March 1999 North American Native Orchid Journal

NORTH AMERICAN
NATIVE ORCHID JOURNAL

______________________________________
Volume 5 March
Number 1 1999
a quarterly devoted to the orchids of North America
published by the
NORTH AMERICAN
NATIVE ORCHID ALLIANCE
* * * * * * *
* * * * * * *
IN THIS ISSUE:
A MID-SUMMER PIPE DREAM
CALYPSO BULBOSA - HYBRIDIZATION AND
CULTIVATION
POLLINATION BIOLOGY OF SOME MEMBERS OF
THE YELLOW FRINGED ORCHID COMPLEX
CONSERVATIONREVIEW……………………….and
more!
1
NORTH AMERICAN NATIVE
ORCHID JOURNAL
(ISSN 1084-7332)
published quarterly in
March June September December
by the
NORTH AMERICAN NATIVE ORCHID ALLIANCE
a group dedicated to the conservation and promotion of our
native orchids
Editor:
Paul Martin Brown
Assistant Editor: Nathaniel E. Conard
Editorial & Production Assistants:
Philip E. Keenan
Stan Folsom
Nancy Webb
The Journal welcomes articles, of any length, of both a scientific

and general interest nature relating to the orchids of North
America. Scientific articles should conform to guidelines such as
those in Lindleyana or Rhodora. General interest articles and notes
may be more informal. Authors may include line drawings
and/or black and white photographs. Color inserts may be
arranged. Please send all inquiries or material for publication to
the Editor at PO Box 772121, Ocala, FL 34477-2121 (late May -
early Oct. Box 759, Acton, ME 04001-0759).
2000 Membership in the North American Native Orchid Alliance,

which includes a subscription to the Journal, is $26 per year in the
United States, $29US in Canada and $32US other foreign
countries. Payment should be sent to Nancy A. Webb, 84 Etna
St., Brighton, MA 02135-2830. Claims for lost issues or canceled
memberships should be made to the editorial office within 30
days.
2
NORTH AMERICAN NATIVE
ORCHID JOURNAL
Volume 5 March
Number 1 1999
CONTENTS
NOTES FROM THE EDITOR
1
RECENT TAXONOMIC AND
DISTRIBUTIONAL NOTES FROM FLORIDA 1.
Paul Martin Brown
3
VALIDATION OF THREE NEW
COMBINATIONS
17
Roger W. Bradley
19
CALYPSO BULBOSA—HYBRIDIZATION AND
CULTIVATION
Stan Ashmore
27
WESTERN PRAIRIE FRINGED ORCHID
TOUR, TOLSTOI, MANITOBA
Christie Borkowsky
34
CONSERVATION REVIEW
Philip E. Keenan
40
INTUITIVENESS
The Slow Empiricist
47
3
POLLINATION BIOLOGY IN SOME
MEMBERS OF THE YELLOW-FRINGED
ORCHID COMPLEX
Part 2. Breeding systems, factors contributing to reproductive
success in the orange fringed orchid, Platanthera ciliaris (L.) Lindley
and the white fringed orchid, P. blephariglottis (Willdenow) Lindley,
and pollinator driven morphological divergence in P. ciliaris
Charles L. Argue
53
BOOK REVIEWS
The Orchid Thief
Collins Photo Guide to the Orchids of Britain & Europe
70
LOOKING FORWARD:
June 1999
74
Unless otherwise credited, all drawings in this issue are by Stan Folsom
Color Plates:
1. page 15 Spiranthes eatonii
2. page 16 Epidendrum conopseum, Triphora trianthophora forma rossii
3. page 29 Calypso bulbosa hybrids
4. page 30 Calypso bulbosa hybrids
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 5, number 1, pages 1-74; issued March 24, 1999.
Copyright 1999 by the North American Native Orchid Alliance, Inc.
Cover: Zeuxine strateumatica by Stan Folsom
4
NOTES FROM THE EDITOR
At long last here is the March issue to start off
our fifth year. Due to an untraceable U S Postal Service
problem with the mailing of the December 1998 issue
of the Journal, many of you did not receive that issue. It
contained the 1999 renewal information. I am sending
the March 1999 to all of our members of record for
1998 with the 1999 renewal envelope. If you do choose
to renew (and I trust you will) please send your renewal
check in the enclosed envelope as soon as you are able.
Many of you have already renewed with out the renewal
notice and that is appreciated. If you have not receive a
copy of the December 1998 issue please indicate with
you renewal and it will be sent out as soon as I receive
that information. My sincere apologies for all of the
confusion and lateness of these two issues. I hope to
back on track with the June issue.
The 4th North American Native Orchid

Conference went well here in Florida and was, as so
aptly put by one of our members, 'a moveable feast' as
we traveled throughout much of the state in search of
those orchids that did manage to flower despite the
drought. More on that in the June issue, as well as the
first set of proceeding from the conference.
1
Articles are still needed for later issues of this
year and next. There have been requests for more
articles from a personal perspective and travel articles.
Please send along your notes and observations and I will
help you put them into a workable format for a journal
article.
The 5th Annual North American Native Orchid

Conference will be in the spectacular mountains of
Olympic Nation Park in the State of Washington from
July 16-20, 2000. Make your travel plans early and send
your reservations and/or registrations in soon. Full
details of speakers and field trips will be in the
September issue.
Paul Martin Brown

Editor
PO Box 772121
Ocala, Florida 34477-2121
(Mid-June-mid-September at PO Box 759, Acton,
Maine 04001-0759)
2
Brown: RECENT TAXONOMIC AND DISTRIBUTIONAL NOTES
FROM FLORIDA 1.
RECENT TAXONOMIC AND

DISTRIBUTIONAL NOTES FROM
FLORIDA 1.
Paul Martin Brown
Epidendrum conopseum R. Brown vat. conopseum

GREEN-FLY ORCHIS Epidendrum conopseum
R. Brown var. mexicanum L.O. Williams
BRONZE GREEN-FLY ORCHIS
In March of 1998 a friend was visiting in Florida

and noted that the green-fly orchid, Epidendrum
conopseum, I showed him was much more highly colored
than what he was used to seeing north of here. This
brought to my attention that there appeared to be two
distinct races of, Epidendrum conopseum growing in
Florida. After careful examination of both 'races' it
appears that in northern Florida (and northward) the
typical E. conopseum with watery green flowers and a
distinct three-lobed lip is the race to be found. This race
ranges southward through Marion County and is
scattered in the counties beyond to Highlands County.
In addition, from Marion County southward a highly
3
FROM FLORIDA 1.
colored race with bronze-pink flowers and a scalloped

lip occurs. In one tree in southwestern Marion County,
at the Ross Prairie, both colorations can be found in
flower at the same time. Upon examination of living
plants of Epidendrum conopseum from Mexico it was
determined that the Mexican plants and the highly
colored, scalloped-lipped plants from central Florida are
the same taxon. This taxon corresponds to the var.
mexicanum described by L.O. Williams. Color on
herbarium specimens is very difficult to ascertain unless
noted by the collector, but the lip shape can be
determined, especially in plants where the lip has been
soften in liquid. Photographs are perhaps the easiest
way to determine the two varieties as both the color and
lip shape is easily seen.
Platythelys querceticola (Lindley) Garay. SYN:

Erythrodes querceticola (Lindley) Ames
LOW GROUND ORCHID
Platythelys sagreana (A. Richard) Garay
CUBAN GROUND ORCHID
It was recently brought to my attention by Dr.

Robert Dressler that Platytheles sagreana was being
recognized in a work in preparation on the Orchidaceae
of the Lesser Antilles. After searching the literature for
the validity of the name and comparing the various
descriptions and keys it became apparent. that this
species should be recognized from Florida. Small
recognized it in his 1933 work as Physurus sagreana and
4
FROM FLORIDA 1.
subsequently it was reduced to the varietal level and

then altogether disappeared within Erythrodes (Physurus)
querceticola. When Garay created the genus Platythelys he
made the transfer of Erythrodes querceticola var. sagreana to
its current status. The two species are not difficult to
identify from herbarium material, photographs or in the
field. They also are geographically distinct in Florida
with P. querceticola being in central and northern Florida
and P. sagreana from Highlands County southward in
Florida. Both are pictures in Luer's 1972 work on the
Native Orchids of Florida (plate 35:1, 2, 4, 6 = P.
querceticola, 3, 5 = P. sagreana). In a recent discussion with
Dr. Luer he told me that he included several
photographs from different locales as they did not look
quite the same!
Triphora trianthophora (Swartz) Rydberg subsp.

trianthophora
THREE BIRD'S ORCHIS;
NODDING POGONIA
trianthophora forma rossii
A massive colony of the three birds orchid was

found in southwestern Marion County, Florida in
September of 1998. I estimate that the colony has in
excess of 250,000 flowering plants and flowered from
late August to mid December!! Some variation was
observed other than number of flowers varying from 2-
14 per plants and colors from palest pink to deep violet
5
FROM FLORIDA 1.
pink. A few individuals of forma albidoflava Keenan were

observed. Occasionally individual colonies would have
very small flowers less than 1.5 cm across when fully
open! The most striking variation was 131 individuals
that were lacking in chlorophyll and produced stems
and leaves that were candy-stripe pink and white with
small white flowers with a yellow crest. This color
variation appears to be undescribed and the following is
proposed.

trianthophora forma rossii P.M. Brown form nov.
Forma folius alba et rosea conspeciebus divers a;

floribus albus et flavibus. TYPE: UNITED STATES:
Florida, Marlin County. Holotype: Ross Prairie. October
23, 1998. (Holotype: FLAS Brown 102398. Photo.
NANO] 19995(1): 16.
Differing from typical Triphora trianthophora subsp.

trianthophora by having its stems and leaves white and
pink with no green present; flowers white with a yellow
crest rather than the typical green; varying from the type
by having white and pink stems and leaves and the
flowers with a yellow crest instead of a green crest.
Spiranthes cernua (Linnaeus) L.C. Richard
6
FROM FLORIDA 1.
All specimens I have examined from a wide

variety of herbaria labeled as nodding ladies'-tresses,
Spiranthes cernua (Linnaeus) L. C. Richard, have proven
to be fragrant ladies'-tresses, Spiranthes odorata (Nuttall)
Lindley, Synonym: Spiranthes cernua var. odorata (Nuttall)
Correll. All sites I have examined in the field that were
reported to be S. cernua have proven to be S. odorata.
Although more northerly populations of S. cernua do not
currently or historically range close to the northern
Florida border it is not impossible that a population of
good S. cernua could still be found in the bordering
counties. Examination of potential sightings and any
additional herbarium specimens will be continued. At
this time I have deleted S. cernua from the Florida list.
Spiranthes eatonii Ames ex P.M. Brown sp. nov.

EATON'S LADIES'-TRESSES
In 1905 A.A. Eaton collected extensively during a

south Florida trip for Oakes Ames. Much of that
expedition is detailed in Ames' Contributions to the
Knowledge of the Orchids of South Florida. Five specimens
from that expedition that are deposited at the Orchid
Herbarium of Oakes Ames (AMES) at the Harvard
University Herbaria were originally labeled Spiranthes
lacera. Ames annotated them as forma angustifolia and
eventually as Spiranthes eatonii sp. nov., even indicating
the type as sheet AMES 6905, but never published such
a name. In the ensuing 94 years this taxon has been
frequently collected and labeled variously as Spiranthes
7
FROM FLORIDA 1.
lacera, Spiranthes gracilis, Spiranthes lacera var. gracilis,

Spiranthes gracilis var. brevilabris, and, not surprisingly,
several specimens from central and northern Florida as
Spiranthes torta. The flowering stems are very slender
with little or no twist to them as in Spiranthes lacera s.s.
and the flowers posses a green throat as in both S. var.
lacera and gracilis as well as S. torta. Although most plants
flower with the distinctive lanceolate to oblanceolate
leaves present, occasional flowering plants are found
with no leaves present. It is this last condition that has
led to misidentifications as Spiranthes torta. Spiranthes
torta has slender, linear leaves that are usually absent at
flowering time.
The plants of Spiranthes eatonii flower from late

February to late April in Florida and into May in the
adjacent Gulf States. Spiranthes lacera var. lacera and S.
lacera var. gracilis both are summer to autumn flowering
species and, when they occur in the same states as S.
eatonii, the flowering time, as well as the geographic
distribution, is well marked.
Very simply put,

• winter/spring flowering white Spiranthes
• with a green throat and
• lanceolate to oblanceolate basal leaves (with a winter
rosette) are S. eatonii.
The lip, sepals and pubescence are all distinctive in

Spiranthes eatonii. Many of these characters are compared
below with the other similar species. Spiranthes eatonii is
8
FROM FLORIDA 1.
easily separated from S. brevilabris by the latter have a

densely pubescent inflorescence as well as creamy-
yellow flowers with deeper yellow lips. Spiranthes torta
flowers later and is confined to the southernmost
counties of Florida. Although the leaves are not present
at flowering, S. torta has very narrow linear leaves.
Spiranthes lacera var. lacera and S. lacera var. gracilis are not
present in Florida. Preliminary DNA comparisons
indicate that S. eatonii is well separated from S. lacera s.l.
as well as from S. brevilabris and S. floridana.
Spiranthes eatonii Ames ex P.M. Brown sp. nov

TYPE: UNITED STATES: Florida. low, sandy pine
woods, Orange Glade, Dade Co., Florida Feb 21. 1905.
Eaton 1200 (Holotype: AMES 6905)
Habitu, forma inflorescenctiae Spiranthes lacera var. lacera

similis sed folia divergens foliae linearis - lanceolatus,
florens vernalis et with spatulatus sepalae, labia alba
cum viridis; sepalae et petalae viridis basi.
Plants 15 - 55 cm tall; leaves basal, oblanceolate to 7.5

cm long x 1cm wide; flowers white, 3-5 mm long with
green throat; petals white with green bases; lip white
with green central area; sepals spatulate with green at
base; leaves appearing in late autumn and withering just
before, at, or immediately after anthesis. Differs from
Spiranthes lacera var. lacera and S. lacera var. gracilis in the
late winter/spring flowering time, longer linear-
lanceolate leaves and smaller flowers without a
pronounced crisped apron on the lip and from S. torta
9
FROM FLORIDA 1.
by the linear-lanceolate leaves rather than narrow-linear

leaves.
Habitat: dry to damp open pinelands often with Aristida

sp., Serenoa sp., Pinguicula sp., Drosera sp., and Polygala sp.
Range: Florida; eastern Texas, southern Louisiana,
southern Alabama, southern and eastern Georgia and
north to Virginia primarily on the coastal plain
ISOTYPES:
AMES 6906, AMES 6907, AMES 6918, AMES 6927
Representative specimens examined:

ALABAMA: Baldwin Co., 10 Jun 1973, R.. Kral 50481. Sandy longleaf
pineland, Seminole. (VDB) Mobile Co., Dry grassy pinebarrens, Mobile Bay.
C. Mohr 1477 (ALU); C. Mohr s.n. (ALU); Lee Co., 21 Jun 1980 Robert R.
Haynes 7862 waste area ca. 2mi. due S of Smiths along Co. Rd. 91; (UNA);
Chilton Co., sandy & gravely pine hills between Adams & Plechter. Roland M.
Harper 4330 (UNA); Mobile Co., Hillside pineland bog by US 45, 3.8 mi se
Chunchula 3 Jun 1970.R. Kral 39610. (UNA); Cadillac Square, oak-pine forest,
infrequent, flowers white. 5-30-64 Rebecca Deramus D95 (UNA);
FLORIDA: Alachua Co.: low, moist sandy pinelands, about 5.5 miles
northwest of Gainesville. Lip green! April 6, 1938 D.S. Correll & H.B. Correll
8926 (AMES); Clay Co.: Jennings State Forest. Dry palmetto scrub with pines.
14 April 1998 P.M. Brown 98-414 (FLAS); De Soto Co.: Ft. Ogdon. March 31,
1905 A. A. Eaton 1458 (AMES); Hernando Co.: lawn on Chinsegut Hill. Nine
plants seen. Corollas white, crystalline, greenish toward base of tube. April 12,
1958. George R. Cooley 5901 (USF); Highlands Co: damp pinelands e. of
Sebring with S. gracilis var. floridana, but is smaller and has more delicate
flowers. Var. floridana now mostly past blooming. 24 March 1948 Ray Garrett
s.n. (FLAS); Moist pinelands, south shore of Bennett Lake, north of Sebring.
Grows with gracilis var. floridana but blooms 2-4 weeks later R.G. 31 March
1948 Ray Garrett s.n. (FLAS); Hillsborough Co., University of South Florida
ecology area, jct of 56th ST & Fletcher. 12 Nov. 1979, R.P. Sauleda & R.P.
Wunderlin. 3527. (GA) (USF); Jackson Co.: flatwoods, Marianna. 22 April
1949 S.C. Hood 1989 (FLAS); Lee Co.: sandy railroad grade, E. Ft. Myers.
March 30, 1905 A. A. Eaton 1441 (AMES); Levy Co.: Rt. 24, 1.25 miles East
of Route 19. Grassy roadside, ditch bank. April 22, 1983 Mary E. Schmid A-
Zil s.n. (USF); Walton Co., 4 mi. S of Freeport in our open grassland and open
10
FROM FLORIDA 1.
pine woods. April 23, 1971 Mr. & Mrs. H. A. Davis 15641 (FLAS); Highlands
Co.: Moist pinelands,s shore of Bennett Lake, n. of Sebring. Grows with S.
gracilis var. floridana but blooms 2-4 weeks later. 31 March 1948 Ray Garrett
s.n. (FLAS); Volusia Co., hammock, New Smyrna April 19, 1910. S.C. Hood
s.n. (FLAS); Open pine woods, Orange City, April 2, 1910. S.C. Hood s.n.
(FLAS); Wakulla Co.: Ochlockonee River State Park. Clay Co.: Jennings State
Forest. Dry palmetto scrub with pines. 10 April 1998 P.M. Brown 98-410
(FLAS);
GEORGIA: Chatham Co., poorly drained meadow, sandy soil, in open area
along railroad 0.4 miles S of St. Augustine Creek along Springfield Hwy., 8.0
mi. w of Savannah City Hall.(GA); Coffee Co., cleared right-of-way along
highway, upland sandy soil 7.9 miles WNW of Douglas. James W. Hardin,
Wilbur H. Duncan 16211 18 May 1953 (NCU; LL); McIntosh Co., pineland
east-central part of Sapelo Island. Williams P. Adams, Wilbur H. Duncan
17866 (GA; LL; NCU); Miller Co. pineland, 2 miles W of Colquitt. April 28,
1947. Robert F. Thorne 3406 (GA) (AMES); Taylor Co., 7 m. SW of Butler on
state highway 137. Long-leaf pine-turkey oak sandhills occasional. 25 May
1991 M.S. Morris 29 (FLAS);
LOUISIANA: Bayou Lacombe R.S. Cocks s.n. May 1909 (NO); Vicinity of
Covington, Sulphur Springs. May 28, 1920. Bro. G. Arsene 11881 (AMES);
NORTH CAROLINA: Beaufort Co.: open pineland on Route 17 about 8
miles south of Washington. 19 June 1947. C.E. Wood & I. D. Clement 6978
(AMES); Perden Co.: Big Savannah. May 1, 1925 (AMES)
SOUTH CAROLINA: Horry Co.: pine barrens south of Socastee. Many 2,
1932. Weatherby & Griscom 16496 (AMES);
TEXAS: Jefferson Co.: 5.5 miles south of Sabine Pass; infrequent on
roadsides. April 18, 1950 V.L. Cory 57167 (SMU);
VIRGINIA: Princess Anne Co: sandy pineland, Cape Henry. June
17, 1935. Fernald, Griscom & Long 4618 (AMES).
Acknowledgments:
The authors thanks Gustavo Romero (AMES), Charles J. Sheviak
(NYS), Paul Catling (DAO), Norris Williams (FLAS) and the
curators of USF, ALA, BRIT, NCU, LL, NO, SMU & TX for the
loan of specimens.
11
FROM FLORIDA 1.
Spiranthes eatonii Spiranthes lacera var. lacera

Range
throughout FL; west to e TX and Nova Scotia west to Alberta
north to seVA; almost south to Missouri and east to
exclusively in the coastal northern Georgia
counties
Habitat
sandy, dry pine barrens; dry, grassy roadsides; old fields;
cemeteries usually associated open woodlands; damp meadows
with pines; drier portions of pine
flatwoods
Flowering period
late February (in southern mid July - mid-August
Florida) to late May in North
Carolina and Virginia
Leaves
usually persistent at anthesis; usually present at anthesis;
petioled, oblanceolate 1.0 x 5.0 obovate 1.0 x 2.5 cm
cm
Floral arrangement
loosely flowered with one to loosely flowered with one to
several twists; lower flowers several twists; lower flowers
widely spaced widely spaced
Color
white with green central portion white with green central portion
on lip and green at base of petals on lip
and sepals
Sepals
white with green on basal white; narrow tapering to a point
portion; 3-5mm long; broadly 3-6mm long
spatulate with the middle pinched
Lip
white; 3-5mm long with basal white; 3-6 mm; broadest at apex
callosities thickened and pointing with a clearly defined, narrow,
outward; lip narrowed to a crisp margin; central portion
rounded apex, central portion green
green, without the crisped apron
found on S. lacera s.l.
Pubescence
capitate glands within the capitate glands within the
inflorescence and scattered on inflorescence and scattered on
the rachis the rachis
12
FROM FLORIDA 1.
Spiranthes lacera var. gracilis Spiranthes floridana

Range
Missouri; Michigan east to Texas east to Florida north to
southern Ontario and Maine and North Carolina - VERY RARE
south to northern Georgia and
west to Texas
Habitat
dry to moist grasslands and open dry to moist roadsides and
woodlands; damp meadows, meadows
cranberry bogs
Flowering period
August - early October March - April
Leaves
usually absent at anthesis; ovate withering at anthesis; yellow-
1.5 x 3.5 cm green; oblanceolate 1.0 x 3.4 cm
Floral arrangement
densely flowered with several flowers approximate with lower
twists; no spacing between lower ones distant
flowers
Color
white with green central portion creamy-white with yellow central
on lip portion; rachis also yellow-green
giving an overall pale yellow cast
to the entire plant
Sepals creamy-white 3-6 mm long

white; narrow tapering to a point
3-6mm long
Lip
white; 4-7.5 mm; rounded at creamy-yellow with darker
apex with a clearly defined, crisp yellow central portion; entire
margin; central portion green flower has a box-like appearance
from the floral parts being of
equal length.
Pubescence
essentially glabrous essentially glabrous
13
FROM FLORIDA 1.
The type specimen of Spiranthes eatonii with Oakes Ames' annotations on

The type the lowermost paragraph reads: Allied to S. gracilis Beck from
the sheet;
which is differs in the persistent, mostly elongated leaves, and in the
oblong-ovate lip which is narrowed toward the apex. O.A.
14
FROM FLORIDA 1.
Spiranthes eatonii Ames ex P.M. Brown

Eaton's ladies'-tresses
Ochlockonee River State Park,
Wakulla County, Florida
April 10, 1998
P.M. Brown
15
FROM FLORIDA 1.
Epidendrum conopseum
var. conopseum var. mexicanum
green-fly orchid bronze green-fly orchid
Triphora trianthophora subsp. trianthophora forma rossii

pink and white multi-color form
Ross Prairie, Marion County, Florida P.M. Brown
16
FROM FLORIDA 1.
VALIDATION OF THREE NEW COMBINATIONS
VALIDATION OF THREE NEW

COMBINATIONS
In the North American Native Orchid Journal [4(1):51-

53. 20 Mar 1998], the following three new combinations
were published: Prosthechea boothiana (Lindl.) W. E.
Higgins var. erythronioides (Small) W.E. Higgins comb.
nov.;
Prosthechea cochleata (L.) W. E. Higgins var. triandra
(Ames) W.E. Higgins comb. nov.;
and Prosthechea cochleata (L.) W. E. Higgins var. triandra
(Ames) W.E. Higgins forma albidoflava (P. M. Brown) P.
M. Brown, comb. nov.
For these three new combinations, the full and direct
references were given on their basionyms.
Even though it may appear that those three new

combinations were validly published, it should be noted
that for an infraspecific name to be valid, its specific
name to which it is assigned is also validly published at
the same time or was validly published before
(International Code of Botanical Nomenclature Art. 43.1).
It is true that Higgins [Phytologia 82(5): 376-377.

1997] did publish the two specific new combinations (P.
boothiana & P. cochleata) validly. However, the Phytologia
vol. 82(5) was issued on 30 Mar 1998 (i.e., 10 days after
the publication of those three infra names). Because of
the delayed publication of the two specific names, those
three infra names were invalid when published.
17
FROM FLORIDA 1.
VALIDATION OF THREE NEW COMBINATIONS
Therefore, Higgins and I re-publish those three new

combinations here for validation.
Prosthechea boothiana (Lindley) W.E. Higgins var.

erythronioides (Small) W.E. Higgins comb. nov.
Basionym: Epidendrum erythronioides Small, Fl. Southeastern
US. 328, 1329. 1903.
Synonym: Encyclia boothiana (Linnaeus) Dressler var.
erythronioides (Small) Luer. Luer, Nat. Orchids of Fl. 204.
1972.
Prosthechea cochleata (Linnaeus) W. E. Higgins

var. triandra (Ames) W.E. Higgins comb. nov.
Basionym: Epidendrum cochleatum Linnaeus var. triandrum
Ames, Contrib. Ames Bot. Lab. 1:16. 1904.
Synonym: Encyclia cochleata (Linnaeus) Dressler var.
triandra (Ames) Dressler. Luer, Nat. Orchids of Fl. 202.
1972.
Prosthechea cochleata (Linnaeus) W. E. Higgins

var. triandra (Ames) W.E. Higgins forma
albidoflava (P. M. Brown) P. M. Brown comb. nov.
Basionym: Encyclia cochleata (Linnaeus) Dressler var.
triandra (Ames) Dressler forma albidoflava P.M. Brown,
NA Native Orchid Journal 1(2): 131. 1995.
Paul Martin Brown.

I wish to thank Dr. K. N. Gandhi, Gray Herbarium Index
Bibliographer & Type Specimens Database Manager, Harvard
University Herbaria, for noting this error and assisting with the
correction.
18
Bradley: A MID-SUMMER PIPE DREAM
Roger W. Bradley
Field trips promising the probability of seeing

several species of orchids are always beckoning, but the
anticipation usually exceeds reality. Offsetting this,
some unplanned trip often more than makes up for
what was lacking on the original field trip. This is what
happened in mid-July in the White Mountains of New
Hampshire.
The high elevation of Frankenstein Cliff delayed

the flowering time compared to the early flowering
season elsewhere in 1998. It was high, very steep and
somewhat treacherous in places, yet we saw lots of good
things such as views of the impressive Presidential
Range and Mt. Washington, highest point in the
northeast. One of the plants seldom seen was
squawroot, Conopholis americana, usually as a single plant
or two. On this occasion there were dozens of them.
About halfway up, there were two pad-leaved

orchids, Platanthera orbiculata, that had passed peak
flowering and that were weather beaten. Last year there
was a colony of them. Above a twisting staircase of
boulders near the base of the cliff was a small purple
fringed orchid, Platanthera psycodes, in good shape
although the single flower was quite small, perhaps
19
attributable to the altitude and lack of soil. There were

also a couple of spotted coralroots, Corallorhiza
maculata var. maculata, well past prime.
The return down the mountain was almost as

arduous as the climb up. We stopped at Arethusa Falls,
the highest in New Hampshire at 200 feet. Seven hours
and five and a half miles later during idle chit-chat at the
end of the day, a fellow amateur botanist invited me to
stay at his place and the next day go over his family's
170 acre woodlot in north central New Hampshire.
There the large pad-leaved orchid, Platanthera
macrophylla, and the pad-leaved orchid, Platanthera
orbiculata, occurred only a few feet apart in perfect
flower for a good comparison. In no way could I pass
up that opportunity!
On a clear mild sunny morning starting at the

hunting cabin, we toured the wood roads past bogs,
fens, and wooded hillsides for 3 glorious hours hunting
orchids. First we came upon the two species of pad-
leaved orchids only a few feet from each other under
tall oaks. The difference was very distinct. The spur on
the Platanthera orbiculata was two centimeters long; the
spur on the Platanthera macrophylla was four and a half
centimeters long. Comparatively, the lateral petals of
the orbiculata flared out sideways while those on the
macrophylla flared upward. The basal leaves of the
macrophylla were slightly larger and rounder than the
more oval leaves of the orbiculata. (Paul Martin Brown
had pointed out these distinctions on a field trip several
years ago in Maine.)
20
Platanthera macrophylla
Goldie's pad-leaved orchis
21
Platanthera orbiculata
pad-leaved orchis
22
Platanthera grandiflora
large purple fringed orchis
23
Platanthera psycodes
small purple fringed orchis
24
Having the woods all to ourselves without

competition from enthusiastic photographers made a
relaxed situation to observe. We had to be very careful
where we stepped, as there were many checkered
rattlesnake orchids, Goodyera tesselata, everywhere
around the Platantheras.
On another woodroad heavily canopied with

oaks, I found two spotted coralroots, Corallorhiza
maculata var. maculata , that my friend had not noticed
before. One was the yellow-stemmed form (flavida)
and the other the red-stemmed form (rubra). Each still
had a few good flowers at the top of its inflorescence.
Finally, a faint path off a branch woodroad over a

low rise led to the "frosting-on-the-cake." There is
surely no more exhilarating sight than to see a brilliant
deep pink colony of large purple fringed orchids,
Platanthera grandiflora, illuminated by a sunbeam
streaming into a small fen within the dark surrounding
forest. A half dozen large size plants were clustered in a
small wet area. Each had a large raceme of flowers
showing the well-defined fringes on the lip.
What a fantastic trip: a most welcome congenial
overnight stay at a friend's place, a tour of a private
wildlife preserve, perfect summer weather, and several
species of wild orchids in excellent condition! "You
can't hardly beat that no more!" Reflecting on the
morning's exceptional good luck made the two hundred
mile drive home no trouble at all.
Roger W. Bradley, 617 Springfield Rd.,
Somers, Connecticut 06071
25
Corallorhiza maculata
spotted coralroot
26
Ashmore: CALYPSO BULBOSA--HYBRIDIZATION AND
CULTIVATION
CALYPSO BULBOSA—HYBRIDIZATION
AND CULTIVATION
Stan Ashmore
No northern terrestrial orchid is more widely

admired than the fairy slipper, Calypso bulbosa.
Exquisitely beautiful and hauntingly fragrant, Calypso
would be a horticultural prize were its cultivation not so
difficult. The reasons for this difficulty are not
completely known, but disease and habitat constraints
appear to be major problems in cultivation attempts.
Hybridization and selection of promising clones

are successful strategies that plant breeders have used to
overcome many disease and adaptation problems such
as those affecting the genus Calypso. By crossing
intraspecific varieties and growing them in artificial
media, I have attempted to apply these strategies to
Calypso bulbosa. Cultivation still presents problems, but
some lovely hybrid plants have resulted.
Hybridization
I first attempted to cross the eastern fairy slipper,

Calypso bulbosa var. americana, plants from south-central
Alaska with the western fairy slipper, Calypso bulbosa
27
CULTIVATION
var. occidentalis, plants from southeast Alaska. Out of

fifteen individual F1 crosses, two produced blooming
plants. One of these was particularly vigorous and
produced blooms in less than 3 years after seeding. This
cross was named Calypso bulbosa Aaron Island after the
island of origin, near Juneau, Alaska, of the pod parent.
Numerous clones of this cross were produced from
protocorms. It has a particularly beautiful flower, with
the yellow beard of Calypso bulbosa var. americana and a
profusion of tiny purple spots (fig. 1).
A characteristic of Calypso bulbosa is its requirement

for cool growing conditions. It does not survive as an
indoor plant at room temperatures. One form of
Calypso bulbosa var. occidentalis, however, grows in
redwood groves as far south as Sonoma County,
California under fairly warm summer temperatures. I
attempted, therefore, to cross the Alaskan Calypso bulbosa
var. americana with the warmer growing Calypso bulbosa
var. occidentalis from California in an effort to introduce
heat resistance into the hybrids. The plants from that
cross are not vigorous and some are deformed. The
Aaron Island hybrid, however, readily crossed with the
California variety. Plants from this cross, between an F1
hybrid and a stable variety (fig. 2), are very vigorous and
produce flowers ranging across the spectrum from
Calypso bulbosa var. occidentalis forms to those resembling
Calypso bulbosa var. americana with spots (fig. 3).
28
CULTIVATION
Left: Fig. 1
Calypso bulbosa Aaron Island
Center: Fig. 2
C. bulbosa var. occidentalis (left)
C. bulbosa Aaron Island (right)
Bottom: Fig. 3
Calypso bulbosa Sonoma hybrids
29
CULTIVATION
Left: Fig. 4
Calypso bulbosa in vitro
Below: Fig. 5
Calypso Bulbosa growing in a hydroponic unit
All photos courtesy of Stan Ashmore
30
CULTIVATION
Cultivation
Numerous Calypso bulbosa hybrids have bloomed in

flasks (fig. 4) under completely sterile conditions,
demonstrating that Calypso bulbosa is a facultative
symbiont and not an obligatory symbiont as is widely
believed. This has important implications for
cultivation, as it suggests that Calypso bulbosa, like most
other autotrophic orchids, can be grown asymbiotically.
Still, growth and survival do not match that found in
many wild populations in Alaska.
I have experimented with all of the commonly

used orchid potting media including rockwool and
perlite, and a variety of organic mixtures. In addition I
inoculated organic media with the mycorrhizal fungus
Epulorhiza which I obtained from Randy Currah of the
University of Alberta in Edmonton. Dr. Currah
extracted Epulorhiza from the roots of Calypso bulbosa
plants. This suggests that it may be a possible Calypso
bulbosa symbiont. No growth or survival advantage
seems to have been imparted by the Epulorhiza in my
experiments.
The most successful artificial growing medium has

been perlite placed in gallon pots in an ebb and flow
hydroponic system. Calypso bulbosa hybrids have been
growing in this system for several years at temperatures
31
CULTIVATION
near freezing in winter to 80° F in summer. Some

plants have bloomed under these conditions. Most
increased in size (fig. 5). Growth was improved by
increasing the hydroponic fertilizer concentration to
that recommended for cymbidiums. Disease was
reduced in this near sterile environment, but still occurs
when the temperature and humidity are high.
The hydroponic growing conditions described

work for Calypso bulbosa propagated asymbiotically and
can not be expected to work for wild plants, which have
invariably been growing in association with symbiotic
fungi. Readers are implored not to remove any Calypso
bulbosa from the wild.
Stanley Ashmore, HC 04, Box 9248-D, Palmer, Alaska 99645

Stan last wrote for the Journal on this same topic in June of 1995.
Calypso bulbosa Calypso bulbosa

var. occidentalis var. americana
32
Borkowsky: WESTERN PRAIRIE FRINGED ORCHID TOUR,
TOLSTOI, MANITOBA
WESTERN PRAIRIE FRINGED ORCHID

TOUR, TOLSTOI, MANITOBA
Christie Borkowsky
Welcome to Manitoba's Tall Grass Prairie

Preserve. In 1987 the Manitoba Naturalist Society
began a systematic search for remnant pieces of prairie
habitat in the province. Several locations were found
but the largest pieces were found here, in southeastern
Manitoba. The Society purchased their first parcel of
land in 1989. A self-guiding trail was designed in 1996,
to encourage the public to visit the Preserve.
At this point, the Preserve consists of over 5,000

acres of land and is divided into three blocks. The
south block includes the self-guiding trail and a
rotational grazing project. The central block, 960 acres
in size, is a designated Wildlife Management Area
(WMA). The headquarters for the Preserve is located in
the north block.
To date, 10 orchid species have been identified

on the Tall-grass Prairie Preserve. Three of the 10 are
listed as endangered species in Manitoba and include the
small white lady's-slipper, Cypripedium candidum, the
western prairie fringed orchid, Platanthera praeclara,
34
TOLSTOI, MANITOBA
and the great plains ladies' tresses, Spiranthes

magnicamporum.
The small white lady's slipper flowers in late

May to early June. The largest known population in
Manitoba is on the Preserve. A total of seven quarter
sections are known to produce this orchid, all in the
south block. Specific counts were started in 1993 and
to date, 1997 has been our peak year with nearly 17,000
flowering stems counted. An additional 18,000
vegetative stems were found for a combined total just
under 35,000 stems.
Great Plains ladies' tresses flower in early

September. It is the smallest endangered orchid in
Manitoba and is known to exist on 17 quarter sections
in both the north and south blocks. Counts began in
1994 and turned up 36 flowering stems, in 1996 the
counts topped 2,800 flowering plants.
The western prairie fringed orchid generally

flowers at the end of June to about the third week of
July, although this year I found three plants flowering
on June 19th. Of the three endangered species, it has
the largest distribution, being found on all the
properties that make up the north block and extending
beyond the Preserve limits. Fringed orchids are found
between the communities of Stuartburn and Vita along
PR201 (14.5 km) and 5.5km north of PR 201 to 2.5km
south of PR 201. Within this area, inventory work has
identified 61 quarter sections where fringed orchids are
found.
35
TOLSTOI, MANITOBA
There are several ongoing projects to monitor

this orchid from year to year. The most intensive
project is the flowering survey. All flowering plants
found are counted. By the time we have completed the
survey we have looked at nearly 60km of ditches
(driving at a speed of 5-10km/h), walked around 21
quarter sections by foot and viewed another 40 quarter
sections of private land from the road. Fortunately
these orchids are highly visible and the use of binoculars
aid in the survey. From the survey we have seen an all
time low of only 1,818 flowering plants in 1995, to an all
time high of just over 21,000 in 1996.
In 1994, two permanent plots were set up to look

at both vegetative and flowering plants. Within each
plot, every fringed orchid that came up in 1994 was
tagged with a numbered, aluminum tag. Just under 200
plants were marked. Each year we record the plant
form (1-, 2-, 3-leaved, flowering or dormant), height,
number of flowers and number of seed pods. When we
returned this year, only a third of the plants came up. In
1997 we found that 66% of the flowering plants
produced at least one seed pod, and of all individual
flowers, 15% produced a seed pod
A third monitoring project resulted when a gravel

road was reconstructed in 1995. The ditch area was
increased and the road built up. This occurred prior to
the official listing of the fringed orchid as endangered.
Summer staff removed 96 plants and transplanted them
to a secured property within the Preserve,
36
TOLSTOI, MANITOBA
approximately 3.5km northwest of the rebuilt road. As

with the permanent plots, each plant was tagged and
data for each plant was recorded. The following year,
nearly 85% came up and in 1997 nearly 67% of the
orchids were growing. This year, while working on the
ditch survey, I saw the first flowering fringed orchid
in the ditch that had been reworked three years earlier.
In 1997, we began a pollinator project for the

western prairie fringed orchid. Research from the
United States, has shown that long-tongued sphinx
moths were pollinating the flowers. There was no such
information for the Manitoba population of fringed
orchids. A moth expert in Manitoba also suggested that
the peak flowering period may not coincide with the
peak flight period of these sphinx moths. Using two
malaise traps and three cone traps, two moth species
with the pollinia structure attached to the eye area were
collected. They were like the wild cherry sphinx,
Sphinx drupiferarium and galium sphinx, Hyles gallii.
This project has been carried over to this field season,
but we have not yet collected moths with the pollinia
attached.
Christie Borkowsky is the biologist for the Tall Grass Prairie in

southern Manitoba and hosted our visit there for the 3rd North
American Native Orchid Conference on July 11, 1998.
Cypripedium candidum
small white lady's-slipper
37
TOLSTOI, MANITOBA
Platanthera praeclara
western prairie fringed
orchis
38
TOLSTOI, MANITOBA
Spiranthes magnicamporum
Great Plains ladies'-tresses
39
Keenan: CONSERVATION REVIEW
CONSERVATION REVIEW
Philip E. Keenan
One has to question the long-term outlook for

our native orchids in North America and throughout
the world —especially with respect to several major
trends that do not bode well for the future of our native
orchids. From the global view, we should face up to the
biggest potential problem, namely the explosive growth
of the world's population. Population demands create
an ongoing and accelerating destruction of the world's
greatest and greenest remaining masterpiece, the
Amazon River Basin. It directly relates to satisfying the
burgeoning population’s demand for fast food
hamburgers, no less! The rainforest’s destruction is
justified because it provides for the ever-expanding
need for homesteads and agriculture to fulfill the fast
food industry’s requirements. This is depleting the
natural resources of one of the largest countries of the
world, namely Brazil.
The forests of Indonesia are even more

devastated. Remember the recent pall of smoke in that
part of the Pacific? It was caused by the uncontrolled
burning by homesteaders and developers in those
fanciful islands, which formerly were known by their
40
Dutch colonial names of Borneo, Sumatra, Java and

New Guinea.
These catastrophes, of course, are directly related

to the spiraling world population. One way the ordinary
citizen might help ease this situation is to become
involved in organizations that encourage sound
ecological policies. It might be smart to support the
United Nations’ initiatives for family planning, which
might successfully defuse the ever-expanding
population in the world through some of their fine
programs.
The developing countries that rationalize their

current situation often do so by citing what Americans
did in the United States and what the Europeans did in
their countries at the turn of the century and earlier. In
the United States of America's case, the destruction of
99% of the virgin white pine forest in the northeast and
97% of the redwood forest in the Pacific Northwest, are
two examples. This wanton destruction can be
attributed to human greed and a failure to correctly
assess the long-term consequences. Human nature
doesn't change much in attempting to make excuses for
our excesses, whatever they may be. And most of us
fail to heed the lessons of history when money is at
stake.
There are many other recent examples such as

the dire threat to the remaining temperate rain forest off
the coast of British Columbia and southeast Alaska
(Tongass National Forest). Eighty-five percent of this
41
priceless, magnificent virgin forest has been destroyed

since World War II.
CITES (Convention on International Trade in

Endangered Species) is an international set of
regulations governing the import and export of plant
materials, such as orchids, signed by 100 countries in
1975. The intent was admirable but the results leave
something to be desired, according to some people.
There are the usual loopholes in the CITES rules, and
not all countries subscribe to them. Political influence
as well as corruption is sometimes a factor in non-
compliance. Unfortunately—human nature again—
when the price is right, some humans will do almost
anything to thwart the law and line their pockets, on
both ends of the transaction. A good example is drug
trafficking and many other illicit activities. If the
demand weren't there, however, the supply and
lawlessness would soon dry up. Some, therefore, argue
that making drugs legal would at least eliminate most of
the criminal aspects of that trade.
Making orchids available freely without

restrictions obviously wouldn't solve the orchid
decimation problem. But, as someone else in the orchid
field has pointed out, why isn't the export of orchids
taxed heavily enough so that it would dramatically
reduce the demand for orchids? Certainly the trade in
wild orchids is not a significant revenue producer for
any country, which would seem to argue in favor (?) of
such a policy.
42
Here in the United States, the country has

increasing amounts of poaching occurring in national
and state parks, as well as digging from private property.
With park personnel drastically reduced in recent years
we can no longer assume populations will remain safe in
these nominally protected environments. Each year one
hears and sees the evidence of what these cutbacks by
the government are doing to the parks. Forty-two
species of mammals have been extirpated in the
National Parks in less than 100 years and far more than
this figure has been lost in plant species. Ninety percent
of the Fraser firs in Smokey Mountains National Park
are dead or dying and the Park Service has no money to
save any or find the reason why this is happening,
although it is a good bet that acid rain is the major
culprit. The park service repair backlog has reached $6
billion dollars. In today's dollars, their annual budget is
$200 million dollars less than it was ten years ago,
despite the fact that visitors have exploded from 70
million in the 1960's to almost 300 million today.
Global warming is considered a fact to be

reckoned with by the vast majority of credible scientists,
yet a minority still manages to sway the United States to
ignore its consequences. Congress has done nothing. It
is reminiscent of the cigarette smoking issue which was
debated for decades before anything was done about it,
primarily because of the resistance and lobbying power
of the tobacco interests.
At the NANOA conference in Minnesota in July,

Welby Smith, author of the excellent Orchids of Minnesota,
43
estimated that thousands of pink lady's-slippers,

Cypripedium acaule, are illicitly dug and exported every
year from Minnesota alone. In the New Hampshire
area, I have personally seen one of the best stations of
pink lady's-slippers pock-marked with several
hundred shovel-holes. The same can be said of the
yellow and showy lady's-slipper, C. parviflorum and C.
reginae, in just the past couple of years. One of these
perpetrators offered the rationalization that this was
merely selective digging which benefits the plants by
thinning them out! Sadly, even in the rare case of
apprehension, the thieves are slapped with a minimal
fine—as in the Upper Michigan case which made the
newspapers a few years ago, where the thieves dug
hundreds of ram's-head lady's-slippers, C. arietinum,
for sale in Europe.
Next, consider the U.S. Forest Service, owners of

more than 200 million acres of United States forests.
Most of us assume the agency protects the forests with
judicious, sustained cutting, as was its original intention.
But, with multiple use in effect, that now means mining,
gas and oil extraction, ski developments, ORV's and lots
of logging, to name a few of those uses that now are
taking place in the forests. In order to harvest the logs,
roads are built, almost 400,000 miles of roads; eight
times more roads than the total miles in the US
interstate highway system. Furthermore, the service is
allowing the "harvest" of trees at a greater rate than
their replacement. At a price 50 percent less than the
cost of building these roads, the Forest Service is losing
44
an average of $240 million dollars a year ($70 million in

1996.)
Most of us know what has to be done on a

global, national, state and local level. But it takes time
and effort on our part—we must be consistent and
persistent. Begin by not supporting nurseries and garden
centers that dig wild plants, hold them for a couple of
years and advertise them as nursery-grown. The law
allows this deception. Instead, buy from those who
propagate from seed or division. There are increasing
numbers out there, many of whom are members of this
organization and some who were participants in the
recent Minnesota meeting, people like Bill Steele of
Spangle Creek Labs (Cypripediums), who recently moved
to Minnesota from Washington.
Writing letters can be effective when they are

written in your own words. Write to garden clubs and
other organizations to nudge them into more activist
positions. Write to companies and businesses that
pollute and otherwise harm the environment. Inform
governments and legislators at all levels who ignore the
environment or do not properly fund the programs that
do work (the National Park Service, Endangered
Species Act, and the Clean Air and Water Act, for a few
examples). Electing environmentally responsible
candidates at all levels is one of the most important
things we can all do. Equally important, don't forget
the thank-you letters—and support—to those
individuals and companies trying to do the right thing.
Many companies and property owners, when alerted to
45
either sensitive areas or rare plants on their property, are

often cooperative and willing to help in one way or
another.
My particular favorite is membership in The

Nature Conservancy, the only conservation organization
in the world, perhaps, whose primary purpose is saving
as much of the remaining great places in the world by
buying them. It is also the highest rated conservation
organization in the United States in terms of giving the
most "bang for the buck" or return on the money that
you contribute. But take your pick. There are many
other good organizations that need your financial
support.
These are some of the things we all can do. The

biggest problem, as I see it, however, is just doing it,
isn't it? So don't rationalize your apathy, get involved for
all our sakes.
Philip E. Keenan, 31 Hillcrest Dr., Dover, NH 03820

Phil is a frequent author for this Journal and most recently
of Wild Orchids Across North America published in 1998 by Timber
Press.
Sources for some of the figures cited in Philip's article
include:
National Park Service, U.S. Forest Service, the National
Audubon Society and the Nature Conservancy.
The Enduring Forests, 1996. Ruth Kirk, editor,
Mountaineers Publishing;
North Woods, 1987. Peter Marchand, AMC Publisher;
Nature Conservancy, Nov.- Dec.1998;
Debbie Sease, legislative director, Sierra Club. And many,
many more sources.
46
MARK YOUR CALENDARS NOW !!!!
5TH NORTH AMERICAN NATIVE

ORCHID CONFERENCE
JULY 16-20, 2000
OLYMPIC NATIONAL PARK,
WASHINGTON
and a special field trip east of Seattle to see
Platanthera chorisiana
MORE DETAILS IN THE JUNE ISSUE
47
Empiricist: INTUITIVENESS
INTUITIVENESS
The Slow Empiricist
Have you ever wondered why some orchid

explorers seem to have fantastic results with their orchid
expeditions? Many novice orchidists and not a few
older, more experienced enthusiasts have wondered
how these successful individuals have such good luck. It
seems as though they are always finding their quarry and
discovering new sites on all their expeditions. I would
like to enlighten and encourage you all, novices and the
less than successful, experienced orchid hunters—
sometimes it is pure luck. I believe, though, that their
successes are more often the result of the "pros" having
spent considerably more time in their quests and having
garnered more general information about the plants.
Having a background of such information and
experience helps them find the plants, ensuring success
time and time again. Keep these ideas in mind as I take
you through the processes that can help you build up
your success rate when you are out exploring.
I feel that a lot of the "pros’" success is skewed
by their sensitivity to the species they are looking to
capture. This sensitivity includes knowing the kind of
habitat the plants prefer, the general appearance of the
plants (especially what they look like out of bloom), a
knowledge of the blooming times and the effects of
weather and climate on the growth cycles. I suspect that
47
when one is armed with such information, he or she will

intuitively select appropriate places and times to realize
his or her quest.
I have been on exploratory searches for new sites
of a particular orchid with such an expert. He does his
homework and has a general idea of what the
requirements are for the orchid in question, such as
expected blooming times, before he starts. This expert
was studying the genus Spiranthes and S. casei in
particular for his Master's thesis. He had been closely
monitoring several sites in northern New Hampshire
for about ten years as part of his study. The sheer length
of time shows you how dedicated the expert can be—
this dedication provides a very good indicator of his
success rate at finding these particular orchids.
He wanted to expand his research to include
other populations to see if his hypothesis about habitat
and characteristics of the individual plants held true.
Among the information about S. casei sites, he found
information about locations in Pennsylvania and New
York State, as well as in Maine and Wisconsin. Using
his extensive knowledge of persons who might know
about the sites he was successful in locating information
about the plants in Pennsylvania and New York. Having
such a network of other enthusiasts to call upon should
help anyone be more successful in locating the plants. It
is especially important when you are traveling in
uncharted territory.
By following their directions to the general areas
for the plants we were able to locate the sites. This was
more easily accomplished because he already had a good
understanding of where these orchids preferred to
48
grow. Since he was familiar with the habitats in New

Hampshire, likely spots that imitated the New
Hampshire sites drew our attention as we traveled
through the new areas. Not only were we able to locate
the described sites, but we also found several large
populations in nearby territory.
As a further example of learning about habitats,
when no expert was found who knew of possible
Spiranthes casei sites in Maine, my companion consulted
the geological maps of the areas in northern Maine. He
was looking for pockets of a certain type of soil
composition in which schist occurs in which the
Spiranthes also tended to thrive in New Hampshire. We
drove several hours to these locations determined by
the maps, areas that neither he nor I had ever been to
before, and sure enough, we soon began to find the
orchid in full bloom.
It takes experience to become more intuitive
about your subject, but as you can see from the
preceding examples, by familiarizing yourself with the
plants and their specifics you can have greater success.
If you arm yourself beforehand, you can have a more
successful time in the field.
Where do you find the ammunition to
accomplish this feat? I would suggest that you avail
yourself of some more experienced person's knowledge.
A good place to start is through enrolling in some
program that will put you in the field with
knowledgeable leaders to acquaint you with the habitats
and the actual plants you want to find. Places like the
New England Wild Flower Society in Framingham,
Massachusetts have excellent educational programs that
49
put you in just such situations. They have instructors

who present information in classroom situations and
extend it through field trip experiences. Their field trips
can range all over New England and surrounding areas.
The Society does not strictly limit their program to
orchids but embraces the broader world of wildflowers,
of which native orchids are an integral part. You can get
more information by contacting the Education
Department at NEWFS.
If you want strictly orchids you will have to try
other types of programs or find people who would be
willing to assist you because they have an abiding love
for orchids. You can find people like this in many
orchid societies in North America. Many people in
these societies, however, are only interested in growing
and hybridizing tropical species. There are usually some
members who like to explore for the native species and,
as you get acquainted, you can learn from them.
Some native plant societies know the native
orchids in their area and can help you become more
familiar with the species. It just takes a little looking into
the possibilities in your area to come up with resources.
The Internet may provide help if you look up orchids in
its listings. Since you already subscribe to or read the
North American Native Orchid Journal, you have more
resources in it to increase your native abilities.
As you become more knowledgeable with your
native orchids you will begin to build up an inner idea
of what to be on the lookout for, and when to get
serious in your search for the particular species you
want to locate. If you have had an expedition with
someone who has taken you to see the plants, and you
50
have actually seen the mature plants, those images will

help you when you explore on your own. If you have
never seen the plant in the wild and you are relying only
on photographs you have seen in books and field guides
you will probably have a harder time locating the plants.
From my own experiences, I have not been able to
locate a particular orchid I have never seen in the wild
because it looks different from what I imagined it was
going to look like from the photo I had seen. I also
have found that once I have located one plant, and have
a visual specimen in front of me, I can find more in the
same habitat that, just seconds before I found the initial
orchid, appeared to be devoid of any of them.
To recap my premises, I cannot encourage you
enough, novice and more experienced alike, to fill your
mind with as much information as you can gather about
the plants you are hoping to find, because it will help
you begin to have a "feel" for the orchid. As you
experience a particular habitat for the orchid you will
suddenly realize that the new habitat you are traipsing
through is exactly like such-and-such habitat that you
encountered on last month's expeditions when you had
the orchid in bloom. Your mind will recall the orchids
you saw and you will intuitively look for evidence of the
plants in this new area. Your search may be successful
by finding the orchids in this new site.
The best explorers also hold in their mind the
various forms that the plants take in their growth cycles.
If you know what the plant looks like in rosette or in
fruit, not only in flower, it won't matter when you come
across new, likely habitat. You will be more apt to find
the orchids because you can identify them in their early
51
stages (rosettes) or past prime (in fruit) or anywhere else

in their growth cycle. Of course there is much more that
goes into knowing a good habitat for orchids than just
looks, but as you gain experience you should begin to
develop a "feel" for a good site from observing the
surroundings in which you find yourself.
I seem to have been debunking the theory that
you can be intuitive and find your orchids, because I
have been pointing out that you need to gain as much
information and experience with the orchids as you can
in order to be successful. I think there are people who
have spent considerable time learning and exploring and
I also feel that they have internalized this information
and experience so that it naturally surfaces when they
may not realize it, hence, intuitiveness. I exhort the
novice and the less successful ‘older hand’ to immerse
themselves in the opportunities that abound for them as
outlined in this column. If they are willing to expend a
little effort in these directions, they will probably
become intuitive and then, magically, experience the
thrills of locating orchids on their own
The Slow Empiricist
52
Empiricist:
Argue: INTUITIVENESS
POLLINATION BIOLOGY
POLLINATION BIOLOGY IN SOME

MEMBERS OF THE YELLOW-FRINGED
ORCHID COMPLEX
Part 2. Breeding systems, factors
contributing to reproductive success in the
orange fringed orchid, Platanthera ciliaris
(L.) Lindley and the white fringed orchid, P.
blephariglottis (Willdenow) Lindley, and
pollinator driven morphological divergence
in P. ciliaris
Charles L. Argue
Part one on the yellow-fringed orchid complex

(Argue, 1998) reviewed recent data on floral
morphology, pollinators, and pollination mechanisms,
chiefly for the white fringed orchid, Platanthera
blephariglottis and the orange fringed, P. ciliaris.
Additional data on pollinators of P. ciliaris were collected
by Dr. Katharine B. Gregg and her student Daniel
Frame from West Virginia Wesleyan College (personal
communication) and published, in part, in abstract form
(Gregg, 1981, 1984; Frame & Gregg, 1981). The list of
pollinators from their study site in northcentral West
Virginia is similar to lists compiled from Michigan
(Smith & Snow, 1976) and North Carolina (Robertson
53
Empiricist:
POLLINATION BIOLOGY
& Wyatt, 1990a,b) except that the primary vector at the

West Virginia site was Papilio glaucus L. (tiger
swallowtail), rather than P. troilus L. (spicebush
swallowtail), and a previously unreported pollinator of
secondary importance, P. polyxenes Scudder (eastern
black swallowtail), was implicated (Frame and Gregg,
1981). The present account provides a summary review
of selected topics from the recent literature on
pollination ecotypes, breeding systems, and
reproductive success in Platanthera blephariglottis and/or
P. ciliaris.
Breeding system--Self-pollination and pollinator

exclusion experiments have revealed that although
Platanthera ciliaris and P. blephariglottis are self-compatible,
they are not autogamous (Faegri & van der Pijl, 1979):
insects are required for pollination (Smith & Snow,
1976; Gregg, 1981; Cole & Firmage, 1984; Robertson &
Wyatt, 1990b).
Experimental transfer of pollen within a single

flower or among flowers on the same raceme
(geitonogamy) both produced 100% seed set in
Platanthera blephariglottis (Smith & Snow, 1976). Smith
and Snow (1976) observed butterflies repeatedly
probing the same flowers of this species and suggested
that removal of a pollinarium on one probing might on
subsequent attempts lead to self-pollination. Based on
these observations and van der Pijl and Dodson’s (1966)
view that pollination mechanisms favor outcrossing
(xenogamy) in most orchids, Smith and Snow (1976)
speculated that seeds of P. blephariglottis are probably the
54
Empiricist:
POLLINATION BIOLOGY
product of geitonogamy and xenogamy and considered

that the former may be more common since many
pollinators visit several flowers on a raceme before
departing.
However, as noted in part one of this review, the

pollinarium of Platanthera blephariglottis usually does not
complete its rotation within the mean time spent by
visitors on a single inflorescence (Cole and Firmage,
1984). This circumstance may provide a mechanism that
reduces the incidence of self-pollination in this species
(meaning either pollinator-induced self-pollination
within a single flower or geitonogamy). Cole and
Firmage (1984) argue that behavior of the pollinators
(see below) and the pollination mechanism in P.
blephariglottis favor outcrossing. Of course, even
assuming that rotation of the caudicle must always
precede pollination, selfing might occur if a pollinator
carrying pollinaria from a given inflorescence returned
to the same inflorescence or if it remained on the same
inflorescence long enough for the rotation to be
completed (Cole & Firmage, 1984).
Cole and Firmage (1984) found that pollinators

usually visit a small number of flowers on several
successive inflorescences of Platanthera blephariglottis
interspersed with rest periods of varying duration. They
measured the amount of nectar remaining after one to a
few pollinator visits and found it differed very little
from that in unvisited flowers. Thus, ample reward
remained to promote the visits of subsequent
pollinators.
55
Empiricist:
POLLINATION BIOLOGY
Pollinators moved an average of about 3 meters

between inflorescences of Platanthera blephariglottis (Cole
& Firmage, 1984). The distance moved and the number
of inflorescences visited in a particular area were related
to the size of the pollinator. Small skippers moved
smaller distances and visited more inflorescences per
unit area than larger Lepidoptera (Cole & Firmage,
1984). This plus territorial behavior of the silverspotted
skipper, Epargyreus clarus Crammer, in areas with high
orchid density (leading to exclusion or reduction in the
number of large butterflies) may result in higher capsule
production (Cole & Firmage, 1984) and, presumably, a
spatial restriction of the pollen component of gene flow
(e.g., Beattie, 1978) in such areas compared to sites with
lower orchid density. Larger butterflies (e.g., Speyeria
cybele Fabricius [great spangled fritillary] and Danaus
plexippus L. [monarch]) behaved differently, landing on
several inflorescences at one location and then moving
10 to 15 meters to another inflorescence (Cole &
Firmage, 1984). Pollinatotors also moved between the
orchid species. Natural hybrids of P. blephariglottis and P.
ciliaris have often been reported (e.g., Hardin, 1961;
Case, 1964; Smith & Snow, 1976; Folsom, 1984), and a
close genetic relationship between these taxa has been
documented (Cowden, 1993, 1998).
Robertson and Wyatt (1990b) discussed factors

favoring outcrossing in Platanthera ciliaris. The flowers
on a single plant vary in the amount of nectar they
produce. Such variability was considered a factor in the
movement of pollinators among tropical mass-flowering
56
Empiricist:
POLLINATION BIOLOGY
trees (Frankie & Haber, 1983) and is thought by

Robertson and Wyatt (1990b) to perhaps play a similar
role in this species. Also according to these authors,
only a few flowers are open at any one time in the
racemes of P. ciliaris. This is believed to result in more
frequent movement of pollinators among plants leading
to a reduction in the amount of geitonogamy
(Robertson & Wyatt, 1990b).
However, Gregg (1983) reported that butterflies

visited from two to 26 flowers of Platanthera ciliaris per
inflorescence in a population where most inflorescences
had between 25 and 35 flowers. Papilio glaucus visited
about three (Frame & Gregg, 1981) to four or five
(Gregg, 1983) with the insertion of pollen occurring in
about one of every six flowers (Frame & Gregg, 1981),
but Papilio polyxenes averaged 13 flowers per
inflorescence during one study year (Gregg, 1983).
As in Platanthera blephariglottis (Smith & Snow,

1984), the pollinaria of P. ciliaris remain attached to a
pollinator for several days (Folsom, 1979), and each
pollinator may carry more than one (Gregg, 1983; Cole
& Firmage, 1984). Robertson and Wyatt (1990b) point
out that the sequential deposition of one to several
massulae on a number of flowers increases cross-
pollination when compared with deposition of the
entire pollinium on a single stigma. Duckett (1983)
found no difference in fruit-set between flowers
pollinated by several massulae and the entire pollinium
in P. lacera, a moth pollinated species. Greenhouse tests
on P. blephariglottis indicated that the pollen remains
57
Empiricist:
POLLINATION BIOLOGY
viable for at least five days (Cole & Firmage, 1981).

Prolonged viability of the pollen and its attachment to a
pollen vector for an extended time period favor
outcrossing and an increase in neighborhood size
through reduction of inbreeding among closely adjacent
plants (Grant, 1977; Cole & Firmage, 1984).
Reproductive success--A number of factors have been

identified which are capable of influencing general
reproductive success or fitness as measured by fruit set.
These include the predation of the fruit and seed, the
weather, the availability and allocation of energy
resources to growth and repair or to flowering, and the
availability of pollinators along with the plant’s ability to
attract and utilize them (Janzen, 1971; Udovic, 1981;
Willson & Price, 1977; Stephenson, 1981; Wyatt, 1982;
Cole & Firmage, 1984).
In Platanthera blephariglottis, Cole and Firmage

(1984) found capsule predation to be uncommon.
However, early frost, occurring in two of the three years
of their study, prevented capsule development in young
flowers of late blooming plants.
These authors also reported no abortion of

capsules in Platanthera blephariglottis, abortion being a
feature associated with resource limitation in other
species (e.g., Willson & Price, 1977; Schemske et al.,
1978; Stephenson, 1981). Rather, they considered
capsule production in P. blephariglottis to be limited by
the availability of pollinators, an interpretation based on
five sets of observations.
58
Empiricist:
POLLINATION BIOLOGY
First, artificial pollination markedly increased the

percentage capsule set compared to control plants.
Ninety eight to 100% of artificially selfed and
outcrossed plants regularly set capsules compared to a
mean of 62.4% in controls. Furthermore, the resulting
capsules were well developed whether from small or
large (more than 40 flowers) inflorescences (Cole &
Firmage, 1984).
Second, a higher percentage of fruit set was

observed in years when relatively few inflorescences
were produced (Cole & Firmage, 1984). Cole and
Firmage (1984) found that the number of inflorescences
in Platanthera blephariglottis increased to a point beyond
which the frequency of pollinator visits to each plant
dropped enough to reduce the percentage of fruit set.
This is consistent with the observation that the year of
their study which produced the greatest number of
flowering plants had a significantly lower percentage of
capsule set. In addition, the same year saw an increase in
the number of plants which set no capsules and a
decrease in the number of plants with 100% capsule set.
When the number of plants with 100% capsule set was
examined over the 3 year period, it suggested that a
decrease in the number of inflorescences was correlated
with an increase in the number of pollinators visiting
each plant (Cole & Firmage, 1984).
Third, a lower percentage of fruit set often

occurred in plants of Platanthera blephariglottis with large
inflorescences (Cole & Firmage, 1984). As inflorescence
59
Empiricist:
POLLINATION BIOLOGY
size increased there was a corresponding increase in the

number of capsules up to an average of 10 to 11 per
plant (Cole & Firmage, 1984). Larger inflorescences
generally produced no mean additional increase in
capsule numbers (Cole & Firmage, 1984). Ten to 11
capsules matches the average capsule number obtained
from inflorescences having the modal number of
flowers (11 to 13). Cole and Firmage (1984) suggest that
this number is fixed by natural selection, as found for
some other species (Willson, Miller, & Rathcke, 1979).
Larger inflorescences presumably did not usually
produce any mean additional increase in capsule
numbers because visitors did not increase the number
of flowers visited per inflorescence with increasing
inflorescence size, and pollinator limitation generally
restricted the number of visits per inflorescence.
Orchids can produce a large number of seeds

following only a single pollinator visit, but few seeds
and seedlings survive, and considerable time is required
to reach reproductive maturity (Cole & Firmage, 1984).
Any increase in the number of progeny produced by
each plant through an increase in the number of
pollinator visits would improve fitness (Cole & Firmage,
1984). Cole and Firmage (1984) demonstrated that large
inflorescences of Platanthera blephariglottis, those with
more than 20 flowers, produce a higher percentage of
capsule set only with repeated pollinator visits. As noted
earlier, they also showed that pollinators remove only a
small percentage of the nectar from the nectar spur on
any one visit. The flowers thus remain attractive to
pollinators and repeat visits were observed. In P.
60
Empiricist:
POLLINATION BIOLOGY
blephariglottis the flowers continue to accumulate nectar

throughout the blooming period so that the older
flowers, those near the bottom of the inflorescence,
contain the most nectar. Cole and Firmage (1984)
believe that this represents an evolutionary adaptation
to promote additional visits prior to senescence of the
older flowers. Such an adaptation should improve
capsule set, particularly if the additional visits follow the
opening of the younger flowers on the inflorescence
(Cole & Firmage, 1984).
Multiple pollinator visits to large inflorescences

are indeed sometimes significant. In favorable years
when pollinators are sufficient, the weather is good, and
resources are abundant, large inflorescences in
Platanthera blephariglottis can improve fitness by
producing an abnormally high number of seeds (Cole
and Firmage, 1984). At the same time, increase in
inflorescence size also provides for higher pollen
production (Cole & Firmage, 1984). Large
inflorescences in P. blephariglottis may also improve
chances for increased fruit set by prolonging the
blooming period (Cole & Firmage, 1984). Stephenson
(1979) and Udovic (1981) consider that large
inflorescences may increase the frequency of pollinator
visits in years when pollinators are scarce. According to
Cole and Firmage (1984) such an interpretation applied
to P. blephariglottis only in years when the number of
plants competing for pollinators was relatively low.
Fourth, Platanthera blephariglottis bloomed late in

the season in the Maine study area and, unlike P. ciliaris
61
Empiricist:
POLLINATION BIOLOGY
in South Carolina (Robertson & Wyatt, 1990a), usually

lacked competition from other plants for its pollinators
(Cole & Firmage, 1984). Comparable sequential patterns
of flowering in bog communities have been related to
limited availability of pollinators (Judd, 1958; Pojar,
1974; Reader, 1975; Heinrich, 1976), and Cole and
Firmage (1984) propose a similar interpretation for P.
blephariglottis.
Finally, Cole and Firmage (1984) gathered data

for 6 to 7 hours per day on 70% of the days when
Platanthera blephariglottis was blooming. No pollinators
were seen on many days, and when present they were
usually sparse (Cole & Firmage, 1984).
Thus, according to Cole and Firmage (1984)

reproduction of Platanthera blephariglottis is limited by the
availability of pollinators and sometimes by bad
weather. Individual plants nevertheless increase their
chance of being visited by producing flowers that
secrete nectar continuously and remain receptive for as
long as 10 days on sequentially flowering inflorescences
that remain in bloom for about two weeks (Willson &
Rathcke, 1974; Cole & Firmage, 1984). High capsule set
can result from just a few pollinator visits over the
course of these two weeks (Cole & Firmage, 1984). In
addition, each plant often blooms over a period of
several years with intervening years of vegetative growth
(Cole & Firmage, 1984). These factors enhance the
reproductive success of individual plants of P.
blephariglottis despite scarce or inactive pollinators and
prolonged inclement weather.
62
Empiricist:
POLLINATION BIOLOGY
As in Platanthera blephariglottis, the number of

capsules produced by P. ciliaris appears to be limited by
pollinators. Robertson and Wyatt (1990b) found that
resources are not limiting and that a higher proportion
of hand pollinated plants set fruit than did plants in the
natural population in three out of four cases. Effective
pollinator activity in disjunct coastal-plain and mountain
populations was highly correlated with fruit production
at these two sites as measured by the percentage fruit set
and the numbers of fruits produced by each plant
(Robertson & Wyatt, 1990b). Twenty percent less fruit
was produced in the coastal-plain populations in both
study years (1983 and 1984), and these figures closely
paralleled data on effective pollinator activity obtained
by a measure of the percentage of flowers that had
pollinia removed from their anthers or deposited on
their stigmas (Robertson & Wyatt, 1990b). No pollen
massulae had been inserted on the stigma and no
pollinia removed from the anthers in about 19% of the
flowers examined on the coastal plain compared to
about 7% in the mountains (Robertson & Wyatt,
1990a,b). However, as discussed below, these
differences relate less to the overall availability of
pollinators than to their effectiveness as pollen vectors
(Robertson & Wyatt, 1990b). Gregg (1981, 1984)
reported additional factors affecting the reproductive
success of this species in West Virginia where 5% of the
capsules were destroyed by fungal infection, and
another 5% by herbivory.
63
Empiricist:
POLLINATION BIOLOGY
Pollinator driven morphological divergence in P.

ciliaris?--To establish the existence of pollination
ecotypes for geographic populations of a plant species a
number of conditions must be demonstrated. According
to Robertson and Wyatt (1990a), the pollinators and
plant species must have evolved together, and their
habitat must have remained relatively unaltered. In
addition, plant morphology and pollinators must differ
among populations, the characters in question must be
inherited, and mechanisms by which natural selection
has been effected by pollinators in bringing about
divergence in the plant characters must be found.
Robertson and Wyatt (1990a) note that the

populations of Platanthera ciliaris and the pollinators in
their study are native to their respective areas (see
above). The study sites have been logged or mowed, but
the effects of these disturbances in some respects are
said to mimic those of fire which has historically been
of common occurrence in both areas (Robertson &
Wyatt 1990a).
Consistent site differences in floral morphology

were evident between the mountain and coastal-plain
populations of Platanthera ciliaris in the diameter of the
nectary orifice, length of the spur, length of the
labellum, and length of the longest lateral fringe
segment, with the larger value in each case occurring in
plants on the coastal plain (Two-way ANOVA by site
and year, P < 0.001). These differences appear to be real
as confirmed by independent measurements of coastal-
plain populations by Folsom (1979) and supplementary
64
Empiricist:
POLLINATION BIOLOGY
sampling of mountain populations by Robertson and

Wyatt (1990a).
Robertson and Wyatt (1990a) experienced

difficulties in reciprocal transplant experiments due to
lack of flowering or low rates of survival of many of the
transplants attempted. The results obtained imply that
both environmentally induced and genetic variation are
present in the flowers of their study material. Thus, for
example, the length of the nectar spurs in transplants
became longer on the coastal plain and shorter in the
mountains, but never reached the lengths found in the
native populations (Robertson & Wyatt, 1990a).
A significant difference in the length of the

proboscis was observed between the primary pollinators
of the coastal-plain population, where both Papilio
palamedes Drury (palamedes swallowtail) and Phoebis
sennae L. (cloudless sulphur) had proboscises which
averaged 28.7 mm in length (even though they differed
greatly in wingspan), and the mountain population,
where the proboscis length in Papilio troilus averaged
23.3 mm. This difference also held for the less
important pollinators at the two sites (Robertson &
Wyatt, 1990a). In the mountains the proboscis of Battus
philenor L. (pipevine swallowtail) measured 24.9 mm and
that of Papilio glaucus, 19.5 mm. (Robertson & Wyatt,
1990a).
The differences in floral morphology between the

mountain and coastal-plain populations show possible
adaptation to morphological differences between the
65
Empiricist:
POLLINATION BIOLOGY
primary pollen vectors at the two sites (Robertson &

Wyatt, 1990a). The larger lips with larger fringes
reported for flowers of the coastal-plain population may
be adapted to the large Papilio palamedes butterflies of the
coastal plain (Robertson & Wyatt, 1990a). Nectar-spur
length in mountain populations (23.8 mm) and
proboscis length of its primary pollinator P. troilus (23.3
mm) are such that the eyes of the pollinator would
contact the viscidia as the insect probed the spur for
nectar. On the other hand, the proboscises of the two
primary pollinators of the coastal-plain populations, P.
palamedes and Phoebis sennae (both 27.8 mm), are distinctly
longer than the mean spur length of this population
(25.6 mm).
This difference correlates with and may explain

the lower count of pollinaria removed or pollinia
inserted on the stigma that was observed on the coastal
plain, since the pollinator can extract nectar from the
bottom of the spur without its eyes contacting the
viscidia (Robertson & Wyatt, 1990a). The coastal-plain
pollinators usually carried fewer pollinaria and fewer of
them carried any pollinaria at all as compared to pollen
vectors in the mountains (Robertson & Wyatt, 1990a).
They were more apt to be seen visiting the flowers of
other plants, and when observed on P. ciliaris spent less
time and visited fewer plants than mountain pollinators
of this species. The differences in pollination rates were
apparent during both years of the study despite the fact
that the number of pollinator visits in relation to the
number of plants was higher on the coastal plain the
first year (Robertson & Wyatt, 1990a).
66
Empiricist:
POLLINATION BIOLOGY
The difference in the “fit” between pollinators

and orchids at the two sites was, as noted, reflected in
reproductive success as measured by fruit production
(Robertson, 1987). In addition, during one year of the
study there was positive correlation between spur length
and the percentage fruit set on the coastal plain
(Robertson & Wyatt, 1990a). Robertson and Wyatt
(1990a) consider that the coastal-plain and mountain
populations of Platanthera ciliaris represent distinct
pollination ecotypes in which the differences in floral
morphology reflect an evolutionary response to
differences among the principal pollinators at the two
localities. They further suggest that ongoing selection
may be occurring for increased spur-length in the
coastal-plain population in response to the longer
proboscises of the pollinators at that site.
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yellow-fringed orchid complex. North Amer. Native Orchid
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Beattie, A. 1978. Plant-animal interactions affecting gene flow in
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Folsom, J. P. 1979. The true nature of the putative natural hybrid
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______. 1984. A reinterpretation of the status and relationships
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Frankie, G. W. and W. A. Haber. 1983. Why bees move among
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Heinrich, B. 1976. Flowering phenologies: bog, woodland, and

disturbed habitats. Ecology 57: 890-899.
Janzen, D. H. 1971. Seed predation by animals. Ann. Rev. Ecol.
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Judd, W. 1958. Studies of the Byron bog in southwestern
Ontario. II. The succession and duration of blooming
plants. Can. Field Nat. 72: 119-121.
Pojar, J. 1974. Reproductive dynamics of four plant communities
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Reader, R. J. 1975. Competitive relationships of some bog ericads
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Robertson, J. L. 1987. Pollination and reproductive biology of Platanthera
ciliaris in disjunct populations in the southeastern United States.
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______ and R. Wyatt. 1990a. Evidence for pollination ecotypes
in the yellow-fringed orchid, Platanthera ciliaris. Evolution
44: 121-133.
______. 1990b. Reproductive biology of the yellow-fringed
orchid, Platanthera ciliaris. Amer. Jour. Bot. 77: 388-398.
Schemske, D. W., M. F. Willson, M. N. Melampy, L. J Miller, L.
Verner, K. M. Schemske, and L. B. Best. 1978. Flowering
ecology of some spring woodlands herbs. Ecology 59: 351-
366.
Smith, G. R. and G. E. Snow. 1976. Pollination ecology of
Platanthera (Habenaria) ciliaris and P. blephariglottis
(Orchidaceae). Bot. Gaz. 137: 133-140.
Stephenson, A. G. 1979. An evolutionary examination of the
floral display of Catalpa speciosa (Bignoniaceae). Evolution
33: 1200-1209.
______. 1981. Flower and fruit abortion; proximate causes and
ultimate functions. Ann. Rev. Ecol. Syst. 12: 253-279.
Udovic, D. 1981. Determinants of fruit set in Yucca whipplei:
reproductive expenditure vs. pollinator availability.
Oecologia 48: 389-399.
69
Empiricist:
POLLINATION BIOLOGY
van der Pijl, L. and C. H. Dodson. 1966. Orchid flowers: their

pollination and evolution. University of Miami Press, Coral
Gables, FL.
Willson, M. F. and B. J. Rathcke. 1974. Adaptive design of the
floral display in Asclepias syriaca. Amer. Midl. Nat. 92: 47-
57.
______ and P. W. Price. 1977. The evolution of inflorescence
size in Asclepias (Asclepiadaceae). Evolution 31: 495-511.
______, L. J. Miller, and B. J. Rathcke. 1979. Floral display in
Phlox and Geranium: adaptive aspects. Evolution 33: 52-63.
Wyatt, R. 1982. Inflorescence architecture: How flower number,
arrangement, and phenology affect pollination and fruit-
set. Amer. Jour. Bot. 69: 585-594.
70
Book Reviews
The Orchid Thief

Susan Orlean
Random House 1998
ISBN0-679-44739-3
$25.00
Please keep in mind the title and subject of this

book as you read this review. The Orchid Thief 's
subject is John Laroche, who was arrested and
convicted of stealing native Florida orchids from the
Fakahatchee Strand State Preserve several years ago
under the guise of working for the Seminole Indians.
The entire tone for the book is set by the cover

illustration - an upside down Phalenopsis, and the title
page of stylized Cymbidiums (?) and Cattleyas(?) -
neither of which appear to be species and most certainly
are not native to North America, let alone the
Fakahatchee! This lack of attention to detail and
concern for accuracy pervades the entire book.
Although Ms. Orleans narrative is fascinating reading,
she was sorely taken in by many of the people she
interviewed and therefore comes up with some really
unbelievable stories. In addition only about 25% of the
70
book is about the title subject - the remainder is an
indictment of the orchid community of south Florida.
But that is not the concern of my review.
I am concerned with the scientific accuracy in

dealing with the orchids and, more so, the intense
maligning of the spectacular Fakahatchee Strand area.
From the very beginning of misstating the number of
species in the Orchidaceae as 60,000(!) - when in fact it
is more closely 30,000, to her to misconception of what
species are and how they relate to the entire hybrid
scheme, Ms. Orlean totally lacks the basic botanical
research for this book. And it is a book that purports to
be about an intense botanical subject.
From the very basic of botanical etiquette that

one does not name a new species after one's self - a new
species is described not named; and the fact that the
author of the species then chooses what or whom to
name it after, to her lack of real experience in the
Fakahatchee, which she describes as 'green hell'-
primarily because she spent a bare minimum of time
and saw next to nothing as she was 'taken for a ride' in
the swamp goes to prove the lack of substantial
research. Those of use who know the Fakahatchee
know it to be just the opposite - an absolute paradise
with an unending number of exciting and beautiful
species. You may get a little hot or wet, but that is no
price to pay for the result of a day in the swamp. It is
unfortunate that Ms. Orlean felt the only flower of any
beauty in the swamp was the elusive ghost orchid
(Polyradicion lindenii) - even though she never did see one
71
in flower. In fact, her description of a plant of ghost
orchid is an excellent description of one of the other
leafless species - the ribbon orchid, Campylocentrum
pachyrhizum! Had she spent more time in research she
would have known that there are dozens of equally as
beautiful orchids species and other flowers well
distributed throughout the swamp. Her statistics for
species of orchids and those, which are either restricted
or endemic in the swamp, are totally off base. There are
really on three species that have been found only in the
Fakahatchee - Bulbophyllum pachyrhizum, Maxillaria
parviflora and Epidendrum blancheanum. All others may be
very restricted in their distribution or presently only
known from the Fakahatchee, but they have been
recorded from outside of the swamp over the past 100
or so years of botanical exploration in south Florida.
Perhaps the greatest injustice in the book is in the

acknowledgments where Ms. Orlean credits the
American Orchid Society for checking botanical
accuracy - something she so obviously did not choose
to use within the final text.
If you like a fast-paced, sensational narrative -

borrow this book form your local library - but do not
waste $25 expecting an accurate story about the
Fakahatchee and it myriad of exciting orchids.
Paul Martin Brown
COLLINS PHOTO GUIDE

ORCHIDS OF BRITAIN & EUROPE
Pierre Delforge
72
Collins Books Ltd. 1995
ISBN 0-002-20024-4
This is an excellent guide to eastern European orchids

with over 800 photos, detailed plant descriptions, and
habitat requirements for the species. This book has
excellent quality photos throughout. It is a useful guide
to us in the USA for the four Dactylorhiza species found
in Timmons, Ontario, Alaska, and Newfoundland. The
guide lets you look at these species in relation to other
closely allied Dactylorhiza. The book is still in stock on
Amazon.com for $21.00. It is a hardcover 8-inch by 5-
inch format book and an excellent buy for that price.
Mark Larocque
73
LOOKING FORWARD
JUNE 1999
PROCEEDINGS OF THE 4TH ANNUAL

NORTH AMERICAN NATIVE ORCHID
CONFERENCE
PART 1
And more…
74

March 1999 North American Native Orchid Journal

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NORTH AMERICAN

NATIVE ORCHID JOURNAL

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

2000 Membership in the North American Native Orchid Alliance,

The 4th North American Native Orchid

The 5th Annual North American Native Orchid

Paul Martin Brown

RECENT TAXONOMIC AND

Epidendrum conopseum R. Brown vat. conopseum

In March of 1998 a friend was visiting in Florida

colored race with bronze-pink flowers and a scalloped

Platythelys querceticola (Lindley) Garay. SYN:

It was recently brought to my attention by Dr.

subsequently it was reduced to the varietal level and

Triphora trianthophora (Swartz) Rydberg subsp.

A massive colony of the three birds orchid was

pink. A few individuals of forma albidoflava Keenan were

Triphora trianthophora (Swartz) Rydberg subsp.

Forma folius alba et rosea conspeciebus divers a;

Differing from typical Triphora trianthophora subsp.

Spiranthes cernua (Linnaeus) L.C. Richard

All specimens I have examined from a wide

Spiranthes eatonii Ames ex P.M. Brown sp. nov.

In 1905 A.A. Eaton collected extensively during a

lacera, Spiranthes gracilis, Spiranthes lacera var. gracilis,

The plants of Spiranthes eatonii flower from late

Very simply put,

The lip, sepals and pubescence are all distinctive in

easily separated from S. brevilabris by the latter have a

Spiranthes eatonii Ames ex P.M. Brown sp. nov

Habitu, forma inflorescenctiae Spiranthes lacera var. lacera

Plants 15 - 55 cm tall; leaves basal, oblanceolate to 7.5

by the linear-lanceolate leaves rather than narrow-linear

Habitat: dry to damp open pinelands often with Aristida

Representative specimens examined:

Spiranthes eatonii Spiranthes lacera var. lacera

Spiranthes lacera var. gracilis Spiranthes floridana

Sepals creamy-white 3-6 mm long

The type specimen of Spiranthes eatonii with Oakes Ames' annotations on

Spiranthes eatonii Ames ex P.M. Brown

Triphora trianthophora subsp. trianthophora forma rossii

VALIDATION OF THREE NEW

In the North American Native Orchid Journal [4(1):51-

Even though it may appear that those three new

It is true that Higgins [Phytologia 82(5): 376-377.

Therefore, Higgins and I re-publish those three new

Prosthechea boothiana (Lindley) W.E. Higgins var.

Prosthechea cochleata (Linnaeus) W. E. Higgins

Prosthechea cochleata (Linnaeus) W. E. Higgins

Paul Martin Brown.

A MID-SUMMER PIPE DREAM

Field trips promising the probability of seeing

The high elevation of Frankenstein Cliff delayed

About halfway up, there were two pad-leaved

attributable to the altitude and lack of soil. There were

The return down the mountain was almost as

On a clear mild sunny morning starting at the

Having the woods all to ourselves without

On another woodroad heavily canopied with

Finally, a faint path off a branch woodroad over a

No northern terrestrial orchid is more widely

Hybridization and selection of promising clones

I first attempted to cross the eastern fairy slipper,