Volume 7, Chapter 7:- More on Haworthia mirabilis and H. mutica from east of Bredasdorp

Posted on May 6, 2012 by Bruce Bayer

The area concerned is the long and wide contact zone between the Limestone stretching from Bredasdorp to Potberg, and the Bokkeveld shale north of that. The soils and vegetation of the two areas are grossly different. The limestones are agriculturally almost useless, while the shales are prime wheat and pasturage producing soils although relatively low yielding. The vegetation of the shales is Renosterveld and there are very few patches left. Large areas resemble ecological deserts with nothing of the original surface intact. Here and there are shale banks and associated quartz outcrops and also some remnants of tertiary deposits that overlie the shale. Under this deposit layer the shale has decomposed to kaolin and in places there are gravel sheets of fine quartz on white clay. The skeletal nature of these remnants is the saving grace but it is unbelievable to what lengths farmers must have gone to make fields arable. Enormous amounts of stone that have been carted away and dumped to make cultivated lands. Sadly the stone is often dumped on exposed rock and prime Haworthia habitat. The remnants are still under threat and a mindset that has developed in the road construction and maintenance arena is that roads must be clean and scraped fence to fence. Similarly there are farmers who want every square inch under control and in subservience to their production needs. Dense vegetation is abhorred and burnt to control predation of sheep by jackal and lynx. Vegetation adjoining crops is treated with weedkiller to minimize crop contamination. Crops are also grown in conjunction with animal production. When crops are in, the animals are on fallow land and on whatever is left of natural vegetation. It is the harsh reality of conservation.

Haworthias exist on those remnants. I have often pondered that maybe they have not been so severely suffered by habitat destruction but now I am not so sure. There are many remnants where nothing is to be found, and others are species rich for no obvious reason. It is therefore difficult to assess.

This chapter explores the occurrences of two species at 11 localities in the Ouplaas vicinity east of Bredadsorp, and records some of their oddities.

The area in question:

This is shown in the map Fig 1 and I need to re-capitulate what I observed on Die Kop because I do not think I have given proper account for the populations I saw and the variance that I encountered in my first exploration there. This is most unfortunate as the name H. hammeri has since been applied to one or more of the variants and without context. Changes have taken place there too, that now compound the issue. I have in Vol. 3 Pt. 1 given an account of H. mirabilis in the wider area and will not repeat all of it, restricting myself just to those few populations that impact directly in these new observations. What is considerably more relevant is the chapter 13 in Part 2 where the main role players are listed and now again emerging in stark reality..

I have changed the arrangement from that in previous accounts to deal with local groups. Thus three sets, H. mirabilis at Die Kop – 5 populations and re-visits, H. mirabilis at Spitzkop, Haarwegskloof and Langvlei – 5 populations, and H. mutica at Spitzkop,- 3 populations.

SET 1 – Haworthia mirabilis Die Kop

Subset 1 MBB7500 Die Kop E. Figs 2-8

Considering the wide array of populations and variants in the greater area, I concluded that these are most probably the result of interaction between H. mirabilis and H. mutica. The surfaces vary from shiny minutely tubercled to smooth, leaf tips rounded or pointed and flowering time is summer. This particular population is in a deposition area off the Silcrete inselberg.

Subset 2 MBB7502 Die Kop N. Figs 9-10

When I first explored here in 2006 I skirted north of the first hill on Die Kop (see fig.1 Map) and observed an array of plants that were more like H. mirabilis and I dismissed them as such. However Jakub Jelimicky and Gerhard Marx both indicated that their observations contradicted mine.

Subset 3 MBB7973 Die Kop N. Figs. 11-27

We went further north on this excursion passing through the area touched on in 2006 that was now dramatically different. The previous shrubbery was absent and the area was trampled and grassy with also Bobartia, a large grass-like Irid. It seemed unlikely that Haworthia previously there could have survived, but we had seen them before still further west and they may still be present. This new area we saw was densely populated with plants similar to but smaller than MBB7500.

Subset 4 MBB7503 Die Kop mid. Figs. 28-34

My memory of these plants, east of those above, was that they inclined more to H. mirabilis than plants from 7500. Fig. 18 is almost identical to a clone photographed at 7500.

Subset 5 MBB7974 Die Kop mid. Figs. 35-46

Change due to heavy rain and erosion was more apparent here. There was less influence of H. mirabilis and the resemblance to 7500 nearly complete although the plants were generally smaller. This may be because the substrate was more skeletal and eroded than at 7500. What was really curious was the plant pictured figs 45-6 with narrow slender leaves. We did not observe any other species there that could have generated this as a hybrid but we did not have opportunity to comb the entire area small as it is.

Subset 6 MBB7504 N Die Kop. Figs 47-55

This population is about 1km north on an isolated inselberg. The plants are small and more typically mirabiloid with rough surfaces and spined margins and keel. Fig. 55 shows a plant with characteristics of the 7500 plants, but otherwise I would say this population perhaps relates better to Tarentaal MBB6539 (see Updates Vol 3.1 note 17).

SET 2 – Haworthia mirabilis Haarwegskloof and Spitzkop.

Subset 7 JDV86-2 Haarwegskloof. Figs 56-63

This subset with subset 11, are included as a reminder of what other populations in the area are like. Originally when so much less was known, this was classified as H. heidelbergensis var. scabra and subsequently so many variants have been found that naming each is not deemed practical by me. It is probably 3-4km west of the following three populations and similar populations occur about the same distance further south at Kathoek and Beyersdal. There are other populations known to me that I have never sampled or photographed and the following task is to explore a large area between Bredasdorp and Spitzkop.

The plants at Haarwegskloof, at the time the pictures were taken in late spring, were both with translucence and without. The plants were small and with leaves erect.

Subset 8 MBB7964 Spitzkop West. Figs 64-81

Fig. 64 indicates how difficult it is to find these small plants until you know exactly what you have to look for. One starts the search walking and eventually end upon all fours! Fig. 65 also shows how cryptic the plants can be. The plants are some of the smallest I have ever seen. Fig.72 shows an unusual plant with rounded tips to the leaves and I will show this again in another subset. Why this is so significant is because if the H. floribunda similarity in so many of the populations in the general area. The photographs disappointment me in that my first reaction on seeing the plants, was the association with images of H. parksiana. The pictures do no seem to fully reflect this. Kobus Venter has related his experience of finding H. parksiana-like plants among seedlings of H. mirabilis ‘magnifica’.

The leaves are reflexed as in H. parksiana as they of course are in many populations of H. mirabilis ‘maraisii’. But it is a curious reminder of the floribunda-like leaves in so many populations and the problem of look-alikes.

The plants were on a south slope in an area with white quartz and we had identified this as the most likely place to find them out of the direct north or northwest aspect. But we then did find them there too.

Subset 9 MBB7965 Spitzkop West. Figs 82-86

The plants were again hard to find over quite an extensive area. Fig. 83 again shows the floribundoid leaf tips.

Subset 10 MBB7966 Spitzkop West. Figs 87-103

It is curious how these small localized quartzitic areas can be so different. This was about 500m further north. There was more kaolin present as well as fine gravel. At an intervening similar habitat we found only Acrodon demidiatus, absent from the previous habitats and also from this one where Gibbaeum austricola now appeared. Again all the plants were small with dark green leaves, virtually no translucence and surfaces ranging from mildly spiculed to only minutely tubercled or just rough. Figs 99-101 are of plants with distinctly floribundoid leaves.

Subset 11 MBB7935 Langvlei. Figs 104-108

These are in white quartz and Aloe ferox is also present. There is moss present which is not the case at Spitzkop but true of Haarwegskloof. Curiously the pinched leaf ends as in fig. 106 have been observed in some plants of H. mutica too.

SET 3 – Haworthia mutica Spitzkop.

Subset 12 MBB7967 Spitzkop (quartz rock). Figs 109-148

Although a small area of quartz in a cultivated field, the plants are densely concentrated in small areas. There were some dense clumps in the rocks and this is unusual for H. mutica. It is the variation in these plants that deserves close attention. Some plants have very rounded leaves, they can be quite elongate or short and squat, others have an end-awn, the leaves may be opaque or even translucent; the surfaces may be shiny and minutely tubercled or smooth. Colour may be reddish, golden, or green. Most striking is perhaps the venation. There may be anything from 3-13 straight lines or very anastomizing (veins joining). There is no sign of either true turbercles or surface spination and also none of marginal spines.

Gasteria carinata is often severely animal grazed.

Subset 13 MBB7968 Spitzkop (pebbles). Figs 149-159

Curiously there is a shale ridge (fig.149) between this population and the first that one would think would be suitable habitat for this species as it is very like the Ouplaas habitat. But they were absent, perhaps because the ridge was lower down the slope. But a little higher where the shale was also exposed in smaller ridges the plants were also absent. Where they did occur was among small quartz pebbles to which the quartz may fragment. Fig. 157 shows a plant that was under the protection of small shrub and shaded.

Subset 14 MBB7969 Spitzkop East. Figs 160-161

This locality is about 3km east of the previous two at a slightly lower altitude. There is very little quartz but well exposed shale creating apparently suitable habitat as at Ouplaas now a short distance away. The area is well utilized by grazing animals and was very weedy, which may account for the fact that we found only two plants and these were not in the shale where we would have expected them.

Conclusion
I must here again relate the story in respect of KG35/70 of which there is only an herbarium specimen. That was found more than 40 years ago at Verfheuwel about 15km eastwards from Spitzkop. I was scrambling up a rough animal track to the top of the Verfheuwel hill when I saw some dried seed stalk less than 10cm high. The plants were tiny, greyish leaved with marginal teeth and rounded tips. I simply did not know what to do with them and identified them tentatively along with a few other populations from that wider area as H. maraisii. I went back to look for them several times with no luck. In 1996 I was with Kobus Venter and we found a strong population of larger plants not far away on the same hill. I recognize now that those first plants I saw were floribunda-like and that this similarity is commonly seen in populations of H. mirabilis in this area.

I speculated in Update 3 about the origins of H. mutica in the context of H. mirabilis and I do contend that these three H. mutica populations show characteristics that support this view. The Die Kop populations suggest to me a more recent amalgamation of H. mutica with H. mirabilis, to virtually reverse the process. The floribunda-like characters and even the parksiana resemblance also support this image of very close common origin and that particularly H. floribunda is simply a polytopic phenomenon in this south western area (polytopic – similar elements arising independently from the same common gene pool) and its genetic resources are embedded in H. mirabilis particularly. This seems to be definitely the case in the greater Swellendam and Heidelberg areas.

What is also so confounding is that now we have H. mutica occurring virtually undifferentiated in several different kinds of habitat. This is quite unlike the Van Reenens Crest area where similar habitat does not even have the same species. The H. mirabilis at Spitzkop west is unlike the other small plants in populations of H. mirabilis in the area and does seem to be differentiated according to habitat.

Acknowledgement:
Mr. Johannes and Pat Uys of Die Kop, Mr. Vlooi and Adele duToit of Die Kop, Mr. Gawie de Wet of Spitzkop (West), Mr. Franko de Wet of Spitzkop (East), Mr. Koos Badenhorst of Langvlei and Mr. Martin Mynhardt of Haarwegskloof. ♦

Haworthia flowers – some comments as a character source, part 1

Posted on May 12, 2012 by Bruce Bayer

M B Bayer, Kuilsriver, RSA

Introduction

The object of this essay is to discuss where we are now with respect to classification of Haworthia. Despite my comments and observations stretching over 50 years, there are still taxonomists writing and arguing on the basis of method and practice that generated the anarchy of names that existed at the start of my involvement. This method is what is probably referred to as “typological” i.e. there is a single herbarium specimen and anything that departs from this in some mind catching way is a different and thus needs a new name. At the generic level, recent DNA studies show that Haworthia is indeed three separate entities (the subgenera), and that these cannot be rationally separated from the aloid genera. Formal classification requires that Haworthia thus be subsumed in Aloe (see Treutlein et al, Rhamdani et al and Daru et al). This is both incomprehensible and anathema to writers and collectors locked into method that does not rest on any insight to what the problem of species actually is, let alone take proper cognizance of the problems that exists at generic level.

In response to some of the expressed anxiety about the proposed changes to Aloe, Dr John Manning, in a personal communication, wrote…”Most folk follow the biological species concept, which in its simplest form proposes that species are groups of interbreeding populations that are more or less reproductively isolated from other species. The existence of infrequent hybrids does not invalidate it. HOWEVER this is just about always a purely theoretical construct because there is no experimental data in most species to back it up. In practice, therefore, folk use morphology as a surrogate for reproductive isolation on the argument that discontinuities in morphology will coincide with these reproductive boundaries. Naturally there are all sorts of natural variabilities within species and so it becomes tricky to decide which of them fall within the boundary of the species and which fall outside of it”.

Two interesting things arise from this. Firstly, a “species” is itself a biological entity and there are not (or should not) be species that are not biological. Secondly there is a difference between a species concept and a definition. The concept is a general idea, while a definition lays down requirements that must be met. It might be better to say…”species determined by their interbreeding potential”, than use the term “biological species”. There is a huge amount of real experience and observation of the interbreeding capabilities of Haworthia both in the field and in cultivation. This suggests that if we really followed this criterion for determining in the genus then most of the Latin species names are superfluous, as I indeed consider them to be. Therefore, while there is a problem with the formal recognition of genera, the seems to be a total vacuum in respect of considerations around the constitution of species and how they can be, or are to, be circumscribed.

I have of course based what I regard as Haworthia species on a view of species as dynamic systems in a geographic framework, because history and my own long experience suggest that there are no morphological (or biological) criteria that produce the kind of classification that writers have produced or that collectors may need or expect. I proved by my studies in Oxalis, that morphological criteria can be extremely misleading, and that characters (or even character sets) thought to be useful in delimiting species, may in fact vary more within species than between species. I observe that this is true in many other genera. Flowers have recently been touted again by aspiring taxonomists as characters that will solve problems and it has even been said that I have ignored them. This is not true. I do not deny their value as characters. My experience simply suggests that variation and similarities in floral characters, mirror the situation with regard to vegetative morphology and may actually provide evidence for further amalgamation of species and reduction of names. This essay thus explores the situation in a data set, fortuitously available by virtue of modern digital photography and opportunity.

The elemental issue underlying all of this is dependent on the species definition. If some floral character is, or characters are, identified and specified as species characters then of course they would acquire significance in that context. What I can only show is that I observe nothing of the kind to relate to such a definition and insist on as species as systems in a geographic framework. Can they be rationally organized in any other system?

The hypothesis this essay addresses is that the floral morphology can be used to identify and circumscribe taxonomic ranks in Haworthia.

Overview

Originally I very laboriously drew flowers – often only one per collection. But one of my projects was trying to find more consequential objective evidence than just subjective vegetative differences to distinguish between H. reticulata and H. herbacea. I used a range of floral measurements but found the variation so great that to get a statistically significant result would have entailed many more samples than the number I used. Doing this would have been both very labour intensive and very destructive to the populations. I must point out that I am confident I could separate H. arachnoidea, H. mirabilis, H. pubescens as well as H. herbacea and H. reticulata (to some degree), all in the Worcester area, on the basis of their flowers. But this is not the level at which any other solution is needed beyond the appearance of the plants. There are only general differences by which one can separate H. herbacea and H. reticulata in respect of both vegetative and floral characters. H. wittebergensis could probably be identified just on the basis of its small white and more recurved flower, but its vegetatitve characters are quite adequate for the purpose.

Photography in those early days was not an option for recording flower detail. The advent of digital photography changes that quite dramatically and this essay is an exercise using this technology to attempt to document field observations. I photographed flowers as available in approximately 30 populations during February 2012, and these are used now in a posteriori manner to test the hypothesis regarding flowers. These images cover three species as I recognize them as systems in a geographic context viz. H. mirabilis (20 populations from three geographic zones), a summer flowering set of H. mutica (3 populations), H. floribunda (5 populations). Several outlier populations are added although sometime by single clones (see Table 1 listing the accessions. Map 1 (at end) indicates relative geographic position). These photographs provide a pictorial record of variation and difference within populations, between populations of the same species and between populations of different species.

Data

1. The plants. Figure Set 1 is a set of to 4 each, of the plants as they are in the populations covered. I have done this as a reference to how the names were derived and as I use them. The plants are far more variable than is generally understood and four images per population is really grossly inadequate as a sample to base a circumscription on. In the case of 7163 H. mirabilis, Frehse Reserve, S Riversdale, I have included pictures of plants (no flowers) because this is the type locality for the name H. magnifica (=H. mirabilis) that is widely used without any consideration of the range of variants there and how they may relate to other populations of H. mirabilis that are equally variable. Curiously there were no flowers and it appeared that the few plants that had flowered had done so earlier than expected. The same applies to a very similar population 6651 from a little to the southwest where there was only a single open flower. But two other populations viz. 7778 and 7818 in the area had many flowers at the time. It was rather similar at Van Reenens Crest where the various populations had not flowered with equal vigour and at some it was not possible to get a respectable number of flowers. It should be noted that in a good season with some summer rain, the plants will flower more successful with more plants flowering and some plants producing successive inflorescences. Thus the flower season could be extended be several weeks. Note that in the case of 7976 H. floribunda I did not take plant pictures for this second population.

Table 1. List of populations and identifications.

1. Haworthia mirabilis – eastern zone, Riversdale area (cf H. magnifica).
6651 H. mirabilis, E Riversdale.
7163 H. mirabilis, S Riversdale.
7778 H. mirabilis, Komserante, E Riversdale.
7818 H. mirabilis, Windsor, SE Riversdale.
7809 H. mirabilis, Koeisekop, Heidelberg.

2. H. mirabilis – central zone, Swellendam area.
7887 H. mirabilis, Rotterdam.
7912 H. mirabilis, Rietkuil.
7913 H. mirabilis, Rietkuil.
7916 H. mirabilis, Van Reenens Crest.
7917 H. mirabilis, NE Dam VR Crest.
7918 H. mirabilis, Van Reenens Crest, SE Dam.
7919 H. mirabilis, Van Reenens Crest, Game camp.
7955 H. mirabilis, Van Reenens Crest.
7959 H. mirabilis, W Van Reenens Crest.
7960 H. mirabilis, Kruiskloof.

3. H. mirabilis – western zone, Greyton area (cf. vars beukmannii, rubrodentata).
7262 H. mirabilis, S Greyton.
7978 H. mirabilis, Schuitsberg.
7979 H. mirabilis, Nethercourt.
7980 H. mirabilis, E Ouplaas.
7981 H. mirabilis, Schuitsberg N.

4. H. mutica – north-eastern zone, Buffeljags (Swellendam area, cf. H. groenewaldii).
7801 H. mutica, Buffeljags.
7888 H. mutica, Rotterdam 1.
7889 H. mutica, Rotterdam 2.

5. H. floribunda – central zone, Swellendam area.
7774 H. floribunda, S Swellendam.
7910 H. floribunda, Rietkuil.
7963 H. floribunda, Niekerkshek.
7975 H. floribunda, Rotterdam 1.
7976 H. floribunda, Rotterdam.

6. H. mirabilis – outliers and ‘outgroups’.
7248 H. mirabilis, Ballyfar, Infanta.
7513 H. mirabilis, Klipfontein N Bromberg.
7612 H. mirabilis, Diamant W.
7780 H. retusa ‘geraldii’, Komserante.
7781 H. retusa ‘foucheii’, Komserante.
7920 H. retusa ‘ nigra’, Van Reenens Crest.
7974 H. mirabilis, Klipfontein, Potberg.
7982 H. mirabilis, Melkhoutkraal, Goukou.
JDV87-132 H. parksiana, Mossel Bay.

The plant images :-

SET 1. H. mirabilis, Eastern Zone.

6651 H. mirabilis, SW Riversdale

7163 H. mirabilis, Frehse Reserve, S Riversdale

7778 H. mirabilis, Komserante

7818 H. mirabilis, Windsor

7809 H. mirabilis, Koeisekop

SET 2. H. mirabilis, Central Zone.

7887 H. mirabilis, Rotterdam

7912 H. mirabilis, Rietkuil

7913 H. mirabilis, Rietkuil

7916 H. mirabilis, Van Reenens Crest

7917 H. mirabilis, Van Reenens Crest

7918 H. mirabilis, Van Reenens Crest

7919 H. mirabilis, Van Reenens Crest

7955 H. mirabilis, Van Reenens Crest

7959 H. mirabilis, Van Reenens Crest

7960 H. mirabilis, Kruiskloof

SET 3. H. mirabilis, Western Zone.

7262 H. mirabilis, S Greyton

7978 H. mirabilis, Schuitsberg

7979 H. mirabilis, Nethercourt

7980 H. mirabilis, E Ouplaas

7981 H. mirabilis, Schuitsberg N

SET 4. H. mutica in its NE zone, Swellendam.

7801 H. mutica, Buffeljags

7888 H. mutica, Rotterdam 1

7889 H. mutica, Rotterdam 2

Set 5. H. floribunda in its western zone, Swellendam.

7774 H. floribunda, S Swellendam

7910 H. floribunda, Rietkuil

7963 H. floribunda, Niekerkshek

7975 H. floribunda, Rotterdam 1

7976 H. floribunda, Rotterdam – not illustrated.

SET 6. Outliers H. mirabilis and some relevant populations.

7248 H. mirabilis, Ballyfar

7513 H. mirabilis, Klipfontein, N Bromberg

7612 H. mirabilis, Diamant

7720 H. retusa ‘nigra’. Van Reenens Crest

7982 H. mirabilis, Goukou

JDV87132 H. parksiana, Mossel Bay

Haworthia flowers – some comments as a character source, part 1

Haworthia flowers – some comments as a character source, part 2

Haworthia flowers – some comments as a character source, part 3

♦

Haworthia flowers – some comments as a character source, part 2

Posted on May 12, 2012 by Bruce Bayer

2. THE RACEME. Figures Set 2 show the bases of the peduncles in several collections to again show how variable they are and not only because of plant vigor and current growth conditions. Diameter can vary by a factor of three. Color is variable and the bract spacing and size of bracts as variables must be noted. Fig 6 is simply a robust spike in a population where the flowers were sparsely spaced on the stem with approximately 15-20 flowers per stem, whereas this raceme had nearly 30 flowers. The number can drop to as low as three. In 7917 we noted a single plant with an inflorescence of over 600mm where the average was below 300. Similarly at 7818 there was one colossal inflorescence of 800mm where again the average was between 300 and 400mm. There is a real problem in that the typological attitude is often adopted when making comparisons like this. An extreme example would be to take H. retusa south of Riversdale as typical of the species. These are massive plants (source of ‘Jolly Green Giant’) and the inflorescences are huge with many flowers. This is not typical for the species and especially so if one takes the mountain cliff populations (H. turgida) to be the same species (as I do) where the plants are proliferous and the inflorescences many and reduced. Plants in poor niches and even poor habitats, flower weakly and the inflorescences are reduced. Figs 7 shows varying capsule positions on the stem. Figs 8 and 9 show a distichous and a secund inflorescence and figs 10 to 15 demonstrate the varied spacing of the flowers that is observable even in any one population although the images are for two different accessions. In Fig. 8 the middle flowers are in a single plane and I regret not having observed the leaf arrangement in that specific plant, because this is a distichous arrangement as the low Fibonacci number of a spiral arrangement of the flowers. This may have been reflected in the leaf arrangement too. The spacing and arrangement of the flowers is also a variable and the number of flowers may also vary. Figs 16 to 18 show bud arrangement and the way in which the fish-tail buds have upturned tips. Although the peduncle does continue to lengthen, most of the lengthening takes place in the flower producing area and towards the upper end of the raceme. The peduncle does not always stay straight and may bend slightly at each floret. The number and distance of the flowers along the peduncle may affect bud packing just as peduncle formation in the rosette results in the appearance of a groove on the leaf face. This is a secondary physical phenomenon and is not an inherent “character”. The leaf keel for example may also be influenced just by physical leaf-packing. A peduncle of a flower from the centre of a plant will have a many angled base, but one arising from a leaf axel only 2-angled. Generally the raceme is indeterminate, meaning that it does not end in a pedicel and flower. But I have seen an individual raceme of H. floribunda with a terminal flower. This exemplifies our problem where characters one might think could be used to determine even genera, are variables at species level.

2.1 7075 H. mutica, Grootvlakte
2.2 7262 H. mirabilis, S Greyton
2.3 7955 H. mirabilis, Van Reenens Crest

2.4 7778 H. mirabilis, Komserant
2.5 7809 H. mirabilis, Koeisekop

2.6 7913 H. mirabilis, Rietkuil
2.7 7919 H. mirabilis, Van Reenens Crest
2.8 7918 H. mirabilis, Van Reenens Crest
2.9 6509 H. mirabilis, W Bonnievale

2.10 7809 H. mirabilis, Koeisekop
2.11 ADH2729 H. mutica, De Hoop
2.12 7959 H. mirabilis, W Van Reenens Crest

2.13 7075 H. mutica, Grootvlakte
2.14 7741 H. mutica, Dankbaar
2.15 7920 H. retusa ‘nigra’, Van Reenens Crest

2.16 7781 H. retusa ‘fouchei’, Komserante
2.17 7778 H. mirabilis, Komserante
2.18 7818 H. mirabilis, Windsor

3. THE BRACTS. Figures Set 3. Bracts occur from the very base of the peduncle and subtend each flower. The bracts vary in density and size and one should recall J.R. Brown’s complaint that the plant he illustrated as H. schuldtiana var. major could not be that variety because the bracts were not the same length specified by G.G. Smith in the description. Although the bracts are attached to the stem, sometimes the stem is slightly swollen as though to form a base for the pedicel and occasionally the bract seems to be attached to that as though recaulescent (Fig.3.1). Usually there is a short pedicel but these do differ in length and width (figs 6 & 7, 8 & 9) within populations so that the florets may sit against the stem and in others away from it on a pedicel 3-4mm long (figs 2&3, figs 3,4 & 5). This is again an example of what might have been a primary character i.e. stipitate vs pedicellate, separating species, as a variable within a species. The bract may be quite narrow but may also be broad and almost encircle the stem (figs 12 & 13). The margins of the bracts may be entire or serrated in the same population and other oddities do occur (figs 8 & 9).

3.1 7262 H. mirabilis, S Greyton
3.2 7818 H. mirabilis, Windsor
3.3 7818 H. mirabilis, Windsor

Bract recaulescence? Note bract positions.

3.4 7918 H. mirabilis, Windsor
3.5 7918 H. mirabilis, Windsor
3.6 7919 H. mirabilis, Van Reenens Crest

Note presence or absence of pedicel.

3.7 7919 H. mirabilis, Van Reenens Crest
3.8 7919 H. mirabilis, Van Reenens Crest
3.9 7963 H. floribunda, Niekerkshek

Double bract

3.10 7818 H. mirabilis, Windsor
3.11 7818 H. mirabilis, Windsor
3.12 7778 H. mirabilis, Komserante

Variation in bract size. Bract almost wrapping peduncle.

3.13 7960 H. mirabilis, Kruiskloof Bract tucked under flower

4. THE BUDS. Figures Set 4. These figures illustrate the bud-tip as it occurs in most of the south-western Cape species. This flattened split tip is most evident in H. herbacea and H. reticulata but also present in H. maculata, H. retusa, H. mirabilis, H. mutica and H. pygmaea, but not in H. floribunda, H. rossouwii and H. variegata. The bud also is often slightly double-arched, bending first downward and the upward again especially evident in the bud of H. mirabilis 7248 Ballyfar. However, this is not quite an absolute and in some buds the character is not clearly flattened or may even be a bit rounded. I am not confident that my observations are entirely correct as I dismissed any floral characters quite early in my exploration when it was apparent to me that using a geographical approach might first tell me something about species from which I could then infer importance of characters of any kind. The effect of the flattened tip is to compress the tips of the upper petals, but in H. herbacea and H. reticulata the petal tips are quite spreading, whereas in H. mirabilis they tend to align vertically even folding behind the upper inner.

SET 4–BUDS

Subset 1. Haworthia mirabilis – eastern zone, Riversdale area (cf. H. magnifica).

7818 H. mirabilis, Windsor, SE Riversdale. (3)

7809 H. mirabilis, Koeisekop, Heidelberg. (3)

Subset 2. H. mirabilis – central zone, Swellendam area.

7916 H. mirabilis, Van Reenens Crest
7917 H. mirabilis, NE Dam VR Crest
7918 H. mirabilis,Van Reenens Crest, SE Dam
7959 H. mirabilis, W Van Reenens Crest

Subset 3. H. mirabilis – western zone, Greyton area
(cf. vars. beukmannii, rubrodentata).

7978 H. mirabilis, Schuitsberg

Subset 4. H. mutica – north-eastern zone, Buffeljags
(Swellendam area, cf. H. groenewaldii).

7801 H. mutica, Buffeljags. (3) 7889 H. mutica, Rotterdam 2.

Subset 5. H. floribunda – central zone, Swellendam area.

7774 H. floribunda, S Swellendam. 7963 H. floribunda, Niekerkshek.

Subset 6. H. mirabilis – outliers and ‘outgroups’.

7248 H. mirabilis, Ballyfar, Infanta. (2) 7612 H. mirabilis, Diamant W. (2)

7780 H. retusa ‘geraldii’, Komserante
7781 H. retusa ‘foucheii’, Komserante
7974 H. mirabilis, Klipfontein, Potberg

7982 H. mirabilis. , Melkhoutkraal, Goukou
JDV87-132 H. parksiana, Mossel Bay

5. THE FLOWERS. Figures Set 5a and 5b

5a. Flower progression. Figures set 5a illustrate the ageing progression in two florets from two populations. One of the biggest hurdles in describing or illustrating the flower is the change that takes place that occurs in the life of the flower from bud opening to flower wilting over a period of 4-6 days. As can be seen in these successive images, the petal spread wider with age. But clearly there is a difference in basic structure of the individual flower that also determines attitude of the petals through the various stages. Colour and texture of the flower may also vary as the flower loses turgidity with aging and wilting.

5a.1 7913 H. mirabilis, Rietkuil

5a.2 7963 H. floribunda, Van Niekerkshek

Flower length in mm

5910	7918	7513	7262
14.3	13.5	11.8	14.5
13	13.5	12.1	13.7
12	13	12	14.6
12.5	13.4	12.5	14.6
14	13.1		13.4
13	13.4		15
13	13.2		15.3
14	13.2		16.8
12.5	13		16.7
14.2	12		15.3
12.5	12.8
13.5	12.5
12	12.4
14	12.6
13.5	12.5
	12.3
	12.5
	12

Fig. 5a.3 Table – headed by collecting nos. Flower length in mm.

5b. Flowers. Figures Set 5. Flowers were photographed as available using fixed lens settings so that images were reasonable comparable. Some were taken and seized against a similarly photographed millimeter rule and these measurements are given in Figure 5a.3 above as a general guide to size and variation. This was only done for 4 sets of flowers because it is very evident that my original observation of variability of a high order is true. The measurement is simply of the maximum length of the flower from attachment to pedicel to furthest upper tip in four collections. Obviously this is not ideal as how the upper petals reflex (and with age too) will distort the result. But I used it, and submit it to provide a scale for any other measurement of those given flower that one would like to make. Smaller dimensions using a millimeter ruler are not remotely accurate enough over the error already present arising from method. The sizes given in species descriptions in the archaic way still practiced, are fairly useless as there is no indication of precision and any statistical measure of inherent deviation. The photographing method, despite changes also inherent in the picture editing process, allows reasonable size comparisons. The fact is that across the three subgenera, tube length is consistently in the area 8-10mm probably corresponding to a common pollinator. An important consideration is also that of the requirements of a sampling process in statistics where randomness is essential. The tendency in the field is to be selective and this generates the problem of bias that is almost unavoidable even when this is considered.

A point that must be stressed is the question of plant condition on whatever structures are used for study. Generally if a plant flowers at all, it can be assumed that the condition of the plant was good enough and the flower appearance would be true. This is an assumption carried into most studies of herbarium material and generally accepted as valid. But when decisions are based on the nature of whole inflorescences, it becomes quite evident that plants in different niches and habitats, and even different seasons, will produce inflorescences according to plant vigour. Over and above this is the variability that derives from habitat adaptation. Proliferating ecotypes on well-drained cliff-faces will produce quite a different inflorescence to that of solitary ecotypes in deeper moisture-holding soils. Flowers on an inflorescence lasting 3-4 days from initial petal spread to wilting also change with time. The position on the flower is also significant as the terminal flowers are invariably smaller. Another problem is the assumption that characters are independent. The fact is that, say, the bud-tip is flattened. Petals contained in that bud-tip are constrained to fold accordingly and if the petals vary in width or length they will display differently simply because of the containment within the bud-tip.

The populations listed in Table 1 were visited during February when they were in flower, and a series of flowers photographed in each population as available. Each flower was photographed in profile and from face view with the camera lens at a fixed focus setting. It is just a fact that all Haworthia flowers are closely the same in respect of size as stated above. I am aware of smaller flowers in H. wittebergensis, as one example, and also very large flowers to nearly 30mm long (against an average of approximately 15-20mm) in H. herbacea from Villiersdorp. The numerical measurements are variable and while there is no doubt that these could be found if sought, the intention here is to show that they are unlikely to help resolve the issues of species differences at the level that they are needed. This is certainly not without very large samples and precise measurement to accuracy of at least 0.1mm. Some observations were recorded as described above of peduncles, capsules and seeds. But only to indicate that they have been considered and also regarded and seen as variables that are unlikely to make any contribution by virtue of the degree of similarity they show even over the few species involved in the observations. What differences are discernible are highly unlikely to have any significance when one thinks that these have to be shown to exist consistently in very similar structures over many populations and many species.

The largest differences one would expect to see should be in the individual flowers (florets). Pictures were thus taken of the flowers in profile and also of the facial aspect. Size is a variable and Fig. 5b.1 shows three flowers from different clones in 7910. Fig. 5b.2 shows two flowers from the base and tip of a single inflorescence in 7980 – the upper floret half the size of the lowest. Fig. 5b.3 is a diagram to show profile and face of a flower (6651) to suggest axes along which a flower could be orientated to obtain proper points of reference for measurements of angles or size. The face and profile are of 6651, but the three part flower bases are all from the same collection. These virtually sum up the entire argument as the diagrams show a floret without a petiole, and various shapes to the base of the tube that I refer to as the perigon – the section before the petals separate. The taper from the pedicel onward (the stipa?) is very variable and may be abrupt or extended. The tube may be uniformly deep or constricted at the middle. A pronounced swelling (gaster) of the tube may occur below near the base. Other variables are the angle at which the flower stands to the peduncle. This one rather complicates the imaging of the face as I tried to photograph holding the camera at right angles to the peduncle and that does not properly capture a flower that is very vertical.

The sequence of illustrations onwards from 5b.4 is first the faces and then the profiles of the flowers according to the listing of the sets of populations in Table 1.

SET 5B.4 – FACES

Subset 1. H. mirabilis – Eastern Zone, Riversdale area.

6651 H. mirabilis, SW Riversdale

7778 H. mirabilis, Komserante

7818 H. mirabilis, Windsor

7809 H. mirabilis, Koeisekop

Subset 2. H. mirabilis – Central Zone, Swellendam area.

7887 H. mirabilis, Rotterdam

7912 H. mirabilis, Rietkuil

7913 H. mirabilis, Rietkuil

7916 H. mirabilis, Van Reenens Crest. 7917 H. mirabilis, Van Reenens Crest.

7918 H. mirabilis, Van Reenens Crest

7919 H. mirabilis, Van Reenens Crest

7955 H. mirabilis, Van Reenens Crest

7959 H. mirabilis, Van Reenens Crest

7960 H. mirabilis, Kruiskloof.

Subset 3. H. mirabilis – Western Zone, Greyton area.

7262 H. mirabilis, S Greyton

7978 H. mirabilis, Schuitsberg

7979 H. mirabilis, Nethercourt

7980 H. mirabilis, E Ouplaas

7981 H. mirabilis, Schuitsberg N

Subset 4. H. mutica – Northeastern Zone, Buffeljags area.

7801 H. mutica, Buffeljags

7888 H. mutica, Rotterdam 1

7889 H. mutica, Rotterdam 2

Subset 5 H. floribunda – Central Zone, Swellendam area.

7774 H. floribunda, S Swellendam

7910 H. floribunda, Rietkuil

7963 H. floribunda, Niekerkshek

7975 H. floribunda, Rotterdam 1, 7976 H. floribunda, Rotterdam

Subset 6. Outliers and ‘outgroups’.

7248 H. mirabilis, Ballyfar

7513 H. mirabilis, Klipfontein, Bromberg

7612 H. mirabilis, Diamant W

7780 H. retusa, Komserante. 7781 H. retusa, Komserante. 7915 H. mirabilis,Van Reenens Crest

7920 H. retusa ‘nigra’, Van Reenens Crest
7974 H. mirabilis (‘floribunda’), Klipfontein, Potberg
7982 H. mirabilis, Goukou

87-132 H. parksiana, Mossel Bay
7937 H. mutica, Spitzkop, Bredasdorp

SET 5B.4- PROFILES

Subset 1. H. mirabilis – Eastern Zone, Riversdale area.

6651 H. mirabilis, SW Riversdale

7778 H. mirabilis, Komserante

7818 H. mirabilis, Windsor

7809 H. mirabilis, Koeisekop.

Subset 2. H. mirabilis – Central Zone, Swellendam area.

7887 H. mirabilis, Rotterdam

7912 H. mirabilis, Rietkuil

7913 H. mirabilis, Rietkuil

7916 H. mirabilis, Van Reenens Crest. 7917 H. mirabilis, Van Reenens Crest.

7918 H. mirabilis, Van Reenens Crest

7919 H. mirabilis, Van Reenens Crest

7955 H. mirabilis, Van Reenens Crest

7959 H. mirabilis, Van Reenens Crest

7960 H. mirabilis, Kruiskloof

Subset 3. H. mirabilis – Western Zone, Greyton area.

7262 H. mirabilis, S Greyton

7978 H. mirabilis, Schuitsberg

7979 H. mirabilis, Nethercourt

7980 H. mirabilis, E Ouplaas

7981 H. mirabilis, Schuitsberg N

Subset 4. H. mutica – Northeastern Zone, Buffeljags area.

7801 H. mutica, Buffeljags

7888 H. mutica, Rotterdam 1

7888 H. mutica, Rotterdam 2

Subset 5 H. floribunda – Central Zone, Swellendam area

7774 H. floribunda, S Swellendam

7910 H. floribunda, Rietkuil

7963 H. floribunda, Niekerkshek

7975 H. floribunda, Rotterdam 1. 7976 H. floribunda, Rotterdam.

Subset 6. Outliers and ‘outgroups’.

7248 H. mirabilis, Ballyfar

7513 H. mirabilis, Klipfontein, Bromberg

7612 H. mirabilis, Diamant W

7780 H. retusa ‘geraldii’, Komserante
7781 H. retusa ‘foucheii’, Komserante
7915 H. mirabilis, Van Reenens Crest

7920 H. retusa ‘nigra’, Van Reenens Crest
7974 H. mirabilis (‘floribunda’), S Klipfontein, Potberg
7982 H. mirabilis, Goukou

JDV87-132 H. parksiana, Mossel Bay

Haworthia flowers – some comments as a character source, part 1

Haworthia flowers – some comments as a character source, part 2

Haworthia flowers – some comments as a character source, part 3

Posted on May 12, 2012 by Bruce Bayer

6. THE CAPSULES. Figs 6.1 first size differences that can be found within individual inflorescences. The remaining images show 8 capsules per population for a few populations to show variation in size and shape. The way the capsule ripens and splits is very variable. In some cases the capsule is pinched near the end but the locule tips flare outwards. This has the effect of seeds being retained in the capsule. In others the locules flare regularly and symmetrically from the base and the seed is all easily released. In the Van Reenen Crest populations the capsules were inclined to be a reddish hue. But colour can vary depending on the ripening process and they also bleach with time. Fig.6.4 7978 shows this in capsules drying after the peduncle was taken, and retaining their greenish colour. In H. floribunda the capsules are smaller and it appears that they may flare at the tips more in splitting and be coarsely crispate. This is not always the case but it is a tendency in the smaller capsules to do this. Fig. 6.7 is of capsules in 7910 H. floribunda, Rietkuil; compared against 7913 H. mirabilis also Rietkuil east of Swellendam. It is quite evident that even a capsule structure as apparently characteristic as in H. floribunda, is replicated in a different species. Fig. 6.8 is of 7955 Van Reenens Crest, and 7262 south of Greyton. I thought the capsules of 7955 were smaller, reddish coloured and slightly rougher than those of 7262. But the capsule structure in the entire genus is very similar to those shown on this figure and it is just not conceivable that some feature will stand out and resolve issues that exist in respect of the entire group.

7163 H. mirabilis, S Riversdale. 7809 H. mirabilis, Koeisekop.
Fig. 6.1 Size variation – range 5-12mm.

6651 H. mirabilis, SW Riversdale
7163 H. mirabilis, S Riversdale
7778 H. mirabilis, Komserante

7818 H. mirabilis, Windsor
7809 H. mirabilis, Koeisekop
Fig. 6.2 H. mirabilis – Eastern Zone, Riversdale area.

7913 H. mirabilis, Rietkuil
7919 H. mirabilis, Van Reenens Crest
7959 H. mirabilis, Van Reenens Crest
Fig. 6.3 H. mirabilis – Central Zone, Swellendam area.

7262 H. mirabilis, S Greyton. 7978 H. mirabilis, Schuitsberg
Fig. 6.4 H. mirabilis – Eastern Zone, Greyton area.

L(1) 7888 H. mutica, Rotterdam.7910 H. floribunda, Rietkuil
R(2) 7889 H. mutica, Rotterdam
Fig. 6.5 H. mutica – Buffeljags.Fig. 6.6 H. floribunda – Swellendam.

Above 7910 H. floribunda, Rietkuil. Above 7955 H. mirabilis, Van Reenens Crest
Below 7913 H. mirabilis, Rietkuil. Below 7262 H. mirabilis, S Greyton
Fig. 6.7 Some capsule comparisons.

7. THE SEEDS. Some seeds are illustrated in figures Set 7. Any differences there might be in the seeds will be extremely difficult to quantify or describe. There is clearly some difference in size but to establish this statistically will be difficult. In the set of pictures provided there is clearly no difference in appearance or size coupled with the fact that the shape beyond “angular” and colour “grayish black” is difficult to articulate. What size difference appears here is due to picture magnification. Again I am quite aware that in some populations seeds are indeed larger than in others, but vegetative characters are already enough to distinguish those. Species recognition would only be facilitated if one based the definition of the species on some formula specifying seed size and shape. In the figure, 7983 is the seed of H. minima to show the quite different flattish shape and marginal wings in that subgenus. My impression was that the seed in H. floribunda may be a little chunkier than in the other samples but it just does so happen that in some capsules there are fewer and larger seed than in others.

SET 7 – SEEDS

1 – 6651 H. mirabilis, 1 – 7163 H. mirabilis, 1 – 7778 H. mirabilis

1 – 7809 H. mirabilis, 1 – 7818 H. mirabilis, 2 – 7912 H. mirabilis

2 – 7913 H. mirabilis, 2 – 7919 H. mirabilis, 2 – 7959 H. mirabilis

3 – 7262 H. mirabilis 3 – 7978 H. mirabilis 3 – 7980 H. mirabilis

4 – 7888 H. mutica, 4 – 7889 H. mutica, 5 – 7910 H. floribunda

5 – 7963 H. floribunda, 6 – 7982 H. mirabilis, L.7262 H. mirabilis, R.7955 H. mirabilis

6 – L.7910 H. floribunda, R.7913 H. mirabilis, 7283 H. minima

DISCUSSION AND CONCLUSIONS
While assembling and sorting the data sets I was made painfully aware of how necessary it was to keep tabs on each item. This because the similarities of between structures in very different populations was so close that without label identification, it was not possible to re-sort images into original order. Starting with flowering time that I have not yet mentioned here, as I have been dealing essentially with populations flowering in February. The southwestern Cape species of the subgenus Haworthia flower either in spring or late summer. This is the essential difference between what I regard as H. retusa (sensu lato – in the broad sense to include H. turgida) and H. mirabilis (also sensu lato to include H. magnifica, H. maraisii and H. heidelbergensis). The former set of populations flowers in the spring and the latter in late summer. But the actual time is a bit variable too as I have seen populations of the spring flowerers doing so as early as June and as late as October, while the summer group can flower from November to April. In addition there are odd plants that just flower ‘out of season’ and field hybrids present within populations of the two sets is evidence of this. It is evident from distribution, vegetative appearance and even flowering time that the two species H. retusa and H. mirabilis are parental in the origins of H. pygmaea in the east (Mossel Bay) and H. mutica in the west (Caledon). So the data here really only addresses the issue of H. mirabilis and its internal relationships, and its relationship to H. mutica in the northeastern populations. This is where one population (viz. 7801) is claimed to be a distinct species viz. H. groenewaldii. The one population 7778 H. mirabilis at Komserante obviously judging from the vegetative appearances, is infused with H. retusa. It is instructive then to look at the only two flowers I have of two nearby populations of H. retusa viz. 7780 and 7781 (see subset 6 of set 5a and 5b)

To what extent this data will influence this last mentioned claim is unknown. I simply have not and cannot discuss the floral structures in detail. This is primarily because they are so incredibly variable. I find virtually the same detail in any of the sets of figures I care to consider. What I would say that my data reflects is the interaction and continuities that I maintain are mirrored in the overall physical appearance of the plants. I have never considered 7801 H. mutica, Buffeljags as anything less than new or as anything more than the expected connection between the original H. mutica var. nigra (now H. retusa var. nigra, south of Heidelberg) and true H. mutica from the wider Bredasdorp and Caledon areas. What I did speculate was the possible involvement of H. floribunda given the distribution of that species and its interaction with both H. retusa and H. mirabilis throughout its distribution range.

I am sure this limited data set, illustrates the possible truth of my rationalization of the relationships of the plants considered. I am simply unable to extract any single point by which I can circumscribe even one population. A possible exception may be 7248 H. mirabilis, Ballyfar in which the colour yellow seems to dominate. But I recall my early involvement in Haworthia when I thought I could possibly separate H. mirabilis and H. maraisii on the basis of their flowers; with the latter having smaller and greener flowers. However, I could not do this an observations on more populations suggested to me this was just geographic variation – NOTE – differences may simply be characeristics of a species and not necessariiy denote different species. This may even apply to flowering time.

My conclusion confirms what originally impressed me. Generally flower characters will not resolve the difficulties of species circumscription based on the method, way of thinking, motives and rationale that has prevailed since Haworthia were first observed and described. Flower morphology is supposed to be central to the Linnaean method in view of their significance in the breeding process. At Buffeljags, the geographic centre of my ‘study’ area, there are close populations of H. mutica, H. mirabilis and H. floribunda across a distance of less than 600m. The plants flower synchronously and the same pollinator is present and was observed at virtually all the populations included in the entire study. H. minima and H. marginata also both occur and hybridize there too. I have up to now concluded that H. floribunda is in any case deeply infused to the point of merging into H. mirabilis both south of Heidelberg and around Swellendam. This new information, including a population pictured and reported by J. Duncan of Kirtsenbosch (nearby but within the Bontebok Park), confirms that H. floribunda is definitely interactive with H. mirabilis. My original perception of 7801 H. mutica, Buffeljags, was that it originates in the interbreeding of the three species H. retusa, H. mirabilis and H. floribunda. BUT this is not in the sense of a post speciation event where the three elements have arrived progressively at individuation as species. It is the result of the large-scale and all-encompassing process by which one gene pool has hived off H. retusa and H. mirabilis and these in turn have hived off H. pygmaea and H. mutica. The argument that 7801 H. mutica, Buffeljags is nearer to H. mirabilis than it is to H. retusa has no merit. I have in the Update Volumes explained that H. mutica is very closely allied to H. mirabilis as evident from variants seen in the field. In any case the argument rests on a typological approach where very narrow limits are attached to species names. My concept of species has developed from the plants as I have experienced them in the field, and I have not imposed any predetermined species concept on them. This study gives strength to my argument that species are complex chaotic fractal systems and that this is the model on which classification and nomenclature must be based.

(Note – a method to get more accurate mensuration would be to photograph at fixed focal length, and photographing also a millimeter rule at the same setting as I have done. However, the pictures must then be used unedited and at higher magnification more precise measurements will be attainable).

ACKNOWLEDGEMENT
I would like to thank Lawrence Loucka for his interest in the subject and his comments and advice on the manuscript. Jannie Groenewald also accompanied and assisted me in the field, as did both Lawrence and Kobus Venter. Landowners were generous with permission and I particularly acknowledge Hesphia and Trevennan Barry of Van Reenens Crest, Nelis Swart of Postdal (Kruiskloof), Leanne and Pieter Urschel of Klipfontein, Johannes and Wilmien van Eeden of Windsor, Mathewis Odendaal of Diamant, Mark and Ryno Stander of Komserante, Wimpie and Nelie Jacobs of Koeisekop, S. Smuts of Nethercourt and P.G. Viljoen of Schuitsberg.

References

Bayer, M.B., & Manning, J.C. Rationalization of species names in Haworthia. 2011. Haworthia Update Vol 7, part 4. Alsterworthia.

Daru, B. H., Manning, J.C., Boatwright, J.S., Maurin, O., Maclean, N., Kuzmina, M., & van der Bank, M. 2011. Phylogeny of the subfamily Alooideae (Asphodelaceae): Paraphylly of Aloe and Haworthia and consequences for classification. In ms.

Treutlein, J., Smith, G.F., van Wyk, B.E. & Wink, M. 2003a. Evidence for the polyphyly of Haworthia (Asphodelaceae subfamily Alooideae: Asparagales) inferred from nucleotide sequences of rbcL, atkK, ITS1 and genome fingerprinting with ISSR PCR. Pl.Biol. 5:513-521.

Treutlein, J., Smith, G.F., van Wyk, B.E. & Wink, M. 2003a. Phylogenetic relationships in Asphodelaceae (ubfamily Alooideae) inferred from chloroplast DNA sequences (rbcL, matK) and from genomic finger-printing (ISSR), Taxon 52:193-207.

Rhamdani, S., Barker, N.P. & Cowling, R.M. 2011. Revisiting monophyly in Haworthia Duval (Asphodelaceae): incongruence, hybridization and contemporary speciation. Taxon 60: 1001-1014.

♦

Haworthia flowers – some comments as a character source, Appendices 1, 2, 3

Posted on May 12, 2012 by Bruce Bayer

Appendix 1.

Haworthia flowers – some comments as a character source.

I am adding images of flowers from 3 populations of H. mirabilis from the Worcester/Robertson Karoo. These are:

6509 H. mirabilis meiringii, W of Bonnievale
7176 H. mirabilis maraisii, Agter Vink
7279 H. mirabilis meiringii, Edendale, E Bonnievale

(The second epithet should not be taken seriously while the omission of a rank viz. var. is abandoned as is often done in zoology – I take the second epithet loosely to refer to a range of variants with some approximate but vague order).

These are to be seen as outgroups to the main body of the report/essay as show the differences of both size and colour that I once thought separated H. maraisii from H. mirabilis. Further exploration and experience have made me change my mind and I cannot see any way in which H. heidelbergensis, H. magnifica and H. maraisii can be visualized or rationalized as distinct from H. mirabilis. The flowering time for these three populations seems to be later than for the eastern and southern populations, but indeed there are populations in the MacGregor area that flower as early as November. Populations towards Montagu may flower in April. While the flowers in the populations of the main set were in seed at end March, capsules were only forming in these added three populations. A population 1708 north of Ashton was just producing buds when these were flowering as fully as could be expected in a very dry season. The flowers are indeed generally smaller than those observed in the main body but I am not sure if this is statistically certain. The same bee pollinator was observed but not the Bombylid flies. The flowers particularly in 7279 appeared to have been punctured or damaged along the sides of the flower tube as though by an insect (pollinator?) of some kind. Damage to the buds and petal ends was also noted.

I would like to explain and emphasize that I never ignored flowers as a character source and am not presenting them now as a solution to anything. What they do provide is another source of information and digital photography offers a means to capture that. It would be very silly to underestimate the volume of data required and ignore the rigours of statistics if sense and information is to be extracted from such a source. There are already problems in similarity of the flowers in what must surely be very divergent (geographically and vegetatively) species and to expect to then find differences between elements that are otherwise similar is very optimistic if not plain fantasizing. The similarity of the flower of what I named H. pulchella globifera to that of H. cymbiformis(cooperi?) var incurvula never ceases to amaze me. Similarly I could not find differences in the flowers of H. angustifolia, H. monticola, H. chloracantha and H. variegate. Another similarity was an exact copy of a flower of H. mutica and that of H. pygmaea. One aspiring taxonomist has claimed that studying the flowers of the Kaboega population set (all H. cooperi to my mind) will resolve that species issue. I think this is also a fantasy.

1. THE PLANTS.

*6509 H. mirabilis meiringii,* W Bonnievale

6509 H. mirabilis meiringii, W Bonnievale

6509 H. mirabilis meiringii, W Bonnievale

7176 H. mirabilis, Agtervink

7176 H. mirabilis, Agtervink

7269 H.mirabilis meiringii, Edendale, Bonnievale

7269 H.mirabilis meiringii, Edendale, Bonnievale

Appendix 2.

This appendix illustrates plants and flowers and buds for the following populations:-

7985 H. mirabilis, Sandberg, S Worcester 6668 H. mirabilis, N Ashton
6672 H. mirabilis, Koningsrivier, Robertson 6875 H. mirabilis, Cogmanskloof
7986 H. mirabilis, E Montagu

Both the Sandberg and Montagu populations flowered very poorly. The Ashton and Cogmanskloof populations are known to be late flowering and I have observed this previously for a Montagu population. The Koningsriver plant is a single clone that I have had for many years.

BUDS:

7985 H. mirabilis Sandberg, 6672 H. mirabilis S Worcester Koningsrivier, 7986 H. mirabilis Robertson E Montagu

6668 H. mirabilis, N Ashton 6875 H. mirabilis, Cogmanskloof

PLANTS:

7985 H. mirabilis, Sandberg, S Worcester

7985 H. mirabilis, Sandberg, S Worcester

6672 H. mirabilis, Koningsrivier, Robertson

6672 H. mirabilis, Koningsrivier, Robertson

7986 H. mirabilis, E Montagu

7986 H. mirabilis, E Montagu

6668 H. mirabilis, N Ashton

6668 H. mirabilis, N Ashton

6875 H. mirabilis, Cogmanskloof

6875 H. mirabilis, Cogmanskloof

Appendix 3.

5b.1 7910a, 5b.1 7910b – H. floribunda, Niekerkshek – size comparisons

5b.2 7980a, 5b.2 7980a – H. mirabilis, S. Greyton – size comparisons

Diagram of flower to show face and profile. The base of the tube is how they can vary in one population. The perigon may narrow gradually stipe-like to the pedicel, or abruptly. The tube may have a pronounced gaster as in Gasteria or the profile may be smoothly curved with no broadening beyond the perigon. The outer leaf margins may meet or be separated to expose the inner midribs. The basal separation of the petals may be quite deeply indented or relatively even.

The dotted lines are suggested as base lines for possible mensuration and computer modeling. ♦

Haworthia flowers – some comments as a character source, part 1

Haworthia flowers – some comments as a character source, part 2

Haworthia flowers – some comments as a character source, part 3

Haworthia flowers – some comments as a character source, Appendix 4

Posted on May 12, 2012 by Bruce Bayer

No conclusion is written to this appendix. It is evident to me that there just is no way for me to generate a composite picture for each set of flowers by which one can adequately compare. In general the summer flowering cf. H. retusa set have flowers that are correspondingly lighter coloured than those of the darker flower of the correspondingly darker H. mirabilis set. While gathering the plant pictures from files it struck me how similar plants are from quite different sets across both groups and persuading me that H. retusa and H. mirabilis are in fact one integral system and H. mutica and H. pygamea are western and eastern offshoots.

Acknowledgement
I must again appreciate the kindness and helpfulness of landowners in allowing access. Neels and Suzanne Smit of Volmoed; Hector Odendaal of Dankbaar; Jon and Cindy Webber of Klipport; Tom,Trevennan and Hesphia Barry of Van Reenens Crest, and Coetzee and Surieta Uys of Morning Star. The present owner of the Bromberg site could not be established. The other plants photographed are not field plants and neither can they be seen as any more than flowers of one or few cultivated specimens with provenance already acknowledged.

The following additional populations and plants were observed:-

1. MBB7954 H. mutica, Volmoed.
2. MBB6512 H. mutica, Klipport.
3. MBB7741 H. mutica, Dankbaar.
4. & 5. MBB7920 and MBB7921 H. retusa (nigra), Van Reenens Crest.
6. MBB7803 H. rossouwii, Morning Star, S. Heidelberg.

In addition I include pictures from plants in cultivation:-

7. MBB7758 H. retusa, Skietbaan, S Riversdale.
8. MBB7780 H. retusa (geraldii), Komserante, E Riversdale.
9. MBB7781 H. retusa (foucheii), Komserante, E Riversdale.
10. MBB7776 H. retusa, Pienaarsrivier, W. Riversdale.
11. MBB6747 H. pygmaea, Vleesbaai.
12. MBB33 H. comptoniana, Georgida.
[ed.] MBB33 doesn’t exist in Bayer’s Accession, correct reference is most likely KG114/72

The population H. rossouwii (elizeae) at Bromberg, Stormsvlei was visited but the plants were only in very early bud.

The Heidelberg H. rossouwii is a small solitary plant and only known to me from two populations. H. rossouwii (elizeae) is also small, proliferous and very problematic. It is so far only known to me as one very restricted population. I doubt if the eastern end of the Riviersonderend Mountains have ever been adequately explored to exclude it from there. Even the Bromberg alone could yield more populations.

MBB7954 H. mutica, Volmoed.

2. MBB6512 H. mutica, Klipport.

3. MBB7741 H. mutica, Dankbaar.

MBB7920 Van Reenens H. retusa (nigra), Crest.

MBB7921 Van Reenens H. retusa (nigra), Crest.

6. MBB7803 H. rossouwii, Morning Star.

7. MBB7758 H. retusa, Skietbaan, S Riversdale.

8. MBB7780 H. retusa (geraldii), Komserante, E Riversdale.

9. MBB7781 H. retusa (foucheii), Komserante, E Riversdale.

10. MBB7776 H. retusa, Pienaarsrivier, W. Riversdale.

11. MBB6747 H. pygmaea, Vleesbaai.

12. MBB33 H. comptoniana, Georgida.

Haworthia flowers – some comments as a character source, Appendix 5

Posted on May 12, 2012 by Bruce Bayer

Haworthia mutica, Sandrift, Drew MBB8018 = KG226/70 = JDV92/64

This is an unusual population in that it is the most northwestern one known although only about 15km from Klipport nearer Stormsvlei. It is the source of the single plant that developed gross milkiness and for which I coined the name “Silver Widow”. The population was first recorded by a Mr. Meiring who worked as a nurseryman for the Bonnievale Nursery of Hurling and Neil in the years at least prior to World War 2. There is a letter in the Compton Herbarium archive which records a transaction between Meiring and Triebner of 100 plants as 1s ea. (i.e. = 10 cents in today’s currency.). I really struggled to find plants again in a very disturbed area and eventually did so after I had placed “Silver Widow” there in her pot to see if the flowers would be pollinated in situ. Indeed they were and then found about 20 plants in a very small area nearby. This visit owes to the report by Jakub Jilemicky of still more plants a very short distance again from these. Here there are about 80 plants in an area of about so many square meters.

Illustrations of the plants and flowers are given. The only comment I can make is to repeat that I never ignored the flowers out of ignorance. The fact is that we have a number of pretentious experts who have no concept of species other than an inherited vague notion that species have something to do with the capacity to interbreed. The fact that Haworthia plants do so freely does not handicap their enthusiasm for new species either. So this is a bit of irony. My contention was that if you did consider flowers there would be a lot less species than even in my conservative approach. In fact I am quite sure a competent professional taxonomist might well consider that I have been too generous with species and too kind to my critics.

(image files mislabeled as 8012, but are 8018 – ed.)

Acknowledgement
I thank Gerhardt Swart for access to the site and to Jakub Jilemicky for informing me of the new find. ♦

Haworthia flowers – some comments as a character source, Appendix 6

Posted on May 12, 2012 by Bruce Bayer

Two further records and data for Haworthia mirabilis.

Kobus Venter drew my attention to a second population of H. mirabilis on the eastern boundary of the farm Schuitsberg, which is the origin of the var. beukmannii of von Poellnitz. But the real motive for exploration in that area was a photograph sent to me by Messrs. Daryl and Priscilla Hackland of a plant on the Zigzag Path just east of the northern end of the town of Greyton. It was their son Andy Hackland who observed the plants and thought I might be interested – indeed. The bright green colour of the plant struck me as most unusual for that area where most of the populations known are south of the river in shale and they are of the brown and red hues. This is not strictly true because records of an expedition by Messrs. Beukman, Otzen and others mentions a small “black” species just before the entrance to Greyton. I have never found that.

However, it is well known that Haworthia mirabilis occurs along the course of the Boesmansrivier on the way to MacGregor. I once observed it there myself hiking part way from the northern end of the walking trail down the river to Greyton. Kobus Venter has also seen it while walking from the southern end. I included this variant under the varietal name ‘consanguinea’ that is actually based on populations on the north side of the mountain in the Dwarswaterkloof west of MacGregor. Ms. Dawn Schwegmann once showed me plants from that general area but I never did establish exact origin. What did strike me about the Boesmansrivier plants is the very close resemblance to Haworthia retusa ‘turgida’ when I still considered populations that occur in Table Mountain Sandstone in the Tradouw Pass and north of Riversdale as a discrete species. The only difference to me was the summer flowering time and the resemblance of the flowers to those of H. mirabilis from the Bredasdorp/Napier area. This observed similarity was largely based on a very small sample, colour, and the fact that the flowers were available simultaneously. I must have assumed that this similarity was unlikely to include ‘turgida’. Unfortunately photography was expensive and almost unmanageable for this sort of detail in that era and I had to observe and record from memory.

While I now have only the memory of the turgida-like element in the pass and a myriad of varying populations of H. mirabilis at the northern end and around MacGregor, I have had no evidence of transition to the red-brown plants (eg ‘rubrodentata’) south of the Riviersonderend River from east of Genadendal to at least Lindashof east of Riviersonderend town. This is as the species occurs in putatively Bokkeveld strata.

The Hacklands saw the plants at the top end of the Zig Zag path and the geology here will need a better description than I can give without a geology map. The vegetation is transitional to Fynbos and the stratum an argillaceous one (clayey) not obviously Sandstone nor shale. But the plants are so different. They are smaller than in populations in the area that I know south of the river, fairly bright green and becoming rather yellow in summer stress dress. A physical description of the plants defies description and I would be a loss to refer to the plants as anything but a variable lot. In the amazing context of Latin names in Haworthia I would like to refer to the plants as Haworthia mirabilis ‘hacklandii’ – if it can be forgotten and forgiven (it unfortunately cannot be) that I question the merit and motives for using personal names for taxa. In any case, the serious limitation is to know how to descriptively and sensible circumscribe such a population. The alternative reference is H. mirabilis MBB8040 or H. mirabilis NE Greyton, or H. mirabilis Zig Zag Path. Whatever name is used, this population is in my opinion quite an awesome demonstration of the morphological continuity that I have come to expect in Haworthia. A wide range of comparisons is possible if one takes single clone after clone. I thought ‘sublineata’, ‘mundula’, ‘ turgida’ (from a range of populations) as well as ‘notabilis’ and even ‘elizeae’ and ‘mutica’). The departure from the wider dark green colour of the mirabiloids is the most unexpected feature although this is just what occurs with ‘consanguinea’. This latter is the ground truth and I fully expect that further populations occur to complete a transition to the even more obviously ‘turgidoid’ variant known from deeper in to the Boesmansriver valley. Later driving eastwards and a little further south towards Schuitsberg, we observed very similar habitats 2-3km further north of the road in foothills of the Riviersonderend Mountains.

Among the pictures of the plants is a view from the top of the Zigzag Path looking north into the Boesmansriver valley (or gorge). The hill on the right front seems to be similar to that of the Path and similar low rocky hillsides occur at several places. Significantly there is one immediately north of Riviersonderend where the river passes by on the north side rather than on the south away from the higher hills away from the main range. Kobus has persistently observed that there must be plants there. The evidence of the Zigzag Path occurrence definitely suggests that this must be so and that many more populations exist unrecorded simply because of lack of exploration specifically for Haworthia. While Andy observed the plants at the top of the Path, they are widespread all the way down to the bottom. It is quite amazing that the presence of the plants has not come in to my ken sooner. It is just an observation of mine that Haworthia distribution may be far wider than we suspect simply because Haworthiophiles have not ventured into unlikely areas.

This is a view northward from the top Zigzag Path to the Boesmanspad (Biushmans Route) that runs towards MacGregor 25km to the NE - and to H. mirabilis 'consanguinea'. The weathered cardboard in the foreground is a very sad but loving tribute by children to their grandfather - messages of love and longing painted on two little stones, on the remains of a shrivelled dried wreath.

Flower profiles, faces, and buds

From Greyton we went to Schuitsberg driving further east than previously. This population seems to have led to the speculation that the type of H. mirabilis as illustrated in Botanical Magazine t1354 (1811) could have come from here. In a recent article in Alsterworthia, Gerhard Marx attempts to ridicule my typification of the name ‘mirabilis’. I frankly think this is much more a point scoring effort than a genuine attempt to arrive a taxonomic truth to which Marx claims to aspire. Apart from speculated similarities of some or other clone to that picture, is the fact that this is an old transport route dating to that era. There is a plaque at that site dated 1988 commemorating a 300year history. The problem for me is the variation among the plants that renders the nomenclatural system and the typological approach to naming dismally doubtful. There is just no Haworthia mirabilis var. mirabilis.

Schuitsberg is the origin of H. emelyae var. beukmannii von Poellnitz .I associate that name and description with an exceptional robust, compact and retused version of H. mirabilis (a version with much longer erect leaves with spined margins viz var. rubrodentata {red teeth} was named from further west along the river). I visited Schuitsberg and adjacent farm Nethercourt in the early 1970s. The plants I saw were indeed robust. However I visited there again 2012 and found the place previously visited destroyed and no plants among the many in the adjoining rocky slopes that resembled what I perceived to be ‘beukmanii’. This is repeated in my experience of this second locality on Schuitsberg east (MBB8041) of where I knew the plants. One can refer to the many variants illustrated with this article. Are these all individually or collectively closer to any early illustration on which the name ‘mirabilis’ can be based? It was only a single large very robust plant with leaves nearly 30mm wide and 20mm think that I would have said also agrees with the original acceptance of the “element”. So what can we say here of all these variants? This is a fact of the retusoid species and close relatives. They are simply highly variable with respect to leaf shape, arrangement, markings, armature, and colour, as well as flower characters. The consequence of these is gross aberration of the formal nomenclatural system and whole host of Latin names that are very useful to collectors, commercial nurseries and any other activity where a formal impressive name adds value. What we have at a place like Schuitsberg is that we have plants that can be labelled ‘beukmani’ because they might look ostensibly like the original described plant or its image, or just because it happens to come from Schuitsberg.

Flower profiles, Faces, and Buds.

Regarding the flowers of both localities that I illustrate here, there is again the high variability that I discuss and illustrate in all the preceding parts of Haworthia Update 8. What we now need is a computer wizard who can perhaps explore the possibility of computerised pattern recognition. In this way to look at trying to arrive at a composite single image that captures the identification of a population. It boggles my mind that someone might think, say ‘beukmannii’, is an array of plants scattered through many populations to so constitute a species.

I have tried to approach this whole problematic, provocative, disputatious subject from a rational objective view based on my own education, experience and acquired knowledge to place it in an environment acceptable to formal botany. That environment has not been all that helpful and conducive to a solution either.

Acknowledgement
I would like to thank Dr. Daryl and Mrs. Priscilla Hackland, and Andy Hackland for information and assistance in visiting the Greyton Zigzag Path population. Mr. P.G. Viljoen not only kindly allowed us access to Schuitsberg but took the trouble to drive us there to show us a wider view of the localities and also the commemorative plaque that mark the old travel route. ♦

Haworthia flowers – some comments as a character source, Appendix 7

Posted on May 12, 2012 by Bruce Bayer

Three further records and data for Haworthia mirabilis.

The populations covered here are:

6631 H. mirabilis ‘mundula’, Mierkraal, SW Bredasdorp.
6635 H. mirabilis ‘badia’, NW Napier.
6639 H. mirabilis ‘sublineata’, S Bredasdorp.

There has been some published comment about the distinctiveness of these three populations stating that I do not recognise this because I treat them as variants of single species. The implication is that I do not see any difference. This is quite bizarre. At the present moment I have a digital library of 165 H. mirabilis populations and this is by no means all there are. There is an enormous amount of variation both in and between these populations. If the specified three are to be recognised as species, it means that there is a wholly unrealistic number of species quite out of keeping even with an already highly diverse set of species of the Cape Flora. Furthermore, as I point out in appendix 6, there is no typical variety of H. mirabilis in any practical sense because the variability in the population designated by me, as well as that of Schuitsberg apparently preferred by another, precludes it.

6631 Haworthia mirabilis ‘mundula’, Mierkraal, SW Bredasdorp.
6631 as a variant ‘mundula’ is only known at this one locale. However, similarities can be found elsewhere. The name was introduced by G.G. Smith without any existing recognition of the prior name H. mirabilis and where that may have been applied. Because only illustration exist for which artistic license must be invoked, there is some doubt about just where the original plant may have originated. Frankly I do not think this even matter because the fact of the individual variation that renders the automatic recognition of typical varieties in Haworthia meaningless.

6635 Haworthia mirabilis ‘badia’, NW Napier.
6635 as a variant is generally thought to be only one locale too. It in fact can be seen to be part of a continuum that extends nearly 4km westwards in the same geological stratum. Furthermore it is complemented by quite different variants in at least seven known populations to the north and east. Exploration is by no means complete.

6639 Haworthia mirabilis ‘sublineata’, S Bredasdorp.
6639 as a variant is known immediately south of Bredasdorp, but has been reported further to the southeast in the same stratum. However, there were two populations north of the town no longer extant. Illustrations are available in the literature and herbarium. Based on these, the same illogic that suggest separation of the three specified above would suggest that these are also discrete species. Further populations exist to the near north and northeast of Bredasdorp that intensify and add to the problem of rank. This population, its variation and the statistical difficulties of explaining this variation is discussed in Haworthia Update Vol.4:62-77.

[ed.] flower files names are 6635, but are 6639.

The present appendix represents another set of plant illustrations and also flower photographs. Unfortunately 6631 did not flower very well and only two photographs could be taken. The habitat has changed in the 43 years since I first observed the population. The plant clumps were considerably smaller and there was none of the colour range that I observed in my first visit in 1969 when the plants showed green, yellow and orange.

Observations
The differences in the plants at the three localities are based on three very simple criteria. ‘Badia’ has fewer leaves and no marginal spines, ‘mundula’ has more abbreviated and also numerically more and smaller leaves with a shorter end-area; and ‘sublineata’ has long slender leaves often with prominent venation and larger marginal spines. By these criteria one can nearly always be sure of identification. Nevertheless wider considerations and similarities of one degree or another are so widespread that I consider a stipulated taxonomic distinction impossible and unrealistic. Flower morphology is as variable as the leaf appearances and it is virtually impossible to arrive at a single word or imaged representation of what represents each population. It in fact suggests the restricted view of formal taxonomic recognition that I suggest.

Acknowledgement
I appreciate the co-operation of Mr. E. Conradie of Mierkraal and Mr. P. deKok of Napier for their forbearance. I would be very remiss not to acknowledge the interest in the subject, and input, by Lawrence Loucka who has generated the HaworthiaUpdates.org website and the ISSN 2306-0956 archive for my writings on Haworthia. ♦

Haworthia flowers – some comments as a character source, Appendix 8

Posted on May 12, 2012 by Bruce Bayer

A report on Haworthia mirabilis ‘badia’ / ‘subtuberculata’

I cannot claim that I have resolved the nomenclature niceties around the use of this name and its many varietal names and synonyms. If critically examined even the use of the name “mirabilis” is in doubt and the very original epithet of “Aloe atrovirens” may be nomenclaturally correct. This would create havoc because then the name H. herbacea as typified by W.T. Stearn may best apply. So I put that all aside and use names as I have in my Revision and subsequent publications. I am well aware that there is a problem with the use of H. mirabilis var. mirabilis where it may be thought that the name “mundula” is therefore redundant. I cover all this up with the explanation that there is no typical variety “mirabilis” and actually use just the name H. mirabilis to cover all those populations and variants to which no Latin names exist. I use names like “badia” and “sublineata” as it is generally possible to recognise all or most of the variants from those populations. But names like “magnifica”, “maraisii”, “heidelbergensis”, “rubrodentata”, “beukmannii” and “depauperata” (among others) have no clear and direct application in respect of a population or even a group of variants of any kind. In my Revision I attempt the use of the name “triebneriana” to cover all the variants that are not included under the typical varietal name “mirabilis”. This does not actually work either because the name has its origins at Stormsvlei and there are other populations that are very different from those occurring there too.

My conclusion was, and is, that it is not possible to formally name and describe all the variants in H. mirabilis. Firstly this is true at population level and very much more so in respect of the incredible variation that exists within those populations.

In this appendix I present images of plants and flowers for just two populations…

MBB6987 Haworthia mirabilis ‘badia’. Sandfontein.
MBB7091 H. mirabilis ‘subtuberculata’. Mierkraal, Napier

(Note: – this is not MBB6631 Bredasdorp ‘Mierkraal’ where ‘mundula’. I use now the name ‘subtuberculata’ as one of two generally for that area Northwest of Napier. These are plants with generally erect to suberect leaves, fairly strongly triangulate and tubercled. The von Poellnitz and Triebner names ‘multituberculata’, ‘napierensis’ and ‘turgida’ are also available).

1. MBB6987 Haworthia mirabilis ‘badia’. Sandfontein.

Flower profiles.

Flower faces.

Bud.

2. MBB7091 H. mirabilis ‘subtuberculata’. Mierkraal, Napier

Flower profiles.

Flower faces.

Buds.

The first population is approximately 2.8km west of the “typical badia” population and in almost the same geological stratum and habitat viz. Table Mountain Sandstone with grassy fynbos. The second population is approximately 2.7km northwest of the first and it is on a small ridge of vertically orientated Bokkeveld shale with Renosterveld. There are several populations west and north from here with similar plants. To the east at Napier and north of Napier the plants vary towards smaller and darker versions of H. mirabilis that very loosely indeed have been covered by the name “maraisii”. This is a complete myth and about as accurate as the use of the name “magnifica” to cover all the H. mirabilis variants around Heidelberg and Riversdale. It must be noted that the name “heidelbergensis” is drawn into the arena too and in Appendix 9 I will explain one aspect of the situation with respect to that name.

What this particular report is intended to convey, is the dramatic change related to firstly location, and secondly habitat. I maintain that there is no prescribed species definition in botany and that this is the prime reason for the confusion that exists at every level in botany. Therefore I have arrived at my own view of species as dynamic systems that are not necessarily recognisable in terms of tangible visible characters. There are two important considerations in the way I classify Haworthia. The first is there geographic spatial positions, and secondly the way in which apparent systems relate to each other with respect to those positions. Therefore what I am showing here is that H. mirabilis ‘badia’ is the sandstone version (ecotype) of the species northwest of Napier, and H. mirabilis ‘subtuberculata‘ the shale version a little to the north and then west. H. mirabilis ‘sublineata‘ is a sandstone ecotype south of Bredasdorp and H. mirabilis ‘mundula’ a fairly unique population on tertiary gravel southwest of Bredasdorp. In Appendix 9 I will give an inkling of what happens to H. mirabilis in the shale and tertiary formations north and then west from Napier.

Acknowledgement
Mr. Wynand Wessels kindly accompanied us to the population at Mierkraal. It was difficult to establish just who owns the property at Sandfontein where H. mirabilis occurs and while we met two landowners in the area we were not sure if we were trespassing or not. ♦

Haworthia flowers – some comments as a character source, Appendix 9

Posted on May 12, 2012 by Bruce Bayer

A report on Haworthia mirabilis and Haworthia rossouwii ‘minor’, Rooivlei, NNE Bredasdorp.

In Appendix 8, I explain some of the rationale of my use of names. A detailed report on H. mirabilis can be found in Haworthia Update Vol.3. and in various chapters of the subsequent Updates. It is shown why I consider the possibility that H. mirabilis and H. retusa are in fact the same species.

In this report I will show just one population of many from Napier northwards and westwards for which no broad name exists. There is simply a transformation from elements that I refer to as H. mirabilis ‘subtuberculata’ in Appendix 8, to a very wide range of populations in the central Southern Cape. Both the names “maraisii” and “heidelbergensis” have been applied to populations in this area. The name H. maraisii var. simplicior from Napky may even be relevant but its application, simply irrational.

The populations for which data is presented here are…

7334 H. miriabilis NE Bredasdorp
6638 H. miriabilis Rooivlei
KG36-70 H. miriabilis ‘minor’ Rooivlei
7821 2013 H. mirabilis x rossouwii ‘minor’
7822 H. miriabilis Rooivlei E of Rd
8044 H. miriabilis with H. rossouwii ‘minor’ KG36-70
8045 H. rossouwii ‘minor’ Rooivlei

The population 7334 H. mirabilis is covered in a general way by the name “maraisoid” that simply denotes smaller darker green versions of H. mirabilis. It is not accurate and the terms “floribundoid” and even “heidelbergensoid” are applicable. A complication among many is “hammeroid” for what appears to me as introgression of H. mutica with H. mirabilis. It is just not possible to simplify and summarise the range of populations and variants that lead on from this close to Bredasdorp (and “sublineata”) on to Rooivlei, then north and eastwards to eventually cover “hammeri”, “bobii”, “paradoxa”, “jakubii”, “Windsor”, “magnifica”, “atrofusca”, “toonensis”, “scabra”, “vernalis” and so on ad infinitum. So these pictures together with 7822 just serve as a reference point for what follows.

1. MBB7334 Haworthia mirabilis, NE Bredasdorp

Flower profiles.

Flower faces.

Bud.

It is Rooivlei approximately 26km NE Bredasdorp that concerns me here. It is quite a high-lying area in the wheat growing area of the southwestern Cape. The geology is primarily Bokkeveld shale with inselbergs of later tertiary origins. At Rooivlei the highpoint has tertiary remnants and the Bokkeveld strata include sandstone among the truly clay derived shales. While the vegetation is broadly renosterveld, there is even a remnant of what Acocks’ described as Valley Bushveld. Curiously is the presence of Acrodon quartzicola first described from there as well as Gibbaeum austricola much more widely distributed even to southwest Heidelberg. From the Haworthia aspect, H. marginata is present. It is the presence of a vicariant population of H. retusa ‘turgida’ that is really odd because it is known again about 40km away to the east and in the Breede River river channel.

What is significant is a plant illustrated as no.12 in 6638. I have seen a plant just like this where H. mutica occurs in the immediate vicinity of H. mirabilis at Klipbankskloof about 20km north of Rooivlei. The illustration referred to my well be a hybrid involving H. retusa ‘turgida’. H. mutica is not known closer than Hasiesdrift (Soesriver) about 6km away.

2. MBB6638 Haworthia mirabilis, Rooivlei

Still more significant now is what I refer to formally as H. rossouwii var. minor. This is KG36/70 originally described by me in my Revision of 1999 but as H. heidelbergensis var. minor. I did explain at the time that there were similarities to H. mirabilis var. sublineata. Since then when I did not know all that much, I have explored a great deal and it is clear that the greater picture is that H. heidelbergensis is truly a variant of H. mirabilis. Because the variant ‘minor’ occurs with H. mirablis it forces a rethink. I did this eventually when H. rossouwii was re-discovered at three locations north-northwest and north of Bredasdorp (forcing the abandonment of the name H. serrata) and not very far from Rooivlei. This is really odd in respect of the distribution and variability of Haworthia because I had described H. serrata from near Heidelberg far to the east. I now have records for 12 populations in that area. This distribution brackets that of ‘minor’. My suggestion thus was that it was better a variant of H. rossouwii than H. mirabilis that now included H. heidelbergensis.

3. KG36/70 Haworthia rossouwii var. minor, type locality

KG36-70 looking northwards

KG36-70 habitat viewed from the west.

4. MBB7821 Haworthia mirabilis x rossouwii ‘minor’

Flower profiles.

Flower faces.

Buds.

View looking east of north

A few years ago I revisited Rooivlei to assess the presence of H. marginata there. I did not find it, but instead noted a population 7821 west of KG36/70 that was very unusual and I suspected it had something to do with both H. mirabilis and H. rossouwii ‘minor’. The object of my recent visit to Rooivlei was to examine this a bit more closely with H. mirabilis in flower. I do not have a record for the flowering time of H. rossouwii ‘minor’ but have memorised it as November. In my visit of 12^th Feb. Almost the first plants we found were 2 plants of H. mirabilis (8044) in flower in the immediate vicinity of KG36/70 that was not in flower. The population (7821) west of this is very extensive and the plants extremely variable. They were mostly in flower and there were odd large plants suggesting hybridization with some other species. To the northwest we again found plants (8045) like those at KG36/70 in the same more prominent vertical shale ridges as at the original locality. There was an indication that these plants were not flowering in synchronicity with 7821 and had in fact already flowered. But the plants overall were intensely variable as can be seen from all the illustrations.

5. MBB7822 H. mirabilis Rooivlei, east of Road

6. MBB8044 H. mirabilis with H. rossouwii ‘minor’ KG36-70

Flower profiles.

Flower faces.

Bud.

7. MBB8045 H. rossouwii ‘minor’ Rooivlei NW of type locality

Flower profiles.

Flower faces.

I therefore hypothesize again that what is occurring at Rooivlei was/is introgression of H. rossouwii and H. mirabilis. The area is quite extensive and I have not explored it in its entirety. I would not be surprised if H. rossouwii in its more “typical” form is found there.

Acknowledgement.
I would like to thank Mr Francois Uys for access to Rooivlei. Florent Grenier accompanied us there. Lawrence Loucka kindly and patiently brought some loose ends together.

Haworthia flowers – some comments as a character source, Appendix 10

Posted on May 12, 2012 by Bruce Bayer

Appendix 10 – An additional report on Haworthia mirabilis and Haworthia rossouwii ‘minor’, Rooivlei and Brakkloof, N and, NNE Bredasdorp.

The previous report indicated the necessity for further exploration of Rooivlei. I had observed H. mirabilis, but not reported it in Update 3, at Brakkloof to the west in 2004. So the object of this appendix is to remedy this oversight and to also cover more area of Rooivlei. At Brakkloof there are several small remnants of rocky shale and we located several populations, while at Rooivlei we actually explored the very western boundary. This constitutes the same topographical area as the Rooivlei populations but the plants we observed were factually on Brakkloof.

The populations reported on here are:-

6537 H. mirabilis, Groudini, W Napier
7285 H. mirabilis, Brakkloof 3
8046 H. mirabilis, Brakkloof 2
8047 H. mirabilis, Brakkloof 1
8048 H. mirabilis, W Rooivlei 1
8049 H. mirabilis, W Rooivlei 2
8050 H. mirabilis, W Rooivlei 3
8051 H. mirabilis, E Rooivlei
8045+ H. rossouwii ‘minor’, NW type locality
8052 H. mirabilis, S.Welgegund
8053 H. mirabilis, Welgegund SE 8052

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A myth corrected to – Haworthia maculata var. livida (Bayer) Bayer – and flowers ignored

Posted on May 18, 2012 by Bruce Bayer

(Haworthia maculata var. livida (Bayer) Bayer, comb.nov. H. pubescens var. livida Bayer in Haworthia Revisited, p.134, 1999, Umdaus) Type: Cape-3319 (Worcester): S Lemoenpoort (-CD), Bayer 1128 (NBG, Holo.).

I described Haworthia pubescens var. livida in Haworthia Revisited (Umdaus, 1999), in the full knowledge that it was in a twilight zone of inadequate information. It is a good example of how Latin names give plants a false reality. The system forces decision making without any slack being cut for doubt. This is thus a good opportunity to demonstrate what inexperience and ignorance add to the process of classification. In the small area along the Breede River north of the Brandvlei Dam near Worcester, the species H. herbacea, H. maculata and H. pubescens grow in close proximity. H. herbacea is ubiquitous throughout the Worcester/Robertson Karoo, while H. maculata has a curious distribution in that area. It occurs at widely separated localities on the western fringe of H. herbacea and I have wondered about its relationship to that species because of the similar flowers and flowering time. H. pubescens is only known from a small set of low ridges east of the Brandvlei Dam where it grows in close proximity to H. herbacea. It also has similar flowers but it flowers a little later in late spring as opposed to early spring.

When I first discovered plants at Lemoenpoort (Hammansberg) about 20km further south I found very few plants and the only thing I could do with them was regard them as in the same context of the Breede River species and hence their description as a variant of that H. pubescens. H. herbacea is very abundant in this area too but almost exclusively in Bokkeveld Shales and Dwyka Tillite. The doubtful plants at Lemoenpoort occur in Witteberg Sandstones. A population of plants was also known from the top of the Moddergat Mountain (Ouhoekberg) to the west that created some problems for me as it seemed to relate to the absence of H. mirabilis from both the Hammansberg and Ouhoekberg. Not only that but it also looked very similar to H. maculata. H. mirabilis is known at its closest east of these two mountains at several localities along the eastern ridges of the Ouhangsberg and Droogerivierberg mountains that are in turn east of Hammansberg. H. maculata is very abundant along the long ridge of low hills north of the Brandvlei Dam. Very similar plants occur high in the Audensberg peaks north of Worcester and also lower down in the Kanetvlei (Sandhills) area west of Heatlie Peak. These hills are very difficult to explore and I speculate a connection to a population at Buitenstekloof to the east. There is a population here that obfuscates the interpretation of the name H. intermedia VPoelln. I lumped this population under H. mirabilis var. notabilis. I have to rely on memory here because having set flower morphology and flowering time aside (without completely ignoring them), I did not have the methodology in place to record detail. The flower was rather similar to H. herbacea and H. maculata and it also flowered late spring. I was not, and am not, sure that there is not a connection between the Buitenstekloof plants and the Kanetvlei plants.

It is unfortunate that Gerhard Marx has expressed his opinion that I have ignored flowers in my classification of Haworthia. While there is no doubt that this is partially true, there are very good reasons why I did this. The main reason is the historic one. It was understood and accepted that the flowers in the species as they were then recognized, were very similar. Even the subgeneric differences were not seen to be significant. My own early attempts at classification also indicated that the flowers were not going to be of assistance at the level at which differences between populations and species were being recognized. In fact I had observed similarities of the flower in populations that were clearly different taxonomic entities. Conversely I observed big floral differences in populations that I imputed to be the same “species”. Because of the added seasonality of flowers, I thus put them aside to focus on exploration and explanation on the basis of the vegetative structures and geographic distribution. Marx’ observation therefore comes at a time when that work is substantial and flowers can now be used to clarify and verify relationships. My conviction is that those relationships are best understood in relation to distribution and we are only now in a position to assess what value the flowers may add to those interpretations.

This article thus reports on the exploration of the Ouhangsberg, Hammansberg and Moddergat (Ouhoekberg) mountains as well as some work done in the Brandvlei Dam area. Again I put flower and flowering time aside – for the moment. This is in the same way I am inclined to the considered view that geographic information and field observation are the essential ingredients whereby ANY method has to provide explanation that satisfies physical experience. The alternative is that what we see and experience physically and mentally, is just the illusion that metaphysicists ascribe to creation.

Populations recorded

The following populations (see map) were found and observed:-

1. KG669/69 N Brandvlei Dam – set 1
2. MBB164 N Brandvlei Dam – set 1
3. MBB1119 (and 6815) Audensberg – set 2
4. MBB1120 (and 7994) Sandhills, Kanetvlei – set 2
5. MBB1128 (and 7066) W Lemoenpoort (type locality var. livida) – set 3
6. MBB1145 Ouhoekberg, Moddergat – set 5
7. MBB2591 NE Brandvlei Dam – set 1
8. MBB4461 (and 6514) Buitenstekloof – set 6
9. MBB7270 Ouhoekberg W – set 5
10. MBB7266 E Lemoenpoort – set 4
11. MBB7271 Cilmor Winery, Dewetsberg – set 7
12. MBB7526 Die Nekkies W. Brandvlei Dam – set 1
13. MBB7991a Ouhoekberg E – set 5
14. MBB7991b west Ouhoekberg E – set 5
15. MBB7992 Hammansberg midpoint – set 8
16. MBB7993 Hammansberg W – set 8
17. MBB7997 Sandberg H. pubescens, type locality – set 9 are for MBB7997 cf H. pubescens N of point 11
18. MBB7995 and 7996 Brickfield H. herbacea ‘submaculata’ – set 10
19. MBB7984, 7985 and 7988 Droogerivierberg, H. mirabilis – set 11
20. Unknowns – set 12

**Map references for Haworthia maculata.**

There are 4 records for the Brandvlei Dam but in Update 6, I report on H. maculata from the pump station east of the Resort, very abundant to the eastern point of Die Nekkies. Three of the records are for west of the Resort while I have not confirmed Etwin Aslander’s observation that he had seen it at the extreme western end of Die Nekkies. G.J. Payne in 1970 communicated to me that he had seen a Haworthia south of the Brandvlei Dam above and west of the Brandvlei Hot Spring. I was unable to confirm that in a single brief visit there ca. 1976. I have not been able to relocate plants at Sandhills but have no doubt that it will be found there if more thorough and extensive fieldwork is done there. In fact a very important consideration is that there is still enormous scope for exploration.

On each visit to the Ouhoekberg I have found plants in different local populations and none of the last four records is the same as seen in my first visit. At the original Lemoenpoort I always struggled to find plants at all, and the same applies to the locality to the southeast. At my last visits to these two places there have been many plants but relatively difficult to find.

H. herbacea is abundant in the Hammansberg and Ouhangsberg area as well as east of Cilmore. It has not been observed near the Ouhoekberg. It occurs between Cilmor and Die Nekkies in a larger form that acquired the name H. submaculata V.Poelln.; and I did suspect that there was a continuity with H. maculata that does not now seem to be the case. I exclude H. pubescens from this report. It occurs with H. herbacea east of Cilmor, but it also occurs on the northeastern slope of Dewetsberg north of the H. maculata recorded habitat. There are/were plants of H. mirabilis vegetatively very similar to H. pubescens at the Breede River Bridge southwest of Roberson. Their flowers and flowering times (March) correspond with H. mirabilis in that area. H. pubescens flowers in November (later than H. maculata at Die Nekkies or of H. herbacea – Sept/Oct) and the flower is similar to that of H. herbacea. It seems fairly obvious that H. pubescens fills the gap for H. mirabilis and the flower and flowering time are problematic.

The plants illustrated:

Set 1. Map points 1, 2, 7, 12. Die Nekkies
Figs. 1.1 to 1.5 7526 H. maculata. Die Nekkies.
Figs 1.6 to 1.10, sn* H. maculata, Die Nekkies – East.

A chapter in Update 6 deals with H.maculata – Die Nekkies Biomes, and Haworthia maculata These few images give scant insight into the variability in respect of rosette and leaf form, colour and marking, and proliferation.

Set 2. Map points 3, 4. Audensberg and Sandhills
Figs. 2.1 to 2.5 MBB6185 H. maculata. Audensberg.
Figs. 2.6 to 2.36 H. maculata, Vreesniet, Kanetvlei.

Unfortunately I found non-digital photography to be costly, cumbersome and unmanageable. So I have no field images for either population on the high Audensberg or at my original Sandhills site. However, I revisited the area and found the plants quite abundant a little further north at Vreesniet that is also Kanetvlei/Sandhills. The plants were almost confined to narrow rock cracks and their presence suggests to me that there is a very probable connection now eastwards to the Unknowns and even further on to the Osplaas H. arachnoidea, Etwin Aslander’s Hex Pass plants, and then still further to the H. marumana ‘dimorpha’ questionable.

Figs. 2.1 to 2.5 MBB6815 H. maculata. Audensberg.

Figs. 2.6 to 2.36 H. maculata, Vreesniet, Kanetvlei.

♦

A myth corrected – part 2

Posted on May 19, 2012 by Bruce Bayer

Set 3. Map point 5.
Figs. 3.1 to 3.34 MBB7066 H. maculata, Lemoenpoort.

This is the type locality (i.e. MBB1128, W. Lemoenpoort) for H. maculata var. livida that I think is largely untenable or unnecessary fragmentation. Lemoenpoort is the valley that separates Hammansberg from Ouhangsberg. The first few plants seen were in exposed situations and had the purplish or bluish-grey colour that prompted the Latin name. This was maintained in cultivation. I linked it at the time to H. pubescens that seemed more probable at the time than to H. maculata and then simply because of the perceived demands of a relatively inflexible nomenclatural system. There are now seven new populations that contribute and improve understanding of H. maculata as a species. All these localities are in Witteberg Sandstone and the type locality is only slightly different in that the stratum of rock is less feldspathic (i.e. mineralized).

Set 4. Map point 10. SE Lemoenpoort
Figs. 4.1 to 4.28 MBB7266 H. maculata SE Lemoenpoort.

This is a more west facing slope and also relatively low down on the mountain side. The plants were in rocks bordering an eroded softer shale band. They probably also occur higher on the slope in less mineralized sandstone. Generally the plants were very cryptic and tended to be solitary.

**MBB7266 SE Lemoenpoort, view to southwest**

♦

A myth corrected – part 3

Posted on May 20, 2012 by Bruce Bayer

Set 5. Map points 6, 9, 13, and 14. Ouhoekberg.
Figs. 5.1 to 5.24 MBB7991a H. maculata, Ouhoekberg E.
Figs. 5.25 to 5.28 Panoramic views.
Figs. 5.29 to 5.33 MBB7991b H. maculata, Ouhoekberg.

These populations were first observed as one on a higher eastern point of the Ouhoekberg above Moddergat in about 1975. George Lombard accompanied Kobus Venter and me there in 1996 and we found them on the western high point. I located them nearby in 2004. We found them again recently as two more populations on the eastern heights near where I must have observed them first. Note the reticulation in the dried fruit capsule. It can be very much more evident in species like H. pulchella but I seriously doubt its diagnostic value. I have included views to… (a) north to show the water of the Brandvlei Dam in the far distance. The area between has not been adequately explored. What is interesting is the geology. The mountains on the left are the Table Mountain Sandstones, nearer is the valley where the soft Bokkeveld Shale has been eroded away, and then comes the Witteberg Sandstone with a neck of soft shale, and then low down is Dwyka tillite. H. herbacea is present on the Dwyka outcrop barely visible in the middle right. (b) The second view is looking east at first the Hammansberg and beyond that Ouhangsberg with H. mirabilis on its eastern flanks. The view looking southeast is over low Bokkeveld shale ridges with an abundance of H. herbacea. The view south is to Villiersdorp where Wolfkloof (not the Robertson Wolfkloof) is a deep valley behind the Table Mountain Sandstone left of the gap through the Rooihoogte Pass. Here is where H. herbacea ‘lupula’ occurs, unusually in sandstone. Altitude and skeletal soils of different origins contribute hugely to the genetic mosaic. Arable depositional soils exclude Haworthia and obviously inhibit contact between populations.

Set 6. Map point 8. MBB4461 and 6514 H. mirabilis ‘notabilis’. Buitenstekloof

I am sorry that I have probably contributed to the confusion here. It started with von Poellnitz with a species he described as H. intermedia from this locality where both H. mirabilis ‘notabilis’ and H. reticulata occur, but he included E. Cape H. cooperi in the synonymy so it is not quite clear what he meant. While H. reticulata can be easily recognized for what it is at Buitenstekloof, the other population I ascribe to H. mirabilis ‘notabilis’. But I am not certain because I recall it flowering in November and with a flower type nearer to H. maculata than to H. mirabilis. It appears to me that further exploration may expose continuity westwards to the Sandhills/Audensberg populations of H. maculata and even northwards to unknowns.

H. mirabilis ‘notabilis’ (with or without the inverted commas) occurs from Klassvoogds between Ashton and Robertson, westwards through Wolfkloof, Robertson to Agtervink and further westwards still to Buitenstekloof. This all along the great Worcester fault line and intrusive igneous rock. The dolomite is apparently difficult to explain geologically and is associated with the granite-like extrusions in the area.

♦ .

A myth corrected – part 4

Posted on May 21, 2012 by Bruce Bayer

Set 7. Map point 11. Cilmor.
Figs. 7.1 to 7.3 MBB7271

An interesting point here that H. herbacea (map point 18) occurs between this point and all the Die Nekkies populations. I am not sure that the area between can be fully explored although I have been into it.

Set 8. Map points 15 and 16. Hammansberg.
Figs. 8.1 to 8.21 MBB7992 H. maculata, Hammansberg.

The first locality 7992 midway between the eastern and western ends of the mountain was very exposed and very sparsely vegetated. The sandstone was very mineralised and fractured. The second locality to the west was well vegetated. In the view northward, the low hill in the near middle is the Draaivlei mountain that is Dwyka Tillite and hosts an abundance of H. herbacea as does the Dwyka strata on the northern side of the Hammansberg. The south side is Witteberg Sandstone with Bokkeveld shale on the lower south-facing slope. The view from the third locality southeastwards, shows massive valleys running into the Table Mountain Sandstone of the Riviersonderend mountains on the right. If one takes in mind that this is a short way east of the Wolfkloof Valley where H. herbacea ‘lupula’ occurs, then this area begs exploration. I do have a recall of H. herbacea in low altitude sandstone either near there or at a similar site at MacGregor to the east. The interesting view is the one looking eastwards that really gives the scale of exploration needed. The furthest mountains are the Langeberg Table Mountain Sandstsones and Buitenstekloof is nested in foothills there and in intrusive igneous rock. The coming closer is the Rooiberg, then Gemsbokberg, then Gannaberg, then Ouhangsberg and Hammansberg itself. Rooiberg is fairly well explored although the significance of H. herabcea ‘flaccida’ is not explained. I doubt if a anything new will crop up on the Hammansberg. But the intervening three definitely hold key populations marginal to these covered in this posting. The Langeberg? I only have the set 12 unknowns and cannot see myself resolving those.

Set 9. Map point 17. H. pubescens. Sandberg.
Figs. 9.1 to 9.20 MBB7997 H. pubescens, Dewetsberg.

I long ago observed H. pubescens southwest of Sandberg and south of the Breede River on the Dewetsberg. This is not much further than 500m north of a population of H. maculata at Cilmor Cellar. Visiting the site again I observe that the plants are slightly different to the Sandberg plants. While they are equally small, cryptic and dark coloured, they have less surface spinuliferation that gives rise to the species name. Also some plants are distinctly spotted in the way that H. maculata are and these spots may give way to translucens when the plants grow in less light. See the images showing leaf surface detail of H. pubescens compared to the two collections of the Brickfields plants (images 21-27) of H. herbacea in Set 10, map point 18.

♦

A myth corrected – part 5

Posted on May 22, 2012 by Bruce Bayer

Set 10. Map point 18. H. herbacea ‘submaculata’. Brickfield, Brandvlei Dam.
Figs. 10.1 to 10.20 MBB7995 H. herbacea, S Brandvlei Brickfield.
Figs. 10.21 to 10.53 MBB7996 H. herbacea, E Brandvlei Brickfield.

I have associated this locality with von Poellnitz H. submaculata and treated it as a synonym of H. herbacea. However, here I first illustrate a population about 600m south of that where the plants are in the usual size range for the species ie.30-40mm diam. At the locality east of the Brickfield and next to the Breede River, the plants are 1/3 to 1/2 as large again and can form huge clumps. North-west from this is a population of H. maculata at the extreme end of Die Nekkies and only about 300m distant, and this population I have always regarded as somewhat intermediate. What is interesting to note is the huge variation in leaf shape and armature within each population. In both cases the plants are in Witteberg Sandstones and at the first site this is both in a very shale-like stratum as well as in a highly quartzitic one. This is unusual for H. herbacea. Both populations wedge in geographically between H. maculata populations and in no known case do both occur.

Set 11. Map point 19. H. mirabilis. Droogerivierberg.
Figs. 11.1 to 11.11 MBB7984 H. mriabilis, Droogerivierberg.
Figs. 11.12 to 11.24 MBB7985 H. mirabilis, Sandberg S.
Figs. 11.25 to 11.31 MBB7988 H. mirabilis, Trappieskraalkloof.

In this area H. herbacea is common in the Dwyka Tillite but not in the Witteberg Sandstone. But instead of H. maculata, H. mirabilis is present there and commonly so. Where at one time I considered that H. pubescens was very probably an extension of H. mirabilis to a western and northern limit, this does not seem to be the case as it is so closely linked to H. maculata and H. herbacea. On the other hand I have wondered about the similarity of the Moddergat H. maculata to H. mirabilis. H. mirabilis is in the eastern areas of the Ouhangsberg and now we know that H. maculata occurs not far away on the western slopes. The centre area remains unexplored and this needs to be done as it may also help understand the idea of two principle role players viz. H. retusa and H. mirabilis as precursors of a very complex Southern Cape assemblage.

Set 12. Map points 20. Unknowns.
Figs. 12.1 to 12.7 MBB7865 H. cf. arachnoidea. Keurkloof, SE Dedoorns.
Figs. 12.9 to 12.15 EA1441 unknown, Hex Pass.

I have only two sets of pictures for this set. These are…(a) MBB7865 that Ernst van Jaarsveld drew to my attention at Keurkloof, south east of De Doorns in the Hex River Valley. There is a continuum of suitable habitat between there and Sandhills/Kanetvlei; (b) EA1441 and this number may be incorrect, but it is at the near-base of the Hex River Pass still further east than Keurkloof. In between is Osplaas Station where there are dramatic forms of H. arachnoidea with very spotted leaves, not to forget the nortieri-like form of H. arachnoidea at Kanetvlei about 200m north of MBB7994 H. maculata.

H. cf. arachnoidea. Keurkloof, SE Dedooorns 008

Conclusion
The nature of the populations of H. maculata on Die Nekkies varies quite considerably in that there is an area east of the Resort where the plants form huge clumps. Individual rosettes can be quite large. But there are also points where the plants tend to be solitary, hidden in rock cracks or even truncated into the soil. The plants in the Southern populations are smaller and tend to be solitary. There may be differences in flowers and flowering time but this will really be significant in relation to H. mirabilis rather than between the populations recorded here or to H. herbacea. Attention must be paid to the way in which H. herbacea is geographically wedged in between populations of H. maculata. Equally significant is the character of H. pubescens southwest of Sandberg (Dewetsberg) where the plants have less spinuliferous leaf surfaces and also spotting (maculation). This is one of the best pieces of evidence I have for the geographic continuities that exist throughout the genus both in respect of species of how I think species can be recognized and of how difficult decision making is. In essence this report only dismisses the idea that the Lemoenpoort population can be assigned to H. pubescens. I doubt if it is rational to even formally recognize it as different to H. maculata as “variety” itself is a mythical status. Fragmentation by names is a rather archaic approach to classification that serves hardly anything but a semi-commercial need.

Acknowledgement
Many people contributed in one way or another to this report … Mr Hentie deWet of Moddergat, Messrs Poffie and Hettie Conradie of Sandberg (South), Mr Pieter Naude of Vreesnicht, Mr Johan and Marie Fourie of Buitenstekloof, Mr Nico Marais of Worcester (Brandvlei) Brickfield, Mr A. Groenewald of Cilmor Cellar and Stefan Hugo of River Farm. Messrs PD and Anso leRoux provided much logistic support, interest and company as did Kobus Venter, Etwin Aslander and Werner Voigt. Lawrence Loucka has managed and facilitated the archiving of publications and pictures.

♦

My view of names

Posted on May 23, 2012 by Bruce Bayer

There seems to be so much harping about my departure from the International Code of Botanical Nomenclature (ICBN) that I obviously need to try and explain myself better. The real issue is that we are dealing with a group of plants that is largely appreciated for its vegetative characters and not for its small and unexceptional flowers. Because the plants are small and so commonly grown by collectors the numbers of plants in cultivation and close observation is large. The plants also do vary in respect of leaf morphology, arrangement, and surfaces to a greater extent than in many other genera. Furthermore, the variation is also exaggerated by growing conditions. The fact that flowers are not used in the classification process beyond the level of sub-genera means that there is an almost total reliance on vegetative characters for classification. The nomenclatural system in botany tends to be a typological one, which means that reliance is placed on descriptions very often derived from single specimens. This is particularly so when the nomenclatural types are simply old illustrations that have been used to arrive at identifications and names by consecutive authors for decades. Thus use of those identifications and names, and their continuing re-interpretation causes a great deal of either grief or great personal satisfaction depending on just who is being affected by the process. The fact that the names should indicate “species” is lost from sight and totally obscured by the additional absence of any good universally accepted explanation for what a species is or might be.

Explanation in and around all this has been a large part of my writing since I started this in 1962. Thus I will not enlarge on the subject but rather try to explain again this way using a species name generated by Col. C.L. Scott with whom I had some considerable altercation. It must be understood that Col. Scott was not a biologist and it is just a simple fact that the problems that the above brief remarks expose, trouble many professional taxonomists to this day. I do not condemn Col. Scott and express my respect and admiration for him and am very grateful to him for the friendship he later extended to me before he died.

The example I will take is that of his species Haworthia geraldii. It comes from a small hillside east of Riversdale running south to north, named Komserante. This is an Afrikaans term meaning the ridge around a small geographic basin. In the way that haworthias are assembled in small habitat defined and localised populations, there are three recognizable populations of plants (of the subgenus Haworthia) along this one ridge about 1km long. The southern population used to be complemented by still another that occurred where the stream bed left the basin at the southern end. A plant from that southern population was named as H. foucheii by K. von Poellnitz. The northern population is a bit problematic and I initially included it in my then concept of H. magnifica. Since then however, it has become clear to me that my idea of species was too conservative and that H. magnifica as described by von Poellnitz from south east of Riversdale (now the Frehse Reserve) is part and parcel of a huge complex that I regard as H. mirabilis. Thus this northern population is seen by me as H. mirabilis but complicated by the fact that it is largely influenced by hybridization with the next population or populations south. A plant (or perhaps, and doubtfully, plants) have been given the name H. vernalis by the Japanese writer M Hayashi. The name H. geraldii is attached to the second population southwards. The plants are very proliferous, form large clumps and the leaves are usually quite strongly retused (“bent back like a thumb”). The name H. foucheii is simply attached to the third because the plants there tend to be solitary and the leaves are also fairly erect and spreading as described and pictured for the original from the fourth population that was off the ridge.

The problem is now that we have descriptions and plants in each population that do not match the descriptions. While I assign the northern population and all the plants in it to H. mirabilis, other writers use the names H. vernalis, H. magnifica and even the confused name H. asperula. What they mean is by no means clear and in actuality a lot more names and descriptions would then be required to name each of the countless variants there for what might be an original pure species and variants or for first second or third generation hybrids with H. geraldii or H. foucheii as those names were applied.

But the problem is far more extensive than just these four populations. After years and years of field exploration it has become evident to me that like H. mirabilis, H. retusa is also highly diverse and I see it to include all the variants of H. turgida. In fact I suggested a long time ago that H. retusa is simply the solitary large form of H. turgida in low-lying level areas as opposed to the more common and extensive clumping cliff dwelling forms of that species. This is also why I object to the requirements of the ICBN that require the name H. retusa to take precedence for the species for historical reasons when it would be far more realistic to take it as a variant of H. turgida for biological reasons.

It has always been simply evident to me that H. geraldii and H. foucheii are variants of H. retusa. The problem now is that the plants in the two populations vary so much that, while the names are indeed useful for commercial and collector use, there can be problems that the use and application of the names will be confused.

Look at it like this. In respect of H. geraldii; it came from one population at Komserante, Riversdale and not all the plants are the same. I think this one population and all those plants in it belong to the species H. retusa, so I called it H. retusa var. geraldii. (var. = variety). Because not all the plants look the same that means that only some are truly geraldii and only the plants that meet Scott’s description are actually var. geraldii. Because the plants multiply vegetatively and Scott only took a piece, the original plant may still be there. This means that we should recognize it as H. retusa var. geraldii forma geraldii and in cultivation as H. retusa ‘Geraldii’ or just as H. ‘Geraldii’. What do we do with the other 100 plants or more in the population that are not H. retusa ‘Geraldii’? I do not think they can all get names so I take away the capital letter, leave out “var.” and use ‘and’ (inverted commas) to show that the identification is, and will, be a bit uncertain unless you know from a label that the plant comes from that particular population at Komserante. The same applies to H. retusa var. foucheii. In this case the original population is gone and I am not sure if H. ‘Foucheii’ is still in cultivation. So I use the name H. retusa ‘foucheii’ for a second population on Komserante. Some of the plants in this population look like some of those in the H. retusa ‘geraldii’ population but not quite like H. ‘Geraldii’ itself. So for me the answer is MBB7780 H. retusa ‘geraldii’, Komserante and MBB7781 H retusa ‘foucheii’, Komserante.

It can be seen that a complication comes in when more than one population is involved. The problem then is that you can put them in a line so that some plants in the first population look like plants in the second, some in the second look like plants in the third, some of those look like those in the fourth until at the end none of the plants in the last population look like any in the first. The trouble in the field is that the line is not straight and it can also go off in different directions. Some of those directions may end up where they started. One simply cannot be truly confident about many of those names that are given to plants. Serious and proper consideration must be given for how different they may be even within the populations. Not to say of the shared similarities and still added variation from geographically adjacent populations. It may happen that I may write H. ‘retusa’ (and add number and place name) for a population that may better (not necessarily correctly) be identified as H. retusaXmirabilis. Other writers who are not biologists and have other considerations in mind may want to give a new name altogether. That new name and description may be again really only for one plant, or maybe a few more in a population, that have real or imagined novelty value and attraction. But they take no account of all of the less attractive variants or other populations that comprise the biological whole by close proximity if for nothing else.

The above discussion is integral to the rationalized list of names that Dr John Manning helped me produce. There are some species names there that I have deliberately presented as synonyms or variants of the species I recognize, because the authors of those names fail to convince me that they have any understanding of the situation at all. The word “species” is apparently simply a convenient naming system for oddities and novelties and not any scientific construct to explain the natural phenomena of living systems and their parts.

To summarize; I have written a report on flower characters in which I refer to the Komserante populations and I have used the following names and my own convention as follows:-

Please note that the way I have omitted the word variety from the following names and also used inverted commas …

7779 H. mirabilis, Komserante (the northern population – I could add ‘vernalis’ but it carries all the baggage of doubt about actual status).
7780 H. retusa ‘geraldii’, Komserante.
7781 H. retusa ‘foucheii’, Komserante.
7920 H. retusa ‘nigra’, Van Reenens Crest.

The omission of the word variety is for two reasons…

1. Economy

2. To convey the idea that the actual indication of status is not certain as I have used the name to indicate a population or populations rather than a single described plant. The prime and overriding uncertainty is that we cannot know what a species is that I have described for Haworthia. Thus how can we possibly rank populations at levels below?

The use of inverted commas reinforces what I want to convey. This is that the individual plants in the populations are variable and it may not be easy to always identify the plants (individual or population) according to a more formal classification.

Any departure from the ICBN or the way the names are treated in formal botany conveys the difficulty that I personally find in trying to reconcile formal nomenclature with names that are so often tied to single plants.

♦

Conclusions and Acknowledgements

Posted on May 23, 2012 by Bruce Bayer

Conclusions.
The nature of the populations of H. maculata on Die Nekkies varies quite considerably in that there is an area east of the Resort where the plants form huge clumps. Individual rosettes can be quite large. But there are also points where the plants tend to be solitary, hidden in rock cracks or even truncated into the soil. The plants in the southern populations are smaller and tend to be solitary. There may be differences in flowers and flowering time but this will really be significant in relation to H. mirabilis rather than between the populations recorded here or to H. herbacea. Attention must be paid to the way in which H. herbacea is geographically wedged in between populations of H. maculata. Equally significant is the character of H. pubescens southwest of Sandberg (Dewtsberg) where the plants have less spinuliferous leaf surfaces and also spotting (maculation). This is one of the best pieces of evidence I have for the geographic continuities that exist throughout the genus both in respect of species of how I think species can be recognized and of how difficult decision making is. In essence this report only dismisses the idea that the Lemoenpoort population can be assigned to H. pubescens. I doubt if it is rational to even formally recognize it as different from H. maculata as “variety” itself is a mythical status. Fragmentation by names is a rather archaic approach to classification that serves hardly anything but a semi-commercial need.

Acknowledgement.
Many people contributed in one way or another to this report … Mr. Hentie deWet of Moddergat, Messrs. Poffie and Hettie Conradie of Sandberg (South), Mr. Pieter Naude of Vreesnicht, Mr. Johan and Marie Fourie of Buitenstekloof, Mr. Nico Marais of Worcester (Brandvlei) Brickfield, Mr. A. Groenewald of Cilmor Cellar and Stefan Hugo of River Farm. Messrs. PD and Anso leRoux provided much logistic support, interest and company as did Kobus Venter, Etwin Aslander and Werner Voigt. Lawrence Loucka has managed and facilitated the archiving of publications and pictures.

Haworthia Update Vol. 9 – Addendum

Posted on May 23, 2012 by Bruce Bayer

The data keep coming …

MBB1128 H. maculata var. livida, Lemoenpoort.

MBB1128 (MBB7066 W. Lemoenpoort) is the type locality for H. maculata var. livida. Plant pics are available in Haworthia Update Vol. 9. A myth corrected to, population 5, set 3. Here are flower photos from 8 December 2012.

MBB4461 (and (MBB6514) H. mirabilis ‘notabilis’ Buitenstekloof

See Haworthia Updates Vol. 9. part 3 – Population 8, Set 6

MBB7266 H. maculata var. livida, E Lemoenpoort.

See Haworthia Updates Vol. 9, part 2 – Population 10, Set 4

MBB7994 Haworthia maculata, Kanetvlei.

In Haworthia Update Vol. 9 and a recent posting on Haworthia nortieri Bruce Bayer makes reference to this Haworthia maculata population, MBB7994 from Kanetvlei.

A myth corrected to – Haworthia maculata var livida (Bayer) Bayer – and flowers ignored.

MBB6694 Kanetvlei, Hex River Valley as a variant of Haworthia nortieri.

MBB7991 Moddergat above Ouhoekberg E, ex hort. single clone

From Updates Volume 9 Set 5 Bayer writes, “These populations were first observed as one on a higher eastern point of the Ouhoekberg above Moddergat in about 1975. George Lombard accompanied Kobus Venter and me there in 1996 and we found them on the western high point.”

These photos were taken 20 December 2012 from a single ex hort. clone and hence consistency, but still illustrative.

8034 Haworthia maculata ‘livida’, Ouhangsberg.

These plants occur in Pteronia communities on very dry and sparsely vegetated north slopes. The geology is Witteberg Sandstone but the particular strata the plants prefer seems to be a softer more erodible and less quartzitic stratum than they occupy at Lemoenpoort to the west. The plants appear to have flowered early in December coinciding with the flowering at Lemoenpoort and southeast Lemoenpoort. The two localities where we observed the plants are approximately midway between H. mirabils at Droogerivierberg to the east and Lemoenpoort to the west. There is no suggestion that they form any continuum and the flowering times are different.

The name of the farm is Iminga Mountain Reserve, a large tract of quite non-arable countryside that is appreciated by the owner as her garden.

Acknowledgement – I deeply appreciate the freedom to explore Ouhangsberg granted by Ms Suzi Broomberg.

MBB8042 Haworthia maculata ‘livida’ – Iminga Mountain Reserve, Ouhangsberg.

Following on from the 8034 record, we returned to explore still further east in a direct line from the 8034 localities to the H. mirabilis population on Droogerivierberg (see map). We found plants at the same altitude as we had observed H. herbacea on the previous visit. The plants were again very cryptic and hard to find. Flowering was also long past and the few capsules present contained very little seed. The situation with respect to habitat was a little different and these plants were just off a truly sandstone stratum and in a narrow band of renosterveld. H. herbacea was within 50m of the nearest H. maculata in a more karoid vegetation similar to that where we had seen H. maculata on the previous visit. This is difficult to understand. Southeast of the Brandvlei Dam, H. herbacea occupies habitat where one would expect to find H. maculata. But it does not do so in the Ouhoekberg nor Hammansberg. Here north of the Ouhangsberg there does not seem to be a visible difference in the choice of habitats for the two species but they cannot be said to truly co-occur i.e. grow together in such close proximity that they intermingle.

The spatial relationships of the various populations are now quite significant. The spread from localities east to west on the map is 5.23km. The distance between known H. mirabilis and nearest known H. maculata is nearly 1.5km. There is no indication of transition and present evidence is that there is no continuity between these two species. There is however still that space between to explore as well as a massive valley south between the two populations in question.

Fig. 1 Image of Iminga Mt Reserve. — Fig. 1 Image of Iminga Mountain Reserve.

♦

Very brief note re Haworthia nortieri flowers

Posted on May 24, 2012 by Bruce Bayer

Some criticism about my supposedly having ignored the flowers in Haworthia comes at a very inopportune time. I set aside flowers for the reasons very obvious from the historical record but also because of the considerable problems of similarity in the appearance of the flowers in apparently quite different species. My priority was a geographic overview and a rational basis on which discussion and decision making could be based. It I just grossly unfortunate that other writers and critics seem to be wedded to a classification paradigm locked into the approach that prevailed 70 years and more ago. This in the total absence of a species definition other than the vague acceptance of a zoological one based on interbreeding capabilities. This ignoring the ease of hybridization among Haworthia variants in general.

While I have written an account of flower appearances in a small selection of populations, I also came across these few images I have of flowers in what I regard as the species H. nortieri. I have also added images of a single flower of H. maculata from a population high in the mountains at Worcester that could be seen as a southern extension of the H. nortieri set of populations. Note must be taken of my early contention that H. nortieri and H. globosiflora were the same species, based on my observation of the intermediate appearance of the flowers of a Vanrhyns Pass population. The H. maculata bud is typical of the species in the Southern Cape, whereas H. nortieri has rounded bud-tips.

The flower of the Trawal plant are dramatically different from that of, say, Sneeuberg. It is very understandable that differences like this lie at the base of all the argumentation and confusion that so despoils the naming and identification of Haworthia. A classification has been needed against which to explore and examine these differences. It seems to me totally unnecessary to try and construct another hierarchy of solely Latin names while so little is still unknown. ♦

♦

Haworthia marxii and H. truteriorum in relation to rational classification.

Posted on May 25, 2012 by Bruce Bayer

Explanatory note: In a rationalized list of names I wrote and had published in Haworthia Update Vol 7. I made two decisions. One of these was in respect of H. marxii and the other in respect of H. truteriorum. In the case of H. marxi I included it under H. emelyae. In the case of H. truteriorum, I placed this under H. bayeri. In neither case did I have good evidence for doing so, much other than my conviction that a classification is intended to reflect origins and relationships. In formalizing names one perforce is pressured into making decisions that you are not informed enough to make. My main defense is that the authors of the two species were not adequately informed either. I think their account of H. mutica (their H. groenewaldii) at Buffeljags demonstrates this. H. marxii presents particular problems that I simply do not have any substantial data that I can process. In the case of H. truteriorum I did have some to which I can now add. This suggests as Breuer and Marx indirectly indicate, that H. truteriorum relates to my concept of H. mirabilis. I concede that I may be quite wrong in attributing it to H. bayeri. I find it very difficult to see the decision to describe it as a distinct species as a logical scientific action. This article does not extol any imagined virtues or skills of mine. It is only intended to further project my opinion that we urgently need to work towards a classification that does satisfy scientific principle and not novelty or commercial ends. It is also a counter to some very negative opinion aimed in my direction.

At a level above classification lies respect for people and their feelings. Therefore this writing should not be seen as anything but a commentary on the classification process and not how this can also denigrate people as much as honor them. H. marxii is described by Sean Geldenhuys in ALOE 44.1:5, 2007 from apparently 2 populations in a confined area in the north and east of the Little Karoo. Placing it with H. emelyae simply reflects doubts that need to be expressed about how this oddity has come about, and respect lost for reasons not necessary to explain. H. truteriorum is described by Ingo Breuer and Gerhard Marx in ALOE 48.3:54, 2012 and refers to a single population of plants southeast of Oudtshoorn. That this latter population is singular and extremely interesting goes without question except for how it is explained. Despite much correspondence with both authors and after reading their published works, I am not aware that either has a concept of what a species is any different than what may have been held by either K. Von Poellnitz or G.G. Smith. Ingo Breuer particularly has published what he refers to as a species concept of Haworthia. This is nothing more than a long list of Latin binomials and we have to assume that this is then also a list of real species whatever they might be. Gerhard Marx maintains that my attempt at a species definition is so broad as to be meaningless. He has not supplied an alternative definition and I am left with the impression that he makes the same assumptions that “character” differences equate species as does the ordinary uninformed mortal.

H. truteriorum is described in a popular journal (Aloe) that has been approached by at least one botanist not to publish new species as there is no official review process. However, the response was that the journal does have its own in-house authorities that cover the possibilities of scientific lapse. I note that the article is specifically foot-noted to indicate an editorial review and I perforce do not see this as a check on the scientific content. There are some lapses that I will deal with but recognizing that these might not be the same ones that a properly qualified botanist (a minimum of a recognized 4-year degree course), nor those that an experienced and knowledgeable botanical taxonomist may have corrected. More important though is the population itself and where it fits into the overall Haworthia picture.

The most substantial gain that we can make is to arrive at a species definition. It is evident to me that species are complex systems in which there are variations that have arisen from to earth differences and must continue to exist to facilitate response. If there is evolution and thus adaptation and selection, then there has to be something to work with. The phylogenetic idiom was “specialization is precursor to extinction”. Geological and habitat diversity result in plant diversity. This diversity is necessary for survival as the habitat changes. Therefore a species will have a geographic distribution across which individuals and populations will vary and be different from one another. Geology and geomorphology will be strong factors influencing plants like Haworthia that are associated with skeletal soils and rocky habitats. This means that we have to look for associations in respect of distributions and the driving forces that affect even vegetation. Thus in the case of H. marxii (known from 2 populations in a small area south of Laingsburg) the vegetative appearance of the plants cannot be seen in any other species geographically closer than H. emelyae, which is quite a long distance away southwards. The presence of H. pumila in the same area as H. marxii suggests that the Haworthia presence there could extend from the Worcester/Robertson Karoo. Hence H. mirabilis, also present in the Montagu area, cannot be ruled out in seeking a relation to H. marxii. There is much more to this issue of H. mirabilis in respect of its variability and its distribution into the Little Karoo that impacts directly on this suggestion. It also impacts on the real identity of H. truteriorum.

Where I see a real problem with H. marxii is in floral morphology. I gather that the flower very much resembles that of H. marumiana dimorpha. In the same way that the flower of H. pulchella globifera is identical to that of H. cymbiformis incurvula at Plutos Vale, there is a massive problem in drawing conclusions from flowers as a character for the level at which all role players are trying to identify species. But so-much for H. marxii and I do not seriously question it.

The case of H. truteriorum is more manageable. I have been speculating for a long time that H. mirabilis and H. emelaye may in fact be the same species. This may be the proper level at which we should be recognizing species as systems. I extend the argument even to say that it is not inconceivable that H retusa and H. mirabilis are one species. This means that I have to bury my long-held objection to the view that “Haworthia is a genus in a state of active evolution”. My objection being that this is an obvious aspect and that in any case all species are faced with the inevitable need for change and adaptation. But this does not weaken my view that species are chaotic fractal systems that vary around a point of attraction according to the stability of their genetic bases and mutating rate. How strong I am on the technicalities of DNA and evolutionary theory may be problematic, but there is no evidence that other authors even contemplate the issues.

Gerhard Marx is a remarkable observer and I have huge respect for his many skills. Breuer is a really competent compiler too. I do not question what they say about the characters of the plants and their observations. What I question is their knowledge and insight into broader botany and the distribution and variation in Haworthia however much more they know than the above average collector.

My prime objection is that the description involves a single localized population. This in itself creates huge doubts in my mind because on this basis, Breuer’s several hundred species is conservative. I will only dwell on three other points. The one is their very trite “Never before have retusoid type Haworthias been found growing in shale in the Little Karoo”. The second point is the habitat description in relation to geology. The third is the flower and flowering time. The fourth is about the illustrations and the art work.

Haworthia mirabilis was recorded in shale at Barrydale by Smith prior to 1947. It was recorded at two places in shale or shale derived soils prior to 1999 and I can add that I have seen it at two new locations in shale in the Montagu area since. The second point is the description where it is referred to H. bayeri and H. emelyae occurring in quartzite and quartzite conglomerates. As in the description of H. groenewaldii, this is simply a very crude and inaccurate account of a very important issue. I am no geologist but I do know what quartz is and that it occurs in both shale and sandstone formations. Furthermore, I do know that the Oudtshoorn area has an incredible geology with ancient and recent geological formations adjoining as a consequence of faulting and folding. Quartz is Silicon Oxide and is apparently soluble in water at high temperature and pressure formation so that it can accumulate in fissures and bands in parent rock. In both sandstone and shale the quartz varies in purity, and the crystals in size. South of the Langeberg the shales are covered in an extensive layer of tertiary gravels that are far less extensive north of the mountains. But the main point is that the vast quartz patches of the Little Karoo are actually associated with quartz existent in the Bokkeveld shale. The “species” need to be properly looked at in their relation to that complex geology that exists there. Marx and Breuer mention the differences north and south of the Outeniqua Mountains. But the Outeniqua Mountains are just an eastern extension of the Langeberg from the Gouritz River gorge. The geology north of that area is quite different from that west of the Gouritz. This suggests an ignorance of geography added to that displayed for the geology.

Checking my own knowledge and experience of the habitats of the species involved (viz. bayeri and emelyae – with the name picta an anomalous insertion), it is very clear that the statement “quartzite and quartzite conglomerates” is erroneous. There are four geological formations involved and these are the Table Mountain Sandstones, Bokkeveld Shales, and then Enon and Tertiary deposits. In the Heimersriver where the H. bayeri, H. emelyae and “H. truteriorum” are reported, lacks Enon presences. The report of the plants in unfragmented, unweathered upright shale (Bokkeveld) needs to note that the quartz patches in the area are the result of fragmented and weathered shale.

Marx and Breuer perhaps should also take account of H. outeniquensis not a great deal further south in an area that I believe is still unexplored. There is also the mystery of a plant found by Avril Schein in that close area that remains unexplained.

We know from the H. retusa – H. mirabilis interaction that flowering time is no barrier to hybridization and that flowering time may not be indicative of a completed speciation process. Ignoring the fact that we can only guess, on the basis of the scientific paradigm, that we are interpreting and trying to understand an evolutionary process. Breuer has attempted to jump this issue by the recognition of “aggregates” and the two authors use the fob of “the mirabilis/maraisii/magnifica complex. This complex I presume is explained as a list of names only. This I do think emanates from a mis- understanding of a real knowledge of field botany generally and of Haworthia in particular. I say this with great emphasis and conviction because it is something I am still working towards. I have just completed a very thorough look at a the flowers of a very small fragment of populations driven by the expressed opinions of both these authors in diverse places, that flowers are significant with the import that I have ignored them. The fact is that if the flowers are considered then we have a bigger problem than before – not a solution. The flowering time of H. truteriorum that Breuer and Marx cite is very possibly an indicator of behavior rather than a species differentiator. Why they emphasize it is most probably, as J. Manning pointed out, most observers (taxonomists) have a subconscious belief that species are things that do not interbreed and so flowering time is seen as such a great barrier to interbreeding that it MUST be a major species indicator. My observation is that this is not true and that species are inherently highly variable systems with great capacity to respond to environmental drama that may threaten their continued existence. The nature of the problem is very well illustrated in the case of H. retusa and H.mirabilis where I contend that a fully developed view of the genus may require that they be seen as one species. Thus the importance of flower and flowering time is part of the myth of a non-existent species definition and is confounded by the concept Breuer and Marx have of the nature of Haworthia species.

There is a very good description of the flower and its character, but the illustration and the art work is weak. The photograph of the flower is of a single dissected flower to show an internal structure that could be of any species in the subgenus and is hardly helpful. That flower does not look to me like the perhaps longer narrower flower of H. mirabilis (real) which is largely recognizable on the arrangement of the petal tips in the bud-stage. This ‘mirabilis’ character should be apparent in the way the petal tips display and we have only a painting to judge this by. Despite Marx’s craftsmanship I am not sure if his flower picture is a true image. I do have an observation on with his outstanding art. Many years ago, photography was a bit of a handicap when it came to illustration of floral and other plant detail. Historically artists were used to capture detail that could not be described or otherwise illustrated. In the present age this is a bit of a myth and it lives on simply because of the skill that is involved in the production of a good piece of botanical art. Marx’s art is up there with the very best. But good art does not equate to good science? Or does it? Botanical art is perhaps not the same as pure art and the intent of either may be quite different. With pure art one could surely be trying to achieve the same goal as science and there must be some trick to understanding the similarity.

My conclusion is to plead for more rational classification and better attention to those things that matter. The fragmentation of a genus by Latin binomials just because of collector and novelty interests, aggravated by commercial implications however slight, is a disservice to all of us. ♦

The reality of Drosanthemum micans L.

Posted on May 26, 2012 by Bruce Bayer

Donald A. Levin (2000) quotes Raven, Berlin, and Breedlove (1971), who wrote…“our system of names appears to achieve a reality which it does not in fact possess”. I find this a curious quote, as Levin was discussing species concepts and we could ask if species themselves have any reality. There is a lack of a universal definition for the word “species” and I find the recurring reference to “concept” as related to the word, very confusing. Why should the species be a concept, subject to individual interpretation? This is of course if they have no reality and we each create our own. How useful is this for science? Donald goes on to generate his own “species concept” in which he proposes…“that each species has a unique way of living in and relating to the environment and has a unique genetic system…”, and he refers to this as the eco-genetic species concept. But still we do not know what he means by “species”. He says that the ecological properties of such species are not uniform within the species and thus not equivalent to the taxonomic properties of species, “which are chosen because they are conservative and stable attributes”. It would be interesting to know just what he means by “taxonomic properties” and I presume he means primarily morphological characters. Certainly there must be taxonomists who utilized or have utilized ecological facets to decide on their species. It must also be recognized that the taxonomist is confined to the material at his disposal for examination and decision making.

My view, to the degree that I can understand Levin’s arguments, is that he has not truly stated the case. The problem concerning the reality of Latin names, is that they have largely been generated by taxonomists who may not have recognized that morphological (“conservative and stable attributes”) properties of a species may vary just as substantially as those of eco-genetic species; and also that these “realms” for each species may be different. From my long experience in taxonomy and the usage of names in communication, it is evident that it is the taxonomic system and taxonomists, which have induced the majority of people who use Latin names in any way, to believe that they do have a reality. The system has been based on the view that Levin’s apparently has, that a taxonomist determines species by characters or character sets which can be quantified and easily (sic!) used. Perhaps also, that the taxonomist has had enough material to make a decision that is universally true. In the absence of a universal definition it is self-evident that taxonomists and persons, who use the names they give and are given, may have quite divergent views on what those names actually mean and what reality they have. This is especially true if sight is lost of the fact that the characters taxonomist use and have used either morphological or ecological, are simply not as “conservative or stable” as believed. Neither may they be adequately circumscribable or quantifiable. The material examined may simply be inadequate to convey the varied characteristics of the species as it occurs throughout its distribution range.

I would like to use an example to illustrate the relationship of names to taxonomic characters and identification. Drosanthemum micans is an old Latin name that was given by Linnaeus for a mesembryanthemum species. The name is apparently based on a Dillenius illustration (fig.1). While the name has been commonly used in the ranks of botanists and horticulturists of my personal acquaintance, there is some doubt that the connection of original illustration and name, to the plants identified as such in recent times may be right. In this case, my use of the name stems from the usage of my predecessor at the Karoo Botanic garden, F.J.Stayner, and he probably obtained the name from identification by Mrs. L. Bolus. I many cases names may come into general use through less authoritative channels. At this point in time the true typification of this name and its correct application is under review, so I will use the name as I know it for a species that to my knowledge occurs both within the Karoo Botanic Garden Reserve, a short distance to the east and also in the developed suburb to the northwest of Worcester, Brandwacht. The name “micans” means “glittering”, and indeed this is the case for this plant (fig.2). The species is characterized by inner and outer rings of bright colour, the calyx bears enlarged bladder cells and the leaves have a grey-blue colour in the summer months. The flowers have outer black staminodes and there is a series of inner smaller petals, often uniformly one-colour, which accentuate the coloured ring effect. There is another species with these black stamindodes viz, D. speciosum, which can be separated from D. micans primarily by the appearance and colour of the leaves. In the former the leaves tend to be slightly more globose and have a greenish-yellow colour. In the latter the leaves are grey-green and tend to be elongated with an uncinate (hooked) end. The black staminodes seem to be unique in the mesembryanthema and who knows if they serve any particular function.

In more recent times, Mrs. Bolus described several other species. One is Drosanthemum bellum that is said to have been collected “near Ceres”, and D. hallii from hills east of Rawsonville. D. bellum was described as pink with black staminodes, and D. hallii as yellow with black staminodes. During a period I which I was interested in these very colourful plants, I came across a population of plants at one spot on the hills “east of Rawsonville” (Die Nekkies, north of the Brandvlei Dam) that demonstrated an array of colour forms. Among these were pink (bellum – fig.3), white, and purple. At the same time I found a population of plants nearby in which all the flowers were bright yellow (hallii – fig.4). More recently I have had occasion to examine this more closely and find many more populations, which indicate the problem that taxonomists have apparently yet to come to grips with if botanists and scientists in other disciplines are to find any reality in among Latin binomials. This is that these characters are not diagnostic for the “species”

On revisiting the “bellum” population I found the same array of colours. There were plants with very pale yellow flowers, pink, pale rose-red, white and white shade with purple (figs.5, 6, 7 & 8). There were no bright yellow flowers despite the fact that such bright yellow flowers characterized plants from every other population along the length of the Nekkies. (Here is where Levins could apply his ecogenetic concept!). What was most dramatic was finding some plants with flowers of the micans type (fig.9 in one population of hallii. I observed colour variants in other populations of hallii further to the south-east where the plants were primarily with bright yellow flowers. On revisiting the D. micans population just east of the Karoo Garden, I found that there were plants with the plain bright yellow flowers of hallii as well as pale rose-red to red flowered plants (figs.10, 11, 12).

The question now arises…”What else?” My own approach to taxonomy and plant identification, is that one has to consider not just the ecological associations of plants, which are mostly more difficult to describe accurately than any morphological property, but geographical distribution. Plant species are just like any other material phenomena. They are distributed in space, and they change with time. This is the determinant of their reality. Taxonomy and the decisions that are made are dependent on the material available to taxonomists and it is often simply not substantial enough to establish a practical reality in the application of their names. Looking further afield, I have three more populations of plants from the southern Worcester area namely at Jonaskop (figs13, 14), Lemoenpoort (figs 15,16) and Droogerivierberg (figs.17, 18,19, 20, 21) which fit the micans/hallii mould in terms of all but colour. The population of D. micans on the Droogerivierberg south of Worcester I thought to be fairly consistent in flower colour. It proved not to be and we found a wide range of colours, which extended to white and also included the more typical bi-colour thought to be the real thing. Curiously Mrs. Bolus also described D. leptum from Stormsvlei Pass as white with black staminodes. I have not been able to find it again, but it does seem, in the light of the distribution and variation of D. micans as I have discussed it here, that it is in fact also this species.

Outside of the Worcester area, there is a population north of the Langeberg mountains in the Keerom Dam area, which is characteristically micans (fig.22 – not that single images can represent the variation in the population). Still further afield is a population north-east of Montagu which suggests affinity with the populations of southern Worcester (fig.23).

Then there is Drosanthemum aureorubrum described from near Drew west of Swellendam. It has the characters of micans except in respect of colour. It is inclined to have flowers rather bronze-red-bronze in colour (fig.24) apparently without much local variation, and it should be noted that this variability can only be assessed in the field or by observation of plants mass propagated and cultivated. I can relate that population to three others, one from the middle Breede River Valley at Napky (fig.25), another from northwest of the Potberg at Potteberg Farm (fig.26) and the third from near the mouth of the Breede River at Infanta (fig.27). All well and good, except that the colour does not agree either with D. micans or D. hallii. Nearly everything else does.

The nail in the coffin of a narrow “character” driven taxonomy is a population northeast of the Bromberg Mountain at Stormsvlei in the Swellendam district. I had found these plants when without flower. Visiting the site in September/October when in flower was quite a revelation. There was a range of colours which included the bronze-red of Drew and the southern Worcester and Breede River populations (figs 28, 29). Very significant were a few plants with flowers coloured precisely the same as micans (fig.30). There were also plants with pure yellow flowers, but the yellow was distinctly on the golden-yellow side. I regret leaving out another dramatically varying population eat of Barrydale.

There is yet another “species” viz. D. lavisii, named by Mrs L. Bolus after Bishop Lavis who collected the original specimens ostensibly from between Struisbaai and Bredasdorp. Although Acocks also deposited a collection, purported to be this species, from Northumberland Point, I have looked for it there in vain. I have found three populations of what must be this species with its red flowers, from Napier (fig.31) from Swartjeskop (fig.32) northwest of Bredasdorp and from Soutkloof (fig.33) still further northwest. At first sight this did seem to be distinctive, but in summation I would opt for the geogenetic option. Looking at all the collections and attempting to relate them to the distribution of other plant species (AND their variants), I suggest that D. lavisii is another name for D. micans. I have seen a red-flowered Drosanthemum in the Bontebok Park at Swellendam (fig.34) and Van Jaarsveld and Pienaar report D. lavisii from the Goukou River in the Riversdale district (fig. 35), and these may be related to what Mrs. Bolus described as D. edwardsii from near George (figs. 36, 37, 38) If one considers the geographic aspect and notes the relationship of the distribution of D. micans and D. speciosum then one must perforce suspect that D. edwardsii is the eastern extension of D. micans. D. micans and D. speciosum do grow in very close association, although very seldom actually sharing habitat as they do on Jonaskop. My observation would be that the distribution of D. micans extends into the Swellendam and southern areas, whereas the distribution of D. speciosum is more karoid and it can be found eastward to Oudsthoorn. An apparently vicariant population near Uniondale has flowers which are golden yellow and with long pedicels (fig.39).

Thus we come to the point where we can ask what these names now mean. An aspect of exploration has been interaction with landowners and other members of the public. In this interaction use is made of names. Botanists (non-taxonomists) who have had reason to explore and report the plant species of the Overberg, have used the names lavisii and speciosum. They seemed not to have the capacity to assess if these identifications were right or wrong, and this is not because they were incompetent. It is essentially because from necessity and habituation they apply a species concept which has no reality. When one tries to convey to a lay-person what one is looking for, names clearly have either no meaning at all, or else are linked tightly to an image and association that the person may have derived from book or contact with someone else. It was most notable that in the Worcester/Robertson area, any red flowered mesem was linked to what I know as Drosanthemum speciosum and people found it difficult to conceive that there was a second species also with red-flowers. It proved well-nigh impossible to convey and communicate that either species could have different coloured flowers or that such variants were not different species. Fig. 40 demonstrate some of the variants in D. speciosum, also at Doogerivierberg, but while D. micans occurred in closer association with Witteberg Sandstanes, D. speciosum was on Dwyka Tillite.

A closing point is to refer again to Levin, where he is copied by Charles Craib in the magnificent work on The Grass Aloes of the South African Veld. Levin writes…”the eco-genetic species concept has utilitarian value, which is important from an operational concept.” The implication is that taxonomists have propounded a system of names which has little operational value. What then is the sense of taxonomy? Certainly this is what Craib conveys where he propounds a choice of a classification that serves the purpose of the user because the officially accepted system does not (meet the need of an informed user). The fact is that no system will have operational value while there is no universal definition in place, and common realization that a binomial system and names will never have any worthwhile reality unless there is a commonality about what names mean. Taxonomists have to start, with self-examination, to teach the user that names are for communication and have a meaning outside of the very narrow confines of obvious morphological difference, guess work based on limited material and knowledge, or idiosyncratic opinion. While it may be argued that the confused names I have quoted are directly related to the absence of a taxonomic revision of Drosanthemum since Mrs. Bolus last described her species, I use this example to reflect my long general experience with plant names and what people make of them.

Acknowledgment
I must acknowledge the association with Dr. H.E.K. Hartmann with whom I have been privileged to communicate over many years, and whose contribution to the knowledge and classification of the Mesembryanthemaceae parallels and possibly exceeds that of Mrs. L. Bolus. Also I would really like to express my appreciation to the many landowners who have taken us on trust and so kindly allowed us access to their property and plants. To list them all would occupy more space than this whole article but I must particularly acknowledge Messrs. Poffie and Hettie Conradie of Worcester (Droogerivierberg), Messrs. Jon and Cindy Webber of Cape Town (Klippoort, Stormsvlei) and Neil and Saartjie Neethling of Swellendam (Potteberg Farm). Jan and AnneLise Vlok very kindly sent me pictures from Riversdale and Mossel Bay. I do not know how to acknowledge Conservation authorities in terms of their permit system and how these are issued and monitored, nor the paranoid response one gets to the term “collecting”. There is in my opinion a total failure to make any distinction between collecting for knowledge gain and satisfaction of natural curiosity, intelligent plucking for financial gain, or crass exploitation that could lead to degradation of the environment, nor to balance this against doubtful control of development and concomitant destruction of natural vegetation by other agencies. A permit applicant seems to be regarded as a confessed reprobate with no sense or sagacity and an immediate threat to biological diversity. At least this is my impression of how I have been seen. Who wants to draw attention to themselves in this way? Steve Hammer kindly commented on the manuscript.

Reference Levin, Donald, A. 2000. The Origin, Expansion, and Demise of Plant Species. Oxford University Press.. ♦

Fig. 01. The possible type of Drosanthemum micans.

New finds in Haworthia

Posted on June 16, 2012 by Bruce Bayer

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

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.

Fig. 6 MBB6666 H. retusa ‘nigra’, Tradouw Pass

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). ♦

What do collectors need?

Posted on July 1, 2012 by Bruce Bayer

Previously published in Alsterworthia International Volume 12, Issue 2. July 2012.

I ask this question because too often the views of the collector are espoused as an excuse or defense for some or other argument about classification. It has often been said to me that collectors are not interested in taxonomy and they are at the most, happy just to have a name. This argument does not impress me because as a society we have a trust and a belief in science and whatever is written, outside of fiction, should seriously address the truth. It should not matter what the reader may want to make of the product other than that the reader may just by chance really want to know and understand something. On reflection, one writes for the reader who must surely be reading because they want to know something, and names are the key to the “something”?

This is why I have responded to reviews of my writing that have been published at various times. I have written as a communication and am glad to know what the reception or rejection has been. Recently Steven Hammer wrote what is listed under the title of “Book Review” comment on a recent book by Ingo Breuer and of Update Vol. 6 by me. It is a wonderful piece of prose and worth every bit of reading and appreciation, but it does not pass as a Review. Or does it? I feel that it has a few mistakes as well as passing over the very real differences between the two publications. So, I wrote a response in the way I treat any publication as an invitation to think and form an opinion; and express it. Passing a draft of this response to a competent observer, I got this reply …”Fortunately, there is little expectation of a review. The point is: was the review positive or negative? Did the reader learn something and gain deeper appreciation, or not? Will they buy the book, or did the review satisfy their curiosity? For most readers, the details are unimportant, as much as you may hate this very concept.”

Why I should hate the concept of most readers regarding the questions of detail unimportant I do not know. But I do think the accuracy, in respect of detail or general, is very important. What my commentator was implying is that the review met the requirements that he was suggesting, and he added that my response was “nit-picking” and would only be seen as criticism of someone who is widely held in high esteem. The fact is that Steven Hammer is also held in very high esteem by me and I am so glad to be able to say that he expressed to me personally that his “review’ was rather a literary fantasy. What Steven does comment on is a view of the needs of collectors. That they care little about schemes of classification and that labels are necessary irritants. I do not question the truth of this view. But would not accept that this is a justification for the imposition of just any kind of scheme because that is what a writer wishes to propound for reasons of his /her own.

These then are the points I made in my response that I think Steven should have addressed. The ‘mistakes’ are … a). The Audensberg population was actually shown to me by Elsie Esterhuizen many years ago and it is not the place where any haworthiophile would ordinarily look for plants. b). The reference to Drosanthemum bellum is odd because Steven describes this as a “niche-sensitive species”. This “species” is at the heart of a very long and detailed story of Drosanthemum micans that I once wrote and lies unpublished. I would surely have used this as an example of the way in which botanical science has also failed us. D. bellum is a pink flowered variant in a much-localized population of D. micans that also has white, purple and red variants. This tiny population sits among a larger widespread population of yellow flowered variants that go by the name of D. hallii. This is turn has variants that include the typical bi-colored flower of the older D. micans that is common north of Worcester. Further variants occur north-east of Montagu, to Oudtshoorn and then south to Mossel Bay (D. edwardsii) back west to Bredasdorp, (D. lavisii, D. aureopurpureum and D. croceum?). The problem here is the failure to establish what is meant by “species”. To refer to D. bellum as a species is a misconstrual of science, or an example of the liberties that are taken with Latin names – botanists and collectors alike. c). Chameleons. Wonderful words of Steven’s, but not quite complete. The story about chameleons’ parallels that of the very low non-tech problem of impossible identification even when there are heaps of “characters” to use. It fortuitously exposes the probability that we are being led up a garden path by high-tech. I have used chameleons in the same way that I studied Oxalis ultimately demonstrating that species are complex systems of variables! d). Kaboega is not the only area I know exceptionally well and it also figures in practically all the other volumes of Updates of which there are five. I doubt if these have ever had much coverage, but they are an account of my voyage of exploration and discovery that is the concern of mine in respect of omission. In the Updates I discuss the populations and their variants as they occur at many different places and show this impact on the application and use of Latin binomials. There is a prevailing misconception that this is only a problem within Haworthia. I show that this is not so. I also make several references to the fact that Haworthia is by no means an integral single genus and that the nature of genera in the Alooideae needs proper attention based on a lot more than the fact that the Haworthia subgenera have small flowers. e.) “Shaggy dog” for H. mirabilis Ballyfar is not a name I coined but Steven himself.

I do think the omission is in the comparison of the books where there is in fact none. My Update Volumes revolve around the way that science has let us down to the extent that any pretender can take up the mantle of taxonomic expert. Botany provides no species definition and hence the Latin binomial is not required to carry any meaning other than a guise of authority. Whatever collectors may require has no import whatsoever in a process of classifying plants as biological entities. They are focal points for the collection and storage of knowledge indicated by Latin binomials and these are not simply and only intended as labels. Even I recognized this as a child when I wanted to know what the plant actually was that my father called H. chalwinii, and where it came from. Every collector who refers to names at all surely expects and believes that there is some mystic or real knowledge associated with the names he is given and uses. It is an injustice to any collector to coin Latin names outside of the context of science where the collector is entitled to believe they belong.

The only predicable thing about Breuer’s system, which is a watered down version of a much more focused and detailed one by Hayashii, is that there are going to be a lot more names. This is not only when someone else climbs the Audensberg, or recollects the Sandhills population, or drifts across into the Heatley Peak area. Throughout the Updates, I warn and demonstrate that character fixated taxonomy may be very misleading. Vavilov was a Russian botanist who pointed out that variations in a genus may be expected in all the members of that genus. Species are therefore to be seen as systems that are natural assemblages of plants that can be associated in respect of ALL the forces, factors and features that generate them – not propagules of single clones that fill availability lists and price catalogues. Drosanthemum bellum is just such an example of how Latin binomial names are used to describe variation within species, rather than to properly organize the basic entities that make up the entire living system.

I have, even in the Updates, shown how the watchdogs of science let us down. I have tried to communicate my experience and observations to a wider and expectedly interested audience. This, in the hopes that it would lead to greater understanding and comprehension of the problems of finding names as the backbone of communication, appreciation and understanding. It is a huge disappointment to me that I have achieved very little other than to grow wiser myself. One of my many critics makes a show of taking up middle ground between me and other Haworthia taxonomists. My response is that taking up middle ground between myself and the ignorant is not going to be very productive. In the first place there is not much space there as I am quite aware of my intellectual limitations. In the second place I have not actually been all that certain that my overview is entirely correct. Despite being credited with a lot of field work (and no good sense to go with it) I am extraordinarily aware of how much I have not seen. This adds to my discomfort as I see a proliferation of new names, gaily forgetting the multitude that I moved aside in my Revision. These are often based on propagules from my own collections (concealed by the creation of new collecting numbers that are not mine). I recognize that the only predictive element in this kind of science is that we can expect many more Latin binomials in a collector driven system rather than one of botanical science. So indeed I see no change from the failed methodology of von Poellnitz, Smith and Scott.

There is no comparison at all between the two books that a true review might have suggested. One book (Breuer) is a collector’s guide to a limited range of variants (albeit 336?), while the other (Bayer) is an account of a very wide range of variants and a hypothesis (not a statement) of how they are related. The latter also throws some light on the universal quandary knowledgeable observers soon come to experience viz. Elton Roberts in the same edition of the Journal where he questions the identity of Mammilaria lasiacantha. The problem he has is a classification fixated on superficial small differences rather than one based on the realities of the variation that should be expected in any system arising from, and actively responding to, differences related to time and space. The essence of science is that an experimental method is applied to a sample and repeated, the result will be the same. If everyone is coming up with a different plant classification, it should occur to us that there is something wrong with the method and perhaps also the hypothesis that is being tested.

I am curious why my commentator dismissed all the above as “nit-picking” when myself I feel that they deal with the most significant elements of writing at all. Especially, they touch on the very core of why we even classify and name plants in the first place. My response should not be seen as a criticism of my commentator, or of Steven, a remarkable man who is also very dear to me. I also respect him enormously for his empathy with plants because if there is anyone who projects my view that this is a conscious creation all the way down to its rocks, it is him. There is surely purpose in creation if only that we should seek and find what that is. ♦

Haworthia Updates

Thoughts and observations on Haworthia

Author Archives: Bruce Bayer

Volume 7, Chapter 7:- More on Haworthia mirabilis and H. mutica from east of Bredasdorp

Haworthia flowers – some comments as a character source, part 1

Haworthia flowers – some comments as a character source, part 2

Haworthia flowers – some comments as a character source, part 3

Haworthia flowers – some comments as a character source, Appendices 1, 2, 3

Haworthia flowers – some comments as a character source, Appendix 4

Haworthia flowers – some comments as a character source, Appendix 5

Haworthia flowers – some comments as a character source, Appendix 6

Haworthia flowers – some comments as a character source, Appendix 7

Haworthia flowers – some comments as a character source, Appendix 8

Haworthia flowers – some comments as a character source, Appendix 9

Haworthia flowers – some comments as a character source, Appendix 10

A myth corrected to – Haworthia maculata var. livida (Bayer) Bayer – and flowers ignored

A myth corrected – part 2

A myth corrected – part 3

A myth corrected – part 4

A myth corrected – part 5

My view of names

Conclusions and Acknowledgements

Haworthia Update Vol. 9 – Addendum

Very brief note re Haworthia nortieri flowers

Haworthia marxii and H. truteriorum in relation to rational classification.

The reality of Drosanthemum micans L.

New finds in Haworthia

What do collectors need?