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ORCHIDS OF CENTRAL (Cuenca Province) A field guide

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Collection Essential Guides of Flora, 2

ORCHIDS OF CENTRAL SPAIN (Cuenca Province) A field guide

Agustín Coronado Martínez Eduardo Soto Pérez www.jolube.es at

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Book

2019 Contents

Prologue...... 6 dyris ...... 1 122 How to use this guide ...... 8 Ophrys incubacea ...... 1 124 The three natural landscapes of Cuenca ...... 20 ...... 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 Key for identifying Orchid genera ...... 50 Ophrys tenthredinifera ...... 140 Aceras anthropophorum ...... 52 ...... 142 pyramidalis ...... 54 Orchis cazorlensis ...... 144 Genus ...... 56 Orchis champagneuxii ...... 146 ...... 58 Orchis fragrans ...... 148 ...... 60 Orchis langei ...... 150 ...... 62 ...... 152 viride ...... 64 Orchis palustris ...... 154 Genus ...... 66 Orchis papilionacea ...... 156 Dactylorhiza elata ...... 68 Orchis picta ...... 158 ...... 70 www.jolube.es ...... 160 ...... 72 Orchis tenera ...... 162 Dactylorhiza insularis ...... 74 at Orchis ustulata ...... 164 ...... 76 Genus ...... 166 Dactylorhiza sambucina ...... 78 Platanthera algeriensis ...... 168 Genus ...... 80 ...... 170 Epipactis cardina ...... 82 sale lingua ...... 172 Epipactis distans ...... 84 aestivalis ...... 174 ...... 86 ...... 176 Epipactis hispanica ...... 88 on Neighbouring ...... 178 Epipactis kleinii ...... 90 Additional Biology ...... 184 Epipactis microphylla ...... 92 Orchids and fungi ...... 184 ...... 94 Orchid cultivation ...... 186 ...... 96 Book Cultivation in pots ...... 192 conopsea ...... 98 Evolution ...... 194 hircinum ...... 100 methods ...... 196 Genus ...... 102 Hybrids ...... 202 Limodorum abortivum ...... 102 Unusual cases ...... 204 Limodorum trabutianum ...... 104 Orchids protection...... 205 Listera ovata ...... 106 Some interesting facts ...... 207 maculata ...... 108 Locations ...... 209 Neottia nidus-avis ...... 110 Spanish-English Lexicon of Habitats ...... 232 Genus Ophrys ...... 112 Glossary of terms ...... 233 ...... 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 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 we began our tentative research, certain that we wouldn’t find more than a dozen different species. With orchids it’s a bit like with mushrooms; hunting for them becomes a hypnotic entertain- 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, 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.

Totally bewitched, we spent the next seven years enjoying one of our favourite pas- times: going out to the country, exploring the terrain, breathing in the pure air, slee- ping rough, venturing into the maze of woodlands, and uncovering the secrets of the 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 www.jolube.es efficiency where each of its elements is provided for and lacks for nothing. at 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, expanded into volumes which filled our bookshelves but which no longer fit into our rucksacks. sale

Since the mid-17th C, and basically stimulated by commercial interests in theiron attrac - tiveness for gardens and greenhouses around the world, research into the family has opened up new fields, techniques and questions for work in the biological sciences: in ecology, physiology, improvements in genetics, molecular genetics, 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 Booktypes 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 of land orchids and natural environment attracts orchid specialists from all over Europe. their specific fungi has permitted extraordinary advances in research into . 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 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.

Spain is to be found between the longitude zones 29 and 31 and latitude bands T and S. The Cuenca province area is within zone 30 and between the parallels T and S. As 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 coordinate is placed in the bottom left corner of the square (not in the centre).

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 reference is first), and from west to east (the horizontal reference goes after). The whole of the Cuenca province is included: the north in the 100 km × 100 km squares VK, 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 kmwww.jolube.es assigned a combination of two numbers. 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 atso on, depending on the desired precision of the 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. sale

LK on UQ CK LJ T UP CJ LH UN CH LG Book UM CG LF FF MF NF PF QF TL UL VL WL XL YL BF CF DF EF LE UK CE LD UJ CD LC UH CC LB UG CB LA UF CA LV S UE CV LU UD CU LT Aguiló Publications UC CT J.E. García Rodríguez

12 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 phical region and would cover it entirely Continental. if they hadn’t been subjected to human intervention. The Holm Oak (Quercus ro- 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- terranean zone on limestone substrate ll parts in the higher altitudes with se- soils. Such are the oak woods of Alcantud, mi-arid conditions. Impoverished by the where we can find Aceras antropophorum continental climate conditions, the oak and Cephalanthera longifolia orchids. In woods in Cuenca are not so rich as in the many parts the oaks have been replaced coastal or the offshore mountain regions by Aleppo (Pinus halepensis), as in of the Mediterranean; there are fewer the Batanejo reservoir and the Camino al plant varieties, as the cold has elimina- www.jolube.esBujoso (the River Cabriel basin) where ted species such as the Butcher’s Broom groups of Limodorum abortivum, Cepha- (Ruscus aculeatus) or the Sarsaparilleat lanthera longifolia, Neotinea maculata (Smilax aspera). and Ophrys dyris are to be found. In the sale on

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24 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. In siliceous parts of the southern Iberian Together with these can be found Philly- System (the region of Alcantud and Sier- 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 gorges) or Ruscus aculeatus (on shaded, Characteristic species: Arbutus unedo, humus-rich soils). In these environments, Erica arborea, Erica scoparia or Phillyrea species of Mediterranean-type orchids angustifolia. In the warmer areas Pistacia such as Ophrys lupercalis or Ophrys dyris lentiscus, Smilax aspera or Erica multi- grow. flora are added, and in the shadier parts Viburnum tinus, Ruscus aculeatus or Bux- Levantine siliceous Maquis us sempervirenswww.jolube.es are also included. shrublands atThe most typical orchids in this type of Tall, generally dense shrubs which act as habitat are: Aceras anthropophorum and the main vegetation stratum or under- Cephalanthera longifolia. growth for the Pinus pinaster pines,sale and on

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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 and second in its number of these are epiphytes, ie. they grow of components within the plant kingdom. on other plants, usually on the trunks of 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 number of species estimated at between gathered between the branches and on 20,000 and 30,000. These are classified the foliage, and take their water from the into 70 subtribes and 22 tribes, divided moisture in the air through tiny absorbent in turn into 5 subfamilies: Apostasiodeae, hanging roots. Cypripedioidae, , Orchi- Antarctica and the arid deserts are the doideae and Spiranthoideae. This division www.jolube.esonly places in the world where orchids do is based mainly on the different number not grow. They have colonized all other of anthers and their arrangement. at habitats, from the tundra in the Arctic Cir- Among this enormous variety of species cle to the plains in , in the Florida there are all sorts: large and beautiful, swamps and in the source of the River Cu- tiny, subtle and delicate, showy,sale fragrant, ervo in Cuenca. All the European orchids odourless or smelling of rotting meat, belong to the Holarctic ecozone. multicoloured, black, white, with mark- The European continent has around 250 ings or designs. on species and numerous subspecies. The There are orchids with tiny flowers which majority belong to the subfamily Orchi- measure only millimetres (Platystele doidae; the genera Cephalanthera, Epi- stenostachyaBook), 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 , 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 38 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 Greek orchis, meaning testicle, due near the main plant and which share to the similarity in appearance of the genetic makeup. In some plants the underground tubers (spherical or this means of plant propagation is more ellipsoid). These tuberous roots store frequent than in others, as for example water and nutrients. in certain species of the genera Epipactis or Orchis champagneuxii, which is why it is common to find them growing packed very closely together.

www.jolube.es Phloem

at Xylem Pericycle Orchis Dactylorhiza Endodermis Starch sale Parenchyma Cortex on Epidermis

Book Ophrys scolopax root Root hair

Platanthera Neottia STEM The genus Dactylorhiza and the genus As with the majority of Monocotyledons, Coeloglossum both have a 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 40 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)

3. Flowers with no spur ...... 4 — Flowers with spur ...... 11

4. Labellum decorated with a shiny central pattern (speculum) surrounded by a more or less hairy band ...... G. Ophrys (112) ­— Labellum with no shiny central marking (speculum) and no obvious hair...... 5

5. Labellum divided by a transverse constriction on the hypochile (inner portion) and the epichile (outer portion)...... 6 — Labellum with no central division ...... www.jolube.es 8 6. Pedunculate ovary. Hypochile concave or cup-shaped, shiny and nectariferous. Flowers patent or pendulous ...... at G. Epipactis (80) — Sessile ovary. Cup-shaped hypochile, no . Flowers horizontal or raised .... 7 7. Green . Hypochile withsale parallel crests. Perianth with free divisions ...... G. Cephalanthera (56) — Bracts of the same colour as the . Epichile without crests. Perianth with sealed divisions...... on G. Serapias (172)

8. White flowers. Spiral axis of or as unilateral sprig...... 9 — Green orBook 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 50 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 at the base of the labellum. The examples of the genus Aceras have two retinacula, or viscidia, sometimes joined together and sometimes separate, surrounded by a simple bursicle. This feature places it between Orchis and Ophrys. In Orchis there are two viscidia enclosed in a bi-lobed bursicle and in Ophrys there are two viscidia but they are enclosed in two separate bursicles.

Etymology. From –phorum, bearer, and Chromosome number. 2n=42. www.jolube.es anthropo–, man. From the shape of a man on Pollination. Various Coleoptera (beetles) the labellum. and some (flying insects), Description. 12-28 cm in height. Hairless unspecified. at cylindrical stem. 5-10 basal leaves, Synonyms. Its closeness to the genus Orchis unmarked but distinctly veined, oblong- has been confirmed by molecular analysis 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 sale 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., ona name 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. 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. Book Habitat. Full sun or shade. On limestone, sand or clay. In dry grasslands and together with thyme, in 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(L.) Rich. rubra

Etymology. From rubra-red; the colour of Pollination. In general, the genus Cephalanthera www.jolube.es the flower. attracts only by its appearance. It attracts Description. 16.5-50 cm in height. Slender, which enter its narrow pathway wavy stem, grey hairs on the upper portion. between the epichile and the column so that at 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 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 appearssale as 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 ontwo veins. compact masses that can be easily extracted Habitat. In shady or semi-shady places. On during flowering, which makes it the only 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. Book 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(Druce) Soó fuchsii

Etymology. The species is attributed to Hybrids. We have found hybrids of this with Leonhart Fuchs, German professor (1501- Dactylorhiza elata. www.jolube.es 1566). Chromosome number. 2n=40. Description. Slender plant of 38-48 cm in Pollination. Beetles (Cerambycidae), insects at height. Thin stem, tinged violet at the top. and bees. 5-11 foliage leaves, keel-shaped and marked, Synonyms. Orchis fuchsii Druce; O. shiny grey-green underside and spotted on maculata var. trilobata De Brévisson; the upper side. Inflorescence compact or Dactylorhiza maculata auct. Non L.; sale 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. meyerion (Rchb. shades of lilac and white. Lateral sepals Fil) Tournay; D. meyeri (Rchb. f.) Aver. 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 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 Bookheavily 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. 70 71 (L.)Epipactis Crantz helleborine

Etymology. From heleboro, the old name for Companion Orchids. Neottia nidus-avis has Epipactis and Veratrum album. the same eco-system preferences. www.jolube.es 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, at ovate or almost round lower leaves, deep 44. shiny green, at an angle, hanging down Pollination. By bees, mainly by Vespidae; and distinctively veined, 4.5 × 7.5 cm; Halicidae and are mere visitors upper leaves smaller and more lanceolate. rather than effective . sale Inflorescence a spiked cluster, one-sided, Synonyms. E. latifolia (L.) All.; E. leutei 5.5-19 cm tall, with 20-36 open flowers. Robatsh. on Sepals green on the outer side and lightly Comments. Epipactis helleborine is a rare 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 strongly tinged with pink or purple. Labellum matter it requires for development. greenish purple, epichile with two bosses. Hypochile deep brownish-grey on inner Book 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. e p

86 87 (L.)Himantoglossum Spreng. hircinum

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. They have numerous large leaves, broadly lanceolate, with no perceptible cross veins. The smaller lower leaves are usually yellowed and withered by flowering time. The flower spike is tall, with numerous large flowers and non-sheathing bracts. The flowers have free sepals which form a hood. The lip broadens at the end, is divided into three lobes, with a very long and twisted central lobe. Short spur, curved downwards and with no nectar.

Etymology. Hircinum refers to the smell ‘of Hybrids. Hybrids have been described with www.jolube.es goats’. the Orchis militaris group, although these Description. Robust plant, 20-110 cm tall, are extremely rare. at 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 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, , bugs, butterfliessale Photo: Óscar García Cardo cm, becoming smaller and clasping up the 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, whichon may tall, very dense, cylindrical and with 20-120 also be potential pollinators. foul-smelling flowers. Sepals oval or broadly Synonyms. Satyrium hircinum L., Aceras 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) inBook 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. e p Companion Orchids. Unknown. Photo: Javier Benito Ayuso Photo: Jolube

100 101 OphrysHuds. apifera

Etymology. From apis-, , and –fera, Flowering Period. End of May and beginning carry or bear, ‘bee-bearing’, due to the lip’s of June. www.jolube.es 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. at 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. flowers. Sepals oval-lanceolate, keel-shaped, Chromosome number. 2n=36. in shades of pink ranging from white to Pollination. Self-pollinating, when the sale purple; obvious green central veins; held flowers open the slightest movement causes horizontally; upper bent backwards. the pollen to fall onto the stigma.on Petals very small, leaning forwards, hairy, Synonyms. O. arachnites Millar; triangular, with margins rolled back. Lip O. aquisgranensis Kaltenb.; O. ripaensis three-lobed, with dark crimson-brown Porta; O. botteronii Chodat; O. jurana velvety hair covering most of the surface, Neuberger; O. trollii Hegetschw.; paler at the tip of the centre lobe, which O. frigurgensis (Freyhold)Book 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 OphrysRisso bilunulata

Etymology. From bilunata, two moons, Pollination. By flying insects of the genus referring to the shape of the markings. (A. nigroaena, A. flavipes) and by www.jolube.es Description. 8-13 cm in height, with oblong- Colletes cunicularius. lanceolate leaves. Inflorescence with very Synonyms. Ophrys flavipes-fuscaauct.; at few – 3-6- obvious, dark flowers. Petals Ophrys leucadica Renz. almost spatula-shaped. Labellum 11.4- Comments. The classification is of western 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 cansale 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 on reach into the mouth of the stigmatic cavity. Habitat. Full sun or shade. On grasslands in pinewood clearings of Pinus halepensis (Aleppo Pine) in full sun, and in open woods of Pinus nigra (European Black Pine) with Book 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- ical habitat of the species, the flowering period and the highest altitude at which the plant has been found.

Dactylorhiza markusii (Tineo) H. Baumann & Kunkele References towww.jolube.es Guadalajara in CARRASCO, MACÍA & VELAYOS (1997) and to Albacete RIBERA & LÓPEZ (1987) underat the name of D. sulphurea subsp. sicilien- sis (Klinge) Franco. It is of Western Mediterranean distribution although more Southern than D.insularis. It growssale on acid substrates in clearings of Pyrenean Oak woods, at altitudes of up to 2000 m. It is quite rare and in localised areas. It flowers during the second on half of May and beginning of June.

Photo: Javier Benito Ayuso.

Book 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 benefit, is known as mycorrhiza: the Germination can only take place in 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 exchange provides essential ions, usually and immediately invades the area of the phosphates and nitrates. embryo. The fungal hyphae penetrate the cell wall and form coils and loops Curiously, however, the mycorrhiza of called pelotons which, when seen under orchids is reversed: the host orchid a microscope, show that the mycorrhizal 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 mycotrophic. Many authors believe this host at controlled intervals to avoid unusual symbiosis should be considered www.jolube.esreinfection, as this would mean it would as a parasitic process, as the fungus does be deprived of nutrition. not seem to benefit from the relationshipat but quite the opposite, although it is The seed undergoes a substantial never completely destroyed by the change. At first the mycotroph or plant. The fact is that the mechanisms protocorm looks like a tiny tuber, an which control the process of infectionsale undistinguishable mass of cells from and how the orchid-fungus relationship which soon appears a shoot at the tip is balanced are still not fully understood. which will produce the leaves. When In fact, orchids produceon phytoalexines, exposed to light this shoot will begin vegetable hormones which control the to pigmentize and presumably begin natural aggressiveness of the fungus, to photosynthesise. The lower part although it is not known to what degree becomes the organs similar to roots it may actuallyBook 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 to be content with the orchid varieties found in garden centres, which, after THE MOST COMMONLY CULTIVATED all, is not bad. ORCHIDS We have been talking about the different methods of multiplying orchids, and Genus Cymbidium because there is a lot of interest, Of Asian origin, it grows in sunny habitats both personal and industrial, in their from to Japan and Australia. It qualities as ornamental plants, we will grows very easily and the genus has a provide some general information on large number of hybrids, which is why it pot cultivation and some indicators on is one of the most cultivated genera. soil, feeding and watering requirements. www.jolube.es Internet has many sites on this hobby at This orchid has a pseudo bulb at the base which has recently become so popular of the leaves comprised of aquiferous around the world. tissues, and it produces from eight to ten leaves and vigorous roots. Some We’ll begin by saying that theresale is a varieties begin flowering in November large number of orchids which, by paying while others flower in February or March. attention to a few specific conditions, The nocturnal temperature should not be can be cultivated at allon latitudes. We higher than 14ºC in summer and in winter should select species which can best it should be between 12º and 15º C. adapt to the predominant climate where we live and, when needed, provide the conditions to make up for any deficienciesBook (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 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 seen, with three sepals, two petals and determined; scientists believe this is why the lip, and a long inferior ovary, dating orchids have taken on such interesting from the Late Miocene period, in other evolutionary adaptations: they are words about 15 million years ago when opportunistic, they produce millions of Europe had a tropical climate. seeds, they need mycorrhizal fungi to germinate and they use unique methods Nevertheless, later DNA studies have to ensure pollination. suggested that the Orchidaceae family could be much older than was originally Depending on the species, each plant thought and that its origins could go can produce from two (eg. Ophrys) to back to the end of the Cretaceous period almost one hundred (some Gymnadenia of circa 65 million years ago. www.jolube.esand Dactylorhiza) mature fertilised ovaries, each containing over a million Be that as it may, it is still a group of at microscopic seeds. recent development compared with the monocotyledons which appeared 100 It should be noted that the evolutionary million years ago, or the dicotyledonssale success of orchids is directly related to of 150 million years ago, or conifers, their capacity to associate with different thought to age from 300 million years ecosystems: on the one hand that of ago. on pollinating insects, with which their relationship is closer and more complex All researchers coincide in that orchids than any other plant’s, and on the other have probably evolved from the hand, with the mycorrhiza, on which botanical family of Liliaceae. They their growth depends to the extent that preserve theBook 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 (beetles) and, more rarely, by slugs and adaptation of the genus Ophrys within snails. the Mediterranean area.

The components of the genus All of this gives us some idea of the Plantanthera create a long spur full of intimate relationship between the nectar which can only be reached by method of fertilisation and evolutionary certain species of moths equipped with development of the Orchidaceae family. long proboscides. At the other extreme, 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 variety of insects can access it. www.jolube.escapacity to adapt to pollinators, and therefore that these pollinators have Cephalanthera longifolia has flowers at driven the differences in flowers and similar to those of Cistus salviifolius species. This has generated much (as seen by insects), and attracts them research which both supports and without needing to give up the pollen refutes the theory. normally given by the rock rose.sale The perianth segments of the genus Serapias form a narrowon tube which THE METHODS USED TO ATTRACT provides a refuge for particular bees at night or in inclement weather POLLINATORS conditions. Book 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 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 themselves. This creates hybridogenetic colonies which are difficult to identify, as the offspring have features of both parents. This is an important detail and should 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 plant should have not just one but many intermediate features. Occasionally the 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 confirmed through repeated observations.

When the crossing is produced between plants of the same genus but of different species, this is called interspecific, or infrageneric;www.jolube.es when it takes place between species belonging to another genus it isat called intergeneric. Intergeneric hybridisation is limited to the genera Dactylorhiza, Coeloglossum, Platanthera, Gymnadenia, Nigritella and Pseudorchis, on the one hand, and to the genera Orchis, Neotinea, Aceras,sale Anacamptis and Serapias, on the other. Ophrys, however, rarely produces interspecific hybrids. The possibility of producing this phenomena in natureon is quite remote; the cases that have been compared (Dactylorhiza × Gymnadenia, Orchis × Dactylorhiza, Gymnadenia × Nigritella, Coeloglossum × Dactylorhiza, among others) have served as a basis for genetic analysis giving rise to a new focus for understanding the development of the Orchidaceae family, and without doubt, will revolutionise the systematics of orchids in the future.Book

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 changes brought about by man. peatbogs, semi-natural dry grassland formations and shrubland on calcareous Uncontrolled urbanization, road soils (Festuco brometalia). building, intensive agriculture etc. all progressively reduce or alter the natural CITES (Convention on International environment and continuously damage Trade in Endangered Species of Wild biodiversity. Fauna and Flora) is an international treaty whose aim is to ensure that Due to their particular biological cycle, international trade in specimens of wild orchids are sometimes subjected to animals and plants does not threaten the other pressures which make them survival of the species. Member states more vulnerable to actions by man. whowww.jolube.es adhere to CITES have to apply the For example the indiscriminate use norms of the convention although these of nitrate fertilizers prevents the atdo not override national legislation. Each development of the mycorrhizal fungi, country must pass its own legislation in pesticides significantly reduce colonies order to guarantee that CITES is applied of potential pollinators and the mass across the nation. 63 species of Spanish picking of attractive species havesale orchids are included in the Convention. brought some orchids to the point of extinction (as in the case ofon Cypripedium At our regional level, the Decree calceolus). In addition to all these 199/2001, dated 6 November 2001, threats is the fact that traditional adds to the Regional Catalogue of farming methods have been abandoned; Threatened Species in Castilla-La the seasonal moving of livestock, the Mancha (CREA) dated 5 May 1998. mowing of fieldsBook 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 for this family who wrote a treaty for the maintained their genus name, to which was 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 intergeneric hybrid crosses produced names In the west, its discovery as a valuable like Catamodes (Catasetum × Mormodes), 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. The desire for new species led to further During this period, Victorian England had complexities – the hybrids themselves were explorers spread all over the planet. In crossed with other species and produced 1818, Swainson, who was in South America hybrids such as Brassiolaeliocattleya collecting mosses and lichens for William (Brassavola × Laeliocattleya), which not Cattley, a British horticulturalist, used the www.jolube.essurprisingly became abbreviated to BLC. When vines of some plants to wrap the moss in, even more hybrid genera were introduced it without realising that he was in fact using at became necessary to create a methodology orchids from the Atlantic Forests of Brazil. exclusively for naming orchids. These vines, or lianas, arrived intact and alive and flowered in England that same year. The complete register of orchid hybrids is held by the Royal Horticultural Society (UK) Cattley was intrigued by their beautysale and and can be accessed on Internet. he took them to John Lindley, an eminent horticulturalist of the time, who named them USES as a genus and provided a ondescription. As their lip was particularly outstanding, Lindley Aromatics included the word ‘labiata’ in the name. 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 Bookand 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.

Aceras anthropophorum 1. 30TWK5488 Alcantud. Dehesa de Alcantud. 800 m. Quejigar. 1-5-02. 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. 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). 6. 30TWK7389 Beteta. Paseo Botánico. 1110 m. Bosque mixto. 20-5-00. 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. 9. 30SWK8316 Almodóvar del Pinar. N-320. Km.www.jolube.es 116. 1050 m. Quejigar. 18-5-00. 10. 30TWK8332 Cuenca. Hoz de San Miguel. 1200 m. Encinar en el calar. 24-5-00. 11. 30TWK8378 Cañizares. Huerta de Marojales.at 1360 m. Pinar de P. sylvestris. 12-6-99. 12. 30TWK8467 Las Majadas. Cobacha del agua. 1440 m. Tomillar. 7-6-02. 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. 15. 30TWK8831 Cuenca. Torca del Agua y Torca del Lobo. 1240 m. Pinar de P. nigra. 8-5-99. 16. 30TWK9378 Cuenca. Los Sabinarejos.sale 1500 m. Pinar de P. sylvestris. 24-6-00 (fruit). 17. 30TWK9476 Cuenca. Nacimiento Río Cuervo. 1500 m. Pinar de Pinus sylvestris. 4-5-02. 18. 30TXK0548 Valdemeca.on 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- 20. 30SWJ7779 Alarcón. Casas del Embalse de Alarcón. 810 m. Coscojar. 22-4-2006. 1. 30TWK629Book3 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. Brezal: Heathland / Moor. Rodenal: Pinus pinaster forest. Bujeda: box forest. Romeral: Rosemary shrubbery. Calar: Calcareous, chalky. Roquedo: crag. Caliza: Limestone. Sabinar: Juniper forest. Cambronal: thomy schrubland. Sauceda: Willow forest. Canchal: Rocky soils, scree. Talud: slope. Cantil: ledge. Terraza: terrace. Carrizal: Reedbed. Toba:petrified lime deposits Cauce seco: dry river bed. Tomillar: Thyme field. Chopera: Poplars. Turbera: peatbog. Claro: clearing. Vaguada: stream bed. Coscojar: Kermes Oak grove. www.jolube.esYeso: Gypsum soil. Cuneta: ditch. Dehesa: Sclerophyllous scrub. at Derrubio: scree / debris. TYPES Encinar: Holm Oak forest. Garriga: Garrigue. Adehesado: turned to scrub. Guijarral: Stony terrain. Arenoso: sandy. Herbazal: meadow. sale Despejado: open, clear. Jaral: Rock rose thicket. Encharcado: swamped. Juncal: Reed bed. on Húmedo: damp. Ladera: hillside. Inundado: flooded. Lastonar: grassland. Mixto: mixed. Marga: Marl (Lime-rich mudstone). Pedregosa: rocky. Matorral: shrubland. Rezumante: seeping. Melojar: PyreneanBook Oak forest. Salino: salty. Olivar: olive grove Umbroso: shady. Orilla: bank border. Yesosa: chalky. Pastizal: grassland.

232 Spanish-English Lexicon of Habitats 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. Basionym: the original, validly published Esparto: tough wiry grasses. name of the taxon. Exogenous: originating externally Biotype: a group of genetically identical Filiform: Very narrow with straight and plants within a species. parallel margins; thread-like. : a leaf-like structure which has no Fusiform: spindle-shaped; widest in the blade or lamina. middle and tapered at each end. Bursicle: A membranous sheath which covers Foliaceous: Leaf-like. the glue in some viscidia. Garrigues: shrubs typically found on calcareous Callus: a raised, fleshy structure found on Mediterranean soils, usually aromatic shrubs the labellum. eg, thyme, rosemary, lavender etc. Calyx: the outer part of the flower, the se- Gibbous: With a pouch-like swelling; humped. pals. www.jolube.es Glabrous: Without hairs. Cambrones: common name for thorny shrubs which grow in the shape of a cushion. Examples atGlaucous: covered with a bloom which lends In Cuenca are Erinacea anthyllis and Genista a bluish lustre. pumilla. Humic: derived from humus. Caudicle: a term used for a pollinium stalk Hymenoptera: flying insects such as bees, derived from the anther; an elastic basalsale wasps and ants. extension of the pollinia. Hypochile: The inner portion of the labellum, Cauline: belonging to a stem, onusually refers often cup-shaped. to leaves. Inflorescence: The flowering structure of a Ciliate: with a fringe of fine hairs. plant. Circumboreal: relating to the northern Labellum: a lip - in orchids it is a highly portion of the Northern Hemisphere. modified that is primarily involved in Cladograms: mapsBook 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 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 which becomes a seed after fertilization. with sessile flowers. Palmate: divided like a hand. Spur: A slender hollow projection from a floral segment (usually the labellum, rarely Pedicel: The stem which supports a single the dorsal sepal). flower in an inflorescence. : male reproductive organs of a peduncle: The main axis of a compound . inflorescence or the stalk of a solitary flower which subtends the pedicel. Staminode: a sterile stamen Pedunculate: Having a peduncle. Stigma: (pl. stigmata) The enlarged sticky area which terminates the pistil and is Perianth: A collective term for the petals and receptive to pollen and allows the pollen sepals of a flower; in orchids this does not grains to germinate. include the labellum. www.jolube.esSupramediterranean: mid-montane zone, Petiole: the stalk of a leaf. 900 - 1200 m altitude. Polje: a large elliptical depression in limestoneat Symbiosis: a close relationship between two regions, sometimes containing a marsh or or more organisms, including parasitism. small lake. Tepal: in Orchidaceae, this term refers to Pollinium: (pl. pollinia) An aggregated any sepal or petal, not the labellum. coherent mass of pollen grains foundsale in the Orchidaceae. Utrillas sands: Albian sands found in the area of Utrillas, in the province of Teruel. Protocorm: Specialised structure that develops after orchid seed germinationon and from which Viscidium (pl. viscidia): A clearly defined a shoot develops. sticky part of the rostellum which is removed together with the pollinia as a unit by a visi- Rhizome: An underground stem with nodes ting insect. and roots, which can form shoots. Xerophyte: a plant adapted to dry growing Rostellum: TheBook 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 American Orchid Society Gold Coast Unlimited Orchid Society, North Miami, FL Amherst Orchid Society, Amherst, MA Grand Valley Orchid Society, Walker, MI 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 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 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 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) Atlanta Orchid Society, Atlanta, GA Houston Orchid Society, Houston, TX Australasian Native Orchid Society Australia Illinois Orchid Society, Chicago, IL Australasian Native Orchid Society, Victoria Group Australia Illowa Orchid Society, Davenport, IA Bankstown Orchid Society, Bankstown, Sydney, NSW, Australia Iowa City Orchid Society, Iowa City, IA Blue Ridge Orchid Society, VA www.jolube.esInternational Phalaenopsis Alliance The Boca Raton Orchid Society, Boca Raton, FL Jupiter/Tequesta Orchid Society, Jupiter, FL Bonsai and Orchid Association International Kansas Orchid Society, Wichita, KS Bournemouth Orchid Society, Bournemouth, UK at Lehigh Valley Orchid Society, Allentown, PA British Orchid Council, UK Les orchidophiles réunis de Belgique, Bélgica British Orchid Growers Association, UK Les Orchiophiles de Montreal, Montreal, PQ, Canadá 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,sale MA Manitoba Orchid Society, Winnipeg, MB, Canadá Cape Fear Orchid Society, Wilmington, NC Maryland Orchid Society, Baltimore, MD Capital City Orchid Society, Wellington, New Zealand Massachusetts Orchid Society, Boston Area Carmel Orchid Society, Monterey, CA The Maxillaria Tribe Catoctin Orchid Society, Frederick,on MD Michigan Orchid Society, Greater Detroit Area, MI 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 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 JudgingBook 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 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, , 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

ABADIE, J-C., PÜTSEPP, Ü, GEBAUER, G., FACCIO, A., BONFANTE, P., SELOSSE, M-A. (2006). Cephalanthera longifolia (Neottieae, Orchidaceae) is mixotrophic: a comparative study between green and nonphotosynthetic individuals. Can. Jor. Bot. (84) 1462-1467. AGUILELLA, A., FOS., S. & LAGUNA, E. (2009). Catálogo Valenciano de Especies de Flora Amenazadas. Conselleria de Medi Ambient, Aigua, Urbanisme i Habitatge, Generalitat Va- lenciana. ÁGREN, L. KULLENBERG, B & SENSENBAUGH, T. (1984). Congruences in Pilosity between Three species or Ophrys (Orchidaceae) and Their Hymenopteran Pollinators. Nova Acta Regiae Societaris Scientiarum Upsaliensis, Serie V: C, 3, 15-25. ARNOLD, J. (1981). Notas para una Revisión del género Ophrys L. (Orchidaceae) en Cataluña, Collectianea Botanica 12(1): 5-61. BATEMAN, R. PRIDGEON, A. & CHASE, M.(1997). Phylogenetics of subtribe Orchidinae (Orchi- doideae, Orchidaceae) based on nuclear ITS sequences. 2. Infrageneric relantionships and reclassification to achieve monophyly of Orchis sensu stricto. Lindleyana 12(3): 113-141. BENITO, J. & AMICH, F. (1999). Orquídeas de La Rioja. Estudio fenológico, ecológico y corológi- co. Catalogación de la especies amenazadas. Zubía 14: 129-131. BENITO, J. ALEJANDRE, J. A. ARIZALETA, J.A. & MEDRANO, L.M. (1998). Epipactis distans Arvet-Touvet en el Sistema Ibérico. Flora Montiberica 8, 55-60. BENITO, J. & HERMOSILLA, C.E. (1998). Dos nuevas especies ibéricas, Epipactis cardina y Epipactis hispanica, más alguno de sus híbridos: Epipactis × conquensis (E. cardina × E. parviflora), y Epipactis × populetorum (E. helleborine × E. hispanica). Est. Mus. Cienc. Nat. de Álava 13: 103-115. BENITO, J., ALEJANDRE, J.A., ARIZALETA, J.A. www.jolube.es(1999). Aproximación al catálogo de las Orquídeas de La Rioja (España). Est. Mus. Cienc. Nat. de Álava 14: 19-64. BENITO, J., ALEJANDRE, J.A., ARIZALETA,at J.A. (1999). Epipactis phyllantes G.E.Smith en la Península Ibérica. Zubía 17: 83-98. BENITO, J., ALEJANDRE, J.A. & ARIZALETA, J.A. (1999). El grupo Ophrys scolopax (Orchi- daceae) en la Península Ibérica. Est. Mus. Cienc. Nat. 14: 65-73. BENITO, J., ALEJANDRE, J.A. & ARIZALETA, J.A.(1999). Aproximación al catálogo de las orquídeas de La Rioja (España).sale Est. Mus. Cienc. Nat. 14: 19-64. BENITO, J. (2000). Platanthera algeriensis Battander & Trab. in the Iberian Peninsula. Jour. Eur. Orch. 32 (3/4): 513-525. BENITO, J. (2000). Plantantheraon algeriensis Battander & Trab. en el Sistema Ibérico. Flora Montiberica 15: 38-41. BENITO, J. & TABUENCA, J.M. (2000). Apuntes sobre orquídeas (principalmente del Sistema Ibérico). Est. Mus. Cienc. Nat. de Álava 15: 103-126. BENITO, J. & TABUENCA, J.M. (2000). El género Dactylorhiza Necker ex Nevsky (Orchidace- ae) en el SistemaBook Ibérico. Est. Mus. Cienc. Nat. de Álava 15: 127-151. BENITO, J. & HERMOSILLA, C.E. (2000). Algunos híbridos de orquídeas nuevos para la Penín- sula Ibérica. Est. Mus. Cienc. Nat. de Alava 15: 185-188. BENITO, J., HERMOSILLA, C.E.& SOCA, R. (2001). Ophrys × bodegomii (Ophrys passionis × Ophrys tenthredinifera) nuevo híbrido de la Península Ibérica. Est. Mus. Cienc. Nat. de Álava 16: 89-92. BENITO, J. & TABUENCA, J.M. (2001). Apuntes sobre Orquídeas ibéricas. Est. Mus. Cienc. Nat. 16: 67-87. BOUILLIE, P. (1985). Aportación al estudio de las orquídeas gienenses. Blancoana 3: 92-94. BOUILLIE, P., BONILLA, J. & FERNÁNDEZ, C. (1992). Orquídeas de la provincia de Jaén. Blancoana 9: 102-111. BUTTLER, K.P. (1991). Field Guide to Orchids of Britain and Europe. The Crowood Press Ltd. British Library Cataloguing in Publication Data. CABALLERO, A. (1941). Apuntes para una flórula de la Serranía de Cuenca, Anales Jard. Bot. 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 154 & Herrero (74) 16, 28, 47, 146, 206, 215 156 — f. bartonii 74 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 — var. brachystachys 54 — subsp. arduennensis 76 Anacamptis tanayensis 54 — subsp. fuchsii 70 Anacamptis urvilleana 54 — subsp. meyeri 70 Anteriorchis coriophora subsp. fragrans 148 Dactylorhiza markusii (Tin.) H. Baumann & Apostasiodeae 38 Kunkele (178) Argorytes mystaceus 198 Dactylorhiza meyeri 70 Cattleya 193, 208 Dactylorhiza munbyana 68 Cephalanthera 38, 40, 43, 50, (56), 62, 197, 239 Dactylorhizawww.jolube.es romana subsp. bartonii 74 Cephalanthera alba 58 Dactylorhiza sambucina (L.) Soó (78) 26, 36, Cephalanthera damasonium (Mill.) Druce (58) at47, 198, 206, 216 25, 30, 44, 47, 48, 60, 62, 88, 102, 210 — subsp. insularis 74 Cephalanthera ensifolia 60 Dactylorhiza sesquipedalis 68 Cephalanthera grandiflora 58 Dactylorhiza viridis 64 Cephalanthera longifolia (L.) Fritsch (60) 24, Epidendroideae 38 25, 26, 28, 33, 47, 48, 52, 58, 62, 90, sale96, 102, Epipactis (80) 17, 38, 40, 41, 42, 43, 50, 66, 196, 104, 108, 196, 197, 211 234, 235 Cephalanthera pallens 58 on Epipactis atrorubens Hoffm. ex Besser (178) Cephalanthera rubra (L.) Rich. (62) 25, 27, 28, — var. tremolsii 96 29, 35, 47, 48, 60, 96, 102, 212 Epipactis campeadorii 88 Cephalanthera xyphophylla 60 Epipactis cardina Benito & C.E. Hermos. (82) 185 17, 29, 47, 48, 84, 90, 217 Coeloglossum 40, 54,Book 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 96 205 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. 68 Epipactis leutei 86 Dactylorhiza fuchsii (Druce) Soó (70) 26, 47, Epipactis microphylla (Ehrh.) Sw. (92) 47, 4 8, 64, 68, 72, 76, 78, 98, 106, 110, 138, 146, 152, 217 199, 203, 204, 206, 214 Epipactis ovata 106 242

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