Volume 5, Chapter 17:- New populations of Haworthia chloracantha, Haworthia parksiana and Haworthia kingiana

In Chapter 1 of  Haworthia Update Vol. 2, I discussed H. chloracantha and H. parksiana in the context of H. floribunda. Fig. 8 in that publication is labelled “North of Herbertsdale” when in fact it is MBB7425 from the Wolwedans Dam north of Great Brak (see fig.1). This was deliberate and not seriously misleading as the plants from the two respective populations are virtually identical. The correct images for that “north of Herbertsdale” are in Haworthia Revisited and labelled JDV87/80 and 97/138.

Recently I had an opportunity to visit Herbertsdale in connection with chameleons and took the opportunity to visit a locality also north of Herbertsdale, but a lot closer (see Figs 2.1-2.11. MBB 7825) reported to me by Mr. Jaap Viljoen of Swellendam.

The plants there are smaller and much less clump-forming than JDV87/80=97/138 and are more reminiscent of H. parksiana. Unlike JDV97/136 (see figs. 3) that are solitary, small, dark-green plants growing in the lichen on a steep east-facing conglomerate slope, the plants are quite green in colour and often the tips of the leaves are slightly rounded and flattened as in H. floribunda. The habitat is west-facing on an old gravel river terrace and much drier and warmer than for JDV97/136, in sharp contrast to the very steep north-facing clay cliff where KDV87/80=97/138 occurs. In neither of these latter two sites are other plants significant, while at MBB7825 the plants are under renosterbush (Elytropappus) and related vegetation. I should mention that my wife on seeing the plants commented on the similarity to H. parksiana and this is not an observation that can simply be dismissed.

About 3km south of Herbertsdale (see fig. 4, MBB7827) we again found very similar plants, perhaps a little more spinose and slightly more clump forming. Neither of these two new populations fit comfortably in the existing formal varieties, as indeed is true for so many populations in most of the other species. I do not think it is rational to attempt formalizing names for them as there is quite substantial variation in each population. Others may do so if these plants ever get into general cultivation from vegetative propagation and onto lists.

From Herbertsdale we went to Klein Brak to investigate a set of four populations on the farm Rooiheuwel as indicated to me by Gerhard Marx. One of the species supposedly there was H. pygmaea and it was rather distressing to find that the Water Affairs Department has seen fit to run a bulldozer over the area  with no indication that there was any need for doing so. However, we did find the other three species indicated by Gerhard. One is H. kingiana (see figs. 5.1-5.4 MBB7835).

The others were H. parksiana (see figs 6 MBB7828)

and H. chloracantha (see Figs 7 MBB7829), but this requires a little bending of the truth.

This H. parksiana site is about 400m southeast of an old locality of mine on an outcrop of rock virtually jutting into the road and now largely obliterated. At Gerhard’s site (viz MBB7828) the plants are small and while there is little doubt that they are generally alike to H. parksiana, there is definitely some degree of hybridization and some of the plants may actually be H. chloracantha. This is a rather difficult decision to make on the basis of stressed plants in habitat. This habitat is very restricted to a patch of fairly level aggregated conglomerate so that the plants are virtually in tiny shallow soil and lichen encrusted patches. A little distance further north there is another population to which we assign the name H. chloracantha. This is above a steep loose conglomerate cliff with a little more soil and immediate bush. The plants have rather abbreviated leaves and a darkish colour. There must be some doubt that they are discretely H. chloracantha. The habitat is not so restricted and one is compelled to ask why the plants then so are.

I try to avoid the unanswerable question of how these two species that flower together and seem to be able to hybridize quite freely, maintain their identity and come to occupy such specific, localized and widely separated sites? It would be nice to think that the technology is now seemingly available in DNA sequencing to begin probing these questions, and I will point, with reference to chameleons (see next chapter), to what I think is a disturbing reliance on that technology without the necessary knowledge and experience of what actually can be seen in the field. ♦

Volume 5, Acknowledgements

In some cases acknowledgements have already been made in respect of some of the populations discussed in these chapters. New acknowledgements are Mr. Artho Saayman of Platkop, Mr. and Mrs. Wilhelm Zietsman of Kruis Rivier, Mr. and Mrs. Coetzee Uys of Morning Star, Messrs. Chris and Pieter van Deventer of Kransriviermond, Johan Groenewald of Buffeljags, Clive and Duprecia Stramrood of Kadies Landing, the Uys brothers  of Sandfontein and Meerlus, the Steenkamps of Appelbos, Annatjie and Jaap Viljoen of Swellendam, Hector Odendaal of Dankbaar, Ryno Stander of KomseRandte, Tineke Kraaij and Carli Venter of Bontebok National Park, Anne Lise Vlok and Rhet Hisemann of W Cape Nature Conservation Board, Stiaan Conradie of Lower Breede Conservancy. Others such as to Mr. and Mrs. Anna and Arno Steenkamp of Anna’s Farm (Oudekraalkop) for their generosity and hospitality, Mr. and Mrs. Hennie van Deventer of Koppies, Mr. Dirk Papendorp of Voorstekop and Uitvlugt,  Mr. Uys Willemse of Goedverwagting, need repeating. We are most grateful to Dr. Paul Taylor for an albeit failed excursion to the west bank of the Duiwenhoks River and similarly to Nico DeJager of Victoriasdale for an interesting but fruitless search of the Wankoe Randte and a little more success on Klipheuwel, Mr. H. Eksteen of Grootkloof and Mr. M. Dippenaar of Diptka for so kindly allowing us access to their properties. There are many other landowners and people we contacted with less significance in respect of places actually visited or in terms of plants found but no less in respect of the universal kindness and helpfulness of landowners.

There are many other peripheral and significant contributions from people (not necessarily agreeing with what I say nor how) such as Prof. Richard Cowling, Dr. Syd Rhamdani, Diederik van den Abbeele and even people who have no connection to Haworthia, but who share the same passion for nature and for plants … Mrs. Hettie Conradie of Worcester who makes us feel as though it is we who own her home and Mrs. Anso LeRoux who may both be housewives, but biologists by nature. Gordon Rowley has demonstrated the value of humour in a field where there seems naught else to do but tear hair out. Etwin Aslander in his quiet and effective way has been a great friend.

Then there is Gerhard Marx, whose dogged refusal to blindly accept anything at face value, has been most helpful in so many respects. Kobus Venter has always been a great friend and mentor in relation to the ways of the world and I am extraordinarily grateful for his unwavering support and kinship. He and Mirna, in graciously allowing us the use of their holiday home at Stilbaai facilitated this work. Steven Hammer has been another remarkable personage in my life and I am most grateful for him being the sensitive, perceptive, empathic and remarkable man he is.

Lastly where I have used the term “we” I have included my wife, Daphne. This is actually with great reluctance because anything to do with taxonomy and nomenclature brings with it a cloud of derogation and negativity – what to say of the negative attitudes that surround the word “collection”. It seems that in past civilizations there was great respect for nature and many of the rites and rituals were an appeasement to nature for the benefits she bestowed in terms of what she provided in the way of food, clothes and shelter – the basic needs of man. Daphne has provided me with the trust and support of a true companion and I regard her very dearly. There is no need for her to remind me, as she seldom does, that few other housewives would traipse so blithely and happily through the wilderness with someone who does not seem to know where he is going or what he is doing next.

I feel a need to explain again that I became a taxonomist by default where there are also systematists and nomenclaturalists and I would not wish to be any of these. We as individuals are all inclined to make bold and general statements based only on what we know and ignore the very much greater body that we do not know at all. I would say that it is depressingly sad that information is processed and distributed as knowledge often from a very limited experience and limited data base. After 45 years of this I actually am painfully aware that there is still a lot that I do not know about. I make no apologies for a revision that is precisely the sort of product that comes from constant testing of a hypothesis. I take it very remiss of a publisher that did not honour the integral part of a contract to further research and validate the work. I am extremely grateful to Harry Mays for publishing a vast series of articles that were originally intended for a home based publication but mindlessly lost in a mire of conflicting interests.

While Alsterworthia so kindly published the Update Volumes 2, 3 and 4, I continually feel that this is not the environment I am comfortable in. Bill Keen was one of the first to publish anything of mine and is the epitome of server to a group. My wish is to see and serve a user group such as the Haworthia Society and see it go from strength to strength. ♦

Aloe pumila, Haworthia pumila; what or who is confused?

With some astonishment I learned recently of an opinion that the name pumila as used in Haworthia is confused and therefore should be discarded. There is no doubt that it is difficult to unravel the literature and the usages of the name, but I think there are a number of important questions which should first be answered. Is the name confused or is it us and others, as individuals, who are confused? It is worth asking such questions because names are words in the process of communication and mutual understanding around which all knowledge and its sharing revolves.

If the facts of the matter are properly examined there is a clear path of events. It may be very complex and take many words to explain, but it is there. If a reason has to be sought for confusion I have no doubt that it can be tracked to the door of the International Code of Botanical Nomenclature. My reasons for saying so should become clear.

Paul Forster, who is himself a professional and highly qualified taxonomist, commented on my treatment of the name H. pumila as follows “We do not live in a vacuum.” Here he suggested that I had failed to consult authority and that there was professional opinion available that would have resolved my confusion.

If one thinks about Paul Foster’s statement it is quite evident that:
1. he does not consider himself able to resolve the problem (this is a valid point because to do so does require knowledge of the elements involved, and also requires insights into the intricate mechanisms of the Code which are beyond most).
2. he simply believes that there is someone in the hierarchy who is competent and able to do so.
3. I had not thought there was.

The latter point is simply not true as I have consulted almost every single botanist and non-botanist (yes, many non-botanists are free to dabble in plant taxonomy because of the non-scientific nature of the activity) available to me. A list of names and also authors and published references would fill a few pages.

Turning to the name “pumila” itself, it can be shown very clearly (botanical terminology to mean something quite the opposite, but which I use here in the true sense of understandable) that Linnaeus used the name for four “varieties” of small Aloe. Confusion certainly followed about which of his names should be used, but there should be no doubt about which four these were.

The problem arises out of the Code and how, relating to names for those four “species”, it is now interpreted and by whom. It becomes a juristic problem requiring a great deal of fancy intellectual footwork, complicated by an approach to the code which largely ignores the basic intention to bring stability and uniformity to plant names. It also involves use of words like “validity”, “legitimacy”, “intention”, “relevance”, “strict interpretation” and “according to article…” with gay abandon.

Without going into the very extensive historical detail which includes the fact that the names margaritifera and pumila were used for the same single thing, it is almost sufficient to say that Col. C.L. Scott took the plunge. Defying the predictable response, he took the assistance of Dr. L.E. Codd, who was then and still remains a highly respected taxonomists. Together they concluded that because of what and despite what, anyone had done with those two names, the name “pumila” was valid and correct for the species represented by the illustration t10 of Commelin 1701.

They argued that despite the fact that the name had already been used in Haworthia for another species, this did not preclude its correct use for the one represented by the Commelin illustration. The fact that confusion may have arisen and continues to this day, is to my mind correctly laid at the door of Dr. W.T. Stearn, paragon and patron saint of plant taxonomy. He wrote a paper in 1938 in which he annotated the names of Salm-Dyck and reconciled them with Berger’s revision of Aloe in 1908. What he did was also to  typify the name Haworthia herbacea (Miller) Stearn. In my opinion, had he known anything about the actual species involved, he would have recognised the problem arising from the inclusion of “pumila” in the history of that name and resolved it accordingly.  He did not.

After Scott (and Dr. Codd) had reached a decision in 1978, I still remained in some doubt using the name “pumila” until forced into a decision when I undertook to write a revision of the genus myself. I had been asked to write a synopsis of the species of Haworthia occurring in the Cape Floral region.  This request was by the taxonomists Dr. J. Manning and Dr. P. Goldblatt and I in turn asked them (as had become my routine practise with taxonomists) to clarify the use of the name “pumila”. They contacted Dr. Fred Barrie who I presume is a figure in the hierarchy alluded to by Paul Foster, and the following was his reply. “According to the Linnaean Typification Project database, Aloe pumila var. pumila was lectotypified by Wijnands (Taxon 34:310, 1985) on Commelin, Hort.Med.Amstel.2:t.10, 1701. Linnaeus cited this figure under A. pumila var. margaritifera in Species Plantarum (p322) as “Comm.hort.2,p.19,t.10.”  Consequently, var. margaritifera and the autonym are synonymous. Var. pumila, as the autonym, has priority.”

There is more to the reply, but it does not address the problem of the different usages of the name “pumila” by various authors until its inclusion in Haworthia by Duval for a species based on a different type.

There is no confusion in this. It is simply a question of how the Code directs that the name should be treated. I was confused over the issue. I did wade through the detail of synonymy with Mr. Larry Leach and subsequently with Dr. Peter Bruyns. It was evident from this arduous process that, was “pumila” not available for the species in question, the name “maxima” should be used. There is absolutely no reason for confusion about which species I am referring to, nor about the names available or the way in which they have been used since Scott 1978. But I decided to treat the name “pumila” according to Dr. Barrie’s response. This suggested to me that Dr. Codd and Co.l Scott were correct and that “pumila” as used by Aiton and Duval was incorrect and did not preclude the use of the name as typified by Wijnands. The essence is that Linnaeus used the name “pumila” as a prime name in Haworthia and it should sensibly remain there.

What is confusing and what should confuse everyone, is that there are persons who feel the need to contest the issue. The need only arises from personal feelings and the fact that the Code has generated this vast arena for endless vain debate. Where a name has so convincingly and obviously been used in the literature of the time, it obviously meets the need. Changing it, or attempting to change it, generates confusion. We do not have to feed on it. ♦

Haworthia mirabilis magnifica, MBB6651, S Riversdale

I think we have to find a way to deal with this issue of names. Let us try H. magnifica. It originated in a population from the Frehse Reserve SE Riversdale and it is truly difficult to circumscribe all those individual variants. The name has also been attached to a number of other plants and populations from various places. The name magnifica is not clearly assignable beyond individuals that can be said to resemble the type (an illustration) and there are individual plants in the Frehse population that do not accord with either picture or description. There are individual plants and populations going all the way to near Caledon that confound the name still further. In my opinion the Frehse reserve plants belong to a single system that I consider to be H. mirabilis (and I am not so sure that it is not bigger still). Do we just drop the name ‘magnifica’ and use locality? So is it better to say H. mirabilis magnifica (Frehse Reserve) and H. mirabilis magnifica (3km S Riversdale) or H. mirabilis magnifica (Windsor) for the variants that occur at each of those places?  Also H. mirabilis jakubii (Goukou) that I think is a connection between H. mirabilis magnifica and H. mirabilis paradoxa (Vermaaklikheid and on to Infanta). We could use this system and also the Breuer and Hayashi names attached to other mirabilis populations in the Riversdale area. The disadvantage of place names is that they convey nothing to people unfamiliar with local geography and to them there may be no difference. On the other hand that the formal Latin names may be restricted by the accompanying description and illustration, and not convey the variations that occur in the various populations.

The Haworthia pollinator

While I have seen and recorded a Solitary Bee species pollinating the flowers of Haworthia, I have never succeeded in photographing this pollinator. Slightly larger than the ordinary Honey Bee, it is a very rapid and busy flyer and does not spend time at any one flower. Here in Cape Town it is relatively rare and there has been an odd season where I have not seen it at all. It makes a nest consisting of a short tunnel dug into the ground where it makes a series of nectar and pollen filled cells in which the eggs are laid. At Worcester these insects were very common and on one occasion I came across them nesting in large numbers in a small patch of bare gravelly clay. It is remarkable how they were obviously able to recognize and home in on their own small tunnels. It does come into relatively insect proof enclosures and can make isolation of plants for pollination difficult. It is unlikely that flying distance has been measured and it seems very unlikely that it will match the observed maximum of the Honey Bee at 13km (8miles). However, I do not know the length of the life cycle and the relation of the feeding/foraging activity to its nesting behavior. It may be possible that feeding but non-nesting bees disperse over greater distances.

The chance to take the two photographs presented here came quite by chance. I was in my plant house that is partly accessible to insects, and saw the bee on a Haworthia flower stalk. Its jaws were clamped on the peduncle and it was totally stationary. I took a few pictures and then attempted one from another angle. In doing so I had to dislodge a second peduncle from nearly under the bee. The slight difficulty in doing so did not register properly. I took a picture and then suddenly the bee came to life and flew off to resume foraging in the house. But it soon settled on another inflorescence (on a plant at another table) again on the peduncle with its jaws clamped on the talk as before. I removed some pollen coated threads hanging from its back legs and left. While downloading the pictures I saw that I had photographed a spider on the peduncle under the bee in my second angle. So went back to see. The bee was still stationary on the flower and then I saw what I thought was the same spider crawling over the bee. Somehow I disturbed the bee again and it flew off up into the shade cloth in seemingly healthy fashion. The next day I thought I would check the original peduncle to see if there was any web. Much to my surprise I saw the original spider and no web. I have seen these small yellowish spiders on flowers before and also noted small amounts of web. Very occasionally there have been dead flies sitting attached to the flower stalks but not with any noticeable web.

So it is still a bit of a mystery if this small spider, or spider pair, could actually capture and immobilize such a bigger potential prey. Certainly the bee I observed had encountered web and may have grasped the stalk to free its hind legs to work the web off. I did not observe anything more than minimal movement to do so.

Regarding pollination. I was not doing any controlled pollination during which I would have been checking for the presence and exclusion of the bee. But what I did observe was seed set on H. limifolia clones that I could not achieve although in desperation I have transferred pollen between different collections. The bee had an advantage in also bringing pollen from unrelated species. This was on H. limifolia ‘gigantea’ and on H. limifolia ‘glaucophylla’. While I have set a few capsules on the former, I have achieved none on the latter. What is interesting is that no seed was set on field collected clones of H. limifolia presumably ‘keithii’ from Isiteki.

During feeding/foraging, the bee holds onto the lower flower limbs and very briefly inserts its ‘tongue’ into the flower and buzzes away. I could not see any deliberate hovering and pollen transfer to the back legs as does the Honey Bee. But pollen is collected and carried in approximately the same way. When I hand-pollinate I adopt the simple approach of approximating the insertion of hair (bees ‘tongue’) into one flower to obtain pollen and then into another to deposit it. Better results are claimed for physically exposing the stigma and transferring pollen using a brush.

A recent trip to photograph flowers resulted in us finding H. floribunda southeast of the Bontebok Park at Swellendam. I have reported elsewhere that I had seen this species within the park but repeated visits had turned up nothing but similar looking H. mirabilis.  On this occasion we were outside the park where we have recorded H. mutica, H. marginata, H. minima, and also similar small forms of H. mirabilis as occur inside the park. We explored a small area we had not covered before and were delighted to find H. floribunda, now easy to see because they were in flower. However, we proceeded to the H. mirabilis locality and on the way again found H. floribunda that we had missed on the previous visit. We also found more H. mirabilis about 60m further along and still about 100m away from the original H. mirabilis locality. These four species were along a stretch of about 5-600m and not occupying shared habitat. The vegetation was fynbos on shallow alluvium, quite stony and well-drained.

It was while photographing the flowers of H. floribunda that a furry fly appeared and ignoring us and the camera, attended to the flowers. While I was trying to get a picture of this fairly fast moving fly, the Anthophorid bee pollinator also appeared but flyng too fast and haphazardly to be pictured. We had seen the same fly on H. mutica at the Buffeljags habitat. The species is Australoechis hirtus known to be a pollinator and visitor to many flower kinds. The flies are nectar feeders and parasitize other insects in their reproductive process.

While we do not know what the pollinating effectiveness of these insects over distance is, it is quite obvious that in this situation there must be substantial transfer of pollen across all the species given the short distances involved. Three of the species were in flower. But H. marginata, curiously, at this locality flowers nearly 5 months earlier than at other known places. Despite that, there are hybrids with H. minima. There did not seem to be hybrids among the H. floribunda plants, but we did think that some of the plants of H. mirabilis could have been hybrid. ♦

Haworthia limifolia – a conundrum or a lesson?

I was contemplating the problem I see in the way we approach classification, contending that there is no resolution to the endless addition of new names and arguments about their validity and usefulness unless we reach some agreement on method and purpose. The confusion about names, descriptions and identifications that had arisen in 1947 came about for four reasons. Small samples and inadequate exploration, too many unqualified experts, lack of insight into what species actually may be, poor methodology.

There is no excuse for this situation to continue. Sampling is at a very high level and because the other three elements have not been removed, confusion is as great as before. Many names may have been lost from sight, but there is no lack of new ones to replace them. Experts are a problem and their qualifications even more so. In my own experience I have communicated with many highly qualified botanists who have contributed absolutely nothing to the resolution of problems as basic as typification of names, or definition of the word ‘species’. So what then contributes to qualification? This is surely an unbridgeable problem when amateurs have dabbled so freely in plant classification with apparent success. But their success has come from the failure of qualified botanists to put something more substantial in place than a nomenclatural code for binomials and nothing to indicate what those should mean, or who is competent to generate or change them.

Neither of those problems can be solved by me. My contribution has been to see as much as I can of what others have seen, and then to myself explore in both the arena of the subject matter and the periphery of biology that touches it. What I have done is to try and relate what I have read and been taught to what I have observed and experienced. In this way “species” have become a reality as systems of populations and individuals that are spread in space and change with time. Whether evolution is a fact or not I cannot say, but change certainly is. Classification since Linnaeus, has always been associated with this change but not necessarily by everyone. But in the time of Linnaeus, the sampling level was very low and anything new could be, or was, interpreted as a new species and diagnostics were put in place to aid identification. Hence even the Latin binomial was a diagnostic. As the sampling level has been raised so has the system come under increasing stress and it is a fallacy to think that only Haworthia is problematic. However, this is the genus under discussion here and we have to admit that the diagnostics and the methodology do not work.

H. limifolia can be used as an example to illustrate the situation.

It is to all intents isolated from the rest of the genus. We do have H. koelmanniorum sitting to the far north-west and then H. tessellata further to the south-west. Otherwise the rest of the genus is in the Cape. The issue of subgenus need not be seen as a consideration here even though the differences there are as great as any elsewhere in the family.

H. limifolia has a very wide distribution from Komatipoort in the north, westwards and southwards to Vryheid, eastwards and southwards to Gollel and then Stanger in KwazuluNatal. The whole of Swaziland and a large part of northern KwazuluNatal may harbor this species. We need to even ask … “Is it a species?”

The fact is that there are very different elements involved in respect of individual appearance and virtually all aspects of that individuality. The possibility is that whole populations may be derived from vegetatively propagating individual clones. Some plants are solitary and propagate with difficulty. Others proliferate as vegetative offsets or by stolons. The leaf shape and surfaces are highly variable. The surfaces may be flat or channeled; absolutely smooth, to having raised wave-like wrinkles, to tuberculate with small individual tubercles, or the tubercles may be fused in groups and ridges. These may be normal leaf-coloured or they may be white. Flowers are extremely variable and so are the seeds and the capsules. Flowering time seems to be opportunistic and in cultivation the plants may flower at any time, usually synchronizing during the summer months.

Do we really have any alternative but to see this as one single very large operating systems, despite all the real problems of isolation by distance and hence the associated problem of gene transfer between individuals and populations? Is it sensible or even practical to insist or suggest that there must be Latin binomials that describe (in the early Linnaean sense) the situation or, more importantly, that there are diagnostic characters by which such single taxa (groups, species, varieties, forms etc) can be identified?

How can the situation we observe in this relatively isolated system and (species?) be extrapolated to the Cape where all three subgenera and the bulk of the species occur?

Recently two authors have proclaimed the value of floral morphology in resolving this ongoing disputation about the species. What their actual view of what species are has not been spelled out but it appears to be a belief that simple structural (morphological) differences are what define them. Behind this must lie the unconscious acceptance of the original definition of species that they were systems of interbreeding or potentially interbreeding organisms. Thus flower structure is assumed to be linked to the breeding system and therefore very important in classification. This view of species still seems to be the power behind classification although it has long been recognized that in plants inter-fertility may extend across generic difference.

In Haworthia the flowers, even across the sub-genera, are fairly similar and especially in regard to size. Within the sub-genera, the similarities between what are considered to be species, is remarkable. There are even exact similarities that can be found in what are truly geographically and vegetative different populations. Flowering time seems to suggest some kind of additional factor but even here it is evident that breeding can occur between populations with flowering times 6 months apart. It has to be considered that flowering time differences like this might even be a characteristic of a species. This has in fact been shown for H. marginata and H. minima that surely can otherwise seen to be different systems.

The pollinating agents are insects and it seems as if flower size may be linked to that fact as it is so consistent across the genera and whatever are thought to be the species. Here I just present a number of pictures of the profiles and faces of flowers from 5 populations H. limifolia of which some are from different clones in those populations. A difficulty in viewing flowers is that they change with age, and place on the peduncle. They only last about three days and it is thus difficult to compare flowers of exactly the same age and condition. Of course these are poor pictures but no essential and detail is lost here by poor focus. But it is quite obvious that there is bigger difference between the two flowers (profiles) of the plants from Ithala than there is between them and other populations. Consider too that it can be very difficult to distinguish any of these flowers from those of other species in the subgenus Hexangulares. This essay is not submitted as conclusive evidence and argument and I will enlarge on this in a following essay. ♦

Volume 6, Introduction

It is a matter of considerable concern to me that the lack of “adequatio” for the understanding of Haworthia is so conspicuous by its absence. Three reviews appeared for 5 small essays published as a supplement to Haworthiad and also as a portion of Haworthia Update Vol. 5. To suggest that these reviews are anything but some sort of myopia is an understatement and I find it difficult to try and explain why. My reason is simply that many criticisms simply arise out of the fact that the reviewers are not familiar with my writing and have no basis for assuming that they are up to the task of reviewing what I have written. Nothing is being learnt from history where the same old arguments about opinions hardly differ from those voiced 60 years ago.

The real issue is that there is confusion in the minds of collectors as the user public, about the uses and application of Latin names in the pursuit of their interests. Professional botany offers no assistance and any numbers of poorly equipped authors of necessity, invent and re-invent types, the interpretation of names and then their application.  This does nothing to assist the collector. The reviews of that small segment of my writing do not help either and certainly do not generate a climate that encourages me to write any further. One review calls for a revision when the message is that a formal revision might just be intellectual vanity when a practical list of names and explanation is presented to add to an existing Revision. A second review seems to tout a mistaken belief that self fertility and polyploidy may be explanations for the intractable problems that we have in the self-sterile non-polyploid haworthias. A third review is seemingly facetious comment that may obscure my opinions as those of the reviewer and acceptable as such, but not if seen to be mine. None of these reviews will be helpful to anyone who may need to decide what set of names makes more sense than another. There is no common voice and any reader is left to make up his own mind in the absence of any good authority. Perhaps this is something I should be personally grateful for because there are noises emanating from various sources that my opinions also threaten understanding rather than enhancing it. I have to accept that it may be difficult for anyone to agree with me who has not had the same exposure to the intractability of biological systems that seems to have been my lot in life.

I regret that I did not, prior to publication, see the forward to Vol. 2 that I hoped would generate “common voice”. It, unfortunately, is just a bit of personalized chitchat about the writer.  It can hardly generate any confidence in what he may have written about and does not attempt to do so.

My revision in 1996 (published 1999) admits the problems of typification and my attempt to stay within the historically recognized interpretations and applications of names. A revision is fundamentally a reconciliation of new information and new “collections” and the Update Volumes have been an ongoing reporting of new finds and observations. More than one reviewer seems to think that a change of mind is indicative of weak argument rather than a product of new and better information.

Update Vol. 1 (2001), published by Umdaus Press includes one essay regarding Haworthia mucronata and five others dealing almost wholly with Haworthia cooperi and its variants.

Subsequent volumes were published by Alsterworthia. Update Vol. 2 appeared in 2006 in two parts comprising a total of 17 chapters and a list of Bayer accessions. Update Vol. 3 was published in 2007, also in two parts and comprising 15 essays. Update Vol. 4 was also published in 2007 in one part comprising 10 essays of which two were intended to close the saga. However, a surge of inquiry and energy made me decide to have a last serious effort to try an answer some outstanding questions in my own mind. So Update Vol. 5 was published in 2009 and comprises 16 essays. Perhaps, if nothing else, these essays just demonstrate how much there is out there in the field that has escaped attention.

Update Vol. 6 is simply a small set of essays that address some loose ends of which there are still many. One essay in particular will remain unwritten because, while I have some information, it is not enough to base any conclusions on. This concerns the area between Oudtshoorn and Uniondale. It is also not the only gap in my experience and knowledge. The chapters in Update Vol. 2 should demonstrate just how intensively the field has to be explored to get to grips with the possible dynamics of the plants. This view should be re-enforced by the subsequent volumes. While I have tried to hold to a belief in “species” and names relevant to some sort of system, it seems to me that as a society we are denied the freedom to really know what the meaning and purpose of creation is. Science itself seems to insist that there is none and botanists largely view the observance of nomenclatural rules as the primary criterion in the application of names.

My experience and observation now suggest to me that creation is purposeful and meaningful. Life is manifested in various distinctive forms and these forms are manifested according to their DNA. This DNA may be the fundamental stuff of consciousness in living things too; flowing from and responsive to the energy fields of physical bodies such as the galaxy and the solar system. Life on earth is driven by catastrophic events of varying degree and varying intervals, so that species simply represent those sets of living things as they come, change and go with successive events. We do not only need to consider what species might be and mean, but also when. Who knows? ♦

Volume 6, Chapter 1:- Haworthia and Chameleons

There is a very curious parallel in the problems and aspects of classification and identification of haworthias and chameleons that I personally find a bit mystifying. Classification of succulent plants and especially Haworthia is often done outside of mainstream science, which means that it is not done by trained professional botanists, who can stand in the intellectual arena with academics and degreed intellectuals. It has been pointed out to me that this is the problem in Haworthia and also why professionals do not want to attempt to resolve the issue because the nomenclature has been so confused and complicated by all the bungling that has taken place. Having been on the fringes, and indeed even failed attempting to cross the bridge into academia, I find it very difficult indeed to reconcile my life experience in the amateur arena with what I encounter from the professional one. This is because I have observed an ever widening gap between the perceptions of the non-science individual compared to that of the professional. My interest in chameleons is simply another aspect of my interest and passion for living things and I am distressed that the knowledge and understanding of these fascinating animals, as with Haworthia, is so clouded by ignorance and confusion.

It is a bit problematic for me to explain why I think classification and plant names is such a big issue and why it is important to me, but fundamental to me is trying to understand and know something about plants and animals as an extension of my life experiences.

Recently a very welcome and valuable booklet, Chameleons of Southern Africa has been authored by Krystal Tolley and Marius Burger and published by Struik Publishers (Cape Town). What makes it so interesting for me beyond my interest in the animals themselves, is the remarks the authors make in the preface in respect of writing the species accounts. The problem of finding identifying characters was a key issue and there is constant reference to confusion. There is of course no statement about what a chameleon species actually is and there is this curious statement, “There are few consistencies of identification characteristics, and we admit that we sometimes initially identify a chameleon by means of ‘gut feeling’.” What makes this so curious is that there is nothing in the booklet to indicate there is any more to the mystery of identification and how a final identification could be made to possibly help the even more than average individual to arrive at own conclusion.

In Haworthia (as indeed is the case in many plant genera) the description of species has been based on a basis of simple visual (morphological) difference, with barely any consideration of variation and in the total absence of a definition of a species beyond the assumption that if they look different they are different (species).

One has to ask what is it that is referred to as ‘gut-feeling’? I have often heard and read that classification and taxonomic statement is a matter of opinion and felt compelled to respond by quoting a writer who made the distinction between mere opinion and informed opinion. “Gut-feel” may vary from the instant response of the ignorant to a fleeting image, to the deeply considered intuitively driven reservation of another who is highly informed and who has a vast bank of experience, images and literature to draw from.

The significant thing is that any decision-making in the chameleon booklet has been facilitated by results from DNA analysis – *please see the footnote. Now I must make it clear that I am not attacking the book or the authors. I want to draw attention to the problems inherent in a methodology which amounts to nothing more than saying “trust me”, and the very serious problem that own judgment is by implication quite useless. As a very knowledgeable observer wrote to me “Our own eyes and brains are seemingly no longer good enough to serve the world – well that is what the RNA-DNA lab technicians are trying to tell us.” There have been several attempts to evaluate the classification of Haworthia by DNA sequencing that do little more than ring alarm bells. Without any pretence at understanding all the convolutions of genetics and the repository of DNA in all its forms, I would have to ask again, what does one do to organize one’s scepticism about the results that are thrust upon one?  If the total facies that an individual organism presents to the eye and senses is not a reasonable representation of the information encoded in the DNA, then we are absolutely helpless.  Could one say that we have been put into this creation without the sensory capabilities to distinguish and discriminate anything?

Recently I submitted photographs of two chameleons to an authority for identification stating that I thought they were not the Karoo Dwarf. The reply I received was “It looks like the Karoo Dwarf. What makes you think it is not?” Apart from the fact that I had already given reasons, the truth is that my correspondent could not possibly have said why he thought it did look like the Karoo Dwarf, thereby shifting the responsibility back on to me.  Are we stuck with the sad and unavoidable fact that unless one can get a DNA result, one cannot know what you are looking at?

What has now happened with respect to Haworthia DNA where we have the same dreadful problem of identification and understanding? I have elsewhere (see Haworthia Update 3 (Ch 3: p92) discussed the papers by Messrs. Treutlein et al  and I summarily dismiss what I think may be unpublished results by another source I must politely keep to myself. There was a rather immodest statement by Messrs. Wallace and Noll published in Haworthiad that promised to unfold the problems of Haworthia classification by DNA analysis. The results have never been disclosed and there is a disconcerting failure by the authors to respond to any inquiry. More recently Dr. Syd Rhamdhani has also been busy sequencing the DNA of a range of Haworthia and species of related genera. The work was initiated in an attempt to relate variability to habitat diversity, but to preclude a total negative result it was deemed advisable to widen the scope to ensure that the methodology was applicable at a broader scale. The results demonstrated, at least from my personal perspective, a complete failure to relate the DNA sequences with the realities of the classification of at least the semi-rational mind of an experienced and informed observer.

Group1A

Group 1Aa

CodeCollector NumberTaxon nameSubgenusLocalitySource
54MBB 7128angustifolia var. baylissiiHaworthiaWellsgate, E. of KirkwoodField
47MBB 6843marumiana var. reddiiHaworthiaInverbolo, StutterheimSheilam
58MBB sntruncata v. maughaniiHaworthia Sheilam

Group 1Ab

CodeCollector NumberTaxon nameSubgenusLocalitySource
35JDV 90/57monticola var. asemaHaworthiaS. of Kruis RiverField
40JDV 97/128mucronata var. mucronataHaworthiaSE of BarrydaleSheilam
6SR&TDsn 4angustifoliaHaworthiaMountain Drive (Gtown)Sheilam

Group 1Ac

Code Collector NumberTaxon nameSubgenusLocalitySource
32EvJ 17548cymbiformis var. transiensHaworthiaHoreeField
52MBB 7032mutica var. muticaHaworthiaRietfontein Sheilam

Group1B

Group 1Ba

CodeCollector NumberTaxon nameSubgenusLocalitySource
30PVB 7047zantnerianaHaworthiaN. of Redcliffe, WillowmoreSheilam
62MBB snreticulataHaworthiaBosfonteinSheilam
48MBB 6875maraisiiHaworthiaCogmanskloof, AshtonShadowlands
51MBB 6985rossouwii var. calcareaHaworthiaDe Hoop 

Group 1Bb

CodeCollector NumberTaxon nameSubgenusLocalitySource
33GM 623marxiiHaworthiaRooi Nek Pass S. of Laingsberg 
36JDV 91/91cymbiformis var. setuliferaHaworthiaRainbow ValleySheilam
4SR & TDsn 2cooperi var. gracilisHaworthiaMayfair (Gtown)Field
18SR & BBsn Site 5- 27/6cooperi var. viridisHaworthiaKaboega, Dassiekop NE of farmhouse 
15SR & BBsn Site 3- 26/6cymbiformis var. obtusaHaworthiaGlen Avon fallsField
16SR & BBsn Site 2- 27/6cooperi var. ‘puberula’HaworthiaKaboega, W of WilgefonteinField
1SR&TD 719cymbiformis var. incurvulaHaworthiaPlatosvaleField
21SR & BBsn Site 3- 18/10cooperi var. viridisHaworthiaKaboega, De PlaatField
24SR & BBsn Site 1- 19/10cooperi var. spec.1HaworthiaKaboega, KaboegapoortField
23SR & BBsn Site 1- 19/10cooperi var. spec.3HaworthiaKaboega, KaboegapoortField
25SR & BBsn Site 1- 19/10cooperi spec. 4HaworthiaKaboega, KaboegapoortField
22SR & BBsn Site 4- 18/10cooperi var. viridis

Grassland form

HaworthiaKaboega, W of WilgefonteinField
38JDV 93/45cymbiformis var. obtusaHaworthiaKagasmondSheilam
27SR & BBsn Site 1- 19/10cooperi spec.2HaworthiaKaboega, KaboegapoortField
39JDV 94/98decipens var. xiphiophyllaHaworthiaW. of CoegaSheilam
64BBsnaristataHaworthiaE of Hopewell farmField

Group 1Bc

CodeCollector NumberTaxon nameSubgenusLocalitySource
13SR & BBsn Site 1- 18/10cooperi typicalHaworthiaKaboega, Gertpiets areaField
26SR & BBsn Site 2- 19/10cooperi var. viridisHaworthiaKaboega, back of the Dam (Spekboomberg)Field
44MBB 6791/4cooperi var. picturataHaworthiaAndrieskraalSheilam
3SR & TDsn 1cooperi var. piliferaHaworthiaMayfair (Gtown)Field

Group 1Bd

CodeCollector NumberTaxon nameSubgenusLocalitySource
63MBB snparksianaHaworthiaMossel baySheilam
50MBB 6983rossouwii var. rossouwiiHaworthiaSoutkloof, NapierSheilam
67MBB snpulchella var. pulchellaHaworthiaConstable StationSheilam
43MBB 6762puchella var. globiferaHaworthiaTouwsberg, Little KarooSheilam

Group2A

Group 2Aa

CodeCollector NumberTaxon nameSubgenusLocalitySource
46MBB 6829Poellnitzia rubifloraNARooikloof Shielam
71AJ290302Poellnitzia rubiflora NA Genbank

Group 2Ab

CodeCollector NumberTaxon nameSubgenusLocalitySource
73MBB snAloe bowiaeNA???Sheilam
9SR&TD S3-29/8 Aloe speciosaNAPlutosvaleField
10SR&TD S6-29/8 Aloe africanaNABothas Hill (Grahamstown)Field
11SR&TD S7-29/9 Aloe lineataNAStones Hill (Grahamstown)Field
72AJ290289Aloe veraNA Genbank

Group 2Ac

CodeCollector NumberTaxon nameSubgenusLocalitySource
7SR & TDsn 6attenuataHexang.PlutosvaleField
34JDV 85/145scabra var. morrisiaeHexang.SchoemanspoortSheilam
31EvJ 16840attenuata var. glabrataHexang.CollywobblesSheilam
56MBB 7179viscosaHexang.Constantia, W. of WillowmoreSheilam
65MBB snvenosa var. venosaHexang.Breede Riv., SwellendamSheilam
41JDV 1942attenuata var. attenuataHexang.Soutkloof, AddoSheilam
28Clark & Pienaar 120venosa var. tessellataHexang.Farm Hughdale 161, Kikvorsberge (Noupoort Region, N. Cape)  

Group 2Ad

CodeCollector NumberTaxon nameSubgenusLocalitySource
66MBB snreinwardtiiHexang.WesleySheilam
68AJ290299kingianaRobusti. Genbank
5SR & TDsn 3coarctataHexang.Ecca Reserve (Gtown) Field
42MBB 6380pungensHexang.Braam RiverSheilam
17SR & BBsn Site 3- 27/6sordidaHexang.Kaboega, E of OlifantskopField

Group 2Ae

CodeCollector NumberTaxon nameSubgenusLocalitySource
19SR & BBsn Site 2- 28/6nigraHexang.DePlaat, KaboegaField
61MBB snvenosa var. woolleyiHexang.SpringbokvlakteSheilam
74AJ290298Gasteria liliputianaHexang. Genbank
45MBB 6809fasciataHexang.GamtoosSelecta Succ.
57MBB 7695glaucaHexang.15 km S of Kaboega Farm house, Darlington Dam to KirkwoodField
20SR & BBsn Site 1- 27/6glaucaHexang.Kaboega, W of WilgefonteinField

Group 2Af

CodeCollector NumberTaxon nameSubgenusLocalitySource
59MBBsnglaucaHexang.???? 
12SR sn 30/6/08glaucaHexang.Gertpiet se Kraal Field

Group 2Ag

CodeCollector NumberTaxon nameSubgenusLocalitySource
60MBB snHybrid

arachnoidea X blackburniae v. graminifolia

HaworthiaSchoemanspoortSheilam
55MBB 7145limifolia var. glaucophyllaHexang.Three Sisters, BarbertonSheilam

Group2B

Group 2Ba

CodeCollector NumberTaxon nameSubgenusLocalitySource
2SR843Astroloba foliolosa Kaboega, De Plaat Field
69MBB 6756Astroloba bullata ??Sheilam
49MBB 6952marginataRobusti.N. of AshtonSheilam
70AJ290292Astroloba foliolosa  Genbank

Group 2Bb

CodeCollector NumberTaxon nameSubgenusLocalitySource
53MBB 7096pumilaRobusti.Konings River Sheilam
37JDV 92/51blackburniaeHaworthiaVoorbaatSheilam
29EA 1211kingianaRobusti.Great BrakSelecta Succ.

Group 2Bc

CodeCollector NumberTaxon nameSubgenusLocalitySource
8SR&TD S2-29/8 Aloe tenuior Committees Drift JunctionField

Dr. Rhamdani has so far generated results from DNA sequencing using the nuclear ITS and chloroplast trnL-F markers. The former, on the evidence of the “species” considered, seems to give a good view of the generic possibilities, with not enough information with respect to the species. The trnL-F spacer results contain several spurious associations if my existing classification (Bayer 1999) and geographic distance are valid considerations. A poster presentation of the results has been published in Alsterworthia International 9(3)13-17.) Dr. Rhamdani has been kind enough to send me the product of an alternative manipulation of his results  that I tend to equate with an attempt to display the relationships of samples as they would be in geographic space. One can see from the groupings (see tables below) that there are gross anomalies that cannot possibly be taken to have classification truth. Group 1Aa for example has H. angustifolia, H. marumiana ‘reddii’ and H. truncata ‘maughanii’ nested together. Group 1Bd includes H. parksiana and H. rossouwii. The presence of H. kingiana in two different groups viz 2Ad and 2Bb has to be dismissed as an identification error where I would stake my reputation on EA1211 rather than AJ290298. Each group can be tediously discussed, but one needs to consider that the actual focus of the exercise was to explore the geographic relationship of DNA sequences below species level. Far too much was expected of a still poorly understood technology.

Coming back to chameleons, I have the problem that, being interested in the creatures, I want to know what they are. Having familiarized ourselves (my wife and I) with the Cape Dwarf (Bradypodium pumilum) we came into contact with the (then) Robertson Dwarf (B. gutturale now Little Karoo dwarf). The distribution given by W R. Branch and the descriptive detail did not seem to match what we were experiencing in the field and we concluded that the Robertson Dwarf and the Karoo Dwarf (B. karroicum) were surely the same species.  We then encountered what we thought was the Western Dwarf (B. occidentale) and were derailed by the opinion given us from a DNA analysis of the “population” observed, that it was a variant of the Cape Dwarf viz. a “renosterveld form”.  What we were told did not satisfy our “gut-feel” however profound and true it may have been or is. Then the booklet was published that confirmed our conviction that the Robertson and Karoo Dwarves were being confused (The true Karoo dwarf is now said to equate the E cape dwarf B. ventrale!), but not throwing any more light on the question of the “Renosterveld form” of the Cape Dwarf.

We then learned of a chameleon in the Gouritz River valley. On seeing these we concluded that on morphological grounds they were not the Little Karoo Dwarf and had to fall within the ambit of the Swartberg and Eastern Cape Dwarves as accepted by Tolley and Burger. These are the chameleons “identified” by an authority as the Karoo Dwarf despite our submission of evidence why it was not. This led us back to the published DNA work and a paper by Tolley et al that discussed specifically the Eastern Cape Dwarves and a number of variants suggested being new species. My experience in Haworthia generated the strong “gut-feel” that this may be quite wrong as wide field experience suggests that species are highly complex systems that defy the conventional taxonomic approach. Further sampling would quite probably generate a host of still other variants and require explanation. It is quite obvious that the chameleons cannot be identified with certainty using morphological characters and this is equally true for Haworthia. My experience is that it is true for many plant genera. Nevertheless, I think that this is because morphological criteria are so heavily relied on with not enough reference to geographic distribution and a rational species definition, as opposed to an open-ended species concept. In another paper by Tolley et al, the two species B. melanocephalum and B. thamnobates are shown to be genetically identical, despite the fact that they are known to be morphologically different and geographically separated. Thus it was a pleasant surprise to find another article (Miller & Alexander (2009) that probes this question and speculates that it is possible that the DNA sequences used in the analysis may not be representative of the full genome. The available results of DNA sequencing for Haworthia suggest to me that those sequences are also not adequately or accurately representing a true or reasonable phylogeny. A problem I then also have is that a phylogram is a simple two-dimensional tree that is being drawn to illustrate the relationship of elements that are spread in at least two-dimensional space derived from change with time. In my opinion far too much is thus inferred from complex statistical procedures that generate a wholly theoretical diagram wholly beyond penetration by ordinary skepticism. I would suggest that the so-named bootstrap and posterior probability values may be subject to the same apparently many vagaries of multivariate statistical procedures that are not perhaps fully understood or even recognized by those who use them.

Dr. Rhamdhani suggested that I mention some of the limitations of the sequencing process that he lists:

1. mostly extant samples are used as these are only available for analysis (except in some cases fossil can be sequenced).

2. small sample size.

3. use of a few selective markers.

4. hybridization and incomplete lineage sorting distorts and complicates patterns.

5. appropriate analytical tools have not been formulated to handle the data and the above issue.

6. manipulation, misuse etc. of species concepts to justify desired results – this can be related to the “gut-feel” issue that this article raises.

I did complain about the problem of probing results published in complex technical papers when the results conflict with my “gut-feel”. The response was that “our eyes and our intuition are frequently tricked by the complexity of the real world…”  This is flatly not true for Haworthia and I strongly suspect that it is not true for chameleons either.  In Haworthia we are tricked by a very simplistic approach and the failure of taxonomic botany to serve interested parties. My kind authority I referred chameleons to also wrote “I know the genetic information may seem very confusing, but it is essential to remember that the genes tell us about genealogy and not morphological or behavioural similarity.” I have said before that the traditional approach of  taxonomy was always to relate morphology, and whatever other information one could acquire, to phylogeny. It is an unlucky fabrication foisted on us by DNA sequencing that this is a new direction for classification. To totally believe that, say, less than a thousand base pairs on a tiny section of mitochondrial DNA is giving what must be seen as a final answer that defies the doubt of an observer is perhaps too much to ask.

An end point in my chameleon inquiry came when Dr. Tolley conceded (as most DNA experts now seem to be doing) “DNA is obviously not a final answer, and everyone that understands well how evolution works, already agrees on that. However, when used wisely, in connection with morphology (which is subject to phenotypic plasticity, strong selection, and local adaptation) it obviously can be useful in understanding the evolutionary history of organisms. We can use DNA to give us clues as to the evolutionary lineages, which sometimes can be the same as what we humans like to call “species”. For me, “species” is partly an artificial construct because evolutionary processes do not suddenly jump, and divide one thing into two species, there are gradual changes, and sometimes (more often than we care to admit), they do not fit in the box we have labelled “species”. She exposes the fundamental problem that no one probably understands properly how evolution works nor do any of us have the wisdom to use DNA information to best advantage.

References

Ashadee, K.M. & Alexander, G. L. 2009. Do dwarf chameleons (Bradypodium) show developmental plasticity.  African Zoology 44:45-54.

Tolley, K.A et al. 2004. Phylogenetics of the southern Africa dwarf chameleons, Bradypodium (Squamata: Chameleionidae).  Molecular Genetics and Evolution 30:354-365

Tolley, K.A. & Burger, M. 2007.  Chameleons of Southern Africa. Struik, Cape Town.

Treutlein, J. et al.  2003a. Phylogenetic relationships in Asphodelaceae (subfamily Alooideae) inferred from chloroplast DNA sequences (rbcL, matK) and from genomic fingerprinting (ISSR). Taxon 52:193 207.

Treutlein, J. et al. 2003b. Evidence for the polyphyly of Haworthia (Asphodelaceae Subfamily Alooideae: Asparagales) inferred from nucleotide sequences of rbcL, matK. ITS1 and genomic fingerprinting with ISSR-PCR. Plant Biology 5:513-521.

Note. The Chamelion DNA study rests on the sequencing of about 40 individual specimens as opposed to the 1000 plus used for the former morphological revision. ♦

Little Karoo Dwarf Chameleon.
Cape Dwarf Chameleon
Chameleon Gouritz – doubtful.
Western Dwarf Chameleon.

Volume 6, Chapter 2:- Towerlands. Haworthia retusa ‘turgida’

In Haworthia Update 4 I wrote an essay about the haworthias east of Albertinia in which I discussed their relation to H. retusa  and H. mirabilis, while generally lumping them largely in H. pygmaea. There are of course real ‘turgida’ populations as far east as near Mossel Bay, so I argued the case for an interplay of the two former species that over the whole distribution range generated two ‘species’ in the east viz. H. pygmaea and H. retusa (to include ‘turgida’), and three ‘species’ in the west adding H. mutica to H. mirabilis and H. retusa.

Recently I had the pleasure of meeting Gregory Nicholson who is studying botany at University, Cape Town. He surprised me by telling me that there was a Haworthia on his parent’s property west of Herbertsdale. It was not in fact so much surprising as confirmation of the belief I formed on a trip a short while before that there must be haworthias in the very suitable terrain of the Jakkals River valley 6km west of Herberstdale. The surprise came when Greg indicated the position of the plants much deeper into the mountains.

We visited the site while exploring H. emelyae north of the Langeberg Mountains and I will report that in an additional short article. We over-nighted east of Barrydale and a puncture made us detour via Riversdale rather than take the northern dirt road. It was raining heavily at Albertinia, but we could see sunshine on the Langeberg in the direction of Herbertsdale. We traveled the dirt road through the upper Gouritz, because we also wanted to gauge the condition of the gorge through the mountain, as Greg reports the presence of H. retusa ‘turgida’ some distance in. Towerlands is some 8km on and it was still just sunny when we arrived. We walked immediately the several 100m to Greg’s site along a good path among true Fynbos vegetation and found the plants very easily in a ferruginous sandstone ridge sloping into the river. Then the rain came and made picture taking a bit difficult (see MBB7489 figs. 1-16).

The plants are quite small and there was not much clustering. Unfortunately the weather restricted us so we could not look for better habitat around the corner where Greg had actually pointed. Greg’s photographs had made me lean to H. pygmaea and indeed I do not think one can deny the reality of this if one also considers the odd plants at Herbertsdale itself (JDV97/135) that I assign to H. pygmaea. However, the similarity to ‘turgida’ is just as striking and Daphne and I concluded that we could recognize similarities to ‘dekenahii’  and even ‘mutica’. To make it even more confusing, I felt, and still do, that the plants had similarities to H. emelyae. This latter was confirmed in my mind when we continued our trip via Cloetes Pass back to the Little Karoo to just west of the Gouritz gorge and a population of H. emelyae known to us from Gerhard Marx. I will submit that in a subsequent short note.

I include a habitat scene (fig. 1 – the arrow marks the locality) and note that looking south east one can see Albertinia and Aasvoelberg in the distance. We know that ‘turgida’ is present in the sandstones of the Langeberg, but this find suggests it is far more abundant and widespread than present records suggest. We need to now what the plants in the upper Gouritz look like.

Acknowledgements Gerhard Marx has been extremely helpful and informative and I am most grateful to Gregory Nicholson for his contribution. ♦

Volume 6, Chapter 3:- Still more about Haworthia on Kaboega

Kaboega is a set of farms on the northeast of the Zuurberg Mountain range, north of Kirkwood and off the Addo National Park. I wrote about the haworthias that occur there in Haworthia Update Vol.1. There is also an article in Aloe 40:10 (2003) in which there is a discussion of the variation of those haworthias as related to geology and topography. My wife and I frequently visit Kaboega to renew relationships with Ian and Sandy Ritchie who live there. Each time we go we try to explore some different area. We generally end-up with something that is notably new.

Part 1.

There is a real problem in trying to reconcile the populations we see with the names that are available and the way in which I have tried to formalize them myself. The problem is that Kaboega seems to occupy some sort of central and neutral position and it is by no means easy to arrive at any clean rational classification. Three of my species are involved, and I have to say they are “mine” because other authors are in strong disagreement. The three species I see are H. cymbiformis, H. cooperi, and H. aristata. It is firstly necessary to explain that I interpret the name H. aristata in Haworthia Revisited quite differently from what I might have done earlier; and quite differently from other authors who have simply taken the easy route and associated the name with Little Karoo elements for which I use the name H. mucronata. My interpretation of the name will be quite evident from my writings and from the pictures submitted with this article. The use of the name H. cymbiformis with respect to Kaboega is a major problem for someone like myself who is firmly convinced that geographical relationships are foremost in the recognition of species as living systems. On Kaboega, plants that look like H. cymbiformis seem to proceed out of a complex that is surely H. cooperi. If one properly considers all the populations that I ascribe to H. aristata one is seriously confronted with the reality that it is also a geographic variant of H. cooperi.

During June 2008 we were again at Kaboega and found two further populations of ostensible H. aristata. One of these viz. MBB7698 (see figs 1a-d) is only about 1km east of Buffelsnek on the western boundary where I recorded MBB7012. We did also see the same plants on a ridge still further east and north of Wilgerfontein where the very green and proliferous cymbiformis-like plants seem to reach their western limit. MBB7698 is effectively on the top of the Spekboomberg ridge, whereas the cymbiformoids are a little to the west and on the south slopes. It would be really interesting to know what occurs further westwards towards Darlington Dam.

We found H. aristata again practically central to the farm and about 1,5km south east of the Weir (MBB7703, see figs 2a-c, and these plants may have been illustrated before elsewhere incorrectly as MBB7697, a Gasteria). This is still about 1km north of the east-facing riverine cliff that houses the very green cymbiformoid plants (MBB7636, see figs 3a-c) of which other populations have already been reported to the east at De Plaat (3 populations) and also to the west at Koks Dam and Spekboomberg.

I have probably argued for recognition of H. aristata as separate from H. cooperi. Such an entity can probably be recognized as occupying the lowlands to the north of the Zuurberg. Both H. cooperi and H. bolusii var. pringlei occur to the north at Ripon and it appears that the latter may be represented by a population northeast of Stonefountain. There is a problem with proper, as opposed to the formal way either Breuer or myself may have done typification of the name H. gracilis, where we have both apparently just guessed that the name applies to the Helspoort variants of H. cooperi. In fact von Poellnitz’ name H. gracilis rest on the citation of a really mixed bag from not just Hellspoort, but from five quite disparate locations across almost the entire Cape. I use the name ‘gracilis’ only in reference to Hellspoort and it is evident to me that H. aristata is in the H. cooperi continuum. Look at the pictures and see what you think. I have added three pictures (MBB7701, see Figs 4a-c) taken of plants above Klipfontein at the eastern end of Kaboega that I refer to as H. cooperi ‘puberula’ and related to another record MBB6908 at Wilgerfontein.

Part 2.

Variants of H. cooperi on Kaboega

The various populations of H. cooperi are discussed or referred to in several of my essays published in the  Haworthia Updates and elsewhere. I have also been privileged to go back to Kaboega several times and further explore there. Each visit generates something new and different and evokes thoughts of what is still unseen, unreported and unknown.

On this occasion Daphne and I were actually hoping to find and photograph perhaps a chameleon, as workers had reported seeing one abut six months previously in the vicinity of the Klipfontein guest house on Kaboega. This is in the northeastern end of the set of farms that now constitute Kaboega. We thought we should explore the upper slopes where there was more sunlight and basking opportunity for the chameleons in the cold winter sun.

We climbed up the mountain via a very overgrown route that served as a passage for stock to access the higher mountain slopes, but also serve as a donkey pack route for the transport of supplies over the mountain in earlier days. This is of course peripheral to plant exploration. Nevertheless it is so fascinating to dwell on the fact that the enlarged game farm is actually comprised of about seven smaller farms that once were home to farmers and family. All abandoned as circumstances have changed over the years. It is as though man invaded this rich treasure house of natural beauty, sucked it dry and abandoned it again leaving nature to now heal the wounds. This is very unlikely because of the game fences and the absence of predators. Rainfall is marginal and the Kudu population is huge. Nature will have to respond in some dramatic and drastic way if she has anything in mind, and she is perfectly capable of that.

We moved east of our original visit to the mountain top where we had seen and photographed a few scattered plants before. We know that if we find one plant, to ask “Now where are they?” This confinement to such discrete, small habitats is so characteristic that we laugh off the notion that to explain the vicarious distributions over great distance, the plants were wider spread at some distant time in the past. Something else is at work and these populations maintain pattern and coherence beyond material means.

Eventually Daphne, whose eyes are now better than mine, did find the first plant and the body of the population we had only touched before. So these are the pictures. The first three pictures are of MBB6901 H. aristata (See figs 5a-c) on the lower country north of the mountain about 3km away on a neighboring farm Doornkloof, the next one would be a cultivated plant of MBB7701 H. cooperi ‘puberula’ nom. nud. (See  fig. 6) in sandstone and Fynbos vegetation in the mountain south of Klipfontein. The rest are H. cooperi (no appellation – MBB6940, See figs 7a-p) just east of my previously reported site north of Klipfontein, also in sandstone and transitional Fynbos. This term is used to simply imply one of these strange anomalies where karoid and Fynbos elements mingle. Many more pictures are really needed to indicate that each plant looks different and we spent a long and enjoyable time calling back and forth to come and observe one more gem.

Some of the clones would correspond to clones in the parent population of H. cooperi ‘viridis’ at Perdepoort about 20km west while others  to H. cooperi ‘gracilis’  from Helspoort north of Grahamstown, 50km east. Thus to my mind there is no satisfactory solution to the naming of these plants in our conventional understanding of Latin names.

The last photograph is a view across the Klipfontein valley and over the old greater farmyard to a mountain ridge and over that to the Zuurberg Mountain itself. Between is the Vyeboomfontein farm. The black arrows mark a point on the near ridge where there is a Bushman painting site and a large quartz rock that is home to plants like MBB7101. That mountain ridge extends westward 1-2km to the abandoned DePlaat farmhouse where there are again populations of H. cooperi on northwest and south aspects, at different altitudes and with their own variants. More conventional H. cooperi ‘pilifera’  can be found on the far mountain while Kaboegapoort is a gorge through the mountain with still another set of populations and variants.

It is all a wondrous never ending drama.

Part 3.

General comments on the plants on Kaboega.

The southern end of Africa is home to one of the six or seven plant kingdoms of the world. This is the so-called Cape Floral Kingdom. Botanists in the past seem to have misinterpreted this flora and not realized that it maintains its integrity for two reasons. The first is that it is on the table Mountain Sandstone formation and the second is that it is in a winter rainfall zone. Its origins are no doubt ancient, but in present day terms it is part of a present day winter rainfall area which includes the Succulent Karoo. Kaboega sits on the eastern verge of this biome and is therefore in a huge tension zone between inland, upland, summer and winter rainfall (and the stress of dryness) vegetation groupings.

Apart from annual rainfall stresses, broken topography, skeletal soils and an enormous range of exposed geological formations and rock series, contribute to an extraordinary range of varied habitats. Thus South Africa and especially the Cape, offer plant species huge opportunities to adapt and change according to ecological differences and stresses.  Kaboega is a wonderful example of the way in which plant species have adapted to these different habitats.

Kaboega has plant species from all of the biomes in the Cape, and the vegetation ranges from upland grassy fynbos, renosterveld, succulent karoo, valley bushveld and countless variation of these. The main rocks are quartzitic, dwyka tillite (of glacial origin) and shales. In addition there is a small amount of alluvial terrace.

What is so dramatic is that the different species occupy sometimes very limited habitats and despite this tight habitat choice (requirement) it is very difficult to specify what those requirements are for any group of species.

The genus Haworthia is represented by two of its subgenera. The Subg. Hexangulares is represented by three species:-

a. H. glauca, which is present mostly on the upper quartzitic ridges where Euphorbia polygona and the asclepiad Huernia brevirostris occur. This form of H. glauca includes some evidence of apparent interaction with H. coarctata which comes from further east.

b. H. sordida is an isolated population south of the Zuurberg and then westwards to Steytlerville. It was known on Kaboega from only one small hill where it grew under renoster bush. It was only while researching this visit that a second group was found.

c. H. nigra, which is very widespread in the greater karoo, but rare this far south and so far it is only known from one small population on the river terrace gravels.

The second subgenus Haworthia is only represented by many varieties of but one species viz. H. cooperi and this recognition of one species is only arrived at by discarding the traditional view of plant taxonomy that has no definite and decisive definition for the word species. The Kaboega plants are known from about 30 populations. These vary dramatically, but continuously, according to habitat and this has to be considered in terms of continuing similarities from Kaboega outwards into the Karoo, the Western Cape and the Eastern Cape. This suggests that there are very complex species systems and species cannot be as easily identified and described by visible characters, as a loose definition has in the past allowed. The Kaboega populations are very different from each other, and could be said to be from four or more species if fancy rules. In fact they are probably adapted by habitat and are evidence of the continuity of them all in one species complex.

It is population structure which is so significant and if one considers all the plant species that occur on Kaboega, one comes to realize that they occupy often unique habitats. There is one level of generalists such as Acacia karoo, Olea africana, Aloe ferox, etc which seem to be ubiquitous. But actually most of the species can be seen to have a “home”. There are about seven species of Aloe on Kaboega and seldom do more than two share habitat.  As one goes through a general list of names, one finds that one has to visit very specific places to find the plants. One can also find places like this one spot at Klipfontein that can be described as a “hotspot” for Kaboega. These are small species rich areas, which host species which do not occur generally and may in fact be very rare for Kaboega.  By also occurring on Kaboega like this, they represent a genetic entity and resource which may be quite unique

Acknowledgement.
I am most grateful to Ian and Sandra Ritchie for their generous hospitality and interest for the many times and hours we have spent fossicking in the wilderness of Kaboega. ♦

Volume 6, Chapter 4:- Haworthia emelyae and some of its variants

Where are we? Geography plays such a critical role in my perception of the species that it is important to try to understand why and how it touches on the issue. The Langeberg Mountains is a Table Mountain Sandstone about 10km wide and 1500m high running east-west and separating the inland Little Karoo from the Lowland Renosterveld of the southern Cape. There are five main travel routes through those mountains of which two are via river gorges viz. Cogmanskloof, Tradouw, and then there is also the Gouritz River Gorge where there is no road. Haworthia is not generally considered a sandstone and high mountain species because, firstly, there are few records to suggest that and, secondly, because they are averse to the higher moisture levels. However, there are many records in the low and close foothills. H. retusa ‘turgida’ is recorded in the higher areas and in the vegetation associated with the sandstone viz. the Fynbos. Fig. 1 is a view taken looking eastwards from Kleindoorn (Kleindoornrivier). This is about 16km east of Barrydale that sits at the northern end of the Tradouw Pass. The next farm is Brandrivier (B) and beyond that is Springfontein. Muiskraal is marked with an “M” and this is at the northern end of Garcia Pass from Riversdale to Ladismith. The “O” marks Oskop which is beyond Zandkraal and about 15km beyond Muiskraal. Another 10km will take you past Waterval and bring you to Aasvoelvallei at the confluence of the Grootriver with the Gouritz. The last stretch is another 10km over the Cloete’s Pass to Herbertsdale from where one travels back westward 7km to Towerlands.

What is there? I speak of species complexes because the classification and naming of Haworthia seems to defy any agreement. I do not think the nomenclatural system allows it either. South of the Langeberg are the H. retusa and H. mirabilis complexes as well as H. minima and H. pumila, as main role players. North of the mountains is H. arachnoidea and H. mucronata. There are incursions and crossing of the mountains by three of these and my contention that they do not all do so is simply because we do not see beyond the narrow confines of our classification system. North of the mountain there is enough evidence to suggest that H. mucronata and H. arachnoidea is very probably the same one complex despite the few situations where they manifest in near proximity. South of the mountains H. retusa and H. mirabilis have an even more complex relationship. It is difficult to begin to realize that there is a very small step between their manifestations south of the mountain to an integration of them both in H. emelyae that occurs north. I have great sympathy for those who contest the issue. I hope that the figures I present here, as a very small set of images from what may be a very small sample of existing populations, will help bridge the difficulty.

My field experience, however much field experience is a measure of any persons competence to process the product, suggests that there is a predictable coherence in the variation that one observes as one moves from discrete population to discrete population. This progressive variation is also driven by surface geology and habitat. It becomes no problem at all but to realize that as another set of samples, the Aasvoelvallei H. emelyae (see MBB 7850 figs 2a-l) is nothing more than the Towerlands H. retusa ‘turgida’ (MBB7849 figs 3a-b) in another guise. It is the habitats that are so vastly different. Towerlands is south facing mountain slope in sandstone and Fynbos vegetation next to a perennial mountain stream. It was no chance that the rain that caught us there stood in such sharp contrast to the warm sunshine that greeted us an hour later at Aasvoelvallei and H. emelyae a mere 10km away as the crow could fly. That habitat is Bokkeveld shale with a heavy overlay of the very white quartz that veins this formation, and the vegetation is Karoid.

H. mirabilis ‘maraisii’ is present north of the mountains at Montagu and towards Montagu from there to Barrydale. It is also present at Barrydale itself. At Kleindoorn we have an enigma (yet another) where the plants (MBB7847 figs. 4a-s) are very similar to the Barrydale plants and they also seem to flower in summer. But these same plants can be likened to H. emelyae ‘major’ at Muiskraal that flowers in the spring as does H. emelyae (and H. retusa). Figs. 5a-c MBB7844 shows plants at Brandrivier. The dramatic thing is that these are recent photographs and very unlike the plants observed there by me 30 years ago and recognized as H. emelyae ‘multifolia’.

Figs 8 are of JDV97-44 (originally GDM429) plants from beyond Muiskraal (where both ‘multifolia’ and ‘major’ occur in disparate habitats) at Zandkraal, where there are three populations to which an appellation ‘H. breueri’ has been fabricated. I downsize that to H. emelyae ‘breueri’ to bring some perspective back to Haworthia nomenclature. I do not include field photographs for Oskop and Waterval, but add two from south of Vanwyksdorp (see figs 9a-c ADH2966) where H. emelyae originated according to the collector (this was a Mrs. le Roux and the plants were sent to Smith via Mrs. Emely Ferguson). The photographs were taken just south of Vanwyksdorp by Adam Harrower and I do hope that readers will realize from this article that it is not at all strange that the von Poellnitz published illustration and type of H. emelyae (figs 10a-b) are so often said to be of H. retusa ‘turgida’. Curiously, H. Fourcade illustrated a plant (his no.144, see fig.11) of H. retusa ‘turgida’ and another (his no. 82, see fig.13) of H. reticulata as H. emelyae. This says a lot for subsequent attempts to interpret and argue vehemently about von Poellnitz’ H. correcta  (see fig.12).

What really struck me on this recent excursion was just how much inviting territory is unexplored. In contrast to that is Kleindoorn where we photographed those plants actually found by Gerhard on a previous visit there with Bob Kent and Kobus Venter many years ago. The plants are on the south side of a sandstone ridge. We could not find them elsewhere on our recent visit and it seems that suitable habitat is just not there for them. There is a most remarkable divide of shale and sandstone on Kleindoorn and while there is/was a form of H. arachnoidea in the sandstone (repeated at Zandkraal), only H. arachnoideanigricans’ is in the shale. At Brandrivier, E. Springfontein and Muiskraal the ‘multifolia’ is in a sandstone pebble layer that overlays the shale in the manner of the Tertiary terraces south of the mountain. West of Springfontein the plants are again in shale, but without the white quartz of other karoid sites mentioned.

Acknowledgement
It is not possible to do this fieldwork without the goodwill of landowners that is threatened by a glut of collectors who often trespass. Therefore I particularly want to thank Bertus le Roux of Kleindoorn, George and Jaco Nel of Brandrivier, Louis Botha of Springfontein, Anton Bredell of Aasvoelvallei, and Gregory Nicholson of Towerlands for access to their property. Gerhard Marx was contributory as was Kobus Venter on a previous excursion. Adam Harrower kindly sent me pictures from his records. ♦

Volume 6, Chapter 5:- Haworthia emelyae ‘major’ and multifolia. New populations.

There used to be a regular bus service between Riversdale and Ladismith and J. Dekenah made use of this for his excursions into the Little Karoo to find plants for G.G. Smith. He thus discovered H. emelyae ‘major’ at the northern mouth of Garcia Pass. He also submitted a single specimen of a plant collected from the karoid veld a little further north. It is that record that suggested to me that ‘major’ was linked to H. emelyae further east and north as well as to ‘multifolia’ to the west. Etwin Aslander found what I regarded as the equivalent of ‘multifolia’ (see figs 2a-i) on a low plateau north of Garcia pass on the farm Muiskraal that is at the foot of the mountains north of Garcia Pass.

Johan Meyer of Barrydale once showed me a cultivated plant that he claimed he collected on the slopes to the north of Etwin’s locality. This was a robust and rather tuberculate plant that was very reminiscent of the forms of H. emelyae at Zandkraal about 12km east. In an earlier chapter I wrote more widely about the problem of land ownership and access and eventually felt that I should look into the situation at Garcia Pass in respect of conservation status of ‘major’, when an opportunity to break the impasse arose.

Fortunately I have quite a long association with Muiskaal and its owners, since I was at the Karoo garden and Mr. Henry Chamberlain, owner of Muiskraal, finally allowed me access.  We visited both Etwin Aslander’s population (MBB7858 see figs 2a-i) as well as locating a population on the northern slopes as indicated by Johan Meyer (MBB7859 see figs 3a-i).

1c. View looking west from north Muiskraal

In addition we also found a population immediately west of the farmstead (MBB7857 see figs. 4a-f). Particularly interesting is the dramatic switches in dominances of shrubby plant species that must be a factor of differences in the very skeletal soils as well as how plants compete with one another (allelopathy). The plants in all three populations are of course similar to the east of Springfontein plants in as much as one generates a mental common denominator for a set of variable plants. This similarity must surely be seen in the similar geologies of the three sites which are on tertiary depositional terraces. We also explored extensively along the foot of the mountain between Muiskraal and Zandkraal without any success.

1f. West Muiskraal

However, we also made contact with the owner of the upper areas around Garcia Pass itself and were able to explore both sides of the road at the ‘major’ locality (see MBB7861 see figs 5a-l) as well as the range eastwards.

We did find a new population some 3km east of the farm Onverwacht (see MBB7860 see figs. 6a-i) and Gerhard Marx reports seeing it immediately north of the same farm.  We did on a subsequent visit briefly get there, but did not find plants. Some of the plants were very ordinary and it is not surprising that J.R. Brown, in commenting on his collection, felt that he had two species viz. H. ‘major’ and H. maraisii as he might have named them. Definitely the plants I saw could have been mistaken for individuals in several populations in the sandstones south of MacGregor or even nearer Villiersdorp (Moddergat).  Unfortunately my camera failed me and I did not get many pictures, but one can see that the plants can have relatively spineless leaf surfaces.

There is something curious about the flowering times. The ‘major’ plants had evidence of recent flower stalks on a few plants while some plants were also beginning to flower. The lower populations at Muiskraal and Zandkraal were already shedding seed. I think it is significant to point out the geological features that I think are the main drivers of ‘difference’. Garcia Pass is a rift through the Langeberg Mountains that are uplifted sand stones. The northern foothills are an eroded remnant of tertiary deposits that overly Bokkeveld shale. The Zandkraal populations are associated with quartz outcrops or variants of such as are common throughout the Little Karoo. The vegetation differences between the sandstones, terrace gravels and shales are astounding and illustrated in the fact that botanists, following their original intention that a biome was a major association of plant species driven by massive and extensive landforms and climate, regard the Karoo vegetation and Fynbos vegetation as belonging to two different biomes. The terrace vegetation is primarily Renosterveld, but it has elements of what is now regarded as the Thicket biome. It would not be a serious overstatement of fact to say that no two plant species occur in each vegetation type. The farmer on Onverwacht (Fynbos) is a flower farmer picking from the field, while the present farmer on Muiskraal (generally Karoo) grows olives and grapes. Muiskraal used to be primarily a sheep farm and Onverwach barely.

Despite this observation on compositional differences of the vegetation, or because of it, my contention is that ‘major’ is a variant in the sandstones while ‘multifolia’ is an expression of the same species, H. emelyae, on the tertiary debris. They both go on to varied forms in quartzitic patches in the shales. In my opinion H. emelyae is in a continuum that is formed out of the H. retusa and H. mirabilis complexes south of the Langeberg Mountains, where broadly speaking H. emelyae moves into sandstone as ‘major’ and the other two species occupy that terrain as ‘turgida’.

Acknowledgement:
Sincere thanks to Mr. Henry Chamberlain of Muiskraal and Mr. Pierre Vermaak of Onverwacht. Gerhard Marx also motivated with interest and information. ♦

Volume 6, Chapter 6:- Die Nekkies and Biomes and Haworthia maculata

It is always assumed that botanists have a good grip of their subject, as one supposes for the scientists in other disciplines. Any science is the advance of knowledge by observation, hypothesis and testing by systematic enumeration and experiment. This is furthered by replication and review by other scientists. Botany is far behind the exact sciences, because of the very nature and complexity of living things and systems, and also behind because zoology where animal life is obviously more organized than is the case with plants and where more attention is focused.

It has always been recognized that South Africa has a unique flora and traditionally the difference from other floras was attributable to the extraordinary richness, diversity and endemism of the so-called Fynbos that came to be regarded as the Cape Flora. Just what the Cape Flora comprised was extraordinarily vague and the early limits were stated in terms of latitude and longitude. In the mid-20century it was recognized that South Africa should be divided into biomes, as major components of the vegetation, and these were defined as units with wide and expansive boundaries. So while the Fynbos biome was recognized among others, no connection seemed to exist between its boundaries and those of the major geological systems. The essential problem is that the boundaries of this biome are by no means wide and expansive, because one can literally step from Fynbos vegetation into Renosterveld, Sandveld and even Karoo. This is no more dramatically demonstrated than at Die Nekkies where it is obvious that the vegetation differences are primarily a result of geological substrate fortuitously linked to rainfall gradients determined by continental land-form, latitude and topography.

The significance of Die Nekkies is that it is formed from a narrow band of the Witteberg rock formation distorted in two transverse planes of the folded Cape Sandstone systems. Die Nekkies hills are directionally east to west and tilted down from the south. Northwards they are lost under the flood plain of the Breede River. Eastwards and southwards there is a wedge of Witteberg and Karoo formations that drives against the Cape Sandstones all the way south west to Villiersdorp. The consequence is a dramatic vegetation barrier related to the very different soil chemistry of Witteberg quartzites as opposed to Cape Sandstone, which is as dramatic where Bokkeveld Shale and Dwyka Tillite meet.

What is in effect the case, and apparently not really conceded by botanists, is that there are primarily two floras in a single winter rainfall biome. Some botanists insist that vegetation differences are simply fire driven. Here it is again difficult to separate the consequences of different fuel loads due to the substrate characteristics and the plants adapted to each substrate from the effects purely due to the fire, but it is nevertheless quite obvious that the major differences are substrate.

This is all a very broad view of a complex phenomenon that emerges when one begins to look at the individual species components of the vegetation. Vegetation classification is really based on dominant species and seldom has any attempt been made to understand species distributions that do not coincide with broad general pattern. When this has been attempt-ed it has been very seriously flawed by the problem of identification. This is a matter that botanists seriously underestimate and consistently fail to address.

It is simply assumed by general botanists that taxonomists in herbaria are competent and able, and that revisionary work and primary taxonomy have kept pace with technological progress and research in all other fields. This is not the case. Taxonomy has for many years been a forgotten science and less and less resources are allocated to either revisionary research or identification. It is simply not possible to collect, mount, store and record the variation of even a fragment of the species flora within any herbarium structure. A related fact is that there is no definition of what a species actually is. Very often it is just assumed that a definition is in place that is satisfied when some, usually morphological, criterion suggests that a specimen and sometimes more than one specimen does not agree with some ill-defined norm dependent on the taxonomist and the herbarium in which he/she is stationed.

It is self-evident that in the nature of evolutionary processes that are presumed to take place, that variation must be intrinsic to any species. This means that species need to be considered as systems that respond and change according to events that also change substrate and climate. It also means that species systems cannot be the same and a model (definition) needs to be in place that recognizes that each species (system) may be in an individual relation to time and space.

Two examples from Haworthia can be given for the significance of Die Nekkies in terms of species diversity and distribution and the difficulties that poor taxonomy introduces to obscure pattern. The first is for Haworthia maculata. Where there is already considerable doubt as to a proper classification of the Alooideae in the Asphodelaceae, Haworthia is persistently seen to be a single natural group. While there is no definition in place for the species category, the genus category is assumed to have some reality. It in fact does not. It is quite evident that some species do have very close affinities that would satisfy the perception that these form good genera, there are many species that elude such simple categorization. In Haworthia, as it is traditionally constituted, there are three distinct sets of species based on a range of characters both vegetative and floral. It is an extraordinary reflection on botanical classification that this has never been properly accepted. The genus Aloe is widely distributed in South Africa and more than 150 species are recognized.  However, only two species can be seen as strictly Cape Sandstone and hence Fynbos “biome” species, although several are closely associated. There is only one species of Haworthia subgenus Haworthia that can truly be said to be a Fynbos species and that is H. nortieri. This subgenus is poorly represented north and west of the Fynbos “biome”, but it is represented by a vast array of variants in the Eastern, Southern Cape and Little Karoo that are difficult to aggregate into species. The variants seem to be ecotypes and it is such ecotypes that do adapt to sandstone habitats. There are none such variants in the sandstone mountains south and west of Die Nekkies where only one “species” viz. H. maculata occurs in localized population at several points along the range. This “species’ is very closely related to H. herbacea that is very common on all the other skeletal soils to the east, both north and south, but very rarely penetrating sandstone habitats. Where H. maculata occurs on the eastern limits of Die Nekkies, there is a subtle change of rock and shallow soil to a more Karoid habitat, and the plants tend to be more clump forming and spinose, suggesting a shift to H. herbacea. This similarity shift is replicated in H. herbacea where it occurs east of the brickworks near the Brandvlei Dam wall. Ecotypes similar to, and classified as, H. maculata occur on the Witteberg formation hills southwards to Villiersdorp, but never in close association with H. herbacea. Curiously, a very similar variant to H. maculata occurs again high in the sandstones north of Worcester on the Audensberg peaks and also lower down in the Sandhill area. Thus H. maculata is a significant marker in the relation of the Asphodelaceae and Haworthia to the distinction of “Succulent biome” and “Fynbos” biome.

Haworthia subgenus Hexangulares is represented by only one species in the south western Cape. This is Haworthia venosa that occurs only in rock faces flanking the lower Breede River and Gouritz River. Its related species H. tessellata is a Karoid element. Haworthia subgenus Robustipedunculares is tightly allied to a winter rainfall “biome”, but consists of only four species. Three of them are endemic to the Southern Cape and three of these do not extend further west than Ashton and Bonnievale. H. pumila is abundant in the Worcester/Robertson Karoo and extends as far afield as Laingsburg, Anysberg and Barrydale, but south of the Langeberg it is not nearer to Swellendam than at Drew Station.  South of the Riviersonderend it is only at Stormsvlei, near Protem and at Vaandrigsdrift. It does not occur on the Witteberg formations south of Die Nekkies, but it does occur on the eastern lower slopes of Die Nekkies itself. So here again Die Nekkies is the evident limit of this species too – the second example.

Possibly the very dramatic case of Drosanthemum halli and D. bellum more clearly suggests the significance of local diversifying factors. D. halli is yellow flowered and abundant whereas D. bellum is pink and localized. They do not co-occur. It can be dramatically shown that D. hallii is nothing but a geographic variant of the bi-coloured D. micans that occurs north and west of Worcester, and that D. bellum can only be different because of very site specific factors.

H. maculata at the western end of Die Nekkies appears to be a solitary plant but, as it occurs more eastwards, it becomes proliferously clumping as the vegetation also becomes increasingly Karoid. Why this should be so is not clear.

The many accompanying illustrations (MBB7526 figs. 8-38) show the scenery and topography and the range of variation in H. maculata. One can see that it is at home in both the Restiods of the Fynbos and under Pteronia paniculata of the Karoid.

All the following photos are MBB 7526 Haworthia maculata. Die Nekkies, west end.

Acknowledgement
I would like to thank Anso Le Roux of Pokkraal, Rawsonville for drawing my attention to H. maculata much further west along Die Nekkies than previously recorded and for inspiring a walk along the full eastern limb. ♦

Volume 6, Chapter 7:- Taking Haworthia cooperi further – Kliprivier

It is very difficult to write about particular plants when one has to contend with the fact that one is not sure what Latin names may mean to the reader. I wrote a piece with closure in mind and thought I would send it to a non-taxonomist botanist whom I think is a tribute to the profession. Extracts from his response are “I am an ardent supporter of your species concept and couldn’t agree more strongly with your statement that without variation, there would be no evolution… I do agree that the pervasive species concepts force us to ignore the most interesting and productive research avenue: documenting and understanding variation in the field.”

It is worth considering what he has said about “pervasive species concepts” and just what he might mean when he says they force us to ignore documentation and understanding of variation in the field. It seems to me that the converse is the truth. Failure to properly understand and document variation has contributed to the pervasiveness of false concepts which fly in the face of science. I keep harping away at this question of lack of definition, because it is not apparent to me that any reader appreciates my point of view. To my mind a key issue is made of nomenclature and the rules that govern it, and very little attention at all is given to whether these Latin names help our understanding at all. I feel that my contribution contributes mostly to documentation of variation and that I cannot do much in respect of understanding. This short article should explain why.

I have tried to explain what I have observed in the three species Haworthia cooperi, H. cymbiformis and H. transiens as I understand the various populations that might be seen to constitute three discrete systems. My problem is that they do not necessarily constitute such discrete systems at all. Even considered together as one system, it is not entirely sure in my mind that they are truly separated from any other species belonging in the same sub-genus Haworthia.  It is not improbable that they diffuse into H. decipiens and into H. mucronata that are in turn diffused into other elements. If one considers that a species definition based on breeding true has been “… used by most taxonomists since 1682.”  (I quote this from a prominent taxonomist), one must recognize that taxonomists have then been ignoring the innate variability which drives change, adaptation and evolution. I am not sure that I have not been making the same mistake although I have consciously tried to recognize that discontinuities of any kind do not necessarily suggest discrete species.

Someone once wrote that “specialization leads to extinction”. If species become too specialized, they lose the capacity to adapt and evolve according to an ever-changing environment. At the other extreme, species that can maximize variation within their individuals and populations will benefit most in those changing circumstances. If taxonomy has been guilty of paying insufficient attention to variation as a concomitant of evolution in present time, it may mean that we may generally be recognizing species at the wrong level. We may collectively be attaching far too much attention to morphological and even genetic differences without due account of distribution and potential for change both internal to the species and external from the environment.

Therefore, I am deeply skeptical that my classification of Haworthia is a true reflection of the situation in respect of actual species. I have been influenced by a paradigm in taxonomy that genera and higher taxa were largely artificial and determined by arbitrary choices that they should be approximately equal in size and number. Without taking this too seriously, there is a huge problem when it comes down to species and to consider that some in a genus may be tightly grouped in terms of limited variation and distribution, while others form vast complexes.

Jan Vlok kindly sent me a picture of a Haworthia from the Little Langkloof. This is the upper reaches of the Keurboomriver Valley which runs inland from Plettenberg Bay, turning west north of the Knysna area and cutting across the Prince Alfred’s Pass road between Knysna and Uniondale. The only Haworthia species in the area are H. transiens and H. scabra and they are abundant there. Jan said the plant was in Karoid vegetation and was generally solitary. I suggested that it was probably an ecotype of H. transiens and that otherwise it may have been H. cooperi var. gordoniana. I have written an account of the interaction of those elements in the Baviaanskloof to the north and west, and it is quite evident to me that there is a close relationship. It should be noted that there is also a major variant of H. scabra closer to the Prince Alfred’s Pass road, so that there is a precedent in adaptive change. I was recently able to visit the area myself and confirm that the plants are in this very same category of interaction between H. cooperi and H. transiens. Thus, in my opinion, they parallel the kind of relationship other variants of H. cooperi has with H. cymbiformis, and in a changed dispensation for taxonomy it may be agreed to regard these two species and H. transiens as one species. It is possible that continuity may still be found with H. mucronata further west. While I write “changed dispensation”, I regard it as essential that it first be understood that there is still a vast amount of exploration and recording to do and that any final solution will only be achieved by collaborative effort and agreement. Professional taxonomists will have to make the decisions, but they will have to first prove themselves competent to do so.

Kliprivier plants in cultivation.

The above was the article as printed in Haworthiad. It deals with my collection MBB7586 (see figs. 1a-e) from east of Die Vlug in the Prince Alfred’s Pass south of Uniondale. My conclusion was that H. transiens was but another expression of H. cooperi at the western end of its distribution zone and possibly transition to H. mucronata.

I had collected seed that I asked Etwin Aslander to grow, which he did very successfully.  The seedlings were initially extraordinarily variable and there were some striking plants that gave indications of reticulate marking that would have shamed H. cooperipicturata’. After Etwin transplanted them the plants looked much more ordinary and I selected a few to grow myself. So this short note is just to present pictures of these and demonstrate the badly beaten horse of variability.

Figs. 1 are a series of pictures of plants in the field that can be said to be representative without leaving out too much. Figs. 2a  and b are of my permit limited collection grown in my outside rockery in relatively full sun. Figs 2c. to 2h are the selected clones that I grew on to show readers what happens between field, nursery and greenhouse table. Any wonder that you can justifiably complain when you do not get what you think you ordered?

I originally omitted names and would now use the name H. cooperitransiens’.

Figs. 1a-l. MBB 7586. Hawortha cooperi ‘transiens’. Kliprivier

Figures 2a and b are of my permit limited collection MBB 7586 of Haworthia cooperi ‘transiens’. Kliprivier grown in my outside rockery in relatively full sun.

Figs. 2c to 2h. Selected clones of plants grown from habitat seed in cultivation, grown on to show readers what happens between field, nursery and greenhouse table.

Acknowledgement
Jan Vlok very kindly informed me of the existence of this locality east of Die Vlug. ♦

Volume 6, Chapter 8:- A fleeting look at Haworthia arachnoidea

How did this start? Is it possible to see anything with such a quick peek? Somewhere in my memory bank is my stated opinion that understanding H. arachnoidea would assist any botanist towards a better understanding of classification. This was long before I came to see that this is a lot more necessary than I thought then. It is not just botanists who seem dumbed down to the reality of a diversity that is necessary for response to change and largely denied by the nomenclatural code and how it is practiced. We have not understood change or what changes can occur. During perhaps only the last 15 to 20 years has it become apparent that changes are cataclysmic and frequent. I will skip the fact that the violence of catastrophe, or the drivers of catastrophe, may also induce change at even a cellular level.

I usually go into the field with a specific goal while also carrying a host of peripheral questions in my mind. While exploring the problem of H. schoemanii, I was thus also thinking of Conophytum. After photographing a species seen at Laingsburg, I mentioned to Steven Hammer that I had also seen it in the southeastern Tanqua Karoo (Bakoven). This was news to him and so in my wish to re-establish the reality of H. venosa subsp. granulata recorded at Patasriver (actually Patatsriver road and that is another story that will be told) as well as the Conophytum for Steven, I undertook an expedition to the Tanqua.

It was co-incidence and luck that brought us into contact with Dr. De Ville Wickens, who is an expert on the geology of the Gondwanaland connection to the Tanqua mountains and who now owns Bizaansgat to the northwest of the farms Droogekloof and Bakoven. Dr. Wickens facilitated our expedition by offering us accommodation at Bizaansgat and we were thus able to spend more time there than otherwise.

Derek Tribble did alert me to a small Haworthia in the Skitterykloof about 20km to the west of Bizaansgat. I had myself seen the plant and concluded that it was H. arachnoidea. Fairly recently Adam Harrower gave me a small plant from the vicinity of Skitterykloof and I confess that I was a bit confounded because it did implicate H. nortieri (See ADH3140, figs 1 & 2) particularly because the leaves were so spotted. Why I say this is because the combined presences of the look-alikes H. arachnoidea and H. nortieri ‘Albispina’ at Koup and elsewhere, as well as their often superficial resemblances, now make me a little wary of snap decisions. I later concluded from the flower that Adam’s plant was in fact H. arachnoidea, and not because I am convinced I could separate the two on the basis of the flower only. So this report covertly deals with the reality of a massive variation within H. nortieri and also H. arachnoidea and hope that readers will generate their own open mindedness to realize that I am drawing conclusions here that could or should be better substantiated. I also hope that this does not mean that a whole new set of Latin names is required to try and do so.

Because my initial field work was to obtain an overview of the genus, I soon stopped paying much attention to H. arachnoidea, because it seemed to be so ubiquitous. A big mistake! Unfortunately digital cameras were then not available and space in the herbarium (contrary to what my learned friend has to say) was and still is at a premium. The consequence is that I do not have a proper physical or mental record of all the plants and populations that I might have seen. In fact I cannot even fully remember how or why I linked H. arachnoidea to H. decipiens the way I did in my Handbook.

What happened on this expedition is that at Droogekloof we were surprised to find a very small Haworthia that I take to be H. arachnoidea (see MBB7644a, figs 3a–j). It is important to note the differences in colour, spination and leaf-spotting and/or reticulation. These plants were single and scattered over a fairly wide area. They varied in size from as little as 20mm to an average of about 30mm diam and become extremely cryptic in the dry summer.

We continued to Bakoven and found it again (see MBB7644, figs 4a-e). Fig. 4c. is a view from a mountain top at Bakoven looking NNW towards Droogekloof and then Bizaansgat in the distance. Fig. 4b illustrates the weak inflourescence with few flowers generally true for many of the populations we observed.  We the retraced our steps and saw similar plants approximately midway back to the Droogekloof population (MBBsn see fig. 5). The next day at Bizaansgat we saw exactly the same thing at our first three stops and then came across an absolute field of them at a fourth. Again at several more stops on Bizaansgat (MBB7646 see figs 6a-h) and yet again on another farm 5km to the east.

The next week took us on the road, the Patatsriver road, from the southern Tanqua to Matjesfontein east of Laingsburg. We stopped on the advice of Gerhard Marx at a well known spot at Perdekraal where Lithops, Didymaotus and Tanquana occur. We saw the same small H. arachnoidea there (MBB7658 see figs 7a-c).

From there we turned north to Bantamsfontein for H. venosa subsp. granulata and we were surprised to find again the same H. arachnoidea at every place we stopped on the farm (MBB 7660 see figs 8a-e).

We travelled on in the direction of Matjiesfontein to Patatsriver and again found it ubiquitous (MBB7666 see fig  9).

9. MBB7666 H. arachnoidea Patatsrivier

We went on to Keurkloof just northwest of Matjesfontein, and found it on the two hillsides we searched (MBB7670 fig.10).

From there we went to Bulhouer, which is north Matjesfontein, and found it at four places (MBB7671 & 7672 see figs. 11a-d  and 12a-e).

It is very important to note that Emil Heunis recorded the form ‘scabrispina’ on this farm Bulhouer and this is what I was expecting to find. However, my four samples were observed at regular intervals of about 1km from south to north. The southern populations were indistinguishable from any we had seen from Perdekraal in the west, but the last population, (MBB7672 figs 12a-e), were as close to as must be possible. My contention is that we thus observed continuity from the west to the east.

On northwards to Dwarsindieweg some 15km further and we found the small forms (MBB7677 see figs 13a-b) en route as well as commonly at Josephkraal another 16km east (MBB7678 see fig. 14). We found it again the next day north of Dwarsindieweg (MBB7686 see figs 15a-d) on the high upper slopes of Klein Tafelkop (MBB7688 see 16 fig 54). Heading for home we stopped on a backroad on the way to the Patatsriver road at Witrantjies  and found it yet again (see figs 17a-c). We had been very cautious all the time to be sure we were dealing with one thing and not two. On this last stop we were confronted with the first drama when it seemed fairly certain that we had also found H. nortieri and this turned out to be false. We did return later to climb a high mountain at Volstruisfontein west of Dwarsindieweg hoping to find H. marumiana. Instead we again found small H. arachnoidea (MBB7802).

It was not flowering time. The pictures that should have accompanied this article will show that the arachnoidea we found were often still accompanied by an odd old inflorescence.  A weak spindly thing with evidence of seldom more than three old capsules.

Generally the plants were very small and we very often came across seemingly mature plants that were down to 20mm in diameter, and seldom exceeding 35mm. Invariably we could find the odd bigger plant in any single population array. The spination was fairly variable in respect of size, spacing and arrangement. The leaves could be short and squat or narrow and elongate. Colour was usually on the bright green side and I do not know what the summer colour would be. Leaf markings were interesting because often there was a slight reticulate pattern to the leaves and translucent dots and stripes and margins could be observed. This seemed to diminish in a cline from west to east. We did conclude, because this is what we also wanted to observe, that there was some evidence for continuity with H. arachnoidea var. scabrispina. There was the one population, however, north of Bulhouer where the plants were indeed bigger (50-60mm diam) and there was a definite tendency to form overarching brown spination so characteristic of scabrispina. There were of course the odd variants elsewhere and one group of four plants east of Dwarsindieweg could have come from the very confounded interface of H. arachnoidea and H. mucronata that is covered in the Latin H. arachnoidea var. nigricans. Readers should recognize that there is no way in which the nomenclatural code can be used to account for this diversity and variation, and it is time that more space and recognition is given to this very common phenomenon in the general context of writing, speech and communication.

The pictures will include one of ‘candida’, an epithet conveniently supplied by M. Hayashi for a variant of H. arachnoidea that occurs very close to Ladismith (KG16/72 see figs 18a-c). The spines in these plants often are black at the tips and the same phenomenon can be observed in massive plants that occur near Worcester in the “typical” variety arachnoidea. But I would like to break away from this usage. The epithet arachnoidea is applied to every single plant of that species wherever it occurs. Varietal names of any kind may be attached and I especially recommend the form and the formalization that Gordon Rowley advocates too; this is the format example…H. arachnoideascabrispina’.  For more precision a whole host of abbreviated prefixes could adorn the label e.g. cf ‘scabrispina’ meaning “compare with” or v ‘Scabrispina’ as a cultivar or var. scabrispina as a generalisation.

There are a set of varietal names in H. arachnoidea already, and the proper address for these small Tanqua plants discussed in this article is in terms of my own Revision H. arachnoidea v. namaquensis.  I do not think that my dispensation is adequate or good enough for the scale of our present knowledge, but neither do I think formalization is going to do much more than arouse an egocentric mass of still more Latin epithets. So I strongly suggest that we as a community simply accept something like H. arachnoidea ‘tanqua’  or ‘minuta’ and drop any pretension that Latinization adds any lustre or glamour to plants that are already fascinating and attractive. One of my objections to what I find a really irksome process of formalization (typification, full citation of basionym and all that stuff), almost senseless, because unless a microchip is also implanted it is unlikely that there will be any chance of associating any plant out of habitat back to its origins.

I am not going to write any further about H. nortieri cf ‘pehlemanniae’ here because I need to know more as well as confirm something about the flowers and flowering times of both sets discussed here. There were the occasional dark coloured plants in the arachnoidea (and excuse me dropping the ‘H’ – it really is not necessary in this informal environment), but I doubt very much that anything further will support the suspicion that arachnoidea will now always be confusable with nortieri. It turns out that what I though might be H. nortieri because of the very grey colour tone of the leaves was in fact also H. arachnoidea. A specimen flowering in cultivation produced non-globose flowers.

Acknowledgement:
Dr. De Ville Wickens, Bizaansgat;  Mr. P Schietekat, Stellenbosch; Mr. M. Jan duToit,  Remhoogte Boerdery;  Mr. Pieter de Graaf,  Patatsriver;  Mr. C.K. Francois, Bulhouer;  Mr. Wynand Theron, Dwarsindieweg; Mr. Johan Kriel, Citruspoort (Volstruisfontein); Messrs. Stephen and Gail Louw, Keurkloof.  Hospitality and access to properties is deeply appreciated and valued. ♦

Volume 6, Chapter 9:- Interesting nursery plants

I have become increasingly concerned about the poor relations that exist between collectors and the authority of Nature Conservation. The argument that collectors threaten and despoil natural populations is very real and I do not dispute at all that Conservation authorities have a very valid complaint. They have a function to perform. On the other hand there is an interaction between human beings and nature in all its forms that should be fostered to the benefit of both sides.

Nurseries, traders and collectors are as much of the picture as are conservationists, institutions, researchers and landowners. It is unfortunate that there is no non-government party that lobbies for the rights and activities of the former group, but it is not my intention nor within my competence to argue all the aspects of the case.

I strongly believe that people have the right of access to nature in all its forms and the issue is one of individual responsibility and proper consideration of consequences. An appreciation of and sensitivity to nature should be reflected in whatever we do in our lives. My own collecting impulses led me to institutional employment where I could exercise my interest to what I thought were efforts more worthy than my personal interests. From that position I also did try to share and extend privileges to a wider circle. It is  in this way that I became involved with Sheilam Nursery. It was not my wish or intention that my collection should have come to be housed there. However, Sheilam has succeeded over a period of nearly 40 years to maintain a fairly true record of my collections obtained as propagated material from the Karoo Garden at Worcester. My offer of permitted collections dating from my revision of Haworthia in 1966 to the Karoo Garden was rejected and for a while resided with Etwin Aslander at Brackenfell. It has since passed to Garth Schwegman at Sheilam who has taken a particular interest in the maintenance and propagation of that collection.

Hand pollination of haworthia has become the norm and Sheilam has made a massive effort to generate plants from this. Of course growing plants from seed is a lengthy process, but is richly rewarded with an astonishing array of variants and beautiful plants.

Need I say how important this activity can be for an understanding of the plants and the way this is expressed in a classification?  A good example is of the hybridization found between H. lockwoodii and H. arachnoideascabrispina’ at the Floriskraal Dam east of Laingsburg (see figs 1-3). The hybrid is very reminiscent of plants that I have seen from between there and Prince Albert; and also southeast of the Rooinek Pass through the Witteberg, and again northeast of Calitzdorp. Roy Mottram underscored the issue of hybridization in Haworthia classification. I have not ignored the issue, but rather viewed it as an imponderable in the continuum between populations that seems to have pattern.

Figures 4 and 5 are seedlings of a collection of mine from north east of Nieuwoudtville of H. nortieri. They could be related to ‘globosiflora’ and it will be interesting to see the flowers. However, it may be foolish to draw any conclusion from whatever flower shape emerges in the broader nature of the variation already observed although not adequately documented, both in H. nortieri and in other species complexes. Plants with these stubby deeply mottled leaves can be the consequence of growing conditions just as much as of genetic variation intrinsic to a ’species’.

Figs 6-12 are seedlings of another collection of mine (MBB7608) from Melkhoutrivier south of the lower Breede River and north of the Potberg Mountain. This collection is reported in Chapter 7 of Update 4, I think very aptly entitled “The brutality of reality in Haworthia”. It is one of 12-15 populations in the greater area that support my suggestion of H. mirabilis as including H. maraisii, H. magnifica and H. heidelbergensis. I suspect that H. mutica is also involved to some degree or other, while it is not difficult to extend comparisons to as far afield as Riversdale and further to H. emelyae in the Little Karoo.

Collectors will no doubt always have a problem with plants that do not match the perceptions they attach to names. Kobus Venter has always expressed anathema for anything but random selection when collecting specimen plants. This is in some degree correct because the specimens are supposed to represent a population. However, the opinion on a population is largely based on field observation. Subsequent cultivation and observation do not greatly influence that, while oddities and unusual forms may help mould opinion. While Nature Conservation may object to the introduction of permitted collections eventually into the collecting arena, I see no ethical grounds for this. I insist that it is an undeniable right that people have right of access to a natural resource with the proviso that there is no pressure on sustainability. ♦

Volume 6, Chapter 10:- Non-pilose "pilosa"

Map of the Potberg
Map of the Potberg

When I was at the Karoo Garden I became a bit befuddled by the way botanists referred to the Cape Floral Kingdom. It seemed to me that they used the term for the vegetation that was on the Table Mountain sandstones and conveniently excluded that which was not. Thus the “Fynbos” vegetation, characterized by its Ericaceae, Proteaceae and Restionacea, was synonymous with this floral kingdom. An official document was published at the time which purported to classify the Southern African vegetation into biomes as major floral assemblages with very broad boundaries. It did not make sense to me because my observations were that the “fynbos”, however different in terms of historical origin, was essentially a flora of the sandstones, and that there was rather a winter rainfall biome which included karoid (Succulent Karoo mainly) flora. The role of geological substrate and skeletal soils seemed to me to be pivotal as there are places where one can virtually take a single step from one vegetation assemblage into another.

My impression is that it is the dramatic range of geological formations and highly eroded and eroding (in geological time) surfaces that is responsible for the extraordinary diversity and variation of the Cape Flora. My experience was also that taxonomists were not really aware that this might be significant in respect of plant classification. Botanising in the north and east of the country is not quite the same as botanising in the Cape and certainly it appears to me that European botanists familiar with the plants of a non-eroding land mass do not, or did not, give enough attention to the relation of substrate to plant variation.  Unfortunately I am not a geologist, but I want nevertheless to illustrate a few plant populations which demonstrate the complexity of the problem where dramatic differences in soils over very small distances are reflected in variation in vegetation and in the plant species themselves.

In the Compton herbarium, there is a specimen of a Haworthia collected by Prof. Compton. It is labeled as from “Potberg sandy flats”. Now I digress because there is a problem of semantics. The word “retuse” is currently defined in botany and in the Collin’s Dictionary as “end rounded with a central depression”. In Smith’s “Dictionary of Plant Names”, the name “retusus” is said to mean “with a rounded slightly notched tip”. But this is not how it is understood nor used in Haworthia. Haworth, and other writers of the time, used the term(s) “retuso-deltoid” to describe leaves that are “bent back like the end of the thumb”. Thus “retuso-“ meaning “bent back” and “deltoid” referring to the triangular end-area. It definitely does not refer to the end notching of a flat leaf face. When we speak of the “Retuse” haworthias, we mean plants with bent back leaves and a flat triangular end-area. Prof. Compton’s specimen was of such plants where in the dried herbarium specimens the margins of the leaf ends seem to have been cartilaginous.

No other collections of Haworthia were known to G.G. Smith from the Potberg area other than perhaps H. minima. The Potberg area (see map) is also quite extensive as it is about 35 kilometers long and about 10km wide. My records at the time of writing Haworthia Revisited were virtually limited to reports of Luckhoff, Otzen and Herre of a small black species (maraisii!) and two collections of such a plant by both Frank Stayner and Peter Bruyns. I had twice been down to Infanta to be shown H. variegata by Chris Burgers on one occasion, and when I collected H. turgida south of Malgas on another. However, Chris Burgers had also reported two different plants, and one of these was from Buffelsfontein, south of the Potberg inside the De Hoop Nature Reserve. It was not possible to form any opinion about Burgers’ specimens without seeing live plants and I assumed that they were probably in the “mirabilis” complex. Adam Harrower recently came to light with a collection from Sandhoogte. This is also south of the Potberg, but about 15km east of Buffelsfontein. I was fortunate enough to see this population and was of the opinion that it was close to H. mirabilis. I was not convinced that this was what Prof Compton had recorded, but the phrase “sandy flats” also stuck in my craw. The Sandhoogte and Buffelsfontein populations (figs 1 to 5) are in very rocky streamside slopes and sandy flats are not the place to look for any Haworthia. The leaves in these plants are sub-erect and this seems to be related to habitat where slope and cliff dwelling plants differ from those on the flats. I regret that the pictures are not strictly comparable. We had spent the day trying to relate the De Hoop Oxalis records and scrappy specimens to correct names, and the final search for Burger’s Haworthia was more difficult than we had anticipated. By the time we found the plants, the light was too poor for photographing them. The photographs of the two cultivated plants obscure the close similarity of the plants in the field populations.

While partly motivated by trying to locate Trichodiadema and Acrodon species for Dr. Heidi Hartmann, my wife and I discovered extraordinarily pilose “retuse” plants along the lower Breede River (figs 6-9)  This is about 4km over the Potberg from Sandhoogte and I surmised it to be the same, as both those and the Buffelsfontein plants evidence incipient spination. All these collections flower in late summer and I consider them to belong in the greater “mirabilis” group as it is becoming increasingly clear that fewer species is more probable than more.

On our most recent visit to the area, we discovered an almost smooth version of the plants being discussed, at a point about 4km over the mountain from Buffelsfontein (figs 10-12). The habitat is extremely rocky and gravelly, as at the lower Breede River, and I am sure that this is now related to Prof. Compton’s collection. The fact that the leaves are retuse in these latter two populations is related to the levelness of the area. I include all four populations in an informal category I refer to as H. mirabilispilosa”, despite the fact that in one population only, the plants are startlingly hairy.

Now I come back to my starting point. The geology of the area is most complex. The Potberg is mostly Table Mountain Sandstone except for the eastern most tip which is Cape Limestone. At the foot of the western end there is Bokkeveld shale which is home to a tiny remnant of renosterveld and all of this vegetation type that is included in the De Hoop Reserve. South of the Potberg it is mostly sandy and the vegetation is “fynbos”. North of the Potberg is a different story all together. While the sandstones meet Bokkeveld shale, the situation is confounded by the extensive and complex tertiary deposits characterized by ferricretes, silcretes, terrace and fluvial gravels and other. This is reflected in the vegetation of the area and close alongside Fynbos genera one can find dense patches of Succulent Karoo species such as Gibbaeum austricola and Acrodon. If one can step from one vegetation type into another like this, it is axiomatic that the complex exposure of this mother material is going to have a dramatic influence on the species that comprise the vegetation.

While I think that the collections I record here are related to what Prof. Compton collected, the habitats can definitely not be described as “sandy flats”. However, I can by no means be said to have explored the area thoroughly. The plants are very cryptic and can be confined to very small and localized habitats so they are very easy to miss. ♦

Volume 6, Chapter 11:- North and Northwest of the Potberg

In an earlier article I described how Haworthia floribunda (at B on map) transmutes eastwards to H. variegata (at A on map) between the localities Klipfontein and Kleinberg, which are north of the Potberg Mountain. This is despite the fact that they both occur in close proximity at the northwestern end of the mountain. Difficulties now arise in the immediate vicinity to the west (at point W) and this extends northwestwards (to points C and D on the map). While we can confidently ascribe names to floribunda and variegata at those particular sites, the plants to the west and northwest are confounding. They fall into a no-man’s-land of these two species with H. mirabilis, H. maraisii, H. heidelbergensis and even H. mutica thrown in.

Localities and their names can also be confusing, so they are listed as follows:

A – the north and western point of the Potberg mountain on the De Hoop Nature Reserve and adjoining both Klipfontein and Juliesfontein farms (Juliesfontein farm has been renamed Poteberg Farm)

B – Byeneskop, a small hill on the western boundary of the farm Klipfontein

C – the farm Brakfontein

D – a hilltop north of Brakfontein named Witklip Kop

W – the two farms at Die Kop, Wydgelee, at the entrance to the De Hoop reserve

Figs 1 to 4 are from immediately west of A and show characteristics of floribunda in respect of the leaf tips. Figs 5 & 6 are of plants also from close by and the influence of H. maraisii is more evident. Generally the leaves are much shorter and stouter than one would expect in H. floribunda and the small size and numerous leaves suggest H. heidelbergensis. Both pointed leaves and leaves with the flattened and rounded tips may occur on the same plant (see figs 7 & 8).

Figs 9 to 13 are of two collections on the same ridge to the south of the Brakfontein homestead and are generally similar to the plants in figs. 1 to 8.

Figs 14 to 22 are plants from a small hill to the north of Brakfontein. There were few plants there as the hill is severely grazed by sheep. There is quite a strong resemblance to H. mirabilis in that the leaves are firmly sub-erect and slightly more robust. The hill is very typical of the ferricrete/silcrete/gravel inselbergs which occur throughout the southern Cape. They are characterized as hard, conglomerated, solid rock that caps the hills, and these are underlain by Bokkeveld shale that has decomposed to kaolin. Exposed Bokkeveld shale is the dominant parent material for the soils of the area and these are arable, whereas the rocky inselbergs are not. The consequence is that habitats suitable for Haworthia are the relatively un-weathered skeletal soils which can only support sparse vegetation. Thus distribution of the plants is island-like and no doubt this plays a significant role in the way the plants relate to each other. ♦

Volume 6, Chapter 12:- A look at Aloe haworthioides

Aloe haworthioides was described by Baker in 1887, moved to a unispecies genus Aloinella by Lemée in 1939 and then into Lemeea by Heath in 1993. This was wisely all undone by G. F Smith et al in 1995. This taxonomic dance is one of those events which probably bring taxonomy into disrepute in the minds of people otherwise respectful of scientific process.

A. haworthioides is rather symbolic of the deeper problem in the genus Aloe and related genera. This problem is that the genera are not firmly embedded in divine dispensation and there is much room for doubt about how they could be classified. Heath’s play with the generic status of the nine small Madagascan aloes rather demonstrates what can pass for well-reasoned, logical argumentation, just as surely as the same arguments were dismembered by Smith et al.

In pondering Haworthia, I have had to consider flower structure in relation to plants which are not Haworthia. So here I present a number of illustrations which show just how different A. haworthioides actually is. To answer “From what?” is a lot more difficult because of all the options available. A. haworthioides is a very small plant and seldom seems to reach a height of more than 80mm. It has many small slender leaves which are quite spinose to the point of “arachnoid’ hairiness and hence its visual similarity to Haworthia (fig.1 above).

The flower is something else and I suppose it could be described as “aloid”. The inflorescence is sub-capitate to cylindric with the flowers congested at the end of the peduncle (fig. 2). The tube of the floret is quite short and the lobes are free to the base (fig. 3). To what extent fusion of florets in the genus is indicative of relationship is a question I wish I could answer. It is nearly as problematic as the question of zygomorphy and actinomorphy, which is exacerbated by the fact that observers will insist there is some sort of twilight zone where the exclusivity of these states dissipates. In bulbs, writers describe the petals as “united into a tube” rather than plump for a word like “fused”. The conditions exist where there is a true tube which has no visible differentiation into tepals, where the tepals are fully fused and yet visible, tightly adnate and almost inseparable, and fully free. All four conditions may occur in the same flower with increasing distance from the base along the “tube”. The use of the word tepal is also prone to some doubt because it is used when the inner and outer petals of a flower are “similar” and it is not certain what constitutes “dissimilar”.

It is instructive to consider that Reynolds abandoned any system of formal sections for his second volume of Aloes of tropical Africa. Where in his first volume, he went to great pains to formally maintain hierarchy. In the second volume, he grouped 13 very small Madagascan species together and wrote “They do not comprise a homogenous group. Species with widely divergent characters are included only on account of their small size to facilitate identification”.

What is quite dramatic about A. haworthioides is the nature of the stamens. The filaments are exserted, but they are very broad so that they form an apparent extension of the floral tube.  It is again interesting to turn to Reynolds. He writes “Perianth white to pale pink, slightly campanulate…Filaments dilated, remarkably thick and fleshy, the 3-inner narrower and lengthening before the three outer, with their anthers in turn exserted.” I am inclined to grasp these words as justification for the poetic license or plain hermetics of my own writing. In the plant I have, the flower is hardly describable as campanulate (bell-shaped), the filaments are certainly subequal in length but very similar in width, while anthers are barely exserted beyond the length of the filaments. Reynolds poetic license seems to extend to the variety aurantiaca where “the whole spike, including the axis, bracts and perianths of a beautiful bright orange-red, with the stamens yellow”. Again in the plant at my disposal it seems as though it is only the filaments which are orange-red and the anthers only are yellow. The style is relatively short and is not exserted like the filaments.

I am sure that many other Aloe species could be analysed in critical fashion and probably more effectively than this, to show that the genus is possibly a very artificial construct. I say this because observers have tended in the past to take the same broad uncritical view of Haworthia, as Dr. L. A. Codd once did, when he wrote to me to say that “we are ignoring the small differences”. This was in response to my complaint that ALL Haworthia flowers were said to be small and the same when his staff lumped Chortolirion into Haworthia.

It seems to me that we are facing an onslaught on classification driven by the novelty and promise of molecular studies where DNA is seen as the ultimate solution to problems of classification. I have my very grave doubts. Aloe haworthioides is an oddity and while DNA studies will surely support this and enable better explanation of its relationships, identification and recognition by name may be better left as it is. Molecular biologists will lead us the way of Reynolds, which is the appreciation of the fact that hierarchical structure in classification is often misconstruction. We also do not all perceive names and their meaning in the same way. If we simply altered our way of thinking by an educational process perhaps we would be better off. The binomial system has worked remarkably well to date, and can continue to do so if we meddle with it with more circumspection. ♦

Volume 7, Chapter 1:- Haworthia retusa ‘nigra’ – Another grand finale

Introduction

I wonder. I have written so many words purporting to be my last that my credibility here too must be under stress. Two very recent articles of mine in Alsterworthia deal essentially with that issue, although they also cover the discovery of Haworthia mutica (Buffeljags) (= H. groenewaldii Breuer). They do not cover my subsequent thoughts on actually reading the description of this new “species” by Breuer, Marx and Groenewald. I hope that the present manuscript will explain why I reject this as a Latin binomial although anyone who is in the least familiar with my writing should already know. Spurred on by that discovery, I instigated a search in another area of the Buffeljags valley adjoining the Bontebok Park accompanied by Jannie Groenewald who informed me of what he had found in still another area I had long wanted to explore. So I instigated another search there too and again with Jannie. A discussion of these new finds is submitted to Cactus and Succulent Journal where I trust it will be published. The essence is already in Alsterworthia and this article is written to widen the readership, submit more pictures and maintain continuity with the 6 volumes of Haworthia Update that Harry Mays has been so conscientiously and determinedly publishing. This is all writing that may not otherwise have seen the light of day. I am personally extremely grateful for that as I have had a mania since writing my revision Haworthia Revisited and Update Vol. 1 (both Umdaus), to set the record straight and explore all the unknowns, or at least some of them. So on with Haworthia retusa ‘nigra’.

The second area opened to me by meeting Jannie, was a farm southeast of Tradouw Pass, Heuningklip. In the Tradouw Pass, H. retusa ’turgida’ is found on the steep cliffs at the southern end. The road has been rebuilt since I first saw the plants and the population is greatly reduced from what it was. I simply do not have a picture handy to show what those plants are like, but that is not the need here and it does appear in Haworthia Revisited.  Immediately south of the pass is another population of plants that puzzled me from the moment I saw them until now. In Haworthia Revisited there is a picture of one clone under J.D. Kobus Venter’s number 93/35. It is named as H. magnifica var. magnifica. The plants are present in a tension zone between Shale and River gravel, and vegetationally between what was Karoo Valley Bushveld and grassy Renosterveld. In April 1997 I collected seed and grew countless seedlings from there, which were all different. Only one picture, as MBB6666) is included in my long treatise on what I discussed under the title of “How to understand H. mutica var. nigra” in Haworthia Update 2.1:50. That picture is also under the name H. magnifica var. magnifica but it was not intended to stay that way. It must be noted that my use of names in the Update volumes was not in a decision-making process so much as a learning and informative one. I read my conclusion to the treatise and note that I made no formal name changes. It is only as further field exploration yielded more and more information that I came to realize that H. magnifica and H. mirabilis, classification-wise, are inseparable. The Tradouw collection still worried me because of the close proximity of H. retusa ‘turgida’. These plants at Tradouw and all the things I referred to as H. mutica ‘nigra’ are in fact better understood as variants of H. retusa in the complexity of its relation to H. mirabilis.

This article thus covers exploration of an area between Swellendam and Heidelberg that I have neglected for too long. It is particularly applied to the problems of classification and naming that have persistently clouded my life and my relations with other writers. But as importantly to me is the unclouding of the way taxonomy and classification is useful and necessary for those interested at all in plants and how it is viewed by them.

 

Consideration and use of formal names
Here I need to explain the naming style I now adopt. I have dropped the use of any rank below that of the species name. I do this because botany has no proper species definition and consequently species descriptions are just based on wild guesses about possible non-similarity and on the flimsiest of supposed character differences. The loosely used word “typical” is only truly useful in respect of the one plant dried as an almost unrecognizable herbarium specimen that is used to anchor the Latin name. The scientific ranks are aggregates and the terms variety and forma are so too. There is simply too much variation between populations and within populations to enable the certain identification of almost any plant that the binomial system suggests. There are no clear entities and ranks and this article should demonstrate that. There is a separate code of nomenclature that covers cultivars. Note that I do not drop names. I only drop the suggestion of status that additional names have, be they single clones or population references for all the different plants within them. I have used names in Haworthia for 70 years and intensively for the last 40 years and I am very familiar with the difficulties of any system. For the field and all the populations and plants I have seen, I can do no better than offer the list of names I updated in February 2009 in Haworthia Update 5.2:192. This, and the set of Updates, comprises a near total overview of the genus and effectively a revision. The attempted revisions by others fall short for me because they do not cover the range of populations that I have seen nor what history tells me, although they may include populations not seen by me. That is not strange because the area to explore is still vast. Another problem with even trying to be formal, is that which G.G. Smith encountered (Mrs. L. Bolus did too) where several people were generating new names at the same time and trying to keep track formally with concurrent name changes would not have been, is not, possible. The point I should make is that in the last ten years, my field work has confirmed the predictive nature of my classification and these latest finds are in extraordinary agreement with that kind of expectation. What I have found essentially agrees with what I would expect. This is dramatically so for H. mirabilis based on a find by J. Jelimicky at Sandhoogte midway geographically and visually between H. mirabilis ‘bobii’ Hayashi (my nom. nuda ‘pilosa’, ’velcro’, Steven Hammer’s ‘Shaggy dog’) along the Breede river, and a collection of mine further south in the DeHoop reserve, MBB7886 H. mirabilis, discussed (p11) and illustrated (p33) in Haworthia Update 5.2.

Prior to extending exploration in the Buffeljags area, I had also been down to the Haworthia populations in the lower Breede River area. This is where H. mirabilis seems to assume a wide range of forms and H. retusa ‘turgida’ and H. variegata’ are also present. I also, in the company of Kobus Venter and Lawrence Loucka, revisited Kransriviermond and Morning Star south of Heidelberg where H. floribunda is in the mix with H. retusa ‘turgida’ and H. mirabilis (in my broad new sense that includes H. maraisii, H. magnifica and H. heidelbergensis). Thus many of those images were very fresh in my mind when I went to Heuningklip southeast of Tradouw. Looking at all the plants and in editing my pictures I was truly impressed with the reality of continuity. The essence of H. retusa and H. mirabilis is in all those plants. A mobius strip is a strip of paper twisted and joined at the ends to produce a shape that only has one surface. What we have in the Southern Cape, and covered or touched on in this article, are interlinking mobius strips with one main surface as well as loose ends that present identifiable end-points. I tried once to illustrate the continuities between populations with single or multiple lines joining the different populations to express degrees of similarity. I can also visualize a pie-chart in which each population is represented by a circular chart with the contribution of main elements drawn as slices proportional to the strength of contribution. Table 1 is a schematic and very crudely suggested enumeration for such a set of charts. I have put the elements retusa and turgida as separate under an H. retusa heading for no other reason that I want to emphasize turgida as a variant at the two geographic extremes. One is eastwards where H. retusa ‘retusa’ is absent (Brandwacht, Mossel Bay) and one westward where it is present (in mutica, Hasiesdrift, Bredasdorp). H. retusa ‘turgida’ is present both east and west. But there are many populations contributing to the H. retusa pie in all the ranges of the two parts, from 0-50% thus making up the continuity from retusa to turgida.

Consideration and characters.

a. The pie chart
Measuring the similarity of any two plants is extraordinarily difficult. Doing this for a population is even more so. Doing so for many populations, including those unseen in order to fulfill the predictive requirement of a proper hypothesis, is next to impossible. Hence all the disputation and name abuse. In table 1, I speculate an arrangement to illustrate the problem. The pie is constructed from the figures in the chart for just that set of genetic characters that are directly relevant to the set of plants in question. The pie is incredibly big and it should be understood that there must surely be pie-parts that spread their influence beyond the restrictions of the slices I conceive. So my table is restricted to the few species that are immediately relevant in respect of geographic association and direct interaction (by possible cross-pollination?). A pie-chart is drawn for each individual population. In the chart I assume that there is a set of genetic information that programs for H. retusa ‘retusa’. I have also suggested a different set for H. retusa ‘turgida’. If one considers only these two elements and all the known populations, my suggestion is that there is a set of charts that moves smoothly from one set to the next with no interruption. There is no discontinuity and the sets average 50:50 for the information. Thus it is one species and H. turgida is noted as H. retusa ‘turgida’.

Table 2 is the complete set of pie-charts that should convey the possible relationships of the different populations and from which can deduce and understand to what degree H. retusa and H. mirabilis can be related. Also it will indicate the similarities between, and the difficulties of, trying to explain H. pygmaea in the east and H. mutica in the west. Both arise from the same genetic source. The problem is compounded by the infiltration of H. floribunda in the area midway between Heidelberg and Swellendam with the re-emergence of H. floribunda in the south at Potberg. H, variegata also enters the picture but from the south and also in the Potberg area. H. emelyae is the very probable combined entity entering the little Karoo. H. rossouwii is almost independent except in the very south-west at Bredasdorp and it may possibly link to other elements such as H. herbacea and H. reticulata.

Table 2. Pie Charts
Table 2. Pie Charts

At Heidelberg there is interaction between an element that is a 50/50 retusa/turgida with a variant of H. mirabilis cf ‘heidelbergensis’. The latter name suggests that H. mirabilis is the general name for all the plants of that species, and ‘cf’ is an abbreviation for ‘compare with’ ‘heidelbergensis’  to refer to the small plants in populations previously regarded as independent species.

b. What really constitutes the pie
The pie is the totality of the genetic material, the DNA that determines all life forms. It is the genes, the materials of characters and inheritance, and the repository of all genetic information. The pie is very large indeed and it is said that the DNA pie of an elephant only differs by 3% from that of a mouse; that of a man by 1% from a chimpanzee. In Haworthia one species from another ‘species’, I could guess at 0.001% or less regardless of whose concept I follow. This difference does also not reside in one consolidated slice that can be reduced to a single small pie as I have done. In DNA analysis, certain segments of DNA from cellular chloroplasts, mitochondria or the chromosomes themselves are isolated and the sequence of the paired amino-acid is analyzed. It requires a formidable statistical process to arrive at a two-dimensional tree (phyllogram) that illustrates a branching relationship with a measure of branch separation and lengths. This can perhaps also be used to generate pie-charts with an innumerable number of slices. I personally consider the phyllogram generated in the DNA sequencing process grossly inadequate.

c.) The characters
This is extraordinarily difficult to specify no matter how obviously the plants seem to differ from one another. Number, size, shape, colour, arrangement, surfaces, spination and attitude of the leaves in the spiral rosette constitute the main things that determine how we see the plants. These are influenced by growing medium and growing conditions so these are complicating factors. Flowers are used in the same way as the leaves but far less so because they are so similar in all the species. Flowers are also not always present either which reduces their usage. Time of flowering is a major factor because it is linked to the reproductive process. It is also difficult to capture any firm picture of the flower because of the aging process in the 3-5 day life of the individual flower.

d.) The relevance here
It simply is not necessary to parade the obviousness of this and the way in which will be attempted anyway in the discussion below. It is flowering time that may be critical beyond, or linked, to the facts of spatial geographic relationships of populations. It is traditionally flower structure that has formed the basis of classification of plants because of the reproductive significance and the role of reproduction in distribution of genetic material between plants and populations. Pollination of Haworthia seems to be primarily by a Solitary bee species. These bees make individual nests and are not communal like the Honey bee. They make tunnels in the soil as nests during the summer months. I have observed them as adults in the winter but then they are less numerous and less active. Few Haworthia flower at the height of winter and do so in two main time frames viz. spring and late summer. The implications are that breeding between isolated populations apart from any other consideration of compatibility, is a question of distance and time. A nest bound Honey bee can forage to 13km from its nest. A free flying Solitary bee can be speculated to cover a considerably greater distance. All these things play on the frequency and power of inheritance. Genetic drift is acknowledged as the genetic movement within a population away from the norm for a species because of isolation, while genetic flow is perhaps the holding together of similarity over geographic distance by a travelling pollinator. Thus the statistical probabilities of breeding across distance are impossible to derive.

I cannot quantify any of these things nor suggest any greater knowledge but this suggests to me, given the freedom with which the various species hybridize in cultivation and even in the field, we need to consider geographic relationships. In my extensive experience with other genera, it appears to me that a key consideration is if things grow together and maintain their difference, or not. Haworthia species are invariably in small isolated populations and seldom do different species grow intermixed. They are also influenced by geology of the substrate and particularly so because they favour the lowered competition from more vigorous and larger plants on the rocky sites or shallow soils they favour. That can be stated in the converse, Haworthia are favoured by conditions poor for other plants.

e.) Flowering time and growing together.
The existence as discrete species can be judged from the way in which populations share geographic space. I discussed this in some detail in Update 2.1:76. If two sets of plants grow close together and flower at the same time, while still maintaining obvious group differences, then they are most probably different species. If they flower at different times while still maintaining difference then the question of being different species is taken as certain. If the groups are separated and flower at different times while the differences are small and apparently within the experienced range of variation observed between populations, then it may be problematic. One would need to resort to the extended distribution and variability of the groups in question. Generally in Haworthia, species belonging in the same sub-genus do not share habitat and this is indicative of fairly rapid absorption of introduced genetic material (by pollinators). Hand-pollination and the ease with which plants of different species hybridize in cultivation suggest that there is practically no breeding isolation between them. In the following discussion, use is thus made of general distribution of species and proximity of populations in assessing to which species they best belong. It should be borne in mind that variability may be clinal. This means that the plants change along a geographic gradient. The direction of the gradient may also not be of increasing distance i.e. linear.  It may also be circular so that ends of a gradient may meet again as two populations that appear wholly different, but linked by an extended set of other populations. A much more complex situation may actually exist where these clinal gradients may be multidirectional and thus involve more than one apparent species. My observation is that populations do differ and differences follow a pattern that becomes predictable i.e. gradual genetic difference over geographic area. This is not just a result of environmental gradient, but also linked to the question of genetic drift (the tendency of isolated populations) to move away from the general norm. It also results from the genetic flow between populations where the change of genetic material between populations at the distribution extremes is only through other populations.

f.) The species definition
This is a subject I have covered many times. My view is that species are dynamic chaotic systems oscillating between different character states with time and with different rates of mutational change. Closely related species may not be recognizable on the basis of single character states and the spatial geographical distributions and local juxtapositions of groups of plants may be the only way to understand what the situation is and so make sensible decisions.

g.) Habitat geology
I contend that species richness in South Africa is largely due to its varied geology, skeletal soils and comparatively mild climate. This present discussion deals with species that are associated with a geological phenomenon that is the Tertiary deposits of the Southern Cape. I am no geologist and am required to make the pretention I am one to explain why plants are distributed the way they are and what influences difference among them. In this case the vegetation at Buffeljags, Swellendam has been described (by Breuer et al. cited below) as on silcrete, and the general vegetation as by vegetation scientists as Ruens Silcrete Renosterveld. But there are two problems. The first is that vegetation classification is not infallible and I have never found it really useful in dealing with a wide range of species that occupy skeletal habitats. Secondly the substrate geology is definitely not simply silcrete nor is the clay soil either simply bentonite or kaolinite. There are many problems in the classification of clays. Bentonite in common terms seems to be the yellowish clay mined in large quantity from Heidelberg eastwards and there is a transition to ochre bearing clay. Bentonite may be aluminium, potassium, sodium or calcium dominated. As a montmorillomite clay it should have expansive cracking properties.  Kaolinite is popularly regarded as the very white clay that is commonly seen under the rocky top layer of the sharp inselbergs that rise above the wheatfields west of Heidelberg.  It seems to be less mineral dominated clay and does not have the expansive/contractive properties of bentonite. The distinction between ferricrete and silcrete seems to be that in ferricrete, iron oxides are infused into the petrified water deposited layers. Both the kaolinitic and bentonitic clays seem to be derived from the weathering of Bokkeveld shale that underlies the silcerte/ferricrete layer. It does not require any stretch of the imagination to see now why any one inselberg may be a little different from the next, and sometimes a lot different. What we do not know is how rapidly the landscape has moved from a broad white silcrete/ferricrete underwater plain to the remnant condition of that rocky plain today.  Neither do we know what it has meant for the plant species that are linked to, and dependent on, the habitats generated in the erosion processes.

Map Legend – east of Swellendam.

1. MBB7899 H. retusa ‘nigra’.  Heuningklip.
2. MBB6666 H. retusa ‘nigra’ ↔ H. mirabilis.  S Tradouw Pass.
3. MBB7896 H. retusa ‘nigra’.  Heuningklip.
4. MBB7897 H. retusa ‘nigra’.  Heuningklip.
5. MBB7898 H. retusa ‘nigra’.  Heuningklip.
6. MBB7888 H. mutica.  Rotterdam.
7. MBB7889 H. mutica.  Rotterdam.
8. MBB7890 H. mutica.  Rotterdam.
9. MBB7801 H. mutica ‘groenewaldii’.  Buffeljags.
10. MBB7741 H. mutica.  Dankbaar.
11. MBB7892 H. marginata.  Rotterdam.
12. MBB7891 H. minima.  Rotterdam.
13. MBB6644 H. mirabilis.  SW Swellendam.
14. MBB7704 H. mirabilis.  Bontebok Park.
15. MBB7805 H. mirabilis.  Bontebok Park.
16. MBB7823 H. mirabilis.  Klipbult.
17. MBB7887 H. mirabilis.  Rotterdam.
18. MBB7876 H. mirabilis.  Disselfontein.
19. MBB7901 H. mirabilis.  Crodini E.
20. MBB7902 H. mirabilis.  Crodini E.
21. MBB7903 H. mirabilis.  Crodini E.
22. MBB7904 H. mirabilis.  Crodini E.
23. MBB7905 H. mirabilis.  Crodini E.
24. MBB7906 H. mirabilis.  Crodini E.
25. MBB7907 H. mirabilis.  Crodini E.
26. MBB7908 H. mirabilis.  Sandkraal W.
27. MBB7909 H. mirabilis.  Sandkraal W.
28. MBB7910 H. floribunda.  Rietkuil.
29. MBB7912 H. mirabilis.  Rietkuil.
30. MBB7913 H. mirabilis.  Rietkuil.
31. JDV93/35 H. retusa ‘turgda’.  Tradouw Pass.

The populations

Set 1 –  THE TRADOUW  PASS AREA (H. retusa ‘nigra’)
This first section deals with populations that complement those in the long discussion under the title “How to understand H. mutica var. nigra” in Haworthia Update 2.1:50.  There I ended with the words … ”The habitat photo (of H. mutica Hasiesdrift) raises such strong images of H. retusa that doubt about the affinity of the var. nigra cannot easily be laid to rest.”  Added were these words … ”There the saga for the moment must rest.” What I had in mind was exploration that no one seems very anxious to do. This has now been done and I would lay the matter to rest as follows:

1. MBB7899. H. mirabilis, Heuningklip. See figs 1-30  Fig. 1 is a scenic view of the area to show the countryside and the type of inselberg the plants are often found on. Fig. 2 is a closer picture of the erodible clay held in place by grass clumps at the base of which the plants often are. Occasionally they are simply out in the open sun. I term these places pressure bursts because water build-up in the soil causes collapse of the small banks and erosion can be quite fast. Several times I have found plants lying in collapsed soil. Such a soil bank near Heidelberg has retreated by nearly 2 meters in about 15 years albeit accelerated by road maintenance, but it does relate to any attempt to explain how much suitable habitat may have disappeared with time.

Figs 3 to 30 is a selection of plants at the site where one is almost compelled to examine each plant separately and easily generate another 28 practically different images. Fig. 3 is of a plant with the surfaces roughened by small tubercles. SEM (scanning electron microscope) images of similar leaf surface often show incipient tiny spines on the tubercles and they appear as spines or hairs in populations like H. mirabilis ‘pilosa’ at Ballyfar, or H. mirabilis at Windsor Riversdale. This surface roughness and all its degrees is fairly characteristic of H. mirabilis but is does occur in H. retusa too. Plants illustrated in Figs 4 and 16 has one muttering Kruisrivier and Komserante, northeast and east of Riversdale respectively, and which I would have allotted to H. magnifica in the early period of my exploration no doubt have Breuer and Hayashi names. Now I simply say H. mirabilis and add the locality to place the plant to at least give it a geographic reality that is better than using the formal Breuer/Hayashi in hanging commas as I also suggest. Fig. 6 is apparently a seedling and demonstrates the leaf-shape of H. floribunda that led me to suggest that H. floribunda is a perpetuated juvenile form. Of course the evidence suggests to me that H. floribunda coalesces with H. mirabilis in this wider area although it does appear at Swellendam to the west of my map as the large form and at the extreme SE corner of the same map as a convincingly normal form. Fig. 9 demonstrates longer pointed leaves and Fig. 12 short more obtuse leaves. Fig. 17 is of a plant that has relatively smooth leaf surfaces rather like H. retusa and the leaves are conspicuously lined. In Fig. 20 there is a suggestion of muticate (without a point) leaves. Fig. 22 is again quite strongly lined. Fig. 22 shows a leaf with a single line. Fig. 25 demonstrates what is termed rotate leaves where the points of the leaves point around as though to encircle the plant. Fig. 28 made me recall plants of H. mirabilis at Melkhoutrivier in the south west near Malgas and further to Ballyfar. The flowering time is late summer.

These images can now be compared with MBB6666 at Tradouw Pass to establish if my assessment is reasonable. My contention is that this MBB7899 population is midway between the other three Heuningklip populations I am about to illustrate and Tradouw Pass. This similarity is repeated widely and eventually contributes to my contention that H. mutica has been generated by drift away from its two contributing parent species viz. H. mirabilis and H. retusa.

2. MBB6666 H. mirabilis. Tradouw Pass See figs 31-71. This population is at the mouth of the Tradouw Pass and relatively low-lying in respect of the previous population. The habitat is quite different being partially coarse river gravels, weathered shale and at one point is an un-weathered shale ridge where the plants are wedged in the cracks. The habitat is of course very different to the steep sandstone cliffs within the pass itself where H. retusa ‘turgida’ occurs. I initially thought these plants fitted with H. magnifica. A decision derived from a very small sample available at the time. Now I looked for the images and photographs in my records under H. mirabilis where they should have been since my inclusion there of ‘magnifica’, and also under H. retusa where I found them in the folder ‘nigra’. The flowering time is late summer. In this respect similar to the previous set MBB7899 H. mirabilis but the plants are more robust. Figs 31-45 were taken in 2009; figs 46-53 in 2011 and figs 54-71 are from seedlings grown by myself. I tried to reduce the pictures and eliminated a few. But I really think they all need to be presented to show again how serious the difficulty of circumscribing them as a single recognizable key-able entity really is. While looking at my pictures I seriously thought that the set taken in 2011 differed from those taken in 2009. The cultivated plants of course are not directly connected to their source and the variation is self-evidently wide. But it is equally so in field plants. There are the smoother surfaces of H. retusa and some very rough surfaces for H. mirabilis. My decision became that they fitted best with the ‘nigra’ populations where my organizing had taken them despite flowering time, but here MBB7899 shows absolute geographic position between H. mirabilis and MBB6666. I consider this key evidence for the basic continuity for H. retusa and H. mirabilis that I have conjectured elsewhere and I would not be able to defend an argument that MBB7889 plants belong better in H. mirabilis and this means MBB6666 belongs there too. I would simply dispute that there is no better solution so stick with history as far as possible and that means H. mirabilis. If Breuer or Hayashi have a name for a variant from this population intended as an aggregate name, I recommend the use in inverted commas. I would actually like the use of the equilibrium diacritic of a double arrow pointing between two names viz. H. mirabilis↔H. retusa ‘nigra’, although this available symbol is not quite the thing.

I come to this vexing question of “typical” that is bandied about even by myself when I liken population to population. There is the same range of variables in MBB6666 noted for MBB7899, but the plants do not darken as much and they are larger. So we need now to look at the other three Heuningklip populations asking what significance is attached to their presence in exactly the same habitat as MBB7899, but geographically separated and by perhaps 500m and 2km respectively. The plants are so variable that it is not possible to arrive at a characterization that separates any two plants, or any two populations in any geographic set for that matter.

3. MBB7896 H. retusa ‘nigra’, Heuningklip 1 See figs 72-89. The plants in this and the next two populations were flowering in September as does H. retusa and they are relatively large, green in colour and generally smooth. Of course both vary enormously due to both intrinsic genetic difference and extrinsic local situation. One plant within an older decaying grass tuft was nearly 200mm diameter, and a healthy green colour, another plant in the open white clay was less than 50mm diam. and dark in colouration.

4. MBB7897 H. retusa ‘nigra’, Heuningklip 2 See figs 90-105. The first two pictures show the general landscape and the local habitat where the plants occur. The clay is highly erodible and often there are no plants at all in the clay. Some of the plants have very acuminate pointed leaves while in others they may be relatively obtuse and rounded.

5. MBB7898 H. retusa ‘nigra’, Heuningklip 3 See figs 106 -132. The great variability is again evident and in this population the plants were observed to be generally without much venation or lines as in fig.112. Then one plant, fig. 119, was so striped as to stand out from all the populations seen.

Set 2 – THE BUFFELJAGS AREA (H. mutica)
This second section follows a recent manuscript submitted to Alsterworthia yet to be published entitled “Some last closing thoughts” written in the hope of finding the closure I seek and that will not come. That article is about a population that has been named as a species viz H. groenewaldii, in an article authored by Breuer, Marx and Groenewald (Alsterworthia 11.2:15) that I only saw after the submission of my manuscript. I also only saw the article after the exploration I would have asked for to verify opinions. The description seems to have been prompted by Marx who refers to similarities to H. springbokvlakensis and to H. magnifia var. atrofusca and raises many issues that my manuscript does not cover. I simply find the whole discussion unrealistic and especially so when no further exploration was done to examine anything. One should refer to Breuer’s World of Haworthias Vol 1 to see that the naive promises in the foreword and introduction of exploration and field study, was never achieved. It is true that the plants have an unusual surface texture and it is also true that the habitat is quite unlike any other H. mutica occupies. But it is Marx’s comment that disturbs me viz. “Unfortunately Bruce Bayer keeps on refusing to see it as something new and calls it simply H. mutica ‘Buffeljags”. This is a distortion and misrepresentation, and whose misfortune this is, not specified. My reticence is reflected in the preceding discussion and Gerhard does not seem to follow, or agree with, what I have been writing about and trying to explain in this general context for so many years. In fact in an unpublished (to my knowledge) manuscript he dismisses all my writing as “pretentious” that no doubt it is, because I have never fancied my self as a taxonomist in the tradition of taxonomists.  I was as fascinated as Marx by the plants and am even more anxious than he is to see just how they connect to the probable geographic counterparts in the context of the species definition that I apply. I was also very curious to know what the connection to H. mutica and the H. retusa ‘nigra’ issue, only now laid to rest in the preceding discussion, actually might be here. This is why I use the word “instigated” in reference to exploration in the very valley where H. mutica ‘groenewaldii’ occurs and to the obvious habitat for similar plants. It is surely unfair iniquitous to so demand that I see this population as “something new” in an article that rather disregards the history of the subject and what I have been trying to understand of it.

I also recently submitted a manuscript to Cactus and Succulent Journal (US) where I wrote of “H. groenewaldii Breuer” “I consider it to be generated from the interaction of H. mirabilis, H. mutica and H. floribunda. I attach no special importance to the fact that it flowers, contrary to H. mutica, in February/March. This is because I have observed many hybrids between patently different species despite a seasonal difference in flowering time”. I have in many places discussed the problem of “hybridization” and it must surely be well-known from my writing that H. floribunda merges into H. mirabilis. Less well-known will be the fact that evidence for H. variegata doing the same is also available. In respect of floral characters, I wrote a chapter in Haworthia Update Vol.3.2:88 explaining the problems of using floral morphology for species recognition. I would challenge Breuer et al. to produce comparative results from studies of the flowers in any of these problematic species to demonstrate difference. Regarding the curious shiny leaf surface, I refer to Haworthia Update Vol.2.2:141, Chapter 15 where I discuss electron scanning photographs of leaf surfaces. This also demonstrates how problematic leaf surface is. It would be tiresome indeed to parade all the evidence in support of these contentions of mine. In Haworthia Update 5.2:108 I also briefly discussed H. mutica from Buffeljags (‘groenewaldii’) and submitted a large number of photographs explaining the habitat and what the possible connection was to the H. retusa ‘nigra’ conundrumI may also have written about the issue in Haworthiad. Suffice to say that the authors of H. mutica ‘groenewaldii have certainly not checked or considered “all the facts” as the principal author claims.  The very opposite seems to be the case.

Considering my oft repeated observation that geological substrate plays a major role in shaping species appearances, the Breuer, Marx and Groenewald description of the habitat for H. mutica ‘groenewaldii’ is a little misleading (see considerations above). Nowhere else is H. mutica on identical substrate and this would just support my contention of ecotypic differentiation that does not justify a Latin binomial. There is considerable doubt about the actual difference between these river boulder situations and the widely distributed terrace gravels elsewhere in the area, notably the Bontebok Park near where the following populations were found:

6. MBB7888 H. mutica, Rotterdam 1 See figs 133-169.

7. MBB7889 H. mutica, Rotterdam 2 See figs 170-188.

8. MBB7890 H. mutica, Rotterdam 3 See figs 189-191
The three populations above are separated spatially by several hundred meters and they are all on remnant river gravels on a more gently sloping bank than the Buffeljags east bank. The vegetation is closer to conventional fynbos as in the adjoining Bontebok Park, but grassy. The pictures are self-explanatory and the leaf surfaces of the plants are also generally as for the Buffeljags population. What is significant is that I cannot say I saw a single plant in these populations that resembles the one chosen and pictured as typical for this highly dubious species, H. groenewaldii.  It makes more sense to me to identify plants from these populations by collecting number than to generate a supposedly descriptive Latin epithet from an imagined typical representative.

9. MBB7801 H. mutica ‘groenewaldii’, Mullershof, Buffeljags See figs 192-215
The following set of photographs has surely seen the light of day somewhere and I repeat them because as far as I am concerned they are H. mutica with the shiny surface claimed for “groenewaldii’. Dankbaar is some 16km west of these Buffeljags populations and the substrate is Table Mountain Sandstone. The flowering time is just normal for H. mutica.  Fig. 215 is a photograph of a plant apparently collected by Jannie Groenewald and it is quite uncharacteristic of the population and would suggest that it is a product of hybridization with H. mirabilis in the vicinity (e.g. Klipbult).

10. MBB7741 H. mutica, Dankbaar See figs 216-235.
My observations in respect of these two sets of plants confirms my widely written convictions about Haworthia taxonomy and the state of classification in the absence of a definition for the concept of “species”, and the intrinsic value of Latin binomials for rational and sensible communication. Of course this Dankbaar population is not identical to other H. mutica populations and I am not driven by fixation on any one character that defines this species. It is a geographic set and the various sets are not simply separable. But I am not finished yet.

Set 3 – THE BUFFELJAGS AREA (Robustipedunculares)
The western Buffeljags area also produced a population of H. marginata and one of a very curious H. minima, that both complement occurrences in the Bontebok Park a little further west. The conventional small, blue-green H. minima is present in the park as well as an unusually green population.

11. MBB7892 H. marginata, Rotterdam See figs 236-243.
This population was quite exceptional and one of the strongest populations known to me. But the plants were in flower in September whereas the flowering time for the species is normally late summer; and the flowers and seed capsules exceptionally large. I do not consider that these factors command another Latin binomial.

Fig. 243 7892 H. marginata.

12. MBB7891 H. minima, Rotterdam See figs 244-252 and Bontebok Park figs 253-4. These were also extraordinary plants, larger than those seen in the Bontebok Park population where the plants were also green viz. figs 253-4. Fig. 252 confuses the issue because, while the other plants seemed alike to H. marginata and distant hybrids, this one plant appears to be a recent hybrid. This occurrence cannot be overlooked in respect of other introgressive (or seemingly introgressive) populations of Robustipedunculares species. The plants were not in flower and again we have a suggestion or direct evidence of hybridization across a difference of seasonal flowering.

Set 4 – THE GREATER BUFFELJAGS AREA (H. mirabilis)
This is a set of 15 populations of H. mirabilis that occur east, west, south north of the H. mutica ‘groenewaldi’ related populations. I exclude from the discussion and  illustrations the several populations of H. floribunda ‘major’ populations to the northwest, while I repeat the statement that H. floribunda contributes to the general gene pool of H. mirabilis and together with H. retusa, they all contribute to the gene pool of H. mutica ‘groenewaldii’. The huge presence of populations more readily assignable to H. mirabilis (sensu lato – in the broadest sense) is used by me to validate my opinion that the “mutica” elements draw most genetic resource from the H. retusa element, and hence why I refer those plants to H. mutica.

The sequence of populations 7901-7908 is located in a horseshoe of the Breede River, southwest of Buffeljags in very promising habitat. Only H. mirabilis was seen. I perhaps need to point out that these populations would have been extraordinarily difficult to name under any previous dispensation of mine where I was still trying to uphold H. magnifica, H. maraisii and H. heidelbergensis as species. These I have since merged in H. mirabilis and I reject the introduction of a group name like “aggregate” as Breuer seems to have done, as a complete fabrication that does nothing to solve the problem other than to highlight it.

The following 6 populations are all associated with tertiary and river gravels. There are differences between most of the populations but of an order that I cannot articulate. It is also not possible to really encapsulate the appearances of the plants in the two photographs for each population that I submit. At the same time it does not seem to me possible to make any communicable distinctions.

13. MBB6644 = 6860 H. mirabilis, SSW SwellendamSee figs 255-256.

14. MBB7704 H. mirabilis, Bontebok Park See figs 257-258.

15. MBB7805 H. mirabilis, Bontebok Park See figs 259-260.

16. MBB7823 H. mirabilis, Klipbult See figs 261-262.

On selecting these two pictures I was reminded of my observation that there was a significant H. floribunda appearance to the plants. I had believed that I had seen H. floribunda in the Bontebok Park nearby many years ago but subsequent exploration has only produced H. mirabilis albeit with some similarities to H. floribunda. My reaction was to suspect that H. mutica ‘groenwaldii’ most probably also then needed to be considered as a product influenced by H. floribunda.

17. MBB7887 H. mirabilis, Rotterdam See figs 263-264.

18. MBB7876 H. mirabilis, Disselfontein See figs 265-266. These plants were a little more robust than the others in this set and rather similar to MBB7901.

The following populations are almost entirely in Bokkeveld shale but on a very close interface with tertiary gravels. In some places there was also some wind-blown sand.

19. MBB7901 H. mirabilis, Crodini 344, S Swellendam See figs 267-278.

20. MBB7908 H. mirabilis, Sandkraal, S Swellendam See figs 279-282.

Set 5 – BACK TO TRADOUW PASS – AS IT HAPPENED
In the process of compiling this manuscript I was bothered by the ever present question of what is still out there that might influence the outcome. My wife, Daphne, was planning an outing to see chameleons at Swellendam while simultaneously contacting landowners for me around the Potberg. Always anxious to get two birds with one stone, I asked her to find out who owned the land west of Heuningklip that we had just explored with Jannie Groenewald. The result was that we ended up spending a day with Odette Curtis of Custodians of Rare and Endangered Wildlife, south of Suurbraak and finding three populations that, for me, put the proverbial cherry on the pie.

All three were on the slopes of silcrete inselbergs and almost the last of this habitat westward from Heidelberg to Buffeljags. The first inselberg was different to the second and from the Heuningklip habitats because it was dominated by Aloe arborescens and A. feroxAloe brevifolia was also present in an unusually small form. The habitat was very impacted by grazing animals and there is an outside possibility that H. retusa ‘nigra’ may once have been present although I personally doubt it. To our surprise we found…

21 MBB7910 H. floribunda, Rietkuil, Suurbraak See figs 283-288.

There were many plants but severely trampled and disturbed. Their survival was obviously aided by the fact that much of the plant is below soil-level. But my experience of this species is that it struggles to maintain its identity in the presence of either H. mirabilis or H. retusa. They never share immediate habitat. There was no hybrid present and this also adds to the problem of interpreting flowering time and distance apart from other species as part of the process. The plants were not the large green variant of the species present to the west at Appelbos, Swellendam; and the leaves were both with and without marginal spines. Eastwards I know the species to be close only at Blackdown north of Heidelberg and at Goedverwagting south of the N highway about 15km east southeast. It definitely seems to be absorbed into H. mirabilis west and north respectively of those two populations. So while this is a slight contradictory element of surprise, it is the next two that really excited me. These were again inselberg habitat but without the strong silcrete upper layer of the former habitat and without the Aloe element. The second ‘population’ was about 200m from the first and comprised four groups of plants along a longer north-facing ridge.

22. MBB7912 H. mirabilis, Rietkuil, Suurbraak See figs 289-306.

23. MBB7913 H. mirabilis, Rietkuil, Suurbraak See figs 307-331
I am very hard-pressed to say much more than that these plants met all my expectation for the very variable set of plants and populations that I ascribe to the one single species H. mirabilis. Who could possibly articulate the differences of the many populations of this species from each other and from this one?  Variable becomes a done-to-death word.   The individual plants, in two populations quite unlike any others I have seen, were generally robust with large specimens up to 100mm. diam. The colour is dark, although green when shaded within or under grass. The leaves were usually sharply pointed but there was always evidence of one or two leaves or odd individuals in which the leaves were muticate i.e. without points.  In some plants the leaves were notably lined in others not. The colour also varied from chestnut to deep reddish-brown when the plants were more exposed. The surface texture was generally tubercled but there was evidence of smoothness of the leaves in many plants. Flowering time was obviously late summer gauged by the absence of flower at the time of the visit viz. September. Except for one oddity. This was a single plant with a weak flower spike. I photographed this for the useful information the flower-bud provides. Where Breuer et al. claim to have considered all the facts, this is a character that is conveniently overlooked when claims are made for the significance of floral characters when paraded to substantiate yet another claim to species significant difference. The bud-tip in all the Southern Cape species of subgenus Haworthia with the exception of perhaps H. rossouwii have this very obvious “fish-tail” bud tip. The tips of the upper two petals are flattened and spreading to resemble a fish’s tail. It is most marked in H. herbacea and H. reticulata where the bud tip curves downward from the semi-erect position of the body of the flower and then up again. I am fearful that my inability and reluctance to consider these plants as “new” may be the motive for someone to rush forward and apply a Latin binomial.

My opinion is that these two populations mirror the paradox in the east where H. pygmaea ‘fusca’ is nearer to H. retusa ‘retusa’ andwhere H. retusa ‘turgida’ is nearby. H. mirabilis ‘splendens’, in the doubt about its affinities, in my opinion remains in the context of H. mirabilis. This is a conclusion I reached after exploration of the area between Riversdale and Albertinia where the same elements noted in this article repeat themselves. I do not for one moment concede that better explanation is available by resort to a multitude of Latin binomials and by an artifact like “aggregate”.

Conclusion
I have been deeply disappointed that so little progress on the classification front has been made among the fraternity writing about Haworthia. In communication with Lawrence Loucka, he wrote …”You might consider saying that a genus revision is needed and that until amateurs depart no proper botanist will make the investment. Until then chaos will rule, names will proliferate.” This is a useful and insightful comment and generally summarizes what I have often said. Unfortunately the position has been rather exacerbated when proper botanists have made their presence known and I can recount many instances to support this statement. My own view has been that botanists have not been able to confront genera that are so publicly and popularly known, and garlanded with names. Thus amateurs, including myself despite at least an iota of formal training in science, simply have a field day generating names like rabbits out of a hat.

Unless some proper thought is given to this whole question of what the names are supposed to mean apart from catalogue items, taxonomy (classification) will never lift out of the disinterest, disrepute and scorn that collectors justifiably seem to have for it. Latin binomials do have a real value and to demean this by one-dimensional derivation and rationalization is intellectually fraudulent. My writing is replete with examples of the variability and interaction of populations over large area. Taking a single population and one flower, and often a single plant, to generate a species description and a Latin binomial is grossly simplistic.

The one change that this article generates is the confirmation that the population MBB6666 from immediately south of Tradouw Pass is better identified by name as an H. mirabilis variant, and not as H. retusa ‘nigra’. MBB7801 at Buffeljags and now also at Rotterdam, remains as H. mutica and the epithet generated by enthusiasts can be added to make the trinomial H. mutica ‘groenewaldii’. For formality, the name for the form illustrated as typical would be H. mutica ‘Groenewaldii’. A few additional names will be needed for similar clones and a lot more for the cultivars that will be derived from this population set.

My conclusions here make no substantial departure from anything that I have observed or written over 50 years. While the subject has been comprehensively covered in the field and in writing, there is a huge amount of exploration that still needs to be done if the genus is to be still better known. My prediction is that the answer will conform very closely to the species-list I finalized in 2009 (see Alsterworthia 5.1:192). The disclaimer is that botanical classification may eventually reach the point where the real nature of species and their purpose comes to be known. In that case what we now may see and persuade ourselves are discrete species, are not. Instead these highly complex chaotic systems may in fact be quantum units of consciousness and whole elements such as some genera and sub-genera we now recognize will be the species.

Acknowledgement
I want particularly to acknowledge Jannie Groenewald whose interest and enthusiasm are boundless. Then Allan Jeptha, an environmental law practitioner with great vision for the Suurbraak community; Odette Curtiss, a consultant for the conservation of Renosterveld vegetation; Lawrence Loucka whose patient guidance and comments I value and who generated the set of pie charts; Kobus Venter and Steven Hammer who have always been supportive and contributory; Max Coetzee of Felix Unite and Samie Lategan of Buffalo Breeding Programme for access to Sandkraal West and Crodini East; Jaap Viljoen for his contribution; my wife Daphne, who trusts me and whom I deeply trust. ♦

Volume 7, Chapter 2:- Further exploration in Haworthia – further to finale

The writing of my grand finale was inspired by several things. One of these was another item of a mind-numbing foray into the classification of Haworthia. So I asked that deep thinker and observer, Gerhard Marx, for a devil’s advocate (abbrev. DA) point of view which he has done with the same competence he has as an artist. I have many times in my writing addressed the issue of a species definition and produced one too. Not surprisingly the first thing the DA does is dismiss my definition without producing one of his own. Simply being able to say that an indeterminate number of plants from some population are sufficiently different in respect of a character or two from other haworthia, is motivation enough for the generation of a new name?

The case of H. groenewaldii Breuer, described in an article authored in Alsterworthia 11.2:13-17 by Breuer, Marx and Groenewald is the case in point. It presents the description of this supposed new species from Buffeljags east of Swellendam. The article is written in the first person (Breuer) who quotes extensively from Gerhard’s e-mails, and includes a piece by Jannie Groenewald under the heading “Description of the Vegetation type and distribution”. The overall impression is of an article that conforms to the style of a forgotten era and it is not possible or sensible to attempt a rational dismissal.  Who is actually responsible for the article and how does one correct misleading statements without giving offence?

There is probably only one main issue that can be made level-headed sense of. This is the statement in the article…”another (it is not clear what the other was) striking feature was the fine but rough leaf texture…”  Then quoted again…”the typical H. mutica grows only 20km to the west at Dankbaar”. But the Dankbaar plants also have the same shiny leaf surface. The word “typical” is generally overused in the article and I would counter claim that the Dankbaar plants are typically H. mutica and reinforce my argument for the similarity to the Buffeljags plants. But this is not the main issue, nor is it the least of many rectifications the article almost demands. In the new-age freedom of communication there are virtually no secrets anymore and this adds to my discomfort because the article only touches on some of the issues clouding Haworthia classification. The DA says I should meet this following argument in support of an H. groenewaldi’…” it is consistently smaller than H. mutica and reminds superficially actually of H. bruynsii or springbokvlakensis with its rounded leaves. Kobus Venter’s feeling was that it is closest to H. magnifica var atrofusca and I agree 100%.” These are subjective statements and I cannot agree about the size as I have seen many H. mutica populations that I do not think supported the opinion. The roundness of the leaves is also subjective. The element H. mirabilis ’atrofusca’ in any case has plants with variably rounded leaves, so which do you mean?.  The name ‘mutica’ means leaves without a point and rounded and it is no co-incidence that Col. Scott initially identified H. springbokvlakensis as H. mutica until I pointed out to him that I thought H. mutica was actually the plants (note plural) in the Bredasdorp area. Scott plagiarized my words describing them alongside old wagon tracks that suggested their discovery in earlier times.

DA lists several more indeterminates…”all the unique morphological features like small size, rounded rough-textured and uniquely flecked leaves”.  Of course there are not several more as I have already dismissed two and this leaves the third viz. the “uniquely flecked leaves”. Firstly not all the plants have flecked leaves and I have shown that these occur in H. mutica at Klipport too. The main issue is flowering time and I agree this is problematic.  But this is actually so in respect of the whole Southern Cape Haworthia complex where it is quite evident that there are two flowering periods, summer and winter and that despite this difference there is hybridization. What happened in the past?

DA writes…”MBB 7801 flowers…with the typical thin tender peduncles and delicate flowers identical to that of H. maraisii and some H. magnifica and H. mirabilis. Normal H. mutica flowers in spring (Sept-Oct) and has more robust peduncles and flowers.” This word “typical” again, does not convey the truthThere are no floral characters by which one can separate the three “species” the sentence suggests. I have also seen small H. mutica  (Crodini) with very small weak inflorescences and comment on this as a general variable.  See fig. 149, a recent picture of peduncles of H. mutica from Grootvlakte, Riviersonderend. There is a population of H. mirabilis (I presume Breuer and Marx would use the name H. magnifica var. magnifica) from south Riversdale MBB6651. The plants are very small and cryptic and when I collected seed it was from small slender inflorescences with 3-4 seed capsules. In cultivation I raised a large number of plants that were all different and some were huge. The inflorescences were robust as well with 15 -20 capsules.  So this reference to floral characters is simply a loose statement and I do not believe that a proper overview of the flowers in H. retusa and H. mirabilis will reveal any dramatic differences. I have taken good note of flowers and in particular the strange fish-tail bud-tip that characterizes the larger set of populations in the southern Cape.

DA states that I… ”made several blatant errors by ignoring the importance of flower features in his studies”. It this one supposed error or several? I have not ignored flowers at all and it is a blatant error to say so. I simply did not and still do not find them useful. It is fraudulent to suggest otherwise. That fish-tail bud is highly significant and I made a great effort to quantify the differences between H. reticulata and H. herbacea. That the attempt failed was only because the sample required to get to statistical difference became too large. Nevertheless I then tackled a good number of H. mirabilis populations that also have the same bud tip. I concluded that there was a difference between the populations south of the Langeberg and those north and then had to discard that as erroneous because the same problem of variability occurred as with the rosette of leaves. Even the general flowering times I gave for the area was quite wrong – flowering time ranged from November to late April. It is odd that the fish-tail bud tip does actually influence the tips of the upper tepals in that the margins tend to be flattened together. But there is a huge difference in that in H. herbacea and H. reticulata the upper tepals are held far out from the mid-axis (hence ‘subregularis’ or ‘cowboy-hat’) while in H. mirabilis the tepals may meet above the middle upper tepal.

More by the DA… ”So, only careful observation and comparison of diagnostic features, starting with flowers, is the solution.” This is a weak sop because it is something that I started to do and found quite unhelpful. The history is that the small aloid genera were virtually recognized by the fact that the flowers were small. The differences between the subgenera were dismissed by Dr L.A. Codd with the statement that “small differences” were being ignored. DA overlooks a single major binding factor between H. retusa and H. mirabilis viz. bud similarity. I have also discussed in detail the whole question of diagnostic features and went to the genus Oxalis to substantiate my observations that small detail is not helpful.  It comes back to the issue of what is a species? I maintain that they are not these artificial elements circumscribed by morphological detail. They are systems set in a geographical framework. The species, whatever they are, have surely not arisen from independent sources. My field work is very extensive and I have shown over and over again that the populations are linked so that when one finds a single population like at Buffeljags, one asks how it fits. It is not any misconstrual by G.G. Smith that a sample from a population south of Heidelberg was likened to H. mutica. In his time it was not known where H. mutica even occurred and hence he made a mistake in describing H. otzenii.  DA is defending a methodology that failed miserably and is not going to provide any solutions in the future.

Well, well, well one might say. The DA maintains that Bayer has lost his way, to support an argument that he had stopped “doing taxonomy”. DA is not strong on logic but it is intended that way and neither are the DA’s observations necessarily correct.  DA writes this … “Bruce’s dilemma is that he spent a lifetime and hundreds of thousands of Rands doing his research and while he did indeed do an invaluable contribution, he is now faced with the fact that he can’t do that full and neat integration of it all because 1) there is more variety in the wild than what his dispensation recognizes and he feels overwhelmed by the idea of having to try and do a real truth-reflecting revision at this ripe age and 2) he made several blatant errors by ignoring the importance of flower features in his studies.” Point two I have already dismissed, but DA cannot be serious when he suggests that there is some kind of dilemma involving  money and time spent. This is an issue that I will have to address when I meet the Angel of Death.

I do recognize that there is great ‘variety in the wild” quite contrary to DA’s claim that here is more variety than what my dispensation recognizes. The main gist of my writing has been this extraordinary variation. I am not overwhelmed by it at all and I maintain that my dispensation is the only way to rationally express it. To say it confuses me is not an overstatement because it is clarification that I have been seeking. Now Steven Hammer puts it very well… ”it’s almost as if you are being blamed for nature’s complexities” and I said the same thing in different words long ago. The Latin binomial system fails and I have argued and shown evidence for the need of a new and different model   What is more I have proposed such a model too. Anyone who contemplates at any age, the “truth reflecting revision” the DA demands will be faced with a daunting task of citing the synonymy of all the species described or recognized by Breuer and Hayashi (let alone cite all the collecting records). That alone could occupy a lifetime. Not only that, but to also cover the usages and references by the many authors who have written about the plants would be a humungous task. I suspect that DA is implying that I have become senile with a “ripe old-age” comment. My response is that at least my senility was preceded by some degree of clarity in contrast to what I am forced to conclude about contemporary Haworthia taxonomists! I see no reason whatsoever to depart from a logical argument about species definition and the significance of geographical distribution from my species list of February 2009.  Effectively this is a revision and it just does not have literature and specimen citations that meet the needs of “a full and neat integration”. Let me state categorically that there is not the slightest hope of such a pie-in-sky solution.

So we ask Lawrence Loucka’s question… ”How best to separate facts and knowledge from personalities and ego?”  First present the facts. I want to add to the information that I presented in my article on “Haworthia retusa ‘nigra’A grand finale”. There I discuss and illustrate plants from the wider general area of Buffeljags and especially from the Tradouw area to the northeast. Covered are four populations from the farm Heuningklip, and three from the farm Rietkuil. Here I want to discuss and illustrate eight more populations from the farm Van Reenens Crest that lies between. The gist of my argument is that classification of Haworthia is far beyond this nonsensical and simplistic generation of new names in exactly the same way that a miasma of names was generated in the years before 1948.

It is not just chance that the area I have recently been exploring has been so neglected for so long.  It is a high-lying and fairly featureless area and very grassy in comparison to the Karoid Broken Veld, Karoid Valley Bushveld remnants, and the rocky hills, streams and river beds of the areas north east and west. The vegetation is a grassy Renosterveld and the species diversity seems extraordinarily low. There is not much there to attract a succulent plant enthusiast and explorer compared to far more enticing and rewarding countryside elsewhere. My exploration there is a direct result of my having exhausted most other options available to me. An added consideration is that of prediction when long experience has proven that connections between populations exist and need to be found.

So it is not surprising that we found 8 new populations, 6 in the context of H. mirabilis as I circumscribe it, and 2 for H. retusa ‘nigra’ in the same vein of personal circumscription. These populations were all new to me and very exciting. Nevertheless I would not dream of seeing them as new species anymore than I did for H. mutica ‘Buffeljags’. The Van Reenen’s Crest populations fit a pattern and need to be considered in the context of the bracketing populations at Heuningklip and Rietkuil discussed in the preceding chapter. The collections and illustrations are mapped in an accompanying map and are identified as follows:-

The populations.

Set 1 –  H. mirabilis.
The plants in the following six populations are collectively different from those at either Rietkuil or Heuningklip. Using the erroneous or absent logic that characterizes the description of “H. groenewaldii”, there is no doubt that at least one new Latin binomial should be generated. Far from being confused by the huge variation I have seen in Haworthia and guided by a systems view of species, I see these as being representative of one species. It would be useful if proponents of some other dispensation take these pictures individually and arrange them according to the characters they so diligently use to denote their many different species. The question of flower character I laugh off because I have tried to use it and know it will fail. I will come to flowering time again. Plant size is barely significant because in such clay soils the plants do not grow to the size that they will in cultivation. Generally the plants in these populations fall in the category 35mm to 800mm diam. But it is difficult to gauge age and the direct effects of niche favourability that will favour growth and size.

1. MBB7914 H. mirabilis, Van Reenens Crest See figs 1-19.
This was a small population as can be judged from fig.1 where the pressure burst and exposed white clay area barely exceeds 100sq m. The plants were only on the perimeter of the area. This is what makes it quite difficult to assess what the population structure may have been in the era before farming and cultivation. Very seldom indeed have I found plants anywhere in pristine arable or even semi-arable landscapes. Fig. 2 is a view looking across to Heuningklip and the locality of MBB7896 H. retusa ‘nigra’.

2. MBB7915 H. mirabilis, Van Reenens Crest See figs 20-27.
Fig. 20 is a view looking south to a very large area that we did not explore because we chose instead a similar valley immediately beyond the one pictured. The MBB7915 locality was rather odd.  In fact the whole exercise was in the respect that we kept being reminded how similar the situation was to Kiewietsvlakte where H. mirabilis and H. retusa play out a similar show.  In this case there was no actual pressure burst. In this case there was less white clay and there were the characteristic platform-like ridges as if made by grazing animals. There were few plants and while rather scattered, were still confined to what I can only describe as low-biomass bearing area i.e. shallow gravelly soil that does not support dense vegetative cover.

3. MBB7916 H. mirabilis, Van Reenens Crest See figs 28-48.
Fig. 28 is a view of the white clay in an area very like that pictured in Fig. 20. Very heavily utilized by grazing animals, it is surprising that there were so many plants. The variability is quite evident from the pictures and some plants stand out as very different.

4. MBB7917 H. mirabilis, Van Reenens Crest See figs 49-70.
This was a dramatic population because the plants were not quite where we expected them to be. Fig. 49 shows a more typical habitat in the distance. Here the plants were rather in a very brown gravelly situation and absent a short distance away where there was more white clay. As for all these populations there was a wide range of surface textures and also leaf shapes and endings. Often the leaves had a very distinct mucro (end-awn or point). Equally often the leaves were without any point and rounded and this was not true for all the leaves on any single plant. It is very problematic to make statements about the plants because when we were looking at Rietkuil plants we were reminded of a host of more easterly populations and plants. In this case even variants of H. emelyae came to mind.

5. MBB7918 H. mirabilis, Van Reenens Crest See figs 71-87.
This population was unusual in that the plants were generally small and proliferous.  Sometimes there were as many as 20 rosettes in a clump.

6. MBB7919 H. mirabilis, Van Reenens Crest See figs 88-116.
One needs to refer to the map to see how the populations now relate to one another.  These were again very variable as I have come to expect in the whole H. retusa/H. mirabilis collective – to coin yet another adjective to do what I do not think the formal system allows. Note carefully fig.100 in which a young bud is clearly visible.

Set 2 – H. retusa ‘nigra’
We found just these two populations.

7. MBB7920 H. retusa ‘nigra’, Van Reenens Crest See figs 117-137.
These plants were in full flower and setting seed as one expects for the summer-flowering H. retusa elements. In fig. 128 a plant is shown with well-developed end-awn and fig. 134 a plant with rounded leaf tips. Fig. 137 is simply a view of the flower that does not even depict the more normal fish-tail bud tip – see next!

8. MBB7921 H. retusa ‘nigra’, Van Reenens Crest See figs 138-148.
Fig. 138 is a plant that could have come from an H. retusa ‘retusa’ population from around Riversdale, while Fig. 146 could have come from H. retusa ‘turgida’ from somewhere around Heidelberg. Fig. 147 clearly shows what I mean by the fish-tail buds. Fig. 148 shows that there has been quite extensive exploitation of the white clay (kaolin). Fortunately (?) unlike the bentonite that is mined on large scale, the kaolin pockets are often small and restricted and apparently not commercially viable.

Conclusion
The plants in the above populations cannot be categorized without reference to the totality of populations and I will try and summarize my point of view. These populations occupy a position very central to the distribution of both H. retusa and H. mirabilis. I need to quote DA again who wrote … ”His (Bayer’s) emphasis is upon phylogeny and understanding the evolutionary processes at work in Haworthia and he loves getting involved with very complicated arguments and theories which (frustratingly or delightedly) to him nobody quite follows or is really interested to follow.”

DA is most fortunate to be able to dismiss the basis of classification and what all biologists involved in classification try to do. It is quite wounding to say that I enjoy this theorizing. I would be far happier if there was no contradiction and I could wander out in to the veld and be assured that there were smart neat boxes and simple ways of filling them from a manageable list of names and unequivocal identifying characters. My observation is that all this so-called “taxonomy” practiced by too many writers is so flawed because the very basis of biological variation is wholly ignored and forgotten. It does not take rocket science to imagine that science was supposed to set us free from superstition and blind belief and allow us to individually question the nature of creation.

Here we have a real problem that is overridden by a basic problem in formal classification.  If I could start now afresh with a clean slate and knowing what I now do, I would still be handicapped by the restrictions of the nomenclatural code. This is that I do not believe that the name H. mirabilis as typified by a plant that comes from an outlying population truly represents the species. Perhaps this is a point not understood by the collector community, and in fact even by botanists in other disciplines. The “type” is only there to serve as a reference point for the NAME and may not be remotely typical of the species that this name is applied to. What is unfortunately true as with the crop of taxonomists who have muddied the classification of Haworthia in more ways than one, is that more often than not the application of names is not to species at all but to an odd assortment of plants that meet no criterion for a basic unit of life forms. In the case of the name and type of H. retusa there has been curiously no problem and the vast variation there is forgotten simply because of familiarity. The interesting thing is that the name H. mirabilis was not even in use before I resurrected it in about 1976, so anything that remotely departs from an unfamiliar norm must be “new”? For H. retusa ‘nigra’, I must have got the name ‘nigra’ from the old literature and reference to the Kransriviermond collection that Smith informally listed as H. mutica. The point is that in my acknowledged ignorance I attached the name to H. mutica because of the absence of an end-awn and a relatively rounded leaf tip to the plants seen. (Looking back to all my pictures I see that in some plants there is definitely an end-awn.) The dark-colour has resulted in a name that is not appropriate for the populations that can be considered to constitute the taxon. Neither is the population at a geographic end-point and as a probable hybrid entity representative. This statement also only scratches at the surface of a real problem of origins and changing faces of the plants with time.

Reflecting about relationship as a taxonomist is required to do, and with due respect to DA and the weird views of my preoccupation with relationships the invitation to him required; it seems to me that the unseen and unknown element, now only evident (H. retusa ‘nigra’) as a widespread and common element in this central area, is important.  It may be the precursor to the real way in which H. retusa and H. mirabilis have developed in close synchrony and association in both easterly and westerly directions.

The really strange occurrences are evident from the map herewith (fig. 150). The two species do not grow in direct close association in the area under consideration. There are places where they do e.g. Skeiding about 20km eastwards; KomseRante, Riversdale; Soetmelksrivier, E Riversdale. Why is this? There is no observable difference in habitat or associated vegetation, and still there is pattern in the distribution.  It appears to me that they are separated by flowering time (that does not preclude hybridization as field hybrids show) and in character by colour and a tendency to greater surface roughness of various kind and degree in H. mirabilis. The mystery deepens when H. floribunda appears in two places but again as sole occupant of habitat seemingly favourable to either H. retusa or H. mirabilis. Pattern is broken here because the evidence seems to be absorption into H. mirabilis in other places where the two species do closely meet. The conclusion has to be that I have come no nearer to a neat solution. The H. retusa ‘nigra’ populations cited above will attract no takers, but the H.mirabilis populations will. I create no new name because my personal view of Haworthia does not call for one.

Acknowledgement
I particularly want to thank Gerhard Marx for his considered points of view he has crafted as my Devils Advocate. He is an extraordinary grower and observer, and an artist of the highest order. He had driven me to more carefully and thoroughly explore and document what I see and think. I am extremely grateful to him. Lawrence Loucka has been a great help as an advisor, and so has Steven who brings something special to the whole field in the way he “communicates” with plants and about them. Kobus Venter is ever supportive. Access to Van Reenens Crest was through kindness of Trevenan Barry unsoured by the pressures of harvest time. ♦

Volume 7, Chapter 3:- A field trip to the Potberg area

Introduction

These field trips, to the Potberg this time, are always made with some objective in mind in respect of new exploration. In this case I wanted to get more pictures of H. mutica as it is a species that I have few digital images of. There were also localities that I remembered from the days when I was sweeping the countryside at a fairly coarse scale and was not much bothered by detail. I confidently expected the number of real species conforming to that in other fields of botany and zoology, to be in the region of 33. I never dreamed that such divergent views would, or even could, arise from less information than even then available to me. So while 450 names were whittled down to the mid-hundreds by me, students of the genus have in recent years pushed that up to the 600 mark. My opinions have been couched in quite conservative terms but it is a problem of the nomenclatural system that an identification in respect of a Latin name evokes a reality that does not exist. I maintain that the problems we experience in Haworthia are no different to that which exists in many animal and plant genera. I think that primarily this is because of the absence of insight into, and understanding of, the actual nature of species and the two dimensional model we use to relate them. Species are very variable systems because they have to be to survive the constantly changing world they occupy. In this article I am just going to present images of plants within populations of four different species viz. H. variegata, H. minima, H. mirabilis and H. mutica.

Set 1  H. variegata
H. variegata has an odd distribution in that it is primarily in the lower Goukou River drainage system southeast of Riversdale.  Plants of H. monticola from the Uniondale and Willowmore areas, and even those of H. angustifolia from the Eastern Cape, can be very similar. So the distribution south and west of the lower Breede River is quite remarkable. Two populations, one high on the western end of the Potberg Mountain and another northwest of this at Luiperd’s Kop are very similar to those in the Goukou area. Chris Burgers of Cape Nature brought me plants from the lower north slopes of the Potberg that were quite different. Small single plants with outcurving leaf ends barely protruding above the pebbles the plants were growing in. I named this H. variegata ‘hemicrypta’ but not without reservation in the same way as I also named the plants in another population on the Potberg and at Kathoek as H. variegata ‘modesta’.

Chris Burgers did take me to his site many years ago and it is one of those places I cannot recall in the pebbly flats and Fynbos vegetation of that area. So the finds here seem to represent that same element but with subtle differences in respect mainly of variegation and spination of the leaf margins. I have mentioned elsewhere that there is an interaction with an element I refer to as H. floribunda although it is not clear if this indeed that species, or if it is an emergent polytopic element of H. mirabilis in consideration of the variation and distribution of these two apparently different species. We may have a truth for one area that is different from a truth in another. Polytopic means similar looking things arising in different places from a common genetic pool.

1. Map of the Diepkloof area (fig.1)

2. MBB7923 H. variegata  S Diepkloof. (figs 2)

3. MBB7924 H. variegata SE Diepkloof (figs 3)

4. MBB7925 H. variegata  SE Diepkloof (figs 4)

5. MBB7927 H. variegata  SSE Diepkloof (figs 5)

6. MBB7928 H. variegata  SSE Diepkloof (figs 6)

7. MBB7515 H. variegata  Stoffelsriver (figs 7)

Applying the available varietal name, I should refer to these populations as H. variegata var. hemicrypta. However, I think this implies fixity of opinion and a certainty or reality that are neither not strictly true. I have added the Stoffelsriver two pictures just as a reference because while the first 5 populations are associated with recent Tertiary deposits and Renosterveld vegetation, the last is in sandstone gravels and in Fynbos.

Set 2  H. minima
This species of course belongs with three others in the Robustipedunculares and in my opinion more closely related to Astroloba than to Haworthia. It is widespread and even occurs in the Little Karoo. None of these populations are significantly different from the general run for the species, but we were awed by the beauty of the plants in their natural settings where exposure results in compactness and beautiful colouration. They can occur in quite different environments and habitats. There are a few aberrant populations and hybrids with other members of the sub-genus.

8. MBB7496a H. minima Diepkloof S (figs 8)

9. MBB7926 H. minima SE Diepkloof (fig. 9)

10. MBB7929 H. minima SE Diepkloof S (figs 10)

11. MBB7930 H. minima S Diepkloof (figs 11)

12. MBB7932 H. minima S Kleinberg (figs 12)

Set 3  H. mirabilis 
H. mirablis is an extremely complex group of many populations and plants and distributed from Albertinia in the east to near Caledon in the west, and from the foothills of the Langeberg to the coast. The relationship with H. retusa hangs on the slender thread of flowering time, with H retusa flowering in late spring and H. mirabilis in late summer. The relationship of these two species to H. pygmaea in the east and H. mutica to the west has been mooted by me elsewhere. A similar scenario can be suggested for the species recognized north of the Langeberg.  Included in the following sets is the population at Die Kop that I surmise has its origins in interaction between H. mutica and H. mirabilis.

There are three populations in these sets (nos 13-15) that are suggestive of H. mirabilis ‘atrofusca’, while others have distinct resemblances to H. floribunda (e.g. 16). It is curious that nearly all the populations of H. mirabilis that occur in the same general geographic area as does H. mutica are of smallish plants. These I originally regarded as a separate species viz. H. heidelbergensis. This should be seen as one of those decisions based on inadequate sampling and knowledge of the field. This phenomenon now suggests to me that H. mutica is occupying middle ground between H. retusa in the ‘turgida’ form and H. mirabilis.

13. MBB7496 H. mirabilis Diepkloof S (figs 13)

14. MBB7497 H. mirabilis S S Diepkloof (figs 14)

15. MBB7933 H. mirabilis  S Diepkloof (figs 15)

16. MBB7922 H. mirabilis Diepkloof (figs 16)
A locality established by Jakub Jelimicky. The similarities to H. floribunda is only apparent in some of the plants where the leaf tips are twisted, slightly flattened and rounded at the ends.

17. MBB7931 H. mirabilis S S Kleinberg (figs 17)

18. MBB7531 H. mirabilis S Kleinberg (figs 18)

19. MBB7935 H. mirabilis Langvlei (figs 19)

20. MBB7938 H. mirabilis SE Stoffelsriver (figs 20)
This locality was recorded when Gerhard Marx and Jakub Jelimicky misunderstood my directions. The plants are incredibly cryptic and very hard to see in a not very well defined habitat. One can only follow my opinion on the classification in the context of 8 other populations in the sandstones and Fynbos, and then relate these to populations in shales, tertiary deposit and transitional vegetation.

21. JDV86/2 H. mirabilis Haarwegskloof (figs 21)
These are small plants in very close association with H. mutica as discussed above and are in quartz outcrops rather than in the shale ridges where H. mutica is found.

Set 4 H. mutica
H. mutica is not a species that is well-known and this is probably because it is not proliferous. The name mutica means without a point and “typically” plants of H. mutica should and do have no point (end-awn). The main object of the trip was to obtain more images of this species to convey a better impression of the many variants and how it can be viewed in relation to other species.

22. MBB7075 H.mutica Grootvlakte (figs 22)
I am not certain of the origins of H. otzenii G G Smith and the locality that I traced was approximately 8km east of Riviersonderend. That site is now very degraded and I could not find plants on a visit many years ago but where I did see them on my first visit there. This Grootvlakte population occupies a similar habitat in an exposed shale ridge south of the Riviersonderend River.

23. MBB7935 H. mutica Crodini (figs 23)
The name Crodini may have been used for the following collection at nearby Witklipkop, and I say this because I did once get seed from there and Jakub Jelimiky informs he has a plant labeled ‘Crodini’. I only saw two plants here at Crodini itself when the site was severely grazed, and we found only 8 plants on this occasion. The plants appear to be in quartz, but this is a bit of an illusion as there are white quartz bands in the shale and the plants prefer the shale.

24. MBB6536 H. mutica Witklipkop (figs 24)
This is not a robust population in shale and also marginal to quartz.

25. MBB7937 H. mutica Platkop, S Napky (figs 25)
The substrate here seems to be silcrete but there is no underlying kaolinite and the area is cultivated, which suggests shale. So that is a bit of an unknown.

26. ADH2729 H. mutica Ouplaas, DeHoop (figs 26)

27. MBB7934 H. mutica Wolwefontein (figs 27)

In early 1970 I took Peter Brandham and David Cutler of Kew to look at Haworthia in the field. In fig.27.12 the view is southward to Kathoek and the cottage in the foreground was occupied. It must have been abandoned soon after and the access road obscured. The result was that the place was later difficult to find, but the plants are still there. It is not far from the problematic Die Kop population to come.

28. JDV85/17 H. mutica Haarwegskloof (figs 28)
This record goes back to 1985. We did not have landowner permission here as we did not know exactly where to go. The area was also unfenced. So we were confronted by the farm manager and never took as many pictures nor explored more widely. It certainly would have been problematic had we not simply being photographing. Anybody else may have ended up at the police station. It is a case of farm land too hilly and rocky to cultivate and bought by apparently wealthy city people with an eye to conservation. This may result in the place being game-fenced, a lodge built for the up-market; and made perhaps even less accessible.

29. MBB6641 H. mutica Hasiesdrift (figs 29 JDV97/26)
This was one of my early encounters in 1969 with Haworthia, where I fortuitously stopped at a river crossing, lifted a branch of a shrublet and saw a magnificent plant just like that illustrated and serving as the type for the name H. mutica (see fig.29.1for an example)). A year later a new road was built and if there had been a population there it was no more. In 1996 when I was writing Haworthia Revisited, I went back there and after a diligent search found a very small group of plants at the base of a small plant of Aloe ferox. The two species seldom grow among each other so this really was odd. I have periodically been back to see what was happening because there were many Aloe seedlings. The very small population has waxed and waned and the mothering Aloe seedling has died. There has been a small spread of plants to occupy an area of about 2 square meters. I know here are more plants about 2km away but have not explored the wider area more thoroughly.

30 MBB7500 H.muticaXmirabilis Die Kop (figs 30)
So we come to the problem child that has acquired the species Latin binomial H. hammeri. I think this is unfortunate as Steven Hammer deserves a great deal better than this. It should not be necessary to have to also manage personal issues in an inquiry into what the name means or if it is a valid statement in science. This population is one I found when I was exploring widely for an understanding of H. mirabilis. It was not growing in conventional habitat and populations occur at many places in the wider vicinity. So I do not comprehend at all why this single population with absolutely nothing to support its designation as a species in respect of anything but the fact that the plants look different from something else. The plants are also very variable as the set of pictures should show, and the plant suggested by the picture accompanying the description is not typical. The proximity of H. mutica and the nuances of these plants with this species and also with that of other nearby populations of H. mirabilis, suggest to me that this population is of hybrid origin. I also use the term hybrid in the sense of genetic similarity because hybridity itself implies separate existence. My feeling is that H. mirabilis and H. retusa have separated in time from a common gene pool and that the process is not complete and nor will it be. My species definition requires that we see all these populations as one system that is spread in geographic space and is changing with time. The rate of change is a fourth dimension.

I suppose one should comment on the shiny leaf surface (fig. 30.3 especially) that is present in H. mutica, as this character is also described as a diagnostic for H. groenewaldii that is as contentious as H. hammeri.


Conclusion
The objectives of the excursion were fully met and I am sure these pictures will contribute to a better view of how these four species vary. Perhaps this will illustrate also the illogic of Latin binomials for every variable, however desirable these names are for traders and collectors.

Acknowledgement
Bennie Viljoen of Grootvlakte, Koos Badenhorst of Langvlei, Maree and Elizabeth Prinsloo of Stoffelsriver, Ferdie of Diepkloof Security Services, Adele and Vlooi DuToit for Die Kop, Martin Mijnhardt of Haarwegskloof, Neil Giliomee of Hasiesdrift. In some cases we simply could not track landowners and trust they will forgive us our trespasses remembering that invasion of ones territory is close to an unforgivable sin. ♦

Volume 7, Chapter 4:- What is typical Haworthia mutica?

A supposed new species of Haworthia viz. H. groenewaldii Breuer, is described in an article authored in Alsterworthia 11.2:13-17 by Breuer, Marx and Groenewald. It presents the description of this supposed new species from Buffeljags east of Swellendam that I would simply have identified as another variant of H. mutica. This is not because I am confounded by the variability among the plants in the genus, or even in any one species. I recognize the species as systems of populations in which the individuals vary from one another as one would expect in any group of living things that maintains the flexibility to adapt to constantly changing world conditions. In this even time becomes a variable. I think species are very important elements if the whole of creation and not just for taxonomist and collector activity. Other people have other ideas of what species are, so my disagreement is hopefully forgivable..

Although H. mutica was described by Haworth in 1821 it was not allied to a South African field population until recognized by Col. Scott in 1985. G.G. Smith had failed to recognize it when he described his H. otzenii in 1945. The type by which the name is supported is a Kew illustration reproduced here as Fig. 1. This then is what one would expect a typical representative of the species to look alike. Now the ever present problem in Haworthia, is that no two plants in a population may look quite the same. Hence my problem with the description in which the word “typical” is rather bandied about. It falls into the first aspect of taxonomy.

Firstly a plant is illustrated on the front cover of the respective Alsterworthia that, presumably the authors, state is a typical specimen of H. groenewaldii. Secondly, Marx is quoted as saying that the “typical H. mutica” grows only 20km west at the farm Dankbaar. Statements like these are used to strengthen and support opinions and generate a reality that Latin binomials sadly lack. It so happens that I know both these populations quite well and these statements are news to me. I do not think the specimen on the front cover is by any means typical of the population at Buffeljags, and certainly not at three sites recently discovered nearby. The plants at Dankbaar also do not in my opinion fully meet the imputed similarity to fig.1. See fig 2. for an image of a plant representative of the Dankbaar population. I would be very hesitant to say that this is typical of Dankbaar plants.

There is a curious problem here in that Scott does not use the type illustration in his Revision and does not state any origin of the plant he uses to illustrate the species i.e. H. mutica. As far as I am aware the type of a synonym that G.G. Smith described viz. H. otzenii, came from east of Riviersonderend, but this is for an Otzen collection no. 6.  The type was cited by Scott as Otzen 10 but this is not in the Compton herbarium where it should be. So I am not sure offhand where that came from. However, this is not really relevant to the discussion. I just want to state that finding a plant that matches the type is no mean feat and that it was by sheer chance that in 1969 I came across a population of plants at Hasiesdrift that did. See fig. 4 and 5. I selected one image and then realized that it did not have round enough leaf tips to meet need, so I selected another. In the first picture the leaves tend to have a “mucro” – a small point to the leaf that looks different to a well developed end-awn (bristle) that the leaves can also have.

Another issue is that Gerhard Marx once argued with me that the mooted H. groenewaldii was much nearer to H. mirabilis than to H. mutica as I had suggested. What he luckily is able to ignore is my observation of the similarity of some plants of H. mutica to H. mirabilis see figs 5 and 6. I never saw a plant quite like fig. 5 in all my exploration at Buffeljags but Jannie Groenewald collected this one there. Fig. 6 is not typical of the population either and I used this same figure somewhere else in my writing to comment on the similarity to H. mirabilis ‘badia’ variants at Sandfontein (east of the “typical”). Gerhard is still more fortunate to be able to ignore the similarity of H. mutica to H. retusa see figs 7 and 8. I even surprised myself when in looking for a suitable picture, from many, I picked this fig. 7 and find it is also Hasiesdrift albeit a cultivated version. (There is such an interesting story around the Hasiesdrift site). Fig. 8 is a representive of H. retusa from Pienaarsriver pictures and I feel sure that readers will agree that the names could be switched. It was very difficult to ignore pictures from Pienaarsriver that I could have used with figs 5 and 6 in the context of H. mirabilis.

It becomes still more interesting (I would have said complicated but my critics maintain that this variation confuses me and it is actually possible to get it all tidy and neat) when one further compares a plant of H. retusa ‘nigra’ ( fig. 9) with both H. mutica (fig. 7); and then H. retusa ‘nigra’ (fig. 10) with H. retusa (fig. 11). The latter is in fact from the population where the variant ‘geraldii’  originates.

A last point I can make is that the leaf flecks said to characterize “H. groenewaldii” do occur in H. mutica at Klipport (see figs 12 & 13). One cannot simply ignore the extraordinary chain of similarities that extends across the entire distribution area producing only a slightly different situation at either extreme.

Floral difference is a great issue that is misused to force an opinion. The flower is extremely difficult to study because the differences across the entire subgenus are so small. There are complications where, as an example, flowers of H. pulchella ‘globifera’ are indistinguishable from those of H. cymbifomis var. incurvula. In the subg. Hexangulares there is an incredible problem where floral differences within species exceeds that between species e.g. a flower of a plant of H. limifolia may more closely resemble that of H. coarctata rather than that of another plant of H. limifolia. It is easy to draw conclusions from small samples of a few flowers from a few populations but one very soon finds that with increasing sample size the most carefully constructed table of differences becomes senseless. Just when is an ever-aging flower on a stalk at the precise same stage of a flower you want to compare it with? How many flowers from how many plants are needed to make a valid statement? Do not forget that the observations must be made on plants on the basis of random selection too. This is a requirement mostly totally ignored when the more serious question of a species difference is being debated. That of course brings us back to this casual use of the word “typical” and its intractability.

The hardest problem to deal with is that of flowering time and on the face of it a winter flowering time versus a summer flowering time can be taken to suggest significant difference. Yet if one considers that a species needs to harbour genetic variation to ensure adaptation to any kind of environmental change and so survival, a different flowering time may be an extremely useful resource. Then we do have the reality of hybrids between species that do flower at these different times. So obviously and self-evidently populations of species may exist that has arisen from such hybridization between plants that flower at different seasons. The capacity of plants and animals to synchronize breeding periodicity is well-known.

I close with my observation that H. mutica is an assorted group of plants that seem to fall in some middle zone between H. mirabilis and H. retusa. Hence in the west we have H. mirabilis/ H. mutica/H. retusa ‘nigra’, while in the east we have H. pygmaea (=H. mirabilis+H.retusa ‘retusa’) and H. retusa ‘turgida’. In the area between there is vast variation of H. mirabilis and H. retusa that get a bit of H. floribunda thrown in too.

So am I confused, or have I confused you? As Steven kindly put it…’it’s almost as if you were being blamed for nature’s complexities’. Of course the ultimate reality is that we each have our own idea of what “species” are, and here I have used my version! ♦

Volume 7, Chapter 5:- Still more Haworthia mutica and Haworthia mirabilis

It so happens that Heidi Hartmann first visited the Karoo Garden more than 35 years ago and it has been very difficult for me to pay attention both to her mesembs and all my other plant interests. In the last few years she has been working on Acrodon. This is a small genus of only 5 to 6 species that occurs in the Southern Cape with much the same distribution and habitat requirements as Haworthia. She had had some second thoughts on a species she had described as Acrodon calcicola and intimated that she needed photographs to show what proves to be detaching fruits (capsules). So off we went to get that northeast of Bredasdorp at Rooivlei. But Nick Helme had about a year before sent me an intriguing picture of a greenish soft looking plant from near the De Hoop Reserve entrance road to the east. I had considered that it might be an equivalent of the H. muticaXmirabilis population at Die Kop (MBB7500) that Ingo Breuer usefully described as H. hammeri. I use the name with great trepidation because to say what is correct usage is difficult. It could pass as a cultivar name, a varietal name or a form name. I am quite sure it has its origins in the interaction of two species and that is what a botanical name should reflect that; thus H. muticaXmirabilis or however else the nomenclaturists may require. So these journeys are never without distractions as Rooivlei itself is a remarkable site. I find that I have few images of the populations of Haworthia that occur there. The product is nearly all pictures/images.

Set 1. MBB7941 H. mutica Ouplaas, Wydgelee (figs 1-45)

This population is just 3km west of the Die Kop MBB7500 population. There are two intervening H. mirabilis populations; one small, one large and scattered, and a third a little to the north of east and more to the north. But that species is well covered in an Update article and my attention here is on H. mutica. Why Nick Helme’s plant was so green and soft-looking is a bit odd because the population of H. mutica there at Ouplaas is quite one of the largest and most dramatic of any that I have seen of the species. Daphne and I spent hours examining, reexamining and exclaiming at the sheer beauty and variation of the plants. There were plants squeezed into cracks and flattened so that one could not see any centre, others in the flat clayish areas between the exposed vertical shale ridges, but mostly in the shale. The site is north-facing and so is very hot and dry in summer. The plants had just finished flowering.

There was one particular image (fig. 24) I have used to show five different leaf-end shapes on the same plant.

Set 2. MBB7943 H. mutica E Klipbankskloof (figs 46-54)

I saw plants and photographed one (MBB7246) south of Klipbankskloof in a valley mapped as Verbrandskloof. There is another farm, also named Klipbankskloof further west where Jakub Jelimicky recorded H. mutica too. I wanted to see that locality and registered the two Klipbankskloof places as the same. Not so. So I intended up exploring further east where we first found this in a west facing ridge. There was no sign of fruits or flower.

Set 3. MBB7945 H. mutica SE Klipbankskloof (figs 55-80)

This was on a north facing slope and at first we were a bit taken aback because the plants we first observed had flowered poorly or some time previously. But that changed as from north facing slope we entered a very minor valley with both northeast and northwest aspects (more to follow). Here there were many plants and all just finishing flowering. There were some outstanding, dramatic plants. I wish I was a more accomplished and better equipped photographer. Some of the images would have been stunning.

Set 4. MBB6638 H. mirabilis Rooivlei (KoeiseKop) (figs 81- 92)

It was fortunate for my story that I took pictures here. As in most of the greater area occupied by H. mutica, H. mirabilis occurs as these smaller, often (how often is often?) dark rough versions of H. mirabilis. Of particular note is the first image (fig. 81). This is of a plant with much smoother textured leaves and had there been and H. mutica in the near vicinity I would surely (mistakenly?) taken this for a hybrid because I know from experience elsewhere that summer and winter flowers can and do hybridize. But wait!  Rooivlei is a unique site because there is a population of H. retusa ‘turgida’ nearby that is prominent in my whole explanation of relationships that my critics so dislike. Outside chance is that hybridisation is an explanation.

Set. 5 MBB7944a H. mirabilis SE Klipbankskloof (figs 93-101)

I have numbered this as ‘a’ and ‘b’ for a reason. The two ‘populations’ are less than 200m apart. In fact the two H. mutica populations are not all that much more distant and there would be some debate as to whether they constitute populations at all. My argument would be that they occupy discrete separated habitats. This mirabilis population falls into middle ground. It is the upper slope where aspect changes from south to north. It is moister and the southern aspect of the small shale ridges provides mirabilis habitat. It is as if H. mirabilis is found where what I can describe is an escape route is present. These dark plants do not readily tolerate a full days exposure to direct sun and I speculate that this is the same kind of thing that happens on north-facing mountain slopes as around MacGregor. Plants are usually where the aspect allows escape from a full days worth of sunlight. The plants are much like at Rooivlei.

[ed.] images mislabeled as 7943, should be 7944.

Set. 6 MBB7944b H. mirabilis SE Klipbankskloof (figs 102-110)

The different number is because the plants are now sharing both habitat and niche with H. mutica. I do not know any other site where this happens. The two species do occur in the close general area but that is then different habitat and the distance between ranges from about 50m (Haarwegskloof) to 400m (Rondeheuwel). What is now fascinating is that some of the plants resemble fig 1 of Rooivlei and we are faced with the probability that there is exchange of genetic material between H. mutica and H. mirabilis. This is what is suggested at Rooivlei too for H. retusa ‘turgida’ and H. mirabilis (see above).


Conclusion
None. This is purely instructive. What I gained from the trip was the realization that H. mutica would be a very good subject by which to explore variability and change with time.  Also to look at conservation and impact of land-use on populations. Over to Jakub Jelimicky.

Acknowledgement
Again a list as long as an arm. Francois Uys for access to Rooivlei. Nick Helme for information in respect of Ouplaas. Odette Curtis for interest in Renosterveld conservation and contact for Nick. Johan Dutoit of Vleitjie and a friendly combine mechanic for access to Ouplaas. Wynand Dutoit of Klipbankskloof for access to what is his wife, Madrẽ’s farm. ♦

Volume 7, Chapter 6:- Field trip to Van Reenens Crest and Niekerkshek

The objective was to explore some likely habitats previously observed at Van Reenens Crest and nearby. We extended the scope to include further exploration for Haworthia mutica as I am still questioning the place of this species in the greater scheme of things. Thus here are four sets of populations that I report on viz. H. mirabilis, H. retusa ‘nigra’, H. floribunda and H. mutica. See maps Figs 1 and 2 for geographical position.

H. mirabilis

Set 1. MBB7955 Van Reenens Crest – mid west. (figs 3-11)

The plants are moderate in size to about 50mm diam. and growing on an erosional slope below tertiary gravels where there is white clay. They vary from densely tubercled to relatively smooth.

Set 2. MBB7956 Van Reenens Crest –  west 7955. (figs 12-34)

Despite apparently favorable habitat these plants were about 300m west of the previous set. Variation is again apparent in respect of presence or absence of surface roughness.  Generally there are plants in which some leaves are without a point (see Fig 1) while others are acute tipped (mucronate – with a short hard point) or even have a long end-awn. Margins may be smooth or very lightly spined. This seems to be more apparent in young plants.

Set 3. MBB7957 Van Reenens Crest – west 7956. (figs 35-41)

Again the plants are very variable with dense surface tubercles to virtually none. The leaf shapes vary and in fig.39 that appears to be a group of seedlings there is a remarkable series of different leaf ends and margins. In some plants there is distinct spination of the leaf margin.

Set 4. MBB7958 Van Reenens Crest – south west 7957. (figs 42-51)

Fig. 43 shows two rosettes of a plant with a “crocodile-skin” surface. The surface tubercles seem to be flattened and the leaves are mucronate. The following photograph, fig.44, is of a plant with acuminate leaves ending in long end-awns, and the surfaces are relatively smooth – retusoid? Unlike the previous records for the area that are north-facing, this is a south-facing “pressure-burst” of white clay. A nearby habitat of vertical shale that might have been considered favorable for H. mirabilis elsewhere was devoid of plants.

Set 5. MBB7959 Van Reenens Crest – west homestead 7956. (figs 52-64)

Figs 53-55 are smallish plants with rounded leaf ends and relatively opaque surfaces.  Fig.58 is remarkably like H. retusa, say, at Pienaarsrivier with very awned leaf ends and a high degree of translucence in the leaves.

Set 6. MBB7960 Kruiskloof, E Van Reenens Crest. (figs 65–78)

This population departs from a very broad norm for Van Reenens Crest and picks up the continuity with populations north, east and south. This is most noticeable in the surface texture where the tubercles are smaller and denser. The leaves can be highly rotate, see fig.68. A feature seen occasionally in the Van Reenens Crest populations and as far afield as the Diepkloof (Malgas) area much further south, is the occasional plant with a very stellate (five –pointed star) arrangement of the leaves that are quite horizontal see fig.70. Among the many oddities is the plant in fig.72 with floribundoid leaf ends and a leaf surface where the tubercles seem to be flattened. In fig.73 the tubercles seem to be spined. In fig.75 tubercles are almost absent.

H. retusa ‘nigra’

Set 7. MBB7961 Kruiskloof, E Van Reenens Crest. (figs 79-90)

These plants reinforce my perception that H. retusa and H. mirabilis are uncomfortably closely related. Figs 79 and 80 have the same general form as H. mirabilis ‘sublineata’.  Fig.82 appears to me as possibly a hybrid as leaf opacity is deeper and there are marginal spines.

H. floribunda

Set 8. MBB7962 Niekerkshek. (figs 91-99)

Set 9. MBB7963 Niekerkshek. (figs 100-111)

The plants vary in size and to some degree in colour. This variation in colour from a deep purple to dark grey is very evident in fig.107. The leaves are much flattened and an occasional leaf is pointed. The plant in fig.102 is unusual in having white spots on the leaves. Marginal spination can be absent or very coarse. The leaf surfaces show no venation or translucens, as is expected in this species.

H. mutica

Set 10. MBB7950 DeDraai Klipbankskloof. (figs 112-123)

Set 11. MBB7951 Klipbankskloof West 1. (figs 124-126)

Set 12. MBB7952 Klipbankskloof West 2. (figs 127-129)

Set 13. MBB7953 Klipbankskloof West 3. (figs 130-144)

Set 14. MBB7954 Rondeheuwel. (figs 145-174)

Set 15. JDV92/64 Sanddrift, Drew. (figs 175-187)

Set 16.  JDV85/17 Haarwegskloof. (figs 188-213)

There is a very wide range of variants among these 7 population sets. Leaf colour and shape vary enormously and it can only be said that generally the leaves are muticate. Almost invariably any one plant has a leaf or leaves that are also mucronate and in some cases even awned. The leaves may be abbreviated and short and squat or drawn out and elongated. Venation is also varied from many to few or only a single vein on the upper leaf surface. These veins may be straight and clearly separated or they may anastomize and link togther in a reticulate pattern. I did include two pictures of flowers, figs 173 and 174 just to indicate that simply taking two flowers at random one can see obvious differences.

Attention can be paid to figs 145 and 146 of Rondeheuwel where I consider these two plants indicate hybridization with H. mirabilis that does occur about 1km to the northwest.  Fig. 147 shows a plant with a rather wrinkled surface.

Conclusion
My opinion is that consideration has to be given to the odd way in which the “species” do not share habitat. It is extremely difficult to rationalize an ‘average’ plant for each population and so establish any norm for proper comparison. The populations vary by degree. It is also clear that if one examines the distribution of the species across the study area, it can be observed that H. mirabilis where it occurs with H. retusa ‘nigra’(at Heuningklip and Kruiskloof in the east), is not the same as plants to the west where H. retusa ‘nigra’ is absent. Consideration has to be given to absorption into H. mirabilis. Why H. floribunda  should make a re-appearance at Rietkuil and Niekerkshek in the west, when everything points to its absorption into H. mirabilis in the east at Skeiding, is a mystery. The fact that at Niekerkshek it also has radically different forms to those found anywhere else adds to the difficulty.

I will conclude by acknowledging that the populations of H. mutica are all summer flowering but I do not think that this is as significant as the authors of “H. groenewaldii” will contend. The Van Reenens Crest complex demonstrated dramatically that H. retusa and H. mirabilis share a very close common history and an equally close contemporary relationship. My conjecture is that H. mutica is linked to that relationship in the same way the complex of populations are that I assign to H. pygmaea. It is painfully obvious that if floral differences that are so easily detected within populations, expecting to find more evidence in the flowers than exists in the vegetative structures across the genera, and especially the subgenus Haworthia is an absolute pipedream. Floral differences between species are minimal and if anything will provide support for a reduction of species rather than add to the. If any small difference is suggested to separate species, then equal weight must be given to similarities that should then unite species. The contention then that flowers are offering characters that I as an author have ignored, while already wholly untrue, is additionally nonsensical.

Acknowledgement
Mr. and Mrs. Trevennan Barry of Van Reenens Crest. Mr. N. Swart of Poststal, Mr. J. duToit of Tarentaal and Wydgelee, Mr. W. duToit of Klipbankskloof, Mr. J. Mathee of Klipbankskloof West, Mr. Vlooi du Toit of Die Kop, Mr. M. Mynhardt of Haarwegskloof, Mr. and Mrs. Nelius Smith of Volmoed, Rondeheuwel, Mr. Adrian Steyn of Wankie Boerdery. Thank you to Kobus Venter and Lawrence Loucka for company and shared observation. ♦