Haworthia Revisited – 60. Haworthia minima

60. Haworthia minima (Ait.) Haw., Syn.Pl.Succ. :92(1812).  Bayer, Cact.Succ.J(U.S.) 43:157(1971).  Bayer :135(1976).  Bayer :81(1982).  Scott :17(1985).  A. margaritifera var. minima (L.) Ait., Hort.Kew. 1:468(1789).  Willd., Spec.Plant. :189(1799).  Haw., Trans.Linn.Soc. :8(1804).  A. margaritifera var. minor (L.) Lam., Encycl. :88(1783).  A. margaritifera (L.) Burm.(f), Prod.Fl.Cap. :10(1768).  A. pumila var. margaritifera beta L., Spec.Pl. :322(1753).  H. granata (Willd.) Haw. Suppl.Pl.Succ. :57(1819).  Haw., Revis. :203(1821).  Apicra granata Willd. Berl.Mag. :269(1811).  A. granata (Willd.) Roem. et Schultes, Syst.Veg. 7:649(1829).  A. granata (Willd.) Salm Dyck, Monogr. 6:t6(1836).  H. margaritifera var. granata (Willd.) Baker, JLinn.Soc.Bot. 43:205(1880).  A. erecta var. laetivirens Salm Dyck, Hort.Dyck. :12((1824).  Type: icon. 20:t16,f18, Dill. Hort.Elth. (1732):  Haworthia minor (Haw.) Duv., Hort.Alenc. :7(1809).  Haw. Syn.Pl.Succ. :92(1812).  Haw. Suppl.Pl.Succ. :53(1819).  A. margaritifera var minor (L.) Ait., Hort.Kew. :468(1789).  Haw., Trans.Linn.Soc. 7:11(1804).  Ker-G., Bot.Mag. :t815(1805).  A. pumila var margaritifera gamma L., Spec.Pl. :322(1753).  A. erecta Salm Dyck, 6:t7(1836)  Type: icon. 20:t16,f17, Dill. Hort.Elth. (1732):  H. margaritifera Haw., Suppl.Pl.Succ. :55(1819).  H. major (Ait.) Duval, Hort.Alenc. :7(1809).  Aloe margaritifera var. major Ait., Hort.Kew. :468(1789).  Haw., Trans.Linn.Soc. :8(1804).  Type: icon. 1:t21, Bradley, Hist.Succ.Pl. 3(1725):  H. erecta Haw. Suppl.Pl.Succ. :56(1819).  Haw., Revis. :55(1821).  H. margaritifera var. erecta (Haw.) Baker, JLinn.Soc.Bot.18:205(1880).  A. erecta (Haw.) Roem. et Schultes :649(1829).  non A. erecta Salm Dyck 6:t7(1836).  A. margaritifera var media DC. Pl.Gr. :57(1799).  A. pumila var. margaritifera gamma L., Spec.Pl. :322(1753).  Type: icon, 21:t11, Comm. Prael.Bot. (1701):  H. brevis Haw., Suppl.Pl.Succ. :57(1819).  A. brevis Roem. et Schultes, Syst.Veg. 7:649(1829).  A. margaritifera var. minor Ker‑G., Bot.Mag. :t.1360(1811).  Type: icon, :t1360, Ker-G., Bot.Mag.:  H. margaritifera var. corallina Baker, JLinn.Soc.Bot. 43:201(1880).  Type: Ex hort. Peacock.  Not preserved:  H. mutabilis V.Poelln., Feddes Repert.Spec.Nov. 44:132(1938).  Type: Cape, Bredasdorp, Payne 23 in Triebn.1110.  Not preserved.

minima: small.

Rosette stemless, slowly proliferous, to 150mm tall.  Leaves to 130 X 15mm, nearly as thick as wide, attenuate, spreading, lanceolate-deltoid, surfaces scabrid with raised flattened non-confluent tubercles, colour bluish-green.  Inflorescence sparsely branched, lax.  Flowers tepals fused, tube straight, lobes abbreviated, veins pinkish.

1982 – H. minima is a small (up to 120mm high), clump forming, blue‑green species, with white tubercles and pink‑tipped florets.  It occurs in Coastal Renosterbos from Bredasdorp eastwards to at least the Gouritz River, and inland to Swellendam and Heidelberg.  Very tuberculate forms are reported in the coastal limestones and a nearly glabrous single plant from Bredasdorp was named as H. mutabilis V.Poelln.  There is no indication of any hybridisation or intergradation with either H. pumila in the west or H. kingiana in the east.  The relationship with H. poellnitziana requires detailed examination.  H. minima hybridises with H. marginata at Heidelberg (Cape).

1999 – The only interesting development concerning this species is the collection from north of the Langeberg Mountains.  It has been found at Brandrivier to the east, and still further east, northeast of Herbertsdale.  H. poellnitziana is now included as a variety of this more generally distributed species simply in recognition of popular opinion.  H. minima does develop long slender leaves, but the fact remains that there seems to be a difference in flower colour, while the distribution is also disjunct.  As is unfortunately usually the case, no herbarium specimens are provided to substantiate opinions.  The synonymy has been substantially altered follow the views expressed concerning the correct naming of H. maxima (Haw.) Duv.  My opinion has been that if the iconotypes had been properly known and understood, the nomenclature of the two species involved would have been much simpler.  H. maxima would have been seen to be the correct name for the “large pearled aloe”, and H. margaritifera for the “lesser pearled aloe”.  The old synonymy follows the types quite closely and unfortunately those who have attempted to pronounce on the nomenclature seem to have been under the impression that the iconotypes were mostly the larger of the species.  The converse is true.

a. var. minima

Distribution: 3321 (Ladismith): Brandrivier (-CC), Laidler 707 (NBG); Perdekop, Bonniedale (-DD), Matthews 1095 (NBG).  3419( Caledon): Wiesdrift (-DB), Smith 5475 (NBG); Voelvlei (-DB), Dymond (BOL); Mierkraal (-DB), Fourcade 199 (NBG)7.  3420 (Bredasdorp): Swellendam (-AA), Rotheno in PRE 39425; Bontebok Park (-AB), Bayer & Fourie 4432 (NBG); NE. Bredasdorp (-AC), Smith 5472 (NBG); Skeiding (-BB), Smith 7220 (NBG); Heidelberg (-BB), Fouche 45 (PRE), Smith 5508, 6569, 7146 (NBG); Bassonskraal (-BC), Burgers 1333 (NBG); Koenskraal (-BC), Venter 4 (NBG); Infanta (-BC), Ross-Frames (BOL); Malgas (-BC), Esterhuysen 5224 (BOL); N. Infanta (-BD), Smith 5474 (NBG); Bontebok Park (-CA), Barker 7232 (NBG); De Mond (-CA), Smith 7352 (NBG); Struisbaai (-CC), Esterhuysen (BOL).  3421 (Riversdale): SW. Riversdale (-AA), Smith 5469 (NBG); Swartheuwel (-AA), Smith 7190 (NBG); Kruisrivier (-AB), Smith 7319 (NBG); About 3km SW. town (-AB), Dekenah 3 (PRE); E. Riversdale (-AB), Smith 5375, 5486, 5757, 6087 (NBG); Vermaaklikheid (-AC), Smith 6108 (NBG); Stilbaai (‑AD), Dekenah 97 (NBG), Smith 5504, 5283 (NBG); E. Albertinia (-BA), van Niekerk 333 (NBG); E. Gouritz Bridge (-BB), Smith 3960, 7354 (NBG); N. Gouritzmond (-BD), Smith 6101, 7518 (NBG).  3422(Mossel Bay): Hartenbos (-AA), Schoeman (NBG).

Inadequately located: Ex hort., Steyn in PRE 8797; Riversdale, Ferguson (BOL), Muir in NBG616/23 (BOL); Kafferkuilsrivier, Bolus (BOL); Bredasdorp, James (BOL).

b. var. poellnitziana (Uitew.) Bayer  Comb.nov.  H. poellnitziana Uitew., Cact.Vetpl. 5:137(1939).  Bayer :146(1976).  Bayer :81(1982).  Type: Cape, Drew, Meiring.  (AMD).

poellnitziana: in honour of Dr. K. von Poellnitz.

1982 – H. poellnitziana occurs on the western extremity of the range for H. pumila, and H. minima does not occur nearer than 30km to the southeast.  H. poellnitziana has slender long leaves up to 180mm long.  They are grey‑green in colour and less blue than in H. minima, while in H. pumila the leaves are dark brownish‑green.  The flower lobes are yellowish in colour.  It is in a winter rainfall area, and grows in fynbos vegetation on old river gravels.

Distribution: 3320 (Montagu): West of Drew, Swellendam (‑CC), Smith 3947 (NBG), Fourcade 185 (NBG), Bayer 154 (NBG); Robertson, van der Merwe 225 (BOL).

Volume 5, Chapter 7:- Haworthia minima

There is not much to write about this less glamorous of plants. It is one of the Robustipedunculares.  It is very widely distributed and even occurs north of the Langeberg Mountains in the Little Karoo – as does its sister species H. pumila. The pictures cover a huge range of variants from DeMond in the southwestern area to Riversdale. I found them all fascinating. Mostly so perhaps the very few large globose plants we saw at Kleinberg near Malgas (Diepkloof). But the plants at Koppies are really interesting because they were overlooked for so long. Koppies is the second known home of H. serrata (now H. rossouwii) and we came to know it also as a refuge for a small population of H. marginata. It was while we were investigating this that Hennie van Deventer casually pointed to plants of H. attenuata var caespitosa in his garden and said they were also on the farm. Because of our surprise and of course doubt, he took us to show the plants that he said he had not looked at nor thought of for 20 years – and there they were. A very small population of large dark coloured plants.

Also interesting were the very small solitary plants on the farm Sandfontein in the Slang River Valley. At Klipheuwel marginal to the coastal calcretes we also came across the species. Initially we saw only two plants while searching for ‘retusa-like’ plants. Failing to find anything else in a very severly grazed field we thought we would return to commiserate with the few H. minima we saw and enjoy the ambience we always feel in the presence of Haworthia. One can barely explain how cryptic these plants are and how easily they avoid detection. We found another 40 plants in the same area abut 10m diameter where we had found but two a short while earlier.

I found it very curious that in our very extensive exploration of the area east of Riversdale, that we saw neither H. minima nor H. marginataGasteria was also conspicuous by its absence and I remark on this because I do not think that taxonomy can remotely afford to ignore fundamental patterns in the general environment. ♦

MBB7737 H. minima, Koppies.

MBB7747 H. minima, Sandfontein.

MBB7757 H. minima, Klipheuwel.

MBBsn H. minima, Bontebok National Park.

MBBsn H. minima, Stoffelsrivier.

Others

Volume 5, Chapter 13:- A February 2009 Miscellany

7807 H.minima.  Swartrug, SE Heidelberg. 002

This chapter is based on recent field exploration and embroiders around many aspects of Haworthia species discussed in earlier chapters. What should be striking is that new populations follow the very predictable geographic pattern that all my earlier exploration has exposed and in my estimation confirm in every way what I consider a sound and satisfactory taxonomic solution and help explain its limitations.

1. Haworthia minima.

Two new populations of this widespread species are recorded:

MBB7807 Swartrug, SE Heidelberg. This is a steepish east facing slope. The plants are small, the leaf length barely exceeding 50mm and with a tendency to form clusters in the shallow clay banks that rift the area. There are many plants in quite a small area and it curious to see them within very large clumps of Euphorbia tridentata. This Euphorbia itself has a strange geographic distribution being known at Riversdale, Calitzdorp and then at Cradock in the Eastern Cape. (see Figs 1)

7807 H.minma. Swartrug, SE Heidelberg 003

MBB7813 NW Skeiding, E Tradouw Pass. This is a high rainfall area and we were expecting to find H. mirabilis in the lower pressure bursts and white clay that can be seen from the farmhouse far away. But instead we came across H. minima at the highest point of the hill among sparse grass. They are rather green in colour and it is interesting to note in the one plant pictured that it has non-tuberculate upper leaf surfaces and white cartilaginous margins and keel of the leaves. (see Figs 2)

2. Haworthia rossouwii.

MBB7803b Morning Star, S Heidelberg. Morning Star was known to J. Dekenah but the area has been very developed since those years. It was very surprising to find this small form in abundance at the top of a high hill in ferricrete pebbles and virtually no vegetation ground cover. However on this subsequent visit we crossed through an internal farm fence and found the plants in abundance about 150m further away and over quite a large area. We later looked at a lower promising hillside and small area nearer the Morning Star homestead and found the plants again. (see Figs 3)

3. Haworthia mirabilis.

This item reports on 6 populations, three of which are new. They confirm the continuity down the Duiwenhoks River and also southwards and westwards from east of Riversdale through to west of Robertson and Greyton and Napier to the south of that. Attention is drawn to south of Heidelberg where the ‘enigma’tic relationship of this species with H. retusa is suspected.

MBB7808 Swartrug,  SE HeidelbergThis farm is well known as the source of ‘Haworthia chromatica’ that I dismissed as an element in the complex surrounding the e H. retusa (mutica) ‘nigra’. The names Droekloof and Doringkloof have been used while the map reference gives Diepkloof for a large area extending to west of the Duiwenhoks River. The plants are on an east facing boulder terrace in a stony low bush site with some grass. The plants are small and show the expected wide range of variation. They are fairly similar to the plants in the lichen patch just east of Heidelberg that I have always mistakenly taken to be the origins of H. heidelbergensis, but here are subtle differences that mark a population difference, but I will come back to this. (see Figs 4)

MBB7809 Koeisekop, SE Heidelberg. This is on the same farm and a similar ferricrete hillside southwards. There are several hilltops by the same name in the general area and I cannot even speculate on the origins of the name (“cowshead”). The plants are larger than in the previous population and very abundant. The similarity to populations in the Swellendam and Riviersonderend area is in my opinion striking. Odd clones are miniature images of H.mirabilis from the southwestern populations west of Napier. (see Figs 5)

MBB7220 Morning Star,  S Heidelberg. I have discussed this population before and I repeat it here because I did not have many field photographs. It is only 300m from the Morning Star MBB7221 H. retusa ‘nigra’’ population and they flower and seed virtually in unison. Therefore they probably also reflect on the strange interaction between the two major systems. What I have surmised is that H. floribunda gets absorbed into H. mirabilis at the southern limits of its distribution and in these pictures there is clear evidence of the rounded and flattened leaf tips of that species. Therefore we may actually here have a three way species interaction. Reinforced by the presence of three different things at Kransriviermond, covered later under 4. H.retusaturgida’.  (see Figs 6a and b)

MBB7221 Morning Star

MBB7811 Kransriviermond, S Heidelberg. Populations further down the Duiwenhoks river have been reported on and this one fall within that mould and the same one as the previous Swartrug and Koeisekop populations. The population is smaller and the plants too are quite small. (see Figs 7)

MBB7814 Skeiding, W Heidelberg. This area is also reported on but digital photography makes it so much easier to record the enormous variability that such populations exhibit. Some of the plants are typically ‘atrofusca’ while others have leaves ended in an awn. Leaf surface may be smooth and there is even an indication of smooth leaves. What is notable is the absence of H. floribunda from what is the sort of habitat it can be associated with and can one can speculate its absorption into H. mirabilis with leaf shapes that demonstrate this. H. floribunda does appear again about 15km to the southwest in association with H. retusa and in the absence of H. mirabilis. (see Figs 8)

MBB7233 Die Plotte, Heidelberg. “H. heidelbergensis” correctly has its origin in this near vicinity and these small summer flowering elements are certainly part of a large assemblage of populations in the central area of what I now come to see as all H. mirabilis. Here discussion must pass to the following. (see Figs 9)

4. Haworthia retusa ‘turgida’.

A difficulty that I experience is in the need to dismiss the impression and general idea we have that there are tiered ranks. H. retusa has many populations and past treatment has placed most of the variants in H. turgida. To suggest that these are really two separate systems is not realistic and confounds any explanation for a few anomalies that exist.

MBB7810 Tierkloof, SE Heidelberg. Tierkloof is quite a deep valley that pushes out of the Duiwenhoks to the east. We found this huge population of large clump forming plants and several such populations are known just north of Heidelberg itself. I have applied the epithet ‘longebracteata’ to these and this is no more satisfactory than generating names for each of a very wide range of populations. There are populations along the Duiwenhoks that are far more inclined to the ‘turgida’ element and Essie Esterhuizen wrote a very good account of the host of variable populations that he encountered. (see Figs 10)

MBB7812 Kransriviermond, S Heidelberg. We only saw three plants at this locality and these plant mirror the preceding, if a bit smaller. The problem here is that we also have the element I myself described and named as H. retusa ‘mutica’  and I originally linked this “variety” with H.. mutica  following what G.G. Smith had done and for predictive reasons. The fact that ‘turgida’ now occurs with it is a huge hurdle to cross.  Think carefully. (see Figs 11)

MBB7804 Kransriviermond. I have written about his population in the chapter “Haworthia enigma”. This explains that the plants are not all leaden-black as perceived. They are spring flowering – as ‘turgida’ is. However, the range of forms and colour virtually replicates the plants in MBB7721 at Morning Star (pictures in the “enigma” chapter) where they are spring flowering. Across the river from both these populations is a weak population of small plants of H. mirabilis (7811 above) summer flowering. We know that this major difference in flowering season is not a total barrier to hybridization. (see Figs 12)

MBB7234 Die Plotte, Heidelberg. This is a population of large clump forming plants on a north slope about 300m south of MBB7233 H. mirabilis ‘heidelbergensis’. It is spring flowering. When I first saw these plants I dismissed (actually I never dismiss anything – I maintain reservations as I do now) these plants as large variants of ‘heidelbergensis’, and actually collected one magnificent specimen that looked to me just like H. mirabilis, say, from west Napier. Looking at the range of variation one is compelled to concede that here again is an infusion of H. mirabilis into a population that is primarily H. retusa in that vast shadowland it casts with ‘turgida’. (see Figs 13)

Conservation
I add these comments for several reasons surrounding the poor image that collectors have in the eyes of conservationists, and the very rickety fence I therefore occupy. There is a note reprinted in a recent issue of the newsletter To the Point. This taken from Times magazine and seems to be a rather hyped up condemnation of widespread destruction based on a view of internet websites. These are also monitored by RSA law enforcement that is rightfully disturbed by the high number of locality and collector citations that are indicative of illegal collecting. I personally no longer apply for a “collecting” permit for the prime reason that I am concentrating on photographic record, as a herbarium record no longer serves any better practical short term purpose. The herbaria are overburdened and I prefer living plants to dead ones. After 70 years of field activity I feel treated like a pariah and reduced to subservience under less-informed inexperienced people operating under a rigid and oppressive legislation aimed at acquiring a prosecution rather than wisely trying to direct activity in a positive direction. This activity of collection is, and I have no doubt about this either, a human right imbedded in the very reason for creation and our place in it. Conservation has its roots in the passion that people have for nature and it is nurtured by experience, contact and knowledge of its varied components. It is just ridiculous that one can go to any animal or plant centre here in South Africa and see plants and animals from anywhere else in the world and none indigenous. It is Ok to, say, keep Madagascan Chameleons in captivity where it is someone else’ responsibility to secure them. But keeping local ones is taboo? It is OK to freely sell cacti that do not occur here but are now being dug out in tons in (if that is really true). Recently it has become illegal to so much as possess Encephalartos latifrons and even the National Botanic gardens can no longer propagate and sell this plant. While we talk “plant indigenous” on all public fronts, the fact is that the acquisition, cultivation and sale of indigenous plants is viewed in the most negative light – a viewpoint encouraged by conservators.

This is control by law, a large degree of pure ignorance and by improper application of power derived from legislation. This is not the way to go. It is quite evident from the Times magazine article that legislation has the negative effect of creating a false value and glamour to discovery, Latin names, acquisition and possession. I believe that knowledge and understanding is true power, not legislation and enforcement.

The collecting numbers, localities and information provided here is to promote understanding and knowledge of plants, and thereby to promote their continued existence against all the vagaries of nature that “threaten” them, least of it which may be actually human greed.

Acknowledgement
I am glad to acknowledge the company, comments and interest of Gerhard Marx and whose organization sceptisism I also value. Landowners always impress me with their generosity and graciousness in allowing us access. On this occasion they include Mr. and Mrs. Nelie and Wimpie Jacobs of Diepkloof, Mr. Cloete and Belinda Fourie of Melkboom (B&B), Mr. and Mrs. Coetzee and Sarita Uys of Morning Star, Messrs. Christo and Pieter van Deventer of Doornvlei, Mr. and Mrs. Neels and Anne-Lise Uys of Skeiding (B&B) and then Ms. Jennifer Steggie of Heidelberg. In view of the comments on conservation noted above, I note that there are several farms that now offer accommodation and an outdoors experience and the opportunity to view these plants in their extraordinary natural surroundings. ♦

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 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. ♦

New finds in Haworthia

Previously published Cact. Succ. J. (Los Angeles) 84(1): 41-50.

Map - east of Swellendam

Map Legend – east of Swellendam.
1. JDV84/75 Haworthia retusa ‘turgida’.
2. MBB6666 H. retusa ‘nigra’↔ H. mirabilis.
3. MBB7898 H. retusa ‘nigra’.
4. MBB7899 H. retusa ‘nigra’.
5. MBB7897 H. retusa ‘nigra’.
6. MBB7896 H. retusa ‘nigra’.
7. MBB7871 H. mirabilis.
8. MBB7823 H. mirabilis.
9. MBB7909 H. mirabilis
10. MBB7805 H. mirabilis.
11. MBB7801 H. mutica ‘groenewaldii’.
12. MBB7886-7889 H. mutica ‘groenewaldii’, H. mirabilis, H. minima, H. marginata.
13. MBB7722 H. floribunda ‘major’

1. Haworthia marginata and H. minima

The Robustipedunculares is quite a distinctive group within the currently recognized genus. While the four species in the group are generally quite distinct, there are some remarkable complexities that rival that elsewhere in the genus. Recognizable and obvious hybrids are found between H. marginata, H. pumila and H. minima, but there are instances of whole populations that appear to consist of such in-between forms e.g. H. Xmortonii. I have speculated that perhaps rather than have just hybridized, the species have never ever really truly separated in the supposed evolutionary process. There is a vast body of variants that still link them in intimately as this piece will show.

I long ago observed that a small remnant population of ostensibly H. minima just south of Swellendam flowered in November as opposed to the general rule for the Robustupedunculares as late summer flowering. A vicarious population at Brandrivier north of the Langeberg (H. minima ‘opalina’) also flowers in November and both populations have fairly large and white flowers for the species.

I have recorded the normal bluish-green H. minima within the Bontebok Park at Swellendam as well as a very green variant. But in very recent exploration to the south-east we found an even more divergent group of plants that, while varying among the plants, seemed to be hybrids of H. marginata and H. minima (fig. 2 as a single sample and not representative of all). It was September and there were no signs of old or new flower spikes. Kobus Venter, who was present, remarked that the first plant seen was reminiscent of the plants once present south of Swellendam. The plants were large and in exposed situations even colored brownish as does H. pumila. No flowers were present and their color may have shown if H. pumila could have been directly involved at all, while it is essentially its distribution restricted eastwards from the Robertson Karoo by some 20km that reduce the possibility.

What makes the situation more interesting is that nearby was a population of H. marginata that was flowering and the flowers were also large and very white for the species (figs 3 & 4). Added to the fun was a smaller probable hybrid (fig. 5).

Differences and complexities like this do not really surprise me because it is what I have come to expect in my many wanderings in the field. The problem is that it certainly makes classification and any agreement on a set of names very difficult. I just accept it as a fact that plant species can exhibit greater differences between individuals of the same species than between individuals of different species, ridiculous as it may sound. This is because I perceive species as systems of individuals in populations with a very strong geographic component. To actually make a decision it is frequently necessary to determine just what else is growing in the vicinity in respect not only of the genus in question, but also that of other plants. Even the habitat factors need to be considered.

In the case of the plants pictured with this, the Bontebok Park terrain is mostly tertiary gravels, while the habitat we found the plants in was more recent riverine boulders. It is very curious that in the description of H. groenewaldii, the habitat is implicitly described as Ruens Silcrete. I do not think this is true. It is in the true Ferricrete – Silcrete that we found the next and it seems to be these differences in substrate geology that play a large role in generating variation and consequent controversy.

2. Haworthia retusa ‘nigra’

I first allied this element with H. mutica because this is how G.G. Smith referred to his collection from Kransriviermond south of Heidelberg. Since then there have been many new collections from which can be gathered that this population is a hybrid one between H. retusa ‘turgida’ and H. mirabilis. There is another collection north of this at Morning Star that appears to have the same parentage but also including H. floribunda. Then there are populations continuing up the Duiwenhoks and then Klip rivers to northwest of Heidelberg, a population between Heidelberg and Tradouw Pass further west and also a population at Goedverwagting south from there. Apart from the Morning Star population (February – March) these are all September/October flowering. There is a population at the southern entrance to Tradouw Pass of the same ilk that is February/March flowering. Then there is quite a distance between these known populations and a remarkable population at Buffeljags south of Tradouw Pass. It is remarkable for the fact that it is lauded as a new species viz. H. groenewaldii when 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.

To explore the realities of the situation we undertook two expeditions, one was to Buffeljags to explore west of H. groenewaldii and the other was to the Tradouw Pass area to explore H. retusa ‘nigra’. The first exploration yielded three populations of ‘groenewaldii’, which convince me that despite its flowering time as for H. marginata above, is simply H. mutica in another guise. I also think far too much is made of superficial and trivial differences that are as much characteristic of the variation in the one original population as they are within the four populations and for H. mutuca in its full sense. I consider it significant that H. mirabilis in its more normal non-retuse and dark green form is present in discrete populations both at and west of Buffeljags.

The second expedition was nearly as fruitful. It showed the Tradouw Pass population to be February/March flowering (see figs 6 & 7), while three new populations we discovered between there and the previous records in the easterly Heidelberg direction were September – October flowering. They link up to the populations elsewhere that I assign to H. retusa ‘nigra’. An additional find by Jannie Groenewald, for whom that H. mutica viz. H. groenewaldii, variant is named, also took us to a population of what is clearly H. mirabilis (see map) as I know it in its many disguises in the white kaolinic/bentonite clays of the silcrete – ferricrete inselbergs throughout the low-lying Southern Cape. There is unquestionably an overlap of characters between what I assign to H. mirabilis and H. retusa and I consider inarguable that H. mutica is a reflection of a shared gene pool.

What this demonstrates again, as does the Kiewietsvlakte populations between Heidelberg and Riversdale, that H. retusa and H. mirabilis are closely intertwined from east to west.  There is an added complexity that H. floribunda is admixed along the northern populations and H. variegata along the southern. The admixture of the two species produces H. retusa ‘turgida’  and H. pygmaea in the east and H. mirabilis, H. mutica and H. retusa ‘turgida’  in the west. This is complicated by the other interactions along the northern and southern areas. In the Potberg area it appears that the genetic material of all five “species” is evident in the populations that I have seen there.

While I would like to explain the situation around H. groenewaldii I. Breuer that I interpret as a variant of H. mutica, this should be left for another occasion as too many images are required to support any argument. As it is, the issue of H. mutica ‘nigra’ occupies 29 pages and 79 illustrations in my book Haworthia Update Vol.2 pt 1:50, where all the above mentioned variants are discussed and illustrated. The naming of Haworthia is highly contentious because the species consist of aggregates of small fairly isolated populations that may differ to large or small degree. The populations are in turn also aggregates of plants that can all be identical from vegetative propagation, similar because of low genetic difference or very different from each other because of large genetic differences. Therefore figures 6-13 simply show just a sample of the variation within these four populations in the Tradouw Pass area. The plants vary quite considerably in size too and the one in Fig 8 is nearly 200mm diameter. I have added to the map locations of the only significant other populations that I know of in the area including H. retusa ‘turgida’ and H. floribunda ‘major’, excluding those within the Bontebok Park viz. H. minima, H. mirabilis and H. marginata.

Fig. 14 The typical ‘pressure burst’ of white kaolinic soil from under more solid ferricrete where H. retusa and H. mirabilis are commonly found.

Perhaps I should close by explaining that I have dropped the use of any rank below that of the species name. I simply am suggesting that we recognize the need for a trinomial system without the pretension of status, and more greatly honour the binomial as an entity of a greater significance than we may know. 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 specimen that is used to anchor the Latin name.

Acknowledgement. Any proper excursion into Haworthia territory always requires acknowledgement of landowners and I thank Jaap Viljoen and Jannie Groenewald for organizing that and for their company. I was also glad to have Kobus Venter along who had persuaded me to show him some of populations known to me on promise of new exploration.

Note.
Cross seasonal hybrids observed are-

H. retusa turgida X H. floribunda Blackdown, Heidelberg.

And also Platjieskop, Riversdale.

H. pygmaea X H. floribunda Coopert Siding, Albertinia.
H. mirabilis
X H. retusa Soetmelksrivier, Riversdale
H. mirabilis X
 H. variegata Stoffelsriver, Swellendam.

References. I need to record that Harry Mays of Alsterworthia kindly undertook the non-profitable publication of 5 volumes of Updates (2-6) between 2006 and 2009. Vol. 1 was published by Umdaus in 2001. These volumes were the product of research to validate or correct what appeared as a formal revision in Haworthia Revisited, published by Umdaus in 1999. The description of H. groenewaldii appears in Alsterworthia 11.2:15 (2011).

Haworthia minima and pumila flowers

6645.1b H. pumila

This H. pumila flower is apparently persistently regarded by botanists as actinomorphic (star-shaped, radially symmetrical) – as though zygomorphy (yoke shaped, bilateral, asymmetrical) in the aloids is an uncommon condition!

Radial symmetry means the flower can be divided into 3 or more identical sectors which are related to each other by rotation about the centre of the flower. Typically, each sector might contain one tepal or one petal and one sepal and so on. It may or may not be possible to divide the flower into symmetrical halves by the same number of longitudinal planes passing through the axis. Zygomorthic flowers can be divided by only a single plane into two mirror-image halves, much like a yoke or a person’s face.

If you see the way the inner upper petal overlaps BOTH the two lower inner petals, you recognise that there can not be actinomorphy in aloid flowers.

Haworthia pumila

Haworthia minima

The Haworthia species of the Bontebok National Park

M. B. Bayer (MSc), Kleinbegin Farm, Kuilsriver, South Africa
Mrs. Carly Cowell (MSc), Regional Ecologist, Cape Research Centre, Conservation Services, South African National Parks, Cape Town, South Africa.


Objective: The significance of the Bontebo National Park occurrences.
The occurrence of members of three aloid “genera” (the three sub-genera of the genus Haworthia could indeed be genera) and the absence of any other member of the Aloids (bar the ubiquitous Aloe ferox) must surely be indicative of the driving forces that determine the flora of the Bonte National Park. This also must surely help establish the significance of the park as a conservancy of considerable merit. The complex interaction of the species enhances even that. The purpose of this report is to examine more closely the variation and nature of a small segment of the Park flora, and demonstrate how much more can be done.

Note: This report has several constraints.  Firstly is the situation in which there is no formal general definition and hence understanding of what a plant species is.  Secondly there is the generally understood view that there is an evolutionary process at work by which organisms evolve from a common distant origin by genetic mutation and adaptation.  Thirdly there are serious flaws in the classification of the Aloid genera.  Several essays dealing with these issues by DNA sequencing are weak because they rest on those flaws and consequently do not address some serious questions of relationships that the results pose.  Fourthly of course is the reality that the knowledge or intellectual capacity to overcome these deficiencies may be absent.  Thus the report is written in the context of all the publications as the original genus revision (Haworthia Revisited, Bayer; Umdaus, 1999) and others available on the internet (HaworthiaUpdates.org).

Introducing the subgenera.
There are 6 species of Haworthia that occur in the Bontebok National Park representing each of the three subgenera.  These subgenera are not real and have no closer relationship than that between any of the other genera in the Asphodelaceae however these are, or have been recognised.

Subgenus Hexangulares.  There are 15 species in this subgenus.  Two are in the northwest of South Africa and the others in the Eastern Cape or across the Karoo.  In respect of other Aloid genera it seems to be close to Chortolirion.  Only one species occurs in the Southern cape viz.H. venosa.

Subgenus Haworthia.  Here there are 37 species by my account.   Non-botanists raise this figure to as much as 600.  By their methodology and ideology it is a great deal more.  This subgenus is at odds with all the other Aloid genera and has an unusual internal diversity suggesting very recent speciation.  Three species occur in the Park viz. H. retusa (turgida), H. mirabilis and H. floribunda.

Subgenus Robustipedunculares.  There are 4 species all in the Southern cape with spill-over into the Little Karoo.  One species, H. pumila also occurs in the southern Great Karoo.  There is a very close affinity with the genus Astroloba.  The four species have a very close affinity and while H. kingiana in the eastern Southern Cape appears to be discrete, the other three definitely are not.  The Park occurrences are evidence of this and these seem to represent two species viz. H. marginata and H. minima.  See more under the treatment of H. minima below.

Map 1. Bontebok Park
Map 1. Bontebok Park

Introducing the species.
Illustrations are of a general form for the species.  Clonal variability is so high that use of the term “typical” can be misleading.Fig. 1 8038 H.venosa, Felix Unite
Haworthia venosa.  (Fig. 1)

This species occurs primarily along the lower course of the Breede River with its upper river limit just beyond the northwestern corner of the Bontebok National Park. Its lower limit is just north of Malgas.  Two populations are known to me off the river and a small way to the west. The species also occurs at Die Eiland in the Gouritz Valley.  Its nearest affinities are with H. tessellata that is very widespread across the northern Karoo and Cape; and with H. granulata.  The latter is certainly more closely allied to H. tessellata and the separation may just be on the basis of geographic separation rather than on the morphological characters that could be cited.   This may be true for H. venosa as well but in my opinion this would conflict with the general evidence of species geography.

Fig. 2 7895 H. retusa 'turgida', Diepkloof, Malgas
Haworthia retusa (turgida). (Fig. 2)

The name is written in this form because in the existing situation where species are confounded by definition, lesser categories are even more so.  The bracketing thus suggests that this is a variant of H. retusa.  This species is very widespread and abundant east of the Park with one vicariant population just north of Bredasdorp.  The (turgida) variant is one I regard as a riverine cliff ecotype and occurs in the park only along the Breede River at the northwest limits of the Park.  It is also downstream along the Breede River.  H. mutica is a closely related species that in my opinion actually just extends the distribution of H. retusa (retusa) westwards.   My opinion further is that an “evolutionary” process is not complete and that H. retusa and H. mirabilis may be the same “species” separated primarily on the basis of a different flowering season.  H. mutica does occur just east of the Park (Rotterdam, Mullershof), and my observation is that H. floribunda is involved with H. retusa and H. mirabilis in this occurrence.   It is in fact one of the objectives in this report; to ask if this might be true.  Flowering time is spring.

This species and its variants appears in various of my publications and often linked to H. mutica and H. pygmaea.

Fig. 3 7903 H. mirabilis, Felix Unite, Swellendam
Haworthia mirabilis. (Fig. 3)

This species is very common and widely distributed from Genadendal, Caledon and Napier all the way to east of Albertinia.  It is generally summer flowering and extremely diverse.  As its interaction with H. retusa in the west generates H. mutica, so I believe the same interaction produces H. pygmaea in the east.  H. retusa is spring flowering but hybrids with H. mirabilis are present in the field and there are populations that suggest that these two species are involved.  Flowering time is summer.

Extensive records are discussed and illustrated in various publications but primarily in the Update series, specifically Update Vol.3, part 1.  The species is incredibly variable and the floral variations are discussed and illustrated in a flower report as Haworthia Update Vol.8  (Alsterworthia 2012).

Fig. 4 7998 H. floribunda, Kruisrivier N, Riversdale
Haworthia floribunda. (Fig. 4)

This species is also summer flowering and has a curious relationship with both H. retusa and with H. mirabilis also with evidence that such “introgression” is evidenced in populations.  It is at its western limit at Swellendam and the variants in this area are extraordinary.  Flowering time is summer.

Also incredibly variable with hybridization with both H. mirabilis and H. retusa and it is hypothesized that some Haworthia populations are derived from such introgression (see also Haworthia Updates Vol.5 ,part 1, Chapter 6).

Fig. 5 7892 H. marginata, Rotterdam, Swellendam.
Haworthia marginata. (Fig. 5)

This species occurs from Ashton eastward to east of Riversdale in a rather narrow eas-west band that widens near Bredasdorp.  It interacts with both H. pumila and H.minima.  The species in this subgenus flower essentially in mid-summer.  The Park population together with that of the adjacent Rotterdam population flowers in spring.  At Rotterdam there is hybridization with a version of H. minima.

Fig. 6 7519 H. minima, S Kleinberg, Malgas.
Haworthia minima. (Fig. 6)

This species distribution generally complements that of H. pumila and then follows, but more widely that of H. marginata.  In the southwest it extends to near Elim and is common along the near-coastal strip as far east as Hartenbos.  The inland distribution is not well known in the area west of the Breede River.  It is probable that it was common in the area now extensively cultivated but it is not known to me inland in that area north of Tarentaal.  West of Swellendam there are only two records known to me at Sanddrift, Drew and again furher west of Drew.  In the one case there is hybridization with H. marginata.   In the second case a short distance west of this is the variant ‘poellnitziana’ and I suspect that introgression with H. pumila may have occurred there  The essential differences are that H. pumila has a colour range in the brown-green range, the flowers are generally olivaceous, and the tubercles are larger and rougher.  The colour range for the smaller H. minima plants is in the blue-green range, the tubercles are smaller and smoother, and the flowers whiter and greener.  H. pumila hybridizes naturally with Astroloba corrugata.  Also at Sanddrift, H. pumila hybridizes with H. marginata .  It is almost inconceivable that it has not hybridized with H. minima and it will be very difficult to superficially judge if plants were indeed hybrids or not without resorting to DNA sequencing (when that technology is advanced enough).   H. minima hybridises with H. marginata  at Heidelberg (S. Hooikraal) and in the past with H. marginata just northeast of Bredasdorp.  Flowering time is summer.

Some variation is presented and discussed in Haworthia Updates Vol.5. part 1, Chapter 7.

The geology of the Park
Bontebok National Park (BNP) is home to three main land types listed in the National Land Type Database, the first land type consists of conglomerate, sandstone and mudstone from the Uitenhage Group (Thamm and Johnson, 2006), which are overlain with Tertiary terrace gravel and can be found on the upper slopes of the north-western section of the park. The second land type is mainly tertiary and quaternary terrace gravels and covers the majority of the park, central and eastern sections. Within this there are high-level gravel terraces with silcrete and ferricrete outcrops, alluvial flats of loam and sandy loam. The final land type consists of siltstone, shales and sandstones of the Bokkeveld and Witteberg Group, these are found along the Breede River section in the western corner of the park. Soils comprise mainly of alluvial sand along the low-lying areas and undeveloped podzols in the main portion of the park. (Chief Director of Survey and Land Information 1993).

BNP_Geology
Map 2 – The geology of the Park

The vegetation and flora of the Park
Originally established to protect the threatened Bontebok antelope the Bontebok National Park is situated in the Cape Flora Region, a Global Biodiversity Hotspot (Goldblatt and Manning, 2000). BNP occurs in an ecotone (transition zone) between Fynbos and Renosterveld and the national vegetation map lists it as Swellendam Silcrete Fynbos (Rebelo et al., 2006) having a mix of both Renosterveld and Fynbos species. The conservation status of this vegetation type is Endangered with the largest portion being formally conserved within BNP. Along the Breede River banks Cape Lowland Alluvial vegetation occurs in the park and is listed as Critically Endangered (Driver et al., 2005), only 1% is formally conserved. A total of 650 floral species are listed as occurring in BNP, from 280 genera and 85 families (Kraaij, 2011). Of these twenty-nine species are listed on the Global threatened species list and a further 23 are recognised as species of conservation concern. The families with the largest species occurrence in the park are the Astercaeae, Iridaceae and Fabaceae (Kraaij, 2011), while the families of Asphodelaceae, Crassulaceae, Poaceae and Cyperaceae were well overrepresented in the park. There are also two species endemic to the park (Aspalathus burchelliana and Diosma fallax) and largest of only 2 populations of Erica filamentosa can be found in Bontebok. Succulents only comprised 8% of the total growth forms within the park with geophtyes being the majority (23%) (Kraaij, 2011). BNP flora is one of the richest in the region and conserves flora that are not conserved elsewhere. It is an area of high conservation priority and diversity and forms a major part in the conservation of Renosterveld in the Overberg region.

BNP_Vegetation_2012_03_22
Map 3 – The vegetation and flora of the Park

The scope and coverage of this report
Three of the Haworthia species are known from single populations, and these are indicated on the distribution map (see map 1).

The report therefore covers those species with multiple records, recording their whereabouts and the pictorial plant body and flower variations. Localities are numbered according to M.B. Bayer accession numbers.

A. Haworthia mirabilis.
Accessions 6644, 7704, 7744, 7805, 8035, 8043.
In Jan 2013 population 7704 was in flower, buds were observed at 8035 but not at 7744.
As in my flower report (Update Vol 8) the flowers are not convincingly different from those of H. floribunda.  The variation of the plants in especially 7704 and 7805 suggest introgression with H. floribunda.

  1. MBB6644 – Haworthia mirabilis

Flower profiles and faces

2. MBB7704 – Haworthia mirabilis

Flower Profiles, and Faces

3. MBB774 – Haworthia mirabilis

4. MBB7805 – Haworthia mirabilis

Flower profiles, faces, and bud

5. MBB8035 – Haworthia mirabilis

6. MBB8043 – Haworthia mirabilis

B. Haworthia floribunda.
Accessions 8017.  In flower Jan 2013.
A record 3439 exists for the Park, but the precise whereabouts is not recorded.  The plant colour and form do not accord with 8017.

7. MBB8017 – Haworthia floribunda

Flower profiles and faces.

Habitat DSCF0041
Habitat requirements of Haworthia can be so specific as to defy my describing them. Here is a picture looking north of the habitat for 8017 H. cf floribunda. The rocky ridge extends a long way further north and then west, as well as a long way south. Much of it is of course unexplored in enough detail to know if there are Haworthia there or not. At the southern end of the ridge about 1km away H. mirabilis occurs.  To the west the ridge continues, rises and then transforms into steep but low cliffs along the Breede River running northwards. On those cliffs one finds H. retusa (turgida). H. mirabilis is again present on north slopes east of the cliffs. H. venosa is on low river cliffs to the south and it may also be where this ridge abuts the river and turns north.

C. Haworthia minima.
Accessions 7743, 8036, 8037

In Jan 2013, a single weak inflorescence was observed at 8036.  8037 was flowering well.  No flower or buds were observed at 7743.  The plants are not the characteristic blue-green colour of regular H. minima, and the tubercles also seem to be rougher.  The flower colour is that of H. minima without the olive-green associated with H. pumila.  Note has to be taken of a now non-existent population just south of Swellendam on the N2 that flowered in late Spring with an abnormally large flower, and then a population to the near east at Rotterdam that appears to flower in Summer and where there are hybrids with H. marginata.  One cannot exclude the possibility that there is a genetic continuity with either or both, H. marginata and H. pumila.

8. MBB7743 – Haworthia minima

9. MBB8035 – Haworthia minima

10. MBB8037 – Haworthia minima

Flower profiles and faces.

D. Haworthia retusa (turgida).

Accession 2420.  This is the cliff-dwelling form of H. retusa and it is only known from the steep riverine cliffs on the northwest boundary of the Park.  Expected to be in flower in September.

The significance of the population is the similarity of some clones to H. reticulata that is only known eastwards along the Breede River until Drew from Worcester.  It also tends to be a clump-forming species favouring steepish rocky slopes as opposed to its geographic sister species H. herbacea that tends to be more solitary and occupying a wider range of habitat.

11. MBB2420 Haworthia retusa ‘turgida’

View downstream from 2420 habitat
View downstream from 2420 habitat
View up stream from 2420 habitat
View up stream from 2420 habitat

References:

DRIVER, A., MAZE, K., ROUGET, M., LOMBARD, A. T., NEL, J., TURPIE, J. K., COWLING, R. M., DESMET, P., GOODMAN, P., HARRIS, J., JONAS, Z., REYERS, B., SINK, K. & STRAUSS, T. (2005) National Spatial Biodiversity Assessment 2004:  priorities for biodiversity conservation in South Africa. . Strelitzia 14. South African National Biodiversity Institute.

GOLDBLATT, P. & MANNING, J. (2000) Cape Plants. A conspectus of the Cape Flora of South Africa, Cape Town, National Botanical Institute of South Africa Missouri Botanical Garden Press.

KRAAIJ, T. (2011) The flora of the Bontebok National Park in regional perspective. South African Journal of Botany, 19.

REBELO, A. G., BOUCHER, C., HELME, N., MUCINA, L. & RUTHERFORD, M. C. (2006) Fynbos Biome. IN MUCINA, L. & RUTHERFORD, M. C. (Eds.) The Vegetation of South Africa, Lesotho and Swaziland. Sterlitzia 19. Pretoria, South African National Biodiversity Institute.

THAMM, A. G. & JOHNSON, M. R. (2006) The Cape Supergroup. IN JOHNSON, M. R., ANHAEUSSER, C. R. & THOMAS, J. R. (Eds.) The Geology of South Africa. Johannesburg, The Geological Society of South Africa.