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FLORA OF TURKEY VOLUME ONE
VOLUME ONE
FLORA OF TURKEY and the East Aegean Islands
edited by
P.H.DA VIS D.Sc
·Senior Lecturer in Taxonomic Botany assisted by J. CULLEN, PHD and M.J.E.COODE,BA DSIR Research Assistants
*
University of Edinburgh
EDINBURGH at the
UNIVERSITY PRESS
© 1965
Edinburgh University Press 22 George Square, Edinburgh Reprinted 1997,2001,2008 Printed and bound by Antony Rowe Ltd, Chippenham, Wiltshire A CIP record is available for this book from the British Library ISBN 0 85224 159 3
PREFACE The last and only Flora to cover Turkey was Boissier's Flora Orienta/is (18671888). Although this monumental work provides as good a foundation as any major Flora of its period, an enormous amount of herbarium material has accumulated since Boissier's time and a great many new species have been described. Our understanding of the Turkish flora has been hampered up to now by the lack of synthesis of all this activity. This is demonstrated by the extreme inaccuracy, as far as Turkey is concerned, of many published distribution maps. Turkey has for several reasons a particularly interesting flora: (1) it is the meeting place of three phytogeographical regions, the Euro-Siberian, Mediterranean, and Irano-Turanian regions, whereas only the first two are represented in the other Mediterranean peninsulas; the Turkish flora is therefore exceptionally rich; (2) Anatolia forms a bridge between Southern Europe and the flora of South-West Asia, and has apparently served as a migration route, particularly for the penetration of Asiatic elements into South Europe; (3) many genera and sections have their centre of diversity in Anatolia, so that their revision in this area leads to a better understanding of their taxonomy elsewhere; (4) species endemism is high- on the evidence of the present volume 20-25 per cent; this factor is presumably connected with the climatic and topographical diversity of the country, and the limited extent of Pleistocene glaciation; (5) many cultivated plants (crop plants, fruit trees, ornamentals) and plants that are weeds in Europe appear to have had their wild origin in Anatolia and adjacent areas; the arid areas produce numerous fodder plants of potential economic importance. For these reasons, the time seems ripe to produce a Flora of Turkey, concurrently with the Flora Europaea and K. H. Rechinger's Flora Iranica. This should provide a sound basis for further research when more material and biosystematic data become available. We have made no attempt to solve all our taxonomic problems; many species are still only known from single gatherings and their status or affinities must remain in doubt until they are collected again. By drawing attention to such cases we hope to spotlight taxonomic problems that need further study. If time were no object, this might have been a more comprehensive volume. It has been essential, however, not to be deflected from our main purpose - the steady progress of the Flora within a limited pericd of time. The work is planned to appear in eight volumes at approximately two-yearly intervals. Volume 2 will include Caryophyllaceae, Illecebraceae, Portulacaceae, Chenopodiaceae, Amaranthaceae, Tamaricaceae, Frankeniaceae, Droseraceae, Elatinaceae, Hypericaceae, Malvaceae, Tiliaceae, Linaceae, Oxalidaceae, Balsaminaceae, Geraniaceae, Zygophyllaceae, Rutaceae, Aquifoliaceae, Rhamnaceae, Celastraceae, Aceraceae, Staphyleaceae, Vitaceae and Anacardiaceae, and Volume 3 Rosaceae and Leguminosae. The editor will be glad to hear from botanists who would be interested in revising genera for Volume 3 and later volumes. P.R. DAVIS Edinburgh, March 1965
ACKNOWLEDGEMENTS The Flora could not have been undertaken without the generous assistance of a grant from the Department of Scientific and Industrial Research. I am extremely grateful for this help which has enabled two research assistants, Dr J. Cullen and Mr M. J. E. Coode, to be employed full-time on the preparation of the Flora. My thanks are also due to the Royal Society for a grant towards the cost of publishing this volume. A number of collaborators have revised genera for the first volume. I am particularly grateful to Mr I. C. Hedge (part of Cruciferae, including the basis of the key to the genera), and Mr D. M. Henderson (Filicales), both of the Royal Botanic Garden, Edinburgh; also to Dr T. R. Dudley (Arnold Arboretum; Alyssum and Aurinia), Dr C. D. K. Cook (Liverpool University; Ranunculus Sect. Batrachium) and Dr A. C. Jermy (British Museum; Isoetes). To Janet Dyer I am indebted for all the botanical illustrations except those of Reseda (by Coode), to Mr I. C. Dunn for drawing the maps, and to Edinburgh University Press and the printers, Cunningham of Alva, for facilitating the speedy production of the book. Among those who have helped us in various ways I should like to thank Mr B. L. Burtt, Dr H. Demiriz, Professor V. H. Heywood, Professor H. Kayacik, Mr R. D. Meikle, Dr H. Runemark, Mr C. Tobey, Professor D. H. Valentine, Professor D. A. Webb and Dr F. Yaltirik. We look forward to increased collaboration with Turkish botanists as the Flora proceeds. Among the numerous herbaria which have sent material on loan or allowed us to work on their collections, we are particularly indebted to the Directors of the following institutions: Royal Botanic Gardens, Kew; British Museum (Natural History); Naturhistorisches Museum, Wien; Conservatoire et Jardin Botanique, Geneve; Botanical Institute of the Academy of Sciences, Leningrad; Museum Nationale d'Histoire Naturelle (Laboratoire de Phanerogamie), Paris; Botanical Museum, Lund; and the Botany Departments of the Ankara and Istanbul herbaria. Dr A. Huber-Morath (Basel) has put much of his important Turkish collection at our disposal and provided us with new records for the Flora. The writing of the Flora has been greatly facilitated by the large collections, which are conserved in the herbaria of The Royal Botanic Gardens of Kew and Edinburgh, made in Turkey by myself, colleagues and research students. For grants to undertake these expeditions I should like to thank the following bodies: The Royal Society, Percy Sladen Memorial Fund, University of Edinburgh, Carnegie Trust for the Universities of Scotland, Leverhulme Trust, Royal Horticultural Society, Godman Fund (British Museum), Mount Everest Foundation, North Atlantic Treaty Organisation, and Commonwealth Scientific and Industrial Organisation. The work has been undertaken at the University Department of Botany, Edinburgh, using the excellent herbarium and library facilities provided by the Royal Botanic Garden. I am indebted to Professor Robert Brown, of the University Department of Botany, for help and encouragement, and to Dr H. R. Fletcher, Regius Keeper of the Royal Botanic Garden, for providing facilities. P.H.D.
CONTENTS List of Maps, p. x List of Figures, p. xi INTRODUCTION
Geographical divisions, p. 1 Topography, p. 3 Climate, p. 5 Plan of the Flora, p. 7 Main sources for Turkish records, p. 14 Phytogeography, p. 16 Euro-Siberian region, p. 16 Mediterranean region, p. 18 lrano-Turanian region, p. 21 Phytogeographical elements, p. 24 Endemism, p. 24 General literature on Turkish vegetation, p. 25 Further collecting, p. 26 Abbreviations, p. 27 FLORA
Pteridophyta, p. 31 Equisetales, p. 31 Lycopodiales, p. 34 Selaginellales, p. 36 Filicales, p. 38 Spermatophyta, p. 67 Gymnospermae, p. 67 Angiospermae, p. 86 Index, p. 541
LIST OF MAPS 1 2 3
Turkish vilayets and Grid system, p. 2 Geographical divisions within Turkey, p. 10 Geographical divisions outside Turkey, p. 12 DISTRIBUTIONS
4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30
Selaginella denticulata, Polystichum lonchitis, p. 69 Abies nordmanniana, A. cilicica, p. 69 Picea, Cedrus, p. 69 Pinus sylvestris, p. 73 Pinus nigra subsp. pallasiana, p. 73 Pinus brutia, p. 73 Pinus pinea, P. halepensis, p. 77 Taxus, Cupressus, p. 77 Juniperus drupacea, J. oblonga, p. 77 Juniperus communis subspp. nana & hemisphaerica, p. 81 Juniperus oxycedrus, p. 81 Juniperus sabina, J. phoenicia, p. 81 Juniperus foetidissima, p. 83 Juniperus excelsa, p. 83 Helleborus orienta/is, H. vesicarius, p. 83 Delphinium albijforum, D. carduchorum, D. bithynicum, D. venulosum, p. 121 Consolida hohenackeri, C. oliveriana, C. stenocarpa, p. 121 Consolida glandulosa, C. anthoroidea, C. axillijfora, p. 121 Consolida raveyi, C. tomentosa subsp. oligantha, p. 141 Actaea spicata, Clematis cirrhosa, C. viticella, p. 141 Papaver triniifolium, P. clavatum, p. 141 Alyssum callichroum, A. caricum, A. huber-morathii, p. 439 Cardamine lazica, Barbarea verna, Rorippa thracica, p. 439 Erysimum crassipes, E. sisymbrioides, p. 439 Cistus creticus, C. taurifolius, p. 509 Cistus parvijforus, C. salviifolius, p. 509 Reseda armena, Viola heldreichiana, V. gracilis, Polygala alpestris, p. 509
LIST OF FIGURES 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Consolida (corollas), p. 127 Consolida (corollas), Delphinium (petals), p. 129 Ranunculus (leaves), p. 155 Ranunculus (leaves), p. 157 Ranunculus ( achenes ), p. 173 Ranunculus (achenes), p. 175 Ranunculus ( achenes ), Adonis ( achenes ), p. 177 Papaver (capsules), p. 223 Papaver (capsules), p. 225 Cruciferae (fruits), p. 251 Cruciferae (fruits), p. 253 Cruciferae (fruits), p. 255 Cruciferae (fruits), p. 257 Cruciferae (fruits), p. 261 Isatis (fruits), p. 291 Isatis (fruits), p. 293 Alyssum (fruits), p. 371 Alyssum (petals, hair-types), p. 373 Barbarea (leaves), Rorippa (fruits), p. 435 Reseda (petals), p. 503
INTRODUCTION This work covers the whole of Turkey as politically constituted today. We have also included most of the eastern islands of the Aegean (part of Greece) for two reasons: because they are floristically closer to Anatolia than to the rest of Greece, and because they are excluded from Flora Europaea; these are the islands shown to the east of the broken line in Map 2 (p. 10). The Flora attempts to include all those species native to Turkey, or which are established aliens; in practice it is often impossible to distinguish between these two groups (for example, with many weeds). Cultivated plants (mostly crop plants or trees) are included only when they have become naturalised, or are so common as to be a feature of the landscape and provide a habitat for the native flora. On the whole we have adopted a fairly broad generic concept for the Flora, although some allowance has had to be made for the preferences of specialists invited to revise particular groups. The species concept adopted is also fairly broad; but as we frequently have only very limited material, many taxa recognised as species may have to be reduced in rank (or included in synonymy) when they become better known. For specific rank we have usually required correlated discontinuous variation in at least two taxonomic characters, although we have sometimes accepted less when traditional usage is strong. We have only recognised two infra-specific categories- subspecies and variety - and these are used somewhat sparingly in accordance with our state of knowledge. Subspecific rank is used for those cases in which variation is only partly discontinuous, or when only a single character is usually coupled with a distinctive distribution (geographical or ecological). Varietal rank is used for two situations: (1) when the taxon is believed to represent a local (rather than a regional) variant; (2) when a striking variant is insufficiently known, so that its status is uncertain. Single-character variants, apparently scattered at random and not forming stable populations (formae), are not formally recognised (unless by accident) in the Flora. It is only in a very few, well-studied groups that cytogenetic information on Turkish material has been sufficient to influence taxonomic treatment. GEOGRAPHICAL DIVISIONS
It was decided to use a grid system, based on two degrees of latitude and longi-
tude, as the primary division for the citation of specimens. The rectangular shape of Turkey means that a grid is particularly convenient, and can be readily memorised. This grid results in Turkey being divided into twenty-nine squares, as shown in Map 1. As can be seen on the same map, we have also used the modem provinces (vilayets) for the citation of specimens. It should be added that some of these vilayets have suffered boundary changes as a result of political vicissitudes, making a grid system particularly useful as a primary reference system. In addition to the squares and vilayets, however, we have recognised certain other, larger divisions that are used for conveying the general distribution of a taxon in Turkey before the citation of specimens. This has been largely based on the editor's field knowledge and our acquaintance with distributional patterns in FOTA
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