Ορχιδέες του Εθνικού Πάρκου Οροσειράς Ροδόπης
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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
ain-
Ran
ge N
atio
nal P
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
. , -
.
, -
,
,
. ,
,
.
-
-
-
,
, , , ,
. , -
,
-
.
.
. .
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
()
.
, ,
,
,
-
.
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
-
-
-
;
( )
,
, , -
. ,
,
. -
.
,
.
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%.
( ), -
,
( ) -
.[20] ,
. -
(..
).
, , .
(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
.
, ,
, ( ) -
-
.
-
, 60
-
, ,
.
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]
(
).
, , .
-
, , -
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
-
, .
.
,
(Juniperus spp.), (Rosa spp.), -
(Crataegus spp.) , , .
, -
.[7] -
, , ,
(
),
.[23]
-
(Fagus sylvatica).
, ,
. ,
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
.
,
,
.
-
-
. , -
, .
, ,
-
. ,
.
, -
.[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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3 W. Greuter, H. M. Burdet, & G. Long, 1986. Med-Checklist. A critical inventory of vascular plants of the circum-mediterranean countries. Vol 3 (Dicotyledones: Convolvulaceae-Labiatae). Conservatoire et Jardin Botanique de Genve, Med-Checklist trust of OPTIMA, , . i-xvi + 1-395 + xvii-cxxix.
4 W. Greuter, H. M. Burdet, & G. Long, 1989. Med-Checklist. A critical inventory of vascular plants of the circum-mediterranean countries. Vol 4 (Dicotyledones: Lauraceae-Rhamnaceae). Conservatoire et Jardin Botanique de Genve, Med-Checklist trust of OPTIMA, , . i-xvi + 1-458 + xvii-cxxix.
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17 E. Eleftheriadou, K. Theodoropoulos, I. Tsiripidis, & N. Athanasiadis, 2001. Pre-liminary results on the classification of the Greek Picea forests : N. zhatay, (.), Plants of the Balkan Peninsula: into the next Millenium. Proccedings of the 2nd Balkan Botanical Congress, Istanbul, 14-18 May 2000. Marmara niver-sitesi, , . 1, . 195-200.
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20 S. T. A. Picket, & M. L. Cadenasso, 2005. Vegetation dynamics. : E. Van Der Maarel (.), Vegetation Ecology, Blackwell Publishing, , . 172-198.
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22 P. A. Keddy, 2007. Plants and Vegetation: Origin, Processes, Consequences. Cam-bridge University Press, , , . 1-680.
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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.
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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.
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-
/ 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