Orchid News # 34

XIX WOC

Another look at Hoffmannseggella H.G.Jones
                                       by Francisco E. Miranda

Why not accepted when first proposed by Jones in 1968:

Hoffmannseggella was established by Henry Gordon Jones (1968), and for reasons mentioned below it was never accepted by taxonomists and growers alike. It is easy to understand growers, as they don’t like and tend not to accept lightly any sort of name changes that will affect well-known cultivated plants. As for the taxonomists, the situation was different but basically boils down to:
First, although the group has always been considered as very distinct from the time Lindley (1842) put the species into Parviflorae and throughout the history of Laelia, Jones did a very poor job describing it as he used features that not all of the species have and also can be readily found in species of other groups. These essentially were:

- Lip much narrower than in Laelia (true to some point).
- Petals similar to sepals (yes, like in several other groups in the Laeliinae).
- Rupicolous or rarely dendricolous habit (too vague, same for many genera in the Laeliinae).
- Laelia (Microlaelia) lundii is not in the group and has all three features, and just this example was enough for his work not to be taken    seriously.

Secondly, there is a natural resistance to separate one group and not do anything about the others (in the case here, the other “Brazilian laelias”).
And finally, the splitting craze that we see today was not yet in the works by then.

Laelia (microlaelia) lundii is a good example of an unrelated species that has all the features mentioned by Jones for Hoffmannseggella. It easily shows why his work was not accepted at the time

Reasons to accept it:

So, considering that Jones’s features were not good enough, let’s try to add other ones and better define the genus:
1- Star-shaped flowers with essentially similar sepals and petals. This rules out most of the large-flowered species from other groups.
2- Flowers have a deeply trilobed lip with well-defined round to elliptic front-lobe. That rules out the hadrolaelias and the purpurata types.
3- Flowers have a lip that is moderately to strongly curved backwards, and this finally rules out Laelia (Microlaelia) lundii and all other “Brazilian laelias”.
4- Inflorescences originating from well-developes sheaths (that can be difficult to see on very small species, but it is there). This alone rules out all of the hadrolaelias.
5- Plants always with very short rhizomes, and this rules out all the other “Brazilian laelias”.

- There are several other features that are unique to the group, but for delimitation purposes these are more than enough. It needs to be mentioned though, that all the species have uniquely large seeds, and this plays an important role in local distribution.

Flower Diagrams:

Flower diagrams of three species of Hoffmannseggella to show how uniform the group is.

Hoffmannseggella species groups

This is a good exercise on judging relevant and irrelevant features. We always have to choose what to use for taxonomy, and in many cases a lot of information about the species is needed to give us a fairly good picture of any group. In the case of Hoffmannseggella, that is what we can bring to the table:

A- One way to organize the species is to group them by color, and this will give us three basic groups:

1- Species with yellow flowers.
2- Species with pink to purple flowers.
3- Species with orange to red flowers.

Advantages:  
- Easy to group because it is very easy to see.

Disadvantages:  
- Doesn’t really group species by affinity (with exceptions), and thus doesn’t help us understand what happens in nature. This is a typical artificial grouping, and can be significantly improved.


B- Another way to organize the species is by flower morphology, which in most cases agree with the plant morphology but not always with flower color. If we want to use more info, we can add inflorescence habit and, why not, plant habit. All this can be crossed with geographic distribution and population variability data, so we can end up with some pretty solid groups.

Advantages:  
- Allow us to group species by theoretical common ancestor,  considering of course that these group ancestors came from one (if we consider this a monophyletic group, something that can be done quite comfortably).

Disadvantages:  
- There is a need for a lot of data to give us a really good picture of   these groups.
- Sometimes it is not easy to place species and affinities, regardless of how much info we have. This means that the more info we gather, the more difficult it can get to organize it.


C- Some features are of little use or we have to be careful with, while others are just plain misconceptions. So, we have to be aware when using them in species keys. Among these are:

1- Size of plants:
This is mostly useless in the group for particular species, as the same species can have a   big variation in size depending on habitat or growing condition. In the habitat, light and wind exposure plays a big role on plant size. The same is true for plants in collections, where the impossibility to give proper exposure as in the habitat produces very different plant. For example, plants of Hoffmannseggella bradei frequently get taller in cultivation as it is difficult to give the high amount of light plants get in the habitat while, on the other hand, plants of Hffgla. cinnabarina tend to get shorter as they don’t have to elongate the pseudobulbs to get light as they won’t be growing inside shrubs.

2- Inflorescence hight:

This have to be used very carefully. Although it is quite obvious that there are species with very short or very long inflorescences, it is also true that some species tend to produce shorter or longer inflorescences in cultivation (e.g. Hoffmannseggella itambana). This is especially problematic as a common trait recently is to do taxonomic work with cultivated plants (for whatever reason). Considering that species keys on the group frequently use inflorescence hight, one can imagine how wrong they can get.

.

Hoffmannseggella itambana, when in cultivation (left), tends to produce much taller inflorescencesa than in the habitat (right).

3- Number of flowers:
We have to be careful with this one. When Pabst described Laelia liliputiana, he wrote that the main feature was that the species ALWAYS produce ONE flower on the inflorescence. Today we know not to make statements like these.

Hoffmannseggella liliputiana, here with three flowers on an inflorescence. This is quite rare but, in the habitat, two flowers is quite common on an inflorescence. If we take literally what the protologue says, we would have another species here.

D- There are other misconceptions that, although not frequently or ever used in keys, give us an incorrect idea of the species of the group. Among these we have:

1- Plants always come from high elevation mountains:                       
At least two species can be found on the coast, and one at around 100 m. elevation (e.g. Hoffmannseggella gloedeniana).

2- Plants are cool growers:
Well, besides the ones on item 1, even the ones growing in the mountains can be subject to very high temperatures during the day, and not always gets cold during the night (e.g. Hoffmannseggella briegeri).


E- Now there are features that are quite stable, besides the flowers of course, and these are very useful for identification. Among these we have:

1- Plant habit:
This is a very solid feature, which means that at least some species can be separted just   by that. This is actually one of the most important features to separate Hoffmannsegella crispata from Hffgla. flavasulina, for example. That means that they can be told apart just by looking at the plants.

2- Inflorescence habit:
Another quite stable feature, if one knows what to look for. One of the most important things here is flower distribution. Species that have the flowers bunched at the top will always be that way.
A good example is in the Hoffmannseggella crispata group.
Hoffmannseggella crispata can be easily separated from Hffgla. endsfeldzii because it has the flowers crowded at the top while Hffgla. endsfeldzii has them spread. Even if everything else was equal, that would be enough to separate them..

Here we have Hoffmannseggella endsfelzii on the left and Hfglla. crispata on the right. It is easy to see the differences in flower placement on the inflorescence.

Natural Hybrids and speciation

This is a group of plants in active speciation, and this can be seen by the quantity of natural hybrids. Whenever more than one species grow together, there is a good possibility that the natural hybrid between them will be found.
If the flowering season of the species involved is different, which is the norm, few hybrid plants are usually found and they usually end up being diluted back into one of the populations (introgression). Good examples are Hoffmannseggella xCristinae and Hffgla. xRaganii.
This also happens if the microhabitats are different, even when flowering season is the same (e.g. Hoffmannseggella xHispidula). It is easy to understand if we remember that seeds are very heavy and thus have a very localized dispersion.
However, when different species share the microhabitat and have similar flowering seasons, full hybrid populations can be found (e.g. Hoffmannseggella xBritoi).

Here we have Hoffmannseggella briegeri (top left) and Hfglla. rupestris (bottom left), that when hibridized produce Hfglla. xCristinae (top and bottom right, flowers from two different plants).

Hoffmannseggella species affinity list

This is tentative grouping of species in alliances within the genus. It is based in plant and flower morphology similarities. Species followed by (*) are still doubtful regarding placement. Species with names between (“”)s need new species names and are referred by their commonly but incorrect known name.

1- Yellow flowers (mostly)

Hoffmannseggella blumenscheinii alliance:

blumenscheinii
gracilis

Hoffmannseggella bradei alliance:

bradei
briegeri
esalqueana
itambana
kleberi
verboonenii

Hoffmannseggella cardimii alliance:

cardimii

Hoffmannseggella crispata alliance:

crispata (flava)
endsfeldzii
flavasulina
milleri
sanguiloba *

Hoffmannseggella mixta alliance:

alvaroana 
bahiensis *
mixta

Hoffmannseggella gloedeniana alliance:

gloedeniana
macrobulbosa

2- Orange to red flowers

Hoffmannseggella angereri alliance: 

angereri

Hoffmannseggella cinnabarina alliance:

cinnabarina
“cowanii”
colnagoi
hegeriana
mirandae

Hoffmannseggella harpophylla alliance:

brevicaulis
harpophylla
kautskyi

3- Pink to purple flowers

Hoffmannseggella caulescens alliance:

caulescens (crispilabia)
pabstii (mantiqueirae)

Hoffmannseggella duveenii alliance:

duveenii

Hoffmannseggella longipes alliance:

fournieri
ghillanyi
kettieana
liliputiana
longipes (lucasiana)
reginae

Hoffmannseggella munchowiana alliance:

munchowiana

Hoffmannseggella pfisteri alliance:

pfisteri

Hoffmannseggella rupestris alliance:

rupestris
tereticaulis

Comments on DNA studies

This is a section where I voice my personal opinion on the very newest trend in orchid science. As with something new, a lot of people jump in and get to the conclusion that this is the solution for all our classification problems. Some people go as far as to say that we should throw all the previous knowledge away and start anew. I lived enough to remember that exactly the same thing happened when cromossome studies were new and also phytochemistry was new. Eventually, all this information was added to our common knowledge, together with morphology, anatomy, phytogeography and several other study areas. The more info we have, the more precise can our conclusions be. DNA studies, being so new, of course will have changing results as a consequence of evolving methodology. Following are parts of Cassio van den Berg’s excellent essay “Considerações sobre as ex-Laelias brasileiras, Sophronitis e outros gêneros”, that was published online (van den Berg, 2003). I choose a few paragraphs that I thought were more relevant for the discussion and commented after each of them. Of course these can be taken out of context so I strongly recommend everyone to read the whole of it, it is excellent reading. So here we go.

“The first is that, although the ITS data show clearly that the Brazilian species of Laelia had to be segregated, the level of variation (the differences) in the DNA within the trees was low, and consequently, the subgroups formed within the tree had little statistic confidence. This means that in a next DNA study (see below for an example), the relationships among the different groups of Laelia could change reasonably. Therefore from a stability of names viewpoint (to avoid later changes) making a single genus was a safer option;”

- Agreed, and this is reason enough to consider any conclusions as premature although as more info is added the better the system works.

“The second important point, is that maintaining these species in a single group we have more information in the system. For example, in the classification system of orchids most largely used today (Dressler, 1993), there are no categories between subtribe and genus, and consequently, the genera are listed in alphabetical order within subtribes. If we had proposed several genera for these species, these genera would be lost among other 50 other genera of Laeliinae, however, the information that all these species are related would have been lost. Our intention is later to propose subgenera which show the different groups of species, which is a formal way of showing the subgroup information. However, because the studies are still in course and we could not achieve conclusive delimitation of the subgroups, these proposals were still not done, in agreement with the stability factor;”

- Agreed with the first part, but this shows how initial the conclusions were at this point.

“The third factor is of historical nature: traditionally it has been maintained in the literature that an average size of a group (family, genus, etc.) is around 40 items, and this was one of the criteria used in the flowering plants classification of APG (1998). The Brazilian species of Laelia were a little above this number, but on the other hand the division into smaller genera would produce many genera with much less species (around 10), and some monotypic genera. Furthermore, the genus Laelia was well accepted in the size it was, only with the questions regarding the two groups (the Brazilian and Mexican). The subgenera of Laelia were always a good solution to show the subgroups, but without losing the coherence of the larger group;”

- Agreed in essence, but keeping polyphylletic groups together doesn’t make much sense. And Hoffmannseggella for example has the perfect 40 species number, give or take a couple.

“And finally, attempts of splitting the subgroups into distinct genera had already been done (ex. Hoffmannseggella, Jones, 1968) and were not well accepted by both botanists (Garay, 1973) or orchid growers. In this way, when we chose to transfer all species of this group to Sophronitis, we chose the simpler, stabler system with the best information content.”

- Yes, Jones didn’t do a good job, but he didn’t know the group and its species very well (or at all). So considering everything, his work would face an uphill battle on taxonomic circles no matter what. As for the amateurs, taxonomists can do whatever they want, Laelia purpurata will always be Laelia purpurata. Until, of course, all the old school growers die and new ones never know about the old names.

“In the work of Chiron and Castro (2002), although a morphological justification is given, a careful reading makes it clear that all groups were delimited exclusively based on the trees of van den Berg et al., (2000), which were reproduced therein, and also the list of species does not differ from the list published in Sophronitis in our adjacent paper. No additional data was presented to justify the decisions. This somehow made us to worry, because we knew the fragility of ITS data to establish groups within Sophronitis (this fragility was stressed in the original study of DNA). This is even more surprising because in the beginning of their article, Chiron and Castro (2002) cited several preliminary cautions about the use of phylogenetic trees for establishing classifications (items A-E), and immediately ignored these criteria (especially item E about robustness of branches) in their proposal.”

- See, Cassio nailed the problem here. For his conclusions to be contested, the work has to be done again, that’s
why we have “Materials and Methods” sections on scientific works – to allow repeatibility. If you don’t redo the
work to get to your own conclusions, you risk basing your arguments in data YOU DON’T HAVE, and in the end
we have an esoteric discussion of sorts. What I tried to do here is circunsbcribe Hoffmannseggella purely on
morphological and ecological grounds, and I think such an uniform group is very easy to deal this way.

“Although studies with several other DNA regions for assessing relationships within Sophronitis are not finished, preliminary data with two plastid regions (Fig. 1) already show that in order to use the system of Chiron and Castro (2002), which was just proposed, several substantial modifications would be necessary”.

“Based on data from the regions trnL-F and matK, the species of Sophronitis in the old sense (before receiving the species of Laelia), constitute a good group (differently from the separation of S. cernua which appeared in the ITS trees), and therefore the placement of the group of S. coccinea in Hadrolaelia is inadequate. Similarly, S. harpophylla goes as sister to the rupicolous species, what makes a dubious situation. At the same time that these species placed in Dungsia have some differences in relation to the rupicolous species (especially size), it is obvious that they share many other morphological characters with this group, and in this sense it appears better than Dungsia would be lumped to Hoffmannseggella in their system. In this case the only real justification that could be made to keep Dungsia separated from Hoffmannseggella would be the will of the authors.”

- Well, that sums it up.

Conclusion 

With the above, I have to conclude that:

1- I cannot accept Dungsia, first because it was described based on the first ITS sequence work that is being contradicted by newer studies, but more importantly because a key morphology feature used to justify it is plain wrong. All the species of the group DO have floral spathe. And based on morphology, just look at the flowers and you know where those species belong.

2- I am glad this is such an easy group to delimit morphologically. Taxonomists have been studying morphology for more than 200 years for the simple reason that it is so easy to study what you can see. This was the only option in the beginning and now there is so much information that it just can’t be dismissed by the shake of the hand to start from square 1. We also have to remember that:
Phenotype = Genotype + Environment, which can be translated as “morphology is a consequence of the DNA plus environmental factors”.

3- DNA studies, at this stage, are very preliminary, and I can’t rely on them to get to conclusions on group separations. I won’t get into detailed discussions for several reasons, including the fact that I didn’t do the work and so have no info to bring to the table and especially because it doesn’t matter as I already made up my mind that they are still preliminary. I took a hard look at all of the DNA work being done and find it extremely interesting, so will get back to it in the future to see how things are going.
You see, the more I think of it I imagine that, even a long time from now, the molecular and morphological info will converge to a point that the results will be essentially the same. This is because both are just two different ways to look at the same thing.

4- Talking with Cassio during the timeframe of the WOC, we discussed his opinion that it will be better to consider the Brazilian Laeliinae as part of Cattleya and the different genera as subgenera of it. I think this is a sensible approach and addresses the main problems created with lumping them with Sophronitis and makes it basically a matter of splitting or lumping genera. In this particular case, lumping is an option but it get at odds with what is happening with other groups as for example the Oncidiinae and Pleurothallidinae where heavy splitting is happening. The rupicolous laelias are a very uniform group that will stay equally well as a separate genus or a subgenus of Cattleya so I will be watching what happens.

5- And finally, at the end of the day, we all have our personal opinions and these reflect on every work we do. Everything I am saying here is just that, my PERSONAL OPINION. The same way I don’t agree with something, you are of course free to disagree with me. Regardless of how much information I tried to put together, it doesn’t change that fact.

Hoffmannseggella harpophylla in the natural habitat, prope Venda Nova do Imigrante, ES. It is easy to see the well-developed inflorescence sheaths, typical of all species in the genus. This by itself renders invalid the morphological justification for Dungsia.

Bibliography:

Jones, H.G. 1968. Acta Botanica Academiae Scientiarum Hungaricae 14: 69.
Lindley, J. 1842. SECT. Parviflorae in Bot. Reg., XXVIII, sub. t. 62.
Pabst, G.F.J. & Dungs, F. 1975. Orchidaceae Brasiliensis, Band I, p. 146. Brucke-Verlag, DE.
Schlechter, R. 1917. Die Einteilung der Gattung Laelia und die Geographische Verbreitung ihrer Gruppen. Orchis 11:87-96.
van den Berg, C. and M.W. Chase. 2000. Nomenclatural notes on Laeliinae 1. Lindleyana 15(2): 115–119.
van den Berg, C. 2003. Considerações sobre as ex-Laelias brasileiras, Sophronitis e outros gêneros. Orchid News 20 in Brazilian Orchids online (http://www.delfinadearaujo.com)
van den Berg, C. and M.W. Chase. 2004. A chronological view of Laeliinae taxonomical history. Orchid Digest 68(4): 226-254.

An exercise in personal opinion

This is a simple random diagram to show how these things go. Depending on the person looking at it, the red line can slide left or right. Depending on where it is positioned, we go from one group to four, six, twelve or seventeen. All this without changing the data, just the person looking at it… And this is valid for morphological data, numeric taxonomy data, and even… DNA data. Just put a real key instead of this one and see people fighting over where to put the line. Human nature, but I digress.

 
Photos: Francisco Miranda

Francisco E. Miranda
4763 Polk City Road,
Haines City, FL 33844, USA
fmiranda@att.net