# Slipper orchid evolution: does anyone care?



## VAAlbert (Dec 6, 2006)

Hi all,

I saw a post somewhere in another thread that stated something to the effect that the various and sundry details of evolution and 'precise' taxonomy of the slippers wasn't so important to him/her, since he/she felt him/herself more of a horticulturalist. Pls. correct me if I'm wrong!

Well, I'll bet that some of you out there are interested in slipper orchid evolution, like I am. I'd love to discuss issues from hobbyist to advanced levels. My previous work with slippers has been in molecular phylogenetics and taxonomy; my present research includes evolutionary developmental genetics (the study of the genes behind morphological features) and comparative genomics (evolutionary studies of large numbers of genes among species and what they tell us). Plus some molecular phylogenetics, currently in the mint family.

Best wishes,

Vic.


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## Mark (Dec 6, 2006)

"does anyone care?"

I do.


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## kentuckiense (Dec 6, 2006)

Evolutionary biology fanatic checking in!


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## NYEric (Dec 6, 2006)

I would be interested in a summation or short version. [I have the attention span of a gnat.] I worked for a religious man who stated that evolution did not happen!!! Scary.


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## slippertalker (Dec 6, 2006)

I would be interested in your ideas. Most of us are not taxonomists or experts at molecular studies but are interested in the concepts of evolution in orchids. Bring it on!


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## Ernie (Dec 6, 2006)

Sign me up too. 

-Ernie


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## gore42 (Dec 6, 2006)

I certainly care! 

- Matthew Gore


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## gonewild (Dec 6, 2006)

VAAlbert said:


> Hi all,
> 
> My previous work with slippers has been in molecular phylogenetics and taxonomy; my present research includes evolutionary developmental genetics (the study of the genes behind morphological features) and comparative genomics (evolutionary studies of large numbers of genes among species and what they tell us). Plus some molecular phylogenetics, currently in the mint family.
> 
> ...



Are your studies and research focused on evolution between species or are you looking at hybrids also?

Can you separate the different species or their hybrids by looking at their genes?


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## VAAlbert (Dec 6, 2006)

Glad to see the interest! Evolution matters, cause its products leave us with plants that want taxonomies.

Re: my present research, it isn't on slippers - I may have given the wrong impression through some twisted English. But the principles are basically the same as if I were studying orchids, the legume family, insects, or mammals.

So, I am doing genomic-scale work at the species level among the Hawaiian endemic mints. I am involved in genomics studies at a much higher level as well, involving the role that whole-genome duplications (all genes at once!, i.e., polyploidy) might have had to do with the evolution of flowers. (For example, it is known that ray-finned fish -- the bony ones -- are 'paleo'polyploids, and this group of fish is the most diverse vertebrate group on Earth; idea = that having extra copies of whole genomes opens up opportunity for evolutionary 'flexibility'). I am also doing molecular developmental work on the cut-flower crop Gerbera, in the sunflower family. In that family, we have the unique opportunity to look at the genetics of flower morphological differences within the SAME GENOTYPE, since the flowering heads of Gerbera bear different flower types (female-only and with a highly expanded petal 'lip', female-only with only moderately expanded petals, and male/female with short petals not really lipped).

Just for fun, you can see some background on these subjects at the following links:

http://www.biomedcentral.com/1471-2229/6/16
folk.uio.no/victoraa/Cui_2006.pdf
folk.uio.no/victoraa/Teeri_BioEssays_2006.pdf

Warning! These are highly technical works! 

*I'm very happy to try to get any thread going that any of you may be interested in re: slipper evolution.* And, no real need for high tech talk at all.

Best wishes,

Vic


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## VAAlbert (Dec 6, 2006)

Forgot to mention that I don't directly intend to work on hybrids, but that I do by necessity among the Hawaiian mints, which like to spread their genes around sometimes.

V


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## gonewild (Dec 6, 2006)

VAAlbert said:


> Forgot to mention that I don't directly intend to work on hybrids, but that I do by necessity among the Hawaiian mints, which like to spread their genes around sometimes.
> 
> V


Is your work on Gerbera also at the specie level?


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## gonewild (Dec 6, 2006)

VAAlbert said:


> *I'm very happy to try to get any thread going that any of you may be interested in re: slipper evolution.* And, no real need for high tech talk at all.
> 
> Best wishes,
> 
> Vic



Great! In low tech talk.....

Are all slipper orchids evolved from a single slipper ancestor?


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## kentuckiense (Dec 6, 2006)

This is awesome! Thanks!

Anyway, my main interests are in the following:

1. The biogeography of the genus, especially in terms of species pairs (reginae and flavum, arietinum and plectrochilum, californicum and subtropicum, etc.)

2. The parallel morphology of the 'basal' clades: Parvisepalum Paphs, Micropetalum Phrags, most Cyps(save guttatum and yatabeanum), Mexipedium, and Selenipedium. You know, the inturned labellum folds, etc.


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## VAAlbert (Dec 6, 2006)

gonewild said:


> Is your work on Gerbera also at the specie level?



No, we use only one variety, Terra Regina, in our work.

V


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## VAAlbert (Dec 6, 2006)

gonewild said:


> Are all slipper orchids evolved from a single slipper ancestor?



Absolutely. For sure. No doubt about it!

If we think about the slipper family tree, we can imagine the slipper 'common ancestor' had leaves something like a Selen or a Cyp, and flowers like most Cyps (inflated pouches), parvi Paphs, micropetalum Phrags, and Mexipedium. Paph, Phrag, and Mex all have a common ancestor of their own, but we're still not clear on the branching order betw. these three genera.

V


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## VAAlbert (Dec 6, 2006)

kentuckiense said:


> This is awesome! Thanks!
> 
> Anyway, my main interests are in the following:
> 
> ...



Great! Re: 1., quite a number of other plants have this N. America/Asia disjunction. No surprises at all there if the species named are really sister pairs.

Re: 2., I've gone into this a little before, and can say a little now based on the anatomical sequence one sees in developing buds of these slips and others that have different pouch types. Suffice it to say that the other pouch types (e.g., Barbata paphs, Lorifolia Phrags, and Cyp guttatum -- each in a different way) can all be pretty easily derived as showing different developmental stages *beyond* what one sees in the inflated-pouch type, the morphological development of which simply *stops* at that stage.

Best again,

Vic


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## silence882 (Dec 6, 2006)

I am interested as well!

One issue I have been wondering about is how molecular phylogenetics deals with placing species that may have arisen from a population of natural hybrids?

For example, Paph. hangianum may have started as a natural hybrid between Paph. emersonii and Paph. malipoense and then evolved into the 'current' species. If this were the case, how would Paph. hangianum fit into a cladogram?

--Stephen


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## kentuckiense (Dec 6, 2006)

VAAlbert said:


> Re: 2., I've gone into this a little before, and can say a little now based on the anatomical sequence one sees in developing buds of these slips and others that have different pouch types. Suffice it to say that the other pouch types (e.g., Barbata paphs, Lorifolia Phrags, and Cyp guttatum -- each in a different way) can all be pretty easily derived as showing different developmental stages *beyond* what one sees in the inflated-pouch type, the morphological development of which simply *stops* at that stage.



So would you tend to chalk that one up to converging/diverging pollination syndromes? IE: the species that have been hypothesized to to mimic other rewarding plants (Paph. micranthum looking like a Rhododendron, etc.) have infolded labellums to trap entering bees while the others (Most non-Parvi Paphs) are structured in order to facilitate flies slipping into the labellum?


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## VAAlbert (Dec 6, 2006)

silence882 said:


> For example, Paph. hangianum may have started as a natural hybrid between Paph. emersonii and Paph. malipoense and then evolved into the 'current' species. If this were the case, how would Paph. hangianum fit into a cladogram?



Perhaps not very well at all, unless the potential hybridization event took place long enough ago for genetic variation to have 'sorted' itself out. And this sorting out can indeed be analyzed at the population genetic level.

V


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## kentuckiense (Dec 6, 2006)

silence882 said:


> I am interested as well!
> 
> One issue I have been wondering about is how molecular phylogenetics deals with placing species that may have arisen from a population of natural hybrids?
> 
> ...


We talked about that in class a while back. That concept was one of the reasons why Cronquist was opposed to a PURE monophyletic taxonomy of plants. Most taxonomists don't even want to think about the amount of plants that arose from polyploid hybrid ancestry, no?


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## VAAlbert (Dec 6, 2006)

kentuckiense said:


> So would you tend to chalk that one up to converging/diverging pollination syndromes? IE: the species that have been hypothesized to to mimic other rewarding plants (Paph. micranthum looking like a Rhododendron, etc.) have infolded labellums to trap entering bees while the others (Most non-Parvi Paphs) are structured in order to facilitate flies slipping into the labellum?



I would say that the inflated pouch character is a 'primitive' one, inherited from the slipper common ancestor, which probably relied on some sort of bee pollination. That would make the occurrence of fly pollinated flowers the subject of convergent evolution -- in Paphs (the lineage including all species with pouch frontal margins that turn outwards, e.g., the barbatas), Phrags (lorifolias et al.), and at least 2 groups of cyps (margaritaceum group; guttatum, e.g.). It is actually quite remarkable to find a group of plants that 'evolves' a fine-tuned trapping/pollination system for bees, which are comparatively 'smart', only to evolve further to trap stupid things like flies. In other words, slippers became opportunists. Another group of plants that had this same transition, i.e., bee-->fly pollination made its way onto the pages of one of the top journals in biology: _Nature_! And I think this other example only showed ONE shift from bees to flies!

BW,

Vic


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## VAAlbert (Dec 6, 2006)

OK, not bees to flies, but still bees (resin collectors) to more generalized pollination syndromes.

Nature 394, 632 (13 August 1998); doi:10.1038/29210

Switch from specialized to generalized pollination

W. SCOTT ARMBRUSTER1 AND BRUCE G. BALDWIN2

1 Institute of Arctic Biology, University of Alaska, Fairbanks, Alaska 99775, USA and Department of Botany, Norwegian University of Science and Technology, N-7034 Trondheim, Norway
e-mail: [email protected]
2 Jepson Herbarium and Department of Integrative Biology, University of California, Berkeley, California 94720, USA

The once prevalent view that the evolution of extreme ecological specialization is accompanied by a loss of potential for adapting to new conditions, and thus is irreversible, has been challenged by several recent examples,,. However, we know of no modern phylogenetic studies showing reversal in pollination relationships from extreme specialization to generalization, although such reversals are theoretically expected,. Here we present molecular phylogenetic evidence for an evolutionary shift in Dalechampia (Euphorbiaceae) vines from a highly specialized relationship (pollination by one or a few animal species,) with resin-collecting bees to generalized pollination by a variety of pollen-feeding insects. This shift was associated with dispersal from Africa to Madagascar, where the specific resin-collecting pollinators are absent. These results show that plants dispersing beyond the range of their specific pollinators may succeed by evolving more generalized pollination systems.


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## Heather (Dec 6, 2006)

I'm late, but I'm definitely interested too! 

So far so good...very glad you joined us Vic!


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## gonewild (Dec 6, 2006)

VAAlbert said:


> Perhaps not very well at all, unless the potential hybridization event took place long enough ago for genetic variation to have 'sorted' itself out. And this sorting out can indeed be analyzed at the population genetic level.
> 
> V



How many generations would "long enough ago" be to sort out a natural hybrid as a distinct specie? Can this happen rapidly or does this require a great span of time?


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## silence882 (Dec 6, 2006)

VAAlbert said:


> Perhaps not very well at all, unless the potential hybridization event took place long enough ago for genetic variation to have 'sorted' itself out. And this sorting out can indeed be analyzed at the population genetic level.
> 
> V



hmmmm

So what would the presence of a natural hybrid in a pool of species do to the cladogram that's generated using molecular phylogenetics? Would the presence of a hybrid mess up the results for the rest of the species?

--Stephen


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## Rick (Dec 6, 2006)

VAAlbert said:


> I would say that the inflated pouch character is a 'primitive' one, inherited from the slipper common ancestor, which probably relied on some sort of bee pollination. That would make the occurrence of fly pollinated flowers the subject of convergent evolution -- in Paphs (the lineage including all species with pouch frontal margins that turn outwards, e.g., the barbatas), Phrags (lorifolias et al.), and at least 2 groups of cyps (margaritaceum group; guttatum, e.g.). It is actually quite remarkable to find a group of plants that 'evolves' a fine-tuned trapping/pollination system for bees, which are comparatively 'smart', only to evolve further to trap stupid things like flies. In other words, slippers became opportunists. Another group of plants that had this same transition, i.e., bee-->fly pollination made its way onto the pages of one of the top journals in biology: _Nature_! And I think this other example only showed ONE shift from bees to flies!
> 
> BW,
> 
> Vic



Ah. We've had this discussion before.:clap: I've found some new info to add to this discussion. The shift to flies could be be construed to actually be an advance into a very specific niche. The type of flies utilized by slippers are hover flies (and there are lots of species to get plant specific associations too), and the mimic is very specific to lure female flies to lay eggs against a potential (fake) food source for its larvae (esentially a predatory flie). Also the visual acuity and discerning capabilties of flies is superior to bees (this is the newer info I read), so bees are more on autopilot than syrphid flies in finding their particular resources. Subsequently it could be construed that its easier to trick a bee than a flie.

Another way to look at this is that bees and plants go a long ways back in development of a cooperative arangement (food for pollen transport). So its not that far of a stretch to see a plant species break into the system ((maybe for energetic reasons (no food production required)) to get some free pollen transport from a pretty dependable engrained mass transit system.

The hover fly system is quite a break from this as there is no preexisting plant -flie mutual arangemnt that I can think of. In the case of fly pollenation its a 2 step process in that plants have a negative association with aphids (fly bait), and flies have an association with aphids (prey). So the plant came up with a way to get the two associations together (with a criter that's not on autopilot with good visual acuity). Now that's advanced.


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## VAAlbert (Dec 7, 2006)

Rick said:


> The shift to flies could be be construed to actually be an advance into a very specific niche. The type of flies utilized by slippers are hover flies (and there are lots of species to get plant specific associations too), and the mimic is very specific to lure female flies to lay eggs against a potential (fake) food source for its larvae (esentially a predatory flie).



This does seem to be the case in Paph rothschildianum, though I'm not aware of similar results from other slippers. I have to go back to the Paph villosum example, but I don't think the conclusion was brood-site deception.



> Also the visual acuity and discerning capabilties of flies is superior to bees (this is the newer info I read), so bees are more on autopilot than syrphid flies in finding their particular resources. Subsequently it could be construed that its easier to trick a bee than a flie.



Certainly, some bees are bumblers, but I think we'll have to look through some of the literature on Cyp to see whether bumble bees are the predominant pollinators! Other bees are far more specific pollinators. It could well be that Paph micranthum, e.g., is pollinated by bumble bees; it has such a huge pouch.



> Another way to look at this is that bees and plants go a long ways back in development of a cooperative arangement (food for pollen transport). So its not that far of a stretch to see a plant species break into the system ((maybe for energetic reasons (no food production required)) to get some free pollen transport from a pretty dependable engrained mass transit system.
> The hover fly system is quite a break from this as there is no preexisting plant -flie mutual arangemnt that I can think of. In the case of fly pollenation its a 2 step process in that plants have a negative association with aphids (fly bait), and flies have an association with aphids (prey). So the plant came up with a way to get the two associations together (with a criter that's not on autopilot with good visual acuity). Now that's advanced.



A great idea! Brood-site deception could certainly be viewed as an advanced trait. However, I question whether brood-site deception occurs with all of the outwardly-turned pouch Paphs, e.g. The staminodia of Paph villosum, e.g., and rothschildianum, e.g., would then seem to me to be different sorts of attractants, perhaps to different sorts of flies.

One problem with brood-site deception being a major force in Paph evolution (e.g.) is that the Barbata slippers, which are by far the most numerous fly-pollinated species, don't have have the roths-type staminode features (aphid nest -like) and they rarely occur in sympatry (in the same place), leading one to suspect that the different floral forms (incl. staminodia) MAY have evolved in allopatry (geographical isolation, e.g., in small populations). Of course, I don't KNOW the latter, but it makes some deductive sense to me.

There are broadly distributed, outwardly-turned pouch Paphs, but we can't yet say whether these distributions result from range expansion, or whether they are relictual. In other words, I wonder whther staminodia of Barbatas eally are finely adapted to different pollinators at all.

But, I digress -- you are perfectly logical at the HIGHER (sectional) level (in taxonomy, which happens to reflect phylogeny), e.g., in the ancestor of Barbatas, if their pollination system could be considered advanced as you intriguingly suggest! There is also a nice paper out recently on deception of Cyp guttatum flowers; I'll look that one over again.

Regardless, I think we can probably agree that the slipper orchids have become opportunists re: fly pollination!

best wishes,

Vic


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## kentuckiense (Dec 7, 2006)

VAAlbert said:


> Certainly, some bees are bumblers, but I think we'll have to look through some of the literature on Cyp to see whether bumble bees are the predominant pollinators! Other bees are far more specific pollinators. It could well be that Paph micranthum, e.g., is pollinated by bumble bees; it has such a huge pouch.



Cypripedium acaule is bumblebee pollinated. I'd wager that C. kentuckiense is as well. The orifice is simply gigantic!


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## kentuckiense (Dec 7, 2006)

Looks like Cypripedium macranthos var. rebunense may be bumblebee pollinated as well. Check out:

"Bumblebee pollination of Cypripedium macranthos var. rebunense (Orchidaceae); a possible case of floral mimicry of Pedicularis schistostegia (Orobanchaceae)" by Sugiura et al.


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## VAAlbert (Dec 7, 2006)

> I wonder whther staminodia of Barbatas eally are finely adapted to different pollinators at all.



Hi again,

Thanks for your input Kentuckiense! BTW, re: the above, 2 things:

- I don't mean to say that Barbata Paphs don't occur in the same geographic regions, but it seems to me that they usually occur isolated from one another in terms of 'microgeography', e.g., even elevation on the same mountain, or on differnt rock substrates on the same little peninsula.

- A 'radiation of forms', such as we do see with the barbatas, and yes, to a degree, with their staminodes, need not be an 'adaptive radiation'. Evolutionary change, and fixation of new morphological forms, can certainly take place in the absence of natural selection on those forms. 

The latter is is in fact exactly what I'm working on with the Hawaiian endemic mints! Such radiations can occur by random chance, when certain alleles (versions; mutants) of different genes become dominant in small populations simply because the small population size favors the fixation of any mutant that comes up, be it a 'good' one or a 'bad' one. Random changes like these can of course become the subject of natural selection LATER, and this has certainly happened in some other systems.

So, while the barbatas are probably pollinated by flies, they need not be pollinated by different species of flies, or even be adapted to different species of flies; the Barbata species could simply be isolated from one another by space or even flowering time, and their distinct (sometimes!!!!) morphologies amongst each other could be 'frozen accidents' of seed dispersal and fixation of new genotypes.

For example, Paph curtisii(/superbiens) can co-occur with Paph tonsum, and the natural hybrid is known. Reproductive isolation can be leaky, and bugs can go after whatever they find. 

Clearly more to talk about here - I'm having fun!

All best,

Vic


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## VAAlbert (Dec 7, 2006)

It seems that Cyp guttatum serendipitously traps bees that are otherwise after other nearby flowers:



> Pollination of a slippery lady slipper orchid in south-west China: Cypripedium guttatum (Orchidaceae)
> HANS BÄNZIGER1,2 , HAIQIN SUN1 and YI-BO LUO1*
> 
> Cypripedium guttatum was studied in north-west Yunnan at 3490 m a.s.l. The flowers are rewardless 'kettle traps'. The structure of the lip, where pollinators are temporarily kept prisoner, and the method of their capture, are unusual in being Paphiopedilum- rather than Cypripedium-like. The deceptive orchid does not mimic any of the diverse flowers concurrently blooming in the habitat, all being visited by the polylectic pollinators of C. guttatum, viz. Lasioglossum virideglaucum, L. clypeinitens and L. sauterum, besides two additional probable pollinators and four non-pollinating visitors (all Halictidae; three new species). The bees got caught when they tried to climb onto the staminode and their forelegs slid down its slippery downward ridges, causing them to tumble to the pouch bottom. To leave, they had to climb a tunnel leading past the stigma to the anthers where a pollen smear was acquired while extruding themselves from the narrow exit. The similarities with myiophylous Paphiopedilum are discussed in view of the possibility that they may foreshadow evolutionary transitions between melittophily and myiophily found in slipper orchids. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 148, 251–264.



Cyp guttatum does have an inwardly folded lip, but the infold is basically fused to the other side of the lip surface, such that the pouch very much resembles that of an (e.g.) Barbata Paph. Cyp guttatum does not come out in a 'primitive' position in the published phylogenetic trees on slippers.

Best,

V


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## VAAlbert (Dec 7, 2006)

Re: Paph villosum:



> The mesmerizing wart: the pollination strategy of epiphytic lady slipper orchidPaphiopedilum villosum(Lindl.) Stein (Orchidaceae)
> 
> HANS BÄNZIGER
> 
> ...



This would seem to support the 'dumb fly' conjecture for many Paph species that have outfolded lips but don't have glandular, rothschildianum-like staminodes.

V


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## VAAlbert (Dec 7, 2006)

BUT, here is more from the Paph villosum paper! -->



> The slippery methods with which C. guttatum captures its pollinators contrasts with those in its congeners C. yunnanense and C. flavum. In these the bees crawl down the wide infolded flaps of the pouch, which are not slippery, and then enter the interior much like they enter a tubular flower such as Salvia brachyloma (unpubl. observ., this study). The staminode plays no direct role in capture, though it may initially function as a false nectar guide in attracting the bees from a distance.
> 
> The pollination strategies of C. guttatum are reminiscent of those seen in Paphiopedilum spp., although in the latter case the pollinators are hoverflies. The staminode has a direct and pivotal role in triggering the pollinator's capture; its slippery surface and adjunct flight interfering devices dispatch pollinators into the pouch. The staminode operates in three different ways.
> 
> In the first type, found in P. rothschildianum and P. callosum, the pollinators manage to grip the staminode for a few seconds while laying an egg before losing their hold. In the second, observed in P. villosum and P. charlesworthii, food-seeking pollinators try to grip a very slippery wart protruding from the staminode, losing their hold instantly. In the third, found in P. parishii and P. bellatulum, they grip the staminode's flattish surface, also without intent of laying, and slide off shortly afterwards. In the first type the pollination syndrome is based on 'perfidious' brood-site deception (leading to the death of the pollinator's progeny); in the second and third types it is based on 'opportunistic' food deception, possibly with a faint brood-site deception also at work (but without deleterious effects)....



Seems there is quite some diversity in lsipper pollination, much of it not published in detail.

Best again,

Vic


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## VAAlbert (Dec 7, 2006)

silence882 said:


> So what would the presence of a natural hybrid in a pool of species do to the cladogram that's generated using molecular phylogenetics? Would the presence of a hybrid mess up the results for the rest of the species?



Well, it can certainly screw things up. For example, when we've looked at the Hawaiian mints, which are a VERY young group of plants, we see what could be termed 'intergeneric hybrids', but still, we are not sure if these genera are natural in the first place. Regardless, the hybrid nature of these plants is real, since we get 2 different genotypes for plants with the same morphology; the cladogram places them on different parts of the tree.

There has been some theoretical work on the influence of hybridization on cladograms, and I am certainly not the first to have seen and published on this in other plants!

Regarding how many generations it would take for a 'hybrid' population to stabilize into separate 'species', that would depend entirely on the impact level of either random processes (e.g., in small populations, as I described in an earlier post) or natural selection. And we can't get a better idea of this without detailed experiments in the field and the lab.

Bottom line, however, is that hybrid speciation is known to occur in plants. It usually is accompanied by polyploidization, but NOT ALWAYS.

Regards,

Vic


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## NYEric (Dec 7, 2006)

kentuckiense said:


> This is awesome! You know, the inturned labellum folds, etc.


STOP! You're making me all sticky! :drool:


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## Rick (Dec 7, 2006)

VAAlbert said:


> BUT, here is more from the Paph villosum paper! -->
> 
> 
> 
> ...



You also need to look at the natural history of the pollinators. The genera mentioned are all Syphidae and all use aphids as a larvae food source. Typically the males are atracted to flowers for nectar and pollen (just like bees). However it was noted that the villosum pollinators were females (ie looking for brood sites). In the case of villosum and species with similar staminodal structures (as opposed to roth and related fuzzy staminodes). The staminode looks like it has a big aphid (or aphid producing honeydew) in the center of it, and would subsequently count as another form of brood dececption as alluded to in posibility in the paper.


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## Rick (Dec 7, 2006)

Another factor in pollinator specificity would be homerange size of pollinator.

For example, many of the native bees in the US are territorial and have relatively small well defined home ranges. (subsequently passing pollen around a small number of plants).

The imported honey bees are far less discriminant, have very large ranges without well defined borders.

This has reshaped some plant demographics in the US since the introduction of hiving honey bees.


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## VAAlbert (Dec 7, 2006)

Thanks, Rick!

There's clearly a lot to learn about slipper natural history.

best,

Vic.


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## Rick (Dec 7, 2006)

What's the weather like in Oslo today?

We had a high of 35 until the front went through, and tonights low is supposed to be 16:sob: 


I'll be burning propane tonight!!


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## Braem (Dec 8, 2006)

Hi Victor,

Evolution is important and off course evolution of orchids is of special importance to us orchid freaks. However, it would be of special interest that you explain the methods used and explain what the results of those methods tell us, and what they don't tell us. 

regards
Guido



VAAlbert said:


> Hi all,
> 
> I saw a post somewhere in another thread that stated something to the effect that the various and sundry details of evolution and 'precise' taxonomy of the slippers wasn't so important to him/her, since he/she felt him/herself more of a horticulturalist. Pls. correct me if I'm wrong!
> 
> ...


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## Braem (Dec 8, 2006)

It is nice and cosy in my newly furbished office and I am getting the heat out of the outside air. No gas, no oil, just air! And it is about 54 F in Lahnau on this 8th of December!

Guido



Rick said:


> What's the weather like in Oslo today?
> 
> We had a high of 35 until the front went through, and tonights low is supposed to be 16:sob:
> 
> ...


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## Heather (Dec 8, 2006)

Finally, it is unpleasantly cold here this morning, only 18°F.


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## NYEric (Dec 8, 2006)

I closed the windows in the front room [with the plants] because of the forecast - teens and windy - and it got so warm from the heat I had to open a window for a while to air out. NYC: Sunny and cold.


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## VAAlbert (Dec 8, 2006)

Braem said:


> ...it would be of special interest that you explain the methods used and explain what the results of those methods tell us, and what they don't tell us.
> Guido



OK, sounds great! I'll explain everything I can, and as simply as possible. It will help me focus this discussion if you all could ask some specific questions that you may be wondering how/if/what can be done. Remember, I am not specifically doing genomic, molecular developmental, or population genetic work on slippers now, but instead on other plants. 

However, just since joining this forum, we have started to try getting some more DNA sequence data to help corroborate / not corroborate our previous phylogenetic work re: the placement of Mexipedium! Results from broad taxonomic sampling have so far been published for only one locus (gene, or if not a gene, some specific DNA region), and the alignment of the DNA sequences (the bases A, T, G, and C -- very important!) was ambiguous & perhaps unintentionally biased, not to mention the fact that the analysis presented in the paper (Cox et al.) was not done as well as it could have been. Specifically, the application of the particular character weighting procedure has now been shown to to be flawed and no longer applicable to any data. Regardless, even without that procedure, Mex stays where Cox et al. put it (just as where I put it in 1992 in my Mex description paper, using yet another locus).

But the jury should remain out, and I hope we can get a few more loci to add to the question.

Regarding more data on species-level issues, this will depend on how much information new sequences could provide. No promises for either the Mex goal or this possibility; we are working on 'side money' from a grant to do other research, so our expenditures will have to be limited. Plus, there's no guarantee we can actually obtain new sequences that will provide meaningful enough amounts of evolutionary divergence between them.

I'll be happy to talk more about other topics; please try to help by throwing some out for me. 

By the way, the weather here is unseasonably mild, and we have had nothing but rain for the last 2 weeks! There is snow in the mountains, of course, but in Oslo we get the 'maritime effect', but usually not this late in the season.

All best,

Vic


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## Braem (Dec 9, 2006)

Victor,

excellent, lets keep this discussion going. I will certainly come up with some questions. I would love to see the new results. 

I think, however, that very few people on this forum are familiar with genetic terminology (please correct me if I am wrong). Therefore, it may be good to explains the basics first. 

regards
Guido



VAAlbert said:


> OK, sounds great! I'll explain everything I can, and as simply as possible. It will help me focus this discussion if you all could ask some specific questions that you may be wondering how/if/what can be done. Remember, I am not specifically doing genomic, molecular developmental, or population genetic work on slippers now, but instead on other plants.
> 
> However, just since joining this forum, we have started to try getting some more DNA sequence data to help corroborate / not corroborate our previous phylogenetic work re: the placement of Mexipedium! Results from broad taxonomic sampling have so far been published for only one locus (gene, or if not a gene, some specific DNA region), and the alignment of the DNA sequences (the bases A, T, G, and C -- very important!) was ambiguous & perhaps unintentionally biased, not to mention the fact that the analysis presented in the paper (Cox et al.) was not done as well as it could have been. Specifically, the application of the particular character weighting procedure has now been shown to to be flawed and no longer applicable to any data. Regardless, even without that procedure, Mex stays where Cox et al. put it (just as where I put it in 1992 in my Mex description paper, using yet another locus).
> 
> ...


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## SlipperFan (Dec 9, 2006)

Braem said:


> ...Therefore, it may be good to explains the basics first.


I, for one, would appreciate that! My background is in art and photography, not science, but I find this interesting.


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## ScottMcC (Dec 10, 2006)

I think that both of the esteemed doctors should make new threads explaining the basics of their respective subjects. I've got a BS in Biochem, an MD, and a wife who's a few months away from her PhD in biochem, and a fair number of the things that have been discussed have been way over my head.

So please, if you could both start new threads explaining the main "tools of the trade" in taxonomy and genetics (at least the parts of both fields in which you're involved), I'd really appreciate it. No need to dumb things down, just start at the beginning--assume that I'm semi-competent at absorbing information.


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## Heather (Dec 10, 2006)

That would be nice, actually, for all of us and especially any members who might be feeling a bit intimidated by these threads.


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## Braem (Dec 10, 2006)

Scott,

the problem is that we certainly have a mixture on the forum (and this is not meant to be an insult to anyone) I don't want to dumb things down for anyone, but to be sure, we might have to start pretty basic. I hope Victor comes in on this, and maybe Victor and I can agree on a procedure off forum. (Victor?) 

The easier thing (I think) would be that you people ask specific questions, and I for my part will then try to be as explicit as possible. 

regards
Guido




ScottMcC said:


> I think that both of the esteemed doctors should make new threads explaining the basics of their respective subjects. I've got a BS in Biochem, an MD, and a wife who's a few months away from her PhD in biochem, and a fair number of the things that have been discussed have been way over my head.
> 
> So please, if you could both start new threads explaining the main "tools of the trade" in taxonomy and genetics (at least the parts of both fields in which you're involved), I'd really appreciate it. No need to dumb things down, just start at the beginning--assume that I'm semi-competent at absorbing information.


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## Braem (Dec 10, 2006)

Heather,

no one should be intimidated by these treads. We alls have different backgrounds and all have different levels of education etc. That is nothing to be ashamed or afraid about. I, for my part am a complete nut when it comes to maths, computers (and many other things). But then. I am a good general biologist, a good botanist, and I am pretty good at Art History and English literature, and I managed to keep a wife over 32 years ... (poor girl). Not bad for a young lad like me . 

Guido



Heather said:


> That would be nice, actually, for all of us and especially any members who might be feeling a bit intimidated by these threads.


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## Rick (Dec 10, 2006)

VAAlbert said:


> Hi again,
> 
> Thanks for your input Kentuckiense! BTW, re: the above, 2 things:
> 
> ...




After checking out a bunch of pictures of aphid colonies it occured to me I can see a clear resemblance of the pimply/warty area of the pouch behind the staminode of many barbata species. This pimply area is especially prominent in wardii, sukhakulii, and dayanum. After spending just a few minutes looking up natural history of aphids, there seems to be allot of aphid species specialized on certain species of plants.

This provides another hypothesis that barbata are still mimiciing other plant species by mimicing an infected segment of these plants.

This is pretty wild!!


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## VAAlbert (Dec 10, 2006)

Fascinating! If you can compile some photos and even try to match them up with a Paph or two, that would be really neat to see.

Re: discussion of evolution, taxonomy, genetics, etc., some of the interchanges we've had on these threads have been a little complex, I admit! But many of these have been specific chats betw. me and certain other folks, and although I can hope you've been interested in reading them over, I surely understand if they haven't given you much to go away with.

As I've indicated before, I think the best way to proceed on a simpler, more explanatory level would be if any of you guys could throw out as specific questions/topics as possible. It'll be hard for me to address questions like 'what can genetics do for understanding slipper orchid evolution?' since this is such a broad, multifaceted topic. If you were to ask something like 'How could you use genetic data to understand why Paph delenatii and Paph wardii petals look different?', even though that's also complicated, I could begin to narrow things down and try to start at the basics for such a point.

I'm very happy to talk about taxonomy and nomenclature as well.

Please don't be intimidated! There's lots to talk about, and lots of levels to talk about stuff at! :rollhappy: 

All best wishes,

Vic 




Rick said:


> After checking out a bunch of pictures of aphid colonies it occured to me I can see a clear resemblance of the pimply/warty area of the pouch behind the staminode of many barbata species. This pimply area is especially prominent in wardii, sukhakulii, and dayanum. After spending just a few minutes looking up natural history of aphids, there seems to be allot of aphid species specialized on certain species of plants.
> 
> This provides another hypothesis that barbata are still mimiciing other plant species by mimicing an infected segment of these plants.
> 
> This is pretty wild!!


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## gonewild (Dec 10, 2006)

Do you have an assumption as to how many years (or time period) orchids with pouches have been evolving. 
How much time is represented in the genetics you are looking at? 
Are there fossil records of orchids with pouches?


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## Braem (Dec 10, 2006)

Lance et alia,

as far as I know there is no fossil whatsoever that can be determined positively as being an orchid. Some decades ago, there were a few things published as "orchid fossils". In fact, they could have been just about anything you want to imagine plant like. 

The (time) origin of the "appearance" of flowering plants is under debate. About five years ago (I am doing this from memory, so don't nail me down on a few years) there was an article in "Nature" or "Science" (I must have it somewhere) about a fossil plant from which the authors deduced that it was a flowering plants more than 100 million years old. 

All about the molecular genetics, I will leave to Victor.

Guido






Maybe Victor has more on this.



gonewild said:


> Do you have an assumption as to how many years (or time period) orchids with pouches have been evolving.
> How much time is represented in the genetics you are looking at?
> Are there fossil records of orchids with pouches?


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## Rick (Dec 10, 2006)

gonewild said:


> Do you have an assumption as to how many years (or time period) orchids with pouches have been evolving.
> How much time is represented in the genetics you are looking at?
> Are there fossil records of orchids with pouches?



I was trying to frame up a similar question, (so I hope I am not coopting or missrepresenting your question Lance).

Human DNA studies have used rates of mutation crossed referenced to the number of mutations in a loci to come up with some dates in human evolution. Has anything comparable done with slippers?


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## Rick (Dec 10, 2006)

VAAlbert said:


> Fascinating! If you can compile some photos and even try to match them up with a Paph or two, that would be really neat to see.
> 
> Vic



This has been fascniating. I'm thinking of putting together a presentation for our orchid society. I'm going to need to do something better than the thumbnails of native bugs though.


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## VAAlbert (Dec 11, 2006)

Rick said:


> I was trying to frame up a similar question, (so I hope I am not coopting or missrepresenting your question Lance).
> 
> Human DNA studies have used rates of mutation crossed referenced to the number of mutations in a loci to come up with some dates in human evolution. Has anything comparable done with slippers?



Hi Rick et al.

Indeed, molecular 'dating' of the origins of plant groups has been attempted, and an estimate for the origin of Orchidaceae is ca. 50 million years ago. A better way of putting this is that the molecular phylogeny suggests that the divergence between the lily relative Astelia and Apostasia (Orchidaceae subfamily Apostasioideae; the 1st branch of the orchid family) occurred around that time.

The reference is:

Kåre Bremer
Early Cretaceous lineages of monocot flowering plants
PNAS 2000 97: 4707-4711; published online before print as 10.1073/pnas.080421597

On the same tree, monocot origins were estimated to have been early Cretaceous, at least 130 million years ago. Fossil angiosperms are known from around the same time. An undisputed monocot fossil dates back about 90 million years:

Gandolfo MA, Nixon KC, Crepet WL
Triuridaceae fossil flowers from the Upper Cretaceous of New Jersey 
AMERICAN JOURNAL OF BOTANY 89 (12): 1940-1957 DEC 2002 

There are no unequivocal orchid fossils that I know of. However, there are fossil euglossine bees in amber, and these are dated to 20-40 million years ago. The euglossines are the resin bees, the ones that pollintae Catasetiinae, etc. -->

Poinar G
Paleoeuglossa melissiflora gen. n., sp. n. (Euglossinae : Apidae), fossil orchid bees in Dominican amber 
JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 71 (1): 29-34 JAN 1998

No fossils of pouches or slipper orchids otherwise, but since the slippers are close to the bottom of the orchid family tree, one could expect them to be almost as old. Maybe about the same age as the orchids as a whole. My own attempts at molecularly dating the origins of the slippers were published here:

Albert, V. A. 1994. Cladistic relationships of the slipper orchids (Cypripedioideae: Orchidaceae) from congruent morphological and molecular data sets. Lindleyana 9: 115-132.

I calculated that the molecular divergence between Apostasia and Selenipedium was in the range of 49.3-69.0 million years ago. This is basically consistent with Bremer's estimate, above.

Furthermore, I calculated the time for the Selen/Cyp divergence to be ca. 15.3-21.4 million years ago. Then from Cyp and any given conduplicate-leaved genus (Paph, Phrag, Mex) --> 18.6-33.8 million years ago. Moreover, between:

Paph and Phrag --> 17.6-24.6 million years
Paph and Mex --> 16.4-23.0 million years
Paph delenatii and Paph glaucophyllum (or sukhakulii) --> 5.4-7.6 million years
Phrag schlimii (or lindenii) and Mex --> 7.7-10.8 million years
Phrag schlimii (or lindenii) and Mex --> Phrag lindleyanum --> 5.4-7.6 million years

So, if we believe these dates for the sake of argument, the slippers may have originated 50 million years ago or more, with all of the genera diversifying from one another by 18-33 million years ago. Furthermore, one can argue that Paphs and Phrags diversified within themselves around the same time in Earth history, some 10 million years or more after the conduplicate-leaved common ancestor evolved.

If you agree that one can infer from the slipper family tree that inflated pouches were the original (primitive) state for the Cypripedioideae (see my earlier posts), then that would mean inflated pouches could be somewhere around 50 million years old.

All best,

Vic


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## kentuckiense (Dec 11, 2006)

I'd like to hear thoughts on my observations about P. bellatulum and P. concolor(and the other Brachys but to a lesser extent).

When I look at the above two species, I see a clutch of bird eggs. The spots, the petal/sepal shapes, and the prostrate/pendant/short inflorescences just really bring that image to mind. In addition, don't the Brachypetalums tend to grow on limestone cliffs or something of that nature? It would seem like such a place would be a popular place for birds to nest.

Anyway, the point is that bird nest are places that I would think a fly would love to visit. There would be fecal matter, possibly broken eggs, and even dead chicks. By mimicing a clutch of eggs, a bloom could attract the same flies that would be attracted the those eggs.

I know it's all simple speculation, but does anyone find this to be plausible at all? Have bellatulum/concolor pollination mechanisms been studied? Does anyone know what the eggs of the local birds look like?


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## Braem (Dec 12, 2006)

Dear Victor,

lovely posting. But that is where I start having problems. Where is the tangible proof for all that. WHAT are you people looking at. WHICH part of the genome. WHY that part of the genome. etc. etc.

All I hear is: We have determined that ... we have calculated that ... etc. Were is your tangible proof that you really are hitting the right things?

Now to the bees. OK, I grant that we have bees that can be dated. But that means that we have bees that can be dated. Nothing more, nothing less. Its the old story about seing a person carrying a pencil. Does it mean that the person can write? NO! It only means that the person has a pensil and is capable of carrying it.
Because those bees NOWADAYS polinate certain orchids does not mean that they have pollinated certain orchids before.

We are running into a serious problem in this discussion. We should learn to walk first and then consider running. 

Victor, in my opinion, there is no point in discussing results until we have discussed the materials amd methods. First we must do the "WHAT", the "HOW" and the "WHY" ... then we can talk about the results and more important about the INTERPRETATION of the results.

regards
Guido




VAAlbert said:


> Hi Rick et al.
> 
> Indeed, molecular 'dating' of the origins of plant groups has been attempted, and an estimate for the origin of Orchidaceae is ca. 50 million years ago. A better way of putting this is that the molecular phylogeny suggests that the divergence between the lily relative Astelia and Apostasia (Orchidaceae subfamily Apostasioideae; the 1st branch of the orchid family) occurred around that time.
> 
> ...


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## Braem (Dec 12, 2006)

*Evolution of Orchids*

Folks,

this is from a recent article in the _Orchid Review_ (Kotsybar, 2006).

"*Orchid origins and adaptations:* Some botanists say the degree to which orchid genera readily hybridise with one another indicates orchids are very recently evolved. They declare orchids must be very young in evolutionary terms as evidenced by the genetic compatibility of so many genera. Others insist the ubiquitous presence of orchids worldwide is best explained if orchid ancestors cam from Pangea - the ancient , single continent from which today's landmasses split, 225 million years ago. Mysteriously, the fossil record can validate neither theory. There simply are no confirmed fossilised orchids - not even seed or pollen microfossils. The prevailing theory that orchids split off from the modern lilies, which some resemble, has been staggered by recent DNA analyses, which designate asparagus as orchids' closest cousin."

Now that paragraph can easily be rephrased as follows:

"*Orchid origins and adaptations:* We don't have a clue. "

Now, this *may* change in the future, and I am happy to accept that molecular genetics *may* be the method of choice, but I would like to see a few facts. See my other post.

Guido


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## VAAlbert (Dec 12, 2006)

Guido, Guido, Guido!

Your comments are as hard to address, in many ways, as the question:

"What proof have we that God did not create the Universe, or the Earth, or orchids, and that He did all of this exactly 4,444 years ago?"

In terms of data and methods, of course I can provide these, but they will never staisfy the eternal skeptic who asks for the ultimate proof. Like nearly every evolutionary biologist, I work in the realm of hypotheses and their evaluation. Not in generating proof.

To say we "don't have a clue" is to maintain that nothing short of The Truth, in the non-agnostic sense, is acceptable. Of course we have clues.

Re:



> "Orchid origins and adaptations: Some botanists say the degree to which orchid genera readily hybridise with one another indicates orchids are very recently evolved. They declare orchids must be very young in evolutionary terms as evidenced by the genetic compatibility of so many genera. Others insist the ubiquitous presence of orchids worldwide is best explained if orchid ancestors cam from Pangea - the ancient , single continent from which today's landmasses split, 225 million years ago. Mysteriously, the fossil record can validate neither theory. There simply are no confirmed fossilised orchids - not even seed or pollen microfossils. The prevailing theory that orchids split off from the modern lilies, which some resemble, has been staggered by recent DNA analyses, which designate asparagus as orchids' closest cousin."



What scientists like to *think* (flail their arms) about hybridization, Pangaea, has nothing to do with *objectively obtained data and analyses* that do indeed suggest that Orchidaceae belongs in the order Asparagales, not next to Asparagus, but too far away from it in phylogenetic terms.

Which parts of genomes used, and how the data are analyzed are of course legitimate, potential shortcomings that can be discussed -- but we are far from not having a clue, far from looking for proof, far from satisfying the eternal skeptic, and I will not present such material if that will be the constant reply.

Best regards,

Vic


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## VAAlbert (Dec 12, 2006)

VAAlbert said:


> but too far away from it in phylogenetic terms.



I meant:

"but NOT too far away from it...."

Sorry for the error.

V


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## VAAlbert (Dec 12, 2006)

kentuckiense said:


> I'd like to hear thoughts on my observations about P. bellatulum and P. concolor(and the other Brachys but to a lesser extent).
> 
> When I look at the above two species, I see a clutch of bird eggs.



I think this is fascinating! I think I recall that Paph concolor has been described by some as having a 'rotting' or 'rancid butter' smell, but I have never sniffed one with this in mind. If my memory served (and I can't remember the source), this could provide circumstantial evidence for your idea. Now, go into the field and look at brachys and birds' nests! Bird nest /egg mimicry would probably be one of the ultimate in terms of wacky pollination syndromes in all flowering plants!

All best,

V


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## Braem (Dec 12, 2006)

Victor,

I knew you would like my post :evil: 

By the way, it was the 15th of October if I am not mistaken, and Usher and the vice-chairman of Cambridge University quarelled about whether it was at 7in the morning or at noon. (If anyone needs the exact date, I will look it up - there is this most magnificent book "Before Darwin").

But lets get serious. I am afraid you are doing what all religious people do: simply saying "Well how do you proof me wrong?" 
That won't do in science. (And by the way, I could bring that proof as far as the creation thing, but then I would get all the religious people on this forum on my back.)

What I am simple asking is: tell me what you are doing. Tell me how you are doing it, and tell me how you come to your interpretations of your results. If you want me (and others) to believe that "molecular systematics" makes sense. Prove it.

If I tell you: "the leaf is 7cm wide by 3 cm long" it is something everyone can measure and say, well, Guido did not have his best day (if my measurements were wrong). Now do the same thing with your DNA studies.

More later, the boss is calling

Guido






VAAlbert said:


> Guido, Guido, Guido!
> 
> Your comments are as hard to address, in many ways, as the question:
> 
> ...


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## kentuckiense (Dec 12, 2006)

VAAlbert said:


> Now, go into the field and look at brachys and birds' nests!



Haha well, umm, I'll try to give it a shot someday!


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## gonewild (Dec 12, 2006)

kentuckiense said:


> I'd like to hear thoughts on my observations about P. bellatulum and P. concolor(and the other Brachys but to a lesser extent).
> 
> When I look at the above two species, I see a clutch of bird eggs. The spots, the petal/sepal shapes, and the prostrate/pendant/short inflorescences just really bring that image to mind. In addition, don't the Brachypetalums tend to grow on limestone cliffs or something of that nature? It would seem like such a place would be a popular place for birds to nest.
> 
> ...



I think your idea is very good. It makes a lot of sense. A bird nest mimic could and would attract many different types of pollinators other than flies.


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## kentuckiense (Dec 12, 2006)

gonewild said:


> I think your idea is very good. It makes a lot of sense. A bird nest mimic could and would attract many different types of pollinators other than flies.



Glad to hear I'm not totally crazy. Another forumite and I are trying to gather some info.


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## gonewild (Dec 12, 2006)

kentuckiense said:


> Glad to hear I'm not totally crazy. Another forumite and I are trying to gather some info.



Yeah! and totally based on reality :rollhappy:

Your bird nest concept struck a familiar thought I had when we first went to Peru. I looked up into a spray of large Catasetum flowers and what I saw looked just like a baby bird with it's mouth open waiting to be fed. It occurred to me that the flower was mimicking a baby bird waiting for it's momma to stuff an insect in. The flowers had perfect baby bird beaks complete with the fleshy side parts. If the flower was to be pollinated by an insect it would need to be a totally stupid bug to go in there! I'm going to go see if I can find a photo to post.


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## Rick (Dec 12, 2006)

kentuckiense said:


> Glad to hear I'm not totally crazy. Another forumite and I are trying to gather some info.



Right now I'm betting that if brachys aren't pollinated by hover flys (which aren't carion feeders) they are pollinated by euglosine bees (like parvies).

Cyp guttatum (pollinated by bees) does not have a strong resemblance to other (non orchid) flowers, but has a nice contrasting pattern that is attractive to bees. I think the brachy pattern would also qualify as bee happy. It also has a general structure that is more paph like than cyp like. 

Within genera there is a strong conservatism for the type of pollinators used by the species within a genus. Within Paphs we are already seeing 2 different groups of pollinators, and adding carrion flys would add a 3rd group (which I think would be 2 more groups than other orchid genera utilize).


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## gonewild (Dec 12, 2006)

OK I found some bird mimic photos. I don't want to change the subject from the Paph bird nest theory but rather offer it some support from another genera on a different Continent. 

Forgive the poor photo, this image was taken before mega pixels were invented. The Catasetums grow in trees and their spikes hang downward. Each flower is about the size of a live baby bird. Many birds in the habitat enter the nest from below to feed the babies so the downward facing flower could be just like a real bird. The person the flower looks exactly like a baby bird begging to be fed.

This is a male flower where the pollen would be picked up.







Here is the female flower where the pollen would be deposited. To me the female flowers look like big spiders which a bird may try to grab to feed to it's young. The act of grabbing the female flower may deposit pollen a bird picked up while trying to feed the male flower.


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## kentuckiense (Dec 12, 2006)

Rick said:


> Within genera there is a strong conservatism for the type of pollinators used by the species within a genus. Within Paphs we are already seeing 2 different groups of pollinators, and adding carrion flys would add a 3rd group (which I think would be 2 more groups than other orchid genera utilize).



Any word on what pollinates section Trigonopedia of Cypripedium? To me, that seems to scream carrion.


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## Rick (Dec 12, 2006)

gonewild said:


> I think your idea is very good. It makes a lot of sense. A bird nest mimic could and would attract many different types of pollinators other than flies.



If you loose your specificity then the odds of pollinaton are very low.

The Cyp guttatum paper by Banziger is very good. Of the 10 bee species found at the site only 3-5 species showed enough interest to enter the flowers, and about 80% of succesful pollen aquisition was by only 1 species. So that's fairly specific, and should ensure that the aquired pollen will end up back in another guttatum.

I should add that all the identified bees visiting the flowers were female too.


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## Rick (Dec 12, 2006)

kentuckiense said:


> Any word on what pollinates section Trigonopedia of Cypripedium? To me, that seems to scream carrion.



Naw!! To me they scream ground bees.oke: oke: 

Actually a couple of them do look alot like brachys as far as spotting goes.


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## kentuckiense (Dec 13, 2006)

Ok then. Are there bees that would be attracted to carrion?


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## Rick (Dec 13, 2006)

kentuckiense said:


> Ok then. Are there bees that would be attracted to carrion?



Holger Perner recently put a couple of articles in Orchids on Chinese Cyps. I'll email him and see if he knows what pollinates these guys.


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## kentuckiense (Dec 13, 2006)

Rick said:


> Holger Perner recently put a couple of articles in Orchids on Chinese Cyps. I'll email him and see if he knows what pollinates these guys.


Now he'd be a good guy to talk to. Good call.


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## VAAlbert (Dec 13, 2006)

Braem said:


> If you want me (and others) to believe that "molecular systematics" makes sense. Prove it.



Really, I could care less. I've been here for fun; this isn't.

Vic


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## Rick (Dec 13, 2006)

kentuckiense said:


> Now he'd be a good guy to talk to. Good call.



His first response was he didn't know, but suspected flys (didn't specify hover or carrion types). I asked back to specify fly guess and whether they had foul odors.


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## Braem (Dec 13, 2006)

Dear Victor,

fine. That is a statement we can all work with. And as far as I know now through your statement, the criticism I have had all along, and phrased as early as 1994 in Fukuoka stands. 

In any scientific field, you have to prove what you are doing. The question now is, why don't you want to? 

I thought this who tread is to clarify questions on taxonomy. Not to amuse anyone. But maybe I was mistaken. 

regards
Guido




VAAlbert said:


> Really, I could care less. I've been here for fun; this isn't.
> 
> Vic


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## Rick (Dec 14, 2006)

kentuckiense said:


> Any word on what pollinates section Trigonopedia of Cypripedium? To me, that seems to scream carrion.



Well Holger wrote back that he has seen "dung flys" using C sichuanense, and a "small black fly going into C. bardolphianum". There is another species in another section that is also an "obligate self polinator". He's got a very cool website on his Cyp preserve in China.

So this group of Cyps may have gone to the dark side for using generic garbage flys.


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## Rick (Dec 15, 2006)

Here's Holger's site I mentioned.

www.cypripedium.de/forum/messages/924.html

This is getting off of the taxonomy/evolution theme, but I think its important to understand pollination relationships with respect to evolution.


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## VAAlbert (Dec 16, 2006)

Rick said:


> Here's Holger's site I mentioned.
> 
> www.cypripedium.de/forum/messages/924.html
> 
> This is getting off of the taxonomy/evolution theme, but I think its important to understand pollination relationships with respect to evolution.




It's great, and very much evolution! Plus, far better than diatribe.

Best regards,

Vic


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## SlipperFan (Dec 16, 2006)

Good link. Thanks.


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