111
Key words: apple culture, chestnut 
culture, feral pigs, Flora Hellenica 
Database, Orchidaceae, overgrazing, 
photovoltaic farms, road verges.
Ključne besede: nasadi jablan, 
nasadi kostanja, divji prašiči, Flora 
Hellenica Database, Orchidaceae, 
prepašenost, fotovoltaične 
elektrarne, cestne brežine.
Corresponding author:
Argyrios Gerakis
E-mail:   
aris.gerakis@ionio.gr
Received: 17. 6. 2024
Accepted: 13. 10. 2024
An Update to the Orchid Flora of Mounts 
Pelion and Mavrovouni, Greece, with 
Notes on Conservation
Abstract
A floristic survey was implemented during 2021–2022 largely within Natura sites 
GR1430008 and GR1420004 in the Region of Thessaly, altogether comprising 
796 km2. While both Natura sites were explored, the sampling effort was 
intensified within a narrow study area of 550 km2 encompassing the N part of 
Mt Pelion and the whole of Mt Mavrovouni. Sampling within the narrow study 
area was one-third guided by previous findings and two-thirds random. In total, 
129 square cells 1 km2 wide were explored, of which 100 yielded orchid findings; 
39 orchid taxa (species and subspecies) plus two nothotaxa were recorded. In 
the narrow study area, 33 orchid taxa and nothotaxa were recorded, of which 
13 for the first time: Anacamptis papilionacea, A. pyramidalis, Himantoglossum 
robertianum, Ophrys × delphinensis, O. sicula, O. speculum, O. sphegodes subsp. 
sphegodes, O. tenthredinifera, O. tenthredinifera × O. sphegodes subsp. taurica (syn. 
O. mammosa), O. umbilicata, Orchis italica, O. simia, and Platanthera bifolia. The 
main threats facing orchids are rooting by feral pigs, overgrazing by ruminants, 
clearing of road verges, intensive arboriculture, and the encroachment of 
photovoltaic farms.
Izvleček
V letih 2021 in 2022 smo v Tesaliji, na 796 km2 velikem območju, izvedli 
floristično raziskavo predvsem na Natura območjih GR1430008 in GR1420004. 
Obe območji smo pregledali, bolj intenzivno pa smo vzorčili na ožjem območju na 
površini 550 km2, ki obsega severni del gore Pelion in celotno goro Mavrovouni. 
Eno tretjino ožjega območja smo vzorčili na osnovi predhodnih raziskav, dve 
tretjini pa naključno. Raziskali smo 129 kvadrantov v velikosti 1 km2, od tega je 
bila v 100 kvadrantih prisotna vsaj ena orhideja. Skupaj smo našli 39 taksonov 
(vrste in podvrste) in dva nototaksona orhidej, od teh jih je bilo 13 najdenih prvič: 
Anacamptis papilionacea, A. pyramidalis, Himantoglossum robertianum, Ophrys × 
delphinensis, O. sicula, O. speculum, O. sphegodes subsp. sphegodes, O. tenthredinifera, 
O. tenthredinifera × O. sphegodes subsp. taurica (syn. O. mammosa), O. umbilicata, 
Orchis italica, O. simia, in Platanthera bifolia. Glavne grožnje, s katerimi se soočajo 
orhideje, so kopanje divjih prašičev, intenzivna paša prežvekovalcev, čiščenje cestnih 
brežin, intenzivno gozdarstvo in posegi zaradi fotovoltaičnih elektrarn.
Argyrios Gerakis1, Ursina Steinemann2, Sofia Drizi3, Vasilis Ioannidis4, 
Eirini Katsalirou1, Athina Klasogianni1 & Kostas Koutis4 
DOI: 10.3986/hacq-2025-001024/1 • 2025, 111–135
1 Department of Food Science and Technology, Ionian University, Argostoli, Greece. 
2 Retired Biologist, Switzerland.
3 Department of Biology, Aristotle University of Thessaloniki, Thessaloniki, Greece.
4 AEGILOPS – Greek Network for Biodiversity and Ecology in Agriculture, Agria Volos, Greece.
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Introduction
The impetus for the study was the initiative by the Man-
agement Unit of Protected Areas of Thessaly (formerly 
Management Body of Karla-Mavrovouni-Kefalovryso 
Velestinou-Delta Piniou) to initiate management actions 
related to protected sites, species, and ecotopes within its 
jurisdiction. This action was focused on two Natura 2000 
sites within the jurisdiction of the Management Unit, 
with considerable diversity in native orchids (family Or-
chidaceae). The two Natura sites are GR1430008 (Oros 
Pilio) and GR1420004 (Karla-Mavrovouni-Kefalovryso 
Velestinou-Neochori), altogether comprising 796 km2. 
The two sites roughly correspond to the geographical ex-
tent of Mounts Pelion and Mavrovouni, respectively. The 
objective reported here was to record the distribution of 
native orchids in the study area with emphasis on five taxa 
of conservation interest, namely Ophrys cornutula (syn. 
O. oestrifera subsp. cornuta, O. scolopax subsp. cornuta), 
Spiranthes spiralis, Ophrys umbilicata subsp. umbilicata, 
Orchis provincialis, and O. simia.
Methods
Initially, we collected data about the terrestrial habitats 
in the study area. The following broad habitat groupings 
derive from the Standard Data Forms of the two Natura 
2000 sites, with contribution in parenthesis (European 
Environment Agency, 2024a, 2024b):
• Broad-leaved deciduous woodland (34,030 ha)
• Shrubland such as maquis, phrygana, etc. (23,005 ha)
• Arable land with herbaceous and woody crops 
(13,784 ha)
• Evergreen woodland (2,500 ha)
• Grassland (1,922 ha)
• Mixed woodland (1,721 ha)
• Coniferous woodland (539 ha)
For Mount Pelion, Fotiades (2005) reports 14 ecoto pes, 
without reference to their respective contribution:
• Precipitous, rocky cliffs with Mediterranean vegeta-
tion
• Short communities of Euphorbia near precipitous, 
rocky cliffs
• Phrygana of Sarcopoterium spinosum
• Sclerophyllous woodland used for grazing (dehesas) 
of Quercus ilex
• Degraded shrubland of kermes oak (garrigues)
• Beech woodland
• Chestnut woodland
• Quercus frainetto woodland
• Platanus orientalis woodland
• Caves not exploited for tourism
• Quercus ilex woodland
• Azonal vegetation along rivers of permanent or in-
terrupted flow, in the form of gallery dominated by 
Salix sp. and Populetea albae
• Thickets of Juniperus communis on calcareous undis-
turbed land
• Thickets of laurel
Evidently, ecotopes such as caves are unlikely to host 
orchids; the remaining ecotopes can be easily transferred 
to the broad habitat groupings derived from the Natura 
2000 Standard Data Forms.
The broader study area belongs to the Meso-mediter-
ranean bioclimatic zone. We witnessed a sharp ecological 
gradient in the overall direction NE to SW. This is the 
result of Mounts Pelion and Mavrovouni catching most 
of the humidity that derives from the Aegean Sea before 
it reaches the hinterland. This ecological gradient results 
in a rich variety of ecosystems unfolding within a span of 
only a few kilometres (for details, see Raus, 1979a: 569 ff. 
and Figs. 5 & 6). 
The Aegean coast is quite forested, with maquis of holly 
oak (Quercus ilex) and strawberry tree (Arbutus unedo) at 
sea level, gradually being replaced by chestnut (Castanea 
sativa) and Hungarian oak (Quercus frainetto), culminat-
ing in beech (Fagus sylvatica), up to the mountain ridge. 
As we descend from the mountain ridge to the drier 
plains of Thessaly, tall deciduous forest is gradually being 
replaced by pseudomaquis of kermes oak (Q. coccifera), 
that may further deteriorate into phrygana or grassland 
(for details, see Raus, 1979b). 
The geology appears to be heterogeneous. The moun-
tain range is predominantly composed of Paleozoic crys-
talline rocks, i.e., metamorphic rocks, especially gneisses 
and mica schists, partly interspersed with marble. The 
crystalline rocks are mostly found in the mid-elevations 
and in the W of Pelion they extend down to the coastal 
zone. Mesozoic limestones and dolomites overlie the crys-
talline rocks, mainly at higher elevations. Additionally, 
these can also appear on a small scale in the coastal zone, 
possibly due to tectonic disturbances. The soils are typi-
cally brown soils and reddish-brown Mediterranean soils 
with highly variable pH values (Biel & Rudolph, 1992).
Since most of the orchid species of conservation inter-
est are photophilic, we concentrated our sampling effort 
on the SW slope of Mounts Pelion and Mavrovouni and 
excluded the extensive tracts of tall, broad-leaved and 
mixed forest in the NE and central part of the study area. 
Broad-leaved and mixed forest, especially of beech and 
Hungarian oak, is so dense that little light reaches the for-
est floor; during our preliminary reconnaissance in these 
ecosystems, we observed the occasional orchids only in 
clearings and along the roadside.
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We tried to find historical surveys of the family Or-
chidaceae within the study area. As a starting point, we 
consulted the Atlas of Greek Orchids (Antonopoulos & 
Tsiftsis, 2017; Tsiftsis & Antonopoulos, 2017). The dis-
tribution maps are microscopic thumbnails where each 
location is represented by a dot 10 km wide, not useful 
for planning a detailed survey.
The useable reference that we could find with respect to 
the orchids of Pelion and Mavrovouni was by Biel & Ru-
dolph (1992), albeit with dated information of 30 years 
ago. A span of 30 years is long enough for cultivated fields 
to be reforested, or, conversely, for forest to be cleared; 
moreover, urban and tourist development encroach on 
natural ecosystems. Still, the work of Biel & Rudolph 
(1992) offered a very useful advantage, in that locations 
were recorded with precision of 0.1 km. Their survey 
covered the entire Natura site GR1430008 (Oros Pilio), 
while from Natura site GR1420004 (Karla-Mavrovouni-
Kefalovryso Velestinou-Neochori) only the SE portion 
was incorporated (Figure 1).
We tallied the findings of Biel & Rudolph (1992). In 
the two Natura 2000 sites, the authors altogether report 
36 orchid taxa (species and subspecies) and one noth-
otaxon (Table1). In general, we follow the orchid nomen-
clature of POWO (2024). In particular, this includes the 
name change of Ophrys sphegodes subsp. mammosa to O. 
sphegodes subsp. taurica. The first use of the epithet taurica 
at subspecific rank in 1890 antedates the first use of the 
epithet mammosa at subspecific rank in 1926 (Art. 11.4 
Shenzhen Code); for details, see Niketić et al. 2018, ac-
cepted by POWO 2024 based on Güner et al. (2022). As 
an exception from POWO, we use Ophrys sicula instead 
of O. lutea subsp. galilaea based on Hennecke (2016).
In the spring season of 2021, we implemented a pre-
liminary reconnaissance. The objectives were (i) to ex-
plore as many different ecosystems as possible in order to 
get a general picture of orchid diversity in the study area 
and (ii) make well founded decisions on the following 
main phase of the survey.
a. Sampling design during preliminary reconnaissance
We adopted a gradient-directed transect sampling or 
“gradsect” (Austin & Heyligers, 1989). In a mountainous 
region, an obvious gradient is that of elevation, which in 
turn determines the gradients of climatic factors and, by 
extension, vegetation. In our study area there is an eleva-
tion gradient with overall direction NE to SW (Figure 2). 
We selected paved roads that more or less followed the 
elevation gradient, meaning that they traversed the two 
Natura sites in the general direction NE to SW. 
The southern boundary of our more narrowly delim-
ited intensive investigation area had been agreed with the 
supervisory authority to coincide with the imaginary line 
Figure 1: Map of the broader study area. With yellow ochre fill the 
prefecture of Larissa, with light blue fill the prefecture of Magnesia, 
with magenta outline the two Natura 2000 sites, with dark blue the 
outline of the narrow study area, and with orange outline the study 
area of Biel & Rudolph (1992). UTM grid spacing 5 km.
Slika 1: Zemljevid širšega preučevanega območja. Z rumeno je označe-
na prefektura Larissa, s svetlomodro pa prefektura Magnesia, s škrlatno 
barvo obe območji Natura 2000, s temno modro barvo je označeno 
ožje preučevano območje in z oranžno preučevano območje v članku 
Biel & Rudolph (1992). Velikost kvadrantov UTM mreže je 5 km.
Figure 2: Elevation map of the two Natura 2000 sites, outlined in 
magenta. Contour interval 200 m, UTM grid spacing 5 km.
Slika 2: Karta nadmorskih višin obeh območij Natura 2000 označenih 
s škrlatno barvo. Izohipse so v intervalu 200 m, velikost kvadrantov 
UTM mreže je 5 km.
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Gerakis et al.
Orchid Flora of Pelion and Mavrovouni
that follows the brook between the villages of Zagora and 
Pouri, passes through Leschiani bridge and ends up at the 
hamlet of Ano Kerasia. The area thus described includes 
the whole of Mount Mavrovouni and the northern slope 
of Mount Pelion and covers 550 km2 in total (Figure 1).
b. Mapping the study area
To map the study area we opted for the Universal 
Transverse Mercator (UTM) projection system (“Univer-
sal Transverse Mercator coordinate system,” 2024) with 
WGS84 as geodetic reference system. UTM coordinates 
facilitated comparison of results with the earlier survey of 
Biel & Rudolph (1992).
As base maps we used digitised versions of georefer-
enced hiking maps (Anavasi, 2008, 2022), the digitised 
administrative boundaries of Greece (Harvard Library, 
2024), and the digitised outlines of Natura 2000 sites 
(Ministry of Environment and Climate Change, 2024).
c. Sampling design during the main phase
According to Delforge (2006), the five taxa of conser-
vation interest prefer open habitats of thin, short vegeta-
tion, including open woodland, except for Orchis provin-
cialis that exhibits a preference for damper environments, 
a fact that was confirmed by our preliminary scouting. 
In Figure 3 we mark the preliminary findings of Orchis 
provincialis on a precipitation map (Hellenic National 
Meteorological Service, 2024). It seems that O. provin-
cialis favours the wetter foothills of E Mount Pelion at 
elevations of 200–600 m asl.
We estimated that project resources (time and money) 
were sufficient to sample 37 square cells 1 km wide or 
nearly 7% of the narrow study area of 550 km2. Thus, 
the narrow study area was divided into 550 square cells 
1 km2. For the main sampling phase, approximately one-
third of the 37 sampling units were allocated according to 
the following criteria:
(i) Findings of the supervisory authority: We assigned 
one sampling unit to each of three biotopes where 
the staff of the supervisory authority had recorded 
specimens of Ophrys cornutula (syn. O. scolopax 
subsp. cornuta), O. umbilicata subsp. umbilicata, 
and Orchis simia.
(ii) Literature: We assigned one sampling unit to each 
of three biotopes where Biel & Rudolph (1992) 
had recorded specimens of Ophrys cornutula (syn. 
O. scolopax subsp. cornuta). The mentioned authors 
did not record Spiranthes spiralis, Ophrys umbilicata 
subsp. umbilicata, or Orchis simia within the nar-
row study area. 
(iii) Most promising habitats: Assuming that the five 
species of interest are of equal importance, six of 
the 31 remaining sampling units were allocated to 
Orchis provincialis in the wetter zone of E Mount 
Pelion at elevations of 200–600 m asl. 
The remaining 25 sampling units (approximately two-
thirds) were randomly assigned to the most promising 
habitats of the other four species of conservation interest. 
These are habitats of thin, short vegetation. The pros and 
cons of random sampling designs for terrestrial surveys 
are analysed by Webster & Oliver (1990).
To identify habitats of thin, short vegetation, we used 
the georeferenced, ortho-corrected maps of Corine Land 
Cover (Hellenic Mapping and Cadastral Organization, 
2015). The CLC classes that correspond to shrub and/or 
herbaceous vegetation associations are:
3.2.1. Natural grassland
3.2.2. Moors and heathland
3.2.3. Sclerophyllous vegetation
3.2.4. Transitional woodland/shrub 
To the above we added “2.2.3 Olive groves,” based on 
our experience that low input olive groves are potentially 
rich orchid habitats, as long as they are not plowed or 
sprayed with herbicides (Katsalirou et al., 2016b), though 
olive groves covered very little of the study area. 
Subsequently, we assigned a unique numerical ID to 
each of 172 square cells within the narrow study area that 
were covered by at least 50% by one or more of the afore-
mentioned CLC classes. Of the 172 cells, 25 were ran-
domly selected by a random number generator (Numbers 
v. 10.3.9, Apple Inc.). Figure 4 depicts the final sampling 
design implemented during the main phase. 
Figure 3: Precipitation map of the two Natura 2000 sites, outlined in 
magenta. Red dots represent findings of Orchis provincialis during the 
preliminary scouting in the spring of 2021. UTM grid spacing 5 km.
Slika 3: Karta padavin dveh območij Natura 2000 označenih s škrlatno 
barvo. Rdeče točke predstavljajo nahajališča vrste Orchis provincialis 
med preliminarnim pregledom spomladi leta 2021. Velikost kvadran-
tov UTM mreže je 5 km.
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d. Sampling schedule
The preliminary reconnaissance phase lasted from 
25.04.2021 to 04.08.2021, with diminished findings to-
ward the end of this period. 
Spiranthes spiralis flowers from August to October 
throughout its broad distribution range, from the At-
lantic coasts of Europe to Iran and from S Sweden to N 
Africa (Delforge, 2006). In the Mediterranean, we would 
expect the onset of flowering to coincide with the onset 
of the wet season, which varies from year to year. In our 
case, we found that the optimum time window was from 
20.10.2021 to 13.11.2021. 
The main sampling phase for the spring-flowering spe-
cies lasted from 16.02.2022 to 04.07.2022, with dimin-
ished findings toward the end of this period. A few sup-
plementary observations were taken on 19.04.2023 and 
20.04.2023.
e. Field procedure
At each location we photographed the specimens and 
filled forms with data such as names of investigators, date, 
nearest settlement, place name, plant community, names 
of taxa, number of specimens, development stage, coor-
dinates, altitude, parent rock, soil type, nearby plant spe-
cies, and main anthropogenic threats. 
Results
In total, 129 square cells 1 km2 wide were explored, mostly 
within the two Natura 2000 sites, of which 100 yielded 
orchid findings (Figure 5). The tally of sampled cells far ex-
ceeds the target of 37 that was decided during the design of 
the main sampling phase. This is because the tally includes:
(i) cells sampled during the preliminary reconnais-
sance phase, 25.04.2021 to 04.08.2021
(ii) circumstantial samplings, i.e., promising biotopes 
explored on the way to and from the 37 designated 
sampling units
(iii) biotopes sampled during the off-hours of the field 
task force
The descriptions of the sites are in Appendix I. The to-
tal number of sites listed are 107, as some sampling units 
contained more than one distinct biotope that have been 
described separately.
In total, we identified 39 orchid taxa (species and sub-
species) plus two nothotaxa (Table 1, nomenclature ac-
cording to POWO, 2024).
Figure 4: Map of the sampling design during the main sampling 
phase, i.e., fall of 2021 (for Spiranthes spiralis) to spring of 2022. Olive 
green fill: Corine Land Cover classes that correspond to open habitats, 
i.e., a combination of scrub and/or herbaceous vegetation. Blue dots: 
Locations suggested by the supervisory authority. Magenta dots: Loca-
tions with Ophrys scolopax subsp. cornuta by Biel & Rudolph (1992). 
Red dots: Locations likely to host Orchis provincialis based on the 
preliminary scouting. Green dots: Sampling units selected at random. 
UTM grid spacing 1 km.
Slika 4: Zemljevid prikazuje način vzorčenja med glavno fazo, to je 
od jeseni 2021 (za vrsto Spiranthes spiralis) do spomladi 2022. Olivno 
zelena: razredi Corine Land Cover, ki predstavljajo odprte habitate, 
to je kombinacijo grmiščne in/ali zeliščne vegetacije. Modre točke: 
lokacije, ki so jih predlagale nadzorne oblasti. Škrlatne točke: lokacije 
z Ophrys scolopax subsp. cornuta iz publikacije Biel & Rudolph (1992). 
Rdeče točke: lokacije, na osnovi predhodnega pregleda terena, kjer se 
verjetno pojavlja Orchis provincialis. Zelene točke: naključne vzorčne 
točke. Velikost kvadrantov UTM mreže je 1 km.
Figure 5: Map with sampling units 1 km wide that yielded orchid 
findings within the broader study area during 2021–2022. With 
magenta outline the two Natura 2000 sites, with dark blue the outline 
of the narrow study area. UTM grid spacing 1 km.
Slika 5: Zemljevid kvadrantov velikosti 1 km, kjer smo v letih 2012–
2022 našli orhideje v večjem preučevanem območju. S škrlatno barvo 
sta označeni območji Natura 2000, s temno modro pa ožje preučevano 
območje. Velikost kvadrantov UTM mreže je 1 km.
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Orchid Flora of Pelion and Mavrovouni
Table 1: List of taxa (species and subspecies) recorded during the sampling period 2021–22 within the broader study area. In bold 
are those recorded within the narrow sampling area. Site numbers refer to Appendix I. Checkmark indicates mention of the taxon 
in the Flora Hellenica Database within the narrow sampling area.
Tabela 1: Seznam taksonov (vrst in podvrst) vzorčenih v obdobju od 2021 do 2022 v preučevanem območju. Številke nahajališč se 
nanašajo na Prilogo I. Kljukica označuje oznako taksona v bazi Flora Hellenica znotraj ožjega preučevanega območja.
S/N TAXON SITES FLORA HELLENICA
1 Anacamptis coriophora 99 ✓
2 Anacamptis morio subsp. caucasica 21, 24, 35, 43, 46, 48, 49, 52, 53, 55, 59, 80, 
99, 103, 104, 105, 106
✓ (sub A. morio)
3 Anacamptis papilionacea subsp. papilionacea 16, 21
4 Anacamptis pyramidalis 27, 79
5 Cephalanthera longifolia 29, 32, 33, 37, 46, 68, 71, 74, 76, 77, 82, 84, 
85, 88, 89, 90, 96, 100, 104
✓
6 Cephalanthera rubra 32, 73, 77 ✓
7 Dactylorhiza romana 10, 21, 45, 53, 54, 83, 87, 89, 90, 106 ✓
8 Dactylorhiza saccifera 67, 69, 70, 72, 73, 75, 76, 82, 89, 90 ✓
9 Epipactis helleborine 64, 68 ✓
10 Epipactis microphylla 23, 32, 36, 73, 82, 89, 91 ✓
11 Himantoglossum calcaratum subsp. rumelicum 62, 66 ✓ (sub. H. jankae)
12 Himantoglossum robertianum 3, 4, 28, 46
13 Limodorum abortivum 3, 5, 8, 11, 14, 43, 66, 77, 84, 90, 94, 100 ✓
14 Neotinea maculata 27, 32, 60, 76, 77, 89, 90, 91, 92, 94 ✓
15 Neotinea tridentata 24, 40 ✓
16 Neottia nidus-avis 73, 77 ✓
17 Ophrys × delphinensis 12
18 Ophrys apifera 99
19 Ophrys ferrum-equinum subsp. ferrum-equinum 33
20 Ophrys reinholdii subsp. reinholdii 13, 43, 44, 53 ✓ (sub O. reinholdii)
21 Ophrys scolopax subsp. cornuta 13, 18, 20, 25, 41, 43, 58, 65, 66, 77, 78, 81, 
99, 103
✓
22 Ophrys sicula (syn. O. lutea subsp. galilaea) 12, 18, 26, 29, 33, 37
23 Ophrys speculum subsp. speculum 6, 7
24 Ophrys sphegodes subsp. sphegodes (syn. O. hebes) 43
25 Ophrys sphegodes subsp. spruneri 26 ✓
26 Ophrys sphegodes subsp. taurica 1, 2, 3, 5, 6, 8, 12, 18, 22, 25, 26, 43, 46, 60, 
66
✓  (sub O. sphegodes 
subsp. mammosa)
27 Ophrys tenthredinifera 1, 2, 3, 6
28 Ophrys tenthredinifera × O. sphegodes subsp. taurica 3
29 Ophrys umbilicata subsp. umbilicata 1, 2, 6, 25, 26
30 Orchis anthropophora 24, 28, 29, 33, 37, 39, 41, 42, 45, 47, 60, 63 ✓
31 Orchis italica 3, 12, 17, 18, 21, 33, 37, 38, 40, 42, 45, 46, 47, 
50, 51, 52, 53, 56, 57, 66, 79, 80, 81, 86, 101
32 Orchis pauciflora 24, 28, 42, 63 ✓
33 Orchis provincialis 9, 35, 60, 77, 83, 89, 90, 91, 92, 93, 95 ✓
34 Orchis quadripunctata 18, 28, 29, 30, 31, 34, 41, 42, 56, 57, 61, 63 ✓
35 Orchis simia 24, 29, 34, 39, 42
36 Platanthera bifolia 76
37 Platanthera chlorantha 32, 77 ✓  (sub P. chlorantha 
subsp. chlorantha)
38 Serapias bergonii 103, 104 ✓  (sub S. vomeracea 
subsp. laxiflora)
39 Serapias lingua 104, 105, 106, 107 ✓
40 Serapias vomeracea 2, 12, 48, 102 ✓  (sub S. vomeracea 
subsp. vomeracea)
41 Spiranthes spiralis 3, 5, 9, 10, 15, 18, 19, 31, 35, 66, 97, 98, 99 ✓
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Discussion
During the final stage of the manuscript, we received no-
tice of the new edition of the Atlas of the Hellenic Flora 
(Strid, 2024). As with the Atlas of Greek Orchids (An-
tonopoulos & Tsiftsis, 2017; Tsiftsis & Antonopoulos, 
2017), the scale of the maps is too coarse to pinpoint the 
exact location of the findings. Fortunately, the author was 
so kind to share the records from the Flora Hellenica Da-
tabase on which the maps of the Atlas are based (Prof. 
Arne Strid, personal communication 14 May 2024). 
Before comparing our findings with the Flora Hellenica 
Database, we processed its records as follows:
(i) We omitted records with only approximate loca-
tion. For example, records in the Database originat-
ing from the Atlas of Greek Orchids were read off 
dot maps of 10 km precision, not useable for our 
purposes. 
(ii) We improved and supplemented the records origi-
nating from the survey of Biel & Rudolph (1992). 
The Flora Hellenica Database listed only approxi-
mate locations read off dot maps. Since we had 
access to the original hard copy, we replaced the 
approximate locations with locations of 0.1 km 
precision and added several missing records. 
The useable records from the Flora Hellenica Database 
that fall within our narrow study area, besides those con-
tributed by Biel & Rudolph (1992), are mostly older 
observations contributed by Thomas Raus (Raus 1979b, 
1980) and Eckhard Willing (Willing & Willing 2003, 
2009). The comparison of our findings to the Flora Helle-
nica Database clearly demonstrates that there are extensive 
tracts of land that our task force explored for the first time 
(Figure 6). Further, the Flora Hellenica Database reports 
only 20 of the 33 taxa that we recorded within the narrow 
study area (Table 1, names in bold). One explanation is 
the strong random component of our sampling design, 
which included tracts of the study area that are normally 
inaccessible.
It is interesting that Orchis simia was missed by pre-
vious surveys although it occurs quite abundantly in its 
respective habitats. We hypothesise that decades of defor-
estation and intensive grazing have thinned out formerly 
forested biotopes, thus attracting photophilic species for 
the first time.
On the other hand, the Flora Hellenica Database reports 
nine taxa within our narrow study area that we did not 
find: Anacamptis coriophora, Epipactis helleborine, Gym-
nadenia conopsea, Himantoglossum calcaratum subsp. ru-
melicum (syn. H. jankae), Neotinea ustulata, Neottia ovata, 
Ophrys sphegodes subsp. spruneri, Serapias bergonii (sub S. 
vomeracea subsp. laxiflora), and S. lingua. Every taxon 
that we missed is reported from only one location each, 
indicating that their presence within the narrow study 
area is rather uncommon. As far as Epipactis helleborine 
and Neottia ovata are concerned, our sampling scheme 
heavily favoured open ecosystems of relatively low veg-
etation, whereas E. helleborine and N. ovata prefer dense 
forests made up by Quercus ilex, Castanea sativa and  Fagus 
sylvatica (Raus, 1979b, 1980); in fact, we were able to 
confirm sizeable populations of E. helleborine further S, 
in chestnut and beech forests.
Proposed conservation 
measures
To improve the conservation status of the family Orchi-
daceae, we propose a series of conservation measures.
a. Containment of wild pigs, domesticated free-range 
pigs, and their hybrids
Herds of wild pigs and their hybrids with domesticated 
pigs roam freely on Mounts Pelion and Mavrovouni (Fig-
ure 7). Feral pigs represent a major threat to biodiversity 
as they dig up extensive tracts of land, uprooting the her-
baceous vegetation (Figure 8). We frequently witnessed 
orchid tubers unearthed by pigs (Figure 9). It is not 
known whether pigs forage on orchid tubers or whether 
their destruction is collateral damage during the search 
for some other food.
Figure 6: Map with square cells 1 km wide that yielded orchid findings 
within the narrow study area during 2021–2022. Green dots represent 
our findings, orange triangles represent records from the Flora Hel-
lenica Database, mostly predating 1992. UTM grid spacing 1 km.
Slika 6: Zemljevid kvadrantov velikosti 1 km, kjer smo v letih 
2012–2022 našli orhideje v večjem preučevanem območju. Zelene 
točke predstavljajo naše najdbe, oranžni trikotniki so podatki iz Flora 
Hellenica Database, predvsem pred letom 1992. Velikost kvadrantov 
UTM mreže je 1 km.
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Orchid Flora of Pelion and Mavrovouni
Besides feral pigs, we encountered herds of domesti-
cated, free-range pigs, that similarly cause major damage 
to the ecosystem. We witnessed sizeable herds roaming 
unattended in the forest ecosystem (Figure 10).
Figure 7: Fearless of humans, feral swine dig up important orchid 
biotopes at site Psili Rachi I on Mount Mavrovouni.
Slika 7: Divji prašiči, ki se ljudi ne bojijo, so prekopali pomembna 
rastišča orhidej na lokaciji Psili Rachi I na gori Mavrovouni.
Figure 8: Feral swine devastate large tracts of land near the hamlet 
of Ano Kerasia on Mount Pelion. In the fall of 2021, the supervisory 
authority tipped us off about a sizeable population of Spiranthes 
spiralis. By the time we got there, the site was all dug up.
Slika 8: Divji prašiči so uničili velike površine zemljišč blizu naselja 
Ano Kerasia na gori Pelion. Jeseni 2021 so nas odgovorni obvestili 
o veliki populaciji vrste Spiranthes spiralis, vendar je bilo rastišče do 
našega prihoda že razrito.
Figure 9: Specimens of Orchis italica dug up by feral pigs at site 
Metochi of S Magnesia.
Slika 9: Primerki vrste Orchis italica, ki so jih izkopali divji prašiči na 
lokaciji Metochi v južnem delu območja Magnesia.
Figure 10: Domesticated, free-range pigs graze unattended at site 
Kalamaki of Larissa.
Slika 10: Udomačeni prašiči, na lokaciji Kalamaki v območju Larissa, 
ki se prosto pasejo.
Recognising the problem, the national authorities have 
instituted an emergency measure to contain the over-
population of feral swine invading rural and suburban ar-
eas. The so-called “persecution crews,” essentially ad hoc 
extermination squads made up of licensed hunters, are 
supposed to hunt feral pigs. This measure is in addition 
to the harvest quota allowed hunters during hunting sea-
son, as it can be invoked by local authorities at any time 
throughout the year. Whether this measure is effective, it 
remains to be seen. We predict that, because the measure 
is implemented on a voluntary basis, without financial 
incentive, its success will be limited. In fact, offering a 
financial incentive could be worse, as part of the public 
may object to tax money subsidising the slaughter of ani-
mals. In addition, there is certainly a liability issue related 
to encouraging consumption of game meat that has not 
been inspected by veterinarians. Feral swine are vectors of 
numerous diseases and parasites infecting livestock and 
humans (Finzel & Baldwin, 2015).
Instead of invoking piecemeal measures such ad hoc 
“persecution crews,” whose success largely depends on 
the goodwill of volunteers, the most successful  feral 
Figure 7
Figure 9
Figure 8
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Orchid Flora of Pelion and Mavrovouni
swine management strategies employ a diversity of tactics 
in a comprehensive, integrated approach. Some solutions 
tried in other parts of the world, other than shooting, 
are trapping (Sharp, 2012) and installation of feeders 
delivering contraceptives (National Hog Farmer, 2021). 
We think that restoring populations of natural predators 
should be considered as an additional measure within 
the framework of an integrated wildlife management 
strategy. 
b. Containment of grazing by ruminants
We witnessed intensive grazing by small ruminants 
(sheep and goats) as well as bovines within the narrow 
study area, especially on Mount Mavrovouni. The dam-
age by ruminants is twofold: (i) direct consumption of 
the aboveground plant parts before they set seed, and (ii) 
inadvertent trampling during the search for some other 
food (Figure 11). Administratively, the narrow study area 
belongs almost entirely to the Municipalities of Ayia, Riga 
Ferreou, and Zagora-Mouresi. From aggregate statistics of 
the Payment and Control Agency for Guidance and Guar-
antee Community Aid (2024) we tallied the populations 
of small ruminants and bovines by municipality. Of the 
bovines, 98–99% are beef cattle, mostly free-range. The 
results for the three Municipalities of interest are mapped 
in Figure 12. The Municipalities of Ayia and Riga Ferre-
ou are grazed by several thousand small ruminants and a 
few thousand cattle; this is consistent with our observa-
tions of severely degraded ecosystems in grazed lands. On 
the other hand, in the Municipality of Zagora-Mouresi, 
where almost no animal husbandry is practised, we have 
not observed degradation of ecosystems due to grazing. 
Unfortunately, we could not find data tallied by Munici-
pal Unit (a subdivision of Municipality) that would yield 
a more accurate picture. 
It is difficult to propose management measures for graz-
ing. A major obstacle is the institutional framework that 
governs EU subsidies for animal husbandry that does not 
hold stockbreeders accountable for their practices. Most 
livestock operations are kept afloat with generous injec-
tions of subsidies, without regard to their inherent eco-
nomic viability or environmental sustainability (Lorent 
et al., 2008; Tsiouni et al., 2021). The root cause of the 
problem is a notorious algorithm concocted by national 
authorities titled the “technical solution.” Simply put, it 
means that stockbreeders earn subsidies based on a hypo-
thetical apportionment of a national inventory of range-
lands that may be located several hundred kilometres 
away, rather than the actual lands they graze. Because the 
subsidies are disjointed from the actual land they graze, 
stockbreeders are not responsible for sound land manage-
ment practices.
At the very least, we propose delimiting biotopes of 
high orchid diversity and enact controls on grazing, so 
that at least some of the populations are protected. Our 
findings can be used as a starting point to define “islands” 
of high orchid diversity that are worth conserving, based 
on the relative rarity and conservation interest for each 
species.
Nevertheless, we do not advocate a general ban on graz-
ing, as grazing is not without its merits:
Figure 11: Beef cattle force their way through the pseudo-maquis, 
trampling numerous orchids in the process. Site Agrioachladia on 
Mount Mavrovouni.
Slika 11: Govedo, ki se prebija skozi pseudomakijo, potepta številne 
orhidje. Lokacija Agrioachladia na gori Mavrovouni.
Figure 12: The narrow sampling area, outlined in dark blue, is shared 
among the Municipalities of Ayia, Riga Ferreou, and Zagora-Mouresi. 
The numbers after each Municipality are tallies of small ruminants/
bovines, in thousand heads.
Slika 12: Ožje preučevano območje, označeno s temno modro, sega 
v občine Ayia, Riga Ferreou in Zagora-Mouresi. Številke ob imenu 
občine so seštevki malih prežvekovalcev/ goveda v tisočih glav.
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Orchid Flora of Pelion and Mavrovouni
(i) Owing to deruralisation, declining human fertility, 
and subsequent human population decline, much 
arable land in the countryside has been abandoned. 
Most orchid species are photophiles, meaning that 
they thrive in open ecosystems with sparse woody 
vegetation, such as thin maquis, phrygana, and 
rangeland. Grazing may contribute to preserving 
these habitats by preventing afforestation. 
(ii) Our research has demonstrated that moderate graz-
ing may preserve and even enhance the soil micro-
bial functions that promote soil nutrient cycling 
and, by extension, soil health (Katsalirou et al., 
2010; Katsalirou et al., 2016a).
In summary, while overgrazing must be contemplated as 
a major driver for declining orchid diversity, moderate graz-
ing may be beneficial to orchid conservation. Therefore, in 
areas of high orchid diversity we propose a ban on graz-
ing during the reproductive stage, from budding to seed 
maturity, which for most species spans the interval from 
March to June. In areas where Spiranthes spiralis abounds, 
we propose a ban on grazing from October to December.
c. Establishment of genetic micro-reserves
To ensure a long-term supply of genetic material for 
future in vitro propagation efforts, we propose the geo-
graphic delineation of genetic micro-reserves for the five 
taxa of conservation interest. These micro-reserves must 
be fenced with solid fences to resist grazing animals. Based 
on our findings, we propose the following locations:
(i) Ophrys scolopax subsp. cornuta: Although it is not an 
altogether unusual taxon within the narrow study 
area, we did not find sizeable populations. The larg-
est population was recorded at location Petralona 
(site 13 in Appendix I).
(ii) Ophrys umbilicata subsp. umbilicata: The largest 
population was recorded near Lake Karla (site 26 in 
Appendix I). Currently, it is under direct and con-
stant threat by a nearby sheep farm. 
(iii) Orchis provincialis: Most locations are unsuitable as 
genetic reserves because they are within inhabited 
areas or logged forests with frequent traffic. The 
best option would be location Stavros II (site 9 in 
 Appendix I).
(iv) Orchis simia: The largest population was recorded at 
location Agrioachladia (site 24 in Appendix I). It is 
under direct and constant threat from grazing and 
trampling by free-range cattle. 
(v) Spiranthes spiralis: Although it is not an altogether 
unusual taxon within the narrow study area, we did 
not find sizeable populations. The largest popula-
tion was recorded at location Skiti II (site 19 in Ap-
pendix I).
d. Management of road verges
Road verges are important wildlife corridors linking 
habitats. Throughout much of the world, road banks and 
medians are respected as valuable genetic reserves of na-
tive flora. If managed correctly, road verges can support 
remarkable collections of orchids. In Mounts Pelion and 
Mavrovouni, spectacular stands of Orchis italica and vari-
ous Ophrys adorn the road banks in springtime. One rea-
son is that roads interrupt the thick, impenetrable forest 
allowing sunlight to reach the ground. Roadside verges 
are sanctuaries where orchids “escape” from intensive 
farming and grazing. Proper management involves simply 
doing less: Give the roadside vegetation some space, avoid 
herbicides, and allow plants to set seed before mowing 
(Fekete et al., 2017).
Unfortunately, local authorities in charge of roadside 
maintenance are not educated about this valuable genetic 
reserve. Along the Aegean coast of Mount Pelion, we 
frequently observed maintenance crews indiscriminately 
mowing the roadsides in early summer in preparation for 
the tourist season. 
On some paved roads, such as from Chania to Kissos, 
clearcutting a wide strip on either side of the road is un-
necessary; a shoulder has been provided between the 
pavement and the kerb that doubles as an emergency 
lane (Figure 13). Mowing the vegetation well beyond this 
shoulder does not confer any additional benefit to traf-
fic safety. The banks of this road are rich in species such 
Figure 13: The verges on either side of the road from Chania to Kissos 
on Mount Pelion are rich in orchids such as Dactylorhiza saccifera 
and Epipactis microphylla. A lone inflorescence of D. saccifera escaped 
mowing.
Slika 13: Na cestnih brežinah na obeh straneh ceste od naselja Chania 
do naselja Kissos na gori Pelion, so bogata nahajališča orhidej, kot 
sta Dactylorhiza saccifera in Epipactis microphylla. Le eno socvetje 
D. saccifera ni bilo pokošeno.
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Orchid Flora of Pelion and Mavrovouni
as Dactylorhiza saccifera and Epipactis microphylla that 
should be protected at least until seed maturity.
A different situation arises along the narrow, convo-
luted roads that connect the traditional villages of Pelion 
along the Aegean coast, such as from Zagora to Chorefto. 
There, the width of the pavement is just enough for two 
cars, and sometimes not even that. While in this case 
mowing of the road banks is unavoidable to maintain 
traffic safety, it should be delayed as much as possible, 
ideally after most orchids have set seed, and certainly not 
before 30 June. 
Finally, along the network of logging trails on the NE 
slope of Mounts Pelion and Mavrovouni, mowing of the 
trail banks can be delayed until well after 30 June, as they 
do not serve tourist traffic.
e. Measures for chestnut culture
Natural chestnut forest can be rich in orchids, as our 
preliminary reconnaissance from the Aegean coast of 
Mount Pelion attests. Chestnut forest often is sparser than 
beech, Hungarian oak, or maquis, so that more sunlight 
reaches the ground. Encouraged by abundant findings in 
natural chestnut woods, our field task force decided to 
expand its search to cultivated chestnut groves on Mount 
Mavrovouni, E of the village of Potamia. During the pre-
liminary reconnaissance phase, we traversed eight square 
cells 1 km2 wide (Figure 14). 
Our survey yielded zero orchid findings, a sober-
ing realisation give the overall high orchid diversity on 
Mount Mavrovouni. We soon realised that, in cultivated 
chestnut groves, the understory is meticulously mowed, 
while pesticides are liberally applied with tractor-drawn 
air-blast sprayers. Under these conditions, survival of 
orchids – or any herbaceous vegetation for that matter 
– is questionable. We recommend that mowing of the 
understory is delayed until after seed set, certainly after 
30 June. Meanwhile, we recommend organic chestnut 
culture as an alternative method to help restore native 
biodiversity. 
f. Measures for apple culture
The N part of the narrow study area that is flatter, and 
the SE part that is centred around the village of Pouri, are 
largely covered by intensely cultivated apple orchards that 
may intermingle with patches of natural vegetation. The 
corresponding Corine Land Cover classes are “2.2.2 Fruit 
trees and berry plantations” and “2.4.3 Land principally 
occupied by agriculture, with significant areas of natural 
vegetation” (Figure 15).
Figure 14: Orange fill indicates chestnut woods (natural and 
cultivated) within the narrow study area. Since Corine Land Cover is 
not so detailed, photo-interpretation was provided by the Management 
Unit of Protected Areas of Thessaly. We surveyed along the green trail 
that crosses the expansive chestnut plantations E and S of the village of 
Potamia. UTM grid spacing 1 km.
Slika 14: Oranžna barva označuje kostanjeve gozdove (naravne in 
kultivirane) v ožjem preučevanem območju. Ker Corine Land Cover 
ni tako natančen, smo uporabili foto-interpretacijo, ki nam jo je 
posredovala Uprava zavarovanih območij Tesalije. Terenski pregled smo 
opravili ob zeleni poti, ki prečka kostanjeve nasade vzhodno in južno 
od vasi Potamia. Velikost kvadrantov UTM mreže je 1 km.
Figure 15: Yellow fill indicates Corine Land Cover class “2.2.2 Fruit 
trees and berry plantations”. Green fill indicates class “2.4.3 Land prin-
cipally occupied by agriculture, with significant areas of natural vegeta-
tion.” At the N and the SE part of the narrow study area, these two 
classes correspond primarily to apple orchards that may be interspersed 
with patches of natural vegetation. UTM grid spacing 1 km.
Slika 15: Z rumeno je označena Corine raba tal “2.2.2 Sadovnjaki in 
nasadi jagodičja”. Zelena predstavlja rabo “2.4.3 Kmetijska zemljišča 
z večjim deležem naravne vegetacije”. Na severnem in jugovzhodnem 
delu ožjega preučevanega območja ta tip rabe predstavljajo predvsem 
sadovnjaki jablan, ki se izmenjujejo s pasovi naravne vegetacije. Ve-
likost kvadrantov UTM mreže je 1 km.
In apple orchards, the ground is meticulously mowed, 
while pesticides are liberally applied with tractor-drawn 
air-blast sprayers. Under these conditions, the occasional 
orchids find refuge within the remnants of native forest 
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Orchid Flora of Pelion and Mavrovouni
(Figure 16). We recommend that borders of native veg-
etation are preserved around the apple orchards. We rec-
ommend organic apple culture as an alternative method 
to help restore native biodiversity. 
g. Photovoltaic parks
A sizeable population of Ophrys umbilicata subsp. um-
bilicata, one of the taxa of conservation interest, was lo-
cated atop the hill NE of the village of Neochori (Plasia) 
of Larissa, along with specimens of Ophrys sphegodes sub-
sp. taurica, Ophrys tenthredinifera, and Serapias vomera-
cea, altogether comprising an important island of orchid 
diversity on a hill surrounded on three sides by intensive 
farming.
We recorded the population of Ophrys umbilicata sub-
sp. umbilicata on 04.04.2022. By 16.05.2022, already 
land surveyors were mapping the hillside. By 29.06.2022, 
the first solar panel mounts were driven to the ground. 
In the process, we witnessed the land being meticulously 
cleared with fire and bulldozers. Currently, a large photo-
voltaic park covers the hillside (Figure 17).
Figure 16: Hillslopes around the 
village of Pouri largely covered 
by intensive apple orchards. The 
occasional orchids find refuge within 
patches of native chestnut forest 
(top).
Slika 16: Pobočja pri vasi Pouri, 
kjer so intenzivni nasadi jablan. 
Orhideje občasno uspevajo v 
zaplatah naravnega kostanjevega 
gozda (zgoraj).
Figure 17: A large photovoltaic 
park on the hill NE of the village of 
Neochori (Plasia) of Larissa, once 
a rich orchid biotope. The native 
vegetation has been cleared with fire 
and bulldozers.
Slika 17: Velik fotovoltaični park na 
SV hribu pri vasi Neochori (Plasia) v 
območju Larissa, kjer je bilo nekoč z 
orhidejami bogato rastišče. Naravno 
vegetacijo so odstranili z ognjem in 
mehanizacijo.
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Orchid Flora of Pelion and Mavrovouni
Projects of such scale effect a dramatic change in the 
landscape and incur massive loss of biodiversity that does 
not seem to matter to policy makers, despite that the or-
chid family has been afforded institutional protection. 
Most solar parks are so remote that they encounter little 
reaction by the public; according to the 2011 census, the 
village of Neochori counts 41 residents. Besides, the pub-
lic has been convinced that renewable energy is good for 
the planet. Actually, these projects stretch the very notion 
of “renewable,” in the sense that the energy itself may be 
renewable yet the ecosystem is irrevocably damaged. At 
the very least, mitigating actions should be considered. A 
small proportion of the budget of such projects could be 
allocated to actions for recording, preserving, and restor-
ing native biodiversity in disturbed ecosystems.
h. Extreme weather
On 5 Sep. 2023, storm Daniel, the fiercest Medi-
terranean hurricane in recorded history, hit the study 
area with full force. Severe erosion and mudslides oc-
curred throughout the study area. It will take years to 
assess the damage to natural resources. We hope that our 
study will serve as baseline to assess the effects of this 
extreme weather event in the years to come.
Conclusions and 
Recommendations
During the fieldwork of the present project 129 square 
cells 1 km2 wide were explored, mostly within the Natura 
sites of Mounts Pelion and Mavrovouni, altogether com-
prising 796 km2. Of the sampled cells, 100 yielded orchid 
findings. In total, 39 orchid taxa (species and subspecies) 
plus two nothotaxa were recorded. While both Natura 
sites were explored, the sampling effort was intensified 
within a narrow study area of 550 km2 encompassing the 
N part of Mount Pelion and the whole of Mount Ma-
vrovouni. In the narrow study area, 33 orchid taxa and 
nothotaxa were recorded. Of those, 13 constitute new 
additions to the Flora Hellenica Database: Anacamptis pa-
pilionacea subsp. papilionacea, A. pyramidalis, Himanto-
glossum robertianum, Ophrys × delphinensis, O. sicula (syn. 
O. lutea subsp. galilaea), O. speculum subsp. speculum, 
O. sphegodes subsp. sphegodes (syn. O. hebes), O. tenthre-
dinifera, O. tenthredinifera × O. sphegodes subsp. taurica, 
O. umbilicata subsp. umbilicata, Orchis italica, O. simia, 
and Platanthera bifolia.
The main environmental threats facing orchids are 
rooting by feral and domesticated free-range pigs, over-
grazing by ruminants, clearing of road verges, intensive 
chestnut and apple culture, and the encroachment of 
photovoltaic farms. A novel threat is extreme weather 
events such as storm Daniel. We recommend contain-
ment of feral and domesticated free-range pigs, limi-
tations on grazing by ruminants, the establishment of 
micro-genetic reserves in orchid-rich areas, restrictions 
on mowing of the road verges until after the orchids have 
set seed, promoting organic chestnut and apple culture, 
and measures mitigating the impact of massive photo-
voltaic projects.
Acknowledgements
The project “Sampling design for the species Ophrys cor-
nutula (Bern Convention), Spiranthes spiralis, Ophrys 
umbilicata, and Orchis simia, in vitro sexual propagation, 
and transplanting in the area Karla-Mavrovouni-Kefal-
ovryso Velestinou-Delta Piniou” materialised within the 
framework of Action “Subsidy toward the Management 
Body of Karla-Mavrovouni-Kefalovryso Velestinou-Del-
ta Piniou for management actions related to protected 
areas, species and ecotopes,” which is part of the Op-
erational Program “Transportation Infrastructure, En-
vironment, and Sustainable Development 2014–2020” 
(MIS: 5032679). The project was funded by the Cohe-
sion Fund of the EU and national funds through the 
Public Investment Programme. The authors are greatly 
indebted to the staff of the Management Unit of Protect-
ed Areas of Thessaly (former Management Body of Kar-
la-Mavrovouni-Kefalovryso Velestinou-Delta Piniou) for 
administrative support. We are particularly indebted to 
staff members Georgia Delivasi, Konstantinos Maggos, 
and Alexandros Chatzigoulas for assistance with the field 
survey. We thank the volunteers who enriched our ranks, 
Eleni Faniadou and Efstathios Koutis. Brian Baer gener-
ously donated the digital camera that documented the 
project. We are grateful to Prof. Arne Strid for gener-
ously sharing records from the Flora Hellenica Database. 
Finally, we thank the Directorate of Forest Protection 
of the Ministry of Environment and Energy for grant-
ing the research and photography permit with protocol 
number ΥΠΕΝ/ΔΠΔ/37540/2100/01-07-2021.
ORCID iDs
Argyrios Gerakis  https://orcid.org/0000-0002-6927-1357
Ursina Steinemann  https://orcid.org/0009-0002-5377-8281
Sofia Drizi  https://orcid.org/0009-0006-9614-9505
Vasilis Ioannidis  https://orcid.org/0000-0003-4860-341X
Eirini Katsalirou  https://orcid.org/0000-0003-1836-8746
Athina Klasogianni  https://orcid.org/0009-0007-4584-5129
Kostas Koutis  https://orcid.org/0000-0002-8591-0227
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S/N UTM SITE DESCRIPTION
1 643 4391 Neochori II Asphodel rangeland dotted with patches of kermes oak on hilly terrrain 0.5 km NE of 
Neochori, 130 m a.s.l.
2 644 4391 Neochori I Asphodel rangeland dotted with patches of kermes oak on hilly terrrain at the fringe of vast 
solar park 0.7–0.9 km NE of Neochori, 100–150 m a.s.l.
3 645 4388 Kastri I Asphodel rangeland interspersed with patches of short kermes oak on rolling terrain 
1.1–1.8 km NE of Kastri, 260–300 m a.s.l.
4 646 4388 Kastri II Short, thin maquis of kermes oak on rolling terrain 1.9 km NE of Kastri, 250 m a.s.l.
5 647 4385 Ano Amygdali Mosaic of maquis, phrygana, farmland and cypress stands on rolling terrain 0.2–0.5 km 
NW of Ano Amygdali, 340–360 m a.s.l.
6 647 4394 Aetolofos II Sparse abandoned olive grove overgrown with phrygana on hillslope 0.1 km NW of 
Aetolofos, 130 m a.s.l.
7 648 4394 Aetolofos I Sparse abandoned olive grove overgrown with phrygana on hillslope 0.3 km N of 
Aetolofos, 180 m a.s.l.
8 649 4383 Kalamaki Larissas Dirt road winding through tall, dense maquis of kermes oak on rolling terrain 1.5–2 km N 
of Kalamaki, 390–490 m a.s.l.
9 649 4388 Stavros II Dense, impenetrable maquis of kermes oak on hilly terrain 4.8 km SW of Potamia, 
220–230 m a.s.l.
10 649 4389 Stavros I Dirt road winding through tall, dense, impenetrable maquis on hilly terrain 3.8 km SW of 
Potamia, 200 m a.s.l.
11 650 4383 Konstadouli  
Rachi I
Dirt road winding through tall, dense maquis of kermes oak on rolling terrain 1.7 km 
N-NE of Kalamaki, 480 m a.s.l.
12 650 4392 Potamia II Tiny patches of rangeland among tall, dense maquis of kermes oak on rolling terrain 
2.7 km W-NW of Potamia, 140 m a.s.l.
13 651 4381 Petralona Paved road through degraded maquis on rolling terrain 1.2 km E of Kalamaki, 300 m a.s.l.
14 651 4383 Konstadouli Rachi 
II
Dirt road winding through tall, dense maquis of kermes oak on rolling terrain 2.4 km NE 
of Kalamaki, 500 m a.s.l.
15 651 4390 Potamia I Dirt road meandering through short, thin maquis interspersed with cultivated fields on 
hilly terrain 1.9 km SW of Potamia, 160–180 m a.s.l.
16 652 4382 Elafos Dirt road through degraded maquis on rolling terrain 0.2 km SW of Elafos, 460 m a.s.l.
17 654 4378 Gianni Palla III Abandoned olive grove overgrown with kermes oak and Jerusalem thorn on hillside 2.2 km 
N-NW of Keramidiotika Kalivia, 220 m a.s.l.
18 654 4379 Gianni Palla I Thin, tall maquis of kermes oak on steep rocky mountainous terrain 2.5–3.1 km N-NW of 
Keramidiotika Kalivia, 120–340 m a.s.l.
19 654 4393 Skiti II Tall maquis at the fringe of cultivated fields on hilly terrain 1.7–2.0 km W of Skiti, 
140–190 m a.s.l.
20 655 4379 Souvalitsa Dirt road winding through maquis of kermes oak on hilly terrain 2.6 km N of 
Keramidiotika Kalivia, 370 m a.s.l.
21 656 4379 Gianni Palla II Vast expanse of phrygana interspersed with dwarf kermes oak and tall Hungarian oak on 
mountainous terrain 2.7–3.5 km N-NE of Keramidiotika Kalivia, 430–540 m a.s.l.
22 656 4395 Skiti I Road winding through gorge with steep slopes lined with maquis forest and orchards 
1.1 km N of Skiti, 250 m a.s.l.
23 657 4384 Koryfi Dirt road through beech forest 3.4 km SW of Sklithro, 770 m a.s.l.
24 658 4380 Agrioachladia Rocky clearings within short, dense maquis of kermes oak on relatively flat terrain, heavily 
grazed by cattle 5.7 km W-SW of Keramidi, 670 m a.s.l.
Appendix I
Surveyed sites: Serial number; UTM easting and northing to 1 km precision (the grid zone designation 34 is omitted); site name; 
site description.
Pregledane lokacije: zaporedna številka; UTM koordinate do 1 km natančno (oznaka mreže 34 je izpuščena); ime lokacije; 
opis lokacije.
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S/N UTM SITE DESCRIPTION
25 659 4370 Gerani Hill Phryganic patches among tall, dense maquis of kermes oak and Jerusalem thorn at the foot 
of Gerani Hill 3.5 km SW of Kanalia, 90–100 m a.s.l.
26 659 4370 Karla Gently sloping rangeland dotted with kermes oak, wild pear and Jerusalem thorn between 
Gerani Hill and Lake Karla 3.4 km SW of Kanalia, 50–60 m a.s.l.
27 659 4391 Ano Polydendri Dirt road through mixed deciduous forest 2.4 km SW of Kato Polydendri, 320 m a.s.l.
28 660 4378 Tsaghiani I Dwarf maquis of kermes oak degraded by heavy grazing on rocky, hilly terrain 4.7–5.0 km 
N-NW of Kanalia, 770–790 m a.s.l.
29 661 4375 Kanalia II Short, thin maquis of kermes oak on rocky hillslope 1.4–1.7 km N-NW of Kanalia, 
360–510 m a.s.l.
30 661 4378 Tsaghiani II Dwarf maquis of kermes oak degraded by heavy grazing on rocky, hilly terrain 4.2–4.3 km 
N of Kanalia, 740–760 m a.s.l.
31 661 4381 Keramidi I Clearings among tall, dense maquis of kermes oak on very rocky ground along a valley 
2-3 km W-SW of Keramidi, 540–620 m a.s.l.
32 661 4386 Sklithro I Mixed chestnut/oak forest on sloping terrain 0.4–0.6 km NE of Sklithro, 230–340 m a.s.l.
33 662 4365 Dog refuge Phryganic patches among maquis of kermes oak at the foot of unnamed hill 1.7 km S-SE 
of Glafyra, 370 m a.s.l.
34 662 4375 Kanalia I Short, dense maquis of kermes oak on rocky hillslope 1.4–1.6 km N of Kanalia, 
480-560 m a.s.l.
35 662 4381 Keramidi II Banks of Kakorema Brook running through patches of tall, dense maquis of kermes oak 
and rangeland dotted with wild pear 1.1–2 km W-SW of Keramidi, 500–560 m a.s.l.
36 662 4386 Sklithro II Mixed forest of chestnut/Hungarian oak on sloping terrain 1.3 km NE of Sklithro, 
230–340 m a.s.l.
37 663 4364 Amygdalia Large expanse of degraded maquis on rolling terrain 2.4 km N-NE of Melissatika, 
300–310 m a.s.l.
38 663 4373 Labitsa III Short, dense maquis of kermes oak on mountainous terrain 1.6 km E of Kanalia, 
210 m a.s.l.
39 663 4376 Psili Rachi I Abandoned olive grove overgrown with juniper, mock privet, and kermes oak on rolling 
terrain 2.5 km NE of Kanalia, 670 m a.s.l.
40 664 4373 Labitsa II Short, dense maquis of kermes oak on mountainous terrain 1.7–2.1 km E of Kanalia, 
250–310 m a.s.l.
41 664 4374 Labitsa I Short, dense maquis of kermes oak on mountainous terrain 2.5 km E of Kanalia, 
370 m a.s.l.
42 664 4376 Psili Rachi II Short, thin maquis of kermes oak and juniper on rocky, rolling terrain 2.8–3.2 km NE of 
Kanalia, 690–720 m a.s.l.
43 664 4381 Keramidi III Tall, thin maquis interspersed with rangeland on hilly terrain 0.5 km SE of Keramidi, 
360–420 m a.s.l.
44 664 4382 Keramidi IV Paved road winding through terraces of chestnut forest, farmland, and rangeland 0.5 km E 
of Keramidi, 290 m a.s.l.
45 665 4368 Paliambela I Patches of Cistus phrygana among maquis on rolling terrain 1.3–1.6 km S of Kerasia, 
320–340 m a.s.l.
46 665 4369 Kerasia I Paved road winding through tall, dense maquis of kermes oak in the valley S of Kerasia, 
150–210 m a.s.l.
47 665 4374 Labitsa IV Short, dense maquis of kermes oak on mountainous terrain 2.8–3.5 km E-NE of Kanalia, 
420–570 m a.s.l. 
48 666 4368 Paliambela II Patches of Cistus phrygana among maquis on rolling terrain 1.6 km SE of Kerasia, 
350 m a.s.l.
49 666 4369 Kerasia II Dirt road winding through tall, dense maquis of kermes oak in the valley 0.7–0.9 km SE of 
Kerasia, 230–260 m a.s.l.
50 667 4365 Lomvarda Dirt road winding through maquis on hill slope 1.2 km N-NE of Gioni, 360 m a.s.l.
51 667 4366 Licopazaro Dirt road winding through maquis on hill slope 2.1 km N-NE of Gioni, 420 m a.s.l.
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S/N UTM SITE DESCRIPTION
52 667 4367 Alevizou Bridge Dirt road winding through maquis on hill slope 2.9 km N-NE of Gioni, 350–380 m a.s.l.
53 668 4369 Panagia 
Paliochoritisa
Dirt road winding through tall, dense maquis of kermes oak on sloping terrain 2.5 km SE 
of Kerasia, 340 m a.s.l.
54 668 4370 Agios Athanasios Dirt road winding through tall, dense maquis of kermes oak on sloping terrain 3.2 km E of 
Kerasia, 550 m a.s.l.
55 669 4369 Moni Sourvias V Clearings amidst regenerated forest of Hungarian oak on hillslope 4 km E-SE of Kerasia, 
480–530 m a.s.l.
56 669 4370 Moni Sourvias I Tall, dense maquis of kermes oak on hilly terrain 3.6 km E-NE of Kerasia, 600 m a.s.l.
57 669 4371 Kastanitsa Tall, dense maquis of kermes oak on hilly terrain 3.6 km E-NE of Kerasia, 
680-700 m a.s.l.
58 669 4373 Kosta Rachi Degraded maquis on steep slope 0.8 km NE of Ano Kerasia, 720 m a.s.l.
59 670 4369 Moni Sourvias IV Clearing amidst regenerated forest of Hungarian oak on hill slope 4.3 km E of Kerasia, 
610 m a.s.l.
60 671 4364 Profitis Ilias 
Makrinitsas
Mosaic of tall, thin maquis of kermes oak and juniper, phrygana, and farmland on hilly 
terrain 0.9–1.2 km N-NE of Makrinitsa, 970–990 m a.s.l.
61 671 4369 Moni Sourvias III Clearing amidst beech forest dotted with dwarf kermes oak on the banks of Kerasiotis 
gorge 6.2 km E of Kerasia, 830 m a.s.l.
62 672 4362 Moni Sarof Paved road through mixed deciduous forest 0.3 km NW of Sarof Monastery, 740 m a.s.l.
63 672 4369 Moni Sourvias II Clearings of rocky rangeland amidst mixed forest of beech and kermes oak along the banks 
of Kerasiotis gorge 6.7 km E-SE of Kerasia, 840–880 m a.s.l.
64 673 4361 Agia Triada 
Portarias
Chestnut forest on mountain slope 1 km E of Portaria, 800 m a.s.l.
65 674 4360 Alikopetra I Phrygana on sloping terrain 1.9 km W of Drakia, 810 m a.s.l.
66 674 4360 Alikopetra II Phrygana on sloping terrain 1.5 km W of Drakia, 970–1000 m a.s.l.
67 677 4362 Chania Paved road through beech forest just E of Chania, 1180 m a.s.l.
68 678 4362 Ski Resort II Beech forest on sloping terrain 1.1 km E-SE of Chania, 1220 m a.s.l.
69 678 4370 Agia Marina Beech forest on steep slope 1.8 km SW of Pouri, 970–1010 m a.s.l.
70 678 4371 Vlachochorafa Chestnut forest on sloping terrain 1.5 km W of Pouri, 810 m a.s.l.
71 678 4372 Pouri VII Terraces of chestnut plantation on steep slope 1.3 km NW of Pouri, 650 m a.s.l.
72 679 4363 Nisaki Dirt road through beech forest on steep mountain slope 1.1 km N of ski resort, 
910-920 m a.s.l.
73 679 4364 Profitis Ilias Beech forest on sloping terrain 0–0.6 km NE of Profitis Ilias church, 780–830 m a.s.l.
74 679 4370 Pouri V Stand of native chestnut forest amidst intensively cultivated apple orchards on hilly terrain 
1.2 km S-SW of Pouri, 580 m a.s.l.
75 679 4371 Melistres Chestnut forest on steep slope 0.9 km SW of Pouri, 840 m a.s.l.
76 679 4371 Pouri VIII Chestnut forest on steep slope 0–0.4 km SW of Pouri, 520–650 m a.s.l.
77 679 4372 Pouri II Dirt road winding through terraces of chestnut plantation on steep slope 0.2–0.8 km NW 
of Pouri, 480–630 m a.s.l.
78 679 4374 Areto Dirt road through maquis dominated by holly oak and strawberry tree 2.1 km N-NE of 
Pouri, 240 m a.s.l.
79 680 4353 Kato Gatzea Paved road running through olive groves and residences on gentle slope 0.5 km NW of 
Kato Gatzea, 30 m a.s.l.
80 680 4355 Agii Apostoli Paved road winding through tall, dense maquis, farmland, and residential areas on steep 
slope 0.7 km NW of Ano Gatzea, 330 m a.s.l.
81 680 4355 Agios Georgios 
Nilias
Trail through abandoned olive grove overgrown with maquis of kermes oak and strawberry 
tree on sloping terrain 0.8 km SE of Agios Georgios Nilias, 470 m a.s.l.
82 680 4362 Ski Resort I Paved road through beech forest on sloping terrain 2.4 km E-SE of Chania, 
1170-1190 m a.s.l.
83 680 4365 Karavoma Chestnuts grown on terraces 2.1 km W-SW of Makrirachi, 630 m a.s.l.
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84 680 4370 Pouri VI Chestnut forest on steep slope 0.8–1.3 km S-SE of Pouri, 300–490 m a.s.l.
85 680 4372 Pouri III Road winding through dense forest of chestnut, strawberry tree, and juniper on hill slope at 
the outskirts of Pouri, 300–340 m a.s.l.
86 681 4354 Agia Triada Ano 
Gatzeas
Paved road winding through farmland and residential areas on steep slope in Agia Triada, 
240 m a.s.l.
87 686 4360 Taxiarches I Chestnut forest on gently sloping terrain 0.5 km NW of Taxiarches, 670 m a.s.l.
88 687 4355 Kefales Dirt road through maquis on sloping terrain 1.9 km E of Milies, 540 m a.s.l.
89 687 4360 Taxiarches II Chestnut forest on gently sloping terrain at the ouskirts of Taxiarches, 530–630 m a.s.l.
90 687 4361 Taxiarches III Chestnut forest on gently sloping terrain at the outskirts of Taxiarches, 570–610 m a.s.l.
91 687 4361 S Agia Paraskevi Private garden in the community of Agia Paraskevi, 510 m a.s.l.
92 687 4361 N Agia Paraskevi Paved road winding through the community of Agia Paraskevi, 450 m a.s.l.
93 687 4362 Agia Kiriaki Paved road through the community of Agia Kiriaki, 360 m a.s.l.
94 687 4363 Mouresi Dirt road through rangeland and olive groves on sloping terrain 0.2–0.5 km E of Mouresi, 
220–240 m a.s.l.
95 688 4361 Fakistra Road winding through olive groves on sloping terrain 0.3 km NE of Agia Paraskevi, 
220–250 m a.s.l.
96 689 4354 Choreftra Dirt road winding through degraded maquis 1.7 km SW of Kalamaki, 520–550 m a.s.l.
97 689 4359 Zoodochos Pigi 
switchbacks I
Olive groves on terraces along the switchbacks of the paved road descending to Zoodochos 
Pigi 0.7–0.9 km E of Xourichti, 220–300 m a.s.l.
98 689 4360 Zoodochos Pigi 
switchbacks II
Olive groves on terraces along the switchbacks of the paved road descending to Zoodochos 
Pigi 1.3 km NE of Xourichti, 80–90 m a.s.l.
99 689 4360 Zoodochos Pigi Seaside olive grove on terraces S of Zoodochos Pigi church, 20–30 m a.s.l.
100 690 4355 Choreftra Dirt road winding through degraded maquis 1.7 km SW of Kalamaki, 520–550 m a.s.l.
101 691 4342 Metochi Abandoned olive grove overgrown with Cistus on hilly terrain 0.3 km N-NW of Metochi, 
250 m a.s.l.
102 691 4357 Kalamaki Magnisias Phrygana on sloping terrain 0.6 km NE of Kalamaki, 230 m a.s.l.
103 692 4338 Milina Abandoned olive groves on steep slope at the outskirts of Milina, 30–70 m a.s.l.
104 693 4338 Lafkos trail to 
Milina
Olive groves sometimes overgrown with phrygana on steep slope 0.2–0.8 km W of Lafkos, 
110–290 m a.s.l.
105 693 4338 Lafkos Olive grove on gentle slope at the outskirts of Lafkos, 290 m a.s.l.
106 693 4339 Chionata Mosaic of tall, dense maquis, rangeland, and untilled olive groves on hilly terrain 
0.2-0.7-km NW of Lafkos, 270–280 m a.s.l.
107 693 4340 Kakouri Brook Untilled olive grove on hilly terrain 1.1 km NW of Lafkos, 220 m a.s.l.