Orchids of Europe, North Africa and the Middle East

640 Pages·2016·104.13 MB·English

by  Delforge P.

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