Natural Variation and Species Delimitation in Haworthia Duval. – Part 6. HAWORTHIA MACULATA (V. Poelln.) Bayer. (1987)

Printed in British Cactus and Succulent Journal 5:13 (1987)
Part 6. HAWORTHIA MACULATA (V. Poelln.) Bayer.
M. B. Bayer, National Botanic Gardens of South Africa, Karoo Gardens, Worcester.

Haworthia maculata (V. Poelln.) Bayer. Haworthia Handbook: 130 (1976);  New Haworthia Handbook: 43 (1982);  J. W. Pilbeam, Haworthia and Astroloba: 89 (1983).  H. schuldtiana var. maculata v. Poelin. in Feddes Repert. Spec. Nov. 49: 25 (1940).  Type: Cape; in the neighbourhood of Worcester, Swellendam, Bredasdorp and Caledon. Major H. Venter No. 6a. No specimen preserved.  Neotype: Cape- 3319 (Worcester): Worcester District, H. Venter No. 6a in G. G. Smith 3912 (NBG)

scans 005

Rosette stemless, 30-70mm in diameter, up to 60 leaves.  Stem thick, white-fleshed, non-fibrous, 1/3 diameter of rosette, never elongate, slowly proliferous from base.  Roots thick, white-fleshed, non-fibrous.  Leaves erect, spreading, slightly incurved at tips, up to 60mm long, 12mm broad, 8mm thick, ovate-lanceolate acuminate, aristate with bristle up to 5mm long, seldom setiferous, green to grey-green, purplish- green in sun, lines distinct with pronounced small longitudinal transparent areas on upper surface, lower surface heavily mottled or with longitudinal markings, spots occasionally with small hairs, face slightly concave at base, convex-turgid towards upper 1/4, back convex, frequently with second keel or with double row of spines on keel, margins sub-acute, lightly spined, spines white, up to 0.5mm long to 1mm apart.  Perianth white, yellowish-green inside, nerves pinkish-brown outside, pinkish inside at tips, tube ascending curved, up to 16mm long, obclavate, bluntly triangular at base, 4.5mm across reducing to 3mm, segments free, regularly stellate, inner lower segments incurved along upper margin.  Buds biarcuate, bifid at tips.  Flowering Oct-Dec.

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Haworthia Revisited – 17. Haworthia maculata

17. Haworthia maculata (V.Poelln.) Bayer :130(1976).  Bayer :43(1982).  H. schuldtiana var. maculata V.Poelln., Feddes Repert.Spec.Nov.49:25(1940).  Type: Cape, Worcester, Swellendam etc. H. Venter 6a.  Not preserved.  Lectotype (B&M): Worcester, Venter 6 (BOL).

maculata: spotted.

Rosette stemless, proliferous, to 8cm φ.  Leaves many, sub-erect to spreading, purplish-green, spotted, short spines on margins and keel.  Inflorescence simple, slender.  Flowers 15-20, few open, white, yellowish in throat, green veined.

1982 – The original locality for this species was not accurately recorded but it was collected by Major H. Venter, and so it is possible from Smith’s and Long’s records to trace it to the Brandvlei Dam south of Worcester.  H. maculata flowers in October/November and the form of the flower is like that of H. herbacea and H. reticulata, rather than like that of H. magnifica (H. schuldtiana).  There is a clear intergradation with H. herbacea in the area, and the plants are very similar to H. herbacea except that there are fewer and more turgid leaves.  H. maculata also occurs further south in a similar quartzitic rock formation to that at Brandvlei Dam.  A problem is the occurrence of populations apparently of H. maculata in the mountains to the north of Worcester (both high altitude ‑ Audenberg Peak, and low altitude ‑ at Sandhills).  A similar element occurs eastward towards Robertson at Buitenstekloof, distinguished again from H. magnifica by an earlier flowering time and the wide spread of the tips of the upper perianth lobes.

1999 – It is difficult to know just what is present on the higher mountains.  Few succulentophiles are also mountaineers, and besides the plants could be expected to be on rocky north faces which may not attract the conventional high altitude botanist.  H. nortieri bears some resemblance to H. maculata and that species is found as far south as Opdieberg (Ceres).  It is quite probable that populations may occur elsewhere in the area between Worcester and Citrusdal.


a. var. maculata
This variety seems to be linked to H. herbacea with possible ties to H. nortieri.  The similarity to other high mountain forms (eg H. vlokii and H. turgida) cannot be overlooked and this has been repeated at several places in this book.

Distribution: 3319 (Worcester): Brandvlei Dam (‑CB), Bayer 164 (NBG), Smith 3912 (NBG); Bayer in KG669/69 (NBG); NE. Brandvlei Dam (-CB), Bayer 2591 (NBG); Audensberg Peak (-CB), Esterhuysen 16706 (BOL), Bayer 1119 (NBG); Moddergat (-CD), Bayer 1145 (NBG); Keeromsberg, Boskloof (-DA), BOL28719; S. Sandhills (-DA), Bayer 1120 (NBG).

b. var. intermedia (V.Poelln.) Bayer stat.nov. 
H. intermedia V.Poelln., Kakteenkunde 9:133(1937).  V.Poell., Feddes Repert.Spec.Nov. 44:233(1938).  Type: Cape, Robertson, McGregor, G.J. Payne  Not preserved.  Lectotype (designated here): Epitype (designated here): CAPE-3319 (Worcester): Buitenstekloof (-DC), Bayer 4461 (NBG).

intermedia: between.

In the case of this variety, Payne (priv. comm.) did indicate the actual origin at Buitenstekloof west of Robertson.  Von Poellnitz’ later citation for Scottburgh, Port Elizabeth, as well, is indicative of the close resemblance of even very different species and the difficulties which arise in trying to identify them consistently and correctly.  In his discussion von Poellnitz concluded that while the plants had the long end-awn of H. mucronata, the reticulated patterning of the leaves was that of the H. reticulata group.  The name suggests the difficulty in deciding just what to do with this element.  It co-occurs with H. reticulata and with H. arachnoidea and bears a very close resemblance to the shale form of H. maraisii var. notabilis.  As already noted it has a different flower and flowering time to that variety.  Nevertheless it may be correct to place them together in one species as there is also a population recorded mid-way between the two at Agtervink.  Possibly a more direct link with H. turgida should be sought as the plants do bear a close resemblance to the montane forms of that species.  Certainly it is possible that there may be a connection somewhere in the mountains between Robertson and Swellendam.

Distribution: 3319 (Worcester): Buitenstekloof (-DC), Bayer 4461 (NBG).

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

1. Looking down the north face Die Nekkies east to west

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

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

(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.

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A myth corrected – part 2

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

This is the type locality (ie. 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).

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A myth corrected – part 3

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.

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A myth corrected – part 5

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 vonPoellnitz 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.

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Haworthia maculata ↔ Haworthia pubescens, MBB8002 Cilmor.

In Haworthia Update Vol 9 there is a report of a population MBB7997 identified as Haworthia pubescens from north of the Cilmor wine cellary. This is approximately 2-3km southwest of the type locality for the species.  I noted that the plants have less spinuliferous leaf surfaces and there is a degree of surface translucens and maculation (spotting). I also presented 3 pictures of MBB7271 of what I identified as H. maculata from south of the Cilmor cellar. When I first visited this locality I had no problem identifying the few plants I saw as H. maculata on account of their marked spotting.  However, on a recent visit we struggled to find plants at all and the few plants we found were too embedded in rock cracks to make any worthwhile identification.  So we revisited the site to explore more extensively and located a large number of plants higher up and slightly west of our first sightings.  These plants are illustrated here. They incline more to H. maculata than the plants at MB7997 and I have accessioned the population as MBB8002. There is the usual expected large variation in respect of superficial and observable characters. The plants can be proliferous and cluster, more so than at MBB7997. Similarly the leaves can have more translucens and even less spinuliferousness of the surfaces. Some plants have few and quite thick swollen leaves while others may have more and very slender pointed leaves. I have not observed the flowers and really do not expect them to make any difference to the problematic classification of populations that again are neither here nor there in a narrow concept of species. H. herbacea occurs at all four geographic positons at a radius of about 2km. At the brickfield to the northwest as well as just northeast of the Brandvlei Dam wall it is evident to me that there is a transition between H. maculata and H. herbacea. I did report the known distribution of H. maculata in Update 9.  While there is no suitable habitat between Die Nekkies hills at the Brandvlei Dam and the Audensburg or Kanetvlei, there is unexplored suitable habitat southwards to Moddergat and Hammansberg.  There is no evidence of H. maculata eastwards to where H. reticulata is known about 15km east on Ribbokkop.  Westwards no Haworthia is known although G.J. Payne did inform me that he had observed plants in the hills immediately southwest of the Dam at the now submerged hot spring in the Brandvlei prison area.

The submitted pictures include two views. View 1 is looking north of east across the Breede River to the Sandberg where H. pubescens occurs. Its full occurrence on those low hills is not known and this I will explore soon. View 2 is looking eastwards looking at a Dwyka Tillite hill across the river in the upper right. We found no Haworthia on that hill although both H. pumila and H. herbacea are present on the smaller rise to the right and behind it – also Dwyka.  H.herbacea is very abundant on a Dwyka tillite hill about 10km to the south. The corresponding hill on the left is Ribbokkop where H. herbaea, H. reticulata and hybrids are present, and H. arachnoidea also occurs. The limits of H. mirabilis are the higher hills in the background viz. Rooiberg, Gemsbokberg and those are Witteberg sandstones.

Addendum Haworthia pubescens MBB8011, SW Sandberg

Addendum.  H. pubescens MBB8011, SW Sandberg.

I need to point out that there is a still earlier article which covers Haworthia maculata (Haworthia maculata <–> Haworthia pubescens) that lays the basis for this discussion.  In that article I note the position of the Sandberg to Cilmor and DeWetsberg and intended to include the Sandberg H. pubescens in that article.  We could not get landowner contact and so that fell away.  However, this problem was overcome and we first explored a Dwyka Tillite outcrop southeast of Sandberg.  There is a vast accumulation of windblown sand on the first hill and we saw no Haworthia.  There is a smaller hill further to the southeast that is also Dwyka and erosion exceeds wind deposition so smaller non-geophytes do quite well.  We found both H. herbacea (see fig.1 MBB8014) and  H. pumila there.  From there we went to the southernmost point of the Sandberg.  A misjudgement landed our vehicle in mud and the drama to get out limited the time we had to explore.   We found a lone H. herbacea (fig. 2 MBB8012).  Returning a week later we approached the Sandberg from the southwest, and almost immediately on reaching the top we found H. pubescent.  Fig. 3 is a view towards Cilmor and DeWetsberg where the plants appear to be intermediate H. pubescens↔H. maculata.  The picture is useful to get some idea of the role of geographic and geological considerations.  The high mountains in the background are Table Mountain Sandstone and no Haworthia is known there.  I am not certain that this is true and G J Payne did tell me that he had seen plants on the extreme lower right and south of the Brandvlei Dam.  But also on the absolute distant and absolute left, is the Riviersonderend Mt.  That is also TMS.  The deep Wolfkloof Valley behind that is the locality for the much unexpected H. herbacea ‘lupula’.  (These inverted single commas are not entirely necessary but I use them to underscore my informal use of names that have less reality.  The var. lupula is real).  The mountains ahead of that last line are Hammansberg on the left and the Moddergat to the right.  Between there and DeWetsberg has not so far turned up Haworthia, but this is an exploration problem.  Behind the DeWetsberg is also underexplored.  H. herbacea does occur between DeWetsberg and behind the mountains on the low right just in the picture and also east of the brickfield out further right.  H. maculata is only known in this area along the Nekkies north (further to the right) of the Brandvlei Dam just visible in the picture.

Prior to this exploration H. pubescens was to me only known from the northern part of the Sandberg that lies south of a road going eastwards to Eilandia between Worcester and Robertson.  Here H. herbacea does occur on the lower northwest warm slopes.  H. pubescens seems to occur only on the upper two ridges and H. herbacea is not known to intermingle with it.  This is all Witteberg Sandstone that as a formation overlies Bokkeveld Shale and underlies TMS.

Coming back to the southwest corner of the Sandberg where we found H. pubescens.  The plants seem very similar to the species as it occurs to the north.  They were very cryptic and often in shady rock retreats where they were really hard to see.  It was mid- to late-morning when we were there and the plants were not going to be better exposed as the sun moved further west.  Although there was very suitable well-drained habitat lower down on the shaded east slopes, there were no plants and I speculate that this may be because the plants may need the cooling effect of wind movement up on the ridge.  The pictures tell the story of variability in respect of a whole range of leaf and rosette characters.

It is worth noting fig. 43 of the dead remains of a plant under a clump of restioid.  It seems that seedling survival is closely coupled to early protection giving rise to the concept of nurse-plants.  Plants are often very difficult to find because they are so hidden beneath accompanying vegetation.  But they do need light and the dynamics of vegetation growth and densification must have quite a big impact on the ageing and survival of plants.  It raises again the question of how long do the plants live?  For plants like Aloe ferox and A. dichotoma I do have a real experience of a lower limit of about 35 years and a top limit in the several hundred.  In  the field, the plants seem comfortably ageless.

The really interesting part is this.  While I was busy tediously cleaning a plant to photograph it, Daphne called to me to come and see a lighter green plant she had seen.  Moving in that direction I saw a plant that registered as H. herbacea but with some hesitation and doubt (see fig.4).  I then went to see what Daphne had observed.  They seemed less obviously H. herbacea but that seemed to be a logical and conservative opinion (see figs. 5-11).  That was until Daphne found two adjacent rosettes at the foot of a restioid clump that left me in no doubt that they were hybrid H. pubescens/maculate (see figs 9-12).  These were in bud whereas H. pubescens plants showed no sign of impending flowering.  Note the buds are less well developed than MBB8014 further east, even if possibly insignificant.  Going back to the other plants we confirmed my doubts.  They were a lighter colour and apparently softer texture that we would have expected in H. herbacea.  These were the only plants we saw in a space bridging the occurrences of plants of H. pubescens.

12-46 MBB8013 H. pubescens, SW Sandberg.

We did not explore the western slopes where habitat would have been more suitable for H. herbacea and I expect it does occur there.  What puzzles me is that so frequently have I found very distinctive hybrids between species in close proximity and very seldom where the species are some distance from one another.  I cannot say I have ever found a hybrid in the clear absence of both parents.  The example of Astroworthia bicarinata at Lemoenkloof, east of Barrydale, may be an exception where only Astroloba corrugata (syn A. muricata, A. aspera) is present but H. pumila apparently not.  Hybridization is thought to be an important element in the “evolution” of new species.  I doubt this as it is quite evident that separation into two species is a pre-requirement.  If new species have evolved in Haworthia by hybridization, how did they evolve as such in the first place?  The answer to me lies in the continuities between populations.  I observe, and have experienced of expected continuity between populations.  While the Cilmor populations are thought to be H. pubescensmaculata it cannot be said anymore that they are hybrid, or populations where the morph or drift to discrete elements has not reached a conclusion.  The latter is more likely.  As there is already apparent geographic continuity of H. maculata and H. herbacea, I was expecting some evidence of a similar relationship between H. herbacea and H. herbacea.  So here it is.  Hybridization as a factor in speciation in Haworthia does not seem to very likely.  It confirms for me that there is a fractal “chaotic” order to species in Haworthia and the reality is that a view of many truly discrete species is a fabrication and a very ill-considered view.


We are always greeted with such kindness and helpfulness that we might have expected this from the Sandberg landowners too.  It came in no small measure.  Driaan Griesel was most enthusiastic and interested and also helped us with extracting our vehicle from the mud on the one occasion, and then jump-starting it after a flat battery on the second.  Our imposition did not so much as touch his view of the day.

(ed. – Bruce made another visit to Southwest Sandberg on 9 December 2012 and includes the following flower pictures.  He makes this comment; personal correspondence 27 December 2012.)

I am actually not sure at all about flowering time now.  I used to be quite sure of being able to collect seed of pubescens mid-Nov.  But I observed at Humansdorp that gordoniana peak flowering could be out by 6 weeks.  In  any case the plants can produce successive spikes so one can get delayed flowering and added to that energy in the first or the second flower set.  I know mirabilis at MacGregor can flower from Nov. thru to March while at Montagu mirabilis can flower as late as April/May.  Retusa and geraldii are quite happy to produce flowers in either Spring or Summer and Kobus observed that splendens did that too.  Maculata can flower from Sept. thru to late Dec.  And each population does its own thing.


Die Nekkies Haworthia – A further visit and updated information

Bruce Bayer has now been back to Die Nekkies and returns with more detail to add to the broader Haworthia maculata picture, and considerable detail for H. maculata at its home range located north of the Brandvlei Dam on a low range of hills named Die Nekkies.  (See Haworthia Updates Volume 9, A myth corrected to – Haworthia maculata var. livida (Bayer) Bayer – and flowers ignored). These low hills extend for 10-12km east to west along the north shore of the Dam. They are geologically Witteberg Sandstone and H. maculata is not known off this formation except in the lower Hex River Pass where it is found in Table Mountain Sandstone.  H. herbacea (and H. reticulata) are more abundant on the Dwyka Tillite, and on Bokkeveld and Ecca shales. The Wiiteberg is sandwiched between Bokkeveld Shale and Dwyka Tillite, and Ecca Shale of the Karoo System overlies Dwyka Tillite.  Because of considerable faulting and folding the Cape Terrain is geologically and topographically very broken and there are a multitude of skeletal rocky habitats – the raw rock is very exposed and the surfaces are erosional. Haworthia are rarely found on depositional sites. The fact that there is also a stable and dependable winter rainfall also contributes greatly to the success of small succulents in situations that cannot maintain high biomass levels. This all drives home the fact that Darwin’s dictum that distribution is the lintel to understanding species is doubly true for Haworthia where morphological differences are minimal.

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Haworthia maculata leaf tips, faces and backs

Die Nekkies – a selection of Haworthia maculata plants with their leaf faces and backs showing variability over this extensive area.

I took the following photographs originally just to show how much the leaves varied in the species. The only locality data record was Die Nekkies with no differentiation into populations. The variability was firmly established and became apparent even within populations where the degree of variation surprised even me. It was also extrapolated across the genus.

As an afterthought, this turned my mind to Farden and Smith’s observation about the idiocy of trying to make varieties in Haworthia attenuata on the basis of leaf characters where the individual leaves varied so much that several varieties could be detected from the same plant. This is exactly what Breuer, Marx and Hayashi are actually doing when they get carried away by their justification for new species, drawing conclusions from single plants because of the inability of the mind to hold many images. This is exactly the way in which “H. magnifica” is maintained as a species when the imagery is based on the original single plant originally described. There can be no such thing as “H. magnifica” because the variation across the range of populations from Riversdale to Riviersonderend is so immense and complex.

The following photographs show a random selection of plants and their leaf faces and backs from Die Nekkies. These were taken some time ago when I did not record actual populations. They do record the variability of plants and leaves over this wide area which I classify as Haworthia maculata. For subsequently photographed plants I have recorded my population collection numbers which are indicated on the map found in Die Nekkies Haworthia – A further visit and updated information.

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