(ORCHIDACEAE) . : ,
.
2009
ARISTOTLE UNIVERSITY OF THESSALONIKIFACULTY OF SCIENCES SCHOOL OF BIOLOGY DEPARTMENT OF BOTANYLABORATORY OF SYSTEMATIC BOTANY AND PHYTOGEOGRAPHY
THE ORCHIDS (ORCHIDACEAE) OF E. MACEDONIA: DISTRIBUTION, ECOLOGY AND HIGH CONSERVATION VALUE AREAS
Ph. D. THESIS SPYRIDON E. TSIFTSIS FORESTER
THESSALONIKI 2009
. ... (Orchidaceae) . : , ISBN:
(. 5343/1932, 202, . 2)
*. ** . & ** . . . & . & . * **
, 2002 ... . . . , . . . , . . . . , . . , . . . ... . . , . .
i
. E. Willing, H. Presser H. Kretzschmar . . . , , .
ii
. ......................................................................................................... II. ORCHIDACEAE .................................................................... 1 .............................................................................. 2 ...................................................................................................... 3 ............................................................................................................ 3.1 .............................................................................. 3.2 ..................................................................... 4 ................................................................................... I. ................................................................................... 1 ....................................... 2 .............................................................. 2.1 .............................................................................................. 2.2 ............................................................................................... 2.3 ......................................................................................... 3 ................................................................................ 4 .............................................................................................................. 5 ...................................................................................... 5.1 ...................................................................................................... 5.2 ...................................................................................................... 5.3 ........................................................................................................ 5.4 ......................................................................................................... 5.4.1 ................................................................................................... 5.4.2 .................................................................................................... 5.4.3 ................................................................................................... 5.4.4 ................................................................................................... 5.4.5 .................................................................................................. 5.4.6 .................................................................................................. 5.5 ....................................................................................................... 5.6 .............................................................................................. 6 ............. V. ................................................................. 1 ............................................................................................ 1.1 taxa ........... 1.2 ............................................ 2 ................................................................................. 3 .................................................................................. 4 ........................................ 4.1 ........................................................ 4.2 .............................. iii 1 7 7 11 14 14 18 22 29 29 32 34 37 39 42 44 46 47 48 50 52 52 53 55 56 56 57 58 59 61 65 65 65 66 69 69 73 74 77
4.3 .......................... V. ......................................................................................... 1 ................................................................ 1.1 ............................................................................................ 1.2 ........................................................................................................ 2 ......................................................................................... 3 ........................................... 4 ................................ 5 .............................................................. 6 ........................................ VI. .................................................................................................... 1 ..................................................................................... 2 .............................................. 2.1 ............................................................................................. 2.2 ................................................... 2.3 ............................................ 3 ................................ 4 - - .................................................................................. VII. ............................................................ SUMMARY - CONCLUSIONS .................................................................. ................................................................................................... I: ...... II: ...... III: .............
78 81 81 82 108 111 115 117 119 154 165 165 169 169 174 179 181 187 195 199 203 221 247 261
iv
.
I. (Orchidaceae) , 800 - - 25.000 (Chase et al. 2003). , , . 193 (Delforge 2006), . . , , , , . , OPTIMA (Organization for the Phyto-Taxonomic Investigation of the Mediterranean Area) 1977 , , OPTIMA . , , Orchidaceae , . , . , , , . , , (Fleischmann 1925, Renz 1928, Renz 1930, Renz 1932). , , ,
1
.
, , , (.. Nelson 1976, Glz & Reinchard 1981, Knkele & Paysan 1981, Willing & Willing 1987, 1988, Willing & Willing 1991, Delforge 1995, Biel et al. 1998, Kretzschmar et al. 2002 .). , Orchidaceae. , , (.. Rechinger 1939, Zaganiaris 1940, Kitanov 1943, Quzel & Contandriopoulos 1968, 1976, Zoller et al. 1977, Papanicolaou 1985, Karagiannakidou & Kokkini 1987, & 1989, 1992, Karagiannakidou et al. 1995, Eleftheriadou & Raus 1996, Schreiber 1998, Petermann 1999, Arampatzis et al. 2001, Eleftheriadou et al. 2001, 2001, Theodoropoulos et al. 2003, & 2003). , , (Hampe 2004). , (Summerhayes 1969, Currie 1991, Huntley et al. 1995, Knollov & Chytr 2004, Pearson et al. 2004). , , , , (Bailey et al. 2002, Knollov & Chytr 2004, Tsiripidis et al. 2007). , - , , (Stewart 1992). , ( 1988).
2
.
, , (Red Data Books) (Phitos et al. 1995, Hgsater & Dumont 1996, .. 1996, Whigham & Willems 2003). , Stewart (1992) Hgsater & Dumont (1996), (, , , ..). . (Breiner 1979, Syska 1995, Wallenwein & Saad 2000), Sundermann (1980) . , (.. 2001, Karagiannakidou et al. 2001) (.. - 1983, 1985, 1991). , , . , (Myers et al. 2000, Heywood & Iriondo (2003), Tilman 1999, Kati et al. 2004). , , (Hgsater & Dumont 1996, Cribb et al. 2003). in situ , . , , (Schuiteman & Vogel 2003). 3
.
, , (Margules & Pressey 2000, Gaston et al. 2002, Williams et al. 2002, Heywood & Iriondo 2003). (Hill et al. 2006). O . , , (Margules & Usher 1981, Prendergast et al. 1993, Myers et al. 2000). , , (Vane-Wright et al. 1991, Arponen et al. 2005). taxa Orchidaceae . , : taxon. taxa . . Orchidaceae , OPTIMA, . , . 4
.
, Orchidaceae, , , , .
5
. ORCHIDACEAE
II. ORCHIDACEAE II.1 - - 800 - 25.000 (Chase et al. 2003). (. 1), . , . , - .. - (Cribb 1999).
1. Orchidaceae (: Pridgeon et al. 1999). Figure 1. Distribution map of Orchidaceae (Source: Pridgeon et al. 1999).
, Orchidaceae . Orchidoideae Epidendroideae. 7
. ORCHIDACEAE
Cypripedioideae, , Cypripedium calceolus L. DNA , . Bateman et al. (1997), Orchis s.l., (Neotinea, Anacamptis Aceras) , (paraphyly). Orchis lactea, Orchis tridentata Orchis ustulata Neotinea, Neotinea maculata. Anacamptis A. pyramidalis, Orchis 2n=36 ( 32) , O. laxiflora, O. palustris, O. collina, O. coriophora subsp. coriophora, O. coriophora subsp. fragrans, O. sancta, O. papilionacea, O. boryi O. morio. Orchis s.s. Orchis ( 2n=42), Aceras anthropophorum Orchis anthropophora. , Coeloglossum viride Dactylorhiza D. viridis. , Pridgeon et al. (1997) Nigritella Gymnadenia, Sundermann (1980), Listera Neottia (Pridgeon et al. 2005). tribes Pridgeon et al. (1999, 2001, 2003, 2005) Chase et al. (2003) : Orchidaceae Cypripedioideae: 155 , , , , , . Apostasioideae: 16 , , , , , , , , , , , , , , .
8
. ORCHIDACEAE
Vanilloideae: tribes (Pogoniieae, Vanillineae) 15 249 , . Orchidoideae: tribes, 3630 , , . , , , , . Tribe Diseae: 400 11 , , . Tribe Diurideae: , , , , . Tribe Orchideae: 62 1800 . - - . Tribe Chloraeeae: (, , , ). Tribe Codonorchideae: . Tribe Cranichideae: 93 600 , , . Tribe Pterostylideae: , , , , , , . Epidendroideae: 18.000 650 , Epidendroideae . , , . , , . 9
. ORCHIDACEAE
, , . Tribe Arethuseae: Arethuseae (, , , , , , , ), , ( ...) ( ). Tribe Calypsoeae: 12 , , , , , . Tribe Collabieae: 18 , . Tribe Cymbidieae. Tribe Dendrobieae. Tribe Epidendreae: 6000 , . ... Tribe Gastrodieae: , , , , , , , . , , , , . Tribe Malaxideae: 13 , , . Tribe Neottieae: , , , , , , . , . Tribe Nervilieae: Nervilieae , 10
. ORCHIDACEAE
, , , Malay, , , . Tribe Podochileae: 28 1200 , , , , Malay, , . . Tribe Sobralieae: 237 . Tribe Triphoreae: Triphoreae ..., , , ( ), . , . Tribe Tropidieae: 30 . Tribe Vandeae. Tribe Xerorchideae: , , .
II.2 . - , , , (Rasmussen 1995). Delforge (1995) (Ophrys, Orchis s.l.), ( Dactylorhiza), (Gymnadenia), (Cephalanthera, Epipactis), (Corallorhiza, Epipogium) (Neottia). Rasmussen (1995) . ,
11
. ORCHIDACEAE
, . . Spiranthes . Spiranthes , . , Orchis s.l., Platanthera Dactylorhiza , , . , , .
2. Epipactis helleborine. : , : (), : , : , : , : , : , : , : (: Cozzolino & Widmer 2005). Figure 2. Flower of Epipactis helleborine. A: anther, : labellum (epichile), : gynostemium, : petals, : pollinia, : rostellum, : lateral sepal, : dorsal sepal, : stigma (Source: Cozzolino & Widmer 2005).
Summerhayes (1969), . . .
12
. ORCHIDACEAE
, , , (Goodyera). Ophrys. . , (Delforge 1995). 180 . ( ). (labellum). . (spur). , , . , , , . ( Epipogium aphyllum Gymnadenia rhellicani) 180 360 ( 1985, 2001). Orchidoideae Epidendroideae . , , ( ). . (column) (gynostegium). () . (.. Cypripedium), (pollinia). ( ) (caudicle) (retincle). 13
. ORCHIDACEAE
. ( ). , ( ). ( ) , (rostellum). . , , . (bursicle). . , , , , ( 1985, 2001, Pridgeon et al. 1999).
II.3 I.3.1 , . . , , . . , . . ,
14
. ORCHIDACEAE
(Rasmussen 1995). Dactylorhiza o Averyanov (1990) 6.000 . - - . Dactylorhiza , . , . . , , . , . , (Grime 1979). , , (1995) , - . Rasmussen (1998) , . . Harvey (1985), . Stoutamire (1985) ( ). . , . 15
. ORCHIDACEAE
, , . , . , . Rasmussen (1995), , , . . 1. (: Rasmussen 1995). Table 1. Time of seed germination (Source: Rasmussen 1995).
Epipactis helleborine Neottia nidus-avis Neottia ovata Platanthera bifolia P. chlorantha Dactylorhiza sambucina Dactylorhiza viridis Orchis militaris O. pallens Ophrys apifera Corallorhiza trifida Epipogium aphyllum + + + + + + +
+ + + + +
, (. 1), , , (Rasmussen 1995). Leake (1994), . , , . , . ,
16
. ORCHIDACEAE
, , . (Rasmussen 1995). , , , . , . () , , . (Delforge 1995). 2. (: Rasmussen 1995). Table 2. Sojourn below ground (Source: Rasmussen 1995).
Epipactis helleborine Limodorum abortivum Neottia nidus-avis Neottia ovata Spiranthes spiralis Orchis simia Ophrys apifera Dactylorhiza sambucina 3 1 1 1-2 1 1-2 1-3 1
, , . , Goodyera repens, , . , . ,
17
. ORCHIDACEAE
, (Peakall 1994). , , (. 2). . , , Summerhayes (1969) 5-8 Ophrys apifera, 9-11 Neottia nidus-avis Cephalanthera damasonium. Neottia ovata, Spiranthes spiralis Neotinea ustulata . I.3.2 (Van der Cingel 1995): - (rewarding orchids), - ( food deception Batesian floral mimicry), - (sexual mimicry) (visual mimicry) - . , (.. Ophrys apifera, Epipactis sp.). , - - . , . . , (Delforge 1995).
18
. ORCHIDACEAE
(2001), : (Basitonae) (Acrotonae). (.. Orchis s.l, Ophrys, Serapias) . . , . (.. Epipactis, Limodorum) , . , , . , , . . Neiland & Wilcock (1998) . , (rewarding) ( Neottia ovata). , (non-rewarding) ( Platanthera). Anacamptis coriophora subsp. coriophora (Dafni 1987), (6070%) (Cozzolino & Widmer 2005). Anacamptis, Orchis Neotinea . , (Dafni 1987). , Anacamptis coriophora subsp. coriophora Anacamptis sancta , Orchis, Neotinea Anacamptis (Cozzolino et al. 2001). Dafni & Ivri (1979) Orchis s.l. . Orchis s.l. . 19
. ORCHIDACEAE
Delforge (1995) Averyanov (1990) Dactylorhiza. Neiland & Wilcock (1998) . , . , , ( ) (Cozzolino & Widmer 2005). Cozzolino et al. (2001). , Orchis s.s. . Orchis mascula, ( ) (Orchis italica), (Orchis anthropophora). (Orchis mascula, Orchis patens, Orchis pallens Orchis spitzelii) . . . ( Ophrys) , (), . , (, ) . , (pseudocopulation) . , , ... (Van der Cingel 1995, Delforge 1995, 2001).
20
. ORCHIDACEAE
. Orchis israelitica Bellevalia flexuosa (Dafni & Ivri 1981). O. israelitica B. flexuosa . . Orchis pallens Lathyrus vernus. O. pallens L. vernus. O. pallens Lathyrus (Dafni 1987). , - . , - . Delforge (1995), Serapias, .. Anacamptis papilionacea A. morio . , . Dafni (1987) Serapias , , , . , . , , (Delforge 1995). Catling (1990), . , . , .
21
. ORCHIDACEAE
. . , . , . Ophrys apifera Epipactis Cephalanthera. , Spiranthes spiralis, Neottia nidus-avis Neottia ovata (Dafni 1987, Catling 1990, Delforge 1995, Summerhayes 1969). Ophrys Ophrys apifera (Dafni 1987). , (Delforge 1995). , Cozzolino et al. (2001), . , .
I.4 Chase (2001) , 110 , . , Orchidaceae , .. Iridaceae, Asparagales. -
22
. ORCHIDACEAE
- - - . , , , , 100-110 . . , Orchidaceae, , . , . Protorchis monorchis Massalongo Palaeorchis rhizoma Massalongo, , (Schmid & Schmid 1977, Jacquet 1994). Eoorchis miocaenica Mehl . 15 ( ) (Mehl 1986). , . 110 , , . (Chase 2001). , , (Aceto et al. 1999). , . , , . , , , . , ( )
23
. ORCHIDACEAE
. , Ophrys (Phitos et al. 1995) Serapias (Del Prete & Mazzola 1995) . Dafni (1987), Ophrys , . , , , . Phitos et al. (1995) Ophrys , 36% . Del Prete (1984) Ophrys , . , , , - . , , .. ( , / ) , . , . Hedrn (Aceto et al. 1999), , , Dactylorhiza. , taxa, Sanford (1985). 24
. ORCHIDACEAE
. , , . , Dafni & Ivri (1981) (interspecific), , . , Orchis s.l. . (Dafni 1987). - - ( , ). , Ophrys, Sundermann (1977), ( Ophrys apifera) (.. O. speculum Campsoscolia ciliata) . Ophrys Orchis Serapias Greilhuber & Ehrendorfer (1975), Orchis s.l., Serapias, Anacamptis, Ophrys Himantoglossum. , Serapias Orchis Ophrys. Ophrys , (Kullenberg & Bergstrm 1973). , Ophrys (), Orchis . . , Ophrys , . 25
. ORCHIDACEAE
Ophrys, Orchis, (Tuttin et al. 1980). (Dafni 1987). Ophrys, Orchis (Baumann & Knkele 1982). Dactylorhiza, Orchis, . (Baumann & Knkele 1982), ( ) . Averyanov (1990) Dactylorhiza . Cephalanthera Orchis s.l., Ophrys, Serapias, Himantoglossum .. - - , . - - (.. D. incarnata) Dactylorhiza (.. D. sambucina) Orchis s.l. , Dactylorhiza Poltava ( ) (Poltava flora), . , , . , , , , . 26
. ORCHIDACEAE
, . , . Dactylorhiza Gymnadenia, Leucorchis Coeloglossum [ Bateman et al. (1997) Dactylorhiza], , () . () , , - . . (Orchis s.l., Ophrys, Himantoglossum, Aceras, Anacamptis, Comperia, Serapias).
27
.
. .1 ( ) 23 25 ( ) 24 45 40 48 41 34 . , . , , . . : , , , , , (. 3). , . 1:50.000 . : , . , - - . . : ) , , ) , ) , ( )
29
.
( ). . ( 2.232 m, 2.194 m, 2.111 m, 2.020 m 1.996 m). 1.768 m. 1.115 ( ).
3. ( ). Figure 3. Map of east Macedonia (NE Greece).
: -, 380 km2. . ,
30
.
. (1.963 m) (1.952 m) (1.820 m). , , , (Karagiannakidou & Kokkini 1987). : . 324 km2 62 % 38 % . , , -. (1.956 m), (1.942 m), (1.835 m), (1.723 m) .. : ( ) , . . 200 m ( ) 1.950 m . (1.814 m), (1.811 m), (1.621 m) .. , : , , , , . , . . . . . ( ) . 31
.
( ) . , - . : . . , . . (2.212 m) (2.172 m) . : . , , . , , . . (1.849 m), (1.641 m), (1.414 m) (1.323 m).
.2 (. 3). , .
32
.
, (7) , (2) , (4) (1) . . 3. (: , : ) . Table 3. Climatic data of the meteorological stations (: rainfall, : temperatre) of the study area.
570 900 1.240 110 610 790 50 40 220 310 250 620 390 660
1977-1989 1950-1964 1978-1990 1960-2002 1965-1981 1964-1998 1990-2000 1931-1940 & 1956-2004 1990-2000 1930-1940 1964-1998 1964-1998 1964-1998 1964-1998
, , , , , , ,
(1993) (, ), (2001) (), (2004) (), .. (2004) (, , , ), (1985) (), ... (, ) .. (: ) (, ).
33
.
.2.1 , , . . 4 . 4. . Table 4. Mean annual and mean monthly rainfall values of the meteorological stations of the study area. 681,0 955,1 980,0 618,1 593,5 618,2 623,5 513,2 642,8 598,8 654,6 799,8 684,3 802,0 54,4 112,2 61 49,9 49,5 97,4 54,4 46,1 69,5 61,1 70,2 81,1 64,1 63,1 71,1 84,1 99 52,7 43,4 110,6 46,9 40,1 85,6 36,0 62,0 78,0 62,8 75,5 51,9 75,1 75 48,1 35,6 95,0 46,9 38,7 58,6 43,9 54,4 65,7 55,9 50,1 53,6 62,5 79 47,3 43,0 73,3 51,4 41,1 48,8 52,1 53,7 57,7 46,7 58,5 69,8 90,6 83 52,4 58,0 90,4 61,6 47,5 57,5 47,0 70,9 75,6 60,9 80,5 63,8 103,0 96 58,6 56,0 67,9 33,6 49,8 31,2 63,3 51,6 57,1 61,6 84,0 59,8 66,0 75 54,1 48,6 61,6 24,9 31,3 21,9 34,6 35,4 56,4 55,6 55,6 47,3 28,7 61 30,7 39,0 57,0 18,9 23,7 18,5 28,7 36,7 43,9 57,0 58,0 24,1 39,1 47 30,0 30,1 39,0 25,5 25,9 25,5 22,6 24,9 38,7 33,4 40,6 31,8 74,6 67 53,9 63,6 69,4 60,7 46,5 43,6 62,8 46,5 58,1 42,2 61,3 89,4 114,0 119 68,3 66,8 120,9 72,9 56,1 82,5 55,2 81,5 87,5 66,2 82,6 64,0 104,6 118 73,3 59,9 130,7 125,7 66,4 99,5 86,6 67,4 99,8 77,9 92,2
4 , , , . , , , , , , , . , , ,
34
.
, , , , , , , , . , (130,7 mm) (18,5 mm), .
100
(mm)
125
(mm)
75 50 25 0
100 75 50 25 0
125
100
(mm)
100 75 50 25 0
(mm)
75 50 25 0
150
150
(mm)
125 100 75 50 25 0
(mm)
125 100 75 50 25 0
35
.
75
125
(mm)
(mm) (mm) 0
100 75 50 25 0
50
25
0
100 75 50 25 0
100
(mm)
75 50 25
100
125
(mm)
100 75 50 25 0
(mm) (mm)
75 50 25 0
75
100
(mm)
75 50 25 0
50
25
0
4. . Figure 4. Mean monthly rainfall diagrams.
36
.
4, , . , 4 . .2.2 ( 1992). , , , , , . 5 . 5. . Table 5. Mean monthly temperature of the study area. 1,5 -1,6 4,5 2,8 4,4 3,9 3,9 2,8 -1,1 6,2 4,5 6,7 6,3 5,3 6,3 2,5 9,3 7,3 8,1 9,7 6,7 11,0 6,5 13,9 11,7 14,9 14,4 12,6 15,2 11,3 19,1 16,9 18,5 19,7 16,2 18,9 15,6 23,5 20,7 21,8 24,4 20,9 21,3 17,4 25,8 23,3 24,1 26,5 24,1 20,9 17,4 25,6 22,3 22,7 25,6 25,1 17,9 14,5 21,7 18,9 21,0 21,6 22,3 12,0 9,2 16,1 13,7 14,2 15,7 15,1 6,6 3,4 10,3 8,9 12,1 9,4 10,0 3,1 0,5 5,6 5,1 6,0 4,8 4,1
5 5, Co, , 3 Co . 6 7 , . , 37
.
. , .
30 25 20 15 10 5 0 -5
(C )
o
5. . Figure 5. Diagrams of mean monthly temperature values. 6. . Table 6. Mean maximum monthly temperature values. 5,7 1,6 9,7 5,7 9,0 8,1 8,0 7,3 2,4 11,6 7,9 11,7 10,7 12,0 11,0 5,8 15,5 10,7 17,1 14,8 15,1 15,7 10,0 20,1 15,3 20,8 19,8 17,0 20,1 14,8 25,7 20,1 25,6 25,3 21,2 23,8 19,5 30,2 24,5 27,9 30,1 27,8 26,3 21,8 31,6 27,1 30,6 32,3 29,6 26,3 21,9 31,9 26,5 26,6 31,8 28,1 23,6 19,1 28,3 23,1 23,4 28,1 25,3 17,7 13,4 22,7 17,6 20,0 21,4 17,8 11,0 6,9 15,9 10,9 15,3 13,9 15,0 7,1 3,7 11,1 8,1 11,1 8,8 11,7
7. . Table 7. Mean minimum monthly temperature values. -3,7 -5,6 0,1 0,4 1,8 -0,1 -0,1 -2,6 -4,6 1,2 1,4 3,6 1,3 1,1 3,2 -1,2 2,5 3,9 4,3 3,9 2,9 4,8 2,2 6,2 7,6 9,9 7,8 6,7 8,6 6,3 11,0 12,2 12,8 12,6 11,3 11,7 9,7 14,3 15,8 16,2 16,6 16,2 13,8 11,5 16,5 17,9 18,7 18,6 18,0 13,3 11,5 16,5 17,3 17,3 17,8 17,0 10,5 9,0 12,9 14,1 13,4 14,3 12,4 5,6 4,1 8,8 9,4 6,8 9,6 8,5 1,1 -0,3 4,1 5,6 5,1 4,7 3,8 -1,5 -2,9 0,8 2,3 2,5 0,8 1,6
38
.
.2.3 - . (Flokas et al. 1983) 8. 8. . Table 8. Mean monthly temperature values of Scaloti. -1,4 0,7 3,8 10,0 15,5 19,2 20,2 20,2 16,0 10,2 5,4 1,0
(. 6) , , . , ( ), . . , ( ), ( ). , . (1980) , . -, -. , -. Emberger (Emberger
39
.
1955). 9, ( , ), ( , ), ( ). 9. Kppen Emberger ( 1980). Table 9. Climatic types (according to Kppen) and bioclimatic zones (according to Emberger).
Kppen Csb Csb Cfb Cfa Csa Csa Csa Cfa
Q2 79,8 126,7 67,21 75,16 74,8 54,75 77,46
m -3,7 -5,6 0,1 -0,1 1,8 -0,1 0,4
, Kppen ( 1992) : .- Csa , ( ), . .- Csb ( ), . .- Cfa ( ), , . .- Cfb ( ), , .
40
.
(mm)
(mm)
80 60 40 20 0
40 30 20 10 0
100 80 60 40 20 0
50 40 30 20 10 0
80 60 40 20 0 40 30 20 10 0 140 120 100 80 60 40 20 0
70 60 50 40 30 20 10 0
(mm)
(mm)
(C )
80 60 40 20 0 40 30 20 10 0 120 100 80 60 40 20 0
60 50 40 30 20 10 0
(mm)
(mm)
80 60 40 20 0 40 30 20 10 0120 100 80 60 40 20 0
60 50 40 30 20 10 0
(mm)
(mm)
(Co)
6. .. . Figure 6. Ombrothermic diagrams of the meteorological stations of the study area.
41
(Co)
(Co)
(Co)
(C )
o
o
(Co)
(Co)
100
50
120
60
.
I.3 , . , . - , , , - . , . , : ) - - ) , . , , , , . , , ( 1985). 10 . 10, . ( ), , , (, , ). , (, ) (, , , , ). , , . . (, , , , , ). ,
42
.
. , ( 1985, 1981, 1965, Meyer & Pilger 1963, Kronberg et al. 1970 ..) . 10. (.... 1983). Table 10. Geological substrates of the study area (I.G.M.E. 1983).
, - , , , - - , - (, , ) - (, ), , - , - , , (, ) (, ) (, ), , , (, , )
43
.
.4 . , , . , , . Ca, Mg, K > 50 % ( 1977, 1992). , , , . . - - , , Ca, Mg, K. , 50 %, 50 % ( 1977). , ( 1977, 1992). ( ) , ( ) , 55-65 %. , , , , . . . , 11 . C/N,
44
.
, . , 11 . , , . ( 1985). .. (1991), () , . (1993), (pH: 5,40-6,20), , . , (pH: 4,20-5,40), , . , & (1996) , (, ), , , N, Ca Mg K P . , (1992) ( ) , , , . Schreiber (1998), pH (5,4-6,6) , pH (6,0-7,7). Nakos (1984) - - pH 6,9 7,3. (2001), , , pH (4,3-5,6). 45
.
2,88-11,62 % 0,6-1,6 mg/100 gr . - - . pH, 3,3-5,3, 2,4-35,0 % 0,8-3,6 mg/100 gr . , pH, 4,3-5,1, 5,9-11,2 % 0,61,4 mg/100 gr . , pH, 3,7-5,6, 8,8-18,8 % 0,8-2,6 mg/100 gr . Karagiannakidou et al. (2001) pH , .
I.5 , , , , . . , , , () ( .. 1996). , - , , , . , .
46
.
.5.1 - - Quercus coccifera 1.000 m. , . , Carpinus orientalis, Fraxinus ornus . Paliurus spina-christi, Cornus mas, Quercus petraea subsp. medwediewii, Fraxinus ornus, Juniperus oxycedrus, Ulmus campestris, Prunus spinosa, Cytisus sp., Chamaecytisus sp. .. Quercus frainetto Q. pubescens . (. 3) , , . (1990), Alnus glutinosa, Tilia tomentosa, Corylus avellana, Juglans regia, Clematis vitalba .. (Fagus sylvatica s.l.) . , . Schreiber (1998). , Pinus nigra subsp. nigra, Quercus coccifera 800 m . , . - - , . Abies borisii-regis
47
.
. , , 36.400 ( & 2003). (Schreiber 1998) . . , (.. , ) (Horvat et al. 1974, & 1992). Quezel (1989) Schreiber (1998). , . Festucetalia valesiacae (Horvat et al. 1974). , . .5.2 , , , . . Quercus coccifera . , . ( ) Q. coccifera 300(-600) 1.300 m. Juniperus oxycedrus subsp. oxycedrus, Crataegus monogyna, Cistus incanus Daphne oleoides. 48
.
Karagiannakidou & Kokkini (1987), Carpinus orientalis. Corylus avellana, Fraxinus ornus, Ostrya carpinifolia, Syringa vulgaris, Ligustrum vulgare, Cornus mas, Lonicera etrusca, Quercus pubescens. . - - , - - . , , Quercus pubescens Quercus frainetto. Juniperus oxycedrus subsp. oxycedrus, Sorbus domestica, Corylus avellana, Acer obtusatum Fraxinus ornus. (Castanea sativa) - ( ) 700 1.000 m. , - , Tilia tomentosa, Ostrya carpinifolia, Cornus mas, Carpinus orientalis, Quercus frainetto Fraxinus ornus. , Carpinus orientalis, Fagus sylvatica s.l. , 600 m - - 1.600 m ( ) 1.350 m ( , ). - - , 900-1.200 m. Quercus sp., Ostrya carpinifolia, Fraxinus ornus, Carpinus orientalis, Juniperus oxycedrus subsp. oxycedrus Sorbus domestica. , Abies borisii-regis.
49
.
, (Populus tremula) . , 1.100 1.963 m ( ). Daphne oleoides, Rosa pimpinelifolia, Genista depressa (Karagiannakidou & Kokkini 1987). Karagiannakidou (1994). .5.3 Papanicolaou (1985), Quercus coccifera, ( - ) 300-700 m ( 900 m, 500 m .. ). , , Q. coccifera Erica manipuliflora. , . - . Juniperus oxycedrus, Erica manipuliflora Phillyrea latifolia, Ostrya carpinifolia, Carpinus orientalis, Pistacia terebinthus, Acer monspessulanum, Fraxinus ornus, Pyrus spinosa Paliurus spina-christi. , - . , , 700 900 m, (Fagus sylvatica s.l.) (50 %) . , , 900 m . , 50
.
, (Abies borisiiregis), - - . , , 700 1.700 m, 1.850 m. , , , , 1.450 m. Schreiber (1998) Symphytum ottomanum-Fagus sylvatica , Primula veris subsp. suaveolens-Fagus sylvatica Vaccinium myrtillus-Fagus sylvatica Fagion sylvaticae. , , , . Moder. , , Populus tremula. , , 500 m, Betula pendula. Betula pendula (Papanicolaou 1985). , , 34 km2 (Karagiannakidou et al. 2001). , . . 10 % ( .. 1997, Karagiannakidou et al. 2001). Karagiannakidou et al. (2001) , . ,
51
.
. .5.4 (. 3). .5.4.1 , 2.657 Ha. , ( .. 1991) 900 1.400 m. Quercus petraea subsp. medwediewii. 1.000 m, 1.100 m. Carpinus orientalis. Pinus sylvestris, 1.000 m, (Picea abies subsp. abies). , - : . 1.000 1.150 m, ( .. 2004). , , , (Mller 1929). ( .. 1991),
52
.
( 2001). , Alnus incana subsp. incana . , , ( .. 1992, 2001). .5.4.2 . . , . Carpinus orientalis, Juniperus oxycedrus subsp. oxycedrus, Fraxinus ornus Quercus frainetto, Quercus petraea subsp. medwediewii Pinus sylvestris. Fagus sylvatica s.l., Pinus sylvestris Picea abies subsp. abies , (1992) Fagetalia. (2001) , . Fagetalia (1992) (Vaccinio-Piceetalia) Vaccinio-Piceion Pinetum sylvestris Piceetum abietis. 1.000 1.600 m. Pinus sylvestris . Juniperus communis subsp. communis, Fraxinus ornus, Rubus idaeus, Salix caprea, Populus tremula
53
.
Betula pendula. Vaccinium myrtillus, V. vitis-idaea, Bruckenthalia spiculifolia. (1990) , , 20 , . , , / , (Zoller et al. 1977, 1990). Zoller et al. (1977) Vaccinium myrtillus, , Luzula , . Gamisans & Hebrard (1980) . Vaccinio-Piceetalia (Piceetum abietis) 1.200 1.800 m. ( 1992). (1992) 2.754 Ha. & (1981) --, . Juniperus communis subsp. communis, Sorbus aucuparia, Salix caprea Corylus avellana, Vaccinium myrtillus, Luzula, Fragaria vesca .. (Abies borisii-regis) ( 2001). , ( moor). Poaceae Cyperaceae 54
.
. (1972) 1.550 m . .5.4.3 . - - Carpinus orientalis, Fraxinus ornus, Juniperus oxycedrus subsp. oxycedrus Corylus avellana. . (Pinus nigra subsp. nigra) . 400 m, 900 1.300 m . Quercion confertae Fagetalia. , . (2001) , , . , , . . -- (Fagus sylvatica s.l. Picea abies subsp. abies Abies borisii-regis). , . , Populus tremula Betula pendula ( 1985). 55
.
Zoller et al. (1977) Luzulo-Fagion , Galio odorati-Fagion . (Pinus peuce) ( 2001). , (Rumex arifolius-Agrostis capillaris, Linum capitatum subsp. capitatum-Festuca varia Silene lerchenfeldiana-Sempervivum heuffelii) ( .. 2001). .5.4.4 Carpinus orientalis, Fraxinus ornus, Juniperus oxycedrus subsp. oxycedrus, Ostrya carpinifolia .. , Quercus petraea subsp. medwediewii Q. frainetto . , Quercion confertae Fagetalia, Pinus nigra subsp. nigra. ( 2001). , , (Fagus sylvatica s.l.) . (2001) . , . 900-1.300 m , 400 m. .5.4.5 . 56
.
. Carpinus orientalis, Fraxinus ornus (Paliurus spina-christi). Quercus petraea subsp. medwediewii Q. frainetto. - - Carpinus orientalis, Ostrya carpinifolia .. 1.100 m , . (2001), , . , (Populus tremula) . , Pinus sylvestris . , Picea abies subsp. abies . Chamaecytisus sp., Rosa sp., Carpinus orientalis .. , . .5.4.6 , . , , . Rosa, Prunus spinosa, Genista sp., Crataegus monogyna Quercus coccifera Juniperus oxycedrus subsp. oxycedrus. 57
.
Carpinus orientalis . , . , Quercus petraea subsp. medwediewii Q. frainetto, (.. Carpinus orientalis, Fraxinus ornus, Cornus mas, Juniperus oxycedrus subsp. oxycedrus). Fagus sylvatica s.l. . (Picea abies subsp. abies), ( 2001). , , (Platanus orientalis). .5.5 Juniperus oxycedrus subsp. oxycedrus. , - - - . - - - . , . , Festuco Brometea, ( & 2003). -, Ostryo-Carpinion, (700-800 m) . 1.160 m, 1.600-1.700 m , . Ostrya carpinifolia, Fraxinus ornus, Carpinus orientalis, Juniperus oxycedrus subsp. oxycedrus, Quercus pubescens, Corylus avellana, Cornus 58
.
mas,
Hippocrepis
emerus
subsp.
emeroides
..
(
1957,
& 2003). Ostryo-Carpinion , 650 m. 800-1.000 m Pinus nigra subsp. nigra, Pinus sylvestris Pinus leucodermis ( ). 1.400 m ( 1.8501.900 m) . . 1.600 m, , . , 1.200-1.500 m (Abies borisii-regis). , Ligustrum vulgare, Pyrus malus, Salix caprea, Quercus dalechampii, Fraxinus excelsior, Tilia vulgaris ., Calamagrostis arundinacea Luzula sylvatica ( .. 1989). , - Juniperus communis subsp. nana ( & 2003). Karagiannakidou et. al. (1999), , . (Populus tremula) Cotoneaster integerrimus. .5.6 (Quercetalia pubescentis). 600-800 m Ostryo-Carpinion Coccifero-Carpinetum. Pinus halepensis subsp. brutia, Cupressus sempervirens Pinus nigra subsp. nigra ( 1985). 59
.
Quercus coccifera, Spartium junceum, Paliurus spina-christi, Fraxinus ornus, Juniperus sp., Quercus pubescens, Carpinus orientalis, Sorbus sp., Salix sp. ( 1976). Robinia pseudacacia . . Quercion confertae. Quercus petraea subsp. medwediewii, 1.000 m, Quercetum montanum (Fagetalia) ( 1985). 800-900 m 1.200-1.600 m. . , . Schreiber (1998) Symphytum ottomanum-Fagus sylvatica Vaccinium myrtillus-Fagus sylvatica. Acer platanoides A. pseudoplatanus, , Sorbus aucuparia, Populus tremula, Quercus petraea subsp. medwediewii, Betula pendula. Salix caprea, Rubus idaeus, Chamaecytisus thessalus, Corylus avellana . ( 1976). ( ) Juniperus communis subsp. nana, Rosa sp., Chamecytisus sp. .. , . , , . Eriophorum latifolium. Cyperaceae, Juncaceae, Orchidaceae Graminae. 60
.
, , Pseudotsuga menziesii, Picea omorica, P. sitchensis, Pinus ponderosa, Larix sp. Abies borisii-regis ( 1976, 1985).
.6 , . 92/43 .. (. 7). ( ) ( ). T 1.500-1.953 m 502 87,2 Ha. 200995/7950/20-5-1980 (... 486, ) , , . 1981 483 Ha ( . 151256/2122/6-8-1981 ), . . ( ) , , . 105497/6459/11-8-1986 (... 656, ) . 3,9 Ha .
61
.
7. Natura 2000. Figure 7. Natura 2000 sites.
, 92/43/ ( . 1996) (. 7). : () (. ), 1.085,56 Ha () (. ), 6.708,90 Ha () (. ), 7.431,51 Ha (. ), 9.845,62 Ha (. ), 10.345,47 Ha (. ), 23.288,69 Ha (. ), 4.914,83 Ha (. ), 6.799,47 Ha Natura 2000, 92/43/ 38 ( 78)
62
.
( ) ( .. 1996).
63
V. -
V. V.1 IV.1.1 taxa , . , , , , , . 662 1.000 m2 , . 2002-2006 taxa . , Spiranthes spiralis L. Chevall., , . , , . , taxa. , , Buttler (1986), Delforge (1995, 2006), Baumann et al. (2006) Kretzschmar et al. (2007). taxa Pridgeon et al. (1999, 2001, 2003, 2005), Bateman et al. (1997), Kreutz (2004), Kretzschmar et al. (2007), Delforge (1995, 2006), Nordstrm (2003) Hedrn et al. (2007). Oberdorfer (1990), Pignatti (1982), Delforge (1995, 2006), Baumann et al. (2006) Kretzschmar et al. (2007).
65
V. -
Epipactis pontica . Helmut Presser (Neumarkt in der Oberpfalz, ). (TAU). taxa . IV.1.2 662 ( ) OPTIMA (Organization for the Phyto-Taxonomic Investigation of the Mediterranean Area), . , , , : 5 UTM (WGS84) . , , (Kitanov 1943), , , , , (. 3). 1:50.000 GPS (Global Positioning System). 66
V. -
11. () . Table 11. Habitat types of the study area.
Corylus avellana (militrisch) UTM (WGS84) MultiTrans AHO V 4.1 (Rauschenberger 2005). , ... . (20) (. 11) . : : (5) : (6) (20) : (21) (50) : (50)
taxon . , : Rechinger (1939), Zaganiaris (1940), Kitanov (1943), Quzel & Contandriopoulos (1968),
67
V. -
(1972), (1976), Zoller et al. (1977), Karagiannakidou & Babalonas (1981), Hlzinger & Knkele (1983), Baumann (1983), Voliotis & Karagiannakidou (1984), (1985), Papanicolaou (1985), Hlzinger & al. (1985), Greuter & Raus (1986), Karagiannakidou & Kokkini (1987), Willing & Willing (1987), Baumann & Baumann (1988), Hlzinger & Knkele (1988), Willing & Willing (1988), & (1989), Quzel (1989), Willing (1990), Willing & Willing (1991), (1992), .. (1992), Kalopissis (1992) Karagiannakidou et al. (1995), Phitos et al. (1995), Raus (1995), Eleftheriadou & Raus (1996), Schreiber (1998), Strid & Tan (1998), .. (1999), Petermann (1999), Athanasiadis et al. (2000), Arampatzis et al. (2001), Eleftheriadou et al. (2001), Karagiannakidou et al. (2001), Presser (2001), .. (2001), (2001), Gembardt & Glz (2002), Theodoropoulos et al. (2003), & (2003), & (2003), .. (2004). , Srensen (Srensen's Quotient of Similarity) : Q/S = 2 J 100 / (a + b), J: taxa , a: taxa b: taxa .
68
V. -
V.2 , ( 1992). , ( 643 662 ) ( 5-15 cm). , , . , , . 2 mm 0,02 mm. : 1. , - 1:1 ( - 1:5). 2. , , (Nelson & Sommers 1982). 3. . 0,5 NaHCO3 pH 8,5 (Olsen & Sommers 1982) ( & 1995).
V.3 . ADE-4 (Thioulouse et al. 1997)
69
V. -
(Outlying Mean Index analysis) (Doldec et al. 2000). OMI , CCA (Canonical Correspondence Analysis) RDA (Redundancy Analysis) . ) , CCA ) (. ) . , CCA RDA , (Doldec et al. 2000). taxa ( taxon), OMI . , . , , (Thuiller et al. 2004). , (niche position) ( ), , . (niche breadth species tolerance) . , (species tolerance) ( specialist species), ( generalist species). . (taxa ) ( ), ( ) (explanatory 70
V. -
variables). , . : pH (%) (mg/100 gr ) . , Pinus halepensis subsp. brutia , . , . - - . . .... 1:50.000. : 71
V. -
- . (Heat Load Index). , , , . ... 16- . , (.. : 0 360, : 90, : 202,5 ...). , (McCune & Keon 2002, Neitlich & McCune 1997) : Heat load index=
1 ( 45) 2
. 0 1 , () (). , ( ) X Y - . (PCA: Principal Component Analysis) OMI.
72
V. -
, , Monte Carlo 10.000 . , Pearson SPSS 12 (Anon. 2003).
IV.4 - - - (Margules et al. 1988, Church et al. 1996, Slymos & Fehr 2005). , . taxa , Ophrys insectifera (Zaganiaris 1940). O. insectifera . , Platanthera bifolia Dactylorhizacordigera
. . UTM (Universe Transverse Mercator WGS 84) [. 8 ()], 10 km 100 km2. UTM.
73
V. -
8. UTM () (). Figure 8. Map of the study area in UTM projection (a) and area of orchid records.
8 (), 100 km2, . /1, /2, /3, /4 (.. GL 36/2). (Troumbis & Dimitrakopoulos 1998). [. 8 ()], . , 207 25 km2 .IV.4.1
: () (species-richness) , () (conservation value) () (rarest species-richness). . . (Williams et al. 1996):
74
V. -
= ()i =1
n
i
n : . . . (0) (1). taxa , taxa , , . (), ( 5 5 km) (Williams et al. 1996) : =1/ci
ci: i () . (.. Selvi, 1997, Sapir et al. 2003, Slymos & Fehr 2005) . , , , ( : , , , ,
75
V. -
) 100 100 km. () : Ci C
= 1
Ci: 100 100 km taxon i C: (100 100 km) (740 ) () :Ti Tmax
=1 -
Ti: (%) i Tmax: (%) 0-1. , . (Species Tolerance) OMI . (Anacamptis palustris subsp. elegans, Dactylorhiza graeca, Gymnadenia frivaldii, Neottia cordata Serapias bergonii) , (). taxa , . , () .= + +
76
V. -
: , : : 0-3. , ( ) . . , . 25 % (Gaston 1994, Williams et al. 1996, Slymos & Fehr 2005). , 25 % .IV.4.2
, (7) . 5 % (Prendergast et al. 1993, Williams et al. 1996, Slymos & Fehr 2005, Traba et al. 2008). , , 207 , 10 . , (complementarity analysis) ( , ). (Vane-Wright et al. 1991) . . 77
V. -
, . , , , . . ( ), - - . , . , . 3 ( ), . , (irreplaceability) (Williams et al. 1996, Rodrigues & Gaston 2002). , (MCA: Mixed Complementarity Analysis), ( ). , , .IV.4.3
, ( ) 78
V. -
Natura 2000, Jaccard. (van Jaarsveld et al. 1998, Dimitrakopoulos et al. 2004, Tognelli et al. 2005) : J = Nc / (N1 + N2 Nc) 100, Nc: , N1: N2: . , , , Spearman. Statistica 6.0 (StatSoft 2001). , Natura 2000 Mann-Whitney U test. : , , , (14, 25 50) .
79
V.
V. V.1 - , taxa . taxon ( taxa ), , , (. IV.1.2). I. taxon . . : : , : -, :-, : -, : , : ,
(): -, (): -, (): -, (): -, (): -, (): - : .
81
V.
V.1.1 -
1. Anacamptis coriophora (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. coriophora (Syn.: Orchis coriophora L. subsp. coriophora) Chor. type: smed(-eurassubozean) : 4 : 119: / -: 403: / -: 566: / : 61:
2. Anacamptis coriophora (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. fragrans(Pollini) R.M. Bateman, Pridgeon & M.W. Chase
(Syn.: Orchis coriophora L. subsp. fragrans (Pollini) K. Richter) Chor. type: smed(-eurassubozean) : 16 : 59: , 126: , 157: , 201: / -: 225: / -: 371: , 372: , 381:
, 385: / -: 450: / : 359: , 361: / : 44: / -: 414: / : 550: , 554:
.. (1999).
3. Anacamptis laxiflora (Lam.) R.M. Bateman, Pridgeon & M.W. Chase (Syn.: Orchis laxiflora Lamarck) Chor. type: med-smed : 6 -: 385: / -: 450: , 510: / -: 461: , 462: / -: 566:
82
V.
4. Anacamptis morio (L.) R.M. Bateman, Pridgeon & M.W. Chase (Syn.: Orchis morio L.) Chor. type: eurassubozean-smed : 38 : 122: , 123: , 125: , 126: , 149: , 154: , 157: , 159: , 161: / : 163: , 245: / -: 364: , 369: , 376: , 384: / -: 387: , 395:
, 431: , 436: , 438: , 443: , 447: , 450: , 516: / -: 332: , 335: , 412: / -: 399: , 463: , 469: / -: 541: / : 70: , 72: , 85: , 166: Baumann & Baumann (1988), Hlzinger & Knkele (1988) Arampatzis et al. (2001), () (Zaganiaris 1940).
5. Anacamptis palustris (Jacq.) R.M. Bateman, Pridgeon & M.W. Chase subsp. elegans(Heuff.) R.M. Bateman, Pridgeon & M.W. Chase
(Syn.: Orchis palustris Jacquin subsp. elegans (Heuffel & Rochel) So) Chor. type: smed-gemkont : 1 -: 227:
6. Anacamptis papilionacea (L.) R.M. Bateman, Pridgeon & M.W. Chase (Syn.: Orchis papilionacea L.) Chor. type: med : 15 : 119: , 125: , 154: , 161: / -: 364: , 369: , 382: , 384: / :
358: / : 178: , 179: / -: 335: / -: 399: / : 65: , 169:
83
V.
7. Anacamptis picta (Loisel.) R.M. Bateman (Syn.: Orchis picta Loisel.) Chor. type: med-smed : 123 : 118: , 119: , 122: , 123: , 125: , 126: , 133: , 149: , 154: , 157:
, 159: , 161: , 202: , 203: / -: 163: , 165: , 215: , 224: , 228: , 230: , 245: , 326: / -: 296: , 312: , 364: , 365: , 369: , 371: , 376: , 378: , 381: , 382: , 383: , 384: , 385: , 394: , 368: / -: 387: , 392: , 395: , 430: , 431: , 433: , 435: , 436: , 438: , 439: , 441: , 442: , 443: , 447: , 449: , 450: , 452: , 453: , 511: , 516: , 517: , 519: , 526: , 652: / : 188: , 190: , 192: , 194: , 247: , 281: , 342: , 350: , 358: , 360: , 361: , 362: / : 45: , 97: , 111: , 178: , 179: / -: 332: , 335: , 397: , 404: , 411: , 412: , 413: , 414: / -: 399: , 400: , 457: , 459: , 461: , 463: , 464: , 469: , 472: , 476: , 477: / -: 545: , 559: / -: 541: , 550: , 554: / : 4: , 8: , 23: , 26: , 28: , 29: , 32: , 38: , 40: , 65: , 70: , 72: , 78: , 85: , 86: , 98: , 166: , 168: , 169: , 613: , 618: Arampatzis et al. (2001) .
8. Anacamptis pyramidalis (L.) L.C.M. Rich. Chor. type: med-smed(-subatl) : 53 : 59: , 123: , 159: , 161: / -: 163: , 174: , 205: , 211: , 215:
, 219: , 222: , 225: , 325: , 326: , 328: , 631: / -: 240: , 249: , 290: , 295: , 330: , 381: , 383: , 630: , 638: / -: 431: , 436: , 526: / : 114: , 184: , 196: , 232: , 247: / : 12: , 44: , 45: , 99: , 612: / -: 332: , 333: , 335: / -: 400: , 457: , 458: , 465: , 476: , 645: / -: 547: , 549: / : 6: , 38: , 70: , 616:
84
V.
Arampatzis et al. (2001) , Rechinger (1939) , Karagiannakidou & Babalonas (1981), Kitanov (1943), Quzel & Contandriopoulos (1968), Baumann & Baumann (1988), Quzel (1989) Schreiber (1998), , Karagiannakidou & Babalonas (1981) Petermann (1999).
9. Cephalanthera damasonium (Mill.) Druce Chor. type: smed-subatl : 79 : 121: , 134: , 139: , 142: , 143: , 152: / -: 171: , 172: , 175:
, 212: , 214: , 216: , 218: , 221: , 228: , 229: , 242: , 328: / : 239: , 241: , 251: , 255: , 297: , 307: , 308: , 316: , 324: , 331:
, 373: , 377: , 379: / -: 454: , 518: / : 231: , 235: , 259: , 266: , 269: , 286: , 343: / : 13: , 47: , 48: , 100: , 103: , 106: , 107: , 108: , 112: , 619: / -: 336: , 406: , 410: / -: 398: , 424: , 465: , 466: , 467: , 468: , 474: , 478: , 480: , 481: , 483: , 497: , 501: , 503: , 508: / -: 536: , 539: , 577: , 581: , 596: /-: 547: / : 62: , 64: , 81: , 87: , 617:
Baumann & Baumann (1988) Arampatzis et al. (2001) , Karagiannakidou & Babalonas (1981) . Kitanov (1943), Schreiber (1998), Baumann & Baumann (1988) Voliotis & Karagiannakidou (1984) Karagiannakidou & Babalonas (1981) . , Karagiannakidou & Babalonas (1981), Rechinger (1939), Schreiber (1998) Voliotis & Karagiannakidou (1984). .. (2004) , (1992) (2001) , (2001) .
85
V.
(Zaganiaris 1940) Cephalanthera cucullata Bois. et Heldr. ( ) , C. cucullata . , ..., C. damasonium C. longifolia . C. epipactoides Fischer & C.A. Meyer (Phitos et al. 1995). - .
10. Cephalanthera longifolia (L.) Fritsch Chor. type: smed-eurassubozean : 61 : 55: , 129: , 144: , 146: , 162: / -: 170: , 172: , 210: , 213:
, 216: , 219: , 229: / -: 237: , 238: , 239: , 241: , 250: , 251: , 297: , 301: , 309: , 365: , 379: , 641: / -: 437: , 445: , 448: , 451: / : 113: , 184: , 185: , 186: , 187: , 267: , 272: , 273: , 284: , 344: , 350: , 351: , 352: , 354: / -: 416: , 636: / -: 402: , 479: , 490: / -: 539: , 576: , 577: , 596: / -: 565: , 567: / -: 552: , 601: , 655: / : 27: , 62: , 64: , 82: , 167: Baumann & Baumann (1988) Arampatzis et al. (2001), Schreiber (1998), Baumann & Baumann (1988) Willing & Willing (1991), .. (2004), (1992) (2001), (2001).
86
V.
11. Cephalanthera rubra (L.) L.C.M. Rich. Chor. type: smed-gemkont : 65 : 121: , 130: , 134: , 136: , 143: , 146: , 147: , 152: / -: 172:
, 212: , 213: , 214: , 216: , 218: , 221: , 229: , 242: / -: 239: , 241: , 254: , 297: , 307: , 324: , 329: , 380: / -: 451: , 520: /: 259: , 260: , 261: , 266: , 269: , 282: , 339: , 343: , 344: / :
47: , 100: , 103: , 107: , 180: / -: 406: / -: 398: , 421: , 424: , 458: , 483: , 487: , 501: , 503: , 508: / -: 535: , 536: , 581: / -: 543: , 547: , 561: , 563: , 565: , 584: / -: 551: , 553: / : 84: , 87: , 617: Baumann & Baumann (1988) Arampatzis et al. (2001), Karagiannakidou & Babalonas (1981), Baumann & Baumann (1988), Rechinger (1939), Kitanov (1943) Schreiber (1998), Petermann (1999), Karagiannakidou & Babalonas (1981), Papanicolaou (1985) Schreiber (1998), () Karagiannakidou & Babalonas (1981) .. (2004), () (1992), Karagiannakidou & Babalonas (1981), Petermann (1999) (2001), (, ) (2001).
12. Corallorhiza trifida Chtel. Chor. type: no(kont), circ : 39 : 134: , 136: , 143: / -: 254: , 298: , 316: / : 269: / :
102: , 106: / -: 415: , 422: / -: 484: , 488: , 489: , 490: , 492: , 493: , 494: , 496: , 497: , 498: , 500: , 501: , 502: , 503: , 506: , 509: , 527: , 587: , 588: , 651: / -: 536: , 575: , 581: , 591: / -: 569: , 584: / -: 604: / : 614:
87
V.
Baumann & Baumann (1988), Schreiber (1998), () .. (2004), (1992), Eleftheriadou et al. (2001), (2001) Zoller et al. (1977), Raus (1995), Eleftheriadou & Raus (1996), Eleftheriadou et al. (2001) (2001), (2001), Karagiannakidou & Babalonas (1981) (2001) (2001). , Baumann & Baumann (1988) Raus (1995).
13. Dactylorhiza cordigera (Fries) So Chor. type: balc : 14 -: 405: , 423: / -: 505: , 507: , 589: , 647: , 648: , 650: / -: 662: / -: 605: , 606: , 609: , 610: / : 41:
() (1992) (1972), Karagiannakidou & Babalonas (1981), Baumann (1983), Hlzinger & Knkele (1983), Baumann & Baumann (1988), Willing & Willing (1991) Strid & Tan (1998).
14. Dactylorhiza graeca H. Baumann Chor. type: balc : 1 : 41:
Phitos et al. (1995), Willing & Willing (1987), Baumann (1983), Hlzinger & Knkele (1983), Baumann & Baumann (1988), Presser (2001) (1976).
88
V.
15. Dactylorhiza incarnata (L.) So Chor. type: no-euras : 4 : 148: , 621: / -: 227: / -: 405:
Baumann (1983), Hlzinger & Knkele (1983), Willing (1990) Willing & Willing (1991), Willing (1990).
16. Dactylorhiza macedonica Hlzinger & Knkele Chor. type: balc : 3 : 148: / -: 405: / -: 462:
Willing & Willing (1988), Hlzinger & Knkele (1988) Willing & Willing (1991), Willing & Willing (1988).
17. Dactylorhiza romana (Sebast.) So Chor. type: omed : 3 -: 390: / -: 388: / : 354:
18. Dactylorhiza saccifera (Brongn.) So Chor. type: osmed : 7 : 155: , 625: / -: 309: / -: 482: / -: 580: , 596: ,
660:
89
V.
Karagiannakidou & Kokkini (1987) Hlzinger & Knkele (1988), Schreiber (1998), Karagiannakidou & Babalonas (1981) Rechinger (1939) (1992) (). , Hlzinger & Knkele (1983) Baumann & Baumann (1988).
19. Dactylorhiza sambucina (L.) So Chor. type: smed-pralp : 98 : 53: , 69: , 75: , 76: , 116: , 118: , 122: , 123: , 135: , 137: ,
141: , 145: , 149: , 153: , 154: , 162: , 199: / -: 206: , 208: , 209: / -: 253: , 292: , 293: , 294: , 295: , 297: , 303: , 305: , 306: , 310: , 314: , 315: , 317: , 318: , 319: , 320: , 321: , 322: , 323: , 329: , 376: , 643: / -: 516: / : 263: , 264: , 265: , 271: , 277: , 285: , 633: / : 47: , 48: , 50: , 51: , 88: , 89: , 93: , 96: , 100: , 102: , 104: , 106: , 620: / -: 407: , 408: , 413: , 416: , 418: , 419: , 420: , 585: / -: 469: , 471: , 475: , 482: , 486: , 490: , 504: , 590: / -: 537: , 539: , 574: , 578: , 579: , 592: , 660: / -: 602: , 611: / : 23: , 26: , 61: , 65: , 68: , 72: , 78: , 79: , 80: , 98:
Karagiannakidou et al. (1995), Baumann & Baumann (1988) Karagiannakidou & Kokkini (1987), Karagiannakidou & Babalonas (1981), Karagiannakidou & Babalonas (1981), & (2003), Karagiannakidou et al. (1995), Kitanov (1943), Willing & Willing (1991), Baumann & Baumann (1988) Schreiber (1998), Karagiannakidou & Babalonas (1981), Karagiannakidou et al. (1995), Karagiannakidou et al. (2001) Schreiber (1998), & (2003), .. (2004), Theodoropoulos et al. (2003) (),
90
V.
(1992) () Eleftheriadou & Raus (1996) (). , Karagiannakidou & Babalonas (1981), Hlzinger & Knkele (1983), Strid & Tan (1998) (1976).
20. Dactylorhiza viridis (L.) R.M. Bateman, Pridgeon & M.W. Chase (Syn.: Coeloglossum viride (L.) Hartman) Chor. type: no-pralp, circ : 15 : 126: , 153: / -: 207: / -: 255: , 298: , 305: , 311: , 315:
, 320: , 322: , 632: / : 263: , 265: , 280: / : 48: Karagiannakidou et al. (1995) Baumann & Baumann (1988) , Karagiannakidou et al. (1995), Karagiannakidou & Babalonas (1981), Kitanov (1943), Willing & Willing (1991), Baumann & Baumann (1988) Schreiber (1998) Karagiannakidou & Babalonas (1981), Karagiannakidou et al. (1995), Rechinger (1939) Papanicolaou (1985) .
21. Epipactis atrorubens Hoffm. ex Besser subsp. atrorubens Chor. type: (no-)eurassubozean-smed : 8 -: 298: , 305: , 311: , 323: , 634: , 635: / : 261: / : 92:
Baumann & Baumann (1988), Karagiannakidou & Babalonas (1981), Karagiannakidou et al. (1995), Kitanov (1943), Baumann & Baumann (1988) Voliotis & Karagiannakidou (1984), Karagiannakidou & Babalonas (1981), Voliotis & Karagiannakidou (1984), () Karagiannakidou & Babalonas (1981).
91
V.
22. Epipactis atrorubens Hoffm. ex Besser subsp. spiridonovii (Devillers & Devillers Terschuren) Kreutz
Chor. type: balc : 1 : 49:
23. Epipactis exilis P. Delforge Chor. type: smed : 17 -: 256: / : 278: , 287: , 347: / -: 483: , 485: , 487: , 491:
/ -: 530: , 579: , 582: / -: 561: , 583: , 584: / -: 603: , 607: / -: 553: (2001) , , , .
24. Epipactis helleborine (L.) Crantz Chor. type: euras(-subozean)-smed : 127 : 121: , 130: , 134: , 136: , 139: , 142: , 143: , 147: , 152: , 162:
, 622: / -: 170: , 212: , 213: , 214: , 216: , 221: , 223: , 226: , 228: , 229: , 242: , 629: / -: 239: , 241: , 251: , 254: , 297: , 308: , 329: , 375: , 379: , 380: , 386: , 391: , 640: / -: 445: , 454: , 522: , 644: / : 234: , 246: , 259: , 260: , 261: , 266: , 269: , 272: , 276: , 282: , 286: , 339: , 343: , 344: / : 13: , 47: , 103: , 107: / -: 334: , 336: , 406: , 408: , 409: , 416: , 637: / : 401: , 402: , 421: , 424: , 458: , 465: , 466: , 467: , 468: , 474:
, 475: , 480: , 481: , 482: , 483: , 487: , 490: , 491: , 497: , 498: , 501: , 503: , 508: / -: 535: , 536: , 539: , 576: , 577: , 581: , 582: , 594: / -: 547: , 559: , 561: , 563: , 565: , 567: , 583: , 92
V.
584: , 657: / -: 546: , 551: , 553: , 555: , 598: , 599: , 601: /: 9: , 10: , 20: , 33: , 34: , 42: , 43: , 62: , 64: , 71: , 74: , 80:
, 87: , 167: , 617: Baumann & Baumann (1988) Arampatzis et al. (2001), Karagiannakidou & Babalonas (1981), Baumann & Baumann (1988), Schreiber (1998) Willing & Willing (1991), Petermann (1999), Karagiannakidou et al. (1995) Schreiber (1998), & (2003), () & (1989), . (1992), .. (2004) Theodoropoulos et al. (2003), () (1992), Eleftheriadou et al. (2001), Petermann (1999) (2001), , () (2001), Karagiannakidou & Babalonas (1981) (2001), Baumann & Baumann (1988), (1985), Karagiannakidou & Babalonas (1981) (1976).
25. Epipactis microphylla (Ehrh.) Sw. Chor. type: smed : 15 : 136: , 152: / -: 254: , 324: / -: 518: / : 279: , 343: ,
344: / -: 424: , 458: , 508: / -: 581: / -: 543: , 547: , 565: Baumann & Baumann (1988), , , (2001).
93
V.
26. Epipactis nauosaensis Robatsch Chor. type: balc : 2 : 136: / -: 254:
Presser (2001) .
27. Epipactis palustris (L.) Crantz Chor. type: euras(-subozean)-smed : 6 : 148: / -: 227: / -: 405: / -: 505: / -: 566: / : 658:
Karagiannakidou & Babalonas (1981) Kitanov (1943), Hlzinger & Knkele (1988), () (1992).
28. Epipactis pontica Taubenheim Chor. type: europkont-balc : 3 -: 530: , 593: , 597:
29. Epipogium aphyllum Sw. Chor. type: no-pralp : 3 -: 483: , 485: , 488:
94
V.
() Athanasiadis et al. (2000) (2001).
30. Goodyera repens (L.) R. Br. Chor. type: nokont, circ : 10 -: 298: / -: 408: , 417: , 418: , 420: , 661: / -: 481: ,
482: , 486: , 495: Schreiber (1998) .. (2004) Eleftheriadou et al. (2001) .
31. Gymnadenia conopsea (L.) R. Br. Chor. type: no-euras(-smed) : 43 : 75: , 76: , 135: , 140: , 141: / -: 206: , 208: , 209: , 210: ,
213: , 225: / -: 255: , 294: , 295: , 298: , 303: , 304: , 305: , 314: , 315: , 318: , 319: , 320: , 323: / -: 450: / : 263: , 264: , 265: , 270: , 271: , 280: / : 44: , 50: , 51: , 92: , 101: / -: 482: , 504: , 649: / -: 539: / -: 568: / -: 602: , 611: Baumann & Baumann (1988) Karagiannakidou & Babalonas (1981), Karagiannakidou & Babalonas (1981), Karagiannakidou & Babalonas (1981), Baumann & Baumann (1988), Voliotis & Karagiannakidou (1984), Willing & Willing (1991), Gembardt & Glz (2002), Schreiber (1998), Papanicolaou (1985) Karagiannakidou et al. (2001), Voliotis & Karagiannakidou (1984). ,
95
V.
(1992) () Eleftheriadou & Raus (1996) .. (2001) .
32. Gymnadenia frivaldii Hampe ex Grieseb. Chor. type: balc : 1 -: 505:
Willing & Willing (1991), Phitos et al. (1995) Strid & Tan (1998).
33. Gymnadenia rhellicani (Teppner & E. Klein) Teppner & Klein (Syn.: Nigritella rhellicani Teppner & E. Klein) Chor. type: alp : 4 -: 318: , 320: , 321: , 322:
Karagiannakidou & Babalonas (1981), Karagiannakidou et al. (1995), Baumann & Baumann (1988), Gembardt & Glz (2002), Voliotis & Karagiannakidou (1984) Schreiber (1998).
34. Himantoglossum caprinum (M. Bieb.) Spreng. Chor. type: omed-osmed : 66 : 59: , 69: , 197: , 200: / -: 174: , 176: , 204: , 205: , 215: ,
219: , 220: , 222: , 224: , 326: , 328: / -: 240: , 250: , 330: , 366: , 371: , 373: , 382: , 384: , 641: / -: 374: , 430: , 433: , 439: , 442: , 446: , 526: / : 113: , 183: , 188: , 191: , 196: , 232: , 247: , 257: , 359: / : 180: / -: 333: , 336: / -: 401: , 96
V.
457: , 458: , 464: , 470: , 476: , 477: , 479: / -: 531: / -: 542: , 544: , 547: , 560: , 567: , 656: / -: 554: / : 2: , 17: , 29: , 30: , 38: , 40: , 615: Karagiannakidou & Babalonas (1981) Baumann & Baumann (1988), Karagiannakidou & Babalonas (1981), Petermann (1999).
35. Limodorum abortivum (L.) Sw. Chor. type: smed : 48 : 162: , 626: / -: 170: , 210: , 211: , 219: , 230: , 327: / : 364: , 384: , 390: / -: 387: , 389: , 395: , 433: , 434: , 441:
, 451: , 452: , 518: , 522: , 523: / : 285: , 351: , 354: / : 12: , 105: , 108: , 112: , 179: / -: 335: , 397: , 636: / -: 477: , 478: , 479: / -: 533: , 654: / -: 549: , 556: , 557: , 558: , 657: / -: 546: , 548: , 550: , 551: , 552: Baumann & Baumann (1988), (1976) Karagiannakidou & Babalonas (1981) .
36. Neotinea tridentata (Scop.) R.M. Bateman, Pridgeon & M.W. Chase (Syn.: Orchis tridentata Scop.) Chor. type: smed : 113 : 53: , 55: , 57: , 59: , 75: , 76: , 122: , 123: , 124: , 125: , 133:
, 135: , 140: , 141: , 144: , 145: , 149: , 151: , 153: , 154: , 157: , 158: , 159: , 161: , 198: , 201: , 202: / -: 163: , 165: , 175: ,
97
V.
204: , 208: , 215: , 217: , 220: , 225: , 230: , 631: / -: 236: , 240: , 244: , 249: , 250: , 252: , 253: , 255: , 289: , 290: , 293: , 294: , 295: , 296: , 300: , 303: , 306: , 312: , 314: , 317: , 319: , 364: , 365: , 369: , 371: , 376: , 378: , 382: , 384: / -: 426: , 428: , 431: , 435: , 436: , 439: , 443: / : 265: , 267: , 268: , 275: , 361: / : 11: , 12: , 15: , 46: , 51: , 89: , 92: , 93: , 94: , 95: , 96: , 97: , 101: , 105: , 110: , 111: / -: 399: , 457: , 463: , 473: , 504: / : 3: , 5: , 6: , 19: , 21: , 26: , 28: , 63: , 66: , 68: , 72: , 617: , 618: Baumann & Baumann (1988) Hlzinger & Knkele (1988), Karagiannakidou et al. (1995), Quzel & Contandriopoulos (1968), Baumann & Baumann (1988), Quzel (1989), Kitanov (1943), Willing & Willing (1991) Schreiber (1998).
37. Neotinea ustulata (L.) R.M. Bateman, Pridgeon & M.W. Chase (Syn.: Orchis ustulata L.) Chor. type: smed(-eurassubozean) : 35 : 75: , 76: , 122: , 135: , 141: , 153: / -: 208: , 217: / -:
236: , 243: , 253: , 294: , 295: , 298: , 303: , 305: , 306: , 314: , 643: / : 263: , 264: / : 46: , 51: , 96: / -: 411: / -: 470: , 490: , 504: / -: 529: , 595: / -: 562: , 568: / -: 611: / : 65: , 68: Baumann & Baumann (1988), Voliotis & Karagiannakidou (1984), Karagiannakidou & Kokkini (1987), Kitanov (1943), Voliotis & Karagiannakidou (1984), Baumann & Baumann (1988), Willing & Willing (1991) Schreiber (1998), Papanicolaou (1985) Karagiannakidou et al. (1995),
98
V.
Karagiannakidou et al. (1995) Voliotis & Karagiannakidou (1984), (1992) () Eleftheriadou & Raus (1996) .
38. Neottia cordata (L.) Rich. (Syn.: Listera cordata (L.) R. Brown) Chor. type: nosubozean-pralp, circ : 1 -: 651:
(1992), Raus (1995), Eleftheriadou et al. (2001) Willing ( ) (Willing: . .). Raus (1995), Greuter & Raus (1986) Eleftheriadou & Raus (1996).
39. Neottia nidus-avis (L.) Rich. Chor. type: euras(-subozean)-smed : 138 : 120: , 127: , 132: , 134: , 136: , 138: , 139: , 142: , 143: , 146:
, 147: , 152: , 155: , 623: / -: 171: , 172: , 212: , 214: , 216: , 218: , 221: , 226: , 228: , 229: , 242: , 628: / -: 239: , 241: , 251: , 297: , 307: , 308: , 309: , 316: , 324: , 329: , 375: , 377: , 379: , 391: , 393: , 396: , 639: / -: 437: , 444: , 451: , 455: , 515: / : 235: , 246: , 259: , 260: , 266: , 269: , 282: , 286: , 339: , 343: , 344: , 352: / : 13: , 100: , 106: , 107: , 112: , 177: /-: 410: , 415: , 586: / -: 421: , 424: , 474: , 475: , 478: ,
480: , 483: , 484: , 487: , 488: , 491: , 494: , 497: , 498: , 500: , 501: , 503: , 508: , 527: , 528: / -: 530: , 532: , 533: , 535: , 572: , 573: , 575: , 581: , 582: , 591: , 597: / -: 561: , 563: , 564: , 565: , 570: , 583: , 584: / -: 571: , 603: , 604: , 607: ,
99
V.
608: / -: 553: , 598: , 599: , 600: / : 20: , 22: , 24: , 25: , 34: , 35: , 37: , 39: , 42: , 60: , 64: , 73: , 77: , 80: , 81: , 82: , 83: , 84: , 87: , 167: , 617: , 627: Baumann & Baumann (1988), Hlzinger & Knkele (1988) Arampatzis et al. (2001), Baumann & Baumann (1988) Schreiber (1998), Schreiber (1998) Rechinger (1939) Papanicolaou (1985), .. (2004), (1992), (2001) Zoller et al. (1977), Eleftheriadou & Raus (1996) (2001), (2001). , (1985), Baumann & Baumann (1988), Karagiannakidou & Babalonas (1981) (1976).
40. Neottia ovata (L.) Bluff & Fingerh. (Syn.: Listera ovata (L.) R. Brown) Chor. type: euras(-subozean)-smed : 30 : 121: / -: 210: , 223: , 225: , 226: , 228: / -: 239: , 329:
, 379: , 391: , 393: / -: 437: , 441: , 515: / : 48: / -: 409: , 413: , 637: / -: 425: , 459: , 464: , 471: , 481: , 482: , 490: / -: 539: , 593: , 596: / -: 562: / -: 602: Baumann & Baumann (1988), & (1989), . (1992), .. (2004) Theodoropoulos et al. (2003), (1992) Karagiannakidou & Babalonas (1981) Voliotis & Karagiannakidou (1984).
100
V.
41. Ophrys apifera Huds. Chor. type: med-smed(-subatl) : 9 -: 363: , 367: , 368: , 371: / -: 523: / : 183: / -: 545:
, 656: / : 18: .. (1999) , Papanicolaou (1985) Hlzinger et al. (1985) , Hlzinger et al. (1985) Baumann & Baumann (1988).
42. Ophrys cornuta Steven Chor. type: omed-osmed : 43 : 54: , 59: , 128: , 144: , 157: , 161: , 202: , 203: / -: 165: ,
215: , 225: / -: 237: , 249: , 252: , 289: , 290: , 296: , 302: , 365: , 368: , 371: , 372: , 380: , 382: , 383: / -: 429: , 440: , 653: / : 114: , 193: , 196: , 247: , 257: , 359: / : 44: , 45: , 109: / -: 400: , 464: / -: 547: / -: 554: / : 3: , 6: .. (1999), , , , Hlzinger et al. (1985), Petermann (1999).
43. Ophrys grammica (B. Willing & E. Willing) Devillers-Terschuren & Devillers Chor. type: balc : 16
101
V.
: 56: , 144: , 150: / -: 164: , 175: , 220: / -: 302: , 319:
, 370: , 383: / -: 431: , 440: / : 194: / : 46: / -: 465: , 467: Arampatzis et al. (2001).
44. Ophrys insectifera L. Chor. type: subatl-smed
(Zaganiaris 1940).
45. Ophrys mammosa Desf. Chor. type: omed : 63 : 55: , 57: , 59: , 69: , 115: , 133: , 161: , 197: , 200: , 202: ,
203: / -: 173: , 176: / -: 237: , 248: , 363: , 364: , 365: , 368: , 369: , 370: , 371: , 383: / -: 431: , 435: , 436: , 440: , 442: , 511: , 522: / : 113: , 183: , 184: , 186: , 188: , 189: , 191: , 195: , 232: , 247: , 257: , 258: , 350: , 355: , 356: , 360: , 361: , 362: / : 99: / -: 542: / : 1: , 2: , 5: , 16: , 17: , 18: , 28: , 30: , 31: , 38: , 40: , 62: , 63: , , Hlzinger et al. (1985), Baumann & Baumann (1988).
46. Ophrys reinholdii H. Fleischm. Chor. type: omed-osmed : 2 : 350: , 355:
102
V.
47. Ophrys zeusii M. Hirth Chor. type: balc : 2 -: 223: / -: 382:
48. Orchis italica Poir. Chor. type: med : 7 -: 365: , 366: / : 113: , 190: , 284: , 349: , 362:
49. Orchis militaris L. Chor. type: smed-euras : 3 -: 294: , 306: , 317:
Kalopissis (1992), Phitos et al. (1995), Willing & Willing (1991) Hlzinger & Knkele (1988), Kalopissis (1992), Phitos et al. (1995), Willing & Willing (1991) Schreiber (1998). , Phitos et al. (1995).
50. Orchis pallens L. Chor. type: smed-pralp : 7 : 135: , 137: / -: 310: , 313: , 314: / : 106: / -: 490:
Baumann & Baumann (1988) Schreiber (1998) (1992) (). 103
V.
51. Orchis pauciflora Ten. Chor. type: med : 5 : 55: , 57: / -: 175: , 209: / -: 427:
Baumann & Baumann (1988).
52. Orchis pinetorum Boiss. & Kotschy Chor. type: med-smed : 73 : 54: , 58: , 67: , 75: , 135: , 137: , 140: , 145: , 153: , 156: , 160:
, 199: / -: 206: , 208: , 209: / -: 236: , 243: , 250: , 253: , 291: , 293: , 294: , 295: , 297: , 300: , 303: , 306: , 308: , 310: , 312: , 313: , 314: , 317: , 319: , 396: / -: 436: , 441: , 514: , 516: / : 233: , 262: , 270: , 280: , 348: , 633: / : 12: , 15: , 50: , 51: , 88: , 89: , 90: , 91: , 92: , 93: , 94: , 95: , 96: , 101: , 104: / -: 475: , 490: / -: 534: , 535: , 539: / -: 524: , 525: , 559: / : 7: , 36: , 61: , 66: , 68: Baumann & Baumann (1988), Kitanov (1943), Baumann & Baumann (1988), Willing & Willing (1991) Schreiber (1998), (1992) .
53. Orchis provincialis Balbis ex DC. Chor. type: med : 3 : 340: , 345: , 346:
104
V.
54. Orchis purpurea Huds. Chor. type: smed : 41 : 140: , 145: , 154: , 159: / -: 206: , 230: , 328: / -: 238:
, 255: , 291: , 292: , 301: , 383: , 384: / -: 432: , 439: , 456: , 520: , 521: , 644: / : 114: , 181: , 357: / : 14: , 89: , 101: , 103: , 105: , 110: , 111: / -: 334: / -: 398: , 458: , 464: , 474: , 490: / -: 535: / -: 559: , 567: / -: 552: / : 64: Baumann & Baumann (1988) Arampatzis et al. (2001), Voliotis & Karagiannakidou (1984), Kitanov (1943), Baumann & Baumann (1988) Voliotis & Karagiannakidou (1984) - (1992).
55. Orchis quadripunctata Cyr. ex Ten. Chor. type: med : 25 : 52: , 55: , 118: , 131: , 140: , 145: , 150: , 151: , 153: , 157: ,
158: , 624: / -: 175: , 220: / -: 236: , 240: , 252: , 253: , 289: , 296: , 300: / : 194: , 274: , 275: , 288: Baumann & Baumann (1988) Arampatzis et al. (2001) , Baumann & Baumann (1988) Strid & Tan (1998) .
56. Orchis simia Lam. Chor. type: (med-orient)smed(-subatl) : 24
105
V.
: 55: , 69: , 117: , 145: , 150: , 158: / -: 211: / -: 244: ,
299: , 302: / -: 427: , 430: / : 113: , 183: , 185: , 270: , 274: , 275: , 283: , 288: / -: 467: / : 6: , 63: , 66: Baumann & Baumann (1988) Arampatzis et al. (2001) - - Kitanov (1943), Voliotis & Karagiannakidou (1984), Baumann & Baumann (1988) Schreiber (1998).
57. Platanthera bifolia (L.) Rich. Chor. type: no-eurassubozean : 4 : 146: / -: 460: , 464: , 466:
Karagiannakidou & Babalonas (1981) , Quzel & Contandriopoulos (1968) .
58. Platanthera chlorantha (Custer) Rchb. Chor. type: eurassubozean(-smed) : 103 : 52: , 56: , 116: , 121: , 123: , 127: , 139: , 162: , 199: / : 170: , 173: , 175: , 213: , 225: , 228: / -: 239: , 244: , 255:
, 291: , 297: , 299: , 301: , 308: , 314: , 319: , 329: , 331: , 379: , 380: , 386: , 388: , 393: , 394: , 635: / -: 434: , 435: , 437: , 441: , 447: , 448: , 450: , 453: , 454: , 456: , 513: , 642: , 644: / : 341: , 348: , 350: , 355: / : 13: , 100: , 103: , 106: , 107: , 108: , 111: / -: 397: , 406: , 407: , 408: , 409: , 410: , 411: , 413: , 415: , 418: , 636: , 637: / -: 398: , 421: , 464: , 465: , 466: , 467: , 471: , 475: , 478: , 479: , 482: , 490: , 499: , 646: /
106
V.
-: 533: , 595: / -: 525: , 538: , 549: , 556: , 558: , 559: ,
562: , 565: / -: 546: , 552: , 555: , 598: , 601: / : 27: , 71: , 72: , 617: Baumann & Baumann (1988), Hlzinger & Knkele (1988), Karagiannakidou & Babalonas (1981), Karagiannakidou & Babalonas (1981), Papanicolaou (1985) Rechinger (1939), Karagiannakidou et al. (1995) & (2003), .. (2004) Theodoropoulos et al. (2003) (), (1992) (2001) () (2001) ().
59. Serapias bergonii E. G. Camus (pro hybr.) Chor. type: omed : 1 -: 384:
60. Serapias cordigera L. Chor. type: med-smed(-subatl) : 2 -: 384: / -: 389:
61. Serapias vomeracea (N.L. Burm.) Briq. Chor. type: (z)med : 3 : 338: , 360: / -: 550:
107
V.
62. Spiranthes spiralis (L.) Chevall. Chor. type: smed-subatl : 41 : 119: / -: 230: / -: 364: , 366: , 367: , 369: , 371: , 372:
, 376: , 382: , 383: , 384: / -: 387: , 395: , 438: , 439: , 442: , 447: , 449: , 450: , 511: , 512: , 519: , 521: / : 188: , 337: , 338: , 342: , 345: , 353: , 361: / -: 332: , 335: / -: 659: / : 540: , 541: , 550: / : 16: , 40: , 166: , 168:
V.1.2
1. Anacamptis coriophora (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. fragrans (Pollini) R.M. Bateman A. laxiflora (Lam.) R.M. Bateman, Pridgeon & M.W. Chase : 1 -: 450:
2. Anacamptis papilionacea (L.) R.M. Bateman, Pridgeon & M.W. Chase A. picta(Loisel.) R.M. Bateman
: 2 : 179: / : 169:
3. Anacamptis picta (Loisel.) R.M. Bateman A. laxiflora (Lam.) R.M. Bateman,Pridgeon & M.W. Chase
: 1 -: 461:
108
V.
4. Dactylorhiza cordigera (Fries) So D. saccifera (Brongn.) So : 1 -: 507:
5. Dactylorhiza graeca H. Baumann D. cordigera (Fries) So (Dactylorhiza serreana H. Baumann) : 1 : 41:
Baumann (1983) Hlzinger & Knkele (1983).
6. Dactylorhiza macedonica Hlzinger & Knkele D. saccifera (Brongn.) So : 1 : 148:
Hlzinger & Knkele (1988) .
7. Gymnadenia rhellicani (Teppner & Klein) Teppner & Klein G. conopsea (L.) R. Br.
Gembardt & Glz (2002).
8. Neotinea tridentata (Scop.) R.M. Bateman, Pridgeon & M.W. Chase N. ustulata (L.) R.M. Bateman, Pridgeon & M.W. Chase [Neotinea dietrichiana (Bogenh.) H. Kretzschmar, Eccarius & H. Dietr.] : 3 : 135: , 153: / -: 253:
109
V.
.. (1999) Willing & Willing (1991).
9. Ophrys mammosa Desf. Ophrys cornuta Steven (Ophrys aetolica H. Baumann & Knkele) : 1 : 114:
10. Orchis italica Poir. O. simia Lam. (Orchis celtiberica Pau) : 1 : 113:
11. Orchis militaris L. O. purpurea Huds. (Orchis hybrida Boenn. ex Rchb.) : 1 -: 306:
12. Orchis pinetorum Boiss. & Kotschy O. pauciflora Ten. (Orchis dicorifiana G. Thiele & W. Thiele) : 4 -: 209: / -: 253: , 293: , 308:
13. Orchis purpurea Huds. O. simia Lam. (Orchis angusticruris Franch. ex Rouy) : 1 : 182:
110
V.
V.2 75 taxa 57 5 - 17 - 13 (Tsiftsis et al. 2007). Orchis (9 taxa) Anacamptis, Dactylorhiza Epipactis (8 taxa ), Ophrys (7 taxa), Cephalanthera, Gymnadenia, Neottia Serapias taxa, Neotinea Platanthera taxa Corallorhiza, Epipogium, Goodyera, Himantoglossum, Limodorum Spiranthes taxon (. 9).
Anacamptis
10 9 8 7 6 5 4 3 2 1 0 1 1 1 1 1 3 3 2 3 8 8 8 7
9
Cephalanthera Corallorhiza Dactylorhiza Epipactis Epipogium Goodyera Gymnadenia Himantoglossum Limodorum
3 2 1
Neotinea Neottia Ophrys Orchis Platanthera Serapias Spiranthes
9. . Figure 9. Number of species and subspecies of the genera of the study area.
15 (Anacamptis coriophora subsp. coriophora, A. laxiflora, A. palustris subsp. elegans, A. papilionacea, Dactylorhiza romana, Epipactis atrorubens subsp. spiridonovii, E. pontica, O. reinholdii, Ophrys zeusii, Orchis italica, O. provincialis, Serapias bergonii, S. cordigera, S. vomeracea Spiranthes spiralis) , (Epipactis atrorubens subsp. spiridonovii Epipactis pontica) . , 46 (Anacamptis coriophora subsp. fragrans, A. morio, A. picta, A. pyramidalis, Cephalanthera
111
V.
damasonium, C. longifolia, C. rubra, Corallorhiza trifida, Dactylorhiza cordigera, D. graeca, D. incarnata, D. macedonica, D. saccifera, D. sambucina, D. viridis, Epipactis atrorubens subsp. atrorubens, E. exilis, E. helleborine, E. microphylla, E. nauosaensis, E. palustris, Epipogium aphyllum, Goodyera repens, Gymnadenia conopsea, G. frivaldii, G. rhellicani, Himantoglossum caprinum, Limodorum abortivum, Neotinea tridentata, N. ustulata, Neottia cordata, N. nidus-avis, N. ovata, Ophrys apifera, O. cornuta, O. grammica, O. mammosa, Orchis militaris, O. pallens, O. pauciflora, O. pinetorum, O. purpurea, O. quadripunctata, O. simia, Platanthera bifolia P. chlorantha) . , . V.1.1 12, . (49 ) , (45), (41), (37), (30) (27). (46), 32 29 . 12. . I: , II: , III: , IV: . Table 12. Total number of species and subspecies in each area and subarea. I: Number
of species and subspecies found only by us, II: number of species and subspecies found by us and other authors, III: species and subspecies not found by us, IV: total number of species and subspecies. - - - I II III IV 9 29 3 41 21 10 1 32 13 31 2 46 26 3 0 29 14 34 1 49 21 14 2 37 - - - - - - 18 8 1 27 15 10 0 25 11 25 1 37 9 11 1 21 16 9 0 25 4 4 1 9 11 5 2 18 18 26 1 45 16 9 5 30
12, 49 , 14 (Anacamptis laxiflora, A. palustris subsp. 112
V.
elegans, A. papilionacea, A. picta, Dactylorhiza incarnata, D. romana, Epipactis exilis, O. grammica, O. mammosa, Ophrys zeusii, Orchis italica, Serapias bergonii, S. cordigera Spiranthes spiralis) , 34 (Anacamptis coriophora subsp. fragrans, A. morio, A. pyramidalis, Cephalanthera damasonium, C. longifolia, C. rubra, Corallorhiza trifida, Dactylorhiza saccifera, D. sambucina, D. viridis, Epipactis atrorubens subsp. atrorubens, E. helleborine, E. microphylla, E. nauosaensis, E. palustris, Goodyera repens, Gymnadenia conopsea, G. rhellicani, Himantoglossum caprinum, Limodorum abortivum, Neotinea tridentata, N. ustulata, Neottia nidus-avis, N. ovata, Ophrys apifera, O. cornuta, Orchis militaris, O. pallens, O. pauciflora, O. pinetorum, O. purpurea, O. quadripunctata, O. simia Platanthera chlorantha) . 18 (Anacamptis coriophora subsp. coriophora, A. coriophora subsp. fragrans, A. laxiflora, A. papilionacea, A. picta, Dactylorhiza incarnata, D. macedonica, Epipactis pontica, Gymnadenia frivaldii, Limodorum abortivum, Neotinea tridentata, Ophrys apifera, O. grammica, O. mammosa, O. simia, Platanthera bifolia, Serapias vomeracea Spiranthes spiralis
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