PART 1

I'd hope this is as good a place as any to mention the conflicting views that arose from the morphotype idea, as I know quite a lot of ferny people are interested in it in this group. I think basically our approaches do differ in a few highly significant ways - which when carried to their logical conclusion have resulted in a major dichotomy with conflicting taxonomies being presented.
These are:
1. In the various genera I've researched I've always had to go through the situation of having certain dubious entities - not sure if they are genuinely distinct or at what level, or not sure if already named or not, which are not ready to publish. As this situation almost always arises when studying a group I just do what most botanists have always done while things remained undecided for them (which I think is your situation as described) - I just give some nick-name such as what I'd been calling the new Irish taxon, like "Torc 1" etc. Then I wouldn't publish on this with a name until I did believe from further study that I was in a position to understand it and to know whether it was new or not and of significance or not, and then, eventually where to place it. This is in order not to clutter up the literature with provisional names any more than has to happen by error or design until I felt confident about them. But the morphotype system uses what look like formal Latin names for what protagonists think of as dubious entities and are being continually used as such, "D. affinis morph. convexa" etc. - so even though evidently not the intention, it has created a kind of alternative nomenclature despite it being said to be undecided. If this happened for many more complex groups (as it could, especially if internationally) we would, as now already, create an intractable mass of alternative names, confusing the issue, just as I believe this did, though unintentionally. I would prefer that if one were undecided one would simply not publish any kind of names like this, or if decided give proper formal names in the usual hierarchies and framework of Botanical nomenclature.

2. Although you say you are yourself undecided or doubtful about the taxa I raised, this was not so much the case with me, as after years of detailed and far-reaching study all over Europe and W. Asia, Macaronesia etc. I became familiar enough with them to feel that I did know what entities there were that were really significant and where they fitted. There was also a great deal more "Hard" or "semi-hard" evidence for them than is usually the case in taxonomic study (considering how it happens world-wide), not only did I have great detail and much material showing the morphological ranges of variation (commonly not the case in most taxonomy) and geographical ranges of the taxa, but also subsidiary proof, even though not "complete" (which, after all it isn't for 90% of named plants). I had not only all the 16-spore-mother-celled chromosome counts showing different pairing behaviour in the different entities, as listed in my unpublished monographic study, but also phytochemical study showing different chemistry for borreri and similar for the others. If you consider the British ferns, the great bulk are without such evidence and quite a few even without such detailed traditional taxonomic study - but lack of so-called "hard" evidence is no grounds for dislodging the various authors' published accounts. The reliability of the study depends on the ability of the various different authors - and from a historically retrospective viewpoint it became well known some were less thorough and successful than others. Their work has been replaced by new evidence, new studies, giving a new taxonomy decided upon by a later author - and that's how it goes on. We do not normally have the situation where an author says in effect, "I am not decided for myself, I feel we need more evidence, therefore I am going to replace the existing taxonomy, not by offering a new one - but by offering a list of any name at any rank, including some unproven new ones, and just call them all morphotypes". To me that is actually retrograde and offers no new findings, expertise or scheme to replace the old one, but creates a parallel, temporary system of dubious "nomenclature" and thus a nasty muddle! No one has actually presented any place where the taxonomy I am putting forward is known to be erroneous - so it's actually just a positive being replaced by a negative without any evidence as to why it should be replaced. I am well aware that this was NOT the intention, as the morphotypes are supposed to be non-judgemental - but the practical result has been to replace the published and researched taxonomic scheme with one that is not actually decided and accepted by the author, but is a list saying "we don't know". Time will tell, of course, but I believe I DO know the things I published! I simply do not see the value of replacing it with what someone else says they don't know and which is only supposed to be temporary until some hypothetical time in the future when that author can say he does know. When you say there isn't much difference between us it is only true in that the morphotypologists do indeed accept the taxa I named, but merely say they don't know where to place them. But that is an important difference!

3. Perhaps the most important difference is that I am quite convinced I can indeed recognise where these taxa fit. It is from that that the hierarchy was deduced. I deliberately distinguish between minor taxa and more major ones - at first I separated the major ones as subspecies (and I remember that at that time it was considered terribly over-splitting!); now after years more study I am placing them as species. It is also clear to me that WITHIN them there are certain geographical entities which are of considerable significance - thus these (which I originally made at the rather unsatisfactory rank of variety) are now subspecies. I can recognise easily from their morphology that subsp. kerryensis and subsp. paleaceolobata are good recognisable entities that obviously belong within D. affinis (in the new, restricted sense of the former subsp. affinis). But beyond that there is some much more minor and morphologically less significant variation that is apparently partially genetically fixed in some populations. To me it is natural and perfectly acceptable that there can be minor, part-recognisable variant forms within any fern-species
I know - unlike the view of a computer-cladistician who says each species must be a single entity only and ends up with only two ranks, the species (actually meaning precisely what morphotype has come to mean) and the genus. That is a place we really differ, as I worked out how they are related from their morphology (inc. spore-size and degree of abortion in a few cases where the basic cytotype has not been found) and produced a hierarchical scheme which reflects it. I do not find it very satisfactory to put this aside without showing any actual errors in it. If, for example, D. affinis subsp. kerryensis (cytologically unknown) turned out to be triploid, not diploid (as its spores and other morphological features strongly indicate), then it should be published and the necessary correction made - one should not deduce from such a question that it indicates the whole taxonomy should be put aside and replaced with a list of unranked names! The vast bulk of taxonomic novelties have not had the luxury of even a basic chromosome-count, especially in the past, but also still today, let alone any other "hard" evidence. It is not thereby invalid or to be replaced by a "don't know" list of unranked names.

4. As morphotypes are unranked this has seriously destroyed one of the major things I was trying to get over to others - the fact that within this apomictic schemozzle some variants are important and of major significance,some less so and obviously fitting within the others, and some of much less significance. Perhaps the most unsatisfactory result of replacing taxonomic names with morphotypes is that it simply places a local form or clone on the same level as a major taxon (and as they are apomictic even a single individual is reproductively isolated and acts as if a "biological species"). As morphotypes they are all at the same level of ignorance about their significance. I cannot accept that a minor local form occurring here and there apparently through cloning should be at the same level as D. affinis (as was the situation when morphotypes were first inserted into this equation), or as subsp. insubrica, for example.

Overall, these various differences in approach have resulted in what must be admitted to be a real problem having been overlaid onto this group. I believe by far the better approach would have been to adjust the taxonomic scheme wherever an error turns up - so far none have that I know of since the earlier nomenclatural/typification adjustment I made - and I really believe now that the morphotype scheme has not only introduced problems unnecessarily, but has really had its day. In my view it was an unfortunate sideline which there is no longer any need for, and I know that the joint author of the original work also regretted having used morphotypes on later reflection.
After all, if you believe "convexa" is an important taxon, then please do publish it now with a proper rank as an additional taxon to those I recognise in the paper on the D. affinis group I have recently submitted for publication. Let's be honest we both have a pretty good idea of where it does belong, don't we? It must be within D. affinis, you may like to have it as an additional subspecies of D. affinis; I would merely disagree on that point and sink it within subsp. affinis. In fact, when you say "Convexa" is in your opinion a distinct and clear form - I agree, but at that very rank! Future botanists can decide for themselves which treatment and rank they felt was more accurate and realistic. If in years to come someone goes into the more minor variants within subsp. affinis (which I would prefer not to name, but which could be varieties or better, forms) I would be able to recognise "f. convexa" at that lower rank along with f. disjuncta, f. atlantica etc. I do not assess it as being a more major taxon, just one of the fairly numerous form-clones, and in my view, not of the same rank as something like subsp. punctata or subsp. paleaceolobata. This sort of doubt was not any reason to replace the whole taxonomy entirely, with morphotypes!

A few comments on questions raised:
(1). Taxon is not a rank, therefore it would never have been a case of referring to "taxon convexa"! I believe an author should have decided where to place it, including at what rank, before publishing a name - which has become apparently formal by usage. Anyway, now, with the treatment of three British species, there should be no real further difficulty in publishing it and saying what species it might be a variety convexa of - as a further subspecies of D. affinis if you think it is worthy of that rank, or as a var. (the Code is so adaptable - nomenclaturally a variety is a variety of its species, not of its subspecies!). Incidentally what is its range in Europe - any idea? I think I saw it in the Schwarzwald in Germany at least, but I did not keep track of every variant of what I now see as subsp. affinis except as can be studied from my many herbarium-collections in NMW, BM (and FR and H).
(2). I think the problem for most people in the earlier stages was not so much that there was little evidence for the taxa (there was actually quite a lot), but that there was quite literally little knowledge of the taxa. I found that many more people were unhappy with a single species divided into subspecies because they felt it was unnecessarily splitting - while only one or two people (I can list them on the fingers of one hand) were unhappy because they felt it an unnatural species-concept, most comments were quite the other way around. Those who didn't like it as one species were only thinking of it theoretically, influenced by cladistics and N. American "taxonomy", and had either very little or zero actual practical knowledge of the group at the time. It is really only after much further practical experience that I have come round to treating species within it, and for quite different reasons from the "biological species" diversion from taxonomic reality!
(3). I am not sure I can agree that naming all significant forms people are likely to come across is so desirable or rather so straightforward - the key word is "significant" and the taxonomic insight to decide practically and meaningfully which are significant and which not. I believe that the two new Morphotypes, "Convexa" and "Insolens" are not so significant - you believe they are. So the way forward is for you to name them formally, then other botanists can decide if they accept those taxa or prefer to sink them. But I would be astonished if anyone wanted to sink paleaceolobata, insubrica etc. - but would also feel they entirely misunderstood them if they didn't place them within D. affinis or D. cambrensis, respectively. It would in my opinion be seriously foolish and display a lack of knowledge of the group to place them all at specific rank, too (assuming we can all now/very soon leave the morphotype scheme behind), but then I don't imagine anyone who knew the group would come to think like that. (4). No such problem exists as being forced to assign disparate elements to inappropriately few taxa. One simply identifies them to subspecific rank - one need not go further if one feels unable to. This is the beauty of standard taxonomic practice and hierarchy. paleaceo-lobata versus cambrensis is nothing to do with that - that is merely a question of misidentification from inadequate keys and description. That occurs in all genera and is much helped to be avoided by the study of authentic herbarium-specimens or guidance from specialists if possible.
(5). I'm afraid any idea of "fixing" the names onto the correct thing can really only be done by means of types - which is why the type system came into existence and widespread use, despite initial opposition. There will surely not be any problem for YOU to select a type that IS representative of your concept - that should be your responsibility to do so! You can then help further by photographing the type itself in your paper. Many of the problem unrepresentative types of the past that you allude to were because the authors had no real knowledge of the genus to allow them to compare their specimen with related species (otherwise through destruction of types, of course), so chose any old specimen at random - if in the 18th C. a plant came from Amboyna it was called "D. amboinensis", with no thought to compare and contrast it with a very closely related species occurring in Sri Lanka as "D. zeylanica", unknown to the author! But this IS another of the great insufficiencies and drawbacks of resorting to morphotypes - there's no formal type system for an informal name - and even if you said it was a type, it would not have to be followed by anyone who decided to formalise the name.
(6). I really do hope and plead that in any publication you may be planning, you do take the plunge and for once and for all tell us precisely what you believe your entities to be - in a formal taxonomic nomenclatural system. I am sure from our conversations etc. that you do indeed have a very good idea now as to how they are most appropriately placed! If it doesn't agree with what I or anyone else thought it doesn't matter - but I feel you really owe botanists the ability to assess your taxa as proper, formally ranked entities. It just would be such a shame to continue to hang on to these "don't know" morphotypes any longer - especially as I am pretty sure you DO know and are well able to make a decision where and how to place them! Then all doubts retypification and meaning of the name can be avoided.

Somehow, I feel, we have to aim to remove the dual, opposing systems for dealing with the Dryopteris affinis group as quickly as appropriately possible - and I feel we can already do so right away. If not, only confusion will continue reign supreme, as it appears to in Britain at the present time. That is why I am quite simply requesting you to abandon the Morphotype system at this point, while continuing your publication and investigation of the group on as wide a scale as possible.
Chris Fraser-Jenkins, Kathmandu.

PART 2 as pdf download

Follow up with comments from Anthony Pigott

RE: [uk-ferns] Re: Dryopteris affinis cx. morph. insolens

Christopher et al.

My comments interspersed below...

Regards

Anthony

 -----Original Message-----
From: uk-ferns@yahoogroups.com [mailto:uk-ferns@yahoogroups.com]On
Behalf Of Christopher Roy Fraser-Jenkins
Sent: 12 November 2006 17:10
To: uk-ferns@yahoogroups.com
Subject: [uk-ferns] Re: Dryopteris affinis cx. morph. insolens

Dear Anthony (and all Borreroids),
Don't worry, I do indeed see the purpose and appreciate the importance of investigating any and every apparently different entity, as it remains possible some might turn out to be more important and significant than they appear at first sight!

Agreed

But I do also see - and this is where I have always differed from the morphotype approach - that there are obviously three major entities, and then a whole lot of others which are clearly much more minor variants, whose morphology does fit into the general pattern of one or other of the three main things. I am pretty confident now that
this is correct and works and I have also found that many others can also see that pattern. It is this that helped convince me to treat three British species at this stage, which I'd previously separated as subspecies. This is on practical, character-based grounds, not because of the biological basis that has influenced some others more.

This is somewhere we fundamentally differ. I believe species are real and either exist or don't, regardless of botanists' ability to identify them.


Ironically, when I first separated the taxa it was against a very uphill struggle to convince anyone (except Hugh Corley) that they existed and should be separated at all! Certain people just couldn't see it in the BM in particular - only to have a turnaround a few years later where suddenly separate species were espoused because the infallible wagnerian school of the USA had made them so, albeit without even seeing the plants, on the principle of biologically different genomic entities (as opposed to any character-based understanding)!

I can't really comment on any initial reaction but I do know that when the D. aff. 'subspecies' were first brought to a wide audience in Chris Page's book, there was much enthusiasm in trying to find and identify them.

I simply don't follow the principles of biological species based on different genomes because, while it may work in some groups, it may not in others. The recent declaration from Gent that all cytotypes in Asplenium will be species, to my mind goes quite against practical common-sense, as well as against the good and carefully rationalised practice of the actual formulator of the research-findings concerned,
the late great Prof. Reichstein.

I wouldn't hold that all cytotypes should be species.

Thus when I mention I now prefer a change to species and am told by a couple of people that, yes, you had chosen the species rank years ago, it must be said that that was not at all for the same reason as why I am now converted to it! - it is,
however, a happy coincidence, that the end result is the same and we are all (or almost all) pleased with it.

Agreed. As you allude to, there are those who would name every different thing as a species, as in brambles. I wouldn't agree with that.

Hugh Corley actually wanted to call AAB (present day D. cambrensis) and AB (present day D. affinis) subspecies of D. affinis, because he felt they were too close
morphologically to be treated as species - in this respect he was more inclined towards the morphological approach I went for, though I do now feel the two can be practicably treated as species. Initially he, too, felt that the three entities I recognised were the three basic things.

Certainly latterly, Hugh wanted affinis, cambrensis and borreri to represent three species - I have an unpublished note from him to that effect.

- but his enormous interest in finding new possible combinations (as also in various other sexual species, which all turned out to be wrong, as did most of his affinis combinations) eventually overtook the practical side - yet on the many occasions I
discussed it with him he definitely didn't plan to make all into species, nor to unrank them, as has in my view so unfortunately been done, albeit temporarily (in which case why publish to say "I don't know"?), by "Morphotypes".

Morphotypes are unranked; see below.

I really think the whole "Morphotype issue" has been entirely unfortunate and a most unusual state of affairs that it was thought appropriate to attempt to replace a clear taxonomic treatment with something designed to state that the subsequent authors concerned felt that they didn't know enough about it. I actually believe that at that time they evidently didn't!

No one was attempting to replace anything else (at least I wasn't). I have always tried to cross-reference my morph. descriptions with formal names where possible. I don't think it's about nor knowing enough - in fact, describing more forms in detail cannot be done without a lot of knowledge.

It seems to me a slightly surprising comparison to make to say the various possible taxa within the general ambit of borreri are not as clear as "'affinis', 'paleaceo-lobata' 'convexa' and kerryensis'.
I do also agree they are generally less distinct than some of those within D. affinis (2x), but what I would definitely have said, as this is what strikes me as more obvious and immediate, is that they are definitely of the same general pattern as D. borreri and do not appear at present to be as clearly distinct as the big three are, inc. D. affinis and D. cambrensis!

Who said they were?

The hierarchy just shouts itself out to me on looking at the whole group. In my view, almost the opposite to yours, kerryensis and convexa, are immediately and obviously of the same general pattern as affinis (as is punctata in Switzerland etc.) -

I was just talking about the ease of circumscribing them - nothing else.

I just can't ignore that natural, visible hierarchy whenever I consider this group as it thrusts itself right at you out of the page/Page(!). With more careful study one can also recognise that paleaceolobata, which looks pretty "abnormal" at first - and that's in a way, how I think of it - is also of the affinis pattern, rather than the cambrensis pattern. This means to me that effectively equal ranking, or unranking, of all these things by "morphotypology" just HAS to be a non-starter right from the word go!

With great respect, I think you're consistently missing the point about morphotypes. They are merely giving labels to the tips of the hierarchy; they say nothing about the hierarchy itself one way or the other. In the same way, all your varieties of the 'old' D. affinis were of equal level; the significance was only implied by the way you grouped them into sub-species and into one species.

Even in your "Morphotypologically" orientated discussions I can't help but detect an inherent, but unsaid flavour of acceptance of the three main things, with others below them - hints that some things are more minor than others and in what general pattern they fit. That's why I can no longer talk about "D. affinis "Insolens" (honest it was just a joke about the name - I actually thought from my poor Latin it must mean becoming in the sun" [preferring sunny localities], interesting to hear it was not that!). To me it has to be shoved over into D. borreri "Insolens", but as you know I prefer not to use a name at all that sounds like a formal name until we find a real name that does match it. In the cases where I have a potentially new taxon that I'm not sure of, as often happens over here, I usually call it after the place, D. wallichiana "Darjeeling form" - or something like that, to ensure there is absolutely no danger of it becoming used like a botanical name - as has unfortunately happened here, with "Morphotype" even appearing to be like a pseudorank, even being abbreviated like ranks, to "mt." - all alarmingly bad, non-botanical practice to me! Only later, when I believe I have got a grip on it and understood it properly, do I change that formula to a Latin epithet and publish it.

Morphotypes are really only like your working titles but made public so people can use them to communicate. They're not meant to be pseudo- formal ranks. I used existing Latin names when they were already in use because it seemed perverse to ignore them and use something completely arbitrary. In practice, I find that in conversation, botanists who know the group just talk about "affinis", "insolens", "kerryensis", etc. and don't actuallymention "morphotypes" or "varieties" very much. There's rarely any confusion.

In a way, I rather feel that it would be a considerable relief for nearly all concerned, whichever side of the fence they currently stand, if one could let one's hair down and almost admit that the basic hierarchy really does exist, even if some adjustments will no doubt become clear and necessary in time.

Of course there's a hierarchy. No, that's not quite true. Of course there's an evolutionary network (not a tree) from which we can almost certainly discern species and where we can probably find meaningful infra-specific divisions.

This would immediately allow the readoption of the universal taxonomic ranks already given them, which not only I find make good practical sense in the field and
herbarium! Do you really think I got it all wrong? - if so, tell me how -

Let me put it this way: I expect that if we both had to draw up the most likely evolutionary network joining these morphs. or varieties or whatever,then we would produce similar results.

or was it more on a principle that modern molecular methods have not yet been applied, so we, i.e. you, should wait? - which is not an established botanical practice by any means and becomes quite irrelevant when one takes a look around the Himalaya etc. at a big fern-flora .....

I would like to see more quantitative evidence - it doesn't have to be molecular although that would be good and probably answer a lot of questions (probably raise a lot more as well!). At the moment it's all too subjective for my liking.

It will be nice to see the photos you are planning - but what will you be using as the basis for the names you use?

I'm just talking about illustrating morphotypes - I wouldn't try to equate them to an existing formal name without being sure of it.

As you know, I mentioned to you when we met recently at the Savill Garden that I was intending to put together my photos (if I can find them all) of the various type-specimens of all the published taxa that I could lay my hands on - and try to get the rest together, too.

I agree. That's why, as you know, I'm keen to get a list and good illustrations of the types widely available.

I explained and I think you agreed, that various names were possibly being used in
mistaken or at least different senses - this particularly applied to the epithets borreri and the silly old mistake of mine, robusta.

I agree that many of the formal names have been wrongly used and I suspect that some of the current formal names may be wrong in the sense that their type is not contained within their circumscription.

They are proper formal names and have types and it is only from these that we can fix the identity and application of the name. So I very much hope that you would apply sufficient caution so as not to disseminate a wrong concept of any names - to get a photo of the type available FIRST, before thinking of posting photos of what you (or I) may THINK is borreri, for example, but which might not be.

As above

One can put a photo of better material (with more developed and less developed forms as well) beside that of the type - which Ken Trewren and I planned to do for the big monograph.

I'm uncomfortable with the idea of 'more developed and less developed forms'. I'd rather think about the range of variation of a natural taxon.

As we see already, it is very much more difficult to get people to unlearn concepts of names than to learn them correctly by typification in the first place. We've got to approach it type first! Anything else is dangerously vague guesswork.

Agreed, but a lot of the earlier formal publication has done just that.

With the nomina nuda "Morphotypes" one can choose whatever one wants to represent the name, though preferably like what you were originally thinking. Even so, trying to formalise an invalid name is a strange exercise in a way.

It's not formalising in that sense, just illustrating to aid understanding of what's being described and talked about.

Bearing in mind that the ICBN (Code of Botanical Nomenclature) makes it clear that any infraspecific rank is a direct subdivision of the species itself, and NOT of any intermediate rank - i.e a variety is a variety of the species, not of the subspecies - could it not now, surely, be more useful to everyone
if you might just take the plunge and describe "Insolens" as a variety, and thus with a type to fix it by? This would agree with your (and my) wish to adopt formal nomenclayure a.s.a.p.! It could always be made a subspecies later if thought to be so - or you could, I suppose, always continue to unrank it in subsequent papers, just as with the previous botanical names that got "Morphotyped" over?

Well, I suppose I might, but I'm uncomfortable about appearing to play nomenclatural games if I'm not really sure about where to place things. (This isn't just being overly pedantic - I've seen unpublished provisional flavonoid analysis that made insolens look more like cambrensis than borreri!) Perversely, having the three at species level, makes things more difficult because one then has to assign all the forms to one species or another, whereas with the old D. affinis they could all be vars. of it. Perhaps that's what I should have done. I was probably too influenced by you always putting the vars. in a hierarchy of subspecies!

Anyway, despite the differences of approach and practice so far, I do think there is a whole lot of common ground - but I just pretend to be a traditional botanist while you wish to discover the Meaning of Life!

Just the Meaning of Affinis!

But I feel that both approaches can actually come together into an honorable and glorious, formal, taxonomic, hierarchical treatment!! Or is that heresy?!

No, I hope so too.

Cheers,
Chris (I can only use 2 fingers on it!)
PS. Excuse my ignorance, but what is that cx., cris-cross thingy?

Do you mean what does cx. mean? - complex.


 Dear All (from Anthony Pigott)

Without wishing to get into the whole affinis issue again here ( I can't type as fast as Christopher!), I thought I'd add a few comments (no particular order).

I'm in the process of putting together a comparative set of images of the distinct morphological forms of the D. affinis complex that have been described previously (in Britain, at least). Hopefully this will help to clarify what's being talked about as even good comparative text descriptions can be difficult to use with these plants. These forms mostly correspond, I believe, on a one to one basis with the varieties (and possibly now subspecies) that Christopher would prefer.

Although I'm comfortable with recognising at least three distinct genetic forms with the 'borreroid' taxa ('borreri', 'robusta' and 'insolens', I must admit that drawing precise morphological boundaries around them is not as easy as with, say, 'affinis', 'paleaceo-lobata' 'convexa' and kerryensis'.
However, I believe that this is probably due to our lack of current knowledge of their environmental variation rather than because they are genetically continuous.

The name 'insolens' does not mean 'insolent' in the usual English sense of rude, cheeky, etc. The Latin word insolens means unusual, unexpected, funny - as in 'funny peculiar'. It came from the time before I realised what it was when often when out in the field with others I would say "I think it's a funny borreri" - usually met with a chorus of "Oh no! Not another funny borreri"! So when I had to think of a name for it, it seemed the obvious choice.

I think it's very good, Christopher, that you now believe that the divisions
with presumed distinct genome combinations should be recognised at species
level. I hope that you won't mind me saying, in the nicest possible way, that some of us have believed that for many years now. I think the original credit for this view has to go to the late (and great) Hugh Corley.
That's why I have described things as, for example D. affinis cx. morph.
insolens (more fully: the morphological form 'insolens' of the Dryopteris affinis
species complex.) rather than D. affinis morph. insolens.

I do think Christopher's point about finding any pre-existing name is very
important from a nomenclatural point of view. I have always said that despite my attachment to the morphotype concept (there - I've said it!) as a useful temporary informal set of handles, I would hope that we can get to a stable set of formal names as quickly as possible.

Regards

Anthony

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Another interesting snippet from Chris giving the background and details of the extant varieties and cv.'s of Dryopteris affinis and Dryopteris x complexa. If you want to use the correct taxonomic names for these 'monstrosities' we grow in our gardens, here they are!! If you want to print this out, download here.

I knew I once knew something about that name Windermere, and your finding it is also a D. affinis agg. thing put me onto it. Here is an extract from the big monograph on D. affinis that I did about 15-20 years ago, but rather gave up on for various reasons (though I'm now very much onto it again, and with Ken Trewren, who has done a huge lot of work on the group). I did a section on each different level of variation, as I saw it, from minor forms up to subspecies - and nowadays it will be up to species, too. I was planning to illustrate them all, but no need, I think, as Martin Rickard did it in his super book, which we discussed together - though I think we still hoped to make a special booklet on D. filix-mas agg. cultivars some day, but then we all only get one life and nowadays it has its non-pteridological diversions, too!
Anyway, I mentioned 'Windermere' in the section on cultivars/abnormal ties, part of which I put here (Ghosh, sorry, I see it's a bit long, unlike most modern papers, which I reckon are usually about as interesting for detail as scaffolding in a cold desert!):

"It is of use to attempt to place the monstrosities into their modern botanical species or subspecies, especially in complexes of related species, though this can be difficult and some cultivars can be quite misleading as almost any parameters of morphology can be altered in them, including those that are diagnostic in normal plants.
In D. affinis a quite large number of monstrosities has been collected in the wild or raised from spores of other monstrosities in gardens, though many of them were originally (and often still are) attributed to D. filix-mas in a wide sense. A number have been separated and assigned to D. affinis (sub. Lastrea pseudomas Wollaston) by Druery (1910), but none had been assigned to the subspecies until investigated by Fraser-Jenkins (summarised by Rickard 2000). However it has proved to be quite easy to place most of the cultivars of D. affinis from their morphology. This has been confirmed by checking the diagnostic difference in spore-size and regularity between subsp. affinis and the other subspecies, in cases where the pinnules have been altered beyond recognition. Several cultivars were also found to belong to the hybrid D. x complexa, especially some of those raised in gardens from spore-sowings and presumed to have arisen from chance hydridisation in the prothallial cultures, which are seldom completely free of contaminants. Some twenty cultivars of D. affinis and its hybrid known to the author to be still in existence in gardens today (many formerly cultivated by him at Bridgend) are listed here in their subspecies. Their survival in cultivation over nearly one and a half centuries in some cases represents a remarkable tribute to both the"Victorian Fern Craze" (see Allen 1969), with the diligent if over-keen collectors and propagators of last century and fine gardens, and to the care of the subsequent generations of British fern-gardeners. Many were lost to horticulture during the Second World War, when many gardens had to be used for vegetables, but some important and fine cultivars have survived and been kept going up to the present day. With the rise in popularity of horticulture over the last twenty years or so several new ones have also been discovered or raised, including in the large commercial centres in Holland. But it must be said that some of them have been unfortunately careless in their nomenclature and have renamed cultivars already known and named in British gardens, which has proved to be very difficult to control as some of the big firms such as Royal Lemkas are almost obsessively secretive and usually prevent anyone at all from having access when the opportunity could be taken to check their nomenclature (but see Rickard 19***). Although a few noticeable monstrosities also occur either wild or cultivated on the continent, the existence of so many British ones and their survival in our gardens is a unique and somewhat unusual situation (as pointed out by Druery 1910). But in Germany etc. most such plants (e.g. the large collection of Dr. E. Rosenstock of Gotha in eastern Germany, sent to Berlin Botanic Garden and a number of others throughout Europe) were lost [***? did Rosenstock ever publish anything on his varieties? If so, cite*****]. This was mainly as a result of the more severe effects there of the Second World War (see Alston 1946). From experience in the field and herbaria, where such plants are often represented, it is confirmed here that they either do not seem to occur in nature on quite such a large scale anywhere else in the world, or have not been so diligently collected and appreciated outside Britain.

Cultivars of D. affinis known to survive in gardens are:
1. D. affinis subsp. affinis var. affinis 'Cristata' (T. Moore & Houlston 1851), originally found wild at Caercleugh Estate, Charleston, near St. Austell, Cornwall by 1850, and other plants found near Ilfracombe, Devon. Often known in gardens as 'Cristata the King', though incorrectly so, as this was only a descriptive remark by Druery (1910: 156), who said it was often termed the "King of the Male Ferns", perhaps following Sim's (1859) description of it as forming a noble mass of fronds. All authors agree that this is one of the finest fern cultivars of all, with regular heavy cresting (multiple bifurcations) at the pinna-tips and frond-apex, forming a fringe around the semi-erect fronds. Interestingly, it was the second apomictic fern discovered (de Bary 1878, and see Manton 1950) and was later investigated by Döpp (1939), who found it to be diploid apomict.

2. D. affinis subsp. affinis var. affinis 'Trippett Windermere' cv. nov. (Fraser-Jenkins 2004, ined.), new cultivar. It was being grown under the name 'Cristata Windermere' [ined.] at Sizergh Castle, Kendal, Cumbria, in 1986. It was originally found near Bowness, Cumbria by Ron Trippett of Leeds. It is generally similar to 'Cristata', but has less tight and regular cresting and a more lax frond. [***? cite type etc.?***]

3. D. affinis subsp. affinis var. affinis 'Cristata angustata' (T. Moore 1859/Sim 1859), originally selected by R. Sim from cultivated sporelings raised from 'Cristata' given him by Wollaston about 1854. Cv. 'Cristata' grown from spores sometimes varies towards narrower forms but the present plant is spectacularly narrow with the same cresting at the margins and is also slightly dwarfed.

4. D. affinis subsp. affinis var. affinis 'Crispa cristata angustata' (Druery 1910), originally raised by W.H. Lang's nursery at Kircaldy, Perthshire, probably from spores of 'Cristata angustata'. It is a very dwarfed, narrow fronded cultivar, with the pinna-apices and elongated frond-apex cristate, somewhat similar to 'Cristata angustata', but smaller and more crowded in all its parts and with crispy-undulated pinnae.

5. D. affinis subsp. affinis var. affinis 'Polydactyla Mapplebeck' (Wollaston ex Phillips 1899, Wollaston ex Druery 1910), one of many 'polydactyla' cultivars, of which Moore's original one (of 1857) is a cultivar of D. filix-mas. First found in 1862 in Westmoreland (now Cumbria) by J.E. Mapplebeck. Rickard (2000) illustrated a plant he obtained under this name from Kaye's nursery in Lancashire, but his plants look very similar indeed to 'Polydactyla Wills' and perhaps do not quite match Druery's photographic impression. Herbarium-material in Moore's herbarium should be examined in order to be sure of the identity of this cultivar. Druery cited the name 'Mapplebeckii' (T. Moore) as a synonym, but if published [**?check*** ] this would presumably be the earliest name and must have been given in order to distinguish it from Moore's 'Polydactyla' . This cultivar is like a fairly normal subsp. affinis but has loose, multiple bifurcations towards the pinna-tips and at the frond-apices, which fork and then bear a small crest at the tip of each fork. 'Polydactyla' cultivars are a step down from 'Cristata' in terms of degree of abnormality and are also not as regular in their characteristics.

6. D. affinis subsp. affinis 'Polydactyla Wills' (Druery 1910), found wild in south Devon "a few years later" than 1862 by J. Wills. Of botanical interest chiefly because it was first investigated by Farmer & Digby (1907), who obtained a very approximate chromosome count for it and described some early details of apomixis, later made more exact by Döpp (1939) and finalised by Manton (1950), all studying this cultivar. It is probably due mostly to Manton's cultivation of this plant, obtained from Prof. W.H. Lang at Manchester University and then cultivated at Leeds, that it still survives today. The plants investigated by Farmer & Digby and Döpp were also obtained from the same source (Manton pers. comm., c. 19***). The normal parts of the pinnae are typical of var. affinis and only the cristate, divided pinna- and frond-apices are different. The difference from 'Polydactyla Mapplebeck' is very small and the two evidently belong to one Cultivar Group (Polydactyla) ; 'P. Wills' is perhaps more compact than 'P. Maplebeck'.

7. D. affinis subsp. affinis var. affinis 'Resendeana' [***? valid, in Latin?***](de Rezende-Pinto 1969) Fras.-Jenk. (2004), discovered at Valongo, Portugal, by Prof. M.C. de Rezende-Pinto in 1944 and named after his mother, though first described as a forma valongensis de Rezende-Pinto. Although the cultivar epithet is in Latin, it is to be retained under the Horticultural Code (1995: Art. 17.3) as the name was formerly published under the Botanical Code. This cultivar is close to 'Polydactyla Wills', but perhaps slightly narrower, towards 'Cristata'. The locality contained several species and other fern-cultivars which may have escaped from past cultivation and it is quite possible that 'Resendeana' may have originated in Britain and might also have been named there previously before being cultivated in Portugal. But it does not quite seem to match any other cultivar yet known to the author. Interestingly de Rezende-Pinto independently made observations on the apparently new pattern of cell-division leading to sporogenesis in this plant without realising he was redescribing some aspects of apogamy, as outlined by Döpp and Manton many years previously, and was surprised to hear of this when the present author met him in Porto in 19***. The plant is cultivated outside the Botany Dept., University of Porto, and by J.I. Lintner's nursery, Nieder-Ofleiden, Marburg in 1987 and original stock was given by the author to Kyre Park, though since lost unless fortunately maintained by some customer of Rickard's Hardy Ferns.

8. D. affinis subsp. affinis var. affinis 'Crispa' (Sim 1859), found in Wales by J.W. Salter in the late 1850s and distributed from spore-sowings by Sim's nursery. It is an attractively dwarfed plant with a crispy, brittle texture and dark-green fronds and has recently been cultivated by Lemkes (Lemkes en Zonen B.V., Koninklijk Tuinbouwbedrijf, Alphen a/d Rijn) on the Continent, as well as occasionally in British gardens.

9. D. affinis subsp. affinis var. affinis 'Crispa gracilis' (Phillips 1899, Druery 1910), occurred spontaneously in a spore-sowing from 'Crispa' grown by Dr. Lyell in 1866. It is closely similar to 'Crispa', but smaller, narrower and more compact. It is rather more frequently grown in British gardens than 'Crispa' and has also been disseminated by Kaye's nursery (see Kaye 1968) as 'Crispa congesta'.

10. D. affinis subsp. affinis var. affinis 'Pinderi' (Moore 1856), discovered near Elterwater, Cumbria by the Rev. G. Pinder in 1855. This is a remarkably narrow-fronded cultivar with erect, linear fronds and the pinnae only shallowly lobed and must have been a great excitement for the Reverend gentleman to discover. It has sometimes been confused in gardens with D. x complexa nothosubsp. complexa 'Stablerii', but can be confirmed from its good spores if in doubt.

11. D. affinis subsp. borreri 'Furcans' (T. Moore 1859), first found near Huddersfield, Yorkshire, by T. Stansfield in the 1850s, and later in Surrey. It resurfaced from Royal Lemkes Nursery, Holland, in the 1980s under a new name 'Terrilis furcata', but was quite possibly from the same origin, as was the 'Furcata' at J.I. Lintner's nursery, Nieder-Ofleiden, Marburg in 1987. The pinna-apices are usually forked, rarely more than once, with divergent tips, as can
be the frond-apex, but in young plants only a few pinnae fork. In other respects it is a normal subsp. borreri.
12. D. affinis subsp. borreri 'Polydactyla Dadds' (Phillips 1899, Druery 1910), grown by J. Dadds, at Ilfracombe, Devon, in 1872, from spores of 'Furcans', from which it is spectacularly different, presumably by further mutation. It is a typical 'Polydactyla' , but laxer and less deeply forked than in 'Polydactyla Wills' and has a wider lamina-base. The normal segments in the lower and mid parts of the pinnae are rather longer and narrower than in normal subsp. borreri and, though less lobed, indicate that it belongs to subsp. borreri. The botanical epithet var. polydactyla was preoccupied by Moore's variety (of D. filix-mas), and 'Polydactyla Dadds' was not published properly as a single word (Dadds was given as the finder, rather than part of a name), but can be used as a cultivar name.

13. D. affinis subsp. borreri 'Schofieldii' (T. Moore 1859, Sim 1859), found near Buxton, Derbyshire, by J. Schofield prior to 1859 and brought into the horticultural trade by Stansfield's nursery, then at Todmorden, Yorkshire. It is a dwarf plant with a once-forked stipe and upper lamina. This cultivar is very nearly extinct and as far as is known to the author survives only in the garden of Bud Lyle, Grange Nurseries, Aloa, Clackmannanshire, Scotland, where he was kindly shown a small, delicate, but typically characteristic plant of it in Sept. 1984, which was of known descent from the original finding.

14. D. affinis subsp. borreri 'Revolvens' (Phillips 1899, Wollaston ex Druery 1910), found in a garden fernery at Iffotsholme Farm, Troutbeck, Cumbria, by F. Clowes in 1868, having been collected locally. It has been preserved mainly due to its propagation and sale by Kaye's nursery. It is not much different from normal subsp. borreri and when pressed can be indistinguishable, but the young fronds have a distinctive tubular shape due to the pinnae curving around to meet their opposite number beneath the frond-axis. The fronds are also markedly erect, so when young are rather outstanding, though adult fronds become flatter.

15. D. x complexa nothosubsp. complexa 'Stablerii' (Druery 1910). This cultivar appeared spontaneously among sporelings from a sowing of the narrow-fronded D. filix-mas 'Barnesii' (Druery), with its incised sides to the pinnae, at H. Stansfield's nursery at Sale, Reading, Berkshire. It can be deduced now that the other parent can only have been D. affinis subsp. affinis presumably var. affinis, whose stray spore must have arrived by chance and the prothallus cross-fertilized. The fronds are very narrow and are generally similar to D. affinis subsp. affinis 'Pinderi', which which it has sometimes been confused, but are considerably taller and rather wider. The pinnae are more deeply, pinnately lobed, the pinnules are longer, with more rounded apices and characteristically lobed sides and the indusia are thinner and lift and shrivel considerably on ripening. A plant obtained by the late Prof. T. Reichstein, at Basel, from Dr. W. Gätzi (originally from Göttingen Botanic Garden), but mistakenly under the name D. affinis 'Pinderi', was found to be tetraploid, with approximately one set of bivalents and two sets of univalents in the 16-s.m.c. sporangia (see below, sub D. x complexa). Its spores were mostly abortive, with some very large, apparently good ones present and it was thus reidentified by the present author to be a cultivar of D. x complexa and was also identified by him as being cv. 'Stablerii' and not 'Pinderi'. [***? check spelling, stablerii?** *]

16. D. x complexa nothosubsp. complexa 'Grandiceps Askew' (Askew c. 1955, Kaye 1968). It is almost sure that Askew raised this cultivar as a hybrid with D. filix-mas from spores of D. affinis subsp. affinis var. affinis 'Cristata'. It is a very tall plant with rather weak "polydactylous" cresting of the pinna-apices and frond-apices, exactly as would be expected from such a mixture; the normal pinnules are typical of the hybrid. The spores in a plant obtained by the author are abortive with some very large, apparently good ones also present, as in normal plants of the hybrid. This plant was one of quite a number of F. Askew's surviving ferns at his garden at Grange, in Borrowdale, Cumbria, given to the author by Mrs. Askew in Sept. 1986. Plants cultivated by the author at Bridgend went to Rickard's Hardy Ferns nursery at Kyre Park, Shropshire. It has also been disseminated from Kaye's nursery and was being grown by Prof. R. Maatsch at Herrenhausen Botanic Garden, Hanover, in 1987, though unlabelled and not mentioned in his book (Maatsch 1980).

17. D. x complexa nothosubsp. complexa 'Atkinson's decomposite' cv. nov. (Fraser-Jenkins 2004), new cultivar. This is a rather stiff-fronded plant with forked pinna- and frond-apices, decomposite, narrowly lobed segments, slightly thick indusia, which then shrivel on ripening, and dense pale scales on the stipe, becoming all narrow, but still dense, on the rachis. The spores are nearly all abortive but with some very large, apparently good ones present. It was almost certainly raised from D. affinis subsp. affinis var. affinis 'Polydactyla Wills' hybridised with D. filix-mas 'Decomposita' (Phillips/Druery) , by F. Atkinson, of Newley Bridge, Staveley, Lancashire. An offset of a clump surviving from his collection was kindly given to the author by the late R. Kaye at Silverdale in Sept. 1986, under an unpublished name of Kaye's "Decomposita polydactyla' , which cannot be used now. It was subsequently passed from Bridgend to Kyre Park, but met its demise there later unless preserved by one of the nursery's customers.

18. D. x complexa nothosubsp. probably critica 'Ramosissima' (Phillips 1899, Moore ex Druery 1910), often listed as "Ramosissima Wright" after its finder, but in error. It was discovered in north Wales by R. Wright of Stone, Staffordshire in 1864. The stipe is forked several times above the base and the frond-apices are cristate. It produces sori which do not ripen fully, but the present author has found that while most sporangia do not develop, a few of the most mature ones contain abortive spores. Although this could be for other reasons, it is likely that this indicates the plant's belonging to D. x complexa and the matt laminar texture, acute teeth and pale stipe-scales tend to suggest nothosubsp. critica more than anything else. However it is not certain that it is not D. affinis subsp.
borreri; only a mitotic chromosome-count could resolve its true identity for sure. As the segments are crowded and reduced in size it is difficult to guess its identity from the normal diagnostic characteristics of pinnule-shape etc. Less branched and crested plants of this, partly reverted to normal, were grown by the late R. Kaye at his Silverdale, Lancashire, nursery and listed by him (Kaye 1968) as D. borreri 'Ramosum furcillans', again having the poorly developing sori.

19. D. x complexa nothosubsp. critica 'Abasipinnula' (E.J. Lowe 1867). This is a very minor cultivar in which the bases of the lowest few pinnules of lower and mid pinnae are narrowed and cuneate. Plants survive from Lowe's old collection in his garden at Shire Newton Hall, near Chepstow, Monmouthshire and were found by the present author in 1986, when he was kindly allowed to cultivate an offset by the Knight family. The spores are highly abortive.

Many other cultivars listed and figured in 19th. or early 20th. Century literature have been lost from cultivation during the two World Wars, and are now known only from herbarium-specimens . A programme of further research is required in the Moore herbarium at Kew (and in Paris for Wollaston's collections etc., and also Berlin for Rosenstock's) to reveal further details of first collections, localities and cultivar groups before a comprehensive account of extant Dryopteris cultivars can be finalised. Subsequent to Moore's more botanically correct accounts, several authors (e.g. especially Phillips 1899, partly Druery 1910, and later works following them) have added the name of the finder as if either a second part of the name, or the authority for the name (rather than the plant). This can be misleading in terms of attribution of the name when it has been published in the form of a botanical name, but is acceptable as a cultivar name."

Hope this might be of use to cultivologists - and I'd be interested to know of any other cvs. of the D. affinis agg., as
obviously this won't be complete. Incidentally there is a small body of literature about how these cultivars (botanical 'Monstrosities') arise - it seems most are genetic, as found by Dr. Andersson-Koto
years ago. I have some information on that, too.
Cheers,
Chris F.-J.
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