June 1997 North American Native Orchid Journal

1
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.
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NORTH AMERICAN NATIVE
ORCHID JOURNAL
Volume 3 June
Number 2 1997
CONTENTS
NOTES FROM THE EDITOR

138
PREPUBLICATION ANNOUNCEMENT
THE GENUS CYPRIPEDIUM
140
WHY DO PLANTS CHANGE NAMES?
P. M. Brown
143
CYPRIPEDIUM PARVIFLORUM VAR. PUBESCENS IN
THE SOUTHWEST
R. A. Coleman
150
SAPROPHYTIC ORCHIDS OF DALLAS
V. S. Engel
157
FIELD TRIP ETIQUETTE
The Slow Empiricist
168
HISTOLOGICAL AND DEVELOPMENTAL STUDIES
OF THE SHOWY LADY'S-SLIPPER
CYPRIPEDIUM REGINAE
K. Sokolski, A. Dovholuk, L. Dovholuk,K. Lavigne, K. Ganey & P. Faletra
198
THE GENUS CYRTOPODIUM IN FLORIDA
R. L. Hammer
176
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ORCHID SEARCH - MANITOBA 1996
L. Heshka
187
ALL TRESSED UP, BUT NO PLACE TO GO
or Local Ladies’-tresses Orchids of Ohio
T. A. Sampliner
215
BOOK REVIEWS
Wild Orchids of the Northeastern United States
Orchids of the Northeast
Orchids of the Ottawa District, Ontario
Orchids of the Bruce and Grey Counties, Ontario
Hidden Orchids
238
LOOKING FORWARD
September 1997
248
Color Plates
1. Cypripedium parviflorum var. pubescens
2. Hexalectris nitida, Hexalectris warnockii
3. Cyrtopodium paranense
4. Platanthera praeclara
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.
Volume 3, number 2, pages 138-250; issued June 16, 1997.
Copyright 1997 North American Native Orchid Alliance, Inc.
COVER: Dactylorhiza aristata var. kodiakensis by Stan Folsom
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NOTES FROM THE EDITOR
As we continue into our third year, this issue

presents a wide variety of articles from all over North
America. Our first article from the Canadian prairie
provinces is included as well as our first from Texas.
Many of you have traveled widely in North America
and I urge you to write something for a future issue.
The March issue devoted to the genus

Cypripedium was very well received and I welcome all
the new NANOA members who joined as a result of
purchasing single issues. We are continuing to reprint
that issue to fill the demand of single orders. Because
of the time demands of filling and printing individual
issues this will be the only single issue available. Full
sets of back issues for both 1995 and 1996 are still
available and will continue to be so. Contact Nancy
Webb for more information on back issues.
Ron Coleman‟s excellent article in Orchids,

March 1997, concerning our first conference in
Pittsburgh was very well received and garnered several
new members. We thank the American Orchid Society
for its generous coverage of the Alliance and our
activities.
Do not forget our second conference to be held

in Arizona this summer. There is still time to register.
138
An information flyer was included with the March
issue.
It is time to prepare for a 1998 calendar. The

subject will be the fringed orchises of North America.
Our new color process will ensure a better quality of
reproduction and this time ONLY vertical, 8X10”
color, glossy, prints will be considered. You may
either send the finished prints or slides. If I am aware
of slides that individuals have, I may be contacting you
for consideration. The calendar will be finished by
September, so all slides or prints to be considered must
reach me before August 1, 1997.
I trust you will continue to have a rewarding and

productive field season and look forward to your many
reports. Please remember the summer address is PO
Box 759, Acton, Maine, 04001-0759 and the
phone/fax is 207/636-3719.
Paul Martin Brown

editor
naorchid@aol.com
139
PREPUBLICATION ANNOUNCEMENT
Coming in August 1997 from Timber Press
THE GENUS CYPRIPEDIUM

by Phillip Cribb
In this monograph, distinguished author Phillip
Cribb has considered the history, biology, evolution,
conservation, cultivation, and classification of the
temperate slipper orchids of the genus Cypripedium.
Orchidists, particularly those interested in slipper
orchids and terrestrial orchids, as well as other
botanists and woodland and alpine gardeners will
appreciate this richly illustrated and highly
authoritative volume.
Cypripediums, popularly called lady's-slippers

or moccasin flowers, are the showiest and most
sought after hardy orchids, collected and grown by
orchid and alpine plant enthusiasts alike. They have
been used medicinally in North America and the Far
East for centuries, having been grown as far back as
2,500 years ago during the time of Confucius in China.
Interest in these plants revived in the nineteenth
century when plants from Siberia, North America, and
the Himalayas were sent to Europe, where the
Industrial Revolution gave people the time and
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resources to build glasshouses in which to grow exotic
plants.
Slipper orchids have fascinated horticulturists

and botanists alike, being set apart from other orchids
most notably by a prominent slipper-shaped lip. Their
pollination biology was first studied by Charles
Darwin. The major source of these plants has been
from the wild; thus many of the more accessible
populations of many species have declined through
overcollection. In Europe and North America, other
factors leading to extinction include habitat
destruction, particularly logging and agricultural
improvement, and drainage of natural habitats. The
only British species, also that country's rarest plant,
has been reduced in its range to a single known
locality and, tragically, to a single plant. This clump
has survived alone in its protected site in northern
England for more than fifty years. Yet the lady's-
slipper was common enough in the area in the
nineteenth century to be collected for table decorations
in a local hostelry and to be dug up to adorn gardens
in the region.
Most of the striking members of the genus are

natives of North America, growing well with ferns in
shaded, damp, acid soils rich in organic matter.
Although several species are thought to be difficult to
grow, the author clearly identifies plants suitable for
beginners and, for all species, provides formulas for
various mixes used successfully by growers of
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Cypripediums. No other book on these important
orchids exists at present.
Phillip Cribb is curator of the Orchid Herbarium

at the Royal Botanic Gardens, Kew. He is the author of
several books on orchids, including The Genus
Paphiopedilum and, with co-author Ian Butterfield,
The Genus Pleione, both in this series. He has traveled
widely in connection with his work and has studied the
genus Cypripedium in the wild on several trips to
China in the past twenty years.
Approx. 300 pp, 26 full-page color paintings,

98 color photographs, 51 black-and-white illustrations,
and 22 maps, 6 x 9", hardcover, ISBN 0-88192-403-2
Price not set, Publication date: August 1997.
Contact: Deborah Garman, Publicity Manager.
Available from Timber Press, Inc., 133 S.W. Second
Avenue, Suite 450, Portland, OR 97204-3527.
Telephone Nos. (800) 327-5680, (503) 227-2878, Fax
(503) 227-3070. http://www.timber-press.com/ and e-
mail at publicity@timber-press.com
142
Brown: WHY DO PLANTS CHANGE NAMES?
WHY DO PLANTS CHANGE NAMES?
Paul Martin Brown
The change of plant names after years of general

usage often confuses plant enthusiasts. Why plants
change names is, perhaps, the most frequently asked
question I receive from members and students alike.
The basic answer is quite simple; the reasons behind it
are not. Plants change names for one of two reasons:
the first is that the currently used name is either
incorrect or invalid—each species has only one validly
published name; the second is that recent research has
shown that the specific species is in need of a new
name. How? When? Why? Where??????
When a specific plant is to be identified, it has

both a genus and a species name given to it. These
names are in Latin and are the same for all botanists
worldwide. This two-part name is known as a
binomial. Binomial nomenclature is the naming of a
plant with two names: a genus name (the group of
similar plants) and a specific epithet (the name which
sets off the species). The name of the plant consists of
both the genus and specific epithet. For brevity the
143
word „species‟ is often used in lieu of „specific

epithet.‟
Prior to the work of Linnaeus in the mid-1700s,

plants (and animals, etc.) had a variety of Latin names
in various combinations, but not in a standard format.
Pre-Linnaean names were often long and cumbersome,
sometimes consisting of 10-15 descriptive terms. In an
effort for uniform classification, Linnaeus, in his
Species Plantarum of 1753, devised a system of
assigning a genus name and a specific epithet to each
distinct individual or group of individuals. The Latin,
or scientific, name of the plant consists of a genus and
a specific epithet followed by the name of the author,
the first person to validly describe the plant.
This is what we know as plant taxonomy: the

classification of plants, including their naming, or
nomenclature. Please keep in mind that we are
discussing plants here, specifically orchids. All of the
following guidelines apply to all plants, as opposed to
other organisms (birds, mammals, etc.) which may
have somewhat different rules.
For the next 150 or more years it was not

unusual for a single species to receive more than one
name, as different botanists each thought they were the
first to describe the plant. This was particularly true of
variable plants of wide distribution. The name that was
used the most became the most widely accepted name
for that species. With the formation of the
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International Association of Plant Taxonomists in the

early 20th century, it became apparent that a set of
well-defined rules were necessary to standardize plant
names. These eventually became the International
Code of Botanical Nomenclature, often referred to
simply as the Code. New editions of the code are
issued following each international conference. The
latest was in 1995. Among many other things, the
Code sets forth a series of universally agreed upon
guidelines for: (a) naming new species, transferring
names from one genus to another, or creating
subspecies, varieties and forms; and (b) more
importantly, what constitutes a valid publication of a
new name or combination.
The Code is very simple: the earliest validly

published name takes precedence. Sounds easy. Very
often it takes years of researching the literature to
determine what is the earliest validly published name.
Rules have been formulated that must be met to insure
a plant name‟s validity.
Since 1935, to be validly published, a species

(or subspecies, variety or form) must have a Latin
diagnosis (a brief statement in Latin stating the
significant differences from similar taxa) and have a
lengthier description in the language of the author; it
must cite a type, which is the specimen from which the
original description is made. The type specimen must
be deposited in a permanent herbarium.
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In addition, it must be effectively published.

That is, it must be published in either a journal or
major work on the genus that is a permanent work (not
a newsletter) and that is distributed and available to
the professional botanical community from which the
species is described. It is unfortunate that several good
species were not validly published and therefore their
names cannot be used (see NANOJ 1(3):254-266
concerning the taxonomy of the Kentucky lady’s-
slipper, Cypripedium kentuckiense). If a Latin
diagnosis is lacking, that omission invalidates the
publication; if the author‟s name is missing, that
omission invalidates the publication; or, if no type is
designated, that omission invalidates the name. If a
Latin name is used in the literature prior to publication
and not identified as in ed. ( a work that is in progress
but not yet published), it too can invalidate that name
for future use.
But back to names changing. Prior to 1900, new

species did not always have a full Latin description or
diagnosis, which complicates choosing the earliest,
valid name for the plant. Careful research of the
literature is the primary culprit for changing the name
of a popular plant to a less well known or even obscure
species name. According to the nomenclature rules, if
a name is correct (i.e. a prior publication that has been
validly published) it must be used. But it can often be
awkward. An example of this can be seen in the genus
Malaxis, the adder’s mouths. Malaxis macrostachya
(Lexara) Kunze is correctly known as M. soulei L.O.
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Williams, because William‟s publication predates

Kunze‟s.
Upon occasion when a species is deemed to be

too broad and two or more species are the result of the
research, the original name can only be applied to the
correctly identified type specimen. The use of the
more familiar name may or may not be employed for
the most widespread and/or familiar resulting species.
Malaxis ehrenbergii (Reichb. f.) Kunze is widely
distributed in Mexico but the plants of northern
Mexico and adjacent Arizona, New Mexico, and west
Texas were determined to be a new species, Malaxis
wendtii Salazar (see NANOJ 2(1):65-66.
Name changes at the generic level have the

same rules but here we have two guiding forces. One
is, of course, the correct or valid name. The other
force is taxonomic opinion. This is the result of
research that some taxonomists clearly feel
demonstrates that the genus is in need of division and
more than one genus is the result. Sometimes, it
simply utilizes prior genera and the species are
grouped according to their technical characters;
alternatively, a new genus is described to
accommodate those species that do not fall into
existing genera. Which genera to use and which
species belong to each genus are very often matters of
opinion. Each taxonomist tries very hard to state his
or her case and some will agree and some will not.
This is a nuisance, especially when a well-known and
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familiar species has been changed. An effort has been

made to reduce this by a list of conserved genera, i.e.
genus names which cannot be changed even if an
earlier name is found. There is however, no provision
for conserving specific epithets.
In the North American native orchids, the two

groups in which we see this segregation happening the
most, are the fringed and rein orchises, Habenaria
and the ladies’-tresses, Spiranthes. For the past half
century the genus Habenaria has been a large, all-
inclusive one that accommodated many species.
Recent works, especially like that of Carlyle Luer,
have demonstrated that several of the original genera
that were merged into Habenaria are more
appropriately retained as separate genera. Hence, we
now see Platanthera, Coeloglossum, Pseudorchis and
Piperia being used. These are not necessarily new
names, but simply the earlier names for smaller groups
of species (see NANOJ 2(1):28-28 for details).
Another interesting example is the genus

Orchis. Two species were always well known from
North America: showy orchis, Orchis spectabilis and
small round-leaved orchis, Orchis rotundifolia.
Again, relatively recent work in the mid-20th century
demonstrated that neither of these species actually had
the critical characters that the Eurasian species of
Orchis possessed. In this case the earlier name of
Galearis accommodated Orchis spectabilis but the
creation of a new genus, Amerorchis, was needed for
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O. rotundifolia. The plants do not change, only their

names.
Paul Martin Brown
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Coleman: CYPRIPEDIUM PARVIFLORUM VAR. PUBSCENS IN THE
SOUTHWEST
CYPRIPEDIUM PARVIFLORUM
VAR. PUBESCENS IN THE SOUTHWEST
Ronald A. Coleman
By the end of the summer of 1995, my struggle

to find the large yellow lady‟s-slipper, Cypripedium
parviflorum var. pubescens, in Arizona had proceeded
in vain for three years, and showed no signs of abating.
In early 1996, I decided to expand my horizons
somewhat and look for it in the neighboring state of
New Mexico. The large yellow lady‟s-slipper has
historically been more numerous in New Mexico than
in Arizona, and might, therefore, prove easier to find.
My motivation was that knowledge gained by studying
the plant in habitat there might yield some clues on
where to look for it in Arizona. Several weeks of
research turned up multiple documented locations
within six to eight hours of driving time from my
home.
Tom Todsen, an orchid researcher who lives in

New Mexico, agreed to go with me. On 25 May we
met for lunch in a town part way to our destination,
and caravaned the rest of the way. We made a brief
stop en route to explore one of Tom‟s favorite orchid
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places, and found spotted coralroot, Corallorhiza

maculata, and a Schiedeelia species in bloom. We also
found Wister’s coralroot, Corallorhiza wisteriana, in
fruit. That evening we camped by a small stream and
planned the route for the following morning.
Tom had visited this lady‟s-slipper colony many

years earlier, and had a pretty good idea of how to get
there. The orchids were in a box canyon, about six
miles from the road. We could get there either by
going cross country along the ridges, or by following
the stream bed. We decided to go in via the ridges,
and come out along the stream. Not too far from
camp, an informal hunter‟s trail headed into the back
country, and we started following it. The first half mile
was up hill, but then the trail leveled off in fairly open
piñon pine and juniper forest. The walking at first was
easy, but after a couple of hours, the trail became faint,
and finally disappeared altogether at the base of a large
hill. We had noticed a canyon coming in from the left
for the last mile or so, and we headed toward it around
the side of the hill.
The easy walking was over, though, because the

hillside was covered with football-sized loose rocks
interspersed with several species of cacti. Eventually,
we saw in the distance a section of canyon wall that
stood out because it was green and lush in sharp
contrast to the rest of the mostly brown terrain; we
assumed that this was our destination. The last two
miles were to take much longer than the first four.
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SOUTHWEST
Between us and the green wall were arms of the

canyon that were too steep to traverse, and several
times we were forced to back-track up the hill looking
for safe passage. Finally, we came to a sheer drop of
over 100 feet, but just beyond lay the green section of
canyon wall.
Off to the side we saw a wooded slope leading

down into the canyon. By hanging onto trees and
bushes we managed to work our way to the bottom,
and then waded and rock-hopped upstream to where
we hoped to find the orchids. The green patch, toward
which we had painfully worked our way, was an
enormous seep about 40 yards wide that reached about
200 yards up the canyon wall. At the stream bed,
sections of the seep ended in straight stream-cut bank,
while other sections were collapsed piles of mud and
muddy boulders. Water was dripping off the entire
length of the seep. Ten or twelve feet up, the slope
was not nearly as steep, and the seep was covered with
grasses and sedges.
We scanned the seep from the bottom, and had

almost decided we were either in the wrong place, or
too early in the season, when I noticed a speck of
yellow out of the comer of my eye. Blooming in the
grasses only a few inches from the edge of the cliff
was a Cypripedium parviflorum var. pubescens.
It took several minutes of wet, muddy climbing,

but we worked our way up to the orchid. First Tom
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pushed me, than I pulled him, and eventually we

reached some soggy but stable ground. The entire
seepage area was wet to the touch, and our feet sank in
a little at each step.
The orchid we had spotted from below was so

close to the edge of an area of recent collapse that it
seemed certain to fall into the stream during the next
major storm. Scattered around the seep were ten
additional large yellow lady’s-slippers; most of these
were a safer distance from the cliff. Seven of the
plants were either in bloom or in bud. The blooming
plants all held single flowers, and were between 15 cm
(7 in.) and 30 cm (12 in.) tall. The pouches were a
bright yellow, and the staminodes were yellow with
red dots. The sepals and petals had a base color of
green, nearly totally covered with dark brown stripe-
like markings that turned to scattered dots near the
column. Pines and firs lined both sides of the seep, but
the orchids were in full sun for portions of the day.
The surrounding grasses were nearly as tall as the
lady‟s-slippers, and it was very difficult to see the ones
that were not in bloom. In an even wetter part of the
seep were hundreds of Platanthera in spike. Tom
speculated they were Thurber’s bog orchis, P.
limosa, because he had seen it elsewhere in the area.
We spent some time photographing the orchids

and exploring the seep, but then realized it was getting
late. The trip back to camp along the stream bed was
as difficult as the way in along the ridges. The canyon
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SOUTHWEST
bottom was littered with a jumble of boulders, and we

had to cross the stream many times. It would have
been impassable in spots had the water level been
higher. We did not find any other orchids along the
stream, although the habitat looked as if it should
support stream orchis, Epipactis gigantea.
The trip to and from the seep was long and

rough, but thanks to Tom Todsen, I achieved my
objective of learning a little more about the habitat of
yellow lady’s-slippers in the Southwest. Shortly
afterwards, I finally found Cypripedium parviflorum
var. pubescens blooming in Arizona. They were
growing in grassy, damp soil by a small stream,
slightly more shaded but very similar to the habitat in
New Mexico. I wish that I could claim the discovery
in Arizona was made easier by the study in New
Mexico, but it was really just a matter of luck (and
perhaps that should be another story).
Ronald A. Coleman
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155
Engel: SAPHROPHYTIC ORCHIDS OF DALLAS
156
SAPROPHYTIC ORCHIDS OF DALLAS
Victor S. Engel
In the summer of 1981, I was wandering around

the escarpment of southwest Dallas, Texas, in search
of something photogenic to put on the last half of a roll
of film so I could send it off to be developed. My
wanderings brought me to a grove of junipers and
oaks. After looking around for a while, I noticed
something that looked at first glance like an orchid. I
did not know there were any orchids in the Dallas area,
so I was quite surprised. On closer inspection, it was
clear that I was looking at the stem of an orchid plant
laden with seed capsules. There were no leaves or
open flowers on the plant. I looked around for others
but found none except a smaller plant growing right
beside the first one. When I got home, I looked
through one of my orchid books and concluded that
what I had found was probably a species of coralroot,
Corallorhiza.
In August of 1982, I went back to the same

place to see if the plant had come up again.
Unfortunately, it had not. So, I looked around the
immediate area for other plants. It was not long before
157
I found about a dozen plants full of flower buds and

developing ovaries. Only one plant had an open
flower, and that flower was open only partially. I
looked through several orchid books to try to identify
the orchid, but I could not find any matches. Any plant
that looked even remotely like the right species had a
range that did not include the Dallas area.
The following year, I started sooner in my

search for the plants in order to catch them in an
earlier stage of development. However, instead of
finding immature inflorescences of the same species, I
found another species in full bloom. I sent a specimen
to the Orchid Identification Center at the Marie Selby
Botanical Gardens in Sarasota, Florida, and the plant
was identified as Wister’s coralroot, Corallorhiza
wisteriana, a fairly widespread species whose range
includes the Dallas area.
The plants I was originally looking for did not

come up that year at all. It wasn‟t until July of 1986
that they came up again. This time, I dissected one of
the flowers to try to make a proper identification. I
noticed that instead of having four pollinia, as
Corallorhiza does, it had eight. By that time, I was
somewhat familiar with the crested coralroots, the
genus Hexalectris, so I concluded that the plant was a
species of that genus. The size and shape of the
flowers when spread out flat indicated that the plant
was the shining crested coralroot, Hexalectris nitida.
However, the range was wrong.
158
Hexalectris nitida had been reported only in

Panther Hill in the Glass Mountains of Brewster
County, Texas, and in the Sierra Mojada in the state of
Coahuila, Mexico. There were some other problems.
In Carlyle A. Luer‟s The Native Orchids of the United
States and Canada excluding Florida, H. nitida was
described as having open flowers with sepals and
petals curving backward toward the stem. The plants
in Dallas very rarely had open flowers. Another
difference was that according to Luer, there were five
ridges on the tip of the lip‟s mid-lobe. The plants in
Dallas had five ridges on the lip, but the ridges were
not at the tip. Because of the differences, I sent a
specimen to the O.I.C. for identification. Soon I got a
reply confirming that the plant was an autogamous
form of Hexalectris nitida.
In Dallas, Hexalectris nitida seems to prefer

living in the leaf litter of a mixed juniper and oak
forest. Although I have seen some plants in oak litter,
the vast majority prefer a bed of decaying juniper
needles below which is limestone bedrock. Although
all the plants seen were growing near the escarpment,
they did not grow past the edge. Presumably, this is
because moisture cannot accumulate very well on the
sloped surfaces of the escarpment. Also, the soil is not
as thick on the slanted surfaces.
Corallorhiza wisteriana grows in the same

forest as Hexalectris nitida. However, C. wisteriana
159
shining crested coralroot

Hexalectris nitida
160
prefers forest dominated by oaks, whereas H. nitida

prefers junipers. Corallorhiza wisteriana also grows in
poorer, sandy soil that is populated by grasses. Plants
growing in these areas were always smaller than the
ones growing in oak litter. Corallorhiza wisteriana
also seems more tolerant of sunlight. Plants frequently
are seen in clearings in the forest.
Another saprophytic species previously

unknown in this area is Texas purple-spike,
Hexalectris warnockii. When I found the original plant
of H. nitida in 1981, I told Dale Williams, an orchid
grower who lived near the location of the orchids. I
asked him if he had ever seen any native orchids in the
area, and he said that he had not. My discovery piqued
his curiosity, and after he saw the plant that I had
found, he kept his eye out for orchids on a neighboring
property that was downwind from the plant that was
blooming. In the summer of 1986, he was rewarded
not only with many plants of H. nitida but also with a
new, showy species, H. warnockii. This species was
known to have occurred in parts of central Texas as
well as the Big Bend area in Brewster County. Thus,
Williams‟ discovery constitutes an extension of the
previous range.
Another saprophytic species reported in the area

in 1986 was the crested coralroot, Hexalectris
spicata. They were seen at the Fort Worth Nature
Center in Fort Worth, Texas. Dale Williams and I have
also seen them on the property of the Dallas Nature
161
Texas purple-spike
Hexalectris warnockii
162
Center (previously known as Greenhills

Environmental Center). In the Dallas area, H. spicata
blooms in May and June, whereas H. nitida blooms in
late June and July.
During our studies of the genus Hexalectris in

Texas, Dale Williams and I found confusing plants
that resembled Hexalectris spicata but were not that
species, nor could they be placed with any other
currently recognized taxon. Among their distinctive
features were relatively small auto-pollinating flowers
that rarely opened fully, their columns lacked a
rostellum and the keels of the lip were relatively low.
Consistently for 10 years, plants of this
distinctive taxon have appeared in the Dallas area, but
in 1992 I found similar plants in the Austin area of
Texas. These plants were eventually recognized as the
Arizona crested coralroot, Hexalectris spicata var.
arizonica Catling & Engel (with a range from central
Texas to southeastern Arizona and adjacent Mexico).
At the Dallas site, Hexalectris spicata var.

arizonica occurs within an area of oak-juniper
woodland that is occupied by both H. nitida and H.
spicata var. spicata, whereas at the Austin site it is
accompanied by H. nitida. Hexalectris spicata var.
arizonica flowers in June and July, later than H.
spicata var. spicata which flowers primarily in May
(occasionally to early June) in eastern Texas.
163
crested coralroot
Hexalectris spicata
164
Arizona crested coralroot

Hexalectris spicata var. arizonica
165
Most of the plants seen in the Dallas area and all

of those seen so far in the Austin area have closed
flowers that never open, but some plants with open
flowers have been found at the Dallas site. The closed
flowers tend to have more truncate perianth parts than
the open flowers.
Athough it may be more widespread than is

currently known, Hexalectris spicata var. arizonica is
nevertheless, based on current information, a
geographically restricted taxon. Fortunately, it is
protected with other native flora in some of the sites
where it occurs.
Editor’s note: This article is based upon two very different

earlier works by Victor Engel in the AOS Bulletin 56(8):831-
835 and by Paul M. Catling and Victor Engel in Lindleyana
8(3):119-126. They both appeared in a different format and
have extracts merged here. The previously published material
is used with the permission of the American Orchid Society.
The latter article is the original publication of Hexalectris
spicata var. arizonica and goes into great detail conerning the
morphology and potential origins of the variety. The reader is
166
urged to consult that reference for full details concerning the

variety.
VICTOR S. ENGEL, 3305 Yellowpine, Austin, Texas 78757
167
FIELD TRIP ETIQUETTE
The Slow Empiricist
This is not an easy article to write, but as the

spring season has nearly ended and orchid enthusiasts
have begun to embark on another series of
explorations into the field it is an article whose time
has come. Let me begin by stating the obvious—that if
everyone treated their fellows with the same courtesies
that they would like to receive, then this article would
be a waste of time. However, that does not seem to
always be the case. If you think the previous statement
about courtesy smacks of Biblical overtones you are
right. Unfortunately, in our hectic modern world little
credence is given to the “golden rule.” You only need
to get in your automobile and drive a little slower than
the speed limit to find out that human nature is not as
tolerant as it perhaps should be. This article has a two-
fold purpose: first, to make the field trip participants
aware of some of their less than admirable behaviors
(which may help them become more sensitive to
others); and second, to encourage the others to find
gentle ways to foster true concern for each other when
sharing orchid expeditions.
168
If you are alone when you are in the field in
your orchid exploration you can have pretty much of a
free rein as to how much time you take with the plants
and you will not be spoiling someone else‟s pleasure.
That is unless you roll on the plants or crush some
under foot in your quest to get the perfect shot for your
slide collection. It is also common sense that you don‟t
disturb the environment so much that you endanger the
plants, such as by weeding out the detritus just so you
can get an unimpeded view of your subject (some
orchids need protective cover). Nor is it such a good
idea to leave some kind of noticeable marker to call
attention to the species lest some less scrupulous
person happen upon the scene and decide to collect the
plants for his/her garden or flower vase.
When you are with others, however, the rules

change. Most people are very considerate of their
companions and try to share the site with their
neighbors. If you recognize yourself in any of the
following situations I would suggest that you might be
considered a tad selfish. Do you always have to be the
first one to take photographs? Do you always push to
the front and hog the best specimens? Are you the first
to complain about anything, be it weather, insects,
plant material, others‟ behavior or whatever? Are you
such an inveterate perfectionist that you have to have
at least an hour or more to achieve a satisfactory
result? It is to these few that I suggest the following
behavioral changes. It would be to everyone‟s benefit
169
if the less generous field trip participant were able to
recognize his or her bad traits and work to modify
them.
First, when you get to the site of one of the

specialties you have been dying to photograph or
commune with ever since you were bitten by the
orchid bug, hang back! Let someone else have first
dibs on the plants! While you are impatiently waiting
for your turn you might try exploring for other
specimens. You might be rewarded for your efforts by
finding even nicer ones than the original ones you
were so eager to get at. If you have really become a
sharing, caring companion you will share your
discovery right away, even to the point of letting
someone else have first go at them. By abandoning
thoughtless me-first behavior you may find the
rewards are greater than you realize. I urge you to
persist.
Second, if you are a complainer, then I suggest

you thank your lucky stars or whatever you like to give
thanks to that you have been able to get out and arrive
at the state of being you are now experiencing. There
are plenty of fellow enthusiasts who through no fault
of their own can‟t be there. They have grown too
elderly to make the trip. They have failing eyesight, a
case of influenza, or even such mundane problems as
job constraints on their time in the field or some other
such problem such as an unsympathetic spouse who
begrudges their time spent on field trips. Looking at
170
your irritations in this way may help you be a better
companion to your fellow sufferers for I acknowledge
that every field trip doesn‟t always result in wonderful
experiences. The weather may be miserable or the
insects intolerable or the habitat daunting, but a good
attitude will help the situation rather than make it
worse.
If you go prepared for the unexpected, you will

have a better time of it. I always carry a little kit of
things that will make my stay a little more pleasurable
and help me survive the more nasty elements of nature.
In my kit I carry some extra food (in case I wander off
the beaten path and believe me I have; or the
expedition lasts longer than my stomach can take). I
pack a small flask of water in case the weather is too
hot for me. I also take a non-aerosol can of bug
repellent for I know the best orchids have guardian
insects who love my tender flesh. I also tuck in some
tissues in case I have a problem that needs wiping or
swabbing. If I know the terrain is going to be rough, I
carry my walking stick to help me up and over
obstacles. I also cover myself with layers of clothing.
If it is too cold my layers help protect me. If it gets
increasingly warmer, I can divest myself of my layers,
draping them about my waist or slinging them over my
arm. I always wear a hat. It keeps low branches from
tangling my hair and bugs have less of an area to
attack. Another item I find indispensable is a small
pair of field glasses. They help me locate plants on
hillsides and off into the distance without having to
171
trudge onto the actual site when I am exploring. There
have been many times when I have given silent thanks
for having remembered my binoculars. It should go
without saying that the avid photographer should not
stint on the amount of film or equipment he needs to
really do the job. When you are miles from anywhere,
running out of film is a real calamity for the
photographer. If all my paraphernalia seems a little
too much, you can tailor your survival kit to your
specific needs.
To the inveterate perfectionist, I suggest that

you wait to be last with the plants so that you can
spend all the time you need to study. One good
example of the perfectionist being aware of his fellows
is this: when he became aware others were waiting, the
photographer allowed others to photograph while he
was setting up or in between takes of his. If this
situation discombobulates you, find a specimen of
your own to work with. If that doesn‟t appeal to you,
consider coming back on the morrow to conduct your
business. Perhaps you should not participate in group
outings if you can‟t speed up your photography time or
make some kind of allowance for the others in your
party. It should not be for your sole benefit and
reward.
Group expeditions should be a celebration of the

group‟s combined experiences that have been shared.
That means accepting the levels of expertise of your
fellow members, their physical abilities (some of us
172
aren‟t as swift of foot as others or have such sharp
eyes as others). It also means being a cheerful
companion even if it sounds Pollyanna-ish to look for
the bright side in less than perfect situations. It means
being helpful and considerate of your fellow
celebrants. With attitudes like that you will have
pleasant memories of your outing even if it was less
than perfect or not up to your standards. You don‟t
need to sacrifice your standards. You just don‟t have to
make everyone else unhappy because you didn‟t
achieve one hundred per cent of your goals.
I hope that these messages have reached you in

a gentle manner and not offended anyone who suspects
that he/she might be seen as a self-centered individual.
If you have been guilty of such behavior in the past,
you might want to take a harder look at the way you
come across. You may find that a few slight changes
in your actions might reward you with more
camaraderie and a happier experience in the field. If
you are one of the patient ones who has been
victimized by someone who has been inconsiderate
while in your presence I urge you to speak up and
assert your rights in a courteous manner—if you can.
This might show the errant one good etiquette in the
field.
If not, you may have to resort to something

more drastic as one lady did who had put up with a
thousand complaints from a fellow member for over a
week. The member had complained about everything.
173
It was too hot! It was too cold or too windy! There
weren‟t enough plants or there were too many plants!
When she complained on the final day that during the
entire trip she had not taken one photograph worth her
time and money, her companion turned to her and
said, “Then you should learn to take better
photographs!”
Need I say more?
The Slow Empiricist
174
A MONOGRAPH OF CORALLORHIZA
(ORCHICACEAE)
by John Freudenstein
Harvard Papers in Botany no. 10 April 1997. ISSN
1043-4534. 51 pp. $27.00 + $2.00 shipping.
For those of us fascinated by the saprophytic

genus Corallorhiza it is a pleasure to seen John
Freudenstein work come together in a publication that
treats the entire genus in greatest detail. Whereas the
majority of species are native to North America it is of
great interest to many NANOA members. Each
species is treated in full detail with line drawings of all
the floral parts, distribution maps and extensive lists of
representative specimens. For those reader‟s who have
been following John‟s work through numerous
publications and his dissertation (Cornell 1992) over
the past 10 years, this publication pulls all of that
work, and more, together in a single highly usable
document. (PMB)
Copies may be ordered from Carolyn

Hesterberg, Harvard University Herbaria, 22 Divinity
Avenue, Cambridge, Massachusetts 02138.
175
Hammer: CYRTOPODIUM IN FLORIDA
THE GENUS CYRTOPODIUM IN

FLORIDA
Roger L. Hammer
There is but a single species of Cyrtopodium

native to the United States, and that species is
Cyrtopodium punctatum (L.) Lindl., colloquially
known as the cowhorn or cigar orchid, in allusion to
the long, cylindrical pseudobulbs. In the United
States, C. punctatum has been recorded on southern,
mainland Florida in Broward, Collier, Dade, Lee,
Martin, Monroe and Palm Beach counties where it
typically inhabits coastal mangrove-buttonwood
forests, cypress swamps and mixed hardwood swamps
and forests. It can be found growing epiphytically on a
variety of trees, especially cypress (Taxodium
distichum) and buttonwood (Conocarpus erectus), and
is also frequently found perched on the trunks of
fallen, dead trees. Although it is commonly found as a
terrestrial species in the tropical Americas, it is almost
always epiphytic in Florida.
Cyrtopodium punctatum, was first discovered in

the United States near Miami, Dade County, Florida in
March, 1877 by Abram Paschall Garber, PhD. The
species was first described by Carl von Linné (who is
better known by his latinized name Carolus Linnaeus)
176
in 1759 as Epidendrum punctatum in the publication

Systema Alaturae. In 1833 John Lindley transferred it
to the genus Cyrtopodium, which he published in his
Genera and Species of Orchidaceous Plants. The type
locality was not stated but the global range of C.
punctatum includes southern Florida, the West Indies,
Mexico, Guatemala and Costa Rica southward to
Brazil and Paraguay in South America.
When in flower, Cyrtopodium punctatum is a

spectacular sight. In late winter each year, new growth
and flower spikes appear at the same time from the
base of the previous year's leafless pseudobulbs. The
inflorescence quickly towers over the leaves and by
late March or early April, showy flowers open. Each
widely-spreading flower bears greenish-yellow sepals
spotted with reddish-brown, bright yellow petals
spotted with reddish-brown, and a reddish-brown lip
with a yellow base. Each inflorescence may be
adorned by thirty or forty flowers and a large specimen
will literally display hundreds of blossoms. It has been
suggested that the flowers of Cyrtopodium mimic
members of the Malpighia Family, Malpighiaceae, and
that native bees, during their search for nectar,
inadvertently pollinate the orchid flowers.
Because of its attractive floral display, the

cowhorn orchid has been highly sought by collectors
over the years. Photos of wagonloads of cowhorn
orchids being brought out of the Everglades just after
177
the turn of the present century reveal how common

this species once was, but today it is rare and found
only in rather remote, protected areas. The historic
abundance of Florida's orchids was documented by the
famed naturalist Charles Torrey Simpson and his
friend, correspondent and renowned botanist John
Kunkel Small. In 1916, Simpson and Small went on
an orchid collecting trip in southwestern Florida and
reported that the orchids were so abundant that their
harvest could scarcely be noticed.
Cyrtopodium punctatum is the only species

mentioned by Dr. Donovan Correll in his publication
Native Orchids of North America north of Mexico
(Stanford University Press, 1950). Dr. Carlyle Luer,
however, in his book The Native Orchids of Florida
(The New York Botanical Garden, 1972) reported a
yellow-flowered, terrestrial species from Florida which
he referred to as Anderson’s cowhorn orchid,
Cyrtopodium andersonii (Lambert ex Andrews) R.
Brown. Although C. andersonii is cultivated in the
Miami area, the species that is naturalized in southern
Florida, which Luer photographed, has been
determined to be another tropical American yellow-
flowered, terrestrial species known as Parana
cowhorn orchid, Cyrtopodium paranaense Schlecter.
The notes of the late field botanist George N.

Avery reveal an interesting account of the original
reports of this plant in Florida. While attending a
Native Plant Workshop meeting in Miami on June 15,
178
1971, a woman named Betty McCormack mentioned

to Avery that a Cyrtopodium had volunteered inside
her screen porch in limestone gravel. Nearly one year
later, on May 12, 1972, McCormack brought Avery a
portion of her flowering Cyrtopodium, which Avery
pressed and deposited in the herbarium at Fairchild
Tropical Garden in Miami (Avery #1155). Although
McCormack told Avery that she had found a picture of
it in Exotica 3 as Cyrtopodium andersonii, Fairchild
Tropical Garden's superintendent Stanley Kiem
correctly identified it as Cyrtopodium paranaense.
Avery pointed out that he had found an article by John
Beckner concerning C. andersonii in the Florida
Orchidist, the publication of the South Florida Orchid
Society (Vol. 7, No. 2, pp. 78-79, 1964), that treated
this species as a native, and an earlier issue of the same
publication (Vol. 2, No. 4, p. 5, 1959) that featured an
article by F. S. Shuttleworth reporting C. andersonii
from Florida's west coast.
Luer (1972) states that Fred Fuchs, a Miami

orchid grower, identified some flowers of an orchid
brought to him "from a plant growing terrestrially deep
within the Everglades" as Cyrtopodium andersonii.
Luer also mentions that "occasionally someone reports
finding a specimen [of Cyrtopodium andersonii] in the
Everglades, but [it] usually turns out to be a yellow
form of Cyrtopodium punctatum." It should be pointed
out that there are no verifiable collections of any
yellow-flowered Cyrtopodium from the "Everglades,"
and that the native C. punctatum is the only species
179
listed in the extensive botanical inventories for

Everglades National Park and the adjacent Big
Cypress National Preserve, together encompassing
over two million acres. Cyrtopodium paranaense does,
however, occur as a naturalized species in at least one
natural area—a sandy pineland preserve in the Kendall
area of Dade County—where it is firmly established
and thriving, even after an intense wildfire in April,
1995. It also readily escapes locally around residential
areas wherever it is cultivated in southern Florida and
perhaps elsewhere.
During several interviews and phone

conversations in June and July, 1972, Stanley Kiem
gave an interesting historical account of Cyrtopodium
paranaense to George Avery and myself. Kiem stated
that it was introduced into Florida in ca. 1949-50 by
John F. Kasper, who operated an orchid nursery at NW
27 Avenue and NW 15 Street in Miami. Kasper had
received a specimen labeled Cyrtopodium paranaense
from Sao Paulo, Brazil. Kiem acquired a plant from
Kasper in 1951 and cultivated it at his residence at SW
118 Street and SW 94 Avenue in Miami. It escaped as
a terrestrial in his yard and as an epiphyte on the
prostrate trunks of saw palmetto (Serenoa repens), as
well as on a dead pine trunk in a nearby pineland. In
September, 1972, Kiem told me that it had spread five
miles around his residence in five years. Kasper closed
his business and retired at SW 112 Street and SW 92
Avenue in 1954, bringing the Cyrtopodium with him.
Three years later, Kiem found plants naturalized at the
180
residence of Ed Jordan who lived just a few houses

away from Kasper.
On May 31, 1976, I sent flowers collected from

Jordan's residence to Dr. Kiat W. Tan, then associated
with Marie Selby Botanical Gardens in Sarasota. Dr.
Tan determined that the flowers belonged to
Cyrtopodium paranaense, and made a voucher
specimen of the flowers, which he deposited in the
Gardens‟ herbarium. A rather painful and
disheartening memory of collecting the flowers from
Jordan was that of while walking across his yard, I
occasionally stepped on crunchy objects beneath my
feet. They were mowed-off pseudobulbs of plants that
had come up in his lawn!
Dr. Gustavo Romero, Research Associate in

Orchidology in the Oakes Ames Orchid Herbarium at
Harvard University, borrowed specimens from the
Fairchild Tropical Garden in May 1991 and reviewed
the Cyrtopodium specimens deposited in the
herbarium. Dr. Romero attached a note to Avery's
1972 Cyrtopodium paranaense specimen stating that
"this material is indistinguishable from Cyrtopodium
glutiniferum Raddi," and a label was placed on the
sheet to reflect this determination.
Cyrtopodium glutiniferum is yet another tropical

American, yellow-flowered terrestrial species. In
December 1995, I telephoned Dr. Romero in an
attempt to clear up this puzzle. Dr. Romero admitted
181
to erring when he made the C. glutiniferum

determination and that he felt that the Florida plants
are correctly C. paranaense.
In May 1996, I sent Dr. Romero, by overnight

mail, a flowering, wild-collected Cyrtopodium from
the Kendall pineland site, as well as flowers of a
cultivated, robust, yellow-flowered Cyrtopodium
obtained in 1993 from Lynn Bretsnyder of Impact
Orchids in Princeton, Florida. Dr. Romero confirmed
by telephone that the wild-collected plant was
Cyrtopodium paranaense, and that the cultivated
specimen was C. andersonii. Dr. Romero has
deposited the wild-collected plant in Harvard
University's herbarium to further voucher its presence
as a wild plant in Florida.
Dr. Romero has in preparation an article

concerning the genus Cyrtopodium that he plans to
submit to the American Orchid Society for inclusion in
the publication, Orchids. His work will hopefully put
an end to the nomenclature confusion of Florida's
naturalized populations of Cyrtopodium, and help
identify yellow-flowered members of this genus in the
tropical Americas and in cultivation as well.
In a letter to me dated 8 April, 1996, Dr.

Romero stated "Cyrtopodium paranaense has smaller
flowers, and the sepals and petals tend to be roundish.
The larger flowers of Cyrtopodium andersonii and
Cyrtopodium glutiniferum overlap in size, but the
182
labellum is indented and the petals are lanceolate,

undulate in C. andersonii versus labellum not indented
and petals with a claw, roundish toward the apex in
Cyrtopodium glutiniferum."
Before finalizing this article for the North

American Native Orchid Journal, various reference
books were checked to determine the natural range of
Cyrtopodium andersonii, and here is the confusing
summary: Hortus Third (Staff of the Liberty Hyde
Bailey Hortorium, 1976) lists it from Venezuela to
Brazil; The Native Orchids of Florida (Luer, 1972)
shows a range that includes southern Florida, Cuba,
the Lesser Antilles and northeastern South America;
The Manual of Cultivated Orchid Species (Bechtel,
Cribb & Launert, 1986) has it occurring in the West
Indies and tropical S. America; and Garay & Sweet in
the Flora of the Lesser Antilles, Orchidaceae(1974)
determined that it is endemic to St. Vincent in the
Lesser Antilles. To defend their determination, Garay
& Sweet state that "Cyrtopodium Andersonii has been
confused with Cyrtopodium flavum and Cyrtopodium
Engelii by all contemporary authors since Lindley,
who united these two different species under
Cyrtopodium Andersonii." It is clearly apparent that
much work needs to be done on this and other species
native to the West Indies and the tropical Americas.
In closing, what has been determined to be

Cyrtopodium paranaense may likely be in Florida to
stay. It has already demonstrated its ability to survive
183
fire and periodic, short-duration freezes, it readily

produces seed pods, and it is capable of spreading
from seed in Florida. All it will probably take is for a
storm to move across southern Dade County when
seed pods are dehiscing to spread this South American
native into other favorable habitats; perhaps even into
the Everglades. It may just be a matter of time.
Roger L. Hammer 17360 Avocado Drive Homestead, FL 33030

Gustavo Romero, Orchid Herbarium of Oakes Ames, made
helpful suggestions to the manuscript.
184
185
186
Heshka: ORCHID SEARCH – MANITOBA 1996
ORCHID SEARCH - MANITOBA 1996

Lorne Heshka
I‟ve had an interest in orchids ever since I, as a

boy, saw my first yellow lady's-slipper back on the
farm in Saskatchewan. At that time there was no
doubt in my mind that this was an "exotic" flower and
later, when I studied botany at University, I discovered
just how unique this family of plants is.
This interest, however, remained at an

"awareness only" level until an assignment to Thailand
with The Food and Agriculture Organization of the
United Nations in 1989 and 1990. There I
"discovered" tropical orchids and was given, as a gift,
several plants. This started me on a new and exciting
hobby of growing orchids and I soon found that the
slipper orchid species became my favourites. These
slipper orchids, the genera Paphiopedilum and
Phragmipedium, reminded me of that first orchid I had
been fascinated with back on the farm those many
years ago.
In comparison to the hundreds of orchid species

found in tropical countries of the world, the 37 species
(and varieties) listed for the Province of Manitoba
appear insignificant. The harshness of our winter
climate is an important controlling factor, with winter
187
temperatures sometimes hovering around -35 degrees

Celsius.
Prior to 1996, I had observed only 20 of the 37

orchid species that can be found in our province.
Although it wasn't a New Year‟s Resolution, early in
1996 I decided to make an effort to find and
photograph as many of our native orchids as I possibly
could through the year.
A couple of weeks of vacation were booked

during peak blooming periods, and then the research
began. From previous experience, I already knew the
location of 20 species. These, however, were the most
common and the easiest to find. It was the other 17
that challenged me.
Sixty-five dollars‟ worth of text books from the

Museum of Man and Nature in Winnipeg ensured that
I at least knew what these flowers looked like and the
habitat they preferred. Fortunately, I also had several
contacts who could give me some specifics as to where
and when certain orchids could be found.
As I did my literature review, I came to the

realization that there were at least three species that I
wasn't going to see. These were ones that could be
found only in the far north and I had no plans to trek
up to Churchill that year. Another species has been
found only once before, in the Duck Mountains, and
188
has never been seen again. The most I could possibly

find, I decided, would be 33 species.
The largest number of species of our native

orchids occur in the southeast corner of the province,
and a large number are found in bogs. As a
consequence, my search was planned in the area with
the Red River as the western boundary, the Winnipeg
River to the north, the United States to the south and
Ontario to the east. This area of approximately 25,000
square kilometres has a variety of bogs and other
habitats suitable to orchids, and also has a number of
Ecological Preserves ensuring minimum human
encroachment.
The late spring, I quickly found, had delayed

flowering and so all of my projected blooming dates
were out between one and two weeks. It was,
however, an "orchid perfect" summer—ample
moisture at the right time, and warm but not scorching
weather. Mosquitoes were bad early in the spring, but
in the bogs where they are usually the worst, there
were few to none of those little pests.
I maintained a diary of our excursions and a

tally showed that my wife Joan and I covered over
5000 kilometres by car, about 75 km on foot (mostly
through bogs), and spent 175 hours searching in every
possible habitat. Joan caught the "orchid fever" just as
I had and was my constant companion. She became an
189
Amerorchis rotundifolia
small round-lef orchis
190
excellent "bog walker" and it was her sharp eyes that

helped us locate some of our rarest finds.
On the best day we located 11 different species,

the worst day, never fewer than 4 or 5. The weather
cooperated with ample periods of overcast, providing
excellent light for flower photography. Remarkably,
we were never "rained out.”
We began our search on June l in the Tall Grass

Prairie near Tolstoi located within a few miles of the
US border and ended our season there on September 7,
coincidentally finding our last species, Great Plains
ladies’-tresses, Spiranthes magnicamporum, within a
hundred meters of where we found the first, the small
white lady’s-slipper, Cypripedium candidum.
The "highs" of this project were unquestionably

the discovery of several new species (new to us, that
is) and meeting some super people along the way.
Finding not only one, but several, very rare albino
(white-flowered) showy lady's-slippers, Cypripedium
reginae forma albolabium, in a localized area had to
be the highlight for us—although my wife would argue
that point, considering the excitement I displayed
when we found for the first time the beautiful and
uncommon rose pogonia, Pogonia ophioglossoides.
The "lows" were finding spade holes where only

a week before a colony of the beautiful stemless
191
(pink) lady's-slipper, Cypripedium acaule, stood.

The "lowest" wad the news that a political decision
had been made to put a well into a bog to raise the
water level in a nearby resort lake, effectively
sounding the death knell for 19 species of orchids
living in that bog.
We never did make it to the far north for

sparrow’s-egg lady’s-slipper, Cypripedium
passerinum and the flat-petalled form of the large
yellow lady’s-slipper, C. parviflorum var. pubescens
(planipetalum form1). We never found the white
adder’s-mouth, Malaxis brachypoda, in the Duck
Mountains and the three other small indiscrete species,
auricled twayblade, Listera auriculata, northern
twayblade, L. borealis, and heart-leaved twayblade,
L. cordata var. cordata, eluded us. We had to satisfy
ourselves with thirty of a possible thirty-seven species.
The new friends we made, the fresh air and

exercise we enjoyed and best of all, a tremendous set
of transparencies to share with others, made this
project worthwhile.
1
The forma planipetalum Fernald is an extreme of the the var. pubescens;
see Charles Sheviak‟s article on the yellow lady‟s-slippers in the December
1996 issue of the Journal. Ed.
192
Platanthera hookeri
Hooker’s orchis
193
rose pogonia
Pogonia ophioglossoides
194
Following is a list of the orchids we observed

along with the date they were first found blooming in
1996:
Scientific Name Common Name First Observed

Amerorchis small round-leaved June 20
rotundifolia orchis
Arethusa bulbosa dragon's-mouth June 29
Calopogon tuberosus grass-pink July 1
var. tuberosus
Calypso bulbosa var. Venus (eastern June 10
americana fairy) slipper
Coeloglossum viride long-bracted green June 29
var. virescens orchid
Corallorhiza maculata western spotted July 7
var. occidentalis coralroot
Corallorhiza striata striped coralroot June 20
var. striata
Corallorhiza trifida early coralroot June 10
Cypripedium acaule stemless (pink) June 10
lady's-slipper
Cypripedium arietinum ram's-head lady's- June 10
slipper
Cypripedium candidum small white lady's- June 8
slipper
Cypripedium northern small June 15
parviflorum var. yellow lady's-
makasin slipper
Cypripedium large yellow lady's- June 15
parviflorum var. slipper
pubescens
Cypripedium reginae showy lady's-slipper June 29
Cypripedium reginae showy lady's-slipper July 7
195
forma albolabium (white flowered

form)
Goodyera repens lesser rattlesnake July 27
orchis
Goodyera tesselata checkered July 20
rattlesnake orchis
Liparis loeselii Loesel’s twayblade July 22
Malaxis unifolia green adder's- July 7
mouth
Platanthera dilatata tall white northern July 7
var. dilatata bog orchis
Platanthera hookeri Hooker's orchis June 10
Platanthera hyperborea tall leafy green June 20
var. hyperborea orchid
Platanthera lacera ragged fringed July 28
orchis
Platanthera obtusata blunt-leaf orchis June 20
forma foliosa
Platanthera orbiculata large round-leaf July 11
orchis
Platanthera praeclara western prairie July 6
fringed orchis
Platanthera psycodes small purple fringed July 27
orchis
Pogonia rose pogonia July 14
ophioglossoides
var.
ophioglossoides
Spiranthes lacera var. northern slender July 28
lacera ladies'-tresses
Spiranthes Great Plains ladies'- September 7
magnicamporum tresses
Spiranthes hooded ladies'- July 31
romanzoffiana tresses
196
Postscript - Because of the efforts of a close friend who was

unafraid to challenge the bureaucracy, the installation of the
well has been delayed until an environmental impact study is
completed. We are hopeful that the public outcry, as a result of
the media attention given to this issue, will result in the
consideration of other environmental-friendly solutions.
Lorne Heshka, 1204 DeGraff Place, Winnipeg, Manitoba, R2G

1Y8 CANADA
notes on the photos:
The western prairie fringed orchis, Platanthera praeclara,

photograph was taken in the Tall Grass Prairie adjacent to the US
boundary southeast of Winnipeg. 1996 was an exceptional year for
this species and the counts in this managed area for this year
exceeded the total for the previous three years together. Over 20,000
plants were counted and this truly is a success story which may be
worthy of follow-up.
The white-flowered form of the showy lady’s-slipper, Cyripedium

reginae forma albolabium, is found in the localized area threatened
by the well mentioned in my article. A rough estimate is that
approximately 5 to 10% of the population of Cypripedium reginae in
this area are the white flowered form. I haven't had the opportunity to
fully explore the extent of this colony with respect to this area.
The blunt-leaved rein orchis, Platanthera obtusata, photograph

demonstrates a characteristic I have observed in this species only
once previously. Part way up the flower stalk is a smaller, thinner
leaf (forma foliosa). This small group of plants exhibit this
characteristic while others in the same location were typical in that
they had only the single leaf.
197
Sokolski et al.: Cypripedium reginae
HISTOLOGICAL AND
DEVELOPMENTAL
STUDIES OF THE
SHOWY LADY'S-SLIPPER
CYPRIPEDIUM REGINAE
Katie Sokolski, April Dovholuk,

Lauren Dovholuk, Kim Lavigne, Karen Ganey
and Peter Faletra
The showy lady's-slipper, Cypripedium

reginae, is a terrestrial orchid found across the
northern regions of central and eastern North America.
In New Hampshire this species commonly grows in
marshy areas or fens preferring the more alkaline soils
found on the border of Vermont and New Hampshire.
Although these plants have never been found in great
numbers in New England, owing to both habitat
restrictions and loss, they are now critically
endangered in much of northeastern North America.
Cypripedium species require the help of insects

for pollination. Orchids, in general, are rather specific
as to the insects which they will accept as pollinators.
In some cases orchids can only be fertilized by the
198
males of a single species of insects. However, this is

not the case in lady's-slippers (Withner, 1974). The
physical arrangement of these flowers creates a tight
space through which the insects (mostly bees) have to
squeeze. A pollinating insect first passes by the
stigma, and upon exiting the trap rubs against the
anther. The insect that now carries pollen must reenter
another lady‟s-slipper to successfully carry out
pollination (Dressler, 1981). One study suggests only
twenty out of one thousand bumble bees are willing to
pass this gauntlet twice. When successfully pollinated,
like many orchids, Cypripedium seed pods contain
thousands of seeds. Because the seeds do not contain
endosperm (Dressler, 1981), they seem to require a
more rigorous set of conditions to germinate and
develop than seeds with endosperm. One of these
conditions is the presence of symbiotic fungi.
Although Cypripedium species, such as C. reginae and
the large yellow lady’s-slipper, C. parviflorum var.
pubescens, are rare, when they do occur, they can
often be found in great numbers, concentrated in a
small area. The habit of the flower to occur in this
colonial fashion seems, in part, owing to their
multiplication via rhizomes. The showy lady’s-slipper
is often regarded as the most regal of the genus with
two flowers (rarely three or four) occurring on three-
foot tall plants.
Cypripedium reginae takes about 8 years to

mature to a flowering state in the wild (Withner,
1974). Therefore, any approach which speeds this
199
maturational process may prove to be an important

adjunct to restoration attempts. In vitro culture could
be of extreme value since it can speed this process
considerably. Nevertheless, for in vitro culture to be
of practical worth a variety of challenges with regard
to efficiency must be met. When considering
efficiency, in vitro methods which affect germination
percent and speed, along with subsequent development
and survival, are central factors. Chu and Mudge
(1994)2 obtained acceptable germination rates for C.
parviflorum (as C. calceolus) when seeds were surface
sterilized in bleach for twelve minutes. Studies on C.
reginae (Dovholuk and Faletra, 1996; Dovholuk et al.,
1997) showed an increase in percent germination when
exposed for up to thirty-six minutes in bleach. Various
other approaches have been attempted to maximize
efficiency (Faletra et al., 1997; Sokolski and Faletra,
1997).
The general goal of this investigation was to

devise a method of efficiently producing large
numbers of Cypripedium reginae through in vitro
culture and micropropagation. Large scale in vitro
culture could then supply sufficient numbers of
seedlings for a restoration attempt. To this end, two
specific approaches were used: 1) Axenic seed
culture, 2) Micropropagation of seedling tissue.
2
Chu and Mudge address the plants they used as Cypriepdium calceolus var.
pubescens, an older and incorrect name for C. parviflorum var. pubescens.
See Sheviak, C.J. 1996. NANOJ 2(4): 319-343 for full taxonomic details.
Ed.
200
Axenic seed culture could supply seedlings for

restoration along with sterile tissue for
micropropagation investigations. Studies of the
morphology, histology, and development were done in
hopes that the normal processes of the seedling
development could serve as a guide to successful
micropropagation. These investigative approaches
were aimed at answering the following questions:
Where are the meristematic tissues?
What, if any, tissues are determined and what are
they destined to become?
At what point can determined meristematic tissue
be manipulated?
What are the areas of the seedlings which can be
micropropagated?
How can in vitro raised seedlings be transplanted
and raised to normal flowering plants?
Can this approach help alleviate the decline of an
endangered species in the face of habitat loss?
Axenic seed culture studies

Conditions that affect speed of germination and
subsequent development affect the efficiency of the
system. With respect to germination, since each pod
produces thousands of seeds, seed supply, although
important, is not a primary factor with regard to
efficiency. With an average plating density of about
201
202
203
45 seeds per culture (although we usually see over

90% germination) a greater than 10% germination rate
should give quite an adequate number of seedlings per
culture vessel. Figure 1 shows results from
experiments on germination rate and subsequent
development in seeds exposed to bleach for increasing
amounts of time. With germination beginning as early
as 12 days, a steady progress in efficiency was seen in
increased rates of germination and speed of
development of the seedlings at longer exposure times
to bleach. The differences among the groups were
significant at p<.05. These and following
experiments (data not shown) indicate that exposures
between 36 and 60 minutes in bleach give optimal
germination. Rates of development reinforce this
trend. The decrease in stage three seedlings seen in
Fig. 1 is most likely owing to their development into
later stages which show increasing numbers with
increased exposure to bleach. Interestingly, this and
other studies had never tried simply soaking seeds in
water as a control, since without any sterilent, cultures
would have little chance of growing free of
contaminants. This is unfortunate, since as the diluent
for bleach, H2O is the appropriate control.
Preliminary results employing a sterile distilled water
(SDH2O) control show extended time in H2O up to
sixty minutes increases germination rate and
development, but it is not yet certain whether the water
alone is as effective as bleach. Further studies are in
progress. An option would be to soak seeds in
204
SDH2O prior to, or after surface sterilization. This is

also being investigated.
An alternative to solid medium is liquid

medium. A disadvantage of liquid medium is the
reputed need for rotation which requires an expensive
orbital mixer. Therefore, it seemed reasonable to
investigate the need for rotation. Seeds placed in non-
rotating liquid culture were not as quick to germinate
and did not have as high a germination percentage as
seeds in rotating liquid culture. This may be attributed
to more effective oxygenation and elimination of
wastes in the rotating cultures in addition to a more
even osmotic pressure. These results suggest there is a
distinct advantage to rotating liquid media.
Our early studies and those of Chu and Mudge

(1994) showed liquid media to support higher
germination rates than solid medium when seeds were
sterilized at 12 minutes in bleach. Since Chu and
Mudge used Cypripedium parviflorum it was of
interest to perform a similar experiment on this
species. Preliminary results indicate that, as with C.
reginae, increased time in bleach up to 36 minutes
increased the germination rate and rate of development
for C. parviflorum.
We have also found that the germination and

development of Cypripedium reginae in solid
medium, for seeds exposed to bleach for more than 36
minutes, is about the same as for seeds in liquid
205
medium. The longer surface sterilization seems to

level the differences between solid and liquid media.
Consequently, although a liquid medium might give
slightly better percent germination, if the speed of
germination and development is not substantially
greater than solid medium there seems little advantage
in the liquid; especially considering the need for an
orbital shaker.
Histological and Developmental Studies

Because axenic culture of Cypripedium species
has only recently been successful, studies of
morphology and development have been somewhat
superficial. This is unfortunate since development is a
key to understanding any plant.
Scanning electron microscopy (SEM)

consistently reveals an orifice at only one end of the
seed (Fig. 2 and 2A). This hole could possibly be a
consequence of the incomplete closure of the
micropyle during seed development. The opposite end
was probably attached to the placenta and bears no
opening. Within the seed coat is contained an embryo
of approximately 500 cells enclosed in a coat possibly
derived from interior integuments (Fig. 3). Seed
germination studies indicate that in order for
germination to proceed this coat needs to be penetrated
by what is most likely water.
206
207
208
It was of interest to determine the fate of seedling

tissues. Figure 4 shows the various tissues of a
seedling approximately ten weeks old. Roots were
revealed by the presence of a root cap and unicellular
root hairs (Fig. 4A). Shoots were indicated by the
presence of stomata (Fig. 4B).
Light microscopy (Fig. 6 and 6A) shows that

early seedlings are composed of undifferentiated
parenchymal cells with no cotyledons which is typical
of orchids but atypical of the class Monocotyledonae
(Scagel, 1965). The cells of the seedling shown in
Figure 6A possess nuclei in a dense arrangement
indicative of meristematic tissue in the region we refer
to as the apex. It is meristematic tissue that is a logical
target for micropropagation. We have monitored this
area over a period of months in a number of seedlings
and in all cases have seen it develop into a shoot. This
has also been supported by SEM which shows this
tissue to contain developing stomata. Cross and
longitudinal sections of root are shown on Figure 7
and 7A respectively. The presence of central vascular
tissue made up of xylem and phloem indicate that this
is root tissue. The unilateral positions of starch
granules and nuclei probably indicate the horizontal
orientation of this root. The presence of large stores of
starch also indicate that these roots store substantial
amounts of energy. Figure 8 shows a longitudinal
section of shoot tissue opposed on either side by a
sheath which is believed to be a coleoptile-like
209
210
211
structure. This reinforces our belief that Cypripedium

grows much like grasses, with the exception of slipper
orchids having successive roots emerging at nodes and
rhizome tissue making up the internodes. This will
occur until the plant is stimulated to produce an aerial
shoot (see drawing).
From these and future studies that localize and

explain the development of both meristematic and
committed tissues, successful micropropagation will
be more probable.
Micropropagation
Unfortunately, the genus Cypripedium has not
yielded to attempts at micropropagation. Hoshi et al.
(1994) recently published results of a fairly exhaustive
micropropagation attempt, revealing marginal success
in the way of limited callus production and no evident
protocorms. Our micropropagation attempts with
similar approaches to that of Hoshi et al. (i.e., matrix
experiments) have not been successful. Histological
sections of early seedlings localized meristematic
tissue (Fig. 6). This seemed important for success in
micropropagation. Unfortunately, when these young
seedlings were sectioned, no success in
micropropagation occurred. This is possibly owing to
the difficulty in cutting such small specimens.
Nevertheless, attempts to micropropagate by
212
sectioning older seedlings were somewhat successful.

As discussed above, scanning electron micrographs
were taken to positively localize different committed
tissues. Knowing exactly where root and shoot tissue
were located helped guide sectioning. When 2-6
month old seedlings were cut approximately in half,
with a piece of a shoot and one or more roots on each
half, both halves developed into healthy seedlings and
eventually matured to the level at which they were
ready to be vernalized (Fig. 9). In some cases, young
seedlings could be cut in more than three pieces.
Older and larger seedlings, being easier to section and
possessing more shoots, often yielded more than three
sections. Although this approach might technically be
considered as micropropagation, it is not the most
desirable situation. Ideally, micropropagation
generates generous numbers of protocorms grown
from tissue randomly cut from a single plant's tissue.
We are now attempting some novel approaches

to micropropagation and are also working to improve
the methods for axenic culture of other endangered
orchid species. We also hope to be planting a
considerable number of showy lady’s-slippers in their
natural habitat this coming year.
We would like to thank Ms. Elaine Faletra for her help with
Light microscopy, Dr. Ezequiel R. Rivera for his histological
assistance and Dr. Charles Daghlian for his help in SEM. This
213
research was funded in part by a grant from the Toyota Motor

Co.
Peter Faletra, PhD, 59 Mountain Meadows Rd,

Warren, NH 03279
Literature Cited
Chu, C. & Mudge, K. 1994. Effects of Prechilling and Liquid

Suspension Culture on Seed Germination of the Yellow
Lady‟s-slipper Orchid. Lindleyana 9(3): 153-159.
Dovholuk, A., Faletra, P. 1996. Development of Cypripedium

Species in Plant Tissue Culture. Bulletin of American
Association for the Advancement of Sciences, Annual
Meeting 1996.
Dressler, R.L. 1981. The Orchids - Natural History and

Classification. Smithsonian Institution, Washington
D.C., Chapters 1-3.
Faletra, P., A. Dovholuk, K. Sokolski & T. King. 1997. Saving

Cypripedium reginae. Orchids 66(2): 138-143.
Hoshi, Yoshiikazu., K. Katsuhiko & H. Shuuichi. 1994. in

Vitro Seed Germination of Four Taxa of Cypripedium
Lindleyana 9(2): 93-97.
Scagel, Robert F. et. al. 1965. An Evolutionary Survey of the

Plant Kingdom. Wadsworth Publishing Company, Inc.
Belmont, California. pp. 584.
Sokolski, K. & P. Faletra. 1997. Growth Studies of the Showy

Lady‟s-slipper (Cypripedium reginae) in Axenic
Culture. Bulletin of American Association for the
Advancement of Sciences , Annual Meeting ,1997.
Withner, Carl L. 1974. The Orchids Scientific Studies. John

Wiley and Sons Inc., New York.
214
Sampline.: All Tressed Up, But No Place to Go
ALL TRESSED UP, BUT NO PLACE TO

GO
or
Local Ladies’-tresses Orchids of Ohio
Thomas A. Sampliner
Editor’s note: Botanical keys are often the bane of the amateur
and professional alike. In all of our native orchids no genus has
more similar species than the genus Spiranthes. In preparation
for a field trip in northern Ohio, Tom had prepared a
comparison of several books and field guides. I asked him to
expand it to make these comparisons more pertinent for many
of our members. It was not practical to reprint the various keys
here, but if you have any or all of the books it would certainly
be helpful to lay them out and follow along with Tom‟s
comparisons. Please keep in mind that regional books cite
habitats and flowering dates for their specific areas and often
that data does not strictly apply elsewhere.
At various points, most of the following authors

have had to fudge their keys by combining many traits
just to make one dichotomous choice. This represents
not only a weakness of a key for practical use but also
violates the concept of what a key is supposed to be.
An even greater failure of some of the keys occurs
when the author not only had to combine many traits
215
to make a separation between species, but had to

require further choices between groupings of opposing
traits before you can separate out another species.
Careful forethought in using the traits is necessary to
construct a key that will only require choices among
one, or maybe two, opposing traits at a time which
immediately separates out species. Admittedly, this is
the ideal. You can judge for yourself how well each
author has done. It must be stated that to have these
experts manifesting conflicts in dividing small from
large florets based upon lip measurement is
inexcusable. The failure of such a trait is
demonstrated by the authors‟ employment of
measurements that not only conflict with each other,
but also require exactness to the millimeter while in
the field. If the experts cannot agree upon, or are not
familiar with, existing writing so they cannot eliminate
such measurements or explain to the lay person how
measurements play a valid role in a key, then the trait
is invalid. There were also some traits used that the
authors felt necessary to caveat. This also is an
inappropriate key trait.
According to Carlyle A. Luer, in his

monumental effort entitled The Native Orchids of the
U. S. & Canada, excluding Florida (1975), the genus
Spiranthes (sensu latu)3 consists of over 300 species
3
Current taxonomic thinking has considerably revised the genus so that
there are 40-45 species and the remaining 300 or so are in a number of
segregate genera. This makes the genus on the whole much more workable.
216
ranging from temperate to tropical regions. Few

species occur in the old world. Only one species each
occurs in Japan, Australia and New Zealand. Africa,
as of 1975, was not known to have any.
In addition to Luer, the other written sources

consulted for this article were: Frederick W. Case Jr.'s
Orchids of the Western Great Lakes Region (1987); R.
T. Whiting & P. M. Catling's Orchids of Ontario
(1986); Paul Martin Brown's A Field and Study Guide
to the Orchids of New England & New York (1993)4;
and Homoya‟s Orchids of Indiana (1995). I had
contemplated using Merritt Lyndon Fernald's key in
the 8th edition of Gray's Manual of Botany (1950).
My decision not to use it was based in large part on (1)
the great differences in coverage, and (2) the traits
employed were so different that a fair comparison with
the others was impossible.
The initial discussion will contrast the keys from

each work with an emphasis on their treatment of local
species. This article is intended to be a preparation
and quick reference source for fall field trips in the
Ohio area when it might be possible to see as many
local species as a one day field trip might permit.
Depending on how the growing season has progressed
and the amount of driving you are willing to do in one
See Garay, L. 1980. A generic revision of the Spiranthinae. Botanical
Museum Leaflets 28(4). Harvard University, Cambridge. ed.
4
Recently republished ( March 1997) in a totally new format with 192 color
plates and new drawings as Wild Orchids of the Northeastern United States.
Cornell University Press, Ithaca, NY. ed.
217
day, you can see six or seven species dawn to dusk.

You can see even more if additional days are available.
It should also be useful for appropriate species in the
entire northeast.
Sadly, I must note that fall is too late in the year

to see shining ladies’-tresses, Spiranthes lucida. In
Ohio this species appears in June. Depending upon
when in the fall you reach Ohio, the same may be true
of northern slender ladies’-tresses, S. lacera var.
lacera, southern slender ladies’-tresses, S. lacera
var. gracilis and grass-leaved ladies’-tresses, S.
vernalis. You should be able to find the other species
in the northeast in accordance with local expected
bloom periods.
Whiting & Catling have selected lip size as the

initial feature for their dichotomous key. The choice is
between florets exhibiting lip size 4-7 mm in length
versus 8-11 mm, with an additional trait for some
species in the first category of being loosely spiraled
while all under the second choice are to be densely
flowered.
Therefore, in our area, Case’s ladies’-tresses,

Spiranthes casei var. casei, S. lucida, northern oval
ladies’-tresses, S. ovalis var. erostellata, and both S.
1acera var. lacera and S. lacera var. gracilis would
fall within the first division. It should be noted that as
of their writing, little ladies’-tresses, S. tuberosa, had
not yet been found in Ontario.
218
Case uses lip size for his first choice as well.

However, he opposes lips less than 6 mm long versus
those 6-11 mm. No companion trait accompanies his
first division. Case does include Spiranthes tuberosa in
his key.
Homoya starts his key by placing stems slender

with small flowers, the lip of the lowest florets 5 mm
or less, in opposition to stems stout with larger
flowers, the lip of the lowest florets 7 to 10 mm. His
key, therefore, also uses lip sizes as a means of
differentiating species.
Luer covers the widest geographical and

climatological areas so his first dichotomy uses such
traits as spiral appearance, ranking of the spirals, the
lip growth (called a tuberosity), and climate.
Obviously, this was done to separate the tropical,
subtropical, southern, and western US species. We
will, therefore, pass over these features and proceed to
the first couplet relevant to the Great Lakes area.
This is an examination of the spiral feature. He

opposes secund (appearing one-sided) to
conspicuously coiled, florets appearing in one rank
versus tightly spiraled flowers appearing in more than
one rank. However, he has a caveat under both
choices that occasional specimens can appear under
the other heading.
219
220
Paul Martin Brown, in his key, has leaves narrow and

grass-like at flowering time versus leaves orbicular or
absent at flowering. For the novice orchidist, this is by
far the easiest observation to use. It has easily
observable characters that the other keys lack at this
juncture.
Based upon the practicality of using a key in the

field in any region, but especially ours, I note a real
problem with Luer's key and an apparent conflict
between the size traits as used by Whiting & Catling
versus Fred Case‟s. I do not mind trying to follow
Luer in his spiral patterns, but if he finds a need to
caveat the separating traits, how useful is it? Keys that
conflict as to how to separate lip size are a real
problem. I do not see how Case‟s 6 mm versus larger
can be helpful if Whiting & Catling can distinguish
lowest florets of an inflorescence of 4-7 mm versus 8-
11. Homoya also uses lip size initially, but he provides
a very practical application by combining size with the
easily observable contrast of slender versus stout
stems. No one else employs this trait. His lip
measurements also avoid the dilemma posed by the
conflicting lip measurements in the keys of Case
versus that of Whiting & Catling. Several of the
authors employ ranking as a trait key. Homoya does as
well, but he goes further than the others by adding the
concept of phyllotaxy to it. Here Homoya cites
Sheviak (Biosytematic Study of the Spiranthes cernua
Complex, 1982) explaining that this is the number of
flowers necessary in the spiral to place the next
221
possible floret in a direct line with the one from which

you started. This is then expressed as a fraction, so that
if it took 5 flowers to get back into perfect alignment,
then the phyllotaxy is 1/5.
Homoya excels in his ability to communicate

various botanical concepts in clear yet precise
language. So that we can more clearly discuss the
keys, I will borrow from Homoya to set forth ranking
and phyllotaxy of the spiral florets. Ranking is the
spiral arrangement of the florets which can be
discernible as single, if so loose that a staircase effect
appears; or multiple to indiscernible, when florets are
so crowded and tight that two, three, or more separate
stairways of florets appear when you are looking up or
down the spike. If you look from the top of the spike
down, you can determine the phyllotaxy. Homoya
contends this concept is more useful than ranking in
separating confusing or atypical specimens. Certainly
this is observable in the field and it avoids the caveats
regarding spiral traits found in several keys.
I must say, I find Brown's first trait the most

highly visible compared to the others, with Homoya a
close second. Shape of and presence or absence of
leaves is readily visible in the field! Let's continue to
see more of who wrote what and how well they serve
our purpose when we are in the field.
Under the small floret species, Whiting &

Catling‟s key uses rostellum and viscidium lacking,
222
cauline (stem) bracts with long, spreading, recurved

blades versus the parts present and bracts being
bladeless. Choice one keys out to Spiranthes ovalis
var. erostellata. The other choice requires further trait
selection for species we will not see on a single field
trip in the month of September in the Cleveland area.
This key is of limited value on the small-flowered
species, unless we find either of the varieties of S.
lacera or S. casei. Frankly, even if we did find these
species, how practical for field use are such esoteric
features?
Case, under small flowered species, next uses

lip color: floor of the lip with bright yellow or green in
the center versus white or pale, creamy yellow
throughout the lip with no strong central coloration.
This is usable. Under lip color he has the two varieties
of Spiranthes lacera as well as the June blooming S.
lucida. Under the
white color we only find S. tuberosa. This latter
species is so distinctive by size alone that either way
we finish up with the locally expected species.
Brown's key ignored size and started with

leaves. Next he uses bright yellow central portions to
the lip, taking you immediately to Spiranthes lucida.
Lip otherwise sends you next to flowers white versus
otherwise. Other than white florets sends you to the
same point in his key that the 2nd choice for his first
trait sent us (leaves orbicular or absent at flowering
time). Therefore, Brown's key will lump: S. lucida,
223
hooded ladies’-tresses, S. romanzoffiana, giant

ladies’-tresses, S. praecox, and nodding ladies’-
tresses, S. cernua, as all having leaves narrow and
grass-like at flower time; S. lucida keys out at strong
yellow in lip central; fiddle (pandurate) shape lip keys
out to S. romanzoffiana; and multiple ranking keys out
to S. cernua from the S. praecox as well as their
habitat differences. Very usable in the field!—
especially since S. cernua is quickly removed from
yellow ladies’-tresses, S. ochroleuca and S. casei with
which it can be confused.
Luer's third key trait under the small flowered

species contrasts leaves that are narrow or linear,
lanceolate to oblanceolate versus leaves ovate, but
often fugacious (withering early). This is a useful trait
for field work. Just to list the species that fall under
linear-like leaves shows how useful: Spiranthes
vernalis, S. praecox, cienegas ladies’-tresses, S.
delitescens (as S. graminea), lace-lipped ladies’-
tresses, S. laciniata, S. casei ( as S. intermedia), long-
lipped ladies’-tresses, S. longilabris, S. ochroleuca
and southern ladies’-tresses, S. torta (syn. S. tortilis).
Sorting through what we may find here in Ohio,

this separation is very helpful. Assuming that S.
vernalis is still possible, S. ochroleuca is the only
other likely local species and lip color in S. vernalis
easily will separate these two. Since S. ochroleuca is
one of those that is in the original S. cernua complex,
224
it is extremely valuable if we can depend upon

fugacious leaves to quickly sort out S. ochroleuca.
Homoya next contrasts bloom time combined

with lip color. His first division is flowering May to
June, with yellow lip, which immediately keys out to
Spiranthes lucida. The contrast has flowering time
July or later combined with lip green or white. At
first, I didn't think the category was contrived;
however, I notice he had three small floret species left;
namely, Spiranthes lacera, S. tuberosa, and S. ovalis.
To progress, he contrasts under the late bloomers,
green versus white florets to separate out S. lacera.
That leaves two late blooming small floret flowers
which viewed in isolation could be difficult for the
amateur. He employs his phyllotaxy concept so that S.
tuberosa keys out with a 1/5 versus the 1/3 to 1/4 of S.
ovalis. He does also use lip shape and leaves absent or
only basal for S. tuberosa while S. ovalis has the
conspicuous stem leaves during bloom time. A nifty
solution for the final small floret species.
Next, let us look at the larger flowered species.

In order to get to Luer‟s treatment of the larger floret
species, we first looked at the spiral traits; then under
tight spiral, multi-ranked if we select narrow leaves,
we find a choice for lip size. Luer divides the species
from those under 9 mm to those above. Just what we
needed! Remember Case versus Whiting & Catling? I
can see it in the field: is it between 4-7 mm? more or
225
226
227
less than 6 mm? or more or less than 9 mm? If the

experts can't agree what can we do?
If we can get beyond this problem, continuing

with Luer, his under 9 mm next requires selection for
lip color and then habitat, including only Spiranthes
ovalis among species we are likely to find. Therefore,
we have a clear shot by using his larger than 9 mm lip
to find our local species. He next opposes lip
constricted near the center and northern or
northeastern distribution versus lip ovate and usually
not constricted near the center. If the lip is oblong to
ovate and only slightly constricted near the center you
are left with S. cernua or S. ochroleuca. If the lip is
pandurate (fiddle-shaped), dilated apex and white
flowered you have S. romanzoffiana (or in Luer a cross
called S. xsteigeri, which has proven to be S.
ochroleuca). In our area we need be concerned only
with the first two. Spiranthes cernua will have the lip
base dilated, tuberosities small and flowers uniformly
all white. Spiranthes ochroleuca will not be dilated at
the base, tuberosities large and flowers will have a
yellowish cast.
Back to Whiting & Catling for their larger

flowered species. Here, the first choice has basal
portion of the rachis glabrous (smooth), hairs sparse
and only 1 mm long: lateral sepals 3 mm or wider, lip
pandurate, perianth veins prominent; which
immediately identifies Spiranthes romanzoffiana. The
opposing features would be: basal portion of rachis
228
pubescent with capitate hairs 0.3 to 0.5 mm, lateral

sepals 2-3 mm wide, lip not pandurate, perianth veins
not prominent.
Next, under the second choice, we select

between: leaves absent at flowering, cauline bracts 4-7
mm, flowers white but central portion of lip is
yellowish and thickened, flowers fragrant, lateral
sepals diverging, calli (small, thickened calluses) 0.5
mm long which keys out Great Plains ladies’-tresses,
Spiranthes magnicamporum. The alternative is: leaves
present, cauline bracts 2-4, flowers white or cream
color, no strong fragrance, lateral sepals appressed,
calli 0.7-1.5 mm. This requires a further choice to
separate S. ochroleuca from S. cernua. If flowers are
cream color, dorsal and ventral curvature of the flower
pronounced, measuring a separation of between 0.5 to
1.5 mm between dorsal and lateral sepals, lip thick,
fleshy centrally, often with slight basal dilation you
have Spiranthes ochroleuca. The opposite choice
requires white flowers, moderate curvature resulting in
dorsal from lateral sepal separation of 0.1 to 0.9 mm,
lip neither thick nor centrally fleshy, and without
distinct lip dilation you are led to Spiranthes cernua.
Rather than looking for these features, which are

somewhat difficult in the field, I find helpful a habitat
observation whereby S. ochroleuca will be in the
sandier more well drained site. This can sometimes be
easily seen above the wetter area, including ditches,
which are preferred by S. cernua. Frankly, considering
229
all the keys‟ attempts to separate these two species,

this is probably the most useful and user-friendly
observation for field work.
Case classifies large-flowered species as lip 6-

11 mm. The first choice is lip pandurate, small basal
callosites, sepals and petals connivent forming a hood,
inflorescence heavy and thick, not strongly spiraled,
secund in rank. This keys to Spiranthes
romanzoffiana. The opposite is lip not pandurate, has
prominent basal callosities, no hood, strongly spiraled
spikes.
Under this latter choice, the next selection in the

key will direct you to Spiranthes magnicamporum
with leaves absent at flowering, lip deflexed giving a
gaping appearance, central lower lip yellow, above
thick white, callosities short, conical, as wide as high,
lateral sepals free and spreading, flowers strongly
fragrant. Its opposite is: leaves present at flowering, lip
color uniformly white or creamy, basal callosities
longer than wide, lateral sepals appressed.
Under this latter category we elect between:

flowers pale cream, yellow or faint green tone, upper
lip surface deeper yellowish cream versus flowers
white, crystalline and in multiple ranks, lip white
throughout, dorsal and lateral sepals and petals
approximate, slightly recurved but gaping or a 2-lipped
aspect, leaves present in good condition at flowering.
These criteria immediately identify Spiranthes cernua.
230
Under the first choice above, he separates the

last two: Spiranthes casei from S. ochroleuca.
Spiranthes casei is lip 6-7 mm dorsal sepal and lateral
petals directed forward, scarcely upcurved, lateral
sepals forward, flowers spaced so that groups appear
as florets of 2-3 in loose spiral; lower leaves shriveled,
upper-most persist at flowering. Finally, S. ochroleuca
under Case‟s key has lip 7-10 mm, dorsal sepal and
lateral petals connivent, recurved providing a 2-lipped
appearance, lateral sepals ascending, prominent basal
lip callosities, lowermost leaves present in good
condition at flowering.
We left Brown‟s key at couplet #6. The choice

is flowers with green central portion versus not. The
aforementioned leads to #7, wherein Spiranthes lacera
var. lacera is identified with loose spike, few twists
and those near the spike summit, basal leaves present
at flowering. Spiranthes lacera var. gracilis calls for
dense florets on the spike, regular twisting on the
spike, basal leaves withered at flowering.
Without green central lip pushes us to #8

wherein flowers purest white is Spiranthes tuberosa
versus pale cream, yellowish white or bicolor. This
moves us onward to choice #9. Here flowers
yellowish/white keys to S. ochroleuca. Flowers white
with yellow/beige lip leads to choice #10. This is the
last couplet: flowers slender, never fully expanded,
plants of northern areas identifies as S. casei. Flowers
231
232
233
fully expanded, lip contrasting to petals and sepals

identifies S. vernalis.
Homoya starts his large-floreted species with a

conglomeration of four widely divergent traits. It must
be said that keys should oppose one, maybe two traits
at each selection point, not up into the four or more
range. The more combined choices the more unwieldy
the key.
His first selection requires: single rank spiral of

1/5 or smaller phyllotaxy; sepals inrolled toward the
base; pubescence pointed; plants of dry grassland;
flowering season from mid-June to July. The
combination is said to isolate Spiranthes vernalis. In
opposition: multi-ranked, phyllotaxy of 1/3 or 1/4;
capitate pubescence; sepals flattened at the base;
blooms September or October with a caveat that it is
even earlier in northern wetlands. I hope you agree
that this is not an effective key since this selection
requires further choices to key out any further species.
Therefore, you must next isolate: lip pandurate;

ascending flower parts; petals and sepals joined to
form a hood which together key out to Spiranthes
romanzoffiana. At this point you still need to make
further choices under the opposing traits before you
get another species. Thus you must look for: lateral
sepals widely spreading with tips arching above the
petals; fragrant flowers, leaves fugacious and, he
claims, calcareous habitat in either dry grassland or
234
dunes for S. magnicamporum. This may be correct for

Indiana but definitely not for Ohio. Ours like wet,
disturbed areas as in the Cuyahoga Valley National
Recreation Area amidst a glaciated plateau which was
disturbed during freeway and other construction and is
mostly connected with borrow pits. Novices could be
confused if Homoya‟s key were followed religiously.
Opposing traits in this key bring you to the final

selection which pits the traits of Spiranthes ochroleuca
against those of S. cernua. For the former, seek lateral
sepals appressed and parallel to petals but ascending
with lip curvature obvious; flowers cream to
yellowish; basal tuberosities prominent. Spiranthes
cernua presents whiter flowers, less prominent
tubercles and a wide assortment of habitats. As
previously mentioned, the observation by Whiting &
Catling, and also Case, that given a wet ditch, S.
cernua likes to be down toward the wet, while S.
ochroleuca will be higher and drier, is the most useful
field trait.
There are obvious strengths and weaknesses in

each key. Comparisons are difficult but some I've
shared with you already. Since we are Ohioans, I
should mention in passing that due to limited species
recognized at their times of publication there is no
point in trying to use either Clara G. Weishaupt's
Vascular Plants of Ohio (1971) or E. Lucy Braun's
The Monocotyledoneae (1967). The only species
covered are: Spiranthes vernalis, S. gracilis, S.
235
tuberosa, S. romanzoffiana, S. lucida, S. cernua and S.

ovalis.
CONCLUSION: Happy hunting and may you now

feel that you will no longer have any trouble with the
tresses; come bloom season they will be awaiting you,
for they are all tressed up with no place to go.
THOMAS A. SAMPLINER
236
237
BOOK REVIEWS
WILD ORCHIDS OF THE

NORTHEASTERN UNITED STATES
A Field Guide by
Paul Martin Brown, drawings by Stan Folsom
Comstock Publishing Associates a division of Cornell
University Press, Ithaca and London. 1997. 236 pp.;
48 pp. color plates. $17.95 paper. ISBN 0-8014-8341-
7.
The last 15 to 20 years have witnessed the

opening of the golden age of the regional field guides.
Regional works have much to recommend them: a
limited geographic area typically limits the number of
species included allowing for more in-depth
treatment, and authorship by someone with extensive
experience within that area assures a level of
practicality of information difficult to achieve in a
national guide. Wild Orchids of the Northeastern
United States, a field guide written by Paul Martin
Brown and illustrated by Stan Folsom, incorporates all
the positive features of a regional guide to make a
significant contribution to the botanical literature of
the northeastern United States.
Some of the special strengths of this work are

the completeness of the species listings, the accuracy
of the local flowering seasons, the author's appraisal of
the current status of each orchid as reflected in the
distribution maps, and frequency of occurrence
comments, and the state by state checklists. Even
238
BOOK REVIEWS
experienced orchidologists will find valuable insights

into little known species such as Epipactis atrorubens
and Gymnadenia conopsea. I found the distribution
maps to be very accurate, reflecting both current
species distribution and important historical
knowledge. By combining these maps with frequency
comments and state checklists, the reader can easily
form an impression of the status of any species in a
given state.
Numerous (192) color photographs contribute to

the beauty and depth of this volume, as they depict
both the typical and variant forms of the orchids. Stan
Folsom's line drawings attractively and accurately
depict each species in this book. If I were to make one
suggestion for future editions, it would be that
whenever possible these drawings be life-size, or if not
then the scale of enlargement or reduction be included.
Wild Orchids measures nine by six inches and is

236 pages long, a convenient size to carry afield or to
"plant" on a personal library shelf. Everyone from
wildflower enthusiasts to orchid experts will want to
add this volume to their collection.
William K. Chapman, PO Box 184, New Hartford, NY 13413.

Bill is the author of Wild Orchids of the Northeast reviewed in
this issue, and numerous other natural history books.
239
BOOK REVIEWS
ORCHIDS OF THE NORTHEAST

by William K. Chapman
Syracuse University Press, Syracuse, NY
1997. 195 pp.
$32.95 ISBN 0-8156-2697--5 cloth ; $17.95 ISBN 0-
8156-0342-8 paper
Spring of 1997 appears to have not only brought

the usual array of wild orchids (and other wild flowers,
too), but also a wave o local native orchid field guides
and publications. My current book (Wild Orchids of
the Northeastern United States) and Bill‟s have been
running parallel for two years now and, as we both
predicted, have become available within a few weeks
of each other. In planning review I thought it only
fitting that we should review each others work. For
those who do not know us we have spent many hours
together in the field and it the two field guides were
not planned to appear together - it just happened!
Bill Chapman‟s work present a very different

aspect of a field guide with extensive text relating to
the historical information of the species treated and
even more information regarding floral measurements
and personal observations. Each species is
accompanied by a color photograph inset and keys are
provided to each multi-species genera as well as to the
entire treatment. The key are innovative and work
well in the field. Chapman‟s treatment of both the
Cypripedium parviflorum complex and The
Platanthera huronensis-hyperborea complex are
240
BOOK REVIEWS
extensive and most welcomed. Any more information

that helps local orchid enthusiast sort out these often
confusing groups is always a major plus. General
ranges are given for each species and an extensive
glossary and bibliography. In comparing it to my book
the complementary aspects would include the section
on entymology for each species and the detailed
information on measurements. As both Bill and I have
known all along the two field guide do complement
each other and together present the most complete
information, photographs distribution maps and
drawings available for the orchids of northeastern
United States. (PMB)
241
BOOK REVIEWS
THE ORCHIDS IN THE OTTAWA DISTRICT:

FLORISTICS, PHYTOGEOGRAPHY,
POPULATION STUDIES AND HISTORICAL
REVIEW
by Joyce M. Reddoch and Allan H. Reddoch
Special Issue of The Canadian Field-Naturalist
VOLUME 111, Number 1, 1997. CAN$10.00 & $2.50
shipping.
This 186-page work describes the 44 orchid species that

have been found within 50 km of Canada's National Parliament
Buildings in Ottawa. It contains information on identification,
past abundance, population changes, development cycles and
relative stability of colonies. It is presented as a baseline study
from which to design further research and prepare effective
planning measures to protect wild orchid populations.
The Introduction describes the history of collecting and

recording since 1856, principal orchid habitats, local
distribution patterns, rare species, colour forms, capsules and
seeds, blooming dates and other topics. Each species account
provides detailed information on the above topics, as well as a
brief description of the plant. A drawing and a spot distribution
map accompany each account. Correlations of some species
with the Canadian Shield or the St. Lawrence Lowlands, or with
calcareous rock, sandstone or sand deposits are shown. Long-
lived colonies of many species are described, and population
studies are included for Corallorhiza striata, Goodyera
pubescens, G. tesselata, Platanthera hookeri, P. orbiculata and
Spiranthes cernua.
With 30 years of field observations amassed the

Reddoch‟s have assembled what is perhaps more
information on a local area than any other local orchid
242
BOOK REVIEWS
243
BOOK REVIEWS
flora. In most instance local floras of such a limited

area have information that is basically of local interest.
Not so in the publication!
Without question this is one of the most

meticulously executed local orchid floras to every be
published in North America. In addition to all of the
botanical information and data, it contains a vast
mount of local natural history. The line-drawing
illustrations by Susan Laurie-Bourque are exquisite
and in great detail. Several species also have fruiting
spikes illustrated. This is to be highly recommend for
all North American native orchids enthusiasts.
Because it is a regular issue of a journal the run may
be limited so do not hesitate to order yours
immediately. (PMB)
To obtain copies of this journal issue, send CAN
$10.00 plus $2.50 (postage and handling) for each
copy to:
The Canadian Field-Naturalist

PO Box 35069, Westgate PO
OTTAWA, CANADA K1Z 1A2
244
BOOK REVIEWS
245
BOOK REVIEWS
246
BOOK REVIEWS
HIDDEN ORCHIDS
Photography by Thomas J. Bulat, Text by Thomas J.
Bulat and Marilyn Bulat
Rudi Publishing, San Francisco and Iowa City 112pp.
ISBN 0-945213-20-4 ISBN 0-945312-19-0 paper.
1995. $44.95 hardcover $32.95 paper
The subtitle of this large-format volume is “a

photographic discovery of the disappearing native
orchids of the United States and Canada” To my
knowledge this is the first large-format photograph
album of native orchids to become available. Thirty-
five species, primarily from eastern North America,
are presented, some with several different views. The
color is lavish and the angles often very dramatic.
Often an extreme close-up is accompanied by a line
drawing of actually size. This is most helpful. The
accompanying text frequently includes a poem by Mrs.
Bulat and narrative or field notes by Dr. Bulat. It is
certainly a welcome addition to the North American
orchid literature and affords an opportunity to view
some dramatic photography.
A few of the photos are ambiguously

identified. Plate 2 - purple fringed orchid is large
purple fringed orchid, Platanthera grandiflora; Plate 3
small yellow lady‟s-slipper is Cypripedium
parviflorum var. makasin; plate 4 is eastern fairy-
slipper, Calypso bulbosa var. americana; Plate 29 &
247
BOOK REVIEWS
30 is Great Plains Ladies-tresses, Spiranthes

magnicamporum, not nodding ladies‟-tresses, S.
cernua; plate 49 & 50 is western prairie fringed orchid,
Platanthera praeclara. These amendments in no way
detract from the beauty of the work and in most cases
are clarifications and/or species which have been
described in the past twenty years and do not appear in
all of the literature.
Appendix A - Identification, Location and

Conservation, presents good basic information on its
subjects but contains the one most often perpetuated
errors in so many native orchid articles and books It
states that epiphytic species are confined to southern
Florida. This is not true as several species range
northward to southeastern North Carolina and
westward through Louisiana. Most notable and wide-
ranging is the green-fly orchis, Epidendrum
conopseum. Appendix B is an annotated list of the
photographs with more details on their locales and
notes the sizes of the plants. Although the book was
not widely distributed upon publication we urge
members to secure a copy while they are still
available. (PMB)
248
BOOK REVIEWS
249
BOOK REVIEWS
250

June 1997 North American Native Orchid Journal

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1

NORTH AMERICAN NATIVE

NOTES FROM THE EDITOR

The opinions expressed in the Journal are those of the authors.

As we continue into our third year, this issue

The March issue devoted to the genus

Ron Coleman‟s excellent article in Orchids,

Do not forget our second conference to be held

It is time to prepare for a 1998 calendar. The

I trust you will continue to have a rewarding and

Paul Martin Brown

THE GENUS CYPRIPEDIUM

Cypripediums, popularly called lady's-slippers

Slipper orchids have fascinated horticulturists

Most of the striking members of the genus are

Phillip Cribb is curator of the Orchid Herbarium

Approx. 300 pp, 26 full-page color paintings,

WHY DO PLANTS CHANGE NAMES?

Paul Martin Brown

The change of plant names after years of general

When a specific plant is to be identified, it has

word „species‟ is often used in lieu of „specific

Prior to the work of Linnaeus in the mid-1700s,

This is what we know as plant taxonomy: the

For the next 150 or more years it was not

International Association of Plant Taxonomists in the

The Code is very simple: the earliest validly

Since 1935, to be validly published, a species

In addition, it must be effectively published.

But back to names changing. Prior to 1900, new

Williams, because William‟s publication predates

Upon occasion when a species is deemed to be

Name changes at the generic level have the

familiar species has been changed. An effort has been

In the North American native orchids, the two

Another interesting example is the genus

O. rotundifolia. The plants do not change, only their

By the end of the summer of 1995, my struggle

Tom Todsen, an orchid researcher who lives in

places, and found spotted coralroot, Corallorhiza

Tom had visited this lady‟s-slipper colony many

The easy walking was over, though, because the

Between us and the green wall were arms of the

Off to the side we saw a wooded slope leading

We scanned the seep from the bottom, and had

It took several minutes of wet, muddy climbing,

pushed me, than I pulled him, and eventually we

The orchid we had spotted from below was so

We spent some time photographing the orchids

bottom was littered with a jumble of boulders, and we

The trip to and from the seep was long and

SAPROPHYTIC ORCHIDS OF DALLAS

In the summer of 1981, I was wandering around

In August of 1982, I went back to the same

I found about a dozen plants full of flower buds and

The following year, I started sooner in my

The plants I was originally looking for did not

Hexalectris nitida had been reported only in

In Dallas, Hexalectris nitida seems to prefer

Corallorhiza wisteriana grows in the same

shining crested coralroot

prefers forest dominated by oaks, whereas H. nitida

Another saprophytic species previously