June 2007 North American Native Orchid Journal

NORTH AMERICAN
NATIVE ORCHID JOURNAL

Volume 13 (2) 2007
IN THIS ISSUE:
A Synopsis of
TWO NEW PLATANTHERA SPECIES FROM THE WESTERN
UNITED STATES
DROUGHT, PERIL, AND SURVIVAL IN THE GREAT PLAINS:
CYPRIPEDIUM CANDIDUM
GOOD THINGS COME IN SMALL PACKAGES or
TINY JEWELS IN THE WILD
and more………….
The North American Native Orchid Journal (ISSN 1084-7332) is a publication
devoted to promoting interest and knowledge of the native orchids of North
America. A limited number of the print version of each issue of the Journal are
available upon request and electronic versions are available to all interested
persons or institutions free of charge. The Journal welcomes article of any
nature that deal with native or introduced orchids that are found growing wild
in North America, primarily north of Mexico, although articles of general
interest concerning Mexican species will always be welcome.
NORTH AMERICAN
NATIVE ORCHID JOURNAL
Volume 13 (2) 2007
CONTENTS
NOTES FROM THE EDITOR
59
Looking Forward
October 2007
60
A Synopsis of
TWO NEW PLATANTHERA SPECIES FROM THE
WESTERN UNITED STATES
Paul Martin Brown
61
DROUGHT, PERIL, AND SURVIVAL
IN THE GREAT PLAINS:
Margaret M. From
66
The Slow Empiricist
75
MY FAVORITE THINGS
A gallery of orchid photos
79
THE EFFECTS OF VARIOUS MEDIA AND ADDITIVES
ON THE GERMINATION AND DEVELOPMENT OF
SEVERAL NORTH AMERICAN NATIVE ORCHIDS
David Niemann
85
Note of Interest:
Epipactis palustris - Another European Visitor
New to the North American Orchid Flora
112
TEXAS LADIES’ TRESSES (SPIRANTHES BREVILABRIS)
REDISCOVERED IN TEXAS
Eric Keith
113
Book Reviews
115
Wild Orchids of the Northeast: New England, New York, Pennsylvania, and New Jersey
Orchids in Hawaii
Vanishing Beauty: Native Costa Rican Orchids 1.
Orchid Arabia
Orchids of Europe, North Africa and the Middle East
Orchidelarium
In Memoriam – T. Sanders McMillan IV (1974-2007)
Jim Fowler
123
ON THE REPORTS OF SPIRANTHES VERNALIS
Engelmann & Gray FROM NEW MEXICO
William F. Jennings & Charles J. Sheviak
124
Unless otherwise credited, all drawings in this issue are by Stan Folsom.
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 13(2) pages 59-126; issued July 5, 2007.
Copyright 2007 by the North American Native Orchid Journal
Cover: Platanthera macrophylla by Stan Folsom
NOTES FROM THE EDITOR
As mentioned in the last issue what had been originally planned for a
single March issue has become two issues. In addition to the articles that were
extracted from the original March issue I have decided to continue the photo
gallery in My Favorite Things as well as several more book reviews. In this issue
David Niemann‘s lengthy article on germination and media of several natives
will be of interest to the specialist and the remaining articles should have an
appeal to a broader audience. Stan Folsom and I will be making a major trip to
the Southwest this summer to photograph a few select species for our latest
book in preparation: Wild Orchids of the Southwestern United States, but should be
able to access email every few days.
I am soliciting additional articles for both the March and October 2008
issues at this time. Please submit both ideas and finished work. I can work with
you on your ideas if you are not quite sure how to develop them.
Paul Martin Brown
Editor
naorchid@aol.com
LOOKING AHEAD……
OCTOBER 2007
THE NATIVE ORCHIDS OF

COLORADO
by
SCOTT F. SMITH
Book Reviews
& Notices
Brown et al.: Synopsis of TWO NEW PLATANTHERA SPECIES
A Synopsis of
TWO NEW PLATANTHERA SPECIES FROM THE
WESTERN UNITED STATES
In a continuing saga of the green-flowered species of Platanthera in the

western United States two new species have recently been published. Platanthera
tescamnis Sheviak & Jennings, the Intermountain rein orchid, in Rhodora 108: 19-
31, 2006 and Platanthera yosemitensis Colwell, Sheviak, & Moore, the Yosemite
bog orchid, in Madrono 54(1): 86-93, 2007. Both of these are segregates from P.
sparsiflora/stricta/hyperborea complex, a group that has previously yielded P.
purpurascens (Rydberg) Sheviak & Jennings (North American Native Orchid Journal
3(4): 444-49. 1997) and P. zothecina (Higgins & Welsh) Kartesz & Gandhi (Great
Basin Naturalist 46: 259. 1986). Due to publication limitations color
photographs and/or distribution maps of both new species were not available
in the original publications. The species have been synopsized here in field-
guide style pages and much of the material has been synthesized from the
original descriptions. The reader is referred to those original publications of all
four species for full details regarding the technical aspects of the plants and
specimen citations.
I am very grateful to Alison Colwell, Peggy Moore, Chuck Sheviak, Bill

Jennings, and Scott Smith for their help in gathering information to put these
pages together.
PMB
Note: Madrono 54(1) Jan.-Mar. 2007 was published on July 3, 2007.
61
Platanthera tescamnis Sheviak & Jennings
Intermountain rein orchid

A new Platanthera (Orchidaceae) from the Intermountain West. Rhodora 108: 19-31. 2006
Range: primarily the Great Basin and Colorado Plateau in western Colorado, Utah, Nevada,
and rare in bordering areas in eastern California and northeastern Arizona
Plants: 29-126 cm tall; leaves usually 4-6 near the base of the stem, lanceolate to 29 cm long,
ascending to spreading, inflorescence 1/3 – ½ the heighth of the plant
Flowers: green to greenish yellow; ca. 1 cm long, the lip greenish yellow, linear-oblong to
oblong, varying to lanceolate at the western margin of the range, to 7 mm long, the spur
generally pendant, slightly clavate or cylindrical to 7 mm long; proportion of spur to lip is
4/5 – 1.2/5; size of the column—small and narrow—and position of the pollinia—close to
the column—is distinctly different from other western species of green-flowered Platanthera.
Habitat: along waterways in the drier areas; the name meaning desert (tesca) and swift-
flowing river (amnis). As with most of the western species of Platanthera habitat can be
variable but this species is distinctive for its preference for essentially mesic to dry areas near
seasonal wetlands at elevations varying from 1825-2950 m
Flowering time: late June to early August
Platanthera tescamnis, the Intermountain rein orchid, has been long

known in the region and usually identified as P. sparsiflora. Although P. sparsiflora
occurs elsewhere in California, Arizona, Utah, and Nevada, with the description
of P. tescamnis, P. sparsiflora it is not known from Colorado. Plants of the
‗hanging gardens‘ from the four corners area are P. zothecina.
Platanthera tescamnis preference for drier areas and at lower elevations than
other related species often aids in identification. Only in the eastern limit of the
range in Colorado does it occur with P. aquilonis.
documented sites
reports
62
Platanthera tescamnis, Colorado

photos by Scott Smith 63
Platanthera yosemitensis Colwell, Sheviak, & Moore
Yosemite bog orchid

A new Platanthera (Orchidaceae) from Yosemite National Park, California. Madrono 54 (1):
86-93. 2007
Range: Mariposa County, California; currently known from few sites a very restricted area in
Yosemite National Park
Plants: 20-80 cm tall; leaves 5-7 on the lower 2/5 of the stem; lanceolate, strongly
ascending and reduced to bracts
Flowers: 30-50, loosely flowered with floral bracts about the same length as the ovary;
sepals green to yellowish-green; petals and lip yellow; lip rhombic-lanceolate somewhat
dilated at the base with the tip usually caught in the dorsal sepal but if free the lip horizontal,
not descending; flowers and fruit diminishing in size upwards and the withered flowers
remaining on the ripening capsules
Habitat: wet, montane meadows and seeps usually in tall(er) grasses and sedges; often found
along turfs and streamlets at an elevation of 2100–2285 meters
Flowering time: July/August
Unquestionably one of the most restricted species of Platanthera in this

section to have been found. Plants were originally collected as Habenaria
hyperborea by G.H. Grinnell in 1923 and then rediscovered by Coleman and
Glicenstein in 1993 and a fragmentary specimen was identified as P.
purpurascens. This placed the latter species well west of all known sites for P.
purpurascens. Subsequent investigation has shown that the Yosemite plants vary
in several critical aspects from other species of Platanthera that are found in the
region. Habitat (slender, open inflorescence) color (yellow corolla) fragrance
(musky) and pollination mechanics all are distinctive in this species. In addition,
the restricted range and specific habitat within that range raises questions as to
whether additional sites will be found.
64
Platanthera yosemitensis, Mariposa Co., California

photos by Peggy Moore
65
From: DROUGHT, PERIL, AND SURVIVAL IN THE GREAT PLAINS:
DROUGHT, PERIL, AND SURVIVAL

IN THE GREAT PLAINS:
Margaret M. From
The small white lady‘s slipper, Cypripedium candidum, is a rare sight

in Nebraska. The shy, diminutive orchid is found in only seven locations
scattered widely across the state, and although some populations may
have a hundred or more blooming individuals, species protection is
critical for its long-term survival. The specific name candidum comes
from the Latin word candere, which means ―to shine or to glow‖. In the
sixteenth century lady slipper orchids were called ―Slipper of Our Lady‖,
which was meant to pay homage to the mother of Christ (Simon, 1975).
The conversion of tall-grass prairies to productive cropland over the last
one hundred years left little more than a few native prairie remnants
intact, so that now the encounter of even one orchid in bloom is a gem
not soon forgotten.
Populations in the state represent some of the most western
locations within the range of Cypripedium candidum, making the
genotype(s) important conservation subjects. Orchids face a precarious
existence in the harsh climate of the Great Plains, where frequent
droughts, bitterly cold, dry, winter winds and blazing summer
temperatures test a species‘ mettle. Drought hit the plains particularly
hard in recent years, and much of Nebraska has now experienced as
much as 8 years of precipitation shortfall. That the orchids survive at all,
under such stress, is simply amazing. Herbivores like to browse the
orchids, often removing the tops before they can produce mature seeds
to ensure a succeeding generation. Human safety takes precedence in
road maintenance, sometimes permanently altering the orchids‘ habitat,
as happened for C. candidum in Nebraska in 2005. When orchids are
66
displaced by human activities, rescue operations are often the only

option left; even if not the ideal. The human aspects are a fact of
modern life, and will continue to put pressure on natural habitats. When
an orchid population is faced with destruction what can orchid
enthusiasts do? Some options are; to notify the appropriate
governmental agencies, discuss with landowners how to protect the
habitat, inquire about a conservation easement for the property,
individuals or organizations can inquire about purchase of the site, one
can move the orchids out of harm‘s way if permission is granted, or
propagate and preserve some germplasm representative of the
population so that the genetic diversity assumed present in the
population does not disappear forever. Ideally, genetic studies should
also be completed before any long-ranging decisions are made, but the
reality is that destruction of habitat often happens with little or no
warning, leaving no time to conduct all the research in advance.
Two of the seven known Cypripedium candidum populations in
Nebraska now face an uncertain future due to land sales and
development, and a third faces environment-altering practices that may
destroy the population. One of those populations is on the edge of a
medium sized city where the orchid‘s habitat is scheduled for
commercial development. All the sites have relatively good soil, which
can easily be converted to crop production.
The species as found in Nebraska, displays a range of variations,
with some individuals possessing yellow sepals and petals with maroon-
brown venations (Fig. 1), some having green sepals with maroon dots
and venations, and some whose petals and sepals are entirely maroon-
brown.
Others have reported the maroon colorations as possible evidence
of the relationship of Cypripedium candidum to Cypripedium parviflorum
(Smith, 1993). An overlap of ranges for C. candidum and C. parviflorum is
believed to result in a natural hybrid known as Cypripedium xandrewsii
Fuller. In Nebraska the nearest population of C. parviflorum is found
within a woodland setting more than one hundred twenty five miles
away from any of the sun loving C. candidum populations, although at
some point in history the two ranges for the species may well have
overlapped. Nebraska‘s C. parviflorum population was just discovered in
67
the early 1990s, approximately 75 years after it was assumed extirpated

from Nebraska, and botanists‘ early accounts of the state do not identify
any overlap of the two species.
The labellum is very similar in each of the variations. It is creamy
white with light markings of magenta pink in lines along the sides and
bright splotches of magenta on the inner rim, as well as on the lower
portion of the column, near the edges. The leaves are reported as
clasping the stem, according to Doherty (1997), but as the growing
season progresses in the Great Plains, the leaves become fully expanded
creating a lovely, open form (Figures 3, 5 and 6).
Figure 1. C. candidum with mostly yellow-green petals and sepals.
Figure 2. Maroon petals and sepals

on C. candidum
68
Figure 3. C. candidum plants in protected habitat at the Omaha‘s Henry Doorly Zoo
In bud, the labellum

is most prominent
(Figure 4), and does
not yet show the
magenta markings
that will be visible
on the opened
flowers later in the
season.
Figure 4. A rescued plant in bud on May 6th, 2006.
As the blooming season comes to a close the flowers senesce and

begin to take on a translucent appearance (Figures 5 and 6), but the
leaves persist and appear to continue photosynthesizing for 3-4 months
longer, producing energy that is translocated to the underground root
69
structures where the flower and leaf meristems develop for the next
growth cycle.
Figure 5. Cypripedium candidum as

the flower senesces.
Rescued plants:
Road work resulted in the displacement and damage to a number
of the plants by the roadside at one of the native habitats. With
permission of the state, some of the rescued plants were placed in a
protected outdoor location and several others were potted up and placed
under greenhouse conditions in order to study the relative merit of
rescue operations if, over time, more plants face eminent threat of
destruction. Rescued plants often die quickly after removal from their
70
habitat (K. Kennedy, personal communication). The orchids were

scraped from the roadside by road grader activity that damaged roots on
most of them, and they were found covered with masses of debris at the
bottom of the ditch (Figure 7). All rescued plants survived and 85%
bloomed the first year, both the individuals planted outdoors as well as
those that were potted up and grown under greenhouse conditions. It
was decided to retain some of the native soil around the roots for those
that were not bare-rooted when found and those specimens produced
somewhat more vigorous growth over the growing season, as compared
to those in the greenhouse.
Figure 6. A rescued plant as the flower begins to fade.
There are risks whenever orchids are removed from their natural
environment, not least of which is that it is difficult to compensate for
local conditions and adaptations, especially if plants are moved any great
71
distance. Those local adaptations are another important reason that

germplasm in the form of plants, seeds or pollen, should be preserved by
long-term means if possible, so that if the need arises and good habitat is
available, preferably in the original area, plants representing the local wild
type could then be used for reintroduction.
Figure 7. C. candidum plant emerging from the roadside debris.
Seed-banking the local wild-type:

Seeds are possibly the most efficient forms of germplasm
preservation because they carry the possibility of genetic diversity and
they are so tiny that great numbers of seeds can be stored within a very
small space. Cryopreservation protocols are being worked out so that the
diversity of Nebraska Cypripedium candidum populations will be
represented and available for future conservation action plans. Until the
time when efforts to keep wild populations safe and healthy are in
practice, and the species‘ propagation and reintroduction methods are
fully outlined, cryopreserved seeds provide materials for future research.
Orchid seed cryopreservation protocols have to be researched on a
species by species basis because each species has its own unique physical
properties and culture requirements. The ability to preserve seeds for
72
years, or even decades, is a powerful tool for conservation, provided that

along with implementation of those protocols we also learn to better
protect remnant habitats, understand the pollinators, and continue to
study the microbial associations that are accepted as critical for the
species. Seeds for C. candidum have already been tested in
cryopreservation and trials are being conducted on their germinability in
post-thaw cultures in the laboratory. The project is preserving seeds
collected from the population facing the most immediate threat of
destruction in the state. Some of the seed testae are damaged in
cryopreservation tests (Figures 8 and 9), but other embryos are able to
withstand the extreme temperatures of liquid nitrogen (-1960C), and still
germinate. Results from the project will determine whether this is a
feasible means to preserve C. candidum seeds for future conservation
actions in the state.
Figure 8. (left) Cryopreserved Platanthera sp. seeds, SEM at 194x.
Figure 9. (right) Scanning electron micrographs (SEM) at 372x, of cryopreserved

tropical orchid seeds.
If practical it would be ideal if each state, province or region could

preserve local genotypes for the long-range future, whether in
cryopreservation or in parent populations grown under protected
conditions. Keeping the populations numbers large is important, since
plant species that maintain large populations stand a better chance of
attracting pollinators and surviving over the long term. In order to
73
maintain healthy populations there need to be increased incentives for

habitat protection by private landowners, and research should be
encouraged about pollination biology, edaphic changes, pollinator
populations, mycorrhizal associations, soil conditions, and plant
communities across the orchid‘s entire range in order to save the
species.. Above all, those who have knowledge about Cypripedium
candidum are encouraged to educate others in the region about the species
itself, and about what can be done to save these beautiful members of
our natural legacy.
Acknowledgements:
My thanks are extended to Jim Pyrzynski and Tim Janssen of the Greater Omaha
Orchid Society for their assistance and perseverance to save the Nebraska
Cypripedium candidum wild-type. Gratitude is also extended to Mike Fritz, biologist for
the Nebraska Natural Heritage Species program at the Nebraska Game & Parks
Commission, and to Dr. Kathryn Kennedy at the Center for Plant Conservation in
St. Louis, for their help and valuable information. Dr. Simmons, director of Omaha‘s
Henry Doorly Zoo, continues to be an inspiration to all of us who know him, and
the motivating force behind conservation efforts for threatened species.
Literature cited:
Doherty, J.W. 1997. The genus Cypripedium; a botanical and horticultural overview.
North American Native Orchid Journal 3(1): 5-120.
Simon, H. 1975. The Private Lives of Orchids. J.B. Lippincott. Philadelphia, Penn.
Smith, W.R. 1993. Orchids of Minnesota. University of Minnesota Press, Minneapolis,
Minn.
Received on 4/23/07: As a postscript to the article about Cypripedium candidum rescued and
planted at the zoo; I am happy to tell you that every one of them has reappeared above ground this
spring. And most of them are displaying more ramets than last year. We are, of course, hoping that
they will persist over the coming years.
Margaret M. From, Plant Conservation Scientist, Omaha‘s Henry Doorly Zoo, 3701 S.
10th St., Omaha, NE 68107 psl@omahazoo.com
Glossary:
cryopreservation: preservation of seeds or tissues at extremely low temperatures
which in effect is supposed to stop metabolism, so that cells don’t age
edaphic: plant communities influenced by the soils, not just the climate (i.e.
effects from sandy soils, acidic soils, or nutrient-poor soils)
germplasm: an organism’s DNA (i.e. seeds are stored DNA in a frozen seedbank)
meristems: clusters of cells at zones that will eventually produce shoots or roots
testae: seedcoat
74
Empiricist: GOOD THINGS COME IN SMALL PACKAGES
above:
Prosthechea
pygmaea
Wally Wilder
left:
Spiranthes eatonii
right
Spiranthes
brevilabris
P.M. Brown
75

The Slow Empiricist
This idea came to me when I heard a fairly new orchid enthusiast talk
about seeing Prosthechea pygmaea. It has only one site that is known for it in
southern Florida. The orchid grows in a large colony on one tree in the
Fakahatchee Swamp. The flower is so very tiny that is hard to see even when
the flower is fully developed. Out of flower most people would not know what
it was. It has been described as being about one quarter of an inch overall. It is
a nice medium shade of yellow but not so brilliant that it catches your eye when
you might be passing by. As the flowers mature and the petals and sepals
disappear as it sets seed and it would be nigh impossible to find the leafy plants
in the tangle of the south Florida wilderness regions except for the fact that it
has colonized this one tree with what appears to be a large single plant.
This experience coupled with the fact that I had been helping Paul
Martin Brown with a project to document and count the extant populations of
Mesadenus lucayanus, which has slender spikes of tiny bronze flowers, got me to
thinking about the other elusive little gems that are out there. Now some of
these tiny wonders might be larger to eye than the aforementioned orchids but
their rarity makes them fit into this topic nicely because if there are only a few
sites for this orchid to be found it should be accorded the same wonder status
as the truly small sized orchids that tantalize the orchid enthusiast to find and
enjoy them.
There are many tiny wonders in the orchid world that grow all over the
United States but Florida seems to have a monopoly on some of them. To
name a few that fit this category you have some of the Spiranthes, especially S.
floridana and S. brevilabris which seem to prefer central Florida for their habitat.
Spiranthes eatonii grows in the Southeastern United States and there are several
sites in Florida. All these Spiranthes share enough characteristics to put them in
the small wonder category. Spiranthes eatonii has tiny little flowers that spiral up a
stem that can be no larger than the lead in a mechanical pencil. Spiranthes
brevilabris and S. floridana are so rare they are easy to miss so even though they
don't officially fulfill the tiny wonder categories they belong in this section, at
least to my way of thinking.
76
Then you have a difficult orchid to even see when you are standing right
next to it. Harrisella is a leafless epiphyte that often likes to grow on old fruit
trees. It is small enough so you can hold the entire plant in the palm of your
hand. The flowers are miniscule but the fruits are at least large enough so you
can see them when you are standing a few feet away. They look like tiny little
bells spraying off the gnarled branches of old orange trees that are one of their
preferred hosts.
Next, we have a family of tiny orchids that are very rare. The smallest in
this group called Triphora are the ones labeled rickettii. They poke up from the
forest floor only about a few inches or so and their tiny blooms are very easy to
overlook. The next in size is craigheadii followed by trianthophora, which is much
larger but has an erratic blooming cycle.
If you live in more northerly climes you have all the twayblades to keep
you searching. These little orchids have flowers that often look like little
dancing men but this entire plant family is usually only a few inches high. Many
of these orchids have a coloring that blends in with their habitat. When you
find a colony, though, you will be richly rewarded by their whimsical
appearance.
Now there are more tiny wonders to find out there so don't be surprised
that I didn't include your favorite or feel miffed because it was left out. Just be
aware that there are a lot more that are just as hard to locate and just as
wonderful when you do find them.
This brings up the first order of business when you set out to locate a
tiny wonder. The best of all possible worlds is to find someone who has already
seen the plants and can lead you right to them. Failing that you would need
accurate directions to be able to have a chance at locating many of the plants.
Lastly, you need a persistent nature and a keen eye for observing these
miniature gems that you would most likely walk right by if you weren't attuned
to looking for them.
Once you gain some familiarity with the plants' habits and their
preferred habitat you might be lucky to encounter a new site for them in your
ordinary scouting for other orchids. All it takes is for you to be sensitive to the
possibilities that the plant may occur somewhere else in similar terrain.
Finding these orchids is probably your first concern but just finding
them, although a great and satisfying experience in itself, should not be your
only goal. You need to learn to look at these orchids and begin to appreciate
the delicate qualities they possess.
This may mean getting down to their level and examining them with a
hand lens. Seeing them magnified shows their beauty much more clearly. You
can see the delicate parts that create the whole and you might even get to see
such tiny parts as their glistening glands or shining hairs. While you are getting
77
down and dirty don't forget to examine the lovely leaves if they are present at
blooming times. Sometimes they are basal rosettes and sometimes they climb
up the stems. Sometimes there is just a single leaf while other orchids have
multiple leaves. Different orchids may have different characteristics even in the
same genus.
This means you have to do some homework if you are going to find these
plants and recognize them for what they are. Reading about them in good
orchid field guides or in journals or other scientific publications can prepare
you enjoy your excursions in your quest of a particular orchid. Then when you
encounter your quarry you will be rewarded with lots of different experiences.
First you should feel exhilaration at having discovered the orchid's hiding
place. Then you should begin to experience the wonder of this tiny creation for
its color, delicacy, shape, scent if it has one, and so forth. These sensory
experiences are what keep orchid enthusiasts out there looking and
experiencing nature's bounty. By using all the aids you need to enjoy the plant
from merely associating with it to really getting up close and personal with it,
you should come away with a deeper understanding of those particular orchids.
I wish you all happiness in your hunting and exquisite experiencing when you
discover them.
Your Slow Empiricist
Harrisella porrecta
Wally Wilder
78
MY FAVORITE THINGS
MY FAVORITE THINGS
A gallery of orchid photos
Calypso bulbosa var. americana

Greg Alikas
June 6, 2004 Golden, Colorado
Digital Konica-Minolta A2
79
MY FAVORITE THINGS
Platanthera leucophaea
Al Menk
June 2006, Michigan,
Digital Nikon D100 with AF28-105 lens
80
MY FAVORITE THINGS
Cypripedium reginae
Tom Nelson
July 8, 2006 Bruce Peninsula, Ont.
Pentax K1000 film
81
MY FAVORITE THINGS
Triphora trianthophora var. trianthophora

Wally Wilder
Marion Co, FL Digital Kodak
82
MY FAVORITE THINGS
Sacoila lanceolata var. lanceolata

Linda Cooper
May 2006 FL Digital Minolta Maxxum 7D SLR
83
MY FAVORITE THINGS
Cypripedium parviflorum var. pubescens lacking anthocyans

Olin Karch
April 28, 2004, Upper Buffalo Wilderness Area, Newton Co., AR Olympus
C2500L digital.
84
Neimann: I. EFFECTS OF VARIOUS MEDIA ON GRMINATION
I. THE EFFECTS OF VARIOUS MEDIA ON THE

GERMINATION AND DEVELOPMENT OF SEVERAL
NORTH AMERICAN NATIVE ORCHIDS
David Niemann
Introduction
Land development and agricultural practices create many side effects, not
the least of which is the destruction of the habitats of many of our rare native
plants. The uncommon species are in especially serious danger because they
often grow only in very special habitats (Niemann, 1986). Once these few
habitats are destroyed, an ecotype or perhaps even the species itself may be lost
forever.
Native North American terrestrial orchids are particularly susceptible to
this type of extinction because they frequently have very narrow ecological
tolerances. There are 78 species of these terrestrial orchids listed in the 8th
edition of Gray’s Manual of Botany for the northeastern United States. Many are
already extinct or extremely rare in parts of their range. With regard to
Cypripedium reginae, Fernald states that ―Plant liable to extinction through raids
by nurserymen and would-be cultivators‖ (Fernald, 1950). Something must be
done if these rare and beautiful plants are to be rescued from extinction.
An intelligent and far-sighted approach to the problem would involve
setting aside the remaining habitats of these and many other rare wildflowers so
that they may continue to grow and reproduce under natural conditions. At the
present time there are too few organizations or individuals willing or able to
undertake this task. Another alternative would involve a thorough study of the
ecological requirements of the mature and seedlings stages of the species most
in danger. With adequate knowledge, suitable habitats could be protected in
areas such as state parks. This is an extremely difficult and time-consuming task
and it has not proved successful in the few cases in which it has been tried. A
third possibility lies in the development of a completely artificial method of
propagation (Niemann, D. 2001). If plants could be propagated in some way
and allowed to reach nearly mature size, they could then be grown in outdoor
habitats on protected land. The test of asymbiotic methods of seed propagation
of these native North American terrestrial orchids is the purpose of this study.
85
Materials and Methods

In speaking of the large number of orchid culture media, Withner (1959)
stated, ―As yet there are no comparative data on all of these media on the same
brood of seedlings…‖. It is unlikely that all of the media recommended in the
literature are suitable for all orchid species. Since the development of a
precisely suited nutrient medium requires a great deal of time, a test of many of
the recommended media could give a more rapid indication of the nutritional
requirements of some of these terrestrial orchids. A test of twenty of the more
common media was set up (See appendix for media and formulation
techniques. (Table 1 shows ion concentrations)
I collected all of the seed used in this experiment. The seeds of Cypripedium
pubescens and C. reginae were collected in my garden in Elmwood Park, Illinois
on September 1, 1968. The capsules were still green at the time of harvest.
Seeds of Calopogon tuberosus were collected in Lake County, Illinois on
September 20, 1968. Plants of Spiranthes cernua were collected at Clark Junction,
Indiana on September 20, 1968 and the plants were grown under artificial light
until October 13, 1968 when the seeds were ripe and harvested. Ripened
capsules of Pogonia ophioglossoides were collected on November 29, 1968 in
McHenry County, Illinois. Soon after being collected, the seeds were removed
from the capsules, air dried, and placed in closed jars containing calcium
chloride. The jars were kept in a refrigerator at 4 degrees C. until the seeds were
planted.
The culture room consisted of a walk-in refrigerator with evaporating coils
mounted in the ceiling. This presented a problem as cold air flowed down over
the culture tables when the refrigeration unit was running. This problem was
solved with a small fan in the growth chamber which maintained enough air
circulation to give a relatively uniform 24 degrees C. +/- 2 degrees C.
The culture tables had plywood surfaces painted with semi-gloss white
enamel. The light source consisted of one General Electric 40 watt daylight
fluorescent per light fixture. Typing paper was placed around the lamp to
reduce the intensity. The fixtures were an industrial type with an enamel
reflector. The fixtures were suspended so that the lamp was 12 inches from the
surface of the table. This arrangement gave relatively uniform light intensity on
each of the six tables. The light intensity varied from 180 to 320 foot candles as
measured by a Weston model 603 light meter at the start of the experiment.
Over most of the surfaces the intensity was 240 foot candles. A preliminary
experiment showed that an intensity of 400 foot candles was not harmful while
an intensity of 600 foot candles was too bright and caused chlorosis of the
seedlings. The intensity of about 250 foot candles was chosen because of the
86
preliminary experiment and previous reports of the necessity of low light

intensities for seedlings (Northen, 1962).
The seeds were sterilized in a 2% Clorox solution (5.25 % sodium
hypochorite) according to Redlinger (1961). The seed was shaken vigorously
for 10 minutes in a 1:50 solution of clorox:water. The seed was then rinsed
twice in sterile distilled water.
The seeds were sown in two ways. The seed of Spiranthes was easy to
suspend in sterile distilled water. The seed of this species was planted with a
dropper pipette. Two drops of seed suspension were placed in each tube. Due
to the number of seeds per unit volume of water this amounted to about 150
seeds per culture tube. The seeds of the other species could not be suspended.
They were planted by removing about 100 seeds from the sterilization vial with
a small spatula. The seeds were then injected into a culture tube with a stream
of sterile distilled water. With some practice, fairly uniform quantities of seed
could be sown in each tube.
Three readings were taken on each tube at 60 day intervals after planting.
The readings were taken using the growth index developed by Curtis (described
by Spoerl, 1948). In previous orchid research results were difficult to interpret
because a uniform method of taking data was not used. Knudson and other
early workers used measurements such as fresh and dry weight of seedlings,
seedling color, protocorm diameter, total leaf and total root length. These
characteristics may be all right for some purposes, but only one reading can be
taken on a group of seedlings. The growth index is probably the best method
of taking data because several readings can be taken on the same group of
seedlings without destroying them. A major advantage is distinguishing degrees
of development. Protocorm diameter is misleading because a large protocorm
is as undeveloped as a small one. The growth index takes into account the
developmental stage of a seedling. There are six stages in my modification of
the index. These may be seen in Figure 1. The index is calculated by counting
the number of seedlings at each stage in a culture tube. From this the percent
of seedlings at each stage is calculated. Then the percent of seedlings at each
stage is multiplied by the stage number. The sum of these products is the
growth index for that tube. The data obtained in this manner are suitable for
statistical analysis (Spoerl, 1948).
87
Table 1.
Chemical Composition of the Media
Medium Nutrient Ions mM/l
NO3 NH4 K PO4 Fe SO4 Ca Mg Cl Mn Sugar Conc.

gm/liter
1 .78 0.0 5.0 1.9 .018 2.5 1.9 3.9 0.0 .022 maltose 20.0
2 4.36 0.0 1.78 .12 0.0 1.48 1.22 1.46 .87 .02 sucrose 20.0
3 12.2 3.79 6.32 1.44 .089 3.0 6.1 1.02 3.34 0.0 glucose 10.0
fructose 10.0
4 12.2 3.79 4.71 3.27 .893 3.72 6.1 1.02 0.0 0.0 sucrose 20.0
5 12.2 0.0 1.84 1.84 .006 1.02 8.26 1.02 2.17 0.0 sucrose 20.0
6 5.2 7.5 7.04 2.48 .137 4.84 1.94 1.02 0.0 .034 sucrose 20.0
7 9.44 5.53 6.15 2.2 .089 .54 0.0 .43 0.0 0.0 glucose 10.0
fructose 10.0
8 12.2 10.6 4.74 2.94 .67 6.52 6.1 1.22 0.0 0.0 glucose 10.0
fructose 10.0
9 11.0 0.0 9.16 .18 .007 .15 1.44 .14 .87 0.0 sucrose 20.0
10 12.2 7.57 1.84 1.84 .112 4.95 6.1 1.02 0.0 .034 sucrose 20.0
11 12.2 7.57 1.84 1.84 .112 4.95 6.1 1.02 0.0 .034 sucrose 20.0
12 4.23 2.75 .88 1.01 .146 1.06 1.48 1.06 0.0 0.0 sucrose 20.0
13 12.2 7.57 1.84 2.10 .267 4.80 4.0 1.02 0.0 0.0 sucrose 20.0
14 7.02 2.75 .88 .88 .016 1.06 2.13 1.06 0.0 0.0 glucose 10.0
15 15.32 3.12 2.62 2.04 0.0 1.02 6.1 1.46 0.0 0.0 glucose 2.0
sucrose 18.0
16 16.7 4.43 6.06 1.27 0.0 0.0 0.0 0.0 .43 0.0 sucrose 35.0
17 sucrose 5.0
18 no
sugar
19 sucrose 5.0
20 no
sugar
All media are in Appendix III of Withner (1959), except Ito (1955), Kano (1968) and Vacin
and Went (1949)
Statistical Design
A 5 x 5 Latin Square was used for the location of the five species involved
on the five tables used in this experiment. Within the species the tubes of the
twenty media were arranged in a completely random manner using the table of
random permutations of twenty in Fisher and Yates (1953). This design
distributes the effect of light variation across the table and temperature
variation with height because all five species are located once in each position
88
across tables and once on each individual table. The ―F‖ test described by
Snedicor and Cochran (1946) was used to determine significance.
Dormancy Test
A preliminary test showed that seed of at lease one of the species being
tested germinated better after a cool period. Seeds of Calopogon tuberosus which
had hardly enlarged in 6 months of culture at warm temperatures resumed
development after a cool period. The preliminary test showed that seeds of
Cypripedium reginae, C. pubescens, and Spiranthes did not react in this way. The
reaction of Pogonia to this treatment was not tested in the preliminary
experiment, but its behavior was similar to Calopogon in some other respects so
it was included in the cool treatment. To further test the effects of a cold
treatment, the tubes of Calopogon tuberosus and Pogonia ophioglossoides were placed
in refrigeration at about 4 degrees C. with no light for 40 days following the
180 degree reading. Though Stoutamire (1964) makes no mention of this
enhancing the germination, he suggests that 4 to 6 weeks of refrigeration are
required to break the dormancy of the corms of Calopogon after they have been
produced by seedlings. The preliminary experiment showed a cool period of 40
days to be very beneficial, and this is why that length of time was chosen. After
removal from refrigeration, the tubes were replaced in their original positions
on the tables and readings were taken after 60 additional days (the 240 day
reading) using the growth index.
Figure 1.
Stage 1 - dead seed or seedling

Stage 2 - ungerminated seed
Stage 3 - embryo breaking through seed coat
Stage 4 - leaf point emerging
Stage 5 - seedling with leaves
Stage 6 - seedling with leaves and one or more roots
89
RESULTS
The results of the test of the nutrient media were surprising in several ways.
The species effect was especially unexpected. Seeds of Cypripedium reginae and C.
pubescens showed no development. Although Curtis (1942) was able to produce
at least protocorms on artificial media, this was not the case though a wide
variety of media were used. At the end of the experiment, all seeds appeared to
still have live embryos, so non-viable seed is probably not the cause.
Approximately 5000 seeds of Pogonia ophioglossoides were planted. One
seedling produced a root and a few rudimentary leaflets, and two others
produced protocorms by the end of the experiment. This is in contrast to
Curtis (1936) and Stoutamire (1964) who described this species as easy to
germinate.
The results for Calopogon tuberosus were not impressive at the 180 day
reading. While Curtis reports obtaining seedlings with leaves 7 cm long after 6
weeks (Curtis, 1936), the longest leaves obtained in this study were about 2 to 3
cm long after the 180 day reading. However, about 50% of the seeds did show
some early stages of germination.
Spiranthes cernua gave somewhat similar results. A large proportion of the
seeds germinated to a small extent. After 180 days, only a few seedlings were
vigorous enough to allow removal from the culture tubes. This species and the
previous one showed a general decline in vigor after 180 days.
The pH of each of the media was checked after the plants were removed
from the culture tubes. In every case, the pH had remained constant, thus
indicating that the media are sufficiently buffered and that the pH was not a
variable factor in the results.
The effect of the time period after planting was significant. In general, one
would expect good development to continue once started. This is generally
what happens in culture of Cattleya and other easily grown genera (Arditti,
1966). Unfortunately, the decline of the vigor of the seedlings in this
experiment is not reflected in the growth index readings. The first reading
consisted, mainly of green protocorms in Spiranthes, although many of the
Calopogon began to develop and already had leaves and roots. The 120 day
reading revealed many chlorotic seedlings while the 180 day reading showed a
still greater decline, especially in the case of Calopogon.
Table 2 shows the analysis of variance for the nutrient medium experiment.
Table 3 shows the treatment means for 5 replications and species and medium
means. Due to the lack of germination of the Cypripedium species, they were not
included in the analysis. Due to the loss of some tubes to contamination, the
experiment had to be analyzed with a non-orthogonal program. The species
effect was significant at the .01 level for all but the 240 day reading. The
90
medium and species x medium effects were significant at the .01 level for all
readings.
Wynd‘s formula gave the best growth index readings. The reading for the
180 day period was 309. Readings increased with time, indicating a continuous
development of the seedlings; however, only one of the seedlings on this
medium developed leaves and a root. Many of the seedlings which developed
without chlorophyll became necrotic at the time of the last reading. It is
impossible to determine exactly when a seedling is dead, so the last reading may
be excessively high due to the inclusion of many seedlings that were on the
brink of death.
The Tsuchiya medium produced the next highest 180 day reading for
Spiranthes cernua with a mean of 279. Again, seedlings increased with each
reading, showing continuous development. In this case, however, not a single
seedling reached stage 6 which can be considered transplanting stage. Many
were very chlorotic and appeared as though they would not live much longer.
Curtis‘ 1949 medium was third best as rated by the growth index of 256.
Again, readings increased for each time period, indicating continued
development, but none of the seedlings reached stage 6. All were small,
undifferentiated protocorms.
Burgeff‘s Eg-1 medium gave a reading of 252 after 180 days. Readings did
not change much in the course of the experiment, but the greatest percentage
of good seedlings were produced, 2.98% of the seeds produced transplanting-
size seedlings (stage 6).
Mariat‘s 1952 medium and Liddell‘s medium both gave final readings of
249. In both cases there were many dead seedlings by the end of the
experiment. The percent of seedlings at stage 6 were .30% and 0.0%
respectively.
Kano‘s medium gave a final reading of 248. Many seedlings enlarged to a
size greater than those on any other medium. Strong roots and root hairs
pushed the seedlings and agar away from the lower side of the culture tubes.
The growth index was much lower than would be expected because of the
death of many of the seedlings which ceased development at an early stage.
Knudson‘s C medium with the addition of 1 mg of niacin per liter of
medium gave a reading of 242 after 180 days. Readings were fairly consistent,
showing not much net development from period to period. Plants reaching
stage 6 was .33%
Knudson‘s B medium and the Curtis 1936 medium gave readings of 235 for
the final period. The readings were fairly consistent, so there was not much
development after the first period. Many seedlings were near death at the end
of the experiment. The percent of seedlings at stage 6 was 1.44% and .43 %
respectively.
91
Table 2
Analysis of Variance
60 Days
__________________________________________________________________
Source d.f. M.S. ―F‖

Blocks 4 67.52 1.27
Species 2 24235.18 455.54 **
Error (a) 8 53.20
Media 19 363.49 15.51 **
Species x Media 38 346.16 14.77 **
Error (b) 184
120 Days
________________________________________________________________
Source df M.S. ―F‖

Blocks 4 204.32 2.37
Species 2 30955.81 359.62 **
Error (a) 8 86.08
Media 19 589.82 4.50 **
Species x Media 38 604.08 4.61 **
Error (b) 184 131.02
180 Days
__________________________________________________________________
Blocks 4 78.67 2.21
Species 2 37973.36 1066.06
Error (a) 8 35.62
Media 19 866.75 12.03 **
Species x Media 38 816.78 11.08 **
Error (b) 184 73.72
240 Days
__________________________________________________________________
Blocks 4 298.85 1.11
Species 1 2554.26 9.51 *
Error (a) 4 268.59
Media 19 1826.59 6.20 **
Species x Media 19 1333.72 4.53 **
Error (b) 160 294.47
* significant at 5% level
** significant at 1% level
92
Table 3
Treatment Means for 5 Replications and Species and Medium Means

60 days 120 days
_______________________________ _______________________________
Species Species
1* 2 3 means 1 2 3 means
1 248.0 209.7 200.0 219.2 267.4 206.1 200.0 224.5
2 250.6 199.3 200.0 216.6 266.4 204.2 200.0 223.5
3 211.1 207.3 200.0 216.1 212.4 210.8 200.0 207.7
4 248.0 205.4 200.0 217.8 255.9 211.0 200.0 222.3
5 255.6 207.8 200.0 221.1 258.9 211.0 200.0 223.6 Me
6 234.8 210.1 200.0 214.9 232.8 213.9 200.0 215.6
di
7 218.8 203.7 200.0 207.5 245.2 204.8 200.0 216.6
um
8 214.0 200.5 200.0 204.8 214.0 202.0 200.0 205.3
9 251.6 202.6 200.0 218.1 L.S.D. 273.8 202.7 200.0 225.5 L.S.D.
10 240.8 206.2 200.0 215.7 3.47 236.6 205.6 200.0 214.1 8.19
11 214.2 202.4 200.0 205.5 211.6 204.0 200.0 205.2
12 213.2 206.2 200.0 206.5 214.8 206.6 200.0 207.1
13 228.6 207.2 200.0 211.9 228.2 205.6 200.0 211.3
14 222.4 213.1 200.0 211.8 225.6 216.8 200.0 214.2
15 238.6 206.0 200.0 214.9 238.0 211.4 200.0 216.5
16 254.4 206.3 200.0 220.2 251.2 204.2 200.0 218.2
17 221.6 200.3 200.0 207.3 223.9 202.2 200.0 208.7
18 224.0 200.8 200.0 208.3 224.0 205.6 200.0 209.9
19 223.6 200.0 200.0 207.9 223.6 200.0 200.0 207.9
20 219.4 200.8 200.0 206.7 219.4 200.0 200.0 206.5
Means231.7 204.8 200.0 236.2 206.5 200.0
L.S.D. 2.04 L.S.D 2.59
*1 = Spiranthes cernua, 2 = Calopogon tuberosus, 3 = Pogonia ophioglossoides
180 Days 240 Days

________________________
Species Species
1 2 3 means 2 3 means
1 309.4 205.9 200.1 238.5 244.0 224.7 234.6

2 279.2 199.3 199.1 225.9 224.2 244.0 234.1
3 214.6 205.1 199.9 206.5 237.2 235.6 236.4
4 251.8 207.8 200.0 219.9 232.4 249.5 241.0
5 250.1 201.0 200.0 217.0 218.0 243.8 230.9
6 228.0 210.3 199.9 212.8 230.8 275.5 253.9
93
7 225.6 204.0 199.9 209.8 269.0 234.7 251.8

8 219.4 201.7 200.0 207.0 204.7 215.4 210.3
9 227.4 201.6 200.0 209.7 208.1 231.0 219.6
10 241.8 203.0 199.9 214.9 L.S.D 244.0 248.3 246.2 L.S.D
11 233.8 203.6 199.9 214.1 6.15 255.4 216.3 235.8 12.29
12 256.4 206.6 200.0 221.0 237.4 236.2 236.8
13 235.4 202.4 199.9 212.6 259.2 231.2 245.2
14 234.6 205.0 199.9 213.4 302.0 236.6 269.3
15 249.2 211.0 200.0 220.1 239.8 218.0 228.9
16 248.1 205.0 202.0 218.4 223.5 226.0 224.8
17 208.4 199.3 200.1 202.6 217.2 204.3 210.8
18 219.4 198.8 200.0 202.6 219.0 205.5 212.0
19 221.4 197.0 200.0 206.1 207.0 200.0 203.5
20 219.0 199.0 199.9 206.0 257.0 199.7 228.4
Means 238.6 203.4 200.1 236.5 228.4
L.S.D. 176 L.S.D. 4.59
Knudson‘s C medium gave a final reading of 234. This medium produced no

seedlings at stage six. A large number of protocorms developed to certain
degree and then stopped developing. This was in contrast to a preliminary
experiment which showed that this medium produced well-developed seedlings
of Spiranthes cernua.
Vacin abd Went‘s 1949 medium gave a final reading of 228. Seedlings which
developed to stage six in this medium was .96%. The index readings varied little
with time.
Cosper‘s medium gave a 180 day reading of 227. The readings fell from a
high value to a low one because of the death of many seedlings toward the end
of the experiment. At first, this medium appeared to be a very promising
medium, but apparently something became a limiting factor near the end of the
experiment. Seedlings with leaves and roots accounted for 2.29% of the
seedlings, although many were chlorotic and appeared to be near death by the
end of the experiment.
The Thomale medium gave a final reading of 226. Development of the
protocorms was poor. Most were colorless and had barely broken through the
seed coat when development stopped. No seedlings with leaves and roots were
produced.
The Vacin and Went tomato juice formulation gave a final reading of 221.
The reading in this case may have been quite inaccurate because of the small
size of the Spiranthes cernua seeds. The seeds blended in with the tomato
fragments in the juice, and were difficult to see. The protocorms were small
and appeared bleached.
94
The Meyer formulation and the Sladden medium modified by Burgeff gave
a final reading of 219. The Meyer tomato juice medium gave results very similar
to those described for the Vacin and Went tomato juice medium. The Burgeff
medium produced one good seedling, but it was unlike those on other media in
that it had very short root hairs and roots. Most of the other protocorms were
very small and were beginning to die by the end of the experiment.
Burgeff‘s N3f medium gave a 180 day reading of 213. One of the reasons
for this low reading is the semi-liquid nature of the medium which made
reading the tubes very difficult. No seedlings developed to stage six. The
protocorms which did develop were very small and were beginning to die.
The germination of Calopogon tuberosus and Pogonia ophioglossoides requires a
cold period. About 10 days after the seeds were planted, the embryos swelled
noticeably and turned greener. Before the embryos broke through the seed
coat, growth ceased in most cases. After 30 days, only a small percentage,
usually less than 1% for Calopogon continued to develop and produce leaves and
roots. This appears to be a case of double dormancy as described by Barton
and Schroeder (1942). In order to overcome this dormancy and get some
readings for these species, the culture tubes were removed from the culture
chamber and given the cold period described earlier.
The results for Calopogon tuberosus were as follows. The 1936 Curtis medium
gave the highest growth index reading of 302 after 240 days. The percentage of
seedlings which reached stage 6 was 17.40%. This was the highest percentage
obtained on any of the media.
The Thomale 1954 medium had the second highest percentage of well
developed seedlings (those at stage six) 11.35%. In this and the previous case,
the growth index was a good indication of which medium produced the most
useful seedlings.
Knudson‘s B medium gave a growth index reading of 260. The percentage
of seedlings at stage 6 was 9.68%.
The Meyer tomato juice medium gave the relatively high growth index
reading of 254 for Calopogon tuberosus. This reading is highly misleading as is
shown by the percentage of seedlings at stage 6, which is 1.18%. As in the case
of the other undefined media, the seeds were difficult to see and only large
embryos were easy to locate, and this would tend to give a higher growth index
reading.
Knudson‘s C medium gave a final reading after 240 days of 253. The
percentage of seedlings at stage six was 10.88%.
Knudson‘s C medium plus 1 mg per liter of niacin gave a final reading of
244 and a percentage of seedlings at stage six of 7.03%. This shows that niacin
was not effective in stimulating the germination of this species.
95
The Liddell medium gave a growth index reading of 240 after 240 days. The
percentage of seedlings at stage six was 11.15%
The 1949 Curtis medium was eighth in rank with a growth index reading of
237. None of the seeds reached stage six.
Burfgeff‘s N3f medium gave a final growth index reading of 233 for
Calopogon tuberosus. This corresponded to 2.82% well developed seedlings at
stage six.
The Vacin and Went medium gave a final reading of 232 as did the Burgeff
Eg-1 medium. This corresponded to a percentage of seedlings at stage six of
4.28% and 5.24% respectively.
Wynd‘s medium was in twelfth place with a growth index reading of 229.
The percentage of seedlings at stage six was .65%. The seedlings of Calopogon
tuberosus on this medium were stunted and most were chlorotic by the end of
the experiment.
Cosper‘s medium gave a growth index reading of 226. Stage six seedlings
represented 6.25% of the total.
The Tsuchiya and the Kano media both gave the final growth index reading
of 224. This corresponds to .74% and 6.25% of the seed which developed to
stage 6. The Calopogon tuberosus seedlings on the Tsuchiya medium were very
chlorotic at the end of the experiment, while those on the Kano medium were
the best looking seedlings produced on any medium. This is an example of how
the growth index values do not necessarily indicate the best seedlings.
The Ito medium gave a final reading of 219, and .45% of the seeds
developed to stage 6. This sugarless medium seems completely inappropriate
for the germination of orchid seed because it does not supply the seeds with a
source of carbon.
Mariat‘s 1952 medium gave a final growth index reading of 218. None of
the seedlings developed to stage six, and all were very chlorotic.
The Chang medium gave a final growth index reading of 217, with 2.72% of
the seedlings reaching stage six. The percentage of well developed seedlings is
undoubtedly high due to the difficulty of seeing ungerminated seeds. The low
growth index reading is due to the poor development of the seedlings. The
protocorms were extremely chlorotic at the end of the experiment.
The Sladden medium modified by Burgeff gave a final growth index reading
of 211, and 1.59% of the seeds developed to stage 6.
The Vacin and Went tomato juice medium gave the poorest results for
Calopogon tuberosus. The growth index reading was 207 with.33% of the seedlings
at stage 6. On this undefined medium, the seedlings became chlorotic, and were
only slightly developed.
The results for Pogonia ophioglossoides are discussed next. The Vacin and
Went 1949 medium gave the highest reading after 240 days. The growth index
96
reading of 276 corresponded to .319 % of the seedlings which reached stage

six. This was the highest percentage for Pogonia ophioglossoides. The reason for
the low number appears to be that the 60 days of growth after the cold
treatment was not sufficient. At a later time, most of the germinated seeds
developed extensive rhizome systems.
The Burgeff Eg-1 medium gave a growth index reading of 250. This
corresponded to.30% of the seedlings at stage six.
Knudson‘s C medium gave the third highest growth index reading of 248,
but none of the seedlings developed leaves and roots.
Tsuchiya‘s medium number 1 had a final growth index reading of 244.
None of the developed seedlings were at stage six, all were at stage three and
were developing well.
Mariat‘s 1952 medium gave a reading slightly under 244, but none of the
seedlings reached stage six.
The Curtis 1949 medium number 5 and the Burgeff N3f both gave final
growth index readings of 236, and no seedlings developed to stage six on either
medium.
The Thomale medium gave a final growth index reading of 235, but gave a
very high percentage of seedlings at stage six.
Both Knudson‘s B and the Cosper medium gave final growth index readings
of 231. In both cases, no seeds on either medium reached stage six.
The Kano medium gave a final growth index reading of 226. This
corresponded to a percentage of well developed seedlings at .176%. Once
again, the growth index value does not indicate the development which
occurred later. After the last reading, seedlings on this medium almost filled the
agar with rhizome systems characteristic of Pogonia ophioglossoides.
Wynd‘s medium gave a growth index reading of 225 and there were no well
developed seedlings. This is in marked contrast to the effects of this medium
on Spiranthes cernua where it gave the highest growth index reading.
Liddell‘s medium gave a final growth index reading of 218, with no
seedlings reaching stage six. Knudson‘s C medium gave a final growth index
reading of 216 with no seedlings at stage six. This is in contrast to the same
medium to which 1 mg per liter of niacin was added. The growth index of the
medium with niacin was much higher than the growth index reading of the
medium without niacin, suggesting that for some orchids, such as Pogonia
ophioglossoides, the addition of 1 mg per liter of niacin may increase the
germination percentage.
The Sladden medium modified by Burgeff gave a reading of 215. No
seedlings reached stage six.
The Ito medium gave a final reading of 205, with no seedlings reaching
stage six. Again, in the case of Pogonia ophioglossoides, we find that the undefined
97
media gave poor results. The seedlings were chlorotic and the seedlings which
did grow developed only slightly.
Chang‘s medium gave a final growth index reading of 204. The seedlings
were chlorotic, and none of the seedlings reached stage six.
The Vacin and Went tomato juice medium gave a final growth index reading
of 200. The same reading applies to Meyer‘s tomato juice medium. No seedling
development could be detected.
Discussion and Conclusions

This discussion involves the results of Spiranthes cernua after 180 days, and
Pogonia ophioglossoides and Calopogon tuberosus after 240 days of growth. The first
observation of importance is that the growth index developed by Curtis is not
always a very practical method of assessing the results of a test involving these
species. The object of this research was to develop a quick method of
germinating the seeds of some of our native North American terrestrial orchids
which could be used by amateurs or professionals to restock natural areas with
native orchid species, and thus preserve the species from extinction. A
germination method could also be used by nurseries to propagate orchids for
sale to the public. The preservation of rare ecotypes of orchid species would
also be possible. Though the growth index is useful for easily grown genera,
such as Cattleya, and does give an indication of the developmental progress of
these species, it is not of much value in determining which medium might
produce the best seedlings, this at stage six. As noted in the results section,
frequently many seeds germinated and broke through the seed coat, but then
stopped developing. A culture tube in which this had taken place would give a
relatively high reading. Another tube in which only a few seeds had germinated
and produced leaves and roots, which is necessary for successful culture out of
the sterile tubes, would give a relatively low reading, even though it contained
useful seedlings while the former tube contained no seedlings of a
transplantable size. A more appropriate method for taking readings on these
seedlings would simply involve a computation of the percent seeds which
produced seedlings of a useful size, those which produce leaves and at least one
root.
An examination of the media on which Spiranthes cernua produced more than
1% seedlings at stage six shows that they all contained sucrose as the carbon
source. This is not true for Calopogon tuberosus and Pogonia ophioglossoides. The
greatest percentage of seedling at stage six for Calopogon tuberosus was on the
1936 Curtis medium which was developed for native orchids. It contained 10
gm of glucose as the carbon source. Pogonia ophioglossoides developed best on the
Thomale 1954 medium which contained 10 gm of glucose and 10 gm of
sucrose. Previous research reviewed by Arditti (1967) shows that for most
98
genera, sucrose seemed to be the best sugar for promoting germination, and
Kano found that sucrose at 2 to 4% concentrations gave the best results on the
genera tested. This appears to be true with Spiranthes cernua, Pogonia
ophioglossoides, and Calopogon tuberosus if size of individual seedlings is used as a
measure because the largest seedlings were produced on the Kano medium.
The results for Spiranthes cernua on Knudson‘s C medium, which contains 20 gm
of sucrose per liter are difficult to interpret. A preliminary experiment showed
that Spiranthes cernua developed quite well on this medium. The explanation may
lie in a physiological change of the seeds during the interval between
experiments. In general the media with a concentration of 20 gm of sucrose per
liter or more showed good germination.
The mineral constituents of the media are quite varied, ranging from
completely defined to completely undefined. In some cases apparently
dissimilar media gave similar results while similar media gave widely varying
results. The media which gave the best results had high levels of nitrate
nitrogen, ranging from 16.37 millimoles per liter for Kano‘s medium to 7.03
millimoles for Curtis‘ 1936 medium (see Table 2). However, other media,
notably Knudson‘s C and Burgeff N3f had equally high nitrate levels of 12.2
millimoles per liter. These media failed to produce any seedlings at stage six for
Spiranthes cernua. Levels of ammonium nitrogen varied from 10.6 millimoles per
liter to no ammonium nitrogen. The effect of the ammonium ion is not clear
because the results are not consistent. The Cosper medium lacked this ion but
produced the highest growth index, and the third highest percentage of
seedlings at stage six for Spiranthes cernua. However the seedlings lacked vigor
and were very chlorotic. This may be due to a lack of ammonium nitrogen, but
according to Raghavan and Torrie (1964), this ion is only beneficial in the early
seedling stages of Cattleya, and once the early stage is passed, it seems to be of
no benefit.
The level of potassium varied from .882 to 9.16 millimoles per liter. The
best development as measured by the percent seedlings reaching stage six, was
at the higher levels of potassium, but again the results were not consistent.
The level of the phosphate ion varied from .112 to 3.27 millimoles per liter.
Again, the best development was associated with high levels of this ion. The
Cosper medium had the lowest level of this ion, and this may have contributed
to the general lack of vigor of the seedlings on this medium. Although it has
been suggested that a high level of the phosphate ion might cause the
precipitation of iron to the point that growth is inhibited (Arditti, 1967) this
appears not to be the case in this experiment.
The level of iron in the media varied from zero to .89 millimoles per liter.
The Kano medium contained no additional iron, but presumably the impure
99
nature of the chemicals which make up this commercial fertilizer contain

sufficient iron because seedlings on this medium showed no sign of chlorosis.
The level of sulphate and calcium ions seems to bear no relation to the
development of the seedlings, however most of the media which produced
more than 1% good Spiranthes cernua. Seedlings contained added magnesium.
The Kano medium was the exception again. Magnesium may have been present
in the impure chemicals in adequate supply. Chlorine and manganese had no
apparent effects.
In summary, the nutritional requirements for a medium to produce vigorous
seedlings of Spiranthes cernua, Calopogon tuberosus, and Pogonia ophioglossoides
probably include sucrose at 20 to 35 gm per liter, nitrate nitrogen at a
concentration of 10 to 16 millimoles per liter, ammonium at 3 to 7.5 millimoles
per liter, phosphate at 1.2 to 3.2 millimoles per liter, iron at .2 to .9 millimoles
per liter, calcium at 1.4 to 3.2 millimoles per liter, and magnesium at 1 to 1.5
millimoles per liter. Presumably a medium with these characteristics should
allow good germination and development of these species, although much
variability is evident in these species, and the exact conditions which trigger the
development of seedlings with leaves and roots not completely clear. The
storage time of the seeds probably had some effect on germination due to a
biological clock which governs the time from capsule dehiscence to
germination.
Acknowledgements
The author wishes to thank Dr. Ervin Denisen for his efforts to secure an NDEA Title
IV teacher training fellowship which allowed the author to perform this research. Dr.
Charles Sherwood provided a review of this paper. Dr. Jowett set up the statistical
experimental designs, and analyzed the data. Dr. L.Y. Quinn supplied some of the seeds
used in these experiments. Thanks also to Chuck Sheviak for thought-provoking discussions
of what might be the causes of poor germination in our native North American terrestrial
orchids.
Literature Cited
Arditti, J. 1966. The effects of niacin, adenine, ribose and niacinamide coenzymes on
germinating orchid seeds and young seedlings. Amer. Orch. Soc. Bul. 35: 892-898.
Arditti, J. 1967. Factors affecting the germination of orchid seeds. Bot. Rev. 33: 1-97.
Barton, L. V. and E. M. Schroder. 1942. Dormancy of seeds of Convallaria majalis L. and
Smilacina racemosa (L.) Desf. Contrib. Boyce Thompson Institute 6: 277-230.
Burgeff, H. 1936. Die Samenkeimung der Orchideen. G. Fischer, Jena.
Curtis, J. T. 1936. The germination of native orchid seeds. Amer. Orch Soc. Bul. 5: 42-47.
Curtis, J. T. 1937. Non specificity of orchid mycorrhizal fungi. Proc. Soc. Exp. Biol. and Med.
36: 43-44.
Curtis, J. T. 1939. The relation of specificity of orchid mycorrhizal fungi to the problem of
symbiosis. Amer. Jour. Bot. 26: 390-399.
100
Curtis, J. T. 1942. Germination and seedling development in five species of Cypripedium L.

Amer. Jour. Bot. 30: 199-205.
Fisher, R. A. and P. Yates. 1953. Statistical Tables. Oliver and Boyd. Edinburgh.
Fernald, M. L. 1950. Gray’s manual of botany. American Book Co. New York.
Galston, A. W. 1961. The life of the green plant. Prentice Hall, Inc. New York.
Israel, H. W. 1963. The production of Dendrobium seedlings by asceptic culture of excised
ovules. Amer. Orch. Soc. Bul. 32: 441-443.
Kano, K. 1968. Acceleration of germination of the so-called ―hard to germinate‖ orchid
seeds. Amer. Orch. Soc. Bul. 37: 690-698.
Knudson, L. 1922. Non symbiotic germination of orchid seeds. Bot. Gaz. 73: 1-25.
Knudson, L. 1924. Further observations on nonsymbiotic germination of orchid seeds. Bot.
Gaz. 77: 212-219.
Knudson, L. 1925. Physiological study of the symbiotic germination of orchid seeds. Bot.
Gaz. 79: 345-379.
Knudson, L. 1930. Flower production by orchid grown non-symbiotically. Bot. Gaz. 89: 192-
199.
Knudson, L. 1941. Germination of seed of Goodyera pubescens. Amer. Orch. Soc. Bul. 9: 199-
201.
Laetsch, W. M. and R. E. Cleland. 1967. Papers on plant growth and development. Little, Brown
and Co, Boston.
Liddell, R. W. 1944. Germinating native orchid seed. Amer. Orch. Soc. Bul.: 12. 344-345.
Miller, C. O. 1956. Similarity of some kinetin and red light effects. Plant Physiol. 31: 318-319.
Niemann, D.A. 2001. Orchid Propagation. Orchids 68: 460-470.
Noggle, C. R. and F. L. Wynd. 1943. Effects of vitamins on germination and growth of
orchids. Bot. Gaz. 104: 455-459.
Northen, R. T. 1962. Home orchid growing., 2nd ed. Van Nostrand, Princeton.
Raghavan, V. and J. G. Torrey. 1964. Inorganic nitrogen nutrition of the seedlings of the
orchid Cattleya. Amer. Jour. Bot. 51: 264-274.
Redlinger, J. R. 1961. Sterilizing agents for orchid seed flasking. Amer. Orch Soc. Bul. 30: 800-
801.
Snedicor, G. W. and W. G. Cochran. 1967. Statistical methods. 6th ed. Iowa State University
Press, Ames.
Spoerl, E. 1948. Amino acids as sources of nitrogen for orchid embryos. Amer. Jour. Bot. 35:
88-95.
Withner, C. L. 1959. The orchids. The Ronald Press. New York.
Stoutamire, W. P. 1964. Seeds and seedlings of native orchids. The Michigan Botanist. 3: 107-
119.
101
Appendix 1.
Composition of the Media
Burgeff EG-1
Dilute solution A and B in 500 ml water, then combine solutions
Solution A
Calcium nitrate Ca(NO3)2.4H20 ...................................... 1.00 gm
Ammonium sulphate (NH 4)2SO4 ................................... .25
Magnesium sulphate MgSO4.7H2O................................... .25
Solution B
Ferrous sulphate FeSO4.7H2O........................................... .02
Potassium phosphate KH2PO4 ........................................ .25
Potassium phosphate K2HPO4 .......................................... .25
Sucrose ............................................................................... 20.00
Agar (Bacto) ....................................................................... 15.00
Burgeff N3f
Part A
Magnesium sulphate MgSO4.7H2O................................... .25 gm
Potassium chloride KCl ........................................................ .25
Ferrous sulphate FeSO4.7H2O ....................................... .02
Calcium nitrate Ca(NO3)2.4H2O .................................... 1.00
Ammonium sulphate (NH4)2SO4 .................................... .25
Part B
Citric acid .............................................................................. .09
Potassium phosphate K2HPO4.3H20 ............................... .25
Dissolve parts A and B in 500 ml water, combine solutions, then add:
Glucose ............................................................................... 10.00
Fructose .............................................................................. 10.00
Agar (Bacto) ....................................................................... 12.00
Adjust pH to 5.0
Chang
Fish emulsion* ..................................................................... 1.5 teaspoons
Agar (Bacto) ......................................................................... 9.0
Sugar (Sucrose was used ) ................................................ 5.5
Peptone (Bacto) ................................................................ 1.0
Water .............................................................................. 1000 ml
* Atlas is recommended, but Ortho was substituted; the analysis is 5-2-2
Cosper
Potassium nitrate KNO3 ................................................ .820 gm
Calcium nitrate Ca(NO3)2 ......................................... .236
Potassium phosphate KH2PO4 .................................. .024
Potassium chloride KCl ............................................... .065
Magnesium sulphate MgSO4.7H2) ............................ .036
Ferric tartrate ..................................................................... .0015
Urea .................................................................................... .050
L asparagines ..................................................................... .050
Sucrose ........................................................................ 20.00
Agar (Bacto) ............................................................. 15.00
Water ............................................................................ 1000ml
pH adjusted to 5.0
102
Curtis 1936
Monopotassium phosphate KH2PO4 ...................... .12 gm
Magnesium sulphate MgSO4.7H2O ........................... .26
Ammonium nitrate NH4NO3 .................................. .22
Calcium nitrate Ca(NO3)2.4H2O ............................... .35
Ferric phosphate FePO4 .............................................. .003
Glucose ............................................................................ 10.00
Agar (Bacto) ................................................................ 14.00
Water .......................................................................... 1000 ml
pH adjusted to 4.7
Curtis 1949
Potassium phosphate KH2PO4 ................................. .112 gm
Magnesium sulphate MgSO4.7H2O ........................ .260
Calcium nitrate Ca(NO3)2.4H2O ............................ .350
Ammonium nitrate NH4NO3 .................................... .220
Ferric phosphate FePO4 ................................................. .027
Yeast extract (Bacto) ........................................................ .10
Agar (Bacto) ............................................................ 15.00
Water ............................................................................ 1000 ml
pH adjusted to 4.8
Ito
Corn starch (Argo) ......................................................... 50.0 gm
Libby‘s tomato juice .....................................................100.0 ml
Add water to make 500 ml
Kano
Hyponex ......................................................................... 3.0 gm
Pepone (Bacto) ............................................................ 2.0
Sucrose .......................................................................... 35.0
Agar (Bacto) ............................................................... 15.0
Water ............................................................................ 1000 ml
pH adjusted to 5.0
Knudson‘s B
Potassium phosphate KH2PO4 ................................. .25 gm
Calcium nitrate Ca(NO3)2 ........................................... 1.00
Ammonium sulphate (NH4)2SO4 .......................... .50
Magnesium sulphate MgSO4.7H2O ....................... .25
Ferric phosphate FePO4 ............................................ .05
Sucrose .......................................................................... 20.00
Agar (Bacto) ............................................................... 17.5
Water ........................................................................... 1000 ml
pH adjusted to 5.0
Mariat 1952
Magnesium sulphate MgSO4.7H2O .......................... .25 gm
Calcium nitrate Ca(NO3)2.4H2O ........................... 1.00
Monopotassium acid phosphate KH2PO4 ............ .25
Calcium chloride CaCl2.2H2O ................................. .12
Ferric chloride FeCl3 ................................................. .001
Agar (Bacto) ............................................................... 16.00
Sucrose ........................................................................ 20.00
Water ........................................................................... 1000 ml
pH 5.0
103
Meyer 1945
Fresh strained tomato juice ..................................... 250 ml
Agar (Bacto) ................................................................ 7.5 gm
Water ......................................................................... 250 ml
pH 4.8
Sladden modified by Burgeff

Calcium nitrate Ca(NO3)2.4H2O ................................ 1.00 gm
Magnesium sulphate MgSO4.7H2O .......................... .30
Monopotassium acid phosphate KH2PO4 ........... .40
Ferric citrate .................................................................... .20
Ammonium sulphate (NH4)2SO4 .......................... .70
Potassium citrate ........................................................... .35
Niacin ............................................................................. .001
Fructose ........................................................................ 10.00
Glucose ......................................................................... 10.00
Agar (Bacto) .............................................................. 12.00
Water .................................................................. 1000 ml
pH 5.0
Thomale 1954
Ammonium sulphate (NH4)2SO4 ....................... .06 gm
Ammonium nitrate NH4NO3 ............................... .37
Potassium nitrate KNO3 ...................................... .40
Potassium phosphate KH2PO4 .......................... .30
Magnesium nitrate Mg(NO3)2.6H2O ............... .11
Ferrous sulphate FeSO4 ...................................... .02
Fructose .................................................................... 10.00
Glucose ..................................................................... 10.00
Agar (Bacto) ............................................................ 16.00
pH 5.0
Tsuchiya 1954
Magnesium sulphate MgSO4.7H2O ...................... .36 gm
Calcium nitrate Ca(NO3)2.4H2O ......................... .20
Potassium nitrate KNO3 ......................................... .08
Sodium sulphate Na2SO4 ........................................ .20
Potassium chloride KCl ............................................ .065
Potassium phosphate KH2PO4 .............................. .0165
Manganese sulphate MnSO4 .................................... .0045
Zinc sulphate ZnSO4.7H2O ................................... .0015
Boric Acid H3BO3 ...................................................... .0015
Potassium iodide KI .................................................. .00075
Glycine ............................................................................. .003
Nicotinic acid ................................................................ .0005
Pyridoxine ..................................................................... .0001
Thiamin ........................................................................ .001
Sucrose ............................................................................ 20.000
Agar (Bacto) ............................................................... 15.00
Water ........................................................................... 1000 ml
pH 5.5
104
Vacin and Went 1949

Tricalcium phosphate Ca3(PO4)2 ................................... .20 gm
Potassium nitrate KNO3..................................................... .525
Monopotassium acid phosphate KH2PO4..................... .25
Magnesium sulphate MgSO4.7H2O ................................ .25
Ammonium sulphate (NH4)2SO4 ................................... .50
Ferric tartrate .......................................................................... .028
Magnesium sulphate MgSO4.7H2O ................................ .0075
Sucrose ................................................................................. 20.00
Agar (Bacto) ....................................................................... 16.00
pH 5.0
Vacin and Went tomato formulation

Fresh tomato juice ..........................................................375 ml
Sucrose .................................................................................. 5.0 gm
Agar (Bacto) ......................................................................... 7.5
Water ..................................................................................125 ml
pH 4.8
Wynd 1933 Type iv R5S1

Potassium sulphate K2SO4 ......................................... .435 gm
Monocalcium acid phosphate CaHPO4 .................... .327
Magnesium nitrate Mg(NO3)2.6H2O ..................... .998
Manganese sulphate MnSO4 ..................................... .000669
Ferric phosphate ............................................................ .000269
Sodium borate Na2B4O7.10H2O ........................... .00143
Maltose ......................................................................... 20.00
Agar (Bacto) ............................................................... 17.5
Water .......................................................................... 1000 ml
pH 4.9
Knudson‘s C
Potassium phosphate KH2PO4 ..................................... .25 gm
Calcium nitrate Ca(NO3)2.4H2O................................... 1.00
Ammonium sulphate (NH4)2SO4 .................................. .50
Magnesium sulphate MgSO4.7H2O ............................... .25
Ferrous sulphate FeSO4.7H2O ....................................... .025
Manganous sulphate MnSO4 ........................................... .0075
Agar (Bacto) ....................................................................... 17.5
Water .............................................................................. 1000 ml
pH adjusted to 5.0
(For Knudson‘s C plus niacin added, 1 mg of niacin was added per liter of the above solution)
Liddell
Calcium nitrate Ca(NO3)2.4H2O ........................... 1.00 gm
Ammonium nitrate NH4NO3 ................................ .25
Magnesium nitrate MgSO4.7H2O .......................... .25
Potassium phosphate KH2PO4 .............................. .20
Potassium phosphate K2HPO4 .............................. .10
Glucose .................................................................. 18.00
Sucrose ................................................................... 2.00
Agar (Bacto) .......................................................... 15.00
Water .................................................................. 1000 ml
pH adjusted to 5.0
105
Niemann: II. EFFECTS OF VARIOUS GROWTH REGULATORS ON THE GERMINATION
II. The Effects of Various Growth Regulators on the

Germination and Development of Several North American
Native Orchids
David Niemann
Introduction
The North American native terrestrial orchids face threats from agriculture
and urbanization. Many habitats have been destroyed resulting in the extinction
of local ecotypes of many of the species. Setting aside areas where these species
can continue to grow would be the best way to preserve the species and
ecotypes, but few individuals or organizations have the financial ability to do
this. As a second choice, these orchids could be grown from seed and kept in
cultivation until suitable areas are set aside where they can be planted and be
allowed to grow Niemann, 2001. Some species of North American orchids do
not germinate well, possibly because of dormancy problems. This paper reports
the results of the use of several growth regulators in an attempt to break
dormancy of these seeds.
Materials and Methods

The seed of Cypripedium reginae were collected in my garden in Elmwood
Park, Illinois on September 1, 1968. The capsules were still green at the time of
harvest. Seeds of Calopogon tuberosus were collected in Lake County, Illinois on
September 20, 1968. Seeds of Cypripedium acaule were donated by Dr. L. Y.
Quinn, of Iowa State University. Seeds were removed from the capsules, air
dried, and placed in a closed jar containing calcium chloride. The jars were kept
in a refrigerator at 4 degrees C until the seeds were planted.
The seeds were sterilized in a 2%Clorox (5.25 % sodium hypochorite)
according to Redlinger (1961). The seeds were shaken vigorously for 10
minutes in a 1:50 solution of clorox:water. The seeds were then rinsed twice in
sterile distilled water.
The seeds these species could not be suspended in water. They were
planted by removing about 100 seeds from the sterilization vial with a small
spatula. The seeds were then injected into a culture tube with a stream of sterile
distilled water. With some practice, fairly uniform quantities of seed could be
sown in each tube.
106
The tubes were placed in a growth chamber with approximately 200 foot-
candles of fluorescent light, and a temperature of approximately 24 degrees C.
Three readings were taken on each tube at 60 day intervals after planting.
The readings were taken using the growth index developed by Curtis (described
by Spoerl, 1948). In previous orchid research results were difficult to interpret
because a uniform method of taking data was not used. Knudson and other
early workers used measurements such as fresh and dry weight of seedlings,
seedling color, protocorm diameter, total leaf and total root length. These
characteristics may be all right for some purposes, but only one reading can be
taken on a group of seedlings. The growth index is probably the best method
of taking data because several readings can be taken on the same group of
seedlings without destroying them. A major advantage is distinguishing degrees
of development. Protocorm diameter is misleading because a large protocorm
is as undeveloped as a small one. The growth index takes into account the
developmental stage of a seedling. There are six stages in my modification of
the index. These may be seen in Figure 1. The index is calculated by counting
the number of seedlings at each stage in a culture tube. From this the percent
of seedlings at each stage is calculated. Then the percent of seedlings at each
stage is multiplied by the stage number. The sum of these products is the
growth index for that tube. The data obtained in this manner are suitable for
statistical analysis (Spoerl, 1948).
The basal medium used in this experiment was White‘s standard with 15 gm
agar and 20 gm of sucrose per liter of medium (see appendix). The growth
regulating substances were prepared by dissolving the proper amount of the
chemical in 95% ethyl alcohol to make a one thousandth molar solution. This
was then added to the nutrient media at a temperature of 45 degrees C. to give
a final concentration of one millionth molar.
Statistical Design
The experiment was set up in a factorial design. There were two blocks of
two species. The treatments were gibberellic acid (GA-3); kinetin; 2,4-D; and
indole-3-acetic acid. They were present alone and in all possible combinations.
This gave a total of 16 treatment combinations. The readings were taken using
the growth index.
Results
The seeds of Cypripedium reginae and C. acaule did not germinate on any
of the media. Calopogon tuberosus germinated on most media, and gave a general
idea of what the effects of these growth regulators are. Statistical analysis was
not possible because of the small number of seeds which actually germinated.
Most of the culture tubes produced only 4 or 5 seedlings, so interpretation
must be tentative until a means to produce more seedlings is developed.
107
The check culture tubes, those with no growth regulators added, produced
seedlings that were abnormal. One tube contained a mass of callus about 1 cm
in diameter. This type of development was not noted in the culture tubes of
any previous experiments. Several small protocorms were produced, but none
developed leaves or roots.
Gibberellic acid alone produced very large protocorms, up to 3 mm in
diameter. Some seedlings produced leaves. In at least one case, a seedling went
dormant as described by Stoutamire (1964). It then resumed growth without
the requirement of a cool period. This had not been observed in any previous
experiments described in Niemann (In press).
The kinetin medium produced one plant with a root, but rhizomes were
very short. The leaves were about one third normal size, although some large
corms up to 3 mm in diameter developed. Kinetin with gibberellic acid also
gave some evidence of a bypass of the dormant period requirement, but the
effect was not as clear as that mentioned for gibberellic acid alone. Some large
corms 3 mm in diameter developed.
The medium with 2,4-D alone caused very short rhizoids less than 1 mm
long developed on Calopogon tuberosus. There was an apparent disorganization
of the shoot apical meristem. The seedlings were not normal, and some
consisted of simple linear tissue masses.
Plants on the medium including 2.4-D and gibberellic acid developed very
little, only to the protocorm stage, and then most of the protocorms died.
The medium with 2,4-D caused the production of one seedling with three
stolon-like growths not characteristic of Calopogon tuberosus. The rest of the
seedlings showed a disruption of the shoot apical meristem. Many of the
seedlings died
The medium with indole acetic acid alone caused poor root growth. The
leaves were about twice the normal length, and many of the seedlings had died
by the end of the experiment.
The medium with indole acetic acid plus gibberellic acid caused some of the
corms to be very large, up to 4 mm in diameter. There was again some
evidence that the dormant period was being overcome, but not so strongly as
on the medium with gibberellic acid alone. Leaf and root growth were normal.
The medium with indole acetic acid plus kinetin prevented roots from
forming. The leaves were very short, only about 20% of the normal length. The
leaves formed a rosette over the plant.
The medium containing indole acetic acid plus gibberellic acid plus kinetin
produced leaves of normal size, but large corms up to 3 mm in diameter.
The medium containing indole acetic acid plus 2.4-D caused very poor
development. Many of the protocorms died and several linear, undifferentiated
seedlings developed.
108
The medium containing indole acetic acid plus 2,4-D plus gibberellic acid
caused many seedlings to die and others became linear seedlings. Development
was not as good as that of the above treatment.
The medium with indole acetic acid plus 2,4-D plus kinetin caused the
development of many protocorms, but they failed to grow. Many died by the
end of the experiment.
The medium with indole acetic acid, gibberellic acid, kinetin and 2,4-D
caused the development of many protocorms, but most had died by the end of
the experiment.
Other Tests
A few other tests were performed and they will be reported briefly. One
experiment involved soaking seed of Calopogon tuberosus for 45 days in sterile
distilled water. Liddell (1944) has suggested that soaking seed of this species for
about a month in nutrient solution enhanced germination. Soaking seed in
sterile distilled water should remove any water-soluble germination inhibitors
which may be present. In this experiment, germination was poorer when the
seed was soaked in water. It appears that Calopogon tuberosus does not possess a
water-soluble germination inhibitor.
In another test, seeds of Cypripedium acaule were sown on media with and
without a saponin extracted from Cypripedium acaule by Dr. L. Y. Quinn of Iowa
State University. The substance was believed to have some stimulatory effect
on the germination of orchid seeds. After 180 days in culture, no development
of protocorms was noted.
In another experiment, indole –3-acetic acid, gibberelic acid and kinetin
were used individually at concentrations of one thousandth, one ten
thousandth, one one-hundred thousandth, and one millionth molar. Seeds of
Spiranthes cernua, and Cypripedium acaule were planted. The experiment was
concluded at the first reading because none of the treatments appeared to give
any beneficial effect on inducing seed germination.
Discussion and Conclusions

Test of Growth Regulators
General trends were observed which seem to follow from previous work on
other species. The discussion will be limited to the main effects because the
interactions appeared to be a simple blend of these effects. Statistical analysis
was not performed because of the small number of seeds which germinated,
Gibberellic Acid
The apparent bypass of the need for a dormant period is in agreement with
the observations in other plants. Sachs (Laetsch and Cleland, 1967) states that
gibberellic acid can substitute for a cold period to induce bolting of the rosette
plant Hyoscyamus niger var. bienne, and states that other workers have made
109
similar discoveries for other plants. Thus it appears that of the growth
regulators tested, gibberellic acid is the only beneficial one in that it allowed
more or less continuous development of Calopogon tuberosus without the
interruption of a cold period. It was hoped that gibberellic acid would also
stimulate ungerminated seeds to germinate. Kahn (1957) found that gibberellic
acid was able to break dormancy of lettuce seeds whose germination was
prevented by temperature inhibition, dark-osmotic inhibition, or far red light
inhibition. Apparently gibberellic acid at the concentrations used in this
experiment is not capable of substituting for the cold period which is required
to bring about the germination of Calopogon tuberosus. Since gibberellic acid gave
no indication of stimulating the germination of Cypripedium seeds, its buildup in
the seeds is probably not required for germination as it is in some seeds, such
as Avena (Galston, 1961).
Kinetin
The effect of kinetin for the most part was inhibitory. Other work has
shown that its effect on tropical orchids is also inhibitory. Miller (1956)
observed that its stimulation of lettuce seed germination in the light and
darkness, however in this experiment, stimulation of orchid seed germination
did not occur.
2,4-D
The auxin 2,4-D was completely non-beneficial in this study. It apparently
induces cell division in Calopogon tuberosus in the concentration used, but cell
division is erratic and abnormal. Differentiation did not take place. 2,4-D also
seems to have a detrimental effect on the metabolism of this monocot species,
indicating that 2,4-D may be a factor in the local extinction of this species and
perhaps other orchids.
Indole acetic acid
This chemical apparently has very little effect on Calopogon tuberosus. In
combination with gibberellic acid it may cause slightly larger seedlings to
develop. It has been noted that indole acetic acid causes an increase in the fresh
weight of the seedlings of some tropical orchids. If enough seeds had
germinated in this experiment an increase in fresh weight might also have
occurred. The general effect of indole acetic acid in the increase in cell wall
plasticity and elasticity in preparation of cell walls before division did not occur
in this experiment.
In conclusion, the only additive used in this experiment which was
beneficial at the one hundred thousandth moles per liter was gibberellic acid. It
helps override the requirement for a cool dormant period.
110
Acknowledgements
The author wishes to thank Dr. Ervin Denisen for his efforts to secure an
NDEA Title IV teacher training fellowship which allowed the author to
perform this research. Dr. Charles Sherwood provided a review of this paper.
Dr. Jowett set up the statistical experimental designs, and analyzed the data. Dr.
L. Y. Quinn supplied some of the seeds used in these experiments. Thanks also
to Chuck Sheviak for thought-provoking discussions of what might be the
causes of poor germination in our native North American terrestrial orchids.
Literature Cited
Galston, A. W. 1961. The life of the green plant. Prentice Hall, Inc. New York.
Kahn, A. J., A. Goss, and D. E.. Smith.1957. Effect of gibberelic acid on germination of
lettuce seeds. Science 125:441-443.
Laetsch, W. M. and R. E. Cleland. 1967. Papers on plant growth and development. Little, Brown
and Co, Boston.
Liddell, R. W. 1944. Germinating native orchid seed. Amer. Orch. Soc. Bul.: 12. 344-345.
Niemann, D. A. 2001. Orchid Propagation. Orchids 68: 460-470.
------. The effects of various media on the germination and development of several North
American native orchids (in press)
Redlinger, J. R. 1961. Sterilizing agents for orchid seed flasking. Amer. Orch Soc. Bul. 30: 800-
801.
Spoerl, E. 1948. Amino acids as sources of nitrogen for orchid embryos. Amer. Jour. Bot. 35:
88-95. Appendix 2.
White‘s standard
(for culturing plant and animal cells)
Calcium nitrate Ca(NO3)2.4H2O ................................................. 12.0 gm
Potassium nitrate KNO3 ................................................................. 3.2
Potassium chloride KCl .................................................................. 2.6
Magnesium sulphate MgSO4.7H2O ............................................ 30.0
Sodium sulphate Na2SO4 ............................................................... 8.0
Sodium phosphate NaH2PO4.H2O ............................................... .76
Manganese sulphate MnSO4.7H2O ................................................ .20
Zinc sulphate ZnSO4.7H2O .............................................................. .12
Boric acid H3BO3................................................................................ .06
Potassium iodide KI ............................................................................ .03
Copper sulphate CuSO4 .................................................................... .0004
Molybdic acid MoO3 (85%)............................................................. .00004
This is added to 4000 ml of water and serves as a stock solution which is further diluted by adding 100 ml of
stock solution to 899 ml water.
The following is dissolved in 100 ml of water and serves as a stock solution: 1 ml is added per liter of final
solution.
Glycine ..................................................................................................... .300 gm
Nicotinic acid........................................................................................... .050
Thiamin hydrochloride .......................................................................... .010
Pyridoxine hydrochloride ...................................................................... .010
Sucrose.................................................................................................. 20.00
Agar (Bacto .......................................................................................... 15.00
Adjust pH to 5.5
David Neimann, 9517 Seeman Road, Union, Illinois 60180

davida@owc.net
111
Epipactis palustris:
Niemann: II. EFFECTS OF VARIOUS GROWTH another European visitorON THE GERMINATION
REGULATORS
Note of Interest:
Epipactis palustris (L.) Crantz another European visitor

new to the North American orchid flora
On June 10, 2006 Mark Larocque
found plants in bud of an Epipactis
which he suspected would be the
European E. palustris growing in a
flooded limestone area near the
Susquehanna River in Lancaster
County, Pennsylvania. When he
checked the site a few weeks later the
plants has finished flowering and were
in fruit. Returning on June 16, 2007
Mark found the plants in bud again.
Due to prior commitments he was to
be unable to return to the site until
July 1. Tom Nelson from New York
City visited the site on June 25 and
found the plants in flower confirming
the identity of the Epipactis. The
population appears to be reproducing
and the plants are growing in a
protected area along with Liparis loeselii and

Spiranthes lucida. This is the first confirmed record
for this species for North America.
In Europe Epipactis palustris occurs rather
frequently in calcareous marshes and is also
popular with native orchid growers. The New
York Flora Atlas lists the plant for New York
based upon a 1990 collection of a specimen taken
from a population in cultivation near Albany that
is spreading in the owner‘s yard, although it has
yet to appear spontaneously in the vicinity.
Weldy, T., R. Mitchell, and R. Ingalls. 2002. New York Flora

Atlas. New York Flora Association, New York State
Museum, Albany, NY. http://nyflora.org/atlas/atlas.htm
Photos by Tom Nelson
112
Keith: TEXAS LADIES’ TRESSES, SPIRANTHES BREVILABRIS, REDISCOVERED IN TEXAS
TEXAS LADIES’ TRESSES, SPIRANTHES BREVILABRIS,

REDISCOVERED IN TEXAS
Eric Keith
On April 11, 2007, while installing vegetation monitoring plots for the
Sam Houston National Forest in Walker County, Texas, I noticed a small
flowered Spiranthes with persistent oval basal leaves. I knew that this
combination of characteristics could be only one of three rare spring flowering
species known from Texas. The densely pubescent rachis and yellowish central
lip distinguished the species as Texas ladies‘-tresses, Spiranthes brevilabris. The
other two species are Florida ladies‘-tresses, Spiranthes floridana, and Eaton‘s
ladies‘-tresses (Spiranthes eatonii) (Brown and Folsom 2003; Sheviak and Brown
2002). On two subsequent visits to the site (April 14 and April 17), I recorded
21 additional plants. The habitat for this population is a remnant black clay
(blackland) prairie approximately five acres in size. Several blackland prairies
are found in Walker County and the Sam Houston National Forest, but no
plants were found on any of the other prairies surveyed. This species was
originally described from southeastern Texas in 1840 by John Lindley based on
a specimen collected by Thomas Drummond (Liggio and Liggio 1999).
Historically, the species has been collected in four other Texas counties with
the most recent collection from Galveston County in 1975 (Liggio and Liggio
1999, Digital Flora of Texas 2003). Until this rediscovery, no surviving
populations were known from Texas, and only two other extant sites are
known for the species, both occurring in Florida (Paul Martin Brown, personal
communication). The habitat for the species has been described as sandy soil
in moist prairies, pine-hardwood forest, and wetland pine savannahs (Liggio
and Liggio, 1999) and dry to moist roadsides and fields (Sheviak and Brown,
2002). Since the species is now known to also occur on blackland prairies, it is
hoped that further surveys in suitable habitat will yield additional populations. I
would like to thank Paul Martin Brown for confirming the identity of the
plants.
Literature Cited:
Brown, P.M. and S.N. Folsom. 2003. The Wild Orchids of North America, North of Mexico.
Gainesville: University Press of Florida.
Digital Flora of Texas. 2003. Herbarium Specimen Browser website.
http://www.csdl.tamu.edu/FLORA/tracy2/main1.html
Liggio, J and A.O. Liggio. 1999. Wild Orchids of Texas. University of Texas Press: Austin.
113
Keith: TEXAS
Niemann: LADIES’ TRESSES
II. EFFECTS , SPIRANTHES
OF VARIOUS BREVILABRIS, ON
GROWTH REGULATORS REDISCOVERED IN TEXAS
THE GERMINATION
Sheviak, C.J. and P.M. Brown 2002. Spiranthes. In: Flora of North America Editorial
Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New
York and Oxford. Vol. 26, pp. 530-545.
Eric Keith, Raven Environmental Services, Inc., P.O. Box 6482, Huntsville, TX 77342,
keith@ravenenviromental.com
Blackland prairie, Walker Co., Texas; with the author‘s sons Lance and Will
Spiranthes brevilabris, Walker Co., Texas April 11, 2007
114
BOOK REVIEWS
BOOK REVIEWS
Orchids of Europe, North Africa and the Middle East
Pierre Delforge
2006
Timber Press
Full color hardcover, 640 pp., 5 x 7.5 in (190 x 125 mm); 1270 color photos plus several line
drawings and watercolors
ISBN-13: 9780881927542; ISBN-10: 0881927546 $39.95
Order from:
http://www.timberpress.com/books/isbn.cfm/0-88192-754-
6/orchids_europe_north_africa_middle_east/delforge
Delforge‘s previous works have always been well received as they should be. He
work is meticulous and although limited to brief entries for each species, as complete as
space permits. The gallery of photographs is mind-boggling. This current book is no
exception. It combines the best of all of his other books and presents a single volume that is
both an update of previous works and much new material as well as needed revisions both
taxonomically and nomenclaturally. Technically a 3rd edition of his original work of the same
name it has more than 200 additional pages than previous French editions.
Introductory chapters on basic orchid information—anatomy, life cycles,
reproduction and orchid identification are detailed and easy to understand with many
excellent graphics. The following chapters of species accounts are broken into four chapters
and arranged by subtribes and the genera arranged systematically. Although this arrangement
is excellent for the botanist it does make it difficult to find a specific genus or species quickly
without consulting the index. Because of the large geographic area covered by the book one
must read through the descriptions very carefully to find those species that might occur
where the reader is exploring. This is not necessarily a negative aspect of the book but does
force the reader into review many more species.
A few synonyms are given with each species entry and fortunately they are in the
index as well. Several taxonomical and nomenclatural points should be mentioned. The
genus Listera is nested into Neottia, Pseudorchis into Gymnadenia; although Coeloglossum is
maintained in its own genus rather than in Dactylorhiza. Hybrids are treated extensively as are
aberrant and unusual forms such as albinos, white-flowered forms and other various color
variants.
The largest genera treated—Epipactis, Dactylorhiza, Orchis, and Ophrys are as complete
as they can possibly be with extensive illustrations to help sort out the oft-confusing species
and forms. Ophrys alone has over 250 pages devoted to the genus!
Pierre Delforge has studied orchids, observed the evolution of their habitats, and
protected them for more than 35 years in Europe, North Africa, and the Middle East and is
an expert on European orchids for the IUCN — the World Conservation Union.
I can recommend this new work with no reservations and at $39.95 it is one of the
best buys in orchid literature available!
PMB
NOTE: This title has been published in Great Britain by A & C Black (ISBN 9780713675252) and
cooperatively with Timber Press in the North America. They have different cover images but the body of the
books is the same.
115
BOOK REVIEWS
ORCHIDS IN HAWAII
Ted Green
$10.95 paper 6" x 9" 130 pp. 175+ color photographs
ISBN: 1-56647-720-4
Mutual Publishing, LLC
1215 Center Street, Suite 210, Honolulu, HI 96816
Hawai‗i‘s love affair with orchids began in the early 1950‘s when a few enterprising
growers made the delicate plants affordable for your everyday, ―garden variety‖ hobbyist.
Today, Hawai‗i is famous for its orchids. With only three of its own native species out of
the more than 35,000 species of orchid world wide, it seems unlikely but true. Orchids in
Hawai‗i offers an introduction to orchids and explains where and when they can be found,
describes species, hybrids, and how to read labels as well as how to take plants to the
mainland. Filled with over 175 stunning photos it is truly the best guide a budding orchid
enthusiast could pick up.
Orchid Arabia
Eric Hansen; illustrated by Barbara Evans
Saudi Aramico World November/December 2006
pp. 10-16
http://www.saudiaramcoworld.com/issue/200606/orchid.arabia.htm
Having nothing to do with North American orchids, but nevertheless a fascinating and
beautifully illustrated article that may be accessed electronically. Hansen is the author of the
popular Orchid Fever.
ORCHIDELIRIUM
Harold Feinstein with an Introduction by Robert H. Hesse
Bulfinch Press
February 2007
$50.00 144 pages 100 full color photographs 13.5 x 11.75‖
ISBN 0-8212-6205-X / 978-0-8212-6205-4
Feinstein‘s‘ assemblage of 100 brilliant, captivating, and exquisite photographs make

Orchidelarium one of the premier orchid ‗coffee table‘ books available on the market today.
The large format and studio photography of primarily single flowers in a variety of views are
multidimensional and jump right off the page. At the conclusion the author presents several
early photographs take out of doors against a brilliant sky. The effect is quite different but
equally as pleasing as the studio shots. Although clearly stated that the book is not intended
to be a botanical account of orchids Bob Hesse‘s Introduction is fascinating reading on its
own. He covers the popular, scientific, and botanical history of orchids and their many uses
as well as the mania that has, and still does, so often surrounds this largest of plant families.
Proportionately no more expensive than a smaller format full color book, the price tag is
well worth it as either (or both) a gift or to grace a personal orchid library.
Ironically the only real error found in the book is the incorrect spelling of
Orchidaceae as Orchiadea on page 12. On page 5 the use of the word Foreword is incorrect
as it is an Author‘s Preface. By definition a Foreword is written by a recognized expert in the
116
BOOK REVIEWS
subject field and usually praises the content of the book and the author. It is never written by
the author.
One of the curious aspects of the book is that several photos that have the flowers
inverted. These are genera/species that normally have the lip in a non-resupinate, or
uppermost, position on the flower, but I will attribute this to the artistic design of the book.
The names of the individual flowers are very useful and synonyms are given for those genera
that have undergone recent revisions--except for the Florida native Prosthechea cochleata that is
listed as Encyclia species. Several photos are simply listed as ‗species‘ that could have easily
been identified to hat rank. But this is the botanist speaking and he needs to be reminded
that the book is a gallery of exception orchid photographs.
Orchidelarium is highly recommended and hopefully it will pique the interest of even
the casual reader to search out and locate local orchid and native plant societies and learn
more about this premier family of plants!
Vanishing Beauties: Native Costa Rican Orchids.

Vol. 1. Acianthera-Kegeliella. F. Pupulin and collaborators (18 collaborators which include
most of the recognized specialists in Neotropical orchids are listed on pp 408-409). 2005.
ISBN 9977-67- 956-8 (Cloth) xxx+421 pp., numerous color photographs, 25×33 cm.
Sistema Editorial y de Difusión Cientifíca de la Investigación, Universidad
de Costa Rica, San Jose, Costa Rica.
The recent publication of yet another large-format spectacular volume on New

World orchids is more than welcome. Given the immense popularity of Costa Rica for
ecotours and orchid exploration both the information and the photographs in this book will
prove to be an excellent resource. Although not all genera are covered, two which stand out
are Beloglottis and Habenaria, those that are treated provide some of the highest quality
photographs of orchid species in the current literature. Bonnie Boutell, a personal friend
who lived in Costa Rica for many years had this to say after examining the book. “There are
no words sufficient to describe the experience of living where wild orchids grow. To watch the
evolution of the ultimate recyler-the rainforest, convert decay into the exquisite beauty of orchids
was life changing. In addition to having them on my own property, I could visit nearby Casa
Orchideas Botanical Gardens. The MacAllister family has devoted their lives to creating this
magical garden filled with the orchids and plants unique to Costa Rica.”
This is the first volume of a series of three and is to be highly recommended and
currently available by inquiry form the Lankester Botanical Garden in Costa Rica
http://www.jardinbotanicolankester.org/ing/books.html
or from OrchidBooks at http://www.orchidsbooks.com/book.asp?id=846.
PMB
117
BOOK REVIEWS
Wild Orchids of the Northeast: New

England, New York, Pennsylvania, and
New Jersey
Paul Martin Brown with original artwork by Stan
Folsom
2007 University Press of Florida.
384 pages 6x9 field guide; 331 full color
photographs, 99 line drawings, 86 maps, keys for
identification; considerable additional informational
material
$29.95 Paper (Flexibind): ISBN: 0-8130-
9780813030340
Do we really need yet another book on the

orchids of the Northeast? As a native orchid
enthusiast, my answer is unequivocally yes. Paul
Martin Brown‘s intense devotion to morphological
detail sets his orchid books apart from most others;
he is almost fanatic in recognizing differences
within a species. While some systematic botanists
may argue that botanical literature is already
overflowing with superfluous plant names, others recognize the value in formally
documenting the wide range of morphological variation within plant species. And in much
the same tradition as M. L. Fernald, Brown does not hesitate to put pen to paper and
describe new taxonomic forms, varieties, and hybrids. In this book, I counted 33
infraspecific orchid taxa from the Northeast originally named by Brown during the past 12
years.
Wild Orchids of the Northeast is the 8th in a series of regional field guides of North
American orchids by Paul Martin Brown and published by University Press of Florida. The
book is designed for anyone interested in orchid identification, taxonomy, distribution, and
conservation, from beginner to expert. It is a complete revision and expansion of Brown‘s
previous work on the subject and now includes all of Pennsylvania and New Jersey, a state
never before covered in its entirety.
The book covers 79 species and varieties, 88 forms, and 15 hybrids. It includes an
illustrated key to genera, workable keys to all species, extensive lists of infraspecific taxa and
full treatment of synonymy, distribution maps, and species accounts extensively illustrated by
high quality photographs and line drawings. The 300+ color photographs are a real strength
of the book.
Another strength of the book is the wide range of supplementary material, including
an extensive section on where and how to find orchids at eight regional hot spots: 1) The
Northwoods: bog, fen, and boreal forest; 2) Connecticut River Valley; 3) Central New York
State; 4) Metropolitan Boston; 5) Cape Cod and Eastern Long Island; 6) The Catskills and
Poconos; 7) The Pinelands of New Jersey and Cape May; 8) The Central Alleghenies. I also
appreciated the well-illustrated section on ―cryptic species, species pairs, and varietal pairs‖.
118
BOOK REVIEWS
The taxonomy and nomenclature followed by Brown closely follows the Flora of
North America treatment of the Orchidaceae published in 2002, with the exception of several
species usually included in Platanthera. Brown splits out P. nivea, P. clavellata, and P. integra and
includes them in Gymnadeniopsis, a transfer originally proposed by Rydberg in 1901 and
favorably noted (but not followed) by FNA author Charles Sheviak. Brown continues to
recognize P. pallida as a valid species endemic to eastern Long Island, New York, whereas
Sheviak placed it in synonomy under P. cristata, but noted its ―distinctive nature‖. Brown
corrected mistakes in previous publications such as the occurrence of P. blephariglottis var.
conspicua in New York.
The author‘s stated purpose of this book is ―to assist the user in identifying the wild
orchids of much of the northeastern United States. It is intended to be used primarily in the
field and is designed for locating information easily while one foot is in the proverbial bog.
The photographs have been taken in the field and are intended to illustrate the species as the
user will see them. They are neither studio shots nor great works of art, just good diagnostic
photos that portray the plants in their habitats.‖ Brown is being too modest in describing his
book and he goes way beyond accomplishing his goals. The Northeast is Brown‘s original
stomping grounds and he knows the territory as well as, if not better than anyone. He is
intimately connected with these orchids and has spent countless hours pursuing and studying
them. Fortunately for us, he enthusiastically shares his vast knowledge and experiences.
I recommend this reasonably priced book for anyone interested in our native
orchids. It has been authored by one of the foremost orchidologists of the region and
contains a wealth of information beautifully illustrated on high quality paper. No library will
be complete without it.
Reviewed by Eric Lamont, Honorary Research Associate, Institute of Systematic Botany, The New York
Botanical Garden, Bronx, NY 10458.
119
BOOK REVIEWS
COMPANION CD TO Please note an update to the following from

what was printed in volume 12: 11,
ORCHIDS OF MEXICO 2006.
LAS ORQUIDEAS DE
MÉXICO
Miguel A. Soto Arenas, Eric Hágsater,
Rolando Jiménez Machorro, Gerardo A.
Salazar, Renato Flores and Ivan González.
Photographs by the authors and 47 additional

photographers. A digital photographic
catalogue with minimal texts in Spanish and
brief comments and instructions in both,
Spanish and English languages.
The number of orchids today recorded from Mexico

has increased in the last years to surpass 1200 taxa
(species and subspecies); in addition, some dozens of
natural hybrids. This CD includes a catalogue with
some 1500 pictures which illustrate 90% of the
Mexican Orchids. It is the complement of the book
Orchids of Mexico (Hágsater et.al. 2005) where only 450
species were illustrated. Some pictures are the first
published for particular taxa. It includes also a checklist
with the orchid names accepted at present and also the
several hundred synonyms used in the last 55 years
(since the publication of Louis O. Williams, The Orchids of Manitoba
Orchidaceae of Mexico in 1951). Synonyms are directly Ames, D., P.B. Acheson, L. Heshka, B.
linked to current names and these to pictures. Voucher
specimens or origin of the plants are given for each Joyce, J. Neufeld, R. Reeves, E.
picture. Reimer, and I. Ward.
2005.
The quality of the photographs, the graphic design, Native Orchid Conservation, Inc. Winnipeg,
user-friendly management, and the cross-references
between accepted names, synonyms and pictures have Manitoba.
been very welcome for those who have used it. 158 pages, 5.5 x 8.5”, full color
photographs, maps; $17.95 CAD Paper
Price US $50.00 plus shipment costs. Orders directly ISBN 0-9734864-0-6.
to: nesponda@chinoin.com.mx; herbamo@prodigy.net.mx The first full-color provincial orchid field
guide to be published. A group effort that has
We are able to accept VISA and MASTERCARD resulted in a workable and usable field guide
credit cards. Charges will be entered in Mexican pesos
and they will appear in your bank statement in your with excellent photographs and a distinct slant
local currency. Please do not forget to include your on conservation. Limited use outside of
credit card number, expiration date, cardholder’s Manitoba; 3 pages of keys for identification.
name and your card code (3 last digits on the back Available through www.nativeorchid.org
of your card). Checks in US Dollars, drawn to
INSTITUTO CHINOIN, A.C. are also accepted.
120
BOOK REVIEWS
ORCHIDS OF EUROPE,
NORTH AFRICA AND THE
MIDDLE EAST
Pierre Delforge
2006
Timber Press
Full color hardcover, 640 pp., 5 x 7.5 in (190 x
125 mm); 1270 color photos plus several line
drawings and watercolors
ISBN-13:9780881927542;
ISBN-10: 0881927546
$39.95
ORCHIDS IN HAWAII
Ted Green
Ted Green
$10.95 paper 6" x 9" 130 pp.
175+ color photographs
ISBN: 1-56647-720-4
Mutual Publishing, LLC
121
BOOK REVIEWS
VANISHING BEAUTIES:
Native Costa Rican Orchids
Vol. 1. Acianthera-Kegeliella. F. Pupulin and
collaborators (18 collaborators which include
most of the recognized specialists in neo-
ORCHIDELIRIUM tropical orchids are listed on pp 408-409).
Harold Feinstein with an Introduction by 2005. ISBN 9977-67- 956-8 (Cloth) xxx+421
Robert H. Hesse pp., numerous color photographs, 25×33 cm.
Bulfinch Press Sistema Editorial y de Difusión Cientifíca de
February 2007 la Investigación, Universidadde Costa Rica,
$50.00 144 pages 100 full color photographs San Jose, Costa Rica. Ordering information
13.5 x 11.75‖ from: librexpo@cariari.ucr.ac.cr
ISBN 0-8212-6205-X / 978-0-8212-6205-4
122
BOOK REVIEWS
In Memoriam – T. Sanders McMillan IV (1974-2007)
Jim Fowler
I met the young man in October of 2005

after he had written Paul Martin Brown in hopes of
identifying a plant that he believed was an orchid.
He had seen Paul‘s book covering Florida‘s orchid
species, and hoped that Paul could help him with
the identification. Knowing that I live in South
Carolina, and that I could possibly visit the site,
Paul gave Sanders my email address and encouraged
him to contact me.
Sanders was a husband, father of a young

son, avid outdoorsman, hunter, and lover of all
things in nature. In addition, he had a job that
consisted of checking out sites to study the possible
environmental impact of proposed development in
the immediate vicinity. What Sanders had seen that
day in October was several plants, each with four or
five lanceolate leaves and a narrow flower spike that
had produced several seed capsules. All flower parts
were long gone.
Both Paul and I were somewhat stumped,

and suggested several possibilities – including (and hoping for) Habenaria quinqueseta. We finally
decided that waiting for next year‘s bloom cycle was the only possibility for an accurate
identification.
In June of 2006, Sanders sent us several photos that allowed us to accurately pin down the
identification to Platanthera lacera – a new species for Richland County, South Carolina. He was quite
pleased with his find, and contacted the curator of the herbarium of the University of South
Carolina in nearby Columbia, South Carolina. This resulted in the site being set aside as a protected
area by the South Carolina Department of Natural Resources – the bureau that is responsible for
protecting native flora and fauna in South Carolina. He was right to be proud of his find.
On April 3, 2007, Sanders took his own life. I guess his reason for this exquisitely sad act is
not important. Paul never had the opportunity to meet Sanders, and I met him only once, but for
some unknown reason, his death has left us both with an unexplained void.
Today, a day after learning of Sanders‘ death, I visited the site once more. It is a small, tree-
filled bottomland that most people wouldn‘t give a second look. But today, I found more than
twenty of Sanders‘ ―mystery plant‖ in full bloom.
Mind you, I‘m not a firm believer in the after life, but maybe, just maybe, one of these days,
I‘ll have another chance to meet this special young man… ―in the stars, my friend."
123
123
Jennings & Sheviak: ON THE REPORTS OF SPIRANTHES
BOOK REVIEWS VERNALIS IN NEW MEXICO
ON THE REPORTS OF SPIRANTHES VERNALIS Engelmann &

Gray FROM NEW MEXICO
William F. Jennings and Charles J. Sheviak
From time to time, there are literature reports that Spiranthes vernalis is or may be in New
Mexico; however, this has been doubted by most authors. Luer (1975) states that "Records of its
occurrence in such areas as Ontario and New Mexico are possibly due to confusion with other species."
The reports of its occurrence in New Mexico seem to have originated with Ames (1905), who
listed Spiranthes vernalis for New Mexico based on a specimen collected by Augustus Fendler, cited by
Ames as: "New Mexico, prairies, August 14, 1847."
Augustus Fendler was a friend of George Engelmann, and apparently it was through the
intervention of Engelmann and Asa Gray that Fendler was allowed to accompany a military detachment
from Fort Leavenworth, Kansas to Santa Fe, New Mexico (Ewan, 1950). Fendler traveled via the Santa
Fe Trail in 1846, following closely behind General Stephen Watts Kearny, who raised the American flag
in Santa Fe on August 18, 1846, completing the bloodless conquest of New Mexico during the Mexican
War. Fendler spent the winter in Santa Fe, collected extensively in the spring and summer of 1847, and
then returned to St. Louis in the fall of 1847, again via the Santa Fe Trail. His collections were
evaluated by Gray (1849) and numerous new species were described. Fendler's collections were some of
the earliest specimens taken in the Southwest, and Santa Fe is the type locality for a large number of
species.
The Santa Fe Trail was used extensively after 1821 to move trade goods by wagon to Santa Fe.
By 1843, about 450,000 pounds of merchandise was carried over the trail in 230 wagons (Brown, 1988).
When the railroad reached Santa Fe in 1880, the trail became a part of Southwest history. In its heyday,
there were actually two trails. Although there were several points of departure in the Kansas City area,
since Fendler was traveling with the military, he started at Fort Leavenworth. The trail cut more or less
diagonally across Kansas to the Arkansas River at Great Bend. West of Fort Dodge, the trail split. The
southern branch (Cimarron Cutoff) continued southwest across the extreme southeast corner of
Colorado and the tip of the Oklahoma panhandle. The northern route (Mountain Branch) followed the
Arkansas River upstream past Bent's Fort to the mouth of Timpas Creek, then angled southwest to
Trinidad, Colorado, and crossed Raton Pass. The trail then followed the base of the mountains,
rejoining the Cimarron Cutoff near today's Watrous, New Mexico. The trail then went through Las
Vegas, San Miguel, and Pecos, New Mexico, almost exactly on the route of today's I-25. There were no
towns along the trail beyond extreme eastern Kansas until Las Vegas, New Mexico was reached, a scant
70 miles from Santa Fe.
Fendler traveled westbound via the northern route (Mountain Branch) of the trail in the period
August 14 to October 11, 1846. Fendler traveled eastbound via the southern route (Cimarron Cutoff) in
the period August 9 to September 24, 1847.
124
Jennings & Sheviak: ON THE REPORTS
BOOK SPIRANTHES VERNALIS IN NEW MEXICO
OF REVIEWS
Based on Ames' report, it would appear that Fendler collected the specimen somewhere
between Las Vegas and Wagon Mound, New Mexico. However, this is not the case. While the
overwhelming majority of Fendler's collections were taken in 1847, some were taken on the trip west in
1846, and this is one of them. The specimen to which Ames referred is at AMES (AMES 71825; GH
1741). It is indeed Spiranthes vernalis, and was annotated so by Ames in 1905, by Correll in 1946, and by
Sheviak in 1990. Fendler's original handwritten col-lection label is with the specimen, and clearly states
"14/8 46, prairie, no. 32." The specimen was taken on August 14, 1846, one year prior to the date cited
by Ames and was taken the first day of the westbound trip, near Fort Leavenworth, Kansas. There is a
printed label on the specimen, provided either by Engelmann or Gray, which reads "Plantae Novo-
Mexicanae, no. 841 [the number handwritten], A. Fendler coll., 14 Aug [the day and month
handwritten], 1847 [printed]. It was common to use a standardized printed label, then to annotate by
hand the pertinent data. It was also common in the nineteenth century for herbarium curators who
distributed large collections of specimens, such as Fendler's, to renumber the specimens in taxonomic
sequence. Thus it was common to see two numbers on old specimens: the collector's number and the
curator's number.
Additional support for the 1846 collection date comes from Shaw (1982). Shaw reported on
Fendler's collection list. Under Orchidaceae, the collection locality of number 841 is given as "low
prairies, 9 miles south of Fort Leavenworth; also about 100 degrees west longitude, not far from the
bed of the Arkansas River; flowers white.‖ Shaw gives the collection period as between August 14 and
September 27, 1846.
Ames (1905) reports that Fendler also collected Spiranthes cernua at: "low prairies, 9 miles south of Fort
Leavenworth, Fendler 119." Fendler likely numbered his collections as he made them (in chronological
order), so Fendler 32 (Spiranthes vernalis) very likely was collected before Fendler 119 (Spiranthes cernua,
according to Ames). It appears most likely that Fend-ler collected the Spiranthes cernua along the
Arkansas River at the end of August or in early September and not near Fort Leavenworth. Likely, it is
the second specimen cited by Shaw.
This position also makes sense from a plant geography and blooming season perspective. Very
few collections of Spiranthes vernalis are from west of the 98th meridian (roughly a line from Oklahoma
City through Wichita and Salina) and the plant is unknown west of the 100th meridian. Based on
Kansas specimens cited by Magrath (1971), Spiranthes vernalis blooms in Kansas from June through mid-
August, with an average date of July 29 for the specimens cited. Latest collection date cited was August
21. Fendler collecting Spiranthes vernalis in northeastern Kansas on August 14 is a reasonable
proposition.
Again based on Magrath (1971), Spiranthes cernua (including Spiranthes magnicamporum, since it was
not described until 1973) is known from Kansas as far west as 99 degrees 30 minutes. Fendler 119
could have been collected along the Arkansas between Great Bend and Larned. Magrath cited one
collection as early as August 27, but all others are after September 13. Average collection date is
October 1. Fendler reached Fort Dodge about September 6, so the Spiranthes cernua specimen would
have been collected in the first week of September, early for the species, but not impossible. It is also
possible that Ames misidentified the specimen, which was not personally seen by me.
In summary, reports of Spiranthes vernalis in New Mexico are incorrect, in our opinion, and are
based on Ames' erroneous citation of Fendler 32 (841) which was collected in eastern Kansas on
125
Jennings & Sheviak: ON THE REPORTS OF SPIRANTHES
BOOK REVIEWS VERNALIS IN NEW MEXICO
August 14, 1846, not 1847. A second white-flowered orchid specimen (Fendler 119), considered by
Ames to be Spiranthes cernua, was collected in Kansas as well.
REFERENCES CITED
Ames, O., 1905. Orchidaceae, Fascicle I, A synopsis of the genus Spiranthes north of Mexico, pp.122-156.
Brown, W. E., 1988. The Santa Fe Trail. St. Louis: The Patrice Press.
Ewan, J.A., 1950. Rocky Mountain Naturalists. Denver: The University of Denver Press.
Gray, A., 1849. Memoirs of the American Academy of Arts & Sciences, v. 4, part 1
Luer, C.A. 1975. The Native Orchids of the United States and Canada, excluding Florida. New York: New York
Botanical Garden.
Magrath, L.K. 1971. Native Orchids of Kansas. Transactions of the Kansas Academy of Science 74: 287-309.
Magrath, L.K., 1973. The Native Orchids of the Prairies and Plains Region of North America, Ph.D. dissertation,
University of Kansas, Lawrence.
Shaw, E.A. 1982. Augustus Fendler's collection list, 1846-1847. Contrib. Gray Herb. 212: 1-70.
Standley, P.C., 1910, Type Localities of Plants First Described from New Mexico, Contrib. U.S. Nat.
Herb. 13: 143-228.
William F. Jennings, P.O. Box 952, Louisville, CO 80027 – wfjenningspe@netscape.net

Charles J. Sheviak, Division of Research and Collections, New York State Museum, Albany, NY 12230 –
csheviak@mail.nysed.gov
126
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June 2007 North American Native Orchid Journal

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NORTH AMERICAN

NATIVE ORCHID JOURNAL

THE NATIVE ORCHIDS OF

In a continuing saga of the green-flowered species of Platanthera in the

I am very grateful to Alison Colwell, Peggy Moore, Chuck Sheviak, Bill

Note: Madrono 54(1) Jan.-Mar. 2007 was published on July 3, 2007.

Platanthera tescamnis Sheviak & Jennings

Intermountain rein orchid

Platanthera tescamnis, the Intermountain rein orchid, has been long

Platanthera tescamnis, Colorado

Platanthera yosemitensis Colwell, Sheviak, & Moore

Yosemite bog orchid

Unquestionably one of the most restricted species of Platanthera in this

Platanthera yosemitensis, Mariposa Co., California

DROUGHT, PERIL, AND SURVIVAL

The small white lady‘s slipper, Cypripedium candidum, is a rare sight

displaced by human activities, rescue operations are often the only

the early 1990s, approximately 75 years after it was assumed extirpated

Figure 2. Maroon petals and sepals

In bud, the labellum

Figure 4. A rescued plant in bud on May 6th, 2006.

As the blooming season comes to a close the flowers senesce and

Figure 5. Cypripedium candidum as

habitat (K. Kennedy, personal communication). The orchids were

Figure 6. A rescued plant as the flower begins to fade.

distance. Those local adaptations are another important reason that

Figure 7. C. candidum plant emerging from the roadside debris.

Seed-banking the local wild-type:

years, or even decades, is a powerful tool for conservation, provided that

Figure 8. (left) Cryopreserved Platanthera sp. seeds, SEM at 194x.

Figure 9. (right) Scanning electron micrographs (SEM) at 372x, of cryopreserved

If practical it would be ideal if each state, province or region could

maintain healthy populations there need to be increased incentives for

GOOD THINGS COME IN SMALL PACKAGES or

The Slow Empiricist

Calypso bulbosa var. americana

Triphora trianthophora var. trianthophora

Sacoila lanceolata var. lanceolata

Cypripedium parviflorum var. pubescens lacking anthocyans

I. THE EFFECTS OF VARIOUS MEDIA ON THE

Materials and Methods

preliminary experiment and previous reports of the necessity of low light

Chemical Composition of the Media

Medium Nutrient Ions mM/l

NO3 NH4 K PO4 Fe SO4 Ca Mg Cl Mn Sugar Conc.

Stage 1 - dead seed or seedling

Source d.f. M.S. ―F‖

Source df M.S. ―F‖

Treatment Means for 5 Replications and Species and Medium Means

180 Days 240 Days

1 309.4 205.9 200.1 238.5 244.0 224.7 234.6

7 225.6 204.0 199.9 209.8 269.0 234.7 251.8

L.S.D. 176 L.S.D. 4.59

Knudson‘s C medium gave a final reading of 234. This medium produced no

reading of 276 corresponded to .319 % of the seedlings which reached stage

Discussion and Conclusions

nature of the chemicals which make up this commercial fertilizer contain

Curtis, J. T. 1942. Germination and seedling development in five species of Cypripedium L.

Sladden modified by Burgeff

Vacin and Went 1949

Vacin and Went tomato formulation

Wynd 1933 Type iv R5S1

II. The Effects of Various Growth Regulators on the