Table Of Content

Orchids of Europe, North Africa and the Middle East 3rd edition (revised and enlarged) Pierre Delforge Edited by Simon Harrap A&C Black • London r Photograph crcdits The 1,270 photographs of this guide have been taken by Gdrard de Belair (Annaba, Algeria) 7, Pierre Delforge (Rhode-Saint-Gcndse, Belgium) 1,003, Pierre Dcvillers (Brussels, Belgium) 1, Jean Devillers-Tersehuren (Brus sels, Belgium) 28, D.M.T. Ettlinger (Great Britain) 3, Wolfram Foelsehe (Graz, Austria) 2, Alain Gevaudan (Villeurbanne, France) 6, Peter Golz (Winterthur, Switzerland) 2, Hmst Giigcl (Munich, Germany) 10, Werner Hahn (Koblenz, Germany) 2, Stefan llertel (Haag, Germany) 11, Jean-Michcl Hervouet (Chatou, France) 3, Alexandre Joukoff (Brussels, Belgium) 2, Erich Klein (Hggersdorf, Austria) 2, Einar Kongshaug (Tiller, Nor way) 3, Horst Kretzschmar (Bad Horsfeld, Germany) I, C.A.J. Krcutz (l.andgraaf, Netherlands) 122, Jurgen Passin (Langenfeld, (icrmany) I, I lolger Peiner ((ieesthaeht, (icrmany) I, I lelmut Presser (Pfiinz, Germany) 3, Hans R. Reinhard (Zurich, Switzerland) 2, Jany Renz (Switzerland) 2, Karl Robatsch (Austria) 2, Dietrich Riiek brodt (Lampertheim, Germany) IS, Manfred Schonlcldcr (I .everkussen, (icrmany) 5, Kurt Seiser (Vienna, Aus tria) 1, Laszlo J. Szentpcteri (Debrecen, I lungary) 2,1 lannelore Spaeth (Germany) 4, Walter Teschner (Velbert, Germany) 1, Daniel Tyteca (Ave-et-AulTe, Belgium) 10, E. Vimba (Riga, Latvia) I, Jurgen Vollmar (Stuken- brock, Germany) 2, Eric Walravens (I lamois, Belgium) 2,1 leinz Werner /aiss (MarlolTstein, Germany) 4. Watercolours, drawings and diagrams Pierre Delforge, Eliza Klopfcnstcin (Brussels, Belgium), Marc Walravens (Bossut-Gotteehain, Belgium). Editor’s Note Pierre Delforge’s Guide des onhidccs d'Europe is a seminal work that is crammed with data. Such large volume of information is hard to present in a readable form in any language, and we hope tha this English translation is at least as readable as the original French. The opportunity has been taken to correct a few mistakes and to bring the taxonomy and nomenclature up to date. Notably, Dactylorhizi longifolia has become D. baltica, Steveniella caucasica is now S. satyrloidcs, Himantoglossum (x samariense now H. samariense and H. (x) montis-tauri now //. Itolleanum. The Glossary has beet amended to better fit an English-language context and as many localities as possible have been angli cised, but it should be noted that it has not been possible to check all place names and some may re main in their French form. I would like to thank Tony Holcombe and Tony Leech for their assistance with translation queries. Simon Harrap, April 2006 Translated from the French by Laurent Penet and Carine Collin. This English edition published in 2006 by A&C Black Publishers Ltd., 38 Soho Square, London W1D 3HB Originally published in 2005 in French by Delachaux et Niestl6, Paris © 1994, 2001, 2005 Delachaux et Niestle, Paris. English translation © 2006 A&C Black Publishers Ltd. The right of Pierre Delforge to be identified as the author of this work has been asserted by him in accordance with the Copyright, Designs and Patents Act 1988 ISBN-10: 0-7136-7525-X ISBN-13: 978-0-7136-7525-2 A CIP catalogue record for this book is available from the British Library All rights reserved. No part of this publication may be reproduced or used in any form or by any means - photographic, electronic or mechanical, including photocopying, recording, taping or information storage and retrieval systems - without permission of the publishers. A&C Black uses paper produced with elemental chlorine-free pulp, harvested from managed sustainable forests. Printed and bound in Italy by Stige www.acblack.com 10 98765432 1 Contents I Foreword 32 Systematic list of species Understanding orchids 35 Key to genera (Jrchids in the plant kingdom Cypripedioideae li ()rchid anatomy 37 Cypripedium The flower I Inderground parts Neottioidae Aerial parts 39 Cephalanthera li Hie life of orchids (Termination 47 Epipactis Annual cycle 116 Limodorum lifestyle 118 Epipogium l I Si-xual reproduction in orchids 119 Neottia The mechanism of fertilisation 122 Spiranthes Methods of attracting insects I nvolution of modes of attraction 126 Goody era Autogamy and cleistogamy Epidendroideae In Apomixis 128 Corallorhiza t >i t liid classification 129 Calypso I ti Syslematics The complexity of the 130 Liparis species concept 131 Malaxis I'he systematics used in this guide Orchidoidae li Nomenclature I'he scientific name 133 Gennaria Problems of nomenclature 134 Habenaria Nomenclatural solutions 135 Herminium adopted in this guide 136 Neottianthe Identifying orchids 137 Traunsteinera IN Looking for orchids 139 Chamorchis l'1 I low to use the descriptions Region covered by the guide 140 Platanlhera l‘i Map 149 Gymnadenia Country abbreviations 171 Coeloglossum Signs, symbols and Latin 172 Dactylorhiza abbreviations used in the text 244 Serapias Using the keys I Jsing the descriptions 276 Anacamptis ' I Aberrant orchids 278 Orchis C ‘olour abnormalities 347 Sleveniella Anomalies of shape 348 Himantoglossum '(i Hybrids 358 Ophrys < >i t liid conservation 364 Pseudophrys 'H Plants in danger "i Possible actions 432 Ophrys Appendices Species accounts 623 Further reading II Systematic arrangement of European orchids 624 Glossary 630 Index 3 Foreword to the first edition professional alike, can judge the adopted Orchids are without doubt one of the most classification for themselves; the character beautiful and dynamic successes of recent istics used for the different species groupings plant evolution; about one flowering plant are given. species in twelve is an orchid. 19th century A work of science is a collective horticulturists spent fortunes bringing back undertaking. I could not have written this book large flowering species from exotic forests that without information, the exchange of ideas were difficult to cultivate. Today, however, or and much encouragement from many chids are no longer a symbol of a person’s botanists, both amateur and professional, in wealth, more of a superb and endangered flora Europe and the Middle East, to whom I very and fauna, smothered by the effects of an ever grateful but who are too numerous to list here. increasing human population. I should, however, especially like to thank my Europeans have turned more and more to colleagues in the ‘Section Orchidees d’Europe wards indigenous species in order to study and des Naturalistes Beiges’, with whom I have protect them and thus try and preserve some of learnt much during the last 13 years. Thanks the natural and semi-natural areas still remain especially to the photographers who have ing on the ancient continent. Over the last allowed the use of rare images, often unique, twenty years, the study of European orchids often accompanied by very valuable has advanced as much as the other scicnces, suggestions and personal observations. The resulting in three times as many species being illustrations would have been far from known in Europe as in 1970. It is therefore complete without the work of Madame Eliza necessary to review our knowledge for the Klopfcnstein (Brussels), who gave generously specialist, amateur and general botanist or for of her precious time. Finally, much thanks to the conservationist who wants to make a pre Monsieur Pierre Devillers (Institut Royal des cise inventory of an interesting habitat that Sciences Naturelles de Belgique, Brussels), could be protected. for having introduced me to modern This identification guide aims to be both as taxonomy, with particular reference to Europe complete as possible and to be understandable orchids, and for generously participating in by all, that is to say to present today’s numerous discussions, lectures and field knowledge of species and problems without outings during the preparation of this guide. avoiding complex questions. The class ification of European orchids, like that of any other family of organisms, is a science; neither finished nor fixed. Everybody, novice and Rhode-Saint-Genese, November 1991 Foreword to the third edition research has uncovered 150 new species, con Thirteen years after writing the first edition tested the validity of others already described and four years after the second edition ap and induced many minor taxonomic changes. peared, much has changed. A lot of work has This third edition, rewritten and enlarged, been done on orchids with the publication of tries again to offer the amateur naturalist and very many articles and monographs concern professional botanist alike, a foil and synthetic ing European orchids, proof of the increasing cross-section of present knowledge, its coher interest shown in this superb family not only ence relying on personal experience of most of by field botanists, but also, and increasingly the taxa in the guide. so, by geneticists and biochemists. Studying orchids by looking at their molecular compo sition is a new science with sometimes dis concerting first results, resulting in some im portant taxonomic modifications of genera; these are, however, often contradictory and may be premature. At the same time classical Rhode-Saint-Genese, December 2004 4 Understanding orchids Orchids in the plant kingdom V i rcently evolved group The systematic position of orchids I Uni ii few years ago, orchids were considered The plant kingdom is divided into branches, i" Iu' ;i very recently evolved family. They then sub-branches, classes, orders (suffix: - «imi' Ihought to have appeared suddenly, their ales), families (- aceae), and genera that i|m i in I characteristics clear-cut, in Asia (prob- groups similar species. Orchids are a part of nl'ly Malaysia) a mere 2 million years ago, as the Spermatophyte (or Phanerogammes) !li> earliest fossils definitely attributed to this branch, plants that reproduce by seed (which ttioup date from that time. However, the lack distinguishes them from Algae, Fungi, ■ il un hid fossils could be due to their way of Lichens, Mosses, Ferns...); the seeds result Ilf I topical plants grow quickly and have from fertilisation of the ovule, the female u iy little chance of being fossilised; nor have organ, by pollen tubes coming from the germi Illimi' growing on sloping or dry ground. Also, nation of pollen, the male cells. All Spermato- uni v macrofossils of orchids are known; the phytes have stems supporting the reproductive Ill11M11e seeds rot quickly and their pollen, organs. Usually the leaves perform the greater iiiii .r.l together in pollinia and transported by part of photosynthesis and the roots provide In ' els, is not scattered over wide areas as stability and nutritive functions but have nei mill wind-pollinated plants. Thus palynology ther leaves nor sexual reproductive organs. 1111 •* revealed nothing for orchids. Within the Spermatophytes, orchids belong to 11 ic recent discovery, in limestone from the the Angiosperm branch, plants with a closed ii|i|"'i Miocene in Germany, of a fossil that ovary containing the ovules (which distin mm obviously that of an orchid, with both a guishes them from the Gymnosperms, plants lip and an inferior ovary, shows that orchids with the ovules not enclosed in an ovary, such Iiiivv existed for at least 15 million years and as in the Ginkgos, Conifers, Yews...). There ii llml lime already occurred in Europe, which are some 170,000 species of Angiosperms lni'1 'i tropical climate. The presence of orchids placed in different families and orders grouped " curly as the Tertiary had already been into two classes. Orchids are members of the ni'l"" lcil by botanists who thought that it was class Monocotyledonae (monocotyledons), Impossible for them to spread throughout the plants with a single embryonic seed leaf h ui lil and adapt to such a wide variety of (cotyledon) at germination, and normally with li il'ii.ii;; hi just 2 million years. Orchids are in simple, entire leaves, with parallel veins, and clIccl cosmopolitan; the only areas where they the flower parts inserted in threes (or multiples il.......I occur are true deserts and arctic regions thereof). They are distinct from class Di- leu ihe soil is constantly frozen. cotyledonae (dicotyledons), which have two ! i' cillicless, even if we place the origin of cotyledons, with branching or fanned veins mi Imis some 20 to 30 million years ago, they and 5- or 4-part flowers (e.g. magnolias, aris mi i recently evolved group if compared to tolochias, buttercups, roses). Within the class llu In .I Monocotyledons (100 million years), Monocotyledonae, orchids are placed in the Hu Iulyledons (150 million years) or the order Orchideae, distinguished from other or i millers l U)() million years). ders, Grasses, Sedges, Rushes, and Wood- rushes, by their flowers with obvious sepals and petals, as in some other orders within the monocotyledons, such as lilies and irises (Liliaceae, Amaryllidaceae, Iridaceae...). 5 Orchid anatomy The flower lip, carries distinctive patterns and is often (Diagrams of flower parts are also given in the intro used as a landing stage by insects; it is ductions to the genera Epipactis, Serapias, Orchis and specialised and different from the two others. Orphrys). Finally, the sexual structures, with the exception of the ovary, are fused into a Orchids are probably derived from lilies. A column (the gynostegium), which faces the comparison between the flower structure of lilies and that of orchids allows an understand lip. These last two transformations give the ing of the similarities that demonstrate their orchid flower a bilateral symmetry, with only relationship and the derived characters that one axis through which it can be cut to give separate them. Lilies (Fig. 1) have hermaphro one half of the flower as the mirror image of dite 3-part flowers, with all floral parts present the other; it is termed zygomorphic. in multiples of 3, and regular or actinomorphic In a final important change, the ovary, which (star-shaped) because, due to their radial sym has become uni-locular, lies below the perianth metry their flower parts are arranged symmet and is thus inferior. It contains a large number rically around the flower centre. They also of minute seeds at maturity. The three sepals possess a superior ovary inserted into the pedi forming the calyx have remained rather simi cel at the same level as the perianth and thus lar, although the two lateral sepals, symmetri visible in its centre. The perianth is composed cal relative to the flower’s axis of bilateral of 6 tepals arranges in two whorls: the external symmetry, have a more or less irregular shape, whorl, the calyx, formed by three sepals, the different to that of the dorsal (or median) sepal, internal, the corolla, composed of three petals. which lies on the axis of the gynostegium. Of The tri-locular (3-chambered) ovary is sur the 6 stamens found in lilies, just 3 remain, mounted by a style, which carries, at its tip, 3 only 1 of which is fertile, the other 2 persisting fertile stigmas. The pollen is grouped in the only in the form of staminodes, fused into the anthers situated at the tip of 6 stamens, also wall of the stigma cavity and sometimes only arranged into 2 whorls. detectable as two small coloured patches, the In the orchids the 3-part structure is always staminodal points. The single fertile stamen present, but adaptation of the flower to has two loculi placed near the tip of the gynos pollinating insects has brought about tegium, each protecting 1 or 2 pollinia (never differentiation and the reduction of 2 petals in more in Europe) constituted of pollen grains comparison to the 3 sepals; the third petal, the assembled in tetrads which are in turn grouped Figure 1. Comparison of Liliaceae and Orchidaceae. A. Liliaceae. 1. Anther. 2. Stamen. 3. Ovary. 4. Stigma. 5. Sepal. 6. Petal. B. Orchidaceae (Ophrys splendida). C. Longitudinal section through an Orchis flower. 7. Dorsal sepal. 8. Lateral sepal. 9. Petal. 10. Labellum. 11. Spur. 12. Gynostegium. 13. Pollinia. 14. Stigmatic surface. into pollen masses sometimes lengthened by a Underground parts stalk, the caudicle. With the exception of 2 genera representing The 3 stigma of lilies are still present; 2 re the sub-tribe Liparinae (Liparis and Malaxis), main fertile and are usually fused into a single which can be considered to be epiphytes, all :;licky stigmatic surface; the third (in orchids, the orchids in Europe are geophytes. Their un the median), is generally transformed into the derground parts comprise various forms of rostellum, a more or less well developed struc- root; roots proper, normally in the form of lure placed between the stigmic surface and slender, cylindrical, unbranched filaments, the pollinia that prevents self-fertilisation. The whitish or brownish, an underground stolonif- lip of the rostellum is normally covered with a erous stem or a rhizome producing aerial slicky substance, contained within 1 or 2 vi- stems, or root-tubers (tuberous roots), organs i.ulia in those species that have pollinia with for storing food that allow the growth of a new i audicles, which sticks the pollinia to the pol plant, but which are not true tubers and cer linating insect’s body. The rostellum is some tainly not bulbs, albeit that these terms are in times elongated into 1 or 2 bursicles, pouch- common use. In the Liparinae, the food store likc membranes that cover the viscidia and organ is a pseudobulb, formed by a swelling preventing them from drying out. of the stem. In European orchids with a very short rhi Among the orchids, the families Apostasi- zome or without tubers, the roots arising from nceae and Cypripediaceae (see Systematic the rhizome can be a dense, tangled ball re miangement of European orchids p.31) show sembling a bird’s nest (Neottia nidus-avis) or mi intermediate evolutionary stage between swollen and club-shaped (Limodorum); the lilies and orchids. The former still have 3 fer rhizome can be branched as in a piece of coral tile stamens, 3 petals and 3 sepals that are sim (Corallorhiza, Epipoguin). In species with ilar, and pollen not gathered together into root-tubers, the roots are always placed at its pollinia. The latter family, represented in Eu tip. Root-tubers are varied: complete and rope by the Lady’s-slipper Cypripedium ovoid (Orchis, Ophrys), palmate (Coeloglos- inlceolus, still has two fertile lateral stamens sum, many Dactylorhiza), flattened and long- mid granular, sticky pollen not formed into digitate (Gymnadenia) or spindle-shaped pollinia. Certain degenerate forms of orchids (many Platanthera). There are generally 2 limy regress towards a flower structure similar tubers on each plant at flowering time; however, in that of the Aspostasiaceae, for example, certain species have 3 or more, often Ii modorum abortivum var. Irabutianum. If attached to the stem by a long underground llie lip is pointing upwards it is less visible to stalk (Orchis champagnewcii, Serapias lingua, living insects and obliges them to settle on it Ophrys bombyliflora). Between the roots npside-down, head and body facing down and aerial parts, there is normally a long wards. It is very likely that mutant plants, in underground stem, the neck, covered in u Inch the lip, turned downwards and more de- whitish, brownish, or sometimes purplish I' i Inble from above, offered a more appropri- scale-leaves. nli' landing stage for the insect's weight and Imvc llius been heavily selected by pollinating Aerial parts Insects. Most European orchids thus have the lip towards the bottom of the open flower, As in most monocotyledons, the stems of Eu w In n as it is found at the top of a flower in ropean orchids are never branched; they are I ii h I. Iliis 180° rotation of the flower, termed nearly always erect, more or less circular in n inpinnlion, takes place as the bud opens via section, rarely angular, solid or hollow, hair Mu twisting of the flower stalk or of the entire less or hairy. nviiiy if stalkless, sometimes via the whole The leaves are like those of other mono llnwei turning (Liparis, Serapias...). A few cotyledons, complete, never composite or di |h i ii :;, however, have their lips turned up- vided, with parallel longitudinal veins, some uin >l.'i Hie lop, either because there is no rota- times with transverse secondary veins forming ........K/iipogium), or by a 360° twisting of the a visible network (Goodyera). "i h v (Malaxis). In most of the saprophytic orchids the 7 leaves, performing no function, have been re near lax or lax, often becoming laxer as flow duced to scales or a sheath. When the leaves ering proceeds. The shape for the inflores are developed, they may be clustered at the cence is variable: cylindrical, conical base of the stem in a basal rosette (as in most (Anacamptis), ovoid, spiral (Spiranthes), uni Ophrys) or spaced out along the stem, in lateral (some Epipactis). which case they can arranged in a spiral (spi Flowers always have bracts, small leaves in rally), in two opposite ranks inserted at the serted into the axis of the inflorescence, at the same level (opposite), or alternately at differ base of the pedicel or ovary; these may be ent levels (alternate); the upper cauline leaves green or coloured as the sepals (Serapias), can be very small, resembling bracts. In some large and leaf-like (Dactylorhiza, Epipactis), genera there are only a few leaves, sometimes or reduced to membranous scales (most only two (Platanthera, Gennaria) or even just Orchis). Floral characters, very varied, are one (Malaxis, Calypso). given in the text describing the genera and The inflorescence is often a spike (flowers species. stalkless) or a raceme (flowers stalked); rarely it comprises a solitary flower (Cypripedium, Calypso). The inflorescence can be dense, The life of orchids The adaptive strategy of orchids may be process, often aborted and thus accentuating judged from their origin and floral characteris the need of a very large number of seeds to as tics. Evolving relatively recently, they sure the species’ survival. Darwin, and Lin emerged into an environment of animals and naeus before him, had already noted that if all plants that was already very diverse with, no the seeds from a single Dactylorhiza maculata tably, numerous potential insect pollinators. germinated as well as those of their descen The aggregation of orchid pollen into heavy dants, the worlds’ surface would be covered pollinia makes carriage by an animal, nearly by D. maculata within 4 generations. always an insect, essential, which permits a concomitant specialisation of the lip. In order Germination that pollination leads to effective cross fertili (Characteristics given in this and following sections sation, it is not only necessary that the insect concern European orchids.) carries off the flower's pollinia but also that Orchid seeds are minute, about 0.5 mm in these are rather quickly deposited onto another length and weighing about 1 Ojj.g (micro flower from a plant of the same species, evi grams). They are probably the smallest Mono dently a complex and infrequent process. cotyledon seeds; in comparison, rice or wheat From this it follows that the pollination mech seeds are enormous. They comprise a fine, anisms must be precise in order that the rare reticulated membrane covering a hardly differ successive visits are to result in fertilisation; entiated, spherical embryo. on the other hand, this problem is somewhat In a pollinated flower, when the capsule is alleviated by the production of a large number mature it splits open to liberate thousands of of seeds at each fertilisation. This necessitates seeds which are carried by the slightest breeze. the production of small seeds, extremely small Most of these will not germinate, either be in orchids, so much so that they totally lack cause they have an abnormality and are not vi the food reserves normally present in mono able, because they are consumed by other or cotyledon seeds in order for them to germi ganisms or because they fall on habitats where nate. This last fact has lead to a complex ger the conditions are unsuitable. Those that end mination mechanism, with the intervention of their journey on suitable terrain must, in order endophytic fungi and the establishment of to germinate, be invaded by microscopic symbiosis (a mycorrhiza), an unpredictable fungi, the Hyphomycetes, often of the genus 8 Lifestyle Ecology As they have significant food reserves, most European orchids can colonise nutrient-poor soils where competition with other plants, which they tolerate poorly, is reduced. Their relatively early flowering and sometimes brief Figure 2. Seed of Orchis lactea X 92. appearance above ground often allows them to (after Lucre) flower and fruit before the remainder of the Hhizoctonia, or Deuteromycetes, rarely the vegetation is at a maximum. They are llnsidiomycetes, living in the soil on roots in therefore, for the most part, favoured by cutting llio form of a mycelium. of meadows and summer grazing, and are well- The fungus penetrates the seed and infects adapted to the stress of dry Mediterranean I lie embryo. If it invades the embryo com summers and the fires that sometimes pletely, it would destroy it; orchid seeds con- accompany them. Some orchids, with an aerial I ii I n fungicides that appear to confine infection life of less than 45 days, have established In a portion of the embryo, the other part de themselves on mountains above the tree-line or veloping rapidly, mainly by taking sugars pro in boreal regions. As soon as conditions are unfavourable, following several poor years or duced by the fungus. Thus a symbiosis is initi- the vegetation closing due to recolonisation by nled, here called a mycorrhiza, beneficial to trees, the underground parts can persist for a both partners and lasting for a long time, for long time, sometimes without producing example, in Cypripedium calceolus, probably leaves. For this reason, the felling of a stand of Hie time necessary to form the plantlet. Sapro trees or a fire on a maquis can result, in the phytic orchids remain dependent on their host spring, in the reappearance in open areas of for their entire life but in most adult orchids orchids that denser and denser shade kept in a I lie endophyte's presence is apparently limited vegetative state underground. lo the peripheral zone of the roots. Due to their slow growth, complex The invaded embryo is transformed; a small germination and symbiotic life style, orchids I uber, the protocorm, forms, containing the generally require stable, sometimes climax, outline of a plantlet at the base of which ad conditions (mature beechwood, peat bog...). ventitious roots form. The young plant Traditional pastoral farming practices in slruightens and elongates vertically, like a Europe over the last few thousand years have minute asparagus covered with scale-leaves considerably extended and stabilised lliut, on emerging from the soil, become very favourable orchid habitats (chalk grasslands, •.mall green leaves normally arranged in a heaths, garrique, hay meadows...). Agricultural rosette. Underground, the protocorm becomes intensification has had a negative effect n root-tuber and enlarges in species with a on these habitats and is very unfavourable root-tuber and elongates in those with a for orchids. i lii/.ome. Two to 15 years pass between germi- Growth i ml ion and production of the first flower, de Most European orchids are autotrophic; with pending on species and soil conditions. green leaves, they manufacture nutrients from soil minerals via their own photosynthesis but, Annual cycle for many, probably also use their mycorrhiza. A few are mycoheterotrophic, feeding on de All orchids are perennial, capable of living composing organic substances [editor’s note; iniiiiy years and flowering each year if condi mycoheterotrophes acquire nutrients from tum:; are favourable. They often appear above fungi, which in turn acquire them from the de I'lduiul only briefly, flowering and fruiting in a composition of organic matter or from other liw weeks and then disappearing; most of their living plants.]; in this situation, considered a I lie cycle takes place underground (see Fig.3, further evolutionary step, the radicular system p. 10). alone is required, the leaves, now unnecessary, 9 Figure 3. Diagram of the annual cycles of Early Spider orchid Ophrys sphegodes (top) and Green-winged orchid Orchis morio (bottom) seen at 850 metres altitude in the Vercors (Ga, Isere, France). Each square represents the sit uation at the start of the month, given in roman numerals. In the Ophrys, a small rosette of leaves pushes out of the ground as early as October and is visible all winter, frost often blackens these first leaves. The second tuber starts to develop in November but does not start swelling until April, when the floral spike grows feeding on the over-winter ing tuber. The plant is in full flower in May; the second tuber is now almost the size of the first which, now emptied of nutritive substances, crumples. In June seeding is nearly completed, only a single discoloured flower remains on the top of the floral spike; the leaves brown, the winter tuber is exhausted, shrivelled; the second tuber, whitish, full of nutrients is at its maximum size; on the spike, seeds are freed from the capsules. In July, the floral spike browns, dries and collapses; the winter tuber is absorbed and disappears, the second tuber inclines upwards; in August a new plant forms at its summit. The Green-winged Orchid has a similar cycle but the winter tuber, at its maximum size in August, emits a new plant bud that does not emerge until April; the plant stays invisible for 6 or 7 months. Orchids with a rhizome have a similar cycle. can be reduced, an advantage as the plant phyll (Epipogium, Neottia nidus-avis) whilst needs less energy for growth. A few sapro- others retain a little (Limodorum, Coral- phytic species are totally devoid of chloro- lorhiza). Certain autotrophic species occasion- 10