abscission scars 0.70-1.28 mm……………………………………………2 2. Type of surface sculpturing reticulate……………………………………………………….3 3. Mericarp size 2.88-3.13 x 2.55-2.86 mm……………………………….………S. pinnata 3. Mericarp size 3.22-5.10 x 3.04-4.50 mm…………………………………...……………4 4. Mericarp size 3.70-5.10 x 3.61-4.50 mm………………...……..……………S. kurdica 4. Mericarp width 3.22-3.68 x 3.04-3.52 mm……….……..…….……S. pseudeuphratica 2. Type of surface sculpturing verrucate or colliculate……………..………………………….5 5. Type of surface sculpturing verrucate…………………….………………………………6

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6. Mericarp size large (4.00-5.38 x 3.56-4.70 mm) …………..……………...…………… ……..………………………...……S. cerino-pruinosa & S. euphratica var. euphratica 6. Mericarp size small (3.17-3.93 x 2.75-3.45 mm) …………….……….………………7 7. Mericarp colour brown, ± spherical mericarps present…………...……S. multicaulis 7. Mericarp colour black, ± spherical mericarps absent…………...….……S. recognita 5. Type of surface sculpturing colliculate……………...……………………………………8 8. Exocarp cells irregular……………………….…………..……………………S. huberi 8. Exocarp cells rounded or pentan-hexangular…………………………………………..9 9. Exocarp cells rounded……………………….…………………………………….10 10. Mericarp colour brown………………………….……………..………………11 11. Mericarp size 3.71-4.25 x 3.00-3.71 mm…………………..…S. suffruticosa 11. Mericarp size 2.77-3.48 x 2.04-2.95 mm…………………S. absconditiflora 10. Mericarp colour black………………….…………………………...…………12 12. Abscission scar diameter 0.93-0.1.28 mm……………..……S. kronenburgii 12. Abscission scar diameter 0.46-0.93 mm……………….…………………13 13. Average mericarp size more than 3.00 mm………...... ……S. divaricata 13. Average mericarp size less than 3.00 mm…………...………S. rosifolia 9. Exocarp cells pentan-hexangular…………………………………………………..14 14. Mericarp colour black…………………………………….……………………15 15. ± Spherical mericarps present………………………………………………16 16. Transverse sections rounded-trigonous………………………S. hedgeana 16. Transverse sections rounded to rounded-trigonous………………..……17 17. Mericarp size 4.22-5.24 x 3.47-4.50 mm………………...…………… …………………………………………S. euphratica var. leiocalycina 17. Mericarp size 2.50-3.32 x 2.20-3.14 mm……………….S. hydrangea 15. ± Spherical mericarps absent………………………………….……………18 18. Mericarp size 2.95-3.32 x 2.20-2.57 mm………….…....……S. tomentosa 18. Mericarp size 3.41-4.11-2.79-3.29 mm…………………………….…...19 19. Mericarp leangth 3.41-3.62 mm, length/width ratio 1.12-1.24…….…. ……………………………………………………………S. anatolica 19. Mericarp length 4.08-4.11 mm, length/width ratio 1.24-1.38….……... ……………………………………………………....………S. pilifera 14. Mericarp colour brown…………………………………………………………20 20. ± Spherical mericarps absent…………..…………………S. blepharochleana

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20. ± Spherical mericarps present………………………………………………21 21. Mericarp size 4.45-5.26 x 3.51-4.15 mm………………………………..22 22. Abscission scar diameter 0.54-0.90 mm…………………..S. ballsiana 22. Abscission scar diameter 0.90-1.25 mm…………….S. macrochlamys 21. Mericarp size 2.95-4.36 x 2.26-3.50 mm………………………………..23 23. Mericarp length 3.55-4.36 mm………………………….S. trichoclada 23. Mericarp length 2.95-3.52 mm………………………………………... …………………………S. bracteata, S. caespitosa & S. pachystachys 1. Largest diameter of abscission scars 0.20-0.65 mm………………..…………………………24 24. Transverse sections of mericarps trigounous……………….……..………………S. viridis 24. Transverse section of mericarps rounded-trigonous………………………….………….25 25. Mericarp length 1.52-2.45……………………………….…………………………..26 26. Abscission scar shape squared……………………………………...…S. nemorosa 26. Abscission scar shape rounded, U-shaped or V-shaped…………………………27 27. Abscission scar shape rounded………………………………………………28 28. Type of surface sculpturing rugose…………….….……………………..29 29. Mericarp length more than 2.00 mm, length/with ratio 1.25-1.40,colour back….….….….….….….….….….….….….….….….….…..S. virgata 29. Mericarp length less than 2.00 mm, length/with ratio 1.07-1.22, colour brown….….….….….….….….….….….….….….….…...S. dicroantha 28. Type of surface sculpturing colliculate…………………………………..30 30. Mericarp size 1.79-2.41 x 1.27-1.71 mm, length/width ratio 1.30-1.61, exocarp cells pentan-hexangular………………..…………S. staminea 30. Mericarp size 2.42-2.84 x 2.12-2.60 mm, length/width ratio 1.09-1.21, exocarp cells irregular……………………………………S. palaestina 27. Abscission scar shape U-shaped or V-shaped………..………………………31 31. Type of surface sculpturing verrucate……………………...…………….32 32. Mericarp size 2.06-2.45 x 1.46-1.97 mm, abscission scar diameter 0.25- 0.35……………..………….……………S. verticillata subsp. amasiaca 32. Mericarp size 1.61-1.96 x 1.02-1.20 mm, abscission scar diameter 0.25- 0.35………..…………...………………S. verticillata subsp. verticillata 31. Type of surface sculpturing colliculate….….….….….….….….….…….33 33. Exocarp cells irregular...………… ...………… …..…….. …………34 34. Mericarp size 2.08-2.29 x 1.14-1.45 mm………………..S. russellii

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34. Mericarp size 2.30-3.10 x 1.59-2.12 mm………………...S. cilicica 33. Exocarp cells pentan-hexangular…………………………………….35 35. Mericarp width 1.27-1.71 mm………………………...S. stamineae 35. Mericarp width 1.79-1.96 mm……………………………S. syriaca 25. Mericarp length 2.45-4.50……………….………………………….……………….36 36. Type of surface sculpturing verrucate……………………………..S. xanthocheila 36. Type of surface sculpturing colliculate, reticulate or rugose……………………37 37. Type of surface sculpturing rugose………………………………………...38 38. Mericarp width 1.95-2.42 mm, abscission scar diameter 0.30-0.44 mm... ………………………………………………………………S. eriophora 38. Mericarp width 2.49-2.92 mm, abscission scar diameter 0.52-0.65 m….. ……………………………………………………………S. ceratophylla 37. Type of surface sculpturing colliculate or reticulate ……………….……...39 39. Type of surface sculpturing reticulate…………………………….……40 40. Mericarps black, abscission scar shape rounded…………….S. indica 40. Mericarps brown, abscission scar shape U-shaped or V-shaped…..41 41. Abscission scar shape V-shaped…………………..…S. glutinosa 41. Abscission scar shape U-shaped………………………….....…42 42. ± Spherical mericarps present…………………...…S. siirtica 42. ± Spherical mericarps absent……………………...………43 43. Abscission scar diameter 0.30-0.37 mm…...S. montbretii 43. Abscission scar diameter 0.40-0.55 mm…………...…… ………………………….S. atropatana & S. microstegia 39. Type of surface sculpturing colliculate…………………………………44 44. Exocarp cells irregular………………………………………...……45 45. Abscission scar shape rounded…………………………………46 46. Mericarp length/width ratio 1.09-1.20……..……S. palaestina 46. Mericarp length/width ratio 1.20-1.40……………………...47 47. Largest diameter of abscission scars more than 0.5 mm…. …………………………………………………S. limbata 47. Largest diameter of abscission scars less than 0.5 mm…... …………………………………………………S. sclarea 45. Abscission scar shape U-shaped……………………….…….…48

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48. Prolate-spheroidal mericarps absent………………………….. ………………………………S. odontochlamys & S. poculata 48. Prolate-spheroidal mericarps present…………...……..……… ...…………………………………...S. cilicica & S. hypargeia 44. Exocarp cells pentan-hexangular…………………………………...49 49. Abscission scar shape V-shaped……………………………..…50 50. Mericarp width 2.30-2.60 mm………………………S. frigida 50. Mericarp width 1.91-2.30 mm………………………...……… ………………….…………S. aethiopis & S. longipedicellata 49. Abscission scar shape rounded or V-shaped……………………51 51. ± Spherical mericarps present…..S. macrosiphon & S. spinosa 51. ± Spherical mericarps absent………………………….……52 52. Mericarp length 3.16-3.31 mm…..…….….S. brachyantha 52. Mericarp length 2.50-3.15 mm………………………53 53. Mericarp length 2.96-3.15 mm…………………….. ………………S. candidissima subsp. candidissima 53. Mericarp length 2.50-2.76 mm………S. cyanescens

Hedge (1982a) reported that some species of the Turkish Salvia had brown or black mericarps and rounded, trigonous or rounded-trigonous mericarps (the most common) in transverse sections. The results of the present study is congruent with those of Hedge (1982a). The abscission scar andmericarp shape were invariable in the tribe Saturejeae while these characters varied significantly in the Westringieae (Husain et al., 1990).

Mericarp data are not only useful as a diagnostic character in delimitation of the species of Salvia, but also they provide support for separating sections of the genus as indicated by some early researchers (Wojciechowska, 1966; MARIN et al., 1994). According to data presented in the Flora of Turkey (Hedge, 1982a), members of the sections Plethiosphace and Hemisphace can be distinguished from those of the other sections by their smaller mericarps. Mericarp size of the section Plethiosphace is 1.5-2.5 x 1.0-2.0 mm and mericarp size of the section Hemisphace is 2.2-2.5 x 1.3-1.5 mm.

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Özkan et al. (2009) reported mericarp properties of twelve Turkish Salvia taxa belonging to different sections. They grouped the species into three types of sculpturing: foveate, reticulate and verrucate. The three types were observed in the section Aethiopis among the examined taxa. While the studied taxa of the sections Salvia and Hymenosphace had the foveate sculpturing pattern, those of the sections Hemisphace and Plethiosphace had verrucate sculpturing pattern. In the present study, the species belonging to sect. Salvia mainly showed the colliculate pattern of mericarp sculpturing. Some discrepancies hamper the usefulness of direct comparisons with the findings of this previous study. In some cases, the SEM images of several taxa do not seem to correspond with surface sculpturing types assigned to the species. In some cases, themagnification seems to be insufficient to show the detail of the sculpturing patterns of exocarp cells.

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Table 11. Quantitative mericarp characteristics of mericarps in the taxa studied of Salvia. All sizes are in mm except for length/with ratio. Numbers refer to minimum-maximum (mean). Taxa and sections Mericarp length Mericarp width Length/width ratio Abscission scar diameter Sect. Salvia S. anatolica 3.41-3.64 (3.52) 2.79-3.15 (3.02) 1.12-1.24 (1.16) 0.70-0.89 (0.79) S. ballsiana 4.45-5.26 (4.95) 3.75-4.15 (3.93) 1.14-1.29 (1.22) 0.54-0.90 (0.77) S. bracteata 3.07-3.45 (3.24) 2.68-3.09 (2.88) 1.01-1.18 (1.12) 0.50-0.86 (0.69) S. caespitosa 2.99-3.38 (3.12) 2.49-2.89 (2.68) 1.09-1.22 (1.16) 0.50-0.79 (0.67) S. divaricata 3.50-4.46 (3.92) 2.67-3.63 (3.07) 1.15-1.37 (1.28) 0.76-0.93 (0.86) S. hedgeana 3.12-4.36 (3.66) 2.26-3.44 (2.88) 1.11-1.38 (1.28) 0.55-0.83 (0.69) S. huberi 2.46-3.54 (2.90) 1.83-2.86 (2.17) 1.20-1.60 (1.34) 0.41-0.75 (0.56) S. kurdica 3.70-5.10 (4.37) 3.61-4.50 (3.78) 1.01-1.12 (1.09) 1.01-1.20 (1.08) S. macrochlamys 4.52-5.20 (4.95) 3.51-4.10 (4.00) 1.14-1.29 (1.23) 0.90-1.25 (1.03) 132 S. pachystachys 2.95-3.52 (3.18) 2.26-2.84 (2.55) 1.11-1.33 (1.25) 0.45-0.70 (0.59)

S. pilifera 4.08-4.11 (4.09) 2.97-3.29 (3.14) 1.24-1.38 (1.30) 0.71-0.86 (0.78) S. pinnata 2.88-3.13 (3.00) 2.55-2.86 (2.71) 1.07-1.14 (1.10) 0.86-1.01 (0.93) S. recognita 3.17-3.89 (3.49) 2.86-3.35 (3.04) 1.13-1.19 (1.17) 0.70-0.88 (0.78) S. rosifolia 2.56-3.63 (2.96) 1.91-2.87 (2.36) 1.12-1.36 (1.25) 0.46-0.79 (0.60) S. suffruticosa 3.71-4.25 (3.95) 3.00-3.71 (3.47) 1.09-1.22 (1.13) 0.70-0.90 (0.78) S. tomentosa 2.95-3.32 (3.20) 2.20-2.57 (2.40) 1.26-1.36 (1.33) 0.50-0.77 (0.63) S. trichocloda 3.55-4.36 (3.92) 2.92-3.50 (3.25) 1.04-1.27 (1.20) 0.55-0.78 (0.70) Sect. Hymenosphace S. absconditiflora 2.77-3.48 (3.06) 2.04-2.95 (2.58) 1.11-1.31 (1.20) 0.70-1.02 (0.90) S. blepharochleana 3.05-3.92 (3.44) 2.40-3.09 (2.72) 1.15-1.40 (1.27) 0.75-1.05 (0.89) S. cerino-pruinosa 4.00-5.27 (4.62) 3.60-4.70 (4.02) 1.10-1.29 (1.15) 1.10-1.19 (1.15) S. euphratica 4.32-5.38 (4.69) 3.56-4.33 (3.83) 1.08-1.27 (1.20) 1.03-1.21 (1.13) var. euphratica S. euphratica 4.22-5.24 (4.63) 3.47-4.50 (3.96) 1.08-1.22 (1.18) 1.08-1.25 (1.17) var. leiocalycina

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Table 11 (continued) S. hydrangea 2.50-3.32 (2.95) 2.20-3.14 (2.62) 1.06-1.18 (1.12) 0.70-0.96 (0.81) S. kronenburgii 4.25-5.13 (4.54) 3.10-3.97 (3.46) 1.26-1.40 (1.31) 0.93-1.28 (1.14) S. multicaulis 3.25-3.93 (3.62) 2.75-3.45 (3.05) 1.12-1.31 (1.21) 0.80-1.05 (0.92) S. pseudeuphratica 3.22-3.68 (3.54) 3.04-3.53 (3.23) 1.02-1.13 (1.08) 0.98-1.13 (1.06) Sect. Horminum S. viridis 2.85-3.34 (3.05) 1.55-1.77 (1.63) 1.70-2.04 (1.86) 0.27-0.36 (0.32) Sect. Aethiopis S. aethiopis 2.67-2.94 (2.78) 1.91-2.28 (2.06) 1.23-1.46 (1.35) 0.28-0.40 (0.35) S. atropatana 3.01-3.51 (3.24) 2.18-2.63 (2.37) 1.25-1.49 (1.37) 0.40-0.55 (0.49) S. brachyantha 3.16-3.31 (3.20) 1.97-2.29 (2.16) 1.39-1.61 (1.49) 0.40-0.49 (0.43) S. candidissima 2.96-3.15 (3.04) 2.09-2.40 (2.24) 1.27-1.45 (1.37) 0.35-0.44 (0.40) subsp. candidissima S. ceratophylla 3.00-3.56 (3.28) 2.49-2.92 (2.66) 1.14-1.27 (1.23) 0.52-0.65 (0.60)

133 S. cilicica 2.30-3.10 (2.66) 1.59-2.12 (1.82) 1.26-1.60 (1.43) 0.30-0.36 (0.33)

S. cyanescens 2.50-2.76 (2.61) 1.80-2.18 (2.02) 1.19-1.40 (1.30) 0.35-0.48 (0.42) S. eriophora 2.70-3.28 (2.91) 1.95-2.42 (2.17) 1.25-1.45 (1.34) 0.30-0.44 (0.37) S. frigida 2.90-3.25 (3.09) 2.30-2.60 (2.47) 1.23-1.34 (1.27) 0.36-0.53 (9.42) S. hypargeia 2.70-3.40 (3.17) 1.91-2.50 (2.31) 1.29-1.50 (1.40) 0.30-0.43 (0.38) S. indica 3.15-4.50 (3.62) 3.00-3.50 (3.33) 1.02-1.21 (1.08) 0.44-0.65 (0.55) S. limbata 2.50-3.18 (2.90) 1.80-2.38 (2.21) 1.23-1.39 (1.31) 0.40-0.60 (0.52) S. longipedicellata 2.62-3.21 (2.95) 1.95-2.30 (2.18) 1.24-1.40 (1.34) 0.26-0.36 (0.32) S. macrosiphon 2.63-2.90 (2.81) 2.25-2.40 (2.32) 1.10-1.22 (1.16) 0.30-0.38 (0.34) S. microstegia 2.80-3.25 (3.09) 2.32-2.64 (2.46) 1.18-1.35 (1.26) 0.40-0.50 (0.45) S. montbretii 3.00-3.22 (3.12) 2.33-2.61 (2.49) 1.19-1.34 (1.25) 0.30-0.37 (0.33) S. odontochlamys 2.75-3.10 (2.91) 2.60-2.99 (2.79) 1.01-1.10 (1.05) 0.35-0.45 (0.40) S. palaestina 2.42-2.84 (2.54) 2.12-2.60 (2.22) 1.09-1.20 (1.14) 0.30-0.47 (0.39) S. poculata 2.82-3.13 (3.00) 2.67-3.00 (2.89) 1.00-1.11 (1.05) 0.35-0.48 (0.43) S. sclarea 2.48-2.90 (2.64) 1.89-2.23 (2.04) 1.20-1.40 (1.30) 0.32-0.44 (0.38) S. siirtica 2.90-3.30 (3.09) 2.40-2.80 (2.61) 1.08-1.25 (1.15) 0.30-0.50 (0.41)

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Table 11 (continued) S. spinosa 2.75-3.04 (2.87) 2.42-2.65 (2.52) 1.09-1.26 (1.14) 0.30-0.40 (0.35) S. syriaca 2.17-2.44 (2.29) 1.79-1.96 (1.86) 1.22-1.34 (1.28) 0.26-0.43 (0.34) S. xanthocheila 3.12-3.64 (3.35) 2.41-2.93 (2.63) 1.21-1.36 (1.27) 0.41-0.55 (0.49) Sect. Drymosphace S. glutinosa 2.47-3.04 (2.74) 1.77-2.02 (1.88) 1.36-1.53 (1.46) 0.29-0.38 (0.34) Sect. Plethiosphace S. dicroantha 1.77-1.98 (1.85) 1.47-1.73 (1.59) 1.07-1.22 (1.16) 0.30-0.40 (0.36) S. nemorosa 1.52-1.88 (1.65) 1.03-1.31 (1.18) 1.40-1.61 (1.51) 0.26-0.35 (0.31) S. staminea 1.79-2.41 (2.01) 1.27-1.71 (1.40) 1.30-1.61 (1.46) 0.27-0.35 (0.31) S. virgata 2.12-2.36 (2.24) 1.61-1.85 (1.71) 1.25-1.40 (1.31) 0.30-0.40 (0.36) Sect. Hemisphace S. russellii 2.08-2.29 (2.19) 1.14-1.45 (1.28) 1.53-1.88 (1.70) 0.25-0.35 (0.30) S. verticillata 2.06-2.45 (2.28) 1.46-1.97 (1.70) 1.19-1.55 (1.35) 0.25-0.35 (0.29)

134 subsp. amasiaca S. verticillata 1.61-1.96 (1.80) 1.02-1.20 (1.15) 1.42-1.67 (1.57) 0.15-0.25 (0.19)

subsp. verticillata

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Table 12. Qualitative mericarp characteristics in the taxa studied of Salvia. S = Near spherical, Pr = Prolate-spheroidal. -, absent, +, rarely present, ++, densely present. Taxa and sections Shape Shape of transverse Colour Shape of abscission Surface sculpturing Shape of exocarp section scar cells S Ps Sect. Salvia S. anatolica - ++ Rounded-trigonous Black Rounded Colliculate Pentan-hexangular S. ballsiana + ++ Rounded-trigonous Brown Rounded Colliculate Pentan-hexangular S. bracteata + ++ Rounded-trigonous Brown Rounded Colliculate Pentan-hexangular S. caespitosa + ++ Rounded-trigonous Brown Rounded Colliculate Pentan-hexangular S. divaricata - ++ Rounded-trigonous Black Rounded Colliculate Rounded S. hedgeana + ++ Rounded-trigonous Black Rounded Colliculate Pentan-hexangular S. huberi - ++ Rounded-trigonous Black Rounded Colliculate Irregular

135 S. kurdica ++ - Rounded Black Rounded Reticulate -

S. macrochlamys + ++ Rounded-trigonous Brown Rounded Colliculate Pentan-hexangular S. pachystachys + ++ Rounded-trigonous Brown Rounded Colliculate Pentan-hexangular S. pilifera - ++ Rounded-trigonous Black Rounded Colliculate Pentan-hexangular S. pinnata ++ - Rounded Black Rounded Reticulate - S. recognita - ++ Rounded-trigonous Black Rounded Verrucate - S. rosifolia + ++ Rounded-trigonous Black Rounded Colliculate Rounded S. suffruticosa + ++ Rounded to rounded- Brown Rounded Colliculate Rounded trigonous S. tomentosa - ++ Rounded-trigonous Black Rounded Colliculate Pentan-hexangular S. trichocloda + ++ Rounded-trigonous Brown Rounded Colliculate Pentan-hexangular Sect. Hymenosphace Rounded S. absconditiflora + ++ Rounded-trigonous Brown Rounded Colliculate Rounded S. blepharochleana - ++ Rounded-trigonous Brown Rounded Colliculate Pentan-hexangular S. cerino-pruinosa + ++ Rounded to rounded- Black Rounded Verrucate - tigonous

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Table 12 (continued) S. euphratica + ++ Rounded to rounded- Black Rounded Verrucate - var. euphratica tigonous S. euphratica + ++ Rounded to rounded- Black Rounded Colliculate Pentan-hexangular var. leiocalycina tigonous S. hydrangea ++ + Rounded to rounded- Black Rounded Colliculate Pentan-hexangular tigonous S. kronenburgii - ++ Rounded-trigonous Black Rounded Colliculate Rounded S. multicaulis + ++ Rounded-trigonous Brown Rounded Verrucate - S. pseudeuphratica ++ - Rounded Black Rounded Reticulate - Sect. Horminum S. viridis - ++ Trigonous Brown V-shaped Reticulate - Sect. Aethiopis S. aethiopis - ++ Rounded-trigonous Brown V-shaped Colliculate Pentan-hexangular

136 S. atropatana - ++ Rounded-trigonous Brown U-shaped Reticulate - S. brachyantha - ++ Rounded-trigonous Brown U-shaped Colliculate Pentan-hexangular

S. candidissima - ++ Rounded-trigonous Brown Rounded/U-shaped Colliculate Pentan-hexangular subsp. candidissima S. ceratophylla + ++ Rounded-trigonous Brown Rounded Rugose - S. cilicica - ++ Rounded-trigonous Brown U-shaped Colliculate Irregular S. cyanescens - ++ Rounded-trigonous Brown Rounded/U-shaped Colliculate Pentan-hexangular S. eriophora - ++ Rounded-trigonous Brown Rounded/U-shaped Rugose - S. frigida - ++ Rounded-trigonous Brown V-shaped Colliculate Pentan-hexangular S. hypargeia - ++ Rounded-trigonous Brown U-shaped Colliculate Irregular S. indica ++ + Rounded-trigonous Black Rounded Reticulate - S. limbata - ++ Rounded-trigonous Brown Rounded Colliculate Irregular S. longipedicellata - ++ Rounded-trigonous Brown V-shaped Colliculate Pentan-hexangular S. macrosiphon + ++ Rounded-trigonous Brown U-shaped Colliculate Pentan-hexangular S. microstegia - ++ Rounded-trigonous Brown U-shaped Reticulate - S. montbretii - ++ Rounded-trigonous Brown U-shaped Reticulate -

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Table 12 (continued) S. odontochlamys ++ - Rounded-trigonous Brown U-shaped Colliculate Irregular S. palaestina + ++ Rounded-trigonous Brown Rounded Colliculate Irregular S. poculata ++ - Rounded-trigonous Brown U-shaped Colliculate Irregular S. sclarea - ++ Rounded-trigonous Brown Rounded Colliculate Irregular S. siirtica + ++ Rounded-trigonous Brown U-shaped Reticulate - S. spinosa + ++ Rounded-trigonous Brown U-shaped Colliculate Pentan-hexangular S. syriaca - ++ Rounded-trigonous Brown U-shaped Colliculate Pentan-hexangular S. xanthocheila - ++ Rounded-trigonous Brown U-shaped Verrucate - Sect. Drymosphace Rounded-trigonous S. glutinosa - ++ Rounded-trigonous Brown V-shaped Reticulate - Sect. Plethiosphace Rounded-trigonous S. dicroantha + ++ Rounded-trigonous Brown Rounded Rugose - S. nemorosa - ++ Rounded-trigonous Black Squared Verrucate -

137 S. staminea - ++ Rounded-trigonous Brown Rounded/U-shaped Colliculate Pentan-hexangular

S. virgata - ++ Rounded-trigonous Black Rounded Rugose - Sect. Hemisphace S. russellii - ++ Rounded-trigonous Brown U-shaped/V-shaped Colliculate Irregular S. verticillata - ++ Rounded-trigonous Brown U-shaped/V-shaped Verrucate - subsp. amasiaca S. verticillata - ++ Rounded-trigonous Brown U-shaped/V-shaped Verrucate - subsp. verticillata

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3.5. Ecology and Distribution

3.5.1. Distribution, Endemism and Phytogeography

The study area contains 59 taxa, 24 (40.7%) of which are endemic and the remaining 35 (59.3%) are non-endemic (Table 13). 7 endemics and 10 non-endemics are distributed only in the area. As the geographical distribution of the taxa is analyzed, East Anatolia seems to have 51 taxa, 22 (43.1%) of which are endemic and Southeast Anatolia has 25 taxa, 3 (12.0%) of which are endemic.

The endemic taxa are concentrated in three main areas within the study area. The first area covers Divriği, Kemaliye and Erzincan, and includes 12 endemic taxa. The second area covers Sarız, Gürün, Malatya, and includes 11 endemic taxa. The third area covers Tercan, Erzurum and Ağrı, and includes 4 endemic taxa. The richest areas for the endemic taxa are shown in Figure 84.

Distribution area of most endemic taxa falls into the Anatolain Diagonal, which is considered one of the most significant areas of endemism in Turkey (Ekim & Güner, 1986; Ekim, 1987) and are under the effect of Irano-Turanian phytogeographical region. The high level of endemic Salvia species in the Anatolian Diagonal can be due to geographical, ecological and climatic factors. For example, verly local endemic species of Salvia, such as S. hedgeana, S. anatolica and S. cerino-pruinosa, is found in large areas of gypseous hills around Divriği, Gürün and Darende. Calcareous, limestone and serpentine slopes in the diagonal are also preferred by most endemic species prefer. Moreover, the diagonal mountains are a speciational barrier for the endemic species. Kemaliye and Erzincan are rich in endemic species because the regions of low altitude and moderate climate on the diagonal.

According to the grid square system used in the Flora of Turkey (sensu Davis, 1965), the B7 square has the highest number of taxa (38) whereas the A8 square has the lowest number of taxa (6). The B7 square has the greatest number of endemic taxa (17), but

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there are no endemic taxa in the B10, C7, C8 and C10 squares. The B9 square has the highest number of non-endemic taxa (24) while the A8 square has the lowest number of non-endemic taxa (5). The number of the endemic and non-endemic taxa is given for each grid square (Table 13, Figure 85).

Phytogeography of the taxa is as follows: 50 taxa (85%) Irano-Turanian, 4 taxa (7%) Mediterranean, 2 taxa (3%) Euro-Siberian, and 3 taxa (5%) unknown or multiregional (Table 13, Figure 85).

Figure 84. The richest three regions in terms of the endemic Salvia taxa in the study area shaded. These regions are indicated by circles. : East Anatolia, : Southeast Anatolia.

Figure 85. The number of the Salvia taxa studied within each square. Endemics/Non- endemics 139

Table 13. The taxa distributed in the study area and their distribution data. Details of the taxa known only within the area, endemism, phtogeographic regions, phenology and altitudinal range of the taxa are also provided. *: taxa known only within the study area in Turkey. Euro-Sib.: Euro-Siberian element, Ir.-Tur.: Irano-Turanian element, Medit.: Mediterranean element, Unk. or Multi.: Unknown or Multiregional. The taxa are listed in alphabetical order. No Taxa Distribution in the study Endemism Phytogeography Phenology Altitudinal range area according to Davis’s grid square system (1965) 1 S. absconditiflora B6-B8 Endemic Ir.-Tur. 4-7 650-2500 2 S. aethiopis A8, A9; B6-B10 Non-endemic Unk. or Multi. 5-8 1-2100 3 S. anatolica B7 Endemic Ir.-Tur. 5-6 1350-1650

140 4 *S. atropatana B9, B10; C9, C10 Non-endemic Ir.-Tur. 6-7 1245-2560 5 *S. ballsiana B7 Endemic Ir.-Tur. 5 1050-1220

6 S. blepharochlaena B6 Endemic Ir.-Tur. 5-6 1000-2000 7 S. brachyantha A9; B8, B9; C7 Non-endemic Ir.-Tur. 5-6 1050-2800 8 S. bracteata B6-B8; C6-C8 Non-endemic Ir.-Tur. 5-7 50-2000 9 S. caespitosa B6, B7 Endemic Ir.-Tur. 5-7 1200-2700 10 S. candidissima B6-B9; C6, C8-C10 Non-endemic Ir.-Tur. 5-9 700-2000 subsp. candidissima 11 S. ceratophylla A9; B6-B10; C6-C8, C10 Non-endemic Ir.-Tur. 4-7 300-2300 12 *S. cerino-pruinosa B7 Endemic Ir.-Tur. 5-6 850-1450 13 S. cilicica B6 Endemic Medit. 7-9 750-2200 14 S. cyanescens B7 Endemic Ir.-Tur. 6-9 400-2300 15 S. dichroantha B6, B7 Endemic Ir.-Tur. 6-9 700-2000

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Table 13 (continued) 16 S. divaricata B7 Endemic Ir.-Tur. 5-6 1030-1870 17 S. eriophora B6 Endemic Ir.-Tur. 5-6 1525-1900 18 S. euphratica B6, B7 Endemic Ir.-Tur. 4-6 800-1800 var. euphratica S. euphratica B6, B7 Endemic Ir.-Tur. 4-6 800-1800 var. leiocalycina 19 S. frigida B6-B9 Non-endemic Ir.-Tur. 5-7 1000-2500 20 S. glutinosa A9 Non-endemic Euro-Sib. 7-10 780-2360 21 *S. hedgeana B7 Endemic Ir.-Tur. (5-) 6 1200-1600 22 S. huberi A8 Endemic Ir.-Tur. 5-8 1100-2200

41 23 S. hypargeia B6, B7 Endemic Ir.-Tur. 6-7 800-2000

24 *S. hydrangea A9; B9, B10 Non-endemic Ir.-Tur. (5-) 6-8 600-2000 25 S. indica C9, C10 Non-endemic Ir.-Tur. 4-5 100-1500 26 *S. kronenburgii B9 Endemic Ir.-Tur. 6-7 1800-2600 27 *S. kurdica C9 Non-endemic Ir.-Tur. 5 900 28 S. limbata A9; B8-B10; C10 Non-endemic Ir.-Tur. 6-8 1060-2340 29 S. longipedicellata B7-B9 Endemic Ir.-Tur. 7-8 1480-2110 30 *S. macrochlamys B7-B9; C8-C10 Non-endemic Ir.-Tur. 6-8 900-2400 31 *S. macrosiphon C8 Non-endemic Ir.-Tur. 5-6 759 32 S. microstegia A9; B6-B9, C7, C9, C10 Non-endemic Ir.-Tur. 6-8 970-3350 33 *S. montbretii C6-C8 Non-endemic Ir.-Tur. 5 700-1150 34 S. multicaulis A8, A9; B6-B9; C6-C10 Non-endemic Ir.-Tur. 4-7 550-2600

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Table 13 (continued) 35 S. nemorosa A9; B8-B10; C9, C10 Non-endemic Unk. or Multi. (5-) 6-9 1000-2300 36 *S. odontochlamys B9 Endemic Ir.-Tur. 6-7 2100-2280 37 S. pachystachys A9; B7, B9, B10 Non-endemic Ir.-Tur. 5-7 1200-3200 38 S. palaestina B6-B8; C6-C9 Non-endemic Ir.-Tur. 5-7 300-1510 39 S. pilifera B7; C6 Endemic Ir.-Tur. 4-5 200-2300 40 S. pinnata C6, C9 Non-endemic Medit. 3-5 1-1060 41 *S. poculata A8; B7-B9; C9, C10 Non-endemic Ir.-Tur. 5-7 1400-2900 42 *S. pseudeuphratica B7 Endemic Ir.-Tur. 5 (-6) 750-900 43 S. recognita B7 Endemic Ir.-Tur. 5-8 900-2200 44 S. rosifolia A9; B7-B9 Endemic Ir.-Tur. 5-8 500-2350 45 S. russellii B7-B9; C6-C9 Non-endemic Ir.-Tur. 5-7 100-1600

142 46 S. sclarea A8; B6-B9; C6, C7, C9, C10 Non-endemic Unk. or Multi. 5-8 1-2000

47 *S. siirtica C9 Endemic Ir.-Tur. 5-6 1450-1500 48 *S. spinosa C6-C9 Non-endemic Ir.-Tur. 5-6 300-900 49 S. staminea A9; B7-B10; C10 Non-endemic Ir.-Tur. 5-8 1700-3150 50 S. suffruticosa B7-B9; C7, C8 Non-endemic Ir.-Tur. 4-6 300-2000 51 S. syriaca A8, A9; B6-B9; C6-C9 Non-endemic Ir.-Tur. 4-7 350-2200 52 S. tomentosa B7 Non-endemic Medit. 4-8 90-2000 53 *S. trichoclada B7-B9; C8-C10 Non-endemic Ir.-Tur. 5-7 300-2440 54 S. verticillata A9; B8, B9; C9 Non-endemic Euro-Sib. 6-8 1-2400 subsp. verticillata S. verticillata A9; B6-B10; C9, C10 Non-endemic Ir.-Tur. 5-9 150-2300 subsp. amasiaca

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Table 13 (continued) 55 S. virgata A9; B6-B10; C9 Non-endemic Ir.-Tur. 5-9 1-2300 56 S. viridis B7; C6-C9 Non-endemic Medit. 3-7 1-2300 57 *S. xanthocheila A9; B8, B9; C9 Non-endemic Ir.-Tur. 5-7 2000-2870

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3.5.2. Habitat, Phenology and Altitudinal Range

Salvia taxa are found in a very wide variety of habitats reaching up to 3350 m elevation. These include limestone slopes (e.g. S. hydrangea), igneous slopes (e.g. S. brachyantha), calcareous slopes (e.g. S. montbretii), serpentine slopes (e.g. S. blepharochleana), rocky and stony slopes (e.g. S. pachystachys), steppe (e.g. S. aethiopis), gypsum areas (e.g. S. euphratica var. euphratica, S. euphratica var. leiocalycina), marly places (e.g. S. hedgeana), roadsides (e.g. S. sclarea), cornfields (e.g. S. limbata), fallow fields (e.g. S. syriaca), cultivated fields (e.g. S. russellii), vineyards (e.g. S. suffruticosa), waste places (e.g. S. bracteata), grassy places (e.g. S. trichoclada), dry meadows (e.g. S. pinnata), macchie (e.g. S. viridis), bushy places (S. candidissima subsp. candidissima), alpine and subalpine meadows (e.g. S. staminea), loamy hills (e.g. S.rosifolia), near permanent snowbeds (e.g. S. microstegia), river shingle (e.g. S. euphratica), Pinus, Quercus, Juniperus, Abies and Pyrus woodland (e.g. S. frigida, S. virgata), and moist places in Fagus, Alnus, Corylus and in Picea forests (e.g. S. glutinosa, S. verticillata subsp. verticillata) (Figures 86-95).

Data concerning the flowering period and altitudinal range compiled from the recent taxonomic revision and herbarium specimens are given in Table 13. Most species have a large altitudinal range, but some of the endemic species, such as S. anatolica, S. ballsiana, S. eriophora, S. hedgeana, S. odontochlamys, S. pseudeuphratica, are restricted to a narrow altitudinal range. Additionally, altitudinal ranges of the endemic species have clearly narrower than the non endemic species. The Irano-Turanian elements are mainly found in higher altitudes than the Mediterranean and Euro-Siberian elements. S. microstegia, S. pachystachys and S. staminea grow at elevations above 3000 m (Table 13).

In the species studied, flowers can apparently be produced between March and October. These data indicate that most species flower between May and July. The earliest part of the flowering is observed in S. pinnata and S. viridis whereas the latest part of flowering is recognized in S. glutinosa. S. absconditiflora, S. aethiopis, S. candidissima subsp.

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candidissima, S. ceratophylla, S. cyanescens, S. dicroantha, S. glutinosa, S. huberi, S. hydrangea, S. multicaulis, S. nemorosa, S. recognita, S. rosifolia, S.sclarea, S. staminea, S. tomentosa, S. verticillata subsp. verticillata and S. virgata have a fairly long flowering time. S. anatolica, S. ballsiana, S. blepharochleana, S. brachyantha, S. cerino-pruinosa, S. divaricata, S. eriophora, S. hedgeana, S. indica, S. kurdica, S. macrosiphon, S. montbretii, S. odontochlamys, S. pilifera, S. pseudeuphratica, S. siirtica and S. spinosa have a short flowering time (Table 13).

Figure 86. S. hydrangea growing on rocky volcanic slopes in Ağrı Mountain.

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Figure 87. S. longipedicellata growing in fields.

Figure 88. S. euphratica growing on marly banks.

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Figure 89. S. caespitosa growing on steppe.

Figure 90. S. divaricata growing on limestone slopes.

147

Figure 91. S. poculata growing in subalpine meadows.

Figure 92. S. multicaulis growing on calcareous rocky slopes.

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Figure 93. S. aethiopis growing along roadsides.

Figure 94. S. glutinosa growing on moist places in Picea orientalis forest.

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Figure 95. S. ballsiana growing on moist places in Quercus scrub.

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3.5.3. Soil Properties

Physical and chemical features of the soil samples taken from the natural habitats of the taxa are given in Table 14. The results show that they grow in different soil types such as clay, loam, sandy and silty soils. These soil types are also composed of different proportions of clay, loam, sand and silt: clayish-loamy, loamy-sandy, sandy-loamy, sandy-clayish-loamy, silty-clayish and silty-clayish-loamy.

Most of the taxa (16) are found in sandy-loamy soils, as in S. blepharochleana, S. brachyantha, S. bracteata, S. candidissima subsp. candidissima, S. ceratophylla, S. euphratica var. euphratica, S. euphratica var. leiocalycina, S. frigida, S. huberi, S. hydrangea, S. indica, S. montbretii, S. pseudeuphratica, S. suffruticosa, S. viridis and S. xanthocheila. 12 taxa are grown in clayish-loamy soils: S. aethiopis, S. anatolica, S. caespitosa, S. cerino-pruinosa, S. cilicica, S. hedgeana, S. limbata, S. poculata, S. syriaca, S. tomentosa, S. verticillata subsp. amasiaca and S. virgata. 11 taxa are grown in clayish soils: such as S. absconditiflora, S. eriophora, S. longipedicellata, S. macrosiphon, S. pinnata, S. recognita, S. rosifolia, S. russellii, S. sclarea, S. spinosa and S. staminea. 10 taxa, such as S. dicroantha, S. divaricata, S. kronenburgii, S. microstegia, S. nemorosa, S. pachystachys, S. palaestina, S. pilifera, S. trichoclada and S. verticillata subsp. verticillata, are found in loamy soils. S. cyanescens, S. hypargeia and S. macrochlamys occur in sandy-clayish-loamy soils. S. atropatana are grown only in silty-clayish soils and S. multicaulis only in silty-clayish-loamy soils.

The taxa generally prefer slightly alkaline soils (pH= 7.07-7.44), but S. eriophora, S. indica, S. longipedicallata, S. pachystachys, S. poculata and S. staminea grow neutral soils (pH= 7.56-8.22). The total salt content of the soil varies between 0.004% as in S. frigida and 0.056% as in S. eriophora and S. pachystachys. The total CaCO3 content ranges from 0.75% in S. eriophora and 35.29% in S. anatolica. The P content varies between 0.82 ppm in S. candidissima subsp. candidissima and S. kronenburgii and 37.41 ppm in S. bracteata. The K content varies between 18.82 ppm in S. candidissima subsp. candidissima and 497.42 ppm in S. aethiopis and S. verticillata subsp. verticillata. The

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organic matter content ranges from 0.19% in S. candidissima subsp. candidissima and 18.70% in S. pachystachys.

Soil features do not play a role in section delimitation, but they show significant differences among some species, particularly morphologically similar species. For example, S. caespitosa prefers slightly alkaline soils, but its closely related species S. pachystachys prefers neutral soils. S. absconditiflora are grown in clayish soils whereas S. multicaulis are found in silty-clayish-loamy soils.

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Table 14. Physical and chemical properties of soil samples taken from the natural habitat of the taxa. No Taxa Total pH Total P (ppm) K (ppm) Organic Sand Silt Clay Texture

salt (%) CaCO3 matter (%) (%) (%) (%) 1 S. absconditiflora 0.039 8.19 15.53 4.47 285.95 0.25 11.9 31.7 56.4 Clay 2 S. aethiopis 0.044 7.70 34.79 13.15 497.42 2.20 33.2 27.8 39.0 Clay-Loam 3 S. anatolica 0.019 7.76 35.29 5.97 150.49 1.07 33.8 35.3 30.9 Clay-Loam 4 S. atropatana 0.039 7.74 5.47 9.48 510.73 4.96 12.2 42.3 45.5 Silt-Clay 5 S. ballsiana ------6 S. blepharochlaena 0.006 7.76 0.94 2.57 46.83 0.82 70.6 17.7 11.7 Sand-Loam 7 S. brachyantha 0.007 8.05 22.27 4.87 63.03 1.57 76.9 14.2 8.9 Sand-Loam 8 S. bracteata 0.012 7.77 4.81 37.41 361.86 2.70 66.2 20.1 13.7 Sand-Loam

153 9 S. caespitosa 0.026 7.59 1.48 15.72 396.85 5.97 25.6 38.5 35.9 Clay-Loam

10 S. candidissima 0.009 7.85 1.48 0.82 18.82 0.19 85.1 9.1 5.7 Loam-Sand subsp. candidissima 11 S. ceratophylla 0.018 8.07 8.88 2.99 296.31 0.69 55.6 24.4 20.0 Sand-Loam 12 S. cerino-pruinosa 0.023 8.20 26.64 1.62 209.11 0.22 29.0 35.2 35.8 Clay-Loam 13 S. cilicica 0.009 7.77 1.30 2.45 142.76 0.53 40.7 23.8 35.5 Clay-Loam 14 S. cyanescens 0.021 7.79 24.16 2.31 227.33 0.82 52.3 15.6 32.1 Sand-Clay-Loam 15 S. dichroantha 0.040 7.59 8.68 2.57 32.06 0.44 47.6 31.9 20.5 Loam 16 S. divaricata 0.013 8.18 7.77 0.94 74.68 0.28 49.9 32.9 17.2 Loam 17 S. eriophora 0.056 7.29 0.75 14.10 396.85 4.49 15.0 29.2 55.8 Clay 18 S. euphratica 0.018 8.07 8.88 2.99 296.31 0.69 55.6 24.4 20.0 Sand-Loam var. euphratica

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Table 14 (continued) S. euphratica 0.018 8.07 8.88 2.99 296.31 0.69 55.6 24.4 20.0 Sand-Loam var. leiocalycina 19 S. frigida 0.004 8.02 2.25 8.26 86.67 0.94 81.3 9.2 9.5 Loam-Sand 20 S. glutinosa ------21 S. hedgeana 0.023 8.20 26.64 1.62 209.11 0.22 29.0 35.2 35.8 Clay-Loam 22 S. huberi 0.006 8.22 2.83 4.47 93.37 0.57 70.5 16.4 13.1 Sand-Loam 23 S. hypargeia 0.006 8.00 0.94 1.08 68.77 1.16 51.9 27.4 20.7 Sand-Clay-Loam 24 S. hydrangea 0.007 8.08 8.12 3.53 32.06 0.25 73.5 17.3 9.2 Sand-Loam 25 S. indica - 7.44 1.06 7.44 215.23 2.62 - - - Sand-Loam 26 S. kronenburgii 0.016 7.95 19.44 0.82 150.49 0.57 27.6 46.5 25.9 Loam

154 27 S. kurdica ------

28 S. limbata 0.014 7.93 16.90 16.00 296.31 2.10 43.9 25.9 30.2 Clay-Loam 29 S. longipedicellata 0.047 7.32 1.11 2.85 227.33 1.70 23.6 24.4 52.0 Clay 30 S. macrochlamys 0.009 7.93 34.91 8.00 106.68 0.57 51.7 27.5 20.8 Sand-Clay-Loam 31 S. macrosiphon 0.024 7.86 34.35 3.13 135.23 0.41 13.4 27.3 59.3 Clay 32 S. microstegia 0.023 7.79 23.50 1.76 52.08 2.83 49.3 28.0 22.7 Loam 33 S. montbretii 0.016 7.74 20.84 6.52 150.49 2.01 54.0 28.1 17.9 Sand-Loam 34 S. multicaulis 0.032 7.82 6.85 8.54 471.30 3.93 17.5 44.7 37.8 Silt-Clay-Loam 35 S. nemorosa 0.016 7.56 0.94 9.36 113.55 2.07 46.6 30.5 22.9 Loam 36 S. odontochlamys ------37 S. pachystachys 0.056 7.38 22.57 21.01 328.34 18.70 41.7 42.8 15.5 Loam 38 S. palaestina 0.026 7.82 3.89 21.01 445.84 2.04 42.8 31.3 25.9 Loam

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Table 14 (continued) 39 S. pilifera - 7.90 22.92 7.90 71.31 0.66 - - - Loam 40 S. pinnata 0.044 7.74 5.63 4.87 433.33 1.88 26.0 19.5 54.5 Clay 41 S. poculata 0.012 7.07 1.13 12.33 396.85 4.77 41.7 29.1 29.2 Clay-Loam 42 S. pseudeuphratica 0.005 8.09 34.79 2.85 127.82 0.63 7.16 14.9 13.5 Sand-Loam 43 S. recognita - 7.84 21.98 2.92 170.41 5.50 25.7 27.4 46.9 Clay 44 S. rosifolia 0.031 7.84 14.62 3.25 239.32 0.25 23.8 27.7 48.5 Clay 45 S. russellii 0.032 7.74 17.17 4.47 285.95 2.10 7.5 38.0 54.5 Clay 46 S. sclarea 0.016 7.92 25.10 3.93 218.14 0.63 27.4 29.6 43.0 Clay 47 S. siirtica ------48 S. spinosa 0.024 7.86 34.35 3.13 135.23 0.41 13.4 27.3 59.3 Clay 155 49 S. staminea 0.047 7.32 1.11 2.85 227.33 1.70 23.6 24.4 52.0 Clay 50 S. suffruticosa 0.018 8.07 8.88 2.99 296.31 0.69 55.6 24.4 20.0 Sand-Loam 51 S. syriaca 0.045 7.75 10.92 1.76 306.83 0.19 36.9 30.7 32.4 Clay-Loam 52 S. tomentosa 0.015 7.56 1.31 4.21 120.62 5.72 24.9 44.6 30.5 Clay-Loam 53 S. trichoclada 0.007 8.15 6.38 2.31 68.77 1.07 44.3 29.8 25.9 Loam 54 S. verticillata 0.007 8.15 6.38 2.31 68.77 1.07 44.3 29.8 25.9 Loam subsp. verticillata S. verticillata 0.044 7.70 34.79 13.15 497.42 2.20 33.2 27.8 39.0 Clay-Loam subsp. amasiaca 55 S. virgata 0.022 7.73 2.64 25.08 652.93 1.79 30.5 34.4 35.1 Clay-Loam 56 S. viridis 0.027 8.04 6.74 2.64 63.00 0.31 76.8 7.5 15.7 Sand-Loam 57 S. xanthocheila 0.007 8.05 22.27 4.87 63.03 1.57 76.9 14.2 8.9 Sand-Loam

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3.6. Assessment of Threat Categories of the Salvia Taxa

Destruction and fragmetation of habitats are the principal threat factors of biodiversity. Due to their destruction and fragmentation, many plant species today occur in smaller and isolated populations, which for a number of reasons are expected to face extinction (Matthies et al., 2004). As a result of rapid human population increase, many of the natural habitats in Turkey have been fragmented and reduced in size or degraded. Establishing monitoring programs and building quantitative databases for preservation programs is a crucial need to achieve future success in maintaining biodiversity in Turkey (Kaya & Raynal, 2001)

The assessment of conservation status of plant species is one of the most significant means in biodiversity conservation. Red Data Lists can play a crucial role by focusing attention on species most in need of conservation action (Balanca et al., 1998; Broughton & McAdam, 2002). The World Conservation Union (IUCN) Red List categories and criteria were preliminary constructed to assess the threat status of species or lower taxa at the global scale. However, many conservation efforts are conducted at national scales and there is consequently a demand for Red Lists at subglobal scales (Gärdenfors, 2001). For the purposes of regional conservation assessments there are significant reasons to assess species‟ extinction risk and publish Red Lists within specific geographically defined areas (IUCN, 2003).

Referring to the IUCN Red List, the threat categories of endemic and rare non-endemic plants in Turkey were first assessed by Ekim et al. (1989). At the national scale the Turkish Endemic Plants Project of 1995-1998 was carried out in order to re-evaluate the conservation status and distribution of the rare and endemic Turkish plant taxa and produce a new Red Data List. In the light of the obtained data, the Turkish Red Data Book (Ekim et al., 2000) was prepared using the 1994 Red List categories and criteria (IUCN, 1994). 33 Salvia taxa were listed under different threat categories in this Red Book. However, our new field observations and current knowledge on the distribution, population sizes and growing environments reveal that the previously assigned threat

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categories of the species are needed further clarifications owing to the lack of detailed field and herbarium studies. Therefore, the current conservation status of the Salvia taxa found in East and Southeast Anatolia are assessed at regional, national and global levels using IUCN Red List categories and criteria (IUCN, 2001), highlighted the principle threats to their survival and recommended priorities and strategies for conservation reasons.

The distribution of the threatened taxa according to regional (the study area), national, and global scale is as follows: at the regional scale: 6 taxa (10%) CR, 9 taxa (15%) EN, 5 taxa (9%) VU, 10 taxa (17%) NT and 29 taxa (49%) LC; at the national scale: 5 taxa (9%) CR, 5 taxa (9%) EN, 7 taxa (12%) VU, 10 taxa (17%) NT and 32 taxa (54%) LC; at the global scale: 5 taxa (9%) CR, 3 taxa (5%) EN, 6 taxa (10%) VU, 10 taxa (17%) NT and 35 taxa (59%) LC (Table 15, Figure 96).

As the distribution of the threatened taxa according to geographic regions is considered, it can be seen that they are classified in the following threat categories at global scale: in East Anatolia: 3 taxa CR, 3 taxa EN, 5 taxa VU, 10 taxa NT and 30 taxa LC; in Southeast Anatolia: 2 taxon CR, 1 taxon VU and 22 taxa LC (Table 15, Figure 96).

The distribution of taxa according to phytogeographical regions is as follows at global scale: 50 taxa Irano-Turanian elements (5 taxa CR, 3 taxa EN, 5 taxa VU, 10 taxon NT, and 27 taxa LC), 4 taxa Mediterranean elements (1 taxon VU and 3 taxa LC), 2 taxa Euro-Siberian elements (LC), and 3 taxa unknown or multiregional elements (LC) (Table 15, Figure 96).

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Figure 96. Distribution of the Salvia taxa studied according to the 2001 IUCN Red List categories at regional, national and global scales, phytogeographic elements, and geographic regions.

The most threatened species at global scale are S. anatolica, S. ballsiana, S. odontochlamys, S. pseudeuphratica and S. siirtica. S. ballsiana was previously categorized as Data Deficiency (DD) in the Turkish Red Data Book (Ekim et al., 2000) because it had not been collected since 1935. We visited its type locality during four years of field surveys, however no specimens were found despite many expeditions to its distribution area. This might be due to extinction of the species in this area. In 2008, our explorations revealed the presence of one population of the species in another location near Gerger (Adıyaman). The novel population was found in natural openings of Quercus scrubs. It was quite localized since it occupied an area of 1050 and 1160 m. According to IUCN (2001) categories and criteria at national scale, S. ballsiana is evaluated as Critically Endangered (CR) based on the B1ab(i, ii, iv) and C2a(ii) criteria (Table 15). The main threat to the species includes habitat destruction caused by overgrazing (Table 16). S. odontochlamys Hedge, known only from the type locality, was treated nationally as Endangered (EN) in the Turkish Red Data Book (Ekim et al., 2000). According to the recent field surveys, however, this species should be evaluated as CR because of smaller population size and narrower geographic ranges (Table 15). 158

Moreover, S. anatolica, S. pseudeuphratica and S. siirtica are categorized nationally as CR (Table 16). The principal threats to the future survival of these species are overgrazing, road construction or fire (Table 16).

S. cerino-pruinosa, S. eriophora, S. hedgeana, S. kurdica and S. macrosiphon, cover area of occupancy less than 500 km2 and are known at no more than 5 locations. Additionally, they are under threat from a wide variety of human activities, including overgrazing, agricultural practices, constructions and urbanization (Table 16). Therefore, they are evaluated nationally as Endangered (EN) based on B2ab(i,ii,iv) criteria (Table 15). 7 taxa are classified nationally as Vulnerable (VU) based on B2ab(i, ii,iv) since they cover area of occupancy less than 2000 km2 and are known at no more than 10 locations (Table 15). They are also subjected to serious threats, such as construction, overgrazing and land clearance (Table 16). 10 taxa are evaluated nationally as Near Threatened (NT) since they may be qualified as threatened in the near future by threats given in Table 16. The remaining 32 non-endemic taxa are Least Concern (LC) as they are widespread or abundant (Table 15).

In comparision with neighbouring countries (Post, 1933; Pobedimova, 1954; Zohary, 1966; Hedge, 1972, 1982a; Mouterde 1983), Turkey seems to have a higher number of Salvia taxa. The research area seems to cover 59 taxa, 24 of which are endemic, 3 of which are rare non-endemic and the remaining 32 taxa are widely distributed. This is more than the half of all Salvia taxa found in Turkey. Thus, there is a necessity to conserve such diversity and the significance of conserving the full extent of natural ranges of the taxa is reflected in the IUCN threat criteria. According to our field observations aiming to determine population size and the possible threats, 27 taxa are evaluated under CR, EN, VU or NT threat categories at national level. 7 endemic and 10 non-endemic taxa are restricted to the research area. Some of the taxa are relatively local and rare endemics, such as S. anatolica, S. ballsiana, S. odontochlamys and S. pseudeuphratica. The remaining 32 taxa are widespread and abundant wherever suitable habitat exists.

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Overgrazing, constructions (e.g. road and dam constructions), land clearing (e.g. agricultural practices), fire, urbanisation and tourism are the main factors threatening the survival of mostSalvia taxa in the study area. They are either affected by a single threat or by the combination of several types of threats. The threats along with some comments for each taxon are included in Table 16. Without the necessary measures taken in time to lessen the effects of the threats for plant conservation, plant diversity may decrease and some of the most threatened taxa, particularly those with small populations and restricted distributions will most likely become extinct in the near future. Therefore, we recommend that the following urgently measures are required for their conservation: (1) The most threatened taxa should be monitored regularly, (2) Special conservation programmes should be established for the richest areas in terms of the number of endemic taxa and ecologically sensitive areas, (3) The areas should be urgently modelled and managed by Geographical Information Systems (GIS), (4) Recovery programmes should be implemented for the establishment of new populations of the most threatened taxa, (5) Habitat destructions due to human impacts, especially including overgrazing, constructions and agriculture activities should be controlled, (6) Damaged habitats should be rehabilitated or restored, (7) The most seriously threatened taxa should be cultivated in the protected areas like national parks and botanical gardens, (8) Public and government should be better inform to protect and manage the threatened taxa and their awareness should be increased.

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Table 15. The recent IUCN threat categories with criteria and an estimated proportion (%) of the global population of the taxa distributed in the research area. Column 1: Taxa, Column 2: Threat categories according to Turkish Red Data Book (Ekim et al., 2000), Column 3: Recommended threat categories for regional scale, Column 4: Recommended threat categories for national scale, Column 5: Recommended threat categories for global scale, Column 6: An estimate of the proportion (%) of the global population occurring within the region, Column 7: IUCN Red List Criteria (2001). The taxa are listed in alphabetical order. +: the taxa endemic, *: taxa known only within the study area in Turkey, euro-sib.: euro-siberian element, ir.-tur.: irano-turanian element, medit.: mediterranean element, unk. or multi.: unknown or multiregional. The taxa are listed in alphabetical order. 1 2 3 4 5 6 7 1 +, ir.-tur.S. absconditiflora LC NT NT NT 20-25 - 2 unk. or multi.S. aethiopis - LC LC LC ? widely distributed 3 +, ir.-tur.S. anatolica CR CR CR CR 30-40 B1ab(i,ii,iv); C2a(ii) 4 * ir.-tur.S. atropatana - LC LC LC ? widely distributed 5 +, *, ir.-tur.S. ballsiana DD CR CR CR 100 B1ab(i,ii,iv); C2a(ii) 6 +, ir.-tur.S. blepharochlaena NT EN VU VU 10-15 B1ab(i,ii,iv) 7 ir.-tur.S. brachyantha - LC LC LC ? widely distributed 8 ir.-tur.S. bracteata - LC LC LC ? widely distributed 9 +, ir.-tur.S. caespitosa LC NT NT NT 50-60 - 10 ir.-tur.S. candidissima - LC LC LC ? widely distributed subsp. candidissima 11 ir.-tur.S. ceratophylla - LC LC LC ? widely distributed 12 +, *, ir.-tur.S. cerino-pruinosa - EN EN EN 100 B1ab(i,ii,iv) 13 +, medit.S. cilicica VU CR VU VU 5-10 B1ab(i,ii,iv) 14 +, ir.-tur.S. cyanescens LC VU NT NT 10-15 - 15 +, ir.-tur.S. dichroantha LC VU NT NT 10-20 - 16 +, ir.-tur.S. divaricata LC NT NT NT 85-90 - 17 +, ir.-tur.S. eriophora VU EN EN EN 50-60 B1ab(i,ii,iv) 18 +, ir.-tur.S. euphratica NT NT NT NT 85-90 - var. euphratica +, ir.-tur.S. euphratica NT NT NT NT 85-90 - var. leiocalycina 19 ir.-tur.S. frigida - LC LC LC ? widely distributed 161

Table 15 (continued) 20 euro-sib.S. glutinosa - NT LC LC ? widely distributed 21 +, *, ir.-tur.S. hedgeana CR EN EN EN 100 B1ab(i,ii,iv) 22 +, ir.-tur.S. huberi NT VU NT NT 10-20 - 23 +, ir.-tur.S. hypargeia LC NT NT NT 30-40 - 24 *, ir.-tur.S. hydrangea - LC LC LC ? widely distributed 25 ir.-tur.S. indica - EN VU LC ? B1ab(i,ii,iv) 26 +, *, ir.-tur.S. kronenburgii VU VU VU VU 100 B1ab(i,ii,iv) 27 *, ir.-tur.S. kurdica VU EN EN LC ? B1ab(i,ii,iv) 28 ir.-tur. S. limbata - LC LC LC ? widely distributed 29 +, ir.-tur.S. longipedicellata NT EN VU VU 10-20 B1ab(i,ii,iv) 30 *, ir.-tur.S. macrochlamys - LC LC LC ? widely distributed 31 *, ir.-tur.S. macrosiphon - EN EN LC 5-10 B1ab(i,ii,iv) 32 ir.-tur.S. microstegia - LC LC LC ? widely distributed 33 *, ir.-tur.S. montbretii - LC LC LC ? widely distributed 34 ir.-tur.S. multicaulis - LC LC LC ? widely distributed 35 unk. or multi.S. nemorosa - LC LC LC ? widely distributed 36 +, *, ir.-tur.S. odontochlamys EN CR CR CR 100 B1ab(i,ii,iv); C2a(ii) 37 ir.-tur.S. pachystachys - LC LC LC ? widely distributed 38 ir.-tur.S. palaestina - LC LC LC ? widely distributed 39 +, ir.-tur.S. pilifera LC VU VU VU 70-80 B1ab(i,ii,iv) 40 medit.S. pinnata - NT LC LC ? widely distributed 41 *, ir.-tur.S. poculata - LC LC LC ? widely distributed 42 +, *, ir.-tur.S. pseudeuphratica - CR CR CR 100 B1ab(i,ii,iv); C2a(ii) 43 +, ir.-tur.S. recognita LC EN VU VU 10-15 B1ab(i,ii,iv) 44 +, ir.-tur.S. rosifolia LC NT NT NT 30-40 - 45 ir.-tur.S. russellii - LC LC LC ? widely distributed 46 unk. or multi.S. sclarea - LC LC LC ? widely distributed 47 +, *, ir.-tur.S. siirtica - CR CR CR 100 48 *, ir.-tur.S. spinosa - LC LC LC ? widely distributed 49 ir.-tur.S. staminea - LC LC LC ? 50 ir.-tur.S. suffruticosa - LC LC LC ? widely distributed 51 ir.-tur.S. syriaca - LC LC LC ? widely distributed 52 medit.S. tomentosa - NT LC LC ? widely distributed 53 *, ir.-tur.S. trichoclada - LC LC LC ? widely distributed

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Table 15 (continued) 54 euro-sib.S. verticillata - LC LC LC ? widely distributed subsp. verticillata ir.-tur.S. verticillata - LC LC LC ? widely distributed subsp. amasiaca 55 ir.-tur.S. virgata - LC LC LC ? widely distributed 56 medit.S. viridis - LC LC LC ? widely distributed 57 *, ir.-tur.S. xanthocheila - LC LC LC ? widely distributed

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Table 16. The main threats to the taxa in the study area. +: the taxa endemic, *: taxa known only within the study area in Turkey. The taxa are listed in alphabetical order. Taxa Overgrazing Building of road, Land Urbanisation dam and other clearing and Tourism structures and Fire 1 +S. absconditiflora + + + 2 S. aethiopis + + 3 +S. anatolica + + 4 *S. atropatana + + 5 +, *S. ballsiana + 6 +S. blepharochlaena + + 7 S. brachyantha + + 8 S. bracteata + + 9 +S. caespitosa + + 10 S. candidissima + + subsp. candidissima 11 S. ceratophylla + + 12 +, *S. cerino-pruinosa + + 13 +S. cilicica + + 14 +S. cyanescens + + + 15 +S. dichroantha + + + 16 +S. divaricata + + 17 +S. eriophora + + 18 +S. euphratica + var. euphratica +S. euphratica + var. leiocalycina 19 S. frigida + + 20 S. glutinosa + + 21 +, *S. hedgeana + + 22 +S. huberi + + 23 +S. hypargeia + 24 *S. hydrangea + 25 S. indica, + + 26 +, *S. kronenburgii + + + 27 *S. kurdica +

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Table 16 (continued) 28 S. limbata + + 29 +S. longipedicellata + + 30 *S. macrochlamys + 31 *S. macrosiphon + 32 S. microstegia + + 33 *S. montbretii + + 34 S. multicaulis + 35 S. nemorosa + + 36 +, *S. odontochlamys + 37 S. pachystachys + 38 S. palaestina + + 39 +S. pilifera + + 40 S. pinnata + + 41 *S. poculata + + 42 +, *S. pseudeuphratica + + + 43 +S. recognita + + 44 +S. rosifolia + 45 S. russellii + + 46 S. sclarea + 47 *S. siirtica + + 48 *S. spinosa + 49 S. staminea + + 50 S. suffruticosa + + 51 S. syriaca + + 52 S. tomentosa + + 53 *S. trichoclada + 54 S. verticillata + + subsp. verticillata S. verticillata + + subsp. amasiaca 55 S. virgata + + 56 S. viridis + 57 *S. xanthocheila + +

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3.7. Numerical taxonomy of Salvia

In this part of the study, I seek to show how numerical taxonomy can aid the infrageneric treatment of Salvia as an addition to classical taxonomy. For the morphometric analysis sixty macromorphological, anatomical, palynological and mericarp micromorphologial characters examined and their codes for numerical analysis are shown in Table 17. Construction of the tree illustrating the relationships between the studied taxa and their infrageneric grouping was performed by using UPGMA.

Table 17. List of macromorphological, anatomical, pollen and mericarp micromorphological characters scored for the numerical taxonomic analysis. No Character Scoring 1 Life forms Annual: 0, perennial or biennial: 1 2 Stem branching Branched above: 0, branched below: 1, unbranched: 2 3 Stem indumentum Mostly antrorse eglangular: 0, mostly retrorsely eglandular: 1, pilose or villous: 2, scabridulous: 3, arachnoid: 4, lanate: 5, tomentosa: 6, pubescent: 7 4 Leaf shape Undivided: 0, pinnatisect: 1, lyrate: 1 5 Leaves Petiolate: 0, sessile or subsessile: 1 6 Petioles Ciliate with long eglandular hairs: 0, without ciliate hairs: 1 7 Bract shape Linear: 0, lanceolate: 1, elliptic: 2, oblong: 3 ovate: 4, absent: 5 8 Bract colour Green:0, pink to mauve: 1 9 Bract texture Membranous: 0, non-membranous: 1 10 Bracts or floral leaves Bracts or floral leaves nearly equal or shorter calyces: 0, bracts or floral leaves longer calyces: 2 11 Inflorescence Candelabriform: 0, non-candelabriform 12 Number of flowers in each 1-2: 0, 2-8: 1, 8-40: 3 verticillaster 13 Distance of verticillasters Distant: 0, close: 1 14 Calyx shape Campanulate: 0, infundibular: 1, tubular: 2, tubular- campanulate: 3, obtriangular: 4 15 Average length of calyx (mm) 16 Calyx texture in fruit Membranous: 0, non-membranous: 1 17 Calyx size in fruit Strongly enlarging: 0, Not or scarcely enlarging: 1 18 Calyx colour Green:0, green to purplish: 1 166

Table 17 (continued) 19 Upper lip of calyx Tri-dentate: 0, flat: 1, actinomorphic: 2, one-dentate: 3, truncate: 4 20 Calyx teeth Spinulose: 1, acute: 1, acuminate: 2, obtuse: 3, absent: 4 21 Average length of corolla (mm) Less than 20 mm: 0, more than 20 mm 22 Corolla colour Lilac: 0, blue: 1, yellow: 2, white: 3, mauve: 4 23 Upper lip colour of corolla Same to the lower: 0, different from the lower: 1 24 Upper lip shape of corolla Straight: 0, falcate: 1 25 Lower lip of corolla Spotted: 0, Unspotted: 1 26 Corolla tube Squamulate: 0, non-squamulate: 1 27 Corolla tube Ventricose: 0, straight: 1 28 Corolla tube Annualate: 0, non-annulate: 1 29 Stamen types A type: 0, B type: 1, C type: 2 30 Stamens Included within upper lip of corolla: 0, clearly exceeding upper lip of corolla: 1 31 Average size of pollen (P x E, 26.8-30.3 x 30.7-34.3: 0, 30.7-59.6 x 34.4-61.6: 1 µm) 32 Pollen sculpturing type Reticulate-perforate: 0, reticulate-granulate: 1, bireticulate: 2 33 Holes on colpus membrane Absent: 0, present: 1 sculpturing of pollen 34 One or several large central Absent: 0, present: 1 lumina in the primary lumen of pollen 35 Mericarp sculpturing type Colliculate: 0, reticulate: 1, verrucate: 2, rugose: 3 36 Shape of exocarp cells Pentan-hexangular: 0, rounded: 1, irregular: 2, absent: 3 37 Abscission scar diameter of 0.7-1.3: 0, more than 1.3: 1 mericarps (mm) 38 Mericarp colour Black: 0, brown: 1 39 Maximum row number of xylem Less than 8: 0, 8-40 rays in roots 40 Average length of xylem ray 14-25: 0, 25-50: 0 cells (µm) 41 Stem contour Quadrangular: 0, Quadrangular to circular: 1, Hexangular: 2 42 Average thickness of Less than 100: 0, more than 100: 1 collenchyma tissue in stems (µm) 43 Average thickness of the cortex Less than 200: 0, more than 200: 1 in stems (µm)

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Table 17 (continued) 44 Average length of vascular Less than 400: 0, more than 400: 1 bundles at the corners in stems (µm) 45 Average thickness of leaf blades Less than 200: 0, more than 200: 1 (µm) 46 Leaves in cross section Amphistomatic: 0, hypostomatic: 1 47 Epidermal cells in both sides Larger upper epidermal cells: 0, equal to each other:1 48 Hypodermis under both Absent: 0, present: 1 epidermis 49 Hypodermis under the upper Absent: 0, present: 1 epidermis 50 Mesophyll Isobilateral: 0, dorsiventral: 1 51 Shape of the adaxial side of Strongly convex: 0, flat: 1, flat to to slightly convex: midrib 2, flat to slightly concave: 3 52 Number of median vascular 1: 0; more than 1 bundle(s) in the midrib 53 Shape of median vascular Open U-shaped: 0, open crescent shaped bundle(s) in the midrib 54 Several inverse vascular bundles Absent: 0, present: 1 on the adaxial side of the midrib 55 Lateral vascular bundle(s) in the Absent: 0, present: 1 midrib 56 Petiole shape Open U-shaped: 0 U-shaped with obtuse or erect margins: 1, open V-shaped: 2, D-shaped with more or less procumbent margins: 3, obtuse triangular: 4, triangular: 5, open crescent shaped: 6 57 Number of median vascular 1: 0; more than 1 bundle(s) in the petiole 58 Shape of median vascular Open U-shaped: 0, U-shaped: 1, open crescent bundle(s) in the petiole shaped: 2, open V-shaped: 3 59 Several inverse vascular bundles Absent: 0, present: 1 on the adaxial side of the petiole 60 Sclerenchyma tissue on vascular Absent: 0, present only on the phloem: 1, present on bundles in the petiole both the phlowm and xylem: 2.

As a result of the analysis, we obtained the phenogram shown in Figure 97, in which we can separate seven major clusters. In the cluster 1, it is remarkable that, for the differentiation of S. russellii, S. verticillata subsp. amasiaca and S. verticillata subsp. verticillata in sect. Hemisphace, the type of stamen, the presence of hypodermis on the

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upper side of leaf blade and small pollen grains are the most valuable characters. In the cluster 2 presenting S. glutinosa in sect. Drymosphace, it clearly distinguished by the strongly falcate upper lip of corollas, hypostomatic leaves, an open V-shaped large vascular bundle in the middle of the petiole in cross section. In the cluster 3 presenting S. dicroantha, S. nemorosa and S. virgata in sect. Plethiosphace except for S. staminea, the clear distinction is the shape of the upper lip of the calyx in fruit. In the cluster 4, for the differentiation of the taxa of sect. Aethiopis, the most significant character is mostly D-shaped petioles with more than two inverse vascular bundles on the adaxial side. In the cluster 5, S. viridis in sect. Horminum is characterized by an annual growing character. In the cluster 6 presenting the taxa of sect. Hymenosphace, the most valuable character is membranous calyces in fruit. In the cluster 7, the species of sect. Salvia are characterized by mostly pinnatisect or trisect leaves.

The first infrageneric classification of Salvia was Bentham‟s 1833 arrangment, which listed 22 species from Turkey and divided them into seven sections based on leaf, calyx, corolla and stamen features: Eusphace (synonym of Salvia), Hymenosphace, Horminum, Aethiopis, Drymosphace, Plethiosphace and Hemisphace. His classification was followed by the 1879 arrangement of Boissier, which recognized 75 species of Salvia in Turkey, dividing them into seven sections. Hedge (1972) also revised Salvia in Europe using Bentham‟s sections. Though Hedge (1957, 1961) placed some Turkish Salvia species under Bentham‟s sections, the species were not referred to the sections in Flora of Turkey and the East Aegean Islands (1982). The results obtained from the present study support Bentham (1833) and Boissiers‟s (1879) taxonomic decisions on infrageneric delimitation.

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Figure 97. Phenogram showing the seven section of Salvia by means of UPGMA algorithm and Gower General Similarity. (1) Sect. Hemisphace, (2) Sect. Drymosphace, (3) Sect. Plethiosphace, (4) Sect. Aethiopis, (5) Sect. Horminum, (6) Sect. Hymenosphace, (7) Sect. Salvia.

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3.8. Systematics

The systematics the genus Salvia and its species is revised and discussed, and a key is given to 57 species, of which S. siiritica are a new species to science, S. macrosiphon is a new record for Turkey and S. cerino-pruinosa and S. pseudeuphratica are re-evaluated as valid species. S. ballsiana is also rediscovered after 73 years. The first sectional key is given. A key to all taxa, synonymy, type citation, amended and expanded descriptions, phenology, distribution and habitats in Turkey, general distribution outside Turkey, phtogeography, specimen citations on a grid square basis, some notes on taxonomy and photographs showing general appearance of the taxa are provided.

Salvia L., Sp. pl. 23. 1753.

Type: Salvia officinalis L., Sp. Pl., 1: 23 (1753).

Description: Shrubs, suffruticose chamaephytes or perennial (rarely biennial or annual) herbs, usually strongly aromatic. Stems erect ascending-erect or procumbent, solitary, several or many, branched or unbranched. Leaves opposite, simple, pinnatisect, trifoliate, pinnatified or lyrate, linear to orbicular (often oblong to ovate); margins very variable, entire, serrate, serrulate, erose, crenulate etc. Inflorescence (1-) 2-10 (40-)- flowered verticillate cymes, positioned along branched or unbranched axes. Calyx campanulate, infundibular or tubular, bilabiate, fruiting calyces slightly or clearly expanded in fruit and then membranous; upper lip tridentate; lower lip bidentate. Corolla bilabiate, tubular, white, pink, purple, blue, violet or yellow; upper lip straight or falcate, lower lip trilobed with a broad concave middle lobe and two small lateral lobes; tube straight or curved, invaginated, or ventricose, annulate or not, squamulate or esquamulate. Stamens 2, included within upper lip of corolla or much exserted, and with a short filament and a ± short or greatly elongated connective bearing at the upper end a fertile theca and, at the lower end, either a smaller fertile or subfertile theca (stamen type

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A) or variously shaped sterile tissue (stamen type B); stamens normally articulating at junction or filament and connective, occasionally not (stamen type C); staminodes (posterior pair stamens) present, small. Style filiform, apically two-lobed, exserted from corolla lips.

Key to sections of Salvia 1. Annuals……………………………………………………………………3. sect. Horminum 1. Biennials or perennials………………………………………..………………………………2 2. Fruiting calyx membranous-reticulate, infundibular with widely spreading rounded lobes.. ……..………………………………………………………………2. sect. Hymenosphace 2. Fruiting calyx thick-textured, tubular to infundibular with more or less dentate lobes…….. ……...……………………………….………………………..……...…….……………….3 3. Corolla tube annulate……………………………………………………………………4 4. Upper lip of corolla strongly falcate………...…………………5. sect. Drymosphace 4. Upper lip of corolla more or less straight………………………….…………………5 5. Staminal connectives shorter than or nearly equal to filaments, upper theca larger than the lower theca which bears at least some fertile pollen………1. sect. Salvia 5. Staminal connectives longer than filaments, the lower theca subulate and sterile… ……………………………………………….………………7. sect. Hemisphace 3. Corolla tube not annulate...……….……….…… …………………………………...6 6. Upper lip of calyx concave and 2-sulcate in fruit………...……6. sect. Plethiosphace 6. Upper lip of calyx not concave and not 2-sulcate in fruit………..…4. sect. Aethiopis

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Key to species of Salvia

1. Annuals……………………………….……………………………………………26. S. viridis 1. Biennials or perennials…………………………………………………………………………2 2. Fruiting calyx membranous-reticulate, infundibular with widely spreading rounded lobes… ………………………………………………………………………………………………3 3. Leaves pinnatisect; petioles often prominently ciliate with long white eglandular hairs…. ....…………………………………………………………………………………………4 4. Stems densely glandular villous, corolla 28-50 mm………….18. S. blepharochleana 4. Stems eglandular pilose, corolla 17-27 mm…………………………19. S. hydrangea 3. Leaves simple, occasionally with 1-2 pairs of small lateral lobes………………………5 5. Shrubs with woody stems at least below………….…………………………..………6 6. Flowers white; bracts and calyces pale yellow-green…………23. S. kronenburgii 6. Flowers pinkish to violet-blue; bracts and calyces green or purplish……………...7 7. Indumentum densely white lanate;inflorescence dense in fruit;corolla 15-20 mm …………………………………………………………..21. S. pseudeuphratica 7. Indumentum glabrous or pilose to villous; inflorescence lax in fruit; corolla 25 35 (-40) mm……………………………………………………………………8 8. Leaves elliptic (oblong to elliptic), always with a pair of small lateral lobes; petiole 2-5 (-7) mm, upper leaves sessile, mainly glabrous-pruinose (rarely pilose); corolla usually violet-blue…………………22. S. cerino-pruinosa 8. Leaves mostly ovate-oblong, rarely with a pair of small lateral lobes; petiole (8-)10-45 mm,upper leaves clearly petiolate,always pilose to villous; corolla usually pinkish……………………………………………20. S. euphratica 5. Herbs with woody rootstock and herbaceous stems………………….…….…………9 9. Calyx often purplish-violet, stem with or without dendroid hairs……..…………… …………………………………………………………………...24. S. multicaulis 9. Calyx often yellowish-green; stem always with dendroid hairs……….…………… ………………………………………………………………25. S. absconditiflora 2. Fruiting calyx thick-textured, tubular to infundibular with more or less dentate lobes...... 10 10. Staminal connectives shorter than or nearly equal to filaments, upper theca larger than the lower theca…………………………………………..…….……………..………11 11. Leaves simple…………………………………………………………………….12 12. Stems glabrous or subglabrous………………….……………………………13 13. Bracts present; pedicels 3-10 mm………………....……….17. S. ballsiana 13. Bracts absent; pedicels 10-35 mm…………………………1. S. divaricata 12. Stems pilose to villous above……………...…………………………………14 14. Bracts clearly longer than calyces………….…………4. S. macrochlamys 14. Bracts shorter than calyces……………………………………………….15 15. Inflorescence paniculate,leafy; verticillasters 1-2-flowered,pedunculate from upper leaf axis………………………………………3. S. kurdica 15. Inflorescence paniculate or not, leafless; verticillasters 4-10-flowered, not pedunculate………………………...…….………..2. S. tomentosa 173

11. Leaves pinnatisect or trisect……………………………………………………...16 16. Terminal leaf segment broadly oblong-ovate; petiole often without prominent ciliate hairs…………………………...………..…………..………...………17 17. Bracts or floral leaves clearly longer than calyces………………………18 18. Stems with very long spreading eglandular hairs;fruiting calyces 18-25 mm………………..….…………………..…….…….9. S. trichoclada 18. Stems with or without long spreading eglandular hairs;fruiting calyces to 18 mm.……………………………...……………….8. S. bracteata 17. Bracts nearly equal to calyces……………………...……………...…….18 19. Stem glabrous below……………………………..………6. S. pilifera 19. Stem glandular villous below and above……………………………20 20. Calyx prominently urceolate, corolla 25-30 mm……7. S. pinnata 20. Calyx ovate, corolla 30-40 mm…………………....5. S. recognita 16. Terminal leaf segment linear-lanceolate; petiole often prominently ciliate with long white eglandular hairs………………………....……………….………21 21. Corolla yellow or yellowish-white………………………………………22 22. Stems glandular and simple patent hairs above..…10. S. anatolica 22. Stems glabrous or almost so………………………...………………23 23. Stems stiffly erect;leaves with 1(-2) or no pairs of leaflets margins often entire, rarely crenulate;calyces with capitate glandular hairs; corolla upper lip yellow, lower lip white……..…...17. S. ballsiana 23. Stems ascending; leaves 2-3 (-4) pairs of leaflets,margins surrelate to serrate; calyces without capitate glandular hairs; corolla sulphur yellow or with lilac coloration…………………16. S. suffruticosa 21. Corolla white to pink, blue or violoet……………………………………24 24. Dwarf mat-forming subshrubs………………………………………25 25. Leaves trifoliate……….…………………………15. S. hedgeana 25. Leaves pinnatisect with 2-4 (-5) pairs of leaflets…………….....26 26. Racemes little exceeding leaf level; terminal leaf segment to 2 x 0.6 mm…………………………………13. S. caespitosa 26. Racemes elongate, clearly exceeding leaf level; terminal leaf segment to 4 x 1 mm…………..…….…14. S. pachystachys 24. Taller growing ascending erect to erect subshrubs………………….27 27. Terminal leaf segment linear-oblong,1-5-2.0 x 0.6-0.9 mm;calyx 7 11 mm; corolla 13-20 mm…………………………..11. S. huberi 27. Terminal leaf segment oblong-elliptic, 2-4 x 0.4-0.7 mm; calyx 10-15 mm; corolla 20-24 mm…………………….12. S. rosifolia 10. Staminal connectives longer than filaments, lower theca reduced…………………...28 28. Upper lip of corolla straight; corolla tube annulate, verticillasters 20-40-flowered; lower theca subulate……………………………………...……………………..29 29. Leaves oblong to ovate; calyx teeth mucronate….…………56. S. verticillata 29. Leaves linear-oblong; calyx teeth not mucronate……………...57. S. russellii

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28. Upper lip of corolla more or less falcate;corolla tube not annulate,verticillasters 2- 10-flowered; lower theca reduced to a usually dolabriform plate of tissue……..30 30. Corolla more than 20………………………………………………………..31 31. Corolla tube not squamulate, ventricose or not…………………...……32 32. Bracts clearly longer than calyces………………………………….33 33. Calyx ca. 10 mm in flower; bracts 14-18 x 11-15 mm...………… ..………………………………...………….….28. S. hypargeia 33. Calyx ca. 13-15 mm in flower; bracts 18-30 x 18-35 mm………. …………………………………………………29. S. monbretii 32. Bracts shorter than or nearly equal to than calyces…………...……34 34. Bracts ovate, yellow white brownish markings…51. S. glutinosa 34. Bracts broadly ovate, flowers white or lilac…………………...35 35. Stems hirsute below with long flattened eglandular hairs; flowers lilac to lilac-white……….……….32. S. palaestina 35. Stems pilose to villous or to tomentose below; flowers white ………………………………………….…………………36 36. Indumentum dense;calyx 15-20 mm in flower,more spiny fruiting calyces; corolla tube to 18 mm………………….. …………………………………………….30. S. spinosa 36.Indumentum sparse;calyx 12-17 mm in flower,less spiny fruiting calyces; corolla tube to 25 mm………………….. ……………………………………...31. S. macrosiphon 31. Corolla tube squamulate, ventricose….………..………………….……37 37. Upper lip of corolla lilac…………………………………………...38 38. Bracts clearly longer than calyces, pink to mauve...35. S. sclarea 38. Bracts not or scarcely longer than calyces, green……………...39 39. Lower lip of corolla dark violet; calyx truncate at apex……... ………………………………………………….49. S. indica 39. Lower lip of corolla lilac to violet………….6. S. cyanescens 37. Upper lip of corolla white………………………………………….40 40. Leaves linear to linear oblong………………..39. S. atropatana 40. Leaves oblong, ovate to orbicular…………………………….41 41. Under-surface leaf glabrous,prominently glandular-punctuate; stem retrorsely scabrid…………...……………39. S. limbata 41. Under-surface leaf pilose to pannose, not or scarcely glandular punctuate; stems not scabrid……………………………….42 42. Calyx pilose to glandular-papillose…………………….43 43. Leaves ± orbicular…….…….…………47. S. cilicica 43. Leaves oblong to broadly ovate….5. S. candidissima 42. Calyx with long spreading capitate glandular hairs……44 44. Leaves to 23 mm wide; verticillasters (4-) 6-12 flowered; calyx 5.5-10 mm in flower………………. ………………………………………..50. S. siirtica 175

44. Leaves to 14 mm wide; verticillasters 4-6- flowered; calyx 9-13 mm in flower…………………………45 45. Calyx campanulate-infundibular, 13-19 mm in fruit; stems to 45 cm………41. S. xanthocheila 45. Calyx campanulate, 11-14 mm in fruit; stems to 125 cm……………………...40. S. microstegia 30. Corolla less than 20 mm………………………..……………….…………..46 46. Flowers lilac, violet, purple or pinkish…………………………………47 47. Corolla tube squamulate……………………………………………48 48. Bracts clearly exceeding calyces, ± amplexicaul…………..…..49 49. Calyx obtriangular; bracts green or purplish, pilose…………. ……………………………………………….43. S. poculata 49. Calyx tubular-infundibular; bracts white-canascent….….. .………………………….……….44. S. odontochlamys 48. Bracts equal or shorter than calyces, not amplexicaul……...….50 50. Inflorescence axis, bracts and calyces dark violet blue; calyx teeth subulate; stems eglandular………….……………….51 51. Stems to 20 cm, densely arachnoid-lanate above; leaves 0.6-1.3 mm wide………..…...…………33. S. eriophora 51. Stems 30-50 cm, sparsely arachnoid-pilose above; leaves 0.8-4 (-11) mm wide……..…………34. S. brachyantha 50. Inflorescence axis, bracts and calyces green; calyx teeth acuminate or spinulose; stems glandular or not………...….52 52. Leaves ± orbicular, pannose……………..….…….……53 53. Cauline leaves ca. 4 pairs;inflorescence few-flowered. …………………………….……………47. S. cilicica 53. Cauline leaves several; inflorescence many-flowered (cf. also S. candidissima)….………46. S. cyanescens 52. Leaves ovate to oblong, pilose, villous or ± pannose….54 54. Leaves pilose to lanate beneath; calyx teeth short spinulose.………………….…..…46. S. cyanescens 54. Leaves arachnoid to sublanate beneath; calyx teeth long-spinulouse………………….……42. S. frigida 47. Corolla tube not squamulate………………………………..………55 55. Bracts strongly coloured violet-bulue, prominent………...……56 56. Inflorescence axis green, bracts and calyces purple; stems glandular or eglandular……………...... ……54. S. nemorosa 56. Inflorescence axis, bracts and calyces dark violet-blue; stems eglandular……………………………...…………………..57 57. Stems to 20 cm, densely arachnoid-lanate above; leaves 0.6-1.3 mm wide……..………...………33. S. eriophora 57. Stems 30-50 cm, sparsely arachnoid-pilose above; leaves 0.8- 4 (-11) mm wide……………….34. S. brachyantha 176

55. Bracts green or lightly tinged purple, ± inconspicuous……...…58 58. Leaves oblong, …………...……….………55. S. dicroantha 58. Leaves ovate oblong to broadly ovate…………53. S. virgata 46. Flowers white, yellow or cream…….………….……..……..…………59 59. Leaves pinnatified with spreading linear segments; biennial…….….. .………………………..………………………..37. S. ceratophylla 59. Leaves not pinnatified; perennial, rarely biennial……...... ………60 60. Inflorescence candelabriform, widely spreading;stems eglandular lanate; biennial or perennial…….….……………36. S. aethiopis 60. Inflorescence not candelabriform; stems pilose or villous; perennials……………….……………………………………61 61. Stamens clearly exserted beyond upper lip of corolla….…… …………………………….……………..…52. S. staminea 61. Stamens included within upper lip of corolla….………….62 62. Pedicels (5-) 6-10 mm…….…….38. S. longipedicellata 62. Pedicels less than 5 mm………………………………63 63. Leaves regularly ovate, cordate, slightly erose to serrulate…………...…………...…….27. S. syriaca 63. Leaves oblong to broadly ovate to orbicular, cordate to cuneate, margins crenate to erose……….…….64 64. Leaves ± orbicular…………….…47. S. cilicica 64. Leaves oblong to broadly ovate or linear to linear-oblong………………………...………65 65. Leaves linear to linear oblong…...………… ..………………………….39. S. atropatana 65. Leaves oblong to broadly ovate………….66 66. Upper lip of calyx ± bisulcate in flower, clearly so in fruit and recurved…..…… ...………………………..53. S. virgata 66. Upper lip of calyx not bisulcate and not recurved in fruit………………….....67 67.Bracts amplexicaul,clearly exceeding calyces…………...…………….…68 68. Calyx obtriangular; bracts green or purplish, pilose….43. S. poculata 68. Calyx tubular-infundibular; bracts white-canscent……………….….. ……………44. S. odontochlamys 67. Bracts not amplexicaul,not or scarcely exceeding calyces…………………69 69.Leaves not densely lanate; upper lip of corolla subfalcate.42. S. frigida

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69.Leaves densely lanate or pilose with glandular and eglandular hairs; upper lip of corolla clearly falcate…….70 70. Leaves pilose with glandular and eglandular hairs….50. S.siirtica 70. Leaves densely lanate…….…71 71. Calyx campanulate , 11-14 mm in fruit; stems to 125 cm…...... …S. microstegia 71. Calyx campanulate to infundibular, 13-19 mm in fruit; stems to 45 cm….…. ……...41. S. xanthocheila

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Section 1: Salvia (Benth.) Hedge, Flora Europaea. 3: 188 (1972). Syn: Sect. Eusphace Benth., Lab. Gen. Sp. 194 (1833).

1. S. divaricata Montbret & Aucher ex Benth. in Ann. Sci. Nat. ser. 2, 6: 37 (1836). (Figure 98). Synonym: S. trigonocalyx Woronow in Monit. Jard. Bot. Tiflis 22: 10 (1912).

Type: [TURKEY B7?] in Cappadocia orientali [ad Euphratem], Aucher 1528 (holo. G!).

Description: Perennial herb with woody rootstock. Stems erect, purplish to green, 30-75 cm high, branched above, shortly eglandular retrorse pilose and sometimes glandular below with some sessile glands, glabrous or subglabrous above. Leaves mostly basal, simple or occasionally with one pair of small lateral lobes, narrowly oblong, (2-) 3-8 (-9) x 0.8-2.5 cm, obtuse, rounded to cuneate at base, adaxial surface sparsely eglandular pilose, abaxial surface densely short tomentose with sessile glands, rugose; margins crenulate; petiole 1-5 (-7) cm long. Inflorescence 17-35 cm long, an elongate, widely branched panicle, glabrous. Verticillasters 2-4 (-6)-flowered, clearly distant. Bracts absent. Pedicels 10-35 mm long, glabrous, rigid, erecto-patent. Calyx tubular- campanulate, purplish to green, 12-18 mm long, densely short glandular pilose and sometimes a few eglandular villous with many sessile glands, striate; upper lip obsoletely tridentate, acuminate. Corolla pink, lilac or white, 25-35 mm long, simple hairs with many sessile glands on outside of upper lip; tube straight below, widening above, 15-25 mm long, incompletely annulate; upper lip ± straight, broad; lower lip unspotted, non-squamulate. Stamens of type A, included within upper lip of corolla. Filaments 5-9 mm long; upper connectives 2.5-6 mm long, lower connectives 1.5-3.5 mm long; fertile anthers ca. 2-4 mm long, sterile anthers ca. 1-2 mm long. Style exserted, 27-37 mm long.

Phenology: Flowering in June and July. 179

Distribution and habitat in Turkey: This species is mainly distributed around Erzincan and Divriği (Sivas) in East Anatolia and rarely around Zara (Sivas) in Central Anatolia where it occurs on limestone slopes, mountain slopes and roadsides, at 1030-1870 m (Figure 102). Specimens examined: B7 Sivas: 26 km from Divriği to Kemaliye, 39o31′14′′N- 38o09′40′′E, 1535 m, 9.vii.2006, A. Kahraman 1260; Erzincan: Çilhoraz Mountain, Karaçamurlu, 39o35′30′′N-38o44′09′′E, 1870 m, 9.vii.2006, A. Kahraman 1267; Erzincan: Çiğdemli Mountain, Hayatlar hill, 39o35′34′′N- 38o44′26′′E, 1775 m, 9.vii.2006, A. Kahraman 1270; Erzincan: Erzincan-Kemah road, 40 km from Erzincan, 39o36′364′′N-39o08′331′′E, 1118 m, 15.vii.2007, A. Kahraman 1478; Erzincan: İliç to Refahiye, 50 km to Refahiye, 39o37′423′′N-38o29′127′′E, 1030 m, 25.vii.2008, A. Kahraman 1590; Erzincan: Mercan Mountains, above Yaylabaşı, 39o38′064′′N- 39o30′637′′E, 1528 m, 26.vii.2008, A. Kahraman 1595.

General distribution and phytogeography: S. divaricata is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. divaricata is similar to S. aucheri, however can easily distinguished from it by the very long pedicels, wide-branching glabrous panicles, mostly oblong basal leaves and different geographic distribution.

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Figure 98. General appearance of S. divaricata in its natural habitat.

2. S. tomentosa Miller, Gard. Dict. ed. 8, no. 2 (1768). (Figure 99). Synonym: S. grandiflora Etl., Salvia 17 (1777); S. grandiflora Etl. subsp. aegaea (Bornm.) Rech. fil. in Bot. Jahrb. 69: 501 (1939); S. grandiflora Etl. subsp. rotundifolia (Vis.) Rech. fil., Fl. Aeg. 519 (1943). Ic: Jacq., Eclog. Pl. Rar. 1: t. 36 (1811-16).

Type: Described from cultivated material, Miller (holo. BM!).

Description: Perennial clump-forming suffruticose herb. Stems several, 30-100 cm high, with vegetative shoots, ascending to erect, often branched above, eglandular tometose with sessile glands, rarely with capiatate glandular hairs above. Leaves simple, narrowly oblong to ovate, 2-12 x 0.8-5.5 cm, subobtuse to acute, rounded to cordate at base, rarely with small basal lobules, rugose; margins entire to crenulate; petiole 1.7-7.3 cm long. 181

Verticillasters 4-10-flowered, distant or condensed above. Bracts broadly ovate, 5-8 x 4- 8 mm, deciduous; bracteoles present. Pedicels (2-) 4.5-10 mm long. Calyx ± tubular, 10- 16 mm long, accrescent to 14-18 mm long in fruit, usually violet, pilose to villous with sessile glands, with or without capitate glandular hairs; upper lip shortly tridentate; lower lip bidentate; teeth shortly subulate-pointed. Corolla lilac, pink violet blue, rarely white, 21-35 mm long; tube straight, 18-22 mm long, incompletely annulate; upper lip straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from April to August.

Distribution and habitat in Turkey: This species is very rarely distributed in East Anatolia (only one record is present in Elazığ province and absent in Southeast Anatolia, but relatively widespread and abundant in the other regions of Turkey (Figure 102). It is often associated with Pinus brutia and P. nigra, Quercus pubescens, in macchie, on limestone and igneous slopes, at 90-2000 m.

Specimens examined: B7 Elazığ: Yaylım Mountain, above Baltaşı village, 1500 m, 29.vi.1983, H. Evren 1582 (AÜ! FUH!).

General distribution and phytogeography: S. tomentosa is a member of the Mediterranean element and is found in Crime, Lebanon, Latakia, Balkans and Armenia.

Taxonomic notes: S. tomentosa resembles to S. aramiensis and S. fruticosa, but differs from them in the stem indumentum above. Leaves of the species are used as herbal tea.

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Figure 99. General appearance of S. tomentosa in its natural habitat.

3. S. kurdica Boiss. & Hohen. ex Benth. in DC., Prodr. 12: 268 (1848). (Figure 100).

Type: [IRAQ] ad rapes prope Amadia, 31.vii.1841, Kotschy 347 (holo. G! E! K! MO!).

Description: Perennial suffruticose herb. Stems 20-50 cm high, brittle, ± erect densely pilose to villous with densely capitate glandular hairs. Leaves simple without lateral segments, broadly ovate to suborbicular, 3.5-7 x 3-6.5 cm, cordate, glandular-pilose and eglandular villous, rugulose; margins crenulate; petiole 2-4 cm 183

long, densely glandular pilose. Inflorescence paniculate, few-flowered, leafy. Verticillasters 1-2-flowered, pedunculate from upper leaf axils, distant. Pedicels 8- 15 mm, pilose. Calyx tubular-infundibular, green to purplish, 10-20 mm long, broadening to 22 in fruit, densely glandular pilose to villous; upper lip shortly tridentate, longer than lower. Corolla bright pink, 30-33 mm long; tube straight below, widening above, 23-25 mm long, incompletely annulate; upper lip ± straight. Stamens of type A. Filaments ca. 6-7 mm long; upper connectives ca. 5-6 mm long, lower connectives ca. 2.5 mm long; fertile anthers ca. 3-3.5 mm long, sterile anthers ca. 1-1.5 mm long. Style ca. 40 mm long, glabrous, exserted from corolla lips and divided in two parts at apex.

Phenology: Flowering in May.

Distribution and habitat in Turkey: This species is only known from one small population at foot of Cudi Mountain in Silopi (Şırnak) in Southeast Anatolia where it grows on limestone cliffs, at ca. 900 m (Figure 102).

Specimens examined: C9 Şırnak: Hessana (Kösreli) at south foot of Cudi Mountain, 900 m, 10.v.1966, D. 42770 (E! K!).

General distribution and phytogeography: S. kurdica is a member of the Irano- Turanian element and present in the northern part of Iraq.

Taxonomic notes: S. kurdica is distinctive in having cordate leaves and short axillary inflorescences.

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Figure 100. Photograph of a herbarium sheet of S. kurdica.

4. S. macrochlamys Boiss. & Kotschy in Boiss., Fl. Or. 4: 595 (1879).

Type: [TURKEY B8 Muş] in schistosis alpium prope Musch Armeniae, 1525 m, [9.ix.1859], Kotschy 508 (holo. G! iso. W!). (Figure 101).

Description: Perennial herb. Stems many, 30-50 cm high, numerous, procumbent to almost erect with sterile shoots, branched above, leafy, densely glandular pilose and sparsely eglandular villous. Leaves simple, ovate, oblong to elliptic, 3.5-9 x 2-7.5 cm,

185

mostly glandular pilose with few sessile glands, without a pair of small lateral lobes, cordate or rounded; margins crenulate to serrulate; petiole 2-5 cm long. Inflorescence 10- 35 cm long, shortly branched paniculate or not, densely glandular pilose and sparsely eglandular villous. Verticillasters 2-flowered, approximating. Bracts longer than calyx, ± enclosing flowers, ovate to ovate-oblong, 25-45 x 15-25 mm, densely glandular pilose and sparsely eglandular villous, membranous, prominently veined, greenish-white; bracteoles present. Pedicels 1-3 (-4) mm long, rigid, erecto-patent. Calyx tubular-ovate, 14-20 mm long, greenish-white, glandular villous to pilose, fruiting calyx thick-textured; upper lip tridentate, acuminate; teeth 2-5 mm long. Corolla upper lip white and lower lip pink, 23- 35 mm long, simple hairs on outside of upper lip; tube straight below, widening above, 18-25 mm long, annulate; upper lip ± straight, truncate; lower lip spotted, non- squamulate. Stamens of type A, included within upper lip of corolla. Filaments 5-9 mm long; upper connectives 3-6 mm long, lower connectives 3-6 mm long; fertile anthers ca. 3-5 mm long, sterile anthers ca. 1-2 mm long. Style 25-40 mm long.

Phenology: Flowering from June to August.

Distribution and habitat in Turkey: This species has very wide distribution in East Anatolia and rare distribution in Southeast Anatolia where it occurs on rocky limestone slopes and open slopes in Quercus scrub, at 900-2400 m (Figure 102).

Specimens examined: B7 Tunceli: Munzur Su, 1 km from Tunceli, 950 m, Hub.-Mor. 10764! B8 Bingöl: Karakoçan to Kiğı, 1060 m, Buttler 15819! B9 Bitlis: Dilektaşı, foots of Kambos Mountain, 38o18′028′N-41o59′015′′E, 1351 m, 8.vii.2007, A. Kahraman 1426; Van: Bahçesaray-Van road, 1-2 km to junction of Gevaş-Çatak road, around Yapılı (Dernis) village, 38o07′622′′N-43o05′997′′E, 2150 m, 9.vii.2007, A. Kahraman 1440; Van: Northwest of Artos Mountain, 38o18′882′′N-42o06′737′′E, 1662 m, 10.vii.2007, A. Kahraman 1444; Muş: Malazgirt, between Aktuzla and Karıncalı, 1600 m, 24.vi.2001, S. 1674 (VANF!). C8 Mardin: Bakakri, Sint. 1888:1200! Siirt: Eruh to Şırnak, 8 km to Şırnak near Yanılmazlar village, 37o42′269′′N-42o15′804′′E,

186

1195 m, 5.vi.2008, A. Kahraman 1540. C9/10 Hakkari: Şırnak to Hakkari, near Cemko bridge, 37o28′496′′N-43o32′427′′E, 1070 m, 7.vi.2008, A. Kahraman 1555. C10 Hakkari: Hakkari-Van road, ca. 5 km from Hakkari, 37o34′208′′N-43o46′057′′E, 1287 m, 7.vi.2008, A. Kahraman 1562; Hakkari: junction of Yüksekova-Hakkari road, 37o42′768′′N-44o03′357′′E, 1625 m, 7.vi.2008, A. Kahraman 1565.

General distribution and phytogeography: S. macrochlamys is a member of the Irano- Turanian element and is distributed in Iraq and Iran.

Taxonomic notes: S. macrochlamys is a very distinct species on account of very large membranous bracts, 2-flowered verticillasters and densely thick glandular villous stems.

Figure 101. General appearance of S. macrochlamys in its natural habitat.

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(1) S. divaricata (3) S. kurdica (2) S. tomentosa (4) S. macrochlamys Figure 102. Distribution of S. divaricata, S. tomentosa, S. kurdica and S. macrochlamys in the study area.

5. S. recognita Fisch. & Mey. in Ann. Sci. Nat. ser. 4, 1:33 (1854). (Figure 103).

Type: [TURKEY B5 Kayseri] Ali-Dagh, prope Kaiseriam, 1 849, Tchihatcheff (holo. P?).

Description: Perennial herb. Sterns 40-100 (-180) cm high, erect, branched above, glandular-villous above and below with long spreading hairs, glabrous and glaucous in middle. Leaves pinnate, occasionally simple with a large oblong-ovate terminal segment, 4-8 (-13) x 2.5-5 cm and 1-2 pairs of small lateral segments, glandular-pilose rugose: margins crenulate. Petiole 4-8 cm long. Verticillasters 4-6-flowered, clearly distant. Bracts up to 17 mm, deciduous. Pedicels 3-6 mm, ± erect. Calyx ovate, 9-15 mm, to 20 mm in fruit, green to purplish, glandular-villous; upper lip straight, very shortly tridentate, not mucronate. Corolla lilac-pink, 30-40 mm; tube straight, widening above, ± annulate ca. 14 mm from base; upper lip straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from May to August.

188

Distribution and habitat in Turkey: This species is very rarely distributed in East Anatolia (only one record present in Tunceli province), however, mainly found in Central and South Anatolia (Figure 106). It grows s on limestone and igneous slopes and screes in steppe and Quercus scrub, at 900-2200 m.

Specimens examined: B7 Tunceli: above Selepur, 1500 m, 23.vii.1957, D. 31607 (E! AÜ! FUH!).

General distribution and phytogeography: S. recognita is endemic to Turkey and is a member of the Irano-Turanian element.

Taxonomic notes: S. recognita is distantly related to S. pilifera and S. pinnata, but can be easily separated from them by the pink and larger corollas.

Figure 103. General appearance of S. recognita in its natural habitat.

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6. S. pilifera Montbret & Aucher ex Benth. in Ann. Sci. Nat. ser. 2, 6: 40 (1836). (Figure 104). Synonym: S. cataonica Boiss. & Hausskn. in Boiss., Fl. Or. 4: 602 (1879); S. purpurascens Post in J. Linn. Soc. (Bot.) 24: 438 (1888); S. amana Bornm. in Notizbl. Bot. Gart. Berlin 7(63): 28 (1917).

Syntypes: [TURKEY C6 Adıyaman] ad Akdağ in Tauro orientali [nr Besni, 1834], Aucher [1927, 1952] (G! W!).

Description: Perennial herb to ca. 1 m high. Stems ascending-erect from creeping rhizome, glabrous below, glandular pilose to villous with eglandular villous hairs above. Leaves pinnatisect with a broad ovate-oblong terminal segment, to 3-8 x 2.5-4 cm, and 1- 2 pairs of smaller lateral segments, submembranous, adpressed glabrous to sparsely pilose, margins irregularly crenulate, upper lateral segments sessile, lower lateral segments onlong to oblong-elliptic; petiole 1.5-5 cm long. Verticillasters 2-15-flowered, distant. Bracts 10-20 x 5-23 mm, long-acuminate, smaller than calyces. Pedicels 4-10 (- 12) mm long, erecto-patent. Calyx campanulate to infundibular, green, rarely purplish, 12-20 mm long, to 23 mm long in fruit, densely glandular villous or pilose with eglandular pilose to villous hairs; upper lip tridentate, median tooth larger, longer than lower. Corolla blue-purple with white markings, 20-35 mm long; tube straight, gradually widening towards throat, slightly pilose within; upper lip ± straight, non-squamulate. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering in April and May.

Distribution and habitat in Turkey: This species is gathered only from several localities in Southeast Anatolia (Figure 106). It is also distributed in the eastern Mediterranean region of Turkey. It is found on limestone slopes, in Quercus scrub and among bushes and macchie, at 200-2300 m.

190

Specimens examined: B7 Adıyaman: East of Adıyaman, Gerger, above Kaşyazı village, 38o03′244′′N-39o04′695′′E, 1210 m, 18.v.2008, A. Kahraman 1504; Malatya: 5 km East of Malatya, 1320 m, Hub.-Mor. 8973! C6 Adıyaman: Gölbaşı, Northeast of Harmanlı, 37o51′328′′N-37o45′478′′E, 946 m, 3.v.2008, F. Celep 1420!; Gaziantep: Sof Mountain, 1450-1500 m, 24.vi.1978, T. Ekim 3729 (ANK!).

General distribution and phytogeography: S. pilifera is endemic to Turkey and is a member of the Irano-Turanian element.

Taxonomic notes: S. pilifera resembles to S. pinnata and S. recognita, but differs from them in having glabrous stems below.

Figure 104. General appearance of S. pilifera in its natural habitat.

191

7. S. pinnata L., Sp. Pl. 27 (1753). Ic: Rouy, 111. Pl. Eur. Rar. 11: t. 269 (1899). (Figure 105).

Lectotype: East Arabia, herb. Clifford: 13.

Description: Perennial herb. Stems to (15-) 25-80 (-100) cm high, procumbent to erect, viscid, densely glandular villous. Leaves irregularly pinnatisect, with an ovate-oblong terminal segment to 4-8 x 2-5.5 cm and 2-5 pairs of sessile or petiolate lateral segments irregularly arranged, submembranous, glandular-pilose, margins crenate-serrate; petiole 4-13 cm long. Verticillasters 4-6 (-9) flowered, distant. Bracts 5-6 mm long, ovate, soon deciduous. Pedicels 5-18 (-20) mm long, erecto-patent. Calyx urceolate, often purplish, 8-15 mm long, scarcely expanding in fruit, densely glandular villous; upper lip truncate, obsoletely tridentate. Corolla mauve-pink, 25-30 mm long; tube gradually widening towards throat, longitudinally pilose within; upper lip ± straight, shorter than lower. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from March to May.

Distribution and habitat in Turkey: This species is found in all regions of Turkey except for East and Central Anatolia. In Southeast Anatolia, it is recorded from only two gatherings (Figure 106). It is found in cornfields, fallow fields and dry meadows, at 1-1060 m.

Specimens examined: C6 Gaziantep: Gaziantep to Nizip, Post. C9 Mardin: 17 km from İdil to Cizre, 600 m, D. 42426 (E!).

General distribution and phytogeography: S. pinnata a member of the Mediterranean element and is distributed in Cyprus, Palestine and Lebanon.

192

Taxonomic notes: S. pinnata resembles S. pilifera and S. recognita, but can be easily distinguished from them by having urseolate calyces.

Figure 105. General appearance of S. pinnata in its natural habitat.

(5) S. recognita (6) S. pilifera (7) S. pinnata Figure 106. Distribution of S. recognita, S. pilifera and S. pinnata in the study area. 193

8. S. bracteata Banks & Sol. in Russell, Aleppo 2(2): 242 (1794). (Figure 107).

Type: [SYRIA] Aleppo, Russell.

Description: Perennial herb, somewhat suffruticose at base. Stems several, often purplish, 20-60 (-80) cm high, often purplish, ascending or erect, densely glandular pilose to villous, sometimes with sometimes long eglandular villous hairs. Leaves pinnatisect, with an ovate to oblong terminal segment 2.5-8 (-11) x 1.5-4 (-6) cm and 1-2 (-5) pairs of smaller lateral segments, ± densely eglandular pilose; margins serrulate; petiole 1-5.5 cm long, sparsely ciliate. Inflorescence paniculate. Verticillasters 5-10-flowered, distant, ± enclosed by sub-membranous floral leaves, 15-30 x 9-17 mm. Bracts many, purplish, 14- 20 x 6-10 mm; bracteoles present. Pedicels 1-5 mm long. Calyx tubular-infundibular, 10- 16 mm long, scarcely expanding in fruit up to 18 mm long, glandular villous. Corolla pink to purplish, 20-35 (-42) mm long; tube 14-25 mm long, irregularly annulate; upper lip straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from May to July.

Distribution and habitat in Turkey: This species is distributed in all regions of Turkey, but relatively widespread and common throughout the East, Southeast and Central Anatolia (Figure 110). It grows in a wide variety of habitats including igneous and calcareous slopes, Quercus brantii, fallow fields, edge of vineyards, roadsides and waste places, at 50-2000 m.

Specimens examined: B6 Malatya: Darende to Malatya, 38o22′116′′N-37o57′105′′E, 1260 m, 17.v.2008, A. Kahraman 1499. B7 Elazığ: Malatya to Elazığ arası, 30 from Kale, after Yalındamlar village, 38o28′47′′N-39o02′23′′E, 1256 m, 17.v.2006, A. Kahraman 1123; Elazığ: Elazığ to Malatya arası, near Hankendi, 38o35′151′′N- 39o04′203′′E, 1125 m, 24.v.2007, A. Kahraman 1386; Erzincan: Divriği to Kemaliye, 15-20 km to Gümüşçeşme village, 39o20′40′′N-38o22′06′′E, 1146 m, 6.vi.2006, A.

194

Kahraman 1191. B8 Erzurum: near Aşkale, 1950 m, M. Zohary 67162. C6 Gaziantep: Karkamış to Nizip, 2 km to Nizip, 36o58′205′′N-37o46′307′′E, 490 m, 21.iv.2007, S. Bagherpour 386! Gaziantep: Nizip to Birecik, 450 m, D. 27924! C7 Şanlıurfa: Karacadağ, Siverek to Diyarbakır, 1250 m, 19.v.1957, D. 28314 (AÜ!). C8 Diyarbakır: 8 km from Diyarbakır to Çınar, D. 28765!

General distribution and phytogeography: S. bracteata is a member of the Irano- Turanian element and present in Palestine, Syria, Iraq and Iran.

Taxonomic notes: S. bracteata is very closely related to S. trichoclada, but differs in the stem indumentum (densely glandular pilose to villous with sometimes long eglandular villous hairs) and smaller fruiting calyces. In the East and Southeast Anatolia, S. bracteata intergrades with the clearly distinct species S. suffruticosa and forms fertile hybrids.

Figure 107. General appearance of S. bracteata in its natural habitat.

195

9. S. trichoclada Benth. in DC., Prodr. 12: 267 (1848). (Figure 108).

Type: [TURKEY B7?] in Mesopotamia, Aucher 1566 (G! K!).

Description: Perennial herb, more or less suffruticose at base. Stems several, often purplish and sometimes green, 20-70 (-85) cm long, with sterile shoots, branched below, ascending-erect, often very long spreading eglandular villous and pilose with rarely glandular pilose hairs. Leaves pinnatisect, with an ovate to oblong terminal segment (1.2-) 1.5-5.7 x (0.6-) 0.8-3.5 cm and 1-2 pairs of small lateral segments, glabrous or eglandular and glandular pilose; margins crenulate to serrulate; petiole 2-6 cm long, sparsely ciliate. Inflorescence 10-40 (-50) cm long, branched, paniculate, densely long eglandular villous with glandular pilose. hairs. Verticillasters 2-8-flowered, distant. Bracts approximately equal to calyx above inflorescence, longer than calyx below inflorescence, ovate- acuminate, (12-) 15-40 x (8-) 10-20 mm, spreading eglandular villous and glandular pilose hairs, purplish; bracteoles present. Pedicels 2-6 (-9) mm long, rigid, erecto-patent. Calyx tubular-infundibular, 10-18 x 8-15 mm, fruiting calyces to (15-) 18-25 mm with diverging lips, purplish, long eglandular villous with glandular pilose hairs; upper lip tridentate, acuminate. Corolla pink to purplish, (20-) 25-40 mm long, short capitate glandular hairs on outside of upper lip, long eglandular villous and short capiate glandular hairs on outside of lower lip; tube 18-30 mm long, irregularly annulate; upper lip straight; lower lip spotted, non-squamulate. Stamens of type A, included within upper lip of corolla. Filaments ca. 6-7 mm long; upper connectives ca. 3-5 mm long, lower connectives ca. 3- 5 mm long; fertile anthers ca. 3-4 mm long, sterile anthers ca. 1-2 mm long. Style 25-42 mm long.

Phenology: Flowering from May to July.

Distribution and habitat in Turkey: This species is distributed in East and Southeast Anatolia where it grows in a wide variety of habitats including rocky limestone slopes, stream, steppe, Quercus scrub, grassy places and fallow fields, at 300-2440 m (Figure 110).

196

Specimens examined: B7 Elazığ: Harput to Buzluk Caves, 38o42′26′′N-39o15′43′′E, 1360 m, 19.vii.2006, A. Kahraman1352B; Elazığ: Yaylım Mountain, near Şemik village, 38o29′671′′N-39o35′031′′E, 1335 m, 3.vi.2008, A. Kahraman 1532; Malatya: Arapkir to Divriği, 1 km after from Çiğnir village, 39o03′829′′N-38o21′419′′E, 1241 m, 25.vii.2008, A. Kahraman 1586. B8 Diyarbakır: N. of Malabad, 800 m, Watson et al. 1117. B9 Bitlis: at foots of Kambos Mountain, 38o18′006′′N-41o59′127′′E, 1282 m, 9.vi.2008, A. Kahraman 1584; Bitlis: Gayda to Bahçesaray, 3 km after from Nazar (Azere Nazer) bridge, 38o07′099′′N-42o30′986′′E, 1256 m, 9.vii.2007, A. Kahraman1428; Van: Soğuksu to Bahçesaray, 11 km to Bahçesaray, near Sisli village, 38o06′254′′N-42o40′874′′E, 1900 m, 9.vii.2007, A. Kahraman 1432. C8 Siirt: Siirt to Pervari, 5 km after from Siirt, 37o59′870′′-41o56′097′′E, 914 m, 5.vi.2008, A. Kahraman 1534; Siirt: Siirt to Eruh, near Demirkaya village, 37o48′871′′N-41o55′025′′E, 523 m, 5.vi.2008, A. Kahraman 1538; Diyarbakır: 10 km S. of Diyarbakir, 750 m, Watson 5588; Mardin: Mardin, Sint. 1888: 956. C9 Siirt: Eruh to Şırnak, near Yanılmazlar village, 37o42′269′′N-42o15′804′′E, 1195 m, 5.vi.2008, A. Kahraman 1541; Şırnak: Şırnak to Hakkari, between Şenoba and Taşdelen villages, 37o23′660′′N-42o51′767′′E, 997 m, 6.vi.2008, A. Kahraman 1545; Hakkari: Şırnak to Hakkari, Güneş to Çığlı, 37o18′709′′N-43o27′791′′E, 1137 m, 7.vi.2008, A. Kahraman 1553; Hakkari: Berçelan, 37o34′684′′N-43o44′206′′E, 1776 m, 7.vi.2008, A. Kahraman 1561; Mardin: Kasrik gorge, 9 km from Cizre, 350 m, D. 42664. C10 Hakkari: Şırnak to Hakkari, near Cemko bridge, 37o28′496′′N-43o32′427′′E, 1070 m, 7.vi.2008, A. Kahraman 1556; Hakkari: Hakkari-Van road, ca. 5 km after from Hakkari, 37o34′208′′N-43o46′057′′E, 1287 m, 7.vi.2008, A. Kahraman 1564.

General distribution and phytogeography: S. trichoclada is a member of the Irano- Turanian element and present in Iraq and Iran.

Taxonomic notes: S. trichoclada is very closely related to S. bracteata, but differs in the stems densely covered with very long spreading eglandular hairs and larger fruiting calyces.

197

Figure 108. General appearance of S. trichoclada in its natural habitat.

10. S. anatolica Hamzaoğlu & A. Duran. in Ann. Bot. Fennici 42: 215-220 (2005) (Figure 109).

Type: [TURKEY] B7 Sivas: 22 km from Divriği to Kemaliye, 1580 m, 30.v.2003, A. Duran 6159, Hamzaoğlu & Sağıroğlu (holo. KNYA; iso. GAZI! ANK! HUM, hb. Yildirimli)

Description: Perennial herb with woody rootstock. Stems few, 25-50 cm high, erect, unbranched, glabrous below, glandular and simple patent hairs above. Leaves mostly basal, pinnantisect, oblong-lanceolate with two or more lateral segments or rarely one pair on sterile shoots, terminal leaf segments ovate-elliptic to lanceolate, 1.6-6.7 x 0.8- 2.0 cm, glandular and sometimes with several simple hairs; margins crenulate, lateral 198

segments 4-24 x 2-8.5 mm; petiole 1-3 mm, densely glandular hairs. Inflorescence 15- 30 cm long, clearly exceeding leaves. Verticillasters 4-6-flowered, distant, internodes 2.5-5 cm long. Bracts narrowly lanceolate to elliptic, 10-20 x (3-) 5-15 mm, greenish; bracteoles present. Pedicels 3-7 mm long, ± erect. Calyx tubular-infundibular, greenish, 14-17 x 10-15 mm, to 19 mm long in fruit, densely glandular and patent simple hairs; upper lip tridentate, ca. 1 mm long, lower lip two teeth ca. 2-3 mm long. Corolla entirely yellow, 35-42 mm long, glandular and long simple hairs on outside of upper lip; tube 20- 25 mm long, non-squamulate, with an annulus in throat; upper lip ± straight, almost equalt to lower lip. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering in May to June.

Distribution and habitat in Turkey: This species is has a very local distribution found only between Divriği (Sivas) and Kemaliye (Erzincan) where it grows on calcareous slopes, in open Quercus scrubs and slightly moist places, at 1350-1650 m (Figure 110).

Specimens examined: B7 Sivas: İliç to Divriği, 39o29′487′′N 38o06′830′′E, 1350 m, 15.vii.2007, A. Kahraman 1482; Sivas: Kemaliye to Divriği, 39o31′286′′N 38o09′536′′E, 1544 m, 2.vi.2008, A.Kahraman 1526; ibid., 6.vi.2006, A.Kahraman & S.Bagherpour 304.

General distribution and phytogeography: S. anatolica is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. anatolica is distantly related to S. bracteata, S. trichocalada and S. blepharochleana, but can be separated from them on account of the entirely yellow corollas.

199

Figure 109. General appearance of S. anatolica in its natural habitat.

(8) S. bracteata (9) S.trichoclada (10) S. anatolica Figure 110. Distribution of S. bracteata, S. trichoclada and S. anatolica in the study area. 200

11. S. rosifolia Sm., Pl. Ic. Ined. 1: 5 (1789). (Figure 111). Type: [TURKEY] in Armenia legit Tournefort ('Salvia armenia foliis alatis crenatis minoribus odores man‟, hole. BM? iso. P-Tourn. 1121).

Description: Perennial herb, suffruticose. Stems many, 20-60 cm high, ascending to erect, unbranched, retrorsely eglandular pilose to pubescent, with or without villous hairs and with glandular hairs. Leaves pinnatisect with an oblong-elliptic terminal segment 2-4 x 0.4-0.7 cm and two smaller pairs of lateral segments, eglandular pilose with sessile glands, not rugose; margins entire to serrulate; petiole 0.6-3.0 cm long long, ciliate. Inflorescence paniculate. Verticillasters (2-) 4-14-flowered, approximating or not. Bracts ovate-acuminate, 6-14 x 3-6 mm, green-viololet; bracteoles present. Pedicels 4-9.5 mm long. Calyx campanulate, 10-15 mm long, to 12-17 mm long in fruit, pilose to villous with numerous sessile glands, often strongly violet-coloured; upper lip tridentate; lower lip bidentate. Corolla lilac-pink to violet, 20-27 mm long; tube ca. 13-16 mm long, annulate; upper lip straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from May to August.

Distribution and habitat in Turkey: This species is distributed in East and Norteast Anatolia where it occurs on rocky limestone and igneous slopes, loamy hills and in Quercetum screes and fieldsides, at 500-2350 m (Figure 116).

Specimens examined: A9 Kars: 22 km from Kağızman to Kars, 40o15′442′′N- 42o58′224′′E, 1581 m, 13.vii.2007, A. Kahraman 1470; 28 km from Kağızman to Kars, 40o16′875′′N-42o57′084′′E, 1834 m, 13.vii.2007, A. Kahraman 1471; Erzurum: Horasan, 1600 m, 8.vi.1957, Davis 29366 (ANK!). B7 Erzincan: Yaylabaşı, 1300-1400 m, 27.v.1995, A.A. Dönmez 6012 (HUB!); Sipikör Mountain, Bendola, Sint. 1890:3230. B8 Erzurum: Ilıca-Tercan road, 1900 m, 10.vii.1957, Davis 30850 (ANK!); Ağrı to Horasan, 1800 m, Lamond 2589; Tercan to Erzurum, 1800 m, Hub.- Mor. 15296. B9 Ağrı: Karaköse, Aktaş village, T.Baytop 4795 (1STE!).

201

General distribution and phytogeography: S. rosifolia is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. rosifolia is very closely related to S. huberi, but differs in the larger terminal leaf segments (2-4 x 0.4-0.7 mm, not 1-5-2.0 x 0.6-0.9 mm), longer calyces (10- 15 mm, not 7-11 mm), corollas (20-27 mm, not 13-20 mm) and tubes (13-16 mm, not 8- 13 mm), longer pedicels (4-9.5 mm, not 2-5 mm) and verticillasters with more flowers.

Figure 111. General appearance of S. rosifolia in its natural habitat.

202

12. S. huberi Hedge in Notes R.B.G. Edinb. 23: 559, t. 24 (1961). (Figure 112).

Type: [TURKEY A8] Erzurum: Yusufeli-Erzurum road, soil slopes, corolla pinkish mauve, 1100 m, 9.vii.1960, Stainton & Henderson 6104 (holo. E! iso. K!).

Description: Perennial herb, suffruticose. Stems many, 15-40 cm high, slightly ascending to erect, unbranched, retrorsely eglandular pilose to pubescent, with or without villous hairs and with glandular hairs. Leaves pinnatisect with a linear-oblong terminal segment 1.5-2.0 x 0.6-0.9 cm and two smaller pairs of lateral segments, eglandular pilose with sessile glands, not rugose; margins serrate; petiole 0.6-2.5 cm long long, ciliate. Inflorescence paniculate. VerticiIlasters 2-5 (-7)-flowered, distant. Bracts ovate- acuminate, 6-12 x 3-5 mm, green-viololet; bracteoles present. Pedicels 2-5 mm long. Calyx campanulate, 7-11 mm long, pilose to villous with numerous sessile glands, often strongly violet-coloured; upper lip tridentate; lower lip bidentate. Corolla pink to violet, 13-20 mm long; tube 8-13 mm long, annulate; upper lip straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from May to August.

Distribution and habitat in Turkey: This species is distributed mainly in Northeast Anatolia and occasionally in East Anatolia where it grows on rocky limestone and igneous slopes and soil slopes with Pinus sylvestris, at 1100-2200 m (Figure 116).

Specimens examined: A8 Erzurum: 41 km from Erzurum to Tortum, 40o13′341′′N- 41o28′545′′E, 2121 m, 14.vii.2006, A. Kahraman 1474; near Tortum, 40o19′629′′N- 41o33′728′′E, 1425-1455 m, 11.7.2006, A. Kahraman 1473; Balıklı to Öğüt, 40o38′32′′N-41o36′50′′E, 1425-1455 m, 11.7.2006, A. Kahraman 1295.

General distribution and phytogeography: S. huberi is endemic to Turkey and a member of the Irano-Turanian element. 203

Taxonomic notes: S. huberi is very closely related to S. rosifolia, but is differentiated by having narrower terminal leaf segments and shorter calyces, corollas and tubes, shorter pedicels and verticillasters with fewer flowers.

Figure 112. General appearance of S. huberi in its natural habitat.

204

13. S. caespitosa Montbret & Aucher ex Benth. in Ann. Sci. Nat. ser. 2, 6: 39 (1836). (Figure 113). Synonym: S. pectinifolia Fisch. & Mey., op. cit. ser. 4, 1: 33 (1854). Ic: Bull. Alp. Card. Soc. 39: 105 (1971).

Type: [TURKEY B6 Sivas] in Monte Saru-tchitchek (Sariçiçek) in Cappadocia Orientali, Montbret 2015 (iso. G! K! W!).

Descriptions: Dwarf suffruticose perennial forming mats to ca. 60 cm diameter. Stems procumbent to ± erect, eglandular pubescent below, pubescent to villous above with some capitate glandular hairs. Leaves pinnatisect, obovate in outline, terminal segments ± lanceolate, 0.6-2 x 0.1-0.6 cm, with 2-4-pairs of lateral ± oblong segments; margins crenate; petiole 0.5-2 cm long, often long-ciliate. Racemes condensed, scarcely exceeding leaf level. Verticillasters 2-6-flowered, approximating to distant. Bracts ovate, acuminate, 11-12 x 5-8 mm; bracteoles present. Pedicels 3-6 mm long. Calyx campanulate, often purplish, 10-14 mm long, to 15 mm long in fruit, indumentum variable, from eglandular pilose to villous with many sessile glands to densely capitates glandular hairs. Corolla often violet-blue to lilac-pinkish, rarely white, 18-40 mm long; tube straight, 11-30 mm long, annulate. Stamens of type A, included within upper lip of corolla

Phenology: Flowering from May to July.

Distribution and habitat in Turkey: This species is distributed in East, Central and South Anatolia where it occurs on rocky limestone and igneous slopes, terraces and steppe, at 1200-2700 m (Figure 116).

Specimens examined: B6 Sivas: Pınarbaşı to Gürün, near Güneşli village, 38o49′947′′N- 42o03′517′′E, 1869 m, 6.vii.2007, A. Kahraman 1404; Sivas: around Yağdonduran Pass, 1650-1850 m, 8.vii.2006, A. Kahraman 1248; Kayseri: 3 km from Sarız to Pınarbaşı,

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38o30′01′′N 36o29′53′′E, 1700-1800 m, 8.vi.2006, A. Kahraman 1233; Kayseri: Sarız, Binboğa Mountain, near Elmaçaz, 38o25′774′′N-36o30′283′′E, 1748 m, 23.vii.2008, A. Kahraman 1584A; Kahramanmaraş: Nurihak Mountain to Elbistan, 2400 m, 17.vi.1960, Stainton & Henderson 5630 (E!). B7 Sivas: Arapkir to Divriği, Sarıçiçek village, 1772 m, 25.vii.2008, A. Kahraman 1587; Malatya: Arapkir, J. Dörfler 916 (E!); Erzincan: Hochadur Mountain (Sernek Mountain nr Gemergöp), Sint. 1890: 2676.

General distribution and phytogeography: S. caespitosa is endemic to Turkey and is a member of the Irano-Turanian element

Taxonomic notes: S. caespitosa is closely related to S. pachstachys and distantly related to S. hedgeana, but differs form the former in the smaller terminal leaf segments and racemes scarcely exceeding leaf level, and can be clearly distinguished from the latter by the pinnatisect leaves (not trifoliate).

Figure 113. General appearance of S. caespitosa in its natural habitat. 206

14. S. pachystachys Trautv. in Bull. Soc. Nat. Mosc. 41(2): 462 (1868). (Figure 114). Synonym: S. caespitosa sensu Boiss., Fl. Or. 4: 599 (1879) p.p. non Montbret & Aucher (1836); S. michajlowskii Sosn. in Monit. Jard. Bot. Tiflis 27: 8 (1913); S. caespitosa Montbret & Aucher subsp. pachystachya (Trautv.) Bordz. in Mém. Soc. Nat. Kiev 25: 110 (1915); S. armenorum Bornm. in Feddes Rep. 38: 158 (1935) e descr.; S. sintenisii Bornm., op cit. 159 (1935).

Type: [USSR, ARMENIA] Alagöz, Lagowski in herb. Trautvetter 4378 (LE, photo E!). Description: Caespitose or mat-forming suffruticose perennial herb. Stems several, green to purplish, up to 40 cm high, branched below, ascending-erect, eglandular pilose to villous hairs below, eglandular pilose to villous with capitate glandular hairs above. Leaves pinnatisect, obovate in outline, terminal segments ± lanceolate, 1.5-4.0 x 0.3-1.0 cm, with 2-4 (-5) pairs of lateral ± oblong segments, eglandular pilose hairs; margins crenate; petiole 2-7 cm, eglandular pilose to villous, often long-ciliate. Inflorescence 3-8 cm long, unbranched, glandular and eglandular hairs. Racemes condensed, clearly exceeding leaf level. Verticillasters 2-8-flowered, ± distant. Bracts ovate, acuminate, 15- 25 x 7-10 mm, eglandular villous with glandular hairs, green to purplish; bracteoles present. Pedicels 2-5 mm long. Calyx campanulate, green to purplish, 10-15 mm long, to 17 mm long in fruit, desnsely capitates glandular and eglandular villous with many sessile glands; upper lip tridentate, acuminate. Corolla white, pale yellow or pinkish, 27- 35 mm long, glandular and long eglandular hairs on outside of upper and lower lips; tube straight, 18-26 mm long, annulate, non-squamulate; upper lip ± straight. Stamens of type A, included within upper lip of corolla. Filaments ca. 8-9 mm long; upper connectives ca. 3-5 mm long, lower connectives ca 3-4 mm long; fertile anthers ca. 3-4 mm long, sterile anthers 1-2 mm long. Style 29-37 mm long.

Phenology: Flowering from May to July.

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Distribution and habitat in Turkey: This species is mainly distributed in East Anatolia and rarely in North Anatolia where it grows on rocky and stony limestone and igneous slopes, screes and subalpine pasture, at 1200-3200 m (Figure 116).

Specimens examined: A9 Kars: Mountains of E. of Kağızman, 20 km from Akçay to Cumaçay, 2200 m, 17.vii.1966, D. 46688 (ANK! E!). B7 Erzincan: Mercan Mountains, 39o37′386′′N-39o29′929′′E, 1877 m, 26.vii.2008, A Kahraman 1597. B9 Van: Gevaş, Northeast of Artos Mountain, 38o15′812′′N-43o07′186′′E, 2639 m, 10.vii.2007, A. Kahraman1442; Artos Mountain, 38o15′639′′N-43o06′997′′E, 2750 m, 10.vii.2007, A. Kahraman1443. B10: Kars: Ağrı Mountain, near Ak Bulak, 3200 m, 25.vi.1934, Koksch (type of S. armenorum).

General distribution and phytogeography: S. pachystachys is a member of the Irano- Turanian element and present in the west of Transcaucasica and the northwest of Iran.

Taxonomic notes: S. pachystachys is closely related to S. caespitosa, however differs in the larger terminal leaf segments and racemes clearly exceeding leaf level.

Figure 114. General appearance of S. pachystachys in its natural habitat. 208

15. S. hedgeana Dönmez in Bot. J. Linn. Soc. 137: 413-416 (2001). (Figure 115).

Type: [TURKEY] B7 Sivas: Divriği, Dumlucadağ, Karasar pass, above Kayaburun village, 1600 m, steppe, 15.6.1995, AAD 4579, slide no: AD 1390, AD 1391 (holo HUB! iso E!).

Description: Perennial suffruticose herb. Stems procumbent to ascending, 5-10 (-17) cm high, branched at base, with sterile shoots, retrorsely eglandular pilose hairs with many sessile glands. Leaves trifoliate, lanceolate in outline, with one pair of lateral segments, terminal leaf segment lanceolate, 2-3.5 (-4) × 0.3-0.5 cm, lateral segments (0.4-) 0.6-0.9 (-1.1) x 0.1-0.2 cm; margins entire, slightly involute; petiole 10-20 mm long, ciliate. Inflorescence racemose, little exceeding leaf level, retsorsely eglandular pilose. Verticillasters 2-4 (-6)-flowered, internodes 7-12 (-15). Bracts lanceolate, 8-15 x 2-3 mm; bracteoles present. Pedicels 5-10 mm long. Calyx campanulate, 10-16 mm, to 18 mm in fruit, retrorsely eglandular pilose with many sessile glands. Corolla 15-22 mm, blue, with a white spot on lower lip, densely villous eglandular hairs outside of upper lip; tube 8-15 mm, non-squamulate, annualate; upper lip ± straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from the end of May through June.

Distribution and habitat in Turkey: This species is represented by two populations in the southwestern of Divriği (Sivas) where it grows on limestone slopes and marly places at 1200-1600 (Figure 116).

Specimens examined: B7 Sivas: Divriği, Karasar Pass, above Kayaburun village, 39o16′53′′N-38o00′00′′E, 1460 m, 6.vi.2006, A. Kahraman 1159; Sivas: Divriği to Mursal, 39o15′889′′N-38o00′466′′E, 1217 m, 2.vi.2008, A. Kahraman 1521A.

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General distribution and phytogeography: S. hedgeana is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. hedgeana is distantly related to S. caespitosa, S. huberi and S. rosifolia, however can be easily separated from them by the trifoliate and longer leaves.

Figure 115. General appearance of S. hedgeana in its natural habitat.

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(11) S. rosifolia (13) S. caespitosa (15) S. hedgeana (12) S.huberi (14) S. pachystachys Figure 116. Distribution of S. rosifolia, S. huberi, S. caespitosa, S. pachystachys and S. hedgeana in the study area.

16. S. suffruticosa Montbret & Aucher ex Benth. in Ann. Sci. Nat. ser. 2, 6: 39 (1836). (Figure 117). Synonym: S. alexandri Pobed. in Fl. URSS 21: 653 (1954).

Type: [TURKEY B6 Kahramanmaraş] ad Akdog in Tauro orientalis, [1834], Aucher 2131 (holo. G, iso. K! W!).

Description: Perennial herb, somewhat suffruticose at base. Stems many, ascending to erect, 30-60 cm high, branched above, often yellowish-green, usually glabrous, occasionally pilose. Leaves pinnatisect, ovate in outline, terminal segment lanceolate, 2.5-5.5 x 0.5-2 cm, with 2-3 (-4) pairs of smaller lateral segments, pilose on veins with some sessile glands; margins serrulate to serrate; petiole 0.5-3 cm long. Verticillasters 2-8 (-10)-flowered, distant. Bracts ovate-acuminate, 8-17 x 4-8 mm; bracteoles present. Pedicels 3-8 mm long. Calyx campanulate, 8-13 mm long, to ca. 15 mm long in fruit and broadening, subglabrous to pilose-villous with dark subsessile glands; upper lip tridentate. Corolla sulphur yellow or with

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lilac coloration, 22-25 (-28) mm long; tube ca. 15 mm long, annulate; upper lip ± straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from April to June.

Distribution and habitat in Turkey: This species is mainly distributed in East, Southeast, Central and Anatolia and rarely in the Mediterranenan region of Turkey (Figure 119). It grows on banks in steppe, fallow fields and vineyards, at 300-2000 m.

Specimens examined: B6 Sivas: Divriği to Kemaliye, 1395 m, 6.vi.2006, A. Kahraman 1180; Malatya: Kangal to Hekimhan, near Hasançelebi village, 2.vi.2008, AKahraman 1527. B7 Elazığ: Elazığ to Pertek, 1300 m, D. 29190! B8 Muş: Muş to Erzurum, 26 km North of Muş, M. Zohary 57121. B9 Van: Van to Değirmenköy, 38o30′773′′N-43o29′572′′E, 1932 m, 8.vi.2008, A. Kahraman 1574; Muş: Malazgirt, Aktuzla village, 1600 m, 21.iv.2001, S. 1243 (VANF!). C7 Şanlıurfa: Şanlıurfa to Hilvan, 600 m, D. 28189 (ANK!). C8 Mardin: 5-10 km from Nusaybin to Mardin, 600 m, 22.v.1957, D. 28402 (ANK!); Siirt: Ramana Mountain, 20 km from Hasankeyf to Batman, 730 m, D. 42978!

General distribution and phytogeography: S. suffruticosa is a member of the Irano- Turanian element and present in Armenia, Syria and Iraq.

Taxonomic notes: S. suffruticosa is related to S. ballsiana, however can be easily distinguished by some morphological characteristics such as the ascending-erect stems, serrulate to serrate leaf margins with 2-3 (4) pairs of lateral segments, shorter and entirely sulphur yellow corollas or with lilac coloration. Some populations of S. suffruticosa in the East and Southeast Anatolia show hybridization with those of S. bracteata. Details of this hybrid are provided below.

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S. suffruticosa x S. bracteata Synonym: S. x spireaefolia Boiss. & Hohen. in Boiss., Diagn. ser. 1(5): 5 (1844); S. spiraeifolia Boiss. & Hohen. var. eriocalycina Bornm. in Beih. Bot. Centr. 28(2): 484 (1911).

Type: [SYRIA] prope Aleppum, Kotschy [256] (G! K!).

Description: Intermediate between the parents; flowers with various combinations of pale yellow, pink and lilac, calyx indumentum varying from pilose to villous, bracts purplish to green, ± enveloping calyces or not.

Phenology: Flowering in May and June.

Specimens examined: B7 Elazığ: Malatya to Elazığ, 30 km from Kale‟den, 38o28′794′′N-39o02′399′′E, 1251 m, 7.vii.2007, A. Kahraman 1414; Elazığ: Elazığ to Pertek, 1300 m, D. 291901 (ANK!); Adıyaman: Gerger, near Kaşyazı, 38o02′04′′N- 39o03′20′′E, 876 m, 19.v.2007, A. Kahraman 1370; Diyarbakır: 5 km North of Ergani, 1000 m, 2.vi.1957, D. 29015 (ANK!). B8 Erzurum (?): 117 km East of Erzincan, 27.v.1962, Andres (E!). C7 Şanlıurfa: Birecik to Şanlıurfa, 305 m, 17.v.1865, Hausskn. C8 Mardin: between Mardin station and Kızıltepe, 600 m, 26.v.1957, D. 28634 (E!); Mardin: Mardin to Savur, 11-12 miles from Mardin, 1200 m, 24.v.1957, D. 28516 (E! ANK!).

Habitat and distribution in Turkey: This hybrid is distributed in East and Southeast Anatolia where it grows on limestone banks, calcareous vineyards and fallow fields, at 305-1300 m.

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Figure 117. General appearance of S. suffruticosa in its natural habitat.

17. S. ballsiana (Rech. f.) Hedge in Notes R.B.G. Edinb. 23: 49 (1959). Synonym: S. suffruticosa Montbret & Aucher ex Benth. var. ballsiana Rech. f. in Ark. Bot. ser. 2, 1(5): 318 (1949). (Figure 118).

Type: TURKEY C6 Malatya: Karanik Dere, Erkenek to Geulbashe (Gölbaşı), Malatya, 1220 m, non-lime screes, 20.v.1935 E.K. Balls 2325 (holo. S! iso. AEF! E! K!).

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Description: Perennial herb, somewhat suffruticose at the base. Stems 35-90 cm high, stiffly erect, branched apically, glabrous or rarely sparsely glandular or eglandular hairy in the apical part. Leaves mostly basal, often pinnatisect with one or no pair of small lateral segments (rarely two pairs) or simple, (2.7-) 3-9 x (0.7-) 1.0-2.5 cm, narrowly oblong lanceolate, all over with a very fine indumentum, often entire-margined (rarely crenulate); petiole 2-7 cm long, with eglandular pilose to sparsely villose hairs. Inflorescence 15-30 cm long, branched or unbranched, paniculate or racemose. Verticillasters 2-10-flowered, clearly distant. Bracts conspicously shorter than calyces, bracteoles absent. Pedicels 3-10 mm long, glabrous, rigid, erecto-patent. Calyx campanulate, 10-13 mm long, up to ca. 15 mm long in fruit and scarcely broadening, densely glandular pilose with dark stalked glands; upper lip tridentate, acuminate; teeth 1-2 mm long. Corolla upper lip yellow, lower lip white, (24-) 26-35 mm long, glandular pilose with rarely sparsely eglandular hairs on outside of upper and lower lip; tube (12-) 14-18 mm long, annulate, non-squamulate; upper lip more or less straight; lower lip unspotted. Stamens of type A, upper connectives longer than filaments. Filaments 7-9 mm long; upper connectives 9-12 mm long, lower connectives 3-5 mm long; fertile anthers 5-6 mm long, sterile anthers 1-2 mm long. Style 35-45 mm long.

Phenology: Flowering in May.

Distribution and habitat in Turkey: This species was first collected from Gölbaşı (Malatya) by E. K. Balls in 1935, however since that time it has not been refound. In spite of many expeditions, it cannot found in the type location. Recently, it was rediscovered from another location (Gerger-Adıyaman) as a very small population (Figure 119). It grows on slopes in Quercus scrub and non-lime screes, at 1050-1220 m.

Specimens examined: B7 Adıyaman: East of Adıyaman, Gerger, above Kaşyazı village, near Dokuzdere, 38o03′277′′N-39o04′695′′E, 1050-1160 m, 18.v.2008, A. Kahraman 1505.

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General distribution and phytogeographyn: S. ballsiana is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. ballsiana is related to S. suffruticosa, but can be easily separated by the stiffly erect habit, taller stems, often entire leaf margins with one or no pair(s) of small lateral segments (rarely two pairs) and very fine decumbent hairs all over, longer petioles, much shorter acute calyx teeth, dense glandular indumentum with stalked glands on calyces and floral leaves, longer corollas, yellow upper lips and white lower lips. Hedge (1982a) reported that S. ballsiana had white upper lips of and yellows lower lips of corollas, however the recent results have revealed that it has yellow upper lips of and white lower lips of corollas.

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Figure 118. General appearance of S. ballsiana in its natural habitat.

(16) S. suffruticosa (17) S. ballsiana Figure 119. Distribution of S. suffruticosa and S. ballsiana in the study area. 217

Section 2: Hymenosphace Benth. Lab. Gen. Sp. 214 (1833). (Figure 120).

18. S. blepharochlaena Hedge & Hub.-Mor. in Notes R.B.G. Edinb. 22: 178 (1957).

Type: [TURKEY B6] Sivas: d. Kangal, Tecer-Gürun, Gipsschutt, 37 km nördlich Tecer, 1570 m, 27.v. 1955, A. Huber-Morath 13048 (holo. G!).

Description: Perennial herb, woody at base. Stems ascending-erect, 10-45 cm high, densely glandular villous, clothed at base with petiolar remains. Leaves mostly basal, pinnatisect, oblong in outline, glandular villous, with 3-5 pairs of lateral segments; terminal segment oblong-elliptic, serrate to serrulate, sessile, 0.1-0.2 x 0.4-0.8 cm; petiole to 30 mm long. Verticillasters (2-) 4-6-flowered, ± distant. Bracts of median verticillasters broadly ovate, ca. 2-3.5 x 1-2 cm, cordate. Pedicels 0-1.5 mm long. Calyx broadly infundibular, green or purplish-suffused, 18-24 mm long, to ca. 26 x 25 mm in fruit, with divergent lips, densely glandular-villous; upper lip subentire or indistinctly tridentate. Corolla white with a pale yellow upper lip and violet veined labellum, 28-50 mm long; tube to 35 mm long, broad, annulate; upper lip straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering in May and June.

Distribution and habitat in Turkey: This species is represented by large populations in Central Anatolia and by very small populations in East Anatolia and the Mediterranean region of Turkey where it grows on serpentine and limestone slopes, at 1000-2000 m (Figure 127).

Specimens examined: B6 Kayseri: Sarız to Pınarbaşı, near Aşağı Beyçayırı, 38o38′25′′N-36o26′28′′E, 1650 m, 20.vii.2006, A. Kahraman 1355; ibid., 17.vi.2008, A.Kahraman 1492A; Sivas: 18 km from Kangal to Gürün, 1564 m, 16.vi.2007, E. Karabacak 5501 (CBB!). 218

General distribution and phytogeography: S. blepharochleana is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. blepharochaleana resembles S. cadmica and S. anatolica, however is differentiated from the former by having pinnatisect basal leaves and from the latter by having smaller terminal leaf segments, white corolla with a pale yellow upper lip and violet veined labellum and larger calyces.

Figure 120. General appearance of S. blepharochleana in its natural habitat.

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19. S. hydrangea DC. ex Benth., Lab. Gen. Sp. 717 (1835). (Figure 121). Synonym: S. dracocephaloides Boiss., Diagn. ser. 1(5): 4 (1844); Schraderia hydrangea (DC. ex Benth.) Pobed. in Fl. URSS 21: 367 (1954) in obs., nom. inval.; S. dracocephaloides (Boiss.) Pobed., op. cit. 368 (1954), Arischrada dracocephaloides (Boiss.) Pobed. in Novit. Syst. Pl. Vasc. (Leningrad) 9: 247 (1972). Ic: Fl. URSS 21: t. 19 f. 2 (1954), as Schraderia dracocephaloides.

Type: PERSIA, Olivier (holo. G-DC).

Description: Tuft-forming suffruticose herb. Stems erect or ascending-erect, 20-60 cm high, branched below, adpressed tomentose-canescent with ± antrorse hairs, eglandular or with scattered sessile glands. Leaves pinnatisect with 2-3 pairs of lateral segments, (1.8-) 2-4 x 0.5-1.2 (1.4) cm, sparsely tomentose with antrorse eglandular hairs; terminal segment linear to oblong elliptic; margins often entire- margined, rarely crenulate; petiole 1-2.5 cm long. Inflorescence racemose. Verticillasters 4-12 (-15)-flowered, distant or approximating. Bracts of upper verticillasters ovate-acuminate, clearly shorter than calyces, 5-25 x 2-7 mm, lowermost transitional to leaves; bracteoles present. Pedicels erecto-patent, 2-7 mm long. Calyx purplish, campanulate, 10-14 x 8-12 mm in flower, 15-18 x 15-23 mm in fruit and membranous with divergent lips, tomentose or with some longer hairs; teeth short; upper lip subentire or 3-lobed. Corolla pink to magenta, 17-23 mm long, simple hairs on outside of upper lip, lower lip spotted; tube 10-15 mm long, annulate, non- squamulate; upper lip ± straight. Stamens of type A, included within upper lip of corolla, staminal connectives shorter than filaments. Filaments ca. 4-6 mm long; upper connectives 2-4 mm long, lower connectives 1.5-3 mm long; fertile anthers ca. 2-3 mm long, sterile anthers ca. 1 mm long. Style 17-27 mm long, exserted from corolla lips and divided in two parts at apex.

Phenology: Flowering from the end of May through August.

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Distribution and habitat in Turkey: This species is distributed in East Anatolia where it occurs on dry rocky volcanic, sandy and limestone slopes, mountain steppe, pastures and roadsides, at 600-2000 m (Figure 127).

Specimens examined: A9 Kars: 1 km from Kağızman to Kars, 40o09′332′′N- 43o06′692′′E, 1315 m, 13.vii.2007, A. Kahraman 1468. B9 Iğdır: foots of Ağrı Mountain, 39o47′17′′N-44o07′41′′E, 1318 m, 14.vii.2006, A. Kahraman 1317. B10 Ağrı: near Doğubayazıt, 14.vii.2006, A. Kahraman 1319B; Doğubayazıt, 1268 m, 16.vi.1971, D.M. Brown 541 (E!); Kars: Pamuk Mountain, 20 km from Iğdır to Doğubayazıt, 1600 m, D. 43875.

General distribution and phytogeography: S. hydrangea is a member of the Irano- Turanian element and present in Armenia and Iran.

Taxonomic notes: S. hydrangea is a very conspicuous and handsome species with large purplish calyces. Hedge (1982a) reported that S. hydrangea may hybridize with S. suffruticosa in the East Anatolia.

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Figure 121. General appearance of S. hydrangea in its natural habitat.

20. S. euphratica Montbret. & Aucher ex Benth. in Ann. Sci. Nat. Ser. 2, 6: 40 (1836). (Figures 122, 123).

Description: Tuft-forming aromatic perennial suffruticose herb. Stems ascending erect, glabrous or glandular pilose to villous, rarely eglandular, 25-50 (-60) cm high, branched below, with sterile shoots. Leaves simple, oblong to ovate-oblong, elliptic, (1.5-) 2-7.5 x (0.7-) 1.3-4 cm, rarely with a pair of small lateral basal lobes, pilose to villous with sessile glands, rugose; margins crenulate to serrate; petiole (8-) 10-25 (-30) mm long. Inflorescence unbranched, glandular pilose to villous, rarely eglandular. Verticillasters (2-) 4-10 (-12)-flowered, clearly distant. Bracts of upper verticillasters broadly ovate, bracts shorter than calyces (in var. euphratica), bracts longer than calyx (in var. leicalycina), 10-45 (- 50) x 10-30 (-35) mm, pilose, green to purplish, lowermost leaf- like. Pedicels 3-7 mm long, densely (e)glandular pilose to villous. Calyx green to purplish, glandular pilose to villous, rarely eglandular, broadly campanulate, (13-) 15-20

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(-25) x (10-)15-20 (-23) mm in flowering, broadening and expanding to 17- 32 (-40) x 17-25 (-45) mm in fruit, fruiting calyces membranous. Corolla violet-blue to pinkish, 25- 35 (-40) mm long, with simple hairs on outside of upper lip; tube 21-30 mm long, slightly curved and widening towards throat ± annulate. Stamens of type A, included within upper lip of corolla, stamina connectives shorter than filaments Filaments 4-6.8 mm long; upper connectives 2.2-4 mm long, lower connectives 2-3.7 mm long; fertile anthers hairless, 2.5-4.1 mm long, sterile anthers 1.2-2.1 mm long. Style (26-) 30-40 (- 44) mm long.

1. Calyx, bract and inflorescence axis with a ± dense indumentum, usually bracts shorter than calyx………………………………………………………………….var. euphratica 1. Calyx, bract and inflorescence axis ± glabrous, usually bracts longer than calyx……………………………………………………………………...var. leiocalycina

var. euphratica (Figure 122).

Syntypes: TURKEY B6/7 Malatya: in Cappadocia orientale (Aucher no. 1516, holotypes G!).

var. leiocalycina (Rech. f.) Hedge in Flora of Turkey, vol. 7: 431 (1982!). (Figure 123). Synonym: S. leiocalycina Rech. f. in Öst. Bot. Zeitschr. 99: 51 (1952!).

Type: Turkey B7 Erzincan: Eğin (Kemaliye), Salihli, in siccis montium, 25.6.1890, Sintenis 1890: 2753 p.p. (holo. W!, iso, LD, WU).

Phenology: Flowering in April and June

Distribution and Habitat: These taxa are mainly distributed in East Anatolia and occasionally in Central Anatolia where they grow on gypsum areas, calcareous slopes, 223

marly banks, river shingle, roadsides and limestone slopes with Quercus scrubs, at 800 and 1800 m (Figure 127).

Specimens examined for var. euphratica: B6 Malatya: 1-1.5 km from Darende to Malatya, 1000-1030 m, 16.v.2006, A. Kahraman 1098; Sivas: Gürün to Kangal, 1455 m, 1.vi.2008, A. Kahraman 1510B. B7 Malatya: Beydağı Mountain, between Gündüzbey and Kozluk, 1400 m, 16.v.2006, A. Kahraman 1116; Sivas: between Karasarbeli pass to Kayaburun village, 1505 m, 6.vi.2006, A. Kahraman 1155A; Erzincan: İliç to Refahiye, before 3 km from Kuzkışla village, 1344 m, 25.vii.2008, A. Kahraman 1592B.

Specimens examined for var. leicalyicana: B6 Malatya: 64 km from Darende to Malatya, near Develi village, 1325 m, 7.vi.2006, A. Kahraman 1216; Sivas: 9 km from Gürün to Gökpınar, 1544 m, 1.vi.2008, A. Kahraman 1511A. B7 Malatya: Arapkir to Divriği, near Çiğnir village, 1268 m, 25.vii.2008, A. Kahraman 1585(A); Sivas: Divriği to Kemaliye, near Demirdağ bridge, 1020 m, 6.vi.2006, A. Kahraman 1172; Erzincan: Kemaliye, near Gümüşçeşme village, river, 1120 m, 13.vii.2005, F. Celep 883A.

General distribution and phytogeography: S. euphratica var. euphratica and var. leiocalycina are endemic to Turkey and members of the Irano-Turanian element.

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Figure 122. General appearance of S. euphratica var. euphratica in its natural habitat.

Figure 123. General appearance of S. euphratica var. euphratica in its natural habitat.

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21. S. pseudeuphratica Rech. f. in Öst. Bot. Zeitschr. 99: 49-50 (1952!). (Figure 124).

Type: TURKEY B7 Elazığ: Keban-Maden, in declivibus saxosis, 20.vi.1889, Sintenis no. 817 (in hb. Mus. Wien, E!).

Description: Tuft-forming aromatic perennial suffruticose herb. Stems ascending to erect, eglandular lanate hairs, 20-30 (-35) cm high, branched below with sterile shoots. Leaves simple, ovate to oblong, 1.5-4 × (0.8-) 1-3.5 cm without any pairs of small lateral basal lobes, white lanate with sessile glands; margins crenulate to serrate; petiole (7-) 10-20 mm long. Inflorescence unbranched, rarely branched, eglandular lanate hairs; verticillasters 4-16-flowered, usually approximating. Bracts of upper verticillasters broadly ovate, shorter than calyces, 5-15 (-25) × 4-10 (-16) mm, lanate, greyish, lowermost leaf-like; bracteoles absent. Pedicels 2-4 (-5) mm long, eglandular lanate hairs. Calyx dark purplish, eglandular sparsely pilose to villous, broadly campanulate, fruiting calyces membranous, 10-15 × 10-16 mm long in flowering, broadening and expanding to (12-) 15-20 × (12-) 15-25 mm long in fruit. Corolla violet-blue, 15-20 mm long, with simple hairs on outside of upper lip; tube 10-15 mm long, slightly curved and widening towards throat ± annulate; upper lip straight. Stamens of type A, included within upper lip of corolla, connectives shorter than filaments. Filaments 2.8-4.7 (-5.2) mm long; upper connectives 1.6-3 mm long, lower connectives (0.8-) 1-2 (-2.3) mm long; fertile anthers hairless, 1.8-2.9 mm long, sterile anthers 0.9-1.3 mm long. Style (15-) 17-23 mm long.

Phenology: Flowering in May and the beginning of June.

Distribution and habitat in Turkey: This species is very local and scarce species recorded from three restricted localities near Keban (Elazığ) in East Anatolia where it grows on calcareous rocks, stony slopes and roadsides, at 750-900 m (Figure 127).

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Specimens examined: B7 Elazığ: 5 km from Keban to Elazığ, 850-900 m, 7.vi.2006, A.Kahraman 1200; Elazığ: 1.5-2 km from Keban to Elazığ, 750-800 m, 24.v.2007, A.Kahraman 1384; Elazığ: 3-4 km from Keban to Elazığ, 800-860 m, 24.v.2007, A.Kahraman 1385.

General distribution and phytogeography: S. pseudeuphratica is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. pseudeuphratica resembles S. euphratica however differs from them in the densely white lanate indumentum, dense inflorescence in fruit, entirely dark purplish calyces and smaller corollas. Its calyces appear more similar to those of S. multicaulis than S. euphratica.

Figure 124. General appearance of S. pseudeuphratica in its natural habitat. 227

22. S. cerino-pruinosa Rech. f. in Öst. Bot. Zeitschr. 99: 51 (1952!). (Figure 125).

Type: TURKEY Kharput, Karatasch, in declivibus supra Pekenik, Sintenis 698 (W!).

Description: Tuft-forming perennial suffruticose herbs. Stems ascending erect, 25-45 cm high, glabrous (rarely sparsely pilose), branched below. Leaves simple, elliptic (oblong to elliptic), 2.3-6 x 1.1-2.4 cm, always with a pair of small lateral basal lobes, glabrous to glaucose with sessile glands (rarely pilose), rugose; margins crenulate to serrate; petiole 2-5 (-7) mm long, upper leaves sessile. Inflorescence usually unbranched, glabrous (rarely pilose to villous), verticillasters 2-8 (-10)-flowered, clearly distant. Bracts of upper verticillasters broadly ovate, always shorter than calyces, 6-15 x 5-16 mm, usually green, glabrous (pilose only in Pertek population), lowermost leaf- like; bracteoles absent. Pedicels (2.5-) 3-7 (-10) mm long, erecto-patent, glabrous. Calyx green to purplish, glabrous, fruiting-calyces membranous, broadly campanulate, 15-20 x 12-20 mm in flowering, broadening and expanding to 19-25 x 18-30 mm in fruit, glabrous (rarely sparsely pilose). Corolla violet-blue, 25-35 mm long, outside of upper lip hairless; tube 20-25 mm long; slightly curved and widening towards throat ± annulate. Stamens of type A, included within upper lip of corolla, staminal connectives shorter than filaments. Filaments 4.3-8 mm long; upper connectives 3.1-4.6 mm long, lower connectives 2.7- 4.3 mm long; fertile anthers with usually long villous hairs, rarely glabrous, 3.0-3.8 mm long, sterile anthers 1.1-1.7 mm long. Style (25-) 27-35 (- 40) mm long.

Phenology: Flowering in May and June.

Distribution and habitat in Turkey: This species is rather local species representing by small isolated populations in Divriği (Sivas), Pertek (Tunceli) and the northern part of Elazığ where it grows on road slopes, field sides and marly banks, at 850 and 1450 m (Figure 127).

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Specimens examined: B7 Sivas: Divriği to Kemaliye, near Çobandurağı village, 1050- 1100 m, 6.vi.2006, A. Kahraman 1188; Sivas: Divriği to Kemaliye, 30 km to Gümüşçeşme, 850-900 m, 6.vi.2006, A. Kahraman 1189; Elazığ: 20 km from Elazığ to Pertek, 850-915 m, 3.vi.2008, A. Kahraman 1530; Tunceli: between Elazığ and Tunceli, before 23 km from Tunceli, 910 m, 10.vi.1981, E. Tuzlacı 46666 (ISTE!).

General distribution and phytogeography: S. cerinopruinosa is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. cerino-pruinosa is closely related to S. euphratica var. leicalycina, but differs in mainly glabrous-pruinose and oblong-lanceolate to elliptic leaves with always one pair of very small lateral lobes, very short petioles, and sessile stem leaves.

Figure 125. General appearance of S. cerino-pruinosa in its natural habitat. 229

23. S. kronenburgii Rech. f. in Öst. Bot. Zeitschr. 99: 50 (1952!). (Figure 126).

Type: TURKEY B9 Van: Wan, auf vulkanischem Boden, 2500 m, 27.vi.1899, Kronenburg 167 (holo. WU).

Description: Tuft-forming suffruticose herb. Stems ascending-erect, 30-45 cm high, branched below, with sterile shoots, villous with sparsely lanate below, glabrous and ± glaucous above. Leaves simple, ovate-oblong to oblong, 2.5-8 x (1.3-) 1.5-3 cm, rarely with a pair of basal lateral lobes, cordate or rounded, pilose pubescent above and below, rugulose; margins crenulate; petiole 1-3.5 cm long. Inflorescence racemose. Verticillasters 2-9-flowered, distant. Bracts of upper verticillasters broadly ovate, clearly shorter than calyces, 12-25 x 16-28 mm, pale gren, lowermost leaf-like. Pedicels 4-10 mm long, erecto-patent. Calyx pale green, broadly campanulate, 17-22 x 17-27 mm in flowering, 22-28 x 25-32 mm in fruit, glabrous, broadening and membranous in fruit; upper lip subentire. Corolla white, 30-50 mm long, simple hairs on outside of upper lip; lower lip unspotted; tube 25-35 mm long, straight below, widening and curved upwards above, annulate, non-squamulate; upper lip ± straight, broad. Stamens of type A, included within upper lip of corolla, staminal connectives shorter than filaments. Filaments 6-9 mm long; upper connectives 4-5 mm long, lower connectives 3-4.5 mm long; fertile anthers ca. 4 mm long, sterile anters ca. 1-2 mm long. Style 35-55 mm long, exserted from corolla lips and divided in two parts at apex.

Phenology: Flowering in June and July.

Distribution and habitat in Turkey: This species is distributed only in Van province in East Anatolia where it occurs on shale hills and step igneous slopes, 1800-2600 m (Figure 127).

Specimens examined: B9 Van: Van to Gürpınar, near Kurubaş pass, 2120 m, 11.vii.2007, A. Kahraman 1454; Van: 5 km from Köşebaşı village to Van, 1975 m,

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8.vi.2008, A. Kahraman 1575; Van: Çatak, 2 km north of Micinger suyu, 1900 m, 25.vi.1954, Davis & Polunin 23234 (ANK!).

General distribution and phytogeography: S. kronenburgii is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. kronenburgii is closely related to S. euphratica, but is differentiated by the entirely white corollas and pale yellowish-green bracts and calyces.

Figure 126. General appearance of S. kronenburgii in its natural habitat.

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(18) S. blepharochleana (21) S. pseudeuphratica (19) S. hydrangea (22) S. cerino-pruinosa (20) S. euphratica var. euphratica (23) S. kronenburgii (20) S. euphratica var. leiocalycina

Figure 127. Distribution of S. blepharochleana, S. hydrangea, S. euphratica var. euphratica, S. euphratica var. leiocalycina, S. pseudeuphratica, S. cerino-pruinosa and S. kronenburgii in the study area.

24. S. multicaulis Vahl, Enum. Pl. 1:225 (1805). (Figure 128). Synonym: S. acetabulosa Vahl, op. cit. 227 (1805) non L. (1767) excl. syn.; S. szovitsiana Bunge, Lab. Pers. 43 (1873); Schraderia ‘acetabulosa’ (Vahl) Pobed. in Fl. URSS 21: 369 (1954); Arischrada multicaulis (Vahl) Pobed. in Novit. Syst. Pl. Vasc. (Leningrad) 9: 247 (1972). Ic: Fl. URSS 21: t.18 f. 1 (1954), as Schraderia acetabulosa.

Type: „Hab. in Oriente. Ex. Herb. Desfontainii‟ (holo. C-Vahl).

Description: Perennial herb, mat-forming, with a woody rootstock. Stems erect, unbranches, 12-60 cm high, usually glandular pilose to villous, especially above, rarely glabrous, occasionally with dendroid hairs. Leaves simple, rarely with 1-2-pairs of small basal lobes, broadly ovate-elliptic to suborbicular, 2-4.5 (-7) x 1-3.5 cm, rugose; margins crenulate, with a dense indumantum of adpressed dendroid to dendroid-stellate hairs; petiole 1.5-6 cm long. Verticillasters 4-10-flowered, usually distant. Bracts broadly

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ovate, ca. 15 x 10 mm; bracteoles present. Pedicels 2-5 mm long, erecto-patent. Calyx campanulate, ca. 15 mm long, to 17-22 mm long in fruit and broadening, sparsely to densely glandular-pilose or villous, purplish-violet, rarely yellow-green; upper lip indistinctly 3-lobed. Corolla purplish-violet, rarely white, 15-22 mm long; tube straight, 11-14 mm long, annulate; upper lip straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from April to July.

Distribution and habitat in Turkey: This species is distributed in all regions of Turkey where it occurs where it occurs in a wide variety of habitats including rocky limestone and igneous slopes, shale and sandy slopes, scree, fallow fields, Quercus scrub, Artemisia stepe and Cedrus, at 550-2600 (Figure 130).

Specimens examined: A8 Erzurum: 30 km from Erzurum to Tortum, 1240 m, 11.vii.2006, A. Kahraman 1294B. A9 Kars: West of Kağızman (Grossheim 7: map 449). B6 Kahramanmaraş: Karaelbiatan to Elbisatan, 38o12′24′′N-37o09′42′′E, 1162 m, 27.iv.2006, A. Kahraman 1000; Kayseri: Pınarbaşı to Sarız, 38o40′58′′N-36o27′57′′E, 1602 m, 16.v.2006, A. Kahraman 1078; Sivas: Pınarbaşı to Gürün, 38o49′947′′N- 42o03′517′′E, 1869 m, 6.vii.2007, A. Kahraman 1405. B6/7 Malatya: Malatya to Adıyaman, around Gözene, 1058 m, 19.vi.2007, A. Kahraman 1367. B7 Malatya: 2 km from Malatya to Elazığ, 38o20′56′′N-38o23′01′′E, 1018 m, 28.iv.2006, A. Kahraman 1013; Sivas: Divriği, foots of Dumlucadağ, 39o21′31′′N-38o05′09′′E, 1177 m, 6.vi.2006, A. Kahraman 1168; Elazığ: Malatya to Elazığ, 30 km after from Kale, 38o28′47′′N- 39o02′23′′E, 1256 m, 17.v.2006, A. Kahraman 1122. B8 Erzincan: ca. 16 km from Tercan to Aşkale, 1631 m, 10.vii.2006, A. Kahraman 1287. B9 Van: 10 km from Van to Özalp, Van, above Çubuklu village, 2250 m, 16.vii.2006, A. Kahraman 1332; Bitlis: Kambos Dağı, Direktaşı (Tito), 2280 m, 9.vi.2008, A. Kahraman 1585C. C6/7 Malatya: ca. 6-7 km from Sürgü to Erkenek, 37o58′40′′N-38o00′48′′E, 1480 m, 28.iv.2006, A. Kahraman 1027. C7 Şanlıurfa: Şanlıurfa to Hilvan arası, 37o22′273′′N-38o34′784′′E,

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583 m, 28.iv.2006, A. Kahraman 1058. C8 Mardin: 16 km from Mardin to Midyat, 37o20′46′′N-40o49′28′′E, 933 m, 23.v.2007, A. Kahraman 1375C; Diyarbakır: Diyarbakır to Elazığ, 38o18′36′′N-39o43′26′′E, 1086 m, 24.v.2007, A. Kahraman 1385A. C9 Siirt: Eruh to Şırnak, 37o41′883′′N-42o16′790′′E, 1276 m, 5.vi.2008, A. Kahraman 1543A. C10 Hakkari: Zap river near turning to Yüksekova, Trelawny 1307.

General distribution and phytogeography: S. multicaulis is a member of the Irano- Turanian element and present in Syria, Iraq, Iran and Egypt.

Taxonomic notes: S. multicaulis is very closely related to S. absconditiflora, but is differentiated by often purplish violet calyces and stems with or without dendroid hairs.

Figure 128. General appearance of S. multicaulis in its natural habitat.

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25. S. absconditiflora Greuter & Burdet in Willdenowia 15: 77 (1985). (Figure 129). Synonym: S. cryptantha Montbret & Aucher ex Benth. in Ann. Sci. Nat. ser. 2, 6: 40 (1836).

Syntypes: [TURKEY B5/6?] in Cappadocia orientali, Montbret 2282 (W), Aucher 1531 (P). Description: Perennial herb, mat-forming, with a woody rootstock. Stems erect, unbranched, 10-40 cm high, ± always of branched to dendroid hairs and sessile glands (without glandular hairs). Leaves simple, without basal lobes, 1.2-6.0 x 0.6-2.5 cm, rugose; margins crenulate, with a dense indumantum of adpressed dendroid to dendroid- stellate hairs; petiole 1.3-6.5 cm long. Verticillasters 4-12-flowered, usually distant. Bracts broadly ovate; bracteoles present. Calyx campanulate, ca. 17 mm long, up to 25 mm long in fruit and broadening, often yellowish-green, rarely tinged purple, with or without some egalndular villous; upper lip generally longer than lower lip. Corolla white to pinkish (to ± purple) purplish-violet, rarely white, 13-21 mm long; tube straight, 11- 15 mm long, annulate; upper lip straight. Stamens of type A, included within upper lip of corolla.

Phenology: Flowering from April to July.

Distribution and habitat in Turkey: This species is mainly distributed in all regions mainly distributed in Central Anatolia and rarely in East and North Anatolia where it grows on rocky limestone slopes, dry steppe, fallow fields and roadsides, at 650-2500 m (Figure 130).

Specimens examined: B6 Malatya: 1 km from Darende to Malatya, 38o30′32′′N- 37o31′24′′E, 1011 m, 16.v.2006, A. Kahraman 1107. B7 Sivas: foots of Dumlucadağ, 39o21′41′′N-38o03′16′′E, 1500-1575 m, 18.v.2006, A. Kahraman 1166. B8 Erzincan: Erzurum to Erzincan, near Tercan, 39o46′211′′N-40o24′437′′E, 1447 m, 14.vii.2007, A.Kahraman 1477A.

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General distribution and phytogeography: S. absconditiflora is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. absconditiflora is very closely related to multicaulis, but differs in the stems with always of branched to dendroid hairs and often yellowish-green calyces.

Figure 129. General appearance of S. absconditiflora in its natural habitat.

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(24) S. multicaulis (25) S. absconditiflora Figure 130. Distribution of S. multicaulis and S. absconditiflora in the study area.

Section 3: Horminum Benth. Lab. Gen. Sp. 220 (1833).

26. S. viridis L., Sp. Pl. 24 (1753). (Figure 131). Synonym: S. horminum L., Sp. Pl. 24(1753)! Ic: Sibth. & Sm., Fl. Graeca 1: t. 19 & t. 20 (1806), as S. horminum; Hegi, Ill. Fl. Mittel-Eur. 5(4): f. 2488, 2489 (1927).

Lectotype: (designeted by Hedge) Herb. Clifford: 12, Salvia No. 3, (Jarvis C.E., Cafferty S. & Forrest L.L. (eds.), Taxon 50(2): 517 (2001), (photo no: BM-000557585 !).

Description: Annual. Stems simple or branched with 1-3 pairs of branches, (7-) 10-40 (- 45) cm high, glandular or eglandular pilose, topped by a usually violet coma or coma absent. Leaves simple, oblong-ovate, (1.5) 2-5 (-7) x (0.8-) 1-3 (-3.5) cm, prominently veined-beneath, rugose; margins finely crenulate; petiole 2-5 cm long. Verticillasters 4-6-flowered, distant or approximating. Bracts broadly ovate, ca. 10x10 mm; bracteoles present. Pedicels 2-3 mm long, erect, flattened. Calyx tubular, 7-11 mm long, to ca. 12-13 mm long in fruit and strongly deflexed, eglandular pilose or

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glandular; upper lip ± truncate. Corolla lilac-purple to white, 12-15 mm long; tube straight, glabrous within; upper lip broad, ± falcate. Stamens of type B.

Phenology: Flowering from March to July.

Distribution and habitat in Turkey: This species is distributed in all regions of Turkey where it grows on rocky slopes, macchie with Cistus, phrygana, sand-dunes, fields and waste ground, at 1-1300 m (Figure 132).

Specimens examined: B7 Erzincan: İliç station, 900 m, 26.vi.1972, T. Gözler & T. Kesercioğlu (EGE!). C6 Gaziantep: Gaziantep to Nurdağı, 550 m, 29.iv.2006, A. Kahraman 1077 ; Fevzipaşa, 600 m, 15.iv.1935, Balls 2130 (ANK!). C7 Adıyaman: Besni to Şanlıurfa, 37o31′645′′N-38o13′121′′E, 625 m, 28.iv.2006, A. Kahraman 1038. C8 Siirt: 3-5 South of Siirt, 700 m, P.H. Davis 43288. C9 Siirt: 26 km from Siirt to Pervari, 38o02′017′′N-42o03′462′′E, 807 m, 5.vi.2008, A. Kahraman 1536.

General distribution and phytogeography: S. viridis is a member of the Mediterranean element and present in most of the Mediterranean area, Crime, Cyprus, Caucasica, Iran (excep Central) and Iraq.

Taxonomic notes: S. viridis is only one annual species in Turkey and also very distinct based on the stems usually topped by a coloured coma of sterile shoots

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Figure 131. General appearance of S. viridis in its natural habitat.

(26) S. viridis Figure 132. Distribution of S. viridis in the study area.

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Section 4: Aethiopis Benth. Lab. Gen. Sp. 222 (1833).

27. S. syriaca L., Systema ed. 10: 854 (1759). Ic: Boul., Fl. Lib. Syr. t. 351 (1930). (Figure 133).

Type: Not designated (Hb. Linn. 42/15!)

Description: Perennial herb; rhizomatous. Stems 30-50 (-80) cm high, yellowish- green, erect, branched or simple, eglandular pubescent to villousbelow, denser above (and rarely glandular). Leaves simple, ovate, cordate, 5-17 x 3-10.5 cm, shortly eglandular or glandular pubescent with sessile glands below, rugose margins slightly erose to serrulate; petiole 3-6.5 cm long. Verticillasters 4-6- flowered, distant. Bracts ovate, 4-5 x 3-5 mm. Pedicels 3-8 mm long, erecto-patent. Calyx tubular, 5-10 mm long, scarcely expanding in fruit, densely eglandular or glandular pubescent; upper lip straight, tridentate. Corolla white, 8-12 mm long; tube straight, glabrous within; upper lip ± straight to slightly falcate. Stamens of type B.

Phenology: Flowering from April to July. Distribution and habitat in Turkey: This species is distributed in all regions of Turkey where it grows in steppe, marly banks, fallow and cultivated fields, at 350- 2200 m (Figure 136).

Specimens examined: A8 Erzurum: around Aziziye, 2074 m, 12.vii.2006, A. Kahraman 1302B. A9 Erzurum: 10 km Horasan to Kars, 1535 m, 12.vii.2006, A. Kahraman 1304B! Kars: East of Kızılcakcak (Grossheim 7: map 434). B6 Kahramanmaraş: Afşin to Tanır, 3 km to Tanır, 38o23′188′′N-36o54′575′′E, 1250 m, 5.vii.2007, A. Kahraman 1393; Malatya: Malatya to Adıyaman, ca. 4 km to Doğanşehir, 38o05′04′′N-37o54′32′′E, 1279 m, 17.v.2006, A. Kahraman 1127. B7 Erzincan: Erzincan to Kemah, ca. 8-9 km to Kemah, 39o38′860′′N-38o49′024′′E, 1282

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m, 15.vii.2007, A. Kahraman 1481; Elazığ: Malatya to Elazığ, 30 km to Kale, after Yalındamlar village, 38o28′47′′N-39o02′23′′E, 1251 m, 17.v.2006, A. Kahraman 1121. B8 Erzurum: 8 km Aşkale to Ilıca, 1700 m, 10.vii.2006, A. Kahraman 1290B; Diyarbakır: Lice to Diyarbakır arası, ca. 65 km to Diyarbakır, 38o18′675′′N- 40o31′895′′E, 789 m, 4.vi.2008, A. Kahraman 1533C. B9 Van: Gürpınar to Van, ca. 2 km to Kurubaş pass ca. 2 km, 38o24′459′′N-43o23′321′′E, 2180 m, 11.vii.2007, A. Kahraman 1453. C6 Gaziantep: Sof Mountain, 37o09′44′′N-37o12′44′′E, 1075 m, 20.v.2007, A. Kahraman 1374; Adıyaman: 15 km from Gölbaşı to Belören, 37o40′157′′N-37o33′115′′E, 1089 m, 3.v.2008, A. Kahraman 1491. C7 Şanlıurfa: Bozova to Şanlıurfa, around Kırkpınar village, 37o16′14′′N-38o41′32′′E, 17.v.2006, A. Kahraman 1128. C8 Diyarbakır: Diyarbakır to Elazığ, 38o18′36′′N-39o43′26′′E, 1086 m, 24.v.2007, A. Kahraman 1385B; Mardin: Mardin to Ömerli, 5 km to Ömerli, 37o22′37′′N-40o52′59′′E, 1123 m, 23.v.2007, A. Kahraman 1376. C9 Şırnak: Şırnak to Hakkari, near Taşdelen village, 37o23′660′′N-42o51′767′′E, 997 m, 6.vi.2008, A. Kahraman 1546!

General distribution and phytogeography: S. syriaca is a member of the Irano- Turanian element and is found in Syria, Iraq, Iran and Armenia.

Taxonomic notes: S. syriaca is a very distinctive oligomorphic species and is characterized by the tidy habit, regularly ovate leaves and small white corollas.

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Figure 133. General appearance of S. syriaca in its natural habitat.

28. S. hypargeia Fisch. & Mey. In Ann. Sci. Nat. Bot. ser. 4(1): 34 (1854). (Figure 134). Synonym: S. montbretii sensu Boiss., Fl. Or. 4: 611 p.p. non Bentham (1836). Ic. Tchihat., Asie Min. Bot. Atlas t. 22 (1860).

Type: [TURKEY B4 Kırşehir] inter Karadjeli (Isahocalı) et Gaman (Kaman), [900 m, 1849], Tchihatcheff (holo. P).

Description: Perennial herb forming tufts to 60 cm diameter. Stems erect, sturdy, 25-40 (-60) cm high, usually unbranched, eglandular arachnoid to lanate below, glandular pilose above. Leaves simple, mostly basal, linear to linear-oblong, 4-9 x 0.5-1 cm including indistinct petiole , greenish above, white lanate below; margins subentire. Verticillasters 4- 8-flowered, clearly distant. Bracts broadly ovate, 14-18 x 11-15 mm, lower surface lanate. Pedicels 2-3 mm long, ± erect. Calyx tubular-ovate, ca. 10 mm long, to ca . 12 mm 242

long in fruit, lanate and glandular; upper lip truncate. Corolla lavender to purplish-blue, ca. 25 mm long; tube 13-15 mm long, straight, slightly ventricose above; upper lip falcate. Stamens of type B.

Phenology: Flowering in June and July.

Distribution and habitat in Turkey: This species is distributed in all regions of Turkey except for Southeast Anatolia and the Aegean geographic region of Turkey. It occurs on limestone slopes and banks with Pinus brutia and fallow fields, at 800-2000 m (Figure 136).

Specimens examined: B6 Sivas: 9 km from Gürün to Gökpınar, 38o39′51′′N- 37o18′05′′E, 1525 m, 7.vi.2006, A. Kahraman 1228; Kayseri: Pınarbaşı to Sarız, 3-4 km to Sarız, 38o30′01′′N-36o26′53′′E, 1692 m, 8.vi.2006, A. Kahraman 1234; Kahramanmaraş: near Afşin, 38o14′814′′N-36o51′472′′E, 1442 m, 5.vii.2007, A.Kahraman 1392; Malatya: 64 km from Darende to Malatya, near Develi village, 38o22′59′′N-37o55′25′′E, 1326 m, 7.vi.2006, A. Kahraman 1215. B7 Sivas: Divriği to Kemaliye, near Demirdağ bridge, 39o25′29′′N-38o06′27′′E, 1017 m, 6.vi.2008, A. Kahraman 1173; Erzincan: Çiğdemli village , 39o35′30′′N-38o44′09′′E, 1268 m, 9.vii.2006, A.Kahraman 1268; Malatya: Arapkir, Sint. 1889:858; Elazığ: Keban Maden, Sint. 1889:844; Tunceli: 10 km N. of Pertek, 1800 m, It. Leyd. 1959:1572.

General distribution and phytogeography: S. hypargeia is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. hypargeia is closely related to S. montbretii, but differs in the narrower leaves, truncate upper lips of calyces, smaller bracts and calyces, and geographic distribution.

243

Figure 134. General appearance of S. hypargeia in its natural habitat.

29. S. montbretii Benth. in Ann. Sci. Nat. ser. 2, 6: 42 (1836). (Figure 135). Synonym: S. montbretii sensu Boiss., Fl. Or. 4: 611 (1879) p.p.

Type: [TURKEY C6 Gaziantep] ad Antab (Gaziantep), [1836], Montbret [1909] (K!).

Description: Perennial herb. Stems several, erect, sturdy, 25-50 (-65) cm high, usually unbranched, white eglandular arachnoid to lanate below, densely glandular pilose with sparsely eglandular villous hairs above. Leaves simple, mostly basal, linear-oblong, 5-17.5 x 1-2.1 cm including indistinct petiole, greenish above, white lanate below; margins sometimes suberose, Inflorescence green to purplish, 15-35 cm long. Verticillasters (2-) 4-8-flowered, clearly distant. Bracts clearly longer than calyces, ovate, 18-30 x 18-35 mm, green to purplish, white lanate with glandular hairs; bracteoles absent. Pedicels 2-6 244

mm long, usually pilose, ± erect. Calyx tubular-ovate, ca. 13-15 mm long, 16-18 mm long in fruit, white lanate with glandular pilose hairs; teeth 2-3 mm long, clearly tridentate upper lips with mucronate teeth; upper lip truncate. Corolla lavender to purplish-blue, 20-25 mm long, simple hairs on outside of upper lip; tube ± straight, 10-15 mm long, slightly ventricose above, incomplete annulate; upper lip ± falcate; , lower lip unspotted. Stamens of type B, upper connectives clearly longer than filaments. Filaments ca. 3 mm long; upper connectives ca. 3.5 mm, lower connectives ca. 3.5 mm long; fertile anthers ca. 2-3 mm long, sterile anthers ca. 1-2 mm long. Style 20-30 mm long, exserted from corolla lips and divided in two parts at apex.

Phenology: Flowering in May.

Distribution and habitat in Turkey: This species is distributed only in Southeast Anatolia where it grows on calcareous slopes, in Quercus brantii scrub, fallow fields and vineyards, at 700-1150 m (Figure 136).

Specimens examined: C6 Gaziantep: 11 East of Gaziantep, 780 m, H. Birand 56 (ANK!). C7 Şanlıurfa: Uetsch-Kara (Üç-Kara), Sint. 1888:804. C8 Mardin: Mardin to Midyat, 37o20′46′′N-40o49′28′′E, 933 m, 23.v.2007, A. Kahraman 1375; Mardin: 10 miles from Mardin to Savur, 1150 m, 24.v.1957, D. 28517 (E! ANK!); Mardin: Mardin to Diyarbakır, 24 km after from Mardin, 1000 m, 27.v.1957, D. 28707 (ANK! E!); Batman: Gercüş, 1003 m, 23.v.2007, A. Kahraman1379.

General distribution and phytogeography: S. montbretii is a member of the Irano- Turanian element and present in Iraq and Syria.

Taxonomic notes: S. montbretii is allied to S. hypargeia, but can be distinguished by the broader linear-oblong leaves, clearly tridentate upper lips of calyces with mucronate teeth, larger bracts and calyces and distinct geographic distribution.

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Figure 135. General appearance of S. montbretii in its natural habitat.

(27) S. syriaca (28) S.hypargeia (29) S. montbretii Figure 136. Distribution of S. syriaca, S. hypargeia and S. montbretii in the study area.

246

30. S. spinosa L., Mant. Alt. 511 (1771). (Figure 137). Synonym: S. aegyptiaca L., Mant. 26 (1767) non L. (1753). Ic: Jacq., Ic. Pl. Rar. 1: t. 7 (1781); Tackh., Stud. Fl. Egypt ed. 2: t. 159 (1974).

Type: Hab. in Aegypto, Forsskål (Hb. Linn. 42/43).

Description: Perennial herb. Stems single, erect, quadrangular, 30-60 cm high, branched above, glandular pilose to villous below, densely glandular above. Leavessimple, ± thin-textured, broadly ovate to ovate-oblong, 6.5-12 x (3-) 4-8 cm, ± tomentose, cordate to rounded; margins subentire to erose; petiole (1.5-) 3-11 cm long. Inflorescence paniculate. Verticillasters 2-6-flowered, distant. Bracts broadly ovate, 15-20 x 12-18 mm, pilose to villous, ± abrupte acuminate-hirsute; bracteoles absent. Pedicels 2-3 mm long, glandular pilose, erecto-patent Calyx tubular, greenish-yellow, 15-20 mm long in flower, slightly expanding and hardening to 21-23 mm long in fruit, glandular hirsute with sparcely eglandular villous; teeth 1-2 mm long; upper lip equally tridentate, spiny in fruit. Corolla white, 20-25 mm long; tube straight, 17-18 mm long, slightly widening above, glabrous within; upper lip falcate. Stamens of type B, staminal connectives longer than filaments, dolabriform.

Phenology: Flowering in May and June.

Distribution and habitat in Turkey: This species is found only in Southeast Anatolia where it ocuurs in disturbed marly steppe, edge of fallow fields and vineyards, at 300-900 m (Figure 140).

Specimens examined: C6 Gaziantep: Nizip to Birecik, ca. 5 km from Euphrates, 400 m, 14.v.1957, D. 27948 (K!); Gaziantep: 5 km Gaziantep to Nizip, 900 m, 29 .v.1978, A Baytop, E. Tuzlacı and K. Alpınar 40049 (ANK! ISTE!). C7 Şanlıurfa: Akçakale to Şanlıurfa, 40 km Şanlıurfa, near Balatlar village, 36o44′940′′N-38o57′171′′E, 356 m, 15.v.2008, S. Bagherpour 536!; Şanlıurfa 32 km Şanlıurfa to Akçakale, 450 m, D.

247

28173 (ANK!); Şanlıurfa between Zeytinbahçe to Abdallı villages, 5. km, 544 m, 13.vi.2006, M.M. Balos 1547 (E!). C8 Diyarbakır: Mardin to Diyarbakır, 46 km to Diyarbakır, 37o39′16′′N-40o28′21′′E, 759 m, 24.v.2007, A. Kahraman 1382; Mardin: Mardin station to Kızıltepe, 600-650 m, 26.v.1957, D. 28632 (ANK! E!). C9 Mardin: 1- 2 km from Cizre‟nin, 350 m, 6.v.1966, D. 42535 (K!).

General distribution and phytogeography: S. spinosa is a member of the Irano- Turanian element and present in Egypt, Palaestina, Syria, Lebanon, Iraq, Iran, Afghanistan, Taranscaucasia and Transcaspia.

Taxonomic notes: S. spinosa is very similar to S. macrosiphon, however can be separated by the denser indumentum, broader leaves and calyces, more spiny fruiting calyces and shorter corolla tubes.

Figure 137. General appearance of S. spinosa in its natural habitat. 248

31. S. macrosiphon Boiss. Diagn. Pl. Or. Nov. Ser. 1, 5: 11 (1844). (Figure 138). Synonym: S. kotschyi Boiss., l.c.: 46 (1846), S. macrosiphon var. cabulica Benth. in D.C., Prodr. 12: 282 (1848); S. macrosiphon var. kotschyi (Boiss) Boiss. Fl. Or. 4: 615 (1879); S. cuspidatissima Pau, Trab. Mus. Nac. Cienc. Nat. Ser. Bot. Madrid 14: 33 (1918); S. albifrons Náb., Publ. Fac. Sci. Univ. Masaryk (Brno) 70:49, tab. 5 (1926); S. macrosiphon var. glandulosissima Bornm., Bot. Jahrb. 62: 238 (1934); S. macrosiphon var. brachycalycina Bornm., l.c.; S. nachiczevanica Pobed., Fl. URSS. 21: 657 (1954), Icon.: Tab. 479, 480, 586 in Flora Iranica.

Type: Persia australi inter Fasa et Schiraz, Aucher-Elloy 5197 (holotype G).

Description: Perennial herbs with a woody rootstock at the base. Stems erect, 15-45 (- 90) cm high, branched above, scarcely eglandular pilose to tomentose below, densely glandular pilose above. Leaves simple, ± thin-textured, elliptical to ovate-oblong, 4-16 (- 22) × 1.5-9.5 (-11) cm, eglandular tomentose; margins irregular, serrate to erose; petiole 2- 10 cm long. Inflorescence panicle, densely glandular pilose. Verticillasters 2-6- flowered, clearly distant, internodes 2-2.5 cm long. Bracts broadly ovate, 10-27 × 7-20 mm, acuminate, eglandular pilose; bracteoles absent. Pedicels 1-3 mm long, glandular, rigid, erecto-patent. Calyx tubular, 12-17 mm long, up to 22 mm long in fruit, 4-5 mm diameter, scarcely expanding in fruit, glandular hirsute with some sessile glands; teeth spiny; upper lip tridentate, less spiny in fruit. Corolla white, 20-35 mm long; tube straight below, slightly widening above, 13-25 mm long; upper lip falcate, non- squamulate. Stamens of type B, staminal connectives clearly longer than filaments. Filaments 2-3 m long, upper connectives 10-15 mm long; fertile anthers hairless, 2-3 mm long. Style glabrous 20-37 mm long, exserted from corolla lips and divided in 2 parts at apex.

Phenology: Flowering in May and June.

249

Distribution and habitat in Turkey: This species is recorded from one small population in Southeast Anatolia and grows at the edges of fields, at 759 m (Figure 140).

Specimens examined: C8 Diyarbakır: Mardin to Diyarbakır, 14 km before Çınar, 759 m, field edges, 37°39′16″N-40°28′21″E, 24.v.2007, A. Kahraman 1382.

General distribution and phytogeography: S. macrosiphon is a member of the Irano- Turanian element and present in Iran, Iraq, Afghanistan, Pakistan and Transcaucasus.

Taxonomic notes: S. macrosiphon is very closely related to S. spinosa, however is differentiate by the less indumentum, eglandular pilose to tomentose hairs stems below, narrower leaves and calyces, less indurate and less spiny fruiting calyces, and longer corolla tubes.

Figure 138. General appearance of S. macrosiphon in its natural habitat. 250

32. S. palaestina Benth., Lab. Gen. Sp. 718 (1835). (Figure 139). Synonym: S. lorentii Hochst. in Lorent, Wanderungen 333 (1845).

Type: [LEBANON] in Palaestinae arvis, N. Bove (holo. G!).

Description: Perennial herb. Stems many, erect, purplish, 20-60 cm high, branched above, hirsute with long flattened eglandular hairs below, densely glandular pilose (sometimes with long flattened eglandular hairs) above. Leaves simple, pinnatilobed below to lyrate, oblong to ovate, 4.5-15 (-20) x 1.5-7 (-8.5) cm, tomentose (sericeous on young growth), rugose; margins erose; petiole 2.5-15 cm long. Inflorescence paniculate, 10-50 cm long. Verticillasters (2-) 3-6-flowered, clearly distant. Bracts broadly ovate, acuminate, 15-25 x 10-20 mm, papillose-glandular with long hairs, often tinged pink or purple. Pedicels 2-5 mm long, erecto-patent. Calyx ± tubular, often pink or purple (rarely green), 12-16 mm long in flower, to 17-25 mm long in fruit, broadening, papillose-glandular with some longer hairs; teeth 2-4 mm long; upper lip equally tridentate, spinulose. Corolla lilac or whitish-lilac, 20-35 mm long, simple hairs on outside of upper lip; tube 10-20 mm long, straight, ventricose above, not squamulate; upper lip ± falcate. Stamens B, included within upper lip of corolla, staminal connectives longer than filaments, dolabriform. Filaments 2-3 mm long; upper connectives 13-16 mm long; fertile anthers 3-4 mm long. Style 25-45 mm long.

Phenology: Flowering in May and July.

Distribution and habitat in Turkey: This species is distributed mostly in East and Southeast Anatolia and very occasionally in the province of Kahramanmaraş in the Mediterranean region of Turkey where it grows on limestone and igneous rocky slopes, cliffs, in Quercus scrub, vineyards and fallow fields, at 300-1510 m (Figure 140).

Specimens examined: B6 Malatya: Kangal to Hekimhan, 39o02′132′′N-37o44′240′′E, 1460 m, 2.vi.2008, A. Kahraman 1528B; Kahramanmaraş: near Afşin,

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38o14′712′′N-36o53′335′′E, 1298 m, 5.vii.2007, A. Kahraman 1298. B7 Elazığ: Keban to Elazığ, 38o42′24′′N-38o58′47′′E, 1151 m, 7.vi.2006, A. Kahraman 1204; Malatya: Malatya to Adıyaman, 38o19′41′′N-38o12′33′′E, 903 m, 17.v.2006, A. Kahraman 1124. B8 Diyarbakır: Lice to Diyarbakır, 65 km to Diyarbakır, 38o18′675′′N-40o31′895′′E, 789 m, 4.vi.2008, A. Kahraman 1533A. C6 Gaziantep: 23 km Gaziantep to Nizip, 37o02′15′′N-37o36′06′′E, 733 m, 18.v.2006, A. Kahraman 1134; Şanlıurfa: near Birecik, 37o00′994′′N-37o53′259′′E, 447 m, 29.iv.2006, A. Kahraman 1063. C7 Şanlıurfa: Tektek Mountain, near İnce village, 37o04′54′′N-38o34′45′′E, 550-700 m, 18.v.2006, A. Kahraman 1137; Adıyaman: 18 km from Adıyaman to Gölbaşı, 37o02′15′′N-37o36′06′′E, 733 m, 18.v.2006, A. Kahraman 1134. C8 Diyarbakır: Mardin to Diyarbakır, near Çınar, 37o49′58′′N-40o15′06′′E, 619 m, 24.v.2007, A. Kahraman 1383A; Mardin: around Mardin station, 37o17′10′′N-40o44′06′′E, 666 m, 23.v.2007, A. Kahraman 1375A; Batman: Batman to Siirt, near Beşiri, 566 m, 5.vi.2008, A. Kahraman 1533B. C8/9 Siirt: Siirt to Pervari, 59 km to Pervari, 38o01′441′′N-42o05′399′′E, 933 m, 5.vi.2008, A. Kahraman 1535. C9 Siirt: Siirt to Güçlükonak, Ormanaradı village, 37o43′389′′N-41o48′117′′E, 5.vi.2008, A. Kahraman 1537B; Şırnak: 5 km from Şırnak to Cizre, 37o30′891′′N-42o25′613′′E, 1090 m, 5.vi.2008, A. Kahraman 1544.

General distribution and phytogeography: S. palaestina is a member of the Irano- Turanian element and present in Egypt, Palaestina, Iraq and Iran.

Taxonomic notes: S. palaestina is variable in the leaves shape and degree of lobbing, indumentums and bract size

252

Figure 139. General appearance of S. palaestina in its natural habitat.

(30) S. spinosa (31) S.macrosiphon (32) S. palaestina Figure 140. Distribution of S. spinosa, S. macrosiphon and S. palaestina in the study area.

253

33. S. eriophora Boiss. & Kotschy ex Boiss., Fl. Or. 4: 611 (1879). (Figure 141).

Type: [TURKEY C5 Adana/Niğde] ab Jool Baatch (Yol Bahçe) ad radices montis Allahdagh (Ala Da.) Ciliciae, 1525 m, 3.vi.1859, Kotschy 229 (holo. G! iso. B! BM! K! W!).

Description: Perennial herb. Stems several from a woody rootstock, 10-20 cm high in flower, to 40 cm high in fruit, erect, widely branched above, eglandular lanate-arachnoid. Leaves mostly basal, linear-oblong, (2-5) 3-6 (-6.5) x 0.6-1.3 cm, narrowing into a (10-) 15-25 mm long petiole, arachnoid, rugose, margins crenulate. Inflorescence 5-25 mm long, paniculate with stiffly erecto-patent branches, arachnoid-lanate, violet-blue. Verticillasters 2-6-flowered, approximating, slightly distant in fruit. Bracts usually almost equal, violet, broadly ovate, 9 - 1 2 x 6 - 8 mm, arachnoid-lanate. Pedicels 2-3 mm long, erect. Calyx tubular-campanulate, 8-11 mm long, to 12-13 mm long in fruit, violet, arachnoid-lanate; teeth long-subulate, ca. 2 mm long. Corolla violet-blue, 8-13 mm long; tube 4-8 mm long, straight, glabrous within; upper lip ± straight; lower lip unspotted. Stamens of type B, included within upper lip of corolla, staminal connectives longer than filaments, dolabriform. Filaments ca. 2 mm long; upper connectives ca. 3 mm long; fertile anthers 2-3 mm long. Style glabrous, 9-15 mm long, slightly exserted from corolla lips and divided in two parts at apex.

Phenology: Flowering in May and June.

Distribution and habitat in Turkey: This species is distributed between Gürün (Sivas) and Pınarbaşı (Kayseri) where it grows on volcanic rocky and limestone slopes, at 1525-1900 m (Figure 143).

Specimens examined: B6 Kayseri/Sivas: near Güneşli village, 38o52′054′′N- 36o51′744′′E, 1878 m, 17.v.2008, A. Kahraman 1495; Kayseri/Sivas: Pınarbaşı to Gürün, 45-50 km to Gürün, 1870 m, 18.v.2007, A.Kahraman 1363.

254

General distribution and phytogeography: S. eriophora is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. eriophora is related to S. brachyantha, but differs by the smaller arachnoid crenulate leaves, smaller stems and geographic distribution.

Figure 141. General appearance of S. eriophora in its natural habitat.

34. S. brachyantha (Bordz.) Pobed. in Fl. URSS 21: 660 (1954). (Figure 142). Synonym: S. modesta Boiss., Fl. Or. 4: 621 (1879) p.p.; S. modesta Boiss. var. brachyantha Bordz. in Mem. Soc. Nat. Kiev 25: 113 (1915). Ic: Fl. URSS 21: t. 15 f. 3 (1954).

Type: [USSR, SOVIET ARMENIA] provincia Eriwán: pagus Mastara inter montes Alagäs (Alagöz) et Bugutlu, vi.1875, G. Radde 195 (holo. LE, photo). 255

Description: Perennial herb. Stems 30-50 cm high from a woody rootstock, erect, sparsely arachnoid-pilose above and below, often purplish-blue or violet above. Leaves simple, ovate to linear-oblong, 5-12 (-20) x 0.8-4 (-11) cm, rugulose, arachnoid-pilose; margins erose; petiole 2-8 cm long. Inflorescence paniculate. Verticillasters 2-8-flowered, distant. Bracts broadly ovate, acuminate, ca. 8-12 (-18) x 8-12 (-20) mm. Pedicels 2-3 mm long, erecto-patent. Calyx tubular-campanulate, ca. 8-10 mm long (including mucros), to ca. 12-13 mm long in fruit, eglandular (rarely with small glandular papillae), sublanate, often violet; upper lip tridentate, median shorter, subulate. Corolla lilac to violet-blue, ca. 12 mm long; tube somewhat ventricose, subsquamulate; upper lip scarcely falcate. Stamens of type B, included within upper lip of corolla.

Phenology: Flofering in May and July.

Distribution and habitat in Turkey: This species is mainly distributed in East Anatolia and also is known from one population (Karacadağ-Şanlıurfa) in Southeast Anatolia where it grows on rocky igneous slopes, in steppe, fallow fields and meadows with often Salvia staminea and Nepeta betonicifolia, at 1050-2800 m (Figure 143).

Specimens examined: A9 Kars: Kars, 30.vi.1912, Lonaczewski-Pietruniaka. B8 Erzurum: Tercan to Aşkale, 1850 m, 8.vi.1957, D. 29321 (ANK! E!). B9 Van: Van to Başkale, Güzeldere, 2791 m, 11.vii.2007, A. Kahraman 1448; ibid., 2800 m, 7.vi.2008, A. Kahraman 1572; Van: Çuh pass, Halanduran Mountain and Güzeldere hill, 2800 m, Watson et al. 1485; Ağrı: 2 km Southwest of Hamur, Murat valley, 1680 m, 3.vi.1966, D. 44084; Bitlis: North of Ahlat, Yuvadamı village, 2300 m, 22.vi.1995, L. Behçet 904 (VANF!). C7 Urfa: Siverek to Diyarbakır, West foot of Karacadağ, 17 miles from Siverek, 1050 m, 19.v.1957, D. 28278 (ANK!).

General distribution and phytogeography: S. brachyantha is a member of the Irano- Turanian element and present in the northwest of Iran and Armenia. 256

Taxonomic notes: S. brachyantha resembles S. eriophora and S. modesta, however, differs from the former in the larger and arachnoid-pilose erose leaves, larger stems and from the latter in the entirely lilac to violet-blue corollas and mostly arachnoid eglandular hairs on inflorescence axes and calyces (sparse in fruit).

Figure 142. General appearance of S. brachyantha in its natural habitat.

257

(33) S. eriophora (34) S. brachyantha Figure 143. Distribution of S. eriophora and S. brachyantha in the study area.

35. S. sclarea L., Sp. Pl. 27 (1753). (Figure 144). Synonym: S. pamirica Gand., Bull. Soc. Bot. Fr. 60: 26 (1913); S. altilabrosa Pau, Trab. Mus. Nac. Scienc. Nat. Ser. Bot. Madrid 14: 33 (1918). Ic: Reichb., Ic. Fl. Germ. 18: t. 1249 (1856); Fl. RPR 8: t. 41 f. 2 (1961); Huxley & Taylor, Fls. Greece t. 239 (1977).

Type: SYRIA and ITALY (Hb. Linn. 42/45).

Description: Biennial or short-lived perennial aromatic herb. Stems to 130 cm high, erect, rather coarse quadrangular, much branched above, pubescent to hirsute below, glandular above. Leaves simple, broadly ovate to ovate-oblong, 6-15 x 5-10 cm, cordate at base, pubescent; margins crenate-erose; petiole 3-9 cm long. Inflorescence paniculate, many-flowered; Verticillasters 2-6-flowered, distant. Bracts ± obscuring flowers, pink to mauve, membranous, ovate, acuminate, 10-35-x 7-25 mm. Pedicels 2-3 mm long, erecto-patent. Calyx ovate-campanulate, ca. 10 mm long, to ca. 15 mm long in fruit, scabrid, glandular punctuate and sparsely white pubescent with sessile glands; upper lip tridentate, mucronate. Corolla with a lilac, purplish pink or pale blue

258

upper lip and cream lower lip 20-30 mm long; tube abruptly ventricose, squamulate; upper lip falcate. Stamens of type B, included within upper lip of corolla.

Phenology: Flowering from May to August.

Distribution and habitat in Turkey: This species is mainly distributed in all region of Turkey where it occurs on rocky igneous slopes, roadsides, shale banks, fields and in mixed deciduous and coniferous woodland, at 1-2000 m (Figure 146).

Specimens examined: A8 Erzurum: Erzurum to Tortum, 40o37′53′′N-41o36′29′′E, 1520 m, 11.vii.2006, A. Kahraman 1299. B6 Kayseri: Sarız to Pınarbaşı, ca. 2-3 km to Sarız, 38o27′51′′N-36o28′24′′E, 1577 m, 20.vii.2006, A. Kahraman 1356. B7 Erzincan: Kemaliye, 39o14′35′′N-38o29′36′′E, 950-1100 m, 6.vi.2006, A. Kahraman 1194; Malatya: Beydağı, Gündüzbey to Kozluk, 38o13′34′′N-38o16′45′′E, 1295 m, 7.vi.2006, A. Kahraman 1214; Tunceli: Tunceli to Pülümür, 1100 m, D. 29227. B8 Erzincan: Erzincan to Erzurum, 39o50′28′′N-40o34′22′′E, 1877 m, 10.vii.2006, A. Kahraman 1288. B9 Van: Soğuksu to Bahçesaray, 11 km to Bahçesaray, 38o06′254′′N- 42o40′580′′E, 1900 m, 9.vii.2007, A. Kahraman 1433. C6 Gaziantep: Gaziantep to Şanlıurfa, 800 m, 7.v.2001, M. Vural 8494 et al. (GAZI!). C7 Şanlıurfa: Ceylanpınar, 510 m, 2.v.1995, N. Adıgüzel 2236 et al. (GAZI!). C9 Siirt: Cevizli to Siirt, 1000 m, 3.vi.1980, M. Koyuncu 3175 & A. Güner (AEF!); Şırnak: 17 km from Şırnak to Hakkari, 1150 m, 5.vi.2001, M. Ekici 2251 (CBB! GAZI!); Hakkari: Zap gorge, 25 km from Hakkari to Çukurca ca. 25. km, 1000 m, D. 44800. C10 Hakkari: 2 km from Şemdinli to Yüksekova, 1550 m, D. 45033.

General distribution and phytogeography: S. sclarea is multiregional and is found throughout most of Europe, Southwest and Central Asia.

Taxonomic notes: S. sclarea is a fairly oligomorphic species and distinctive on account of the habit, large coloured bracts and often exceeding the flowers.

259

Figure 144. General appearance of S. sclarea in its natural habitat.

36. S. aethiopis L., Sp. PI. 27 (1753). Ic: Jáv. & Csap., Ic. Fl. Hung. 432, t. 2931 (1934); F1. RPR 8: t. 42 f. 1 (1961). (Figure 145).

Type: ILLYRIA (COASTAL JUGOSLAVIA), GREECE & AFRICA (Hb. Linn. 42/48).

Description: Perennial or biennial herb. Stems erect, sturdy, 23-130 cm high, eglandular lanate, hairs denser below, with sessile glands. Leaves simple, mostly basal, ovate-elliptic to oblong, 7-25(-32) x 4-15 cm, ± lanate, rugose, acute to obtuse, cauline leaves sessile; margins erose; petiole 4-15.5 cm long. Inflorescence widely branched, candelabriform, many-flowered. Verticillasters 4-10-flowered, approximating above. Bracts broadly obovate, 10-15 x 12-18 mm, cuspidate, sparsely pilose. Calyx tubular-ovate, 10-15 mm long, to ca. 17 mm in fruit, densely 260

lanate; upper lip tridentate, median tooth shorter. Corolla white, 12-20 mm long, often with a pale yellow lip; tube abruptly ventricose, squamulate; upper lip falcate (sometimes subfalcate). Stamens of type B, included within upper lip of corolla.

Phenology: Flowering from May to August.

Distribution and habitat in Turkey: This species is mainly distributed in all region of Turkey except for Southeast Anatolia where it occurs on steppe, igneous and limestone slopes, fallow fields, roadside banks, at 1-2100 m (Figure 146).

Specimens examined: A8 Erzurum: 30 km from Erzurum to Tortum, 40o11′07′′N- 41o27′36′′E, 1239 m, 11.vii.2006, A Kahraman 1294A. A9 Erzurum: 10 km from Horasan to Kars, 40o04′58′′N-42o18′30′′E, 1535 m, 12.vii.2006, A. Kahraman 1304A; Kars: near Selim, 40o17′40′′N-42o39′21′′E, 2012 m, 12.vii.2006, A. Kahraman 1309. B6 Malatya: Malatya to Darende, Yukarı Ulupınar village, 38o23′34′′N-37o34′58′′E, 1408 m, 7.vi.2006, A. Kahraman 1222; Sivas: Kangal to Divriği, 1420-1500 m, 6.vi.2006, A. Kahraman 1142. B7 Elazığ: Elazığ to Baskil, above Kayabeli village, 38o35′35′′N- 38o52′26′′E, 1600 m, 7.vi.2006, A. Kahraman 1209A. B8 Erzurum: Ilıca to Erzurum, 30 km to Erzurum, 39o55′00′′N-41o12′42′′E, 1817 m, 10.vii.2006, A. Kahraman 1292; Erzincan: Erzurum to Erzincan, 500 m to Tercan, 39o46′211′′N-40o24′437′′E, 1447 m, 14.vii.2007, A. Kahraman 1477B. B9 Ağrı: Ağrı to Horasan, near Tahir village, 39o50′12′′N-42o24′34′′E, 2110 m, 15.vii.2006, A. Kahraman 1323; Van: 3 km from Van to Özalp, 1896 m, 16.vii.2006, A. Kahraman 1330. B10 Ağrı: Doğubeyazıt, near Telçeken village, 1600 m, 24.vi.1972, T. Gözler & T. Kesercioğlu (EGE 19701!).

General distribution and phytogeography: S. aethiopis is multiregional and is found in Central and South Europe, Caucasiaca, Crime, Iran, and is introduced North America.

Taxonomic notes: S. aethiopis is a distinct species based on the sturdy stems, lanate indumentum and candelabriform inflorescence. 261

Figure 145. General appearance of S. aethiopis in its natural habitat.

(35) S. sclarea (36) S. aethiopis Figure 146. Distribution of S. sclarea and S. aethiopis in the study area. 262

37. S. ceratophylla L., Sp. Pl. 27 (1753). (Figure 147).

Type: PERSIA (Hb. Linn. 42/47, B!).

Description: Biennial or perennial herb, lemon-scented. Stems sturdy, 25-70 cm high, glandular villous, densely above. Leaves mostly basal, deeply pinnatifid, oblong in outline, with spreading linear segments, white lanate when young, terminal segments 10-30 x 3-10 cm, often several times as long as lateral segments, rachis winged; petiole subabsent. Inflorescence pyramidal and spreading paniculate, yellowish-green. Verticillasters 2-8-flowered, distant. Bracts ovate, cuspidate, ca. 15 x 12 mm Pedicels erecto-patent, 2-4 mm long. Calyx ovate- campanulate, 10-12 mm long, to ca. 18 mm long in fruit with diverging lips, glandular villous; upper lip tridentate with cuspidate teeth. Corolla cream or sulphur, rarely with lilac upper lip, 15-20 mm long; tube ca. 4-5 mm long, abruptly ventricose, squamulate; upper lip falcate. Stamens of type B, included within upper lip of corolla.

Phenology: Flowering from April to July.

Distribution and habitat in Turkey: This species is mainly distributed in all region of Turkey except for the Aegean geographic region where it grows on limestone, igneous and gypsum slopes, waste ground, corn and fallow fields, at 300-2300 m (Figure 149).

Specimens examined: A9 Kars: 23 km from Kağızman to Karakurt, 1350 m, Rechinger 57430. B6 Malatya: Darende to Malatya, 38o22′116′′N-37o57′105′′E, 1260 m, 17.v.2008, A. Kahraman 1500A & B; Sivas: Kangal to Divriği, 38o16′376′′N- 38o01′536′′E, 1169 m, 2.vi.2008, A. Kahraman 1520. B7 Adıyaman: Gerger to Kaşyazı village, 3 km to Kaşyazı, 38o02′04′′N-39o03′20′′E, 876 m, 19.v.2007, A. Kahraman 1370; Erzincan: Divriği to Kemaliye, 2 km to Gümüşçeşme village, 39o19′27′′N- 38o23′23′′E, 963 m, 6.vi.2006, A. Kahraman 1193; Sivas: Divriği to Kemaliye, ca. 5-10

263

km Gedikbaşı, 39o28′53′′N-38o14′55′′E, 1273 m, 6.vi.2006, A. Kahraman 1186. B8 Erzincan: Tercan to Aşkale, 1600 m, D. 29343. B9 Van: 5 km from Hoşap to Başkale, 2250 m, 17.vii.2006, A. Kahraman 1361. B10 Ağrı: Karabulak to Doğubayazıt, 1554 m, 14.vii.2006, A. Kahraman 1319C. C6 Gaziantep: 5 km from Nizip to Gaziantep, 36o58′761′′N-37o42′895′′E, 572 m, 29.iv.2006, A. Kahraman 1075. C7 Şanlıurfa: Şanlıurfa to Hilvan, 700 m, D. 28241. C8 Mardin: Mardin to Gercüş, 37o26′16′′N-41o21′59′′E, 955 m, 23.v.2007, A. Kahraman 1378; Siirt: 5 km South of Siirt, 700 m, D. 43214. C10 Hakkari: Zap river valley, Trelawny 1319.

General distribution and phytogeography: S. ceratophylla is a member of the Irano- Turanian element and is present in Palestine, Syria, Afghanistan, Transcaucasia, Iraq, Iran, Talish and Turcomania.

Taxonomic notes: S. ceratopylla is a very distinct and oligomorphic species on account of the deeply pinnatified leaves, white lanate when young and yellow or cream corollas.

Figure 147. General appearance of S. ceratophylla in its natural habitat. 264

38. S. longipedicellata Hedge in Notes R.B.G. Edinb. 23: 57 (1959). (Figure 148).

Type: TURKEY B8 Erzurum: between Ilica and Tercan, near the turning to Aşkale, 1850 m, 10.vii.1957, Davis & Hedge, D. 30875 (holo. E! iso. ANK! BM‟ K!)..

Description: Perennial herb. Stems erect, solitary or 2-3, 60-90 cm high, sturdy, lanate below, densely glandulai-pilose above with capitate glandular hairs. Leaves mostly basal, ovale-lanccolale lo elliptic, 6.5-20 x 2.5-7.5 cm, lanate to pilose below, with a thinner indumentum above; margins irregularly lobulate serrate, truncate or attenuate; petiole 3-12 cm long, with lanate hairs. Inflorescence paniculate, densely glandular pilose. Verticillasters 4-6-flowered, clearly distant. Bracts clearly shorter than calyces, broadly ovate-acuminate, 7-18 x 6-15 mm. Pedicels (5-) 6-12 mm long. Calyx tubular-campanulate, 9-11 mm long, to 11-14 mm long in fruit, densely glandular pilose with numerous sessile glands. Corolla white, 15-20 mm long; tube ventricose, squamulate, 8-9 mm long; upper lip falcate. Stamens of type B, included within upper lip of corolla, upper connectives clearly longer than filaments. Filaments 2- 4 mm long; upper connectives 10-20 mm long; fertile anthers 2-5 mm long Style 20-45 mm long, slightly exserted from corolla lips and divided in two parts at apex.

Phenology: Flowering in July and August.

Distribution and habitat in Turkey: This species is distributed in East, Central and North Anatolia, however populations in Central and North Anatolia are rather smaller than those in the East Anatolia (Figure 149). It occurs on disturbed steppe, meadows and fallow fields, at 1480-2110 m.

Specimens examined: B7 Erzincan: 2 km from Refahiye to Erzincan, 1540 m, 9.vii.1979, B. Çubukçu, E. Tuzlacı & M. Saraçoğlu 42564 (ISTE!); B8 Erzurum: near Erzurum, 40o04′487′′N-42o07′471′′E, 1779 m, 29.7.2008, A. Kahraman 1608; near

265

Aşkale, 39o58′729′′N-40o33′729′′E, 1742 m, 29.vii.2008, A. Kahraman 1609. B9 Ağrı: 60 km from Ağrı to Horasan, 2081 m, 12.vii.2007, A. Kahraman 1459.

General distribution and phytogeography: S. longipedicallata is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. longipedicellata is distantly related to S. chionantha, but can be easily distinguished by the broader leaves, longer pedicels and smaller calyces and corollas.

Figure 148. General appearance of S. longipedicellata in its natural habitat.

266

(37) S. ceratophylla (38) S. longipedicellata Figure 149. Distribution of S. ceratophylla and S. longipedicellata in the study area.

39. S. atropatana Bunge, Lab. Pers. 47 (1873). (Figure 150). Synonym: S. montbretii Benth. var. virescens Freyn in Bull. Herb. Boiss. ser. 2, 1: 278 (1901); S. linguifolia Hedge & Hub.-Mor. in Notes R.B.G. Edinb. 22: 181 (1957).

Type: [IRAN] prope Tikmedescht ad orientem ab urbe Tabris, Bienert (holo. LE? fragment P).

Description: Perennial herb with a woody rootstock. Stems several, 25-70 cm high, erect, usually densely glandular pilose or eglandular arachneid with sessile glands. Leaves mostly basal, linear to linear-oblong, (5-) 10-22 x 0.5-4 (-6.5) cm, arachnoid- tomentose with sessile glands to ± lanate; margins entire (on linear leaves), serrate to suberose, cuneate; petiole (1.5-) 3-11 cm long. Inflorescence paniculate, narrowly obpyramidal. Verticillasters (2-) 4-6-flowered, approximating above. Bracts ovate- acuminate, 8-25 x 7-15 mm. Pedicels 2-4 mm long. Calyx ± campanulate, 8-10 mm long, to 10-15 mm long in fruit, densely glandular; upper lip indistinctly tridentate. Corolla white, or with a yellow lip, 18-25 mm long; tube 7-9 mm long, ventricose,

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squamulate; upper lip strongly falcate. Stamens of type B, included within upper lip of corolla, dolabriform.

Phenology: Flowering in June and July.

Distribution and habitat in Turkey: This species is found only in East Anatolia where it grows on rocky slopes in steppe, sloping meadow, fallow fields and roadsides, at 1245-2560 m (Figure 153).

Specimens examined: B9 Van: Gürpınar to Van, 38o24′459′′N-43o23′321′′E, 2110 m, 15.vii.2006, A. Kahraman 1452; Bitlis: Hizan to Sağıllı, north of Armut hill, E. Tuzlacı 47330 (ISTE!); Bitlis: Hizan, above Karbastı village, 1600 m, 26.vi.1990, L. Behçet 2189 (VANF!); Ağrı: Ağrı and Doğubayazıt, 17 km to Doğubayazıt, 1780 m, 12.vi.1979, E. Tuzlacı 42773 (ISTE!). B10 Van: 3-4 km Northeast of Başkale, 2300 m, D. 44544. C9 Şırnak: Şırnak and Hakkari, Yemişli village, 1245 m, 6.vi.2008, A. Kahraman 1548. C10 Hakkari: Hakkari and Van arası, 1839 m, 7.vi.2008, A. Kahraman 1570.

General distribution and phytogeography: S. atropatana a member of the Irano- Turanian element and present in Iraq, Iran and Turcomania.

Taxonomic notes: S. atropatana is similar to S. chionantha and S. microstegia but is differentiated from the latter by the linear to linear-oblong leaves, shorter pedicels and smaller calyces and corollas and from by the former in the much narrower leaves with a thin indumentum.

268

Figure 150. General appearance of S. atropatana in its natural habitat.

40. S. microstegia Boiss. & Bal. in Boiss., Diagn. ser. 2(4): 17 (1859). (Figure 151). Synonym: S. verbascifolia Bieb. var. cana Boiss., Fl. Or. 4: 619 (1879); S. chnoodes Stapf in Denkschr. Akad. Wiss. Wien, Math.-Nat. Kl. 50: 98 (1885). Ic: J. Roy. Hort. Soc. 85(10): t. 140 (1960).

Lectotype: [TURKEY B5 Kayseri] in monte Alidagh prope Caesaream (Kayseri) Cappadociae sito, 1400m, 9.viii.1856, Balansa 1083 (holo. G! iso. W).

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Description: Perennial herb with a woody rootstock, Stems few or several, erect, 20- 125 cm high, densely pilose to villous glandular, often eglandular lanate below. Leaves mostly basal, variable in size and shape, ovate to oblong, (3.5-) 6-18 (-20) x 3-9 (-14) cm, white or grey lanate, subcordate; margins obtusely lobed or irregularly serrate; petiole 3-17 (-20) cm long. Inflorescence usually a widely spreading panicle, often yellowish-green. Verticillasters 4-6-flowered, usually distant. Bracts variable, broadly ovate, 9-17 x 8-14 mm. Pedicels 2-5 mm long. Calyx ± campanulate, 9-13 mm long, to 11-14 mm long in fruit, densely capitate glandular with sessile glands; upper lip shortly tridentate, median tooth much shorter; lips ± equal. Corolla white, lower lip fading yellow, 17-30 mm long, tube 5-11 mm long, ventricose, squamulate; upper lip compressed, strongly falcate. Stamens of type B, included within upper lip of corolla.

Phenology: Flowering from June to August.

Distribution and habitat in Turkey: This species is distributed in all regions of Turkey except for the Aegenan and Marmara geographic regions where it grows on rocky limestone and igneous slopes, screes, fieldsides, in Quercetum and Pinus woodland and near permanent snowbeds, at 970-3350 m (Figure 153).

Specimens examined: A9 Kars: Kağızman, Turkewicz 627. B6 Kayseri: Sarız to Pınarbaşı, 38o30′01′′N-36o26′53′′E, 1692 m, 5.vii.2007, A. Kahraman 1399; Sivas: Pınarbaşı to Gürün, near Güneşli village, 38o50′417′′N-36o53′296′′E, 1868 m, 17.v.2008, A. Kahraman 1493. B7 Erzincan: around Sipikör pass, 2410 m, 10.vii.2006, A. Kahraman 1280. B8 Erzurum: 14 km from Erzurum to Pasinler, 2000 m, D. 47452. B9 Bitlis: Kambos Mountain, above Hürmüz, 38o19′408′′N-41o58′079′′E, 2280 m, 9.vi.2008, A. Kahraman 1423; Van: Northwest of Artos Mountain, 38o18′882′′N- 42o06′737′′E, 1662 m, 10.vii.2008, A. Kahraman 1447; Ağrı: Ağrı to Horasan, near Tahir, 2100 m, 15.vii.2006, A. Kahraman 1321A. C7 Adıyaman: Nemrut Mountain, 2000-2100 m, 20.vi.1987, N. Tanker & M. Koyuncu 8167 (AEF!). C9 Hakkari: Hakkari

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to Kaval (Piyannis), 2500 m, D. 45395; Berçelan, 3000 m, 21.vii.1974, M. Koyuncu (AEF!). C10 Hakkari: Sat Mountain, 2950 m, D. 45702.

General distribution and phytogeography: S. microstegia a member of the Irano- Turanian element and present in Syria and Lebanon.

Taxonomic notes: S. microstegia is closely related to S. argentea, but differs in the stems with lanate hairs at base, obtusely lobed to serrate margins

Figure 151. General appearance of S. microstegia in its natural habitat.

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41. S. xanthocheila Boiss. ex Benth. in DC., Prodr. 12: 284 (1848). (Figure 152).

Type: [IRAN] in monte Ellburz Persiae borealis, Kotschy 128 (iso. K!).

Description: Perennial herb with a thick woody rootstock. Stems 15-45 cm high, sturdy, erect densely glandular pilose. Leaves mostly basal, broadly ovate to ovate-oblong, 2.3- 9 (-11) x 1.5-6 cm, lanate to pannose; margins irregularly lobed or dentate to subentire, ± cordate; petiole 1-7 cm long. Inflorescence 10-35 cm long, a short sturdy panicle. Verticillasters 4-6-flowered, crowded. Bracts broadly ovate, acuminate, 12-21 x 10-19 mm. Pedicels 2-4 mm long. Calyx campanulate-infundibular, 9-13 mm long, to 13-19 mm long in fruit with widely diverging lips, densely glandular; upper lip shortly tridentate. Corolla white, 17-27 mm long; tube 8-10 mm long, ventricose, squamulate; upper lip clearly falcate. Stamens of type B, included within upper lip of corolla. Filaments 2-4 mm long; upper connectives 10-20 mm long; fertile anthers yellow, 2-4 mm long. Style glabrous, 30-40 mm long, slightly exserted from corolla lips and divided in two parts at apex.

Phenology: Flowering from May to July.

Distribution and habitat in Turkey: This species is distributed only in East Anatolia where it grows on rocky igneous slopes and steppe, at 2000-2870 m (Figure 153).

Specimens examined: A9 Erzurum: Horasan to Karaurgan, 2000 m, D. 29498; Kars: Kağızman, Turkevicz 114. B8 Erzurum: around Palandöken, 39o51′077′′N-41o17′020′′E, 2417 m, 14.vii.2007, A. Kahraman 1475. B9 Van: Van to Başkale, 38o10′55′′N- 43o54′39′′E, 2870 m, 17.vii.2006, A. Kahraman 1340; Ağrı: Aşağı Mountain, Northwest of Cumaçay, 2000 m, Watson 5641. C9 Hakkari: Berçelan, 2661 m, 7.vi.2008, A. Kahraman 1559.

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General distribution and phytogeography: S. xanthocheila is a member of the Irano- Turanian element and present in Talish and Iran.

Taxonomic notes: S. xanthocheila is related to S. microstegia, but differs in the larger bracts and fruiting calyces and and and shorter stems.

Figure 152. General appearance of S. xanthocheila in its natural habitat.

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39) S. atropatana (40) S. microstegia (41) S. xanthocheila Figure 153. Distribution of S. atropatana, S. microstegia and S. xanthocheila in the study area.

42. S. frigida Boiss., Diagn. ser. 1(5): 10 (1844). (Figure 154). Synonym: S. spinulosa Montbret & Aucher ex Benth. in DC., Prodr. 12: 283 (1848); S. frigida Boiss. var. oblongifolia Boiss., Fl. Or. 4: 621 (1879); S. frigida Boiss. var. albiflora Bornm. in Beih. Bot. Centr. 24(2): 487 (1909).

Type: [TURKEY C2 Denizli] in regione alpina excelsiori montium Cariae, Cadmi orientalis (Honaz Mountain) supra Colossam (Honaz) et occidentalis (Baba Mountain) supra Gheyra (Göyre), vi.1842, Boissier (holo. G!).

Description: Perennial herb with a thick woody rootstock. Stems solitary or several, erect, 10-30 (-50) cm high, below pilose to villous with sessile glands, densely glandular above with capitate glands. Leaves mostly basal, variable, ovate to narrowly oblong, 2-15 x 1.2-5 cm, rugulose, arachnoid to ± lanate with many sessile glands; margins crenulate to erose; petiole 1.5-8 cm long. Inflorescence little branched to widely paniculate. Verticillasters 2-6-flowered, usually distant. Bracts ovate to orbicular, (6-) 8-12 x (6-) 7-11 mm. Pedicels 2-3 mm long. Calyx campanulate to infundibular, 7-10 mm long, to 10-12 mm long in fruit and widening, densely capitate-glandular; teeth usually

274

prominently spinulose; upper lip tridentate, median tooth much shorter, truncate or not. Corolla white to lilac, 12-16 mm long; tube ca. 8 mm long, ventricose, squamulate; upper lip narrow, somewhat falcate, scarcely bifid. Stamens of type B, included within upper lip of corolla.

Phenology: Flowering from May to July.

Distribution and habitat in Turkey: This species is distributed in East, Central Anatolian, Mediterranean and Aegean regions of Turkey where it grows in Pinus nigra, Juniperus, Abies and Pyrus woodland, on limestone slopes, crevices and meadows, at 1000-2500 m (Figure 157).

Specimens examined: B6 Sivas: Pınarbaşı to Gürün, ca. 42 km to Gürün, around Güneşlli village, 38o50′09′′N-36o53′22′′E, 1880 m, 16.v.2006, A. Kahraman 1092; Kayseri: ca. 3-4 km from Sarız to Pınarbaşı, 1690 m, 8.vi.2006, A. Kahraman 1235B. B7 Sivas: Karasar to Kayaburun, 1650 m, 6.vi.2006, A. Kahraman 1450; Malatya: 5 km East of Malatya, 1300 m, Hub.-Mor. 8988. B8 Erzincan: 17 km from Tercan to Aşkale, 1680 m, Hub.-Mor. 10768. B9 Bitlis: Bitlis to Tatvan, 1900 m, D. 22354.

General distribution and phytogeography: S. frigida is a member of the Irano-Turanian element and is found in Iran.

Taxonomic notes: S. frigida is closely related to S. yosgadensis and S. modesta, but is differentiated from the former by the glandular pilose to villous stems above and from the latter by the smaller corollas and larger calyces.

275

Figure 154. General appearance of S. frigida in its natural habitat.

43. S. poculata Náb. in Publ. Fac. Sci. Univ. Masaryk Brno 70: 50, f. 14 (1926). (Figure 155). Synonym: S. brevidens Hedge & Hub.-Mor. in Notes R.B.G. Edinb. 22: 183 (1957).

Lectotype: [IRAQ] distr. Serizor, Handrian Dar (Handren Dagh) supra Rewanduz dit. Erbil, in humosis, c. 1700 m, 23.v.1910, Nábělek 1570 (BRA).

Description: Perennial herb. Stems erect, 25-50 cm high, branched above, glandular- pilose below, densely glandular pilose to villous above. Leaves simple, ovate to oblong, 6-15 x 3-7 cm, arachnoid floccose (± lanate on young growth); margins crenate to erose; petiole 2.5-9 cm long. Inflorescence paniculate. Verticillasters 2-8-flowered, clearly distant. Bracts broadly ovate to ± semicircular, abruptly acuminate, 17-22 x 20- 25 mm. Pedicels 1-3.5 mm long, erecto-patent. Calyx obtriangular, 8-10 mm long, to 12- 276

14 mm long in fruit and broadening, sometimes tinged purplish, glandular villous; upper lip tridentate, median tooth clearly shorter than laterals. Corolla white or pale lilac, 12- 16 mm long; tube ± straight, ca. 6 mm long, squamulate, ventricose above; upper lip scarcely falcate. Stamens of type B, included within upper lip of corolla.

Phenology: Flowering from May to July.

Distribution and habitat in Turkey: This species is distributed only East Anatolia where it occurs on rocky limestone and igneous slopes, sloping meadows and near melting snow, at 1400-2900 m (Figure 157).

Specimens examined: A8 Erzurum: Erzurum to Tortum, 2030 m, Hub.-Mor. 15307. B7 Erzincan: Keşiş Mountain, above Cimin, 2500-2600 m, 27.vii.1957, Davis & Hedge 31670 (E! ANK!). B8 Erzurum: 12 km North of Hınıs, 1950 m, Zohary 571622, Siirt: Sason Mountain, Halkis Mountain above Sason, 1700 m, Watson et al. 1217. B9 Van: Erek Mountain, 38o29′43′′N-43o29′09′′E, 2052 m, 16.vii.2006, A. Kahraman 1325B; Van: Soğuksu to Bahçesaray, 20 km to Bahçesaray, 2218 m, 9.vii.2007, A. Kahraman1431; Bitlis: Kambos Mountain, Direktaşı (Tito), 38o19′408′′N-41o58′079′′E, 2280 m, 9.vi.2008, A. Kahraman 1585B; Ağrı: Tutak to Hamur, 39o36′392′′N- 42o57′767′′E, 1610 m, 12.vii.2007, A. Kahraman1457. C9 Hakkari: Berçelan, 37o38′129′′N-43o45′064′′E, 2661 m, 7.vi.2008, A. Kahraman 1558; Siirt: Eruh, Yassıdağ (Serikur Mountain), above Meşindağı pass (Birini pass), 1800 m, 18.vii.1981, E. Tuzlacı 47348 (ISTE!). C10 Hakkari: Şat Mountain, 2850 m, D. 45498.

General distribution and phytogeography: S. poculata is a member of the Irano- Turanian element and is found in Iran and Iraq.

Taxonomic notes: S. poculata is rather closely related to odontochlamys, but differs in the obtriangular and green calyces.

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Figure 155. General appearance of S. poculata in its natural habitat.

44. S. odontochalmys Hedge in Notes R.B.G. Edinb. 22: 184 (1957). (Figure 156).

Type: [TURKEY B9] Bitlis: Kambos Dağ above Hürmüz, 2100 m, fls. white tinged lilac, 30.vi.1954, Davis & O. Polunin, D. 23385 (holo. K!).

Description: Perennial herb. Stems erect, ca. 30 cm high, branched above, glandular- pilose below, densely glandular pilose to villous above. Leaves simple, oblong-lanceolate, 5-13 x 2.5-5 cm, arachnoid floccose; margins crenate to erose; petiole 2.5-8.5 cm long. Inflorescence paniculate. Verticillasters 3-7-flowered, clearly distant. Bracts broadly ovate to ± semicircular, densely white-canascent, gradually acuminate; upper to 2 x 1.5

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cm, lower to ca. 4.5 x 1.3 cm. Pedicels 1-3.5 mm long, erecto-patent. Calyx tubular- infundibular, short bilabiate, 9-11 mm long, up to ca. 13 mm long in fruit and broadening, sometimes tinged purplish, glandular villous; upper lip tridentate, median tooth clearly shorter than laterals. Corolla white or lilac, ca. 17 mm long; tube ± straight, ca. 7-8 mm long, squamulate, ventricose above; upper lip scarcely falcate. Stamens of type B, included within upper lip of corolla.

Phenology: Flowering in June and July.

Distribution and habitat in Turkey: This species has a rather local distribution known only from the type location (Figure 157). It grows on sloping meadows and near melting snow, at 2100-2280 m.

Specimens examined: B9 Bitlis: Kambos Mountain, above Hürmüz, 38o19′408′′N- 41o58′079′′E, 2280 m, 9.vi.2008, A.Kahraman 1579H.

General distribution and phytogeography: S. odontochlamys is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. odontochlamys is very close to odontochlamys which grows on the same mountain at a similar altitude, but differs in the tubular-infundibular calyces and densely white-canascent bracts.

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Figure 156. General appearance of S. odontochlamys in its natural habitat.

(42) S. frigida (43) S. poculata (44) S. odontochlamys Figure 157. Distribution of S. frigida, S. poculata and S. odontochlamys in the study area.

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45. S. candidissima Vahl. subsp. candidissima (Figure 158). Synonym : S. candidissima Vahl. var. cordifolia Náb. in Publ. Fac. Sci. Univ. Masaryk Brno 70: 52, t. 4 f. 3 (1926). Ic: Jacq., Eclog. Pl. Rar. t. 16 (1811), as S. odorata Willd.

Description: Perennial herb. Stems 30-90 cm high, erect, branched above, pilose to lanate below with few sessile glands, densely pilose to glandular-papillose above. Leaves simple, oblong to broadly ovate, 5-10 (-14) x 3-10 cm, pubescent to densely pannose, cordate to rounded; margins serrulate to erose; petiole 3-11 cm long. Inflorescence paniculate, often yellowish-green; Verticillasters 2-6-flowered, distant. Bracts ovate- acuminate, 4-10 x 3-6 mm. Pedicels 2-4 mm long. Calyx tubular-campanulate, 12-15 mm, to 18 mm and widening in fruit, densely pilose to glandular-papillose; upper lip with 3 closely connivent mucronate teeth. Corolla white with a yellow lower lip, 20-30 mm long; tube ca. 12-13 mm long, ventricose, squamulate; upper lip strongly falcate. Stamens of type B, included within upper lip of corolla.

Phenology: Flowering from May to September.

Distribution and habitat in Turkey: This species is distributed in all regions of Turkey except for Marmara anad Aegean geographic regions where it occurs on rocky limestone and shale slopes, bushy places, fallow fields and amongst Pinus, Quercus, Abies and Cedrus, at 700-2000 m (Figure 161).

Specimens examined: B6 Sivas: Gürün, Gökpınar, 1525 m, 7.vi.2006, A.Kahraman 1227. B7 Erzincan: Kemaliye, 977 m, 6.vi.2006, A. Kahraman 1390; Malatya: Beydağı, 1300 m, 7.vi.2006, A. Kahraman 1213; Elazığ: Elazığ to Keban, ca. 2 km to Keban, 760 m, 7.vi.2006, A. Kahraman 1198. B8 Erzincan: Tercan to Aşkale, 23 km East of Tercan, 1680 m, Hub.-Mor. 10767. B9 Van: Van to Gürpınar, 1798 m, 16.vii.2006, A. Kahraman 1327; Bitlis: Gayda to Bahçesaray, near Nazar bridge, 1256 m, 9.vii.2007, A. Kahraman 1429. C6 Adıyaman: Besni to Adıyaman, 700 m, E. Tuzlacı 40102 (ISTE!). C8 Mardin: 11 km from Gercüş to Hasankeyf, 750 m, D.

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43042. C9 Hakkari: Şırnak to Hakkari, Güneş to Çığlı, 1137 m, 7.vi.2008, A. Kahraman 1554. C10 Hakkari: 30 km from Başkale to Hakkari, 1900 m, D. 44677.

General distribution and phytogeography: S. candidissima subsp. candidissima is a member of the Irano-Turanian element and is found in Iraq.

Taxonomic notes: S. candidissima subsp. candidissima differs S. candidissima subsp. occidentalis in the large and often discolorous leaves with a thin (to thick) indumentum, serrulate to erose margins and white corollas with a yellow lip.

Figure 158. General appearance of S. candidissima subsp. candiddissima in its natural habitat. 282

46. S. cyanescens Boiss. & Bal. in Boiss., Diagn. ser. 2(4): 19 (1859). (Figure 159).

Type: [TURKEY C5 Niğde] in collibus secus fluvium Kamechly Tchai (Kameçli çay) ad basin fluvii Masmeneu Dagh Cappadociae inter Tarsous et Caesaream, 14.vi.1856, Balansa 233 (holo. G! photo E!).

Description: Perennial herb. Stems erect, 25-85 cm high, branched above, pilose to lanate below, pilose with sessile glands to ± densely papillose glandular above. Leaves simple, oblong to broadly ovate to suborbicular, 3-9 x 2-6.5 cm, pannose, serrulate to subentire, ± cordate; petiole 1-7 cm long. Inflorescence paniculate with slender branches. Verticillasters (1-) 2-6-flowered, distant. Bracts broadly ovate-acuminate, 2.5-6 x 2-5 mm. Pedicels 1.5-3 mm long. Calyx tubular-campanulate, usually purple-tinted, 7-12 mm long, to 15 mm long in fruit, shortly pilose or papillose glandular; upper lip tridentate, median tooth longer or shorter than, laterals, scarcely mucronate. Corolla lilac to violet, 16-33 mm long; tube 7-20 mm long, ventricose, squamulate (rarely not); upper lip strongly falcate. Stamens of type B, included within upper lip of corolla.

Phenology: Flowering from June to September.

Distribution and habitat in Turkey: This species is distributed in all regions of Turkey except for Southeast Anatolia where it grows on limestone and igneous slopes, shale banks, gravel river beds and in Pinus nigra forest, fallow fields and vineyards, at 400-2300 m (Figure 161).

Specimens examined: B7 Erzincan: İliç to Refahiye, 3 km from Koçkaya village, 39o46′249′′N-38o38′638′′E, 1609 m, 25.vii.2008, A. Kahraman 1593.

General distribution and phytogeography: S. cyanescens is endemic to Turkey and a member of the Irano-Turanian element.

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Taxonomic notes: S. cyanescens is close to S. candidissima and S. cilicica, but is differentiated from the former by the smaller lilac-tinted calyces and corollas and from the latter by the several pairs of cauline leaves and many-flowered inflorescences.

Figure 159. General appearance of S. cyanescens in its natural habitat.

47. S. cilicica Boiss. & Kotschy in Boiss., Diagn. ser. 2(4): 19 (1859). (Figure 160). Synonym: S. crastfolia sensu Boiss., Fl. Or. 4: 622 (1879) p.p. non Sibth. & Sm. (1806) nee Desf. (1809).

Type: [TURKEY C5 Niğde] in schistosis vinetorum pagi Anadscha Tauri Cilicici, in calcareis ad fluvium Bosante tchai (Pozanti Cay) in via inter Tar-sous et Caesaream (Kayseri), 1220 m,viii.1855, Balansa 546 (holo. G!). 284

Description: Perennial herb. Stems erect, 30-130 cm high, branched above and slender, pan-nose to ± lanate below, very shortly pilose with capitate glandular hails above. Leaves simple, broadly ovate to orbicular, 2-11 x 3-11 cm, pannose, serrulate to subentire, cordate; petiole on lower leaves 1.5-7 cm long, upper leaves sessile. Inflorescence paniculate, few-flowered. Verticillasters (1-) 2-6-flowered, clearly distant. Bracts broadly ovate-acuminate, 2.5-4.5 x 2-3 mm. Pedicels 1-3 mm long. Calyx tubular- campanulate, striate, 5-8 mm long, to ca. 9.5 mm long in fruit, shortly pilose with sessile glands; upper lip scarcely tridentate, recurved in fruit. Corolla lilac-white with a yellow lower lip, 14-25 mm long; tube 5-8 mm long, ventricose, squamulate; upper lip clearly falcate. Stamens of type B, included within upper lip of corolla.

Phenology: Flowering from July to September.

Distribution and habitat in Turkey: This species is distributed mainly in the eastern Mediterranean region of Turkey and occasionally in East Anatolia where it grows on limestone and granite slopes, in phrygana and among Pinus, at 750-2200 m (Figure 161).

Specimens examined: B6 Kahramanmaraş: Göksun, Ericek to Yeşilköy, 2-3 to Yeşilköy, 38o00′721′′N-36o44′568′′E, 1411 m, 5.vii.2007, A. Kahraman 1390.

General distribution and phytogeography: S. cilicica is endemic to Turkey and a member of the Mediterranean element.

Taxonomic notes: S. cilicica is close to S. cyanescens, but differs in four pairs of cauline leaves and few-flowered inflorescences.

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Figure 160. General appearance of S. cilicica in its natural habitat.

(45) S. candidissima subsp. candidissima (46) S. cyanescens (47) S. cilicica Figure 161. Distribution of S. candidissima subsp. candidissima, S. cyanescens and S. cilicica in the study area.

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48. S. limbata C.A. Meyer, Verz. Pfl. Cauc. 86 (1831). (Figure 162). Synonym: S. chrysadenia Freyn in Bull. Herb. Boiss. ser. 2, 1: 279 (1901). Ic: Takht. & Fedorov, Fl. Erevana ed. 2, f. 82 (1972).

Type: [USSR, AZERBAIJAN] in locis siccis lapidosis montium Talüsch (Talysh) prope pagurn Swant, c. 1280 m, C.A. Meyer 140 (holo. LE).

Description: Perennial herb with erect stems much branched above, 30-120 cm long, retrorsely scabridulous below and with sessile glands above. Leaves simple, ovate-oblong, cordate at base, (3.5-) 5-16 x (2-) 4-10 cm, glabrous and prominently glandular- punctate below, sparsely pilose above; margins erose-dentate; petiole 2-12 cm long. Verticillasters 2-6 (-8)-flowered, distant. Bracts ovate-acuminate, soon deciduous. 2-7 mm long. Pedicels 2-6 mm long, erecto-patent. Calyx ovate-campanulate, 6-10 mm long, to 13 mm long in fruit, with scabrid eglandular hairs and sessile glands; upper lip shortly tridentate, mucronate, recurved. Corolla white with pale yellow lip, 20-25 mm long; tube short, 5-9 mm long, abruptly ventricose, squamulate; upper lip falcate, villous at tip. Stamens of type B, included within upper lip of corolla. Filaments 3-4 mm long; upper connectives 15-20 mm long; fertile athers 3-4 mm long. Style 35-40 mm long.

Phenology: Flowering from June to August.

Distribution and habitat in Turkey: This species is distributed in East and Southeast Anatolia where it grows on stony slopes, steppe, sandy loam, roadsides and cornfields, 1060-2340 m (Figure 165).

Specimens examined: A9 Kars: Kötek to Kars, 62 km to Kars, 1480 m, 15.vi.1979, A. Baytop, B. Çubukçu, E. Tuzlacı and M. Saraçoğlu 42990 (ISTE!). B8 Erzurum: Ilıca to Erzurum, ca. 5 km to Erzurum, 38o55′00′′N-41o12′42′′E, 1817 m, 10.vii.2006, A. Kahraman 1293. B9 Van: Van to Gürpınar, 38o22′31′′N-43o23′30′′E, 2220 m, 16.vii.2006, A. Kahraman 1329; Bitlis: Van to Bitlis, 38o21′268′′N-42o40′981′′E, 1959

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m, 8.vi.2008, A. Kahraman 1582. B10 Kars: 32 km from Doğubayazıt to Iğdır, 1420 m, 13.vii.1979, A. Baytop, B. Çubukçu, E. Tuzlacı and M. Saraçoğlu 42824 (ISTE!). C10 Hakkari: Hakkari to Van, 1839 m, 7.vi.2008, A. Kahraman 1571.

General distribution and phytogeography: S. limbata is a member of the Irano- Turanian element and present in Iran and Transcaucasia.

Taxonomic notes: S. limbata is an unusual species on account ofthe absence of capitate glandular hairs.

Figure 162. General appearance of S. limbata in its natural habitat.

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49. S. indica L., Sp. Pl. 26 (1753). (Figure 163). Synonym: S. brachycalyx Boiss., Fl. Or. 4: 625 (1879). Ic: Bot. Mag. t. 395 (1798).

Type: Described from INDIA. [No specimen in the Linnean herbaria has been found; no. 1079 in Hb: Toumefbrt (Paris) could possibly be selected as type.] Description: Perennial herb. Stems 80-150 cm high, erect, branched above, sparsely eglandular pilose below, densely or sparsely glandular or eglandular pilose above. Leaves simple, 15-30 x 10-24 cm, broadly ovate, truncate, reticulate, erose-dentate, glaucous or eglandular-pilose especially on the veins, cordate; petiole 2-12 cm long; cauline leaves ± sessile. Inflorescence paniculate. Verticillasters 4-8-flowered, clearly distant. Bracts 8-10 x 6-8 mm, ovate-acuminate, green. Pedicels 2-5 (-7) mm long, erect. Calyx campanulate, truncate at apex, 10-12 x 7-9 mm, slightly broadening and up to 15 mm long in fruit, densely glandular-villous and sparsely eglandular; upper lip shortly tridentate; calyx teeth mucronate, connivent. Corolla upper lip lilac, lower lip dark violet, spotted with purple, 25-32 mm long, strongly compressed laterally; tube 9-13 mm long, whitish, short, squamulate within, abruptly ventricose above. Stamens of type B, included within upper lip of corolla, upper connectives clearly longer than filaments. Filaments 3-4 mm long; upper connectives 17-21 mm long; fertile anthers 4-5 mm long. Style glabrous, 38-45 mm long, exserted from corolla lips and divided into two parts at apex.

Phenology: Flowering in April and May.

Distribution and habitat in Turkey: This species is distributed in East, South and Southeast Anatolia where it grows on on rocky limestone slopes and moist places in Quercus scrub, at 100-1500 m (Figure 165).

Specimens examined: C9 Siirt: Eruh to Şırnak, near Yanılmazlar village, 37o42′269′′N- 42o15′804′′E, 1195 m, 5.vi.2008, A. Kahraman 1539; Şırnak: Hessana (Kösreli) at foot

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of Cudi Mountain, 900 m, 10.v.1966, D. 42804 (E!). C10 Hakkari: below Oramar in gorge running, Southeast of Ziri, 1300 m, 17.v.1978, Trelawny & McPhail 2040 (E!).

General distribution and phytogeography: S. indica is a member of the Irano-Turanian element and present in Palaestina, Iran, Iraq, Syria and Jordan.

Taxonomic notes: S. indica is a very handsome species and close to the recently described endemic species S. siirtica, however can be distinguished by the stem and petiole indumentum, distant verticillasters, larger flowers, truncate calyces at apex and a lilac upper lips and dark violet lower lips of corollas spotted with purple.

Figure 163. General appearance of S. indica in its natural habitat.

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50. S. siirtica Kahraman, Celep & Doğan (Figure 164).

Type: TURKEY Siirt: North of Siirt, Aydınlar, Çatılı region, 38o1'23.9988''N, 42o16'0.9984''E, 1473 m, open Quercus forest, 3.vi.2009, Kahraman 1650 (holotype: ANK, isotypes: E, GAZI). Description: Herbaceous perennials. Stems 40-120 cm high, 0.4-1.0 mm wide at base, erect, quadrangular, green, branched above or unbranched, densely glandular pilose to villous. Leaves simple, green, 7-25 x 5.5-23 cm, broadly ovate, reticulate, erose-dentate to lobed, cordate or not, pilose with glandular and eglandular hairs, especially on veins and margins; petiole 2-14 cm long, densely glandular pilose to villous; cauline leaves ± sessile. Inflorescence paniculate, densely pilose to villous with mostly glandular hairs. Verticillasters close, 6-12-flowered, rarely 4-flowered at apex. Bracts ovate-acuminate, 6-12 x 5-12 mm, green, bracts of upper verticillasters shorter than calyx; pedicels 2-4 mm long, rigid, densely pilose to villous with glandular and eglandular hairs. Calyx campanulate, 5.5-10 x 5-8 mm, clearly concave at apex, 13-15-veined, green, thick- textured, densely pilose to villous with glandular and eglandular hairs, slightly broadened and up to 12 mm in fruit; upper lip tridentate; calyx teeth mucronate. Corolla entirely white, 13-25 mm long, upper lip densely villous with eglandular purple hairs and sparse glandular hairs; tube 5-8 mm long, glabrous, squamulate, ventricose above. Stamens of type B, included within upper lip of corolla, clearly longer than filaments. Filaments glabrous, 3-4 mm long; upper connectives 11-22 mm long, lower connectives 3-5 mm long, sterile, reduced to a dolabriform plate, with a tooth running down the plate; fertile anthers yellow, glabrous, 3-5 mm long. Styles white at base, violet at apex, glabrous, up to 35 mm long, exserted from corolla lips and divided in two parts at apex.

Phenology: Flowering in May and June.

291

Distribution and habitat in Turkey: This species is distributed only in Çatılı region in Siirt province, the southeastern part of Turkey where it grows in open forests of Quercus, at 1450-1500 m (Figure 165).

General distribution and phytogeography: S. siirtica is endemic to Turkey and a member of the Irano-Turanian element.

Taxonomic notes: S. siirtica is closely related to S. indica, but differs in the densely glandular stems and petioles indumentum, close verticillasters, smaller flowers, clearly concave calyces at apex and entirely white corollas.

Figure 164. General appearance of S. siirtica in its natural habitat.

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(48) S. limbata (49) S. indica (50) S. siirtica Figure 165. Distribution of S. limbata, S. indica and S. siirtica in the study area.

Section 5: Drymosphace Benth. Lab. Gen. Sp. 218 (1833).

51. S. glutinosa L., Sp. Pl. 26 (1753). Ic: Fl. RPR 8: t. 39 f. 1 (1961); Huxley & Taylor, Fls. Greece t. 237 (1977). (Figure 166).

Type: Described from EUROPE (lecto. Hb. Linn. 42/34 photo! Microslayt: IDC 11.18).

Description: Perennial herb. Stems to ca. 130 cm high, erect, branched above, ± densely glandular and sparsely eglandular villous. Leaves simple, ovate-triangular, 4-20 x 3-12 cm, sagittate-hastate, serrate, subglabrous to sparsely pilose on veins and with sessile glands. Petiole 4-11 cm long. Verticillasters 4-6-flowered, distant. Bracts ovate, 6-14 x 4-8 mm; bracteoles present. Pedicels up to 5 mm, erecto-patent, elongating in fruit. Calyx tubular to campanulate, 10-14 mm long, up to 20 mm long in fruit, densely glandular- villous; upper lip 1-dentate, ± straight. Corolla yellow with brownish markings, 30-40 mm long; tube exserted, straight for ca. 15 mm, villous within; upper lip falcate. Stamens of type B, included within upper lip of corolla. Filaments 4-6 mm long; upper connectives 10-15 mm long, lower connectives 4-5 mm long; fertile anthers 4-5 long mm. 293

Phenology: Flowering from July to October.

Distribution and habitat in Turkey: This species is distributed mainly in North Anatolia and rarely in East Anatolian, Mediterranean and Marmara geographic regions of Turkey (Figure 167). It occurs on moist places in deciduous (Fagus, Alnus, Corylus) forest and scrub and in Picea forest, at 780-2360 mm.

Specimens examined: A9 Ardahan: Posof to Derindere, 41o28'597''N-42o46'655''E, 1660 m, 28.vii.2008, A. Kahraman 1606.

General distribution and phytogeography: S. glutinosa is a member of the Euro- Siberian element and is found in Europe, Caucasia and Iran.

Taxonomic notes: S. glutinosa is a very distinctive species on account of the ovate- triangular and sagittate-hastate leaves.

Figure 166. General appearance of S. glutinosa in its natural habitat. 294

(51) S. glutinosa Figure 167. Distribution of S. glutinosa in the study area.

Section 6: Plethiosphace Benth. Lab. Gen. Sp. 230 (1833).

52. S. staminea Montbret & Aucher ex Benth. in Ann. Sci. Nat. ser. 2, 6: 41 (1836). Pobedimova in Not. Syst. (Leningrad) 21: 320-324 (1961). (Figure 168). Synonym: S. staminea Montbret & Aucher ex Benth. var. persica Bornm. in Bull. Herb. Boiss. ser. 2, 8: 110 (1908); S. staminea Montbret & Aucher ex Benth. subsp. armeniaca Bordz. in Acta Horti Jurjev. 13: 22 (1912); S. staminea Montbret & Aucher ex Benth. var. latifolia Bornm. in Monit. Jard. Bot. Tiflis 32: 7 (1914); S. armeniaca (Bordz.) Grossh. in Beih. Bot. Centr. 44(2): 237 (1927); S. transcaucasica Pobed. in Not. Syst. (Leningrad) 21: 321 & fig. p. 323 (1961). Ic: Fl. URSS 21: t. 16 f. 3 (1956), as S. armeniaca. Type: [TURKEY B7] Cappadocia orientali, Aucher 1526 (holo. G!).

Description: Perennial herb. Stems erect, 20-70 cm high, branched above, with a variable indumentum, pilose to villous eglandular below and glandular above. Leaves distributed over stem or rosette-forming, linear-oblong to ovate, 2.5-15 x 1-6 cm, subglabrous to shortly tomentose with sessile glands; margins subentire to erose; petiole 2-9 cm long. Inflorescence up to 40 cm long, paniculate, eglandular tomentose to villous 295

sometimes with glandular pilose to villous. Verticillasters 2-6 (-8)-flowered, approximating above. Bracts broadly ovate, 2-15 x 4-10 mm. Pedicels 2-3 mm long. Calyx tubular-campanulate, 6-8 mm long, up to 12 mm in fruit, yellowish-green or with violet blue striations, densely glandular with or without long flattened eglandular hairs; upper lip tridentate, equal to or shorter than lower. Corolla white to pale yellow, 12-16 mm long; tube ca. 5 mm long, ventricose, squamulate, upper lip ± straight, narrow. Stamens of type B, much exserted. Filaments ca. 1-2 mm long, upper connectives (5-) 9- 15 (-18) mm long, lower connectives ca. 1-2 mm long; fertile anthers ca 1-1.5 long.

Phenology: Flowering from May to August.

Distribution and habitat in Turkey: This species is distributed in East and Northeast Anatolia where it grows on limestone and igneous rocky slopes, cliffs, screes, in Quercus scrub and alpine meadows, at 1700-3150 m (Figure 172).

Specimens examined: A9 Erzurum: Şenkaya, Allahuekber Mountain, 40o33'307''N- 42o26'780''E, 2643 m, 27.vii.2008, A. Kahraman 1601; Kars: Susuz to Ardahan, 31 km to Ardahan, 40o58'17''N-42o53'50''E, 2012 m, 13.vii.2006, A. Kahraman 1310. B7 Elazığ: Elazığ to Baskil, above Kayabeli village, 38o35'35''N-38o52'26''E, 1600 m, 7.vi.2006, A. Kahraman 1209B; Erzincan: Sipikör Mountain, 39o52'39''N-39o33'36''E, 2270 m, 10.vii.2006, A. Kahraman 1277. B8 Erzurum: 29 km from Hınıs to Pasinler, 1800 m, 12.vii.1966, D. 46372 (E!); Bingöl: Bingöl Mountain, above Karlıova, 2000 m, 10.vii.1966, D. 46185 (E!); Siirt: Meleto (Meretuğ) Mountain, ca. 2800 m, Hand.- Mazz. 2786. B9 Van: Bahçesaray to Van, Sarı (Kilazer) hill, 38o08'618''N-42o51'618''E, 2503 m, 9.vii.2007, A. Kahraman 1436; Ağrı: Ağrı to Horasan arası, near Tahir village, 39o50'13''N-42o25'275''E, 2081 m, 12.vii.2007, A. Kahraman 1460; Ağrı: Cumaçay to Akçay, ca. 1-2 km to Sarıbulak village, 39o59'507''N-43o19'953''E, 2200 m, 13.vii.2007, A. Kahraman 1460; Bitlis: Nemrut Mountain, 38o33'14''N-42o15'06''E, 1976 m, 18.vii.2006, A. Kahraman 1349B. B10 Van: Güzeldere pass, 34 km from Başkale to

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Hoşap, 2800 m, 3.vii.1977, D. 45946 (E!). C10 Hakkari: Cilo Mountain, 3150 m, D. 24140 (K!).

General distribution and phytogeography: S. staminea is a member of the Irano- Turanian element and present in Armenia, Georgia and Iran.

Taxonomic notes: S. staminea is very distinct species on account of the creamy corollas with a narrow upper lip and the long exserted stamens beyond upper lip of corolla.

Figure 168. General appearance of S. staminea in its natural habitat.

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53. S. virgata Jacq., Hort. Vindob. 1: 14, t. 37 (1770). (Figure 169). Synonym: S. sibthorpii Sibth. & Sm., Fl. Graeca 1: 17, t. 22 (1806); S. similata Hausskn. in Mitt. Thür. Bot.Ver. N.F. 11:36 (1897); S. virgata Jacq. var. densiflora Náb. in Publ. Fac. Sci. Univ. Masaryk Brno 70: 52, t. 15 f. 4 (1926); S. virgata Jacq. var. canovelutina Rech. f. in Ann. Naturh. Mus. Wien 51: 420 (1941). Ic: Sibth. & Sm., Fl. Graeca 1: t. 22 (1806), as S. sibthorpii.

Type: Described from cultivated material; no type indicated.

Description: Perennial usually coarse herb. Stems erect, (10-) 20-130 cm high, much branched above or not, indumentum variable, pilose to tomentose, glandular or eglandular. Leaves simple, distributed over stem or rarely restricted to basal rosettes, ovate- oblong to broadly ovate, 5-30 x 2-15 cm, eglandular-pilose with numerous sessile glands, cordate, rugulose; margins erose, crenate, serrate to subentire; petiole 1-15 cm long. Inflorescence a widely branched panicle with long ± slender secondary branches. Verticillasters 2-6-flowered, distant, rarely condensed. Bracts ovate-acuminate, 3-9 x 3- 7 mm. Pedicels 1-3 mm long. Calyx ± tubular campanulate, 6-10 mm long, to 10-12 mm long in fruit with a strongly recurved bisulcate upper lip, glandular or eglandular pilose. Corolla violet-blue to lilac, rarely white, 10-18 mm long; tube 6-13 mm, ventricose, not squamulate; upper lip falcate. Stamens of type B.

Phenology: Flowering from May to September.

Distribution and habitat in Turkey: This species is distributed in all regions of Turkey where it occurs on scrub, woodland, meadows, fallow fields and roadsides, at 1- 2300 m (Figure 172).

Specimens examined: A9 Kars: Kars to Kağızman, around Kötek village, 1275 m, 14.vii.2006, A. Kahraman 1313. B6 Malatya: Malatya to Darende, 1410 m, 7.vi.2006, A. Kahraman 1218. B7 Erzincan: İliç to Kemah, 1306 m, 9.vii.2006, A. Kahraman

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1261. B8 Muş: Muş to Bingöl, 20 km West of Muş, 1420 m, Hub.-Mor. 10760. B9 Bitlis: near Kuşkuran pass, 2156 m, 18.vii.2006, A. Kahraman 1341. B10 Ağrı: Karabulak to Doğubayazıt, 1554 m, 14.vii.2006, A. Kahraman 1318B. C9 Şırnak: Şırnak to Hakkari, near Yemişli village, 1137 m, 6.vi.2008, A. Kahraman 1549.

General distribution and phytogeography: S. virgata is a member of the Irano-Turanian element and present in Crimea, Cyprus, Balkans, Italy, Caucasia, Talish, Iran, Iraq, Turcomania and Afghanistan.

Taxonomic notes: Size, degree of branching, leaf size and shape, flower colour and indumentum are very variable in S. virgata. The species is close to S. nemorosa and S. dicroantha, but differs from them by the larger ovate-oblong to broadly ovate leaves.

Figure 169. General appearance of S. virgata in its natural habitat.

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54. S. nemorosa L., Sp. Pl. ed. 2, 35 (1762). (Figure 170). Synonym: S. sylvestris sensu Boiss., Fl. Or. 4: 628 (1879) p.p. non L. (1753); S. tesquicola Klokov & Pobed. in Fl. URSS 21: 662 (1954) or at least sensu Grossh., Fl. Kavk. ed. 2, 7: 394 (1967). Ic: Bot. Mag. 153: t. 9169 (1929), as S. x superba Stapf; Fl. RPR 8: t. 44 f. 2 (1961). Type: TARTARIA (TARTARY), Gerber (?) (Hb. Linn. 42/14). For a note on nomenclature see Notes R.B.G. Edinb. 23: 563-566 (1961).

Description: Perennial herb. Stems 30-80 (-100) cm high, many, erect, branched above, eglandular pilose to pubescent with short hairs, occasionally retrorse, or longer spreading hairs. Leaves simple, oblong-lanceolate, cordate, 3-10 x 1.2-4 cm, pubescent with few to many sessile glands; margins serrulate to crenate; petiole l-4 (-4.5) cm long. Verticillasters many, ca. 6-flowered, approximating, showy. Bracts ovate, acuminate, 4-10 x 2.5-6 mm, purplish or violet. Pedicels 2-3 mm long. Calyx rubular-campanulate, ca. 6-7 mm long, to 8-8.5 mm long in fruit, pilose to villous with sessile glands; upper lip strongly recurved in fruit. Corolla purple to violet-blue, 10-12 mm long; tube ventricose, not squamulate; upper lip slightly falcate. Stamens of type B.

Phenology: Flowering from the end of May to September.

Distribution and habitat in Turkey: This species is distributed mainly in East Anatolia and rarely in Northeast Anatolia where it grows on rocky slopes in steppe, fallow fields, sloping meadows and waste ground, at 1000-2300 m (Figure 172).

Specimens examined: A9 Kars: near Selim, 40o17'40''N-42o39'21''E, 2012 m, 12.vii.2006, A. Kahraman 1308; Ardahan: Posof, Türkgözü, 28.vii.2008, A. Kahraman 1603; Erzurum: Şenkaya, Allahuekber Mountain, 40o33'212''N-42o22'188''E, 2025 m, 27.vii.2008, A. Kahraman 1600. B8 Erzurum: 8 km from Aşkale to Ilıca, 39o56'12''N-40o47'09''E, 1700 m, 10.vii.2006, A. Kahraman 1290A. B9 Ağrı: Tutak to

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Hamur, 2 km to Hamur, 39o36'392''N-42o57'767''E, 1610 m, 12.vii.2007, A. Kahraman 1456; Bitlis: Kotum to Reşadiye, D. 22384; Van: 5 km North of Çatak, 1800 m, D. 23245. B10 Ağrı: near Doğubayazıt, 14.vii.2006, A. Kahraman 1319A. C9/10 Hakkari: Hakkari to Van, 37o41'990''N-44o05'292''E, 1767 m, 7.vi.2008, A. Kahraman 1568. C10 Hakkari: Yüksekova, 1950 m, 7.ix.1967, Duncan & Tait (E!).

General distribution and phytogeography: S. nemorosa is a member of the Irano- Turanian element and is found in Europe, Balkans, Russia, Crimea, Caucasia, Iran and Afghanistan.

Taxonomic notes: S. nemorosa is close to S. dicroantha and S. virgata, but differs them from them in the prominent (always) purplish or violet bracts.

Figure 170. General appearance of S. nemorosa in its natural habitat.

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55. S. dicroantha Stapf in Denkschr. Akad. Wiss. Wien, Math.-Nat. Kl. 50(2): 96 (1885). (Figure 171). Synonym: S. sylvestris sensu Boiss., Fl. Or. 4: 628 (1879) p.p. non L.; S. nemorosa auct. p.p. non L. (1753).

Type: [TURKEY C2 Muğla: inter Baschibunar (Başpınar) et Gurdef (Girdev), 19.vii.1882, Luschan] (holo. WU, photo E!).

Description: Perennial herb. Stems 20-60 (-80) cm high, ascending-erect, branched above or not, eglandular pubescent. Leaves simple, narrowly oblong to oblong- lanceolate, ± cordate, 3-14 x 0.5-4 cm, rugulose, pubescent with numerous sessile glands; margins crenulate; petiole 1-5 cm long. Inflorescence paniculate or not, up to 40 cm long. Verticillasters 2-6-flowered, approximating. Bracts ovate-acuminate, 1-6 x 1-4 mm. Pedicels 0.5-2 mm long. Calyx tubular-campanulate, 4-7 mm long, to ca. 8 mm long in fruit, green to purple, pubescent with numerous sessile glands and rarely with some capitate glandular hairs. Corolla pale lilac or blue, 9-15 mm long; tube ventricose, not squamulate; upper lip falcate. Stamens of type B.

Phenology: Flowering from June to September.

Distribution and habitat in Turkey: This species is distributed all regions of Turkey except for Southeast Anatolia where it grows on Steppe, clearings in forest, fallow fields and roadsides, at 700-2000 m (Figure 172).

Specimens examined: B6 Sivas: Kangal to Gürün, 38o53′400′′N-37o16′068′′E, 1765 m, 16.vii.2007, A. Kahraman 1487; Kayseri: Sarız, Yukarı Kırkısrak village, 1680 m, 23.vii.2008, A. Kahraman 1582. B7 Elazığ: Elazığ, 28.x.1964, F. Sayi (E!).

General distribution and phytogeography: S. dicroantha is endemic to Turkey and a member of the Irano-Turanian element.

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Taxonomic notes: S. dicroantha resembles S. nemorosa and S. adenocaulon, but differs from the former by the absence of the prominent violoet bracts and from the latter by the egalandular stems.

Figure 171. General appearance of S. dicroantha in its natural habitat.

(52) S. staminea (54) S. nemorosa (53) S. virgata (55) S. dicroantha

Figure 172. Distribution of S. staminea, S. virgata, S. nemorosa and S. dicroantha in the study area. 303

Section 7: Hemisphace Benth. Lab. Gen. Sp. 310 (1833).

56. S. verticillata L., Sp. PI. 26 (1753). Rech. f. in Bot. Jahrb. 71: 539-544 (1941); Pénzes in Borbasia 5-6 (1-3): 1-31 (1944). (Figures 173, 174).

Description: Perennial herb; stems erect or ascending, 15-70 cm high, many, branched above or not, pilose to villous below with sessile glands. Leaves simple, oblong to ovate, 2.5-13 x 2-9 cm, or lyrate with one or two pairs of unequal basal lobes, pilose to villous with many sessile glands, subentire to serrate, rounded to cordate, petiole 1.5-7 (- 9) cm long. Verticillasters (8-) 20-40-flowered, clearly distant. Bracts ovate-acuminate, 5-7 x 2.5-3 mm, deciduous. Pedicels 2-8 (-10) mm, some ± deflexed. Calyx tubular, 5-6 mm long, to ca. 7 mm long in fruit with a bisulcate upper lip, violet-blue, pilose to villous with sessile glands, teeth mucronate. Corolla violet-blue, lilac, rarely white, ca. 12 mm; tube straight, ca. 8 mm with a V-shaped annulus; upper lip straight, narrowed at base. Stamens of type C.

1. Leaves broadly ovate, clearly cordate; stem, leaves and calyx indumentum of villous soft hairs; plant to ca. 70 cm, much branched………………...…...…….subsp. verticillata 1. Leaves oblong, elliptic, or oblong-ovate, rounded or subcordate; indumentum of short ± scabridulous hairs; plant 15-50 cm, little branched………………….….subsp. amasiaca subsp. verticillata (Figure 173) Synonym: S. uberrima Rech. f.: in Bot. Jahrb. 71: 543 (1941). Ic: Reichb., Ic. Fl. Germ. 18: t. 1255 (1856); Fl. RPR 8: t. 41 f. 1 (1961).

Type: Australia et Misnia (Meissen, Germania), Herb. Linn. 42/30.

Phenology: Flowering from June to August.

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Distribution and habitat in Turkey: This species is distributed in the East Anatolian, Black Sea and Maramara geographic regions of Turkey where it grows in a wide variety of habitats including Pinus, Quercus, Fagus and Corylus woods, meadows, gravel beds and roadsides, at 1-2400 m (Figure 176).

Specimens examined: A9 Ardahan: Posof, Türkgözü, 28.vii.2008, A. Kahraman 1604; Erzurum: Şenkaya, Allahuekber Mountain, 2643 m, 27.vii.2008, A. Kahraman 1599. B8 Erzurum: Ilıca to Tercan, 1850 m, 10.vii.1957, D. 30862 (ANK!); Muş: South slopes of Bingöl Mountain, near Gümgüm (Varto), 1710 m, Kotschy 332 (type of S. uberrima). B9 Bitlis: Kambos Mountain, 1282 m, 9.vi.2008, A. Kahraman 1583A; Van: Çatak, Kavuşşahap Mountain, 2150 m, 22.vii.1954, D. 22990 (ANK!). C9 Şırnak: Şırnak to Hakkari, near Yemişli village, 1245 m, 6.vi.2008, A. Kahraman 1551; Hakkari: Hakkari to Uludere, 2400 m, 22.vii.1974, M. Koyuncu (AEF!).

General distribution and phytogeography: Subsp. verticillata is a member of the Euro- Siberian element and is found in Europe, Caucasia, Iran, Iraq and Talish.

Figure 173. General appearance of S. verticillata subsp. verticillata in its natural habitat. 305

subsp. amasiaca (Freyn & Bornm.) Bornm. in Bull. Herb. Boiss. ser. 2, 8: 110 (1908). (Figure 174). Synonym: S. amasiaca Freyn & Bornm. In Ost. Bot. Zeitschr. 41: 58 (1891); S. paalii Pénzes in Borbásia 5-6(1-3): 14 (1946). Ic: Fl. URSS 21: t. 17 f. 1 (1954).

Type: [TURKEY A5 Amasya] Amasia: in rupestribus ad 4-600 m, 11.vi.1899, Bornmüller 611.

Phenology: Flowering from May to September.

Distribution and habitat in Turkey: This species is distributed in all regions of Turkey where it grows in a wide variety of habitats including rocky slopes, Stipa steppe, sandy banks, meadows, Quercus and Pinus woods, fields and roadsides, at 150- 2300 m (Figure 176).

Specimens examined: A9 Kars: Horasan to Sarıkamış, 20 km to Sarıkamış, 1500 m, 12.vii.2006, A.Kahraman 1305. B6 Malatya: Malatya to Darende, near Yukarı Ulupınar village, 1410 m, 7.vi.2006, A. Kahraman 1217. B7 Sivas: Dumlucadağ, 1500-1575 m, 6.vi.2006, A. Kahraman 1169. B8 Erzurum: Ilıca to Erzurum, 1817 m, 10.vii.2006, A. Kahraman 1817. B9 Bitlis: Kambos Mountain, 1282 m, 9.vi.2008, A. Kahraman 1583B. B10 Iğdır: Ağrı Mountain, 1318 m, 14.vi.2007, A. Kahraman 1318A. C9 Hakkari: Cilo Mountain, 2280 m, D. 23946. C10 Hakkari: Bacirge, 1750 m, D. 45256.

General distribution: Subsp. amasiaca is a member of the Irano-Turanian element and is found in Bulgaria, Caucasia, Iran and Iraq.

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Figure 174. General appearance of S. verticillata subsp. amasiaca in its natural habitat.

57. S. russellii Benth. in DC., Prodr. 12: 357 (1848). (Figure 175).

Type: [SYRIA] prope Aleppo, Russell (holo. BM).

Description: Perennial herb. Stems 20-60 cm high, erect, usually simple, many, arising from a woody rootstock, eglandular-pubescent. Leaves simple, linear-oblong, sometimes sublyrate or deeply lobed at base, 5-8 x 1-1.5 cm, eglandular pilose especially on veins and with numerous sessile glands, rugulose; margins cuneate; petiole 2-8 (-10) cm long. Verticillasters 20-30-flowered, clearly distant. Bracts 5-6 x 1.8-2 mm, ovate-acuminate. Pedicels 2-6 mm long, erecto-patent. Calyx tubular, 5-7 mm long, purplish-blue, eglandular spreading pilose and with sessile glands, ovate in fruit with a concave-bisulcate upper lip, teeth not mucronate. Corolla violet-blue, 8-10 mm long; tube straight wit an 307

inverted V-shaped annulus; upper lip straight, compressed, narrowed at base. Stamens of type C.

Phenology: Flowering from May to July.

Distribution and habitat in Turkey: This species is distributed in the Central, East, Southeast Anatolian, Mediterranean and Marmara geographic regions of Turkey where it occurs on rocky slopes, grassy meadows amongst Quercus, fallow and in cultivated fields, 100-1600 m (Figure 176).

Specimens examined: B7 Tunceli: above Pertek, 1600 m, D. 31054 (ANK!). B8 Batman: North of Sason, Sason Mountain, Batman, 1000 m, Watson 5597. B9 Van: Van to Gürpınar, 2230 m, 11.vii.2007, A. Kahraman 1451. C6 Gaziantep: Gaziantep, 850 m, Balls 1166. C7 Şanlıurfa: North of Şanlıurfa, Hand.-Mazz. 1911. C8 Mardin: Mardin station to Kızıltepe, 600 m, D. 28637. C9 Hakkari: Şırnak to Hakkari, Güneş to Çığlı, 37o18′709′′N-43o27′791′′E, 1137 m, 7.vi.2008, A. Kahraman 1553.

General distribution and phytogeography: S. russellii is a member of the Irano- Turanian element and present in Syria, Iran and Iraq.

Taxonomic notes: S. russellii is close to S. verticillata, but can beeasily distinguished by the habit, narrowly linear-oblong leaves and acuminate (not mucronate) calyx teeth.

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Figure 175. General appearance of S. russellii in its natural habitat.

(56) S. verticillata (56) S. verticillata (57) S. russellii subsp. amasiaca subsp. verticillata

Figure 176. Distribution of S. verticillata subsp. amasiaca, S. verticillata subsp. verticillata and S. russellii in the study area.

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CHAPTER 4

CONCLUSION

A taxonomic revision of the genus Salvia is provided based on study of plant specimens collected from about 600 different populations in East and Southeast Anatolia (Turkey) in April-August, both in the flowering and fruiting period, between 2005 and 2009 and a large number of herbarium speciemens in AEF, ANK, B, BM, CBB, E, FUH, G, GAZI, HUB, ISTE, ISTF, K, KNYA, LE, MO, VANF and W. The revision also includes macromorphological, anatomical, palynological, mericarp micromorphological, ecological, phytogeographical and numerical analysis. During the extensive field studies, the data on distribution and habitat, GPS coordinates, population sizes, the number of mature individuals, phenological and ecological features and threat factors were recorded. Numerous new localities, often far from the previously known distribution area, are reported.

According to the results, the research area has 59 taxa, 24 (40.7%) of which are endemic and the remaining 35 (59.3%) are non-endemic. East Anatolia includes 51 taxa, 22 (43.1%) of which are endemic, while Southeast Anatolia includes 25 taxa, 3 (12.0%) of which are endemic. The endemic taxa are distributed in three main areas within the study area. The first area covered by 12 endemic taxa is Divriği, Kemaliye and Erzincan. The second area covered by 12 endemic taxa is Sarız, Gürün and Malatya. The third area covered by 4 endemic taxa is Tercan, Erzurum and Ağrı. Distribution of the taxa according to phytogeogaphical regions are as follows: 50 taxa Irano-Turanian element with 85%, 4 taxa Mediterranean element with 7%, 2 taxa Euro-Siberian element with 3%, and 3 taxa unknown or multiregional with 5%.

According to the results of the taxonomic revision, one new species, S. siirtica, one new record, S. macrosiphon, and two species are re-evaluated as valid species, S. cerino- 310

pruinosa and S. pseudeuphratica, are described. S. ballsiana is rediscovered. The species was first collected in Gölbaşi (Malatya) in 1935, and was not collected again until 2008, when it was found in Gerger (Adıyaman).

Macromorphological characters that were considered to have taxonomically diagnostic value are investigated and their possible variations are evaluated. Habit, stem, leaf, inflorescence, bract, calyx, corolla and stamen properties of the genus are compared at both infrageneric and species level. The results showed that some characters such as shape of leaf, number of flowers in each verticillasters, length, shape and colour of calyces and corollas, calyx structure and stamen type are are of taxonomic significance.

The first comprehensive evaluation of the systematic value of anatomy (root, stem, leaf blade and petiole), palynology and mericarp micromorphology of Salvia is presented. The usefulness of anatomical properties of the vegetative organs in species identification and infrageneric delimitation is discussed. Significant variation, mainly useful at the infrageneric level, is found in row numbers and length of ray cells in the root, type of structure of the leaf blade, presence of hypodermis, presence of stomata of the upper and lower sides, shape of cross section of the petiole, shape and number of median vascular bundles. The other diagnostic characters, such as number of cell layers of collenchyma and cortex, vascular bundles in the stem, number of cell layers of palisade parenchyma, number of median and lateral vascular bundles in the midrib, size of midrib, size of petiole, number of lateral vascular bundles and number of cell layer of collenchyma in the petiole, are helpful in the identification of species. A key to the sections is given based on anatomy.

The variation in pollen morphological characteristics is discussed with respect to their potential systematic value. The main pollen characters, such as pollen size and shape, exine sculpturing type (reticulate-perforate, reticulate-granulate, bireticulate), the number of perforations, the number and size of secondary lumina, the presence of one or several central large lumina in the primary lumen, the shape of primary lumen and the

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ratio of the muri thickness are, are useful particularly in separation of the species studied. Several pollen characters are of restricted value for infrageneric classification. For example, the species of the section Hemisphace differs from the others in its smaller pollen grains. Moreover, S. viridis in sect. Horminum, S. staminea in sect. Plethiosphace and most of the species in sect. Aethiopis differ from the members of the other sections in having one or several large lumina in the middle of the primary lumen. A key to the taxa is given based on pollen morphology.

Major mericarp morphological characteristics of the taxa studied are compared and discussed on the basis of taxonomical concepts. Mericarp morphological characters are of taxonomical value. Mericarp size and shape, mericarp length/width ratio, surface sculpturing type (colliculate, reticulate, verrucate, rugose), abscission scar diameter and shape and exocarp cell shape, show significant variation among the taxa. Mericarp size and abscission scar diameter are mainly helpful at infrageneric level. For example, sect. Hymenosphace can be clearly differentiated by its large abscission scars from all the other sections except for sect. Salvia. Sect. Plethiosphace and Hemisphace are distinguished from sect. Hymenosphace and Salvia by their smaller mericarps. A key to the taxa is provided based on mericarp morphology.

Macromorphological (e.g. habit, leaf, calyx, corolla and stamen) and anatomical (e.g. root, leaf blade and petiole) properties of Salvia are useful at both infrageneric and species level, but pollen and mericarp micromorphological properties are mainly useful for species discrimination in the genus.

The taxa studied are found in a very wide variety of habitats reaching up to 3350 m elevation: limestone slopes, igneous slopes, calcareous slopes, serpentine slopes, rocky and stony slopes, steppe, gypsum areas, marly places, roadsides, cornfields, fallow fields, cultivated fields, vineyards, waste places, grassy places, dry meadows, macchie, bushy places, alpine and subalpine meadows, loamy hills, near permanent snowbeds, river shingle, Pinus, Quercus, Juniperus, Abies and Pyrus woodland and moist places in Fagus,

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Alnus, Corylus and in Picea forests. The flowering time of the taxa ranges from March and October.

The results of soil analysis of the samples obtained from suitable habitats of the taxa revealed that they grow in a variety of soils, such as such as clay, loam, sandy and silty soils in their habitats. These soil types are also composed of different proportions of clay (C), loam (L), sand (S) and silt (Si): clayish-loamy, loamy-sandy, sandy-loamy, sandy- clayish-loamy, silty-clayish and silty-clayish-loamy. Most taxa prefer sandy-loamy soils.

Based on the current geographical and ecological distribution, conservation status of the taxa is reassessed at regional, national and global levels using IUCN Red List categories and criteria. Critically Endangered (CR), Endangered (EN), Vulnerable (VU), Near Threatened (NT) and Least Concern (LC) categories are recognized. At the regional scale, the distribution is CR for 6 taxa (10%), EN for 9 taxa (15%), VU for 5 taxa (9%), NT for 10 taxa (17%) and LC for 29 taxa (49%). At the national scale, the distribution is CR for 5 taxa (9%), EN for 5 taxa (9%), VU for 7 taxa (12%), NT for 10 taxa (17%) and LC for 32 taxa (54%). At the global scale, the distribution is CR for 5 taxa (9%), EN for 3 taxa (5%), VU for 6 taxa (10%), NT for 10 taxa (17%) NT and LC for 35 taxa (59%).

The threatened taxa are under pressure from human activities such as overgrazing, constructions, land clearing and urbanization. Some significant measures are recommended for the conservation and management of the high number of endemic taxa under threat in the study area.

The infrageneric delimitation of Salvia is performed using multivariate analysis. As a result of the analysis, the phenogram presents seven major clusters showing sections. Identification keys to sections and species are given. Synonymy, type citation, amended and expanded descriptions, phenology, distribution and habitats in Turkey, general distribution outside Turkey, phtogeography, specimen citations on a grid square basis,

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some notes on taxonomy and photographs showing general appearance of the taxa are provided.

During the timeline of this thesis, about 30 papers and inter(national) presentations including new taxon, new records, rediscovery, numeric taxonomy, pollen, mericarp micromorphology, ecology and conservation status of Salvia were published, submitted prepared for submission.

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CURRUCULUM VITAE

PERSONAL INFORMATION

Surname, Name: Kahraman, Ahmet Nationality: Turkish (TC) Date and Place of Birth: 18 August 1980, Denizli Phone: +90 505 8036273 E-mail: [email protected], [email protected]

EDUCATION Degree Institution Year of Graduation MS Dumlupınar University 2004 BS Dumlupınar University 2002 High School Eşme High School, Uşak 1997

WORK EXPERIENCE Year Place Enrollment 2004-Present METU Research Assistant

FOREIGN LANGUAGES English

PUBLICATIONS Papers published in international journals 1. Kahraman A, Celep F & Doğan M. 2009. A new record for the Flora of Turkey: Salvia macrosiphon Boiss. (Labiatae). Turkish Journal of Botany 33: 53-55.

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2. Kahraman A, Celep F & Doğan M. 2009. Comparative morphology, anatomy and palynology of two Salvia L. species (Lamiaceae). Bangladesh Journal of Plant Taxonomy 16(1): 73-82.

3. Kahraman A, Celep F & Doğan M. 2009. Morphology, anatomy and palynology of Salvia indica L. (Labiatae). World Applied Sciences Journal 6(2): 289-296.

4. Celep F, Doğan M, Bagherpour S & Kahraman A. 2009. A new variety of Salvia sericeotomentosa (Lamiaceae) from South Anatolia, Turkey. Novon 19(4): 432-435.

5. Bagherpour S, Celep F, Doğan M & Kahraman, A. 2009. Rediscovery of Salvia freyniana Bornm. (Lamiaceae), A critically endangered species in Turkey. Bangladesh Journal of Botany 38(2): 189-201.

6. Kahraman A, Celep F & Dogan M. 2010. Anatomy, trichome morphology and palynology of Salvia chrysophylla Stapf (Lamiaceae). South African Journal of Botany 76: 187-195.

7. Kahraman A & Doğan M. 2010. Comparative study of Salvia limbata C.A. and S. palaestina Bentham (sect. Aethiopis Bentham, Labiatae) from East Anatolia, Turkey. Acta Botanica Croatica 69(1): 47-64.

8. Kahraman A, Celep F, Doğan M & Bagherpour S. 2010. A taxonomic revision of Salvia euphratica sensu lato and its closely related species (sect. Hymenosphace, Lamiaceae) by using multivariety analysis. Turkish Journal of Botany 34: 261-276.

9. Kahraman A, Celep F & Doğan M. 2010. Morphology, anatomy, palynology and nutlet micromorphology of Salvia macrochlamys (Labiatae). Biologia 65(2): 219-227.

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10. Bagherpour S, Kahraman A, Doğan M, Celep F, Başer B & Pehlivan S. 2010. Anatomical and micromorphological characteristics of Salvia vermifolia Hedge & Hub.- Mor. (Sect. Aethiopis Bentham, Lamiaceae). Central European Journal of Biology 5(6): 872-879.

11. Celep F, Doğan M & Kahraman A. 2010. Re-evaluated conservation status of Salvia L. (Sage) in Turkey I: The Mediterranean and the Aegean geographic regions. Turkish Journal of Botany 34: 201-214.

12. Şenol FZ., Orhan İ, Celep F, Kahraman A, Doğan M, Yılmaz G & Şener B. 2010. Survey of 55 Turkish Salvia taxa for their acetylcholinesterase inhibitory and antioxidant activities. Food Chemistry 120: 34-43.

13. Kahraman A, Celep F, Guerin GR & Doğan M. 2011. Mericarp morphology and its systematic implications for the genus Salvia L. section Hymenosphace Bentham (Lamiaceae) in Turkey. Plant Systematics and Evolution 292: 33-39.

14. Kahraman A, Celep F & Doğan M. 2011. A new endangered species of Salvia L. (Lamiaceae) from Turkey. Nordic Journal of Botany (in press).

15. Kahraman A, Bagherpour S, Karabacak E, Doğan M, Doğan HM, Uysal İ & Celep F. Reassessment of conservation status of Salvia L. (Lamiaceae) in Turkey II. Turkish Journal of Botany (in press).

16. Büyükkartan HN, Kahraman A, Öölgeçen H, Doğan M & Karabacak E. 2011. Mericarp micromorphology and anatomy in Salvia hedgeana Dönmez, S. huberi Hedge and S. rosifolia Sm. (section Salvia Hedge, Lamiaceae). Acta Botanica Croatica 70(1): 65-80.

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17. Özler H, Pehlivan S, Kahraman A, Doğan M, Celep F, Yavru A, Başer B & Bagherpour S. 2011. Pollen morphology of the genus Salvia L. (Lamiaceae) in Turkey. Flora 206(4): 316-327.

18. Celep F, Kahraman A, Doğan M. 2011. Taxonomic notes for Salvia aucheri (Lamiaceae) from Southern Anaolia. Novon 21(1): 34-35.18.

19. Celep F, Kahraman A, Doğan M. 2011. A new taxo of the genus Salvia (Lamiaceae) from Turkey. Plant Ecology and Evolution 144(1): 111-114.

20. Şenol FS, Orhan IE, Erdem SA, Kartal M, Şener B, Kan Y, Celep F, Kahraman A & Doğan M. Evaluation of cholinesterase inhibitory and antioxidant activities of wild and cultivated samples of sage (Salvia fruticosa) by activity-guided fractionation. Journal of Medicinal Food (accepted).

Abstracts published in international and national conferences 1. Kahraman A, Doğan M, Celep F & Bagherpour S. Comparative anatomy and trichome morphology of some Salvia L. (Lamiaceae) taxa and their systematic significance. XIII. OPTIMA Conference, 22-26 March 2010, Belek, Antalya.

2. Celep F, Kahraman A, Bagherpour S, Karabacak E, Başer B, Doğan M, Akaydın G, Pehlivan S & Uysal İ. Revision of the genus Salvia L. (Lamiaceae) in Turkey. XIII. OPTIMA Conference, 22-26 March 2010, Belek, Antalya.

3. Bagherpour A, Kahraman A, Doğan M & Celep F. Anatomy, trichome and nutlet micromorphology of Salvia vermifolia Hedge & Hub.-Mor. (sect. Aethiopis, Lamiaceae) endemic to Turkey. XIII. OPTIMA Conference, 22-26 March 2010, Belek, Antalya.

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4. Kahraman A, Doğan, M, Celep F, Bagherpour S & Akaydın G. Rare non-endemic species of the genus Salvia L. (Lamiaceae) in Turkey. International Symposium on Biology of Rare and Endemic Plant Species (BIORARE), 26-29 May 2010, Fethiye, Muğla.

5. Celep F, Kahraman A, Bagherpour S, Karabacak E, Doğan M & Akaydın G. Conservation status of rare endemic species of Salvia L. (Lamiaceae) in Turkey. International Symposium on Biology of Rare and Endemic Plant Species (BIORARE), 26-29 May 2010, Fethiye, Muğla.

6. Çetiner Ç, Kaya Z, Dizkırıcı A, Kahraman A, Celep F & Doğan M. Determination of phylogenetic relationships within Salvia euphratica complex & its closely related species by using internal transcribed spacer (its) region of nuclear ribosomal DNA. International Symposium on Biology of Rare and Endemic Plant Species (BIORARE), 26-29 May 2010, Fethiye, Muğla.

7. Dizkırıcı A, Kaya Z, Çetiner Ç, Celep F, Kahraman A & Doğan M. Phylogenetic relationships between some taxa of Salvia genus by usİng cpDNA regİon from trnT (UGU) to trnF (GAA). International Symposium on Biology of Rare and Endemic Plant Species (BIORARE), 26-29 May 2010, Fethiye, Muğla.

8. Demirci B, Temel HE, Demirci F, Celep F, Kahraman A, Dogan M, Baser KHC. The chemical caracterization of some salvia essential oil and their inhibitory effects on acetylcholinesterase and butrylcholinesterase. 41st International Symposium on Essential Oils. 5-8 September 2010, Wroclaw, Poland.

9. Şenol FS, Orhan İ, Yılmaz G, Celep F, Kahraman A, Doğan M & Şener B. Survey of various Turkish Salvia taxa for their acetylcholinesterase inhibitory and antioxidant activities, 9th International Symposium on Pharmaceutical Sciences (ISOPS-9), 23-26 June, 2009, Ankara, Turkey,

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10. Doğan M, Akaydın G, Celep F, Bagherpour S, Kahraman A & Karabacak E. Infrageneric delimitation of Salvia L. (Labiateae) in Turkey. International Symposium 7th Plant Life of Southwest Asia, 25-29 June 2007, Eskişehir.

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