Flora of Turkey, Volume 9 9781474466141

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FLORA OF TURKEY VOLUME NINE

to

FRIEDERIKE SORGER

VOLUME NINE

FLORA OF TURKEY and the East Aegean Islands

edited by

P.H.DAVIS

D.SC

Professor of Taxonomic Botany assisted by R. R. Mill PhD and Kit Tan PhD s E Rc Research Assistants

*

EDINBURGH at the

UNIVERSITY PRESS

1985

© 1985

Transferred to Digital Print 2010 Edinburgh University Press 22 George Square, Edinburgh Reprinted 1997,2001,2008

Printed and bound in Great Britain by CPI Antony Rowe, Chippenham and Eastbourne A CIP record is available for this book from the British Library ISBN 0 85224 516 5

PREFACE Volume IX marks the completion of the Flora of Turkey, with the exception of the Supplement. The format of this volume, and the method of production, are the same as those adopted for Volumes VII and VIII. This volume completes the treatment of the Monocotyledons. Three important families, Juncaceae, Cyperaceae and Gramineae, are covered. Together these comprise 165 genera and 692 species, of which 2 small genera and 53 species (7 .8%) are endemic to the Flora area. The much lower endemism in the 'Glumiflorae', relative to families treated in earlier volumes, is largely explained by the fact that the group is mainly wind-pollinated. The Gramineae (512 species in 142 genera) is by far the largest family of Monocotyledons in Turkey, and the fourth largest angiosperm family after Compositae, Leguminosae and Labiatae. The Supplement volume will contain descriptions of new taxa and much additional information. P. H. DAVIS University Department of Botany Edinburgh, August 1985

[v]

ACKNOWLEDGEMENTS We are most grateful to the Science and Engineering Research Council (u .K.) for their continued generous financial support for the Flora of Turkey project. Their grant provides for the salaries of our two loyal Research Assistants, Dr Robert R. Mill and Dr Kit Tan; Aileen Robertson, our Senior Secretary, who has again skilfully set the manuscript on the IBM Composer; and our technician, Margaret Glancey, who once more has dealt with the administrationofincoming and outgoing loans with quiet efficiency. The following botanists (not on the current Flora of Turkey team) have contributed accounts to the present volume: Professor A. Baytop (Istanbul Arundo, Cortaderia, Molinia, Phalaris, Phragmites); Dr M. Dotan (Ankara Gramineae tribes Agrostideae, Aveneae, Phalarideae (excl. Phalaris) & Phleeae, Milium; Zingeria, with Mill); Dr J.R. Edmondson (Liverpool - Poa); Dr I. Markgraf-Dannenberg (Zurich - Festuca); A. Melderis (London- Gramineae tribe Triticeae, excl. Aegilops, Secale & Triticum); Professor 0. Nilsson (Uppsala- Carex); Dr A. Oteng-Yeboah (Sokoto, Nigeria- various genera of Cyperaceae, with Davis or Kit Tan); Professor H. Scholz (Berlin- Gramineae tribe Paniceae; Stipa); Dr P. Smith (Edinburgh- Bromus); DrS. Snogerup (Lund - Juncaceae); and Dr C.A. Stace (Leicester - Gramineae: several genera of tribe Poeae). To all the above I am most grateful. I am indebted to the following artists for their illustrations: Annette Carlstrom (Figures 1, 2), Dr Kit Tan (Figures 3, 4, 7-10, 13-18 (re-arranged and redrawn from drafts provided by Dr I. Markgraf-Dannenberg), 19-21, 24 and 26), Dr Frances K. Hibberd (Figures S, 6), Dr Katherine Kay (Figures 11, 12), Helen Southworth (Figure 22), Dr R.R. Mill (Figure 23) and Christine Huber (Figure 25). The Directors/Keepers of the following herbaria (cited by their standard abbreviations following Index Herbariorum ed. 7) have assisted us by lending material for thepreparationofVolume IX: ANK, B, BM, BRA, BRNM, EGE, G, H, HUB, HUJ, ISTE, ISTF, ISTO, K, L, LD, LE, M, P, TUR, W, WU, and ZT. Dr K.P. Buttler, Dr. A. Huber-Morath and Dr F. Sorger have loaned specimens from their own collections. Other collections made available to us, often by way of gifts to the Edinburgh Herbarium in exchange for names, include those made by the following botanists: Professor Y. Akman, Dr R. Alava, Dr R. An§.in, Y. Aya~igil,ProfessorsA. & T. Baytop,Dr R. vonBothmer, Dr A. ~irpici, Dr T. Ekim, Dr A. Engin, Dr A. Giiner, H. Kehl, Dr N. Kilin~, Dr H. Kiirschner, E. Leblebici, S. Oflas, Drs N. & E. Ozhatay, Professor K.H. Rechinger, Dr E. Se~men, Professor A. Strid, J. Trelawny, Dr E. Tuzlaci, Dr P. Uotila, Dr M. Vural, J. Watson, Dr E. Yurdakulol and the late P. Aellen and 0. Polunin. This is the third volume to have been prepared under the aegis of Professor M.M Yeoman, Head of the Department of Botany at the University of Edinburgh. To Professor D.M. Henderson, Regius Keeper of the Royal Botanic [vii]

Acknowledgements

Garden, particular thanks are due for allowing the continued co-operation of his scientific staff in so many different ways, and for the excellent facilities of the herbarium, library and garden, which combine to provide a first-class base for floristic research. P.H. DAVIS

[viii]

CONTENTS List of Maps, p. x List of Figures, p. xv Abbreviations, p. xix

FLORA

143 Juncaceae, p. 1 144 Cyperaceae, p. 32 145 Gramineae, p. 158 Addenda,p.622 Map Section, p. 627 Index to Volume IX, p. 665 Index to Families and Genera in Volumes I-IX, p. 705

[ix]

LIST OF MAPS 1 Turkish vilayets and grid system, p. xvi 2 Geographical divisions within Turkey, p. xvii DISTRIBUTIONS

Note: To facilitate reference to the main text, the species number is cited in brackets after each binomial in the map captions. 3 !uncus acutus, J. heldreichianus subspp. heldreichianus & orienta/is, J. littoralis, J. maritimus, J. rigidus, p. 627 4 !uncus balticus, J. conglomeratus, J. effusus, J. filiformis, J. inflexus, p.627 5 !uncus compressus, J. gerardi subspp. gerardi & libanoticus, J. tenuis, p.627 6 !uncus capitatus, J. sparganiifolius, p. 628 7 !uncus atratus, J. fontanesii subspp. fontanesii & pyramidatus, J. pygmaeus, J. striatus, J. subnodulosus, p. 628 8 !uncus alpigenus, J. alpinus, J. anatolicus, J. articulatus, J. bulbosus, J. thomasii, p. 628 9 Luzula forsteri, L. sylvatica, L. turcica, p. 629 10 Luzula luzuloides, L. nodulosa, L. spicata, L. sudetica, p. 629 11 Luzula campestris, L. multiflora, L. pallescens, L. pseudosudetica, p.629 12 Bulbostylis tene"ima, B. woronowii, Cyperus esculentus, C. rotundus, Juncellus laevigatus subsp. distachyos, J. pannonicus, p. 630 13 Cyperus fuscus, C. serotinus, Pycreus flavidus, p. 630 14 Fimbristylis annua, F. bisumbellata, F. sieberiana, Pycreus [lavescens, P. sanguinolentus, p. 630 15 · Eleocharisacicularis, E. atropurpurea, E. quinque[lora, p. 631 16 Eleocharis carniolica, E. uniglumis, p. 631 17 Eleocharis mitracarpa, p.631 18 Isolepis cernua, I. setacea, p. 632 19 Schoenoplectus litoralis, S. mucronatus, S. triqueter, p. 632 20 Fuirena pubescens, Schoenoplectus supinus, Scirpus sylvaticus, p. 632 21 Eriophorella pumila, Eriophorum angustifolium, E.latifolium, E. vaginatum, p. 633 22 Blysmus compressus, p. 633 23 Kobresia simpliuscula, Rhynchospora alba, Schoenus nigricans, p. 633 24 Carex appropinquata, C. diandra, C. distachya vars. distachya & phyllostachioidea, C. paniculata subspp. paniculata & szovitsii, p. 634 25 Carex divulsa subspp. coriogyne, divulsa & leersii, p. 634 26 Carex brizoides, C. colchica, C. disticha, C. iraquensis, C. muricata, C. praecox, C. spicata, p. 634

[x]

List of Maps 27 Carex cilicica subspp. cilicica, muglaica & muratica, C. cretica, C. diluta, C. extensa, C. [lava, C. punctata, p. 635 28 Carex eriocarpa, C. kitaibeliana, C. macrolepis, C. pontica, C. tristis, p.635 29 Carex atrata subspp. aterrima & atrata, C. caucasica, C. hartmanii, C. kukkonenii, C. oligantha, p. 635 30 Carex acuta, C. caespitosa, p. 636 31 Carex nigra subspp. alpina, dacica & nigra, p. 636 32 Carex elata subspp. elata & omskiana, C. kurdica, C. orbicularis subsp. kotschyana vars. caucasica & kotschyana, p. 636 33 Brachypodium kotschyi, B. pinnatum, B. retusum, B. sylvaticum, p. 637 34 Trachynia distachya, p. 637 35 Amblyopyrum muticum sensu lato, p. 637 36 Elymus caninus, E. libanoticus, E. longearistatus subsp. sintenisii, E. panormitanus, E. sosnowskyi, E. transhyrcanus, p. 638 37 Elymus erosiglumis, E. [laccidifolius, E. gentryi, E. koshaninii, E. lazicus subspp. divaricatus, lazicus & lomatolepis, E. tauri subsp. tauri, p. 638 38 Elymus elongatus subspp. elongatus, ponticus, salsus & turcicus, E. nodosus subspp. caespitosus, gypsicolus, platyphyllus & sinuatus, p.638 39 Elymus pycnanthus, E. repens subspp. elongatiformis & repens, p. 639 40 Elymus hispidus subspp. barbulatus, hispidus, podpyerae & pulcherrimus, p. 639 41 Elymus clivorum, E. farctus subsp. bessarabicus vars. bessarabicus & striatulus, subsp. farctus vars. farctus & sartorii, subsp. rechingeri, p.639 42 Aegilops cylindrica, A e. speltoides sensu lato, A e. umbellulata, p. 640 43 Aegilops crassa, A e. markgrafii, Ae. triuncialis, p. 640 44 Aegilops biuncialis, A e. columnaris, A e. neglecta, p. 640 45 Triticum baeoticum subspp. baeoticum & thaoudar, T. dicoccoides, T. timopheevi var. araraticum, p. 641 46 Secale anatolicum, S. montanum, S. sylvestre, p. 641 4 7 Secale cereale vars. ancestrale & vavilovii, S. ciliatoglume, Triticum monococcum, p. 641 48 Hordeum gentculatum, H. marinum vars. marinum & pubescens, H. spontaneum, H. violaceum, p. 642 49 Hordeum murinum subsp. glaucum, subsp. leporinum vars. leporinum & simulans, subsp. murinum sensu lato, p. 642 50 Hordeum bulbosum, p. 642 51 Bromus commutatus, B. pseudobrachystachys, B. pumilio, B. racemosus, B. secalinus, p. 643 52 Bromus cappadocicus subspp. cappadocicus & sclerophyllus, B. riparius, B. tomentellus, p. 643 53 Bromus benekenii, B. erectus, B. inermis, B. ramosus, B. sipyleus, p.643 54 Avena barbata subspp. atherantha & barbata, A. clauda, A. eriantha, A. fatua vars. fatua & glabrata, A. wiestii, p. 644 55 Avena sterilis subspp. ludoviciana & sterilis, Helictotrichon pubescens subspp.longifolia & pubescens, p. 644

[xi]

List of Maps 56 Helictotrichon argaeum, H. armeniacum, H. compressum, H. convolutum, H. planiculme, H. pratense, H. versicolor, p. 644 57 Arrhenatherum elatius subsp. elatius, A. kotschyi, A. palaestinum, p. 645 58 Gaudinia fragilis, Gaudiniopsis macra subspp. antalyensis, macra & micropyroides, p. 645

59 Gaudiniopsis huber-morathii, G. quercetorum, G. sorgerae, Ventenata dubia, V. eigiana, V. subenervis, p. 645 60 Parvotrisetum myrianthum, Trisetaria aurea, T. loeflingiana, Trisetum flavescens, T. rigidum, T. sibiricum, T. thospiticum, T. turcicum, p. 646 61 Rostraria berythea, R. cristata vars. cristata & glabriflora, R. hispida, R. obtusiflora subspp. amblyantha & obtusiflora, p. 646 62 Koeleria brevis, K. cristata, K. eriostachya, K. lobata, K. nitidula, K. pyramidata, p. 646 63 Deschampsia caespitosa, D. flexuosa, p. 647 64 Aira caryophyllea, A. elegantissima subspp. ambigua & elegantissima, A. praecox, p. 647 65 Holcus annuus, H. lanatus, p. 647 66 Calamagrostis arundinaceus, C. epigejos, C. pseudophragmites, p. 648 67 Apera baytopiana, A. intermedia, A. interrupta, A. spica-venti, A. triaristata, p. 648 68 Agrostis canina, A. castellana subsp. byzantina, A. gigantea, A. olympica, A. planifolia, p. 648 69 Agrostis balansae, A. capillaris vars. aristata & capillaris, A. lazica, A. stolonifera, p. 649 70 Milium effusum, M. pedicel/are, M. schmidtianum, M. vernale subspp. montianum & vernale, p. 649 71 Zingeria biebersteiniana subspp. biebersteiniana & trichopoda, Z. pisidica, Z. verticillata, p. 649 72 Alopecurus aequalis, A. arundinaceus, A. bulbosus, A. geniculatus, A. pratensis, p. 650 73 Alopecurus aucheri, A. davisii, A. gerardii vars. cassius & gerardii, A. glacialis, A. textilis subspp. textilis & tiflisiensis, A. vaginatus, p. 650 74 Alopecurus creticus, A. laguroides, A. lanatus, A. rendlei, A. setarioides, Phleum crypsoides, P. paniculatum subsp. ciliatum, Pseudophleum gibbum, p. 650 75 Alopecurus myosuroidesvars.latialatus, myosuroides & tonsus, A, utriculatus subspp. anthoxanthoides & utriculatus, Phleum arenarium, P. echinatum, p. 651 76 Phleum alpinum, P. boissieri, P. pratense, P. subulatum subsp. ciliatum & subulatum, p. 651 77 Phleum bertolonii, P. exaratum subspp. aegaeum & exaratum, P. montanum subspp. montanum & serrulatum, P. phleoides, p. 651 78 Festuca arundinacea, F. drymeja, F. gigantea, F. pratensis, F. sclerophylla, p. 652 79 Festuca anatolica subspp. anatolica & borealis, F. artvinensis, F. cyllenica subsp. uluana, F. woronowii subspp. argaea, caucasica, turcica & woronowii, p. 652 80 Festuca cappadocica, F. chalcophaea subspp. chalcophaea & euryphylla,

[xii]

List of Maps

81 82 83 84 85 86 87 88 89 90 91

92

93 94 95 96 97 98 99 100 101 102 103 104 105 106 107

F. decolorata, F. glaucispicula, F. longipanicula, F. rubra subsp. pseudorivularis, F. ziganensis, p. 65 2 Festuca brunnescens, F. bushiana, F. cratericola, F. elwendiana, F. jeanpertii, F. karsiana, F. oreophila, F. xenophontis, p. 653 Festuca cataonica, F. lazistanica subspp. giresunica & lazistanica, F. pinifolia vars. phrygia & pinifolia, F. punctoria, F. ustulata, p. 653 Festuca callieri subspp. callieri & zederbaueri, F. valesiaca, Intermediates, p.653 Festucaadanensis, F. airoides, F. beckeri, F. ilgazensis, F. pseudosupina, F. sipylea, p. 654 Festuca paphlagonica subspp. paphlagonica & villosula, F. polita, F. pontica, p. 654 Bellardiochloa argaea, B. carica, B. polychroa, B. violacea, p. 654 Lolium persicum, L. subulatum, L. temulentum vars. arvense & temulentum, p. 655 Lolium rigidum vars. rigidum & rottbollioides, p. 655 Poa araratica, P. caucasica, P. cenisia, P. davisii, P. jubata, P. longifolia, P. psychrophila, P. speluncarum, p. 655 Eremopoa attalica, E. capillaris, E. mardinensis, E. multiradiata, E. persica, E. songarica, Hyalopoa pontica, Nephelochloa orienta/is, p. 656 Catabrosella araratica, C. fibrosa, C. gillettii, C. parviflora subspp. calvertii & parviflora, C. variegata vars. leiantha & variegata, Paracolpodium colchicum, p. 656 Puccinellia bulbosa subspp. bulbosa & caesarea, P. distans subspp. distans & sevangensis, P. festuciformis, P. grossheimiana, P. intermedia, p.656 Puccinellia ciliata, P. convoluta, P. gigantea, P. koeieana subsp.anatolica, p.657 Catabrosa capusii, Colpodium versicolor, Cynosurus cristatus, C. echin· atus, C. effusus, p. 657 Briza humilis, B. marcowiczii, B. maxima, B. media, p. 657 Parapholis filiformis, P. incurva, p. 658 Ammochloa palaestina, Parapholis marginata, P. pycnantha, Pholiurus pannonicus, p. 658 Hainardia cylindrica, Sesleria alba, S. araratica, S. phleoides, p. 658 Melica eligulata, M. min uta, M. persica subspp. canescens, inaequiglumis, jacquemontii & persica, M. uniflora, p. 659 Melica ciliata subspp. ciliata sensu lato, magnolii & transsilvanica, p. 659 Melica altissima, M. penicillaris, M. picta, p. 659 Glyceria arundinacea, G. fluitans, G. maxima, G. nemoralis, p. 660 Stipa bromoides, S. caragana, S. kurdistanica, S. pontica, S. tirsa, p. 660 Stipa capensis, S. ehrenbergiana, S. hohenackeriana vars. assyriaca & hohenackeriana, p. 660 Stipa capillata, S. holosericea, p. 661 Piptatherum blancheanum, P. laterale subsp. laterale, P. miliaceum subspp. miliaceum & thomasii, P. virescens, p. 661 Piptatherum coerulescens, P. holciforme subsp. holciforme vars.glabrum & holciforme & subsp.longiglume var. longiglume, p. 661

[xiii]

List of Maps

108 Aristida adscensionis, A. caerulescens, Danthonia alpina, D. decumbens, Schism us arabicusf Stipagrostis plumosa, p. 662 109 Eragrostis cilianensis, E. collina, E. pilosa, p. 662 110 Eragrostis barrelieri, E. curvula, E. minor, p. 662 111 Cleistogenes serotina subsp. serotina, Dactyloctenium aegyptium, Eleusine indica, Sporobolus virginicus, p. 663 112 Crypsis aculeata, C. acuminata subspp. acuminata, ambigua & borszczowii, C. alopecuroides, Tragus racemosus, p. 663 113 Crypsis faktorovskyi, C. schoenoides, p. 663 114 Saccharum ravennae, S. strictum, Spodiopogon pogonanthus, p. 664 115 Andropogon distachyos, Bothriochloa bladhii, B. ischaemum, Dichanthium annulatum, Hyparrhenia hirta, p. 664 116 Arthraxon hispidus, Chrysopogon gryllus subsp. gryllus, Hemarthria altissima, Phacelurus digitatus, Themeda triandra, p. 664

[xiv]

LIST OF FIGURES 1 Juncaceae: flowers with ripe fruit, p. 7 2 Juncaceae: flowers with ripe fruit, p. 28 3 Cyperaceae: glumes and spikelets, p. 41 4 Cyperaceae: glumes, perianths and nuts, p. 57 5 Carex utricles and glumes, p. 83 6 Carex utricles and glumes, p. 123 7 Character states in Turkish Gramineae (inflorescence types), p. 177 8 Character states in Turkish Gramineae (vegetative and spikelet characters), p.l79 9 Elymus spikelets, p. 210 10 Elymus spikelets, p. 215 11 Bromus panicles and spikelet morphology, p. 281 12 Bromus panicles and vegetative morphology, p. 293 13 Festuca: transverse sections of leaves, p. 405 14 Festuca: transverse sections of leaves, p. 413 15 Festuca: transverse sections of leaves, p. 417 16 Festuca: transverse sections of leaves, p. 427 17 Festuca: transverse sections of leaves, p. 429 18 Festuca: transverse sections of leaves, p. 439 19 Vulpia spikelets, p. 453 20 Catapodium inflorescences, p. 463 21 Catapodium and Cutandia spikelets, p. 465 22 Poa lemmas, p. 477 23 Gramineae: spikelet morphology, p. 489 24 Puccinellia spikelets, p. 507 25 Stipa lemmas, p. 549 26 Gramineae: spikelet morphology, p. 571

[xv]

I

s

4

3

2

.........

•

I

1

'

r-

8

200

26° MAP

2..

3ft'

1 Turkish vilayets and grid system

Jr

34.

36.

38.

.o~o·

km

.ol2"

I

.....

1361

........,

~. ....

'--'

MAP

2 Geographical divisions within Turkey (seep. xviii)

GENERAL DISTRIBUTION IN TURKEY

areas numbered as on Map 2, p. xvii Turkey-in-Europe (1) North Anatolia (2-4) West Anatolia (5-7)

South Anatolia (8-15) Central Anatolia (16-20) East Anatolia (21-25)

The following terms are also used: Outer Anatolia: areas 2-15 Inner Anatolia: areas 16-25 North-West Turkey: areas 1, 2 & 5, together with vilayet Bilecik; shaded on map North-East Anatolia: areas 4 & 24, together with part of vilayet Giimii~ane in area 21; shaded on map South-West Anatolia: areas 7, 8, 9, 12 & 17; shaded on map South-East Anatolia: area 23, together with parts of vilayets Van and Bitlis in area 25 ; shaded on map North Turkey: N. Anatolia & Turkey-in-Europe West Turkey: W. Anatolia & Turkey-in-Europe Islands: E. Aegean Islands; E. of dotted line on Map 2 For treatment of external distributions, see Volume I, Introduction, p.l3 and Map 3.

[xviii]

ABBREVIATIONS This list does not include abbreviations of authorities, books and journals, or of herbaria; for the first we have followed general usage, for the last Index Herbariorum, and for the remainder we have used abbreviations that should be readily comprehensible. As asterisk '*' before a generic or species name indicates that the taxon only occurs in Turkey as an introduced or cultivated plant. An exclamation mark '!' following a cited specimen shows that the specimen has been seen by the author of the account; following a synonym it indicates that type material of the synonymous taxon has been seen. A bracketed exclamation mark '(!)' following a cited specimen is used to show that a specimen has been identified by the editorial team at Edinburgh, and was not seen by the author of the account (see Prefaces to Volumes VI and VII). agg. Alt. anon. Azn. Bal. Boiss. Bornm. Cand. C. c. cf. Contandr. cv. D. d. Da. De. diam. E. e.g. Ehrend. Euro-Sib. excl. f. Fl.

Fors.-Maj. Fr.

fr. G.

aggregate Altitude collector unknown G.V. Aznavour B. Balansa P.E.E. Boissier J .F .N. Bornmiiller C.A. Candargy Central circa (about) confer, conferatur (compare) J. Contandriopoulos cultivar (cultivated variety) P.H. Davis district Da~, Dati, or Datlari (Turkish for mountain or mountain range) Dere, Deresi (Turkish for valley or small river) diameter East exempli gratia (for example) F. Ehrendorfer Euro-Siberian excluding figure; forma Flowering period (in months) C.J. Forsyth-Major Fruiting period (in months) fruit Gol, Golii (Turkish for lake)

[xix]

Abbreviations F .H. von Randel-Mazzetti M. Haradjian C. Haussknecht T. von Heldreich holotype A. Huber-Morath in the same place Icon (illustration) including lrano-Turanian Islands isotype Iter Leydenense lectotype on the page previously cited A. Manissadjian Mediterranean mountain North near new series observed, observation in the work previously cited photograph or microfiche U. Plitmann pro parte (in part) pro synonymum (as a synonym) quod vide (which see) K.H. Rechinger fil. H. Runemark South P .E.E. Sintenis sea level sine numero (without a number) S. Snogerup F. Spitzenberger J.D.A. Stainton & D.M. Henderson tabula (plate) unlocalised J.S.C.D. d'Urville West Yayla, Yaylasi (Turkish for summer pasture) F. Yaltirik more or less x 'times' (e.g. 3 x 4 mm = 3 mm long by 4 mm wide) or denoting a hybrid IJ.m micron(= O.OOlmm)

Hand.-Mazz. Haradj. Hausskn. Heldr. holo. Hub.-Mor. ibid. Ic incl. Ir.-Tur. Is. iso. It. Leyd. lecto. loc. cit. Maniss. Medit. mt. N. nr n.s. obs. op. cit. photo Plitm. p.p. pro syn. q.v. Rech. Runem. S. Sint. s.l. s.n. Snog. Spitz. Stn. & Hend. t. unloc. d'Urv. W. Y. Yalt. ±

[xx]

SPERMATOPHYTA (concluded)

ANGIOSPERMAE

MONOCOTYLEDONES concluded 143. J U N C A C E A E Perennial or annual herbs. Leaves long and narrow; phyllotaxis 1/3. Inflorescence of monochasial cymes (sometimes condensed into heads), usually many-flowered, rarely flowers solitary. Flowers actinomorphic, hermaphrodite, anemophilous, usually protogynous. Perianth segments 6, in 2 whorls, scarious, light to dark brown or herbaceous (in Turkey). Stamens free, in (1-)2 whorls of 3; anthers basifixed, introrse. Ovary superior, of 3 fused carpels, 1- or 3-locular; stigmas 3; ovules 3 or numerous. Fruit a dehiscent, loculicidal capsule; embryo small; seeds endospermic, often with a basal appendage; endosperm helo bial. Buchenau, F. 1880. K.ritisches Verzeichniss aller bis jetzt beschriebenen Juncaceen nebstDiagnosenneuer Arten. Bremen. Buchenau, F.l890. Monographia Juncacearum. Leipzig. Buchenau, F. 1906. Juncaceae. In Engler, Pflanzenreich 25 (IV.36):1-284. 1. Leaves not ciliate; capsule many-seeded 1. Leaves ciliate, at least when young; capsule 3-seeded

1. Juncus 2. Luzula

1. JUNCUS L. S.

S N 0 G E R UP

Perennials, usually rhizomatous, or annuals. Leaves glabrous, like the rest of the plant; sheaths normally open. Flowers either ebracteolate and crowded in monochasial cymes (often condensed into heads), or-solitary and subtended by one 2-fid adaxial bracteole and two simple abaxial bracteoles forming a basal involucre. Pollen shed in tetrads. Fruit a many-seeded capsule. Seeds numerous, without or with 1 or 2 scarious appendages. Nilsson, 5. & Snogerup, S. 1971-2. Drawings of Scandinavian plants 45-80: !uncus L. Bot. Not. 124:1-8, 179-186,311-316, 443-441; op. cit. 125:131-138, 203-211. Snogerup, S. 1978. A revision of the !uncus atratus group. Bot. Not. 131:189-196. Novikov, V. 1978. Synopsis specierum generisJuncus L. ex Asiae Mediae. Novit. Syst. Pl. Vase. 15:77-93. Fernandez-Carvajal,M.C. 1981. Revisi6n del genero !uncus L. en la Peninsula Iberica I. Categonas supraespeciffcas y claves para las especies. Anales Jard. Bot. Madrid 38:79-89. Fermindez-Carvajal, M.C. 1981. Revisi6n del genero Juncus L. en la Peninsula Iberica II. Subgeneros !uncus y Genuini Buchenau. Op. cit. 38:417-467. Fernandez-Carvajal, M.C. 1982. Revisi6n del genero Juncus L. en la Peninsula Iberica III. Subgeneros Subulati Buchenau, Pseudotenageia K.recz. & Gontsch. y Poiophylli Buchenau. Op. cit. 39:79-151. Very variable in vegetative characters. Most specimens are easily referred to their subgenus by the different leaf structure and arrangement of the leaves.

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143. J U N C A C E A E The relative lengths of filaments and anthers given in the following account refer to specimens in late flowering or fruiting states. Herbarium material has been very frequently misidentified. 1. Inflorescence branches ending in single flowers, each with two involucra! bracteoles Group A 1. Inflorescence branches ending in dense, few-to many-flowered heads; each flower with a bract but lacking bracteoles Group B Group A 1. Leaves of adult plants terete; perennial 2. Flowering stems with several cauline leaves (Subgen. Subulati) 6. subulatus 2. Leaves solitary on specialised shoots from rhizome, and as bracts of inflorescence (Subgen. Genuini) 3. Plant often slender, not or laxly caespitose;inflorescence not more than I-S-flowered 4. Ripe capsule equalling perianth; bract at least 1/3 as long as stem 10. filiformis 4. Ripe capsule exceeding perianth; bract less than 1/3 as long as stem 11. balticus 3. Plant robust, densely caespitose; inflorescence many-flo)Vered 5. Stem almost smooth, with 40-70 weak striae below inflorescence 8. effusus 5. Stem with 10-30 ridges below inflorescence 6. Capsule acute to obtuse; pith interrupted 7. inflexus 9. conglomeratus 6. Capsule retuse to obtuse; pith continuous 1. Leaves flat or rarely subterete and then canaliculate or with much enlarged cells of dorsal epidermis; annual or perennial 7. Perennials with a usually strong rhizome (Subgen. Pseudotenageia) 8. Perianth segments apiculate to aristate 14. tenuis 8. Perianth segments obtuse 9. Anthers not more than 2 x filaments; capsule usually exceeding perianth 12. compressus 13. gerardi 9. Anthers 2.5-6 x filaments; capsule± equalling perianth 7. Annuals without a rhizome, !-stemmed or usually tufted (Subgen. Poiophylli) 10. Capsule spherical 11. Inflorescence occupying less than 1/2 stem height; auricles well developed 15. tenageia 11. Inflorescence occupying more than 1/2 stem height; auricles absent on lower leaves 16. sphaerocarpus 10. Capsule ellipsoid to ovoid 12. Seeds minutely but markedly reticulate (microscope!); inner perianth segments scarious except for midrib 17. rechingeri 12. Seeds smooth to finely striate (microscope!); inner perianth segments herbaceous in central part

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1. Juncus 13. Capsule ellipsoidal to prismatic; inner perianth segments obtuse to acute 14. Plant usually 10-20 em tall with some leaves 5-10 em; outer perianth segments 5-8.5 mm, often much exceeding inner; seeds 0.3-0.4 20. hybridus mm 14. Plant usually 3-15 em tall with all leaves rather short; outer perianth segments 3.5-6 mm, slightly exceeding inner; seeds 0.35-0.5 mm 21. turkestanicus 13. Capsule ovoid;inner perianth segments apiculate to acute 15. Capsule 3-5 mm; anthers 1/3-1 x filaments; plant very variable in 18. bufonius size and habit 15. Capsule 2.5-3 mm; anthers 1/4-1/3 x filaments; plant small, thinstemmed or with flowers aggregated near ground 19. minutulus Group B 1. Small annuals, !-stemmed or tufted 22. capitatus 2. Capsule ovoid-globose; leaves without auricles 26. pygmaeus 2. Capsule narrowly oblong; leaves with auricles 1. Perennials with a rhizome or rarely bulb-like stem bases, or long floating stems, or tall and caespitose 23. sparganiifolius 3. Leaves flat, broad, entirely non-septate 3. Leaves terete 4. Leaves non-septate, pungent, with several centrally placed vascular bundles; auricles absent (Subgen. !uncus) 5. Inner perianth segments without apical auricles;rhizome strong, creeping 4. maritimus 6. Anthers c. 2 x filaments; capsule equalling perianth 5. rigidus 6. Anthers 2.5-5 x filaments; capsule exceeding perianth 5. Inner perianth segments with broad apical auricles; rhizome absent or inconspicuous, plant densely caespitose 1. acutus 7. Capsule 4-6 mm, with 80-120 seeds, conical at apex 7. Capsule 2.5-4 mm, with 25-75 seeds, conical or trigonous at apex 2.littoralis 8. Capsule trigonous at apex; flowers dark brown 3. heldreichianus 8. Capsule conical at apex; flowers light brown 4. Leaves septate, not pungent, uni- or pluritubulose, tubes empty or with loose aerenchyma; auricles present (Subgen. Septati) 9. Plant without a rhizome, with bulb-like stem bases or forming long 25. bulbosus floating branches only 9. Plant rhizomatous, without bulb-like stem bases 10. Leaves pluritubulose, both cauline and solitary from rhizome 24. subnodulosus 10. Leaves unitubulose, all cauline 11. Perianth segments narrow, acuminate-cuspidate, dark brown to blackish; leaves with 7-17 longitudinal ridges 12. Capsule acutely tapering, its beak 0.4-1.2 mm 13. Inflorescence consisting of one dense cluster of heads or flowers; 34. 'anatolicus perianth segments equal or subequal

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143. J U N C A C E A E 13. Inflorescence consisting of 15-50(-200) usually quite separate heads; inner perianth segments longer 31. atratus 12. Capsule broadly obtuse, its beak 0.2-0.4 mm 14. Leaves usually 1-2 mm diam.; stamens 3 or 6; inflorescence 33. alpigenus usually of one cluster 14. Leaves usually 2-4 mm diam.; stamens 6; inflorescence of several clusters or many discrete heads 32. thomasii 11. Perianth segments ovate or narrowly ovate to lanceolate, light to dark brown, greenish or reddish, obtuse to acuminate; leaves smooth or with many small ridges 15. Capsule obtuse with short mucro (0.1-0.4 mm) 16. Perianth segments 2.5-3.5 mm, outer ones acute 29. articulatus 16. Perianth segments 2-2.5 mm, outer ones obtuse though often with a mucro 30. alpinus 15. Capsule attenuate, or if abruptly contracted apically, then with a 0.5-1.5 mm mucro 17. Rhizome weakly developed and often inconspicuous; creeping stems frequent 28. fontanesii 17. Rhizome well developed; creeping stems absent or occasional 18. Anthers 2-3 x filaments; leaves finely but conspicuously striate 27. striatus 18. Anthers 1-2 x filaments; leaves smooth or when dry weakly and irregularly striate 29. articulatus Subgen. Juncus. Perennials. All leaves basal, terete, pungent; auricles absent. Involucral bracteoles absent; flowers in heads. Seeds with appendages.

1. J. acutus L., Sp. Pl. 325 (1753). Ic: Roles, Brit. Fl. ill. 4:25,f.l548 (1965); Rech. fil., Fl. Iranica 75: t. 2 (1971). Figure 1. Map 3. Plant densely caespitose with poorly developed rhizome; intravaginal shoots present. Stems (15-)50-150 em, rigid, with 2-5 basal leaves. Inflorescence many-flowered, usually dense, globose; lowest 2 bracts well developed though usually short. Perianth segments subequal, oblong; outer obtuse or acute; inner with broad scarious apical auricles. Stamens equalling or slightly shorter than perianth; anthers 1.5-2 mm, several times longer than filaments. Capsule ovoid, 4-6 mm, much exceeding perianth, with broadly conical apex, shortly mucronate. Seeds 80-120 per capsule, 1.5-2.5 mm, with equal appendages. Fl. 3-5. Coastal sands, salt or freshwater marshes, s.l.-150 m. Described from seaside marshes in England, France & Italy. Lectotype designated here: Hb. Linn. 449/1, left-hand specimen! Turkey-in-Europe, Outer Anatolia, Islands. A2(E) Istanbul: Kii~k