June 1999 North American Native Orchid Journal

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NORTH AMERICANNATIVE ORCHID JOURNAL

______________________________________

Volume 5 JuneNumber 2 1999a quarterly devoted to the orchids of North America

published by the

NORTH AMERICANNATIVE ORCHID ALLIANCE

* * * * * * *

* * * * * * *IN THIS ISSUE:PROCEEDINGS OF THE 4thANNUAL NORTHAMERICAN NATIVE ORCHID CONFERENCETHE UNPREDICTABLE WEATHERPOLLINATOR OF CLUSTERED LADYS-SLIPPERORCHID IN OREGONA STRATEGY FOR COPING WITH RARITY IN THEORCHID CYPRIPEDIUM FASCICULATUM.and more


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NORTH AMERICAN NATIVEORCHID JOURNAL

(ISSN 1084-7332)published quarterly in

March June September December

by theNORTH AMERICAN NATIVE ORCHID ALLIANCEa group dedicated to the conservation and promotion of our

native orchids

Editor:Paul Martin Brown

Assistant Editor: Nathaniel E. ConardEditorial & Production Assistants:

Philip E. KeenanStan Folsom

Nancy Webb

The Journalwelcomes articles, of any length, of both a scientificand general interest nature relating to the orchids of NorthAmerica. Scientific articles should conform to guidelines such asthose in Lindleyana or Rhodora. General interest articles and notesmay be more informal. Authors may include line drawingsand/or black and white photographs. Color inserts may bearranged. Please send all inquiries or material for publication tothe Editor at PO Box 772121, Ocala, FL 34477-2121 (late May -early Oct. Box 759, Acton, ME 04001-0759).

2000 Membership in theNorth American Native Orchid Alliance,which includes a subscription to theJournal, is $26 per year in theUnited States, $29US in Canada and $32US other foreigncountries. Payment should be sent to Nancy A. Webb, 84 EtnaSt., Brighton, MA 02135-2830. Claims for lost issues or canceledmemberships should be made to the editorial office within 30days.


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NORTH AMERICAN NATIVEORCHID JOURNAL

Volume 5 June

Number 2 1999CONTENTS

NOTES FROM THE EDITOR75

THE UNPREDICTABLE WEATHERThe Slow Empiricist

774TH ANNUAL NORTH AMERICAN NATIVE ORCHID

CONFERENCEPhilip E. Keenan

84

ORCHIDS AT RABENAU CAMPChuck McCartney90

PLATANTHERA HYPERBOREAAND AREAPPRAISAL OF SOME BASIC GREEN

PLATANTHERASCharles J. Sheviak

115ORCHIDS OF MEXICO

Eric A. Christenson142

PHYLOGENY AND BIOGEOGRAPHY OFPOGONIINAE (VANILLOIDEAE: ORCHIDACEAE)

Kenneth M. Cameron151

KEYING OUT A RED MITSUBISHIor the Trials, Travails, and Tears of an Orchid Hunter

Tom Sampliner164


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RECENT TAXONOMIC AND DISTRIBUTIONALNOTES FROM FLORIDA 2.

A Striking New Color Form ofSacoila lanceolata

Paul Martin Brown169HOW IT ALL CAME ABOUT

Stanley N. Folsom174

POLLINATOR OF CLUSTERED LADYS-SLIPPERORCHID, CYPRIPEDIUM

FASCICULATUM, (ORCHIDACEAE) IN OREGONC. S. Ferguson & K. Donham

180A STRATEGY FOR COPING WITH RARITY IN THE

ORCHID CYPRIPEDIUM FASCICULATUM

Penelope A. Latham185

LOOKING FORWARD:September 1999inside back cover

Color Plates:1. p. 197 Habenaria quinqueseta

2 p. 198 Basic Green Platantheras3. p. 199 Sacoila lanceoaltavar. lanceolataformafolsomii

4. p. 200 Cypripedium. fasciculatum

Unless otherwise credited, all drawings in this issue are by Stan Folsom

Unless otherwise credited graphics (charts, maps etc. ) are created by theindividual authors.The opinions expressed in theJournalare those of the authors. Scientific

articles may be subject to peer review and popular articles will be examined forboth accuracy and scientific content.

Volume 5, number 2, pages 75-200; issued June 24, 1999.Copyright 1999 by theNorth American Native Orchid Alliance, Inc.

Cover: Cleistes bifariaby Stan Folsom


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NOTES FROM THE EDITOR

With this the June issue for 1999 I trust the

Journal is now back on track. If you are still missingany issues, please let me know so they can be replaced.Several changes are being made in the Journal - all of

which I truly hope will improve both the quality andservice.

Because of the labor involve, and the lack of anyhelp assembling the Journal, all of the color willappear as the last set of pages in each issue.

Starting with this issue all mailings will be by BookRate rather than by Bulk Mail. We are losing toomany issues with the Bulk Mail and the service hasbeen unreliable.

The subscription rate for the Journal will not change,although the cost of postage will increaseconsiderably. I will make every effort not todecrease the annual size of the Journal. Severalmembers have already offered to include extra withthe renewals to help cover the additional postage -this is most appreciated.

Foreign mailing to Canada and Europe will begreatly improved by sending all of the Journals toone address and then they will be mailed out withthat area from there. We have a member in Canadaand one in England who have offered to assist withthis.


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There are many interesting and exciting articles in thisissue, not all of which are from the 1999 conference. Iam especially pleased with the pair of article onpollinators and rarity of Cypripedium fasciculatum fromOregon. Although they are brief, please take time toread them carefully.Plans are well underway for the 5th Annual North

American Native Conference to be held in the majesticOlympic Mountains and Cascade Range of Washingtonin July of 2000. Registration information is within thisissue.

The September issue will be printed and assembled inlate September when we return to Florida

Paul Martin BrownEditorPO Box 759,

Acton Maine 04001207/[email protected]

after September 15:PO Box 772121

Ocala, Florida 34477352/861-2565
mailto:[email protected]:[email protected]:[email protected]


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Empiricist: THE UNPREDICTABLE WEATHER

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THE UNPREDICTABLE WEATHER

The Slow Empiricist

It seems that the weather patterns change almostas often as years go by. This variability then affects thegrowing and blooming times of all natural floras in agiven area. Controlled conditions in the home orgreenhouse can overcome much of the vagaries that anordinary year or two of weather can bring to theoutdoors. In a controlled situation, you can compensatefor long periods of overcast skies with lightingconditions. Periods of drought or prolonged wet spells

won't affect indoor specimens that have carefullyregulated watering times and amounts. This is not so innature, however, and it can produce a variety of results.

My first year in retirement in Florida in 1997 was

marked by periods of heavy and frequent rain most ofthe winter season. I was beginning to wonder if Floridawas aptly nicknamed, The Sunshine State, we had so muchinclement weather. Many areas were threatened withflooding. The weather people blamed the conditions onEl Nino, and excessive warming of part of the Pacific

waters that seemed to alter the normal rhythms of thejet stream. That was followed by a hot and very dry


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summer, which produced many wild fires that burnedover 700,000 acres of land in the state. The fall seasoncontinued the drought and it has persisted into thespring of 1999. The current weather has been hotterthan normal as well. According to the weather expertsLa Ninahas replaced El Nino and we can expect moredrought conditions until it abates.

How has this impacted the world of nativeorchids? It has been a curious mix of events. The North

American Native Orchid Alliance's annual conferencewas held in Florida in the month of April. Specifically, it was held April 8, 1999 to April 13, 1999. There werefield trips set up to catch as many species in bloom as

possible. The weather was not cooperating. Floridaladies'-tresses, Spiranthes floridana was blooming earlyand going to fruit. In 1998, it was holding bloom wellinto the middle of April. Eaton's'-tresses, Spirantheseatonii, a newly described species1was not even showingitself in the Jennings Forest State Forest were there

were several dozen plants last year. The short-lippedladies'-tresses, S. brevilabris was also blooming earlierbut it at least had several plants that would hold for thegroup that hoped to take photographs. To compound

the problems, many of the blooming plants of theseSpirantheswere smaller than usual.

The prolonged dry spell and the warmer thannormal conditions had pushed the plants into a seemingrush-mode or a non-mode of blooming. Green

1See article by P.M. Brown in March 1999 issue of NANOJ p. 3-15.


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adder's-mouth,Malaxis unifolia, which was abundant in1998, had not even shown itself by April 8, 1999,joining those orchids that chose the non-mode style ofbloom. The search for it at the SR 200 site in Ocalaturned up a new species for that particular area, though.English friends, Mike and Carol Parsons, whileexploring the area found the speckled ladies'-tresses,Cyclopogon cranichoides, blooming in late March. They

were smaller than the ones at Alexander SpringsRecreation Area on the other side of Ocala but they

were there and gave the state a fourth extant for thespecies.

Paul Martin Brown, who organized the

conference, spent many hours scouting the projectedfield trip sites and scoured the territory to find viableplants for the group. Contingency plans changed as theplants followed their own paths for bloom. The groupended up going into southern Georgia to observeSpiranthes eatonii. It was in spike there and just cominginto flower. They were also able to see bearded grass-

pink, Calopogon barbatus and pale grass-pink, pale grass-pink, C. pallidus, along the roadsides on the way to theS. eatoniisite.

Just the week before, in preparation for theconference field trip there, Paul and I were in the

Jennings State Forest scouting for small spreadingpogonia, Cleistes bifaria, and Spiranthes eatonii, when Ifound several plants of the many-flowered grass-pink,C. multiflorus. Two were only in bud. When Paul wentback the following Tuesday, two days before the


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scheduled field trip as a last minute check, they had allgone to fruit.

Weather has certainly played a major factor inman's plans for viewing orchids or for that matter, anyplant. Being flexible helps a person weather the vagariesthat effect flowering time. Unfortunately in this day andage man can do very little to alter weather conditions. Itis probably a bad idea to even venture to try, as natureseems to know what its doing even if we don'tparticularly care for the current status of things. The

wild fires of last year are a good example of how goodcan come from a bad situation. This is not to trivializethe loss of property and income that these fires created

but many Florida orchids are respondant to fire as ameans of flowering. Calopogon multiflorus, Cleistes bifariaand the large spreading pogonia, C. divaricata allappear in greater numbers in areas that have beenburned over in the year previous. In May we found newsites for both species of Cleistes, one is in Cary StateForest and the other in the Raiford WildlifeManagement area, where we had only found C.divaricata last year. Fire, if it is not all consuming, seemsto cut back the competition, open up the forest floor

and allow these plants to flourish.

My best advice is to persist in your quests andadjust to peculiarities of nature that you find yourself in,in any given year. Fretting over a current condition thatis frustrating can be alleviated if you consider that thisgives you an opportunity to revisit this site on a morefavorable year and season. The Slow Empiricist


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many-flowered grass-pinkCalopogon multiflorus


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pale grass-pinkCalopogon pallidus


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small spreading pogoniaCleistes bifaria


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Keenan: 4thANNUAL NORTH AMERICAN NATIVE ORCHIDCONFERENCE

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4TH ANNUAL NORTH AMERICANNATIVE ORCHID CONFERENCE

Philip E. Keenan

Although the University of South Florida in Tampa,Florida hosted the weekend of meetings during the 4th

Annual North American Native Orchid Conference, it would

be fair to say that the four days of field trips ranged allthe way from the Georgia border to the Big Cypressregion of extreme south Florida. As usual at theseannual meetings, Tampa featured some of the biggestnames in the orchid world.

John Beckner of the Orchid IdentificationCenter, Marie Selby Botanical Gardens, led things off

with a provocative talk entitled Florida - the Rules areDifferent Here!, emphasizing some of the extremes of the

state: hurricanes, droughts, fires, "terminal" watersupply situation, over-development, over-population(now the fourth largest state in population!, behind onlyNew York, Texas and California), illegal trade in drugs,plants and people, and the dangers of attempting theFakahatchee Swamp alone. If all this wasn't bad enough,and if the pollinators go!


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Roger Hammer of the Dade County Park Servicefollowed with a slide presentation on the Orchids of SouthFlorida, especially in the same Fakahatchee Strand,featuring a marvelous slide of the incredible night-fragrant epidendrum, Epidendrum nocturnum, and hisfavorite of all, the ghost orchid, Polyradicion lindenii.Charles J. Sheviak of the Biological Survey, New YorkState Museum, attempted to ascertain, explain, andrationalize some of the uncertainties that plaguePlatanthera hyperborea and other green platantheras thatmost of us prefer to just ignore as a "hybrids of what"problem as he so aptly put it.

Larry Zettler, a biologist at Illinois College,

presented an update on the symbiotic seed germinationofeastern prairie fringed orchis, Platanthera leucophaea,yellow fringeless orchis, P. integra, and northern ovalladies'-tresses, Spiranthes ovalisvar. erostellata with hisusual precise graphics and impeccable speech. PaulMartin Brown filled us in on "a new name for an olddiscovery" a reference to Eaton's ladies'-tresses,Spiranthes eatonii, in his usual professional and forthrightmanner.

On Sunday, Eric Christenson, a Ph.D. researchscientist in Florida indulged in a change of pace with anerudite review of the Orchids of Mexico. ChuckMcCartney, former editor of the AOS Bulletin, followed

with an excellent slide show on the Orchids of RabenauCamp featuring everyone's favorite Polyradicion lindenii,the sensational ghostly flying frog of folklore. His threeslides of this remarkable orchid were easily "the best in


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show." Wendell Vaught and Mike Owen, both of theFlorida Park Service, teamed up with a history of theFakahatchee Strand.

Two members presented some of there favoriteswith Dennis Horn showed his slides of a trip with Willand Hazel Caster to Gtland, Sweden and theabundance of orchids to be found there while DennisMaleug shared some of his striking photos of the

various fringed orchids.

Ken Cameron of the New York BotanicalGarden, spoke on his specialty, the Phylogeny andBiogeography of Pogoniinae, the bottomline for which is still

based on educated guesses, in terms of "how did theyget where they are now," for one example, but we arestill trying to put the puzzle together.

The concluding speaker was Robert Dressler who"regaled" us with The Theory and Practice of Classification,relating how several individual plant characters havebeen termed the "best" by many different taxonomistsover the centuries, in determing species, but in hisopinion, a variety of characters are the key in the final

analysis. Incidentally, Dr. Dressler had the best joke ofall the speakers; one scientist to the other: "what do youthink of God?" "Well, he must have had a greatfondness for beetles!" (largest number of species of anyplant or animal --- as we all knew, right?)

Mike Owen, biologist for the Fakahatchee StrandState Preserve deservedly received the 1999 Alliance


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Conservation & Education Award for his meticulousrecord keeping and generous sharing of his knowledgeof the native orchid world with his devoted work at theFakahatchee Strand.

Two days of field trips before the two-dayweekend meetings were followed by two more days onMonday and Tuesday. Because of severe droughtconditions in all of Florida for several weeks prior to thesecond week of April conference dates, the scorecard oforchids in bloom fell short of the norm. Nevertheless,there were several excellent orchids in prime conditionto satisfy most of us, even though it required travelingthe first day up to St.George in Georgia (where the state

line does a little belly flop into Florida just west of Jacksonville). Here we picked up three grass-pinks -bearded grass-pink, Calopogon barbatus, commongrass-pink, C. tuberosus, andpale grass-pink, C. pallidusin the same roadside "bog" which was perfectly dry onthis day. Lesser pitcher plant, Sarracenia minor, the mostcommon pitcher-plant in Florida, and parrot pitcher-plant, S. psittacina, whose leaves lie nearly flat on theground, were bonus plants.

Back in Florida, near Middleburg and Starke weadded the very rare and tiny Florida ladies'-tresses,Spiranthes floridana and Eaton's ladies'-tresses, S.eatonii, the latter reminding of an even smaller southernslender ladies'-tresses Spiranthes gracilis. The premierorchid for me turned out to be the diminutive speckledladies'-tresses, Cyclopogon cranichoides, in peak bloom atthe Alexander Springs Recreation Area in the Ocala


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National Forest. A hand lens made the difference onthis one as it does with so many of our native orchids.

West of Ocala, the short-lipped ladies'-tresses,Spiranthes brevilabris was in good bloom, apparently ononly one road shoulder.

The excessive heat of an early Spring knockedout a remote cemetery site for the bronze green-flyorchid, Epidendrum conopseumvar. mexicanum. But furthersouth near Avon Park, we hit the jackpot with hundredsof primewater spider orchids,Habenaria repens, prolificalong the water-less ditches. In one of the dry ditchesseveral plants of the white-lipped form ofearly ladies'-tresses, S. praecox forma albolabia, were flowering. On

the dry road shoulders above these same ditches werehundreds of grass-leaved ladies'-tresses, Spiranthesvernalis, also in prime condition.

In the Disney Preserve near Kissimmee, wecaught the last of the many-flowered grass-pink,Calopogon mutiflorus, in bloom along with a couple ofrattlesnakes for good measure, putting to good use the

walking sticks we carried on the trip, much to thepuzzlement of the non-believers who passed us by.

Spiranthes praecox - without the green stippling - (formaalbolabia ) waved in the breezes along equally dried uproadsides not too far out of the Disney Preserve.

At Highlands Hammock State Park, we saw thesame colony of jingle bell orchid, Harrisella porrecta,

which Luer used in his classic volume of the nineteenseventies, in the expected fruiting stage, supposedly


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superior to the flowers. A big disappointment for mewas the absence of anySacoila lanceolata, the red ladies'-tresses, which I had remembered on my only other tripto Florida more than forty years before!

Our last day, after a brief stop at the airport bythe visitors center for the Big Cypress Preserve, where

we sawSimpson's grass-pink, Calopogon tuberosusvar.simpsonii, featured the long anticipated FakahatcheeSwamp and everything the folklorist had conjured up inour minds over these many years of anticipation. Butalas, the drought had pretty much dried up even thisbastion of the subaqueous. We did see one dingy starorchid, Epidendrum amphistomum, in full flower with its

dingy coppery flowers. However, rather than harborfeelings of defeat and dismay, we went away, knowingthat the future would surely present an opportunity toexperience this wilderness kingdom in all its glory.

Philip E. Keenan, 31 Hillcrest Dr., Dover, NH. [email protected]

Phil has written frequently for the journal and annually recaps theconferences for us. He is the author of the recently publishedWild Orchids Across North Americapublished by Timber Press.
mailto:[email protected]:[email protected]:[email protected]


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McCartney: ORCHIDS AT RABENAU CAMP

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ORCHIDS AT RABENAU CAMP

Chuck McCartney

In March of 1979, while on a botanical tour inColombia, I met a man named Bill Rabenau and his 16-year-old son, William. They lived in Davie, Florida, a

western suburb of Fort Lauderdale in Broward County,the same county where I lived and worked.

We got to be friends, and in September of thatyear, Bill invited me to spend the weekend at hishunting camp in the Big Cypress Swamp. Although Iam a third-generation South Floridian, I had spentrelatively little time in the Big Cypress when Billextended his invitation, so I wasn't quite sure what toexpect. As it turned out, I found something rather closeto paradise for an amateur botanist/naturalist.

The word "camp" made me envision rather crude

living conditions. But Bill's place had most of thecomforts of home -- minus the annoyance of aconstantly ringing telephone.

But even better than the accommodations was thesetting. Bill's main camp occupied a small oak-shadedtree island surrounded by a rich mosaic of plant


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communities that included pinelands, open glades,cypress forests, and, best of all to my orchid-loving soul,deep, mysterious swamps of pond apple, Annona glabraand pop-ash, Fraxinus caroliniana.

Although this central camp encompassed only 90acres, Bill also owned about 20 acres of mostly pinelandconverted to pasture about a half-mile to the east and50 acres of mixed hammock, pineland and swamp amile to the west of his main property. In my mind, this

whole area came to be known as Rabenau Camp, and Istill use that designation for the region.

Rabenau Camp is in the northeastern Big Cypress

Swamp in Collier County, Florida, near where it comestogether with Broward and Hendry counties. Being atthe edge of the Big Cypress, the area seems to havemore varied plant communities than are encountereddeeper into this vast tract of wilderness making upmuch of the southwestern tip of the Florida Peninsula.

The diversity of habitats makes for a rich and variedflora, and that diversity extends to the orchids of thearea.

In the company of Bill's son, William (who was,like this father, an excellent woodsman), I started toexplore the fascinating area around Rabenau Camp.Later, when William went away to college, I continuedthese explorations on my own.

In the nearly 20 years that I have been botanizingin the area of Rabenau Camp, William and I have


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encountered 29 species of orchids. All of these speciesoccurred in an area of only about six square miles. Thisis more than one-fourth of the orchids reported forFlorida and it represents one of the highestconcentrations of orchid species to be found anyplacein North American north of Mexico. The only possiblecontender for this distinction might be the fabledFakahatchee Strand, which lies along the opposite

western border of the Big Cypress from Rabenau Camp. The orchid list for the Fakahatchee (Austin, undated)numbers 45 taxa (some questionable). But the speciesthere are spread out over a geographic area of morethan 126 square miles. Still, there may be some six-square-mile area within the Fakahatchee that could

harbor as many species as are found in the region ofRabenau Camp.

Of the 29 species of orchids discovered in the vicinity of Rabenau Camp, 11 are epiphytes. Thisnumber has remained constant since our first years ofexploration in the area. However, the number ofterrestrial species discovered keeps growing, with thelatest two species recorded as recently as June 1996 and

June 1997. To date, the number of terrestrial orchid

species observed in the Rabenau Camp area stands at18.

The following is a quick look at the orchidsdiscovered in this fascinating area of southern Florida.

The nomenclature is conservative, with proposed namechanges discussed where necessary:


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THE EPIPHYTES

Epidendrum anceps:A fairly common species ofthe pond apple swamps, where some plants grow to bequite large. A few plants produce reddish-purple ratherthan green leaves. A winter/spring bloomer withhemispherical clusters of small flowers, ranging in colorfrom bronzy brown to green. Reblooms from the sameinflorescence. Epidendrum monographer Eric Hagsaterof Mexico has proposed the name Epidendrumamphistomum for these Florida plants (personalcommunication), but I can't see how he distinguishesour plants from Epidendrum anceps.

Epidendrum difforme: Rare in this sectionof the Big Cypress. Found in cypress swamps andadjacent pond apple/pop-ash sloughs. Blooms in

August and September with umbellate clusters oftranslucent concolor chartreuse flowers. The taxonomyat present is very confused for these plants. Thesegregation into the genus Neolehmannia has prettygenerally been rejected. But Hagsater (1993) says theFlorida plants are different from the type form ofEpidendrum difforme, so he has renamed our plants

Epidendrum floridense.

Epidendrum nocturnum: Relatively common inthe pond apple swamps of the region, with some plantsgrowing to nearly three feet tall. Flowers in fall/winter,producing surprisingly large, although spidery, flowers

with greenish-brown sepals and petals and a glisteningwhite, three-lobed lip with two bright yellow calluses at


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the entrance to the tube formed by the adnation of thelip and column. Typical of many Florida plants of thisspecies, plants at Rabenau Camp most often producecleistogamous flowers, meaning they fertilize themselves

within the maturing bud and go directly to seed withoutever opening. This doesn't seem to be entirely genetic,though, because plants with cleistogamous flowersoccasionally will produce a flower that opens normally.

Epidendrum rigidum: A sprawling rhizomatousminiature species, which sometimes forms largecolonies. Fairly common at Rabenau Camp. It is, intruth, the kind of orchid that only an orchid lover couldlove. The tiny green flowers appear most commonly in

the fall and winter months, although they may be foundsporadically at other times.Encyclia tampensis: The only Encyclia in the

strictest sense verifiably reported for Florida2. Oncecommon in the Rabenau Camp area, where it grew inhabitats ranging from sunny, open forests of pondcypress, Taxodium ascendens, to tree island hammocks andpond apple swamps. Seems less frequent in the areanow, with many dead plants found attached to the trees.Flowers primarily from late spring through summer,

producing flowers with a honey-sweet fragrance, mainlyin the afternoon. Flower color is variable, but mostly

with brownish-yellow sepals and petals and a white lip

2 A specimen ofEncyclia rufacollected by Small from 'near Eau Gallie' BrevardCounty has been found at the New York Botanical Garden. No other recordor report of this species has ever been found for Florida. It must remain a bitof a mystery. PMB


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with a purplish-red spot or stripes in the middle of themidlobe.

Encyclia cochleata: Regularly encountered inthe pond apple swamps, where it blooms mostly in thefall and winter. Flowers typical of the triandrousFlorida mutation, with narrow, twisting greenish-yellowsepals and petals dangling beneath the non-resupinate,yellow-striped purple clamshell lip. There has been lotsof taxonomic juggling with this species. It doesn't fit inthe genus Encyclia in the strictest sense. The latestproposal (Higgins, 1997) is to move it to the unwieldyand unpronounceable genus Prosthechea. But that genus,too, seems a catchall grouping, so the genus Anacheilium

eventually may prevail for the clamshell encyclias.

Polystachya concreta: Fairly common in pondapple swamps and occasionally found in hammocks andcypress forests. The bloom peak seems to be fall, butsome plants flower in the summer. The tiny, non-resupinate flowers are conical and yellowish-green. Thisspecies, now considered pantropical in distribution,produces multiple secondary flower clusters along therachis of the inflorescence, accounting for the generic

name, which means "many spikes."

Harrisella porrecta: One of Florida's three African-angraecoid-related leafless orchid species andone of South Florida's two smallest orchids (the otherbeing the rare Lepanthopsis melanantha from theFakahatchee). A common small-twig epiphyte mostoften seen on cypress trees, but also growing on a


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number of other hosts, ranging from oaks to hog plum, Ximenia americana, to old, woody stems of the large-flowered climbing or swamp aster. Aster carolinianus. Afew tiny gray-green roots radiate from a short central"stem," from which the thread-like inflorescences areborne, mainly from late summer well into autumn. Thetiny yellowish flowers with their saccate nectar spurappear quite complex under sufficient magnification.

The easiest way to search for this diminutive species isto stand under a likely host tree and hunt for orchid-looking roots. But plants are much easier to find infruit rather than in flower. The little globose greencapsules split open at the end to dehisce their seeds.

The walls of the capsules turn an orangy brown as they

mature, making the old capsules the prettiest part of theplant to the naked eye. Because of the shape of theseed capsules, this little orchid recently has acquired the"common name" jingle bell orchid. At least it's acommon name that makes sense (compared to manylisted in Wunderlin, 1998). Some authors (Wunderlin,1998) are using the name Harrisella filiformis for thisorchid, but Dr. James D. Ackerman in Puerto Rico, athorough botanist whose taxonomic judgments are tobe trusted when it comes to orchids, maintains that

Harrisella filiformis, is, in fact, a Campylocentrum speciesdifferent from the monotypic Harrisella porrecta(Ackerman, 1995).

Polyrrhiza lindenii: Florida's legendary ghostorchid. Another of our leafless African angraecoidrelatives. Surprisingly frequent in the pond appleswamps around Rabenau Camp, in contrast to its


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reported rarity elsewhere in and around the Big Cypress. This species is amazing for the relatively large size ofthe flower produced from such minimal plant mass.

The plant consists of a spidery array of fat, gray-greenroots with light green growing tips. These roots radiateoutward from the short central stem, which producesthe wiry inflorescence. In the Rabenau Camp region,this species generally blooms from late June intoSeptember. Because of the large white flower withnocturnal fragrance and a very long nectar spur, thepollinator is assumed to be a large, long-tongued night-flying sphinx moth. Many authors are now using thename Polyradicion lindenii for this species (Wunderlin,1998.) Both names refer to the numerous roots often

produced by these orchids.

Ionopsis utricularioides: The onlyOncidiumrelative found at Rabenau Camp, where it is not verycommon. These little twig epiphytes are most closelyrelated to the West Indian "equitant oncidiums" nowrightly segregated as the genus Tolumnia. The littleplants, sometimes reddish in intense sunlight, are foundmost often on cypress trees, sometimes dangling offtheir host by a few thread-like roots. The panicles of

pretty little white, purple-striped flowers generally areproduced from February through April.

Cyrtopodium punctatum: This is the mostspectacular orchid of Rabenau Camp, considering plantsize and the number of showy flowers produced. It isfound only sparingly in this area. It once may havebeen more frequent in the region, but because of its


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large size and showy bloom displays, collectors havetargeted it for years. Most of the plants seen aroundRabenau Camp have been growing low on cypress treesin fairly sunny areas. It blooms primarily from Februarythrough April. The flowers, to two inches across, arecrowded on panicles with large yellowish-green bractsbarred with reddish brown that mimic the sepals of theflowers. The petals are yellowish with a scattering of afew rust-red spots (accounting for the species epithet,

which refers to puncture wounds). The rather waxy lipis a beautiful reddish mahogany color with a yellowcrest and center to the midlobe.

THE TERRESTRIALS

Spiranthes vernalis:The most common ladies'-tresses orchid at Rabenau Camp, where it is at its bloompeak in February and March. Flowering plants are oftenfound by the dozens on roadsides and in old pasturesand open prairies.

Spiranthes brevilabris: One small colony ofthis rare species (rare for South Florida, at any rate)

persisted for a few years in an area of rough pastureconverted from pineland. First found on March 27,1981, it was blooming at the same time as Spiranthesvernalis, but the flowers seemed different, being smaller,more yellowish and with an obviously shorter lip. Theplant, too, was quite different. Whereas Spiranthesvernalis produces ascending, narrow, rather grass-likeleaves, those of this orchid were broad, rounded, and


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formed a basal rosette, although they were starting tofade at anthesis.

Using Dr. Carlyle A. Luer's distribution maps inThe Native Orchids of Florida (1972) as a guide, I deducedthat this orchid must be what he called Spiranthesbrevilabrisvar.floridana. But the flower description didn'tquite match up. That variety (now called Spiranthes

floridana ) is essentially glabrous. But the flowers andinflorescence of the plants near Rabenau Camp werehairy, as clearly showed in the few rather poor-qualityclose-up color slides I took on February 19, 1982,before I "lost" this colony. Clearly, this must beSpiranthes brevilabris var. brevilabris instead -- and the

Rabenau Camp-area population may be a first recordfor Collier County, Florida3.

The only problem: Since I realized what thisorchid really is and sought to take better photographsand collect a voucher specimen, I have been unable tofind it! I know the precise location, so that is not theproblem. The colony just seems to have "disappeared."I'm hoping that, like isotrias in the North, the plants aresimply "taking a rest" (of many years duration!). But the

area were these plants occurred is becoming overgrown,especially so since cattle were excluded from theproperty. I'm wondering whether, if this pinelandburned (as is a common occurrence with this sort ofhabitat), perhaps this colony might make a

2. This indeed is the first record for Collier County. PMB


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reappearance. Nevertheless, year after year, the searchgoes on ....

Spiranthes laciniata: This species is a near twinof Spiranthes vernalis, with only minor floral detailsseparating the two. But what does separate them in the

vicinity of Rabenau Camp is a marked hiatus of abouttwo months between their bloom times, with Spirantheslaciniatastarting to flower in late May and extending into

July.

Flowering as they do at the start of the rainyseason in South Florida, the plants ofSpiranthes laciniataoften bloom out of a few inches of standing water.

This species is common in the open glades of theregion, often becoming abundant along the edges wherethe glades grade into open cypress forests.

Spiranthes torta: First found in flower onJune 23, 1996, this is the 28th orchid species discoveredin the Rabenau Camp region. It was blooming at thesame time as Spiranthes laciniata, which grew very nearbyin the same rough pasture succeeding to pineland. Butthe center of the lip on flowers of this plant was

greenish, matching the plants of Spiranthes torta as Iknew it from verified populations on Big Pine Key inthe rock pinelands of the Lower Florida Keys. Bycontrast, the lip ofSpiranthes laciniata is yellowish in themiddle. Also, the flowers of this greenish-lipped plantnear Rabenau Camp were smaller, somewhat nodding,

with the lower flowers somewhat separated from theothers in the loosely spiraled raceme. The leaves were


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grass-like, similar to Spiranthes laciniata, and not like PaulMartin Brown's recently described Spiranthes eatonii(Brown, 1999). A specimen was collected anddeposited in the herbarium at the Marie Selby BotanicalGardens in Sarasota, Florida. But there is still somesmall degree of doubt in my mind whether this is trulyS. torta. I will look for these plants in flower again thisyear in the hope of alleviating these lingering doubts.

Spiranthes odorata: The only fall-bloomingladies'-tresses we have found at Rabenau Camp, this oneis the whopper of the genus. Plants can grow to nearlyfour feet tall. Luer (1972) treated it as a robust southern

variant of the northern nodding ladies'-tresses,

Spiranthes cernua, but it is now pretty well accepted as a valid species in its own right. The tall stems of large(well, large for a Spiranthes ) white flowers occur mostfrequently in open cypress forests, often growing out ofseveral inches of standing water, making this one ofFlorida's "aquatic" orchids. The flowers, as the speciesname implies, are quite fragrant, exuding a sweet, citrus-like aroma during the day.

Stenorrhynchos lanceolatum: The only

segregate that has shown up at Rabenau Camp, thisspecies appears to be rare in that area. I have seen itonly once, and a plant collected by William Rabenau at asecond, widely separated station, serves as the voucherfor this species for area. This species is much moreabundant, especially along roadsides, slightly farthernorth in peninsular Florida. Sometimes called the terra-cotta orchid, that "common name" does not do justice


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to the brilliant red-orange color of this species' large,showy flowers (although, of course, color variationsoccur, with even a green-flowered form). Both the

voucher specimen and the plant I photographed on BillRabenau's western 50 acres were in bloom in lateMarch. Dr. Leslie A. Garay (1980), in his rather finesplitting of the red-flowered Spiranthes, called this orchidSacoila lanceolata, and most authors now seem to befollowing that usage.

Ponthieva racemosa: This intriguing littlespiranthoid orchid with the mysterious common nameshadow witch grows on a few tree islands, often nearthe edge, where the island grades down into thesurrounding swamp. Perhaps they prefer this locationbecause of the greater abundance of moisture orsunlight -- or both. However, the first plants of thisspecies recorded for the Rabenau Camp area grewunder cabbage palms (Sabal palmetto ) at the edge of apineland. But, again, it was a fairly moist habitat. Irecently returned to this spot after many years, only tofind the whole understory rooted up by feral hogs.Plants of this species produce basal rosettes of beautiful

velvety green leaves. They always remind me of the

northern goodyeras - minus the beautiful tracery of veins in the leaves. In the Rabenau Camp region, thisdelightful little species produces its intricate white,green-striped, upward-facing flowers primarily in

Januaryand February.


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Zeuxine strateumatica:: This little naturalizedexotic orchid "weed" is found at Rabenau Camp mostlyin disturbed areas, such as roadsides and old pastures.It is native to southeastern Asia and is thought to havecome into Florida with seeds of centipede grass,Eremochloa ophiuroides, imported in the early part of thiscentury for use as a grass for lawns (hence, the ratherprosaic common name lawn orchid applied to theZeuxine ). The orchid apparently liked its new homebecause it is now reported from most of Florida's 67counties, as well as from areas of a few adjacent states.

The little pointed clusters of glistening white flowers with their golden yellow, spade-shaped lips are quitepretty under magnification. They are borne mostly in

the winter months.

Oeceoclades maculata: Called Eulophidiummaculatum when it was first detected in Florida in theearly 1970s, this recent arrival is now so happy that isgrowing like a weed in the southern part of the state.Unfortunately, that proliferation has gotten it intotrouble, earning this species the label "exotic pest plant"from a quasi-governmental group called the Exotic PestPlant Council. Because the group has put this orchid

on their hit list of "bad" plants, well-meaning butmisguided "environmentalists" are ripping this orchidout of the ground wherever they find it and destroyingthe plants. The laugh is on them, though, because

whether this orchid got here by natural means orthrough man's intervention (a point still much debated),Oeceoclades maculata is firmly entrenched in our state's


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flora. It is here to stay, and these environmental "do-gooders" will never eradicate it.

This orchid of African origin is the 29th andlatest species recorded in the vicinity of Rabenau Camp.

A few young plants were discovered in June of 1997 ona tree island that a group of us have to cross to get to adeep swamp harboring a healthy population of ghostorchids, and I wonder whether we inadvertently mayhave spread the seeds ofOeceocladesthere on our muddyboots from other sites where this species is plentiful.No sexually mature plants ofOeceocladeshave turned upon this island yet, but it's only a matter of time until wefind plants in flower.

Elsewhere in South Florida, this species bloomsfrom late summer into autumn. This is the last part ofthe rainy season, and it is theorized (Gonzalez-Diaz and

Ackerman, 1988) that the weight of raindrops hittingthe flowers jars the pollinia loose and onto the flower'sown stigma. Our plants of this species, then, would beself-pollinating, thus accounting for the heavy fruit setand resulting rapid territorial expansion of this orchid.

Eulophia alta: Related to Oeceoclades maculataand, like it, a plant of African origin that is now

widespread in the Western Hemisphere, this species isfairly common at Rabenau Camp, growing in disturbedareas, such as roadsides, was well as in open naturalhabitats. The sterile plants resemble palm seedlings,accounting for the common name wild coco (as incoconut).


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This fall-blooming species is one of our showiestnative terrestrial orchids, producing inflorescences to sixfeet tall with dozens of flowers. The species epithet, infact, means "tall." The three sepals, distinctivelypointing upward, range in color from greenish to honeygold to mahogany. The lateral petals are usuallysimilarly colored and converge with the crested lip toform a tunnel, presumably for pollinator alignment.

The lip can be purplish to pure white in color (the lattercolor form being extremely rare).

Triphora gentianoides: Aesthetically one of theleast appealing orchids of the Rabenau Camp area, this

species was first discovered there in early July of 1984on the same tree island where we later found Oeceocladesmaculata. Sizable Triphoracolonies persist on this island,and, in the past, a small colony was found on a nearbyisland, but that spot hasn't been revisited in a number ofyears. Although this species is frequent in yards and themulch around landscape plantings at spots along thesoutheast Florida coast, the 1984 collection of thisspecies on the tree island near Rabenau Camp mayrepresent the first record ofTriphora gentianoidesfor

Collier County Florida4.

Malaxis spicata: This delightful miniatureproduces the second smallest flowers of any orchidspecies encountered in the vicinity of Rabenau Camp.

To call this species a miniature may be a little misleading

4Also a first record for Collier County. PMB


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because robust inflorescences can approach nearly afoot in height at the end of a long period of blooming,although most plants are considerably smaller. Plentifulin the Rabenau Camp region, this species generallygrows in swamps under quasi-epiphytic conditions inthe accumulated detritus onstumps, fallen logs, and the knees and buttressed basesof cypress trees, often just above the high water line. Itblooms primarily in autumn. In the Rabenau Camparea, it also has been found rooted in the sandy humusunder wax myrtles,Myrica cerifera, in areas that have onlya short hydroperiod.

The two leaves and inflorescence emerge from a

small cormous base. The inflorescence continues toelongate as more and more of the tiny greenish tobronzy orange flowers are produced over a long periodof time. The pointed lip is uppermost due to a 360-degree twist. Dr. Robert L. Dressler (1981) calls theflower "hyper-resupinate," and Luer (1972) theorizesthat this unusual occurrence may be due to adaptationto a changing pollinator.

Habenaria quinqueseta: Four true habenarias

occur in Florida (or five if you support the separation ofHabenaria macroceratitis ). Of the three that occur atRabenau Camp, Habenaria quinqueseta is arguably theprettiest. There, it mostly grows in slightly higher, drierareas of South Florida slash pine, Pinus elliottiivar. densa,

where it generally is at its bloom peak from late Augustinto September.


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The flowers, ranging from white to greenish white, are typically shaped for most of the genusHabenaria in the New World. Although the speciesname means "five bristles," referring to what at firstappears to be a lip of five narrow, sweeping segments,the labellum, in fact, is only three-lobed. Theuppermost segment on each side of the "five-lobed" lipis actually the lower lobe of each deeply bilobed lateralpetal. The upper lobe of each lateral petal tucks underthe edge of the concave dorsal sepal to form a hoodover the entrance to the inch-long (or longer) nectarspur growing from the back of the lip. Habenariaquinqueseta is not common at Rabenau Camp, perhapsdue to the limited amount of pineland in the area.

Habenaria odontopetala: This is the mostcommon Habenaria species at the southern end ofFlorida, where it generally blooms later than Habenariaquinqueseta, with flowering starting in midautumn. Someof the geenish-yellow flowers persist nearly through the

winter, although they don't appear to be reproductivelyactive by that time.)

The flowers, at first glance, appear to be

differently formed from those of Habenaria quinqueseta,but on closer examination, the common Habenariaflower pattern is evident. The typically long side lobeson the lip of most habenarias here are reduced to nubsat the top of the slightly broader midlobe. The upperlobe of each lateral petal is broadly toothed at the apex(accounting for the species epithet and the commonname, tooth-petaled rein orchid ), and these lateral


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petal lobes often are more erect, standing out somewhatfrom the concave dorsal sepal. The lower lobe of eachlateral petal, like the side lobes of the lip, is reduced to atiny lobule, sometimes barely noticeable.

At Rabenau Camp, plants ofHabenariaodontopetala can be quite robust in sunny optimumconditions, occasionally growing to four feet tall withinflorescences densely packed with flowers. But in thespecies' more common shady hammock-type habitats,plants are much smaller, producing only scatteredflowers on the inflorescence. Wunderlin (1998) uses thename Habenaria floribundafor this orchid.

Habenaria repens: This is the most commonHabenaria in Florida (and perhaps all of the WesternHemisphere). Ironically, it is found only sparingly insouthern Florida. I have seen it only once in theRabenau Camp region, when I photographed a stemfrom a single colony on November 1, 1980. I havesearched for it in that area ever since, but to no avail.

This species is, however, encountered occasionally inthe Fakahatchee Strand to the west.

The plants seen near Rabenau Camp grew amongpickerelweed, Pontederia cordata, in an open swampy areaat the edge of a tree island far to the south of camp.

That is a clue to the habitat of this species, the mostaquatic of Florida's native orchids. The plants oftengrow and bloom in floating mats of vegetation in lakesand roadside ditches. That habitat, coupled with themyriad little green, typical Habenaria-shaped flowers,


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accounts for the apt common name water spiders.Elsewhere in the state, this species appears to bloomsporadically throughout the year.

Platanthera nivea: This attractive and sweetlyfragrant little orchid is found sparingly in the region ofRabenau Camp, where it grows in open glades, oftennear the edges of cypress forests. All the plants we haveencountered have been well to the southwest of the

main camp, closer to typical Big Cypress habitat.

Called the snowy orchid for good reason (evenits species name alludes to this), the glistening little blue-

white flowers are borne in conical clusters at the tip of a

thin blue-green stem. The lip is uppermost in thisspecies, although this can be hard to detect at firstglance. The dorsal sepal, lowermost in these is slightlybroader and imitates the pollinator "landing pad"function normally performed by the labellum. Thelateral sepals, located where you'd expect the lateralpetals to be, have an odd twist to them so that for mostof their length, you are looking at their reverse side.

The long nectar spur juts straight out horizontallybehind each slower. At the center of the flower is a

yellow spot on the column.

In the Big Cypress, this species seems to have arather restricted bloom period from late May into June.Blooming at the outset of the rainy season, the plantsoften grow out of a few inches of water, much likeSpiranthes laciniata, which flowers at the same time insimilar habitats.


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Dr. Richard P. Wunderlin (1998) has, ratherinexplicably, returned this orchid to the genus Habenaria,

where it is clearly out of place. Even in Platanthera, it isa bit of an oddball, so we eventually may see anothergeneric name for this pretty orchid.

Calopogon tuberosus: Like Platanthera nivea,this orchid is found sparingly in the vicinity of RabenauCamp, usually well to the southwest of camp, where itgrows in open glades. It is well represented in similarhabitats in the heart of the Big Cypress. This is easilyone of our prettiest native terrestrial orchids and is ashame it lost the apt species name pulchellus, which

means "pretty." This is probably the most widespreadorchid of eastern North America, growing fromNewfoundland south to Florida, Cuba and theBahamas, and from the Atlantic westward to beyond theMississippi. Because this orchid is so well known, itdoes not need any further description here. Plants canbe found in flower in the Big Cypress anywhere fromMarch to June. The southern form in these prairies hasbeen called var. simpsoniiby some authors.

Bletia purpurea:A common species at RabenauCamp, where it grows on canal spoils banks, alongroadsides, in old pastures, on floating logs in swamps,and even on cypress knees and the buttressed bases ofcypress trees.

Blooming from winter into spring, it producestall, sometimes branched, inflorescences of pretty pink


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flowers. When the flowers are open, the sepals arearranged in a typical triangular orchid pattern. Thepetals, however, arch over the lip, forming a tube,probably for pollinator alignment, much as in Eulophiaaltain South Florida or Cleistes, Isotriaor Pogoniaspecieselsewhere. The magenta-edged lip has beautiful raisedgolden keels running down much of the length of themidlobe. In some Florida mutations of this species, atleast one of these yellow keels is repeated in the lateralsepals.

Because this tropical orchid is at the northernend of its range in southern Florida and sufficientsuitable pollinators may not be available, many, many

plants of this species in Florida produce cleistogamousflowers, that is, the buds mature and self-pollinate without ever opening to reveal their beauty to the world. What a waste, aesthetically. This shortcut tocolonization may be successful at first. But will thelong-term results be beneficial genetically?

Although he was a good steward of the land, BillRabenau was forced to sell his camp property when theBig Cypress National Preserve expanded its

northeastern borders. He bought new property inadjacent Hendry County, west of the Big Cypress IndianReservation, and built a sumptuous new "camp" in ashady hammock of live oaks, Quercus virginianaandcabbage palms. But it was never the same to him. Heloved his old camp down in "the Cypress," as he calledit, and its loss through the forced buyout broke hisheart. He lived to enjoy his new camp only a short


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while. Bill Rabenau died on April 16, 1991, and thispaper is dedicated to his memory.

After Bill's eviction from Rabenau Camp, Idespaired of ever having the pleasure of botanizingthere again because one of the first things governmentbureaucracies do when they buy public land is deny thepublic access to it -- even though it was bought with thepublic's money. Luckily, that did not occur with theproperty around Rabenau Camp, so in July of 1993, werevisited the area after several years. Since then, I have

visited frequently. Of course, the camp structures aregone. They were bulldozed into a pile and burned. Butthe natural areas still gloriously intact, and I have been

able to add Spiranthes tortaand Oeceoclades maculatato ourlist of orchid species for the region since then.

Also, the Big Cypress National Preserve has beenkind enough to grant to me a limited collecting permitso I can complete my herbarium voucher series for theorchids of Rabenau Camp. Of the 29 species we havediscovered there over the past 20 years, I still needherbarium specimens for just six: Epidendrum difforme,Epidendrum nocturnum, Habenaria repens, Harrisella porrecta,

Oeceoclades maculata and Spiranthes brevilabris. I hope toadd some of these this fall. For the moment, though, Ifear I'll never get my voucher specimens of Habenariarepens and Spiranthes brevilabris, for reasons discussedearlier in this article. All voucher specimens are ondeposit in the herbarium at the Marie Selby BotanicalGardens in Sarasota, Florida.


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And who knows? Perhaps as I continue tosearch for these "missing" voucher specimens, I mayencounter still more orchid species for Rabenau Camp.FOR FURTHER READING:Ackerman, James D. 1995. An Orchid Flora of Puerto Rico and the

Islands. The New York Botanical Garden, Bronx, NewYork.

Austin, Daniel F. (undated). Vascular Plants of Fakahatchee StrandState Preserve. Naples Chapter, Florida Native PlantSociety.

Brown, Paul Martin. 1999. Spiranthes eatoniiAmes ex P.M. Brown:Eaton's Ladies'-Tresses, North American Native OrchidJournal5(1):3-15.

Dressler, Robert L. 1981. The Orchids: Natural History andClassification. Harvard University Press, Cambridge,Massachusetts.

Garay, Leslie A. 1980. A Generic Revision of the Spiranthinae,Botanical Museum Leaflets(Harvard University), 28(4): 277-425

Gonzalez-Diaz, Nancy, and James D. Ackerman. 1988."Pollination, Fruit Set, and Seed Production in the OrchidOeceoclades maculata, Lindleyana3(3):150-155.

Hagsater, Eric. 1993. Epidendrum floridense Hagsater, IconesOrchidacearum2, Plate 133.

Higgins, Wesley E. 1997. A Reconsideration of theGenus.Prosthechea(Orchidaceae). Phytologia, 82(5):370-383.

Luer, Carlyle A. 1972. The Native Orchids of Florida. The NewYork Botanical Garden, Bronx, New York.

McCartney, Chuck. 1981. The Orchids of Rabenau Camp: 1. TheEpiphytes. American Orchid Society Bulletin50(5):527-535.

1981. The Orchids of Rabenau Camp: 2. The Terrestrials.American Orchid Society Bulletin.50(6):653-660.

1986. More Orchids at Rabenau Camp. American Orchid SocietyBulletin, 55(3):232-241.

Wunderlin, Richard P. 1998. Guide to the Vascular Plants of Florida.University of Florida Press, Gainesville.


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Chuck McCartney, 2226 Lincoln St., #3, Hollywood FL 33020Chuck is a prolific writer and explorer of the orchids of theSoutheastern United States. He is a former editor of theAmerican Orchid Society Bulletin and a popular speaker and field

trip leader with the various natural history organizationthroughout Florida.


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PLATANTHERA HYPERBOREAAND AREAPPRAISAL OF GREEN

PLATANTHERAS

Charles J. Sheviak

The American members ofPlatanthera that wereassigned to the genus Limnorchis by Rydberg (1901)

constitute some of the most baffling taxonomic subjectsin our flora. Portions of the group seem to defyreasonable delimitation of species, and it is indeedpossible that no truly acceptable species concept will befound to be applicable to them. Foremost among theproblematical plants are those comprising the so-calledPlatanthera hyperborea - P. dilatata complex. I've beenstudying these plants rather intensively for 20 years, andin a more general way for over 15 before that. I'vestudied populations across most of the range of the

complex from northwest Alaska to the Mexican borderto the Atlantic Coast. Representative specimens havebeen maintained in cultivation for periods ranging toover 15 years. Cytology and pollination have beeninvestigated. A synthesis of the information gained hasresulted in some progress, and a series of publicationson the group is now beginning to appear. The present

Sheviak: PLATANTHERA HYPERBOREAAND A REAPPRAISAL OFGREEN PLATANTHERAS


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paper presents an overview of certain patterns ofvariation that have become apparent.

For 25 years now we've been using Carlyle Luer'streatment (Luer, 1975) of the group as the basicreference for the complex. I t has indeed been veryhelpful, and Luer brought attention to various taxa thathad been all but lost in Correll's earlier treatment thatincluded fewer, more broadly delimited taxa (Correll,1950). Now, with considerably more information andexperience, further revisions are warranted. Luer'sframework is still a sound foundation upon which tobuild, however, and it will be used here as an outline forthe present paper.

I want to introduce the group with its mostunmistakable member, Platanthera dilatata (Pursh) Lindl.ex Beck. This species is distinctive in its brilliantly pure

white flowers; in all other species they are more or lessgreen. Platanthera dilatatafurthermore has a strong clovescent that is characteristic and apparently limited to thisspecies. The lip typically is marked by a broad, roundedbasal dilation, although the degree to which it isexpressed varies; most commonly it is suborbicular to

transversely elliptical. The species is transcontinental inthe north and widespread in the western cordillera. Likemuch of the rest of the group, it is quite variable, andtraditionally three varieties have been recognized.



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In the boreal forest east of the cordillera, rangingsouth through the Great Lakes Region and eastward tothe Coast, is the nominate Platanthera dilatatavar. dilatata.

This variety is distinguished from other varieties by aslender, mostly white spur about the length of the lip.

The viscidia are linear.

From southern Alaska southward through theRocky Mountains to Colorado, we find Platantheradilatata var. albiflora (Cham.) Ledeb. This variety ischaracterized by a short, clavate, generally green spur.

The viscidia are a bit broader than in var. dilatata, linear-oblong to oblong. The lip is also often broader than inthe nominate race, and in Alaska the basal dilation is

sometimes so broadly elaborated that the base of the lipactually bears two forward-directed lobules.

To the west of the Rocky Mountains is Platantheradilatata var. leucostachys (Lindl.) Luer. This variety isdistinguished by a slender white spur markedly longerthan the lip. Often the lip is very slender, sometimesnarrowly lanceolate and undilated, and the viscidia arelinear. Most significantly, in extreme form, such as inthe Sierra Nevada, var. leucostachysis scentless during the

day and fragrant at night.

Platanthera dilatata var. leucostachys is obviouslyspecialized for moth pollination, whereas the shortspurred, diurnally fragrant var. albifloraseems to be


Sheviak: PLATANTHERA HYPERBOREAAND A REAPPRAISAL OFGREEN PLATANTHERASSheviak: PLATANTHERA HYPERBOREAAND A REAPPRAISAL OFGREEN PLATANTHERAS







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specialized for diurnal, shorter-tongued insects. Theshort, fat spurs of the latter variety reach their greatestreduction in the southern Rockies of Colorado, wherethey are sometimes reduced to a small green sac.Northward they are less reduced. This is a recurringtheme in the group: local specialization and variation.

Although in extreme form Platanthera dilatatavar.leucostachys is far-western, plants referable to it rangeeastward across the Great Basin to western Utah. I say'referable', because these more eastern plants often don'texhibit the pronounced nocturnal fragrance, it beingmore variably and weakly expressed. Indeed, eastwardfrom the Sierra the plants become less distinctive. For

instance, the Ruby Range in northeastern Nevadasupports the species in moist subalpine meadows andalong streams at lower elevations. Spur length in thesepopulations is variable, and many plants wouldarbitrarily be assigned on this basis to P. dilatatavar.ideally, whereas others are referable to var. leucostachys.

Varieties leucostachysand albiflora therefore appear to begeographically variable, reaching extreme form inrestricted geographic areas. In fact, northward thesituation becomes quite complex.

In interior British Columbia most plants bearspurs that are short, stout, and green when the flowersopen. In cultivation, however, as flowers age the spurscommonly elongate, becoming slender, white, andabout equaling the length of the lip. Varietyalbiflorawas



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described from Unalaska, and indeed, most Alaskanplants seem referable to this variety, with short, fat,green spurs. A few herbarium specimens show spurssuggesting var. dilatata, but it's not clear that that is whatthey are. I've grown a few plants of varietyalbiflorafromthe Kenai Peninsula. These were all representative var.albiflorawhen collected, and that is what they looked likein subsequent years. One year, however, one plantopened its characteristic var. albiflora flowers, and thenafter an interval of a few days, the spurs abruptlyelongated until they were slender and markedly longerthan the lip. Essentially the plant changed from var.albiflorato var. leucostachys. How then do we interpret thelong-spurred plants from the Alaskan Panhandle, where

var. leucostachysseems to reach its northern range limit?

Basically, it seems that the Northwest is the centerof variability of Platanthera dilatata, and as the speciesranges eastward through the boreal forest, andsouthward down the Rockies and the more westernranges, variability is reduced in response tospecialization for different pollinators. It seems thenthat the recognized varieties of P. dilatataare really justendpoints in a very complex variation pattern. They

have some utility for discussion purposes, but they are very simplistic representations of the underlyingsituation.

The very distinct Platanthera dilatatais therefore



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quite complex, perhaps unexpectedly so. The rest of themembers of the complex are the green-flowered plantsthat I refer to as "basic green platantheras," or "BGP's"for short. These are the plants that are so oftentroublesome, although some are actually very distinctiveand readily determined once they, and theircharacteristics, are recognized.

Platanthera hyperborea (L.) Lindl. has the mostextensive range of any species in the group, rangingfrom northwestern Alaska southward to New Mexicoand southeastward to central Illinois and New England.

Although this appears to be a very nondescript plant, itis actually one of the most distinctive members of the

group. The markedly yellowish lip is rhombic-lanceolatein outline, not at all dilated. The spur is decidedlyclavate and slightly shorter than the lip, but it appearsshorter because of a sharp curve. The column of thisspecies is very distinctive. It is proportionately verysmall, with the anther sacs lying at low angles atop thestigma. Most remarkably, the flowers are self-pollinating: the pollinia fall free of the anther sacs andeventually contact the stigma. Perhaps because of this,across most of its range the flowers are scentless. The

pollinaria are functional, however, and the viscidia areorbicular. None of our other recognized species havesuch a column nor pollination mechanism. Thecombination of column structure, pollinationmechanism, lip color and lip and spur shape, and usuallyscentless flowers combine to make P. hyperborea verydistinctive.



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It is ironic, then, that two of the most distinctiveand well-defined members of the group have been usedto designate the so-called hyperborea-dilatata complex.

The complex is actually composed of other species andplants.

Luer's treatment recognized five varieties ofPlatanthera hyperborea, and they form a useful frameworkfor presentation of some of the remaining plants ofinterest here. Varietyhyperboreahas just been introduced.

Platanthera huronensis(Nutt.) Lindl. has been one ofthe most vaguely-defined concepts in the group. Luertreated it as a variety of P. hyperborea, in keeping withmost earlier workers who recognized it. This plantimmediately differs from P. hyperborea in its whitishgreen flowers. The lip is generally rounded-dilated at thebase, but generally less so than is characteristic of P.dilatata.The spur is slender and about the length of thelip. The flowers are intensely sweetly pungent-scented

with a scent that to me suggests a sweet version of the

laundry detergent isle at the grocery store. The columnis larger and the anther sacs much higher than in P.hyperborea; the flowers are not self-pollinating. The

viscidia are oblong.



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This species has been the source of a lot ofconfusion, stemming in part from its color and thedilation at the base of the lip. It looks like a hybrid ofPlatanthera dilatataand P. hyperborea, and very likely that'show it arose. The species is a tetraploid, and tetraploidspecies in nature are almost always of hybrid origin.

Very likely, then, P. huronensis arose as a P. dilatatahybrid, and Catling and Catling (1997) presented resultsof a numerical analysis indicating that P. hyperboreamightbe the other parent.

The species ranges southward in the cordillerathrough British Columbia to Colorado and Utah, and

across the Great Lakes region to the Atlantic Coast. Theplants vary greatly in stature and can be quite robust,reaching over a meter in height. The density of theinflorescence varies from lax to very dense.

Two other points can be made here. First, in thenortheast this species is commonly confused withputative hybrids of Platanthera dilatata and P. hyperboreaand called P. xmedia (Rydb.) Luer. Secondly, the plantsin western Canada and Alaska that Luer called P.

hyperboreavar. viridiflora(Cham.) Luer are in all respectsreferable to P. huronensis.

The next species I want to discuss is Platantherapurpurascens (Rydb.) Sheviak & Jennings. This is thefourth taxon treated by Luer as a variety ofP. hyperborea.Rydberg's name had been ignored for most of thiscentury, and we owe Luer a lot for introducing the



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plant, which had been quite problematical for manyyears. Again, as BGP's go, P. purpurascensis quite easy torecognize once you learn what to look for. The

vegetative habit is distinctive, with rather short, blunt,abruptly wide-spreading leaves. This spreading leaf habitis a significant feature; it doesn't seem to vary with theamount of exposure to sun and wind. The flowers tendto cluster in fascicles in a long, slender raceme, aparticularly unusual and significant feature. Lips vary inshape from narrowly lanceolate to broadly dilated, andin color from watery greenish yellow to intensely bluishgreen. Sometimes reddish blotches occur on the lip orother floral segments; these are contributed by thecoloring of dense networks of minute veins and

evidently are responsible for the specific epithet.Regardless of the actual color, the intense, wateryquality is characteristic. The spur is variously short andinflated, usually strongly inflated-clavate to saccate andscrotiform. The flowers are strongly and uniquelyscented with a marked musty, semen-like scent.Columns are rather typical of the group, with the rathersmall anther sacs well-elevated above the stigma and therostellum lobes somewhat spreading. The viscidia aretypically orbicular, but vary to oblong. There is quite a

bit of variability in these features, but the uniformlyshort spur and peculiar coloring, abruptly spreadingleaves, and singular scent hold the group together andindeed make it quite obvious.



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I really don't think Platanthera purpurascens hasmuch to do with P. hyperborea, but it shares a fewfeatures with P. strictaLindl. Platanthera stricta is a morenorthern species, ranging from Alaska southward tonorthernmost California and about YellowstoneNational Park. Throughout this range the plant ismarked by short, abruptly spreading leaves. The spur isrelatively short and inflated, and varies from inflated-clavate to very short and saccate. Lips are generallyrather long and linear to merely lanceolate. The color,however, is a flat green, not the watery, deep colorationof P. purpurascens, and the column is very different thanin that species, with prominent, more or less parallelanther sacs. The flowers are, to my nose, virtually

scentless, sometimes with just the slightest spicy quality.Viscidia are orbicular.

Taxa in this group are obviously capable ofconsiderable variation, regardless of how strictly theymight be delimited, and interpretation of limits isconfounded by convergence under similar pollinationpressures. The similar range of spur forms in Platanthera

purpurascens and P. stricta is consequently of uncertainsignificance. Their vegetative habits might be more

significant, but can a single vegetative feature define aspecies? I think that the very different columns of P.

purpurascens and P. strict a, in contrast, lie outside therange of variation seen in other species in the groupand, when taken together with the other differencesbetween them, indicate that, indeed, we are dealing withtwo distinct species.



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That said, I might point out yet another problem.From southern British Columbia to northern California,at least, occur very tall, slender plants with less abruptlyspreading leaves. The spurs on these plants areuniformly inflated-clavate, not at all saccate.

They are similar to those often seen onuppermost portions of the inflorescences of the moretypical Platanthera strict adiscussed above, so they are notunique. However, in addition to these morphologicaldifferences, these slender plants have proven to beexceedingly difficult to cultivate under my conditions,

whereas the more typical form grows reasonably wellfor me. Hence, these two groups may be different at

some level.

That brings us to Luer's Platanthera hyperboreavar.gracilis (Lindl.) Luer, the last of his varieties of thisspecies. He illustrated this variety with a single photofrom Yellowstone. Comparison of Luer's photo withLindley's type at Kew, collected near the coast of BritishColumbia on the border with the end of the Alaskanpanhandle, discloses very different plants. The holotypesheet includes three specimens, all with very narrow

leaves that in one specimen are quite long. The plantsotherwise seem to fall nicely into P. stricta. The leavesevidently have a tendency to spread abruptly. Lips arelong and slender and the spurs are entirely comparableto those of the slender, problematical plants of P. strictamentioned above. I think P.gracilisLindl. is refereable toP. stricta, at least at some level.



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This may dispense with the name, but what wasLuer's plant? His published photo shows a slenderinflorescence, lanceolate lips, and blunt, clavate spurs abit shorter than the lip. I have found entirelycomparable plants in northern Colorado, geographicallynot too far removed from Luer's locality. From whatcan be seen in Luer's photo, the match between the lipand spur of these plants is precise. These plants alsobear 'normal' columns: they are not merely Platantherahyperboreawith slender inflorescences.

After years of looking for, and at, these things inthe field, I've become convinced that the plantsrepresented by Luer's photo are products ofhybridization of Platanthera dilatatavar. albiflora and P.

purpurascens. The plants designatedgracilis Luercommonly occur in complex populations with these twospecies and often with a variety of other peculiar plantsthat exhibit considerable variation in lip shape and inflower color, varying from deep green to dull white.

They are uniformly fragrant, usually with thepronounced sweet-pungent scent of P. huronensis,sometimes a bit more musty as in P. purpurascens.Flowers often tend to be grouped in fascicles. Viscidiaare oblong. Individual plants furthermore may varygreatly from year to year. One plant that I maintained incultivation for some years was essentially identical to theplant in Luer's photo when collected and subsequently



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in cultivation, except for one year when it produced a very dense inflorescence and virtually duplicated P.huronensis in all salient characters. Unlike that species,however, 'this plant, like the others discussed here, wasdiploid. These plants quite obviously constitute hybridsof the two diploid species with which they commonlygrow, and between which they vary.

To date I have found only a single specimen that was not diploid, and this was triploid, as reportedpreviously (Sheviak & Bracht 1998). This plant, growing

with various diploid intermediates and the parentalspecies, suggested Platanthera huronensis and differedfrom that species primarily in a very short spur and an

abruptly dilated lip with an eroded margin. Themorphology and cytology of this plant suggest a hybridof P. huronensis and P. dilatatavar. albiflora, but theformer was not evident at the site. Of course, it mayhave been present but vegetative, or it may haveoccurred there in the past. Indeed, it would have takenonly a pollen grain brought in from another colony tohave given rise to this plant.

Demography, then, is a complicating factor that

contributes to the difficulty of interpreting populations. The plants blooming one season do not necessarilyrepresent the total set of variants present then or in thepast. Indeed, by revisiting populations over periods ofup to fifteen years, I've noted marked changes in



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abundance ofPlatanthera dilatata, P. purpurascens, and theevident hybrids. In some years, colonies which normallysupported a complete range of parentals andintermediates were wholly lacking in blooming plants ofone of the parents or the hybrids. Without such atemporal sample, the total variation pattern and it'ssignificance can easily be missed.

The evidence suggests that the plants identified byLuer as Platanthera hyperboreavar. gracilisare products ofhybridization. If instead they are some distinct entity oftheir own, they are so thoroughly involved inhybridization with other species that it is difficult tojustify maintaining them as distinct. With that in mind,

the situation on Mt. Graham in southeastern Arizona,where P. purpurascens occurs at perhaps itssouthwestemmost station, may be significant. Also onMt. Graham are plants with the same morphology andsweet scent as the Colorado plants discussed above. Ifthese things are hybrids, what are they hybrids of?

There isn't another species on Mt. Graham, noranywhere remotely near it. The nearest P. dilatata is innorthernmost New Mexico. These Arizona plants might

suggest that Luer's plants are indeed a distinct species(or whatever), and in portions of their range they occur

with other species, perhaps hybridizing with them or atany rate being confused with hybrids. On the otherhand, two other explanations are equally plausible. First,these southern populations could represent the effectsof gene flow in the past, when another species waspresent.



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Certainly the Pleistocene provides a readyexplanation: P. dilatata almost certainly occurred in thearea under the cooler, more moist climate that prevailedin the Southwest at that time. Or, secondly, perhaps P.

purpurascens simply varies more than I've realized.Perhaps these indeterminable BGP's, or some of them,represent merely extreme forms ofP.purpurascens.

A population of similar plants in the central SierraNevada Mountains provides further ambiguous butprovocative evidence. I was directed to this site byRonald Coleman, who, following Luer, referred theplants growing there to Platanthera hyperboreavar.gracilis.

Throughout the area P. dilatatavar. leucostachys was a

common plant, but in only one small seep occurredgreen-flowered plants with lanceolate lips, oblongviscidia, and slenderly clavate spurs about the length ofthe lip. The plants appeared very similar to the plantsdiscussed above, except that the spurs were slightlylonger and the flowers wholly scentless. Subsequently incultivation, the lips were longer and proportionatelymore slender, and the spurs proportionately longer andmore pointed. Lip color, too, changed from a medium,flat green to a watery yellowish green.

What are these plants? This colony is not unique,for I have seen similar herbarium material from a fewother sites in the region. At the station I visited, theindeterminable green-flowered plants seemed to behybridizing with Platanthera dilatata var. leucostachys,producing greenish white flowered plants with longerspurs. Platanthera purpurascens is known from a few



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disjunct populations in the same area; its occurrencethere may be significant, or it may be coincidental.Despite the ambiguity, the distribution pattern of theseindeterminable BGP's suggests a plausible origin.

An hypothesis that accommodates all of theproblematical plants resembling the plant figured byLuer as Platanthera hyperboreavar. gracilis rests on onequestion: what is the most generalized flower in thisgroup, one without any particular specializations? It ismedium green with lanceolate lip, clavate spur a bitshorter than the lip, and probably oblong viscidia.Deviation from this norm in any of these features, ascharacterizes the various species in the complex, thus

represents a derived condition. For the most part thisprobably is due to specialization for a particular type ofpollinator, but in some cases it might result merely fromdrift. Anything that disrupts the genetic balance thatunderlies the specializations characterizing the variousspecies is apt to produce a nondescript, basic flower,like the one figured by Luer. This can be hybridization,because genetic recombination may simply erase derivedstates based on homozygous conditions.

Furthermore, the physiographic complexity ofthe cordilleran region provides a mechanism forgenerating considerable variability within species. Moregenerally, then, as species reach their range limits oroccupy habitats outside the norm, the selectivepressures that maintain their distinctive features may belessened or absent, and populations then may drift backto a basic floral form. Hence, across the region,



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different processes acting on different species may havegiven rise to very similar plants.

Considerable progress has been made in ourunderstanding of the so-called Platanthera hyperborea-dilatata complex, and this gives hope that we mayultimately arrive at a satisfactory taxonomy.Nonetheless, the remaining known problems aredaunting, and each year in the field brings forth newones. Whatever the practical taxonomic productsultimately prove to be, however, th

June 1999 North American Native Orchid Journal

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Transcript of June 1999 North American Native Orchid Journal