June 1997 North American Native Orchid Journal

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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

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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

138

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.

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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

[email protected]

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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 [email protected]

Brown: WHY DO PLANTS CHANGE NAMES?

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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


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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

Coleman: CYPRIPEDIUM PARVIFLORUM VAR. PUBSCENS IN THE

SOUTHWEST

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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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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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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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Engel: SAPHROPHYTIC ORCHIDS OF DALLAS

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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


158

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.


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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


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shining crested coralroot

Hexalectris nitida


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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


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Texas purple-spike Hexalectris warnockii


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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.


164

crested coralroot

Hexalectris spicata


165

Arizona crested coralroot Hexalectris spicata var. arizonica


166

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


167

urged to consult that reference for full details concerning the

variety.

VICTOR S. ENGEL, 3305 Yellowpine, Austin, Texas 78757

168

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.

169

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

170

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

171

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

172

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

173

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.

174

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

175

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.

Hammer: CYRTOPODIUM IN FLORIDA

176

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)


177

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


178

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,


179

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


180

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


181

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


182

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


183

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


184

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.


185


186

Heshka: ORCHID SEARCH – MANITOBA 1996

187

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


188

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


189

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


190

Amerorchis rotundifolia

small round-lef orchis


191

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


192

(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.


193

Platanthera hookeri

Hooker’s orchis


194

rose pogonia

Pogonia ophioglossoides


195

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

rotundifolia small round-leaved

orchis

June 20

Arethusa bulbosa dragon's-mouth June 29

Calopogon tuberosus

var. tuberosus grass-pink July 1

Calypso bulbosa var.

americana Venus (eastern

fairy) slipper

June 10

Coeloglossum viride

var. virescens long-bracted green

orchid

June 29

Corallorhiza maculata

var. occidentalis western spotted

coralroot

July 7

Corallorhiza striata

var. striata striped coralroot June 20

Corallorhiza trifida early coralroot June 10

Cypripedium acaule stemless (pink)

lady's-slipper

June 10

Cypripedium arietinum ram's-head lady's-

slipper

June 10

Cypripedium candidum small white lady's-

slipper

June 8

Cypripedium

parviflorum var.

makasin

northern small

yellow lady's-

slipper

June 15

Cypripedium

parviflorum var.

pubescens

large yellow lady's-

slipper

June 15

Cypripedium reginae showy lady's-slipper June 29

Cypripedium reginae showy lady's-slipper July 7


196

forma albolabium (white flowered

form)

Goodyera repens lesser rattlesnake

orchis

July 27

Goodyera tesselata checkered

rattlesnake orchis

July 20

Liparis loeselii Loesel’s twayblade July 22

Malaxis unifolia green adder's-

mouth

July 7

Platanthera dilatata

var. dilatata tall white northern

bog orchis

July 7

Platanthera hookeri Hooker's orchis June 10

Platanthera hyperborea

var. hyperborea tall leafy green

orchid

June 20

Platanthera lacera ragged fringed

orchis

July 28

Platanthera obtusata

forma foliosa blunt-leaf orchis June 20

Platanthera orbiculata large round-leaf

orchis

July 11

Platanthera praeclara western prairie

fringed orchis

July 6

Platanthera psycodes small purple fringed

orchis

July 27

Pogonia

ophioglossoides

var.

ophioglossoides

rose pogonia July 14

Spiranthes lacera var.

lacera northern slender

ladies'-tresses

July 28

Spiranthes

magnicamporum Great Plains ladies'-

tresses

September 7

Spiranthes

romanzoffiana hooded ladies'-

tresses

July 31


197

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.

Sokolski et al.: Cypripedium reginae

198

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


199

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


200

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.


201

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


202


203


204

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


205

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


206

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.


207


208


209

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


210


211


212

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


213

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


214

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.

Sampline.: All Tressed Up, But No Place to Go

215

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


216

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.


217

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.


218

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.


219

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.


220


221

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


222

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,


223

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,


224

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,


225

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


226


227


228

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


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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


230

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.


231

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


232


233


234

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


235

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.


236

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


237

BOOK REVIEWS

238

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

BOOK REVIEWS

239

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.

BOOK REVIEWS

240

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

BOOK REVIEWS

241

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)

BOOK REVIEWS

242

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

BOOK REVIEWS

243

BOOK REVIEWS

244

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

BOOK REVIEWS

245

BOOK REVIEWS

246

BOOK REVIEWS

247

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 &

BOOK REVIEWS

248

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)

BOOK REVIEWS

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BOOK REVIEWS

250

Documents

June 1997 North American Native Orchid Journal