Ορχιδέες του Εθνικού Πάρκου Οροσειράς Ροδόπης

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Ορχιδέες του Εθνικού Πάρκου Οροσειράς Ροδόπης Orchids of Rodopi Mountain-Range National Park ΣΠΥΡΟΣ ΤΣΙΦΤΣΗΣ ΙΩΑΝΝΗΣ ΤΣΙΡΙΠΙΔΗΣ ΚΩΣΤΑΣ ΒΙΔΑΚΗΣ ΦΟΡΕΑΣ ΔΙΑΧΕΙΡΙΣΗΣ ΟΡΟΣΕΙΡΑΣ ΡΟΔΟΠΗΣ - MANAGEMENT BODY OF RODOPI MOUNTAIN-RANGE NATIONAL PARK

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Transcript of Ορχιδέες του Εθνικού Πάρκου Οροσειράς Ροδόπης

  • ISBN978-618-80276-0-2

    Co-Financed By Greece and the European Union

    European UnionEuropean Regional Development Fund

    Operational Programme Environment and Sustainable

    Development

    . - , , . This book presents fifty nine native orchid species and subspecies of Rodopi Mountain-Range National Park. These tiny plants form a natural reserve of great ecological importance for the region. Getting to know them is our duty but also a good opportunity for aesthetic education.

    Orchids of Rodopi Mountain-Range National Park

    - MANAGEMENT BODY OF RODOPI MOUNTAIN-RANGE NATIONAL PARK

    O

    rchi

    ds o

    f Rod

    opi M

    ount

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    Ran

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    ark

  • Orchids of Rodopi Mountain-Range National Park

    - TEXTS

    - SPYROS TSIFTSIS

    - IOANNIS TSIRIPIDIS

    - PHOTOGRAPHS

    - COSTAS VIDAKIS

    - MANAGEMENT BODY OF RODOPI MOUNTAIN-RANGE NATIONAL PARK

    - MESOCHORI, PARANESTI GREECE

    2012

  • , , , , , , ,

    , , ,

    221/1993 , .

    All rights reserved. No text, images or other part of this publication may be copied, reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic,

    mechanical, photocopying, theatrical, cinematographic, radio, television or otherwise, without the prior written permission of the authors, the photographer and the publisher.

    - EDITORIAL COORDINATION

    & .. - PHOTO/GRAPHS STUDIO E.E.

    - CHIEF SCIENTIFIC ADVISER

    - SPYROS TSIFTSIS

    Y - SCIENTIFIC ADVISER

    - IOANNIS TSIRIPIDIS

    - GREEK TEXT EDITING

    . - CHRISTOS P. PHARACLAS

    - ENGLISH TEXT EDITING

    - NIKOS THOMAIDIS

    H - ART DIRECTOR

    - SOFIA SIGGIRIDOU

    - TEXTS

    - SPYROS TSIFTSIS - IOANNIS TSIRIPIDIS

    - PHOTOGRAPHS

    - COSTAS VIDAKIS (PHOTO/GRAPHS STUDIO E.E.)

    - PRINTING

    GRAPHIC ARTS - BAXAS GRAPHIC ARTS

    2012 MANAGEMENT BODY OF RODOPI MOUNTAIN-RANGE NATIONAL PARK 660 35 Mesochori, Paranesti, Greece | Tel: +30 2524 021030 - Fax: +302524022165 e-mail: [email protected] | Website: http://www.fdor.gr

    Co-Financed By Greece and the European Union

    European UnionEuropean Regional Development Fund

    Operational Programme Environment and Sustainable

    Development

    ISBN978-618-80276-0-2

  • - Contents

    Foreword by the president 5

    The Rodopi Mountain-Range National Park 7

    Vegetation and diversity in Rodopi Mountain-Range National Park 13

    Vegetation types reaching their southernmost distribution in Rodopi Mountain-Range

    National Park 15

    Vegetation zonation in Rodopi Mountain-Range National Park 26

    Grassland vegetation in Rodopi Mountain-Range National Park 36

    The importance of vegetation conservation in Rodopi Mountain-Range National Park 43

    H

    The Orchid Family 47

    The evolution of orchids 49

    The morphology of orchids 51

  • The biology of orchids 56

    The importance of Rodopi Mountain-Range National Park for orchids 63

    Orchids of Rodopi Mountain-Range National Park 69

    Index of Greek common names 190

    Index of English common names 191

    Index of scientific names 192

    Selected bibliography 196

  • Foreword

    . , -

    .

    , -

    ,

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    . ,

    ,

    .

    -

    -

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    ,

    , , , ,

    . , -

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    . .

    The Rodopi Mountain-Range National Park is, in every

    respect, a treasury of natural wealth; a wealth whose

    ecological and aesthetic values outweigh its economic.

    Given that it has preserved unique flora species of ear-

    lier geological periods and that it is the southernmost

    area of distribution of many other species, the National

    Park can be considered as a natural museum. However,

    the exhibits of this museum are living and trigger a

    deep interest in protecting and preserving them. Getting

    to know the National Park could convince even the

    most sceptical or indifferent visitor that this is a paradise

    whose beauty is undeniably exceptional.

    Undoubtedly, one of the most interesting and fascinat-

    ing flora representatives of Rodopi Mountain-Range

    -both from an aesthetic point of view and a scientific

    perspective- is the orchid family. The plants of this fam-

    ily exhibit a wide diversity in form, colour, scent, site

    preference, reproductive methods and relationship with

    the environment. This publication is the result of long

    and laborious cooperation of specialists and aims to fill

    a bibliographic gap in academic research, yet also to

    provide to the general public a comprehensive guide

    for the identification of local orchids and an oppor-

    tunity for raising environmental awareness. I sincerely

    wish and certainly hope that this remarkable book will

    fulfil its objectives.

    Dr Nikolaos D. Avtzis

    President of the Board of DirectorsManagement Body of Rodopi Mountain-Range National Park

  • / Orchids of Rodopi Mountain-Range National Park | 7

    ()

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    173.115 ha

    .

    , ,

    -

    .

    -

    , (

    ) .

    (7)

    NATURA 2000 (2)

    -

    . , :

    - .

    .

    he Rodopi Mountain-Range National Park (RMRNP) extends in parts of the Regional Units of Drama and Xanthi in the Region of Eastern Macedonia and Thrace,

    Greece.

    The Park, as its name suggests, is located in Rodopi

    Mountain Range, the second longest in Greece (after

    Pindos). It extends from Mt. Falakron and River Nestos in

    the northeast to the Bulgarian border and the mountain-

    ous area of Xanthi. Its total area is 173,115 ha; it is one of

    the largest National Parks in mainland Greece.

    The special features of the area, the geomorphologic limita-

    tions and legal restrictions as well as the lack of large-scale

    investment were the reasons why the Parks ecosystems

    have remained relatively undisturbed until today.

    The designation of the area as a National Park is a recogni-

    tion of its ecological and aesthetic value. In fact, the Park

    comprises of a number of protected sites that have been

    designated at different times during the past decades, the

    main of which are seven sites of the European Ecological

    Network NATURA 2000 and two sites that have been

    designated as Preserved Monuments of Nature under

    national Law. These main sites are:

    Preserved Monuments of Nature

    TSICHLA - DRIMOS

    PARTHENO DASOS FRAKTOU KENTRIKIS RODOPIS

    N. DRAMAS

    The Rodopi Mountain-Range National Park

  • NATURA 2000

    (SPAs)

    (GR1140008)

    (GR1140009)

    (SCI)

    - (GR1120003, .

    3.492 ha)

    (GR1140001, . 1.090 ha)

    () (GR1140002, . 6.715 ha)

    , (GR1140003, . 7.447 ha)

    (GR1140004, . 9.845 ha)

    . 40379/2009

    (...)

    NATURA 2000 sites

    Special Protection Areas (SPAs)

    GR1140008: KENTRIKI RODOPI KAI KOILADA NESTOU

    GR1140009: OROS FALAKRO.

    Sites of Community Importance (SCI)

    OROS CHAINTOU - KOULA KAI GYRO KORYFES

    (GR1120003, c. 3,492 ha)

    DASOS FRAKTOU (GR1140001, c. 1,090 ha)

    RODOPI (SIMYDA) (GR1140002, c. 6,715 ha)

    PERIOCHI ELATIA, PYRAMIS KOUTRA (GR1140003, c.

    7,447 ha)

    KORYFES OROUS FALAKRO (GR1140004, c. 9,845 ha)

    Under Joint Ministerial Decision Number 40379/2009, four

    wider zones are applied in Rodopi Mountain-Range National

    Park. These are:

  • 10 | / Orchids of Rodopi Mountain-Range National Park

  • / Orchids of Rodopi Mountain-Range National Park | 11

    ( -

    ):

    () ,

    () ,

    ()

    .

    (1, 2 3)

    , -

    - ,

    .

    (7 )

    ,

    , (Alnus

    incana subsp. incana),

    ,

    .

    , (5)

    (1)

    .

    ,

    ,

    -

    .

    (a) Absolute Nature Reserve Areas,

    (b) Nature Reserve Areas, and

    (c) Special Management Areas, Sustainable Use and

    Development Areas.

    In each zone, a different protection regime is applied. The

    stricter restrictions apply to zone (a) and the laxer to zone

    (c).

    Zone (a) (Absolute Nature Reserve Areas) comprises of

    three sub-zones (A1, A2 and A3). These include the Virgin

    Forest of Frakto, the Preserved Monument of Nature

    Tsichla, and the area Gyftokastro - Koula. These sites

    were already designated under previous national Laws.

    The Parks Nature Reserve Areas (wider zone b) include

    sub-zones that surround the above-mentioned Absolute

    Nature Reserve Areas. These include the two shallow and

    flat bogs located near the forestry station of Elatia, the

    riparian forest of Grey or Speckled Alder (Alnus incana

    subsp. incana) in the area called Ruins of Vathyrema vil-

    lage, the bed of Stravorema stream and the wet meadows

    along it.

    Finally, the third wider zone (c) includes five Special

    Management Areas and a Sustainable Use and Development

    Area.

    The unique natural elements and the landscape of the

    National Park could trigger sustainable development with

    great benefits for the wider region.

  • (i) taxa Euro+Med,1 Greuter et al.2, Greuter et al.3, Greuter et al.4, Strid5 Strid & Tan6.

    (i) The nomenclature of plant taxa follows Euro+Med,1 Greuter et al.2, Greuter et al.3, Greuter et al.4,

    Strid5 and Strid & Tan6.

    - -

    Vegetation and diversity in Rodopi Mountain -Range National Park

    -

    .

    -

    . -

    (Picea abies subsp. abies(i)), (Betula pendula)

    (Alnus incana subsp. incana), -

    . [7, 8, 9, 10, 11, 12]

    he Rodopi Mountain-Range National Park is highly diverse, both

    in terms of plant species and vegetation typology. Some plant

    species -and the vegetation types they form or participate in- are

    rare or unique to Greece and the Mediterranean. Examples are the

    forests of Norway spruce (Picea abies subsp. abies(i)), birch (Betula

    pendula) and grey alder (Alnus incana subsp. incana). All the

    above-mentioned species -and the forests they form- are mainly or

    exclusively found in the mountain range of Rodopi.[7, 8, 9, 10, 11, 12]

    T

  • -

    -

    ;

    ( )

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    ,

    .

    What are the reasons behind this extreme diversity of the park

    in the context of Greece and the wider Mediterranean area?

    The answer lies in three major factors: (i) the geographical

    position of Rodopi (latitude and distance from the sea), (ii)

    the geological history of the area, and (iii) the migration

    history and the routes of plant species (especially during

    the glacial periods). To the above factors one can add the

    history of human presence. Indeed, different types of land

    use and management practices and, subsequently, the almost

    complete desertion of the area by its people have played

    a decisive role in the enhancement of vegetation and the

    enrichment of diversity.

  • / Orchids of Rodopi Mountain-Range National Park | 15

    -

    (41 )

    , (..

    , , ).

    ,

    .

    .

    (Picea abies subsp. abies)

    , .[13,10,11]

    he middle geographical latitude (41 north) and its long

    distance from the sea (in comparison to other Greek

    mountains, e.g. Athos, Paggaio, Olympos) position Rodopi

    next to a biogeographical border, which marks the transi-

    tion from a Mediterranean to a more or less continental

    climate. This climatic transition is the major reason why

    many plant species and vegetation types reach their south-

    ernmost distribution area in Rodopi. One of the most

    characteristic example is the Norway spruce forest at the

    northernmost part of Greek Rodopi in the areas of Elatia,

    Frakto and Lepida.[13,10,11]

    -

    Vegetation types reaching their south-ernmost distribution in Rodopi Moun-tain-Range National Park

    T T

  • [14,15] . - . - , .

    Norway spruce extends in large parts of Eurasia[14,15] with continental climate. Thus, the occurrence of this species in Rodopi is an indication of the transitional climate (from Mediterranean to continental). Several species, found in Norway spruce forests in other parts of its worldwide distribution, are also found in Rodopi.

  • / Orchids of Rodopi Mountain-Range National Park | 17

    , Vaccinium myrtillus, V. vitis-idaea, Avenella flexuosa,

    Luzula luzulina, Pyrola minor, Pyrola media, Pyrola rotundifolia, Sorbus aucuparia subsp. aucuparia,

    Corallorhiza trifida, Rosa pendulina, Moneses uniflora, Neottia cordata, Goodyera repens ..[11,16,17] -

    (ii) .

    Typical species of Norway spruce forests, found all over its distribution space, such as Vaccinium myrtil-

    lus, V. vitis-idaea, Avenella flexuosa, Luzula luzulina, Pyrola minor, Pyrola media, Pyrola rotundifolia, Sorbus

    aucuparia subsp. aucuparia, Corallorhiza trifida, Rosa pendulina, Moneses uniflora, Neottia cordata,

    Goodyera repens etc.[11,16,17] are also found in plant societies of Norway spruce forests (ii) in Rodopi.

    (ii) .

    (ii) A plant society is a special vegetation type, which consists of particular flora species and occurs in various areas with similar ecological conditions.

    Moneses uniflora Rosa pendulina

    Vaccinium myrtillus Vaccinium vitis-idaea

  • 18 | / Orchids of Rodopi Mountain-Range National Park

    ,

    ,

    (Betula pendula).

    ,

    .[14,18]

    , ,

    , -

    .12]

    ,

    .

    -

    .

    ,

    , -

    , (iii).

    Another example of a vegetation type that is rare to the

    Mediterranean area and Greece comprises of birch (Betula

    pendula) forests. Birch extends from central and northern

    Europe up to central Asia. In the Mediterranean, and in

    the northern mountainous areas of Greece in particular, it

    occurs sporadically.[14,18] However, there is only one site in

    which this species forms extended forest stands and this

    is the so-called Kalyvia Karyotou in Rodopi.[12] A unique

    morphological characteristic of Betula pendula is the white

    colour of its bark, which makes it distinct from any other

    tree species occurring naturally in Greece. The presence

    of birch forests in Rodopi may be attributed to the areas

    climate (transitional, as mentioned above) as well as to

    the management practices in the past century. Birch is a

    pioneer species, establishing itself in areas not covered by

    forests. It is also a species favoured by disturbances(iii).

  • / Orchids of Rodopi Mountain-Range National Park | 19

    ;

    -

    20

    , .[19]

    20 .

    1945

    . -

    50-60 -

    85%.

    ( ), -

    ,

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    .[20] ,

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    (iii) . (.. , ), (.. , ) (.. , ).

    Factors favouring the distribution of birch in Rodopi

    Mountain-Range National Park

    The occurence of birch forests in Rodopi Mountain-Range

    National Park may be attributed to the favourable climate

    as well as to the periodic disturbances caused by the

    nomadic type of pastoralism practiced by the Sarakatsani

    people before the middle of the 20th century.[19]

    Livestock grazing and grassland burning (practiced by the

    Sarakatsani people every autumn in order to improve their

    productivity) are the two major reasons for the preserva-

    tion of grasslands in Rodopi up to the middle of the 20th

    century.

    However, animal husbandry in the area has been heavily

    reduced after 1945 and many of the villages have been

    deserted. The significant reduction of disturbances during

    the last 50-60 years allowed reforestation of many naked

    (open) areas and led to the gradual increase of forest cover

    to a percentage above 85%.

    Reforestation of naked areas usually occurs in accordance

    to a process known as ecological (or vegetation) succes-

    sion. This succession process depends on the time period

    elapsed after the disturbance event, the species occurring

    in the wider area (the so-called available species pool)

    and the ecological conditions of the site.[20] At the initial

    stages of succession pioneer species establish first; these

    form a forest and improve the ecological conditions of

    the site (e.g. enrichment of the sites soil with nutrients

    and improvement of moisture conditions through canopy

    (iii) Disturbance is called any event that causes the destruction of living plant biomass. The reasons of such events may be abiotic (e.g. fire, wind), zoogenic (e.g. grazing, trampling) or anthropogenic (e.g. clearings, wood-cutting).

  • 20 | / Orchids of Rodopi Mountain-Range National Park

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    , 60

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    shading). This forest is usually preserved for not more than

    a few decades, paving the way for the establishment of

    more competitive species. When these (more competitive

    species) succeed to reach and establish themselves, it is

    only a matter of a few decades for them to outcompete

    and replace the pioneer ones by taking away the available

    resources.

    It became clear to scientists that birch in Rodopi has reached

    its peak of distribution sometime in the past. Currently

    -almost sixty years after the end of intense and extended

    disturbances- this species tends to occupy fewer areas and

    withdraw from more sites than in the past, evident of its

    replacement by other more competitive species.

  • / Orchids of Rodopi Mountain-Range National Park | 21

    ;

    .[21]

    -

    . -

    . ,

    , ,

    ,

    , -

    . ,

    . -

    (Quercus dalechampii)

    (1000-1200 m) (Pinus sylvestris)

    (1200-1400 m).

    , , -

    .

    .

    ,

    .[14,22]

    (

    ).

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    -

    , , -

    Which species are more successful in competition?

    Within the plant kingdom exists the so called competitive

    hierarchy,[21] meaning that some species are more competi-

    tive than others, i.e. they are more capable of taking away

    (from less competitive species) fundamental life-support

    resources (e.g. light, water, nutrients). Despite the existence

    of this hierarchy, there may instances when less competitive

    species -under particular ecological conditions- succeed

    in outcompeting more competitive ones. However, these

    instances are rare and cannot occur among species with

    major differences in the competitive hierarchy. In any case,

    and according to this hierarchy, birch in Rodopi is gradually

    replaced by Quercus dalechampii in the lower altitudes

    (1,000-1,200 m) and by Pinus sylvestris in the higher ones

    (1,200-1,400 m).

    The occurrence of Scotch pine forests (Pinus sylvestris) in

    Rodopi is another fact that marks the areas biogeographical

    importance. The world distribution of this species overlaps

    to a high extent with that of Norway spruce. Scotch pine

    is a cold-living conifer, distributed in central and northern

    Europe and central Asia.[14,22] In southern Europe, it is found

    in relict, geographically isolated populations, i.e. popula-

    tions which remained in the southern refugia of the spe-

    cies after the glacial periods. In Greece, Scotch pine forms

    natural forests in three areas, i.e. in Mt. Rodopi, Mt. Voras

    and Mt. Pieria. The story of Scotch pines establishment is

    similar to the one of birch, as they are both pioneer spe-

    cies. Scotch pine has been distributed at the Mediterranean

    area for different reasons than those applying to the case

    of its distribution in central and northern Europe. In the

    Mediterranean, this species distribution was a result of

    vegetation succession that followed extended disturbances.

    In Rodopi, Scotch pine replaced birch forests or established

  • , .

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    (Juniperus spp.), (Rosa spp.), -

    (Crataegus spp.) , , .

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    .[7] -

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    ),

    .[23]

    -

    (Fagus sylvatica).

    , ,

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    itself in non-forested areas, invading pre-forest communities

    comprising of Juniperus spp., Rosa spp., Crataegus spp. and

    other scrub species. The establishment of Scotch pine is also

    triggered by fire, as this species is able to successfully regener-

    ate in sites where forest litter is removed (e.g. burnt) and the

    inorganic is revealed.[7] However, similarly to the case of birch,

    in sites with ecological conditions favouring the growth of

    broadleaved species (i.e. in almost every part of Rodopi), Scotch

    pine is sooner or later bound to be replaced.[23]

    The main competitor and replacement species of Scotch pine

    is beech (Fagus sylvatica). Yet, in sites favouring its growth,

    Norway spruce could also replace Scotch pine. Up to date the

    distribution of Scotch pine in Rodopi has been largely limited,

  • / Orchids of Rodopi Mountain-Range National Park | 23

    .

    ,

    ,

    .

    -

    -

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    , .

    , ,

    -

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    .[7]

    mainly due to vegetation succession. This gradual replace-

    ment of this species does not only affect the distribution

    of dominant species but also of the understory ones. The

    floristic composition of Scotch pine forests depends heavily

    on moisture conditions and reflects the succession stage as

    well as the plant community that emerges after the replace-

    ment of the existing vegetation. Thus, in sites with higher

    moisture and rich in nutrients, the floristic composition

    is similar to the one observed in beech forests. In sites of

    higher altitude (e.g. Elatia and Frakto) the floristic composi-

    tion is similar to the one of Norway spruce forests. On the

    other hand, at drier sites the floristic composition is similar

    to the one observed in oak forests. These latter sites are the

    ones where pure Scotch pine stands are found. It is believed

    that these pure stands will last longer in the future as the

  • 24 | / Orchids of Rodopi Mountain-Range National Park

    -

    Alnus

    incana subsp. incana. -

    -

    .[14,24]

    .[25]

    .

    ,

    . Alnus incana

    ,

    , Alnus glutinosa,

    ,

    last remaining patches of this vegetation type in the area.[7]

    A rare and important, from a biogeographical point of view,

    vegetation type consists of the riparian forests of European

    gray alder (Alnus incana subsp. incana). Alnus incana

    is a Nordic species with scattered distribution in central

    Europe and sporadic occurrence in the mountainous areas

    of southern Europe (up to northern Apennines) and in the

    northwestern and central Balkans.[14,24] In Greece, it is dis-

    tributed exclusively at the area of Kalyvia Karyotou and the

    south-western part of the Elatia Forest.[25] In addition to the

    above-mentioned areas, few individuals of this taxon have

    been found at the area of Lepida forest. An ecological char-

    acteristic of the sites where Alnus incana subsp. incana

    occurs is that they are periodically flooded, especially in

    winter and early summer. Alnus incana can be easily dis-

  • / Orchids of Rodopi Mountain-Range National Park | 25

    , .[26] -

    .

    Alnus incana 1200 m,

    , Alnus glutinosa

    1000 m 800 m ,

    .

    ,

    , (Platanus orientalis),

    (.. Salix alba, S. x fragilis, S. xanthicola).

    ( ),

    .

    Alnus incana subsp. incana, Aegopodium podagraria, Lamium maculatum, Elymus caninus, Carex remota,

    Geum urbanum, Paris quadrifolia, Scrophularia nodosa, Galium aparine, Geranium robertianum, Urtica dioica, Oxalis acetosella, Stellaria nemo-

    rum, Thalictrum aquilegiifolium, Stellaria media, Impatiens noli-tangere, Ranunculus repens, Stachys sylvatica, Moehringia trinervia ..[9]

    Typical species of Alnus incana subsp. incana forests are the Aegopodium podagraria, Lamium maculatum, Elymus caninus, Carex remota, Geum

    urbanum, Paris quadrifolia, Scrophularia nodosa, Galium aparine, Geranium robertianum, Urtica dioica, Oxalis acetosella, Stellaria nemorum,

    Thalictrum aquilegiifolium, Stellaria media, Impatiens noli-tangere, Ranunculus repens, Stachys sylvatica, Moehringia trinervia etc.[9]

    Lamium maculatum Oxalis acetosella Impatiens noli-tangere

    tinguished from the other species of the genus, i.e. Alnus

    glutinosa, by its acute leaf apex, its viscid young shoots and

    its smooth and without cortex bark.[26] Both species occur

    in Rodopi Mountain-Range National Park, yet in different

    altitudes. Alnus incana appears in altitudes above 1,200

    m, where air is colder. Alnus glutinosa occurs in altitudes

    lower than 1,000 m (more often lower than 800 m), where

    the climate is more or less characterised as Mediterranean.

    The riparian vegetation of Rodopi also comprises of stands

    of plane (Platanus orientalis) and willow (e.g. Salix alba,

    S. x fragilis, S. xanthicola). These occur in the vicinity of

    riverbeds, as they are well-adapted to intense flood events

    and prefer to grow in sandy soils.

  • 26 | / Orchids of Rodopi Mountain-Range National Park

    -

    -

    ,

    -

    . , ,

    , -

    (

    )

    .

    -

    M

    Vegetation zonation in Rodopi Mountain-Range National Park

    characteristic ecological feature of the Park is the

    zonation of its vegetation, i.e. the differentiation of

    vegetation types and the spatial arrangement of plant

    species in accordance to altitude. In the mountains of the

    mid-latitudes (such as Rodopi and Falakron) an increase

    in altitude is accompanied by a change in climate (it

    becomes colder and wetter) and also by a corresponding

    differentiation in vegetation. This change can be paralleled

    to latitude increase; this is the reason why those travelling

    within the Park from areas of lower altitude to those of

    A

  • / Orchids of Rodopi Mountain-Range National Park | 27

    . ,

    , ,

    , ,

    .

    ,

    , .

    , , , -

    . ,

    ,

    , (Olea

    europaea), (Pistacia lentiscus), (Arbutus

    spp.) ..

    ,

    ,

    (Quercus coccifera)

    (..

    )

    .

    , -

    (.. , ),

    (Carpinus orientalis) -

    . -

    , ( 1000

    m) (Quercus pubescens).

    -

    , Quercus

    frainetto Q. dalechampii. -

    higher altitude have the impression that they travel from

    the Mediterranean to central and northern Europe.

    In the previous paragraphs, we described certain vegeta-

    tion types that are typical in central or northern Europe

    yet also appearing in higher altitudes in Rodopi. However,

    in the lower altitudes of Rodopi Mountain-Range National

    Park one can observe vegetation types of relatively

    Mediterranean character, though not strictly Mediterranean

    due to the large distance of Rodopi from the sea as well as

    to its relatively high latitude. It is also noted that no typi-

    cal Mediterranean vegetation (e.g. plant communities with

    Olea europaea, Pistacia lentiscus, Arbutus spp. and other

    typical Mediterranean species) occurs in the area.

    Not far from the city of Drama, at the foothills of Mt.

    Falakron, kermes oak (Quercus coccifera) is found. The

    structure and the floristic composition (e.g. dominance of

    evergreen broadleaved species) of this vegetation resem-

    ble those of Mediterranean vegetation types.

    In the lower altitudinal zone of Rodopi, particularly in

    areas with siliceous geological substrate, the dominant

    vegetation comprises of Carpinus orientalis and decidu-

    ous oaks. Areas of higher altitude (up to 1,000 m) host

    forests of downy oak (Quercus pubescens).

    In sites with relatively low disturbance, two other oak

    species appear i.e. Quercus frainetto and Q. dalechampii.

    Quercus frainetto prevails in lower sites and Q. dale-

    champii in higher sites.[27] Oak forests occur usually in

    altitudes ranging from 800 to 1,100 m. In higher altitudes

    they are replaced by beech forests.

    Beech (Fagus sylvatica) is a typical oceanic species, which

    demands sites with high moisture and mild climate with-

    out harsh winters.[28] Greece is the southernmost area of

  • , -

    .[27]

    800 1100 m

    , .

    ,

    .[28]

    , ,

    ,

    .

    .

    , ,

    [29] Horvat et al.,[30]

    . ,

    ,

    . , -

    ,

    (

    ).

    its distribution. It is very competitive, especially in those sites

    where climatic and soil conditions favour its growth. As the

    major factor limiting its distribution is moisture deficiency, in

    Greece it usually forms fragmented forests only in sites with

    adequate moisture.[29,30] In Rodopi, this species forms extended

    forests, which constitute a distinct vegetation zone (character-

    ised by high precipitation and siliceous geological substrate).

    Only a handful of other species can survive and grow within

    beech stands, mainly due to the unfavourable light conditions.

    Indeed, only a small percent of sun radiation can penetrate the

    crown of beech trees.

  • / Orchids of Rodopi Mountain-Range National Park | 29

    . ,

    . -

    ,

    -

    .

    ,

    ,

    (Carpinus betulus). -

    , , -

    (Corylus avellana). -

    In the lower altitudes of beech occurrence, this species

    mixes with Quercus frainetto and Q. dalechampii. These

    mixed forests appear very diverse when compared to

    other plant communities of beech forests. This is due to

    the fact that the participation of oak species in the tree

    layer allows a high amount of radiation to penetrate and

    reach the understory layer.

    In the same altitudinal zone and in sites with concave relief,

    another -relatively rare- vegetation type occurs. This type is

    characterised by the dominance of European Hornbeam

    (Carpinus betulus). In sites with similar relief, yet with cal-

    careous substrate, small patches of scrub vegetation com-

    prising of common hazel (Corylus avellana) can be found.

    This vegetation type is more or less common in sites of

    middle altitudes (e.g. 400-1,000 m) in Mt. Falakron.

  • 30 | / Orchids of Rodopi Mountain-Range National Park

    (.. 400-1000 m) .

    -

    .

    , -

    pH, .[16,31] 1000 m

    .

    -

    (, , ),

    . -

    Epipactis

    Cephalanthera.

    In higher altitudes, beech forests may vary according to

    soil conditions and geological substrate. More specifically,

    in sites with siliceous soils and relatively low pH, we

    observe acidic beech forests.[16,31] These forests occur

    above 1,000 m and they are considered to be less diverse

    in terms of species.

    On the contrary, in sites with adequate moisture and

    calcareous substrate (i.e. Stamna, Elatia, Frakto and Mt.

    Falakron) we observe calcareous beech forests. Their flo-

    ristic composition is characterised by the frequent occur-

    rence of orchids (in particular of the genera Epipactis and

    Cephalanthera).

    However, in sites with calcareous substrate but with insuf-

    ficient moisture or soil depth, beech forests cannot grow

  • / Orchids of Rodopi Mountain-Range National Park | 31

    ,

    (Pinus nigra subsp. nigra).

    .

    ( 1400 m)

    ( 30 m).

    .

    ,

    (Abies borisii-regis). Zoller et

    and they are gradually being replaced by black pine for-

    ests (Pinus nigra subsp. nigra). The communities of black

    pine forests depend heavily on soil properties and appear

    diverse in terms of species numbers.

    A typical characteristic of beech forests that grow on flat

    or concave relief in high altitudes (usually above 1,400 m)

    is the large height of beech trees (in many cases exceeding

    30 m). These forests grow in moist, rich soils and they can

    be found either pure or mixed with Norway spruce (Picea

    abies subsp. abies) and Bulgarian fir (Abies borisii-regis).

    This vegetation type (considered to be representative of

    the so-called climax community) comprises of mesophile

    species, i.e. species with high demands on moisture and

    nutrients.[32,13,33]

  • -

    , Soldanella rhodopaea

    Lathraea rodopaea.[34,35,36,37,38] Lathraea

    rodopaea

    ,

    ,

    -

    ( )

    , .[38]

    Beech forests host common species to the

    central-European area, as well as specialist

    species like the Balkan endemics Soldanella

    rhodopaea and Lathraea rodopaea.[34,35,36,37,38]

    Lathraea rodopaea is a species without

    chlorophyll, which parasitizes on the roots

    of broadleaved tree species (such as beech

    and common hazel). It flowers relatively

    early (until the end of May) and disappears

    soon afterwards (through a swift decay and

    decomposition process).[38]

  • al.,[32] & [13] ,[33]

    (-

    ).

    ,

    .

    ()

    . , ,

    .

    , ,

    . , ,

    .

    .

    , -

    ,

    . ,

    1700 m,

    In the mesophile beech forests, fir is rather frequent. Fir does

    not form extended forests in Rodopi (as it does in other areas

    in Greece) but it appears in mixed stands together with beech

    and spruce. Only in few sites, e.g. Drymos, pure fir stands are

    found. However, it is estimated that its distribution in the area

    is expanding, probably due to the fact that it is one of the spe-

    cies that appear last in the ecological succession process.

    The so-called subalpine beech forests (ecologically similar to

    the mesophile beech forests) constitute a very rare vegetation

    type that occurs in high altitudes (usually above 1,700 m) in

  • 34 | / Orchids of Rodopi Mountain-Range National Park

    , , .[32]

    , , -

    . Schreiber,[39] ,

    , -

    -

    ,

    Geranium

    macrorrhizum.

    , -

    , . -

    .

    ,

    . Tzedakis[40]

    -

    .

    ,

    . -

    , -

    ,

    .

    (..

    ), -

    , .

    (.. Tilia tomentosa), -

    (.. Acer pseudoplatanus A. platanoides)

    (.. Ulmus glabra).

    Greece. It is characterised by the presence of subalpine,

    usually tall herbs.[32] The most common habitat of these

    species is the banks of small subalpine streams. According

    to Schreiber,[39] the forest limits in Mt. Falakron are formed

    by beech forests, which represent a particular plant com-

    munity, classified under subalpine beech forests.

    Another rare vegetation type occurring sporadically in

    Rodopi is the ravine forests. These forests are character-

    ised by particular ecological features and occur within the

    zone of oak forests as well as within the lower part of the

    beech zone. Due to their favourable moisture and tem-

    perature, ravines host habitats that have potentially played

    the role of refugia for plant species. Indeed, according to

    Tzedakis,[40] areas with geomorphological characteristics

    similar to those of ravines have not only offered significant

    protection against unfavourable climatic conditions but

    also adequate moisture for several species during the gla-

    cial periods. Such ravines are home (even up to our days)

    to many relict species, i.e. remnants of past geological

    eras. The existence of such relict -sometimes rare- species

    and the favourable soil and micro-climatic conditions are

    the reasons behind the ravine forests rich biodiversity. In

    Rodopi Mountain-Range National Park, such forests can be

    found along the valley of Nestos River (e.g. downstream of

    the Thysavros Dam), as well as along the valleys of other

    smaller rivers or streams, such as Diavolorema Stream

    and Despatis River. Typical tree species of this vegetation

    type are lime (e.g. Tilia tomentosa), sycamore (e.g. Acer

    pseudoplatanus and A. platanoides) and elm (e.g. Ulmus

    glabra).

  • / Orchids of Rodopi Mountain-Range National Park | 35

    (

    65 2,5 )

    Haberlea rhodopensis. Gesneriaceae

    , .[6] , , -

    . -

    .

    Haberlea rhodopensis is a typical relict species of the tertiary period (the geologic period between 65 and 2.5 million years

    ago). It occurs along the ravines of Rodopi Mountain-Range National Park. This species is one of the five belonging to the

    Gesneriaceae family (a typical family of the Tropics) that have remained in Europe.[6] The plants of this species grow in rock

    cliffs with high moisture and form a characteristic green carpet. The occurrence of such relict species, with such narrow

    ecological niche, is a clear indication of the great conservation importance of ravine forests.

    Haberlea rhodopensis

  • 36 | / Orchids of Rodopi Mountain-Range National Park

    ( 85%) .

    ,

    , -

    ,

    ( ) .

    ,

    , -

    .

    Grassland vegetation in Rodopi Mountain-Range National Park

    he Rodopi Mountain-Range National Park is mainly

    covered by forests (85% of the Parks total area).

    Notwithstanding its limited extent, the Parks non-forest

    vegetation triggers interest, mainly due to the fact that its

    conservation is threatened by swift reforestation. Indeed,

    the few remaining shepherds say that the left-overs of

    grasslands will not remain much longer as the forest runs

    within them.

    Most of the grasslands in the Park are of semi-natural form,

    created after the destruction of forests. In addition, most of

    them grow on a siliceous substrate. Grasslands are usually

  • / Orchids of Rodopi Mountain-Range National Park | 37

    .

    .

    -

    .

    (1800-2000 m)

    .[30]

    (.. -

    , ), (.. ,

    ) (..

    ) . ,

    .

    .

    .

    , -

    (.. )

    . ,

    .

    -

    , -

    .

    , ,

    (Pteridium aquilinum).

    , ,

    rich in terms of species and their floristic composition dif-

    ferentiates according to altitude and soil. In general, and

    due to climate, even the grasslands occurring above the

    current forest limits (1,800-2,000 m) should be covered

    by forests;[30] their current occurrence is merely attributed

    to past land use (animal husbandry, agriculture, etc.), the

    disturbances (e.g. fires and grazing) and soil degradation

    (e.g. erosion and reduction of soil depth).

    However, there are cases where the growth of forest veg-

    etation can be suppressed by special soil conditions or

    even periodic flooding. In most cases, the occurrence of

    grasslands in the National Park may be attributed to the

    land use practice before the Second World War. In some

    cases, soil has been degraded to such extent that the estab-

    lishment of tree species is impossible. In others, degrada-

    tion is not that intense thus allowing reforestation. Another

    important factor that affected vegetation succession is graz-

    ing, which was much more intense two-three decades ago

    and allowed the preservation of grasslands.

    The most typical subalpine-alpine grasslands of the Park

    are found in Mt. Falakron. These are based on a calcareous

    substrate and they are highly diverse in terms of species

    number.

    In many sites within the Park (especially in forest open-

    ings) vegetation is dominated by bracken (Pteridium

    aquilinum), a species widespread among forest plant

    communities. Under certain ecological conditions it forms

    dense thickets, also called bracken. Pteridium aquilinum

    is a rather competitive species, which expands rapidly

    through its rhizomes. However, its dominance among

    other plants usually ends at some point. According to

    Marrs & Watt,[41] the ecology of this species follows a cir-

    cular succession process with phases of maximum and

  • 38 | / Orchids of Rodopi Mountain-Range National Park

    , Sesleria rigida, Asperula

    aristata, Asperula purpurea, Dianthus gracilis, Festuca varia, Bromopsis cappadocica, Thymus thracicus,

    Trifolium pratense, Trifolium campestre, Vincetoxicum hirundinaria subsp. nivale, Thymus praecox, Achillea

    millefolium, Potentilla detommasii, Potentilla cinerea, Scabiosa ochroleuca, Festuca valesiaca ..

    Of the species appearing in subalpine and alpine grasslands of Mt. Falakron most common are: Sesleria

    rigida, Asperula aristata, Asperula purpurea, Dianthus gracilis, Festuca varia, Bromopsis cappadocica, Thymus

    thracicus, Trifolium pratense, Trifolium campestre, Vincetoxicum hirundinaria subsp. nivale, Thymus praecox,

    Achillea millefolium, Potentilla detommasii, Potentilla cinerea, Scabiosa ochroleuca, Festuca valesiaca etc.

    Trifolium alpestre Anthyllis aurea Sideritis scardica

    Centaurea napulifera subsp. napulifera Viola perinensis Viola delphinantha

  • / Orchids of Rodopi Mountain-Range National Park | 39

    , .

    .[41] ,

    , Marrs & Watt[41]

    .

    , -

    .

    , 1500 m, -

    -

    () , Juniperus

    communis subsp. hemisphaerica, J. communis subsp. alpina,

    Vaccinium myrtillus Vaccinium vitis-idaea.

    Crocus flavus

    minimum growth. At the phase of its minimum growth,

    when its cover is less dense, it is replaced by forest veg-

    etation.

    In relatively high altitudes (usually above 1,500 m) vegeta-

    tion is dominated by dwarf shrubs (chamephytes) such

    as Juniperus communis subsp. hemisphaerica, J. commu-

    nis subsp. alpina, Vaccinium myrtillus and Vaccinium

    vitis-idaea. This vegetation type is called subalpine heath,

    because of the occurrence or even dominance of heaths

    (Ericaceae species such as Vaccinium spp.). These heaths

    are not widely distributed in the Greek part of Rodopi

    Mountain Range, as this includes only a small area of sub-

    alpine grasslands.

  • 40 | / Orchids of Rodopi Mountain-Range National Park

    ,

    / Ericaceae (..

    Vaccinium).

    ,

    .

    (

    1700 m) -

    ,

    Nardus stricta. -

    .

    .[42]

    -

    .

    1300 m

    .

    .

    , -

    , . ,

    , -

    .

    .

    Sphagnum.

    . -

    (iv).

    -

    (iv) Sphagnum.

    Another rare vegetation type of high importance (from a

    conservation point of view) occurs in areas with siliceous

    substrates and usually in high altitudes (above 1,700 m).

    This vegetation type is dominated by Nardus stricta, a per-

    ennial species that grows in sites with high precipitation

    and acidic soil. This species is described as competitive in

    soils relatively poor in nutrients.[42]

    In flat and concave sites adjacent to streams -which

    receive water from higher grounds- wet meadows can be

    found. These meadows occur usually above 1,300 m and

    are dominated by tall herbs. Typical wet meadows can be

    found in Rodopi Mountain-Range National Park along the

    Stravorema stream (Elatia area).

    Another soil-dependant vegetation type (very rare in

    the Mediterranean) consists of bogs. In the area of the

    Park bogs are found in the areas of Elatia and Lepidas.

    Their major ecological characteristic is that they grow on

    organic soil that is formed by the accumulation of dead

    organic matter (mainly of plants). Another characteristic

    of this vegetation type is the high cover of a dense moss

    carpet composed by species of the genus Sphagnum. Bogs

    are also known as peat-lands as they produce peat. The

    organic substrate of bogs remains wet all over the year;

    this is the reason why forest vegetation is usually absent

    in these sites. The wet state of the organic substrate is

    also the reason for the accumulation of the organic mat-

    ter, which slowly decomposes under oxygen lack. Several

    species of wet grasslands are also found in bogs with the

    most typical being Sphagnum contortum,[43] Eriophorum

    latifolium, E. angustifolium, Dactylorhiza cordigera, Carex

    (iv) Peat is composed by dead organic remnants of plants and mainly of those of Sphagnum species.

  • / Orchids of Rodopi Mountain-Range National Park | 41

    ,

    , , ,

    , ,

    .

    Sphagnum contortum,[43] Eriophorum latifolium, E.

    angustifolium, Dactylorhiza cordigera, Carex hita, C. remota, C.

    canescens, Juncus filiformis, J. effusus, J. bufonius, J. thomasii ..

    ,

    Drosera rotundifolia.

    Drosera rotundifolia

    hita, C. remota, C. canescens, Juncus filiformis, J. effusus, J.

    bufonius, J. thomasii etc. Finally, it must be noted that in

    very few patches within the bogs of the Park, the carnivo-

    rous plant species Drosera rotundifolia is also hosted.

  • / Orchids of Rodopi Mountain-Range National Park | 43

    .

    -

    ( ) .

    .

    -

    .

    . -

    , ,

    ,

    . , -

    .

    , .

    , -

    , -

    -

    ,

    -

    ,

    .[44]

    -

    he Rodopi Mountain-Range National Park is characterised by its high vegetation diversity as well as by the occur-

    rence of rare vegetation types. The latter include vegetation

    types that reach their southernmost distribution in the area

    of Rodopi or consist of rare and endemic (mainly Balkan)

    species.

    The ecosystems of Rodopi are subject to sustainable

    management. This type of management favours vegetation

    types which, depending on site conditions, form the so-

    called climax communities, i.e. plant communities at the

    latter stages of succession. Vegetation types that comprise

    of pioneer species may be significantly limited in size,

    due to the reduction and the intensity of disturbances. In

    order to conserve such pioneer vegetation types, spe-

    cial management practices that simulate the impacts of

    disturbances should be applied. This kind of management

    in certain sites could positively affect regeneration and

    growth of pioneer species.

    Management should also focus on the control of grass-

    lands size. A potential reduction of their area could even-

    tually lead to the loss of potential niches of plant and

    animal species. Surveying and monitoring of plant species

    (and vegetation types), setting conservation targets and

    implementing conservation measures are the manage-

    ment actions that need to be taken in order to conserve

    and further enhance biodiversity in the ecosystems of Mts.

    Rodopi and Falakron.[44]

    The importance of vegetation conser-vation in Rodopi Mountain-Range Na-tional Park

  • 44 | / Orchids of Rodopi Mountain-Range National Park

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  • / Orchids of Rodopi Mountain-Range National Park | 45

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    32 H. Zoller, P. Geissler, & N. Athanasiadis, 1977. Beitrge zur Kenntnis der Wlder, Moos- und Flechtenassoziationen in den Gebirgen Nordgriechenlands. Bau-hinia, (), . 6(1), . 215-255 + . 1-14 + .

    33 . , 1985. . , ..., . 29, . 595-670.

    34 E. Eleftheriadou, & T. Raus, 1996. The vascular flora of the nature reserve Frakto Virgin Forest of Nomos Dramas (E Makedonia, Greece). Willdenowia, . 25(2), . 455-485.

    35 . Eleftheriadou, K. Theodoropoulos, N. Athanasiadis, & I. Tsiripidis, 1998. Geo-graphical distribution of Arabis procurrens Waldst. & Kit, Lilium rhodopaeum Delip. and Lathraea rhodopea Dingler in Greece. : I. Tsekos, & M. Mous-takas (.), Progress in botanical research. Proceedings of the 1st Balkan Botanical Congress, Kluwer Academic Publishers, , . 57-60.

    36 I. Tsiripidis, & N. Athanasiadis, 2003. Contribution to the knowledge of the vascular flora of northeast Greece: Floristic composition of the beech (Fagus sylvatica L.) forests in Greek Rodopi. Willdenowia, . 33(2), . 273-297.

    37 . , . , . , . , & . ,

    2009. Lathraea rhodopea Dingl. - (VU) : . , . - & . (.), & - , . 2, , , . 114-116.

    38 . , . , & . , 2009. Soldanella rhodop-aea F. K. Meyer - (NT) : . , . , & . , (.), & , . 2, , , . 328-329.

    39 H. J. Schreiber, 1998. Waldgrenznahe Buchenwlder und Graslnder des Fala-kro und Pangon in Nordostgriechenland. Syntaxonomie. Struktur und Dyna-mik. Arbeiten aus dem Institut fr Landschaftskologie. Westflische Wilhelms-Universitt, , . 4, . 1-171.

    40 P. C. Tzedakis, I. T. Lawson, M. R. Frogley, G. M. Hewitt, & R. C. Preece, 2002. Buffered Tree population changes in a Quaternary refugium: evolutionary implications. Science, . 297, . 2044-2047.

    41 R. H. Marrs, & A. S. Watt, 2006. Biological Flora of the British Isles: Pteridium aquilinum (L.) Kuhn. Journal of Ecology, . 94, . 12721321.

    42 M. J. Chadwick, 1960. Biological Flora of the British Isles: Nardus stricta (L.). Journal of Ecology, . 48, . 255267.

    43 . . , 1977. Sphagnum contortum Schultz, Sph. subsecundum Nees, Sph. palustre L. Sph. squarrosum Pers. (Cr.). . , ..., . 10, . 263-281.

    44 C. R. Margules, L. R. Pressey, 2000. Systematic conservation planning. Nature, . 405, . 243-253.

  • / Orchids of Rodopi Mountain-Range National Park | 47

    (Orchidaceae)

    ,

    800 , , 25.000

    . ,

    ,

    . -

    ,

    . , ..

    .

    .

    ,

    , ,

    -

    ,

    - .

    . -

    ,

    , -

    . ,

    , (1 .. )

    ( Anacamptis

    papilionacea):

    ,

    ( , 3, 126, 2, 1).

    H

    he orchid family (Orchidaceae) is one of the richest families of the Plant Kingdom, comprising of approxi-mately 800 genera and of more than 25,000 species.

    Though orchid species are distributed all over the world

    (except Antarctica), most of them are found in the humid

    tropical and subtropical regions. In the tropical regions the

    majority of orchids are either epiphytic on forest trees or

    lithophytic on rocks and cliffs; only few of them are ter-

    restrial. On the contrary, in temperate regions (e.g. Europe)

    all orchids are terrestrial and widespread in almost all

    habitat types.

    Orchids have traditionally triggered experts interest and

    amateurs love. This interest derives from their complex

    biology, spectacular beauty, eccentric flowers and, last but

    not least, by the bizarre ways through which pollination

    takes place. Indeed, the pollination mechanisms of orchids

    are a great example of flower - insect adaptation.

    Many orchid species stimulated the imagination of peo-

    ple around the world and gave rise to superstitions and

    legends. The medicine of antiquity attributed aphrodisiac

    properties to orchids, because of the similarity of the root

    system of several species with testicles. It was also consid-

    ered that they could determine the gender of the children.

    With regard to the last point in particular, Dioskorides (1st

    century AD) wrote about the bulbous root of the genus

    orchis (probably Anacamptis papilionacea): It is said, in

    respect to this orchid, that when a man eats its bigger root

    he begets boys; when its smaller root is eaten by a woman

    she gives birth to girls.

    Their name derives from the shape of their bulbs, for

    The Orchid Family

    T

  • 48 | / Orchids of Rodopi Mountain-Range National Park

    :

    , ( )

    , ( ) , ()

    , , ( )

    , , , ,

    , (

    ) , (v).

    ,

    , -

    .

    Orchidaceae -

    ,

    Orchidoideae Epidendroideae. ,

    Cypripedioideae, ,

    Cypripedium calceolus L.,

    (,1988), .

    Orchis simiaNeottia ovata Orchis militaris

    (v) Sed inter pauca mirabilis est orchis herba sive serapis, foliis porri, caule palmeo, flore purpureo, gemina radice testiculis simili, ita ut maior sive, ut aliqui dicunt, tenuior ex aqua pota excitet libidinem, minor sive mollior e lacte caprino inhibeat (Plin., Hist. Nat., 26, 62).

    which Pliny the Elder (23 AD - 79 AD) wrote: The plant

    named Orchis or Serapias is admirable (together with few

    other plants); its leaves resemble those of leek, its stem is as

    tall as the palm of a hand, its flower is purple and its root

    consists of two testicle-like bulbs; drinking the larger -or

    harder as some say- bulb stimulates libido, whereas drink-

    ing the smaller or softer with goat milk suppresses it(v). Due

    to the form of the flowers of several species, orchids were

    dedicated to Satyrs and Sileni, i.e. Greek mythical lustful

    creatures that were part man and part beast and always in

    the company of Dionysus.

    The orchid family is further classified into five subfamilies,

    with two of them distributing in Greece (Orchidoideae and

    Epidendroideae). The presence of the third subfamily called

    Cypripedioideae (consisting of the species Cypripedium

    calceolus L.) has not been yet confirmed in Greece.

  • / Orchids of Rodopi Mountain-Range National Park | 49

    110

    , -

    ,

    . ,

    Orchidaceae ,

    .. Iridaceae, Asparagales.

    , ,

    . , ,

    -

    ,

    .

    ,

    ,

    100-110 . .

    Orchidaceae

    , -

    .

    ,

    .

    ,

    ,

    . Eoorchis miocaenica Mehl,

    15 (

    ), . ,

    Protorchis monorchis Massalongo

    Palaeorchis rhizoma Massalongo,

    , Eoorchis miocaenica Mehl

    ( ), -

    .

    he evolution of orchids must have started 110 million years ago, almost simultaneously to the evolution of

    other plant families. In particular, the Orchidaceae fam-

    ily (as well as other highly evolved families including

    Iridaceae) is classified under the Asparagales order. It is

    believed that this order evolved before the end of the

    Cretaceous period, i.e. relatively early. The above view is

    further supported by the fact that complex pollinating

    insects such as Hymenoptera, regarded to be responsible

    for the high diversity of the orchid family, do not appear

    in fossils dating 100-110 million years ago.

    The estimation of the time orchids first appeared on earth

    is further hindered by the fact that no fossils of their

    early species have been yet found. The earliest orchid

    fossil found comes from Germany; this is the Eoorchis

    miocaenica Mehl and it was taken out by a layer dating

    back 15 million years (Upper Miocene), i.e. in a period

    when the climate of Europe was tropical. The fossils of the

    plants Protorchis monorchis Massalongo and Palaeorchis

    rhizoma Massalongo, found in the region of Verona in

    Italy, are earlier than the one of Eoorchis miocaenica Mehl

    (dating back in the Italian Eocene period), yet difficult to

    attest whether they are orchids or not despite the fact that

    they are considered to be monocotyledons.

    rchid seeds are capable of dispersing at a long distance.

    However, their geographic distribution is primarily attrib-

    uted to plate tectonics. Indeed, c. 110 million years ago,

    when orchids first appeared, the Earths continents were

    relatively close to one another thus allowing orchids to

    spread. Yet, the exact location of their first appearance

    The evolution of orchids

    T

  • 50 | / Orchids of Rodopi Mountain-Range National Park

    T , ,

    . -

    . ,

    110 , -

    ,

    .

    .

    ,

    ,

    .

    ,

    . , -

    ,

    . , , -

    ,

    . , -

    ( -

    , )

    .

    remains a question unanswered.

    The orchid family is considered to be the prime example

    of monocotyledons evolution. This is due to the unique

    attributes of its species flowers such as the variation in

    their colour and the construction of their pollinia. These,

    among other, are the mechanisms which are responsible

    for orchid speciation. Speciation, as a consequence of iso-

    lation, is associated to pollinators specialisation, namely

    the dependence of orchids in certain insect species. The

    attraction of new pollinators (or the beginning of a com-

    plex mechanism of deception which will be herein ana-

    lysed) can result in divergent floral morphologies that lead

    to the creation of new species or species groups.

  • / Orchids of Rodopi Mountain-Range National Park | 51

    -

    . , ,

    , ,

    .

    (Ophrys, Orchis

    s.l.), ( Dactylorhiza),

    (Gymnadenia), -

    (Cephalanthera, Epipactis),

    (Corallorhiza, Epipogium) (Neottia).

    -

    . (. ),

    ,

    .

    . Spiranthes

    Ophrys mammosaNeottia nidus-avis Anacamptis pyramidalis

    errestrial orchids have a very simple root system. Each

    plant, except tubers, often develops unbranched, fleshy

    and thick roots, typical of the species that have mycor-

    rhizae. Their root system consists of tubers whose shape

    differs among species, i.e. oval (Ophrys, Orchis s.l.), pal-

    mate (several species of Dactylorhiza) or flattened and

    digitate (Gymnadenia). Other genera have branched

    (Cephalanthera, Epipactis), coral-like (Corallorhiza,

    Epipogium) or nest-like (Neottia nidus-avis ) rhizomes.

    The thick and fleshy roots serve a variety of functions

    including mycotrophy, the absorption and storage of nutri-

    ents and the anchoring of the plant body to the ground.

    Roots can last up to three years. In the genus Spiranthes

    the root system consists of cylindrical or spindle-shaped

    tubers, which have mycotrophic and storage functions. On

    The morphology of orchids

  • 52 | / Orchids of Rodopi Mountain-Range National Park

    ,

    . ,

    Orchis s.l., Platanthera Dactylorhiza -

    ,

    .

    the contrary, species of the genera Orchis s.l., Platanthera

    and Dactylorhiza have both tubers and thin roots that

    support mycotrophy. The latter (thin roots) develop during

    autumn or spring and disappear next summer along with

    the leaves. Their life span is shorter than one year.

    Pollinia

    Rostellum

    Hypochile

    Anther

    Dorsal sepal

    Lateral sepal

    Biscidium

    Epichile

    Spur

    Sepal

    Pollinaria

    Petal

    Lip

  • / Orchids of Rodopi Mountain-Range National Park | 53

    -

    , ,

    .

    -

    ,

    In tuber-bearing species, the oldest tuber decays during

    flowering. During winter, simultaneously to the develop-

    ment of leaves and the re-launch of the photosynthesis

    process, nutrient storage in the young tuber begins. This

    emerging tuber will be preserved until the following year

    Hypochile

    Bract

    Epichile

    Sepals

    Ovary

    Dorsal sepal

    Petal

    Gynostegium

    Stigmatic cavity

    Basal lip area

    Speculum

    Lateral sepal

    Labellum

    Appendage

  • 54 | / Orchids of Rodopi Mountain-Range National Park

    . -

    .

    , ,

    ,

    (Goodyera).

    (Ophrys).

    . , -

    .

    180

    . (

    ). -

    (labellum).

    .

    (spur). , ,

    . , -

    , ,

    . Epipogium

    aphyllum Gymnadenia rhellicani

    180 360

    .

    (Orchidoideae Epidendroideae)

    .

    , ,

    ( ).

    .

    and will eventually replace the old tuber.

    Leaves, as in most monocotyledons, are entire with parallel

    veins; sometimes they form a net-like mesh with the sec-

    ondary veins (Goodyera). The basal leaves usually form a

    rosette (Ophrys). The leaves of the stem are opposite or spi-

    rally arranged and the upper ones are very small and bract-

    like. In most saprophytic orchids, leaves are scale-like.

    Orchids form spikes that consist of resupinate flowers.

    A resupinate flower is one in which the pedicel does a

    180-degree twist while it develops, bringing what would

    be the bottom of the flower to the top. The perigon is

    formed by two circles, one on the exterior and one on the

    interior. A part of the inner circle forms a lip known as

    labellum, which appears diverse among different species.

    The labellum is the part on which insects sit during the

    pollination process. In several genera, the back of the label-

    lum is elongated and forms a spur. The labellum is usually

    positioned at the bottom of the flower facing at the front.

    In closed buds the labellum is positioned at the top but

    it turns down while flowers develop. In some cases (e.g.

    in species such as Epipogium aphyllum and Gymnadenia

    rhellicani) the degree of the twist during the resupination

    process may not be 180o. In other cases the degree of the

    twist may reach 360o, leaving the labellum on top.

    The reproductive organs of the orchids found in Greece

    (Orchidoideae and Epidendroideae) consist of a fertile sta-

    men in the outer circle of each flower. On either side of

    the fertile stamen develops an incomplete and sterile sta-

    men (i.e. not producing pollen), i.e. two in total. These two

    sterile stamens, known as staminodes, do not participate

    in the pollination process. The stigma and the style are

    joined together with the fertile stamen forming the gynos-

  • / Orchids of Rodopi Mountain-Range National Park | 55

    (column),

    (gynostegium).

    () ,

    . ,

    ()

    -

    (pollinia).

    ( ) -

    (caudicle), -

    (retincles).

    .

    ( )

    () ,

    () ( ).

    ( ) -

    ,

    (rostellum).

    ()

    () .

    ,

    -

    .

    - (bursicles).

    - ,

    . , ,

    , , ,

    , .

    tegium, a special, column-like structure situated above the

    (inferior) ovary. The anthers rarely release pollen-grains;

    these grains are glued together by a sticky (viscous) sub-

    stance and are bound by elastic threads in packets called

    pollinia. In most cases, the pollinia have small stalk-like

    caudicles, which in turn are joined together with viscous

    glands called retincles. The overall formation consisting

    of the pollinia, the caudicle and the retincle is called pol-

    linarium. The orchid stigma is 3-lobed with only two lobes

    being functional, i.e. capable of pollination. These lobes

    form a seemingly single surface coated by a more or less

    () adhesive substance at the base of the gynostegium, in

    a more or less () shallow cavity (stigmatic cavity). The

    middle sterile lobe of the stigma called rostellum evolves

    into a special area of the gynostegium above the fertile

    part of the stigma and beneath the anther. Sometimes, the

    rostellum is limited to a small, sticky and rather globular

    gland projecting from the stigmatic cavity. Key parts of the

    rostellum are the viscous glands (or a single gland), where

    the stalks of the pollinaria are joined. The viscous glands

    are either exposed or enclosed in small bursicles. A key

    function of the rostellum is to produce a sticky-viscosity

    substance, which enables the pollinaria to stick to and

    transferred by insect pollinators; the rostellum also pre-

    vents pollen to fall onto the fertile stigma, thus preventing

    self-pollination and autogamy.

  • 56 | / Orchids of Rodopi Mountain-Range National Park

    ,

    .

    . ,

    ,

    .

    , ,

    .

    -

    , , ,

    .

    Orchis quadripunctata

    Germination MycorrhizaUnlike the seeds of other plant families, orchid seeds are

    very small in size. A typical feature of them is that they

    have no endosperm and that their embryo is undiffer-

    entiated. When the mature capsule opens, usually in dry

    weather conditions, the seeds are released and dispersed

    by the wind. The seeds consist of an outer membrane

    surrounding the embryo in such a manner that allows a

    considerable amount of air within them. This, in turn, ena-

    bles seeds to float on air longer, travel further and disperse

    more widely than any other plant species. The seeds can

    achieve extensive coverage of the area around the mother

    The biology of orchids

  • / Orchids of Rodopi Mountain-Range National Park | 57

    (

    ) (6.000 -

    10.000 ).

    , -

    , ,

    .

    .

    , -

    ,

    . ,

    .

    . -

    -.

    ,

    , .

    ,

    , -

    ,

    .

    , ,

    ,

    ,

    .

    , -

    , -

    plant, as their size is small (allowing them to travel far) and

    their number is large (6,000 to 10,000 per capsule).

    Notwithstanding the very large number of seeds released,

    only a small percentage eventually germinates. Their

    majority will either never reach the ground or not germi-

    nate due to unsuitable substrate. Other factors affecting

    germination are the consumption of seeds by soil fauna

    and fungi infection, usually by Basidiomycetes, the main

    suppliers of seeds with water and minerals. Fungi infec-

    tion, in particular, can be highly specialised, as some orchid

    species are infected by only certain species of fungi. At

    the beginning, fungi penetrate seed cells; soon after, they

    penetrate root cells and form hyphae that nourish the

    plant. This heterotrophic nourishment of orchids is called

    mycotrophy or mycoheterotrophy. There are times, when

    orchids rely entirely on mycotrophy; at other times they

    either alternate to or combine mycotrophy with photo-

    synthesis. Mycotrophy is a process that allows orchids to

    adapt to a wide variety of habitats, even in those with

    extreme conditions (e.g. in sites with little soil or lack of

    light). Indeed, when conditions are unsuitable, the under-

    ground organs of orchids can remain alive and stagnant

    for several years; when conditions improve these organs

    start growing.

    As mentioned above, orchid seeds lack endosperm, thus

    making them completely dependent on mycorrhizal

    symbiosis for germination and growth. Mycorrhiza helps

    plants to grow through the absorption of water, nutrients

    and carbohydrates. The interval between the underground

    infection of the seeds until the emergence of the first over-

    ground section ranges from one to three years, depending

    on the species.

    After seed germination, a certain period until the first

  • 58 | / Orchids of Rodopi Mountain-Range National Park

    .

    , .

    -

    , .

    .

    5-8 Ophrys apifera, 9-11 Neottia nidus-

    avis Cephalanthera damasonium, Neottia ovata,

    Spiranthes spiralis Neotinea ustulata 13-15

    .

    -

    , .

    :

    (rewarding orchids),

    ,

    ( - food

    deception Batesian floral mimicry),

    (sexual

    mimicry) (visual mimicry),

    .

    , (.. Ophrys apifera,

    Epipactis sp.).

    flowering of the plants should elapse. This period var-

    ies widely, ranging from 5-8 years in Ophrys apifera, 9-11

    years in Neottia nidus-avis and Cephalanthera damaso-

    nium and 13-15 years in Neottia ovata, Spiranthes spiralis

    and Neotinea ustulata.

    Pollination No other plant family, in its evolutionary history, has

    developed such an impressive and complex reproduction

    mechanism, as the orchid family did.

    Orchids are insect pollinated species. Their pollinators are

    attracted:

    by nectar (rewarding orchids);

    due to deception (mimicry of nectariferous plants by

    non-nectariferous plants, also known as food decep-

    tion or Batesian floral mimicry;

    by sexual and visual mimicry;

    because they find shelter against harsh conditions.

    Some orchids are self-pollinated (e.g. Ophrys apifera,

    Epipactis sp.).

    The pollinaria are attached to the bodies (particularly to

    the heads) of pollinators, allowing the latter to visit and

    pollinate a reasonable number of flowers. At the begin-

    ning, the pollinaria stand upright in the heads of the

    insects; their stalks later bend and twist forward, along the

    axis of the insects bodies. When a pollinator visits a flow-

    er, the attached pollinaria come in contact with the viscous

    (sticky) surface of the stigma and pollination takes place.

    A very important and remarkable mechanism of attraction

    by orchids is sexual and visual mimicry. Sexual mimicry

    is a complex process during which the flowers of several

  • / Orchids of Rodopi Mountain-Range National Park | 59

    .

    .

    ,

    ,

    . ,

    ,

    ()

    .

    Dactylorhiza saccifera Corallorhiza trifida

    orchids (mainly species of the genus Ophrys) emit chemi-

    cals that are similar to pheromones emitted by female

    insects (Hymenoptera). In most cases, flower-emitted

    pheromones vary between different species. Therefore,

    orchid species are pollinated only by specific insects. Male

    insects are further deceived by the orchid flowers shape

    and colour that give the impression of female insects;

    under this false impression, male insects proceed to

    pseudocopulation on the labellum of the flowers. During

    pseudocopulation, the pollinaria are attached to the head

    or the abdomen of the insects and later transferred to a

  • 60 | / Orchids of Rodopi Mountain-Range National Park

    -

    . -

    . (

    Ophrys) ,

    ().

    , -

    . -

    Epipactis helleborine

    flower, either of the same or of another individual. This

    recurrent process results in the pollination and fertilization

    of flowers by insects.

    Visual mimicry is a mechanism which orchid species

    adopted in order to imitate species of other families (e.g.

    Orchis israelitica imitates Bellevalia flexuosa and Orchis

    pallens imitates Lathyrus vernus). This mimicry mechanism

    allows plants to share pollinators. In the second example

    mentioned above, insects are deceived by the colour of

    the flowers which is similar in both species, as well as by

  • / Orchids of Rodopi Mountain-Range National Park | 61

    ,

    (, )

    . ,

    -

    (pseudocopulation).

    , . -

    , -

    .

    Epipactis leptochila subsp. neglecta

    the similarity of the spur of O. pallens with the calyx of L.

    vernus.

    Non-deceived pollinators may visit some orchid species

    for other reasons, e.g. shelter. Indeed, some orchid flow-

    ers form a hood, in which insects can find shelter. This

    need for shelter is the main reason behind the attraction

    of pollinators by orchid species of the genus Serapias, as

    well as by species such as Anacamptis papilionacea and

    Anacamptis morio. Finally, it must be noted that some

    orchid species are self-pollinated. These species do not

  • 62 | / Orchids of Rodopi Mountain-Range National Park

    ,

    (..

    Orchis israelitica Bellevalia flexuosa,

    Orchis pallens Lathyrus vernus),

    . -

    , . Lathyrus vernus

    Orchis pallens,

    O. pallens L. vernus..

    O. pallens

    Lathyrus.

    , -

    . ,

    -

    ,

    . -

    Serapias,

    , .. Anacamptis

    papilionacea A. morio, . ,

    ,

    .

    Ophrys apifera,

    Epipactis Cephalanthera.

    Spiranthes spiralis, Neottia nidus-avis Neottia ovata.

    need insects to transfer the pollinaria from one flower

    to another. Self-pollination is the primary reproduction

    method of Ophrys apifera; it can also happen in species of

    the genera Epipactis and Cephalanthera. A similar pollina-

    tion method can also take place in the species Spiranthes

    spiralis, Neottia nidus-avis and Neottia ovata.

  • / Orchids of Rodopi Mountain-Range National Park | 63

    ,

    ( 1/3

    ).

    .

    ,

    ,

    ,

    -

    Dactylorhiza cordigera

    oth Rodopi Mountain-Range National Park and Mt.

    Falakron are characterised by their rich flora, which

    includes a significant number of orchids (about 1/3 of

    the species found in Greece). Several of them have been

    included in the Red Data Book of Rare and Threatened

    Plants of Greece, mainly due to their limited geographical

    distribution and their unique ecology.

    Orchids are considered to be fragile and vulnerable spe-

    cies due to (i) their dependence on specific species of

    fungi and insects, (ii) their great variation in population

    size (a consequence of climatic variability), and (iii) their

    The importance of Rodopi Mountain-Range National Park for orchids

    B

  • 64 | / Orchids of Rodopi Mountain-Range National Park

    ,

    -

    .

    ,

    , -

    ,

    . , ,

    ,

    .

    . , -

    Neottia cordata

    sensitivity to environmental changes.

    The aesthetic attributes and legendary properties of these

    species have historically attracted a dangerous enemy,

    which is no other than man. Natural and manmade threats

    of increasing intensity endanger the very existence of

    orchids and lead them to rarity.

    rchids are important not only because of their complex

    biology but also because of the phytogeographic and

    ecological interest they trigger. Indeed, in the wider area

    of Rodopi and Falakro, such interest-triggering species

    occur in wet meadows and bogs (e.g. Dactylorhiza mac-

    edonica, D. incarnata, D. cordigera, Epipactis palustris and

    Gymnadenia frivaldii). The above species are distributed

  • / Orchids of Rodopi Mountain-Range National Park | 65

    (Dactylorhiza macedonica,

    D. incarnata, D. cordigera, Epipactis palustris Gymnadenia

    frivaldii).

    ,

    ,

    , . -

    ,

    .

    -

    (.. Epipactis pontica, Gymnadenia rhellicani,

    Goodyera repens, Neottia cordata, Orchis italica, O. militaris).

    Goodyera repens

    within a rather limited geographic range. Yet, their habi-

    tats are among the rarest and most vulnerable habitats in

    Rodopi, Greece, and even Europe. The protection and con-

    servation of these species depends on adequate feeding of

    their habitats with water. Any interruption on water flow

    would lead these species to extinction.

    Equally important are the species that form their south-

    ernmost or northernmost distribution limits in Rodopi

    and Mt. Falakron (e.g. Epipactis pontica, Gymnadenia

    rhellicani,Goodyera repens, Neottia cordata, Orchis italica

    and O. militaris). These sensitive species are threatened

    by climate change, thus making them ideal indicators for

    monitoring global warming. Gymnadenia rhellicani, in

  • 66 | / Orchids of Rodopi Mountain-Range National Park

    ,

    -

    . Gymnadenia rhellicani

    , -

    .

    Orchis militaris,

    -

    200 .

    Goodyera repens, ,

    Orchis militaris

    particular, faces an additional threat, i.e. the one of devel-

    opment activity in the vicinity of the Falakro Skiing Centre,

    where it occurs. Orchis militaris is, in turn, threatened

    by extensive grazing, a threat that has to be immediately

    managed as th