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Collection Essential Guides of Flora, 2

ORCHIDS OF CENTRAL SPAIN


(Cuenca Province)

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A field guide

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Agustín Coronado Martínez
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Eduardo Soto Pérez


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2019
Contents

Prologue................................................ 6 Ophrys dyris ............................................1 122


How to use this guide ................................ 8 Ophrys incubacea ......................................1 124
The three natural landscapes of Cuenca ......... 20 Ophrys insectifera .....................................1 126
Types of Habitat in Cuenca ........................ 24 Ophrys lupercalis ......................................1 128
Rare and protected habitats ......................... 32 Ophrys lutea ............................................1 130
General Biology ...................................... 38 Ophrys scolopax ........................................1 132
Morphology and Anatomy ............................ 40 Ophrys speculum .......................................1 134
Phenology ............................................. 46 Ophrys sphegodes ...................................... 136
Orchids found in Cuenca ............................ 48 Ophrys subinsectifera ................................. 138

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Key for identifying Orchid genera ................. 50 Ophrys tenthredinifera ............................... 140

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Aceras anthropophorum ............................. 52 Genus Orchis ........................................... 142

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Anacamptis pyramidalis ............................. 54 Orchis cazorlensis ...................................... 144
Genus Cephalanthera ............................... 56 Orchis champagneuxii ................................. 146

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Cephalanthera damasonium ........................ 58 Orchis fragrans ......................................... 148
Cephalanthera longifolia ............................ 60 Orchis langei ............................................ 150

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Cephalanthera rubra ................................. 62 Orchis militaris ......................................... 152

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Coeloglossum viride .................................. 64 Orchis palustris ......................................... 154
Genus Dactylorhiza ................................. 66 Orchis papilionacea .................................... 156

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Dactylorhiza elata .................................... 68 Orchis picta ............................................. 158

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Dactylorhiza fuchsii .................................. 70 Orchis purpurea ........................................ 160
Dactylorhiza incarnata .............................. 72 Orchis tenera ........................................... 162

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Dactylorhiza insularis ................................ 74 Orchis ustulata ......................................... 164
Dactylorhiza maculata ............................... 76 Genus Platanthera ................................... 166

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Dactylorhiza sambucina ............................. 78 Platanthera algeriensis ............................... 168

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Genus Epipactis ...................................... 80 Platanthera bifolia .................................... 170
Epipactis cardina ..................................... 82 Serapias lingua ......................................... 172
Epipactis distans ...................................... 84 Spiranthes aestivalis .................................. 174
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Epipactis helleborine ................................ 86 Spiranthes spiralis ..................................... 176
Epipactis hispanica ................................... 88 Neighbouring species ................................ 178
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Epipactis kleinii ....................................... 90 Additional Biology .................................... 184


Epipactis microphylla ................................ 92 Orchids and fungi ...................................... 184
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Epipactis palustris .................................... 94 Orchid cultivation .................................... 186


Epipactis tremolsii ................................... 96 Cultivation in pots ..................................... 192
Gymnadenia conopsea ............................... 98 Evolution ................................................ 194
Himantoglossum hircinum .......................... 100 Pollination methods ................................... 196
Genus Limodorum.................................... 102 Hybrids .................................................. 202
Limodorum abortivum ............................... 102 Unusual cases .......................................... 204
Limodorum trabutianum ............................ 104 Orchids protection..................................... 205
Listera ovata .......................................... 106 Some interesting facts ................................ 207
Neotinea maculata ................................... 108 Locations ............................................... 209
Neottia nidus-avis .................................... 110 Spanish-English Lexicon of Habitats ................. 232
Genus Ophrys ........................................ 112 Glossary of terms ...................................... 233
Ophrys apifera ........................................ 114 Useful links ............................................. 235
Ophrys arnoldii ....................................... 116 Bibliography ........................................... 238
Ophrys bilunulata .................................... 118 Index of scientific names ............................. 242
Ophrys castellana .................................... 120
4 5
Prologue

We imagine that many of our readers will have come across this book with surprise to
discover that orchids are to be found in Central Spain (Cuenca province). It was the
same for us, in fact the first time we tried to identify one we had trouble in determi-
ning which family it belonged to, let alone believe it! We were astonished that these
wonderful flowers that we thought belonged exclusively to the tropical forests could
also form part of the habitat of the mountains, woods and river banks of our own
province.

Once we had confirmed the existence of orchids in Europe we began our tentative

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research, certain that we wouldn’t find more than a dozen different species. With

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orchids it’s a bit like with mushrooms; hunting for them becomes a hypnotic entertain-

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ment which you can’t stop even though the sun is going down. When we’d identified
twenty different species we just had an uncontrollable need to carry on finding more,

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to explore every corner of our land and discover where their hiding places were, which
were their favourite landscapes, and in which months they were flowering.

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Totally bewitched, we spent the next seven years enjoying one of our favourite pas-

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times: going out to the country, exploring the terrain, breathing in the pure air, slee-

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ping rough, venturing into the maze of woodlands, and uncovering the secrets of the

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natural world. We discovered the subtle relationships of an ecosystem which permits
such an incredible ability to be so amazingly in harmony, such detailed complexity and

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efficiency where each of its elements is provided for and lacks for nothing.

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At the beginning of this field work we started with a small reference material which,
in line with our growing interest into the various aspects of the ecology of orchids,

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expanded into volumes which filled our bookshelves but which no longer fit into our
rucksacks.
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Since the mid-17th C, and basically stimulated by commercial interests in their attrac-
tiveness for gardens and greenhouses around the world, research into the Orchidaceae
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family has opened up new fields, techniques and questions for work in the biological
sciences: in ecology, plant physiology, improvements in genetics, molecular genetics,
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etc. Morel’s work in the cultivation of orchid meristems set the foundations for the So why a guide to Cuenca province in Central Spain? Our Serranía mountain range is
cloning of plant species. Kullenberg’s work on orchid pollinator types and adaptations one of the most densely populated in orchid species in the whole of the Iberian Pe-
established the methods for developing numerous subsequent studies on habitat and ninsula, and we are delighted to be able to demonstrate that Cuenca’s well-preserved
pollination. The work by Zettler and Rasmussen on the symbiosis of land orchids and natural environment attracts orchid specialists from all over Europe.
their specific fungi has permitted extraordinary advances in research into mycorrhiza.
While Chase, Bateman, Pridgeon and Qamariz-Zaman, together with their respective But the main reason for publishing this guide is that we are convinced that this floral
collaborators, have achieved one of the first refined evaluations of the genetic diver- wealth should be considered as a common heritage for all to know and admire. Curio-
sity of a whole botanical family group. sity incites knowledge; it is the unconditional seed of appreciation, and in the end, the
only kind that places respect above threats or coercion, the ideal way to preserve that
Through their efforts during the past thirty years, the depth of knowledge of their natural beauty which we must protect for the future.
native orchids by European botanists has increased, and the number of recognized
species in Europe has almost tripled since 1970.

6 7
Locations

The examples of the plants are positioned according to the UTM coordinates (Uni-
versal Transvers Mercator) used in cartography and European plant distribution in the
OPTIMA project. Before the description of each species you will find a map of distribu-
tion in which you will see the province divided into squares. Let’s explain what they
correspond to.

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Spain is to be found between the longitude zones 29 and 31 and latitude bands T and

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S. The Cuenca province area is within zone 30 and between the parallels T and S. As

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can be seen, the UTM coordinates do not specify a single point but a square grid zone
which can measure 1, 10, 100 or 100.000 m. The reference value defined within the

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coordinate is placed in the bottom left corner of the square (not in the centre).

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In order to simplify this, sometimes squares are also defined as 100 × 100 km areas with
a combination of capital letters in alphabetical order, from south to north (this vertical
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reference is first), and from west to east (the horizontal reference goes after). The
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whole of the Cuenca province is included: the north in the 100 km × 100 km squares VK,
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WK, and XK, and the south in the squares VJ, WJ and XJ. Each of these large squares can
be divided into 100 squares of 10 km × 10 km assigned a combination of two numbers.
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These, in turn, can be divided into a further 100: 1 km × 1 km, and these also into ano-
ther 100: measuring 100 m × 100 m, and so on, depending on the desired precision of the
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location. For this Guide, we have used squares of an area of 1 km × 1 km, but the squares
on the map of distribution correspond to 10 km × 10 km.
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LK
UQ CK
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LJ
T UP CJ
LH
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UN CH
LG
UM CG
LF MF NF PF QF TL FF
UL VL WL XL YL BF CF DF EF
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UK CE
LD UJ CD
LC UH CC
LB UG CB
LA UF CA
LV
S UE CV
LU UD CU
LT UC CT
Aguiló Publications
J.E. García Rodríguez

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Types of Habitat in Cuenca

What follows is a description of the main types of woods and woody


shrublands to be found in Cuenca. We indicate the species of orchids that
belong to each type.

Oak Forests In Cuenca we find two types of oak


woods which, depending mainly on the
These make up a large part of our geogra- altitude, we will call Mediterranean and

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phical region and would cover it entirely Continental.

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if they hadn’t been subjected to human
intervention. The Holm Oak (Quercus ro-

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Below 1000 m we have the Mediterranean type
tundifolia) covers the whole of the area of oak woods, situated in the mesomedi-
of Castilla-La Mancha except for sma-

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terranean zone on limestone substrate
ll parts in the higher altitudes with se- soils. Such are the oak woods of Alcantud,

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mi-arid conditions. Impoverished by the where we can find Aceras antropophorum
continental climate conditions, the oak and Cephalanthera longifolia orchids. In
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woods in Cuenca are not so rich as in the many parts the oaks have been replaced
coastal or the offshore mountain regions
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by Aleppo pines (Pinus halepensis), as in
of the Mediterranean; there are fewer
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the Batanejo reservoir and the Camino al


plant varieties, as the cold has elimina- Bujoso (the River Cabriel basin) where
ted species such as the Butcher’s Broom
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groups of Limodorum abortivum, Cepha-


(Ruscus aculeatus) or the Sarsaparille lanthera longifolia, Neotinea maculata
(Smilax aspera).
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and Ophrys dyris are to be found. In the


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Arbutus unedo, Viburnum tinus and even occasionally for the Pinus halepensis or
Phillyrea latifolia. Quercus rotundifolia (= Q. ilex subsp. bal-
lota, on siliceous soils (sandstone and ter-
In the Sierra de Altomira region the char- tiary siliceous, Utrillas sands, Buntsand-
acteristic species are: Pinus halepensis, stein sands, etc.) in the mesomediterra-
Quercus coccifera, Juniperus oxycedrus, nean zone (under 1,000 m altitude) with
Juniperus phoenicea, Rhamnus alater- a subhumid ombrotrophic climate in the
nus, Rosmarinus officinalis and Cistus lower areas or dry in the higher zones.
albidus.

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In siliceous parts of the southern Iberian

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Together with these can be found Philly- System (the region of Alcantud and Sier-

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rea angustifolia, Pistacia terebinthus, ras de Mira and Talayuelas) with a meso-
Arbutus unedo, Viburnum tinus and Bux- mediterranean climate.
us sempervirens (in shaded areas and

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gorges) or Ruscus aculeatus (on shaded, Characteristic species: Arbutus unedo,
humus-rich soils). In these environments,
species of Mediterranean-type orchids .jo
Erica arborea, Erica scoparia or Phillyrea
angustifolia. In the warmer areas Pistacia
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such as Ophrys lupercalis or Ophrys dyris lentiscus, Smilax aspera or Erica multi-
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grow. flora are added, and in the shadier parts
Viburnum tinus, Ruscus aculeatus or Bux-
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Levantine siliceous Maquis us sempervirens are also included.


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shrublands The most typical orchids in this type of


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Tall, generally dense shrubs which act as habitat are: Aceras anthropophorum and
Cephalanthera longifolia.
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the main vegetation stratum or under-


growth for the Pinus pinaster pines, and
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Arbutus unedo 33
General Biology

Among the many herbaceous species which grow in our province there is
one little known botanical group in spite of it belonging to one of the most
beautiful and sophisticated families of the plant kingdom: the orchids.

The Orchidaceae is one of the most high- rain forests, where the classic orchids
ly-evolved families within the Angiosperm boxed for exotic gifts come from. Half

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Monocotyledons and second in its number of these are epiphytes, ie. they grow

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of components within the plant kingdom. on other plants, usually on the trunks of

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One out of every twelve species is an or- large trees which they cling to with their
chid. There are over 850 genera and a embracing roots. They feed on the humus

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number of species estimated at between gathered between the branches and on
20,000 and 30,000. These are classified
into 70 subtribes and 22 tribes, divided .jo
the foliage, and take their water from the
moisture in the air through tiny absorbent
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in turn into 5 subfamilies: Apostasiodeae, hanging roots.
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Cypripedioidae, Epidendroideae, Orchi-
Antarctica and the arid deserts are the
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doideae and Spiranthoideae. This division


only places in the world where orchids do
is based mainly on the different number
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not grow. They have colonized all other


of anthers and their arrangement.
habitats, from the tundra in the Arctic Cir-
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Among this enormous variety of species cle to the plains in China, in the Florida
there are all sorts: large and beautiful, swamps and in the source of the River Cu-
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tiny, subtle and delicate, showy, fragrant, ervo in Cuenca. All the European orchids
odourless or smelling of rotting meat, belong to the Holarctic ecozone.
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multicoloured, black, white, with mark-


The European continent has around 250
ings or designs.
species and numerous subspecies. The
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There are orchids with tiny flowers which majority belong to the subfamily Orchi-
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measure only millimetres (Platystele doidae; the genera Cephalanthera, Epi-


stenostachya), and plants in the Nation- tactis and Listera belong to the subfamily
al Park of Machu Picchu with 3-4 metre Epidendroideae, and the genus Spiranthes
stalks which produce dozens of flowers. is assigned to the subfamily Spiranthoide-
Orchids can be found in all kinds of hab- ae. The European orchids are all terrestri-
itats, from the ones which grow on rocks al and smaller than the tropical orchids,
to the underground orchids of Australia a fact which requires closer attention in
(Rhizanthella gardneri) which only see order to discover them and admire their
the sun when they flower, or the sapro- beauty. The enthusiast’s efforts to see
phyte Neottia nidus-avis, which we can them will be rewarded and quickly will
find in the Oak forests in Cuenca and begin to recognize the characteristics
which spends years without surfacing and which differentiate the genera. With a lit-
flowers underground. Most orchid species tle practice, a linen tester magnifier will
are found in the tropical and subtropical be found to be a most useful tool and will

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Morphology and Anatomy

ROOTS thick rhizome with many branches. The


tubers in the genus Gymnadenia are
The European orchids are robust flat and deeply dividing. The rhizome of
herbaceous plants. Their underground Neottia nidus-avis sends out numerous
parts, rhizomes or pseudotubers, can roots in a tangled ball which looks like
remain alive for several years, annually a bird’s nest. All the genera are able to
replacing the aerial or epigeous organs. send out stolonoid roots, or root-like
The name orchid derives from the stems which can produce new plants

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Greek orchis, meaning testicle, due near the main plant and which share

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to the similarity in appearance of the genetic makeup. In some plants

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the underground tubers (spherical or this means of plant propagation is more
ellipsoid). These tuberous roots store frequent than in others, as for example

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water and nutrients. in certain species of the genera Epipactis
or Orchis champagneuxii, which is why it
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is common to find them growing packed
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very closely together.
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Phloem
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Xylem
Pericycle
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Orchis Dactylorhiza Endodermis


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Starch
Parenchyma
Cortex
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Epidermis
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Ophrys scolopax root Root hair

Platanthera Neottia
STEM
The genus Dactylorhiza and the genus As with the majority of Monocotyledons,
Coeloglossum both have a tuber divided the stem is straight, single and
into finger-like lobes, rather like a unbranched. In general it has a circular
duck’s webfoot, and their roots grow section and may be hollow or filled;
from these fingers and also from the hairy, generally ash-grey in colour,
base of the stem. On the other hand, the also completely hairless, smooth and
genus Orchis has spherical or ellipsoid shiny. Usually green but can sometimes
tubers and the roots only grow from adopt reddish, yellowish or dun colours
the base of the stem. In the genera depending on the species.
Epipactis and Cephalanthera we find a
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Key for identifying Orchid Genera

1. Plants without green leaves, no chlorophyll ........................................... 2


— Plants with leaves, or green scales along the stem................................... 3

2. Long, thin spur. Stem pink or violet colour ...................... G. Limodorum (102)
— No spur. Stem light brown or cream colour.......................... G. Neottia (110)

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3. Flowers with no spur ....................................................................... 4

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— Flowers with spur .......................................................................... 11

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4. Labellum decorated with a shiny central pattern (speculum) surrounded by a

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more or less hairy band ............................................... G. Ophrys (112)
­— Labellum with no shiny central marking (speculum) and no obvious hair........... 5
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5. Labellum divided by a transverse constriction on the hypochile (inner portion)
and the epichile (outer portion)............................................................... 6
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— Labellum with no central division ........................................................ 8


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6. Pedunculate ovary. Hypochile concave or cup-shaped, shiny and nectariferous.


Flowers patent or pendulous ........................................ G. Epipactis (80)
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— Sessile ovary. Cup-shaped hypochile, no nectar. Flowers horizontal or raised .... 7


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7. Green bracts. Hypochile with parallel crests. Perianth with free divisions ..........
..................................................................... G. Cephalanthera (56)
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— Bracts of the same colour as the sepals. Epichile without crests. Perianth with
sealed divisions.............................................................. G. Serapias (172)
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8. White flowers. Spiral axis of inflorescence or as unilateral sprig..................... 9


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— Green or yellow flowers. Cylindrical inflorescence ................................... 10

9. Ovate basal leaves, with five clearly marked veins, joined together by highly visible
cross-veins .................................................................... G. Goodyera (179)
— Linear or lanceolate leaves, with no marked transverse nerves .......
............................................................................ G. Spiranthes (174)

10. Plant with two opposite leaves. Green flowers....................... G. Listera (106)
— Plant with more than two alternate leaves. Yellow flowers.......... G. Aceras (52)

11. Labellum with the central lobe considerably longer than the lateral lobes and
twisted in a spiral .................................................. G. Himantoglossum (100)
— Whole labellum or with the central lobe slightly longer than the lateral ones; no
torsion ........................................................................................... 12
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Aceras anthropophorum
(L.) W.T. Aiton

Genus Aceras R. Br.


Etymology: from a, without, and ceras, horn
or spur –Without-a-horn.

Monospecific genus of Mediterranean-


Atlantic distribution close to the genus
Orchis. Plants with nectar and without a
spur. The nectar is contained in a small cup

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at the base of the labellum. The examples

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of the genus Aceras have two retinacula,
or viscidia, sometimes joined together and

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sometimes separate, surrounded by a simple
bursicle. This feature places it between

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Orchis and Ophrys. In Orchis there are two
viscidia enclosed in a bi-lobed bursicle and

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in Ophrys there are two viscidia but they are
enclosed in two separate bursicles.

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Etymology. From –phorum, bearer, and Chromosome number. 2n=42.
anthropo–, man. From the shape of a man on Pollination. Various Coleoptera (beetles)

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the labellum. and some Hymenoptera (flying insects),
Description. 12-28 cm in height. Hairless unspecified.

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cylindrical stem. 5-10 basal leaves, Synonyms. Its closeness to the genus Orchis
unmarked but distinctly veined, oblong- has been confirmed by molecular analysis

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lanceolate, from 5-15 cm × 1-4 cm. techniques (BATEMAN TR.M. et. al. 1997),
Lengthened inflorescence, dense flowering which has led some authors to include it
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towards the tip, from 5-20 cm tall and with within this genus, very close to the group of
between 22 and 60 flowers. Closely packed Orchis militaris, and it has come to be known
sepals forming a hood. Pale green, narrow as Orchis anthropophora (L.) All., a name
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petals under the hood. Labellum greenish- synonymous with its previous name. It has
yellow or orange with darker edges, no spur, also been called Ophrys anthropophora L.
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hanging downwards with three lobes; two Comments. Catalogued as of special


long narrow side lobes. The whole shape is interest.
reminiscent of a hanging man.
Habitat. Full sun or shade. On limestone,
sand or clay. In dry grasslands and together
with thyme, in pine woods of all types, and
oak woods (Portuguese Oak and Holm Oak).
Flowering period. During May and the first
week of June.
Companion Orchids. Shares the ecotope
with other forest species such as
Cephalanthera longifolia, Limodorum p
abortivum and Orchis tenera.
Hybrids. Hybrids are common with species d e
from the Orchis militaris group (O. militaris,
O. simia and O. purpurea).

52 53
Cephalanthera rubra
(L.) Rich.

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Etymology. From rubra-red; the colour of Pollination. In general, the genus Cephalanthera
the flower. attracts only by its appearance. It attracts

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Description. 16.5-50 cm in height. Slender, bees which enter its narrow pathway
wavy stem, grey hairs on the upper portion. between the epichile and the column so that

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Basal leaves reduced to scale leaves, 3-8 the pollen sticks to the thorax, but when
foliage leaves, lanceolate or oblong- the insect departs it is unaware that it has

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lanceolate. Inflorescence lax, of up to 27 not been rewarded, as there is no nectar.
cm, spike with 2-8 flowers, pale pink or deep Synonyms. In the bibliography it appears as
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purple-pink. Labellum white with concave, basionym Serapias rubra L.
white hypochile; epichile lanceolate, white, Comments. It is the only species of the
with pink, wavy edges and small yellow genus in which the pollinia form two
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veins. compact masses that can be easily extracted


Habitat. In shady or semi-shady places. On during flowering, which makes it the only
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limestone soils, rocky or base-rich; damp or species of the genus clearly adapted to
dry. In pinewoods, Portuguese Oak and Holm pollination by another flower (allogamy).
Oak woods.
Flowering period. From the end of May until
the end of June.
Companion Orchids. Species which grow
on impoverished soils which are undergoing
reforestation, such as Cephalanthera longifolia
and Limodorum abortivum. Also forest
species such as Orchis tenera or Orchis
langei.
Hybrids. Hybrids have been recorded with d
Cephalanthera damasonium and Cephalanthera
longifolia, but we have not been able to
confirm this in our findings on site. e p
Chromosome number. 2n=36, 44, 48. Photo: Jolube

62 63
Dactylorhiza fuchsii
(Druce) Soó

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Etymology. The species is attributed to Hybrids. We have found hybrids of this with
Leonhart Fuchs, German professor (1501- Dactylorhiza elata.

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1566). Chromosome number. 2n=40.
Description. Slender plant of 38-48 cm in Pollination. Beetles (Cerambycidae), insects

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height. Thin stem, tinged violet at the top. and bees.
5-11 foliage leaves, keel-shaped and marked, Synonyms. Orchis fuchsii Druce; O.

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shiny grey-green underside and spotted on maculata var. trilobata De Brévisson;
the upper side. Inflorescence compact or Dactylorhiza maculata auct. Non L.;
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lax, conical or cylindrical, of 5.5-7 cm in Dactylorhiza maculata subsp. fuchsii (Druce)
height, with 14 - 30 flowers ranging between Hylander; D. maculata subsp. meyeri (Rchb.
shades of lilac and white. Lateral sepals Fil) Tournay; D. meyeri (Rchb. f.) Aver.
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are raised and tinged with violet. Labellum Comments. The species is very close to
deep three-lobe incisions, the central lobe Dactylorhiza maculata, but D. fuchsii is
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is cupped and decorated with violet stripes, associated with limestone areas, its upper
rings, lines and loops; the lateral lobes are leaves are broader than the lower leaves
wavy at the edges and can hide the divisions and decorated more heavily in the centre
on the lip. Conical spur, 6-10 mm, white or of the lip. In the Sierra de Valdemeca there
pink, horizontal or downward facing. are colonies which show features halfway
Habitat. Shade or semi-shade. On humus- between the two species.
rich soils; damp meadows and hedgerows,
swamps, cool shady woods of Portuguese and It is catalogued as being of special interest.
Pyrenean Oaks and Pines.
Flowering period. During June and into the
beginning of July.
Companion Orchids. In shady wet areas it is d
likely to find Listera ovata nearby. In semi-
shade it can be together with Dactylorhiza
elata and Dactylorhiza incarnata. In damp e p
mountain clearings it can often be found Photo: Jolube
growing together with Orchis champagneuxii.
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Epipactis helleborine
(L.) Crantz

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Etymology. From heleboro, the old name for Companion Orchids. Neottia nidus-avis has
Epipactis and Veratrum album. the same eco-system preferences.

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Description. Greenish stem, 28-45 cm in Hybrids. None are mentioned in Cuenca.
height, upper part hairy. With 4-15 broad, Chromosome Number. 2n=18, 32, 36, 38,40,

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ovate or almost round lower leaves, deep 44.
shiny green, at an angle, hanging down Pollination. By bees, mainly by Vespidae;

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and distinctively veined, 4.5 × 7.5 cm; Halicidae and Apidae are mere visitors
upper leaves smaller and more lanceolate. rather than effective pollinators.
on
Inflorescence a spiked cluster, one-sided, Synonyms. E. latifolia (L.) All.; E. leutei
5.5-19 cm tall, with 20-36 open flowers. Robatsh.
Sepals green on the outer side and lightly Comments. Epipactis helleborine is a rare
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tinged purple or pink on the inside. Petals, plant in our latitudes due to its shade-loving
of a similar size to the sepals, are more nature and the large quantity of organic
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strongly tinged with pink or purple. Labellum matter it requires for development.
greenish purple, epichile with two bosses.
Hypochile deep brownish-grey on inner
surface and green on outer surface. There is
a conical-shaped gland on the central lobe
of the stigma, or rostellum. The ovary is
smooth and club-shaped.
Habitat. Shade, on deep nutrient-rich soils.
On riverbanks, it grows alongside Salix sp.
(Willows) and Populus sp. (Poplars), on valley
floors and dense woods of Quercus faginea,
(Portuguese Oak), Corylus avellana (Hazel) d
and Tilia platyphyllos (Linden / Lime).
Flowering period. In flower during the
second half of June and beginning of July.
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86 87
Himantoglossum hircinum
(L.) Spreng.

Genus Himantoglossum Spreng.


Etymology: from the Greek himas meaning
leather strap or thong and glossa, meaning
tongue, hence ‘strap-tongued’ in reference
to the long, narrow lip.

They are autotrophic plants with two tubers.


A straight, smooth and very tall stem.

s
They have numerous large leaves, broadly

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lanceolate, with no perceptible cross veins.
The smaller lower leaves are usually yellowed

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and withered by flowering time. The flower
spike is tall, with numerous large flowers and

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non-sheathing bracts. The flowers have free
sepals which form a hood. The lip broadens

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at the end, is divided into three lobes, with
a very long and twisted central lobe. Short

w
spur, curved downwards and with no nectar.

w
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Etymology. Hircinum refers to the smell ‘of Hybrids. Hybrids have been described with

at
goats’. the Orchis militaris group, although these
Description. Robust plant, 20-110 cm tall, are extremely rare.

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overall colouring yellowish-green to white. Chromosome number. 2n=18, 24,36.
green stem, sometimes washed with purple- Pollination. The strong smell of its flowers

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violet tinge, with 4-10 oval-lanceolate attracts a wide variety of flying insects,
leaves, the lower leaves 6-25 cm × 2.4 – 6.2 including bees, wasps, flies, bugs, butterflies Photo: Óscar García Cardo
cm, becoming smaller and clasping up the
on
and moths and beetles. More specifically
stem. Herbaceous bracts, up to twice the Cerambycidae, or Long-horned beetles, and
length of the ovary. Flower spike 14-26 cm Scarabaeidae, or Scarab beetles, which may
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tall, very dense, cylindrical and with 20-120 also be potential pollinators.
foul-smelling flowers. Sepals oval or broadly Synonyms. Satyrium hircinum L., Aceras
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lanceolate, 9-13 mm × 4-6.2 mm, pale green hircinum (L.) Lindl.


in colour with some purple blotches. Side Comments. In Cuenca it has been recorded
petals linear, greenish, sometimes notched by Delforge (1989) in the Serranía and more
into three, 8-10 mm × 1-2 mm. Very long lip, recently in the municipality of Torrenjoncillo
41-60 mm × 5-8.8 mm, with a wavy border, del Rey (Aureliano, environmental officer,
three-lobed, central lobe linear and twisted, commented personally). Given that Cuenca
with a deep notch at the tip 2-7 mm long, province is part of its area of distribution
side-lobes linear and pointed, sickle-shaped, and that it chooses a wide range of
shorter than the middle lobe. Spur is curved environments in which to grow, its presence
and cone-shaped, 2-6 mm long. may be more plentiful than these recordings
Habitat. On dry limestone substrates, grass, suggest.
pastureland, sloping banks at the edge of d
pine and oak forests.
Flowering period. From the end of May and
throughout June.
Companion Orchids. Unknown.
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Photo: Javier Benito Ayuso Photo: Jolube

100 101
Ophrys apifera
Huds.

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Etymology. From apis-, bee, and –fera, Flowering Period. End of May and beginning
carry or bear, ‘bee-bearing’, due to the lip’s of June.

at
appearance which attracts bees. Companion orchids. The orchid best suited
Description. 24.5-31.5 cm in height. With to its habitat is Ophrys castellana.

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3-4 oval-lanceolate basal leaves of up to Hybrids. Hybrids with O. scolopax have been
25 cm long. Inflorescence lax, with 2-4 described.

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flowers. Sepals oval-lanceolate, keel-shaped, Chromosome number. 2n=36.
in shades of pink ranging from white to Pollination. Self-pollinating, when the
on
purple; obvious green central veins; held flowers open the slightest movement causes
horizontally; upper sepal bent backwards. the pollen to fall onto the stigma.
Petals very small, leaning forwards, hairy, Synonyms. O. arachnites Millar;
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triangular, with margins rolled back. Lip O. aquisgranensis Kaltenb.; O. ripaensis


three-lobed, with dark crimson-brown Porta; O. botteronii Chodat; O. jurana
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velvety hair covering most of the surface, Neuberger; O. trollii Hegetschw.;


paler at the tip of the centre lobe, which O. frigurgensis (Freyhold) Nägeli; O. bicolor
is rounded, with green edges curled under; Nägeli.
side lobes pointed, conical, short, held Comments. Of this genus it is the orchid
forwards, very hairy, greenish or white, on which best adapts to damp soils. We consider
the outer side; variable speculum, bluish or it to be rare in our province. We have
greyish and edged with yellow; basal area seen samples with white sepals, due to a
light brown U-shaped, spreading to below deficiency of pigmentation (hypochromia).
and merging with or forming separate rings.
Conspicuous appendage, triangular, green,
sometimes divided into three, with a deep
notch which curls up underneath the lip. d
Deep stigmatic cavity, paler than the lip.
Habitat. In full sun or semi-shade, on d e
limestone soils. In grasslands and damp
meadows, in areas of Portuguese Oaks or
Aleppo Pines.
114 115
Ophrys bilunulata
Risso

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Etymology. From bilunata, two moons, Pollination. By flying insects of the genus
referring to the shape of the markings. Andrena (A. nigroaena, A. flavipes) and by

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Description. 8-13 cm in height, with oblong- Colletes cunicularius.
lanceolate leaves. Inflorescence with very Synonyms. Ophrys flavipes-fusca auct.;

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few – 3-6- obvious, dark flowers. Petals Ophrys leucadica Renz.
almost spatula-shaped. Labellum 11.4- Comments. The classification is of western

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13.1 mm × 7.4-9.3 mm, dark brown with Mediterranean distribution. In the province
small white dots, three-lobed, flat and of Cuenca it is highly localised and we can
on
spread out on cross-section, with a fine yet find it in areas with a clear Mediterranean
distinct yellow border around the edge. The influence.
markings on the lip are omega-shaped and
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reach into the mouth of the stigmatic cavity.


Habitat. Full sun or shade. On grasslands
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in pinewood clearings of Pinus halepensis


(Aleppo Pine) in full sun, and in open woods
of Pinus nigra (European Black Pine) with
Rosemary (Rosmarinus officinalis).
Flowering period. This orchid flowers early
for our latitudes: the end of March and April.
Companion Orchids. The colony in
Monteagudo de las Salinas grows together
with Ophrys dyris and Ophrys lupercalis.
Hybrids. Given its flowering period we think
it is possible that hybrids can come from
Ophrys dyris and Ophrys lupercalis, even d
though the pollinators are different.
Chromosome number. Unknown.
d
118 119
Neighbouring species

In this section we have included species which we have not found personally
in the Cuenca province but which have been recorded by other authors. We
also mention species from the neighbouring provinces which may appear
occasionally in ours.

We also provide the bibliographical references of the recordings, the typ-

s
ical habitat of the species, the flowering period and the highest altitude

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at which the plant has been found.

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Dactylorhiza markusii (Tineo) H. Baumann &
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Kunkele
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References to Guadalajara in CARRASCO, MACÍA


& VELAYOS (1997) and to Albacete RIBERA & LÓPEZ
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(1987) under the name of D. sulphurea subsp. sicilien-


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sis (Klinge) Franco. It is of Western Mediterranean


distribution although more Southern than D.insularis. It
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grows on acid substrates in clearings of Pyrenean Oak


woods, at altitudes of up to 2000 m. It is quite rare
on

and in localised areas. It flowers during the second


half of May and beginning of June.
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Photo: Javier Benito Ayuso.


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Epipactis atrorubens Hoffm. ex Besser


References to Guadalajara in CARRASCO, MACÍA
& VELAYOS (1997) and to Teruel: MATEO (1992) and
BENITO & TABUENCA (2001). On the Iberian Peninsula
its distribution is almost exclusive to the Cantabrian
Mountains and the Pyrenees. It grows in any type of
lime substrate, on sandy soils, screes, in meadows and
woods (Poplars and xerophyte pines), at altitudes of
up to 2400 m. It is rare in the Mediterranean area.
It flowers during the second half of July.
Photo: Javier Benito Ayuso.

178
Additional Biology

Orchids and fungi


Fungi are associated with root systems have shown that many orchids require
of more than 90% of all species of plants. a prolonged period of cold for optimum
This relationship, usually symbiotic, in germination to take place.
which the fungus and the plant both

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benefit, is known as mycorrhiza: the Germination can only take place in

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fungus receives sugars produced through the presence of fungi. The fungus
photosynthesis by the host plant and in penetrates the testa of the orchid seed

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exchange provides essential ions, usually and immediately invades the area of the
phosphates and nitrates. embryo. The fungal hyphae penetrate

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the cell wall and form coils and loops

.jo
Curiously, however, the mycorrhiza of called pelotons which, when seen under
orchids is reversed: the host orchid a microscope, show that the mycorrhizal
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feeds off the fungus as its source of process has been successful. These
energy, the process of which is called pelotons are digested by the orchid
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mycotrophic. Many authors believe this host at controlled intervals to avoid


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unusual symbiosis should be considered reinfection, as this would mean it would


as a parasitic process, as the fungus does be deprived of nutrition.
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not seem to benefit from the relationship


but quite the opposite, although it is The seed undergoes a substantial
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never completely destroyed by the change. At first the mycotroph or


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plant. The fact is that the mechanisms protocorm looks like a tiny tuber, an
which control the process of infection undistinguishable mass of cells from
and how the orchid-fungus relationship which soon appears a shoot at the tip
on

is balanced are still not fully understood. which will produce the leaves. When
In fact, orchids produce phytoalexines, exposed to light this shoot will begin
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vegetable hormones which control the to pigmentize and presumably begin


natural aggressiveness of the fungus, to photosynthesise. The lower part
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although it is not known to what degree becomes the organs similar to roots
it may actually be the fungus which which do not pigmentize. From this point
alters its own metabolism once infection onwards the whole plant develops.
by the orchid is produced.
The orchids preserve the mycotroph into
As we have said, orchid seeds are the adult phase, perhaps as an additional
extremely small and have no food mechanism for obtaining energy during
reserve substances. Thousands of them the cold months of dormancy and as a
are released from the mature seedpod nutritional supplement during the growth
and are dispersed great distances (even period. It is quite likely that the orchids
thousands of kilometres) by the wind. have developed this ability so that it can
When the seed falls to the ground it does be used it if the stem parts are eaten by
not germinate straight away. In darkness herbivores.
it can stay dormant for between 2 and 15
years depending on the species and the This heterotrophic capacity is an
soil conditions. Laboratory experiments important factor to consider in
184
Cultivation in pots

First a word of warning: commercial drainage and water retention and helps
cultivation of terrestrial orchids does not the fleshy, fibrous roots to take hold
exist. It is impossible to cultivate the easily.
orchids found among the Cuenca flora in
pots, and it is a criminal offence to take In general they require regular watering
plants from their natural habitat. Maybe for constant moisture, taking care not to
in a few years’ time forced cultivation over-water. Periodically they should be
will enable their commercialisation, but given plant food and microelements.
for now garden enthusiasts will have

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to be content with the orchid varieties

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found in garden centres, which, after THE MOST COMMONLY CULTIVATED

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all, is not bad.
ORCHIDS

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We have been talking about the different
methods of multiplying orchids, and
Genus Cymbidium
because there is a lot of interest,
both personal and industrial, in their .jo
Of Asian origin, it grows in sunny habitats
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from India to Japan and Australia. It
qualities as ornamental plants, we will
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grows very easily and the genus has a
provide some general information on
large number of hybrids, which is why it
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pot cultivation and some indicators on is one of the most cultivated genera.
soil, feeding and watering requirements.
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Internet has many sites on this hobby This orchid has a pseudo bulb at the base
which has recently become so popular of the leaves comprised of aquiferous
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around the world. tissues, and it produces from eight to


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ten leaves and vigorous roots. Some


We’ll begin by saying that there is a varieties begin flowering in November
large number of orchids which, by paying while others flower in February or March.
on

attention to a few specific conditions, The nocturnal temperature should not be


can be cultivated at all latitudes. We higher than 14ºC in summer and in winter
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should select species which can best it should be between 12º and 15º C.
adapt to the predominant climate
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where we live and, when needed,


provide the conditions to make up for
any deficiencies (usually related to
temperature). Commercial exploitation
for cut flowers and pot plants covers
about fifty genera (none of which exist
naturally in Cuenca) and florists have
catalogues with many other hybrids.
The main countries producing orchids are
Brazil, China, Costa Rica, the U.S, the
Philippines, Indonesia, the Low Countries
and Thailand.

The ideal substrate for orchid cultivation


is obtained by mixing pine bark, peat
and perlite, which provides good
192
Evolution

The Orchidaceae family is a recent Of the six stamen in Liliaceae only


addition to plant genealogy, and as three remain in Orchidaceae, and only
a botanical group was distinguished one is fertile. The other two are sterile
not so long ago, when the majority of stamenoids, often appearing as two
ecological niches were taken up. The spots on the wall of the stigmatic cavity.
oldest known orchid fossil was found in
Bodensee (Germany) in strata derived The fact that orchids are in their early
from sedimentation, at the bottom of stages of evolution means that many
a shallow lake. A flower can clearly be of their peculiar traits are still to be

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seen, with three sepals, two petals and determined; scientists believe this is why

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the lip, and a long inferior ovary, dating orchids have taken on such interesting

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from the Late Miocene period, in other evolutionary adaptations: they are
words about 15 million years ago when opportunistic, they produce millions of

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Europe had a tropical climate. seeds, they need mycorrhizal fungi to
germinate and they use unique methods
Nevertheless, later DNA studies have
suggested that the Orchidaceae family .jo
to ensure pollination.
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could be much older than was originally Depending on the species, each plant
w
thought and that its origins could go can produce from two (eg. Ophrys) to
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back to the end of the Cretaceous period almost one hundred (some Gymnadenia
of circa 65 million years ago. and Dactylorhiza) mature fertilised
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ovaries, each containing over a million


Be that as it may, it is still a group of microscopic seeds.
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recent development compared with the


monocotyledons which appeared 100
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It should be noted that the evolutionary


million years ago, or the dicotyledons success of orchids is directly related to
of 150 million years ago, or conifers, their capacity to associate with different
on

thought to age from 300 million years ecosystems: on the one hand that of
ago. pollinating insects, with which their
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relationship is closer and more complex


All researchers coincide in that orchids than any other plant’s, and on the other
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have probably evolved from the hand, with the mycorrhiza, on which
botanical family of Liliaceae. They their growth depends to the extent that
preserve the 3-part structure of three without this relationship germination
petals and three sepals. The flower’s is compromised. Other species also
adaptation to pollinating insects is maintain this type of dependence, but as
the hypothesis that explains the slight a collaborator, not as a trigger. This close
modification to one of the three petals relationship, so vital for evolutionary
that we know as the lip, or labellum, success, reminds us of Nobel candidate
which means the flower loses its Lynn Margulis’ theory of evolutionary
actinomorphic, or radial, symmetry in symbiosis.
favour of a zygomorphic, or bilateral,
symmetry such as we humans have: one As orchids only produce two pollinia
half (right) is a mirror image of the other masses, rather than large quantities of
(left). pollen grains as most other plants do,
this has led to other adaptations for
fertilisation to be guaranteed. Of course,
194
Pollination methods

Orchids are probably the plants best its systematic position, although this is
adapted to entomophily (pollination not always the case. In general, various
by insects), and because of this there species of orchid can be visited by the
is a wide variety of mechanisms for same pollinator species and produce
attracting insects within the orchid hybrids, which, in turn, may attract a
family. They are pollinated by several new specific pollinator and may produce
species of Hymenoptera (bees, wasps, a new isolating barrier. This evolutionary
ants), also by Diptera (flies), Lepidoptera strategy is responsible, for example,
(butterflies and moths), Coleoptera for the huge propagation through

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(beetles) and, more rarely, by slugs and

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adaptation of the genus Ophrys within
snails. the Mediterranean area.

be
The components of the genus All of this gives us some idea of the

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Plantanthera create a long spur full of intimate relationship between the

.jo
nectar which can only be reached by method of fertilisation and evolutionary
certain species of moths equipped with development of the Orchidaceae family.
w
long proboscides. At the other extreme,
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in some species of the genus Epipactis, It is assumed that the great diversity
the nectar is easily accessible and a wide of orchid species is largely due to its
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variety of insects can access it. capacity to adapt to pollinators, and


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therefore that these pollinators have


Cephalanthera longifolia has flowers driven the differences in flowers and
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similar to those of Cistus salviifolius species. This has generated much


(as seen by insects), and attracts them research which both supports and
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without needing to give up the pollen refutes the theory.


normally given by the rock rose.
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The perianth segments of the genus


Serapias form a narrow tube which
THE METHODS USED TO ATTRACT
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provides a refuge for particular bees


at night or in inclement weather POLLINATORS
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conditions.
Nectar
Some species of the genus Ophrys are In some plants the nectar gives off a
able to imitate the female members strong scent and is usually found inside
of the insects which pollinate them, the bottom of the spur, such as in
and can also synthesise substances that Gymnadenia, Anacamptis, Platanthera
imitate the female sexual hormones, and some Orchis. In Epipactis and
thereby confusing the male insect who Listera, it is found in grooves on the surface
‘copulates’ with the lip of the orchid. and although it has no perceptible smell
Apparently, Ophrys speculum can only for humans, evidently insects find it very
be pollinated by one species of insect: attractive. The nectar prize that the
flower offers the pollinating insect is the
Campsoscolia ciliata. Due to all this
orchids’ most successful tool for achieving
specialisation, sometimes knowing
fertilisation.
an orchid’s pollinator can provide us
with valuable information regarding
196
Hybrids

Due also to their relatively recent emergence, Orchids form the group of plants
with the greatest number of hybrids, that is to say, descendants from fertilisation
with a different plant species. Horticulturalists and enthusiasts in orchid cultivation
have exploited this fully and forced crossings between different genera and species,
bringing about over 150,000 different cultivars (the CITES register records over one
thousand new cultivars per year).

The hybrid is usually sterile and remains isolated, but exceptionally they can cross-
fertilize with one, or both, of their original parent plants, or even reproduce among

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themselves. This creates hybridogenetic colonies which are difficult to identify, as

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the offspring have features of both parents. This is an important detail and should

be
be helpful for us to distinguish between a hybrid and an abnormal plant which
simply shows a variation of the species. One essential detail to observe is whether
the hybrid plant is growing among colonies of its supposed parent plants. The hybrid

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plant should have not just one but many intermediate features. Occasionally the

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hybrid can adapt more easily to unfavourable conditions than the parent plants and
appear during a year when they are absent, although this is uncommon and should be
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confirmed through repeated observations.
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When the crossing is produced between plants of the same genus but of different
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species, this is called interspecific, or infrageneric; when it takes place between


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species belonging to another genus it is called intergeneric.


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Intergeneric hybridisation is limited to the genera Dactylorhiza, Coeloglossum,


Platanthera, Gymnadenia, Nigritella and Pseudorchis, on the one hand, and to the
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genera Orchis, Neotinea, Aceras, Anacamptis and Serapias, on the other.


on

Ophrys, however, rarely produces interspecific hybrids. The possibility of producing


this phenomena in nature is quite remote; the cases that have been compared
(Dactylorhiza × Gymnadenia, Orchis × Dactylorhiza, Gymnadenia × Nigritella,
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Coeloglossum × Dactylorhiza, among others) have served as a basis for genetic


analysis giving rise to a new focus for understanding the development of the
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Orchidaceae family, and without doubt, will revolutionise the systematics of orchids
in the future.

202
Orchid protection

The orchid family has a very flora and fauna. Three species of orchid
cosmopolitan distribution and in are included, but within its protection
many cases new species are capable a range of habitats are covered that
of colonising altered substrates. constitute the ideal environment for the
Nevertheless, we should not be fooled growth of threatened colonies of orchids
by this ability to adapt. As in all wildlife, in the Iberian System: sclerophyllus
orchids are subject to a range of forests of pastureland (dehesa),
adversities which make it hard for their moorland, Mediterranean pastures of tall
biological rhythms to keep ahead of the grasses and reeds, wetlands, calcareous

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changes brought about by man. peatbogs, semi-natural dry grassland

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formations and shrubland on calcareous

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Uncontrolled urbanization, road soils (Festuco brometalia).
building, intensive agriculture etc. all
progressively reduce or alter the natural CITES (Convention on International

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environment and continuously damage Trade in Endangered Species of Wild
biodiversity.
.jo
Fauna and Flora) is an international
treaty whose aim is to ensure that
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Due to their particular biological cycle, international trade in specimens of wild
w
orchids are sometimes subjected to animals and plants does not threaten the
other pressures which make them survival of the species. Member states
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more vulnerable to actions by man. who adhere to CITES have to apply the
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For example the indiscriminate use norms of the convention although these
of nitrate fertilizers prevents the do not override national legislation. Each
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development of the mycorrhizal fungi, country must pass its own legislation in
pesticides significantly reduce colonies order to guarantee that CITES is applied
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of potential pollinators and the mass across the nation. 63 species of Spanish
picking of attractive species have orchids are included in the Convention.
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brought some orchids to the point of


extinction (as in the case of Cypripedium At our regional level, the Decree
calceolus). In addition to all these 199/2001, dated 6 November 2001,
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threats is the fact that traditional adds to the Regional Catalogue of


farming methods have been abandoned; Threatened Species in Castilla-La
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the seasonal moving of livestock, the Mancha (CREA) dated 5 May 1998.
mowing of fields for feeding livestock This extension of CREA has allowed
and clearing the forests for collecting the inclusion of 20 species of the
firewood were all advantageous Orchidaceae family (the earlier version
practices for orchids when compared to only included one species) which are
their closest competitors. protected under different sections
depending on the frailty of their colonies
There are international, national and in the region. From greater to lesser
regional plans for protection which can degree of frailty there are the following
help to reduce the negative impact of categories: ‘in danger of extinction’,
human activities on orchid colonies. ‘sensitive to changes in their habitat’,
‘vulnerable’ and ‘of special interest’.
The European Council directive 92/43/ The criteria used for determining the
CEE, dated 21 May 1992 relates to the inclusion of orchid especies in each
conservation of the natural habitats of category are as follows:
205
Some interesting facts

A BIT OF HISTORY them for a mixture of sugars and nutritious


minerals that the fungi normally provide.
As is the case with so many other matters, it Thus commenced the expansion of forced
was the Chinese who were the first to leave cultivation.
written records on their impressions and
knowledge of orchids. Chinese records with The first artificial orchid to flower was in
references to orchids go back 1500 years, 1856, the Calanthe Dominyi (furcate ×
although the first monographic record dealing masuca), a cross achieved by John Dominy.
exclusively with orchids was written by Chao
Shih-Keng (1233), a botanist with a passion The first hybrids were intrageneric, and so

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for this family who wrote a treaty for the maintained their genus name, to which was

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orchids of Fukien, giving all sorts of details on added a new, un-italicised name, as seen
methods for their cultivation. in this example of Calanthe Dominyi. New

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intergeneric hybrid crosses produced names
In the west, its discovery as a valuable like Catamodes (Catasetum × Mormodes),

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ornamental flower took place at the turn of Laeliocattleya (Laelia × Cattleya), etc.
the 19th Century, when by chance the first
plants of Cattleya labiata arrived in Europe.
During this period, Victorian England had .jo
The desire for new species led to further
complexities – the hybrids themselves were
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explorers spread all over the planet. In crossed with other species and produced
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1818, Swainson, who was in South America hybrids such as Brassiolaeliocattleya
collecting mosses and lichens for William (Brassavola × Laeliocattleya), which not
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Cattley, a British horticulturalist, used the surprisingly became abbreviated to BLC. When
vines of some plants to wrap the moss in, even more hybrid genera were introduced it
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without realising that he was in fact using became necessary to create a methodology
orchids from the Atlantic Forests of Brazil. exclusively for naming orchids.
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These vines, or lianas, arrived intact and alive


and flowered in England that same year. The complete register of orchid hybrids is
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held by the Royal Horticultural Society (UK)


Cattley was intrigued by their beauty and and can be accessed on Internet.
on

he took them to John Lindley, an eminent


horticulturalist of the time, who named them USES
as a genus and provided a description. As
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their lip was particularly outstanding, Lindley Aromatics


included the word ‘labiata’ in the name.
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It is no less anecdotal but somewhat more You may be surprised to learn that when you
relevant to know that thirty years previously, eat ice-creams and desserts you are actually
Hipólito Ruiz and José Pavón, Spanish eating seeds from orchids. Vanilla is extracted
botanists who had been sent to America by from the pods of orchids from the genus
the King of Spain, Carlos III, described more Vainilla. Of course, the etymological root of
than 600 orchids in Peru during an exploratory the term comes from the Spanish ‘vainilla’
journey which lasted eleven years (1777- which actually means ‘small pod’. Vainilla
1788). fragrans is an epiphytic plant, a long, twining,
creeping liana from the forests in the east
At the beginning of the 20th Century the of Mexico where it has been cultivated since
Frenchman Noel Bernard discovered the the time of the pre-Columbian Mayans and
mycorrhizal phenomenon. At around the Aztecs. They named it tlilxochiti and used it
same time, the German, Hans Burgeff, to flavour their favourite drink: chocolate.
demonstrated that orchid seeds could
germinate in agar with the mycorrhizal fungus In Peru it has been used since antiquity
in the lab. And in 1922, the North American, to flavour tobacco, and also in food and
Lewis Knudson got orchid seeds to germinate perfumery.
in agar without the fungi, substituting
207
Locations

In order to locate the species more easily they have been arranged in alphabetical
order. The recorded sitings are organised from east to west using the UTM coordinate
system. This is followed by the name of the municipality and the place name of the
location where the specimen was found. When kilometres are included, these refer
to the distance between the reference and the first place mentioned; the second
place specifies the road which joins the two. This information is followed by the
altitude taken from the map, then the type of habitat and finally the date on which
the find was recorded. This always coincides with when the plant is found in full
flower, unless otherwise indicated. The spanish name for the type of habitat has been
retained, for which a Spanish-English lexicon has been provided at the end of the chapter.

s
.e
Aceras anthropophorum

be
1. 30TWK5488 Alcantud. Dehesa de Alcantud. 800 m. Quejigar. 1-5-02.

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2. 30TWK5633 Cuenca. Puerto de Cabrejas. 1160 m. Quejigar. 23-5-99.
3. 30TWK6040 Navalón. Pinar al sur del pueblo. 1050 m. Bosque de P. nigra. 10-6-99.

.jo
4. 30TWK6579 Cañamares. Pto. Monsaete. 900 m. Bosque de P. nigra. 30-4-99.
5. 30TWK6580 Cañamares. Pto. Monsaete. 950 m. Pinar de P. pinaster. 22-6-99 (fruit).
w
6. 30TWK7389 Beteta. Paseo Botánico. 1110 m. Bosque mixto. 20-5-00.
w
7. 30TWK7491 Beteta. Fuente de la Carrera. 1110 m. Bosque mixto. 12-6-99.
8. 30TWK7660 Zarzuela. Dehesa Boyal. 1200 m. Bosque mixto. 1-6-00.
w

9. 30SWK8316 Almodóvar del Pinar. N-320. Km. 116. 1050 m. Quejigar. 18-5-00.
10. 30TWK8332 Cuenca. Hoz de San Miguel. 1200 m. Encinar en el calar. 24-5-00.
at

11. 30TWK8378 Cañizares. Huerta de Marojales. 1360 m. Pinar de P. sylvestris. 12-6-99.


12. 30TWK8467 Las Majadas. Cobacha del agua. 1440 m. Tomillar. 7-6-02.
le

13. 30SWK8707 Almodóvar del Pinar. Arroyo Fte. del Pocico. 1050 m. Bosque mixto. 25-5-99.
14. 30TWK8730 Cuenca. Loma de las Yeguas. 1230 m. Pinar de P. nigra. 8-5-99.
sa

15. 30TWK8831 Cuenca. Torca del Agua y Torca del Lobo. 1240 m. Pinar de P. nigra. 8-5-99.
16. 30TWK9378 Cuenca. Los Sabinarejos. 1500 m. Pinar de P. sylvestris. 24-6-00 (fruit).
on

17. 30TWK9476 Cuenca. Nacimiento Río Cuervo. 1500 m. Pinar de Pinus sylvestris. 4-5-02.
18. 30TXK0548 Valdemeca. Fte Avellaneda. 1680 m. Pinar de P. sylvestris. 19-6-99 (fruit).
19. 30SXK4408 Talayuelas. Ladera N. del Pico Ranera. 1080 m. Pinar de P .pinaster. 28-5-
ok

20. 30SWJ7779 Alarcón. Casas del Embalse de Alarcón. 810 m. Coscojar. 22-4-2006.
Bo

Anacamptis pyramidalis
1. 30TWK6293 El Pozuelo. Camino al refugio de El Pozuelo. 1010 m. Aliagar. 15-6-99.
2. 30TWK6480 Cañamares. Cima del Pto. Monsaete. 960 m. 30-4-99.
3. 30TWK6580 Cañamares. Pto. Monsaete. 950 m. Pinar denso de P. pinaster. 22-6-99.
4. 30TWK7179 Fuertescusa. Ctra. Fuertescusa-Poyatos. 1100 m. Juncal con Salix sp. 8-6-02
5. 30TWK7343 Cuenca. El Chantre. 970 m. 25-5-99.
6. 30TWK7379 Fuertescusa. Fuente de la Dehesa. 1100 m. Pinar. 10-6-00.
7. 30TWK7439 Cuenca. Fuente de Martín Alhaja. 960 m. Chopera. 1-6-99.
8. 30TWK7491 Beteta. Fuente de la Carrera. 1110 m. Bosque mixto. 12-6-99.
9. 30TWK7577 Fuertescusa. Fuente del Mostajo. 1100 m. Pendiente rezumante. 16-6-00.
10. 30TWK7660 Portilla. Dehesa Boyal. 1200 m. Bosque mixto. 1-6-00.
11. 30TWK8085 Santa Mª del Val. Pantano de la Tosca. 1200 m. Arenas albenses. 22-6-99.
12. 30TWK8141 Buenache. El Rollo. 1000 m. Carrizal. 10-7-02
13. 30TWK8176 Poyatos. Cima de El Cuerno. 1490 m. Aliagar. 10-6-00.
14. 30TWK8245 Cuenca. Arrollo Vegalindo. 1090 m. Aliagar. 29-5-98.
15. 30TWK8245 Cuenca. Ladera este del Marimorena. Herbazal rezumante. 1150 m. 2-6-02.

209
Spanish-English Lexicon of Habitats

HABITATS Pasto: pasture.


Pedregal: scree.
Alameda: Poplar grove. Pendiente: slope.
Aliagar: Genista scorpius shrubs. Pendiente rezumante: Seepage slope.
Arcilla: clay. Pinar: Pine forest.
Arenas albenses: Albian sands. Quejigar: Portuguese Oak forest.
Arroyo: stream Regueros: irrigation ditches.
Barranco: ravine. Rendzina: type of limestone soil.
Borde de camino: wayside. Rezumadero: swamp.
Bosque mixto: mixed woodland. Ribera: Riverbank.

s
Brezal: Heathland / Moor. Rodenal: Pinus pinaster forest.

.e
Bujeda: box forest. Romeral: Rosemary shrubbery.

be
Calar: Calcareous, chalky. Roquedo: crag.
Caliza: Limestone. Sabinar: Juniper forest.

lu
Cambronal: thomy schrubland. Sauceda: Willow forest.
Canchal: Rocky soils, scree. Talud: slope.
Cantil: ledge.
Carrizal: Reedbed. .jo
Terraza: terrace.
Toba:petrified lime deposits
w
Cauce seco: dry river bed. Tomillar: Thyme field.
w
Chopera: Poplars. Turbera: peatbog.
Claro: clearing. Vaguada: stream bed.
w

Coscojar: Kermes Oak grove. Yeso: Gypsum soil.


Cuneta: ditch.
at

Dehesa: Sclerophyllous scrub.


Derrubio: scree / debris. TYPES
le

Encinar: Holm Oak forest.


Garriga: Garrigue. Adehesado: turned to scrub.
sa

Guijarral: Stony terrain. Arenoso: sandy.


Herbazal: meadow. Despejado: open, clear.
on

Jaral: Rock rose thicket. Encharcado: swamped.


Juncal: Reed bed. Húmedo: damp.
Ladera: hillside. Inundado: flooded.
ok

Lastonar: grassland. Mixto: mixed.


Marga: Marl (Lime-rich mudstone). Pedregosa: rocky.
Bo

Matorral: shrubland. Rezumante: seeping.


Melojar: Pyrenean Oak forest. Salino: salty.
Olivar: olive grove Umbroso: shady.
Orilla: bank border. Yesosa: chalky.
Pastizal: grassland.

232
Glossary of terms

Albian sands: The Albian is an age of the Ecotope: the smallest ecologically distinct
geological timescale which corresponds to landscape feature in a landscape mapping
the youngest or uppermost subdivision of the and classification system.
Early/Lower Cretaceous epoch/series. Its Entire: whole; not toothed, nor lobed or
approximate time range is 113.0 Ma to 100.5 divided in any way.
Ma (million years ago).
Epichile: The apex portion of the labellum,
Amplexicaul: clasping the stem but not enti- often heart-shaped.
rely circling it.
Epeirogenetic: type of movement of the
Anther: the pollen-bearing part of a stamen. earth’s crust affecting large areas of land or
Autogamy: the process of self-pollination. ocean bottom.

s
Basionym: the original, validly published Esparto: tough wiry grasses.

.e
name of the taxon. Exogenous: originating externally
Biotype: a group of genetically identical

be
Filiform: Very narrow with straight and
plants within a species. parallel margins; thread-like.
Bract: a leaf-like structure which has no

lu
Fusiform: spindle-shaped; widest in the
blade or lamina. middle and tapered at each end.
Bursicle: A membranous sheath which covers
the glue in some viscidia. .jo
Foliaceous: Leaf-like.
w
Garrigues: shrubs typically found on calcareous
Callus: a raised, fleshy structure found on Mediterranean soils, usually aromatic shrubs
w
the labellum. eg, thyme, rosemary, lavender etc.
w

Calyx: the outer part of the flower, the se- Gibbous: With a pouch-like swelling; humped.
pals.
Glabrous: Without hairs.
at

Cambrones: common name for thorny shrubs


which grow in the shape of a cushion. Examples Glaucous: covered with a bloom which lends
a bluish lustre.
le

In Cuenca are Erinacea anthyllis and Genista


pumilla. Humic: derived from humus.
sa

Caudicle: a term used for a pollinium stalk Hymenoptera: flying insects such as bees,
derived from the anther; an elastic basal wasps and ants.
on

extension of the pollinia. Hypochile: The inner portion of the labellum,


Cauline: belonging to a stem, usually refers often cup-shaped.
to leaves. Inflorescence: The flowering structure of a
ok

Ciliate: with a fringe of fine hairs. plant.


Circumboreal: relating to the northern Labellum: a lip - in orchids it is a highly
Bo

portion of the Northern Hemisphere. modified petal that is primarily involved in


Cladograms: maps of genetic similarities pollination.
between species. Lanceolate: lance-shaped; narrowly oval,
Claviform: club-shaped, clavate. longer than wide and tapering at each end,
especially the apex.
Cleistogamy: the process of self-pollination
occurring without the flowers opening. Lax: loose, not dense.
Clinandrium: The top margins of the column Meristem: plant tissue from which new cells
or the cavity below the anther and behind are formed, e.g. the tips of roots or stems;
the stigmatic cavity, in which the pollinia lie. the growing tip.
Corolla: a group of petals, the inner whorl of Mesomediterranean: sub-montane zone, 600
the perianth. – 900 m altitude.
Cuneiform: wedge-shaped. Mycorrhiza: A beneficial relationship between
the roots of a vascular plant and fungi resulting
Decurved: curved downwards. in nutrient exchange.
Dichlamydeous: flowers which have a corolla
and calyx.
233
Mycotrophic: Plant that obtains some or all Saprophyte: An almost leafless plant lacking
of its nutrition through mycorrhizal fungi; chlorophyll that derives sustenance from
another term for saprophytic. decaying wood or other plant parts, in
Nectariferous: Bearing nectar-secreting association with a symbiotic fungus.
glands. Sessile: Without a stalk, pedicel or petiole.
Ombrotrophic: typically rainy climate. Speculum: a pattern on the labellum, often
Oromediterranean: montane zone, 1200 – with a shiny metallic lustre, of the bee and
1900 m altitude. spider orchids (genus Ophrys).
Ovule: The small structure within the ovary Spike: A simple unbranched inflorescence

s
which becomes a seed after fertilization. with sessile flowers.

.e
Palmate: divided like a hand. Spur: A slender hollow projection from a
floral segment (usually the labellum, rarely

be
Pedicel: The stem which supports a single the dorsal sepal).
flower in an inflorescence.
Stamens: male reproductive organs of a

lu
peduncle: The main axis of a compound flowering plant.

.jo
inflorescence or the stalk of a solitary flower
which subtends the pedicel. Staminode: a sterile stamen
Stigma: (pl. stigmata) The enlarged sticky
w
Pedunculate: Having a peduncle.
area which terminates the pistil and is
Perianth: A collective term for the petals and
w
receptive to pollen and allows the pollen
sepals of a flower; in orchids this does not grains to germinate.
w

include the labellum.


Supramediterranean: mid-montane zone,
Petiole: the stalk of a leaf.
at

900 - 1200 m altitude.


Polje: a large elliptical depression in limestone Symbiosis: a close relationship between two
regions, sometimes containing a marsh or
le

or more organisms, including parasitism.


small lake.
Tepal: in Orchidaceae, this term refers to
sa

Pollinium: (pl. pollinia) An aggregated any sepal or petal, not the labellum.
coherent mass of pollen grains found in the
Orchidaceae. Utrillas sands: Albian sands found in the area
on

of Utrillas, in the province of Teruel.


Protocorm: Specialised structure that develops
after orchid seed germination and from which Viscidium (pl. viscidia): A clearly defined
sticky part of the rostellum which is removed
ok

a shoot develops.
together with the pollinia as a unit by a visi-
Rhizome: An underground stem with nodes ting insect.
Bo

and roots, which can form shoots.


Xerophyte: a plant adapted to dry growing
Rostellum: The area of tissue that separates conditions.
the stigma from the anther; an adhesive portion
of the stigma which aids pollen transfer.

234
Useful links

Orchid plants can claim to have the greatest number of enthusiasts and collectors in
the world and a large number of clubs and societies exist as a result.

What follows are several different types of lists which provide ample proof of their
popularity and which may be of use to the interested reader.

Associations with websites Five Cities Orchid Society, Arroyo Grande, CA


Aberdeen Branch, Scottish Orchid Society, UK Florida North Central (AOS) Judging Center, Tampa, FL
Akashi Orchid Society, Japón Ft. Lauderdale Orchid Society, Ft. Lauderdale, FL
Alamo Judging Center, San Antonio, TX Fort Worth Orchid Society, Fort Worth, TX
All Japan Orchid Society Gainesville Orchid Society, Gainesville, FL

s
American Orchid Society Gold Coast Unlimited Orchid Society, North Miami, FL
Amherst Orchid Society, Amherst, MA Grand Valley Orchid Society, Walker, MI

.e
Ann Arbor Orchid Society, Ann Arbor, MI The Greater Cleveland Orchid Society, Cleveland, OH
ANOS Illawarra, Wollongong, Australia Greater North Texas Orchid Society, Dallas, TX

be
Asociación de Orquideología de Quito, Ecuador Greater Omaha Orchid Society, Omaha, NE
Associacao Orquidofila de Divinopolis, Divinopolis, MG, Brasil Groupement Midi-Pyrénées des Amateurs d’Orchidées, Francia

lu
Associació Catalana d’Amics de les Orquídies, España Heart O’ Texas Orchid Society, Austin,TX
Association des Naturalistes de l’Ariège The (UK) Hardy Orchid Society

.jo
Associazione Lombarda Amatori Orchidee (ALAO), Hill Country Orchid Society, New Braunfels, TX
Varese, Italia Himeji Orchid Society, Japón
Associazione Triveneta Amatori Orchidee (ATAO), Firenze, Italia Houston Judging Center (Texas)
w
Atlanta Orchid Society, Atlanta, GA Houston Orchid Society, Houston, TX
Australasian Native Orchid Society Australia Illinois Orchid Society, Chicago, IL
w
Australasian Native Orchid Society, Victoria Group Australia Illowa Orchid Society, Davenport, IA
w

Bankstown Orchid Society, Bankstown, Sydney, NSW, Australia Iowa City Orchid Society, Iowa City, IA
Blue Ridge Orchid Society, VA International Phalaenopsis Alliance
The Boca Raton Orchid Society, Boca Raton, FL Jupiter/Tequesta Orchid Society, Jupiter, FL
at

Bonsai and Orchid Association International Kansas Orchid Society, Wichita, KS


Bournemouth Orchid Society, Bournemouth, UK Lehigh Valley Orchid Society, Allentown, PA
British Orchid Council, UK Les orchidophiles réunis de Belgique, Bélgica
le

British Orchid Growers Association, UK Les Orchiophiles de Montreal, Montreal, PQ, Canadá
sa

British Paphiopedilum Society, UK Lincoln Orchid Society, Lincoln, NE


Canadian Orchid Congress, Canadá. Malihini Orchid Society, Cupertino, CA
Cape and Islands Orchid Society, Cape Cod, MA Manitoba Orchid Society, Winnipeg, MB, Canadá
Cape Fear Orchid Society, Wilmington, NC Maryland Orchid Society, Baltimore, MD
on

Capital City Orchid Society, Wellington, New Zealand Massachusetts Orchid Society, Boston Area
Carmel Orchid Society, Monterey, CA The Maxillaria Tribe
Catoctin Orchid Society, Frederick, MD Michigan Orchid Society, Greater Detroit Area, MI
ok

Central East Texas Orchid Society, Tyler, TX Mobile Area Orchid Society, Mobile, AL
Central Iowa Orchid Society, Des Moines, IA MoriokaOrchid Society, Japón
Central Ohio Orchid Society, Columbus, OH Mt. Baker Orchid Society, Mt. Vernon, WA
Bo

Central Ontario Orchid Society, Waterloo, ON, Canadá Na Okika O Hawaii


Central Vancouver Island Orchid Society, Nanaimo, BC, Canadá National Capital Orchid Society, Washington, D.C. Area
Cincinnati Judging Center, Cincinnati, OH Native Orchid Conservation Inc., Manitoba, Canadá
Círculo Orquidófilo de Juiz de Fora, Juiz de Fora, MG, Brasil De Nederlandse Orchideeën Vereniging /The Dutch
Círculo Orquidófilo Regional de Timbó, Timbó, SC, Brasil Orchid Society, Países bajos.
Círculo Paulista de Orquidófilos, São Paulo, SP, Brasil Werkgroep Masdevallia
Club Amigos de las Orquídeas, Madrid, España New Hampshire Orchid Society, Bedford, NH
Commercial Orchid Growers Guild New Orleans Orchid Society, New Orleans, LA
Connecticut Orchid Society, Rocky Hill, CT The New Zealand Native Orchid Society
Cymbidium Society of America Norsk Orkideforening (NOF)/The Norwegian Orchid Soc.
Dallas Judging Center (Texas), and Society Information North American Native Orchid Alliance
Deep Cut Orchid Society, Colts Neck, NJ North American Regional Orchid Specialist Group
Delaware Orchid Society, Wilmington, DE North Jersey Orchid Society, East Hanovoer, NJ
Delray Beach Orchid Society, Delray Beach, FL North of England Orchid Society
Desert Valley Orchid Society, Phoenix, AZ North Shore Orchid Society, Sydney, Australia
Deutsche Orchideen-Gesellschaft Northeastern Wisconsin Orchid Society, Green Bay, WI
Diablo View Orchid Society, East San Francisco Bay Area, CA Northwest Orchid Society, Seattle, WA
East Everglades Orchid Society, Homestead, FL The Odontoglossum Alliance
Eastbourne Orchid Society, Sussex, United Kingdom Oklahoma Orchid Society, Oklahoma City, OK
Eastern Canada Orchid Society, Montreal, PQ, Canadá Orchid Badge Club International
The Eastern Orchid Congress The Orchid Digest
235
Bibliography

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s
Collectianea Botanica 12(1): 5-61.

.e
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reclassification to achieve monophyly of Orchis sensu stricto. Lindleyana 12(3): 113-141.
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co. Catalogación de la especies amenazadas. Zubía 14: 129-131.
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Arvet-Touvet en el Sistema Ibérico. Flora Montiberica 8, 55-60.
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w
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w

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sa

BENITO, J., ALEJANDRE, J.A. & ARIZALETA, J.A.(1999). Aproximación al catálogo de las
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on

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ok

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British Library Cataloguing in Publication Data.
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Madrid 2: 236-265.

238
Index of Scientific names

Names in bold refer to the accepted names for the species detailed in this guide.
Aceras anthropophorum (L.) W. T. Aiton (52) Dactylorhiza gemmana 72
24, 33, 42, 47, 48, 52, 108, 152, 199, 206, 209 Dactylorhyza incarnata (L.) Soó (72), 16, 31,
Aceras densiflora 108 36, 42, 44, 47, 68, 70, 76, 78, 206, 215
Aceras intacta 108 — f. ochrantha 72
Aceras hircinum 100 — var. altísima 72
Anacamptis 42, 51, 142, 172, 196, 202 — var. drudei 72
Anacamptis brachystachys 54 — var. hyphaemetodes 72
— var. tanayensis 54 — var. lobelii 72
Anacamptis champagneuxii 146 — var. reichenbachii 72
Anacamptis fragrans 148 Dactylorhiza insularis (Sommier) Ó. Sánchez

s
Anacamptis palustris 154 & Herrero (74) 16, 28, 47, 146, 206, 215

.e
Anacamptis papilionacea 156 — f. bartonii 74

be
Anacamptis pyramidalis (L.) Rich. (54) 25, 28, Dactylorhiza maculata (L.) Soó (76) 26, 37, 47,
48, 90, 132, 136, 148, 204, 209 70, 194, 203, 206, 216

lu
— var. brachystachys 54 — subsp. arduennensis 76
Anacamptis tanayensis 54 — subsp. fuchsii 70
Anacamptis urvilleana 54
Anteriorchis coriophora subsp. fragrans 148 .jo
— subsp. meyeri 70
Dactylorhiza markusii (Tin.) H. Baumann &
w
Apostasiodeae 38 Kunkele (178)
w
Argorytes mystaceus 198 Dactylorhiza meyeri 70
Cattleya 193, 208 Dactylorhiza munbyana 68
w

Cephalanthera 38, 40, 43, 50, (56), 62, 197, 239 Dactylorhiza romana subsp. bartonii 74
Cephalanthera alba 58 Dactylorhiza sambucina (L.) Soó (78) 26, 36,
at

Cephalanthera damasonium (Mill.) Druce (58) 47, 198, 206, 216


25, 30, 44, 47, 48, 60, 62, 88, 102, 210 — subsp. insularis 74
le

Cephalanthera ensifolia 60 Dactylorhiza sesquipedalis 68


Cephalanthera grandiflora 58 Dactylorhiza viridis 64
sa

Cephalanthera longifolia (L.) Fritsch (60) 24, Epidendroideae 38


25, 26, 28, 33, 47, 48, 52, 58, 62, 90, 96, 102, Epipactis (80) 17, 38, 40, 41, 42, 43, 50, 66, 196,
on

104, 108, 196, 197, 211 234, 235


Cephalanthera pallens 58 Epipactis atrorubens Hoffm. ex Besser (178)
Cephalanthera rubra (L.) Rich. (62) 25, 27, 28, — var. tremolsii 96
ok

29, 35, 47, 48, 60, 96, 102, 212 Epipactis campeadorii 88
Cephalanthera xyphophylla 60 Epipactis cardina Benito & C.E. Hermos. (82)
Bo

Ceratobasidium 185 17, 29, 47, 48, 84, 90, 217


Coeloglossum 40, 54, 202 Epipactis distans Arv.-Touv. (84) 29, 47, 48,
Coeloglossum alpinum 64 82, 206, 216
Coeloglossum viride (L.) Hartm. (64) 29, 31, Epipactis fageticola 179
36, 47, 48, 72, 206, 213 Epipactis helleborine (L.) Crantz (86) 30, 31,
Composcolia ciliata 196 34, 47, 48, 216
Cymbidium 188, 192, 244, 246, 248 — subsp. distans 84
Cypripedioidae 38, 248 — subsp. tremolsii 96
Cypripedium calceolus 205 Epipactis hispanica Benito & C.E. Hermos.
Dactylorhiza 40, 42, 51, (66), 98, 172, 194, 198, 202 (88) 30, 31, 47, 48, 216
Dactylorhiza elata (Poir.) Soó (68) 30, 36, 37, Epipactis kleinii M.B. Crespo, M.R. Lowe & Pie-
47, 70, 72, 76, 94, 148, 154, 168, 174, 203, ra (90) 17, 25, 47, 48, 58, 60, 82, 84, 96, 217
204, 206, 213 Epipactis latifolia 86
— var. corsica 68 Epipactis leutei 86
Dactylorhiza fuchsii (Druce) Soó (70) 26, 47, Epipactis microphylla (Ehrh.) Sw. (92) 47, 48,
64, 68, 72, 76, 78, 98, 106, 110, 138, 146, 152, 217
199, 203, 204, 206, 214 Epipactis ovata 106
242