june 2001 north american native orchid journal

8/8/2019 June 2001 North American Native Orchid Journal

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NORTH AMERICANNATIVE ORCHID JOURNAL____________________________________

Volume 7 JuneNumber 2 2001a quarterly devoted to the orchids of North America

published by the

NORTH AMERICANNATIVE ORCHID ALLIANCE

* * * * * * *

* * * * * * *IN THIS ISSUE:

REPRODUCTIVE BIOLOGY OF THE LADY'S-SLIPPERS.UPDATE ON CORALLORHIZA BENTLEYIBOOKS AND PAMPHLETS ON THE ORCHIDS OFTHE UNITED STATES AND CANADAOUT OF THE LOOP.andmore


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The North American Native Orchid Journal(ISSN 1084-7332) is a publication devoted to promoting interest andknowledge of the native orchids of North America. Alimited number of the print version of each issue of the

Journal are available upon request and electronic versions are available to all interested persons orinstitutions free of charge. The Journalwelcomes articleof any nature that deal with native or introduced

orchids that are found growing wild in North America,primarily north of Mexico, although articles of generalinterest concerning Mexican species will welcome.

Requests for either print or electronic copies should besent to the editor:Paul Martin Brown, 10896 SW 90th Terrace, Ocala, FL34481 or via email at [email protected].


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NORTH AMERICAN NATIVE

ORCHID JOURNAL

Volume 7 JuneNumber 2 2001

CONTENTSNOTES FROM THE EDITOR

115

REPRODUCTIVE BIOLOGY OF THE LADY'S-SLIPPERS.

I. Introduction and the Pink Lady's-slipper,Cypripedium acauleAit.Charles L. Argue

117UPDATE ON CORALLORHIZA BENTLEYI

BENTLEY'S CORALROOTStan Bentley

140BOOKS AND PAMPHLETS ON THE ORCHIDS

OF THE UNITED STATES AND CANADARonald A. Coleman

143OUT OF THE LOOP

The Slow Empiricist154

A NEW ZEALANDER LOOKS AT ORCHIDS INTHE WILD WEST

Ian M. St George159


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THE ARCTIC ORCHIDS OF ADELBERT VON

CHAMISSOMark Nir

169RECENT TAXONOMIC AND

DISTRIBUTIONAL NOTES FROM FLORIDA10.

Paul Martin Brown176

6th ANNUAL NORTH AMERICAN NATIVEORCHID CONFERENCE

181

NEW TAXA AND COMBINATIONSPaul Martin Brown

186Book Reviews:

Native Orchids of Nova ScotiaCarl Munden

187LOOKING FORWARD:

September 2001

189

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

The opinions expressed in theJournalare those of the authors. Scientificarticles may be subject to peer review and popular articles will be examined for

both accuracy and scientific content.Volume 7, number 2, pages 115-189; issued June 31, 2001.

Copyright 2001 by theNorth American Native Orchid Alliance, Inc.Cover: Platanthera hookeriby Stan Folsom


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especially note the extensive bibliography at the end. Itis one of the most complete on the subject I have everseen.

We will be in Maine through early October.Please note the telephone number change below. This

will be a quiet summer for us, as we do not have anytrips planned, although getting ready for the conferencehere in September will be busy enough. Our new book,The Wild Orchids of Florida, is scheduled for release late inthe year so final editing and proofing is well underway.More about that in the next issue.

Paul Martin Brown

Editor

PO Box 772121Ocala, Florida 34477-2121352-861-2565 October - May

PO Box 759Acton, Maine 04001-0759Late May - September207-636-1107

Email: [email protected]


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Argue: REPRODUCTIVE BIOLOGY OF THE LADY'S-SLIPPERS

117

REPRODUCTIVE BIOLOGY OF THELADY'S-SLIPPERS.

II.

Introduction and the Pink Lady's-slipper,Cypripedium acauleAit.

Charles L. Argue

So different are lady's-slippers from otherorchids that botanists sometimes place them in a familyof their own. They are distinguished by a number ofcharacters, but the most conspicuous difference is the

deeply saccate, pouch-like lip of the flower, from whichthe plants take their name.

This lip plays an important role in pollination.Species ofCypripediumhave flowers of a type known astrap or semi-trap blossoms (Pijl and Dodson 1966,Dressler 1981). The flowers temporarily imprison theirinsect pollinators and force them to follow a prescribedsequence of behaviors in order to obtain their release.

An insect of the appropriate size, usually a bee, entersthe lip through the obvious large opening at its top. Theslippery inner surface and the in-folded margins of thelip supposedly prevent it from leaving by the same route(Summerhayes 1951, Proctor & Yeo 1973), but Knoll(1922) and Daumann (1968) have shown that bees areunable to escape from the lip of the European C.calceoluseven after the in-folded margins are cut away.


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Figure 1. Sketches of the flower ofCypripedium acaule. A. Side view

of flower. B. Gynostemium and staminode as seen from

inside the flower. The gynostemium is comprised of theunited style and the filaments of the two fertile anthers.The staminode is modified from a third sterile anther. C.Flower in partial longitudinal section showing path ofbumblebee queen. a = anther, b = bract, e = escape orexit hole, gy = gynostemium, l = labellum, lo = labellaropening, o = ovary, s = sepals, sd = staminodium, st =stigma.


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Although bees have been reported to sometimeschew through the labellum in the showy lady's-slipper,C. reginae Walt. (Guignard 1886) and pink lady's-slipper, C. acaule (Stoutamire 1971), most find adifferent way out. A foothold is provided by tightlypacked, long hairs (trichomes) on the bottom of the lip(Ziegenspeck 1936, Summerhayes 1951, Stoutamire1967, Proctor & Yeo 1973). These lead along a pathway("haarstrassen") toward escape holes at the base or heelof the slipper (Fig. 1). The escaping bee may also beattracted along this path by colored lines on the lip'sinner surface (Artz 1954) and by light coming from theescape holes or, in some species, from translucent areas("light windows") in the side of the lip near its base

(Webster 1886, Troll 1951).

Daumann (1968) found that bees are well able tofind their way out when these "windows" are covered,but that light gradients are nevertheless a definiteorienting factor. Thus it took a bee an average of 11minutes to find its way out of the labellum ofEuropean yellow lady's-slipper, Cypripedium calceolusunder natural light conditions, but only two or threeminutes when an external light source was focused on

the base of the labellum. If the light was focused on theapex the time was increased to 30 minutes.

Nilsson (1981) believes that the hairs inside thelabellum may have an additional function. Droplets ofoil which are present on their distal tips could absorbbody odors (pheromones) from visiting bees and theseodors may serve to attract additional pollinators (see


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below). Because these hairs are strongly light refractive,Ziegenspeck (1936) speculated that they might alsostimulate a phototactic response complementing that ofthe light coming from the base of the labellum.

In its escape the bee must pass two points wherethe passageway is narrowed. At the first of these it isforced to squeeze under and rub its back against thesurface of the stigma (Fig. 1B, C) (e.g., Stoutamire1967). The stigma provides leverage that allows the beeto push down on and slightly depress the elasticlabellum, thus enlarging the passageway for its escape(e.g., Wright 1975, Nilsson 1979). Ziegenspeck (1936)considered that the basal trichomes might reduce

friction between the base of the labellum and the insectat this point. The second narrow passage is the exit holeitself (Fig. 1A, C). One exit hole is located on either sideof the base of the flower. An anther is so positionedbeside each exit that a bee of the proper size cannotforce its way out without contacting the anther andcarrying away a mass of sticky pollen on the dorsalsurface of its thorax (Fig. 1B, C) (Pijl and Dodson 1966,Stoutamire 1967). Since the bee contacts the stigmabefore the anther and usually does not reverse

directions, it does not ordinarily transfer pollen to thestigma of the same flower (functional protogynyaccording to Faegri and Pijl (1971)). Rather, pollinationis effected when and if the bee, upon escaping from thefirst flower, is subsequently trapped again, usually in adifferent flower, and the escape procedure is repeated.


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The same bees are often trapped more than oncein flowers of the California lady's-slipper, Cypripediumcalifornicum (Kipping 1971) and other species ofCypripedium (Nilsson 1979). Davis (1986), for example,observed five bumblebees follow the prescribed courseof entry and exit from flowers of C. acaule at a site inMassachusetts. Two were carrying pollen masses ontheir thoraxes when they entered the flower and eachdeposited the pollen on the stigmas of these flowers. Ina study in Nova Scotia, O'Connell and Johnston (1998)found a greater than 90% correlation in male and femalereproductive success. In other words, over 90% of theflowers that had a pollinium removed also received one.

Visitation rates were low, but once removed a pollinium

had a 36% to 51% chance of being transferred to thestigma of another flower. These and similarobservations in other taxa seem to refute the contentionthat once having endured the ordeals of entrapment andthe subsequent lack of reward, bees avoid repeating theexperience (Webster 1886, Baxter 1889, but see Gill1989).

Botanists are uncertain just why bees enter theflowers in the first place. In some cases the entry is

inadvertent. The bees, exploring the outer surface of thelip, tumble into the trap. This has been associated withan inflected rim near the staminode, the so-called slidingzone (Nilsson 1981). In other cases entry appears quitedeliberate (e.g., Nilsson 1981). Daumann (1968) believesthat insects may collect oil from the hairs on the insideof the lip. This has yet to be confirmed. It has also beenconjectured that small amounts of nectar are present or


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that the bees feed on the hairs in the labellum (Nilsson1979 and references therein). But it now appears likelythat the flower provides no food, and the bees aresimply deceived by false nectar guides, color, and theodor of the blossoms, which promise nectar where noneis available (Nilsson 1979). In addition, the staminode,

which projects into the labellum, is often bright yellowin color with contrasting spots and may appear to be asource of pollen (Vogt 1990).

Stoutamire (1971) believes that bees must learnby experience which flowers offer the best nectarreward (orchid pollen can't usually be deliberatelycollected by bees because of the anthers' position on the

flower) and will avoid flowers that offer no reward at all. According to this view seed production in thelady's-slippers may be dependent upon nave or possibly"forgetful" pollinators, insects that are newly hatched,new to the area, or shifting from a depleted to a newfood source (Delpino 1874, Pijl 1966, Dressler 1981,

Ackerman 1986).

However, Nilsson (1979) contends that, inaddition to general food deception, the floral attractions

in the European C. calceolus are attuned to otherinstinctive responses in bees and that very little learningis involved. The composition of the floral fragrance inthis orchid is uncommon, which suggests that any allureit may have differs from the usual advertisement of afood (nectar) source (Nilsson 1979). In addition to amonoterpene alcohol called linalool, which may elicit afeeding response, the orchid's fragrance contains


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acetates and farnesene. The acetates are found incephalic pheromone secretions ofAndrena bees, andfarnesene is a component of the abdominal Dufourgland in female Andrenas (Nilsson 1979, Bergstrom and

Tengo 1974). Pheromones are used to odor-markobjects which then attract bees of the same species.Farnesene is found as a lining in the nest and alsosignals the location of the nest site (Bergstrom and

Tengo 1974). According to Butler (1965), females ofAndrena flavipesdeposit farnesene in the soil surroundingthe nest site, and the odor provokes instinctive landingresponses in both sexes. The opening in the labellummay mimic the opening of the nest tunnel (Catling andCatling 1991). In addition, as already noted, several

features of the hairs within the labellum suggest thatthey may be well adapted to the absorption ofpheromones directly from visiting bees as a supplementto the artificial pheromones produced by the orchid(Nilsson 1979).

A survey of 8 species of North American orchidsfound comparable acetates present only in the ivorylipped lady's-slipper, Cypripedium kentuckiense and one

variety ofyellow lady's-slipper, C. parviflorum(probably

var. makasin based on provenance) (Barkman et al.1997). Contrary to Bergstrom et al. (1992), Barkman etal. (1997) found no farnesene in variety pubescence.Nilsson (1981) notes additional differences in thechemical composition of the floral fragrances in C.calceolus and varietyparviflorum. These differences maycorrelate with differences in primary pollinator species,as pheromone profiles can differ intragenerically among


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pollinators (Tengo 1979). A varying blend of odorconstituents may have evolved to stimulate aninstinctive or learned pattern of response on the part ofa range of pollinators to food, sexual reproduction, ornest location (Nilsson 1979, Gregg 1983). Much workremains to be done in this area, especially on Americanrepresentatives.

Among species ofCypripedium the size of theflower, the width of the entrance, and especially thediameter of the anther-exit holes and the space betweenthe labellar floor and the stigma determine the size ofthe insect involved in the pollination (Stoutamire 1967,Catling and Knerer 1980). The dorsal-ventral thickness

of the thorax appears to be of critical importance(Catling and Knerer 1980). In addition, Nilsson (1981)noted that the depth of the labellum in the European C.calceolusmust exceed the length of the pollinating bee bya minimum of 3 to 4 mm or the bee can simply crawlback out through the labellar opening. All of these floralcharacters are clearly under strong selection pressure inrelation to the primary pollen vectors. Research is still atan early stage, but the pollinators of some species oflady's-slippers can now be tentatively identified.

The largest exit holes are found in the pinklady's-slipper (Cypripedium acauleAit.) (Stoutamire 1967).

This orchid can occur in clusters of two to ten plants(Wright 1975). Vegetative propagation is by rhizomes,and adjacent plants may be from the same genet (e.g.,

Wright 1975, Cochran 1986). A single flower with adisproportionately large, bladder-like and pendant


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labellum is produced on a short scape from a pair ofbasal leaves (Cribb 1997, Luer 1975, Gill 1989). Theflower blooms for about three weeks but senesces inless than six days when pollinated (O'Connell and

Johnston 1998). The labellum is pink to more or lesspurple with a parallel ridge on either side of theopening. This ridge differs in color and vein patternfrom other parts of the labellum and is basically whitemarked by pink to purple lines (Stoutamire 1971). Theopening itself is a longitudinal, usually closed slit 3-4 cmlong. A pollinator must deliberately push the edges ofthe lip apart in order to enter, with the openingsubsequently closing behind it (Wright 1975). Any lightcoming from the base of the labellum must be largely

from the exit holes, as no translucent areas ("light windows") are present in this species (Wright 1975).However, the base may be lighter or may contrast incolor from the rest of the lip (Wright 1975), and theorienting pathway of tightly packed hairs on the bottomof the lip, referred to earlier, is present (Wright 1975).

The stigma is sticky and grooved rather than simplypapillate as in other species of Cypripedium (Luer 1975,Cochran 1986). The lateral petals and the sepals varyfrom yellow-green to greenish brown, usually with a

purple tinge (Luer 1975). Along with the labellum, theyproduce a sweet odor, which can be detected 2-3 inchesaway from the flower (Stoutamire 1967, 1971). Longdistance attraction is visual and is related to reflectanceof ultraviolet and blue-violet by the labellum andstaminode (Wright 1975).


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In keeping with the size of its exit holes, thisorchid appears to be pollinated by bumblebee queens inthe spring prior to the emergence of the workers(Stoutamire 1971), a time of the year when pollinatorsare usually competing for nectar (e.g., Cochran 1986).

Although Gray (1862) and Muller (1883) basicallyunderstood the pollination mechanism in Cypripediumcalceolus, the process in C. acaule took longer to clarify.Correll, as late as 1950, considered the chewing orpiercing activity of insects such as beetles and fliessomehow necessary to its pollination, and Bingham(1939) thought the exit holes were too small toaccommodate the escape of bumblebees.

The establishment of bumblebee queens asprimary pollinators is based, in part, on observationsmade in a northern Michigan spruce-blueberry bog by

Warren Stoutamire (1967). After much effort, a foragingbumblebee queen, Bombus vagans, common in the bog,

was captured in the lip of one flower. It carried pollenof the pink (stemless) lady's-slipper on its thoraxfrom earlier visits to other flowers in the area. (Theloose pollen ofCypripediummay adhere more effectivelyto hairy insects such as Bombus than would the viscidia

of more organized pollinia (Stoutamire 1971).) Bombusvaganshas also been collected carrying pollen ofC. acaulein New Brunswick (Plowright et al. 1980, Barrett andHelenurm 1987). Subsequent examination ofbumblebees in the collection at Michigan StateUniversity disclosed the apparent presence ofCypripedium pollen on several specimens of Bombusborealis (Stoutamire 1967). Wright (1975) notes that no


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other Cypripedium is primarily dependent on bumblebeequeens for pollen transport. Further work hasimplicated queens of additional species of Bombus andfemales of Xenoglossodes and Psithyrus as possiblepollinators (e.g., Stoutamire 1967, Plowright et al. 1980,Nilsson 1981, Davis 1986, Catling and Catling 1991).

A study in Ohio supports the role ofBombusandpossiblyPsithyrusbut casts doubt on the effectiveness of

Xenoglossodes sp. (Wright 1975). Four species of Bombusand a Psithyrus female, all of about the same size, wereobserved to traverse the labellum and remove pollen.

The relative sizes of the exit holes and the bees allowedescape, but with sufficient difficulty to insure contact

between the thorax and the anther (Wright 1975).Similarly, the hinge construction of the labellum allowedthe bees to force their way beneath the stigma, while theopposing pressure of the labellar hinge assured contactbetween the stigma and the dorsal thorax of the bees(Wright 1975). A female Xenoglossodes sp., althoughslightly smaller than the exit hole, was also observed toremove pollen. However, the mean distance betweenthe labellar floor and the stigma exceeded the height ofthis bee by several millimeters, and Wright (1975) did

not consider it an effective pollinator of Cypripediumacaule.

Unlike the pollen vectors of most other lady's-slippers, the bumblebees that pollinate Cypripedium acaulemay reverse direction after reaching the anther, causingself-pollination (Macior 1974, Wright 1975). However,self-pollination does not appear to occur unless


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pollinators are present, and development of seed in theabsence of fertilization is probably absent (Davis 1986).

Fruiting success in Cypripediumand other orchidswhich offer no reward is consistently much lower thanin orchids that provide nectar or that mimic plants thatdo (Gill 1989 and references therein). Insects visit C.acaule infrequently (e.g., Plowright et al. 1980, Barrettand Helenurm 1987, Gill 1989): in some studies lessthan 20% of the pollen masses are removed or flowerspollinated or fruit set under natural conditions(Plowright et al. 1980, Cochran 1986, Davis 1986, Gill1989, Gill and Stoutamire 1990, O'Connell and

Johnston 1998). On the other hand, artificial

pollinations have been found to produce high fruit-setin selfed and cross-pollinated flowers (70%-100%; Wright 1975, Cochran 1986, Davis 1986, Gill 1989,O'Connell and Johnston 1998). Thus, Cypripedium acauleis highly fertile, but short-term seed production appearsto be limited by the effective activity or availability ofpollinators (e.g., Cochran 1986). A similar pattern hasbeen reported in other orchids that rely on deception(e.g., Plowright et al. 1980, Dafni 1984, Davis 1986,Barrett and Helenurm 1987, Gill 1989, Primack and Hall

1990, Nilsson 1992).

Many authors consider pollinator limitation to beevolutionarily unstable. According to this view, naturalselection should maximize reproduction withinparameters set by the availability of resources with atrade-off between current reproduction and futurereproductive success and/or growth (e.g., Cole 1954,


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Williams 1966, Charnov 1982, Lloyd 1980, Waser and Jones 1991). Various strategies to increase pollinationsuccess to the level where resources are limiting shouldbe favored and should spread through the population.In Cypripedium acaule these might include a modificationin floral anatomy that would allow autogamy to occur inthis already self-compatible species (Gill 1989).

Alternatively, flowers might develop that provide anectar reward (Gill 1989). Such an invader could beexpected to reverse the negative reinforcement ofrepeated visitor behavior in non-rewarding flowers(Dafni 1987, Gill 1989). Although the presence or theartificial addition of nectar does not always mitigatepollinator limitation (e.g., Ackerman 1986, Burd 1995,

Johnson and Nilsson 1999), it had a significant effect onboth pollinia removal (male function) and fruit set(female function) in C. acaule (Cochran 1986). Finally,according to Gill (1989), a delay in flowering wouldincrease the abundance of newly emerging, navebumblebees available as prospective pollinators, butmight be ineffective due to a mid-summer reduction ofsunlight in the understory and the bumblebee'spreference for well exposed foraging sites. Gill (1989)finds it surprising that no such modifications have

arisen. Given the abundance of C. acaule and its longhistory, why has the predicted increase in individualreproductive fitness not resulted in the spread orfixation of such forms in any current population of thisorchid?

Data based on hand pollinations suggest that anincrease in fruit production can have a negative impact


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on subsequent growth and reproduction (e.g., Lloyd1980, Janzen et al. 1980, Montalvo and Ackerman 1987,Primack and Hall 1990, Ackerman 1989, Zimmermanand Aide 1989, Snow and Whigham 1989, Ackermanand Montalvo 1990, Primack and Stacy 1998).

Therefore, even though fruit production within seasonsmay be pollinator limited, an increase in pollination ratemight not significantly improve overall reproductivesuccess because such an increase could have an adverseeffect on lifetime fecundity (Janzen et al. 1980,Montalvo and Ackerman 1987, Primack and Hall 1990,Snow and Whigham 1989, Zimmerman and Aide 1989,

Ackerman 1989).

In Cypripedium acaule resource limitation resultedin a decrease in both plant size and the probability offlowering following repeated episodes of sexualreproduction and fruiting (Primack and Stacy 1998).However, according to Cochran (1986), the effects ofresource limitation are subordinate to pollinatorlimitation in this species: two years of completepollination are equivalent to ten to twenty years ofnormal fruit set, and long life spans and low mortality

would, in time, permit resource-depleted individuals to

resume reproduction.

Calvo and Horvitz (1990) also consider the costsof reproduction secondary to pollinator limitation.

According to their model, increased fitness resultingfrom higher levels of pollination and fruit set wouldovercome the relatively low cost of fruiting. Studies ofreproduction in the orchid Tolumnia variegata were


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consistent with these predictions (Calvo 1993). Astatistically significant reduction in future growth andflowering was observed only in plants subjected to ahigh pollination intensity treatment (viz., all the flowersin the inflorescence were pollinated resulting in a meanfruit set about 88 times greater than in open pollinatedplants). Simulations revealed that the production of onlya few seedlings per fruit could more than compensatethe cost of fruiting and that therefore selection forhigher levels of pollination should be favored. Calvo(1993) believes that the low level of pollination and fruitproduction frequently observed in nonautogamousorchids may be due to a low correlation between fruit orseed production and seedling recruitment. Selection for

increased levels of pollination would be ineffectively lowif an increase in seed production is not translated intoan increase in fitness. Under such circumstancespollinator limitation may be evolutionarily stable (Calvo1993).

Kull (1998) has reported observations onseedlings in Estonian populations ofCypripediumcalceolus

which are consistent with this view. In this studyseedling recruitment was limited by the availability of

suitable microsite conditions. Fruit-set, which appearedto be pollinator limited, had no observed effect onsubsequent flower and fruit production (resourcelimitation) and was uncorrelated with seedlingrecruitment. It therefore had no significant effect onpopulation fitness. Similarly, Keddy et al. (1983)observed a scarcity of microsite conditions suitable forseedling establishment of the sparrows egg ladys-


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slipper, C. passerinum among a mosaic of seres on thenorth shore of Lake Superior. On the other hand, Gill(1989) reported high levels of seedling recruitment forC. acaulein Virginia.

According to Primack and Hall (1990) andPrimack et al. (1994), Cypripedium acaule may perhapsbest utilize its reproductive resources over a protractedtime period by maximizing seed production duringperiods when habitat conditions are favorable. Byroutinely allocating resources toward the production ofmany more flowers or ovules than mature fruits andseeds, it could benefit from chance variation inpollinator behavior or seasons when pollinators or

resources required for fruit maturation are present inabundance (e.g., Lloyd 1980, Ackerman 1986, Cohenand Dukas 1990, Burd 1994). Under this view, theexpenditure of resources over the lifetime of the orchidmay involve a strategy adjusted to both pollinator andresource limitations (Ackerman 1986, 1989, Montalvoand Ackerman 1987, Haig and Westoby 1988, Nilsson1992, Burd 1994, but see Morgan 1993).

References Ackerman, J. D. 1986. Mechanisms and evolution of food-

deceptive pollination systems in orchids. Lindleyana 1:108-113.

Ackerman, J. D. 1989. Limitations to sexual reproduction inEncyclia brugii(Orchidaceae). Syst. Bot. 14: 101-109.

Ackerman, J. D. and A. M. Montalvo. 1990. Short- and long-termlimitations to fruit production in a tropical orchid. Ecology71: 263-272.


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Artz, T. 1954. Frauenschuh und Spinne.Natur. Volk 84: 421-425.Barkman, T. J., J. H. Bearnan, and D. A. Gage. 1997. Floral

fagrance variation in Cypripedium: Implications forevolutionary and ecological studies. Phytochemistry 44(5):875-882.

Barrett, S. C. H. and K. Helenurm. 1987. The reproductivebiology of boreal forest herbs. I. Breeding systems andpollination. Canad. Jour. Bot. 65: 2036-2046.

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Bergstrom, G. and J. Tengo. 1974. Farnesyl and geranyl esters asmain constituents of the secretion from Dufour gland in 6species ofAndrena (Hymenopter, Apidae). Chem. Scr. 5:28-38.

Bergstrom, G., G. Birgersson, J. Groth, and L. A. Nilsson.1992.Floral fragrance disparity between three taxa of lady's-slipper Cypripedium calceolus (Orchidaceae). Phytochemistry

(Oxford) 31(7): 2315-2319.Bingham, M. 1939. Orchids of Michigan. Cranbrook Inst Sci. Bull.

15: 26-34.Butler, C. G. 1965. Sex attraction in Andrena flavipesPanz. (Hym.

Apidae) with some observations on nest site restriction.Proc. R. Ent. Soc. Lond. (A) 40: 77-80.

Burd, M. 1994. Bateman's principle and plant reproduction: Therole of pollen limitation in fruit and seed set. Bot. Rev. 60:83-139.

Burd, M. 1995. Pollinator behavioral responses to reward size inLobelia deckenii: no escape from pollen limitation of seed

set.Jour. Ecol. 83: 865-872.Calvo, R. N. 1993. Evolutionary demography of orchids-Intensity

and frequency of pollination and the cost of fruiting.Ecology74 (4): 1033-1042.

Calvo, R. N. and C. C. Horvitz. 1990. Pollinator limitation, costof reproduction, and fitness in plants: a transition matrixdemographic approach.Amer. Naturalist136(4): 499-516.


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Catling, P. M. and V. R. Catling. 1991. A synopsis of breedingsystems and pollination in North American orchids.Lindlyana6 (4): 187-210.

Catling, P. M. and G. Knerer. 1980. Pollination of the Small White Lady's-slipper (Cypripedium candidum ) in Lambton

County, southern Ontario. Canad. Field-Nat. 94(4): 435-438.Charnov, E. L. 1982. The theory of sex allocation. Princeton

University Press, Princeton, New Jersey.Cochran, M. E. 1986. Consequences of pollination by chance in

the pink lady's-slipper, Cypripedium acaule. Ph.D.Dissertation, The University of Tennessee, Knoxville.

Cohen, D. and R. Dukas. 1990. The optimal number of femaleflowers and the fruits-to-flowers ratio in plants underpollination and resources limitation.Amer. Naturalist. 135:218-241.

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phenomena.Quarterly Review of Biology29:102-137.Correll, D. S. 1950. Native orchids of North America. Chronica

Botanica Co., Waltham, Mass.Cribb, P. 1997. The genus Cypripedium. Timber Press, Portland,

Oregon.Dafni, A. 1984. Mimicry and deception in pollination. Ann. Rev.

Ecol. Syst. 15: 258-278.Dafni, A. 1987. Pollination in Orchisand related genera: evolution

from reward to deception. InJ. Arditti (ed.), Orchid biology,reviews and perspectives. IV, pp. 80-104. Cornell Univ. Press,Ithaca, NY.

Daumann, E. 1968. Zur Bestaulringsokologie von Cypripediumcalceolus. L. Osterr. Bot. Z. 115: 434-446.

Davis, R. W. 1986. The pollination biology of Cypripedium acaule(Orchidaceae). Rhodora88: 445-450.

Delpino, F. 1874. Sugli apparecchi della fecondazione nelle piante.Firenze.

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Faegri, K. and L. van der Pijl. 1971. The principals of pollinationecology. 2nd ed. Pergamon Press, New York.

Gill, D. E. 1989. Fruiting failure, pollinator inefficiency andspeciation in orchids. Pages 458-481 inD. Otte and J. A.Ender, eds. Speciation and its consequences. Sinauer,

Sunderland, Mass.Gill, D. E. and W. P. Stoutamire. 1990. Are the breeding ofdeceptive orchids evolutionarily stable? Experimental testof inbreeding depression in Cypripedium acaule. Aiton.Amer. Jour. Bot. 77(6): 45.

Gray, A. 1862. Fertilization of orchids through the agency ofinsects.Amer. Jour. Sci. and Arts34: 420-429.

Gregg, K. B. 1983. Variation in floral fragrances and morphology:Incipient speciation in Cycnoches. Bot. Gaz. 144: 566-576.

Guignard, J. A. 1886. Insects and orchids. Ent. Soc. Ontario, 16thAnn. Rep.: 39-48.

Haig, D. and M. Westoby. 1988. On limits to seed production.

Amer. Naturalist131: 757-759.Janzen, D. H., P. De Vries, D. E. Gladstone, M. L. Higgins, and

T. M. Lewinsohn. 1980. Self- and cross-pollination ofEncyclia cordigera (Orchidaceae) in Santa Rosa NationalPark, Costa Rica. Biotropica12: 72-74.

Johnson, S. D. and L. A. Nilsson. 1999. Pollen carryover,geitonogamy and evolution of deception in orchids.Ecology80: 2607-2619.

Keddy, C. J., P. A. Keddy, and R. J. Plank. 1983. An Ecologicalstudy ofCypripedium passerinumRich. (Sparrows Egg Lady-Slipper, Orchidaceae) on the North Shore of Lake

Superior. Canad. Field-Nat. 97: 268-274.Kipping, J. L. 1971. Pollination studies of native orchids. M. S.

Thesis, San Francisco State College.Knoll, F. 1922. Fettes Ol auf den Blutenepidermen der

Cypripedilinae. Osterr. Bot. Z. 71: 120-129.Kull, T. 1998. Fruit-set and recruitment in populations of

Cypripedium calceolus L. in Estonia. Bot. Jour. Linn. Soc.126(1-2): 27-38.


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Lloyd, D. G. 1980. Sexual strategies in plants I. An hypothesis ofserial adjustment of maternal investment during onereproductive session.New Phytol. 86: 69-79.

Luer, C. 1975. The native orchids of the United States and Canada. NewYork.

Macior, L. W. 1974. Biological aspects of coadaptations betweenflowers and insect pollinators.Ann. Missouri Bot. Gard. 61:760-769.

Montalvo, A. M. and J. D. Ackerman. 1987. Limitations to fruitproduction in Ionopsis utricularioides (Orchidaceae).Biotropica19: 24-31.

Morgan, M. 1993. Fruit to flower ratios and trade-offs in size andnumber. Evol. Ecol. 7: 219-232.

Muller, H. 1883. The fertilization of flowers. Trans. D'ArcyThompson. Macmillan Co., London.

Nilsson, L. A. 1979. Anthecological studies on the Lady's-slipper,Cypripedium calceolus (Orchidaceae). Bot. Notiser 132: 329-

347.Nilsson, L. A. 1981. Pollination ecology and evolutionary

processes in six species of orchids.Acta Univ. Ups. 593: 1-40.

Nilsson, L. A. 1992. Orchid pollination biology. Trends Ecol. Evol.7: 255-259.

O'Connell, L. M. and M. O. Johnston. 1998. Male and femalepollination success in a deceptive orchid, a selectionstudy. Ecology79(4): 1246-1260.

Pijl, L van der. 1966. Pollination mechanisms in orchids. pp. 61-75 inJ. G. Hawkes (ed.) Reproductive biology and taxonomy of

higher plants. Pergamon Press, Oxford.Pijl, L. van der and C. H. Dodson. 1966. Orchid flowers, their

pollination and evolution. University of Miami Press, CoralGables.

Plowright, R. C., J. D. Thomson, and G. R. Thaler. 1980. Pollenremoval in Cypripedium acaule (Orchidaceae) in relation toaerial fenitrothion spraying in New Brunswick. CanadianEnt. 112: 765-769.


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Primack. R. B. and P. Hall. 1990. Costs of reproduction in thepink lady's slipper orchid: A four-year experimental study.Amer. Naturalist136 (5): 638-656.

Primack, R. B., S. L. Miao, and K. R. Becker. 1994. Costs ofreproduction in the pink lady's-slipper orchid (Cypripedium

acaule ): Defoliation, increased fruit production, and fire.Amer. Jour. Bot. 81 (9): 1083-1090.Primack, R. and E. Stacy. 1998. Cost of reproduction in the pink

ladys-slipper orchid (Cypripedium acaule, Orchidaceae): aneleven year experimental study of three populations.Amer. Jour. Bot. 85: 1672-1679.

Proctor, M. and P. Yeo. 1973. The pollination of flowers. London.Snow, A. A. and D. F. Whigham. 1989. Costs of flower and fruit

production in Tipularia discolor (Orchidaceae). Ecology 70:1286-1293.

Stoutamire, W. P. 1967. Flower biology of the Lady's- slippers.The Michigan Botanist6: 159-175.

Stoutamire, W. P. 1971. Pollination in temperate Americanorchids. Pp. 233-243 inM. J. G. Coorrigan [ed.], Proceedingsof the 6th World Orchid Conference. Sydney, Australia. HalsteadPress, Sydney.

Summerhayes, V. S. 1951. Wild orchids of Britain. London.Tenago, J. 1979. Odour-released behavior inAndrenamale bees

(Apoidea, Hymenoptera). Zoon7: 15-48.Troll, W. 1951. Botanische notizen II.Abhandlung Math. Naturwiss.

Kl. Akad. Wiss. Mainz 1951(2). Vogt, C. A. 1990. Pollination in Cypripedium reginae. Lindleyana 5

(3): 145-150.

Waser, P. M. and W. T. Jones. 1991. Survival and reproductiveeffort in banner-tailed kangaroo rats. Ecology72: 771-777.

Webster, A. D. 1886. On the growth and fertilization ofCypripedium calceolus. Trans. Proc. Bot. Soc. Edinb. 16: 357-360.

Williams, G. C. 1966.Adaptation and natural selection. PrincetonUniversity Press, Princeton, N.J.


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Wright, L. W. 1975. The pollination ecology ofCypripedium acaule Ait. (Orchidaceae). M.S. Thesis. University of Akron,Akron, Ohio.

peck, H. 1936. Orchidaceae. InO. von. Kirchner, E. Loew & C.Schroter, Lebensgeschichte der Blutenpflanzen Mitteleuropas.

Stuttgart.Zimmerman, J. K. and T. M. Aide. 1989. Patterns of fruitproduction in a neotropical orchid: pollinator vs. resourcelimitation.Amer. Jour. Bot. 76: 67-73.

Charles L. Argue, Department of Plant Biology, College ofBiological Sciences, 220 Biological Sciences Center, 1445 GortnerAvenue, St. Paul, MN 55108-1095. email: [email protected]. Argue is a regular contributor to the Journal and has writtenpapers on the pollination of Listera cordata, Pogonia ophioglossoides,the yellow fringed orchis complex, and many other native orchidspecies.


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Bentley: UPDATE ON CORALLORHIZA BENTLEYI

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UPDATE ON CORALLORHIZA

BENTLEYI J. V. FREUDENSTEIN,BENTLEY'S CORALROOT

Stan Bentley

Since press time for Native Orchids of the Southern Appalachian Mountains(June 2000), several discoveries

have greatly expanded our knowledge of Corallorhizabentleyi, Bentley's coralroot.

Until August 2000, this new species was knownonly from one location, which is in Monroe County,

West Virginia. I first discovered the plants in capsule inthe autumn of 1996. There were only four capsules -two from the 1996 season and two from the previousyear. This singular population has grown in number ofplants seen each subsequent season. In 1997, there were

eight plants found. But that number jumped to twentyplants in 1998, twenty-seven in 1999, and forty-oneplants in mid-July 2000.

On August 19, 2000, I returned to a site in GilesCounty, Virginia where I had, the previous year, foundsome unusually tall plants of autumn coralroot,Corallorhiza odontorhiza. In the area, I discovered forty-


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four plants of Bentley's coralroot. This discoveryestablished the species in Virginia and, at the same time,doubled the then known population for the entirespecies. This Virginia site is some eight miles removedfrom the original discovery site and is approximatelyeight hundred feet higher in elevation. A mile from theabove site, I discovered another new Virginia site, thisone with fourteen plants.

On August 26, 2000, friends from Pittsburg,Clete Smith and Scott Shriver, and Doug Jolley from

West Virginia accompanied me on another search formore populations ofBentley's coralroot. A completelynew area was found to contain nearly sixty plants. Jolley

spotted the first plants (large, healthy beauties). Whilestill in Giles County, Virginia, this area consisted ofseveral small sites scattered over roughly a half-mile offorest roadside. The most exciting discovery was severalplants in more than one site that had flowers with redlips rather than the typically described yellow.

Searching later in September 2000, I discoveredthree more Giles County, Virginia sites. Also found wasa new site in Monroe County, West Virginia which had

twenty-nine large healthy plants. Fortuitously, this newWest Virginia site also contained plants ofoval ladies'-tresses, Spiranthes ovalis, a rare species in this part of thecountry. This find is a Monroe County record.

At present then, there are six known Virginiasites and two known West Virginia sites for Bentley'scoralroot. The total population count for the year 2000


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exceeded two hundred individual plants. Adding to theexcitement also is several promising dried capsulesfound in this past winter of 2000-2001. I am mosteagerly awaiting the arrival of the 2001 flowering seasonfor Bentley's coralroot.

Stan Bentley, 1201 MacGill St. Pulaski, VA 24301Stan is a well-known native orchid enthusiast and photographer as well as theauthor ofNative Orchids of the Southern Appalachian Mountainspublished by theUniversity of North Carolina Press in 2001.

Corallorhiza bentleyiBentley's coralroot


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Coleman: BOOKS AND PAMPHLETS

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BOOKS AND PAMPHLETS ON THE

ORCHIDS OF THE UNITED STATESAND CANADA

Ronald A. Coleman

The search for native orchids is a worthy goal, whether for personal or scientific purposes. Thisfascination, even addition to wild orchids that we shareis not a recent phenomenon. A valuable historic recordsremains because many orchid students of the past tookthe time to capture for posterity the fruits of their laborin the form of articles, pamphlets and books. For those

with access to those sources, a wealth of knowledgeawaits. Collecting orchid literature can also be a passion,and is a worthwhile adjunct to the search for plants inthe wild.

Almost from the time I started studying wild

orchids Ive also been searching for and collectingliterature on them. The orchid literature is immense, andit would be an expensive hobby obtaining all the orchidbooks published each year, much less trying to obtainrare and out of print publications. Just as Ive limitedmy study of wild orchids to those in United States andCanada, so too I specialize in the acquisition of booksfrom that region. The list below includes all the books


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and pamphlets on wild orchids of the United States andCanada that I have been able to identify. All but two arein my personal collection.

Many articles and papers on native orchidsappear each year in lay and scientific publications.Keeping track of them is beyond the scope of thisarticle, although many papers are referenced in thebibliographies of the books listed. However, some ofthe pamphlets in the list first appeared in a scientificjournal and were subsequently issued as stand-alonedocuments.

The publications in the list are segregated based

on their geographic coverage. Those classified asNational in scope cover all or major portions of theUnited States and Canada. Those classified as Regionalcover areas equivalent to one or more states orprovinces. Those classified as Local cover a portion of astate or province, or a specialized area such as a specificpark or district. Some of these publications are fairlytechnical and are classified as Scientific. Two books didnot seem to fit any of the previous categories and arecalled Special.

Some of these books, especially the regionaltreatments, are still in print, and more are being added.It is gratifying to see the number of native orchid bookscovering a state or region that have been published inthe last decade, with others in press or planning stages.Books still in print can be ordered by most bookstores.

An excellent source of in-print books is the American


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Orchid Society. The AOS BookShop can be reached viathe internet at orchidweb.org.

Out of print books are more of a challenge, butmultiple resources are available. Two that I often useare Raymond M. Sutton Books reachable via email [email protected]; and McQuerry Orchid Books at

www.mcquerryorchidbooks.com. The EBAY bookauction often has out of print orchid books. Go to theEBAY web site at EBAY.com and search for bookusing the wildcard orchi* as the title. Several out ofprint books have been re-issued. In those cases thedates and publishers of the original and reprint are inthe table.

Should the reader know of any books coveringthe orchids of the United States and Canada that are notin the below list, please contact the author. Author Year Title Publisher Coverage

Ames, O. 1924 An enumeration ofthe Orchids of theUnited States andCanada

AmericanOrchidSociety

National

Ames, O. 1910(1979)

The GenusHabenaria in NorthAmerica (reprint)

Merry-mountPress;(Earl M.Coleman)

National

Correll,D. S.

1950(1972)

Native Orchids ofNorth America

ChronicaBotanicaCompany

National


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(StanfordUniversityPress)

Luer, C. A. 1975 The Native Orchidsof the United Statesand Canada

New York

BotanicGarden

National

Petrie, W. 1981 Guide to theOrchids of NorthAmerica

HancockHouse

National

Williams J.G. &Williams, A.E.

1983 Field Guide toOrchids of NorthAmerica

UniverseBooks

National

Ames, B. 1947(1979)

Drawings of FloridaOrchids

BotanicalMuseum(Earl M.Coleman)

Regional

Baldwin, H. 1884 The Orchids of NewEngland

John Wiley& Sons

Regional

Bentley,Stanley L.

2000 Native Orchids ofthe SouthernAppalachian

Mountains

Universityof NorthCarolina

Press

Regional

Bingham,M. T.

1939 Orchids of MichiganCranbrookInstitute ofScience

Regional

Brackley, F 1985 The Orchids of NewHampshire(Rhodora No. 849)

Rhodora Regional


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Brown,P. M. 1993

A Field and StudyGuide to theOrchids of NewEngland and NewYork

OrchisPress

Regional

Brown,P. M.

1997 Wild Orchids of theNortheastern UnitedStates

CornellUniversityPress

Regional

Burian, R. 2000 Native Orchids ofOregon

OregonOrchidSociety

Regional

Cameron,J. W.

1976 The Orchids ofMaine

Universityof Maine

at Orono

Regional

Case,F. W.

1964(1987)

Orchids of theWestern Great LakesRegion

CranbrookInstitute ofScience

Regional

Chapman,William K.

1997 Orchids of theNortheast

SyracuseUniversityPress

Regional

Coleman,

R. A.

1995 The Wild Orchids of

California

Cornell

UniversityPress

Regional

Donly,J. F.

1963 The Orchids ofNova Scotia

Regional

Ettman,J,

1979 An AnnotatedCatalogue andDistribution

(selfpublishedpamphlet)

Regional


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Account of theKentuckyOrchidaceae

Fuller, A 1933 Orchids ofWisconsin

North

AmericanPress

Regional

Gibson, W.H.

1905 Our Native Orchids DoubledayPage andCo.

Regional

Gupton, O.W. &Swope, F.C.

1986 Wild Orchids of theMiddle AtlanticStates

UniversityofTennessee

Regional

Henry, L.K., WernerE. Buker,and D. L.Pearth

1975 WesternPennsylvaniaOrchids

CarnegieMuseumof NaturalHistoryreprint ofCastaneaSpecialIssue,

Regional

Homoya,M. A.

1993 Orchids of Indiana IndianaAcademy

of Science

Regional

Keenan, P.E.

1983 A Complete Guideto Maines Orchids

DeLormePublishingCo.

Regional

Liggo, Joeand AnnOrto Liggo

1999 Wild Orchids ofTexas

Universityof TexasPress

Regional


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Long, J. C. 1970 Native Orchids of

ColoradoDenverMuseumof NaturalHistory

Regional

Luer, C. A. 1972 The Native Orchidsof Florida

New YorkBotanicGarden

Regional

Morris, F.& Eames,E. A.

1929 Our Wild Orchids CharlesScribnersSons

Regional

Munden, C.

Niles, G. G.

2001

904

Native Orchids ofNova Scotia

Bog Trotting forOrchids

UniversityCollege ofCape

BretonPress

G. P.PutnamsSons

Regional

Regional

Nylander,O. O.

1935 Our NorthernOrchids

Regional


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SzczawinskiA.E.

Sheviak, C.J.

1975

1974

The Orchids ofBritish Columbia

An Introduction tothe Ecology of theIllinois Orchidaceae

BritishColumbiaProvincialMuseum

IllinoisStateMuseum

Regional

Regional

Slaughter,C. R.

1993 Wild Orchids ofArkansas

Slaughter Regional

Smith, W.R.

1993 Orchids ofMinnesota

MinnesotaDepart-ment of

NaturalResources

Regional

Smreciu,E.A. & R.S.Currah

Summers,B.

1989

1981(1987)

A Guide to theNative Orchids ofAlberta

Missouri Orchids

Universityof AlbertaDevonianBotanicalGarden

MissouriDept. of

Conserva-tion

Regional

Regional

Wallace, M.E.

1951 The Orchids ofMaine(earlier version ofabove)

Universityof Maineat Orono

Regional


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Whiting, R.E. &Catling, P.M.

1986 Orchids of Ontario CanacollFounda-tion

Regional

Winterringer,G. S.

1967 Wild Orchids ofIllinois

IllinoisStateMuseum

Regional

Ames, O 1904 A Contribution toOur Knowledge ofthe Orchid Flora ofSouthern Florida

E. W. Wheeler Local

Bruce-GreyPlantCommittee

Busswell,W.

1997

1945

The Orchids ofBruce and Grey

Native orchids ofsouth Florida

StanBrown

Bulletin oftheUniversityof Miami

Local

Local

Craighead,F. C.

1963 Orchids and OtherAir Plants ofEverglades NationalPark

Universityof MiamiPress

Local

Fisher, R.M. 1980 The Orchids of theCypress Hills Fisher Local

Fultz, F. M. 1928 Lily, Iris, andOrchid of SouthernCalifornia

SpanishInstitutePress

Local


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Lamont, E.E.

1996 Atlas of the Orchidsof Long Island, NewYork

TorreyBotanicalClubreprint

Local

Reddoch,Joyce M.and AllanH. Reddoch

1997 The Orchids of theOttawa District(Canadian Field-Naturalist 111(1) )

CanadianField-Naturalist

Local

Allen, Caroled.

1996 North AmericanNative TerrestrialOrchidsPropagation andBreeding

NorthAmericanNativeTerrestrialOrchidConfer-

ence

Scientific

Brower, A.E.

1977 The Prairie WhiteFringed Orchid inMaine

MaineStatePlanningOffice

Scientific

Plaxton, E.H. ed

1981 North AmericanTerrestrial Orchids

MichiganOrchidSociety

Scientific

Sheviak, C.J. 1982 Biosystematic Studyof the Spiranthescernua Complex

New YorkStateMuseum

Scientific

Bulat, T. 1995 Hidden Orchids RudiPublishing

Special

Keenan, P.E.

1999 Wild Orchids AcrossNorth America

TimberPress

Special


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Ronald Coleman, 11520 E. Calle del Valle, Tucson, AZ 85749email: [email protected] is the author ofWild Orchids of Californiafrom CornellUniversity Press and many articles over the years in both this

journal and the American Orchid Society's publications. He iscurrently working on a new book on the wild orchids of Arizonaand New Mexico.


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Empiricist: OUT OF THE LOOP

154

OUT OF THE LOOP

The Slow Empiricist

Have you ever felt out of the loop when you aretrying to keep abreast of the current names andinformation about native orchids? There is much newmaterial currently being published and researched. The

North American Native Orchid Journalhas tried to keep youabreast of this constantly changing information witharticles and news bites. It sometimes seems like adauntless task.

Right now there are several new publicationsabout to hit the bookstores that will effect yourinformation and knowledge about orchids. Thepublishers of Flora of North AmericaNorth of Mexico areabout to issue volume 26, which contains the orchids.

There are many changes in nomenclature and alliances.

If you are a relatively new orchid enthusiast youmight feel lost amidst the myriad names and labelsattached to each orchid. Especially if you are a novice,you will need to be apprised of the procedures used toaffect these changes as well as why orchids and in fact,all plants, have a rigorous and scientific method of


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being identified and an appropriate name appended toeach specimen.

Let us begin with a short foray into the namingof an orchid. When a new species is discovered it needsa name. Science has found that common names do notdo the job as there can be too many commonappellations for a particular plant. This can result inmisapplications of names to the wrong plants or overthe years of a particular plant having several namesattached to it that are not valid. These last are calledsynonyms. Since this is confusing and can result inmisidentification, science came up with a strict code(International Code of Botanical Nomenclature) for the proper

naming of a new species. It goes something like this: theplant has to have a Latin name. This will ensure thatpeople all over the world will have a common language.

There can also be affixed a common name to the orchidin whatever language the information is beingpublished. There can be further information in thename such as forms added to the description. Then thebotanist who publishes this information will havehis/her name attached to the end of the orchid's name.

This is about as simple as I can make it. There are a lot

of nuances that I have not included like a new speciesbeing separated from an existing species cannot have aname too close to its original name as it would be tooconfusing. Believe me there are more steps andprocedures before a new species can be properlynamed.


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Maybe it would be a good idea if I took youthrough the naming of a new orchid that I was privy tolast year. On an excursion along Florida's Turnpike tocount the number of Sacoila lanceolata plants that weregrowing along the grassy edges of the road, Paul MartinBrown and his cohort, Stan Folsom, discovered a newcolor form (gold) for the red spiranthes. The newname of the new color form was named for Folsom.

Therefore S. lanceolata, the red spiranthes had a newrelative, S. lanceolata forma folsomiiP.M. Brown. It waspublished last year in this journal and has joined theother orchid names in the world.

The other things that have been happening all

center around advances in the scientific tools forascertaining what each orchid really is and in whichfamily it belongs. As simply put as I can describe it theyhave been testing orchids for DNA and finding thatcertain orchids don't belong with certain other orchidsas was originally thought.

They have also been most diligent in tracing theorigins of the names applied to the orchids to insurethat the proper designation has been supplied. Couple

these changes with the constant ongoing searches fornew species and you have a large and sometimesbewildering array of new descriptions, or families, ornew orchids to digest and learn about.

What brought me to write this column was alecture I attended recently in which the speaker referredto several of the orchid slides he was presenting by


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names that were no longer considered correct for theplants. Since he was a retired botanist of some years ofage it could be forgiven him that he was not cognizantof the current appellations for those orchids.

As I stated in an earlier column, nomenclature isgoverned by a strict set of rules. This publication tries tofollow the proper procedures when plant names areused. The current researchers in the field of nativeorchids are looking at all the previously cited names foreach particular orchid described to determine whichname is the earliest and correctly published name forthe orchid. That means that some commonly heldnames for orchids are in error as there were previously

validly published names for these plants. Hence theconfusion when someone refers to a plant by its old,commonly accepted name that is no longer valid.

I thought I would try to make up a list of someof the changes that I have become aware of as I haveread about their occurrence. I'm sure that I will not haveall the current changes as they are happening nearlyevery day somewhere in the world. I hope that PaulMartin Brown will enrich the selections with more if he

has knowledge of the changes coming into effect.

The red Spiranthes, is now known as Sacoilalanceolata. It used to be Spirantheslanceolata. There are alsoseveral related color forms like the green form and thegold form.


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The entire genus of Spiranthes has seen its ranksswell with the inclusion of S. eatonii, which had beenhiding under the names of S. laceravar. gracilis and S.torta. P. M. Brown discovered the plant specimenscollected by Ames in an herbarium with all thenomenclature present but it was never validly publisheduntil Brown published it the March 1999Journal.

Platantheras have been very carefully analyzed bypeople like Chuck Sheviak in New York State and havehad extensive changes wrought by his findings.

Epidendrum conopseum is now known as E.magnoliae since that name was published a scant two

weeks before the other more familiar name waspublished.

These are just a few of the many changes thatyou all will have to become cognizant of as the newpublications come on line. I hope you will look at thechanges as a chance to enrich your knowledge and notas an onerous chore. You wouldn't want to be called bya misapplied name or synonym if it really didn't fit you.Speaking for the orchids, neither do they.

Your Slow Empiricist


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St. George:A NEW ZEALANDER LOOKS AT ORCHIDS IN THEWILD WEST

159

A NEW ZEALANDER LOOKS ATORCHIDS IN THE WILD WEST

Ian M St George

One of the books from his own childhood that my fatherpassed on to us was called something like Buffalo Bills WildWest. It had a lot of Buffalo Bill Cody stories, and I loved them.With a cry, Cody sprang from the trees and Codys riflecracked and another redskin bit the dust were strong fuel for a

boys imagination. The first western movie I ever saw wasColorado Territory.

So the Rocky Mountains of Colorado have a nostalgicpull for me. This is Arapaho, Pawnee and Comanche country.Cody is buried near Denver. Wild Bill Hickock, on the otherhand, is buried at Deadwood in South Dakota. Ive been theretoo. The unsinkable Molly Brown, Titanic survivor, lived in

Denver, about as high above and as far from the sea as she couldrun.

Bill Jennings is a mining engineer who lives at thefoot of the Rockies in Louisville, Colorado. He has adetailed knowledge of Colorado orchids. I phone himfrom Wellington, and when I reach Colorado in early

July, he is at my hotel an hour later in his pickup (I cantell youre from Colorado he had been told on a recentorchid jaunt to Arizona: the cracked windscreen is a


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dead giveaway, a sign of the gravel laid on Coloradoroads in winter to improve traction). We head for thehills, through the city of Boulder and up FlagstaffMountain Road to an area among the red rocks and theponderosa pines, where we walk down a trail.Theres a plaque on the 15th step of the State Capitol

building in Denver: it marks altitude 5280 feet: one mile.Denver is Mile High City. Up here we are at 7500 feet,and it makes you a little breathless. First stop is a dampcreekside where the green bog orchid, Platantherahuronensis,is in full spike. We ascend further, pause to leta grey/green broad-tailed hummingbird, little biggerthan a bumble-bee, take nectar from trackside flowers.

The flowers are like our New Zealand weeds, a legacy of

the miners who came and went last century. A few areunfamiliar: the scarlet tufts of Indian Paintbrush forinstance. Further up we find giant rattlesnake orchis,Goodyera oblongifolia,in bud, its green leaves marbled with

white. The coral-roots: spotted, Corallorhiza maculataandWister's, C. wisteriana,are in fruit, but by a creek is thebroad-lipped twayblade, Listera convallarioides, itsflowers translucent brownish green mothwings. Bill tellsme he has never seen a rattlesnake around here and onlyonce in 30 years has he seen a black bear and I am

reassured. If you come from a country with nodangerous animals (except the domestic ones) you dolike to ask.

On the way home we stop in the Bouldersuburbs, for there between two new office carparks isanother orchid site; the developers are aware of thetreasure they guard, and have limited the size of the


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carparks to preserve it. This is Ute ladies'-tresses,Spiranthes diluvialis, a nationally threatened orchid, andone of them has a few white flowers fully open. I returna few days later to find several plants in full flower.

Ten thousand acres of this forest were razed byfire last July 4, Independence Day: why do we marksummer celebrations with fireworks? Bill has suppliedmaps and printed an itinerary for me so the next day Iam awake at 5.30, and soon on my way. Haybales in thepaddocks remind me this is summer, despite the winterI have left behind in New Zealand. The air is so dry Iget a static electric shock every time I step out of the car

at this time of year the relative humidity is in single

figures here. Names like Old Stage Road, Big ElkMeadow and Beaver Meadow remind me I am in theWild West. Fifty miles later I am among the high peaksin Rocky Mountain National Park. The north faces andflat areas under steep banks still carry snowdrifts. AStellers jay saunters off as my car approaches thecarrion he is eating; a chipmunk races across in front ofme. I am at the carpark near Bear Lake at 7.30, here tolook at the clustered lady's-slipper, Cypripedium

fasciculatum, a ladys slipper with two round glossy leaves,

and a couple of brownish flowers. Alas, there has beena mild winter, an early spring and a hot dry summer, sothey are over: I sadly photograph the spent heads. But

whoa! Beside them are what the British call the lessertwayblade (heart leaved twayblade), Listera cordata, andnearby, in the damp creeksides, the porcelain-white

white bog orchid, Platanthera dilatata, the yellow-flowered P. huronensis, some other Platanthera, some


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coral-roots past flowering, and then suddenly, in fullflower, spotted coralroot, Corallorhiza maculata, its redstems and flower parts contrasting strongly with the

white, red-speckled labellum.

Chipmunks chatter their warnings as I pass, andsquirrels scuttle up the far side of trees, away from mycamera, chirruping their distaste for my presence. In acarpark traffic island a chipmunk delicately eats wildstrawberries, pauses, darts, pauses, darts: she sits stilljust long enough for me to photograph the rock shedbeen sitting on a moment earlier. Silver birches amongthe pines; spruce, aspen, juniper.

A doe feeds in the lush green of a swamp nearCub Lake. A coyote lopes across the road, untidy as heloses his winter coat. A marmot runs for cover underhis rocks, then bold as brass, sneaks flattened to theground, clear in his own mind that he is invisible, backtowards me. I take time to approach a chipmunk,camera at eye, clicking as I come closer; I finish and rise,smiling smugly to myself in anticipation of thephotographs, to find I am being watched by a group of

American trampers, pitying, but entertained by my

interest in such an everyday creature. Out on the flat ofthe glacial moraine the members of a group of butterflycollectors huddle, nets like triangular pennants in thebreeze, looking at their treasures. In the warmth of analpine summer morning the air is fragrant with theperfume of the pines.


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I drive further up. Two elk bulls in velvet graze ahigh grassland, all power and majesty. Above them theconifers become stunted, twisted, and suddenly they arefinished and I am in open alpine meadowland, the

wildflowers colouring patches of white, yellow, orange,pink to scarlet through crimson to mauve, purple in allits shades, sky to dark blue to black. At the top it is11,796 feet, a tad shorter than Mount Cook.

There is something exhilarating about treasurehunts, and orchid forays in foreign countries oftenprovide that sense: Continue east on US36 to FishCreek Road, go south for three miles till road curves

west, then turn south along gravel road past Fish Creek

Ranch (Its a dude ranch, my papers saydisparagingly), enter Camp Cheley under a woodenarch, drive a further 250 yards until you reach a point

where the power lines on your left send a branch southacross the road: there is a sign on the power pole thatreads Watch out for future world leaders at work andplay: The large yellow lady's-slipper, Cypripedium

parviflorumvar.pubescens,is about 50-100 yards up the hillon the south side of the road: there are two extra largeponderosas right there. (I should tell you Camp

Cheley takes young people and trains them in leadershipskills.)

Dang. The Cypripedium is also over, the remnantsof its flowers browning in the afternoon heat. Nearbythough, I am compensated with what the British call thefrog orchid, Coeloglossum viride, in full flower. At anothersite I seek the flamboyant eastern fairy slipper, Calypso


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bulbosa. In 1979 I took a sabbattical leave in the American West and it was not all work. Before I lefthome I had been trying some rather pretentiousarchitectural photography in black and white, and whento my surprise I found a lovely pink orchid under thegiant Californian redwoods I forgot I didnt have colourin the camera and took a dismal series of pictures of thisfabulous flower in shades of grey. This time I am ready

with colour film, but sadlyCalypso is well in fruit by earlyJuly.

Near Red Rock Lake I find more Listera cordata,some plants with brown flowers, some with green. Thelake is one third covered with a beautiful native yellow

water lily, but the Indian Peaks of the Great Divide arereflected in the stillness of the other two thirds. Bill hasmapped out a number of further stops in case I havemissed orchids, but I am replete and head back to myhotel. Hard physical exercise is good for jetlag, but at5pm I am flagging (its 11 am tomorrow in NewZealand and its 12 hours since I slept).

I descend through Boulder Canyon: tannedbareheaded denim-clad couples on Harley-Davidsonszoom by, blonde hair flying, dangerously yet somehowenviably unencumbered with crash helmets. A Denverradio station plays continuous country music. Later,down on the flatland below the foothills, prairie dogshave turned the paddocks into a moonscape, theentrances to their underground cities looking likecraters; one stands upright keeping watch as the others


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feed nearby. Their habitat is shrinking as the cities ofmen expand.

I spend the working week sweltering in thesummer heat of the plains, and marveling at the earlymonsoon, the evening thunderstorms that bring rain,spectacular displays of lightning, and death. Coloradohas the second highest death rate from lightning strikein the US: three children taking an outdoor spa bathdied recently in Boulder; mountaineers will abort theirclimb if they cant reach the summit by midday.

A week later I am on Interstate 70, heading upinto the mountains again, this time toward the great ski

resorts of Aspen, Vail and Breckenridge. Theres a deadporcupine on the road, a bison herd grazing to the rightof the highway. The yellow tailings of gold and silvermines streak the hillsides. Higher up a couple of stonesfalling down a bank above the road catch my eye: sureenough, two bighorn sheep are grazing near the top,their grey fleeces perfect camouflage against the rocks.

My orchid stop is above the historic gold andsilver-mining Georgetown, its Victorian weatherboard

houses like Wellingtons. Above the town is GuanellaPass, and the road zigzags up the hill. My guide has

written, The road is narrow and potholed continueto Clear Lake and at the upper end turn left into theparking lots. Go to the lower parking lot. The wet seepyhillside between the parking lot and the road is full ofhooded ladies'-tresses, Spiranthes romanzoffiana, andPlatanthera huronensis. Indeed it is, and though I spot


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the tall Platantheraquite quickly, and though I recognizethe suitability of this habitat for Spiranthes, it takes mesome time to spot the first. Spiranthes romanzoffiana istiny, but I spot many as I begin to get my eye in andappreciate the tallest of them is 10cm, and most less:5cm or so. It is vanishingly rare in Britain, reportedfrom only one site.

In Georgetown for the night I discover arestaurant called Prague. On his menu the Czech chefhas (pause, then hushed, reverent voice), rack ofNew Zealand lamb, 21 dollars 95. Thats aboutNZD50. I settle for pork and sauerkraut, congratulatethe chef on the food, congratulate him doubly on his

choice of a smooth Californian merlot, and tip himgenerously in my postprandial largesse.

Next day I drive the highest (12,095ft)continuous paved road in the world, called (of course)Independence Pass. The roadside ditch west of thecontinental divide is full ofPlatanthera dilatataat 11,000feet. A doe, ears out and looking, makes sure it is acamera and not a rifle I am taking from my pack, thencontinues grazing. I sneak a quick look at the

pretentious tourist town of Aspen and pass on (Vail Igo past: I am told it is even worse). At GlenwoodSprings I visit the grave of Doc Holliday. Doc wastrained as a dentist, but when it dawned on him thatbettin and shootin were more fun than tooth pullin hebecame a gunman and gambler. He must have beenquite good at the shootin for he died of pulmonarytuberculosis.


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I have come home intending to revise my way of

helping orchid-minded visitors to New Zealand. I thinkI will do what Bill did for me: guide them to orchidsthat grow in wonderfully scenic sites with a set oftreasure hunt instructions, setting them free,unencumbered by the obligation of my company, to feellike adventurous pioneering children seeking gold in a

wild new country.

Ian St George is editor of theNew Zealand Native Orchid GroupJournal.22 Orchard St, Wadestown, Wellington, New Zealand, phone0064 4 499 4227, fax 0064 4 3894178, email [email protected].

eastern fairy slipperCalypso bulbosavar. americana


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THE ARCTIC ORCHIDS OF ADELBERT

VON CHAMISSO

Mark Nir

The purpose of this paper is to tell the story ofthe origin of some orchid names encountered in theflora of North America and of their progenitor.

The brig "Rurik," a smallish two-master of 180tons, under the command of the Lieutenant of theRussian Imperial Navy Otto von Kotzebue, left the portof Copenhagen on August 17, 1815. Well financed andfor that time relatively lavishly provisioned by Admiral

Adam von Krusenstiern, one of the earlycircumnavigators, it was commissioned by CountNikolaj Romanzoff, Chancellor of the Russian Empire,to undertake "A Voyage of Discovery, into the South Sea andBeering Straits, for the Purpose of Exploring a North-East

Passage." It was granted the privilege of displaying theRussian imperial naval flag and bore a crew of thirtysailors and eight small cannon. The first officer wasGleb Simonovich Shishmareff; the ship surgeon JohannFriedrich (a.ka. Ivan Ivanovich) Eschscholtz, physician,entomologist and botanist; the painter Louis (Login

Andrejevich) Choris; and the naturalist Adelbert

Nir:THE ARCTIC ORCHIDS OF ADELBERT VON CHAMISSO


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(Loginovich) von Chamisso. The patronymics wereindispensable on a Russian ship, although of thosementioned only Cleb Simonovich was Russian.

Chamisso was 34 years old when he joined theexpedition. He was already prominent in Germanliterary circles as a poet who had achieved internationalfame with the novel "Peter Schlemihl," the story of aman who sold his shadow. Born Charles Louis

Adelaide Count Chamisso de Boncourt, he was elevenyears old when during the French revolution his father,confronted with the choice between decapitation and ahasty exile, unsurprisingly chose the latter. Twocenturies ago life as a refugee appears to be no different

from what it is today. The family wandered for fouryears in the Low Countries and in several German statesuntil permitted to settle in Berlin in 1796. At the age offifteen he helped support his family by paintingminiatures at the royal porcelain factory. His talent as apainter is probably best attested to by the beautifulcolor illustration of Eschscholzia1 californensis Cham.,

which accompanies his description of the YellowCalifornia Poppy, the California stare flower. Followingseveral years of military service and travel, Adelbert

became a student of natural history in Berlin. It was atthis point in his career that due to ill health thenaturalist originally intended for the voyage of the Rurikhad to be replaced. Through well-placed connectionsand good luck Chamisso became the replacement.

1Dr. Eschscholtz spelled his name with a "t", but Chamisso omitted it fromhis new genus Eschscholzia.



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The first leg of the voyage from Plymouth to

Tenerife resulted in a major scientific find: studying thediverse marine creatures collected floating close to thesurface, Chamisso and Eschscholtz discovered themetamorphosis of the tunicates.2

The crossing of the equator for the first time onthe 23rd of November was' accompanied by the thencustomary celebrations, which nowadays arc reduced toa terse announcement (if at all) by the pilot. Theyreached Santa Cruz (off Brazil), continued southwards,rounded Cape Horn, and after a short stay in Chilesailed due west skirting Sala-y-Gomes, visited Easter

Island, passed between the Marquesas and FrenchPolynesia, then gradually turned north, revisitingRomanzoff Island (now Wotje) in the Marshall

Archipelago, discovered by yon Krusenstiern in anearlier expedition, and finally reached Petropavlovsk onKamchatka on June 19th.

The first period of exploration of the Bering Seaand the Aleutians between July 27th and September14th 1816, resulted in many geographic discoveries,

among them the Kotzebue Sound and in it the smallChamisso Island, Choris Peninsula and EschscholrzBay. Exploring the area provided part of the materialfor Chamisso's publication in Linnaea(1828).

2http://www.uinla6.kI2.wy.us/WWW/MS/8grnddlnlo%20Acccss/SP

ANTLGY/urochord.htm



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The Rurik then turned southwards and after a

stay in San Francisco continued towards the SandwichIslands. Arriving at the harbor of Hana-ruru on theisland of O'Wahu they met king Tameiameia. It is notthe purpose of this paper to summarize the rest of the"Voyage" but the preceding sentence should serve to

whet the appetite of those so inclined, to read the wholething (although calling the English version a"translation" would be too kind). A lengthy explorationof Hawaii and the Marshalls gave Chamisso theopportunity to make observations and measurementspublished in his "Notices sur les iles de corail de GrandOcean" (1821), which provided much of the material

and crucial information for Darwin's "The Structure andDistribution of Coral Reefs" (1842). The Rurik thenheaded northward again, back to the Aleutians.

The second stay of the Rurik in Unalaska lastedfrom April 24th to June 29th. It served mostly toprovision and equip the vessel for a second attempt tosail farther north in search of the northeast passage, butfor Chamisso it provided the opportunity to botanizeduring the period of maximum flowering. Anemones

and orchids were starting to appear towards the end ofMay and flowering into June, providing material for therest of his Orchidaceae Arcticae.

The trip northward failed again and the Rurikreturned to Unalaska for the last time on July 22nd. The



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ship left the Aleutians on August 181h and returnedthrough the by then well-established route through the

Marshalls, Hawaii and Guam to Manilla, thenceto Capetown and back to Europe.

Orchidaceae Arcticae contains the following taxa (themost informative reference follows in smaller font):

1. OrchisJ latifolia L.(?) var. beeringiana Cham. =Dactylorhiza aristata (Fisch. ex Lindl.) Soo. Hult. Fl.Kamtch. 259

2. Orchis camtschatica Cham. = Platanthera camtschatica(Cham.) Makino. Hult. Fl. Kamtch. 262

3. Habenaria borealisvar. albiflora Cham. = Platantheradilatata (Pursh.) Lindl. ex Beck. Correll, Nat. Orch. N. A. 714. Habenaria borealisvar. viridifloraCham. = Platantheraconvalliaefolia(Fisch.) Lindl. Hult. FI.Aleut. 1515. Habenaria schischmareffiana Cham. = Piperia unalascensis(Spreng.) Rydb. Hult. Fl. Aleul. 1486. Habenaria chorisiana Cham. = Platantehra chorisiana(Cham.) Rchb. f. Hult.. Fl. Aleut. 1487. Habenaria viridis R. Br. = Coelogloglossum bracteatum(Muhlenb. ex Willd.) Parl. Correll, Nat. Orch. N. A.: 114

8. Spiranthes romanzoffianaCham. Correll, Nat. Orch. NA 2209. Listera cordata R. Br. Correll, Nat. Orch. N.A.: 12610. Listera eschscholziana Cham. = Listera convalllarioides(Sw.) Torrey. Hult. Fl. Aleut. 15411. Ca!ypso borealis Rich. = Calypso bulbosa Correll, Nat.Orch. N.A.: 28012. Cypripedium macranthumSw. Hult. Fl. Kamtch. P.25713. Cypripedium guttaumSw. Hult. Fl. Kamtch. P. 256



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14. Malaxis diphyllosCham.15. Corallorhiza innata (sphalmate intacta) R. Br. =Corallorbiza trfiidaChat. Hult. Fl. Aleut. 156

After his return Chamisso worked as associatecurator at the botanical garden and herbarium at Berlin-Dahlem under von Schlechtendal and later as itsdirector. He settled down to a rather unsettled familylife, producing offspring both marital and extramarital,as well as a large body of poetry and a wealth oftaxonomic publications. Allthese before he died at age57.

Adelbert von Chamisso drawn by E.T.A. (TheNutcracker) Hoffmann, 1805.

References:Darwin, C. 1842. The Structure and Distribution of Coral Reefs.Chamisso,. A. 1819. De animalibus quibusdam e classe vermium

Lineana: De Salpa. F. Diimmler Verag, Berlin.



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Chamisso, A. 1828. Orchideae Arcticae. Linnaea25-35.Chamisso,A. 1975. Werke in Einem Band.Winkler Verlag, Munich.

Correll, D. S. 1978 Native Orchids of North America. StanfordUniversity Press.

Hulten, E. 1960. Flora of the Aleutian Islands.J. Cramer, New York

Hulten, E. 1927. Flora of the Kamchatka and the Adjacent Islands.Almqluist & Wiksells, StockholmLahnstein, P. 1987.Adelbert von Chamisso. Ullstein Verlag, Berlin.

Acknowledgment:I thank Alfred von Krusenstiern and Dr. Frank Axelrodfor valuable help.

Mark Nir, M.D.. 18 Ashford Ave., Dobbs Ferry, NY 10522;email: [email protected] is the author of the exhaustive Orchidaceae Antillanae

published in 2000 by DAG Media. It includes all of the orchids inthe Caribbean and many in southern Florida.

Platanthera chorisiana Malaxis diphyllos


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RECENT TAXONOMIC AND

DISTRIBUTIONAL NOTES FROMFLORIDA 10.

Paul Martin Brown

Just Visiting or Permanent Resident?

Florida has several species of orchids that have

been found in a wide variety of counties in the state thatare generally considered not to be part of the nativeflora. In many instances that determination is clear-cutbecause the first occurrence of the species can bedocumented and then the spread and establishment ofthe species has been noted. These species should beclassified as Naturalized: well established andreproducing in the wild. If the original plants werepurposefully planted out in the wild they would beconsidered Introduced.

In other cases the species may be a populargarden subject and subsequently found in one or twonearby localities, but rarely persisting. If sporadicsightings are made then these plants could be classifiedasAdventive: occurring in the wild (or at least withoutcultivation) but not necessarily reproducing orpersisting.


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The third category is a catchall for those species

that have been found but a single time, are notreproducing and are usually short-lived. These would beconsidered as Waifs.

Because of Florida's geographic position so nearthe West Indies, there are several records of speciesmore frequently seen southward that have had one ortwo Florida records. Whether to consider them as trulynative or as a casual introduction (a waif?) requirescarefully assessing the habitat in which they have beenfound and were they found in areas that are infrequentlybotanized.

The above categories certainly are not cast instone, and it is conceivable that a species that isconsidered a waif or garden escape could eventuallybecome thoroughly naturalized. As far as some of theCaribbean species are concerned there may never be asatisfactory answer.

june 2001 north american native orchid journal

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