A NUMERICAL TAXONOMIC STUDY ON THE FAMILY PLUMBAGINACEAE IN TURKEY

Transcription

1 A NUMERICAL TAXONOMIC STUDY ON THE FAMILY PLUMBAGINACEAE IN TURKEY A THESIS SUBMITTED TO THE GRADUATE SCHOOL OF NATURAL AND APPLIED SCIENCES OF MIDDLE EAST TECHNICAL UNIVERSITY BY JELENA ERDAL IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN BIOLOGY SEPTEMBER 2015

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3 Approval of the thesis: A NUMERICAL TAXONOMIC STUDY ON THE FAMILY PLUMBAGINACEAE IN TURKEY submitted by JELENA ERDAL in partial fulfillment of the requirements for the degree of Master of Science in Biology Department, Middle East Technical University by, Prof. Dr. Gülbin Dural Ünver Dean, Graduate School of Natural and Applied Sciences Prof. Dr. Orhan Adalı Head of Department, Biology Prof. Dr. Musa Doğan Supervisor, Biology Dept., METU Examining Committee Members: Prof. Dr. Zeki Kaya Biology Dept., METU Prof. Dr. Musa Doğan Biology Dept., METU Prof. Dr. Osman Ketenoğlu Biology Dept., Ankara University Prof. Dr. Hayri Duman Biology Dept., Gazi University Assoc. Prof. Dr. Nursen Çoruh Chemistry Dept., METU Date:

4 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: JELENA ERDAL Signature : iv

5 ABSTRACT A NUMERICAL TAXONOMIC STUDY ON THE FAMILY PLUMBAGINACEAE IN TURKEY Erdal, Jelena M.S., Department of Biology Supervisor: Prof. Dr. Musa Doğan September 2015, 119 pages The aim of this study was to make evaluation of existing structure of Plumbaginaceae in Turkey in order to see infrafamilial grouping as well as infrageneric arrangement of the big genera (Limonium and Acantholimon). For this purpose, numerical taxonomic method and clustering analysis were used. Also, this was the first time that all the genera of Plumbaginaceae found in Turkey were examined together, within the same study. Herbarium specimens of 61 species were evaluated on the base of 57 morphological characters to construct data matrix. Data matrix was analysed by Multi Variate Statistical Package (MVSP) software. First, Unweighted Pair Group Method with Arithmetic Mean (UPGMA) analysis was used to construct phenograms. Later, Principal Component Analysis (PCA) was performed to indicate which characters were the most significant for the outcome of the analysis. Obtained results were discussed, and compared with previous findings. According to UPGMA phenogram, the genera in our study were well separated from one another, exhibiting quite natural structure. Plumbago and Ceratostigma formed one cluster that correspond to the Plumbaginoideae subfamily. The other cluster analogous to subfamily Staticoidea was formed by Goniolimon, v

6 Limoniopsis, Armeria, Psylliostachys, Acantholimon and Limonium. Within genera Acantholimon and Limonium clusters equivalent to the main sections were observed. Furthermore, it was seen that our findings were largely supported the taxonomic treatments given in the Flora of Turkey and the subsequent revisional studies. Eventually, a new generic identification key for the Plumbaginaceae in Turkey including the newly recorded genus Psylliostachys is provided. Keywords: Plumbaginaceae, numerical taxonomy, cluster analysis, Turkey vi

7 ÖZ TÜRKİYE DEKİ PLUMBAGİNACEAE FAMİLYASI ÜZERİNDE BİR NÜMERİK TAXONOMİK ÇALIŞMA Erdal, Jelena Yüksek Lisans, Biyoloji Bölümü Tez Yöneticisi: Prof. Dr. Musa Doğan Eylül 2015, 119 sayfa Bu çalışmanın amacı, Türkiye deki Plumbaginaceae familyasındaki mevcut familya içi sınıflandırmanın yanısıra büyük cinslere (Limonium ve Acantholimon) ait cins içi sınıflandırmayı ortaya koymaktır. Bu amaçla, nümerik taksonomi metodu ve kümeleme analizi kullanılmıştır. Ayrıca, bu çalışma Türkiye de bulunan tüm Plumbaginaceae cinslerini kapsayacak şekilde ilk kez aynı çalışma içerisinde ele alınmıştır. Data matriksi oluşturmak için 61 türe ait herbaryum örnekleri, 57 morfolojik karakter tabanında değerlendirilmiştir. Veri matrisi MVSP yazılımı kullanılarak analiz edilmiştir. İlk olarak, UPGMA analizi fenogramlar oluşturmak için kullanılmıştır. Daha sonra, analizin en belirgin karakterlerinin hangileri olduğunu belirlemek için PCA kullanılmıştır. Elde edilen sonuçlar tartışılmış ve daha önceki bulgular ile karşılaştırılmıştır. UPGMA fenogramına göre çalışmadaki cinsler birbirinden açık bir şekilde ayrılarak neredeyse doğal bir yapı göstermektedir. Plumbago ve Ceratostigma, Plumbaginoideae alt familyasına karşılık gelen tek bir kümelenme oluşturmuştur. Staticoidea alt familyasına ait diğer kümelenme analojisi Goniolimon, Limoniopsis, vii

8 Armeria, Psylliostachys, Acantholimon ve Limonium tarafından oluşturulmuştur. Acantholimon ve Limonium cinslerinde cins içi sınıflandırma daha önceki değerlendirmeleri desteklemektedir. Ayrıca, bulgularımızın Türkiye Flora sında verilen taksonomik düzenlemelere ve sonrasındaki yeniden düzenleme çalışmalarıyla büyük ölçüde uyumlu olduğu görülmüştür. Sonuç olarak, Plumbaginaceae familyasına ilişkin yeni bir cins anahtarı yakın zamanda Türkiye den kaydedilmiş olan Psylliostschys cinsinide içerecek şekilde verilmiştir. Anahtar Kelimeler: Plumbaginaceae, nümerik taksonomi, kümelenme analizi, Türkiye viii

9 To my beloved husband and my families ix

10 ACKNOWLEDGEMENTS I would like to express my deepest gratitude to my supervisor Prof. Dr. Musa Doğan for his guidance, advices, encouragements, patience, efforst and suggestions throughout the research. It is great honor and pleasure for me to be his student. I appreciate for the patience and efforts of my thesis examining committee members: Prof. Dr. Zeki Kaya, Prof. Dr. Osman Ketenoğlu, Prof. Dr. Hayri Duman and Assoc. Prof. Dr. Nursen Çoruh, during reading and commenting my thesis study. I would also like to thank to my former supervisor Assoc. Prof. Dr. Mayda Gürsel for her advices, assistance and personal attention. I would like to express my special thanks to my friend Deniz Tiambeng for her help, advices and kindness. I present my thanks to the Graduate School of Natural and Applied Sciences of Middle East Technical University for supporting this study with grants for graduate students. At last, I owe great gratitude to my husband Serkan Erdal for his unconditional support, encouragement, help, attention, understanding, and care. Also, I would like to thank to my families, Erdal and Obradović, for their kindness, efforts, concern and support. x

11 TABLE OF CONTENTS ABSTRACT... v ÖZ vii ACKNOWLEDGEMENTS... x TABLE OF CONTENTS... xi LIST OF TABLES... xiii LIST OF FIGURES... xiv ABBREVIATIONS... xvi CHAPTERS 1. INTRODUCTION Systematic Studies on Plumbaginaceae Juss General Characteristics of Plumbaginaceae Juss The Genus Plumbago L The Genus Limonium Mill The Genus Goniolimon Boiss The Genus Limoniopsis Lincz The Genus Acantholimon Bioss The Genus Armeria Willd The Genus Ceratostigma Bunge The Genus Psylliostachys (Jaub. & Spach.) Nevski Ecological Features of the Plumbaginaceae Juss. in Turkey Phytogeographic distribution of the Plumbaginaceae in Turkey Habitat preferences of the Plumbaginaceae in Turkey Endemism Conservation Status of Plumbaginaceae in Turkey Numerical Taxonomy Objectives of the Study MATERIAL AND METHOD Material of the Study Method of the Study...35 xi

12 Selection of Operational Taxonomic Units (OTU s) Selection of characters and coding of character states Construction of data matrix and its use The comparison of character states RESULT AND DISCUSSION Result Result of the Cluster Analysis and UPGMA phenogram Result of the Principal Component Analysis Infrafamilial Grouping of the Plumbaginaceae Juss. in Turkey CONCLUSIONS REFERENCES APPENDICES A: A NEW IDENTIFICATION KEY TO THE GENERA OF PLUMBAGINACEAE IN TURKEY B: THE LIST OF THE OPERATIONAL TAXONOMIC UNITS C: CHARACTERS VS OTUs DATA MATRIX xii

13 LIST OF TABLES TABLES Table 1: The list of Plumbaginaceae native to Turkey and their general ecological information...18 Table 2: The list of Plumbaginaceae that are present in Turkey as ornamental plants and their general ecological information...29 Table 3: Character table...37 Table 4: Eigenvalues, percentages and cumulative percentages (Eigenvalues >1)...49 Table 5: The summary of the highest character loadings at the first 6 axis...51 Table 6: The most significant diagnostic features according to PCA.52 Table B1: List of Operational Taxonomic Units and general information about specimens...98 Table C1: Data matrix xiii

14 LIST OF FIGURES FIGURES Figure 1: Plumbago europaea L Figure 2: Plumbago capensis Thunb Figure 3: Limonium bellidifolium Dumort... 6 Figure 4: Goniolimon Boiss Figure 5: Limoniopsis owerinii (Boiss.) Lincz... 9 Figure 6: Acantholimon glumaceum (Jaub. & Spach) Boiss Figure 7: Armeria maritima Willd Figure 8: Ceratostigma plumbaginoides Bunge Figure 9: Psylliostachys (Jaub. & Spach.) Nevski Figure 10: Phytogeographic regions of Turkey Figure 11: Acantholimon avanosicum...33 Figure 12: UPGMA phenogram: Plumbaginaceae in Turkey. 44 Figure 13: UPGMA phenogram: Infrafamilial grouping of the Plumbaginaceae in Turkey Figure 14: UPGMA phenogram: Infrageneric grouping of the Acantholimon Figure 15: UPGMA phenogram: Infrageneric grouping of the Limonium Figure 16: Principal component analysis of the Plumbaginaceae in Turkey with 2 axis...50 Figure 17: Ceratostigma plumbaginoides Figure 18: Plumbago europaea Figure 19: Plumbago capensis Figure 20: Psylliostachys spicata Figure 21: Goniolimon incanum xiv

15 Figure 22: Limoniopsis davisii...58 Figure 23: Limonium sinuatum section Pteroclados...60 Figure 24: Limonium echioides, section Schizyhymenium...62 Figure 25: Limonium lilacinum, section Sphaerostachys...63 Figure 26: Limonium gmelini, section Limonium...64 Figure 27: Limonium graecum, section Limonium...65 Figure 28: Limonium anatolicum, section Sarcophyllum...65 Figure 29: Limonium bellidifolium, section Limonium...67 Figure 30: Limonium gueneri, section Limonium...68 Figure 31: Limonium smithii, section Limonium...69 Figure 32: Limonium vanense, section Limonium...70 Figure 33: Armeria trojana...71 Figure 34: Armeria cariensis var. cariensis...72 Figure 35: Armeria cariensis var. rumelica...72 Figure 36: Geographic distribution of A.cariensis var. cariensis and A. cariensis var. rumelica in Turkey...73 Figure 37: Acantholimon bracteatum, section Acantholimon...74 Figure 38: Acantholimon quinquelobum, section Tragacanthina...75 Figure 39: Acantholimon laxiflorum, section Staticopsis...76 Figure 40: Acantholimon köycegizicum, section Staticopsis...77 Figure 41: Acantolimon calvertii, section Staticopsis...78 Figure 42: Acantolimon caryophyllaceum, section Staticopsis...78 Figure 43: Acantolimon ulicinum, section Staticopsis...79 xv

16 ABBREVIATIONS A. Huet... Pavillon, Alfred Huet du APG II system.. Angiosperm Phylogeny Group II system Berthel.... Berthelot, Sabin Boiss.. Boissier, François de Sauvages de Lacroix Brongn. Brongniart, Adolphe Theodore cm.... centimeter C... Central CR... Critically Endangered Dumort Dumortier, Barthélemy Charles Joseph E East (E)... Royal Botanic Garden, Edinburgh EN.. Endangered et al. et alii (and others) Fourr.... Fourreau, JulesPierre Griseb... Grisebach, August Heinrich Rudolf Hausskn.... Haussknecht, Heinrich Carl Heldr.... Heldreich, Theodor Heinrich Hermann von IUCN International Union for Conservation of Nature and Natural Resources Jaub.. Jaubert, Hippolyte Francois Juss Jussieu, Antoine Laurent de K. Koch.. Koch, Karl Heinrich Emil Kusn.. Kusnezow, Ivan V. L... Linnaeus, Carolus Lam.... Lamarck, Jean-Baptiste Lestib.... Lestiboudois, Gaspard Thémistocle Lincz Linczevski, Igor Alexandrovich LC.. Least Concerned Lindl... Lindley, John xvi

17 m. meter Mill.. Miller, Philip mm millimeter MVSP... Multi Variate Statistical Package N North NT. Near Threatened Species OTU... Operational Taxonomic Unit PCA Principal Component Analysis Poir.... Poiret, Jean Louis Marie Rech. Rechinger, Karl Reichenb Reichenbach, Heinrich Gustav S... South Sauv... Sauvage, Charles Philippe Félix Sect.... section Sint... Sintenis, Paul Ernst Emil Sosn.. Sosnowsky, Dmitrii Ivanovich sp. nov.... species nova (new species) stat. nov. status novus (new status) subsp.... subspecies syn.. synonym Takht.... Takhtajan, Armen Thunb Thunberg, Carl Peter UPGMA Unweighted Pair Group Method with Arithmetic Mean var variety Viv.. Viviani, Domenico VU... Vulnerable W.. West Willd..... Willdenow, Carl Ludwig Zimmerm..... Zimmerman, Walter Max xvii

18 xviii

19 CHAPTER 1 INTRODUCTION 1.1. Systematic Studies on Plumbaginaceae Juss. A detailed worldwide revisional study of Plumbaginaceaae as a whole has not been done so far. Most of the studies in past were done in the regional level, as a part of the floristic studies. The main focus of the research was usually on the biggest genera Acantholimon Boiss. and Limonium Mill. The first revision of the Plumbaginaceae was done by Boissier in his 4 th volume of Flora Orientalis, in which he described 7 genera and 121 species (Boissier, 1879). Flora of USSR, 18 th volume, accepted 11 genera and 131 species (Komarov, 1967). In the 3 rd volume of Flora Europaea, 8 genera and 146 species were recognized (Tutin et al, 1972). Flora Iranica includes, 8 genera and 192 species (Rechinger and Schiman-Czeika, 1974). In the 15 th volume of Flora of China, 7 genera and 46 species were accepted (Wu and Raven, 1996). The major contribution in studying and describing Plumbaginaceae in Turkey was made in P.H. Davis, 7 th volume of Flora of Turkey and the East Aegean Islands, revised by M.H. Bokhari and J.R. Edmonsdon. In this flora, 6 genera were accepted, namely: Plumbago L., Limonium Mill., Goniolimon Boiss., Limoniopsis Lincz., Acantholimon Boiss. and Armeria Willd. In this family they recognized 52 species within those genera. (Davis et al., 1982, 1988). Also, Bokhari had made a taxonomic study on the South West Asian Plumbaginaceae. Beside revision and description of new taxa, he also recognized the taxonomic values of stigma, pollen types and anatomical features (Bokhari, 1970). The most recent revisional study of the Plumbaginaceae in Turkey was conducted by Doğan and Akaydın. In six years, they carried out extensive field 1

20 studies, gathered specimen all over Turkey and examined material housed in various herbaria in Turkey, as well as abroad. Specimen were studied for assembling morphological, anatomical, palynological, ecological and biogeographic information. The study included the genera: Plumbago L., Limonium Mill., Goniolimon Boiss., Limoniopsis Lincz. and Armeria Willd. This project was preceded with profound revision of Acantholimon in Hence, the final result of those studies was 13 new species of Acantholimon, 2 new species of Limonium, and a few new subspecies and varieties. Also, the presence of Ceratostigma plumbaginoides Bunge, Plumbago capensis Thunb and Goniolimon besseranum (Shultes ex. Reichenb.) Kusn. as ornamental plants was recorded from Turkey (Doğan and Akaydın, 2003a, 2006) General Characteristics of Plumbaginaceae Juss. Plumbaginaceae Juss. was described first time by Antoine Laurent de Jussieu in 1789, in his book Genera Plantarum (Jussieu, 1789). It was the only family in the order Plumbaginales Lindl. However, APG II system (Angiosperm Phylogeny Group II system) of plant classification that was published in 2003 by the Angiosperm Phylogeny Group, placed Plumbaginaceae in the order Caryophyllales Perleb. Hence, the most recent systematic classification of the Plumbaginaceae family is given below: Kingdom: Plantae Subkingdom: Tracheobionta Division: Magnoliophyta Cronquist, Takht. & Zimmerm. ex Reveal Class: Magnoliopsida Brongn. Subclass: Caryophyllidae Takht. Order: Caryophyllales Perleb Family: Plumbaginaceae Juss. Common name of the Plumbaginaceae is leadworts, from the Latin words plumbum ("lead") and agere ("to resemble"). Family includes 27 genera and species (Simpson, 2010). Based on morphological and chemical features this family was divided in two subfamilies Plumbaginoideae and Staticoideae 2

21 (Kubitzki, 1993; Simpson, 2010). This was also supported by phylogenetic studies using plastid DNA sequences (Moharrek et al., 2014). The genera of Plumbaginaceae are as follows: Acantholimon, Aegialitis, Afrolimon, Armeria, Bakerolimon, Bamiania, Bukiniczia, Cephalorrhizum, Ceratostigma, Chaetolimon, Dictyolimon, Dyerophytum, Eremolimon, Ghaznianthus, Gladiolimon, Goniolimon, Ikonnikovia, Limoniastrum, Limoniopsis, Limonium, Muellerolimon, Neogontscharovia, Plumbagella, Plumbago, Popoviolimon, Psylliostachys, Vasssilczenkoa (Kubitzki, 1993). Genus Myriolimon that was previously treated as a section Myriolepis Boiss. in Limonium, was segregated from it and recognized as a genus (Lledó, et al, 2003, 2005). Thus, currently family has 28 genera. The taxa of Plumbaginaceae are mostly perennial (with few exceptions of annual). They are in a form of shrubs, subshrubs, lianas or herbs. The leaves are alternately arranged, in basal rosettes, or in fascicular rosettes on annual shoots below scapes. Foliage is simple with entire margin, rarely pinnatifid. Usually leaves are with short or without petiole. Chalk glands that exude calcium salts and water for neutralizing salt from ground, are often present on the leaves and stem. Inflorescence is spicate, paniculate, subcapitate or capitate. Flowers are bisexual, actinomorphic, mostly arranged into bracteate spikelets. Calyx is gamosepalous. It can be tubular, subtubular, obconical or infundibular. It has 5 lobes and 5 or 10 ribs. In some taxa calyx is hairy and/ or glandular. Calyx lip is often hyalinated. Corolla consists of 5 petals which are completely free or connected at the base only. Exception are genera Plumbago and Psylliostachys with gamopetalous corolla. Flower has five stamens. Ovary has superior position, one loculus and it is connected to ovule with long stalk. The number of styles is 5 or 1. If it is 1 than it has five-lobbed stigma (genera Plumbago and Ceratostigma). Stigma is capitate to cylindrically filiform. Fruit has dry, membranous cover and contains a single seed. (Davis, et al., 1982, Kubitzki, 1993, Simpson, 2010). Even though Plumbaginaceae members seems to have cosmopolitan distribution, the center of diversity is in the Mediterranean area and central and 3

22 western parts of Asia. Members of this family especially prefer coastal habitats, inland salty area, steppes, or rocky highlands The Genus Plumbago L. Plumbago species are shrubs, herbs or lianas. Their leaves are alternate and simple. Inflorescence are terminal, capitate spikes, compound of spikelets with 3 bracts and 1 flower. Calyx is tubular, with dentate lip and large stalked glands. Corolla is gamopetalous. There is one glandular style and stigma is filiform (Davis et al, 1982). This genus is widely spread through the tropic regions, especially Africa. In the Flora of Tropical East Africa 8 indigenous species of Plumbago were recorded (Friss et al, 2012). Also, species of this genus are often cultivated as ornamentals. Figure 1: Plumbago europaea L. (edited from the Plant illustrations webpage; Sibthrop, J., Smith, J.E., Flora Graeca, vol. 2, p. 75, 1813) 4

23 Plumbago europaea L. is the only native species of Plumbago in Europe as well as in Turkey (Tutin et al, 1972, Davis et al, 1982). Another species that is common at the coast of Turkey is Plumbago capensis Thunb. (syn. Plumbago auriculata Lam., due to its auriculate leaves). This species is native to South Africa, but it is also cultivated as a garden plant, in all warm parts of the world. Occasional escapes of cultivation and naturalization are recorded, as well. Figure 2: Plumbago capensis Thunb. (edited from the Plant illustrations webpage; d Orbigny, C.V.D., Dictionnaire universel d histoire naturelle, plates vol. 3, t. 44, Dicotylédones 13, ) The Genus Limonium Mill. Plants from Limonium genus are subshrubs or herbs, mostly perennieal with few exceptions of annuals. Leaves are usually arranged in basal rosettes. In some of the species leaves dry up before the end of the flowering time. Scapes are often with 5

24 paniculate or subcorymbose branching, and may have sterile branches, as well. Inflorescence is spicate, spikes consist of many spikelets, with 1 up to (8-12) florets. Calyx is infundibuliform, obconical or tubular, with hyaline lip. Stigma is cylindrically filiform (Davis et al, 1982, Kubitzki, 1993). The very center of Limonium diversity is in the Mediterranean region, but its species might be found in other parts of the world with Mediterranean-like climate. Limonium species prefer coastal habitats or salty marches, as well as inland salty areas. That is where their common name Sea Lavenders came from. Figure 3: Limonium bellidifolium Dumort (edited from the Plant illustrations webpage; Sibthrop, J., Smith, J.E., Flora Graeca, vol. 3: p. 90, t. 295, 1819) This is one of the largest genus of Plumbaginaceae, roughly it has around 350 species (Kubitzki, 1993). Also, it is quite complex genus and there are constant changes within it. Since new species are being described and some of the species are being segregated from Limonium. Even though the first detailed study of Limonium was done by Boissier in 1848, it was segregated from genus Statice L. later, by 6

25 Miller. As previously mentioned, variety of species within this genus is very wide and complex that makes establishing firm and objective sectional separation quite hard. For example, in the flora of U.S.S.R. 32 species are separated into 5 sections: Pteroclados Sauv. & Vindt, Platyhymenium Boiss., Limonium Boiss., Sarcophyllum (Boiss.) Lincz. (this section was previously considered as subsection of the section Limonium by Boissier) and new section Siphonocalyx Lincz. (Komarov, 1967). Separation is mostly done on account of scape and calyx features. Flora of Iran contains almost same organization of sections with omission of Siphonocalyx and establishment of new section Nephrophylum Rech. (Rechinger and Schiman-Czeika, 1974). Approach to intrageneric structure of Limonium in Flora Europea is different in a sense that all species are divided into 3 subgenera by: Pteroclados, Myriolepis (previously it was one of the section of the Statice that Boissier had described) and Limonium (Tutin et al, 1972). The latest change in structure of Limonium genus is that subgenus Myriolepis is segregated and raised into generic level, under the name Myriolimon (Lledo et al, 2003, 2005). In the Flora of Turkey, 17 species of the Limonium had been described and splitted into 5 sections, as: Pteroclados, Limonium, Sarcophyllum, Sphaerostachys and Schizyhymenium. The last two sections were established by Bokhari (Davis et al, 1982). Later in the supplement volume of the Flora of Turkey two new species were added, as Limonium vanense Kit Tan & Sorger and Limonium caspium (Willd.) Gams (Davis et al, 1988). Recently, two more species were described and published as Limonium smithii Akaydın (Akaydın, 2007) and Limonium gueneri Doğan, Duman & Akaydın (Doğan et al, 2008). Beside these, in the METU herbarium there are specimens named as Limonium davisii, Limonium didimense and Limonium marmarisense that are suspected to be a new species. Those specimens are included in this study in order to compare them with other species and to determine their potential placement within the sections of Limonium The Genus Goniolimon Boiss. All the species of Goniolmon are herbs, with short woody caudex and leaves in basal rosettes. Scapes are winged, panicled or sybcorymbose. Calyx is 7

26 infundıbular, plicate. Styles are with long hairs at lower part. Stigma is depressed capitate (Davis et al, 1982). Distribution range of the genus spreads from the North Africa and east Mediterranean till to China and Mongolia. The highest number of species is described from Europe and Russia. Flora of U.S.S.R mentions 12 species, divided in two sections, Unicuspidaria and Tricuspidaria according to the number of cusps on the 2 nd inner bract. Within section Tricuspidaria additional subsection and series are also described (Komarov, 1967). In the European flora 11 species are present, without sectional differentiation (Tutin et al, 1972). Flora of China contains only 4 Goniolimon species (Wu and Raven, 1996). Figure 4: Goniolimon Boiss. (edited from the Muséum national d histoire naturelle webpage, Collection des vélins du Muséum national d histoire naturelle, vol.16: t.78) In Turkey only one Goniolimon species is present. This species was described in the Flora of Turkey as a Goniolimon collinum (Griseb.) Boiss. (Davis et al, 1982). However, in later revision this species was referred as Goniolimon incanum (L.) Hepper (Davis et al, 1988). Doğan and Akaydın (2006) in their revision of 8

27 Plumbaginaceae in Turkey, recorded presence of the Goniolimon besseranum, as a cultivated garden plant. Normally this species grows in east parts of Europe The Genus Limoniopsis Lincz. Plants with short woody caudex and leaves arranged in rosettes. Scape is panicled, spiklets are with remote spacing and 1-3 flowered. Calyx is subtubular. Stigma is capitate (Komarov, 1967). This genus has only two species. Limoniopsis owerini (Boiss.) Lincz. was described by Boissier as a Statice owerinii. Later Limoniopsis was segregated from Statice and established as a monotypic genus by Linczevski. It mostly grows at eastern Caucasus. In Turkey it is found around Kemaliye in Erzincan province. Limoniopsis davisii Bokhari, is endemic species of Van province in South East Anatolia (Davis et al, 1982). Figure 5: Limoniopsis owerinii (Boiss.) Lincz (edited from the Russian Federation Plant s Redlist webpage (Красная Книга Российской Федерации (Растения)) 9

28 The Genus Acantholimon Bioss. Subshrubs or shrublets with cushion formation. Leaves are rigid, linear, triquetrous and pungent, homomorphic or heteomorphic. Scape is with scales. Inflorescense is in a form of branched or not branched spikes, in some cases capitate or paniculate. Spiklets are 1-flowered or 2-5 (6) flowered, with 3 or more bracts. Calyx infundibular, with hyaline lip. Stigma is oblong-capitate. (Davis et al, 1982). The genus contains about 165 species (Kubitzki, 1993). Its distribution range stretches form the southern Greece, over Turkey, Iran, Afghanistan, to the west Tibet. Most of the species are confined to mountainous regions, usually in dry, stony soils and exposed rocky slopes (Komarov, 1967). The rate of endemism in this genus is very high. Most of the species are local endemics, with narrow distribution range. After Limonium, this is the second most species rich genus in Plumbaginaceae. Same as Limonium, Acantholimon genus is very complex and shows great variation in characters between species, which makes grouping within genus and sectional delimitation quite complicated. Acantholimon was first known as subgenus Armeriastrum Jaub. & Spach in Statice L. genus (Lledo et al, 2003). Boissier, was the first who described the Acantholimon, as a separate genus and provided its earliest classification, in He divided genus into following sections: Armeriopsis, Staticopsis and Glumaria (Moharrek et al, 2014). Russian botanist, Bunge had published in 1872 a very first detail revision of the genus and organized 83 known species into 7 sections: Armeriopsis, Staticopsis, Glumaria, Acmostegia, Cymaria, Pterostegia and Tragacanthina (Doğan and Akaydın, 2003a). This Bunges classification was revised by Boissier in by addition of the section Pulvinaria. Mobayen recognized 112 species and made a change in this classification by removing Cymaria and Pulvinaria sections and describing new Dracogina section (Moharrek et al, 2014). Linczevski, had accepted sections: Armeriopsis, Glumaria, Pterostegia, Staticopsis and Tragacanthina, and added section Gontscharovia, for the grouping of around 70 species included in flora of U.S.S.R. (Komarov, 1967). Flora of Iran contains 153 Acantholimon species, separated into 15 sections (8 old and 7 new). Old sections 10

29 established by previously mentioned authors are: Acmostegia, Acantholimon, Dracogina, Glumaria, Gontscharovia, Pterostegia, Staticopsis and Tragacanthina, New sections that had been described, are: Bromeliopsis, Inermia, Physostegia, Platystegia, Poicilocephala, Schizostegia and Stereophylla (Reichinger and Schiman-Czeika, 1974). This classification had been revised by Assadi in 2005, by reducing number of sections into 7. He accpeted old ones: Acmostegia, Acantholimon, Platystegia, Pterostegia, Staticopsis, Tragacanthina and added one new, Microstegia (Moharrek et al, 2014). Figure 6: Acantholimon glumaceum (Jaub. & Spach) Boiss. (edited from the Plant illustrations webpage; Houtte, L. van, Flore des serres et des jardin de l Europe, vol.7: t. 677, 1852) In the Flora of Turkey 25 species had been described, along with 11 doubtfully recorded and 2 imperfectly known species. Those species are divided into 11

30 3 main sections: Acantholimon, Tragacanthina and Staticopsis. Within section Staticopsis there are 3 subsections: Caryophyllacea, Microcalycina and Androsacea. (Davis et al, 1982). Muvaffak and Doğan had described additional subsection Muvaffak and Doğan had described additional subsection Halophiliacea (Muvaffak et al, 2001). In a period of three years a very profound study was conducted by Doğan and Akaydın with aim to revise genus Acantholimon in Turkey. It included collecting specimen all around Turkey, examining existing specimens at different herbaria, including material cited in Flora of Turkey and East Aegean Islands, 7 th volume (Dogan and Akaydın, 2003a). Result of this study showed that there were 52 species, 10 subspecies, and 17 varieties in Turkey. From those 13 of the species were described for the first time: A.birandii (Dogan and Akaydın, 2001), A.avanosicum (Dogan and Akaydın, 2002a), A.karamanicum (Akaydın and Doğan, 2002), A.anatolicum (Dogan and Akaydın, 2002b), A.yildizelicum (Akaydın,, 2002), A.köycegizicum and A.göksunicum (Dogan and Akaydın, 2003c), A.turcicum (Dogan and Akaydın, 2003b), A.hoshapicum, A.artosense and A.bashkaleicum (Dogan and Akaydın, 2004), A.evrenii (Dogan and Akaydın, 2005), and A.ekimii (Doğan and Akaydın, 2007a). Species A.laxiflorum, was first described by Boissier. However, probably by mistake, Bokhari and Edmondson, labeled this species as a variety of the A.venustum, in the Flora of Turkey. Later, newest research findings showed that species is pretty different, both from A.venustum, and the rest of Acantholimon species (Doğan et al., 2003d). Presence of the species A.araxanum Bunge, A.hohenackeri (Jaub. &Spach) Boiss., A.latifolium Boiss. and A.lepturoides (Jaub. & Spach) Boiss. that were marked as a doubtfully recorded species in the Flora of Turkey, was confirmed, too. Also, species A.senangense, A.fominii and A.tragacanthinum, previously known from other countries, had been found to grow in Turkey, as well (Doğan and Akaydın, 2003a). As a part of this revision, 3 sections within Acantholimon genus were accepted: Acantholimon, Tragacanthina and Staticopsis. The species A.evrenii, seems to be the only representative of the section Glumaria Boiss. in Turkey (Dogan and Akaydın, 2005). Furthermore, Staticopsis section was reorganized and the following subsections are suggested: Robustea Dogan and Akaydın subsect. nov., Diantifoliea Dogan and Akaydın subsect. nov., Circinnatea Dogan and Akaydın subsect. nov., Androsacea Bunge, Caryophyllacea 12

31 Bunge (Doğan et al, 2007). As a final result of this study Synopsis of Turkish Acantholimon Boiss. (Plumbaginaceae) had been published (Doğan and Akaydın, 2007). In the following years there were two more publications about Acantholimon species. First species is A.doganii placed under section Staticopsis and subsection Androsacea (Bağcı et al, 2009). Second species is A.riyatguelii, included in section Staticopsis. Because of its specific morphology that differs from other species in this section, a new subsection Exacantha, was described (Yıldırım and Crespo, 2014) The Genus Armeria Willd. Armeria species are herbs, with branched woody root stock, and leafless scape. Leaves are linear, arranged in basal rosettes. Inflorescence is capitate, composed of cymose, bracteate, spikelets. Calyx is infundibular. Styles are hairy in lower third. Stigma is filiform (Davis et al, 1982). Figure 7: Armeria maritima Willd. (edited from the Plant illustrations webpage; Lindman, C.A.M., Bilder ur Nordens Flora, vol. 1: t. 140, ) 13

32 Armeria is distributed in north temperate regions, mostly in Europe, as well as some parts of Chile and Tierra del Fuego in South America. However, center of its diversity is west Mediterranean, especially Iberian Peninsula, where almost 60% of the species are present (Aguilar, 1999). This genus is taxonomically very complex. Since hybridization occurs often, it increases variation of diagnostic characters, and sometimes make boundaries between species hard for determining (Selvi, 2009). Genus was the first time described by Linnaeus as a part of Statice L. genus, as a single species Statice armeria. It was recognized as a separate genus by Willdenow (Yeşil et al, 2014). In European flora 43 species were accepted and divided in two sections, Armeria and Macrocentron, based on calyx features (Tutin et al, 1972). In Turkey there are two native species. Armeria trojana is a local endemic, known only from Mount Ida (Kaz Dağı) in Balıkesir province. Armeria cariensis, according to the Flora of Turkey, consist of two varieties Armeria cariensis var. cariensis and Armeria cariensis var. rumelica that was also supported by later revision (Dogan and Akaydın, 2006). Also, there are opinions that those varieties should be recognized as separate species Armeria cariensis and Armeria rumelica (Yeşil et al, 2014). Armeria maritima is not native to Turkey but it may occur as a garden plant The Genus Ceratostigma Bunge Plants of this genus are mostly shrubs and herbs. Leaves have ciliate margin. Inflorescences are terminal, subcapitate to capitate spikes, with 3 bracts and 1 flowered spikeletes. Calyx is tubular. Corolla is gamopetalous (Wu and Raven, 1996). Ceratostigma is native to China, with wide distribution range from temperate and tropical regions up to Himalayas. One species is also known from east Africa. Ceratostigma plumbaginoides Bunge, is grown worldwide as a garden plant. Also, there are records that it has been locally naturalized in North West France and North West Italy (Tutin et al, 1972). This species is present in Turkey as well, especially in gardens of coastal area. Due to that we decided to include this species in 14

33 the study, in order to compare and contrast it with native Turkish genera of the Plumbaginaceae. Figure 8: Ceratostigma plumbaginoides Bunge (edited from the Plant illustrations webpage; Curtis s Botanical Magazine, vol.76, t.4487, 1850, drawing by W.H. Fitch) The Genus Psylliostachys (Jaub. & Spach.) Nevski Psylliostachys consist of annual herbs. Leaves are simple or pinnatifid. Inflorescences are terminal and lateral, cylindrical spikes, composed of many spikelets, with 2 bracts and 2-4 florets. Calyx is infundibular. Corolla is gamopetalous. Glandular hairs are present on calyx, leaves and scape (Kubitzki, 1993; Tutin et al, 1972). Boissier considered Psylliostachys as a section of the genus Statice, and part of the Corolla gamopetala group (Lledó et al, 2003). Jaubert and Spach raised it at the level of subgenus. Finally, it was recognized as a genus by the Russian botanist Nevski (Rechinger and Schiman-Czeika, 1974). Genus has a 15

34 distribution range spreading from Russia and Central Asia, over Caucasus and Iran till to Syria and Palestine. Figure 9: Psylliostachys (Jaub. & Spach.) Nevski (edited from the Plant illustrations webpage; Curtis s Botanical Magazine, vol. 113, t. 6959, 1887, drawing by M.Smith) In the summer of 2014, research expedition made in the south east Turkey revealed the presence of the species Phylliostachys spicata, in the saline areas around the Siirt city. This was the first record of the Psylliostachys in Turkey Ecological Features of the Plumbaginaceae Juss. in Turkey Currently, 83 species, 8 subspecies and 23 varieties of Plumbaginaceae are known from Turkey. They are given in the Table 1. Also, there are 4 nonnative species present as ornamentals, given in a Table 2. Furthermore, both tables contain 16

35 additional information about species habitat preferences, elevation, geographic distribution range, endemism, phytogeography, and IUCN threat categories (Davis et al, 1982; IUCN, 2001; Doğan and Akaydın, 2003a, 2006). Each of those topics will be discussed in details, as follows. 17

36 18 Table 1: The list of Plumbaginaceae native to Turkey and their general ecological information Species Habitat type Elevation range (m) Geographic Range Phytogeography IUCN status 1. Acantholimon acerosum subsp. acerosum var. acerosum (Willd.) Boiss. subsp. acerosum var. parvifolium Bokhari Rocky igneous slopes, calcareous and sandy soils, steppe Rocky igneous slopes, calcareous and sandy soils, steppe Anatolia, except S. E., (Turkey), W. Syria, N. Iraq, Iran, Armenia Endemic from Göksun, Kahramanmaraş province, Turkey LC VU subsp. brachystachyum (Boiss.)Doğan & Akyadın Rocky igneous slopes, calcareous and sandy soils, steppe Endemic of W. and C. Anatolia, Turkey LC subsp. longibracteolorum Doğan & Akyadın Acantholimon anatolicum Doğan & Akaydın Acantholimon araxanum Bunge Acantholimon armenum var.armenum Boiss. & A.Huet. var. balansae Boiss. & A.Huet. Rocky igneous slopes, calcareous and sandy soils, steppe Gypseous, saline, soil on mountain slopes Sandy slopes, steppes Sandstone and calcareous slopes, steppes Sandstone and calcareous slopes, steppes Endemic of N.W. and W. Anatolia, Turkey Endemic of Nalıhan, Ankara province, Turkey E. Anatolia (Turkey), Armenia, Caucasus, Iran S. and W. Anatolia (Turkey), W. Syria, N. Iraq, Armenia S. and W. Anatolia (Turkey), Armenia LC CR NT LC LC

37 19 Table 1. (Continued) Acantholimon artosense Doğan & Akaydın Acantholimon avanosicum Doğan & Akaydın Acantholimon bashkaleicum Doğan & Akaydın Acantholimon birandii Doğan & Akaydın Acantholimon bracteatum (Girard) Boiss. Acantholimon caesareum Boiss. & Balansa Acantholimon calvertii var. calvertii Boiss. var. glabrum Doğan & Akaydın Acantholimon capitatum subsp.capitatum Sosn. subsp. sivasicum Doğan &Akaydın Calcareous mountain slopes 2500 Calcareous slopes and steppes 950 Sedimentary rocks and montane steppes 2100 Calcareous mountain slopes Calcareous, rocky slopes and montane steppes Volcanic slopes, steppes Calcareous and volcanic slopes, steppes Calcareous and volcanic slopes, steppes Calcareous and dry slopes Serpentine rocks, steppe Endemic of Artos Mountine, Van province, Turkey Endemic from Avanos, Nevşehir province, Turkey Endemic from Başkale, Van province, Turkey Endemic from Karaman province, Turkey S. E. Anatolia (Turkey), Armenia, N.W. Iran Endemic of C. and N. Anatolia, Turkey Endemic of E. Anatolia, Turkey Endemic of E. Anatolia, Turkey Endemic of E. Anatolia, Turkey Endemic of C. Anatolia, Turkey CR CR CR CR VU CR LC VU VU CR

38 20 Table 1. (Continued) 13. Acantholimon caryophyllaceum Boiss. Volcanic, calcareous slopes and steppes E. Anatolia (Turkey), N. Iraq, Armenia, N. and W. Iran LC 14. Acantholimon confertiflorum Bokhari Calcareous, clay slopes or steppes Endemic of C. Anatolia, Turkey EN 15. Acantholimon damassanum Mobayen Calcareous steppe, stony hills, dry slopes E. Anatolia (Turkey), Syria VU Acantholimon dianthifolium Bokhari Acantholimon doganii Y.Bağcı, Doğu& Akaydın Acantholimon ekimii Doğan & Akaydın Acantholimon evrenii Doğan & Akaydın Acantholimon fominii Kusn. Acantholimon glumaceum (Jaub.& Spach.) Boiss. Acantholimon göksunicum Doğan & Akaydın Rocky, high montane steppes Cedrus forest glades 1640 High mountain steppes 2300 Rocky mountain steppes Sandstone mountain slopes, steppes Limestone rocks, clay slopes, chalky hills and steppes Calcareous slopes, steppes 2100 S. E. Anatolia (Turkey), N. Iraq Endemic from Sarıveliler, Karaman province, Turkey Endemic from Akçay, Ağrı province, Turkey Endemic from Baltaşı, Elazığ province, Turkey E. Anatolia (Turkey),Caucasus, Iran, Azerbaijan E. Anatolia (Turkey), Armenia Endemic from Göksun, Kahramanmaraş province, Turkey VU CR CR CR VU VU CR

39 21 Table 1. (Continued) Acantholimon halophilum Bokhari Acantholimon hohenackerii (Jaub.&Spach)Boiss. Acantholimon hoshapicum Doğan & Akaydın Acantholimon huetii var. huetii Boiss. var. breviscapum Doğan & Akaydın Acantholimon hypochaerum Bokhari Acantholimon iconicum (Boiss.) Boiss. & Heldr. Acantholimon karamanicum Akaydın & Doğan Acantholimon kotschyi (Jaub. & Spach.) Boiss. Acantholimon köycegizicum Doğan & Akaydın Acantholimon latifolium Boiss. Salt steppes, saline clay hillocks, open Pinus nigra forest Rocky mountain slopes Dry montane steppes 2050 Calcareous slopes and steppes Calcareous slopes and steppes Volcanic, limestone or clay slopes Sedimentary and calcareous rocks, steppes Calcareous mountain slopes, Quercus glades, Astragalus steppe Salt steppes, sandy banks, calcareous soils Serpentine, rocky stream beds Calcareous steppe Endemic of C. Anatolia, Turkey E. Anatolia, (Turkey), Iran, Armenia Endemic from Hoşap, Van province,turkey Endemic of S. Anatolia, Turkey Endemic of S. Anatolia, Turkey Endemic of E. Anatolia, Turkey Endemic of N. and C. Anatolia, Turkey Endemic of Karaman province, Turkey Endemic of S. and C. Anatolia, Turkey Endemic from Köyceğiz, Muğla province, Turkey S. E. Anatolia (Turkey), N. Iraq, W. Iran Mediterranean CR VU CR CR CR EN EN CR LC CR CR

40 22 Table 1. (Continued) 33. Acantholimon laxiflorum Boiss.ex Bunge Serpentine, rocky stream beds 0-50 Endemic from Hatay province, Turkey Mediterranean CR 34. Acantholimon lepturoides (Jaub.& Spach) Boiss. Limestone and rocky slopes 1400 E. Anatolia (Turkey), Armenia NT Acantholimon libanoticum Boiss. Acantholimon lycaonicum var. lycaonicum (Boiss.) Boiss. &Heldr. var. cappadocicum Akaydın & Doğan Acantholimon multiflorum (Bokhari) Doğan &Akaydın Calcareous, mountain slopes, steppes Quercus glades, calcareous and serpentine slopes, sediment rocks, steppes Quercus glades, calcareous and serpentine slopes, sediment rocks, steppes Dry stony hills S. Anatolia (Turkey), Lebanon Endemic of S. and C. Anatolia, Turkey Endemic of C. Anatolia, Turkey Endemic of E. Anatolia, Turkey Mediterranean Mediterranean VU NT NT VU 38. Acantholimon parviflorum (Bokhari) Akaydın &Doğan Eroded hills and gypsum slopes Endemic C. Anatolia, Turkey CR 39. Acantholimon petraeum Boiss. & Hausskn. ex Bunge Volcanic rocks, montane steppes E. Anatolia (Turkey), Iraq NT Acantholimon petuniiflorum Mobayen Acantholimon puberulum subsp. puberulum Boiss. & Balansa Rocky, dry slopes and calcareous montane steppes Calcareous and serpentine slopes, arid steppes S. E. Anatolia (Turkey), N. Iraq S. and C. Anatolia (Turkey), N. Iraq CR LC

41 23 Table 1. (Continued) subsp. longiscapum (Bokhari) Doğan &Akaydın subsp. peroninii (Boiss.) Doğan &Akaydın Acantholimon quinquelobum var. quinquelobum Bunge var. curviflorum (Bunge) Doğan & Akaydın Acantholimon reflexifolium Bokhari Acantholimon riyatguelii Yıldırım & Crespo Acantholimon saxifragiforme (Hausskn. & Sint.) Bokhari Acantholimon senganense Bunge Acantholimon spirizianum Mobayen Acantholimon strigillosum Bokhari Calcareous slopes, Astragalus steppes Calcareous slopes, steppes Saline, dry, sandy hills Saline, dry, sandy hills Volcanic and limestone rocks, dry stony slopes Gypsum soils Calcareous montane steppes, rocky hills Volcanic rocks, montane steppes 2200 Calcareous slopes and steppes Eroded hills, serpentine rocks Endemic of C. Anatolia, Turkey Endemic of C. Anatolia, Turkey E. Anatolia (Turkey), Armenia, N. W. Iran E. Anatolia (Turkey), W. Iran Endemic of C. Anatolia, Turkey Endemic from Eskişehir province, Turkey Endemic of E. Anatolia Turkey E. Anatolia (Turkey), Syria, Iraq, Iran Endemic of E. Anatolia, Turkey Endemic of E. Anatolia, Turkey Mediterranean LC LC EN EN VU CR CR VU VU VU 49. Acantholimon tragacanthinum (Jaub. & Spach.) Boiss. Saline, dry, sandy hills 1200 E. Anatolia (Turkey), W. Iran NT

42 24 Table 1. (Continued) Acantholimon turcicum Doğan & Akaydın Acantholimon ulicinum var. ulicinum (Willd. ex Schultes) Boiss var. purpurascens (Bokhari) Bokhari & Edmondson var. creticum (Boiss.) Bokhari & Edmondson Acantholimon venustum var. venustum Boiss. var. assyriacum (Boiss.) Boiss. Acantholimon wiedemannii Bunge Acantholimon yıldıelicum Akaydın Armeria cariensis Boiss. Armeria rumelica Boiss Calcareous mountain scree Calcareous, serpentine or exposed rocky slopes Calcareous, serpentine or exposed rocky slopes Calcareous, serpentine or exposed rocky slopes Rocky volcanic slopes, limestone hills, mountain steppes, Pinus and mixed forests Rocky volcanic slopes, limestone hills, mountain steppes, Pinus and mixed forests Claystone slopes, steppes Calcareous and volcanic, dry mountain slopes and steppes 1700 Grasslands and montane steppes Endemic of E. Anatolia, Turkey Albania, Greece, W. and S. Anatolia (Turkey) Endemic of S. W. Anatolia Turkey C. and S. Anatolia (Turkey), Greece C., E. and S. Anatolia (Turkey), W. Syria, N. Iraq, W. Iran Endemic of E. Anatolia, Turkey Endemic of C. Anatolia, Turkey Endemic of Yıldızeli, Sivas province, Turkey N. E. Greece, N.W. and W. Turkey Grasslands and montane steppes S. E. Balkans Mediterranean Mediterranean Mediterranean Mediterranean Mediterranean CR LC CR LC LC VU VU CR EN EN

43 25 Table 1. (Continued) Armeria trojana Bokhari & Quézel Goniolimon incanum (L.) Hepper Limoniopsis davisii Bokhari Siliceous rocks, stony places Steppe, fallow fields, chalky and serpentine hills, stream beds, rocky sea shores Limestone mountain slopes Endemic of Kaz Mountain, Turkey From South Balkan to Syria Endemic of S. E. Anatolia, Turkey Mediterranean Mediterranean CR LC CR 60. Limoniopsis owerinii (Boiss.) Lincz. Rocky mountain slopes E. Anatolia (Turkey), E. Caucasus CR 61. Limonium anatolicum Hedge Continental saline areas Endemic of C. Anatolia, Turkey CR 62. Limonium angustifolium (Tausch) Turrill Seaside salt-flats, rocky limestone slopes 0-10 Mediterranean area Mediterranean EN 63. Limonium bellidifolium (Gouan) Dumort. Coastal and inland saline areas W. Turkey, S. and C. Europe, Crimea, C.Asia Euro-Siberian EN 64. Limonium caspium (Willd.) Gams Continental salty areas C. Anatolia, (Turkey), N.W. Iran, Azerbaijan, Siberia, C. Asia CR 65. Limonium effusum (Boiss.) Kuntze Salty and arid areas 0-10 Endemic of W. and S.W. Anatolia, Turkey Mediterranean EN 66. Limonium echioides (L.) Miller Muddy salt-flats, sandy areas near sea shore 0-5 W. and S. Anatolia (Turkey), Mediterranean Mediterranean VU

44 26 Table 1. (Continued) 67. Limonium globuliferum var. globuliferum (Boiss. & Heldr.) Kuntze Continental salty areas C. Anatolia (Turkey), Syrian Desert CR var. subglobosum Doğan & Akaydın Continental salty areas Endemic of C. Anatolia, Turkey CR 68. Limonium gmelinii (Willd.) Kuntze Sea shore, saline or gypsuous inland places Turkey, C. Europe, Balkans, Caucasus, Crimea, Iran, C. Asia Euro-Siberian LC 69. Limonium graecum var. graecum (Poir.) Rech. Sea shores, sands, volcanic and limestone rocks 0-5 W. Anatolia (Turkey), Greece, Aegean islands Mediterranean CR var.hyssopifolium (Girard) Bokhari Sea shores, sands, volcanic and limestone rocks 0-5 W. Anatolia (Turkey), S. Greece, Aegean islands Mediterranean CR 70. Limonium guenerii Doğan, Duman & Akaydın Steep coastal calcareous cliffs 20 Endemic from Kaş, Antalia province, Turkey Mediterranean CR 71. Limonium iconicum (Boiss. & Heldr.) Kuntze Continental salt steppes and carbonate rocks Endemic of C. Anatolia (Turkey) EN 72. Limonium lilacinum var. lilacinum (Boiss.& Balansa) Wagenitz Continental saline areas Endemic of C. Anatolia, Turkey CR

45 27 Table 1 (Continued) var. laxiflorum Doğan & Akaydın Continental saline areas Endemic of C. Anatolia, Turkey CR 73. Limonium meyeri (Boiss.) Kuntze Continental saline areas N.E. Anatolia (Turkey), Russia, Crimea, C. Asia Caucasus VU 74. Limonium ocymifolium (Poir.) Kuntze Sandy shores, calcareous and metamorphic littoral rocks 0-10 W. Anatolia (Turkey), Greece and Aegean Islands Mediterranean CR 75. Limonium pycnanthum (K.Koch) Kuntze Continental saline areas Endemic of C. Anatolia, Turkey CR 76. Limonium sieberi (Boiss.) Kuntze Sea shores and salt marches 0-5 S. and S.W. Anatolia (Turkey), S. Greece, Aegean islands Mediterranean EN 77. Limonium sinuatum (L.) Miller Coastal cliffs, beaches, sandy shores, pastures W. and S. Anatolia (Turkey), Mediterranean, Georgia Mediterranean EN 78. Limonium smithii Doğan & Akaydın Continental salty areas 1085 Endemic from Seyfe lake area, Kırşehir province,turkey CR 79. Limonium tamaricoides Bokhari Continental salty areas Endemic of C. Anatolia, Turkey CR

46 28 Table 1 (Continued) 80. Limonium vanense Kit Tan & Sorger Saline meadows and marches Endemic of E. Anatolia, Turkey CR 81. Limonium virgatum (Willd.) Fourr. 82. Plumbago europaea L. 83. Psylliostachys spicata (Willd.) Nevski Sea shores, maritime rocks, saline waste fields Dry gravelly slopes, limestone, igneous slopes, fallow fields and waste places Saline clays and sandy soils, in low foothills, on plains, along sea coast W. Turkey, Mediterranean area, W.Europe Mediterranean, Central Europe, Armenia, N. Iran, Azerbaijan, Iraq Russia, C. Asia, Caucasus, Iran, Syria, Palestine, S. E. Anatolia (Turkey) Mediterranean Euro-Siberian EN LC LC

47 29 Table 2: The list of Plumbaginaceae that are present in Turkey as ornamental plants and their general ecological information 1. Species Armeria maritima Willd. Habitat type Elevation range (m) Sea shore, sandy soils 0 Geographic Range Most of the Europe, except S. E. and most of the islands IUCN status LC 2. Ceratostigma plumbaginoides Bunge Rocky places, often in foothills Native to E. China, in other places present as ornamental plant LC 3. Goniolimon besseranum (Schult. ex Rchb.) Kusn. Steppes, on fine earth, chalky, and stony slopes, saline soils Native to area from N.E. Bulgaria to S. and C. Ukraine EN 4. Plumbago capensis Thunb. Streambeds, canyons in coastal scrubs Native to South Africa, in other places present as ornamental plant LC

48 Phytogeographic distribution of the Plumbaginaceae in Turkey Turkey is divided in 3 phytogeographic regions as a consequence of wide variation in climate and topography. These are the Euro-Siberian, the Mediterranean and the region (Davis at all, 1965). Figure 10: Phytogeographic regions of Turkey (edited from the Ecopangea webpage) The Euro-Siberian region includes the major part of the North Anatolia and the narrow area of the Black Sea coast of the European part of Turkey. Climate in this region is very humid and has high rainfall. Vegetation consists of deciduous forest that is gradually passing into conifer forest with increasment of elevation (Davis at all, 1965). Since this type of habitat is not suitable for the growth of Plumbaginaceae they are vary rare in this region and represented only with 2 Limonium and 1 Plumbago species. This is only 3.61 % of the current total number of Plumbaginaceae in Turkey. The Mediterranean region of Turkey consists of the Gallipoli peninsula and West and South Anatolia. The climate is typical Mediterranean, with dry, hot summers, with drought and mild, rainy winters, without frost. The dominant 30

49 vegetation are evergreen, mostly aromatic shrubs, called macchie, which are replaced by conifers at higher elevation (Davis at all, 1965). There are 5 species of Acantholimon, 3 species of Armeria, 1 species of Goniolimon, and 9 species of Limonium in this region, which is forming 21.69% of Turkish Plumbaginaceae. region is the largest from all and richest in species content. It covers Central and East Anatolia, and consist of mountains and highland plateaus in between. Climate in this region is typical continental, in some areas, gradually passing into semi desert. Summers are hot and dry. Winters are cold, with very low temperatures and high snow cover, which is the main source of precipitation. Rain is mostly present during spring. Also, open plateau areas are often very windy which increases dryness and affects tree growth. Temperature oscillation are high, both at annual and daily base. This kind of harsh climate is favoring vegetation dominant by herbaceous plants, especially grasses that form steppes and shrubs that occasionally may pass into rare, open conifer forest, in the areas where amount of moisture allows that. Vegetation growth period is during late spring and early summer. Except at high altitudes, the region in Turkey is divided in two vegetation areas: 1. a broad outer area of deciduous scrub and rare park-like forest; 2. inner areas of the real steppe without trees. The main part of steppe area is the Central Anatolian steppe that eastward, gradually passes into montane steppes or mountain cushion vegetation of the East Anatolian highlands (Davis at all, 1965). This is the most suitable region for the growth of Plumbaginaceae in Turkey. Hence, 2 Limoniopsis, 10 Limonium, 49 Acantholimon and 1 Psylliostachys species are present in this region. In total that is 62 species or 74.7 % Habitat preferences of the Plumbaginaceae in Turkey Plumbaginaceae prefers two main types of habitats. First type is the coastal areas (seaside cliffs, sandy shores, or salty marches), where around half of the Limonium and all of the Armeria species are present. Also, Phylliostachys spicata is found on saline clay and sandy soil on planes or along sea coasts. Second type includes continental habitats that can be: typical steppe or montane steppe, inland salty areas, or dry, rocky, exposed mountain slopes. The other half of Limonium 31

50 species prefer inland salty areas, as well as some species of Acanthalimon. The majority of Acantholimon is confined to highlands and mountain area, and dry, rocky or limestone slopes. Some are present in steppes and mountain steppes, too. Plumbago europea may grow at volcanic or limestone slopes and uncultivated fields or waste areas. Goniolimon incanum is present in the steppes, uncultivated fields, or on the serpentine and chalky hills Endemism From the currently accepted 83 species of Plumbaginaceae in Turkey 44 species are endemics (53.01%). Also there are 6 endemic subspecies and 8 endemic varieties. The highest number of endemics are present in Acantholimon, 33 species (61% of the Acantholimon species growing in Turkey), from which most are very narrowly distributed local endemics. In Limonium there are 9 endemic species (42.86%). Also, there is 1 endemic in Limoniopsis (L.davisii) and 1 in Armeria (A.trojana). The highest number of endemics is present in region, especially mountain areas of South East Anatolia. The number of endemic species in this region is 38 or 86.36% of all Turkish Plumbaginaceae endemics. From those 30 species are Acantholimon, 7 are Limonium and 1 is Limoniopsis. The number of endemics is lower in the Mediterranean region. In here, there are 3 Acantholimon, 1 Armeria and 9 Limonium endemic species, which is 13.64% of all endemics in this family. So far there are no records of endemic Plumbaginaceae from Euro-Siberian region (Davis et all, 1982, 1988; Dogan and Akaydın, 2003a, 2006) Conservation Status of the Plumbaginaceae in Turkey According to the IUCN (International Union for Conservation of Nature and Natural Resources) threat categories, in Turkey: 37 Plumbaginaceae species (44.58%), are in CR (Critically Endangered) category, 13 species (15.66%) are EN (Endangered), 17 species (20.48%) are VU (Vulnerable), 5 species (6.02 %) are NT (Near Threatened) and 12 species (14.46%) are LC (Least Concern) (Doğan & Akaydın, 2003a, 2006). All of the threat categories of the taxa are given on the basis of IUCN. Red List Categories: Version 3.1. (IUCN, 2001). 32

51 The most threatened species are the local endemics with very narrow distribution range and small populations. The main threat to them is habitat destruction caused by human effect, erosion and extensive grazing. Consequently, the highest number of species in CR category is among Acantholimon species. Figure 11: Acantholimon avanosicum, Critically Endangered species, local endemic of Avanos, Nevşehir province (edited from the article, Doğan and Akaydın, 2002a) 1.4. Numerical taxonomy Taxonomy is a part of systematics that includes four main components: description, identification, nomenclature and classification of taxa (Simpson, 2010). Taxa (singular taxon) is a defined or delimited groups of organisms. The very first classification system used since the time of Ancient Greece was the artificial system of classification. This kind of system makes comparison of objects depending on, only one or at most few characters. It was replaced by natural system of classification, built upon several to many characters that bring together organisms into natural groups that have highest number of shared features. Further, Darwin s theory of evolution brought new approach to classification, phyletic or evolutionary classification. Although, it offered explanation for homogeneity of classified groups, through the idea that similarity of organisms was a result of 33

52 descent from the common ancestor, it did not bring new methods to taxonomy. All those approaches share the common problem that they are highly subjective. In the sense that selection of characters and their comparison and evaluation depend largely on individual approach of taxonomists. Numerical taxonomy (phenetics), as a new approach, was created in 1960 s by Robert R. Sokal and Peter H.A. Sneath as a reaction to that situation. Their main aim was to classify organisms based on their overall similarity and all available characters without any weighting, by clearly defined numerical procedures, in order to avoid high level of subjectivity that exists in previous systems of classification. Therefore, numerical taxonomy was defined as a: Grouping by numerical methods of taxonomic units on the basis of their character states (Sneath and Sokal, 1973) Objectives of the Study There seems to be only a few taxonomic studies ever done on Plumbaginaceae in Turkey and they always gave highest attention to Acantholimon and Limonium. Furthermore, in the recent years, some new species were described, a few imperfectly known, potential species still exist, and even a new genus, Psylliostachys (Jaub. & Spach.) Nevski, had been recorded in Turkey. Also, some of the species such as, Armeria cariensis and the genera Limonium and Acantholimon have unresolved taxonomic structure. Hence, we believe that, there is a need for reexamination of the taxa in order to construct more natural classification. For the first time all the genera of Plumbaginaceae, existing in Turkey, have been evaluated all together, by the use of numerical taxonomy. Our main aim is to see linear arrangement of genera within family. Also, to see infrageneric grouping in the big genera like Limonium and Acantholimon. As well as, to discover potential placement, of the new species and genus in existing taxonomic structure. The highest possible number of the character and character states will be used, in order to obtain the most objective result and to produce long lasting natural grouping in the family. At the end obtained result will be discussed, and compared with previous findings. Also, a potential changes in the structure of genera or the family will be suggested. 34

53 CHAPTER 2 MATERIAL AND METHOD 2.1. Material of the Study Majority of the samples examined in this study had been collected as a part of two projects supported by The Scientific and Research Council of Turkey (TÜBITAK): Türkiye nin Acantholimon Boiss. (Plumbaginaceae) Türlerinin Revizyonu (199T011) (Doğan and Akaydın, 2003a) and Türkiye'deki Plumbaginaceae Juss. Familyası Üzerinde Revizyonel Çalışmalar (102T088) (Doğan and Akaydın, 2006). Those studies had been conducted by Prof. Dr. Musa Doğan and Prof. Dr. Galip Akaydın, between 2000 and Those specimens are kept as a herbarium material in the Plant Systematics Laboratory, Middle East Technical University (METU). Some of the duplicate specimens of Acantholimon from the Royal Botanic Garden, Edinburgh (E), are part of the collection, as well. The specimens had been identified by the use of the relevant floras, as Flora of Turkey and the East Aegean Islands (Davis at all,1982, 1988), then Flora Iranica (Rechinger and Schiman-Czeika, 1974), Flora Europaea (Tutin et al, 1972), Flora of USSR (Komarov, 1967), etc Method of the Study As a method of the study numerical taxonomic analysis has been performed, following the procedure steps suggested by Stussey (Stuessy, 2009), which are based on methods proposed by Sneath and Sokal (1963,1973) in their books, Principles of Numerical Taxonomy and Numerical Taxonomy. 35

54 Selection of Operational Taxonomic Units (OTU s) First step in numerical taxonomic analysis is to select Operational Taxonomic Units (OTUs). OTUs could be individuals, populations, species, genera, etc. OTUs in this study are species of the genera of Plumbaginaceae present in Turkey. In total 61 OTUs (species) was used. They belong to genera Plumbago (2), Ceratostigma (1), Goniolimon (1), Limoniopsis (1), Armeria (3), Psylliostachys (1), Limonium (24) and Acantholimon (28). In the case of Acantholimon, species for the study had been selected with intention to include a few representatives of all section and subsections, as well as to include all available specimen of the species that are newly described. For the rest of genera all of the species available in METU herbarium were used. For each species at least two of the specimen had been examined. All specimens are supplied with identification number and additional information about locality (from where it was collected and position of the locality in the grid square system in Turkey defined by Davis (Davis et al, 1965). As well as, habitat type, elevation, date (when it was collected) person who collected and /or identified specimens. Complete list of the used OTUs, is given in the Table B1 in the Appendix B Selection of characters and coding of character states Second step is selection of characters and coding of the character states. In numerical taxonomy, the conventional approach is to use as wide as possible range of the characters, choosen in an objective way, with avoidance of overlapping. It is advisable to select characters from all parts of the OTUs. Characters are usually combination of quantitative and qualitative features. They may have two opposite states or characters may have multi states. About number of selected characters, it is considered that it is ideal to choose as many characters as possible, by expecting that resulted correlations between OTUs will be more reliable if the number of character is higher. Also, one of the basic ideas of numerical taxonomy is that all characters should be unweighted or equally weighted, without giving any priority to some of the features. In accordance with those principles 57 characters were selected for this study. Characters are morphological and chosen from all part of OTUs: 5 general characters (life form, life length, etc.), 6 scape characters, 14 leaf characters, 7 36

55 inflorescence character, 9 bracts characters, 10 calyx characters, 2 corolla characters and 4 characters regarding stigma and style features. Data are mixed types, and include 46 qualitative characters and 11 quantitative (from those 3 characters were coded and the rest were used as numeric value). In the following step character states of selected characters are described, measured and coded. Complete list of characters and their states and codes is given below in the Table 3. Table 3: Character table No Characters: Character states: General characters: 1. Life length: Perennial (0) Annual (1) 2. Life form: Shrub /subshrub/ shrublet (0) Herb (1) 3. Habit forms: Cushion formation (0) Not cushion formation (1) 4. Stem is: Branched (0) Not branched (1) 5. Woody caudex: With woody caudex (0) Without woody caudex (1) Scape characters: 6. Scape is: With scales (0) Without scales (1) 7. Scape is: With wings (0) Without wings (1) 8. Scape is: Branched (0) Not branched (1) 9. Scape is: With sterile branched (0) Without sterile branches (1) 10. Scape is: Glabrous (0) Without aglandular hairs (1) With glandular hairs (2) 11. Scape length: cm Leaf characters: 12. Leaves are: Rigid, linear, triquetrous, and pungent (0) Fleshy or leathery, never pungent (1) 13. Duration of the leaves: Leaves dry up before ending of flowering (0) Leaves do not dry up (1) 14. Leaves are: Arranged into basal rosettes (0) Arranged into fascicular rosettes on shoots below the scape (1) Not arranged into rosettes (2) 37

56 Table 3. (Continued) 15. Shape of the leaves: Linear (0) Lanceolate (1) Lanceolate to oblong - lancelolate (2) Broadly ovate to obovate (3) Elliptic (4) Elliptic to oblong obovate (5) Spathulate (6) Oblong spathulate (7) Obovate - spathulate (8) Polymorphic leaves (9) 16. Leaves are: With petiole (0) Without petiole (1) Usually down leaves are with petiole, medium and upper leaves are sessile or auricular (2) 17. Leaf margin is: Entire (0) Pinnatifid (1) 18. Leaves are: Glabrous (0) Glabrous with scabridulous margin (1) Glabrous with ciliate/ciliolate margin (2) With aglandular hairs (3) With aglandular hairs and scabridulous margin (4) With glandular hairs (5) 19. Leaf tip is: Acute (0) Obtuse to rounded (1) Both type of the leaves are present (2) 20. Leaf tip is: Without mucro (0) Mucronate (1) Apiculate (2) Spinose or pungent (3) 21. Average leaf length: mm 22. Average leaf width: mm 23. Length/Width ratio: ratio 24. Leaves are: Homomorphic (0) Heteromorphic (1) 25. Previous year s leaf bases are: Inflorescence characters: Circinnate (0) Not circinnate (1) 26. Inflorescence is: Spicate (0) Terminal and lateral cylindrical spikes (1) Terminal, subcapitate to capitate spikes (2) Capitate condensed spikes (3) Capitate (4) Paniculate (5) Corymbose to paniculate (6) Globose (7) 38

57 Table 3. (Continued) 27. Inflorescence is: Unilateral (0) Bilateral (1) Both unilateral and bilateral (2) Not as above (3) 28. Average inflorescence length: cm 29. Average number of spikelet per inflorescence: number 30. Number of florets per spikelet is: 1 floret per spiklet (0) 1-(2)-(3)-(4) florets per spikelet (1) 2-(3)-(4)-(5) florets per spiklet (2) 3-(4)-(6) florets per spiklet (3) 4-5 florets per spiklet (4) 8-12 florets per spiklet (5) 31. Average spikelet length: mm st internode is: Longer than spikelet (0) Shorter than spikelet (1) Bract characters: 33. Number of bracts is: Less than three (0) Three (1) More than three (2) 34. Outer bract s tip is: Acute (0) Obtuse (1) 35. Outer bract s vein is: Not excurrent (0) Excurrent (1) 36. Outer bract s margin is: Not hyalinated (0) Fully hyalinated (1) Fully hyalinated except vein (2) 37. Outer bract is: Glabrous (0) With aglandular hairs (1) With glandular hairs (2) st inner bract s tip is: Acute (0) Obtuse (1) Emarginate (2) st inner bract s vein is: Not excurrent (0) Excurrent (1) st inner bract s margin is: Not hyalinated (0) Fully hyalinated (1) Fully hyalinated except vein (2) st inner bract is: Glabrous (0) With aglandular hairs (1) With glandular hairs (2) Calyx characters: 42. Sepals are: Connate (0) Free (1) 43. Calyx shape is: Tubular (0) Infundibular (1) Subtubular to tubular (2) Obconical (3) 39

58 Table 3. (Continued) 44. Calyx duration: Calyx is permanent (0) Calyx is not permanent (1) 45. Calyx is: Glabrous (0) With aglandular hairs (1) With glandular hairs (2) With glandular and aglandular hairs (3) 46. Calyx is: Not hyalinated (0) Hyalinated (1) Partially hyalinated (2) 47. Calyx veins are: Not excurrent (0) Excurrent (1) 48. Calyx lip is: With lobes (0) Without lobes (1) 49. Calyx lip margin is: Dentate (0) Not dentate (1) Plicate (2) Undulate (3) 50. Calyx lip color is: White (0) White to pink (1) White to violet (2) White to purple (3) Pink to purple (4) Pink to clatter red (5) Violet (6) Brownish to pink-purple (7) Brownish (8) Green-whitish (9) 51. Average calyx length: mm Corolla characters: 52. Corolla: Corolla is gamopetalous (0) Petals are connate at the base only (1) 53. Corolla color is: White (0) White to cream (1) White to pale pink (2) Pale pink (3) Pink (4) Pale violet (5) Violet (6) Bluish violet (7) Pale blue (8) Blue (9) Cream to yellow (10) Stigma and style characters: 54. Stigma is: Filiform cylindrical (0) Capitate,oblong-capitate,oblique or maleiform (1) 55. Style number is: One (0) Five (5) 56. Styles are: United throughout their length, except for the fairly long stigmas (0) Free from the base (1) 57. Styles are: Glabrous (0) Hairy (1) 40

59 After selection of characters and coding of their states, the numerical data are placed into basic data matrix, which will be used for computing similarities or dissimilarities between OTUs, by comparison of character states Construction of data matrix and its use Data matrix was constructed based on 61 operational taxonomic units indicated in the Table B1 in the Appendix B and 57 characters indicated in the Table 3. Complete data matrix is given in the Table C1 in the Appendix C. Qualitative data were coded into binary states (0 or 1) for the presence or absence of character state, or multi states (from 0 to n states). Quantitative characters counted by number of flowers, spikelets, bracts, etc. are coded as well, because there are specific patterns within genera that allow clear delimitation of groups with specific number of flowers, bracts, etc. Quantitative data such as length, width and ratios are represented by arithmetic mean of at least 10 measurements of particular plant part and from at least 2 specimen from each species. Those data are not converted into codes and they are kept as numerical value within 0.01 scale The comparison of character states Comparison of the character states has two main steps. First, is to select particular statistics for measuring similarity within OTUs, by using some of the coefficients of similarity. They all serve for calculations of affinity between OTUs. In this study Gower General Similarity Coefficient, a type of association coefficient, was used. Because it is the most suitable and most commonly used for the mixed data (such as using both qualitative and quantitative characters that have binary states or multistate, and missing data are acceptable as well). Second, is to select an algorithm to determine phenetic relationship among all the OTUs. For the second step there are two main approach: clustering and ordination. Clustering is the most commonly used method and there are three basic types: single linkage (nearest neighbor technique), complete linkage (farest neighbor technique) and average linkage. Average linkage is effort to connect new OTU to an average value of the particular group rather than to the extreme similarity or 41

60 difference within it, like with previous two types. Average linkage has two basic types: arithmetic average (it can be unweighted UPGMA and weighted WPGMA) and centroid (unweighted and weighted). In this study UPGMA (Unweighted Pair Group Method with Arithmetic Mean) was used to calculate the average similarity or dissimilarty of an OUT to a cluster. In other words, the distance between two clusters is depend on the mean distance between all objects in these two clusters. Results of phenetic relationships are the most commonly presented graphically, in the form of phenograms or cluster diagrams. Phenogram is constructed to expresses graphically relationships among all OTUs and to reveals the taxonomic structure of groups and subgroups among the OTUs (Stuessy, 2009). It is important to mention that phenograms simply show phenetic similarity based on comparison of character states and that evolutionary pathways are not represented or inferred in any way. Ordination method contains few different approaches which all have the same goal, to calculate and show multidimensional relationships of OTUs. In this study Principal Component Analysis (PCA) was used to reveal multidimensional relationships of OTUs and to indicate which characters are the most important in forming particular clusters. For the purpose of data evaluation and determination of relationships between OTUs a software program was applied on the previously constructed data matrix. Software used for this purpose was MVSP A Multi-Variate Statistical Package by Kovach Computing Services. As it was mentioned before, Gower General Similarity Coefficient was used in this study. Since it was the most suitable for our case with mixed data and few missing data (in the case where character was not applicable to the particular OTU). After the phenetic relationships had been obtained the next step is to determine ranks of OTUs. Ranking can be done by placing OTUs into informal categories (groups, subgroups, clusters, etc.) or into formal taxonomic categories or subcategories (sections, genera, species, etc.). 42

61 CHAPTER 3 RESULT AND DISCUSSION 3.1. Result Result of the Cluster Analysis and UPGMA phenogram Result of the cluster analysis performed by MVSP software with Gower Similarity Coefficient was given in the form of UPGMA phenogram in Figure 12. As it is shown in Figure 13, a line drawn at 0.66 separates tree in 2 main subgoups that may correspond to 2 subfamilies: Plumbaginoideae and Staticoideae. Also, drawing a line at 0.85 divides tree into 7 clusters that are equal to genera of Plumbagianceae. The only exception is Psylliostachys which shows earlier separation from the rest of genera at 0.71 similarity coefficient. Sections within Acantholimon and Limonium are observed as well around 0.9 cut off line, Figure 14 and Figure

62 44 Figure 12: UPGMA phenogram: Plumbaginaceae in Turkey

63 The list of the species, according to their ordinal numbers in the Figure 12: 1. Plumbago europaea 2. Plumbago capensis 3. Ceratodtigma plumbaginoides 4. Goniolimon incanum 5. Limoniopsis davisii 6. Armeria trojana 7. Armeria cariensis var.cariensis 8. Armeria cariensis var. rumelica 9. Psylliostachys spicata 10. Acantholimon bracteatum 11. Acantholimon capitatum subsp.capitatum 12. Acantholimon hoshapicum 13. Acantholimon tragacanthinum 14. Acantholimon quinquelobum var. quinquelobum 15. Acantholimon araxanum 16. Acantholimon laxiflorum 17. Acantholimon dianthifolium 18. Acantholimon calvertii var. calvertii 19. Acantholimon hypochaerum 20. Acantholimon köycegizicum 21. Acantholimon huetii var. huetii 22. Acantholimon ulicinum var.ulicinum 23. Acantholimon anatolicum 24. Acantholimon yıldıelicum 25. Acantholimon karamanicum 26. Acantholimon confertiflorum 27. Acantholimon puberulum subsp. puberulum 28. Acantholimon glumaceum 29. Acantholimon caryophyllaceum 30. Acantholimon avanosicum 31. Acantholimon venustum var. venustum 32. Acantholimon petraeum 33. Acantholimon spirizianum 34. Acantholimon kotschyi 35. Acantholimon libanoticum 36. Acantholimon halophilum 37. Acantholimon acerosum subsp. acerosum var. acerosum 38. Limonium sinuatum 39. Limonium echioides 40. Limonium anatolicum 41. Limonium ocymifolium 42. Limonium meyeri 43. Limonium lilacinum var. lilacinum 44. Limonium pycnanthum 45. Limonium marmarisense 46. Limonium angustifolium 47. Limonium effusum 48. Limonium gmelinni 49. Limonium guenerii 50. Limonium bellidifolium 51. Limonium vanense 52. Limonium davisii 53. Limonium caspium 54. Limonium smithii 55. Limonium graecum var. graecum 56. Limonium virgatum 57. Limonium sieberi 58. Limonium didimense 59. Limonium globuliferum var. globuliferum 60. Limonium tamaricoides 61. Limonium iconicum 45

64 46 Figure13: UPGMA phenogram: Infrafamilial grouping of the Plumbaginaceae in Turkey

65 47 Figure14: UPGMA phenogram: Infrageneric grouping of the Acantholimon

66 48 Figure 15: UPGMA phenogram: Infrageneric grouping of the Limonium

67 Result of the Principal Component Analysis In addition, Principal Component Analysis in MVSP was carried out to indicate the most important characters for the forming of different clusters. Table 4: Eigenvalues, percentages and cumulative percentages (Eigenvalues > 1) Eigenvalues Axis 1 Axis 2 Axis 3 Axis 4 Axis 5 Axis 6 Eigenvalues Percentage Cum. Percentage

68 50 Figure 16: Principal Component Analysis of the Plumbaginaceae in Turkey with 2 axix

69 The Principal Component Analysis (PCA) exhibited similar result as our phenogram and separation within genera (Figure 16). The first 6 axis of the principal component analysis, resolved % of total morphometric variation. The 1 st axis explained %; the 2 nd axis %; the 3 rd axis %; the 4 th axis %; the 5 th axis 4.433% and the 6 th axis 2.633%; of the total variation within Plumbaginaceae (Table 4). The summary of the character loadings on the first 6 axis are given in the Table 5. Table 5: The summary of the highest character loadings at the first 6 axis Character No: Axis 1 Axis 2 Axis 3 Axis 4 Axis 5 Axis

70 Accordingly, the most important characters, found on the first 6 axis that explained most of the variation in clustering of Plumbaginaceae in Turkey are given in the Table 6: Table 6: The most significant diagnostic features according to PCA No: Character: 2. Life form: shrub /subshrub/ shrublet or herb 3. Habit is: cushion formation or not cushion formation 8. Scape is: branched or not branched 10. Scape is: glabrous, without aglandular hairs or with glandular hairs Leaves are: rigid, linear, triquetrous, and pungent or fleshy or leathery, never pungent Leaves are: arranged into basal rosettes, arranged into fascicular rosettes on shoots below the scape or not arranged into rosettes Shape of the leaves: linear, lanceolate, lanceolate to oblong-lancelolate, broadly ovate to obovate, elliptic, elliptic to oblong obovate, spathulate, oblong-spathulate, obovate-spathulate or polymorphic leaves Leaves are: with petiole, without petiole or usually down leaves are with petiole, medium and upper leaves are sessile or auricular Leaves are: glabrous, glabrous with scabridulous margin, glabrous with ciliate/ciliolate margin, with aglandular hairs, with aglandular hairs and scabridulous margin or with glandular hairs 19. Leaf tip is: acute, obtuse to rounded or both type of the leaves are present 20. Leaf tip is: without mucro, mucronate, apiculate, or spinose/pungent 26. Inflorescence is: spicate, terminal and lateral cylindrical spikes, terminal, subcapitate to capitate spikes, capitate condensed spikes, capitate, paniculate, corymbose to paniculate or globose 27. Inflorescence is: unilateral, bilateral, both unilateral and bilateral or not as above 30. Number of florets per spikelet is: 1 floret per spiklet, 1-(2)-(3)-(4) florets per spikelet, 2-(3)-(4)-(5) florets per spiklet, 3-(4)-(6) florets per spiklet, 4-5 florets per spiklet or 8-12 florets per spiklet 33. Number of bracts is: less than three, three or more than three 37. Outer bract is: glabrous, with aglandular hairs or with glandular hairs; 38. 1st inner bract s tip is: acute, obtuse or emarginated 39. 1st inner bract s vein is: not excurrent or excurrent 40. 1st inner bract s margin is: not hyalinated, fully hyalinated or fully hyalinated except vein 52

71 Table 6 (Continued) 41. 1st inner bract is: glabrous, with aglandular hairs or with glandular hairs 43. Calyx shape is: tubular, infundibular, subtubular to tubular or obconical 47. Calyx veins are: not excurrent or excurrent 49. Calyx lip margin is: dentate, not dentate, plicate or undulate 50. Calyx lip color is: white, white to pink, white to violet, white to purple, pink to purple, pink to claret red, violet, brownish to pink-purple, brownish or greenwhitish 53. Corolla color is: white, white to cream, white to pale pink, pale pink, pink, pale violet, violet, bluish violet, pale blue, blue or cream to yellow 54. Stigma is: filiform-cylindrical or capitate, oblong-capitate, oblique or maleiform 3.2. Infrafamilial Grouping of the Plumbginaceae Juss. in Turkey The infrafamilial grouping of Plumbaginaceae Juss. In Turkey, as the final result of this study, was indicated in the UPGMA phenogram in the Figure 12 and the Figure 13. First of all, in the Figure 13, a cut-off line at 0.66 separated the family into two main subgroups. Those two subgroups were considered as tribes by Boissier, and this approach was also accepted in the Flora Europaea, and the tribes were named as Plumbaginae and Staticea (Tutin et al, 1972). From the other point of view, some authors treated these subgroups as subfamilies: Plumbaginoideae and Staticoideae (Komarov, 1967; Kubitzki, 1993; Lledo et al, 2001). Obvious differences between subfamilies in morphology, were also supported by molecular analyses based on plastid DNA sequences and nuclear sequences (Lledo et al, 2001, 2005; Moharrek et al, 2014). Differences in chemical compounds were proven to be usefull for this subgrouping, too. (Hanson et al, 1994; Lledo et al, 2001). In this study, the following morphological characters typical for the Plumbaginoideae are observed. Inflorescences are terminal spikes (in a form of cluster that moslty consist of flowers); corolla is gamopetalous, calyx is tubular, herbaceous and usually glandular, calyx lip is dentate, leaves are usually polymorphic within the same species, with or without petiole, mucronate, with or 53

72 without glandular hairs, and do not form rosette. Stigma is filiform, number of the styles is 1 and styles are usually hairy. Only two genera are present in Turkey from this subfamily, as Plumbago (P.europaea, P.capensis, former as a native and latter as ornamental plant) and Ceratostigma, (C.plumbaginoides, as ornamental). In our phenogram (Figure 13) the separation line between them is at 0.76, and it is based on the character mentioned below. Figure 17: Ceratostigma plumbaginoides: A. inflorescence; B. flower; C.stem; D.leaf base; E.leaf; F.leaf tip and margin. (edited from Discover life webpage and Wikimedia commons webpage) Ceratostigma has glabrous leaves, without petiole and with ciliate margin; bracts with excurrent veins and fully hyalinated margin; free sepals; permanent, not glandular calyx, with pink to clatter red lip; glabrous style. Plumbago has polymorphic leaves, with or without petiole, with glandular or aglandular hairs; bracts without excurrent veins and not hyalinated; connate sepals; not permanent, glandular calyx, with green- white lip color; hairy style. 54

73 Figure 18: Plumbago europaea: A, B. inflorescence; C.stem and leaves; D. calyx and bracts (edited from Acta Plantarum webpage) Figure 19: Plumbago capensis: A, B. inflorescence; C.stem and leaves; D. calyx and bracts. (edited from Phyto Images, International Carnivorous Plants Society; Toowoomba Plants and Vascular Plants Image Library webpages) 55

74 Beside genera separation, we also got insight into differences between P.europea (the only Plumbago that is present in Europe and Turkey) and P.capensis (a representative of the rest of Plumbago that is mostly confined to tropical regions of Africa and Asia), that split at 0.91 line. First one is herb, with glandular hairs present on leaves and scape and violet rose flowers. Second is a shrub, with long branches terminating with many flowered inflorescences (20-40 light blue flowers); with aglandular hairs on the scape, leaves and calyx. All of the rest genera included in this study belong to subfamily Staticoideae, which is also more variable in all of the characters we applied for numeric taxonomic analysis. Cut off line between those genera is drawn at 0.85, on the UPGMA phenogram (Figure 13). The only exceptions are Psylliostachys and the species L.sinuatum that are separated from the rest of genera at 0.67 point, and one from the other at 0.71 point. They seemed to be morphologically quite distinctive from the rest of Staticoideae in Turkey. Psyllioststachys (P.spicata) was for the first time recorded from Turkey in Also, this is the first time that it is compared with the rest of Plumbaginaceae in Turkey, trough a numerical taxonomic analysis. Characters that showed this species to be very different from the others are: these are annual plants; glandular hairs are present on scape, leaves, and bracts; leaves are with pinnatifit margin; inflorescences are terminal or lateral, cylindrical spikes compound from many (up to 50 or more) spikelets with 2-4 flowers, and only 2 bract; calyx is glandular; corolla is gamopetalous (it is the only genus in Staticoideae with gamopetalous corolla). Parallel molecular and morphological studies of Plumbaginaceae in Spain including Psylliostachys gave different outcomes. Molecular analyses based on plastid DNA sequences showed that Psylliostachys if forming a clade with Armeria (with 100% probability), but this was supported by morphology with 69% (Lledo et al, 2001). Another molecular work showed existence of this clade, too. This was supported by 0.69 when plastid DNA and 0.96 when nuclear DNA sequences were used (Moharrek et al, 2014). What L.sinuatum brought close to Psylliostachys spicata are numerous differences that both species show comparing with other 56

75 Plumbaginaceae in Turkey. However, these species do not share many common characters, except pinnatifid leaves and hairiness of the scape, leaf and bracts, but type of the hairs is different. Due to that, they split early, at 0.71 point. Figure 20: Psylliostachys spicata: A. habit; B. spikelet. (edited from the Plantarium webpage and Forestry Images webpage) As it is mentioned before, the rest of genera within Staticoideae are proved to be well separated and defined clusters. Limonium, Limioniopsis and Goniolimon look more close to each other than Armeria and Acantholimon. Goniolimon and Limoniopsis are closest to each other according to the results of this study. Goniolimon (G.incanum) differ from the Limoniopsis (L.davisi) in: winged, glabrous scape; lanceolate to oblong-lanceolate, glabrous leaves, with ciliolate margin, acute and mucronate leaf tip, and bigger leaf size; corymbose to paniculate inflorescence; 1 flower per spikelet; obtuse outer bract and obtuse and tricuspidate 2 nd inner bract; infundibular calyx; white corolla; styles with long hairs. 57

76 Figure 21: Goniolimon incanum: A. habit; B. leaves; C. inflorescence (METU herbarium, photo by Jelena Erdal) Figure 22: Limoniopsis davisii: A. habit; B. leaves; C. inflorescence; (METU herbarium, photo by Jelena Erdal) 58

77 On the other side, L.davisii, has: scape hairy at base and without wings; oblong-spathulate leaves, glabrous, obtuse and apiculate leaf tip, smaller leaf size; paniculate inflorescence; 1 (occasionally 2) flower per spikelet, and large spacing between spikelets; acute outer bract and 1st inner bract, non excurrent vein of 1st inner bract; subtubular to tubular calyx; pale pink corolla, and glabrous styles. They both differ from Limonium in capitate stigma; not hyalinated calyx lip, with plicate margin; numbers of spikelet per inflorescence and flowers per spikelet. Limoniopsis species have not been used in the molecular based research so far. Goniolimon was included in the previously mentioned molecular work and it was forming clade together with Acanholimon and Dyctiolimon, which supposed to be a sister clade of clade formed by Psylliostachys and Armeria (Lledo et al, 2001). Limonium is well separated from the rest of genera in this study with exception of the species L.sinuatum (Figure 13, Figure 15). However, this kind of result was not a surprise, because taxonomic status of L.sinuatum and section Pteroclados, where it belongs, are not clearly defined, and different points of view exist about this issue. Also, within the cluster representing Limonium, a smaller clusters corresponding to the sections were observed. Beside that within section Limonium a possible subsections are indicated, at the 0.92 cut off line. The typical features of the genus Limonium are: herb or shrub life form; leaves mostly in basal rosettes; sterile branches present or not; spikelets grouped terminally (sometimes in capitate like structures); 3 bracts usually present; calyx infundibular, obconical or tubular; petals are connate at the base only; stigma is cylindrically filiform. L.sinuatum is the only representative of the section Pteroclados Sauv. & Vindt. and it has very specific morphology comparing with the rest of the Limonium species, in Turkey. It has winged, hairy scape; pinnatifit, hairy leaves arranged into basal rosettes; 3-4(6) flowered spikelets; more than 3 bracts; very characteristic 1 st inner bract, which is herbaceous, hairy and 2-3 toothed, with long exccurent veins; truncate calyx lip margin; white to cream flowers. For this reason, L.sinuatum appeared on phenogram as a completely separated branch, not only from the rest of Limonium species, but also from the rest of Plumbaginaceae in Turkey. Similar 59

78 results regarding L.sinuatum were obtained in the past (Ingrouille, 1984; Doğan and Akaydın, 2006; Fazlıoğlu, 2011) which justifies need for reconsideration of taxonomic status of section Pteroclados. Figure 23: Limonium sinuatum, section Pteroclados: A. leaves; B, C. stem; D, E. inflorescence; F. flowers (edited from Wikimedia Commons and Cut Flowers Exports of Africa webpages) Considering that this section consists of more species outside Turkey, beside L.sinuatum, it is necessary to look at the broader picture. In the Flora Europaea this section has status of subgenus Pteroclados (Boiss.) Pignatti, which consists of species: Limonium sinuatum Miller, Limonium bonduellei (Lestib.) O.Kuntze, Limonium thouinii (Viv.) O.Kuntze, Limonium brassicifolium (Webb & Berthel.) O.Kuntze. Those species are annual or perennial herbs, with sinuate or 60

79 lobed leaves, and winged scape (Tutin et al, 1972). Another subgenus recognized in the same flora as the subgenus Myriolepis (Boiss.) Pignatti was segregated from Limonium as separate genus, Myriolimon (Boiss.) Lledó, Erben & M. B. Crespo (Tutin et al, 1972; Lledo et al, 2003, 2005a). From that point of view it might be possible to do the same with section Pteroclados. Section Pteroclados has in total around 18 species. Beside L.sinuaum and European species mentioned above, there are L.beaumieranum, L.mouretii and L.bonduellei from Marocoo, L.lobatum (wide spread From Canary Island over North Africa till to Iran) and the rest of species are endemics to the Canary Islands. Molecular studies also support clear separation of this group from the rest of Limonium species (Lledo et al, 2005, 2011; Akhani et al, 2013). What makes researches hesitant in segregation of this section from the Limonium into a separate genus is the lack of unique characters of this group (Karis, 2003; Lledo et al, 2005, 2011). Because when we observe section Pteroclados next to the rest of Limonium it looks undoubtedly different, but the characters that make it unique within Limonium, are not unique comparing with other genera of Plumbaginaceae. For example, the typical features of this section are also present in another genera, such as: winged scape in Goniolimon, or sinuate leaves in Psyllisotachys. Even though, this section was subject of various studies, final solution for its taxonomic position has not been found so far. Limonium echioides is a single species representing the section Schizyhymenium (Boiss.) Bokhari, and within our Limonium cluster it is segregated from the rest of the species as a single branch. This is the only annual Limonium in Turkey with very distinguishing morphology. It has tuberculate leaves organized into basal rosette; very unusual calyx with long exccurent calyx veins forming hooks; and characteristic large, tuberculate, 3-veined, 2 nd inner bract. Recent molecular study revealed a moderately supported connection of L.echioides as a single branch and clade formed by Mediterranean species as L.delicatulum, L.furfuraceum, L.cymanum, L.minutum (Akhani et al, 2013). Nonetheless, further and different types of analyses are needed to reveal the most suitable taxonomic status of this peculiar species, and show its potential to be regarded as a subgenus of Limonium or an independent genus of its own. 61

80 Figure 24: Limonium echioides, section Schizyhymenium: A. habit; B. leaves; C.spikelet. (edited from the Plantes del Litoral webpage and Flora silvestre del Mediterráneo by Jose Quiles webpage) The cluster that consists of the species: L.lilacinum, L.globuliferum and L.pycnanthum is matching with the section Sphaerostachys (Boiss.) Bokhari. One of the imperfectly known specimens from our study L.marmarisense seems to belong together with this group, as particularly close to L.pycnanthum. Distinct morphological characters of this cluster are: herbaceous habit; eliptic or spathulate leaves, with obtuse, mucronate tip; inflorescence corymbose-paniculate, both unilateral and bilateral; more than 3 bracts; infundibular calyx (exception is L.globuliferum that has globose inflorescence; 3 bracts and obconical calyx). Another imperfectly known species, L.davisii, also share many common characters with this group, except that it has 3 bracts; emarginate 1 st inner bract, and obconical, glabrous, calyx. Previously mentioned, species are s, mostly from Central Anatolia, with exeption of L.marmarisense which is found in coast of Marmaris in Muğla province. For this reason, it is necessary to give more attention to this imperfectly known species in order to resolve its potential taxonomic status. 62

81 Figure 25: Limonium lilacinum, section Sphaerostachys: A. habit; B. leaves; C.inflorescence. (METU herbarium, photo by Jelena Erdal) The rest of the Limonium species belong to the section Limonium that seems quite heterogeneous. Because it did not appear as a single cluster in this study, instead four well defined clusters were formed (Figure 15). Those clusters are morphologically and ecologically different and may have potential to be recognized as a subsections within section Limonium. Species L.meyeri, L.effusum, L.gmelinii and L.angustifolium form cluster 1. They are all herbs, with large, eliptic to oblong-obovate leaves arranged in basal rosettes, quite tall scapes, paniculate inflorescence, and obconical calyx. Species L.gmelinii and L.meyeri have wide distribution ranges spreading from Europe till to Central Asia, both in coastal and inland salty areas. L.effusum is endemics of South and South West Anatolia and L.angustifolium grows all around the Mediterranean. 63

82 Figure 26: Limonium gmelini, section Limonium: A. habit; B. inflorescence. (METU herbarium, photo by Jelena Erdal) Another clearly distinguished cluster is cluster 2, formed by L.graecum, L.virgatum and L.sieberi. One of the specimen, suspected to be a new species, named as L.didimense, is shown to belong next to those. Same as, L.ocymifolium that have different position in various studies. In the Flora of Turkey L.ocymifolium is accepted to belong to section Limonium, in the later studies it was appearing in section Sarcophyllum along with L.anatolicum (Doğan and Akaydın, 2006; Fazlıoğlu, 2011). Our result is in accordance with the first opinion that L.ocymifolium is a part of section Limonium, because at phenogram, it seems closer to L.greacum, L.virgatum, L.sieberi and L.didimense than to L.anatolicum. Those species are differing from the other species in the following morphological characters: subshrub form, with woody caudex; sterile branches on the scape; small spathulate leaves, without mucro, organized in fascicular rosettes; inflorescence paniculate, usually unilateral; calyx infundibular; corolla pale violet. They are all Meditteranean s, found in similar types of habitat and form one well circumscribed group. 64

83 Figure 27: Limonium graecum, section Limonium: A. habit; B. inflorescence; C.leaves. (METU herbarium, photo by Jelena Erdal) Figure 28: Limonium anatolicum, section Sarcophyllum: A. habit; B. branch; C.inflorescence; D. leaves. (METU herbarium, photo by Jelena Erdal) 65

84 L.anatolicum is the only representative of the section Sarcophyllum (Boiss.) Lincz in Turkey and it seems to be closest to the previous group. Since it appears as a joint branch of the cluster 2, according to our phenogram (Figure 15). L.anatolicum, also, has some specific morphological features such as: very narrowly spathulate leaves, with short, flat, narrow petiole, forming numerous dense fascicular rosettes; capitate-like inflorescence, formed by condensed spikes; tubular calyx; whitish to pale blue corolla. Beside distinctive morphology this species is also, an endemic of Central Anatolia, confined to continental saline areas, that again supports its segregation from the previous group. L.bellidifolium, L.caspium and L.iconicum form cluster 3. The main morphological characters of this groups are: subshrub form with woody caudex; sterile branches on the scape; leaves that dry up before the end of the flowering; inflorescence bilateral, paniculate; calyx obconical; corolla color bluish-violet. L.tamaricoides seems closest to this group according to our tree, what makes it different from the other three species is: hairy scape, with paniculate branching and globose, many flowered inflorescence. Those species are s with exception of L.bellidifolium which is known as a Euro-Siberian in Turkey, but it has wide distribution range from South and Central Europe till to Central Asia. Accordingly, it is regarded as Mediterranean or in other regions or countries. Furthermore, all the species share same habitat preferences and grow in salty inland areas. Ecological features of this group also support its separation from other previously mentioned Mediterranean group. Connection between L.bellidifolium and L.caspium as well as few more Irano Turanian species was revealed by molecular study, where authors assumed two possible pathways by which L.bellidifolium was related to similar species. One hyposthesis is origin of the group with secondary distribution into other regions. Another hypothesis is a migration of a common ancestor from another region to region (Akhani et al, 2013). 66

85 Figure 29: Limonium bellidifolium, section Limonium: A. habit; B. inflorescence; C.leaves and sterile branches. (METU herbarium, photo by Jelena Erdal) There are three species, L.gueneri, L.smithii and L.vanense on our phenogram that seem to be out of the main clusters that appeared within section Limonim. Even though, L. gueneri and L. smithii form small cluster 4, they look distant one from the other. Also, L.vanense appears as a single branch not closely related to the rest of the clusters. Actually, those three species could be taken as an example how number and weigthing of the characters may affect classification. Because if we classify those species on the base of a few the most important diagnostic characters, they would be easily placed within the cluster 2 (L.gueneri) and cluster 3 (L.smithii and L.vanense). However, when we use 57 characters that are all equally important, differences between clusters are becoming more obvious and they are placed out of the main clusters. L.gueneri is one of the recently described species. It is a Mediterranean, a local endemic growing around ancient city of Patara in Antalya province. This species is a maritime, morphologically closest to L.ocymifolium (Doğan et al, 2008). From that point of view, it might be expected to see this species together 67

86 in cluster with the L.ocymifolium and joined with L.greacum, L.virgatum, L.sieberi and L.didimense. Still, there are significant differences in morphological characters between this cluster and L.guenrerii. First of all, is absence of the sterile branches on the scape, further, L.guenerii has herbaceous scape on woody caudex; obtuse to acute, mucronate leaves; inflorescence is corymbose-paniculate, bilateral, with a very specific number of flowers per spiklet (4-5). Figure 30: Limonium gueneri, section Limonium: A.habit; B. leaves; C.inflorescence. (METU herbarium, photo by Jelena Erdal) L.smithii is an, an endemic of Seyfe lake area in Kırşehir province and it grows naturally together with L.tamaricoides (Akaydın, 2007). Hence, it is supposed to appear together in cluster with L.bellidifolium, L.caspium, L.iconicum and L.tamaricoides, as the most closely resembleing L.caspium (Akaydın, 2007). However, there are few differences in characthers with the species forming this cluster L.smithii that caused this kind of outcome, such as: average scape length, leaf shape, length, width and ratio; average number of spikelets per inflorescence; number of florets per spikelet; 1 st inner bract not emarginated; average calyx length and corolla color. 68

87 Figure 31: Limonium smithii, section Limonium: A. habit; B. leaves; C.inflorescence. (METU herbarium, photo by Jelena Erdal) According to our phenogram L.vanense look quite far from the clusters mentioned above. It is an, endemic to East Anatolia and one of the species with leaves that dry up before end of the flowering. For this reason, it was expected that this species appears together with the L.bellidifolium, L.caspium, L.iconicum and L.tamaricoides. Nevertheless, closer examination of the morphological characters provided better insight into their differences and showed that L.vanense has: herbaceous form, with woody caudex, others are subshrubs; missing sterile branches on the scape; eliptic to oblong-obovate leaves with, acute and apiculate leaf tip, others have spathulate leaves, with obtuse and mucronate leaf tip; calyx is infundibular, obconical in others. It would be interesting to study further this group of species with the leaves drying up before flowering, in order to see weather this character is result of their common origin or it is just parallel structure independently developed in different species. 69

88 Figure 32: Limonium vanense, section Limonium: A. habit; B. leaves; C.inflorescence. (METU herbarium, photo by Jelena Erdal) The cluster representing the genus Armeria is well isolated in our phenogram, regarding to its distinctive features: unbranched, leafless scape, without scaly leaves; growing from branched woody rootstock; fleshy, linear leaves; capitate inflorescence, with many 2-4 flowered spikelets; more than 3 bracts (6); emarginate 1 st inner bract and styles that are hairy at lower third. Armeria in Turkey is represented by 2 native species, A.trojana and A.cariensis. A.trojana is a local endemic confined to Mount Ida (Kaz Dağı) in Balıkesir province. Because of its specific dwarf habit, hairy base of the scape and ciliolate leaf margin and 2-3 flowers per spikelet, it is clearly separated from A.cariensis. 70

89 Figure 33: Armeria trojana (METU herbarium, photo by Jelena Erdal) Taxonomic status of the A.cariensis with its two varieties A.cariensis var. cariensis and A.cariensis var. rumelica, is a source of long lasting debate. Some of the authors accept these varieties (Davis et all, 1982; Doğan and Akaydın, 2006), while the others treat those varieties as a separate species A.cariensis and A.rumelica (Tutin et all, 1972; Yeşil et al, 2014). For this reason we included both of the varieties in this study. As a result, our phenogram did not show overlapping in morphological characters, and separate this two with the cut off line through 0.97 similarity. Characters that were indicating differences are: scape length, heteromorphic leaves in A.cariensis cariensis and calyx length and structure. However, situation is not simple how it seems at the first glance. In general, the whole structure of Armeria is pretty complicated. Beacause of the often hybridization, mostly sympatric distribution and occasionally uncertain borders between species. A.cariensis cariensis has range covering west Turkey and western and eastern parts of Trace region. A.cariensis rumelica is present in the whole Thrace region covering southeast Balkans. Consequently, those species show sympatric distribution in the Thrace region which includes European part of Turkey as we can see in the Figure

90 Figure 34: Armeria cariensis var. cariensis: A.habit; B.leaves; C. inflorescence (METU herbarium, photo by Jelena Erdal) Figure 35: Armeria cariensis var. rumelica: A. habit; B. leaves; C. inflorescence (METU herbarium, photo by Jelena Erdal) 72

91 Figure 36: Geographic distribution of A.cariensis var. cariensis and A. cariensis var. rumelica in Turkey (edited from TÜBİVES website) Since most of the studies, regarding this issue, are done on the specimen only from Turkey (Davis at all, 1982; Doğan and Akaydın, 2006), including this one, there is not enough evidence to make firm separation of these two varieties into species. Even though, this has been done by the group that also had available some of the Bulgarian specimen and did anatomical studies as support of the claim (Yeşil et al, 2014). Nevertheless, we believe that without profound study based on the specimen from all of the distribution range, including Greece, Bulgaria and Turkey, and different types of analyses, it is not possible to give a final word about this issue. Acanholimon is clearly split from rest of the genera according to phenogram (Figure 13). Characters that keep it apart from other genera are: cushion formation; rigid, linear, triquetrous, pungent leaves, without petiole and with glabrous or scabridulous margin; usually spicate inflorescense; bracts with excurrent veins and fully hyalinated except the vein part; influndibular, hairy, hyanlinated calyx, with mostly white calyx lip; corolla usually in different nuances of pink color; capitate stigma; glabrous styles. Three section are present in Turkey: Acantholimon, Tragacanthina Bunge and Staticopsis Boiss. They were accepted in the Flora of Turkey (Davis et al, 1982) and confirmed by the later revision of Acantholimon 73

92 (Doğan & Akaydın, 2003a). Those sections seem well circumscribed on our tree and separated from each other by the 0.93 cut off line (Figure 14). Section Acantholimon is represented by 3 species in this study: A.bracteatum, A.capitatum and A. hoshapicum. It has quite sharp character difference comparing to rest of the Acantholimon, such as: heteromorphic leaves and persistent spring leaves; unbranched scape; capitate inflorescence formed by condensed spikes, with 2-5 flowered spikelets and 2-6 obtuse, bracts; infundibular, hairy calyx. These species are s confined to East Anatolia. Figure 37: Acantholimon bracteatum, section Acantholimon: A. habit; B.inflorescence. (METU herbarium, photo by Jelena Erdal) Second section Tragacanthina included 3 species in this study: A.tragacanthinum, A.quinquelobum and A.araxanum. It also has specific morphological characters: heteromorphic leaves and deciduous spring leaves; branched scape; paniculate diffuse infloresecense, with 1- flowered spikelets, and 3 bracts; tubular calyx. A.tragacanthinum and A.quinquelobum are typical representatives of the section and they appear close to each other on phenogram. From the other hand A.araxanum look distant, but still belonging to same cluster. Reason for that is that it shares many common morphological characters with the group, but it has homomorphic leaves and infundibular calyx. All, species are Irano- 74

93 Turanian s from East Anatolia and west to north west Iran, and have preferences to dry sandy slopes, as a habitat. Figure 38: Acantholimon quinquelobum, section Tragacanthina: A. habit; B.inflorescence; C. spikelet. (METU herbarium, photo by Jelena Erdal) The last section Staticopsis is separated from the others by the following characters: monomorphic leaves; spicate, simple or branched inflorescence, with 1- flowered spikelet; 3 bracts and infundibular calyx. Within this section a few subclusters equate to subsection are observable at the 0.94 cut off line. First subcluster consist of A.laxiflorum and A.köycegizicum and they look pretty isolated from the rest of the species in this section. Considering that both species have circinnate previous year s leaf bases, it might be expected to see them within subsection Circinnatea Doğan & Akaydın. However, in the case of A.laxiflorum this result was not surprising, since it differs a lot from the other species with circinnate leaf base, and the special subsection Robustea Doğan and Akaydın, was established for it (Doğan et al, 2007). Characters shown to be specific for A.laxiflorum are: glabrous, really long (average 29 cm) scape; long spikes (average 22.5 cm), with average 26 spikelets, 1 st internode shorter than spikelet; non excurrent calyx vein; brownish calyx lip. At the other side A.köycegizicum, morphologically resembles other members of Circinnatea, but there are also significant difference that bring it close to 75

94 A.laxiflorum. First of all, A.köycegizicum has quite robust habit, too. It has: long scape (22cm average), 12 spikelets per infloescence in average; 1 st internode longer than spikelet; nonexcurrent calyx vein and brownish calyx lip color. Important difference between these two species is that: in A.laxiflorum spikes are unbranched, laxly distichous, in A.köycegizicum spikes are branched and they consist of 2-5 distichous branches (Dogan and Akaydın, 2003c, 2003d). Figure 39: Acantholimon laxiflorum, section Staticopsis: A. habit; B. inflorescence (METU herbarium, photo by Jelena Erdal) Both of the species are Mediterranean s, preferring serpentine, rocky stream beds as a habitat, which also clearly separates them form the rest of the subsection Circinnatea that are species, occurring in highlands and mountains of South East Anatolia. Accordingly, taxonomic status of A.köycegizicum, should be more closely examined. Even though, it belongs to same floristic region, share same habitat demands, and robustness of the appearance as A.laxiflorum, still it does not seem suitable to include A.köycegizicum it in the subsection Robustea. Because of the very obvious difference in inflorescence morphology. From the other hand, as a pure coastal, Mediterranean it should not be kept together with the 76

95 highland, mountain s of region in the subsection Circinnatea. Establishing of two series within subsection Circinnatea one including A.köycegizicum and another including the rest of the species with circinnate leaf bases, might be one of the possible solutions. Also, in this case different type of analyses, beside morphology and ecology, would possibly offer better understanding of this issue. Figure 40: Acantholimon köycegizicum, section Staticopsis: A. habit; B. inflorescence (METU herbarium, photo by Jelena Erdal) Second subcluster within section Staticopsis was formed by the A.calvertii and A.huetii, matching with the subsection Circinnatea Doğan & Akaydın. This subcluster is separated on the account of the following morphological characters: hairy scape; previous year's leaf bases circinnate; hairy leaves with scabridulous margin; branched spicate inflorescence; 1 st internode shorter than spikelet; excurrent calyx veins; average calyx length 12,5 mm; calyx lip color white to purple. Both species are s, endemics, restricted to South East Anatolia. They grow at highland steppes and calcareous or volcanic slopes, at high elevations (from 1200 up to 3500 m). 77

96 Figure 41: Acantolimon calverti, section Staticopsis: A. habit; B. inflorescence. (METU herbarium, photo by Jelena Erdal) The rest of the Acantholimon species form two big clusters associated to the subsections Androsacea Bunge and Caryophyllacea Bunge. First cluster, analogous to subsection Caryophyllacea include species: A.caryophyllaceum, A.confertiflorum, A.avanosicum, A.venustum, A.petraeum, A.acerosum, A.halophilum and A.kotschyi. Figure 42: Acantolimon caryophyllaceum, section Staticopsis: A. habit; B.inflorescence. (METU herbarium, photo by Jelena Erdal) 78

97 Second cluster regarding subsection Androsacea consists of the species: A.ulicinum, A.anatolicum, A.glumaceum, A.yıldızelicum, A.hypochareum, A.puberulum, A.spirizianum, A.karamanicum, A.libanoticum and A.diantifolium. Figure 43: Acantolimon ulicinum, section Staticopsis: A. habit; B.inflorescence. (METU herbarium, photo by Jelena Erdal) All of those species are s (except A.libanoticum), growing on different types of rocks or montane steppes. Difference between those two groups lays in inflorescence structure. Androsacea has simple or shorlty branched spikes, and terminally densely congested spikelets and Caryophyllacea has, simple, lax, 2-ranked spikes (Doğan et al, 2007). Beside this, characters shown to be responsible for the separation of those groups are: scape length; inflorescence length; texture of outer and 1 st inner bract and calyx lip color. Structure of the groups is mostly in harmony with the ones accepted in the Flora of Turkey and the later revision of the Acantholimon (Davis et al, 1982; Doğan & Akaydın, 2003a; Doğan et al, 2007). Nevertheless, there are few exceptions. One of those is A.diantifolium. This species was segregated in subsection Diantifoliea Doğan & Akaydın (Doğan et al, 2007). However, in our phenogram it appeared as a part of subsection Caryophyllacea, which is in accordance with the old position of the species suggested by Flora of Turkey (Davis et al, 1982). Characters that caused this kind of placement are: scape and inflorescence length; texture of outer bract; shape of the 1 st inner bract and calyx lip color. Another species with unexpected placement is 79