March 1997 North American Native Orchid Journal

NORTH AMERICAN
NATIVE
ORCHID JOURNAL
______________________________________
Volume 3 March
Number 1 1997
a quarterly devoted to the orchids of North America
published by the
NORTH AMERICAN NATIVE ORCHID
ALLIANCE
* * * * * *
* * * * * *
IN THIS ISSUE:
The Genus Cypripedium
NORTH AMERICAN NATIVE
ORCHID JOURNAL
(ISSN 1084-7332)
published quarterly in
March June September December
by the
NORTH AMERICAN NATIVE ORCHID ALLIANCE,
Inc.
a group dedicated to the conservation and promotion of our
native orchids
Editor: Paul Martin Brown

Assistant Editor: Nathaniel E. Conard
Editorial Consultants:
Philip E. Keenan
Stan Folsom
Production Assistant:
Nancy A. Webb
The Journal welcomes articles, of any length, of both a scientific

and general interest nature relating to the orchids of North
America. Scientific articles should conform to guidelines such as
those in Lindleyana or Rhodora. General interest articles and notes
may be more informal. Authors may include line drawings,
and/or black and white photographs. Color inserts may be
arranged. Please send all inquiries or material for publication to
the Editor at PO Box 772121, Ocala, FL 34477-2121 (mid June -
August: PO Box 759, Acton, ME 04001-0759).
1999 Membership in the North American Native Orchid Alliance,

which includes a subscription to the Journal, is $26 per year for
United States addresses, $29US in Canada and $32US other
foreign countries. Payment should be sent to Nancy A. Webb, 84
Etna St. Brighton, MA 02135-2830 USA. Claims for lost issues or
cancelled memberships should be made within 30 days.
NORTH AMERICAN NATIVE
ORCHID JOURNAL
Volume 3 March
Number 1 1997
CONTENTS
NOTES FROM THE EDITOR
1
THE NEED FOR GLOSSARIES
The Slow Empiricist
3
THE GENUS CYPRIPEDIUM
John W. Doherty
Guest Editor: William Steele
5
GLOSSARY
121
LOST & FOUND
129
LOOKING FORWARD
June 1997
134
Color Plates
Fig. 1 Cypripedium arietinum; Fig. 2 Cypripedium candidum p.105
Fig. 3 Cypripedium xandrewsii nm. andrewsii; Fig. 4
Cypripedium xandrewsii nm. Favillianum p.105
Fig. 5 Cypripedium parviflorum var. makasin; Fig. 6
Cypripedium parviflorum var. pubescens p.107
Fig. 7 Cypripedium kentuckiense p.108
Fig. 8 Cypripedium montanum); Fig. 9 Cypripedium
parviflorum var. pubescens flat-petalled form from Red Deer,
Alberta p.109
Fig. 10 Cypripedium reginae; Fig. 11 Cypripedium reginae pale
form p.110
Fig. 12 Cypripedium arietinum; Fig. 13 Cypripedium
xandrewsii ‗Gunter Preusse‘ CCM/AOS p.111
Fig. 14 Cypripedium xandrewsii ;Fig. 15 Cypripedium
parviflorum var. pubescens p.112
Fig. 16 Cypripedium parviflorum var. makasin ‗Sara Cond‘s
Flower‘ CCM/AOS p.113
Fig. 17 Cypripedium reginae p.114
The opinions expressed in the Journal are those of the authors. Scientific
articles may be subject to peer review and popular articles will be examined
for both accuracy and scientific content.
All drawings inthis issue are by Stan Folsom
Volume 3, number 1, pages 1-137; issued March 1, 1997.
Copyright 1997 North American Native Orchid Alliance, Inc.
COVER: Cypripedium reginae by Stan Folsom

NOTES FROM THE EDITOR
This issue is a major departure from our

previous issues, in that we are devoting the entire
Journal to one comprehensive work on the genus
Cypripedium. All of the North America species and
many additional species from other parts of the world
are included. To better understand the scientific terms
in the article, the Slow Empiricist suggested that a
glossary might be helpful. After careful consideration,
and with the author‘s permission, I encouraged the
Slow Empiricist to supply this issue with such a
glossary. You will find it at the end of the major
article, but I have included the Slow Empiricist‘s
explanations at the conclusion of this introduction. In
addition there are several longer footnotes to assist in
clarifying some of the terms and concepts.
This issue would not have been possible

without the help of several people. Bill Steele
generously gave of his time as ―guest editor‖ and went
through John‘s manuscript in fine detail, making
several helpful suggestions. Stan Folsom and assistant
editor Nat Conard both spent considerable time
helping to edit and format the issue. Stan‘s line
drawings have been used throughout as well as John‘s
color photographs. It has been an enormous task but
1
well worthwhile. I have not quite decided if one
article is simpler to prepare for the Journal as opposed
to the usual number of articles and authors the Journal
includes, but I enjoyed the challenge!
I trust you will find the work on the genus

Cypripedium elucidating and the glossary helpful.
Paul Martin Brown

15 Dresden Street
Jamaica Plain, MA 02130-4407
e-mail: NAORCHID@aol.com
2
THE NEED FOR GLOSSARIES
The Slow Empiricist
A few articles back (Jargon? 2(3):228-238), I

covered many terms to help beginners understand
some of the words that the more scientific members of
the orchid world use in their regular conversation. As
you become more familiar with the language that is
used, the meanings become much clearer and a
specificity tightens the descriptions. When the
following splendid article came in about the entire
range of the genus Cypripedium in the world, and
North America in particular, I was excited about the
possibilities for adding new knowledge to my
understanding. As I read through the article, I realized
that, to fully appreciate it, a glossary seemed
invaluable. I pulled out the words or terms that I
needed to understand better, let the computer
alphabetize them, and used the built-in dictionary to
give me precise definitions for the words. With Paul
Martin Brown‘s help, we further pulled out words and
phrases to send to the author so he could explain
exactly what he had in mind with his use of the terms.
(Incidentally, he was thrilled that people wanted to
3
have a glossary so that more people would be able to
use his valuable work.) Ginny Magee, a retired
biology professor, proofread the glossary and
provided me with further information which has been
added to the definitions.
The result is the glossary which is appended to

this work. If you see an asterisk before a particular
word or phrase, you should be able to find its
definition by consulting the appropriate page of the
glossary. (It is in alphabetical order, remember?)
I was very wary of injecting my ideas into this

fine piece of work; however, I have found that many
people have welcomed the idea, and since this
included the author, I persevered. I hope the glossary
is helpful to you and increases your understanding and
enjoyment of this comprehensive work.
The Slow Empiricist
4
THE GENUS CYPRIPEDIUM
a botanical and horticultural overview
John W. Doherty
Introduction
Terrestrial orchids are currently enjoying a

surge in popularity among horticulturists and
orchidophiles around the world. This is due in large
part to increased accessibility to the ever expanding
information base regarding this intriguing group.
However, it is to advances in propagative technology
that must be given most of the credit for making
terrestrials so much more widely available. The
*genus Cypripedium remains one of the most beloved,
interesting, beautiful and sought after members of this
decreasingly exclusive group and it shall be the
subject of this paper.
A circumglobal genus of the subfamily

Cypripedioideae, species can be found in the majority
of *temperate climes in the northern hemisphere, with
the exception of the Cypripedium irapeanum group,
Doherty: The Genus Cypripedium
6
Mexican natives, and the recently described C.

subtropicum from Tibet. With as few as 30 different
species, or as many as 50, all remain *deciduous
during a pronounced cold period in the winter months.
The majority of species withstand below freezing
temperatures for a prolonged period. *Tropical
relatives include the other slipper orchids, in the
*genera Paphiopedilum, Mexipedium,
Phragmipedium, and Selenipedilum.
Pollination in North American Cypripedium has

been extensively studied by Nilsson (1979), among
others, who showed that it is achieved through
deceptive floral lures, as opposed to floral rewards
which are conspicuously absent in this genus. Flower
colour and false nectar guides surely aid in deception
of the various species of bees that are known
pollinators; however, the most intriguing adaptation is
the use of odour. Fragrance emitted by mature flowers
is surely of a complex and incompletely understood
nature; however, it is likely that it is partially
composed of food-based attractants. Furthermore, it
has been found that a series of acetates dominates the
fragrance and bears significant chemical *homology to
certain bee *pheromones. Honeybees (Apis mellifera)
have no fewer than 36 documented pheromones, with
others remaining unknown, each with a specific
purpose in the bee communication process. It is
reasonable to assume that other bee species, and even
other genera, have a comparably complex method of
7
communication. Besides foraging for food, bees also

seek out potential future nest sites. Potential nest sites
tend to be somewhat cavernous, as may the lip of a
slipper orchid appear to be. These potential sites are
marked with a particular pheromone indicating such,
which would then work to attract *conspecifics,
prompting further investigation. It is suggested that
this genus, and therefore perhaps other slipper orchids,
have ―tapped‖ into this behavioral pattern of these
bees as manifested by their floral structure as well as
by the pheromonal composition of their fragrance.
Such would truly be a marvel of evolutionary
processes, as several bee species have been shown
responsible for pollination: Bombus and Psithyrus
ashtoni queens, *halictid bees, Andrena spp. and
Lasioglossum spp., as well as Megachile (see Catling
and Catling, 1991, for an excellent review of all
documented North American orchid pollinators), each
probably with its own unique pheromone system.
This adaptation would necessitate fine tuning at the
species level (for which orchids are famous) to ensure
efficacy of prolonged attraction to both the structure
of the lip, and the pheromone-tainted fragrance.
Unfortunately, this tidy hypothesis relies

partially upon conjecture and definitely requires
further investigation, at a molecular level, of the
orchids as well as of the bees in question, in order to
confirm pheromone/fragrance homology, in addition
to the behavioral role of the pheromone in question.
8
There are a great many misconceptions

regarding this genus as a whole; this paper is an
attempt to clarify some of those misconceptions and
hopefully further the general understanding of this
group. To achieve this, there will be three sections:
Part I will provide a brief *taxonomic overview of the
genus; Part II will give a brief explanation of
*mycorrhizal fungus and its integral role in the
development of these orchids; finally, Part III will
look at some current stances on propagative
technology and *cultural practices.
This paper is by no means intended to act as a

definitive reference, but rather as a botanical,
*mycological, and horticultural overview. The
references listed at the end of each section are felt to
be some of the best available, and are recommended to
any reader with a specific interest in that area.
References
Catling, P.M., and V.R. Catling. 1991. Pollination in North

American orchids. Lindleyana. 6:187-210.
Nilsson, L.A. 1979. Anthecological studies on the lady‘s-

slipper, Cypripedium. Bot. notiser. 132:329-347.
9
Part I
A Brief Taxonomic Overview
The most recent treatments of this genus are by

Cash (1991), and Perner (1996), who provide a
detailed and thorough *infrastructure, into which they
have, for the most part, neatly fit the various species.
Cash‘s outline will be followed here. Species listed
are those with which I have had some sort of first hand
experience, preferably those that I have observed in
the wild. The kind of knowledge to be gained from
direct habitat observation cannot be surpassed by any
number of books. For species not listed, readers are
directed to Cash (1991).
Subgenus Criosanthes
Cypripedium arietinum R. Brown (Fig. 1)
Ram’s-head lady’s-slipper
This species has been the source of some
taxonomic controversy as it was once thought to differ
significantly enough from other species to warrant its
own genus, proposed to be Criosanthes. Reasons for
this exclusion include entirely unfused *sepals, a
*spurred lip, and *anther-like *staminode. These
factors alone do not warrant a separate distinction;
however, it has been suggested that its ephemeral
bloom of only a day or two represents a significant
evolutionary step in pollination biology and therefore
Doherty: The Genus Cypripedium: Part I
justifies taxonomic distinction. In reality, the bloom is

not at all ephemeral. Under cultivation, I have found
that flowers remain open for up to a week under
average to above average temperatures; even longer if
kept cool.
Cypripedium arietinum has a limited range

centred about the Great Lakes region. The species
occupies two distinct habitats, the first being cool
cedar swamps, often floored with sphagnum moss.
Here it is usually found singly and infrequently.
However, extremely large individuals can develop.
The second habitat can be either sandy dunes or
limestone *barrens near shorelines, where offshore
breezes provide necessary cooling. It is in this habitat
that C. arietinum will develop huge colonies with
individual, smaller-statured and infrequently clumped
plants numbering into the hundreds.
Cypripedium plectrochilon Franchet is a

vicarious species1, native to a small region in Asia,
1
Cypripedium arietinum and Cypripedium plectrochilon are vicarious
species. They are remarkably similar in nearly every respect, except that they
are found in drastically different parts of the world. They do NOT result from
convergent evolution, as they most likely arose from common stock. Nor do
they exhibit divergent evolution, as they have not significantly diverged, or
become different. Rather, they have evolved as parallel species, and
maintained their similarities, most likely due to very similar environmental
pressures experienced in each of their geographically distinct habitats.
11
Cypripedium arietinum R. Brown

Ram’s-head lady’s-slipper
12
and is felt by some to be conspecific with C.

arietinum. Sing-Chi (1983) and Perner (1995) have
reviewed the taxonomic issues concerning this pair (as
well as another vicarious pair, C. flavum and C.
reginae) and feel C. plectrochilon to be sufficiently
different to warrant species level distinction. Perhaps
the closely matching distribution patterns, as well as
very similar ecological preferences shared by each
pair (but not between pairs), combined with the
relatively primitive structure of these four species,
suggest evolution from a common stock, which
formerly had a common *areal range. Furthermore,
―The modern distribution patterns of the vicarious
species are by no means fortuitous, but result from
similar migration patterns.‖ (Sing-Chi, 1983). The
*phytogeographic implication of these relationships is
a fascinating topic on which much light could be shed
through use of modern molecular genetic analyses.
Subgenus Cypripedium *Alliance Cypripedium

Cypripedium calceolus L.
Eurasian yellow lady’s-slipper
Cash (1991) lists all of the North American
variants, as well as the Eurasian plants, under this
species. Sheviak (1994, 1995, 1996a, 1996b) has
since reclassified the North American species as a
separate *complex under C. parviflorum;
consequently they shall be discussed under that
heading. Cypripedium calceolus, however, remains
the name for the Eurasian species. This species is
13
readily distinguished in the trade from its North

American relatives by its white staminode. Preferring
*calcareous soils, this species can become locally
abundant and enjoys a variety of shaded or semi-
shaded habitats; however, the one common feature
appears to be a preference for an alkaline *substrate.
Cypripedium candidum Muhl. ex Willd.2 (Fig. 2)

Small white lady’s-slipper
This species is native to northeastern North
America and probably most commonly distributed in
Michigan. Its reputation for sunny habitats is well
deserved as it is never found in much shade, unless at
the border of a mucky meadow that is gradually being
overgrown, where it will not flourish for much longer.
Easily the most sun-loving of all our native species, C.
candidum consequently has a high moisture
requirement. Water can often be found in a small hole
dug adjacent to healthy plants. Black, mucky soil
seems to be preferred; however, it has also been found
in heavy *clay substrates. Native to prairie-type
environments, this species can thrive under controlled
burn techniques used in management of prairie
conservation areas. If allowed to grow continually,
prairie grasses can choke out thousands of plants to a
handful in a matter of one season. Carefully timed
2
Muhlenberg was the first to name this species but did not publish a valid
description. Willdenow was the first to validly publish the name according to
the International Rules of Botanical Nomenclature. His name is preceded
by an ―ex‖ to indicate that it was taken ―from‖ Muhlenberg‘s description.
14
burns can astonishingly replenish such populations in

as fast a time.
It is suggested by Cash (1991) that this species

―...is to all appearances an *alba (white flowered) form
of the small yellow lady‘s slipper, Cypripedium
calceolus var. parviflorum [parviflorum var. makasin
or parviflorum var. parviflorum].‖ Unfortunately
there are a number of *morphological and ecological
differences between the two that point to the contrary.
The leaves of C. candidum at flowering remain furled
and held tightly around the stem. This may be in an
effort to reduce high *transpiration rates that would
surely be encountered at such an early stage in the
season where neighbouring grasses are not providing
much-needed shelter from direct sun. The leaves of C.
parviflorum are much more expanded at flowering. C.
candidum is very rarely two flowered, whereas C.
parviflorum is frequently so. C. candidum does not
produce *albinistic flowers. Petals are often
pigmented with browny greens; the interior of the
pouch is often marked with red. It is seen, however,
that plants in extremely high light conditions have
―bleached‖ looking flowers where normally pigmented
flowers are pure white and yellow, mimicking
*albinism. The greatest reason to discount this
hypothesis, however, is that of habitat differences.
Neither species is ever prevalent in the other‘s habitat.
The only time one could be found in the habitat of the
other is in an overlap situation, where a much shadier
15
region, such as a woodlot, is found immediately

adjacent to an open type prairie/meadow. Such a
pronounced habitat contrast is the strongest argument
against the alba-race hypothesis.
In addition to floral pigment, plant stature also

varies according to levels of sunlight. The shortest
plants are always found in sun, whereas taller plants
are found in shadier transition zones. This is not, as
one might expect, genetically fixed; rather plant height
is an extremely variable characteristic highly
dependent on a number of environmental variables,
sunlight being one of them. This principle is not at all
unique to this particular species, but rather a
characteristic that can be generalized to most of the
genus. The environmental factor certainly confounds
taxonomic treatises that attempt to use plant stature as
a distinguishing character for a certain species,
particularly in the Cypripedium parviflorum complex.
There are, of course, limits to a plant‘s potential height
response, not entirely discounting the use of height in
*taxonomy.
Cypripedium xandrewsii Fuller3 (Fig. 3, Fig. 4)
is a natural *hybrid (*nothospecies) that occurs in the
previously described situation where there is habitat
overlap between C. candidum and C. parviflorum and
its varieties, var. makasin and var. pubescens. The net
3
When a hybrid has been given a name, this is indicated by placing an x
(denoting multiplication) before the specific epithet, as in Cypripedium
xandrewsii.
16
result is a blend of both parents that can sometimes

yield striking combinations, the most notable being
white-lipped multiple flowers with dark chocolate
brown petals, held above the foliage. There are,
however, all types of intermediates with pouch colours
ranging from pure white to pale yellow, and petals
from yellow green to chocolate brown. Plant stature
and *floral stance are comparably variable.
Cypripedium xfavillianum is the previous name used
to describe the nothospecies resulting from a cross
between C. candidum and C. parviflorum var.
pubescens before the yellow complex was reclassified
to a single species, necessitating the use of a single
name. The correct current nomenclature for the
hybrids would be Cypripedium xandrewsii Fuller
*nm. xandrewsii and Cypripedium xandrewsii Fuller
nm. favillianum (Curtis) Boivin. A third *nothomorph
has also been described, Cypripedium xandrewsii
Fuller nm. landonii (Garay) Boivin (C. xandrewsii
nm. xandrewsii x C. parviflorum var. makasin).
Cypripedium cordigerum D. Don

Of all the Asian species, this high-elevation
native of India is most closely related to the
Cypripedium parviflorum complex. Its floral structure
appears to be quite similar; however readers are
referred to Cash (1991) for specifics regarding habitat
information and distribution.
17
Cypripedium kentuckiense
Kentucky lady’s-slipper
18
Cypripedium kentuckiense C. Reed (Fig. 7)

Kentucky lady’s-slipper
Cash‘s structure lists this as a *variety of
Cypripedium calceolus (C. parviflorum); however,
there are a number of morphological characteristics
that distinguish it as a separate species in addition to,
most importantly, its discrete habitat range preference.
C. kentuckiense is a southern species, with its range
dipping south into Texas, with Kentucky actually
representing the northern limit of its distribution. It is
a warm, long-season grower and cannot be forced into
the shorter seasons experienced up north; it is the first
up and last down. It can be found in shady locations,
often near a source of running water, and apparently it
is often inundated with water. The ability to survive
submergence alone likely constitutes reason enough
for species distinction; however, floral characteristics
lend further support.
Plants are always large statured. Flowers are

always very large, the smallest being as large as, if not
larger than, the biggest of the C. parviflorum complex.
The pouch is, on average, the size of a chicken egg
and ranges in colour from butter yellow to pure white.
There are seldom two flowers, and floral stance is
characteristic. Stems often appear as though they are
―bowing‖ in an effort to support the flowers. The
*dorsal sepal is rarely vertical, usually drooping far
over the pouch. Flowers give the impression that they
19
are hanging. Petals can become quite long, thin, and

corkscrewed.
Cypripedium montanum Dougl. ex Lindl. Fig. 8)

Mountain lady’s-slipper
This is a montane species with a specific
habitat: it prefers open woods, either moist or dry, at
high elevations. This results in its restricted range in
the Rocky Mountains in British Columbia south into
California. Plants are frequently two-flowered and
sometimes three-flowered, with a white lip and petals
of a softer brownish green than that seen in C.
candidum or the C. parviflorum complex.
Again, Cash (1991) suggests this to be an alba

race of Cypripedium parviflorum. Unfortunately, the
previously mentioned characteristics make this
impossible. First of all, it is not albinistic, for the
petals show significant brown colour in addition to
red/purple pigment inside the pouch. It is rarely single
flowered. However, most importantly, its specific
habitat preference distinguishes it from C.
parviflorum. There is occasional habitat overlap,
however, resulting in the production of C.
xcolombianum Sheviak, a natural hybrid between the
two.
One would imagine a white-flowered race of a

particular species to exist somewhere within the range
of that species. Originating from a single albinistic
20
mutant, differing from the rest of the population

significantly enough to warrant attention from a
different pollinator, that individual would proliferate
and eventually establish its own separate population.
Such cannot be said for Cypripedium montanum nor
for C. candidum, as they both exhibit a high degree of
ecospecificity such that their habitat of choice is
normally very different from that of C. parviflorum. It
is possible, and likely probable, that these two species
did initially arise from an older precursor of the C.
parviflorum complex; however, they have been
through the evolutionary gears of *speciation long
enough to become two very distinct and different taxa.
Cypripedium parviflorum Salisb.

Yellow lady’s-slipper
Without a doubt this complex has attracted the
greatest amount of taxonomic attention of any in the
entire genus. This is due in large part to its huge
North American distribution, and consequent
variability. It is likely, however, that the astounding
ability of plants to change morphologically depending
on environmental conditions is the single greatest
confounding factor. Traditionally, classification is
based on characteristics such as plant stature, leaf size
and stance, floral size, colour and morphology, and
most importantly with the slipper orchids, staminode
shape. In subspecies where this last characteristic
remains fairly consistent throughout the entire range
21
of this complex, one is required to rely on the

previously listed features. Unfortunately, these
features can vary tremendously within an individual.
Plants from open, sunny situations tend to be shorter
statured, with smaller, narrower leaves and smaller,
paler flowers (some may have larger flowers in
sunnier locations). Conversely, the same plant facing
shady woodland conditions might have larger, broader
leaves on a tall stem. Flowers may be much larger and
darker coloured. This stunning environmental
adaptability has clearly provided this complex with a
strong competitive advantage, perhaps explaining the
wide geographic distribution. It makes excellent
reproductive sense to be able to adapt to an
immediately open situation as the result of a
disturbance, or a gradually shaded situation, as the
majority of orchids are not found in *static, climax
stage habitats4. It is, therefore, the competitive
strength of this complex that weakens taxonomic
treatments. Sheviak (1994, 1995, 1996a, 1996b)
appears to be the first to demonstrate this principle in
4
A climax stage habitat is any habitat occupied by species characteristic of
that stage, their most notable characteristic being the ability to survive in the
shade of their ―parents‖ until they are large enough to displace or replace
their forefathers. This is in particular reference to tree species. Herbaceous
species, such as orchids, can be found in these habitats as well. Climax stage
habitats are very static (as opposed to dynamic) and represent the top of the
―habitat evolutionary ladder,‖ assuming some extreme force, such as a forest
fire, earthquake, or human encroachment does not act on the habitat and
change its ―balances‖.
22
cultivated plants; his treatment of the complex is

outlined here.
Cypripedium parviflorum var. parviflorum

Southern small yellow lady’s-slipper
This variety is restricted in range to the eastern
United States and is found in the driest habitat of the
three varieties. It has medium size flowers
characterized by dark brown petals that tend to hold
their colour along the entire length.
Cypripedium parviflorum var. makasin (Farwell)

Sheviak (Fig. 5)
Northern small yellow lady’s-slipper
This is the northern, smaller flowered variety
that prefers wetter sites, often in black, mucky soil.
Again, the petals tend to be dark brown, perhaps
breaking towards the middle of the flower into
blotches, revealing the greeny-yellow basal colour of
the petals.
Cypripedium parviflorum var. pubescens (Willd.)

Knight (Fig. 6)
Large yellow lady’s-slipper
This is the most variable of the three varieties.
It can have dark brown petals, however, usually not of
the intensity seen in the other two varieties. Colour
ranges from pure yellow, to yellow green, to green
brown. Flowers tend to be significantly larger. Plant
23
stature has the greatest potential here to be large;

however, very diminutive plants can also be found.
An apparently dwarf strain within the complex

found on windswept shorelines in Newfoundland was
at one point given the varietal status var.
planipetalum, on account of its flat petals, thought to
be unique to that population. Based on the previously
discussed environmental adaptability of this complex,
this varietal distinction has been abandoned. A
number of sources have reported flat-petalled plants
far removed from the Newfoundland populations, in
places such as New York. Experience with cultivated
plants confirms this. Figure 9 shows a flat-petalled
plant, very similar to those thought to be var.
planipetalum. This particular individual was collected
in Alberta amongst a normal-flowered population.
Strangely enough, at the time of collection, it was no
different from any others in that population as its
petals were twisted. The flower came up flat-petalled
the following season, likely the result of an altered
environmental factor.
Please see Cash (1991) for additional members

of this alliance: Cypripedium henryi Rolfe, C.
microsaccus Kraenzl. and C. shanxiense S.C. Chen.
24
Cypripedium fasciculatum
Clustered lady’s-slipper
25
Alliance Corymbosa
Cypripedium fasciculatum Kellogg ex S. Watson
Clustered lady’s-slipper
This native of the northwestern United States
can be found at high elevations in *coniferous forests
and shrubby thickets. Its two to four plus flowers are
small, greenish-brown and, according to Fred Case,
look like peeled grapes. They tend to droop, pointing
towards the ground, making observation difficult;
however this is likely in an effort to attract specific
pollinators. Successful pollination results in
substantial straightening and lengthening of the stem,
thereby enhancing seed dispersal.
Alliance Macrantha
Cypripedium macranthos5 Sw.
Cypripedium macranthos is a widespread
species typifying a very large complex requiring
further taxonomic clarification, as there are a number
of closely related species whose interrelationships
remain to be fully worked out. A major barrier to
clarification is a political one, as much of its range,
along with related species, is within China. This
complicates and reduces the opportunity for field
observation. As a result, there is much to be learned
about this entire group.
5
Cypripedium macranthos Sw. is a synonym for C. macranthum Sw. and
the preferred spelling. See Cribb (1994) and Perner (1966). PMB
26
Preferring cool climates, Cypripedium

macranthos can be found in high-light situations in
either deciduous shade or *montane grasslands. Once
considered a separate species, var. rebunense is a
closely guarded Japanese population of pure yellow
flowers. A deep, solid, purple-red form is known as
var. hotei-atsumorianum. The Asian range of C.
calceolus overlaps in some regions with C.
macranthos; the natural hybrid between the two, C.
xbarbeyi, occurs in these situations. Apparently this
hybrid exhibits extensive variability, with colours
ranging from pure white to deep wine red.
Please see Cash (1991) for remaining members

of this alliance: Cypripedium amesianum Schltr., C.
corrugatum Franch., C. fasciolatum Franch., C.
himalaicum Rolfe, C. lanuginosum Schltr., C.
speciosum Rolfe, C. thunbergii Bl., C. tibeticum King
ex Hemal., C. ventricosum Sw., C. wilsoni Rolfe, and
C. yunnanense Franch.
Alliance Flabellifolia
Cypripedium acaule Ait.
Pink lady’s-slipper; stemless moccasin flower
Likely the most *acid-loving of all our native
species, C. acaule has a very wide distribution from
northeastern Alberta, into the Northwest Territories,
over to the Atlantic coast and as far south as Georgia.
There are two general and very different trends with
respect to habitat. The first is high and dry woods
27
Cypripedium acaule
Pink lady’s-slipper; stemless moccasin flower
28
which are often sandy. The second habitat is

somewhat boggy, usually with some type of sphagnum
nearby. These two apparently divergent habitats
maintain an acid medium as their common
denominator. White-flowered forms of this species
exist and are most prevalent in the northeast.
Cypripedium farreri W.W. Smith

This large-flowered dwarf species *colonizes
limestone cliffs in the Yunnan province of China and
perhaps represents an interesting evolutionary link
between members of the genus. Unlike other
members of this alliance, a number of leaves are
produced alternately on a vertical stem, as opposed to
two *sub-*opposite leaves close to the ground (as in
Cypripedium acaule) or a short distance from it (as in
C. formosanum and C. japonicum). In addition, the
flowers completely lack the central *labellar fissure
common to these three species. Interestingly enough,
the pouch *orifice is toothed and somewhat *everted,
reminiscent of C. guttatum, yet there is sufficient
*infolding to link it to the alliance Cypripedium.
These differences urge its inclusion in a different
alliance. However, it has been kept here for lack of a
more appropriate place, as opposed to a distinction all
its own.
Cypripedium formosanum Hay.

Closely allied to C. japonicum, this species is
found in constant moisture on steep *shale slopes in
29
Taiwan, where it grows in bright shade. Conditions

apparently are acid. The paired leaves are unique in
that they appear as though someone neatly trimmed
the ends off with a pair of scissors.
Cypripedium margaritaceum Franch.

The paired leaves of this species closely hug the
ground and are marked with purple-brown spots, a
characteristic unique to this species and its closest
relative, Cypripedium lichiangense (previously C.
daliense before Cribb‘s recent reclassification). It is a
montane species preferring moist, well drained slopes
in pine or mixed pine forests (Cribb & Chen 1994).
See Cash (1991) for remaining members of this

alliance: Cypripedium bardolphianum W.W. Sm. &
Farrer, C. debile Reichb. f., C. ebracteatum Rolfe, C.
elegans Reichb. f., C. japonicum Thunb., C.
micranthum Franch., C. palangshanense Tang &
Wang, and C. wumengense S.C. Chen.
Alliance Guttatum
Cypripedium guttatum Sw.
Spotted lady’s-slipper
Cypripedium yatabeanum Makino
Yellow spotted lady’s-slipper
There exists some confusion in regard to these
closely allied species. This can be largely attributed to
the natural hybrid formed between the two,
Cypripedium xalaskanum P.M. Brown. Information
30
recently provided by Paul Martin Brown, based on

extensive field experience, has clarified this situation.
Cypripedium guttatum is a small woodland species,
producing one (or rarely two) white flowers with
cranberry blotching. Flowers are held above two or
three *alternate leaves. The underground *rhizome
can be quite extensive and wandering, producing a
large number of stems, or individual leaves over a
large area. In contrast, C. yatabeanum is a much
larger-statured plant in every respect. Flowers are
larger and basically yellow, with rusty-brown to green
markings. Petals are shaped differently, appearing
somewhat pinched near the tips. In addition to
morphological differences, habitat ranges and
preferences are distinct as well. Cypripedium
yatabeanum is primarily an Asian species, found in
North America only on Kodiak and neighboring
islands. It has apparently never been found on the
mainland, and prefers much sunnier locations.
Cypripedium xalaskanum, and the results of its

subsequent interhybridization, can be found on both
the Alaskan mainland and the islands. It exhibits the
entire range of intermediates between the two extreme
forms, represented by the two different species.
Floral structure of these species is unique in the

genus. The front margins of the lip are polished and
everted, without any infolding characteristic of the
genus. This feature is very reminiscent of
31
Cypripedium guttatum
Spotted lady’s-slipper
32
Cypripedium yatabeanum
Yellow spotted lady’s-slipper
33
Paphiopedilum, leading some to suggest that this

species might represent a possible evolutionary link
between the two genera. When considering
*vegetative and geographic similarities, this linkage
becomes very unlikely. It is more reasonable to
assume that this is an example of convergent
evolution6.
Plant characteristics having a pronounced effect

on reproductive strategies of a species, such as floral
traits, tend to be rather *dynamic as minor alterations
can have drastic outcomes on population trends. It is,
therefore, highly unlikely that a dynamic feature of a
population7 such as ―lip curvature‖ would be
maintained over the extremely long evolutionary
period where vegetative changes might occur from the
tropical to temperate plant habit (not temperate to
6
This is an evolutionary process whereby two organisms of differing origins
evolve towards a similar feature. A good example of this is the eye of the
octopus in comparison to the human eye. The eye structure in an octopus is
remarkably similar to that in humans. It is obvious that humans did not
evolve from octopi and maintain this trait. Millions of years ago the
respective ancestors branched off of the evolutionary path in response to
different environmental pressures. At some point, both "lineages" were faced
with similar environmental pressures in some certain, particular aspect,
resulting in the gradual development of very similar structures.
7
Some features of a population change very little over time, such as stem
thickness (assuming constant environmental conditions!), leaf shape, or rate
at which a particular enzyme functions. These can be loosely referred to as
static or very slow moving traits. A dynamic trait is something that changes
relatively rapidly from generation to generation, such as floral colour
patterns. Dynamic traits are not as "fixed‖ as static traits.
34
tropical). If one were to search for relatedness

between Cypripedium and other genera based entirely
on floral structure (a hazardous approach), it would be
best to look to members of the genus Phragmipedium,
as they exhibit very similar infolding of the lip so
prevalent in Cypripedium. It is most advisable
however, to look to tropical intermediates such as C.
irapeanum or C. subtropicum and their relationship
with other genera, including Selenipedilum, in order to
understand fully the genetic ―organization‖ within this
subfamily.
Alliance Obtusipetala
Cypripedium californicum A. Gray
California lady’s-slipper
The most stunning feature of this yellow and
white species is its production of small flowers at
*leaf bracts along the length of the stem, a feature
unique in the entire genus. It has a very small range in
northern California and southern Oregon and is found
in open woods, usually near streams, often in the
company of the cobra lily (pitcher plant),
Darlingtonia californica. Plants tend to form clumps
and exhibit very little floral variation.
Cypripedium dickinsonianum Hagsater

Dickinson’s lady’s-slipper
Cribb (1996) has apparently cleared up a degree
of confusion regarding this Central American group
that has long been considered a single species, the
35
Cypripedium californicum
California lady’s-slipper
36
Mexican yellow lady’s-slipper, Cypripedium

irapeanum Llave & Lex. Morphological and
geographic differences, however, pointed towards the
delineation of three different species: C. irapeanum,
C. dickinsonianum and the downy yellow lady’s-
slipper, C. molle Lindl. Distinctions between the
three are based on plant stature, leaf size and shape,
floral differences and geographic distribution. The
entire group is found throughout southern Mexico and
into Guatemala and Honduras. C. irapeanum can be
found in the drier parts of valleys, going without rain
from October to May and experiencing summer
temperatures from 30-35 C, dipping to less than 5 C in
the winter. It prefers clayey soils in pine/oak forests.
The flowers of this group are truly stunning. Medium
sized to extremely large bright yellow flowers are
produced consecutively, atop showy stems that can
vary in height from 30 cm to 1.5 m. Flowers are very
reminiscent of Paphiopedilum armeniacum, whereas
flowering habit, unique in this genus, is characteristic
of species in the Cochlopetalum group of
Paphiopedilum, such as P. primulinum or P.
glaucophyllum.
Cypripedium passerinum Richards

Sparrow’s-egg lady’s-slipper
This is a northerly species, found from Alaska
eastward into Quebec, remaining in the northern
regions of the provinces along the way, with the
exception of Alberta and British Columbia, where the
37
cooler temperatures afforded by the increased

elevation of the Rockies permits its occurrence at
lower latitudes. The plant can be found in bright,
open clay woods, usually with some source of water
nearby, or in gravelly outwash plains. Flowers are
small and self-pollinating, the only North American
species in the genus with such a reproductive strategy.
Consequently, there is very little variation within the
species. *Ovaries are very swollen at blooming.
Shoots are produced along a rhizome with a strong
tendency to branch, and sprout from *dormant eyes.
Frequently found accompanying this orchid in the
western part of its range is the small round-leaved
orchis, Amerorchis rotundifolia, often numbering into
the hundreds.
Cypripedium reginae Walter (Fig. 10)

Showy lady’s-slipper
This moisture-loving native of the northeast is
often prevalent in the Great Lakes region. It can be
found in boggy, open forest edges in *neutral to
alkaline black mucky soils. Sun is a critical factor in
the *population ecology of this species. It seems to
require direct sun for only a portion of the day, as it is
seldom found in constant sun or full shade. It prefers
intermediate areas, such as boggy forest borders, or
clearings within such a forest. It is this type of
environment that Cypripedium reginae will quickly
colonize, forming very large groups of stunning
plants. There is a direct correlation between the
38
amount of sun received and population size; as soon

as its clearing or edge begins to grow in, the plants get
taller from season to season, until eventually
individuals begin to disappear completely. Copious
moisture is an additional requirement of this species,
and it is often found accompanied by tamarack, Larix
decidua. This orchid is arguably the largest and
showiest of the North American species. Leaves are
long, broad, and relatively massive, resulting in very
high rates of transpiration. This physiological luxury,
likely resulting in its increased overall stature, is
afforded by its wet habitats. It is possible that the
exceptionally *hirsute leaves are designed to be so in
an effort to combat further transpirational water loss.
Fred Case (1987) suggests that deer play an

important role in the reproductive biology of this
species.
Firstly, they tend to graze tree branches near

forest edges. This allows for greater sun penetration,
creating the optimal blend of sun and shade for C.
reginae, which can be found in large numbers in such
situations. Secondly, C. reginae inhabitations tend to
serve as winter deer yards, where the action of hooves
buries the orchid seed to an ideal depth for
germination. One to four flowers are produced, and
white-flowered variants are known. Cash (1991) lists
a variant, var. albolabium, lacking pigment in the lip.
My experiences flowering numerous *clones of this
39
species under cultivation makes such a distinction

questionable. The flower seen in Figure 11, although
slightly overexposed, could very easily be interpreted
to be such a variant as there is indeed very little
pigment on the lip. This flower is from the same plant
seen in Figure 10, but from the following season.
There is no genetic difference between the two. The
difference lies in the environmental conditions at the
time of flowering. The plant in Figure 10 experienced
what will be called a ―normal‖ spring, in Windsor,
Ontario, resulting in ―normal‖ flower development.
The following season, that same plant experienced
another ―normal‖ spring, this time in Guelph, Ontario.
If anything, the plant was kept a little cooler during its
development as Guelph is usually cooler than
Windsor. The week before flowers were to open, the
plant was brought to Windsor, which was then
experiencing a heat wave, resulting in abnormally
high temperatures for development. The result is the
flower in Figure 11. Extent of floral *pigmentation
may, therefore, be partly dependent on *ambient
temperature during floral development. The cooler the

temperature, the darker the flowers. That is not,
however, to say that genetic factors do not play a role.
They can indeed figure prominently into the extent of
pigmentation. The small plant in the bottom left of
Figure 17 illustrates this point, as all flowers seen
experienced the same environmental factors, however,
that particular clone appears to be significantly darker.
40
The principle of floral pigment intensity varying with

environmental factors (including *soil *pH) is not a
new one; however, this species appears to be
especially sensitive, a notion confirmed by other
growers.8
Cypripedium flavum Ward is a vicarious

species, with limited distribution in two regions of
Asia (Sing-Chi, 1983). Unlike the other vicarious pair
discussed (C. arietinum and C. plectrochilon) there is
little question of the two being different species.
Staminode shape and flower colour are quite different,
eliminating any such concerns. Apparently, however,
habitat preference and vegetative characteristics are
remarkably similar, confounding previous
*herbarium-based analyses.
Please see Cash (1991) regarding remaining

members of this alliance: C. flavum and C.
subtropicum Chen & Lang.
8
White-flowered forms of Cypripedium reginae are correctly known as
forma albolabium and occur sparingly within normally colored populations.
These plants, in addition to having pure white flowers, have paler green
leaves and retain this coloration for the life of the plants. I too have seen
pale colored flowers on plants that otherwise have had full colored flowers.
Often these pale flowers appear to be white but upon close examination show
slight traces of pink within the lip. Aside from producing a genetic sport, a
given plant does not change its color from year to year. Intensity of color
certainly can be variable. PMB
41
References and Recommended Reading
Case, F.W. 1987. Orchids of the Western Great Lakes Region.

Cranbrook Institute of Science, Bulletin 48.
Cash, C. 1991. The Slipper Orchids. Timber Press, Portland,

Oregon.
Cribb, P. 1996. The Genus Cypripedium in Mexico and Central

America. North American Native Orchid Journal. 2: 3-
18.
Cribb, P. & S.C. Chen. 1994. Chinese spotted-leaved

Cypripediums. Orchid Review 102: 321-323.
Luer, C.A. 1975. The Native Orchids of the United States and
Canada Excluding Florida. New York Botanical
Garden. NY.
Perner, H. 1995. Winterharte Fraunschue (1): Cypripedium

arietinum R. Brown und Cypripedium plectrochilum
Franchet. Die Orchidee 46(3): 103-108.
——. 1996. Die Gattung Cypripedium L. in einer

stammesgeschichtlichen Ubersicht. Jour. Eur. Orch.
28(1): 84-107.
Sheviak, C.J. 1994. Cypripedium parviflorum Salisb. I: The

Small-flowered Varieties. American Orchid Society
Bulletin. 63: 664-669.
——. 1995. Cypripedium parviflorum Salisb. II: The Larger-

flowered plants and Patterns of Variation. American
Orchid Society Bulletin. 64: 606-612.
42
——. 1996a. The Cypripedium parviflorum complex in North

America. North American Native Orchid Journal 2(4):
319-343.
——. 1996b. Cypripedium hybrids in the Russian Far East: The

Red Influence. part 1. Orchids 65(11): 1170-1175. part
2. Orchids 65(12): 1294-1299.
Sing-Chi, C. 1983. Two pairs of vicarious species of

Cypripedium (Orchidaceae) from eastern Asia and
eastern North America. Brittonia. 35: 159-163.
43
Part II
Mycorrhizal Fungus
Undoubtedly, the relationship orchids maintain

with their mycorrhizal partners is the most poorly
understood aspect of their entire biology. Research in
this area was quickly abandoned as Knudson
pioneered the techniques for *asymbiotic germination
of tropical orchids, completely eliminating any need
for mycorrhizal fungus in seed-sowing practices.
Interest was renewed in mycorrhizae as greater
attention was drawn to temperate terrestrial orchids,
due in part to increased horticultural pressures, as well
as the ecological threats, which put the continued
existence of these plants into question. *Temperate
terrestrials are usually resistant to conventional seed
propagation techniques. Effort has therefore been
made to understand better their mycological partners
and thus their reproductive biology, perhaps to
provide some of the keys to stubborn propagative
temperament.
In the orchid trade, both temperate and tropical,

there are an astounding number of misconceptions
regarding mycorrhizal fungus. Rather than list and
individually address them, a brief overview of what is
Doherty: The Genus Cypripedium: Part II
currently known, with specific emphasis on temperate

terrestrials, will be given.
Seed Germination
One of the most important characteristic
features of orchids is the minute seed they produce.
Their small size effectively facilitates dispersal over
long distances by wind. One of the disadvantages of
having such a small seed, however, is the inability to
store significant amounts of nutrients. An orchid seed
can consist of as few as 50 different cells, with storage
reserves being primarily proteins and *lipids. It has
been shown that in nature, seeds cannot *germinate
without the help of a mycorrhizal fungus.
The fungus usually enters the seed through what

is called the suspensor end. The testa, or seed coat,
completely envelopes the *embryo; with the exception
of the point at which it was attached to the ovary wall
throughout its development. This ―hole‖ in the seed
coat (Fig. 19) is at the suspensor end of the seed. It is
where the fungus usually enters and first comes into
contact with the embryo. The suspensor end of the
embryo differentiates to form what is called
mycotrophic tissue. The opposite end of the embryo,
called the chalazal end or chalazal pole, eventually
forms *meristematic tissue which will then develop
into the plant organs.
45
Once the fungus has established this *interface

with the embryo, germination begins. Technically
speaking, the fungus never is actually inside the
individual cells. To understand better this interface, a
plant cell will be simplified to the following: a square
water balloon inside a rigid wooden box with holes in
the side. The water balloon is the cell itself, the
balloon forming the cell membrane or plasmalemma,
maintaining the cell contents in one place. The
wooden box represents the cell wall, providing the cell
with structural integrity. Without their rigid cell
walls, plants would be *amorphous blobs, like people
without skeletons. The holes in the wooden box
represent what are called pit fields, or thin areas in the
cell wall. These permit intercellular communication,
as well as the conduction of numerous substances
from one cell to another. When the fungus interfaces
with the cell, it never actually pierces the water
balloon. Rather, it inserts the equivalent of its arms
and fingers into the holes in the side of the box. This
greatly alters the structure of the water balloon inside.
However, it never actually breaks it. The more ―arms‖
the fungus inserts into the cell, the greater the surface
area between the two, allowing for more rapid and
efficient exchange of materials.
As mycorrhizal fungus lacks a ―body‖ (i.e., no

central unit more developed than another), it is said to
consist of many individual strands called *hyphae. It
is these hyphae that branch off in all directions (from
46
no real centre) and act as ―arms‖ of the fungus and

grow through the pit fields to interface with the cell.
The extent to which hyphae are present within a cell
can vary. Sometimes structures called pelotons are
formed that consist of extensive hyphae forming a
complex coil within a single cell. This maximizes
shared surface area and therefore exchange.
As the embryo germinates and grows, structures

called rhizoids may be formed. They are simple
extensions of *epidermal cells and are formed on the
basal part of the *protocorm. Their function is to
facilitate mycorrhizal infection.
The germinated embryo will first grow into a

mass of relatively undifferentiated tissue called a
protocorm. Once of sufficient size, the meristematic
region of the protocorm will develop a shoot and
roots. The fungus remains in the mycotrophic tissue
throughout this entire development process, supplying
the growing plant with an array of required
substances.
The Exchanged Materials

Very little is understood with respect to exactly
what chemical exchanges occur between the fungus
and the orchid. Lipids in seed reserves can be
converted to starch and then glucose, providing the
necessary energy. It is suggested, however, that the
fungus facilitates mobilization of these reserves that
47
otherwise would not be possible. In effect, the fungus

is actually providing the embryo with access to its
own reserves.
The portion of the fungus within the orchid

tissue is only a small part of its whole. It remains
attached to its hyphal network within the soil. This
hyphal network functions as an extended root system
with a large surface area, affording the germinating
orchid with tremendously greater access to soil
nutrients than would otherwise be possible with its
own small surface area. Substances assimilated have
been shown to include water, nitrogen, phosphorous,
mineral salts, carbohydrates and other organics
(Rasmussen, 1995). In an elegant *in vitro
experiment, carbon and phosphorous *isotopes were
used to trace movement of these elements from the
surrounding medium into the fungus, where they were
transferred to the orchid (Smith, 1996; Alexander et
al., 1984).
Very little is known about what the fungus

receives from the orchid. In fact, it is suspected that it
might not receive anything. This would, surprisingly
enough, make the orchid a *parasite on the fungus.
Clarification of this issue will come with time as
experimental technology improves. Experiments, such
as that described, involving the tracing of materials
from the environment into the plant, are currently
much simpler to execute than those attempting to trace
48
material originating within the plant and its suspected

transfer to its inhabitant fungus.
Life Cycle of the Fungus

Once inside the embryo, it is likely that the
fungus will remain within the plant for a number of
years, perhaps in some cases for the entire life of the
plant. The mycorrhizal interface previously described,
however, is not a static relationship. Rather, it could
be described as a recurring cycle, starting with initial
infection of the embryo. This infection leads to the
formation of extensive hyphal connections, and
perhaps peloton formation. These connections are
subsequently the victims of digestion by the plant,
followed by reinfection of adjacent cells by the same
fungus to complete the cycle.
Peloton digestion is a process through which the

plant utilizes its own digestive *enzymes to kill and
assimilate the portion of the fungus within the cells.
Non-absorbed remains of the fungus are *vacuolated
or compartmentalized within the cell where they
remain. *Chitin is a substance the fungus
manufactures as one of its own cell wall components
(it also is the main component of insect
*exoskeletons) and it cannot be digested by plant
enzymes (nor by human enzymes). Consequently, it is
a primary constituent of the vacuolated fungal
remains. The digestion of the fungus is of particular
benefit to the orchid as it obtains a great number of
49
chemical components from within the fungus.

Unfortunately, the chemical makeup of assimilated
components is quite poorly understood. It is likely
that somewhere within this chemical soup lie key
components that are integral to the orchid‘s
dependence on the fungus. The purification and
analysis of such constituents would be a long, tedious,
but extremely worthwhile process that would surely
serve to further our understanding of the biochemical
factors affecting the interaction between fungus and
orchid.
Besides entering the orchid at the seed stage

through the suspensor end, mycorrhizal fungus can
also enter mature plants, presumably through root
hairs. Figure 20 is a cross sectional view of a cleared
root of Spiranthes odorata. The particular section of
root chosen shows no signs of infection, which would
be seen as black regions within the cells as a chitin-
specific stain was used. Figure 21 however, clearly
shows a region from the same root that has undergone
extensive colonization, as seen by the black bodies
within the cells. Very few of these black bodies
represent living pelotons; the majority of them have
been digested, nor were very many alive at the time of
preparation. The actual hyphal coil can be
distinguished in living cells, as well as its hyphal
connection to adjacent cells (Figure 22). It can be
seen that the only living coils are to be found at the
edge of the mycorrhizal colony. Figure 23 shows the
50
edge of the root with attached and cleared root hairs.

The black lines within the hairs are a fungal hypha,
presumably demonstrating their pathway into the
roots. The problem, however, with this type of
evidence is that there is no direction implied in the
photograph. The hyphae could be coming into the
root from the soil, or out of the root into the soil.
Experiments with live material have shown the former
to be more probable. Figure 25 shows a group of root
cortical cells pricked out9 and isolated, showing black,
vacuolated fungal remains. It should be noted that in
Figures 20-22, no colonization is observed within the
*stele (the central cylinder of conductive tissue). This
is an important aspect of the orchid/fungus
relationship. If the fungus were to venture into the
stele, the result would be clogging of the conductive
tissue (xylem and phloem). Consequently, the plant
would die in a fashion very similar to those plants
infected by Verticillium wilt, a fungal *pathogen that
infects plant roots and does indeed inhabit the stele,
thereby causing a slow death by dehydration.
Host-fungus Specificity
There has been a recent change in the school of
thought concerning the specificity of mycological
partners. It was once believed that only a single
9
Cortical cells are the "filler" storage cells located in the roots, between the
epidermis and vascular cylinder, or stele. Pricking them out is just their
removal during dissection with a fine needle, or something similar.
51
species of fungus was capable of forming this

*symbiosis (perhaps more appropriately parasitism)
with its orchid partner. It stood to reason, therefore,
that for every orchid species there was a single
compatible species of fungus. Extensive
experimentation has shown otherwise. Rather, a
single species of fungus can infect an array of orchid
hosts. What varies sometimes, but not always, is the
extent of its compatibility. Certain ―groups‖ of fungus
may form a positive relationship with certain ―groups‖
of orchids.
This type of research has encountered numerous

difficulties including the formation of pure fungal
cultures. Mycorrhizal fungus can be very fussy in
vitro, as it tends to resist culturing. The only source of
mycorrhizal fungus until now has been infected plants.
Successful isolations from the soil have not been
achieved. It is quite difficult to isolate a pure culture
from a living plant. Very often plants are infected
with other non-mycorrhizal entities, making
purification very difficult. Sterilization of root tissue
cannot be performed without killing the sought-after
fungus as well. As shown by previous photographs,
living pelotons can be notoriously rare in some species
(Curtis, 1939; Harvais, 1974; and Stoutamire, 1990),
Cypripedium spp. in particular, making isolation
nearly impossible. The problems encountered with
fungal cultures could be attributed to either or both of
two factors: (1) once within the plant, the fungus
52
undergoes irreversible alteration making its in vitro

growth extremely difficult, or (2) the fungus has
specialized nutritional requirements that remain to be
optimized.
To complicate matters further, it has been

suggested that the fungus that infects seedlings may,
in fact, be a completely different species from that
which infects mature plants (p. 241 in Rasmussen,
1995). This stands to reason as the mode of entry in
each situation is very different. As an
embryo/seedling, the orchid undergoes morphological
changes to facilitate infection. As a mature plant,
there are very few, if any, such structures. Entry is
through the root hairs, a specialized cell designed for
altogether different purposes (water absorption).
Colonization of mature roots could not be the result of
the fungus‘s originating from embryo infection and
gradually moving through the plant and down into the
root *cortex. First of all, colonies found within
mature roots are not continuous with older portions of
the rhizome, nor can the continuity of colonies be
traced through the root with digested pelotons. In
addition, roots produced from mature plants arise
*endogenously, from the *pericycle surrounding the
stele. Therefore, the fungus that may already exist
within the rhizome never has access to newly formed
roots. Infection of new roots, therefore, must be from
the soil, and through a much less ―welcoming‖
pathway.
53
This difference in mycological partners at

different stages of growth would easily explain why
initial experiments, testing symbiotic germination by
using isolates from mature plants, met with little
success. It is very clear that further research needs to
be done in this area.
One final point to be made is with respect to the

ability of the fungus to sustain itself in the soil without
any apparent orchid partners nearby. Orchid seed
could be germinated in the wild in a situation where
there are no single species of orchid within kilometres,
yet the seed will be colonized. This gives rise to many
questions: Where did the fungus come from? How
could it sustain itself in the soil? This group of fungi
can exist in the soil as *saprophytes; however, can it
rely exclusively on this method of survival? How do
orchids factor into this balance? Is there really a
mycorrhizal fungus per se, or is it some type of
ubiquitous species that is irreversibly altered once in
contact with the orchid? A thorough and clear
understanding of terrestrial orchid biology cannot be
attained until these issues are addressed.
Specifics Regarding Cypripedium

Different orchids vary in their dependency on
their mycological partners as mature plants (Sadovsky,
1968; Rasmussen, 1995, Chapter 4). Without a doubt
every orchid needs its mycorrhizal fungus to reach a
54
*photosynthetic stage where it can begin to provide

for itself; however, once this stage is achieved its
dependence can change greatly. Some species remain
highly dependent on their fungi, perhaps represented
in the extreme by the *achlorophyllous species, such
as the phantom orchid, Cephelanthra austiniae. The
other end of the extreme, in my mind, is represented
by the genus Cypripedium. As previously mentioned,
living pelotons, as well as digested pelotons, are
notoriously difficult to find in mature roots. For lack
of sufficient material with Cypripedium, the fragrant
ladies’-tresses, Spiranthes odorata had to be used for
Figures 20-23, 25. It is the following experiment,
which I conducted, that was found to be most
convincing for *autotrophism in adult Cypripedium
plants:
Numerous populations of Cypripedium parviflorum var.

pubescens were observed at various locations on the Bruce
Peninsula in Ontario. A few healthy plants were seen; however,
nearly all plants encountered (at least 1000 in small groups, or
as individuals over a large area) showed some degree of floral
*mutation or abortion, as well as leaf *necrosis. This affliction
was not limited to individuals. What had clearly been at one
time sizable clumps were also affected, showing mutated
flowers and stems greatly reduced to a few leaves. These
symptoms were indicative of some sort of root pathogen.
Sample plants were collected and examined by an
Agriculture Canada plant pathologist. Remaining samples were
placed under pot cultivation for experimentation. One pot was
treated with a systemic fungicide (until this point, a cultural
55
taboo). The fungicide is toxic to mycorrhizal fungus. The other

pot was not treated.
The plant in the untreated pot died by the end of the
season, failed to make new growth for the following season and
was discarded. The plants in the treated pot came up the next
season with as many growths as the previous year, and produced
two flowers on two of three stems. The analysis by the
pathologist revealed that Rhizoctonia solani was present, as well
as something he could not identify. Rhizoctonia solani is a
common root pathogen and likely the cause of the symptoms.
The unknown fungus was possibly mycorrhizal.
These results were quite surprising. Depending on the
fungicide used, one would expect the treated individual to
immediately lose its mycorrhizal partner and therefore perish.
Such was clearly not the case, putting into question the value of
the fungus to a mature plant. Admittedly, mammalian factors (a
squirrel) affected long term study of the treated plants. It was
impossible to assess adequately things such as recolonization, or
the possibility that such a dramatic recouperation was in the
short term only. It would have been ideal to observe the plant in
the long term to see if its vigour was maintained.
The genus Cypripedium is not any different

from other genera during young stages with respect to
its fungal dependency, as the first root (of exogenous
origin, i.e. from the protocorm cortex) is mycotrophic
from the beginning (p. 239 in Rasmussen, 1995).
However, the following roots become less specialized
for mycotrophy as there are fewer root hairs formed
(Rasmussen, 1995). ―The extent of mycorrhiza
declines from year to year, while the above ground
organs increase in volume.‖ (Fuchs and Ziegenspeck,
1925 in Rasmussen, 1995).
56
Two-year old roots show patches of infection

that are mainly dead pelotons. These patches could be
the remains of temporary mycorrhizal infection during
the underground season; however, they were
interpreted to be fortuitous and possibly pathogenic
(Harvais, 1974).
Despite these findings, mature Cypripedium

plants have been known to remain dormant
underground for one to five years (Gill in Rasmussen,
1995). It is suggested that this might be impossible
without the aid of the fungus. It is possible that under
such conditions, the plant may drastically increase its
dependence on the fungus. It is equally possible,
however, that the plant is relying on the significant
stored energy potential within the rhizome and roots
during this time, eliminating the need for the fungus
during its dormant period.
The fact remains that the importance of

mycorrhizal fungus to mature plants is still
incompletely understood. Further experimentation
needs to be done on a larger and long term scale with
mature plants.
References
Alexander, C., I.J. Alexander, and G. Hadley. 1984. Phosphate

uptake in relation to mycorrhizal infection. New
Phytologist 97: 401-11.
57
Curtis, J.T. 1939. The relation of specificity of orchid

mycorrhizal fungi to the problem of symbiosis. Am. J.
Bot. 26:390-9.
Harvais, G. 1974. Notes on the biology of some native orchids

of Thunder Bay, their endophytes and symbionts.
Canadian Journal of Botany. 52:451-60
Rasmussen, H.N. 1995. Terrestrial orchids: from seed to

mycotrophic plant. Cambridge University Press.
Sadovsky, O. 1968. Orchideen im eigenen garten. BLV

Munchen Basel Wien.
Smith, S.E. 1966. Physiology and ecology of orchid mycorrhizal

fungi with reference to seedling nutrition. New
Phytologist 65:488-99.
Stoutamire, W.P. 1990. Eastern American Cypripedium species

and the biology of Cypripedium candidum. Pp. 40-48 in
North American Native Terrestrial Orchid Propagation
and Production Conference Proceedings, March 1989.
Brandywine Conservancy, Chadds Ford, Pennsylvania.
58
Part III
Propagation and Cultivation
Mature plants are readily propagated by

division, ideally in the fall or spring while growth is at
a minimum; however, this can be accomplished quite
successfully at any time of the season assuming proper
care is taken to minimize root damage. Roots as old
as three or four years remain active so should not be
sacrificed if possible. New roots are formed shortly
after flowering at the base of the new growth. Single
bud or growth divisions should be avoided, but are
sometimes unavoidable with the *stoloniferous
species such as Cypripedium formosanum or C.
guttatum. Division is practical and effective for small
scale propagation, but is limiting because of the few
*propagules produced over a relatively long period, as
well as their genetic uniformity.
Without a doubt, seed propagation is the most

desirable method of propagation. Large numbers of
genetically variable seedlings can be produced in a
short period of time. Unfortunately, this line of
research has historically been met with repeated
failure or astronomically mediocre results. It is only
within the last decade that significant refinement of
techniques has been achieved so as to overcome
previously insurmountable hurdles. We are not, as
many predicted years ago, on the verge of large scale
commercialization of this genus. Propagative
Doherty: The Genus Cypripedium: Part III
mysteries remain, and further refinement of techniques

is necessary. However, the long and slow path has
finally yielded some encouraging and viable results.
Numerous growers around the world have

unlocked the mysteries of propagating this genus from
seed and are consequently offering a vast array of
species and hybrid seedlings for sale (see Appendix I
for a listing). Others have outlined their highly
successful procedures in excruciating detail for public
consumption, specifically Bill Steele of Spangle Creek
Labs (Steele, 1995, 1996) and Svante Malmgren, a
surgeon in Sweden (Malmgren, 1993, 1996), as well
as Chu and Mudge (1996). Their work has provided
the basis for the following summary.
Considerations to be made when propagating

from seed can be loosely divided into seven different
categories, each to be considered individually:
(1) collection of seed,
(2) sowing and pretreatment of mature seed,
(3) cold stratification/dormancy requirements,
(4) light conditions,
(5) substrate composition,
(6) replating10
(7) *deflasking and planting out.
10
Replating is the procedure of transferring in vitro growing plantlets to a
different, sterile container. This is usually done to space out the seedlings;
however, it can also be performed to change the nutrient medium or remove
seedlings from an infected culture.
60
(1) Collection of Seed

As with tropical orchids, seed of Cypripedium
can be sown as dry, mature seed, or as unripe,
immature seed (―greenpodding‖). Each method has its
own advantages. ―Greenpodding‖ requires unripe
seed pods to be harvested prematurely. Ideal harvest
date depends on a number of factors, including species
(which will likely exhibit clonal differences), as well
as environmental conditions experienced during
ripening. Harvest dates cited in the literature vary
from four weeks after pollination to as many as eight
or nine; five or six may be a good starting point.
Unfortunately, optimal harvest dates must currently be
*empirically determined as there is no known way to
consistently predict when the seed is ideal for
germination. Efforts have been made to correlate
ovary size and swelling in Cypripedium parviflorum
to suitability for harvest (Light, 1989); however
generalizing to other species would not be appropriate.
The major advantage to ―greenpodding‖ is the
circumvention of seed dormancy. Dormancy factors
that are incorporated into mature seed are not yet
developed in green seed. The ideal pod is one which
has been allowed to develop for as long as possible
without the onset of dormancy, which will greatly
reduce subsequent germination. The two major
disadvantages to ―greenpodding‖ are the uncertain
timing of fruit collection and the need to flask it very
shortly thereafter. Green pods will not keep, making
their storage impossible and mailing to other parties
61
Cypripedium parviflorum var. pubescens

large yellow lady’s-slipper
62
very difficult. Given that dormancy factors in mature

seed can be overcome through various techniques,
sowing and storing mature seed is usually the method
of choice.
Mature seed is harvested in the fall, ideally just

before the pod cracks, thereby minimizing
contamination from exterior sources as well as seed
loss to the wind. Sometimes this later harvest cannot
be avoided, and it is usually of no great consequence.
(2) Sowing and Pretreatment of Mature Seed

Whole green pods are surface-sterilized shortly
after harvesting. Under *aseptic conditions, the pod is
then cut open and the immature seed scraped out and
sown directly into culture vessels. Immature seed
usually is still attached to the ovary wall, and is white
to yellow in colour.
Mature seed differs significantly in its

colouring. It usually turns to some shade of brown;
however, in some species its colour transformation is
not so pronounced and it appears very similar to the
juvenile yellow-white seed.
As previously mentioned, mature seed contains
dormancy factors not found in immature seed.
Unattended, these factors can severely inhibit
germination processes, sometimes completely
arresting them. This natural biochemical mechanism
surely acts as an environmental sensor to cue the plant
63
to germinate at the most appropriate time; however,

under artificial propagation regimes, control of these
processes is necessary. It has been found that
extended exposure to bleach before sowing greatly
reduces the action of these dormancy factors.
Sterilization times for orchid seed have conventionally
been in the 10-15 minute range for a 10% solution of
commercial bleach, (i.e., 0.5% NaOCl); however,
Anderson (1990) found by accident that leaving seed
in the solution for two hours significantly enhanced
germination rates. Subsequent analysis showed some
species to prefer soaking times of up to six hours!
Malmgren (1993) has discovered this same principle
and has found that treating the seed with acid before
bleach achieves the same result without the lengthy
soaking time. It is likely that the brief acid treatment
weakens the seed coat, facilitating more effective
action of the bleach. Regardless of the technique, the
net effect is the oxidation of the dormancy factors by
the bleach and thus their relative inactivation. In
addition to extensive bleaching of mature seed,
germination can be expedited further in some species
by adding a *cytokinin to the germination medium.
(3) Cold Stratification/Dormancy Requirements

Both mature plants and seedlings have
dormancy requirements that have to be met. Ideally,
mature plants should be given sub-zero temperatures
for at least three months, if not longer. Colder-climate
species such as Cypripedium guttatum and C.
64
passerinum usually prefer longer and deeper cold.

Seed of some species survives freezing, though
seedlings, unlike mature plants, will die if frozen and
therefore must be kept between 0 and 5 C during
winter months. This is achieved in a cold frame,
refrigerator, or similar setup.11
A great concern when vernalizing seedlings is

timing. Seedlings will not leaf out until given an
appropriate cold period. Until that point, the only
shoot structure that forms is the small overwintering
bud, easily seen on mature plants in the fall. Ideally,
plantlets with fully developed buds should be
vernalized in the fall, concurrent with their natural
development cycle. This means that seed must be
sown at an appropriate time to allow for full
development of the seedlings prior to *vernalization,
however, not too soon so as to force the plantlet to
wait too long before its required cooler temperatures.
Seedlings are usually deflasked and sealed in airtight
dishes or plastic freezer bags with a little extra
moisture to prevent desiccation during vernalization.
Again, determination of these procedures remains
empirical as considerable variations exist from species
to species.
11
Seed of at least some species survives freezing quite well. I‘ve had
enhanced germination of C. parviflorum seed from capsules left on the plant
all winter and subjected to temperatures in the range of -15 C, as compared
with refrigerator-stored seed maintained at 2–3 C. WS
65
(4) Light Conditions

Seed must be placed in darkness immediately
after sowing. Light is not absolutely necessary until
plantlets complete with roots and bud have been
deflasked and vernalized; however, it can be given as
soon as the first initial shoots are evident.
(5) Substrate Composition

Harvais (1982) was the first to extensively
investigate the precise chemical requirements for seed
germination of Cypripedium reginae. His medium,
therefore, was optimized for that particular species.
Bill Steele (1995, 1996) has quite successfully
duplicated and even improved Harvais‘ results, and
subsequently modified his recipe to accommodate
other seemingly pickier species. Apparently the
element of interest is nitrogen. If provided in
*inorganic form, notably ammonium nitrate, it can be
detrimental to many species. Steele has reduced its
concentration in Harvais‘ medium from 1400 mg/l to
500 mg/l with greatly improved results for a number
of species.
Svante Malmgren (1993, 1996) has also

achieved success by supplying nitrogen in the form of
*amino acids, a completely organic form. This could
be correlated to a natural situation where the plant
derives nutrients from digested fungal hyphae,
containing *metabolized nitrogen in the form of amino
acids. The glaring absence of hyphae in roots of
66
mature plants suggests that if the plant were to depend

solely on fungus as its nitrogen source, it would surely
starve. The reasons for improved germination and
growth with reduced ammonium nitrate or complete
substitution are not completely understood; however,
answers could lie in potential sensitivity to ammonium
nitrogen, or an optimal ratio of nitrate nitrogen to
ammonium nitrogen.
Additional components of Malmgren‘s medium

are *kinetin, pineapple juice (likely containing a
substance with plant growth regulator properties,
probably analogous to the potato cubes used by
Steele) and a basic vitamin solution. Readers are
directed to the references for specific formulations.
(6) Replating
Cytokinin in the germination medium will cause
abnormal growth as the protocorms begin to
differentiate. Consequently, as soon as the first root
initials are evident, developing protocorms must be
transferred to fresh medium. It should be identical in
composition to the germination medium, with the
exception of kinetin, which must be excluded.
(7) Deflasking and Planting Out

The ideal time for deflasking is in the fall or
winter in preparation for vernalization. Attempting
vernalization on the flasking medium causes high
mortality in some species, e.g., C. reginae.
67
Choice of substrate for outplanting varies.

Malmgren uses a combination of three parts woods
soil, one part peat and one part lime soil. Other
growers use inorganic particulate media that are sandy
or gravelly, providing excellent drainage. Seedlings
must never be allowed to dry out and should be
watered frequently. Humidity is equally important as
it directly affects drying out. Malmgren uses clear
plastic covers over peat-filled trays into which clay
pots are sunk. This keeps humidity at a good level,
and also keeps the pots cool. Clay pots likely provide
some gas exchange near the root zone as well as
additional cooling. Direct sun can often lead to
dehydration problems, as can excessive wind. A
sheltered, shady location is therefore recommended.
Tissue Culture
There currently is no published literature
concerning the tissue culture propagation of
Cypripedium. Chin Chu (1996), a researcher at
Cornell University, has experimented with this
technique and achieved encouraging results. I have
conducted research in this area as well, and feel there
to be significant propagative potential.
In my work mature plants of Cypripedium

parviflorum var. pubescens were cultivated in pots and
kept in a cooler over winter. Plants were removed
from their containers and washed of all soil in early
68
spring, just as the overwintering bud began to break

(Fig. 24). This overwintering bud (to be referred to as
the primary bud) was removed and dissected in an
effort to locate a shoot meristem for tissue culturing.
Dissection was performed at this time of year in an
effort to induce proliferation of putative meristems
during a period of active growth, coinciding with their
natural cycle. Well developed *flower primordia were
found inside the primary bud. At the base of the
primary bud were found one or two smaller, secondary
buds (Fig. 26), the *vegetative primordia for the
season one year away. These secondary buds were
removed, fixed, embedded in *LR White resin and
sectioned with a microtome12. Sections revealed
differentiated regions within the buds, corresponding
to flower primordia. Tissue, therefore, was
inappropriate for tissue culture as differentiation had
already proceeded too far. At the base of the
secondary buds were found additional,
undifferentiated dome shaped tissues (Fig. 27, Fig. 28)
which likely were tertiary buds, or the secondary bud
initials for the following season. These tissues proved
to be the least differentiated and were selected for
tissue culture.
12
The process whereby very thin slices (micrometres thick) are sliced off of
a prepared block of material that has been hardened with a compound such as
LR White resin. Knives used to slice material are made of fractured glass, the
edge of the fracture acting as the very fine blade.
69
Excised meristematic regions initially showed

little response to various hormonal treatments in liquid
culture. Subsequent formulations, however, did
indeed induce proliferation; however, *explants did
not survive long enough for sustained proliferation.
The results of this experiment were promising;

potential sources for meristematic tissue were located
within a mature plant. It was also shown that
proliferation can be induced. Unfortunately it is likely
that the next course of action has to be a highly
empirical approach, to formulate the optimal medium
for proliferation and sustained growth. This will
likely result in the sacrifice of a large number of
mature plants, as they cannot survive the *excision of
the meristematic region.
Ethical Considerations
Before any discussion regarding cultivation of
these plants can begin, the crucial topic of ethical
issues must be addressed. Ultimately, all cultivated
plants come from the wild; consequently, the means
through which we obtain plants for cultivation has a
direct impact on wild populations. Unfortunately,
these wild populations are sensitive to the
encroachment of man into their homes, making some
(but not all) of them increasingly rare. A degree of
caution must be exercised when engaging in any
activity that affects these delicate populations.
70
Whenever possible, artificially propagated

plants should be purchased from reputable dealers.
There are a multitude of advantages to this approach:
most obviously, plants are not removed from the wild,
and thus can remain there, happy (for the most part)
and reproductive. Financial support of artificial
propagators provides the necessary funds for
additional optimization of propagative techniques.
Artificially propagated plants are better under
cultivation as they have been through a sort of
selection process. They are available to you because
they represent the most genetically vigorous
specimens, surviving the rigours of mass production.
In addition, they likely have been selected for floral
characteristics that are most appealing to us. These
principles can be demonstrated by Paphiopedilum
rothschildianum. Currently, after generations of
breeding and selection, artificially propagated plants
(really, all that is available) are more vigorous than
wild collected plants as the former have been selected
for vigour under greenhouse conditions. In addition,
flower quality is drastically improved over wild
counterparts. The increased appeal of artificially
propagated plants virtually eliminates collection
pressure on wild populations. Unfortunately, there is
a sector of orchid hobbyists that, despite the
aforementioned points, feels jungle plants to be
superior. They claim jungle plants provide ―fresh‖
genes to a stagnating gene pool resulting from
generations of inbreeding. This valid argument loses
71
merit when weighed against the benefits seen from

such an intensive breeding effort, as well as the
inevitable detriments from further depleting this
already highly threatened species in the wild. For
these reasons, there should be no motivation to go to
the wild for plant material. There are exceptions,
however.
In two circumstances collecting from the wild is

completely legitimate and conservationally sound.
The first instance is obvious. If a plant or group of
plants is in imminent danger of destruction, due to
construction or other artificial activities of man, then
collection is beneficial. Save the plant and try to grow
it or pass it on to someone else who will.
The second instance is much more gray on the

surface, yet after careful examination is just as black
and white as the first circumstance. Many
Cypripedium species are successional plants.
Succession is the gradual transformation of one initial
habitat into another different habitat, due to the
modifying effect that growing vegetation has on its
microclimate. These changes have a pronounced
effect on the species composition of the habitat. A
hypothetical example that realistically illustrates the
point would be a newly formed island in an ocean.
The island is initially composed of sand. A seed may
arrive and germinate, leading to formation of a grassy
area. Gradually, as this grass grows and accumulates
72
its own dead organic matter over time, new species

that couldn‘t previously survive do. There they
germinate and grow, creating a little bit more shade,
and a little bit more *humus. This makes way for a
different species that needs a bit more shade, and a bit
more humus. Unfortunately for the grass, the new
shade created by this new species starves it of sun and
the grass eventually perishes. However, a different
species finds these conditions ideal and colonizes.
This process continues until the climax stage is
reached. At this stage, the overstory species creating
all of the shade (Quercus spp., for example) are able to
reproduce and sustain themselves. Their seedlings can
tolerate the shade of their parents and grow and
replace them. Along the way to this final climax
stage, an array of plants have come and gone. It is
only the species that can survive in this shaded, climax
stage that remain.
Many Cypripedium species are not climax stage

plants. They are a *transient part of their landscape
with bell shaped population curves over time. It is
important to note, however, that some are climax
species. They are perfectly happy in shaded
woodlands and reproduce there effectively. It is the
plants occupying diminishingly suitable habitats that
can be removed without detriment to the reproductive
population. They will subsequently reward their
rescuer with dramatically increased growth the
73
following season, if provided with higher light levels

under cultivation (see Initial Successes).
This kind of ecological understanding requires

extensive field experience, enabling one to read the
landscape and determine if indeed a habitat is
changing to the detriment of the orchid, and not to
misinterpret natural fluctuations. Such a skill can be
learned, and should not remain within the domain of
the advanced botanist. Rather, such understanding
should be spread and shared, most importantly with
the horticulturist to encourage ecologically sound
practices, before stupid and selfish decisions are made
at a cost to other naturalists and, more importantly, to
the orchids themselves.
Cultivation
The most valuable cultural advice comes from

those growing a particular species outside of its
natural range. This is because the grower is required
to deal with every variable affecting the plant‘s
growth. Seasonal fluctuations of things such as *solar
radiation, precipitation patterns, and temperatures are
all foreign to the plant being grown and must be paid
special attention by the grower, over and above
considerations being made if the species is grown
within its own range. It is much easier to go wrong
outside of the range, leaving more room to learn.
Having said this, I shall provide cultural
74
recommendations for at least five different species,

living very close to the hearts of their ranges. Despite
having much in my favour growing within the natural
range of many species, a great deal has been learned. I
have received and used valuable information from
other growers in similar situations, so I shall not
hesitate to provide it.
One hundred percent of my plants are grown in

pots, and have been that way from the beginning. This
situation initially arose out of necessity. It was
impractical (and not allowed by my father) to convert
the flower beds at our city home to the necessary, and
highly variable, specifications demanded by different
species. Pots provided flexibility in growing medium,
as well as portability. It was not until a number of
years later that it became apparent that pots, in many
ways, are superior to in-ground beds. Growing in pots
allows for superior cultural flexibility. Growth
medium can be changed quickly and easily if found to
be inappropriate. Moisture levels are easily
controlled, and not affected by invading tree roots.
*Herbivory is somewhat reduced, depending on where
the pot garden is kept. Root and bud growth is easily
monitored. Most importantly, plants can be well
admired while standing, and even brought indoors to a
prominent location in the house while flowering.
Disadvantages of *pot culture are encountered

when growing wandering species such as
75
Cypripedium formosanum or C. guttatum. They can

find their pots restrictive, necessitating annual
repotting. Repotting is not normally a detriment for
other species, as it is only required every few years
provided all other factors are optimal. The biggest
disadvantage of pot culture is the warming of the root
zone. It is of considerable importance to maintain a
cool root zone. This is achieved in a number of ways,
to be discussed shortly.
All things considered, pot culture appears an

effective approach to cultivation. All
recommendations here are the result of personal
experience with this technique; however, the majority
of principles are general and can be applied with equal
effectiveness to bed-cultivated plants.
Choice of Container
The underground structure of Cypripedium
species is usually very horizontal. Roots tend to
spread outwards from the rhizome and do not go as
deep as they do long. As a result, initial attempts to
grow plants in conventionally dimensioned pots were
met with some difficulty as a good portion of the
medium was not being occupied by the roots. This is
detrimental for a number of reasons. Excessive
moisture can collect and lead to rot problems. In
addition, medium outside of the *rhizosphere is
beyond the effects of root exudates, potentially
leading to the eventual buildup of pathogens.
76
The solution was to use *bulb pans, or similarly

shaped pots that are broader than they are deep. Until
now plastic pots have been used; however,
Malmgren‘s success with seedlings in clay pots
suggests clay pans to be comparable to plastic, if not
better. Porous clay would facilitate root oxygenation,
as well as (and most importantly) decreasing root zone
temperature through the continual evaporation of
water through the clay. This would mean more
frequent watering and the associated complications;
however, it is likely to be worth the trouble.
Not all species require bulb pans. Vigorous and

fast growing species such as Cypripedium parviflorum
grown in normal pots will send roots down and
throughout the entire medium. Because it is so
massive, C. reginae will similarly occupy huge barrel-
like pots, but will do equally well in dishpans or kitty
litter tubs.
Substrate
Substrate is likely the one cultural consideration
paid the most attention by growers, perhaps
excessively so. Although this genus is notorious for
its species-specific habitats and corresponding media
requirements, there are a number of properties that are
common to most species. Medium pH is a critical
factor to growth as it directly affects nutrient uptake.
For this genus it can be generalized that pH is optimal
77
from 6.0–7.0. The most notable exception is

Cypripedium acaule, which prefers a distinctly acid
medium. The pH of the growing medium can be easily
increased with a light sprinkling of lime at the
beginning of each season, or decreased if necessary
through the addition of phosphoric or nitric acid to
irrigation water. If appropriate, acid humus such as
oak leaves or rotting pine needles or wood can be
included in the substrate to achieve a similar effect.
*Aeration of the medium is without a doubt the

most important factor, with medium pH close behind.
Roots demand a highly aerated, well drained mix.
This is achieved with a number of widely available
media components such as perlite, *turface,
vermiculite, coarse grades of sand and gravel, and
even *rockwool (some growers use pure rockwool).
Drainage is enhanced by putting a layer of stones or
something comparable in the bottom of the pot.
Pot ―partners‖ is an important concept that

should be a part of any pot growing regime. In the
wild, there is no species that exclusively occupies the
soil surrounding the plant. There is a very active root
zone, composed of many different plant species with
their roots closely interacting with one another, as
well as the native *microbial and fungal populations.
Rearing the plant in an environment devoid of this
type of interaction can be done, successfully so;
however, the incorporation of wild neighbouring
78
grasses into the growing medium significantly

increases the chances of success and acts as an
effective buffer to varying environmental conditions.
Maintaining grasses and other neighbouring plants in
the same pot necessitates additional watering due to
the increased transpirational load on the medium, yet
makes sense for a number of reasons. It is more
difficult to overwater pots with companion plants,
thereby reducing problems with rot. The roots of
companion plants aid in the modification of the
rhizosphere by producing additional and appreciated
root exudates, thereby reducing potential pathogen
proliferation. Most importantly, leaves of companion
plants keep sun off of the soil surface, keeping it
cooler. Cool roots equal happy plants. If plants are
not collected in the wild and companion species are
unavailable, any type of lawn grass seed sown into the
top of the pot will effectively provide the necessary
company in the rhizosphere.
Zonation of the root atmosphere is an important

consideration particularly relevant to the cultivation of
Asian species. The underground bud and subsequent
underground stem section of Asian species appear to
be extremely sensitive to moisture throughout the
year. I have sent an uncomfortable number of Asian
plants to the big pot garden in the sky due to bud or
stem rot which did not seem to be any sort of a
problem in similarly grown North American species.
Apparently many Asian species encounter very little
79
winter moisture and, therefore, have little tolerance for

it. The solution under cultivation is to divide the
underground region into two distinct zones, their
junction running horizontally through the middle of
the rhizome. The bottom zone is occupied by the
roots and should be a well aerated mix with good
drainage, as well as with varying humus components.
The top layer, covering the top half of the rhizome and
the entire underground dormant bud, should be
inorganic and extremely well draining. The purpose
of this is to allow zero moisture to collect around the
bud (or stem in the growing season) thereby avoiding
rot. North American species tend to be much less
sensitive to this cause of rot, tolerating significant
winter moisture caused by the repeated thawing and
freezing with little or no snow blanket that is
encountered in much of our climate.
Temperature
Many species are tolerant of a wide range of
temperatures; however, there are definite preferences,
under which optimal growth will occur. The
temperature under which a species is kept is closely
tied to the light levels it encounters. Cypripedium
guttatum grows in the wide open in Alaska. It would
last three days if it were grown in the wide open in
Windsor (the warmest spot in all of Canada).
Apparently in Alaska there are few days with full sun;
most are overcast, with a maximum temperature in the
low seventies (Fahrenheit). Therefore, in Windsor,
80
where there can be more than twenty days a summer

over one hundred degrees, it is given significantly less
sun than it would receive under a cooler temperature.
Ideally, one would maximize solar radiation under the
coolest of temperatures, resulting in increased
photosynthetic capacity; however, such cannot be
achieved in outdoor conditions. Since we have little
control of temperature, the best approach is to modify
light levels, giving less light to the cooler-growing
plants. The specifics of achieving this are dependent
on the species and where it is being grown. Coolness-
loving plants under heat stress caused by sun will
initially show slight necrosis at the leaf tips. If
temperature or light levels are not reduced, the leaf
will quickly blacken and die for the season. Despite
reaching its heat threshold in August under full shade
in Windsor and completely blackening off, C.
guttatum returns year after year. My oldest plant is
now seven years old.
Root zone temperature is a concern that requires

careful consideration if plants are to thrive. Many
horticulturally popular plants experience optimal
growth under elevated root zone temperatures. In fact,
artificially elevated root zone temperatures can result
in greater nightly cold tolerance, resulting in
decreased heating costs for greenhouse growers. Such
is not the case with Cypripedium species. The lower
the root temperature, the better. Cool roots provide
greater heat tolerance during warm summer months.
81
In the spring, warmed roots will lead to faster and,

therefore, softer shoot growth. Stems become weak
and wobbly, requiring staking. Cool spring roots
make shoot growth slower and stronger.
Because plants are above the ground with pot

culture, they are exposed to higher root zone
temperatures from the beginning. There are a number
of measures, some previously mentioned, that can be
taken to keep root zone temperature at a minimum.
Clay pots can be used which remain cooler because of
the evaporative action of medium moisture through
the walls of the pot. Companion grasses can be kept
in the pots to block sun from hitting the medium
surface. If companion grasses are not used, thick,
insulative mulches can be provided. Finally, plants
can be grouped in such a way to shade each other,
with the only pots receiving direct sun on their sides
being the ones along the edge of the grouping.
Light
Light is a fairly simple requirement to meet,
especially with pot culture. Under unsuitable light
conditions, the pot is easily moved to a more
appropriate location. If growing on a centralized
bench, one should select an area receiving direct sun
for approximately half of the day. From that point,
shade cloth can be added to different sections
depending on the species housed. Not even
82
Cypripedium candidum will take full sun for an entire

day.
Too much sun is easily seen on the plant as

yellowing leaves, or slight browning of the leaf tips.
This signals a hasty move to a shadier location.
Inadequate light is seen as dark green leaves, or long,
stretched out growths. There is a bit of a dilemma
when selecting light levels. The higher the light level,
the more growths will be produced the following
season. Unfortunately, growths tend to be short with
pale (sometimes not) flowers. Conversely, the taller,
showier, and darker flowered plants resulting from
reduced light levels tend to make fewer growths for
the following season.
As a general rule, most species will appreciate

deciduous shade. Very high light levels at the
beginning of the season (when lower temperatures and
smaller leaf surface area make this possible) gradually
diminish to high shade. Prairie shade is another term
mostly applicable to Cypripedium candidum. I had
previously observed this species in the wild a number
of times during its flowering period. It was seen in
very open, exposed locations. The light levels seen at
these locations were duplicated under cultivation,
leaving me at a loss when plants would prematurely
blacken near the end of the summer. The problem was
solved when plants were visited in the wild much later
in the season, revealing neighbouring grasses over one
83
Cypripedium candidum Muhl. ex Willd.

Small white lady’s-slipper
84
metre tall! Subsequently plants were given increased

shade as the season progressed, and they appeared
much happier.
Water
The highest quality water should always be
used, rainwater being at the top of the scale and city
water or tap water at the bottom. Media should be
kept continually moist and never be allowed to dry
out, although there are a few species that do indeed
prefer this. Despite being found in seemingly sopping
wet conditions, species such as Cypripedium reginae
need only be kept moist, and not soaked.
Traditionally, fertilizers have not been a part of

watering regimes for terrestrial orchids; however,
these plants do not differ greatly from any others with
respect to their nutritional requirements. Perhaps
initial attempts at fertilization resulted in damaged
plants, discouraging further efforts. Currently many
successful growers are using fertilizers, resulting in
increased vigour. Formulations used vary, as do brand
names. An all purpose 20-20-20 formulation is a good
starting point, never to be used at full strength.
Applications should be at ¼ to ½ strength at regular
intervals throughout the vegetative season, depending
upon the concentration used. Cypripedium guttatum
and C. henryi are two species that have proved to be
extremely sensitive to fertilizer, and should not be
given any, or else severe root burning and subsequent
85
leaf blackening will occur. It should be noted that

fertilization is especially important if using an entirely
(or even partially) inorganic substrate. Without this
external source of nutrition, plants will deplete limited
root reserves and gradually diminish.
*Fungicides have been recommended by a few

growers as well. These are of particular importance
during continually wet periods where plants are most
susceptible, as well as the deflasking stage where
sterile plants can be quickly overtaken by undesirable
pathogens. Label application instructions should be
followed, applying when necessary. Due to the low
dependence on mycorrhizae of this genus, detrimental
effects of fungicides on mycorrhizal populations are of
little consequence in such artificial cultural conditions
where complete nutrition can be supplied by the
grower.
As previously discussed, Asian species tend to

be very sensitive to winter moisture. In climates
receiving a thick and continuous snow blanket
throughout the winter months, this is not a great
concern. In warmer climates where repeated winter
thaws result in constant exposure to moisture, bud rot
is a serious threat. I have lost a number of Asian
species to excessive winter moisture. The solution is
very simple, yet perhaps somewhat difficult to initially
accept. Sensitive species are grouped together and
covered with a layer of opaque plastic as soon as they
86
are sunk into the ground. The plastic should be right

on top of the pots and anchored to the ground.
Tomato stakes stuck into the comers of the plastic
sheet work well, as they serve an equally important
function of signaling the location of the pots to any
potential human traffic through the area. This
protective layer should remain in place until the spring
when its removal will reveal many happy, dry and pale
green sprouts ready to grow. Apparently any water
requirements the plants might have over the winter are
met by existing media moisture, as well as by that
which is stored.
Dormancy Requirements
Nearly all species require a period of winter
dormancy where air temperatures drop below freezing
for at least three months. The ideal duration of
dormancy is dependent on species; northerly species
such as Cypripedium guttatum and C. passerinum
prefer a longer period. Degree of coldness is not fully
understood, as scientific comparisons have not been
made between the responses to, for example, -5 C and
-15 C. For general purposes, dormancy requirements
appear to be met at any temperature below zero.
An important point to be made regarding winter

temperatures is that air temperature is by no means a
direct reflection of soil temperature. The ground is
almost always warmer than the air. This effect is
enhanced if there is an insulative blanket of snow
87
Cypripedium passerinum
Sparrow’s-egg lady’s-slipper
88
covering the ground. Because of this, a plant in

Windsor, where there are repeated winter thaws and
fairly insignificant snow cover, may spend most of its
winter colder than a plant in the Yukon, that receives a
very thick layer of snow that remains for the entire
season.
If pots were left on their growing benches over

winter, they would freeze solid and more than likely
kill their contents. Because of this, pots must be
sunken into the ground in the fall to ensure proper
chilling. This can be done as late as shortly after the
first frosts; however, the task is most comfortable in
warmer weather. They can then be removed from the
ground in the spring, hosed off to remove soil clinging
to the sides of the pots, and returned to the growing
bench. There are two problems with this: (1) it is very
labour intensive, and (2) it requires a permanently
empty location in the yard where nothing is grown,
with the exception of annuals that can be removed in
the fall. Without a doubt these difficulties are the
greatest disadvantages of pot culture.
Initial Successes
Removing plants from the wild may result in
tremendous growth response in following seasons,
especially if plants are treated to significantly
increased light levels. This rapid response exhibited
by the plant is a trait that has been built-in through
89
years of evolution in response to what is termed a

disturbance. A disturbance is the result of an external
factor acting on an ecosystem, causing some sort of
environmental change. Such disturbances can be
caused by a number of factors, either natural or
imposed by man. Lightning may eliminate a large
branch from the forest *canopy. A prairie fire may
occur, eliminating much of the covering brush.
Forests may be selectively logged, drastically
changing incoming light levels. Strips of land
adjacent to railroads may be mowed to the ground in
an effort to control invading brush. All of these things
significantly increase light available to various
*microclimates within the habitat. The rapid growth
response of the orchid to this new and valuable
resource is in an effort to maximize its use and thus
reproductive potential before the resource is utilized
by competing species. Orchids are well known for
this ability and are highly competitive in such a
disturbance situation. They can quickly colonize and
even take over such an area.
Because of this effect, recent transplants may

exhibit exceptional growth in the first year or two
under culture, even if cultural conditions are not
optimal. A true estimate of cultural success cannot be
made until several years after transplanting.
Greenhouse Culture
90
A number of factors exacerbated by pot culture

point towards greenhouse or cold frame culture.
Without a doubt, greenhouse culture would maximize
environmental control and therefore cultural success.
Temperature control is one obvious advantage.
Evaporative cooling pads in the summer would give
plants greater tolerance to light by decreasing air
temperature (and therefore root zone temperature),
increasing photosynthetic efficiency. The temperature
gradient through the greenhouse created by the
cooling system would allow for selective placement of
plants at suitable locations within the greenhouse.
Increased humidity as a result of this type of cooling is
an additional benefit. Minimal cost heating in the
winter could maintain temperatures close to zero,
satisfying dormancy requirements yet eliminating the
need to bury pots. The roof of the greenhouse could
act as a collecting agent for rainwater. The potential
for cultural optimization is great; it is clear that
additional exploration of this possibility is necessary.
Specific Recommendations
Cypripedium arietinum (Fig. 12).

I have had the most success with transplants
originating from cool shoreline limestone barrens, as
opposed to the sand dune or cool swamp type habitats.
In the barrens, the plants grow singly, with roots in a
thin layer of black, organic, sandy soil that is
somewhat moist, on top of solid limestone. The stems
91
grow through a very thick layer of very dry mosses,

likely absorbing much overhead moisture as well as
insulating the root zone from warming sun. Plants are
therefore grown in a mix of black topsoil and coarse
grade turface, with grass immediately sown into the
medium surface. Grass should be allowed to grow no
higher than the plant. Light is medium to low to keep
plants cool. An annual sprinkling of lime is a must.
Cypripedium candidum
Because this species is found in open, moist and
sunny conditions, it should be given full light for a
good portion of the day. As previously described,
prairie shade is essential as tolerance of the plants to
the sun decreases as the season progresses. Grasses
should definitely be included in the pot and allowed to
slowly overgrow the plant. Substrate should be black
and mucky, kept very moist. Although the species can
be found in clayey soils affording little drainage,
perlite, turface, coarse gravel or something
comparable should be included for aeration and
drainage. The pH should not be allowed to drop below
7.0.
Cypripedium xandrewsii (Fig. 13, 14)

Cultural requirements of this nothospecies are
nearly identical to Cypripedium candidum, with the
exception of light. Plants are usually larger statured
than C. candidum, with broader leaves. This would
point to less intense light; however, such is not always
92
the case as hybrids can be found adjacent to the

species. As a general rule, light levels should be
intermediate between the two parents; however,
increased levels can be tried easily enough with little
risk to the plants. Overexposed plants will blacken
slightly at the leaf tips within a day or two and can be
readily moved to a shadier location.
Cypripedium cordigerum
This montane species from India prefers cool
summers but will do well if provided with enough
shade, in a cool pot. It is very sensitive to winter
moisture, and rotted flower primordia within dormant
buds result from plants stressed during shipping,
aborting their flowers. Any soft portions of rhizomes
or buds should be removed before fall planting to
reduce chances of rot. During the dormancy period,
all efforts should be made to keep plants continually
cold, and free from any moisture. Substrate should
consist of an aerating component such as perlite, etc.,
a moisture retentive component such as chopped
sphagnum, a drainage component such as gravel or
coarse sand, and finally clay or loam as the soil base.
Lime should be given annually.
93
Cypripedium montanum
Mountain lady’s-slipper
94
Cypripedium kentuckiense
This is likely the easiest to grow of all
Cypripedium species, surpassing even Cypripedium
parviflorum in its amenability to cultivation. Light
should be low to moderate; however, higher levels
will encourage faster growth. Substrate can be
completely inorganic, composed entirely of well
drained sand and gravel. This would, therefore,
require regular fertilizer application. It is said that this
species enjoys seasonal inundation with water, as well
as frequent repotting; however, success can be
achieved without much of either.
Cypripedium montanum
Apparently C. montanum can be found in forest
clearings and edges, though not in direct sun, very
reminiscent of C. reginae. Substrate for this selective
species should be a silty soil, with gravel, lots of
perlite, and conifer needles. A top-dressing of lime is
95
Cypripedium parviflorum var. parviflorum

Southern small yellow lady’s-slipper
96
recommended. Very sensitive to excessive moisture,

especially in the winter, this species apparently does
not enjoy continual moisture and appreciates drying
out. Excellent drainage is essential.
Cypripedium parviflorum (Fig. 16)

With such a hugely variable complex, it is
difficult to give specific cultural recommendations as
the variety of habitats occupied is vast. Plants can be
found on dry limestone barrens or in black, mucky
acidic bogs, thereby requiring different cultural
practices. If habitat information is available for a
particular plant, then attempts should be made to
duplicate those conditions. Because artificially
propagated divisions or seedlings of this species are so
readily available, specific habitat information is not
always accessible (a fortunate situation in a
roundabout way). The following general
recommendations can be followed: substrate can
consist of basic black garden soil (or peat or clay),
amended with perlite, gravel, etc. for aeration and
drainage. Humus can be incorporated, especially for
woodland plants, however, it is not necessary.
Watering should be regular, and light levels from low
to high. Annual sprinkling of lime is a good idea but
may not be appreciated by all strains. Grasses planted
into the pots are very effective. Because this complex
is so highly adaptable, it can thrive under new
conditions that may be very different from its initial
97
abitat, greatly reducing the need for extremely precise

habitat duplication under cultivation.
Cypripedium macranthos and relatives

This species can be difficult to establish from
poorly rooted imported rhizomes, and can be resentful
of division. However, it will thrive under the right
circumstances. As recommended for Asian species,
the top layer above the rhizome should be well
drained, but not necessarily inorganic. Pumice stones,
coarse sand, gravel or fine bark can all be used
individually or in combination. Rooting substrate can
be entirely inorganic gravel, sand, etc., or with a
significant humus component such as peat humus,
mulched oak leaves, or chopped sphagnum.
Calcareous clay can also be included. The pH should
be kept between 6.0–7.0. Sowing grasses into the
medium is recommended.
Cypripedium acaule
Without a doubt Cypripedium acaule has
proven to be the most difficult species to maintain
under pot cultivation. Numerous plants have been
victim to overwatering, or unexplained decline over
the years. An acid substrate is essential, as well as the
excellent drainage and aeration that a pine needle
mulch provides, however additional factors are not
fully understood. A number of growers have had
success raising it in beds, however, cultivation
principles applied to pots do not seem to carry over.
98
Perhaps increased root zone temperature encountered

with pot culture is too much for this species to contend
with, or there is some vital mycorrhizal interaction not
accommodated in pot culture on which this species
depends heavily.
Cypripedium formosanum
This Taiwan native is one of the few species
requiring winter dormancy that usually cannot stand
freezing, although some growers have had no
problems with hardiness. Because of the need for
winter dormancy, plants should be kept in a cold
room, ideally with temperatures between 1–5 C. Plants
can also be unpotted annually and kept in a plastic bag
with some damp sphagnum moss, and stored in a
refrigerator. Substrate should be moisture retentive
with a high peat component, with cut up moss, sand,
perlite, or gravel for aeration. The pH should be
neutral to acid, and light levels very low.
Cypripedium guttatum and C. yatabeanum

Under cultivation, Cypripedium yatabeanum
appears to be quite vigorous, and not ideally suited to
pot culture because of its wandering, stoloniferous
rhizome. Substrate should be stony or sandy, with a
high humus content, particularly for C. guttatum as it
is more of a woodland species. The pH does not seem
to be critical. Light levels can vary with different
strains, therefore, some experimentation is usually
99
required. Fertilizer should never be given to this

group as it is very sensitive and will burn easily.
Cypripedium passerinum
Under cool northern temperatures, this species
enjoys a period of full sun in early morning or late
day. However, under warmer conditions it must be
kept in deeper shade to keep it cool. The surface of
the medium can be mulched with moss to keep cool,
or planted with grass. Substrate should be clay-based
with lots of sand or similarly draining material.
Cypripedium reginae (Fig. 17)

Full sun is appreciated by this stately species for
a few morning hours. Plants like to be very moist, but
not sopping. If using primarily organic substrate, one
should resist the urge to keep pots in a tray of water
for continual moisture. It does not like this, and will
not send any roots down into the saturated portion of
the medium. Apparently this approach is successful,
however, when very well-drained inorganic medium is
used, likely due to its increased aeration. Black,
mucky soil can be used with sand or perlite to loosen
it. The pH should be adjusted to 6.5–7.0. Some
growers mulch with dry grass clippings. Pot partners
usually are not necessary as the leaves tend to block
out most of the sun before reaching the pot surface,
eliminating the need for the cooling effect of the
grasses, as well as reducing the chances of their
successful establishment.
100
References and Recommended Reading
Anderson, A.B. 1990. Asymbiotic germination of seeds of

some North American orchids. Pp. 75-80 in North
American Native Terrestrial Orchid Propagation and
Production Conference Proceedings, March 1989.
Brandywine Conservancy, Chadds Ford, Pennsylvania.
Chu, C.C. and K.W. Mudge. 1996. Propagation and

conservation of native lady‘s-slipper orchids
(Cypripedium acaule, C. calceolus (C. parviflorum) and
C. reginae). Pp. 107-112 in North American Native
Terrestrial Orchid Propagation and Production
Conference Proceedings, March 16 & 17, 1996. North
American Native Terrestrial Orchid Conference,
Germantown, Maryland.
Cribb, P., and C. Bailes. 1989. Hardy Orchids. Timber Press.

Portland, Oregon.
Harvais, G. 1982. An improved culture medium for growing

the orchid Cypripedium reginae axenically. Canadian
Journal of Botany. 60:2547-2555.
Malmgren, S. 1993. Orchids from seed? Large scale

propagation of Cypripedium calceolus. Wildflower.
Summer 1993:32-34.
——. 1996. Orchid propagation: theory and practice. Pp. 62-71

in North American Native Terrestrial Orchid
Propagation and Production Conference Proceedings,
March 16 & 17, 1996. North American Native
Terrestrial Orchid Conference, Germantown, Maryland.
101
Steele, W. K. 1995. Growing Cypripedium reginae from seed.

American Orchid Society Bulletin. 64:382-381.
——. 1996. Large scale seedling production of North American

Cypripedium species. Pp. 11-26 in North American
Native Terrestrial Orchid Propagation and Production
Conference Proceedings, March 16 & 17, 1996. North
American Native Terrestrial Orchid Conference,
Germantown, Maryland.
Acknowledgments
This document and all it represents would not have been
possible without the continued support, generosity, and
friendship of the following people. Thank you all very much.
Gunter Preusse, Svante Malmgren, Fred and Boots Case, Carson

Whitlow, Allan Anderson, Bill Steele, Larry Peterson, John M.
Doherty
John W. Doherty, 23711 32nd Ave., Langley, British Columbia,

Canada V3S 7B9. John became interested in orchids when he
was 16, based on an introduction by his father. Since then he
has studied and practiced their propagation and production, with
a specific interest in Cypripediums. His interests led him to the
Univeristy of Guelph, where he recently graduated with a B.Sc.
in Plant Biology. He is entering his fourth year of the judging
program of the American Orchid Society and is currently
working for Brookside Greenhouses, Langley, British
Columbia, in commercial orchid production.
102
Appendix I
Commercial Sources for Artificially Propagated
Cypripedium
Spangle Creek Labs

West 2802 Depot Springs Road
Spangle, Washington 99031
USA
Walter Bongers
Kolner Strasse 30
D-45145 Essen
Germany
Cyp Haven
2291 - 280th Street
Adel, Iowa 50003-9417
USA
Paget‘s Orchids
P.O. Box 119, Mirboo North
Victoria 3871
Australia
Raising Rarities
PO Box 405
Jacksonville, Vermont 05342
USA
106
107
Fig. 7 C. kentuckiemse
108
109
110
111
112
Fig. 16 Cypripedium parviflorum var. makasin ‗Sara Cond‘s
Flower‘ CCM/AOS, grown in plastic kitty littler pan with
appropriate holes drilled for drainage.
113
Fig. 17 Growing bench with various clones of Cypripedium
reginae.
114
115
116
117
118
119
120
GLOSSARY
achlorophyllous = without chlorophyll
acid = a pH of less than 7; a high concentration of hydrogen
ions
aeration = the supplying of air; the providing of oxygen
alba = refers to white-flowered forms
albinism = the lack of all color
albinistic = lacking all color or pigment
alkaline = sweet or basic as opposed to acidic; with a pH of
more than 7
alkaline substrate = soil with a pH of greater than 7; often
limestone
alliance = a group of loosely related species
alternate leaves = leaves that are arranged on a stem in uneven
steps; with only one leaf at a node
ambient temperature = surrounding; encircling temperature
amino acid = an organic compound which contains both an
amino group (NH 2) and a carboxylic acid group (COOH),
especially any of the 20 compounds that have the basic formula
NH2CHRCOOH, and that link together by peptide bonds to
form proteins
amorphous = lacking definite form; shapeless
anther = the pollen-bearing/male reproductive organ; at the tip
of a stamen
areal = of or pertaining to an area
aseptic = free of pathogenic microorganisms:
asymbiotic germination = the germination of seed without the
benefit of mycorrhizae
autotrophism = (autotrophic) able to obtain its food by
photosynthesis (rather then by feeding on something else)
barrens = areas with very little vegetation
bulb pans = shallow pots, broader than deep
121
calcareous = calcareous areas are areas with a limy or
calcareous substrate (usually limestone) and with a relatively
high pH. Calcareous species are species which grow best on
alkaline substrates with a high pH (calciphiles).
canopy = upper story of the forest
chitin = a tough, protective, semi-transparent substance,
primarily a nitrogen-containing polysaccharide, forming the
principal component of many arthropod exoskeletons and the
cell walls of certain fungi
clay substrates = clay soils
climax vegetation = the vegetation of an area that will be
present ultimately after a number of successional changes; the
plants that will be present in an area after it has been left alone
for an indefinite period of time
clone = individuals of identical genetic make-up which result
from asexual reproduction
colonized = said of an area on which have developed sizable
groups of given species
complex = a group of closely related species
coniferous = cone-bearing; most are evergreen
conspecific = belonging to the same species
cortex = the region of tissue in a root or stem lying between the
epidermis and the vascular tissue.
cultural = refers to methods used for cultivating plants
cytokinin = any of a class of plant hormones that promote cell
division and growth and delay the senescence (aging) of leaves.
deciduous = annually losing and replacing its leaves; falling
early
deflasking = removing from a flask
dormant eyes = growth buds that have not yet begun to grow
dormant = not actively growing
dorsal sepal = the uppermost part of the outer perianth
dynamic = continually altered or changing
embryo = the initial stage of a developing organism
empirical = knowledge gained from observation
122
endogenously = produced or growing from within
enzymes = any of numerous proteins or conjugated proteins
produced by living organisms and functioning as biochemical
catalysts; they speed up the rate of chemical reactions without
being used up themselves
epidermal = on the surface
everted = turned inside out or outward
excision = removal by or as if by cutting
exoskeleton = a hard outer structure, such as the shell of an
insect or a crustacean, that provides protection or support for an
organism; an outer skeleton
explant = living tissue which is removed from the natural site
of growth and placed in a medium for culture
floral stance = the position of the flower
flower primordia = flowers or parts in their most rudimentary
form or stage of development
fungicides = chemicals used to kill fungi
genera = plural of genus
genus = a taxonomic term for a group of closely related species
germinate = to start opening or growing (a seed)
halictid = a specific taxonomic group of bees
herbarium = a collection of dried, pressed plant specimens
herbivory = the eating of plants [eaten by deer!] ( as compared
with carnivory, the eating of animals)
hirsute = with long, soft hairs
homology = the relation of comparable structures, with a
common origin
humus = a growing medium rich in organic matter
hybrid = the progeny of two different species
hyphae = thread-like filaments which make up the vegetative
part of fungi (mycelium = a mass of hyphae)
in vitro = in an artificial environment outside the living
organism
infolding = with the edges folded inward
123
infrastructure = an underlying base or foundation especially
for an organization or a system
inorganic = a growing medium lacking organic matter; not
produced by living things; lacks carbon
interface = a surface forming a common boundary between
adjacent regions, bodies, substances, or phases
isotopes = one of two or more atoms having the same atomic
number but different mass numbers; atoms having the same
number of protons but a different number of neutrons and a
different weight
kinetin = a plant hormone that promotes cell division
LR White resin = a plastic-like compound used in microscopy
to make material very strong for sectioning with a microtome
labellar fissure = the opening in the pouch
leaf bracts = reduced leaf-like structures
lipids = any of a group of organic compounds, including the
fats, oils, waxes, sterols, and triglycerides, that are insoluble in
water but soluble in common organic solvents, are oily to the
touch, and together with carbohydrates and proteins constitute
the principal structural material of living cells
meristematic = the undifferentiated plant tissue from which
new cells are formed, as that at the tip of a stem or root; plant
tissue in which the cells divide and form new growth
metabolized nitrogen = nitrogen that has been absorbed by
living organisms
microbial = having to do with microscopic organisms
microclimate = localized habitat; the climate of a small,
specific place within an area as contrasted with the climate of
the entire area
montane grasslands = mountain meadows
montane species = high elevation species
morphological = size and shape
mutant = an individual in which a new characteristic appears
resulting from a mutation
124
mutation = a sudden and permanent change in the DNA which
makes up a gene
mycological = having to do with fungi
mycorrhizal = the symbiotic association of the mycelium of a
fungus with the roots of certain plants, such as conifers,
beeches, or orchids
necrosis = death
neutral = having a pH of 7 or close thereto; neither acid nor
alkaline
nothomorph (nm.) = taxonomic term for different forms of a
nothospecies derived from different varieties of the parent
species
nothospecies = taxonomic term for a hybrid that has received a
name
opposite leaves = the arrangement of leaves such that two
leaves are arranged evenly across from each other on the stem
orifice = the opening of the spur at the base of the lip
ovaries = the seed bearing organs at the base of the pistil(s);
female reproductive organs
parasites = organisms that live on and at the expense of other
organisms
pathogen = an agent that causes disease, especially a living
microorganism such as a bacterium or fungus
pericycle = a plant tissue characteristic of the roots, located
between the endodermis and phloem; it is from this layer that
root branching takes place
pH = the pH scale is based on the concentration of hydrogen
ions and gives the acidity or alkalinity; a pH of 1-6 is acid, 7 is
neutral, and 7 - almost 14 is alkaline ( basic)
pheromone = a chemical secreted by an animal, especially an
insect, that influences the behavior or development of others of
the same species
photosynthetic = capable of photosynthesis, the production of
food by green plants. In the presence of chlorophyll and light
125
energy from the sun, carbon dioxide and water are converted
into glucose (food) with the release of oxygen.
phytogeographic = the study of the geographic distribution of
plants
pigmentation = density of color
population ecology = study of the distribution, environment
and habitat; the relationships of plants to each other
pot culture = the growing of plants in pots
propagules = any of various usually vegetative portions of a
plant, such as a bud or other offshoot, that aid in dispersal of the
species and from which a new individual may develop
protocorm = the initial stage of development for every orchid;
a small ―ball‖ of cells
rhizome = a thickened surficial or underground stem from
which
roots and growth buds emerge
rhizosphere = within the area of the roots
rockwool = an artificial growing medium, spun from melted
rocks and then cooled, resulting in a structure very similar to
fibreglass
saprophytes = plants, fungi, etc. that feed on dead organic
matter
sepal = the outer portion of the perianth of the flower
shale = a metamorphic rock formed from clay or mud, which
splits into thin layers
soil pH = a scale for describing the relative acidity and/or
alkalinity of the soil
solar radiation = warmth and light, etc., from the sun;
whatever is radiated by the sun
speciation = evolutionary process that creates new species
spur = slender, tube-like appendage at the base of the lip
staminode = sterile stamen; shield-shaped structure found
within the flower of Cypripedium; the pollinia are found
beneath (there are no true stamens as such)
static = not changing
126
stele = the central core of vascular tissue in a plant stem or root;
it includes the xylem (conducts water) and phloem (conducts
dissolved foods)
stoloniferous = reproducing vegetatively by means of stolons,
creeping stems which run near the surface or along the ground
and send forth new roots and shoots
sub-opposite = appearing to be opposite because they are so
close together
substrate = the soil, sand, planting medium, etc., on which a
plant grows; also, the substance acted on or changed by an
enzyme
sucrose = a crystalline disaccharide carbohydrate, C12H22O11,
found in many plants (table sugar)
symbiosis = a relationship of mutual benefit or dependence
systemic fungicide = a fungicide absorbed through the roots or
leaves and then throughout the plant
taxonomic = having to do with classification
taxonomy = system of classification
temperate = a climate that has all four seasons; the temperate
regions are the regions between the tropical regions and the
poles
temperate terrestrials = plants that grow in the ground in areas
that experience the four seasons; land plants
transient = remaining in a place for a brief time only
transpiration = the loss of water by plants through the stomata,
or openings on the lower surfaces of leaves
tropical = a climate that is uniformly warm with minimal
seasonal variation
turface = an artificial growing medium consisting of clay
pellets, coming in a wide range of shapes and pellet sizes
vacuolated = containing vacuoles; small cavities in the
cytoplasm of cells, bound by a single membrane and containing
water, food, or metabolic waste
variety = taxonomic rank below species; a species may be
divided into varieties
127
vegetative = refers to the non-reproductive parts of a plant, such
as roots, stems, and leaves
vegetative primordia = organs or parts in their most
rudimentary form or stage of development
vernalization = chilling plants to induce them to grow
128
LOST & FOUND
from Richard Manuel in England:
Dear NANOA,
As a recent subscriber to your alliance, and your

Journal - congratulations on getting such a necessary
and interesting publication off the ground and flying!
I am writing to make known to you the

existence of the Hardy Orchid Society (of the UK),
an organization with similar interests and aims to your
own. We have been in existence for 31/2 years now
and our membership, understandably smaller than
yours, has stabilized at around 150. Our starting base
was rather different. Whereas it seems that NANOA
grew initially from a number of people mainly
interested in field work and study of orchids in their
habitats, HOS started largely as a grower's society
(hence the name) and has since adopted many
members who study and photograph orchids in the
wild, as well as a solid cadre of members who are very
much involved in growing native orchids from seed.
Our current chairman is the well known photographer
and writer on European orchids, Paul Harcourt Davies,
and we have many other members who are equally
well known in this country for their knowledge of wild
orchids in Europe. Some of us are interested in all
aspects of the subject; others concentrate on just one.
We currently hold two meetings during the year,
one in November and the other, an AGM (which we
manage to keep down to only a small part of the
proceedings) in May. We get a good turn-out for
these meetings even though, as a national society,
many members have to travel considerable distances;
an encouraging demonstration of the level of interest
in our subject! Meetings usually feature several talks
on the subjects already mentioned. Inevitably the
'travelogue' type of presentation is prominent, as many
of us go off on vacation somewhere in Europe that just
happens to have orchids in flower at the time.
Another popular feature is the 'culture forum', in
which a member gives a presentation on some aspect
of culture followed by a general discussion involving
the whole membership. It is a gratifying feature of our
society that we are blessed with many knowledgable
and articulate members, so these sessions become both
informative and enjoyable.
In the summer we try to run two or more Field Trips to

either one large venue, or a small number of localities
in the same area. We are hoping to become a little
more ambitious and try at least one society trip abroad
next year
We have just initiated a quarterly Society Newsletter

which is proving popular after three editions.
130
I hope that our two societies will be able to set
up and maintain some sort of permanent contact. For
instance, if any NANOA members are visiting the UK
or other parts of Europe and maybe hoping to visit
orchid-rich sites, we almost certainly have the
expertise somewhere in our membership to advise on
when and where to look in most parts of the UK and
continental Europe, and perhaps you could reciprocate
in this. And of course, if any of your members
happen to be in England in November or May, they
would be most welcome to attend our meetings.
Richard Manuel, Honorary Secretary, the Hardy

Orchid Society, 45 Thorncliffe Road, Oxford OX2
7BA, ENGLAND.
richard.manuel@zoology.oxford.ac.uk
from Aaron Hicks in New Mexico:
Quite recently, I have started on a new

endeavor: the Orchid Seedbank Project. It is, as the
name implies, a seedbank for orchid species. I would
like to know if members of your organization would
be interested in the existence of such a seedbank.
Something I would like to stress would be the

native North American species; obviously, there is
considerable interest in this realm - but an absence of
seed for people to exchange in order to develop
techniques towards propagating the native species.
The seedbank would serve as a sort of "catalog" of
131
seed that is available from different collectors (and,
therefore, representations of many different
individual colonies), such that conservation and
preservation may take place.
This is a completely new project. I have been

testing the waters, as well as acquiring lab supplies
to assure quality and viability of the seed we are
receiving. It's not a large project as of yet, but we
want to start it out right.
I hope that we can work together towards

conservation of the species of orchids that exist in
the United States and Canada. This is a project that
could easily become a life's work, and quite frankly, I
hope it does so.
Members that are interested in this project

should contact me at ahicks@nmt.edu or write me at
the address below.
Aaron J. Hicks, Director

Orchid Seedbank Project
PO Box 1873
Socorro, NM 87801
132
PUBLICATION ANNOUNCEMENT
MISSOURI ORCHIDS by Bill Summers

Third Edition now available from Missouri
Department of Conservation, Jefferson City, Missouri
$5.00 Telephone 573-751-4115 ext. 335 to order
Several revisions and new color photos. If you have

the previous edition you will want this one for the
additional information and photos, otherwise you need
this new edition to cover the orchids in this part of the
country. It is a real bargain!
CORRECTION
On page 344 of the December issue in the article by
Ed. Greenwood The Florida Govenia, the common
name of bottle flowered govenia was added by the
editorial staff. The Journal has a policy of adding
common names to each Latin name whereever
possible. This common name, although in use in
Florida, is inaccurate and was not included by the
author. Greenwood makes this abundantly clear in
subsequent paragraphs. The editor takes full
responsibility for adding this common name.
133
LOOKING FORWARD
June 1997
Orchid Search - Manitoba 1996
Cyrtopodium in Florida
Texas Saprophytes
All Tressed Up and No Place To Go -

Ladies‘-tresses in the Northeast
Cypripedium parviflorum var. pubescens in New

Mexico
Why Plants Change Names
Field Trip Etiquette
and much more!
134
135
136
137

March 1997 North American Native Orchid Journal

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March 1997 North American Native Orchid Journal

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NORTH AMERICAN

Editor: Paul Martin Brown

The Journal welcomes articles, of any length, of both a scientific

1999 Membership in the North American Native Orchid Alliance,

COVER: Cypripedium reginae by Stan Folsom

This issue is a major departure from our

This issue would not have been possible

I trust you will find the work on the genus

Paul Martin Brown

The Slow Empiricist

A few articles back (Jargon? 2(3):228-238), I

The result is the glossary which is appended to

I was very wary of injecting my ideas into this

The Slow Empiricist

Terrestrial orchids are currently enjoying a

A circumglobal genus of the subfamily

Mexican natives, and the recently described C.

Pollination in North American Cypripedium has

communication. Besides foraging for food, bees also

Unfortunately, this tidy hypothesis relies

There are a great many misconceptions

This paper is by no means intended to act as a

Catling, P.M., and V.R. Catling. 1991. Pollination in North

Nilsson, L.A. 1979. Anthecological studies on the lady‘s-

The most recent treatments of this genus are by

justifies taxonomic distinction. In reality, the bloom is

Cypripedium arietinum has a limited range

Cypripedium plectrochilon Franchet is a

Cypripedium arietinum R. Brown

and is felt by some to be conspecific with C.

Subgenus Cypripedium *Alliance Cypripedium

readily distinguished in the trade from its North

Cypripedium candidum Muhl. ex Willd.2 (Fig. 2)

burns can astonishingly replenish such populations in

It is suggested by Cash (1991) that this species

region, such as a woodlot, is found immediately

In addition to floral pigment, plant stature also

result is a blend of both parents that can sometimes

Cypripedium cordigerum D. Don

Cypripedium kentuckiense C. Reed (Fig. 7)

Plants are always large statured. Flowers are

are hanging. Petals can become quite long, thin, and

Cypripedium montanum Dougl. ex Lindl. Fig. 8)

Again, Cash (1991) suggests this to be an alba

One would imagine a white-flowered race of a

mutant, differing from the rest of the population

Cypripedium parviflorum Salisb.

of this complex, one is required to rely on the

cultivated plants; his treatment of the complex is

Cypripedium parviflorum var. parviflorum

Cypripedium parviflorum var. makasin (Farwell)

Cypripedium parviflorum var. pubescens (Willd.)

stature has the greatest potential here to be large;

An apparently dwarf strain within the complex

Please see Cash (1991) for additional members

Preferring cool climates, Cypripedium

Please see Cash (1991) for remaining members

which are often sandy. The second habitat is

Cypripedium farreri W.W. Smith

Cypripedium formosanum Hay.

Taiwan, where it grows in bright shade. Conditions

Cypripedium margaritaceum Franch.

See Cash (1991) for remaining members of this