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Post by Dave Evans on Aug 10, 2007 22:05:33 GMT
What species should go into this group? I have listed:
N. alata N. boschiana (= N. faizaliana?) ?? N. burbidgeae N. chaniana N. clipeata ?? N. copelandii N. eymae N. fallax/stenophylla N. fusca N. glandulifera ?? N. hurrelliana N. maxima N. pilosa N. playchila N. truncata ?? N. veitchii ?? N. vogelii ?? N. zakriana
?? = not sure if these are actually closely related to N. maxima.
Is this list complete? Should any be removed?
The main characteristics I am using in assigning species into the maxima group are: petiolate leaves; hairyness; more or less flat and or expanded peristomes; peristomes which form a column under the lid; having lids with basal appendages (minus the plants from Sumatra based on many other characteristics which clearly show they are not close to N. maxima), and sometimes apical appendages; also most members have infundibular form upper pitchers.
I am having a difficult time find good example of flowers, both males and females, so this feature is not taken into to account.
The list of plants at the start of this thread, those which do not have "??" next to them all share these traits and most are highlanders too, which helped to keep their original populations separated and allowed them to evolve independently.
edit 1/14/09 added N. copelandii as a separate species. Still not convinced N. lowii and N. ephippiata belong in the N. maxima group--maybe in their own sister grouping apart of all other species--they are just too different and bizarre.
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Clint
Full Member
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Post by Clint on Aug 11, 2007 0:21:06 GMT
I always thought alata should have it's own complex.
I think that N. clipeata, N. truncata, and maybe even N. chaniana are distinct enough (there's the big arguement!) that they should not be included. If some of the others like N. platychila and N. vogelii are included, then certainly N. spectabilis should be included.
I also push for more use of subspecies status. I think the whole genus needs a good cleaning and reorganizing.
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Post by Dave Evans on Aug 11, 2007 1:02:05 GMT
Hello Cllint, Well, please go into to detail about it... I only have a one or two clones of each and that is a really small sample so making sense of it will be a group effort.
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Post by phissionkorps on Aug 11, 2007 1:16:37 GMT
I wouldn't include burbidgeae. It seems that would fit more into a group with rajah, mira, etc, but thats just my opinion. I feel alata is distinct enough to have its own group, which is certainly needs. I wouldn't put the alata group as a subgroup of maxima, unless you are talking about making some sort of tree with it splitting off quite a long time ago. I don't know how I feel about chaniana and pilosa...I probably wouldn't include them unless again, they are grouped together and split off earlier than things like eymae. I'd also take out truncata, and probably veitchii too...I'm not really drawing any similarities between it (veitchii) and maxima at all. Maybe you can clarify? So we're pretending zakriana is a species now?
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Post by Dave Evans on Aug 11, 2007 3:22:58 GMT
Hi Ron,
Well, N. pilosa and N. burbidgeae appear to be each other's closest relatives, but since it is from the remote areas of Kalimantan, it has not been recollected and there is no living material of it in cultivation... There is not much info about it other than the type specimen.
I can't see how either of these two could be put into a group with the other plants, like N. rajah and N. villosa... What are the affinities, other than where they grow being physically closer?
Hmm... What about N. paniculata, that one looks rather like N. eymae... No?
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Post by phissionkorps on Aug 11, 2007 3:32:54 GMT
Burbidgeae and rajah occupy very similar and sometimes the same habitat, which I think is important to consider. Also is the fact that they readily hybridize with each other. I have said it before, and I will say it again: I think too often we classify Nepenthes solely on morphology, and while providing a good background, I don't think its the best way to go. Especially since the genus is polymorphic, especially within the maxima group. But on the other hand, burbidgeae's lower pitchers are pretty mira-esque IMO. You do agree that mira and rajah are very close, right?
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Post by glider14 on Aug 11, 2007 6:15:30 GMT
hmmmm....what about...N. boschiana, talangensis(meh??), or even macfarlanei? Alex
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Post by Dave Evans on Aug 11, 2007 7:58:04 GMT
Hello Ron and Glider,
I have included some of these plants into the maxima group as per research done by other authors. My idea is to dispute what has been proposed and see if it actually makes sense to people. I too have a difficult time putting N. truncata into the M-G, but people who have done a lot more research than I have have done just that...
I'll put in N. boschiana and N. faizaliana (each other's closest relatives and possibly the same species), but with reservations, marked by "??". I cannot tell if these are superficially similar in appearance to N. maxima, or if they are actually close relatives of N. maxima.
I am not sure if there are any members of the maxima group east of Borneo... There are species in the singalana-spathulata group from Sumatra that have basal appendages on the lids, but these don't seem to have anything else in common with N. maxima. N. talangensis and N. aristolochioides appear very close to each other, and neither shows much in common with N. maxima--the leaves are sessile (M-G are petiolate), completely different hairs and general structure of the plants indicate affinities with species like N. inermis and N. jacquelineae. N. macfarlanei has me stumped as to where it might go on the Nepenthes family tree, but it doesn't really look anything like N. maxima.
I am completely unfamiliar with N. mira. While N. burbidgeae and N. rajah do occasionally hybridize, I don't see how that makes them closer relatives...? If they were really close, wouldn't hybridization keep them from being so distinct? I suspect N. rajah is actually closer to N. lowii and N. ephippiata...
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Post by phissionkorps on Aug 11, 2007 8:30:19 GMT
Not in this genus. There are no pre-zygotic and apparently no post-zygotic mechanisms to prevent hybridization between any of the species (in cultivation). In situ, temporal and spatial isolation play a role, but obviously not between burbidgeae and rajah as evidenced by alisaputrana. Rajah and mira IIRC are supposed to be each other's closest relatives. I find rajah completely different in all respects from lowii and ephipiiata, except that they are all highland plants.
Could you give some clarification as well on the relation of vetichii to the maxima group? I think some of the others listed may be a bit of a stretch, but it's possible to include them after using one's imagination a bit. Veitchii though is much different in almost all respects, and putting it in with maxima makes about as much sense to me as putting in something like burkei.
I've not heard this. Could you please clarify a bit? I thought faizaliana and fusca were closest as well?
I just looked at your amended 1st post. Isn't copelandii closest to alata and not fallax/stenophylla? IIRC, wasn't it you that told me they were very close and there has no doubt been some hybridization between the two at least along alata's southern range?
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Post by rsivertsen on Aug 11, 2007 13:32:43 GMT
According to Danser, N. rajah, N. villosa and N. burbidgeae are in their own Urceolate group, with a very limited distribution in northern Borneo, and all three are found growing exclusively on serpentine rock outcrops.
N. alata is its own group, with distribution throughout the Philippines and Palawan, and are lowland species for the most part. While the species in Sumatra evolved, in my opinion, from the N. gymnamphora complex and are highlanders.
The Malayan highland species also seems to have evolved on their own, outside the Maxima group; just too many significant differences in their leaf/petiole structure.
N. lowii is actually very close to N. maxima in their leaf and petiole shape and connection to the stem; the floral parts needs to be compared too, as these things have far more botanical significance, but that large vaulted lid certainly is far from anything that N. maxima displays. Danser also included N. stenophylla and N. pilosa in the Maxima group, and again, the leaf/petiole shape and connection are very close.
N. truncata is also very close, and I'm not familiar with N. boschiana enough to comment on it. N. clipeata is another one that's close, but again, the vaulted lid, and the peristome seem too different to me, but floral parts may show otherwise. N. faizaliana is certainly part of the Maxima group as is N. hurrelliana and N. chaniana, while N. eymae is just a form of N. maxima proper in Sulawesi.
The distribution of the Maxima group, according to Danser extends from northern Borneo, to Sulawesi, and parts of West Irian Jaya, and Sulawesi seems to be the epicenter for this group, and almost all species are highlanders.
I’m not familiar enough to comment on the other species listed previously here.
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Clint
Full Member
Posts: 808
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Post by Clint on Aug 11, 2007 14:42:54 GMT
Is the N. hurrelliana debate over? Hybrid or species or species evolved from a hybrid?
Dave, my reasoning for N. spectabilis to be included is it's similarity to maxima/fusca. While it's closely related to N. lavicola, I still say if we are going to start complexes it should be in the Maxima complex. If there's a gymnamphora complex, N. lavicola should go there with xiphioides and pectinata.
As far as N. copelandii, I believe it's SO similar to N. alata that it doesn't even deserve species status. A Rajah complex/group sounds reasonable. Could include N. rajah, N. Sp. palawan. N. Sp. DA, N. villosa, N. burbidgeae, and maybe N. mira.
I also think that if a complex/group system were to be in place, care should be taken to try and make everything "fit".
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Post by Dave Evans on Aug 11, 2007 20:18:07 GMT
Hello Ron,
You're talking about other people's confusions, about how to interpret specimens, not the actual species. For example: Authors which didn't review some species (or thought they were, but later we find out they were looking at something else) then went ahead and combined them into one name, but these combinations are artificial and have been subsequently disputed. N. fusca is a highland, mainly an epiphytic species, while N. faizaliana is lowland-intermediate and is strictly terrestrial on limestone, like like it's twin N. boschiana and the completely unrelated N. campanulata. Just because plants that later proved to be close to N. fusca used to be called by the name N. faizaliana doesn't make the actual species closer, it only means people were confused about the identification.
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Post by Dave Evans on Aug 11, 2007 20:35:13 GMT
Re: Hybrids, just because some species hybridize, for the hybrids to contribute back to one or both of the parent species, they have to be viable long term. I think hybrids can act as bridges between species, but not always. Sometimes the hybrids die out, sometimes they can out number and take over areas and then become species in their own right. If N. rajah and N. burbidgeae have always been physically close to each other and have always interbred, then they wouldn't look very different. To me, they are clearly from two separate lineages that now happen to be in the same place, Mount Kinabula. I'm not saying subsequent hybridization cannot have made them closer, but is there anything to indicate that N. alisaputrana is contributing genes back to either parent?? I don't see it in the plants, but this does look like it could be the case between N. rajah and N. villosa.
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Post by Dave Evans on Aug 11, 2007 21:04:55 GMT
Hi Ron and Clint,
As per your advice, I placed ?? next to N. veitchii. What do you suppose its closest relative is, if not other members of the N. maxima group, like N. fusca?
The hispid hairs which cover the plant, the lack a basal crest an no apical appendage on the lid and the way it grows are the only reliable methods to distinguish it from N. maxima...
Have you looked at a large plant of N. copelandii? It is very similar to N. fallax, much moreso than it looks like N. alata. The confusion I was referring to regarding plants from Mindanao, is that both N. copelandii and another species identified as N. alata grow there together. The N. alata plants are widespread, while N. copelandii is a highland plant restricted to three mountains. Hybrids that occur between these two species on these mountains have blurred these plants together as one, but only in some people's minds (For reference to how weak some interpretations are: Some folks think nepenthes pitchers look like bananas) The easiest way to tell them apart: N. c. is a fuzzy plant with "v" shaped pitchers. "N. alata" on Mindanao is glabrous and never make "v" shaped pitchers, they always have a swollen base.
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Post by phissionkorps on Aug 11, 2007 22:07:26 GMT
I've never been to Kinabalu, so I couldn't tell you, but one woudl have to assume its totally possible. The "problem" I think, is that there are not many alisaputrana plants, so any burbidgeae x alisaputrana or rajah x alisaputrana hybrid will be washed out relatively quickly. This I would guess is due to temporal isolation, as it is certainly not geographic isolation. Then again it could be a pollinator "problem". I wrote a paper about the pollination syndrome of sexually deceptive orchids once. While each species mostly had one euglossine species act as a pollinator, there was a wide range of euglossine species that would pollinate some orchid species on occasion. Some orchid species also shared pollinators. If species A pollinates only rajah and there are 100 of them in a given system, and species B pollinates only burbidgeae and there are also 100 of them, but species C can pollinate either/or, but there are only 5 of them, we certainly will not see hybrids as often. I don't know if that's the case or if its temporal isolation, as I don't know when each species flowers. Obviously there is a reason that there aren't so many alisaputrana, but there are large, variable, stable populations of things like kuchingensis. I think though, that since rajah and burbidgeae do indeeed hybridize, one cannot completely discount the possibility of genetic drift.
No, I have not have a chance to look at a large copelandii. Even if I had one, all I would have to compare it to are my 2 forms of alata as I don't grow fallax, and one of my forms is boschiana mimic, so I don't think that would do me much good.
ATM I have no idea what to put with veitchii. I'll have to think it over for a bit...
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