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EVOLUTION IN POLLINATION OF ORCHIDS

Edited by Sysser Waspe
ICS, Wed, 7 May 97


I have moved questions and answers around to group them. There were so many "right" answers, that I didn't want to delete any. You will find that conferees quite often summed up the subjects which were discussed. I have, of course, also added my comments here and there.

INDEX

  1. ONC. ORNITHORHYNCHUM

  2. CHANGE IN POLLINATORS

  3. HAND POLLINATING AND HYBRIDS

  4. UV-REFLECTANCE

  5. POLLINATORS IN THE WILD

  6. FRAGRANCE

  7. PREDATORS

  8. DARWIN

  9. MADAGASCAR POLLINATION EXPERIMENT

  10. VANILLA planifolia

  11. POLLINATION ILLUSTRATIONS

  12. ORANGE JUICE

  13. BIBLIOGRAPHY

marylois
Let's talk POLLINATION....please, one and all. Let's do go over some points in Sysser's paper, add, ask questions - she has kindly offered to edit our discussion and make comments for distribution... Guess we are all agreed on the wonderful job Sysser did on her monumental thesis on orchid pollination. I believe it has something new in it for everyone! Let's each tell something we found "NEW TO US" in Sysser's paper.

ONC. ORNITHORHYNCHUM

prankster d(Susan from Oregon)
Does anyone have any idea what sort of beast pollinates Onc. ornithorhynchum? (Inquiring minds want to know!)

It's the smell of O. ornith. that makes me wonder. Unusual but lovely! Temperate zone plants just don't smell that way!

nodosa
Re O. ornithorhynchum. Certainly not a bird (re the name) unless it is a hummingbird. My guess would be one of the smaller bees, judging from the look of the flowers.

marilyn001
I did some work with Onc. ornithorhynchum last year. Presented to the Odontoglossum Alliance. I do not know which pollinator is usual. I do know that flowers must be pollinated almost as soon as they have opened if one wants to get seed.

nodosa
Marilyn, I think it is good practice to use a very new receptor and to use pollen from a more mature flower.

graphicgreg
Onc. bahamense is pollinated by territorial bees that see the flowers as invaders (in UV) and I would suspect ornithoryhnchum to have a similar pollinator. I have also seen bumble bees copulating with the Onc, sphacelatum here in our garden

Sysser
In the book "Orchid Flowers, Their Pollination and Evolution by L. van der Pijl and Calaway H. Dodson," page 181, there is a long list of Orchids and their KNOWN pollinators. The 6 different Oncidiums mentioned there are all pollinated by BEES (mainly from the Centris sp.)

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CHANGE IN POLLINATORS

(John in Arcadia CA)
I was wondering if the pollinators might change as the plants have changed. Is it a two-way street?

marylois
Seems more like the flowers are the sellers and the pollinators to buyers - so the former has to adapt to the latter...too many other flowers out there...and there is a reference to the fact that a relatively small segment of a population will be worked by the pollinators - they don't have to go far.

nodosa
Lois - I don't think there would be much incentive for a pollinator to evolve to service a species. Sort of a chicken and egg thing, but a pollinator must exist before the species can reproduce in quantities needed for success.

marylois
Ed, Marilyn - either aware of an instance where the pollinator evolved to meet the need of the orchid???

Sysser
According to Dressler's and Phil.'s comments the pollinators of today have hardly changed compared with fossil finds. Pollinators are MOBILE, and can fly wherever they want to and can therefore be "choosy".

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HAND POLLINATING AND HYBRIDS

Fleur
I have a question, are we changing evolution of orchids by hand pollinating them and preventing natural pollinators from doing their work?

nodosa
Fleur - I don't think our "in captivity" breeding has much effect on the evolutionary picture. Sure, we selectively breed and in time change many aspects of the subject. Still, we breed only a tiny number of any one species so true natural selection has the overwhelming influence from a statistical point of view.

marilyn001
Good question, Fleur. I am investigating population behaviour in Cyp. parviflorum. There are some plants which act primarily as pollen donors and do not set seed whatever the pollen used. There are other plants which are superior seed parents, at least horticulturally speaking. Seeds of different plants germinate differently on various medium formulations. Distant outcrosses germinate on a wide variety of media whereas near neighbor crosses are much more fussy.

The only challenge comes if we intend to reintroduce orchids to the wild.

HelenH
Fleur, it is hard for me to get too many natural pollinators on the 11th floor although we have had praying mantis and lady bugs fly in...

(Susan from Oregon)
Fleur, we already crossed that bridge when we hybridized 'em. (Do hybrid orchids attract hybrid pollinators?) As long as we leave a few species plants in their rightful homes and let nature take it's course with them, their natural evolution should prevail, as long as the integrity of the habitat holds out, that is! Once a plant comes into our greenhouses, it's not evolving naturally. Does nature ALWAYS select polyploid flowers?

Sysser
Good question, I have not come across any literature, which covers that subject. Perhaps it have not been studied yet?

Fleur
My point was that our hybrids may HAVE to be hand pollinated in future as they will no longer be able to attract a suitable pollinator.

Sysser
Fleur - Yes in a small way, but I also strongly believe that many strange man-made hybrids wouldn't stand a chance in the wild.

(Magda from Miami)
About hybrids, I would think most need to be hand-pollinated now, not just in the future, because who knows whether they can reliably attract a pollinator. And besides, the hybrids are probably living far away from their parents' home and therefore their pollinators.

Fleur
There is a spider in Australia somewhere that puts out a perfume to attract moths to its web, so I suspect that smell is a great attraction.

nodosa
Forgive me, Fleur, but isn't that Aussie spider dispensing ersatz pheromones to attract prey?

(John in Arcadia CA)
Thanks, Marilyn. I assume then that this predation does affect evolution as any mutaions might not even survive to reproduce.

nodosa
Fleur - I wouldn't give up on natural pollinators yet. We have a ground-dwelling black bee (about half the size of a bumble bee) that can and will pollinate anything and everything. It also eats holes in cedar siding and is generally obnoxious. It pollinates hybrid or species in our outside growing area with no difficulty.

(John in Arcadia CA)
Fleur - I believe that we may have eliminated some pollinators so that we will have to do it from now on.

Magda - that makes a great deal of good sense!

graphicgreg
We have a similar bumble bee here that will pollinate almost anything...also nest in the drainage holes of pots

(John in Arcadia CA)
Hybrids may not have a natural pollinator as each parent may have different pollinators and thus their "off-spring" would not have any natural one.

HelenH
It is possible that hybrids will attract a similar pollinator as one of the parents.....

sysser.
Why not, if it is visiting anyhow. But even if it visited your area, the pollinator would not go out of its way to pollinate your delicious flowers. No known pollinator only services orchids. The pollinators have a lot of other kinds of flowers to choose from.

marylois
There just aren't too many "natural hybrids" ... Ed, explain hybrid swarms. Magda - that section is the one I referred to unknowingly -- it is very few of the dominant flowers -- those who sell their wares the best -- who are responsible for reproduction, according to that study. The pollinators didn't go much further than those.

marilyn001
Actually, hybrids might be worthwhile when establishing plants in the outdoor tropical landscape. If they do not set seed, they are unlikely to escape cultivation and overwhelm local flora.

(John in Arcadia CA)
Marilyn - that is a very interesting theory and one that appears to be correct as far as I can see.

graphicgreg
That's very interesting Marilyn. Magda, what you say about the hybrids not attracting pollinators, but as Ed pointed out, our orchids often get pollinated. I was getting ready to take a C. amethystoglossa to judging a few years ago and the day before a bee hit it and closed up three flowers. Surely not the same bee as in Brazil...but a bee that found the flower attractive for whatever reason. I think the odds would be low but one of the things that interests me most about orchids is the fluidity of their breeding, They are evolving before our very eyes as countless natural hybrids become species unto their own.

sysser
Yes, so true, Greg. We just have to wait to see if those hybrids will make it - unless they happen to be self-pollinated.

HelenH
Fleur, at the turn of the century colonies of Catasetum were gathered - actually wiped out and brought to England. As they bloomed out, there were natural hybrids among the species found all within the same colony of plants..... so this might also apply to other species, besides orchids....

(Magda from Miami)
Marilyn, I thought once you got a pollinium stuck to the stigmatic surface, there was no room for more. So what's the advantage of getting insects to revisit pollinated flowers?

marilyn001
I think we need to separate hybrids as in hybrid swarms of naturally evolving orchids with hybrids as in multi and intergeneric hybrids. While complex hybrid flowers will undoubtedly attract potential pollinators, there will be less of a chance of successful pollination and seed production.

While most orchids have solid pollinia, several have granular pollen, sectile pollen or sticky pollen that can be transferred in part to many flowers in turn. Disa pollen is of the sectile type. Each tiny packet of pollen (massula) containing 100 or more pollen grains. Cypripedium pollen is sticky or granular while pollen of Epipactis is downright powdery. Some or all of a pollinium is deposited by a pollinator.

I saw Masd. peristeria 'capture' a nosey housefly. The fly entered, got 'pollinated' but could not get out because the thorax got caught.

prankster d(Susan from Oregon)
Pollinator flys that can't exit sounds like an evolutionary dead end for both plant and insect!

mplewinska
Ah, now it makes sense.Thanks.

mplewinska
Maybe they are on the way to evolving into carnivorous plants :)

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UV-REFLECTANCE - a very important and interesting subject

marilyn001
We have done UV-reflectance photography of Epipactis helleborine flowers. Interestingly, virgin flowers have UV-reflective stigmas but within 15 minutes after pollination, the stigmatic surface becomes UV-absorbent, resembling the nectar-laced lip surface. Wasps are first attracted to previously visited flowers then later visit virgin flowers. In this orchid, pollen does not always germinate well so the flowers invite wasps to revisit!

Fleur
So, as one pollinator is eliminated another one comes along.

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POLLINATORS IN THE WILD

nodosa.
Pollinators in the wild are much less efficient than us toothpick toters in the greenhouse. You rarely see more than a handful of seed capsules on a wild spike - even something like O. sphacelatum or altissumum where there might have been hundreds of flowers.

Sysser
Exactly

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FRAGRANCE

(John in Arcadia CA)
Marilyn - does the fragrance come out as the flower opens then?

Sysser
That would defeat the idea and the flower would use too much energy. All fragrant flowers will only produce an odor, when they are ready for pollination (like dogs on heat).

graphicgreg
Personally, I think fragrance to be less important in many genera. Several Brassias are pollinated by predatory wasps that see the flower as a spider in a web. They normally would attack the spider and narcotize it, then lay an egg in it for food supply for the larva. In stinging the Brassia they effect pollination

sysser
Odor is a long distance attractant compared with, e.g., colour, patterns and size.

marilyn001
When we consider fragrance, I believe we must consider the overall effect that many flowers may have on attracting a pollinator. Once amongst the flowers, pollinators may not be so choosy. I do not know if this has ever been tested.

nodosa
Greg - I think fragrance may be very important where a moth is the primary pollinator.

Sysser
And colour (from white to pale green)

Pollinators follow a trained pattern. In the beginning, when in their "teens", they are perhaps clumsy and make mistakes, but soon learn which flowers offer the reward they are looking for.

graphicgreg
True, Fleur. The Angraecums and other night-blooming orchids especially (Brassavola for instance). Also I'm sure you all must be familiar with the involved fragrance-related pollination of Catasetums.

Fleur
Isn't that a perfume/smell of some sort, Ed?

nodosa
A pheromone is a perfume only if attenuated to your personal frequency. I suppose if your personal frequency is frequent enough, other conditions might apply.

bmtorchids
Fleur, I was told long time ago, old socks smell, orchids are fragrant.

Fleur
Barbara, some of my orchids smell. *grin*

marilyn001
Fleur, I wonder if you grow Dendrobium kingianum? If so, are some clones more fragrant than others?

Fleur
I have about 10 kingianum, and they all have a strong perfume, hard to tell if some are stonger than othes.

(John in Arcadia CA)
Fleur - smell, but one "smell" is fragrance to others. Some people would not like the smell of cloves for instance and that one would smell to them.

Fleur
Only a dung beatle or a carrion lover would go near some of my Masd.

Tarwood(Karen, NY)
I presently have a Phrag. Grande in bloom, quite stunning, but it's not fragrant...it SMELLS like old sneakers.

(John in Arcadia CA)
Fleur - I believe that I will take what you say on trust. Those are the ones I would NOT be interested in. I suppose, however, some of them are really very beautiful.

graphicgreg
What about the Draculas and Bulbos that smell like carrion and are pollinated by flies???

(John in Arcadia CA)
Are any of the phragmipediums fragrant?

Fleur
And a blooming Masd. civilis will have a large blow fly in each flower within seconds of opening a door.

nodosa
Fleur, will your new greenhouse need an embalming room for those things you grow?

HelenH
Yes Bulbos have wonderful, ahemmm, smells......often like a toilet bowl not been cleaned for a while...

mplewinska(Magda from Miami)
(What about the Draculas and Bulbos that smell like carrion and are pollinated by flies) - I'm sure they are lovely.

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PREDATORS

(John in Arcadia CA)
In the wild, are there any predators on the ripening seed pods?

Sparky-Steve
John, I have seen a racoon eating pods of a Enc. tanpensis---although they will eat ANYTHING!

marilyn001
John asks about predation. Capsules are eaten by deer, squirrels or the tropical equivalent. Some plants are more heavily attacked by insects than others. Cyps and many other native orchids are fed upon by weevils. Larvae feed on the developing seeds

Predation can affect evolution in that predation-prone plants are rarely successful producing seed. They do however continue to be pollen donors.

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DARWIN

graphicgreg
No mention of Darwin finding Angcm. sesquipedale and stating that there must be a moth with a foot-and-a-half proboscis ???

Sysser
I read about it, but didn't think it worth my while to mention that, as Darwin's find is always mentioned, nearly no matter where one looks. I had to fit this project into only 12 A4 papers. The whole subject is so vast that it was an absolute nightmare for me to "cut a toe here and a heal there" to make it fit and still try to cover as much as possible. My personal aim was also to write something new, something that would be worthwhile for others to read and enjoy. I thoroughly enjoyed every minute of it and learned an awful lot, not just about orchids, but about the TOTAL picture in evolution.

HelenH
Greg, I believe it is 13 inches...and ten years after Darwin saw the flower they did find the moth!

prankster d(Susan from Oregon)
If pollinators came before their orchids, what did Darwin's moth
pollinate before they learned to enjoy Angraecum? I always thought
this was THE classic case of pollinator/flower co-dependance.

sysser
L. Anders Nilsson wrote: "As certain moths of Madagascar became larger through natural selection in relation to their general conditions of life, either in the larval or mature state, or as the proboscis alone was lenghtened to obtain honey from the Angraecum and "other" deep tubular flowers, those individual plants of the Angraecum, which had the longest nectariees (and the nectary varies much in length in some orchids), and which, consequently, compelled the moths to insert probosces up to the very base, would be fertilized. Those plants would yield "most seed" and the seedlings would generally inherit long nectaries; and so it would be in successive generations of the plant and of the moth. A candidate hawk-moth was discovered 40 years later: Xanthopan morgani praedicta. Darwin believed that plant fittness strongly correlated with the length of the nectary exceeding the pollinator tongue length, and this was corroborated in this case. A. sesquipedale exhibits an average floral-tube length of 28-32 cm (L.A.N. et al., unpublished results), while tongue length in X. praedicta reaches ~25 cm. It was found that insects do indeed insert their probosces no further than necessary to obtain nectar".

The moths were trained by similar looking plants before that specific Angraecum type evolved. The Angrecums or other orchids with long spurs succeeded to be pollinated by moths and a trend was therefore created. The unsuccessful Angraecum which the moths didn't fancy - didn't manage to get pollinated and that was eventually the end of their appearance on earth. Orchids are "copy-cats". They either copy flowers nearby (remember the ultra violet patterns, which we cannot see) or simulate odors or look like their pollinator, plant themselves on another plant and share the host plants pollinators (and benefit from pollinators making mistakes), etc., etc.

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MADAGASCAR POLLINATION EXPERIMENT

mplewinska
I have a question about the Madagascar pollination experiment. I don't understand whether the statement that only a few individuals were responsible for reproduction refers to the flowers or the insects.

(John in Arcadia CA
mplewinska - understood that to mean only a few individuals insects

sysser
Yes, John, that is so.

marylois
Going back to the Madagascar experiment: I cannot imagine hand labelling pollinia! Wonders of science.

mplewinska(Magda)
Re madagascar, what I really can't imagine is going back and reading all the labels!

Fleur( Tasmania)
I wonder how they did it, Lois. Microdots?

Sysser
"The 3.000 lables we used were small individually numbered pieces of thin microfilm, which we cut into 0,4 x 0,6 mm. The result was numbered labels of the size of 1/4 of a mm2. Later we dissolved the glue on ordinary lables with Cloroform and got a thinly running glue, which could be applied as microdrops on the film pieces. The lables were placed in 3.000 flowers' pollinia, 2 in each, without destroying or disturbing the flowers functions...L.A.N."

marylois
I'm hoping Marilyn could tell us -- I'd almost wager some atomic marking and reading???

marilyn001
I have colored Epipactis pollen with food coloring and watched the colored pollen deposited in dribs and dabs from flower to flower and plant to plant. It takes a steady hand to place 2 microlitres of dye on the pollinium without disturbing it.

nodosa
Greg - sesquipedale translates from Latin as foot and a half. I think the moth was Xanthopan morganii praedicta.

prankster d(Susan from Oregon)
Theres a guy here in town that did his thesis on ants, by gluing little nametags on each one.

graphicgreg
Helen, I love the name they gave the moth; Xanthopan morgani praedicta....yep, ole' Charlie sure did praedict it

marilyn001
I was so excited the first time the coloring experiment worked. Now the problem is finding which flower has been visited with what colored pollen: I use several colors.

Sparky-Steve
Color experiment sounds very interesting!!!!!

marilyn001
I wonder how they attached the microtag without disturbing the anther cap? No mention of that in the paper.

sysser
It is not mentioned in L. Anders Nilsson and Borge Pettersson paper. L. Anders Nilsson's address is: Department of Systematic Botany University of Uppsala - PO Box 541- S-751 21Uppsala, Sweden (they might be on Internet - worth a search]. Many universities are now linked up to the Net)

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VANILLA planifolia

Sparky-Steve
How about some feedback on pollinating vanilla. And is their pollinator extinct??

Sysser
It is pollinated by a bee (Melipona beechii) (Phil & Dodson)

nodosa
Steve - no mystery to vanilla. In Vanilla planifolia, the plant of commerce, there is a double septum. If one addresses the wrong one, the tube simply goes to a cul de sac. Do a transverse section of the flower and you will see the two chambers and immediately understand which is the proper one. The right one is further obscured by a membrane or flap over the stigmatic area. All is compounded by the short (6-8 hours) life of the flower. I've seen girls in the vanilla fields (it is a terrestrial vine) pollinate hundreds of flowers an hour by hand. There's bound to be a natural pollinator, but not one as fast as the girls and their "needles" as they call them.

Sparky-Steve
Ed, I do have a large planifolia, maybe 50 or so spikes this year. I have only gotten 3 mature pods that I did myself, and none done by any other pollinators. The funny thing is they're native to Florida!

nodosa
Steve - I've heard the native to Florida thing about vanilla, but I have more confidence in the "escaped from cultivation" stories. Seems some entrepreneurs set up an extensive V. planifolia range in S. Florida back in the 20's. The product was not of good quality and the processing cost (always labor intensive) was high. Add to this Watkins and others came out with some really good synthetics. The vines were abandoned and naturalized in a number of areas.

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POLLINATION ILLUSTRATIONS

graphicgreg
Steve, I have tried several times to photograph the sex-crazed bees on the sphacelatum but they are just too frenzied and move too fast

marylois
Are you speaking of those pesky "carpenter bees"?

JCY8S(John in Arcadia CA)
I have taken a catasetum to school so that I could illustrate the "shooting" of the pollinia

marylois
Phrag. caudatum petals grow 'til they touch something, I'm told, then stop. Reason being they grow 'til they touch the ground and then crawling insects use the petals to climb up....sort of a Rapunzel story. What crawling insects???

marilyn001
GraphicGreg! I am reading you loud and clear. I understand that Catasetums are visited by Euglossine bees which are perhaps orchid species specific. Right?

graphicgreg
According to Dr. Adams, Marilyn...right. What I find interesting is that these Euglossine bees collect Ctsm fragrance as an ingredient of their OWN sex lure

marilyn001
I have never had Catasetum flowers visited by local insects such as bumblebees and the like although these insects visit many other flowers such as cattleyas and oncidiums.

Sysser
All Catasetum (21 different kinds) mentioned by Phil & Dodson are all pollinated by bees (dif. sp. of Euglossa, Eulaema and Euplusia).

HelenH
John, that is a good thing to show kids...when I have one in bloom and Charlie has a bad day I let him vent pressure by triggering all the flowers...sometimes my cats get too close to the Catasetums and walk around with pollen all over them as it is very sticky.

JCY8S(John in Arcadia CA)
Helen - I am sure that it is very good therapy for Charlie!! I know that it is for me!

graphicgreg
More intriguing if you get too close !!! did you know that super glue is made from Catasetums ??? (just kidding)

JCY8S(John in Arcadia CA)
Greg, you may be kidding, but that Catasetum "glue" really is strong!

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ORANGE JUICE - pollination aid

nodosa
For what it is worth, when we find a plant that is difficult to pollinate, we try fresh squeezed orange juice on the stigmatic surface. Ups the take rate, as a rule. Probably as a result of ascorbic acid, but may have something to do with comets and spiral nebulae.

Sparky-Steve
Ed, you use that before you pollinate? what is the time frame?

nodosa
Steve - you apply the orange juice immediately before you apply the pollen. If there is any fresh orange juice left, you mix it with an equal amount of gin and take that yourself to steady your toothpick hand. Works every time.

Tarwood(Karen, NY)
Ed, does the acid attract the pollinator or does the acid change the pH and allow the take?

marylois
Good talk this evening, thanks to Sysser -- and good points
brought up by EVERYONE!
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BIBLIOGRAPHY

Behar, MD, Moises (1995). Evolution and Orchids. A.O.S. Bulletin. Dec. issue.

Dodson, Calaway H. (1991). Ecology and Pollination of the Orchids . Native Columbian Orchids, Vol. 3, Columbian Orchid Society, Compania Litografica National S.A., Medellin.

Dressler, Robert L. (1981). The Orchids - Natural History and Classification. Harvard Univ. Press.

Garay, Leslie A. (1974). The enigmatic Power of Orchids. A.O.S.Bulletin, May issue.

Herdon, Christopher N. (1996). Eulophiella. Jewels of Madagascar. A. O. S. Orchids, Feb. issue.

Johnson, Steve (1996). Pollination of satyrium species: lastest findings S.A.Orchid Journal, Volume: 27, no.3

Jones, David L. (1993). Native Orchids of Australia. The Biology of Native Orchids Reed/William Heinemann, Australia.

Manuel, R., Warren, R., Miller, D., Discussion by (1996) Sophronitis coccinea: A Pollination Study. A. O. S. Orchids, Jun. issue.

Morgan, Peter (1986/7) Aspects of Orchid Pollination nos. I, II, III. - 17, no. 4 - 18, no. 1.

Nilsson, L. Anders (1988). The evolution of flowers with deep corolla tubes Nature, Volume: 334, no. 14.

Nilsson, L. Anders, Elisabeth Rabakonandrianina & Borge Pettersson (1992). Exact tracking of pollen transfer and mating in plants. Nature, Volume: 360, no. 17.

Nilsson, L. Anders (1992). Orchid Pollination Biology. TREE vol. 7, no. 8, August.

Nilsson, L. Anders and Pettersson, Börge (1995) Vem parer sig med vem blandt växterna?
Dansk Orkide Klub, Volume 16, no. 3. (translation:" Who mates with who amongst the plants?")

Northen, Rebecca T. (1990). Home Orchid Growing. Simon & Schuster.

Pijl , L. van der and Dodson, C. H. (1969 ). Orchid Flowers, Their Pollination and Evolution Univ. of. Miami Press.

Pridgeon, Dr. Alec M. (1989) Lips. S.A.Orchid Journal, Volume: 20, no. 4.

Withner, Carl L. (1974). The Orchids - Scientific Studies. John Wiley & Sons. Inc.

Personal conversation and correspondence:

Behar, Moises, Guatemala
Dressler, Robert L., Florida
Garay, Leslie A., Blacksburg, VA
Johnson, Steve, Rondebosch, Cape Town
Light, Marilyn H.S., Ottawa, Canada

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