12. Haworthia floribunda V.Poelln., Feddes Repert.Spec.Nov. 40:149(1936). idem. 44:228(1938). Bayer :117(1976). Bayer :39(1982). Scott :58(1985). Type: Cape, Heidelberg, Mrs E. Ferguson. Not preserved. Lectotype: icon (B). Epitype (B&M): Blackdown, N Heidelberg, Bayer 158 (NBG).
floribunda: many flowered.
Rosette stemless, up to 3cm φ, slowly proliferous. Leaves 20-30 dark green opaque, upto ovate-lanceolate, spreading, twisted with flattened, rounded tip, margins scabrid to dentate. Inflorescence simple, to 250mm. Flowers 10-15, greenish-white, few open together.
1982 – This is a very interesting small species with twisted lanceolate leaves with blunt rounded tips. It was described from plants collected north of Heidelberg (Cape) where they are all glabrous and where hybridisation with H. turgida also occurs. It has not been collected further west although there is a very old collection in the Botanical Research Institute (PRE) from Swellendam. Around Riversdale the plants may have scabrous leaves with denticulate leaf margins. There is a known population north of Albertinia in which the plants have more and shorter leaves, as well as another similar population near Gouritzmond. These two populations may suggest an affinity with H. chloracantha, as put forward by A.E. Speechley (unpublished). There may be such a relationship, but it seems likely that H. floribunda and H. parksiana are in fact related. They both tend to grow well‑shaded and in moss and lichen. Further exploration of the Gouritz River tributaries may produce an answer to this puzzle.
1999 – The Gouritz valley has provided no answers to the above puzzle but there have been a number of other significant collections. There is a population south of Swellendam of which specimens in cultivation have been very robust. They are relatively light green in cultivation and also individual plants are smooth as was the original type. Odd specimens have pointed leaves. Northwest of Swellendam is a similar but darker green plant also with pointed leaves and this is taken to be H. variegata. Further south from Swellendam, H. floribunda assumes the same form as around Riversdale. At Great Brak there is a population of plants growing with H. pygmaea which were assumed to be H. floribunda, but they are probably simply H. chloracantha var. denticulifera. This is true also of the populations south of Albertinia and also of one population south of Heidelberg which has pointed leaves. The flattened leaf-tip is not confined to this species, and appears in H. magnifica and H. maraisii. A relationship with H. variegata is probable through the two populations in the vicinity of Swellendam.
a. var. floribunda. The typical variety is not typical of the species at all and this glabrous variety is only known from the type locality at Heidelberg where it is very scarce. Hybrids with H. turgida are also present.
Distribution: 3420 (Bredasdorp): 6km N. Heidelberg (-BB), Smith 5545 (NBG, PRE), Bayer 158 (NBG); Heidelberg commonage (-BB), Ferguson in BOL20507.
b. var. dentata var. nov. Type: CAPE-3421 (Riversdale): W. Riversdale, Dekenah 90 in Smith 5502 (NBG, Holo.).
dentata: toothed.
Differs in being smaller, to 4cmφ, the leaves very dark green, slightly scabrid and with spined margins. (A var. floribunda foliis subtiliter scabridis et denti-marginatis differt).
This variety describes the smaller form which has distinct and widely spaced marginal spines. The leaves are very dark green and also slightly scabrid. It occurs from the Bontebok Park at Swellendam to northwest of Riversdale, and includes larger forms east of Riversdale also with pronounced marginal spines.
Distribution: 3420 (Bredasdorp): Bontebok Park (-BA), Bayer 3439 (NBG); E. Buffeljachts (-BA), Viviers 156 (NBG). 3421(Riversdale): W. Riversdale (-AA), Dekenah 90 in Smith 5502 (NBG): 5km N. Riversdale (-AB), Smith 5381 (NBG, PRE); 15km E. Riversdale (-AB), Smith 5758 (NBG); Dassieklip (-AC), Venter 92/31 (NBG); Wydersrivier (-BA), Smith 5491, 6781 (NBG), Bayer 2311.
c. var. viridescens var.nov. Type: CAPE-3420 (Swellendam): S. Swellendam (-AB), De Kok (NBG, Holo.).
viridescens: becoming green.
Very green plants with darker coloration at the basal leaf margins. Relatively glabrous and more robust in cultivation. (A var. floribunda habitu robusto et colore viridi vivido ad margines basales foliorum atranti differt).
This is a large robust plant in cultivation and includes two forms, one of which has pointed leaves. The coloration is greener than normal for even cultivated plants of the species.
Distribution: 3420 (Bredasdorp): Below Swellendam Stn. (-AB)., C.A. Smith 2724a (PRE); S. Swellendam (-AB), D.De Kok (NBG).
Introduction Haworthia Revisited was drafted in 1996, and since then the first author has undertaken a number of field excursions in an attempt to clarify uncertainties. The putative nature of species of Haworthia as recognised by Bayer (listed in Haworthia Revisited, Umdaus 1999) and the importance he attached to geographic distribution are stressed in all his publications. This is because these so-called species seem to vary continuously with one another in that context of geography. Classification seeks to portray relationships and origins. Hence when a species has been recognised, a cognitive attempt has been made to speculate on phylogenetics, where distribution must be significant. In the case of Haworthia floribunda this proves rather difficult, and this article is a discussion of the relationship of this species to its possible relatives. The point we do make is that the Linnaean binomial system, as well as cladistic methods, seem neither to deal with nor portray the problem of reticulate relationships. In other words, the nomenclatural system and the way we classify plants and analyse their relationships assumes linear dichotomy in those relationships.
Considerations H. floribunda was described by von Poellnitz in 1938. It was preceded by H. parksiana in 1936. Other species which need to be considered are H. chloracantha Haw. (1821), H. variegata Bol. (1929), H. magnifica VPoelln. (1933) and H. maraisii VPoelln. (1935). Each of these species, although H. parksiana to a lesser degree, has complex variability within assignable geographic space. The difference with H. floribunda is that it is peripheral to all of them.
Bayer, in his Haworthia Handbook (1976), writes of H. floribunda ” … It is not certain what the relationship is between this species and H. chloracantha var. denticulifera which is found north and south of Albertinia on the west of its distribution range. Thus H. chloracantha abuts on H. floribunda and may be geographically continuous with it.” In the New Handbook (1982) Bayer writes “There is a known population north of Albertinia in which the plants have more and shorter leaves, as well as another similar population near Gouritzmond. These two populations may suggest an affinity with H. chloracantha (here an unpublished comment by A.E Speechley is added which probably has its origins in Bayer’s note of 1976, or in private communication between the two persons). There may be such a relationship, but it seems likely that H. floribunda and H. parksiana are in fact related. They both tend to grow well‑shaded and in moss and lichen.”
The problem is not resolved in Bayer’s more recent revision (1999). He does recognise three varieties of H. floribunda. These are:
a. the typical one comprising a single population north of Heidelberg (Fig.1 MBB158).
b. a similar single population near Swellendam representing the var. major (Figs.2a & b MBB6859).
Fig. 02a. MBB6859 H. floribunda var major. S Swellendam.Fig. 02b. MBB6859 H. floribunda var major. S. Swellendam.Fig. 03b. JDV89-17 H. floribunda var dentata. NE Riversdale.Fig. 04. MBB2311 H. floribunda var dentata. N Albertinia.
c. the remainder, the var. dentata (Figs.3a & b JDV89/17).
In the discussion of H. chloracantha, it is also stated that a population north of Albertinia (i.e. Draaihoek, Fig.4 MBB2311) considered as chloracantha in 1976, considered non-commitally as chloracantha in 1982; ” … includes plants which resemble H. parksiana and it may best be related to H. floribunda“. This collection is cited under H. floribunda var. dentata. Thus there is a degree of uncertainty for which the author offers an apology, and here attempt to summarise the situation a little more comprehensively.
Fig. 04. MBB2311 H. floribunda var dentata. N Albertinia.
There are some red herrings in various comments made by Bayer. These include reference on p48 of Haworthia Revisited to the lower Gouritz population (Fig.5 MBB5460) “previously being assigned to H. floribunda“, possibly referring to a Smith identification on a photograph and cited under H. chloracantha var. denticulifera. There is a similar population nearer Albertinia (Fig.6 MBBsn). Also there is a citation of a specimen from the Duiwenhoks Causeway (Muller-Doblies 89/098) under H. chloracantha var. denticulifera, which may be the same population sampled by Venter and Esterhuizen (Figs.7a & b JDV92/31) from a site named as Dassieklip also on the Duiwenhoks River (the geographical grid references differ and these may be incorrect), but cited under H. floribunda var. dentata.
Fig. 05. MBB5460 H. chloracantha var denticulifera. NW Gouritzmond.Fig. 06. MBBsn H. chloracantha var denticulifera. S Albertinia.Fig. 07a. JDV92-31 H. floribunda var dentata. Dassieklip, S Heidelberg.Fig. 07b. JDV92-31 H. floribunda var dentata. Dassieklip, S Heidelberg.
Haworthia parksiana is known from at least five different sites of which two are recorded by Mr Jan Vlok, viz. at Outeniqua Siding, and north-east of Brandwacht, Mossel Bay. No herbarium specimens of these are known to have been lodged. J.D Venter has commented that parksiana-like seedlings were grown from H. magnifica seed (parent plants ex near Heidelberg). It is doubtful if this can be used to support a claim that this is the origin or relationship of the two species. Bayer has also grown small stunted forms of magnifica from seed of an H. magnifica population south of Riversdale. Such dwarfed individuals have been observed among seedlings derived from other populations. The connection of H. magnifica to the Mossel Bay area is just too tenuous. At Albertinia itself we have the variation to H. magnifica vars. dekenahii and splendens, while to the south west we have the var. acuminata. This is assuming that Bayer is correct in making these combinations which are known to be complicated by consideration (among many) of H. emelyae north of the Langeberg mountains.
H. chloracantha is treated in Haworthia Revisited, as three varieties. The typical variety is considered to consist of the known single population north of Herbertsdale (Bayer in KG411/75). However, Paul Vorster drew my attention to a population at the Wolwedans Dam north of Great Brak which is almost identical and which invalidates varietal distinction defined by geographical considerations (Fig.8 MBB7425). A specimen of this collected by C Burgers is also cited for the Gouritz Gorge where it exits the Langeberg Mountains. P.V.Bruyns and E.A.van Jaarsveld have also made collection from further north in the same gorge, which are uncited. These three populations cannot be categorically included in the typical variety and could be just as at home in the var. denticulifera. The only consideration is probably size and the greenish colour – as opposed to the usually smaller and purplish-green of the var. denticulifera. The var. sub-glauca also does not have a clear geographical base in terms of difference or distribution range, and may just be a much localised ecotype in the vicinity of Great Brak. It occurs close to the Wolwedans population both to the south and to the west. The recognition of these varieties makes it a little difficult also to include the Albertinia and Cooper Siding populations as cited in Haworthia Revisited under H. chloracantha as they are so closely similar, and located, to H. floribunda. The geographical considerations, however, argue for their inclusion.
Fig. 08. MBB7425 H. chloracantha var chloracantha. Wolwedans, Great Brak. MBB7425
The Great Brak/Mossel Bay area contains some endemic species which suggest the operation of a mechanism which could have isolated the vegetation of that area at some time in the past. H. parksiana, H. kingiana, H. pygmaea, H. chloracantha and species like Euphorbia bayeri and Duvalia immaculata suggest this. The Langeberg mountains are an east/west barrier separating the more arid Little Karoo and its succulent Karoid vegetation from the Southern Cape where Renosterveld and Karoid Valley Bushveld are more strongly represented. The Gouritz River Gorge may have been, or is, a similar north/south divide. Westwards, the Swellendam area seems to provide a vegetation interval that H. turgida seems to bridge with difficulty, and which also marks some kind of a break for the H. magnifica/H. maraisii complex. This divide may be a function of inadequate collecting or the lack of adequate habitat. The Breede River valley may, like the Gouritz, present a north/south barrier. It is the home for H. venosa subsp. venosa and also for an endemic asclepiad Stapeliopsis stayneri. West of Swellendam is also the start of the Worcester Robertson Karoo which is vegetatively much closer to the Little Karoo than is the eastern part of the Southern Cape. This area also has its own Haworthia endemics or near-endemics.
The physical commonalities of the three species floribunda, chloracantha and parksiana, are that the leaves are in a stemless rosette, having from 20-40 leaves per rossete. The lowest numbers are in floribunda with parksiana and chloracantha generally having more. The leaves are firm and slightly scabrid. H. chloracantha is more spinose than the other two species and the leaves are triangular in cross-section with a conspicuous keel. In H. parksiana the leaves tend to be short and sharply recurved. In H. floribunda the leaves are more erect, although spreading. Characteristically the leaves are twisted and there is no keel, so the leaf is more effectively strap-shaped particularly towards the apex. The leaf-tip is also rounded with a short point. In H. chloracantha, the leaves are erect and spreading, not twisted. They are keeled and thus triangular in cross section. The leaf tip is attenuated and pointed. The leaf surfaces of the three species are variable. Both H. floribunda and H. parksiana can exhibit quite tuberculate surfaces, although parksiana never has the relatively glabrous surface that may occur in either chloracantha or floribunda. Colouration is also variable across the three species, although parksiana does not occur in the lighter colours that the other two species may exhibit. A hatched pattern of the under-leaf surface may appear. All three species make offsets although parksiana is slower and more reluctant to do so – an observation which may just be peculiar to the individual grower and the clones he has under his conditions.
New material Regarding H. parksiana, the only new records are those of Jan Vlok mentioned above. The writers assume that the identifications are correct and that they establish the discrete identity of that species. Regarding H. chloracantha, there is only one significant new collection. This is one of the var. denticulifera made by Bayer, Kent and Venter. It was made at the same time as a visit to the site of the typical var. chloracantha north of Herbertsdale. Here the plants are on a very steep clay slope (in fact recent rains had caused massive mud-slides). The plants are quite large and form immense chlorotic-green clumps. We were at Herbertsdale to verify a collection of H. pygmaea made by Ernst van Jaarsveld in the same vicinity. This is a short distance away on a steep conglomerate slope. H. pygmaea was present on the dry northern edge. But in the moss and lichen of the cooler eastern overhang were the small single plants of H. chloracantha (Figs.9a, b & c JDV97/136). These were so evidently different from the typical form that we accepted (with reservation) the identification as H. floribunda. One reason for such identification is simply one of informal communication. A similar “floribunda“-like plant grows with H. pygmaea and H. parksiana virtually within Great Brak Township (Fig.10 JBouwer sn). This was collected and referred to as H. floribunda by both messrs. H. Gie and J. Bouwer. It is possible, and probable, that it is in fact H. chloracantha var. denticulifera, as is the new population JDV97/136.
Fig. 09a. JDV97-136 H. chloracantha var denticulifera. Herbertsdale.Fig. 09b. JDV97-136 H. chloracantha var denticulifera. Herbertsdale.Fig. 09c. JDV97-136 H. chloracantha var denticulifera. Herbertsdale.Fig. 10. JBouwer sn H. chloracantha var denticulifera. Great Brak township.
The Dassieklip population (Fig.7) remains somewhat of an enigma. A good comparison can even be made with collection of H. monticola from Trompetters Poort, north of Willowmore, as the plants also have an apparently smoother epidermis than appears evident in chloracantha. This is a highly improbable relationship, but it is necessary to make this comparison in view of what has been speculated elsewhere about the possibility of a continuation between H. chloracantha, H. variegata and H. monticola (H. divergens Bayer!). This speculation has been by Bayer and is not significant enough to cite in detail. What is important is that the Dassieklip population may in fact be closer to H. variegata. Hence its citation under both H. chloracantha and H. floribunda is unavoidably misleading. However, that can only be considered in a detailed evaluation of H. variegata in its relationship to the coastal limestones and re-occurrence in shales both west and south of Swellendam.
The newest finds relating to H. floribunda are in the broad area southwest of Heidelberg, and west of Bredasdorp. However, there is one new record by E. Aslander of plants from northeast of Albertinia (Figs.11a, b & c EA1238) in which plants very clearly have the characteristics of H. floribunda. There are also individuals with the leaves of H. chloracantha. There is another collection by Aslander from Snymanskraal west of Albertinia (Figs.12a & b JDV92/2) which resembles the Draaihoek sample and thus may also be H. chloracantha var. denticulifera. A further collection by J.D. Venter is midway between this collection and Riversdale (Figs.13a, b, & c JDV93/56), and is clearly H. floribunda. A feature of this collection is the striated rugosity of the leaf surface in some clones and also the occurrence of short, almost terete, leaves that are evident in a clone of H. parksiana in a JDV collection.
Fig. 13c. H. floribunda var dentata. Plattekop, E Riversdale.Fig. 15a. MBB6881 H. floribunda var dentata. S Oudekraal, Heidelberg.Fig. 11a. EA1238 H. floribunda var dentata. NE Albertinia.Fig. 11b. EA1238 H. floribunda var dentata. NE Albertinia.Fig. 11c. EA1238 H. floribunda var dentata. NE Albertinia.Fig. 12a. JDV99-1 H. floriibunda var dentata. Snymanskraal, E Riversdale.Fig. 12b. JDV99-1 H. floribunda var dentata. Snymanskraal, E Riversdale.Fig. 13a. JDV93-56 H. floribunda var dentata. Plattekop, E Riversdale.Fig. 13b. JDV99-1 H. floribunda var dentata. Plattekop, E Riversdale.
Collections from the western areas include:- from a population reported by E. Esterhuizen on the farm Koppies, southeast of Swellendam (Figs.14a, b & c MBB6879) where the plants tend to H. maraisii; from slightly to the east and south of this at Oudekraal (Figs.15a, b, c & d MBB6881) the plants are very like H. floribunda var. dentata as it occurs at Buffeljachts and at the Bontebok Park south of Swellendam. Esterhuizen commented on the appearance of the twisted and flattened leaf-end of H. maraisii north of Bredasdorp. This is confirmed in populations at Napky (MBB7030) and at Adoonskop (Figs.16a, b & c MBB6640) Adoonskop (maraisii) where the plants do indeed look like very robust forms of floribunda. The same characteristic twisted and rounded leaf-tip is evident in maraisii at Napier (Figs.17a, B & c MBB6973) and especially in the seedlings.
Fig. 14a. MBB6879 H. floribunda var dentata. Koppies, Heidelberg.Fig. 14b. MBB6879 H. floribunda var dentata. Koppies, HeidelbergFig. 14c. MBB6879 H. floribunda var dentata. Koppies, Heidelberg.Fig. 15a. MBB6881 H. floribunda var dentata. S Oudekraal, Heidelberg.Fig. 15b. MBB6881 H. floribunda var dentata. S Oudekraal, Heidelberg.Fig. 15c. MBB6881 H. floribunda var dentata. S Oudekraal, Heidelberg.Fig. 15d. MBB6881 H. floribunda var dentata. S Oudekraal, Heidelberg.Fig. 16a. MBB6640 H. maraisii var maraisii. Adoonskop, N Bredasdorp.Fig. 16b. MBB6640 H. maraisii var maraisii. Adoonskop, N Bredasdorp.Fig. 16c. MBB6640 H. maraisii var maraisii. Adoonskop, N Bredasdorp.Fig. 17a. MBB6973 H. maraisii var maraisii. N Napier.Fig. 17b. MBB6973 H. maraisii var maraisii. N Napier.Fig. 17c. MBB6973 H. maraisii var maraisii= floribunda var dentata! N Napier.
Several other collections confound the picture completely either because of intrinsic variability or because they make it difficult to uphold any geographical recognition of variation. Firstly there is a collection attributable to P.V. Bruyns, north of DeHoop (Figs.18a, b & c MBB6539) which has strong resemblance to H. magnifica var. atrofusca.
Fig. 18a. MBB6539 H. maraisii var atrofusca! Tarentaal, N DeHoop.Fig. 18b. MBB6539 H. maraisii var atrofusca! Tarentaal, N DeHoop.Fig. 18c. MBB6539 H. maraisii var atrofusca! Tarentaal, N DeHoop.
Secondly there is a series of populations from the northwestern end of the Potberg, south of Swellendam and east of Bredasdorp. These are:- Juliesfontein (Figs.19a, b, c & d MBB6882); Brakfontein (Figs.20a, b, c, d & e MBB6886); northern Potberg slopes (Figs.21a, b & c MBB6889); and north of Brakfontein (Figs.22a & b MBB6890). These complement collections Burgers 2506, Bayer in KG35/70 and Bruyns in KG49/76 cited in Haworthia Revisited under H. maraisii, as well as MBB(PVB)6544 (Figs.23a & b), and MBB6545 (Figs.24a & b) cited under H. heidelbergensis var. scabra.
Fig. 19a. MBB6882 H. floribunda var dentata. W Juliesfontein, S Swellendam.Fig. 19b. MBB6882 H. floribunda var dentata. W Juliesfontein, S Swellendam.Fig. 19c. MBB6882 H. floribunda var dentata. W Juliesfontein, S Swellendam.Fig. 19d. MBB6882 H. floribunda var dentata! NW Potberg, S Swellendam.Fig. 20a. MBB6886 H. floribunda var dentata. Brakfontein, S Swellendam.Fig. 20b. MBB6886 H. floribunda var dentata. Brakfontein, S Swellendam.Fig. 20c. MBB6886 H. floribunda var dentata. Brakfontein, S Swellendam.Fig. 20d. MBB6886 H. floribunda var dentata. Brakfontein, S Swellendam.Fig. 20e. MBB6882 H. floribunda var dentata. Brakfontein, S Swellendam.Fig. 21a. MBB6889 H. floribunda var dentata. NW Potberg.Fig. 21b. MBB6889 H. floribunda var dentata. NW Potberg.Fig. 21c. MBB6889 H. floribunda var dentata. NW Potberg.Fig. 22a. MBB6890 H. heidelbergensis var scabra! Witklipkop, S Swellendam.Fig. 22b. MBB6890 H. heidelbergensis var scabra! Witklipkop, S Swellendam.Fig.23a. MBB6544 H. heidelbergensis var scabra NW Potberg.Fig. 23b. MBB6544 H. heidelbergensisv var scabra NW Potberg.Fig. 24a. MBB6545 H. heidelbergensis var scabra! Juliesfontein, Potberg.Fig. 24b. MBB6545 H. heidelbergensis var scabra! Juliesfontein, Potberg.
The influence of other species such as H. mirabilis, H. variegata and H. serrata are in evidence. It can be noted that KG35/70 was of very small plants at Juliesfontein. Returning to the same site 30 years later I could not find these small plants at the original site, and instead found the bigger plants of MBB6882 a short distance away. There are three more collections to be considered:- by Denis DeKok near Swellendam (Figs25a, b & c MBB6644), about 10km west of Swellendam (MBB6861), and from the farm Rondeheuwel south of Stormsvlei (Figs.26a, b & c MBB6882, Bayer in KG326/71). This latter collection has previously been reported (Haworthia Handbook 1976) under one of the populations intermediate with H. mirabilis, and is cited in Haworthia Revisited under H. maraisii. These populations, as well as that southwest of Heidelberg (KG107/74 cited under H. magnifica (Figs.27a,b & c MBB6663, Bayer in KG107/74) confound the issue enormously as we actually have four populations which cannot with confidence be allied with either H. magnifica, H. maraisii or with H. mirabilis. The reality is that neither H. heidelbergensis nor H. floribunda can be excluded from the consideration of these populations. The case for each “species” needs to be dealt with separately. In this case, H. floribunda is reflected in the nature of the leaf shapes. In all the four latter populations given, there are individuals which have the same characteristic leaf shape of H. floribunda, although not necessarily the elongated strap-like leaves of the typical form. In addition to this, seedlings of many different collections of H. magnifica (Kweekkraal, Figs.28a & b MBB6817 – and the reader should refer to the chapter where interplay between H. magnifica and H. floribunda is reported), H. maraisii, H. heidelbergensis, H. mirabilis (Goudini, Caledon, Fig.29 MBB6537), and even H. mutica (Hasiesdrift, Fig.30 MBB6982), have the floribunda-like shape evident in the young leaves. Kobus Venter was especially struck by this phenomenon while photographing H. mirabilis on the Bromberg Mountain near Stormsvlei. These rounded obtuse leaf-tips are maintained until at least the three to five-leaf stages and then largely disappear. Thus there seems to be a distinct sign that the leaf-type is juvenile and that H. floribunda represents a “species” with retained juvenile characters. This may extend to the fact that the seedlings of this species also seem to remain slightly distichous for longer than is the case in other species. In the case of H. floribunda var. major from southwest of Swellendam, the leaves in some individuals may be fully pointed and triangular in cross-section and thus more closely resemble plants in the population of H. variegata var. hemicrypta west of Swellendam. This is the same kind of variation found in the populations about Albertinia, where the speculated differentiation to H. chloracantha occurs.
Fig. 25a. MBB6644 H. maraisii=mirabilis! SW Swellendam.Fig. 25b. MBB6644 H. maraisii=mirabilis! SW Swellendam.Fig. 26a. MBB6862 H. maraisii=mirabilis. Rondeheuwel, Stormsvlei.Fig. 26b. MBB6862 H. maraisii=mirabilis. Rondeheuwel, Stormsvlei.Fig. 27. MBB6644 H. maraisii=mirabilis. SW Swellendam.Fig. 27a. MBB6663 H. magnifica var magnifica. SW Heidelberg.Fig. 27b. MBB6663 H. magnifica var magnifica. SW Heidelberg.Fig. 27c. MBB6663 H. magnifica var magnifica. SW Heidelberg.Fig. 28a. MBB6817 H. magnifica var atrofusca. Kweekkraal, W Riversdale.Fig. 28b. MBB6817 H. magnifica var atrofusca. Kweekkraal, W Riversdale.Fig. 29. MBB6537 H. mirabilis var triebneriana. Goudini, Caledon.Fig. 30. MBB6982 H. mutica. Hasiesdrift, N Bredasdorp.
Flower morphology and flowering time does not appear to offer any solutions. As it is, the flowers have so far been shown in Haworthia to be useful only in so far as recognising the sub-genera is concerned. It can be shown that even at that level, the distinction causes problems for botanists. Most of the species discussed above flower in late summer. While the epithet ‘floribunda‘ was chosen to suggest many-flowered, this is true for other populations considered to be H. magnifica.
Conclusion The circumscription of H. floribunda remains obscure and the situation is in fact exacerbated by new samples which indicate the labyrynthine relationships with several species previously excluded from debate. H. floribunda may have juvenile characters. It does have distinctive populations in a recognisable geographic area. It does not directly share habitat with any species although it does occur very near to H. turgida (north of Heidelberg, where hybrids are also recorded). Where the chloracantha-like equivalent occurs with H. pygmaea var. argenteo-maculosa at Cooper Siding, hybridisation is also evident. It occurs very near to H. magnifica var. atrofusca northwest of Riversdale, growing on a cool southern slope as opposed to the latter on a hotter north slope. The situation northwest of the Potberg, where as many as seven otherwise apparently discrete species need to be included in the discussion, will be very difficult to explain.
It should be noted in closing that the Heidelberg population MBB6663 (as KG107/74) was instrumental in the initial decision by Bayer in 1976 when H. maraisii and H. magnifica were treated as one under H. maraisii. This was repeated in 1982 when H. magnifica was given its chronological priority over H. maraisii. In Haworthia Revisited these two elements are separated as discrete species and the Heidelberg collection is cited under H. magnifica. This decision was made rather to accommodate the varieties which are attached to them. Thus the additional three collections noted in connection with it viz. MBB6860, MBB6861 and MBB6862 all fall within this same indecisive category.
M.B. Bayer, Cape Town, South Africa R.W. Kent, Poway, California
Introduction After writing Haworthia Revisited in 1996, I became aware of just how inadequate readers seem to be to the task of assimilating all the available literature on Haworthia, in the botanical and intellectual climate in which we live. It seems as though the more information we have the more confused we become. In order to generate the material needed to disprove or fortify my classification hypothesis, I have spent a further considerable amount of time in the field and in cultivating plants from seed. Unfortunately the editorial support and speed of publication has not kept pace with my own effort and much of my writing and my evidence is still in manuscript form. This short essay was therefore to put forward only a little more evidence to show just how complex plant species are – not necessarily only in Haworthia.
In my first Handbook (1976), I anticipated H. maraisii and H. magnifica to be separate species – the former west of, and the latter east of Heidelberg, Cape. In the second book (1982), I felt that I was dealing with a single species and referred to a population just west of Heidelberg as H. magnifica var. magnifica (instead of as H. maraisii). I did this transposition quite deliberately in order to suggest that the distinction between the two species was very arbitrary. In my later revision (1999) I separated H. magnifica and H. maraisii again. The rationalisation is given there and I just need to explain that I thought this was a better way of communicating the nature of the variable populations attached to each of two elements. It was then evident that populations relevant to magnifica and maraisii were proving to be more radically different than a single species hypothesis could comfortably accommodate. Also new evidence was accumulating for the nature of:
1. H. heidelbergensis, which was not even included in the previous debates. 2. H. floribunda, also proving to be more variable than predicted. 3. H. mirabilis.
The overall relationship of these three species is far closer to both H. magnifica and H. maraisii than even my original conservative approach suggests.
In the essay which deals with H. floribunda I make two points particularly relevant to this article:
1. The geographic facts, viz..”Westwards (from Gouritz River), the Swellendam area seems to provide a vegetation interval which H. turgida seems to bridge with difficulty, and which also marks some kind of a break for the H. magnifica/H. maraisii complex. This divide may be a function of inadequate collecting or the lack of adequate habitat”.
2. The populations known to me. To quote from the floribunda manuscript “There are three more collections to be considered:- by Denis DeKok near Swellendam (Figs.25a, b & c MBB6644), about 10km west of Swellendam (MBB6861 not illustrated), and from the farm Rondeheuwel south of Stormsvlei (Figs.26a, b & c MBB6882, (Bayer in KG326/71). This latter collection has previously been reported (Haworthia Handbook 1976) under one of the populations intermediate with H. mirabilis, and is cited in Haworthia Revisited under H. maraisii. These populations, as well as that southwest of Heidelberg (Figs27a, b & c MBB6663, Bayer in KG107/74 – cited under H. magnifica, Bayer, 1999) confound the issue enormously. Thus we actually have four populations which cannot with confidence be allied with either H. magnifica, H. maraisii or with H. mirabilis. The reality is that neither H. heidelbergensis nor H. floribunda can be excluded from the consideration of these populations. The case for each “species” needs to be dealt with separately.”
Primarily illustrated here are plants from the population MBB6644 which occurs west of Swellendam. The population is not remarkable for the variation observable there among the individual plants, and I have taken it to demonstrate that there was, and is, predictive value in my classification hypotheses. Much of my recent writing has been directed at the very weak concepts and perceptions that appear in our general understanding of “species” and their variability. This has an impact on individual credibility, plausibility and the truth with which we examine our human condition and pursue our hobbies.
Results The photographs depicted are all taken at virtually the same distance so that the size of the plants is relatively correct; with the largest plants being near 90mm diam. (The pots are 90mm square.)
The first illustration depicts a conventional form of H. maraisii from a population from north of McGregor Fig. 1 MBB6646 H. maraisii var. maraisii). The species was first described from Stormsvlei where the plants are a little larger and more tuberculate than these pictured. Actually it has become difficult to say just where this species stops and starts, and this will become obvious. There is a degree of translucence in the leaves and there is a conspicuous vein down the centre of the leaf. This is by no means a ‘character’ for the species, and an article could be written about the variation in venation as well as about this single population (MBB6646) too.
The Swellendam plants have been named as H. maraisii=mirabilis in the preceding chapter and are illustrated as follows:
Fig.2, 3 and 4 are the clones 3, 8 and 17 of MBB6644. The surfaces are a little less tuberculate than the McGregor plant shown, but the only significant difference is actually that they are slightly bigger plants, the spination is more obvious and the leaves tend to be fewer and more erect. These are also by no means diagnostic characters. There is little translucence and the venation is inconspicuous. The identification of the population as H. maraisii thus seems obvious.
However, clones 2 and 4 illustrate a narrowing of the leaf (Figs.5 & 6 MBB6644). Clone 15 is a small plant also with smaller narrower leaves (Fig. 7 MBB6644). Were its leaves more erect instead of so recurved, it could be perhaps be taken to be H. heidelbergensis var. scabra.
Clones 1 and 3 (Figs.8 & 9) of MBB6663 are from west of Heidelberg and would easily be lost among the Swellendam plants if unlabeled. In the preceding chapter the population is identified as H. magnifica var magnifica. Clone 2 of this population (Fig.10 MBB6663) is a plant with more erect leaves – but …
Clones 7, 10, 11, 14 and 16 of MBB6644 (Figs.11 to 15) have a similar narrowing and erect bearing of the leaves, not to say that the plants are otherwise identical. This condition of the more erect leaves seems to be the more general one in both populations, MBB6644 and MBB6663. This is almost the crux of the problem in the classification of Haworthia, and in my experience it is not a problem unique to Haworthia. In all of biology there is this variability that requires statistical method to establish what “average” is. When one is dealing with different growing condition, and the nuances of texture, colour, shape, leaf recurvature and number, it is in fact not possible to generate such a “mean”.
Clones 9 and 13 of MBB6644 (Figs.16 & 17) are unusual in either H. maraisii or H. magnifica. The leaves tend to narrow quite considerably, but the leaves can also be unusually narrow and erect in populations of H. maraisii as in a population west of Robertson (Fig.18 MBB6647.8).
I include an illustration of a plant from a population of H. floribunda var. dentata from south of Swellendam (Fig. 19 MBB6881), – a population that tends to link floribunda with either maraisii or heidelbergensis to expose the reality that the link extends to H. mirabilis. (Apparently in a Dutch journal a writer has suggested that H. floribunda var. dentata is a form between H. floribunda and H. chlorocantha var. subglauca! This would be a complete misrepresentation of my classification hypothesis which has been fully explained and there is no reason for such a mistake. The var. dentata is represented by many populations and Chapter 3 details the possible relationship of the species floribunda and chlorocantha without any frivolity about the varieties and forms.)
Fig. 19. MBB6881.6 H. floribunda var dentata. Oudekraal, Heidelberg.
Also illustrated is a plant from a population very close (NW Kweekkraal), both geographically and in appearance, to H. magnifica var. atrofusca west of Riversdale (Fig.20 MBB6817). I have included it here because general colouration and texture are the same and it evidences the typical round-tipped leaves of floribunda which occurs so frequently in maraisii and magnifica (see older leaves in Fig.2 or Fig.8).
Fig. 20. MBB6817.1 H. magnifica var atrofusca. NW Kweekkraal, Riversdale.
Clones 1, 5 and 18 of MBB6644 (Figs.21, 22 & 23) are small rather nondescript specimens, while is clone 64 of MBB6639 (Fig.24) is a plant of H. mirabilis var. sublineata from south of Bredasdorp. These last four illustrations depict a series of smaller plants which bear a resemblance to H. heidelbergensis. The difference in the latter is the lighter green colouration, slightly more translucence and more conspicuous spination. These are significant in the context of these collections and these photographs. However, put into the context of all the populations of H. heidelbergensis, H. floribunda, H. mirabilis, H. maraisii, H. magnifica, also H. emelyae and the total variability among these, it is impossible to stipulate difference.
The four problem populations, MBB6644, MBB6861, MBB6882 and MBB6663 are not as distinctive as suggested by the way in which I have specified them. They have also to be seen in the light of the variability of the individuals in those populations, and the many other populations which abut geographically onto them. Thus where I suggest they confound a clear difference between the two species H. maraisii and H. magnifica, this is not only how it is to be understood. The essay concerning H. floribunda will illustrate how extensive the problem actually is. These four populations can be discussed in terms of each of the species named in the previous paragraph.
One has to come back to biological variation and consider how one is to circumscribe a species in such a way as to facilitate identification. Here we have a single population in which the basic features such as colour, size, texture, spination, venation and translucence, are so variable that it is impossible to establish what the mean or average plant is. The problem is compounded by the fact that the variables can also not be broken down to discrete quantities. I have shown only the whole plant and thus the vegetative features. The flowers are less variable both within and between populations and ‘species’. It is already clear from the literature that species in the sub-genera of Haworthia can barely be separated on the basis of their flowers. To expect that the flower can then provide a ‘signature’ or character by which population variability can be understood, is fallacious. Floral character in fact must persuade one that there are less species than we wish to find from the vegetative characters which attract us to the plants in the first place. Flowering time is useful but it is also not diagnostic. Generally flowering time in the species mentioned in this article lies between September and May. It is no more useful than the geographic location of the population. H. maraisii tends to flower from March to May and H. mirabilis is generally earlier. However, to establish the actual flowering times for each population in terms of different seasons and years, and as a mean for the individuals in each population, must be a herculean task.
What I have done is to take geographic distribution as a key element in the classification. My experience is that distribution reflects the relationship between populations and between species. This is just a logical extension of the processes we should expect from speciation processes associated with genetic interbreeding, and isolation which obviates it. “Genetic drift” should reflect the probabilities associated with pollination across distance and with the nature of the pollinating agent. Similarities between populations should reflect this, as well as the probabilities of distribution by whatever agent such as wind or water. Naturally these will not be absolute, and the classification hypothesis will only be as good as the information allows, with the proviso that it is more information we are going to require if a better hypothesis is to be formulated.
What is striking in all my formal and informal communication with interested parties is the failure to appreciate what judgmental factors may underlie the vexing thing we regard as a “species”. I need to make it quite clear that I take a species to be a system of individuals judged on a very wide variety of factors – including geographical relationships and the influence these seem to have on other similar systems belonging to other genera.
In Haworthia, I do not deceive myself that these systems are as discrete as my classification may suggest. That is why I wrote that I was recognising nodes in a complex interconnected array. Someone put forward their own solution to one such array by suggesting that the intermediate element be recognised as a species. This is particularly unacceptable when one considers the full implication. All the intermediates could be recognised as species and the present species then as intermediates! The position is that my classification was undertaken as a proper revision when a considerable amount of new data was available; sufficient to consider it really representative and adequate on which to base an overview of the genus. For persons now to suggest an alternative classification without a proper and thorough insight into all that old material, and without adding adequately to the known record, will border on the irresponsible and inconsiderate of the purpose of classification.
[1] Breckenridge (Haworthiad 19:4, 2005) for an example among others, admits to being wholly lost in respect of a remark I made (Haworthiad 18:44, 2004) in respect of varietal names. I wrote “… (variety) is a useful rank only if there is no typical variety, and if it is recognized that the species name is used for all the variants both formally named AND unnamed.” I was referring to the practice in nomenclature where the description of an inferior rank (variety in this case) automatically creates a “typical” rank. It is obvious that if a species is described from limited material, it is improbable that the description will include all the variants. Thus any new material of that species may differ from the described. Formally giving this new material a name should in fact broaden the initial description. If the ramifications of this fact are not apparent to the reader, then no amount of writing and explanation will lift the fog of misunderstanding. ♦
This appeared as an article in ALOE 38:31 (2001). Unfortunately there was a problem with illustrations and captions and these are corrected here. A comment is also added as an addendum to respond to criticism by I. Breuer published in Alsterworthia 1:13(2002).
Introduction:
I described Haworthia serrata in 1973 (Jl S.AFr.Bot.39:249, see Figs.1) from Oudekraal, southwest of Heidelberg. I commented then on the wisdom of describing a new species when “the recognition, estimation of taxonomic rank and circumscription of elements in Haworthia…” was so problematic. The new species was said to resemble H. emelyae var. multifolia (Figs.2). In respect of its distribution, I said it was closest to H. heidelbergensis at Heidelberg (Figs.3 JDV87/1) and as at Matjestoon (Fig.4 JDV87/3), and also to H. sublimpidula at Swellendam (now known to be H. floribunda var. major (Fig.5 MBB6859, taxonomically with little connection to H. rossouwii). The implication was that it could have been taxonomically related to those elements in terms of geographic distribution. I was still puzzled by the relationships of H. serrata when I wrote (New Haworthia Handbook :55, 1982) that collections by C. Burgers from the coastal limestones might throw more light on the matter (Fig.6 MBB6985 H. mirabilis var. calcarea).
Fig. 01a. JDV87-47 H. rossouwii. Oudekraal, SW Heidelberg.rossouwii 1 043Fig. 01b. JDV87-47 H. rossouwii. Oudekraal, SW Heidelberg.Fig. 01c JDV87-47 H. rossouwii, Oudekraal, Heidelberg.Fig. 01d. JDV88-53 H. rossouwii. Oudekraal, Heidelberg.Fig. 02a. JDV87-162 H. emelyae var multifolia. Springfontein, Karoo.Fig. 02b JDV87-162 H. emelyae var multifolia. Springfontein, Karoo.Fig. 03a. JDV87-1 H. heidelbergensis. E Heidelberg.Fig. 03b. JDV87-1 H. heidelbergensis. E Heidelberg.Fig. 03c. MBBsn H. heidelbergensis. E Heidelberg.Fig. 04. JDV87-3 H. heidelbergensis var toonensis. Matjestoon, SW Heidelberg.Fig. 05. MBB6859 H. floribunda var major. S. Swellendam.
In his revision of Haworthia (1983), C.L. Scott recognised my species. He confused it, and illustrated it, with a photograph of H. mucronata var. mucronata sensu Bayer (1999), which he called H. mclarenii. This does exemplify the problem of look-alikes in the subgenus, and this curiously does have some connection to the actual problem.
When I wrote Haworthia Revisited (1999), I was still not sure what the real affinities of H. serrata were. I mentioned that my 1982 observations were vague because of the unusual elements which occurred in the coastal limestones. I described these as new: H. mirabilis var. calcarea from De Hoop Nature Reserve (see Fig.6), H. variegata var. petrophila (CBurgers2158) from Karsriver and H. heidelbergensis var. minor from N Bredasdorp (Figs.7 JDV86/5). In my 1999 classification, I placed H. magnifica var. paradoxa (Figs.8 JDV86/78 & 94/89) in H. mirabilis, suggesting that it may have a connection there via the var. calcarea and an element from the Potberg (CBurgers 2018) which I have not yet seen in the field myself (since I wrote the article I have in fact seen this population, MBB7356, and it is discussed in the Chapter re H. mirabilis). The connections of paradoxa must be viewed in its relationship to H. emelyae var. major, in the same way that serrata is compared with H. emelyae var. multifolia. I also suggested that H. serrata had a very close resemblance to some forms of H. mirabilis var. sublineata from south of Bredasdorp. This present paper addresses some of these observations again.
Fig. 17b. Fig. 06a. MBB6985 H. rossouwii var calcarea. DeHoop.Fig. 06b. MBB6985 H. rossouwii var calcarea. DeHoop.Fig. 07a. JDV86-5 H. heidelbergenis var minor. Rooivlei, Bredasdorp.Fig. 07b. JDV86-5 H. heidelbergensis. Rooivlei, N Bredasdorp.Fig. 07c. JDV86-5 H. heidelbergensis var minor. Rooivlei, Bredasdorp.Fig. 08a. JDV86-78 H. mirabilis var paradoxa. Vermaaklikheid.Fig. 08b. JDV86-78 H. mirabilis var paradoxa. Vermaaklikheid.Fig. 08c. JDV98-89 H. mirabilis var paradoxa.
New Records In June 1999, I revisited H. serrata at Oudekraalkop (MBB166) again, this time together with J.D.Venter. Despite rumours of over-collection, it was still very abundant, although localised. It was also still present at Koppies (Fig.9 JDV99/6) a short distance to the west, but in very low numbers at a badly burnt site. P.V. Bruyns (priv. comm.) has reported it at a more northerly site as well. This could be where J.N.O. Uys also showed it to me at a second site on Oudekraal when he first introduced me to this species. It struck me again how similar the plants were to H. mirabilis (particularly the var. sublineata, Figs.10 MBB6639, bearing in mind the vast variation that exists there) and to H. heidelbergensis var. minor. The visit in June ’99 was a general excursion to explore the reported presence of H. maraisii at Koppies (ex E. Esterhuizen), in relation to that species and heidelbergensis as I know it in that general area. Our finds and observations led us later in the year to explore the area north of the Potberg. The extraordinary finds there led me to further explore westward to Bredasdorp itself and to see for myself the var. calcarea in the De Hoop. Two visits for that purpose were fruitless, but thanks to Dennis DeKok, I did locate it in the company of Lawrence Loucka in July 2000 (MBB6985-fig.11). This latter excursion was particularly fruitful because, by great good fortune, we discovered H. serrata in unmistakable form north of Bredasdorp (Soutkloof, Fig.12a & b MBB6983), and Nooitgedacht, MBB6984). Both sites are west of Rooivlei where heidelbergensis var. minor occurs. We also collected variegata var. petrophila (Fig. 13 MBB6986) west of the Karsriver limeworks. For that variety I have C. Burger’s record (CB2158) from east of the limeworks, and a collection I made at Renosterfontein (Fig.14 MBB6632) on the east bank of the Karsriver in 1997. This latter collection, in particular, reminded me in both form and colour of H. emelyae var. multifolia (see fig.2).
Fig. 10a. MBB6639 H. mirabilis var sublineata. S Bredasdorp.Fig. 10b. MBB6639 H. mirabilis var sublineata. S Bredasdorp.Fig. 10c. MBB6639 H. mirabilis var sublineata. S Bredasdorp.Fig. 10d. MBB6639 H. mirabilis var sublineata. S Bredasdorp.Fig. 10e.Fig. 10f.Fig. 10g.Fig. 11a. MBB6985 H. rossouwii var calcarea. DeHoopFig. 11b.Fig. 12a. MBB6983 H. rossouwii. Soutkloof. Fig. 12b.Fig. 13a. MBB6986 H. petrophila KarsrivierFig. 13b.Fig. 14. MBB6632 H. rossouwii var petrophila. Renosterfontein, Karsriver.
The conclusion from seeing H. serrata north of Bredasdorp, and both var. calcarea and var. petrophila, is that this is the prime connection. It seems now more improbable that the var. petrophila is as close to H. variegata as I had supposed, and that the var. calcarea is closely connected to both H. serrata and to var. petrophila. The other observations remain valid viz. that there is a strong resemblance between serrata and mirabilis var. sublineata; but particularly that serrata may be connected to heidelbergensis via the var. minor. However, heidelbergensis in its general relationships is extremely complex. It is interfused with H. maraisii, it may be the direct (if complex) geographic extension of H. mirabilis in the context of the relationship between that species and maraisii, and it may have affinities and continuity with both H. variegata and H. floribunda. There is still another problem: H. variegata var. modesta at Kathoek (Figs.15 JDV97/24) and Potberg (Figs.16 MBB6542). My collection of this element at Kathoek (Figs.17 MBB2551) suggests that it is possibly a small very proliferous sand dwelling serrata. J.D. Venter’s collection from the same farm is a more robust element with rather longer, more erect leaves which suggests H. variegata. My Potberg collection (fig.16) is another small element which I relate to variegata although the leaves are shorter and more turgid. It must be borne in mind that H. variegata var. hemicrypta occurs north of the Potberg on the gravelly flat so that there is ecotypic variation in evidence. Further work will have to be done to elucidate what could be an unresolvable situation i.e. the Potberg is home to variants which link all the elements so far named. The evidence for this is already available, but insufficient to crystallise a solution.
Fig. 15a. JDV97-24 H. vaiegata var modesta. Kathoek.Fig. 15b.Fig. 16a. MBB6542 H. variegata var modesta. Potberg.Fig. 16b.Fig. 16c.Fig. 17a. MBB2551 H. variegata var modesta. Kathoek.Fig. 17b.
There will be difficulty in considering the extent and the degree of similarity of H. serrata to other populations. Some of these which include serrata-look-alikes are:
Fig. 18. JDV96-52 H. heidelbergensis var . N Kathoek.Fig. 19. JDV93-51 H. heidelbergensis. Melkboom.Fig. 20a. JDV87-4 H. heidelbergensis var scabra, Leeurivier, Drew.Fig. 20b.Fig. 20c.Fig. 20d.Fig. 21. JDV97-49 H. mirabilis var consanguinea. Dwarswaterkloof, MacGregor.Fig. 22. JDV88-83 H. mirabilis var consanguinea. Dwarswaterkloof, Macgregor.Fig. 23a. JDV96-38 H. maraiss var notabilis. Agtervink, Robertson.Fig. 23b.Fig. 23c.Fig. 23d.Fig. 24a. JDV98-1 H. maraisii var notabilis, Wolfkloof, Robertson.Fig. 24b.Fig. 25. JDV97-30 H. cf. heidelbergensis. N Ashton.Fig. 26. JDV86-1 H. mirabilis var triebneriana. Stormsvlei.
Some of these populations and their variants are dealt with in other chapters (Particular note must be made of H. elizeae which has to be considered in relation to H. rossouwii).
H. serrata was coming into bud in July a good five months after mirabilis had started to flower. This shows that my predictive thoughts about the relationship with calcarea and petrophila may not be particularly sound if flowering time is considered. Calcarea flowered a little before mirabilis var. sublineata in January, petrophila flowered in February while Roovlei heidelbergensis and Leeurivier (Drew) heidelbergensis flowered in December and March respectively. Flowering time for other collections needs to be observed, noting that this is not necessarily definitive either (for example while H. mirabilis var. badia is essentially late-summer flowering, single plants may flower in winter.
When I consider how much time I have spent in the area, it was really surprising to find serrata so far west. It means that something very interesting can still be found in the large triangle Swellendam/Bredasdorp/Riviersonderend, and that exploration will have to be done on a very small scale. The Karsriver and Renosterfontein petrophila are just smaller more spinose versions of Oudekraal, and the leaves are more spotted on the reverse. Calcarea was previously known from two clones which had rather shorter leaves and, perhaps, fewer leaves per rosette. This, the locality, and the spotting on the back of the leaves would have been the reason for opting for a relationship with mirabilis. We only found it with quite detailed directions because it was such a small population and so cryptic, consisting of three tiny locations under bush with a total of about 15 clumps. One location could have been the product of vegetative propagation with six to seven clumps. The second was similar with only two clumps and evidence of porcupine activity around the clumps. The third covered about 15sqm and there were another 15 or so plants as clumps or as single rosettes. These did vary and only one clump really resembles the two I have seen in cultivation viz. the Karoo Garden plant of C. Burgers, and a plant I received from Dennis DeKok. Burgers, however, appears to have made two collections from De Hoop; this needs to be confirmed. The plants seen by me in habitat do vary quite a bit and the range of variation equates that of both serrata ex Oudekraal (the Koppies population is reduced to about five plants) and north of Bredasdorp. The mean length of the leaves and leaf number per rosette may be shifted to the shorter and fewer, respectively.
The two Bredasdorp populations of H. serrata are remarkable for the similarity of the plants to serrata at the type locality. The habitat is a little different and at the westernmost of the populations, the plants cluster much more freely than the Heidelberg populations. At Koppies the plants are on a Witteberg substrate, whereas at Oudekraal and N Bredasdorp, it is Bokkeveld shale. We may have a slightly biased image of serrata with turned out leaves; the plants can be quite globose, and I did illustrate a plant like that in my original description when I drew the comparison with herbacea.
Where serrata really differs from mirabilis is by the greater number of leaves per rosette, the thinness of the leaves, narrowness at their bases, lack of a mid‑leaf swelling and by the incurving of the leaf tip; that character is fairly significant. H. mirabilis var. sublineata grown recently from seed has proved to be remarkably variable, but none of the plants have incurving leaves. Some of the clones are very similar to serrata, but the flowering time is earlier in the year and more in keeping with H. mirabilis in that respect.
Discussion H. variegata var. petrophila and H. mirabilis var. calcarea should be transferred to the species H. serrata. The same may be true for H. heidelbergensis var. minor. However, H. heidelbergensis has a complex distribution and there are other complexities in variation that preclude a definitive statement. It is quite probable in the light of the findings discussed in Haworthia Update Vol.1, that there is no discontinuity which satisfactorily and convincingly can be claimed to distinguish species. This is probably just another example of a more general problem which I have experienced with the identification and classification of many plant genera.
I am conscious that I have not dealt fully with the problem in omitting mention of co-occurrence (I mean sympatry – when two obviously different elements grow together at the same site or very closely adjacent). I see this as the best measure of discontinuity. Thus if two elements grow in close proximity, even if not sharing the same habitat, they are probably discrete. This does not preclude the possibility that they are simply ecotypes and this is taken into consideration. However, I have before pointed out that a taxonomic solution which can be deduced in this way for populations might not be true for the entire distribution of the species concerned. It is evident that serrata generally occurs alone. At Oudekraal and at Koppies, there is a floribunda/maraisii element in the vicinity. In the case of heidelbergensis var minor, both turgida and maraisii are present. I have always been struck by the similarity of mirabilis and serrata, and by the fact that they do not co-occur. Mirabilis, in the context of this paper, is presently not known east of Napier except at Bredasdorp and Mierkraal south of Bredasdorp. This is as the var. sublineata and var. mirabilis respectively, which differ from serrata in respect of flowering time, less leaves per rosette, and the non-incurving leaves. Curiously, it appears that mutica, at least in the Bredasdorp area, grows generally alone too, although at Haarwegskloof (Bredasdorp) it does grow with heidelbergensis. At Drew it is in the close company of both maraisii and heidelbergensis, and south of Stormsvlei is near to a maraisii/mirabilis intermediate. Thus even an element as apparently different as H. mutica may presumed to be, cannot be ignored.
However, another consideration now enters the picture with the discovery of serrata so far west. There are three poorly understood elements which were reported to have come from Bredasdorp and Napier respectively. The first of these was H. bijliana von Poelln. that does, however, seem to belong rather with H. arachnoidea than with anything else. G.G. Smith appears to have had a plant of this ilk which he received from Maj. Venter 9, from Caledon. The other two, however, are H. altilinea var. bicarinata Triebner and H. rossouwii v.Poelln. They were both described from the same collection viz. Napier at Bredasdorp, Rossouw in Triebner 1059. The former was never illustrated and the description is rather brief … “Backs of leaves always with two keels and with teeth. Leaves 3-5cm long, 6-8mm broad, with 4-6 dark lines in lighter part, with two keels at back, keels with up to 2mm long denticula”. This description is in the joint paper by Triebner and von Poellnitz, but authorship is attributed to Triebner alone. von Poellnitz later retained this variety when replacing the name altilinea with mucronata (note that in the context of the names which I use, Scott confused my serrata with mucronata as well). This is very curious because it is evident from the description of H. rossouwii, which is very much more detailed, that another plant from the same collection is in fact again described. The differences are essentially that von Poellnitz gives the leaves as 5-6cm long, and 8-10mm wide as opposed to the respective 3-5cm and 6-8mm by Triebner; and with three lines on the ‘end surface’ as opposed to four to six by Triebner. This says a great deal about the detailed descriptions which are routinely called for, and which in Haworthia seldom amount to much because of the problem of look-alikes, and the variability within and between populations.
I repeat the von Poellnitz description here to illustrate that it is a very good description that nevertheless omits some key information:
H. rossouwii von Poellnitz spec.nov. (Stemless rosette, spirally many-leaved. Leaves rather pale green, nearly oblong lanceolate, acuminate, on the face somewhat retused near the apex thus forming a triangular face; on the back, towards the apex, with sub-pellucid tubercle-like spots, denticulate on the margins and keels; the triangular face strongly acuminate, sub-translucent, not tubercled, but carrying some sub-pellucid small teeth, and traversed by a few longitudinal lines).
Stemless many-leaved rosette about 5-6cm in diameter. Leaves more or less erect, 5-6cm long, 8-10mm wide, about oblong-lanceolate, tapering gradually, pointed, slightly widened at the base, almost straight or a little bent to one side near the tip, rather light green, dull. On the face flat or usually somewhat convex, near the tip slightly bent backwards (retused) at an angle of 20 degrees; on the back, from the middle outward or near the tip obliquely keeled, or somewhat more rarely with two keels, near the tip with a few, often elongated, very often raised slightly pellucid spots, and frequently with a few leaf-coloured tubercles. The end-surface half-pellucid, 20mm long or somewhat longer, 7-9mm wide at the base, triangular, tapering very gradually, pointed at the tip often slightly curved inward, somewhat convex, smooth, rarely with a few indications of tubercles, but with some semi-translucent teeth (these teeth loosely arranged in a lengthwise middle line, now and then also in another line nearer the margin and very rarely scattered about in small numbers) with three long, occasionally interrupted, green lengthwise lines, of which only the middle one may reach almost to the apex, and usually with a few very short line; leaf margins and keel with pale teeth, or near the base often leaf-coloured, teeth, up to barely 2mm long; end-bristle pale, with small side bristles, 5-7mm long.
… On account of the teeth on the end surface stands near to H. schuldtiana and to H. paradoxa (I treat these respectively as H. maraisii and H. mirabilis var. paradoxa) but is easily distinguished from both. Its flowers are thus far unknown”.
The omissions are the flower of course, and there is no stipulation of the approximate number of leaves, and leaf thickness. The weakness is that it is a single plant description, evidenced by the fact that another plant from the same collection is accepted as a variant of a totally different complex.
In the past I have I regarded rossouwii speculatively as a synonym of H. mirabilis, due largely to the fact that it was recorded from the Bredasdorp area (and to the ambiguity of the type). Another reason was the similarity I observed between the illustration of H. rossouwii in Kakteenkunde with H. heidelbergensis var. minor and with H. mirabilis var sublineata. I opted for the latter (with which Scott was in accord) and the similarities are also recorded in the discussion above. I dismissed bicarinata as insufficiently known because I assumed that Triebner may have had some better reason to have equated his specimen with the Little Karoo H. mucronata. Possibly there has been a mistake in the Triebner numbers. It is this discovery of serrata so close to Bredasdorp, and equidistant from Napier, which revives and also casts new light on the problem. I conclude that my H. serrata is a later synonym of H. rossouwii of von Poellnitz.
There are still two other considerations. In my 1999 revision, the fact is that I placed paradoxa as a variety of mirabilis while discussing the possible connection of these elements via calcarea. In the light of this paper it is evident to me that paradoxa could also be linked to serrata rather than with mirabilis. The same is true for emelyae var. multifolia. I would not like to commit myself to this probable solution at this stage, as I think more evidence should be obtained. It is also improbable that, however attractive, the solution will not avoid impacting on H. emelyae, H. magnifica and H. heidelbergensis. It is also evident that the requirements of the ICBN are that should paradoxa indeed need to be incorporated into serrata, more name changes will result, as paradoxa will have priority over rossouwii. My opinion is that this is a fundamental weakness in the code, which inhibits speculative treatments, and also which results in smaller taxa becoming the typical main element of a species. My further opinion is that it is a weakness of the Code that the description of an element automatically creates an equivalent taxon. This may be acceptable and desirable where the elements are clearly separable and geographically defined, but not where the typical element is required to absorb any other variants which are not so defined. Thus H. paradoxa var. paradoxa could become the main component of a species where it is actually a geographical isolate. In a system which does not have an adequate general definition for ‘species’, this probably does not matter; whereas it might if there is a definition which suggests that the ‘species’ is an operating living system (see Bayer in Asklepios 77:3, 1999).
The formal statement of this paper, which would better be regarded as an hypothesis subject to test, is thus as follows:
H. rossouwiivon Poelln. in Kakteenk.7:75 (1938). Type: Napier near Bredasdorp, collected by Mr Rossouw, South Africa Police in Napier, Triebner 1059. Not preserved. Lectotype: designated by Breuer & Metzing 1998, icon in Kakteenk. non syn. H. mirabilis sensu Scott 1985, Bayer 1976, 1982, 1999.
H. altilinea var. bicarinata Triebn. in Triebner & von Poellnitz in Feddes Repert.Spec.Nov. 45:170 (1938). Type: Napier at Bredasdorp leg. Mr Rossouw (=Triebn.1059). Not preserved.
H. serrata Bayer, Jl S.Afr.Bot. 39:249(1973). Bayer :55(1976). Bayer :147(1999). non Scott :62(1985). Type: CAPE‑3420(Bredasdorp): Oudekraalkop, Heidelberg (‑BA), Bayer 166 (NBG).
H. rossouwii var. calcarea (Bayer) comb.nov. H. mirabilis var. calcarea Bayer in Haworthia Revisited :110(1999). Type: CAPE-3420(Bredasdorp): De Hoop (-AD), C. Burgers 1648 (NBG, Holo.).
Material seen:- 3420(Bredasdorp): NNE. Buffelsfontein, Potberg (-BC), Burgers 2018 (NBG). De Hoop (-AD), C. Burgers 1648 (NBG), Bayer 6985.
H. rossouwii var. petrophila (Bayer) comb.nov. H. variegata var. petrophila Bayer in Haworthia Revisited:159(1999). Type: CAPE-3420(Bredasdorp): Renosterfontein (-AC), Burgers 2158 (NBG, Holo.).
The provisions of this solution are that the epithet paradoxa may be instated preferentially, and that emelyae var. multifolia and variegata var. modesta also be included.
A second provision is that the species H. bijliana v.Poelln. be rejected as insufficiently described and sourced, and that the type (the illustration in the Berlin herbarium) is ambiguous – as indeed are many Haworthia types and descriptions based on single plants and not correctly tied to geographical origin. This is demonstrated by the citation by VPoelln. (Feddes Rep. 1936) of this species viz. bijliana from Klawer and Eenriet and also Feddes Rep. in 1937 from Springbok – which cannot be anything other than H. arachnoidea. In this last publication he cites a specimen from Bonnievale, another from Ezeljacht Oudtshoorn and also sinks fergusoniae here which in fact was supposed to have come from ‘near’ Grahamstown.
H. bijliana Poelln. in Feddes Repert. 27:134 (1929); Bredasdorp, Mrs va der Bijl. Type van derBijl s.n. in (B). Not preserved. Lectotype desig. Breuer 1999, unpublished photo. icon. (B). Species rejecienda.
Acknowledgement I am grateful for the company on the field trip of Lawrence Loucka of Connecticut (USA), and that of my wife, Daphne. The assistance of Paul Botma of Cape Nature Conservation is acknowledged over and above the general acknowledgement to that Department. Mr. C.L. Neethling, Mr. Thys Swart and Mr. Pieter deKok were most kind and helpful in respect of access to sites. I am also grateful to Kobus Venter for support, advice, and constructive exchange of ideas. Not least to Steven Hammer, with comment from Bob Kent, who was so kind as to read this manuscript critically and make corrections. Residual errors will be mine.
Addendum Alsterworthia Vol 1:13(2002) “A comment on Bayer’s latest taxonomic contribution” by Ingo Breuer. [link] With reference to the comments in Alsterworthia 1:13(2002) regarding the identity of H. rossouwii and H. serrata:
Notes
Fig. 27 H. rossouwii. Vpoelln. type.Fig. 28a. MBB6984.1 H. rossouwii. Black and white profiles from two angles, of the same one plant actively growing, undessicated and turgid. The plant is a clone from Bayer 6984 Nooitgedacht, NW Bredasdorp. This is correctly H. rossouwii, and correctly H. serrata.Fig. 28b.Fig. 29a. MBB6984.1 H. rossouwii. Colour photo originals of the preceding. Fig. 29b. Fig. 30. MBB6984.2 H. rossouwii. Nooitgedacht, Bredasdorp. A second plant of MBB6984. This is a more vigorous grower and it shows the differences that obviously confound the uninitiated. There were not many plants at Nooitgedacht. I only collected from two clones and this is a photograph of the second. Fig. 31. MBB6983 H. rossouwii. Soutkloof, N Bredasdorp. Showing a plant in which the convexity of the mid-upper surface (and concavity of the lower surface) is virtually absent. I could have, from this sample (MBB6983), produced pictures to match Figs.27 to 29. So no one should claim that Fig.30 is a different species as someone now well might. Fig. 33 Flowers of MBB6672 H. maraisii Koningsrivier, SW Robertson.
Fig. 27 A copy of the von Poellnitz lectotype of H. rossouwii.
Figs 28a & b Black and white profiles from two angles, of the same one plant actively growing, undessicated and turgid. The plant is a clone from Bayer 6984 Nooitgedacht, NW Bredasdorp. This is correctly H. rossouwii, and correctly H. serrata.
Figs 29a
& b. The colour originals of the preceding.
Fig.30 A second plant of MBB6984. This is a more vigorous grower and it shows the differences that obviously confound the uninitiated. There were not many plants at Nooitgedacht. I only collected from two clones and this is a photograph of the second.
Fig.31 A photograph of a plant of MBB6983 Soutkloof, north of Bredasdorp. Showing a plant in which the convexity of the mid-upper surface (and concavity of the lower surface) is virtually absent. I could have, from this sample (MBB6983), produced pictures to match Figs.27 to 29. So no one should claim that Fig.30 is a different species as someone now well might.
Figs 32a
& b. Scanned and manipulated copies of figs 28a & b.
Fig.33
Flowers of MBB6672 H. maraisii Koningsrivier, SW Robertson.
Figs.7a, b & c. These are clones of JDV86/5 H. heidelbergensis var. minor, Rooivlei, NNE Bredasdorp. This is a small population, of also small plants, from northeast of the collection MBB6984, Nooitgedacht. The plants partly resemble H. rossouwii from there, and partly the var. petrophila from Karsriver. Classifying them as a variety of H. heidelbergensis is not necessarily correct. Considering habitat and substrate, one must consider that ecotypification is playing a role, and that was habitat available, there would be further populations which would confirm continuity with these known collections.
Regarding Figs.28a & b. A bit of play with light and focus and the keel and under-leaf spotting could be varied. I would like to suggest that they be considered them faded, less sharply focused, printed with less contrast and on a coarser grained paper. Figs.32a & 32b are in fact such scanned copies of Figs.29a & b. I do not think it even necessary to consider that the original rossouwii of Fig.1 endured a long journey in the post to a foreign land. The assertion by Breuer in Alsterworthia is that not only are the plants I figured as H. rossouwii different from Fig.1, but that this Fig.1 lectotype is “probably” of the maraisii-complex. This is at variance with…
1. the Triebner description of the same collection as a variety of H. mucronata;
2. with Col. Scott who regarded it as H. mirabilis. What to say of von Poellnitz’ own assessment – the man who described both species i.e. maraisii and rossouwii in the first instance.
3. with the facts of the pictures before you.
What should an observer make of maraisii growing very near to rossouwii at Oudekraal and near to rossouwii at Napier? Can we really see a closer resemblance of Figs 27 & 28 with those long attenuate, double-curved, thin leaves – to H. maraisii rather than to H. serrata. I truly do not see that such a judgement can be seriously considered in the light of Breuer’s interpretations. It is of course valid when one considers all the variants that I illustrated in my article above and in terms of my over all classification.
Generally these plants of the SW Cape tend to have the double curvature of the leaves as is evident in the VonPoellnitz lectotype, but one can find in the same population and/or taxon, clones with incurved, outcurved or erect leaves (see fig.30). Broadly speaking H. maraisii has thick recurved leaves. But this is a non-argument even if one (outside of a “lumping vs. splitting” red-herring) considers the complexity of the so-called maraisii-complex, and I am sure that Breuer does not. I frankly think that most observers cannot even dream how complex it is and that it is just braggadocio to write as though anyone can. Breuer implies that one can make the inference of similarity between rossouwii and maraisii from the illustration (Fig. 27), the description and its given locality. This is patently and self-evidently false as is evident from the fact that I used this same information to reasonably deduce the similarity of rossouwii and serrata.
I did not originally submit a copy of the von Poellnitz lectotype, perhaps because I felt that the response “you will see that rossouwii is quite a different plant from serrata“, is what I could have expected from an uninformed audience! The fact is that all the evidence, circumstantial and descriptive, show that they are the same. Including the lectotype would have just been “overkill” and I have never before felt required to write for a readership with less than common sense or basic knowledge, nor for obstructive readers.
It should not be necessary to say this, but the character of the flowers cannot be specified in the way that writers are inclined to suggest. They have this same enormous range of variability and it is just fraudulent to say that there are floral characters by which these taxa (the varieties of H. rossouwii which I now recognise) can be unmistakeably differentiated. It is the typological concept which drives people to think there is a single description which can encompass the description of a taxon. This is nonsense. It is a statistical problem and we have the difficulty of establishing the probabilities associated with “decision”. Breuer writes “… von Poellnitz’ description probably depends only on 1 or very few selected plants and no other records are reported for this taxon”. This is circa 1945 circumlocution, it is unnecessary and it is untrue. There was never a question of “selection”, and furthermore we have the Triebner description of the same collection as H. altilinea var. bicarinata. Why this sudden doubt about the interpretation of types. It is a shifting of goalposts.
There are few collections in flower now in April, but I could take this picture of three flowers from the one collection MBB6672 H. maraisii – Koningsriver, SW Robertson (Fig.33). It is not an exceptional case or an exceptional collection. But one can see the differences in thickness of peduncle and variation in the individual flowers. It can happen like this, albeit to lesser degree, on the same plant. Colour is also frequently quite variable between and often within collections. Many years ago I undertook to illustrate all the flowers in my maraisii collections. But they were so variable that I just gave it up. The variability in the flowers mirrors the variability of the rosette and the latter is just a little easier to deal with. One has the added problem with flowers, of the changes which take place during the opening and aging process. So then how does one define the flower, say, of H. mirabilis. It cannot even be done for the range of leaf rosettes, and this has now been shown in the literature many times.
May I also point out that one of my understandings of botany and plant classification is that there is a tendency to overreach ones talents. I have freely admitted that I am there because of the paucity of talent. I frankly do not think the environment is enriched by the particular article I now refer to in these added notes.
I can point out that there are writers who really persist in flogging dead horses. The result is phantasmagoric fanfaronades. The article concerning the name H. correcta (Alsterworthia 1:6, 2001) is not particularly accurate either and indeed it may be my own fault. Smith explained that Mrs. Blackburn did not in fact collect correcta ‑ it was a Mrs. le Roux and the plants came from near Vanwyksdorp. Later Mrs. Ferguson sent a plant from 26 miles east of the town. Thus from this information H. correcta has to be a later synonym for H. emelyae.
Regarding types:- the painful realisation was actually expressed by Schelpe in the distant past, and brought home to me many times. These old types are doubtful and ambiguous. I was faced with that option long ago – just forget them all, or try to reach a compromise with the understanding and information available at the time. H. rossouwii is a case where there is real new information which would have made me use the name rossouwii in 1976 and before, had it been available. But here, had I persisted with the name serrata for this new material MBB6983 and 6984, there is absolutely no doubt in my mind that I would have been criticised for failing to see the connection to rossouwii. Not that this would concern me unduly, for I think there is actually very little benefit in this mindless re-interpretation of types which takes us nowhere. Notwithstanding, and without compromising, anything so far written:- the really disconcerting thing about the whole matter is that without the locality data, it will be possible to generate an argument bolstered by illustrations, to show that rossouwii was in fact H. decipiens, H. cooperi or H. herbacea (among others), of the option less improbable than H. maraisii.
The closing paragraphs of the article are a rather hackneyed parody sung by most “wannabe” taxonomists. Of course it is right when Breuer says if we lump it means we would have to lump many other things. It is just as true to say, if we split we have to split many other things. That does not make sense either. This is ancient stuff of 60 years ago and it seems strange that it still provides so much fodder for writers, who have in fact no rational species concept to begin with. The Linnaean system was based very largely on floral characters and hence taxa at all levels are based on “related floral characters”. To suggest that this pertains in some special way to “series” is not correct. This sums up the article in Alsterworthia. ♦
In a very interesting book by Stephen Gould entitled “Rock of Ages”, in which he propounds his principle of NOMA – non-overlapping magisteria. This states that science and religion should not be confused nor mixed.
So this is not a confession of confusion – you do not confess to what is obvious. It is an admission, and an admission can be construed as an apology. But, as a rhetorical question, how can one apologize and expect forgiveness when one continues to walk the errant path?
I started to write about Haworthia to dispel confusion, and yet more than 40 years on, this confusion has not become any less. The conclusion I have come to (and I wish it was a closure) is that the prime source of confusion is simply the human condition. In mystic philosophy one can read… “Born in ignorance, we live in ignorance and we die in ignorance.”
I think that my interest in Haworthia stems from my conscious effort to dispel this primal confusion and find some of the order in my view of creation. The classification of plants suggested just one small piece of my world which was available to me, and Haworthia as one group which no one else could explain to me. What have I now learned and what contribution does this make to dispel confusion?
My courage to now say something more directly arises from a recent request by SANBI to write a synopsis of Haworthia for an E. Cape Flora. I feel that I have done that fairly successfully. The problem is now to produce a similar product for the SW Cape and this is considerably more difficult.
To explain Haworthia one has to first cross the hurdle of the present classification of the Asphodelaceae and the perceptions that Aloe, Haworthia and other genera are clear discrete groups that the classification suggests. Ignore completely the issue of whether the so-called species are clearly separable. Haworthia consists of three quite discrete groups which have no closer relation to each other than do any other of the extant Asphodeloid genera. They each have their own problems and idiosyncrasies but it is the subgenus Haworthia which concerns me most.
In 1975 I prepared a map to illustrate the relationship of the SW Cape species (primarily south of the Langeberg Mts.) and this was published in Excelsa (5, 1975). In this map I tried to illustrate the problem of continuity which simply makes it impossible to recognize clear closed groups of plants which we can with any truthfulness say are species. It is the reality for Haworthia, and my conviction is that the model which underlies the Latin binomial system is fundamentally flawed. It works because our ignorance clouds its shortcomings.
Map 1 Distribution of SW Cape species of Haworthia
Updating my map may show why (Map.1). It is perhaps possible to identify three main elements that are nevertheless extremely difficult to map. Very seldom do we find all three growing in close proximity and never do we find any of the components I suggest doing so either. We often struggle to suggest which of the three we are being faced with.
The names we can use for these three are:-
H. retusa – to include or cover mutica, turgida and pygmaea
H. mirabilis – to include or cover maraisii, magnifica, emelyae and heidelbergensis
H. floribunda – to include or cover parksiana, chloracantha, floribunda and variegata.
There are other elements viz:-
H. herbacea – to include or cover reticulata, maculata and pubescens
H. rossouwii –
But this is a simplification and not a solution.
What simply has to be recognized as a fact of life that this problem is not of my creation and I do not believe that the use of any technical sophistry, trick of classification or nomenclature will possibly dispel it. These “species” are as interconnected as the strands of a complex web as intricate as any a confused spider could construct.
Essentially one has a mid- to late-summer flowering group and a spring flowering group, but they are linked. H. pygmaea, in the retusa complex, actually springs from the mirabilis complex; while its counterpart mutica in the west, seems to spring from the retusa complex. Emelyae, in the mirabilis complex, seems to be linked back to rossouwii via multifolia. These are the kinds of realities one has to deal with.
The relationship of the three main elements I have mentioned differs from point to point on the map, and to describe these relationships means that one has to virtually deal with populations and groups of populations one-by-one. I have done that, particularly in the book Haworthia Update Vol. 1n and again in Vol. 2. But since the drafting of the last manuscript, I have been to the field again and found more material, which I will show now.
Essentially there are three areas which I will cover:- a. the southern Potberg and where the Breede River enters the sea south of Swellendam (Map 2); b. secondly Klipfontein at the northwest of the Potberg (Map 3); c. thirdly an area just west of that which I also call Die Kop, Wydgelee (Map 4). The localities are not indicated precisely on these maps but the detail is in archived and available information.
Two of three relevant populations are discussed in Chapter 15. These are ADH594 Sandhoogte, and MBB7248 Lower Breede River H. mirabilis (pilosa). I refer to CB2018 and I can now report and illustrate it myself. This is MBB7251 (Figs 2a & b) from Buffelsfontein within the DeHoop Nature reserve. In the field the plants are not distinguishable from ADH 594 and grow in exactly the same Fynbos habitat viz a very rocky and grassy streamside slope with a dry northern aspect. The illustrations are of plants cultivated and hence far greener and softer than in habitat. What is striking is the tendency to surface spination which is such a feature of MBB7248.
b. Northwestern Potberg.
Some of these populations are also covered in Chapter 15. These are MBB6890, MBB6886, MBB6882, MBB6889 and MBB6544 (figs A18…22). My comment that H. variegata occurs here too should be borne in mind. This is on the upper slopes of the extreme western end of the Potberg. There is another record of C Burger’s from further east and south of Diepkloof for which I coined the name H. variegata var hemicrypta. Some doubt will now be thrown on the wisdom of that decision. The mirabilis populations referred to above are on the farm Juliesfontein and my new records are now east of that towards Diepkloof along the Breede River.
1. MBB7487 H. floridunda. Byeneskop (figs 3a to g). No good opinion is ventured in Chapter 15 concerning the populations noted above. Those populations seem to fall in the context of an interaction between H. maraisii/heidelbergensis/floribunda and I warn of infusion of H. variegata. The Byeneskop is on the very eastern border of the farm Juliesfontein and the plants are surprisingly and confidently comparable with the typical form of H. floribunda. The leaves are generally fairly strap like and flattened to the characteristic twisted ends as evidenced in floribunda. A few plants (see fig.3f) do have shorter and stubbier suberect leaves which characterize so many collections that confound the recognition of the relevant species. This particular habitat can be described as Renosterveld on silcrete with dominance of white clay and much grass and other succulents. Similar plants were also seen about a kilometer eastwards on the verge of vegetation transitional to Fynbos.
2. MBB7492 Klipfontein East (figs 4a to d). This is again a typically H. floribunda-like population. Whereas the preceding tends to be slightly tuberculate with the rougher greyer texture and colour of H. maraisii which would suggest floribunda var dentata, these plants tend to have a smoother leaf surface. Pictures of the flower are included although of very little diagnostic significance. The habitat is a steep rocky sandstone hillside vegetated with low Fynbos species.
3. MBB7494 Klipfontein East (figs 5a to d). The plants suggest H. floribunda except that the colour of the leaves is greener and there is a tendency to translucence, or less opacity of the leaves. The white flecks in the leaves are quite conspicuous and are more often encountered in H. turgida. Their significance taxonomically is probably nil as these markings/inclusions may appear in plants as remote as H. angustifolia var baylissii on the Zuurberg, E. Cape. The habitat is again short grassy Fynbos but unusual because of the presence of finely textured white quartz rock.
4. MBB7495 Klipfontein East (figs 6a to c). While we have been looking at plants on a series of buttresses of the Potberg, these plants are now closely associated with large white quartz boulders. They are small and although very similar to the preceding, the leaves seem to have lost the characteristic rounded ends and the margins are relatively strongly toothed.
5. MBB7496 Kleinberg (figs.7a to g). This population is only about 1km east of the preceding but there is a transition to Renosterveld. The habitat is again silcretious with the typical white clay substrate. This seems to be a very erodable surface and plant cover is quite low. The plants are surprising because they depart quite radically from the floribunda-like mode of the preceding populations. They are now distinctly magnifica-like and particularly reminiscent of var atrofusca except much smaller. Curiously a population across a small valley southwards in sandstone and Fynbos is in keeping with the floribunda-like finds westwards.
6. MBB7499 Juliesfontein (figs 8a & b). We returned to Juliesfontein to look at a promising rocky slope between Byeneskop and our previous closest collection on Juliesfontein. These are almost directly north of H. variegata and as can be seen from the pictures they revert to the forms described in Chapter 15 for that area viz. there is a similarity to heidelbergensis. The more leaves are more frequently pointed rather than rounded and translucence and venation is more conspicuous.
c. Die Kop, Wydgelee.
The nearest plants dealt with in other chapters are MBB7539 at Tarentaal that I liken to magnifica var atrofusca and MBB6030 at Napky which is maraisoid with floribunda-like character (type locality for H. maraisii var simplicior with rotate leaf-tips). It is important to note that the Tarentaal plants are not uniformly larger that my identification suggests and also that H. mutica is also present at its most southeasterly locality known south of Napky. Tarentaal only just appears on the map and the mutica locality is just off the map to the upper right (northeast) corner. It should be noted that populations assigned to H. heidelbergensis occur to the west at Kathoek, Beyersdal, N. Windheuwel and terminating in the contentious var minor at Rooivlei north of Bredasdorp. I think if an overview was now made of all these populations I have assigned to heidelbergensis in different chapters and in different publications, it will have to be concluded that it may represent the pivotal element in the whole classification process. Where it was the least represented species in the period preceding publication of Haworthia Revisited it is now far better known and any hypothesis prepared to test relationships in these southwestern Cape species Haworthia will need to question this centrality. Die Kop is a set of 5-6 silcrete and ferricrete hilltops of varying size and height but otherwise not significantly different from a large number of such habitats extending from Stormsvlei in the northwest and terminating against the Potberg and Breede River in the southeast. Collections from several of these have already been reported on in Chapter 15 with the general observation that their identification is in the uncertain linking maraisii and mirabilis. I hope, in the minds of readers, that I have not excluded heidelbergensis from this milieu.
1. MBB7500 Die Kop East (figs 9a to v). I doubt if any collection has generated such reservations in my mind as this one has and many illustrations are used to express the problem. The plants are large and up to 80-100mm in diameter. They are closely spaced in a small grassy, non-rocky area which seems to be geomorphically stable. They are extremely variable and resemblances extend from badia to retusa to mutica. The leaves may be short and rounded or fairly long and attenuate, closely adpressed to the ground (recurved) or sub-erect. One notable feature is that the plants have a milky colouration to the leaves that Seven Hammer once remarked on as a feature of H. mutica. While variability is known to be great, it is seldom that it is as dramatic as this in a population of plants as they occur in the field.
2. MBB7502 Die Kop West (figs 10a to c). The plants here are smaller and more widely distributed and scattered over a larger area. The habitat is skeletal and rocky and the vegetation more herbaceous than at the eastern site. The similarity of the plants to those of 7500 is close but the variation is less extreme. There is less evidence of a mutica influence and a return to more or less the maraisii/floribunda/heidelbergensis archetype.
Fig. 9 7502 Die Kop W
3. MBB7503 Die Kop Mid (figs. 11a to i). The first impression I had was that the plants included more mirabiloid influence. Not as big as 7500 and also scattered on a skeletal slope. At one point there was a gravel patch with Gibbaeum austricola which is commonly found on such gravel patches associated with ferricrete inselbergs.
4. MBB7504 Die Kop North (figs 12a to k). The plants are smaller and there is a more evident floribunda influence. I did not think that these plants were significantly different from the many populations discussed in Chapter 15 between Tarentaal and Stormsvlei.
Conclusions: It should be very evident from all the information provided so far that more information is not producing a better picture of discrete elements. To the contrary and this reinforces the conclusion already mooted, that movement to a new view of “species” in the subgenus Haworthia is necessary, but is it possible?
Acknowledgement: Mr. Keith Spencer, DeHoop Nature Reserve. Messrs. Christine & Malcolm Wallace, Ziekenhuis. Mr. John Douglas-Hamilton, Ballyfar. Mr. Johan Groenewald, Klipfontein, Potberg. Messrs. Saartjie and Neil Neetling, Juliesfontein. Messrs. Adele & Vlooi DuToit, Die Kop & Frederickskraal. Messrs. Pet & Johannes Uys, Die Kop. Messrs. Sofia & Jurie Vermeulen, Tarentaal
Again this piece is written against the background of a detailed discussion in Haworthia UpdateVol. 2. Again I am not able to say the situation is fully comprehensible and neither do I want to encourage the daffy view that nature is just too much for us all. It is really curious that this species is woven into the fabric of H. mirabilis and also into that of H. chloracantha, H. parksiana and H. variegata. New finds have not clarified the picture so much as added another dimension to an extraordinary display. Not that H. floribunda is a spectacular species. In the field it can be extraordinarily cryptic and obscure while in cultivation it is an unlikely favourite. I do not want to repeat what I have already written while I hope that this will not contradict that either. H. floribunda seems to occupy a clear niche along the base of the mountains between Albertinia in the east and Swellendam in the west. It occurs as discrete from any other species although hybrids with both H. retusa and H. mirabilis do occur. South and west of Heidelberg it seems to lose its identity within H. mirabilis and then emerges briefly in a limited area near the Potberg in the southwest in a ‘mirabilis’ context as well as in H. variegata context. At Klipfontein farm at the western end of the Potberg it seems to be recognizable in relatively the same form as the very original description. But let us look at new information.
1. MBB7722 H. floribunda ‘major’. Appelbos. This is just west of Swellendam and we came across this locality while trying to confirm a very old record of the form ‘dentata’ once brought to me from Buffeljagts very nearby. The plants are quite large and have a clear green colour quite unlike the normal dark green or purplish hue of ‘typical’ floribunda. It should not be lost to the reader that this same colour difference characterizes the difference between H. retusa and H. mirabilis. Or that both colour variation is apparent in H. mirabilis ‘paradoxa’. Kobus Venter found a similar population northwest of Swellendam and further so than my rather poorly noted collection of proliferous dark plants with predominantly pointed leaves. In fact this population is hidden in its citation in my Revision under H. variegata. This will perturb those readers who are imprisoned in the paradigm of rigid similarities that define species. The Appelbos plants generally have the straight acuminate leaves that H. variegata has, but before arguing the issue and without actually properly looking at the whole H. variegata milieu, here are two connected records …
2. MBB7738 H. floribunda ‘major’. Swellendam. These plants were in fact small when first collected and in cultivation grew so large that I coined the name ‘major’ for them.They do still exist in a very small and disturbed area close to gum trees but curiously in moss free of leaf litter. I did also find them a little further away in a more grassy area where they are/were more typically small and dark coloured. I should note that I also recorded this ’dentata’-like version within the Bontebok Park close to where H. mirabilis occurs and I am still committed to again finding that population in the light of this new material.
3.MBB7774 H. floribunda ‘major’. Swellendam. We were concerned about the disappearance of an interesting form of H. minima from the area and in extending our search for plants came across H. floribunda in gumtree leaf litter about 400m east of the previous H. floribunda population. At first glance I did a double-take and then another because I actually thought I was looking at a slender form of H. marginata as occurs at Drew. The plants we saw were really large forms of H. floribunda with pointed leaves exceeding 100mm in length. They were quite proliferous and not all nearly as big as the first surprise ones. We also located smaller plants in numbers on the opposite side of the road also under gumtrees.
4. MBB7708 H. floribunda ‘dentata’. Goedverwagting. Apart from the Buffeljagts and Bontebok Park records there is an inexplicable hiatus in the appearance of the species west of Heidelberg. This is probably simply due to inadequate exploration that these new records now expose. It is quite evident that in the general area north of an imaginary line connecting Swellendam and Albertinia, H. floribunda does occur independently and in near company of H. mirabilis and the latter invariably in the form of ‘atrofusca’. This Goedverwagting record fills this gap as an association with ’atrofusca’ that pushes southwards.
5. JDV93/58 H. floribunda ‘dentata’. RoseInnis Drive; JDV93/56 do. Plattekop; MBB7760 do. Witkleikop; MBB7764 do. NW Platkop; MBB7767 do. Plaatjieskop. Being so cryptic may be a reason for the non-record of this species. It is associated with the ferricrete inselbergs and has the really curious association with H. mirabilis already remarked upon. In these four records, three of which are new, the plants occur independently of other Haworthia species and there is no obvious habitat preference that suggests why this is so. I also think that there is significance in which the characteristic rounded leaf tip is replicated especially in the juvenile leaves of H. mirabilis. The last cited record is the dramatic one. The plants are in very close association with H. retusa in two small geomorphologically different and adjacent sites, and we observed what seem to be reciprocal hybrids.
JDV93/58 H. floribunda ‘dentata’. RoseInnis Drive
JDV93/56 H. floribunda ‘dentata’. Plattekop.
MBB7760 H. floribunda ‘dentata’. Witkleikop.
MBB7764 H. floribunda ‘dentata’. NW Platkop.
MBB7767 H. floribunda ‘dentata’. Plaatjieskop.
Hybrid MBB7768 H. floribunda x retusa, Platjieskop
In a recent set of articles published by the Haworthia Society I wrote the following in connection with H. floribunda… “MBB7738 H. floribunda ‘major’. Swellendam: These plants were in fact small when first collected and in cultivation grew so large that I coined the name ‘major’ for them. They do still exist in a very small and disturbed area close to gum trees but curiously in moss free of leaf litter. I did also find them a little further away in a more grassy area where they are/were more typically small and dark coloured. I should note that I also recorded this ’dentata’-like version within the Bontebok Park close to where H. mirabilis occurs and I am still committed to again finding that population in the light of this new material”.
In connection with H. mirabilis, I wrote … ”The Dankbaar plants are small versions of this and of course tie up with both older and newer (MBB7704) records for the Bontebok National Park. 2. MBB7743 H. mirabilis. Bontebok Park: Having written that, we did in fact locate still another population and of course it looked different as the area where it occurs had been recently burned and being on a northwest aspect the plants were very exposed and even more cryptic than usual.”
Since those finds and comments, we have again been back to the Bontebok Park. It is of course a large area of largely recent geological origins and therefore mostly ferricrete and riverine gravels. The lower lying areas are seasonally flooded. Being unsuitable for cropping or grazing, the area was handed over to the conservation cause (thank goodness for non-arable land). H. venosa occurs on the exposed Bokkeveld Shale cliffs along the Breede River, and H. minima is present in small scattered populations. H. marginata has virtually disappeared from thePark.
It is quite difficult to identify suitable Haworthia habitats as even in the seasonally flooded areas there is much gravel and consequently good drainage. It was in such a site that I seemed to remember seeing the record for H. floribunda cited in my Revision. Our searching has been very limited as it is quite difficult to explore the park with the restraints of accompanying rangers however willing or enthusiastic and knowledgeable. This is because I personally need mental space and physical freedom to walk where my intuition takes me. On this occasion we first went to an old entrance to the Park from the town where we thought H. marginata was first recorded. I was very disappointed to find that the hillside there had suffered the same mindless engineering fate that hit the gravel hills around Worcester. These were ”strip-mined” to obtain road materials. Of course this may have been necessary, but not the mindless extent of the surface scraping when some consideration for simply good housekeeping would have limited the desecration of the landscape and demanded some degree of resortaion.
We moved some way west to avoid the disturbed area and eventually found H. mirabilis (see MBB7805 Figs 1) about 500m east of the MBB6513 record of 1996 when I was writing the Revision, and illustrated elsewhere. That record came from a more distinct interface with the riverine shale while the new record was in more sandstone gravels from an old river bank. The plants were rather smaller than in 7704 with quite slender leaves which were also fairly toothed. There was little evidence of the flattened rounded leaf tips that characterize H. floribunda. We drove around the park trying while I was wracking my memory banks for where I might have seen H. floribunda so many years ago and eventually was drawn to where Tineke Kraaij had on a previous visit shown us H. mirabilis on a burnt zone (see MBB7704 Figs 2). These are new images to the original published ones, because after 6 months the plants had largely lost the burnt look but also being in flower they were far more visible. The leaves were more elongate and pointed than suggested by the photographs we took on the first visit and the plants reminded me very strongly of what I named as H. floribunda var. dentata. I also add here images for H. mirabilis (see MBB7744 Figs 3) from another population approximately midway between the previous two. Here the area had also been burnt as part of the management plan for the Park, but the plants themselves do not seem to have been affected possibly because there was a lighter fuel load that at the 7704 locality at the time of burning. The plants were more “maraisoid” and similar to the very first record of mine for this species in the Park (MBB6513). That collection I often noted among a set of 4 or more populations extending from Riversdale westwards to Riviersonderend that confounded any distinction between H. magnifica, H. maraisii and H. mirabilis.
All things considered, it seems highly unlikely that H. floribunda could possibly co-exist with H. mirabilis other than in the present known configuration where we have H. floribunda var major close to and north of the N1 highway near the town and inland, whereas H. mirabilis is further south. This geographic and presumed taxonomic relationship seems to hold at Heidelberg and Riversdale with some anomaly at Albertinia down the Gouritz river. This is where H. mirabilis is absent (with consideration for the retusa/pygmaea conundrum) and where H. chlrorocantha or H. variegata may be confounded in H. floribunda (or vice versa!). ♦
I wonder. I have written so many words purporting to be my last that my credibility here too must be under stress. Two very recent articles of mine in Alsterworthia deal essentially with that issue, although they also cover the discovery of Haworthia mutica (Buffeljags) (= H. groenewaldii Breuer). They do not cover my subsequent thoughts on actually reading the description of this new “species” by Breuer, Marx and Groenewald. I hope that the present manuscript will explain why I reject this as a Latin binomial although anyone who is in the least familiar with my writing should already know. Spurred on by that discovery, I instigated a search in another area of the Buffeljags valley adjoining the Bontebok Park accompanied by Jannie Groenewald who informed me of what he had found in still another area I had long wanted to explore. So I instigated another search there too and again with Jannie. A discussion of these new finds is submitted to Cactus and SucculentJournal where I trust it will be published. The essence is already in Alsterworthia and this article is written to widen the readership, submit more pictures and maintain continuity with the 6 volumes of HaworthiaUpdate that Harry Mays has been so conscientiously and determinedly publishing. This is all writing that may not otherwise have seen the light of day. I am personally extremely grateful for that as I have had a mania since writing my revision Haworthia Revisited and Update Vol. 1 (both Umdaus), to set the record straight and explore all the unknowns, or at least some of them. So on with Haworthia retusa ‘nigra’.
The second area opened to me by meeting Jannie, was a farm southeast of Tradouw Pass, Heuningklip. In the Tradouw Pass, H. retusa ’turgida’ is found on the steep cliffs at the southern end. The road has been rebuilt since I first saw the plants and the population is greatly reduced from what it was. I simply do not have a picture handy to show what those plants are like, but that is not the need here and it does appear in Haworthia Revisited. Immediately south of the pass is another population of plants that puzzled me from the moment I saw them until now. In Haworthia Revisited there is a picture of one clone under J.D. Kobus Venter’s number 93/35. It is named as H. magnifica var. magnifica. The plants are present in a tension zone between Shale and River gravel, and vegetationally between what was Karoo Valley Bushveld and grassy Renosterveld. In April 1997 I collected seed and grew countless seedlings from there, which were all different. Only one picture, as MBB6666) is included in my long treatise on what I discussed under the title of “How to understand H. mutica var. nigra” in Haworthia Update 2.1:50.That picture is also under the name H. magnifica var. magnifica but it was not intended to stay that way. It must be noted that my use of names in the Update volumes was not in a decision-making process so much as a learning and informative one. I read my conclusion to the treatise and note that I made no formal name changes. It is only as further field exploration yielded more and more information that I came to realize that H. magnifica and H. mirabilis, classification-wise, are inseparable. The Tradouw collection still worried me because of the close proximity of H. retusa ‘turgida’. These plants at Tradouw and all the things I referred to as H. mutica ‘nigra’ are in fact better understood as variants of H. retusa in the complexity of its relation to H. mirabilis.
This article thus covers exploration of an area between Swellendam and Heidelberg that I have neglected for too long. It is particularly applied to the problems of classification and naming that have persistently clouded my life and my relations with other writers. But as importantly to me is the unclouding of the way taxonomy and classification is useful and necessary for those interested at all in plants and how it is viewed by them.
Consideration and use of formal names Here I need to explain the naming style I now adopt. I have dropped the use of any rank below that of the species name. I do this because botany has no proper species definition and consequently species descriptions are just based on wild guesses about possible non-similarity and on the flimsiest of supposed character differences. The loosely used word “typical” is only truly useful in respect of the one plant dried as an almost unrecognizable herbarium specimen that is used to anchor the Latin name. The scientific ranks are aggregates and the terms variety and forma are so too. There is simply too much variation between populations and within populations to enable the certain identification of almost any plant that the binomial system suggests. There are no clear entities and ranks and this article should demonstrate that. There is a separate code of nomenclature that covers cultivars. Note that I do not drop names. I only drop the suggestion of status that additional names have, be they single clones or population references for all the different plants within them. I have used names in Haworthia for 70 years and intensively for the last 40 years and I am very familiar with the difficulties of any system. For the field and all the populations and plants I have seen, I can do no better than offer the list of names I updated in February 2009 in Haworthia Update 5.2:192. This, and the set of Updates, comprises a near total overview of the genus and effectively a revision. The attempted revisions by others fall short for me because they do not cover the range of populations that I have seen nor what history tells me, although they may include populations not seen by me. That is not strange because the area to explore is still vast. Another problem with even trying to be formal, is that which G.G. Smith encountered (Mrs. L. Bolus did too) where several people were generating new names at the same time and trying to keep track formally with concurrent name changes would not have been, is not, possible. The point I should make is that in the last ten years, my field work has confirmed the predictive nature of my classification and these latest finds are in extraordinary agreement with that kind of expectation. What I have found essentially agrees with what I would expect. This is dramatically so for H. mirabilis based on a find by J. Jelimicky at Sandhoogte midway geographically and visually between H. mirabilis ‘bobii’ Hayashi (my nom. nuda ‘pilosa’, ’velcro’, Steven Hammer’s ‘Shaggy dog’) along the Breede river, and a collection of mine further south in the DeHoop reserve, MBB7886 H. mirabilis, discussed (p11) and illustrated (p33) in Haworthia Update 5.2.
Prior to extending exploration in the Buffeljags area, I had also been down to the Haworthia populations in the lower Breede River area. This is where H. mirabilis seems to assume a wide range of forms and H. retusa ‘turgida’ and H. variegata’ are also present. I also, in the company of Kobus Venter and Lawrence Loucka, revisited Kransriviermond and Morning Star south of Heidelberg where H. floribunda is in the mix with H. retusa ‘turgida’ and H. mirabilis (in my broad new sense that includes H. maraisii,H. magnifica and H. heidelbergensis). Thus many of those images were very fresh in my mind when I went to Heuningklip southeast of Tradouw. Looking at all the plants and in editing my pictures I was truly impressed with the reality of continuity. The essence of H. retusa and H. mirabilis is in all those plants. A mobius strip is a strip of paper twisted and joined at the ends to produce a shape that only has one surface. What we have in the Southern Cape, and covered or touched on in this article, are interlinking mobius strips with one main surface as well as loose ends that present identifiable end-points. I tried once to illustrate the continuities between populations with single or multiple lines joining the different populations to express degrees of similarity. I can also visualize a pie-chart in which each population is represented by a circular chart with the contribution of main elements drawn as slices proportional to the strength of contribution. Table 1 is a schematic and very crudely suggested enumeration for such a set of charts. I have put the elements retusa and turgida as separate under an H.retusa heading for no other reason that I want to emphasize turgida as a variant at the two geographic extremes. One is eastwards where H.retusa ‘retusa’ is absent (Brandwacht, Mossel Bay) and one westward where it is present (in mutica, Hasiesdrift, Bredasdorp). H. retusa ‘turgida’ is present both east and west. But there are many populations contributing to the H. retusa pie in all the ranges of the two parts, from 0-50% thus making up the continuity from retusa to turgida.
Consideration and characters.
a. The pie chart Measuring the similarity of any two plants is extraordinarily difficult. Doing this for a population is even more so. Doing so for many populations, including those unseen in order to fulfill the predictive requirement of a proper hypothesis, is next to impossible. Hence all the disputation and name abuse. In table 1, I speculate an arrangement to illustrate the problem. The pie is constructed from the figures in the chart for just that set of genetic characters that are directly relevant to the set of plants in question. The pie is incredibly big and it should be understood that there must surely be pie-parts that spread their influence beyond the restrictions of the slices I conceive. So my table is restricted to the few species that are immediately relevant in respect of geographic association and direct interaction (by possible cross-pollination?). A pie-chart is drawn for each individual population. In the chart I assume that there is a set of genetic information that programs for H. retusa ‘retusa’. I have also suggested a different set for H. retusa ‘turgida’. If one considers only these two elements and all the known populations, my suggestion is that there is a set of charts that moves smoothly from one set to the next with no interruption. There is no discontinuity and the sets average 50:50 for the information. Thus it is one species and H. turgida is noted as H. retusa ‘turgida’.
Table 1. A schematic distribution of similarity
Table 2 is the complete set of pie-charts that should convey the possible relationships of the different populations and from which can deduce and understand to what degree H. retusa and H. mirabilis can be related. Also it will indicate the similarities between, and the difficulties of, trying to explain H. pygmaea in the east and H. mutica in the west. Both arise from the same genetic source. The problem is compounded by the infiltration of H. floribunda in the area midway between Heidelberg and Swellendam with the re-emergence of H. floribunda in the south at Potberg. H, variegata also enters the picture but from the south and also in the Potberg area. H. emelyae is the very probable combined entity entering the little Karoo. H. rossouwii is almost independent except in the very south-west at Bredasdorp and it may possibly link to other elements such as H. herbacea and H. reticulata.
Table 2. Pie Charts
At Heidelberg there is interaction between an element that is a 50/50 retusa/turgida with a variant of H. mirabilis cf ‘heidelbergensis’. The latter name suggests that H. mirabilis is the general name for all the plants of that species, and ‘cf’ is an abbreviation for ‘compare with’ ‘heidelbergensis’ to refer to the small plants in populations previously regarded as independent species.
b. What really constitutes the pie The pie is the totality of the genetic material, the DNA that determines all life forms. It is the genes, the materials of characters and inheritance, and the repository of all genetic information. The pie is very large indeed and it is said that the DNA pie of an elephant only differs by 3% from that of a mouse; that of a man by 1% from a chimpanzee. In Haworthia one species from another ‘species’, I could guess at 0.001% or less regardless of whose concept I follow. This difference does also not reside in one consolidated slice that can be reduced to a single small pie as I have done. In DNA analysis, certain segments of DNA from cellular chloroplasts, mitochondria or the chromosomes themselves are isolated and the sequence of the paired amino-acid is analyzed. It requires a formidable statistical process to arrive at a two-dimensional tree (phyllogram) that illustrates a branching relationship with a measure of branch separation and lengths. This can perhaps also be used to generate pie-charts with an innumerable number of slices. I personally consider the phyllogram generated in the DNA sequencing process grossly inadequate.
c.) The characters This is extraordinarily difficult to specify no matter how obviously the plants seem to differ from one another. Number, size, shape, colour, arrangement, surfaces, spination and attitude of the leaves in the spiral rosette constitute the main things that determine how we see the plants. These are influenced by growing medium and growing conditions so these are complicating factors. Flowers are used in the same way as the leaves but far less so because they are so similar in all the species. Flowers are also not always present either which reduces their usage. Time of flowering is a major factor because it is linked to the reproductive process. It is also difficult to capture any firm picture of the flower because of the aging process in the 3-5 day life of the individual flower.
d.) The relevance here It simply is not necessary to parade the obviousness of this and the way in which will be attempted anyway in the discussion below. It is flowering time that may be critical beyond, or linked, to the facts of spatial geographic relationships of populations. It is traditionally flower structure that has formed the basis of classification of plants because of the reproductive significance and the role of reproduction in distribution of genetic material between plants and populations. Pollination of Haworthia seems to be primarily by a Solitary bee species. These bees make individual nests and are not communal like the Honey bee. They make tunnels in the soil as nests during the summer months. I have observed them as adults in the winter but then they are less numerous and less active. Few Haworthia flower at the height of winter and do so in two main time frames viz. spring and late summer. The implications are that breeding between isolated populations apart from any other consideration of compatibility, is a question of distance and time. A nest bound Honey bee can forage to 13km from its nest. A free flying Solitary bee can be speculated to cover a considerably greater distance. All these things play on the frequency and power of inheritance. Genetic drift is acknowledged as the genetic movement within a population away from the norm for a species because of isolation, while genetic flow is perhaps the holding together of similarity over geographic distance by a travelling pollinator. Thus the statistical probabilities of breeding across distance are impossible to derive.
I cannot quantify any of these things nor suggest any greater knowledge but this suggests to me, given the freedom with which the various species hybridize in cultivation and even in the field, we need to consider geographic relationships. In my extensive experience with other genera, it appears to me that a key consideration is if things grow together and maintain their difference, or not. Haworthia species are invariably in small isolated populations and seldom do different species grow intermixed. They are also influenced by geology of the substrate and particularly so because they favour the lowered competition from more vigorous and larger plants on the rocky sites or shallow soils they favour. That can be stated in the converse, Haworthia are favoured by conditions poor for other plants.
e.) Flowering time and growing together. The existence as discrete species can be judged from the way in which populations share geographic space. I discussed this in some detail in Update 2.1:76. If two sets of plants grow close together and flower at the same time, while still maintaining obvious group differences, then they are most probably different species. If they flower at different times while still maintaining difference then the question of being different species is taken as certain. If the groups are separated and flower at different times while the differences are small and apparently within the experienced range of variation observed between populations, then it may be problematic. One would need to resort to the extended distribution and variability of the groups in question. Generally in Haworthia, species belonging in the same sub-genus do not share habitat and this is indicative of fairly rapid absorption of introduced genetic material (by pollinators). Hand-pollination and the ease with which plants of different species hybridize in cultivation suggest that there is practically no breeding isolation between them. In the following discussion, use is thus made of general distribution of species and proximity of populations in assessing to which species they best belong. It should be borne in mind that variability may be clinal. This means that the plants change along a geographic gradient. The direction of the gradient may also not be of increasing distance i.e. linear. It may also be circular so that ends of a gradient may meet again as two populations that appear wholly different, but linked by an extended set of other populations. A much more complex situation may actually exist where these clinal gradients may be multidirectional and thus involve more than one apparent species. My observation is that populations do differ and differences follow a pattern that becomes predictable i.e. gradual genetic difference over geographic area. This is not just a result of environmental gradient, but also linked to the question of genetic drift (the tendency of isolated populations) to move away from the general norm. It also results from the genetic flow between populations where the change of genetic material between populations at the distribution extremes is only through other populations.
f.) The species definition This is a subject I have covered many times. My view is that species are dynamic chaotic systems oscillating between different character states with time and with different rates of mutational change. Closely related species may not be recognizable on the basis of single character states and the spatial geographical distributions and local juxtapositions of groups of plants may be the only way to understand what the situation is and so make sensible decisions.
g.) Habitat geology I contend that species richness in South Africa is largely due to its varied geology, skeletal soils and comparatively mild climate. This present discussion deals with species that are associated with a geological phenomenon that is the Tertiary deposits of the Southern Cape. I am no geologist and am required to make the pretention I am one to explain why plants are distributed the way they are and what influences difference among them. In this case the vegetation at Buffeljags, Swellendam has been described (by Breuer et al. cited below) as on silcrete, and the general vegetation as by vegetation scientists as Ruens Silcrete Renosterveld. But there are two problems. The first is that vegetation classification is not infallible and I have never found it really useful in dealing with a wide range of species that occupy skeletal habitats. Secondly the substrate geology is definitely not simply silcrete nor is the clay soil either simply bentonite or kaolinite. There are many problems in the classification of clays. Bentonite in common terms seems to be the yellowish clay mined in large quantity from Heidelberg eastwards and there is a transition to ochre bearing clay. Bentonite may be aluminium, potassium, sodium or calcium dominated. As a montmorillomite clay it should have expansive cracking properties. Kaolinite is popularly regarded as the very white clay that is commonly seen under the rocky top layer of the sharp inselbergs that rise above the wheatfields west of Heidelberg. It seems to be less mineral dominated clay and does not have the expansive/contractive properties of bentonite. The distinction between ferricrete and silcrete seems to be that in ferricrete, iron oxides are infused into the petrified water deposited layers. Both the kaolinitic and bentonitic clays seem to be derived from the weathering of Bokkeveld shale that underlies the silcerte/ferricrete layer. It does not require any stretch of the imagination to see now why any one inselberg may be a little different from the next, and sometimes a lot different. What we do not know is how rapidly the landscape has moved from a broad white silcrete/ferricrete underwater plain to the remnant condition of that rocky plain today. Neither do we know what it has meant for the plant species that are linked to, and dependent on, the habitats generated in the erosion processes.
Map of Swellendam to Suurbraak area.
Map Legend – east of Swellendam.
1. MBB7899 H. retusa ‘nigra’. Heuningklip. 2. MBB6666 H. retusa ‘nigra’ ↔ H. mirabilis. S Tradouw Pass. 3. MBB7896 H. retusa ‘nigra’. Heuningklip. 4. MBB7897 H. retusa ‘nigra’. Heuningklip. 5. MBB7898 H. retusa ‘nigra’. Heuningklip. 6. MBB7888 H. mutica. Rotterdam. 7. MBB7889 H. mutica. Rotterdam. 8. MBB7890 H. mutica. Rotterdam. 9. MBB7801 H. mutica ‘groenewaldii’. Buffeljags. 10. MBB7741 H. mutica. Dankbaar. 11. MBB7892 H. marginata. Rotterdam. 12. MBB7891 H. minima. Rotterdam. 13. MBB6644 H. mirabilis. SW Swellendam. 14. MBB7704 H. mirabilis. Bontebok Park. 15. MBB7805 H. mirabilis. Bontebok Park. 16. MBB7823 H. mirabilis. Klipbult. 17. MBB7887 H. mirabilis. Rotterdam. 18. MBB7876 H. mirabilis. Disselfontein. 19. MBB7901 H. mirabilis. Crodini E. 20. MBB7902 H. mirabilis. Crodini E. 21. MBB7903 H. mirabilis. Crodini E. 22. MBB7904 H. mirabilis. Crodini E. 23. MBB7905 H. mirabilis. Crodini E. 24. MBB7906 H. mirabilis. Crodini E. 25. MBB7907 H. mirabilis. Crodini E. 26. MBB7908 H. mirabilis. Sandkraal W. 27. MBB7909 H. mirabilis. Sandkraal W. 28. MBB7910 H. floribunda. Rietkuil. 29. MBB7912 H. mirabilis. Rietkuil. 30. MBB7913 H. mirabilis. Rietkuil. 31. JDV93/35 H. retusa ‘turgda’. Tradouw Pass.
The populations
Set 1 – THE TRADOUW PASS AREA (H. retusa ‘nigra’) This first section deals with populations that complement those in the long discussion under the title “How to understand H. mutica var. nigra” in Haworthia Update 2.1:50.There I ended with the words … ”The habitat photo (of H. mutica Hasiesdrift) raises such strong images of H. retusa that doubt about the affinity of the var. nigra cannot easily be laid to rest.” Added were these words … ”There the saga for the moment must rest.” What I had in mind was exploration that no one seems very anxious to do. This has now been done and I would lay the matter to rest as follows:
1. MBB7899. H. mirabilis, Heuningklip. See figs 1-30 Fig. 1 is a scenic view of the area to show the countryside and the type of inselberg the plants are often found on. Fig. 2 is a closer picture of the erodible clay held in place by grass clumps at the base of which the plants often are. Occasionally they are simply out in the open sun. I term these places pressure bursts because water build-up in the soil causes collapse of the small banks and erosion can be quite fast. Several times I have found plants lying in collapsed soil. Such a soil bank near Heidelberg has retreated by nearly 2 meters in about 15 years albeit accelerated by road maintenance, but it does relate to any attempt to explain how much suitable habitat may have disappeared with time.
Fig. 1 A view looking north from Heuningklip to Tradouw. MBB6666 at arrow and MBB7899 at 6pt star – bFig. 2. A photograph of the habitat of MBB7899.
Figs 3 to 30 is a selection of plants at the site where one is almost compelled to examine each plant separately and easily generate another 28 practically different images. Fig. 3 is of a plant with the surfaces roughened by small tubercles. SEM (scanning electron microscope) images of similar leaf surface often show incipient tiny spines on the tubercles and they appear as spines or hairs in populations like H. mirabilis ‘pilosa’ at Ballyfar, or H. mirabilis at Windsor Riversdale. This surface roughness and all its degrees is fairly characteristic of H. mirabilis but is does occur in H. retusa too. Plants illustrated in Figs 4 and 16 has one muttering Kruisrivier and Komserante, northeast and east of Riversdale respectively, and which I would have allotted to H. magnifica in the early period of my exploration no doubt have Breuer and Hayashi names. Now I simply say H. mirabilis and add the locality to place the plant to at least give it a geographic reality that is better than using the formal Breuer/Hayashi in hanging commas as I also suggest. Fig. 6 is apparently a seedling and demonstrates the leaf-shape of H. floribunda that led me to suggest that H. floribunda is a perpetuated juvenile form. Of course the evidence suggests to me that H. floribunda coalesces with H. mirabilis in this wider area although it does appear at Swellendam to the west of my map as the large form and at the extreme SE corner of the same map as a convincingly normal form. Fig. 9 demonstrates longer pointed leaves and Fig. 12 short more obtuse leaves. Fig. 17 is of a plant that has relatively smooth leaf surfaces rather like H. retusa and the leaves are conspicuously lined. In Fig. 20 there is a suggestion of muticate (without a point) leaves. Fig. 22 is again quite strongly lined. Fig. 22 shows a leaf with a single line. Fig. 25 demonstrates what is termed rotate leaves where the points of the leaves point around as though to encircle the plant. Fig. 28 made me recall plants of H. mirabilis at Melkhoutrivier in the south west near Malgas and further to Ballyfar. The flowering time is late summer.
Fig. 3. MBB7896 H. retusa ‘nigra’. Heuningklip.Fig. 4. MBB7897 H. retusa ‘nigra’. Heuningklip.Fig. 5. MBB7898 H. retusa ‘nigra’. Heuningklip.Fig. 6. MBB7888 H. mutica. Rotterdam.Fig. 7. MBB7889 H. mutica. Rotterdam.Fig. 8. MBB7890 H. mutica. Rotterdam.Fig. 9. MBB7801 H. mutica ‘groenewaldii’. Buffeljags.Fig. 10. MBB7741 H. mutica. Dankbaar.Fig. 11. MBB7892 H. marginata. Rotterdam.Fig. 12. MBB7891 H. minima. Rotterdam.Fig. 13. 13. MBB6644 H. mirabilis. SW Swellendam.Fig. 14. MBB7704 H. mirabilis. Bontebok Park.Fig. 15. MBB7805 H. mirabilis. Bontebok Park.Fig. 16. MBB7823 H. mirabilis. Klipbult.Fig. 17. MBB7887 H. mirabilis. Rotterdam.Fig. 18. MBB7876 H. mirabilis. Disselfontein.Fig. 19. MBB7901 H. mirabilis. Crodini E.Fig. 20. MBB7902 H. mirabilis. Crodini E.Fig. 21. MBB7903 H. mirabilis. Crodini E.Fig. 22. MBB7904 H. mirabilis. Crodini E.Fig. 23. MBB7905 H. mirabilis. Crodini E.Fig. 24. MBB7906 H. mirabilis. Crodini E.Fig. 25. MBB7907 H. mirabilis. Crodini E.Fig. 26. MBB7908 H. mirabilis. Sandkraal W.Fig. 27. MBB7909 H. mirabilis. Sandkraal W.Fig. 28. MBB7910 H. floribunda. Rietkuil.Fig. 29. MBB7912 H. mirabilis. Rietkuil.Fig. 30. MBB7913 H. mirabilis. Rietkuil.
These images can now be compared with MBB6666 at Tradouw Pass to establish if my assessment is reasonable. My contention is that this MBB7899 population is midway between the other three Heuningklip populations I am about to illustrate and Tradouw Pass. This similarity is repeated widely and eventually contributes to my contention that H. mutica has been generated by drift away from its two contributing parent species viz. H. mirabilis and H. retusa.
2. MBB6666 H. mirabilis. Tradouw Pass See figs 31-71. This population is at the mouth of the Tradouw Pass and relatively low-lying in respect of the previous population. The habitat is quite different being partially coarse river gravels, weathered shale and at one point is an un-weathered shale ridge where the plants are wedged in the cracks. The habitat is of course very different to the steep sandstone cliffs within the pass itself where H. retusa ‘turgida’ occurs. I initially thought these plants fitted with H. magnifica. A decision derived from a very small sample available at the time. Now I looked for the images and photographs in my records under H. mirabilis where they should have been since my inclusion there of ‘magnifica’, and also under H. retusa where I found them in the folder ‘nigra’. The flowering time is late summer. In this respect similar to the previous set MBB7899 H. mirabilis but the plants are more robust. Figs 31-45 were taken in 2009; figs 46-53 in 2011 and figs 54-71 are from seedlings grown by myself. I tried to reduce the pictures and eliminated a few. But I really think they all need to be presented to show again how serious the difficulty of circumscribing them as a single recognizable key-able entity really is. While looking at my pictures I seriously thought that the set taken in 2011 differed from those taken in 2009. The cultivated plants of course are not directly connected to their source and the variation is self-evidently wide. But it is equally so in field plants. There are the smoother surfaces of H. retusa and some very rough surfaces for H. mirabilis. My decision became that they fitted best with the ‘nigra’ populations where my organizing had taken them despite flowering time, but here MBB7899 shows absolute geographic position between H. mirabilis and MBB6666. I consider this key evidence for the basic continuity for H. retusa and H. mirabilis that I have conjectured elsewhere and I would not be able to defend an argument that MBB7889 plants belong better in H. mirabilis and this means MBB6666 belongs there too. I would simply dispute that there is no better solution so stick with history as far as possible and that means H. mirabilis. If Breuer or Hayashi have a name for a variant from this population intended as an aggregate name, I recommend the use in inverted commas. I would actually like the use of the equilibrium diacritic of a double arrow pointing between two names viz. H. mirabilis↔H. retusa ‘nigra’, although this available symbol is not quite the thing.
Fig. 31 6666 H. retusa ‘nigra’
I come to this vexing question of “typical” that is bandied about even by myself when I liken population to population. There is the same range of variables in MBB6666 noted for MBB7899, but the plants do not darken as much and they are larger. So we need now to look at the other three Heuningklip populations asking what significance is attached to their presence in exactly the same habitat as MBB7899, but geographically separated and by perhaps 500m and 2km respectively. The plants are so variable that it is not possible to arrive at a characterization that separates any two plants, or any two populations in any geographic set for that matter.
3. MBB7896 H. retusa ‘nigra’, Heuningklip 1 See figs 72-89. The plants in this and the next two populations were flowering in September as does H. retusa and they are relatively large, green in colour and generally smooth. Of course both vary enormously due to both intrinsic genetic difference and extrinsic local situation. One plant within an older decaying grass tuft was nearly 200mm diameter, and a healthy green colour, another plant in the open white clay was less than 50mm diam. and dark in colouration.
Fig. 72 7896 H. retusa ‘nigra’
4. MBB7897 H. retusa ‘nigra’, Heuningklip 2 See figs 90-105. The first two pictures show the general landscape and the local habitat where the plants occur. The clay is highly erodible and often there are no plants at all in the clay. Some of the plants have very acuminate pointed leaves while in others they may be relatively obtuse and rounded.
Fig. 90 The countryside looking towards the Tradouw Pass.Fig. 91 The ‘white clay pressure burst’
5. MBB7898 H. retusa ‘nigra’, Heuningklip 3 See figs 106 -132. The great variability is again evident and in this population the plants were observed to be generally without much venation or lines as in fig.112. Then one plant, fig. 119, was so striped as to stand out from all the populations seen.
Fig. 106 Silcrete hilltopsFig. 107 7898 H. retusa ‘nigra’.
Set 2 – THE BUFFELJAGS AREA (H. mutica) This second section follows a recent manuscript submitted to Alsterworthia yet to be published entitled “Some last closing thoughts” written in the hope of finding the closure I seek and that will not come. That article is about a population that has been named as a species viz H. groenewaldii, in an article authored by Breuer, Marx and Groenewald (Alsterworthia 11.2:15) that I only saw after the submission of my manuscript. I also only saw the article after the exploration I would have asked for to verify opinions. The description seems to have been prompted by Marx who refers to similarities to H. springbokvlakensis and to H. magnifia var. atrofusca and raises many issues that my manuscript does not cover. I simply find the whole discussion unrealistic and especially so when no further exploration was done to examine anything. One should refer to Breuer’s World of Haworthias Vol 1 to see that the naive promises in the foreword and introduction of exploration and field study, was never achieved. It is true that the plants have an unusual surface texture and it is also true that the habitat is quite unlike any other H. mutica occupies. But it is Marx’s comment that disturbs me viz. “Unfortunately Bruce Bayer keeps on refusing to see it as something new and calls it simply H. mutica ‘Buffeljags”. This is a distortion and misrepresentation, and whose misfortune this is, not specified. My reticence is reflected in the preceding discussion and Gerhard does not seem to follow, or agree with, what I have been writing about and trying to explain in this general context for so many years. In fact in an unpublished (to my knowledge) manuscript he dismisses all my writing as “pretentious” that no doubt it is, because I have never fancied my self as a taxonomist in the tradition of taxonomists. I was as fascinated as Marx by the plants and am even more anxious than he is to see just how they connect to the probable geographic counterparts in the context of the species definition that I apply. I was also very curious to know what the connection to H. mutica and the H. retusa ‘nigra’ issue, only now laid to rest in the preceding discussion, actually might be here. This is why I use the word “instigated” in reference to exploration in the very valley where H. mutica ‘groenewaldii’ occurs and to the obvious habitat for similar plants. It is surely unfair iniquitous to so demand that I see this population as “something new” in an article that rather disregards the history of the subject and what I have been trying to understand of it.
I also recently submitted a manuscript to Cactus and Succulent Journal (US) where I wrote of “H. groenewaldii Breuer”… “I consider it to be generated from the interaction of H. mirabilis, H. mutica and H. floribunda. I attach no special importance to the fact that it flowers, contrary to H. mutica, in February/March. This is because I have observed many hybrids between patently different species despite a seasonal difference in flowering time”. I have in many places discussed the problem of “hybridization” and it must surely be well-known from my writing that H. floribunda merges into H. mirabilis. Less well-known will be the fact that evidence for H. variegata doing the same is also available. In respect of floral characters, I wrote a chapter in Haworthia Update Vol.3.2:88 explaining the problems of using floral morphology for species recognition. I would challenge Breuer et al. to produce comparative results from studies of the flowers in any of these problematic species to demonstrate difference. Regarding the curious shiny leaf surface, I refer to Haworthia Update Vol.2.2:141, Chapter 15 where I discuss electron scanning photographs of leaf surfaces. This also demonstrates how problematic leaf surface is. It would be tiresome indeed to parade all the evidence in support of these contentions of mine. In Haworthia Update 5.2:108 I also briefly discussed H. mutica from Buffeljags (‘groenewaldii’) and submitted a large number of photographs explaining the habitat and what the possible connection was to the H. retusa ‘nigra’ conundrum. I may also have written about the issue in Haworthiad. Suffice to say that the authors of H. mutica ‘groenewaldii have certainly not checked or considered “all the facts” as the principal author claims. The very opposite seems to be the case.
Considering my oft repeated observation that geological substrate plays a major role in shaping species appearances, the Breuer, Marx and Groenewald description of the habitat for H. mutica ‘groenewaldii’ is a little misleading (see considerations above). Nowhere else is H. mutica on identical substrate and this would just support my contention of ecotypic differentiation that does not justify a Latin binomial. There is considerable doubt about the actual difference between these river boulder situations and the widely distributed terrace gravels elsewhere in the area, notably the Bontebok Park near where the following populations were found:
6. MBB7888 H. mutica, Rotterdam 1 See figs 133-169.
Fig. 133 7888 H. mutica.
7. MBB7889 H. mutica, Rotterdam 2 See figs 170-188.
Fig. 170 7889 H. mutica.Fig. 185
8. MBB7890 H. mutica, Rotterdam 3 See figs 189-191 The three populations above are separated spatially by several hundred meters and they are all on remnant river gravels on a more gently sloping bank than the Buffeljags east bank. The vegetation is closer to conventional fynbos as in the adjoining Bontebok Park, but grassy. The pictures are self-explanatory and the leaf surfaces of the plants are also generally as for the Buffeljags population. What is significant is that I cannot say I saw a single plant in these populations that resembles the one chosen and pictured as typical for this highly dubious species, H. groenewaldii. It makes more sense to me to identify plants from these populations by collecting number than to generate a supposedly descriptive Latin epithet from an imagined typical representative.
Fig. 189 7890 H. mutica.
9. MBB7801 H. mutica ‘groenewaldii’, Mullershof, Buffeljags See figs 192-215 The following set of photographs has surely seen the light of day somewhere and I repeat them because as far as I am concerned they are H. mutica with the shiny surface claimed for “groenewaldii’. Dankbaar is some 16km west of these Buffeljags populations and the substrate is Table Mountain Sandstone. The flowering time is just normal for H. mutica. Fig. 215 is a photograph of a plant apparently collected by Jannie Groenewald and it is quite uncharacteristic of the population and would suggest that it is a product of hybridization with H. mirabilis in the vicinity (e.g. Klipbult).
Fig. 192 7801 H. mutica ‘groenewaldii’.Fig. 215 7801 H. mutica ‘groenewaldii’.
10. MBB7741 H. mutica, DankbaarSee figs 216-235. My observations in respect of these two sets of plants confirms my widely written convictions about Haworthia taxonomy and the state of classification in the absence of a definition for the concept of “species”, and the intrinsic value of Latin binomials for rational and sensible communication. Of course this Dankbaar population is not identical to other H. mutica populations and I am not driven by fixation on any one character that defines this species. It is a geographic set and the various sets are not simply separable. But I am not finished yet.
Fig. 216 7741 H. mutica.
Set 3 – THE BUFFELJAGS AREA (Robustipedunculares) The western Buffeljags area also produced a population of H. marginata and one of a very curious H. minima, that both complement occurrences in the Bontebok Park a little further west. The conventional small, blue-green H. minima is present in the park as well as an unusually green population.
11. MBB7892 H. marginata, Rotterdam See figs 236-243. This population was quite exceptional and one of the strongest populations known to me. But the plants were in flower in September whereas the flowering time for the species is normally late summer; and the flowers and seed capsules exceptionally large. I do not consider that these factors command another Latin binomial.
12. MBB7891 H. minima, Rotterdam See figs 244-252 and Bontebok Park figs 253-4. These were also extraordinary plants, larger than those seen in the Bontebok Park population where the plants were also green viz. figs 253-4. Fig. 252 confuses the issue because, while the other plants seemed alike to H. marginata and distant hybrids, this one plant appears to be a recent hybrid. This occurrence cannot be overlooked in respect of other introgressive (or seemingly introgressive) populations of Robustipedunculares species. The plants were not in flower and again we have a suggestion or direct evidence of hybridization across a difference of seasonal flowering.
Fig. 244 7891 H. minima.Fig. 254 sn H. minima, Bontebok Park
Set 4 – THE GREATER BUFFELJAGS AREA (H. mirabilis) This is a set of 15 populations of H. mirabilis that occur east, west, south north of the H. mutica ‘groenewaldi’ related populations. I exclude from the discussion and illustrations the several populations of H. floribunda ‘major’ populations to the northwest, while I repeat the statement that H. floribunda contributes to the general gene pool of H. mirabilis and together with H. retusa, they all contribute to the gene pool of H. mutica ‘groenewaldii’. The huge presence of populations more readily assignable to H. mirabilis (sensu lato – in the broadest sense) is used by me to validate my opinion that the “mutica” elements draw most genetic resource from the H. retusa element, and hence why I refer those plants to H. mutica.
The sequence of populations 7901-7908 is located in a horseshoe of the Breede River, southwest of Buffeljags in very promising habitat. Only H. mirabilis was seen. I perhaps need to point out that these populations would have been extraordinarily difficult to name under any previous dispensation of mine where I was still trying to uphold H. magnifica,H. maraisii and H. heidelbergensis as species. These I have since merged in H. mirabilis and I reject the introduction of a group name like “aggregate” as Breuer seems to have done, as a complete fabrication that does nothing to solve the problem other than to highlight it.
The following 6 populations are all associated with tertiary and river gravels. There are differences between most of the populations but of an order that I cannot articulate. It is also not possible to really encapsulate the appearances of the plants in the two photographs for each population that I submit. At the same time it does not seem to me possible to make any communicable distinctions.
13. MBB6644 = 6860 H. mirabilis, SSW Swellendam. See figs 255-256.
14. MBB7704 H. mirabilis, Bontebok ParkSee figs 257-258.
15. MBB7805 H. mirabilis, Bontebok Park See figs 259-260.
16. MBB7823 H. mirabilis, KlipbultSee figs 261-262.
On selecting these two pictures I was reminded of my observation that there was a significant H. floribunda appearance to the plants. I had believed that I had seen H. floribunda in the Bontebok Park nearby many years ago but subsequent exploration has only produced H. mirabilis albeit with some similarities to H. floribunda. My reaction was to suspect that H. mutica ‘groenwaldii’ most probably also then needed to be considered as a product influenced by H. floribunda.
17. MBB7887 H. mirabilis, RotterdamSee figs 263-264.
18. MBB7876 H. mirabilis, DisselfonteinSee figs 265-266. These plants were a little more robust than the others in this set and rather similar to MBB7901.
The following populations are almost entirely in Bokkeveld shale but on a very close interface with tertiary gravels. In some places there was also some wind-blown sand.
19. MBB7901 H. mirabilis, Crodini 344, S Swellendam See figs 267-278.
20. MBB7908 H. mirabilis, Sandkraal, S SwellendamSee figs 279-282.
Set 5 – BACK TO TRADOUW PASS – AS IT HAPPENED In the process of compiling this manuscript I was bothered by the ever present question of what is still out there that might influence the outcome. My wife, Daphne, was planning an outing to see chameleons at Swellendam while simultaneously contacting landowners for me around the Potberg. Always anxious to get two birds with one stone, I asked her to find out who owned the land west of Heuningklip that we had just explored with Jannie Groenewald. The result was that we ended up spending a day with Odette Curtis of Custodians of Rare and Endangered Wildlife, south of Suurbraak and finding three populations that, for me, put the proverbial cherry on the pie.
All three were on the slopes of silcrete inselbergs and almost the last of this habitat westward from Heidelberg to Buffeljags. The first inselberg was different to the second and from the Heuningklip habitats because it was dominated by Aloe arborescens and A. ferox. Aloe brevifolia was also present in an unusually small form. The habitat was very impacted by grazing animals and there is an outside possibility that H. retusa ‘nigra’ may once have been present although I personally doubt it. To our surprise we found…
21 MBB7910 H. floribunda, Rietkuil, Suurbraak See figs 283-288.
There were many plants but severely trampled and disturbed. Their survival was obviously aided by the fact that much of the plant is below soil-level. But my experience of this species is that it struggles to maintain its identity in the presence of either H. mirabilis or H. retusa. They never share immediate habitat. There was no hybrid present and this also adds to the problem of interpreting flowering time and distance apart from other species as part of the process. The plants were not the large green variant of the species present to the west at Appelbos, Swellendam; and the leaves were both with and without marginal spines. Eastwards I know the species to be close only at Blackdown north of Heidelberg and at Goedverwagting south of the N highway about 15km east southeast. It definitely seems to be absorbed into H. mirabilis west and north respectively of those two populations. So while this is a slight contradictory element of surprise, it is the next two that really excited me. These were again inselberg habitat but without the strong silcrete upper layer of the former habitat and without the Aloe element. The second ‘population’ was about 200m from the first and comprised four groups of plants along a longer north-facing ridge.
22. MBB7912 H. mirabilis, Rietkuil, Suurbraak See figs 289-306.
23. MBB7913 H. mirabilis, Rietkuil, SuurbraakSee figs 307-331 I am very hard-pressed to say much more than that these plants met all my expectation for the very variable set of plants and populations that I ascribe to the one single species H. mirabilis. Who could possibly articulate the differences of the many populations of this species from each other and from this one? Variable becomes a done-to-death word. The individual plants, in two populations quite unlike any others I have seen, were generally robust with large specimens up to 100mm. diam. The colour is dark, although green when shaded within or under grass. The leaves were usually sharply pointed but there was always evidence of one or two leaves or odd individuals in which the leaves were muticate i.e. without points. In some plants the leaves were notably lined in others not. The colour also varied from chestnut to deep reddish-brown when the plants were more exposed. The surface texture was generally tubercled but there was evidence of smoothness of the leaves in many plants. Flowering time was obviously late summer gauged by the absence of flower at the time of the visit viz. September. Except for one oddity. This was a single plant with a weak flower spike. I photographed this for the useful information the flower-bud provides. Where Breuer et al. claim to have considered all the facts, this is a character that is conveniently overlooked when claims are made for the significance of floral characters when paraded to substantiate yet another claim to species significant difference. The bud-tip in all the Southern Cape species of subgenus Haworthia with the exception of perhaps H. rossouwii have this very obvious “fish-tail” bud tip. The tips of the upper two petals are flattened and spreading to resemble a fish’s tail. It is most marked in H. herbacea and H. reticulata where the bud tip curves downward from the semi-erect position of the body of the flower and then up again. I am fearful that my inability and reluctance to consider these plants as “new” may be the motive for someone to rush forward and apply a Latin binomial.
My opinion is that these two populations mirror the paradox in the east where H. pygmaea ‘fusca’ is nearer to H. retusa ‘retusa’ andwhere H. retusa ‘turgida’ is nearby. H. mirabilis ‘splendens’, in the doubt about its affinities, in my opinion remains in the context of H. mirabilis. This is a conclusion I reached after exploration of the area between Riversdale and Albertinia where the same elements noted in this article repeat themselves. I do not for one moment concede that better explanation is available by resort to a multitude of Latin binomials and by an artifact like “aggregate”.
Conclusion I have been deeply disappointed that so little progress on the classification front has been made among the fraternity writing about Haworthia. In communication with Lawrence Loucka, he wrote …”You might consider saying that a genus revision is needed and that until amateurs depart no proper botanist will make the investment. Until then chaos will rule, names will proliferate.” This is a useful and insightful comment and generally summarizes what I have often said. Unfortunately the position has been rather exacerbated when proper botanists have made their presence known and I can recount many instances to support this statement. My own view has been that botanists have not been able to confront genera that are so publicly and popularly known, and garlanded with names. Thus amateurs, including myself despite at least an iota of formal training in science, simply have a field day generating names like rabbits out of a hat.
Unless some proper thought is given to this whole question of what the names are supposed to mean apart from catalogue items, taxonomy (classification) will never lift out of the disinterest, disrepute and scorn that collectors justifiably seem to have for it. Latin binomials do have a real value and to demean this by one-dimensional derivation and rationalization is intellectually fraudulent. My writing is replete with examples of the variability and interaction of populations over large area. Taking a single population and one flower, and often a single plant, to generate a species description and a Latin binomial is grossly simplistic.
The one change that this article generates is the confirmation that the population MBB6666 from immediately south of Tradouw Pass is better identified by name as an H. mirabilis variant, and not as H. retusa ‘nigra’. MBB7801 at Buffeljags and now also at Rotterdam, remains as H. mutica and the epithet generated by enthusiasts can be added to make the trinomial H. mutica ‘groenewaldii’. For formality, the name for the form illustrated as typical would be H. mutica ‘Groenewaldii’. A few additional names will be needed for similar clones and a lot more for the cultivars that will be derived from this population set.
My conclusions here make no substantial departure from anything that I have observed or written over 50 years. While the subject has been comprehensively covered in the field and in writing, there is a huge amount of exploration that still needs to be done if the genus is to be still better known. My prediction is that the answer will conform very closely to the species-list I finalized in 2009 (see Alsterworthia 5.1:192). The disclaimer is that botanical classification may eventually reach the point where the real nature of species and their purpose comes to be known. In that case what we now may see and persuade ourselves are discrete species, are not. Instead these highly complex chaotic systems may in fact be quantum units of consciousness and whole elements such as some genera and sub-genera we now recognize will be the species.
Acknowledgement I want particularly to acknowledge Jannie Groenewald whose interest and enthusiasm are boundless. Then Allan Jeptha, an environmental law practitioner with great vision for the Suurbraak community; Odette Curtiss, a consultant for the conservation of Renosterveld vegetation; Lawrence Loucka whose patient guidance and comments I value and who generated the set of pie charts; Kobus Venter and Steven Hammer who have always been supportive and contributory; Max Coetzee of Felix Unite and Samie Lategan of Buffalo Breeding Programme for access to Sandkraal West and Crodini East; Jaap Viljoen for his contribution; my wife Daphne, who trusts me and whom I deeply trust. ♦
The objective was to explore some likely habitats previously observed at Van Reenens Crest and nearby. We extended the scope to include further exploration for Haworthia mutica as I am still questioning the place of this species in the greater scheme of things. Thus here are four sets of populations that I report on viz. H. mirabilis, H. retusa ‘nigra’, H. floribunda and H. mutica. See maps Figs 1 and 2 for geographical position.
Van Reenens CrestFig. 1 Van Reenens Crest. M = H. mirabilis, RN = H. retusa ‘nigra’, F. = H . floribunda.Fig. 2 Known distribution of H. mutica
H. mirabilis
Set 1. MBB7955 Van Reenens Crest – mid west. (figs 3-11)
The plants are moderate in size to about 50mm diam. and growing on an erosional slope below tertiary gravels where there is white clay. They vary from densely tubercled to relatively smooth.
Fig. 3 7955 Van Reenens Crest
Set 2. MBB7956 Van Reenens Crest – west 7955. (figs 12-34)
Despite apparently favorable habitat these plants were about 300m west of the previous set. Variation is again apparent in respect of presence or absence of surface roughness. Generally there are plants in which some leaves are without a point (see Fig 1) while others are acute tipped (mucronate – with a short hard point) or even have a long end-awn. Margins may be smooth or very lightly spined. This seems to be more apparent in young plants.
Fig. 12 View looking east to Set 1.Fig. 13 7956 Van Reenens Crest
Set 3. MBB7957 Van Reenens Crest – west 7956. (figs 35-41)
Again the plants are very variable with dense surface tubercles to virtually none. The leaf shapes vary and in fig.39 that appears to be a group of seedlings there is a remarkable series of different leaf ends and margins. In some plants there is distinct spination of the leaf margin.
Fig. 35 View looking south to Tradouw PassFig. 36 7957 Van Reenens Crest
Set 4. MBB7958 Van Reenens Crest – south west 7957. (figs 42-51)
Fig. 43 shows two rosettes of a plant with a “crocodile-skin” surface. The surface tubercles seem to be flattened and the leaves are mucronate. The following photograph, fig.44, is of a plant with acuminate leaves ending in long end-awns, and the surfaces are relatively smooth – retusoid? Unlike the previous records for the area that are north-facing, this is a south-facing “pressure-burst” of white clay. A nearby habitat of vertical shale that might have been considered favorable for H. mirabilis elsewhere was devoid of plants.
Fig. 42 7958 Van Reenens Crest
Set 5. MBB7959 Van Reenens Crest – west homestead 7956. (figs 52-64)
Figs 53-55 are smallish plants with rounded leaf ends and relatively opaque surfaces. Fig.58 is remarkably like H. retusa, say, at Pienaarsrivier with very awned leaf ends and a high degree of translucence in the leaves.
Fig. 52 MBB7958 View to MBB7957 a. Daphne inspecting, Lawrence checking the GPS.Fig. 53 7959 Van Reenens Crest
Set 6. MBB7960 Kruiskloof, E Van Reenens Crest. (figs 65–78)
This population departs from a very broad norm for Van Reenens Crest and picks up the continuity with populations north, east and south. This is most noticeable in the surface texture where the tubercles are smaller and denser. The leaves can be highly rotate, see fig.68. A feature seen occasionally in the Van Reenens Crest populations and as far afield as the Diepkloof (Malgas) area much further south, is the occasional plant with a very stellate (five –pointed star) arrangement of the leaves that are quite horizontal see fig.70. Among the many oddities is the plant in fig.72 with floribundoid leaf ends and a leaf surface where the tubercles seem to be flattened. In fig.73 the tubercles seem to be spined. In fig.75 tubercles are almost absent.
Fig. 65 7960 Kruiskloof
H. retusa ‘nigra’
Set 7. MBB7961 Kruiskloof, E Van Reenens Crest. (figs 79-90)
These plants reinforce my perception that H. retusa and H. mirabilis are uncomfortably closely related. Figs 79 and 80 have the same general form as H. mirabilis ‘sublineata’. Fig.82 appears to me as possibly a hybrid as leaf opacity is deeper and there are marginal spines.
Fig. 79 7961 KruiskloofFig. 82
H. floribunda
Set 8. MBB7962 Niekerkshek. (figs 91-99)
Fig. 91 7962 Niekerkshek
Set 9. MBB7963 Niekerkshek. (figs 100-111)
The plants vary in size and to some degree in colour. This variation in colour from a deep purple to dark grey is very evident in fig.107. The leaves are much flattened and an occasional leaf is pointed. The plant in fig.102 is unusual in having white spots on the leaves. Marginal spination can be absent or very coarse. The leaf surfaces show no venation or translucens, as is expected in this species.
Fig.100 View looking south to Potberg.Fig. 101 7963 Niekerkshek
H. mutica
Set 10. MBB7950 DeDraai Klipbankskloof. (figs 112-123)
Fig. 112 – 7950 Die Draai, E Klipbankskloof
Set 11. MBB7951 Klipbankskloof West 1. (figs 124-126)
Fig. 124 – 7951 Klipbankskloof West 1
Set 12. MBB7952 Klipbankskloof West 2. (figs 127-129)
Fig. 127 – 7952 N Klipbankskloof West 2
Set 13. MBB7953 Klipbankskloof West 3. (figs 130-144)
Fig. 130 – 7953 Klipbankskloof West 3 View westwardsFig. 131 – 7953 N Klipbankskloof westward view of habitatFig. 132- 7953 N Klipbankskloof West 3
Fig. 175 JDV92-64 Very localised.Fig. 176 JDV92-64 Sanddrift
Set 16. JDV85/17 Haarwegskloof. (figs 188-213)
Fig. 188 JDV85-17 Haarwegskloof
There is a very wide range of variants among these 7 population sets. Leaf colour and shape vary enormously and it can only be said that generally the leaves are muticate. Almost invariably any one plant has a leaf or leaves that are also mucronate and in some cases even awned. The leaves may be abbreviated and short and squat or drawn out and elongated. Venation is also varied from many to few or only a single vein on the upper leaf surface. These veins may be straight and clearly separated or they may anastomize and link togther in a reticulate pattern. I did include two pictures of flowers, figs 173 and 174 just to indicate that simply taking two flowers at random one can see obvious differences.
Attention can be paid to figs 145 and 146 of Rondeheuwel where I consider these two plants indicate hybridization with H. mirabilis that does occur about 1km to the northwest. Fig. 147 shows a plant with a rather wrinkled surface.
Conclusion My opinion is that consideration has to be given to the odd way in which the “species” do not share habitat. It is extremely difficult to rationalize an ‘average’ plant for each population and so establish any norm for proper comparison. The populations vary by degree. It is also clear that if one examines the distribution of the species across the study area, it can be observed that H. mirabilis where it occurs with H. retusa ‘nigra’(at Heuningklip and Kruiskloof in the east), is not the same as plants to the west where H. retusa ‘nigra’ is absent. Consideration has to be given to absorption into H. mirabilis. Why H. floribunda should make a re-appearance at Rietkuil and Niekerkshek in the west, when everything points to its absorption into H. mirabilis in the east at Skeiding, is a mystery. The fact that at Niekerkshek it also has radically different forms to those found anywhere else adds to the difficulty.
I will conclude by acknowledging that the populations of H. mutica are all summer flowering but I do not think that this is as significant as the authors of “H. groenewaldii” will contend. The Van Reenens Crest complex demonstrated dramatically that H. retusa and H. mirabilis share a very close common history and an equally close contemporary relationship. My conjecture is that H. mutica is linked to that relationship in the same way the complex of populations are that I assign to H. pygmaea. It is painfully obvious that if floral differences that are so easily detected within populations, expecting to find more evidence in the flowers than exists in the vegetative structures across the genera, and especially the subgenus Haworthia is an absolute pipedream. Floral differences between species are minimal and if anything will provide support for a reduction of species rather than add to the. If any small difference is suggested to separate species, then equal weight must be given to similarities that should then unite species. The contention then that flowers are offering characters that I as an author have ignored, while already wholly untrue, is additionally nonsensical.
Acknowledgement Mr. and Mrs. Trevennan Barry of Van Reenens Crest. Mr. N. Swart of Poststal, Mr. J. duToit of Tarentaal and Wydgelee, Mr. W. duToit of Klipbankskloof, Mr. J. Mathee of Klipbankskloof West, Mr. Vlooi du Toit of Die Kop, Mr. M. Mynhardt of Haarwegskloof, Mr. and Mrs. Nelius Smith of Volmoed, Rondeheuwel, Mr. Adrian Steyn of Wankie Boerdery. Thank you to Kobus Venter and Lawrence Loucka for company and shared observation. ♦
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.
Fig 1. Map of area and points covered
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.
Fig. 2 7500 Die Kop E
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.
Fig. 9 7502 Die Kop W
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.
Fig. 11 7973 Die Kop N
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.
Fig. 28 7503 Die Kop mid
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.
Fig. 36 7974 Die Kop mid
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).
Fig. 47 7504 N Die Kop
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.
Fig. 56 JDV86-2 Haarwegskloof
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.
Fig. 64 7964 Spitzkop West
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.
Fig. 82 7965 Spitzkop West
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.
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.
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.
Fig. 109 7967 Spitzkop (quartz rock)Gasteria carinata is often severely animal grazed.Looking SE towards Die Kop and PotbergAlso looking SE towards Die Kop and PotbergViewing site from the north
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.
Plants absentPebbles and plants presentThis is not how one expects to see this species.Fig. 152 7968 Spitzkop (pebbles)Showing the small shale ridgesLooking to the previous habitat of larger quartz rock
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.
Fig. 160 7969 Spitzkop East
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. ♦
I recently wrote an essay on the situation between Haworthia retusa and Haworthia mirabilis at Komserante east of Riversdale. The essay was entitled “My view of names” and is posted on the HaworthiaUpdates.org web site. Etwin Aslander posted some pictures from what he called Kruisrivier. These caught my eye because they did not look like the plants I know from a place of the same name. My known population is JDV95/62 and generally these plants have the dark colour and rough surface texture of H. mirabilis. The issue is that they are spring flowering whereas H. mirabilis is generally considered and observed to be late summer flowering. Etwin indicated to me where he had found his plants and I duly went to look.
In the process I incidentally called on a well known H. retusa population at the Skietbaan locality south of Riversdale. There has been a dramatic turnabout in the appearances of these plants since I last looked there 2 years ago. Whereas there were then huge clones well above ground level, the plants were now again smaller and drawn into the ground. I experienced this dramatic shift in plant appearances just west of the Frehse Reserve many years ago when there were giant size plants as opposed to my first visit when the plants were really small and withdrawn.
Kobus accompanied Daphne and I to Kruisrivier where the owners Wilhelm and Mandi Zietsman were extremely helpful. They told us also of a neighbor, Gert van Rensburg, who had also seen the same plants on his farm to the west. Mandi accompanied us on a jaunt to find that farmer and failing that we explored north of the original Kruisrivier locality. There we found another population of plants as well as H. floribunda (see Set 1 MBB7998). These two species H. retusa and H. floribunda were occupying different habitat and spaced about 100m apart. The H. floribunda was numerous and rather smooth leaved as well as paler green in colour than I expect from that species. The H. retusa-like plants were much smoother in surface texture than the original known population and they were in bud (see Set 2 MBB7999). We went back to the older population just to confirm that they were in bud too as we expected. Just so and the buds were just emerging from the rosettes. The plants were generally smaller than they were at a previous visit (see set 3 JDV95/62).
We parted company with Mandi Zietsman, and went off westwards intended to explore the Klein Kruisrivier area that seemed to better fit Etwin’s site indicator. By good fortune we ran into Gert van Rensburg of Wegwysersrivier. He eyed us very suspiciously indeed and obviously very reluctant to show anyone the plants. However, he very kindly relented, took us to the spot and left us to freely photograph and explore (see set 4 MBB8000). The plants can be described as midway between the generally rougher surfaces of JDV95-62 and the smooth surfaces of MBB7999. What was more dramatic is that there were six flower spikes so that flowering is possible as early as July 6th.
We returned via another route regretting leaving distant habitat unexplored. But we did find another population of H. floribunda, a little more toothed and perhaps brighter green than at Kruisrivier.
I also note that I long ago confirmed Smith’s record for H. retusa ‘turgida’ at Klein Kruisrivier in the upper Wegwysersrivier Gorge. This is the small spinose proliferous version known elsewhere from the Langeberg Mts.
Digesting this new information is a bit difficult in view of the very opposed views of what names mean and how they should be applied. Taking all the populations that I have explored and written about, my perspective is further to a view expressed in Haworthia Update 7. This is that H. retusa and H. mirabilis are uncomfortably close. The only thing that appears to separate them is the yellowish green and smooth tendency in H. retusa and the darkish green and surface rough tendency in H. mirabilis. Further to that is of course the question of spring flowering versus late summer flowering. But I have already reported several case of hybridization across this divide as well as the Komserante situation. Here we now have plants in three populations that occupy middle ground and one of these populations has a significant degree of a winter flowering capacity. The identification should perhaps utilize the chemical equilibrium symbol. This is not quite it “↔” as the better symbol comprises halved arrows pointing in opposite directions.
I wish to add that in the case of plants I attribute to H. ‘turgida’ at Towerlands, I commented on the very real possibility of a close connection to H. emelyae. There is also evidence for this elsewhere. I use the name ‘turgida’ like this because of the uncertainty of it really being H. retusa var. turgida or perhaps H. pygmaea.
My experience in other situations viz. H. limifolia, H. herbacea/H. reticulata, H. arachnoidea/H. mucronata, H. cymbiformis/H. cooperi, Kiewietsvlakte etc. all suggests to me that the view of species is grossly distorted in the splitter direction. It is clear to me, if to no one else, that H. retusa and H. mirabilis form a very cohesive entity with ramifying oddities the length and breadth of the distribution range. I do not cover this issue here, but there is the added complication of the involvement of H. floribunda. It seems to be very discrete in most places, whereas at others it seems to get lost mainly (only?) in H. mirabilis. This may be because the introgression is favoured by the same flowering season. H. retusa and H. mirabilis are drifted apart by the difference in flowering season but it is by no means anything more than a general observation.
I have added the images of the available flowers as well as that of a bud to show the flared fishtail bud-tip that the southern Cape species tend to have. The flowers are variable and it is difficult to make a statement that characterizes them i.e. no composite image forms.
Acknowledgement I would like to acknowledge Etwin Aslander’s input. Wilhelm and Mandi Zietsman were extremely helpful. Gert van Rensburg was surprisingly well informed about the plants too and very generous in his attitude to us. Part of the pleasure of field work is meeting people like this.
Set 1. MBB7998 H. floribunda Kruisrivier
Set 2. MBB7999 H. retusa Kruisrivier
Set 3. JDV92/65 H. retusa Kruisrivier
Set 4. MBB8000 H. retusa Wegwyserivier
MBB8000 flower faces
MBB8000 flower profiles
MBB8000 flower bud
Addendum To demonstrate the problem of similar looking plants that appear in different populations, I take 3 plants from the original Kruisrivier population (JDV92/65) see figs 1 to 3. Fig. 1 is obviously a mirabiloid plant and if this population flowered in late summer it would probably be identified as H. mirabilis. The figs 2 and 3 are more retusoid. I leave out plants from the newer Kruisrivier population (MBB7999) because none of the plants have the rougher mirabiloid leaf surfaces. I add a Wegwysersrivier (fig. 4 MBB8000) plant that is again mirabiloid and like Fig. 1 except that it appears to be a spring flowering population with a significant number of plants in flower in early July. From there I take a plant from Komserante (MBB7779) that flowers in late summer but is apparently generally hybrid with H. retusa. Moving eastwards from Riversdale and impinging on H. mirabilis splendens, I show a plant MBB7762 from Platkop (fig. 6) where both H. mirabilis and H. retusa occur with occasional hybrids. Fig. 7 is MBB7818 H. mirabilis Windsor SE Riversdale, where the plants frequently have a frosted appearance because of minute surface spines.
There is a significant geographic jump with fig. 8 MBB7850 H. emelyae north of the Langeberg at Aasvoelvallei. This is a population that I have noted elsewhere that highlights the probable relationship of H. emelyae with the H. retusa turgida and pymaeaoid elements from Herbertsdale eastwards. Fig. 9 is a plant of MBB6666 Tradouw Pass that I recognize as a hybrid population H. mirabilisXretusa. Inland from there are several populations, MBB7899 is H. mirabilis, Heuningklip (fig. 10) and MBB7896 H. retusa nigra also Heuningklip (fig. 11). East of that are three populations of H. mirabilis, MBB7912 and MBB7913 Rietkuil and MBB7919 Van Reenens Crest (figs 12 to 14).
As only single plant comparisons, it seems fairly safe to say that, bar flowering time and figs 2 and 3, they are all similar. However, the variability in each of these populations is great and this has been reported elsewhere in the Update volumes. If one had to now take figs 2 and 3 and look for similarities in other populations, it would be very easy to demonstrate a complete gradation from what could be construed as typical H. mirabilis through to typical H. retusa through a large array of populations.
Fig. 1. JDV95-62.9 H. retusa, KruisrivierFig. 2. JDV92-65 H. retusa, KruisrivierFig. 3. JDV92-65 H. retusa, KruisrivierFig. 4. MBB8000.5 H. retusa, WegwysersrivierFig. 5. MBB7779 H. mirabilis. KomseranteFig. 6. MBB7762, H. mirabilis, W Platkop 4616Fig. 7. MBB7818, H. retusa, WindsorFig. 8. MBB7850, H. emelyae, AasvoelvalleiFig. 9. MBB6666 H. mirabilisX retusa, TradouwFig. 10. MBB7899 H. mirabilis, HeuningklipFig. 11. MBB7896 H. retusa nigra, HeuningklipFig. 12. MBB7919 H. mirabilis, Van Reenens CrestFig. 13. MBB7912, H. mirabilis, RietkuilFig. 14. MBB7913 H. mirabilis, Rietkuil
It has long been my contention that there is no separation between Haworthia retusa and Haworthia turgida. It is one very variable system viz H. retusa, with a larger fairly non-proliferous plants tending to level areas and then smaller proliferous plants on steeper habitats. There is huge variability among members of any one population and of course much more between populations. Over and above this is the relationship of this apparently one single system, with H. mirabilis that is probably even more complex and varied. If one takes all the known populations and variants into consideration it become necessary to ask if H. retusa and H. mirabilis are also not just elements of one system, and one species. If all the considerations are summed and referral is made to vegetation and speciation drivers; what constitutes an area of endemism, then I am sure the answer will be “Yes”! What seems to have happened is a natural sequence. As sampling has progressed so has there been recognition of differences. The logical outcome is that sampling progression should lead to understanding and synthesis by reduction. Unfortunately there will be diehards that stay with the differences syndrome and cannot see the similarities.
There is surely no longer any doubt that Haworthia classification has been confounded. There are several factors. One is the historical one of sampling and naming. Sampling dates from the seventeenth century and description based on a few words and weak illustration. The second problem has been a nomenclatural system that revolves around the single types and assumes that departures from that still allow comfortable accommodation of all other departures (variants) under the primary name. The third problem is the absence of a species definition deriving from a lack of knowledge and hence understanding of what species are. The fourth problem is enthusiasts, writers, collectors, editors, reader etc. who generate and propagate within the confines of their own needs, limitations, knowledge and understanding. Too often they are not adequately informed to undertake something that really should be the task of professional botanists. Of course it is also true that professionals have not proved to be faultless either, as simply the lack of a species definition alone indicates.
From my personal point of view, I consider the sampling history and the nomenclatural constrictions of priority and automatic creation of type elements, among the main obstacles to a classification solution. Perhaps it is only secondary to the human factor where writers become anxious to establish their own opinions, based on who knows what, crowing from the top of a metaphorical farm dungheap. I have paid a lot of attention to an element that I name as H. retusa ‘nigra’. It is based on a very unrepresentative specimen from Kransriviermond where it seems as if it is a product of hybridization between H. retusa and H. mirabilis. But this situation, and the name, is inextricably involved in a series of populations grouped in areas like Van Reenens Crest (Swellendam), Klip River (W Heidelberg) and Kiewietsvlakte (W Riversdale). These populations do not seem to figure in any earlier exploration and if looked at objectively may now perhaps be seen as the mother lode from which H. retusa, H. mirabilis, H. pygmaea, H. mutica and even H. emelyae as species may emerge. The reality is that all these may not in fact even be discrete species.
In the forerunning article I discussed some new populations at Kruisrivier northeast of Riversdale and where they occur along the Kruis River. This river partially follows the interface of the Cape Sandstone Fold Mountains in an east/west direction and the situation is replicated to some measure elsewhere between Worcester and George. Most relevant is the similarity of the Kruis River valley and its south banks to the situation along the Klipriver west of Heidelberg. But it is not in the scope of this discussion to cover that now and I would in any case need to research the geology of both areas to do so. I will just present images to cover firstly the area, then to allow creation of a composite image of what the small sandstone H. retusa looks like, then images from just two populations of H. retusa to show how shadowy our image of this species is; and then I will show the images of plants from five points down a distance of about 4km along the south bank of the Kruis River to a point south and west of the better known Kruisrivier plants.
Setting the scene. The Kruis River Valley is slightly east of north from Riversdale and the Kruis River is a headwater of the Goukou River that enters the sea at Stilbay. The views I have (figs. 1-4) are of a satellite image of the Valley from where the Kruis River exits the Langeberg Range to where it joins the Goukou. Three images are from the south bank looking northwest and northeast. The area is very heavily impacted by alien vegetation. The grazing animals are mainly cattle (beed and dairy) and hoof impact on the slopes is severe. The underlying shale along the south bank is not exposed and there is both boulder deposit, river alluvium and even some marine terrace in evidence. Nevertheless the area is not so dramatically varied geologically as is the area around Heidelberg to the west. Hence the transitions and interactions between species and their variants are both similar but very different. The initial conditions will be very different too.
Fig. 1 An overview of the Kruis River Valley.
Fig. 2 Some indication of the topography of the south bank.Fig. 3 View northwest from a central position.Fig. 4 View to the northwest across the flood plain.
The populations. The first three sets are to briefly review H. retusa. The general perception of H. retusa is very restrictive and similarly the case with what was H. turgida. I consider them together. Nevertheless I have to sympathize with the splitter approach because I have to make a concession for the sake of discussion. Thus…
Set 1 MBB7895 H. retusa ‘caespitosa’, Diepkloof, Heidelberg (figs 5-9). I explored the upper Kruis River in early 1970 and found the small sandstone version of what was then recognised as H. caespitosa. Later seen as H. turgida, it is known at many places from the Tradouw Pass, north of Heidelberg, Garcia Pass and Kok’s peak. Small and very proliferous, its leaves are also quite spinose on margins and keel and also flecked with reticulate and longitudinal translucence. The summer coloration tends to reds and yellows. I do not have images of this actual form but show five from a population south of Heidelberg that is very similar. But it should not be assumed that the transitional flow in the Heideberg area equates that at Kruis River. The variants down the length of the Klip and Duiwenhoks Rivers is another profound story.
Set 2 MBB7758 H. retusa, Skietbaan, S Riversdale (figs 10-19 ). I show images of this species very recently taken. This is because I do not think that as a species H. retusa is properly appreciated for all its variants. It is this reality that we are confronted with when considering populations as at Komserante and now Kruisrivier where there is an evidence that it cannot any longer be seen as completely removed from H. mirabilis.
If images 10a and b appear to be different from the rest of this wet, they should. They are plants from the same locality but they were taken 4 years ago when most of the plants were very much larger. They illustrate a very useful point – the plants do NOT necessarily look the same all of the time.
Set 3 MBB8010 H. retusa, 8km S Riversdale (figs 20-28 ). Primarily one should consider surface roughness and colouration as main separators from H. mirabilis. Flowering time is the prime differentiator and this is what is now to be seen in another light.
Set 4 MBB8004 H. retusa↔mirabilis, Bloekombos, Kruisriver (figs 29-42). Reference can perhaps be made to the three sets illustrated in the preceding article. Here is will just re-capitulate to say that there is some doubt in respect of plant appearance and colouration, to examine the view that the represent H. retusa or H. mirabilis. (I dismiss totally and absolutely the notion that a third species needs to be fabricated to accommodate differences impossible to itemize or catalogue across the range of variants within and between populations.) I do not think that the similarity of surface roughness to that of the H. mirabilis ‘atrofuscoids’ should be overlooked – even some of the leaves show the tendency to roundness at the ends. This particular population was in full flower at the end of July.
Subset 4 The flowers (figs 43-45). I do not see anything to distinguish these flowers from those photographed for the Wegwysersrivier population and given in the preceding article. It is difficult to imagine that the flowers may offer any distinctive features in the direction of difference that are not already offered by vegetative and geographic considerations. There may of course be direction towards similarity.
Set 4 H. retusa-mirabilis, MBB 8004 Bloekombos, Kruisriver – Subset 4a Flower profiles
Set 4 H. retusa-mirabilis, MBB8004 Bloekombos, Kruisriver – 45. Subset 4b Flower faces
Set 4 H. retusa-mirabilis, MBB8004 Bloekombos, Kruisriver – 45. Subset 4c
Set 5 MBB8003 H. retusa↔mirabilis, W Bloekombos, Kruisriver (figs 46-73). Curiously this population was only in early bud although only a few hundred meters away from the preceding. I do think there is significant difference in the appearances of the plants with more translucence of narrower leaves. However, plants from this and the preceding population could very easily be said to identify with those in any of the H. mirabilis populations southeast, south and west of Riversdale. It is also obvious that there is a degree of similarity to the rougher plants of H. retusa illustrated in sets 2 and 3, and also to the Komserante H. retusa↔mirabilis plants.
Set 6 MBB8007 H. retusa↔mirabilis, 600m W Bloekombos, Kruisriver (figs 74-81). The banks west of set 5 are less steep and the terrain not clearly differentiated into discrete habitats. Heavy cattle trampling would severely impact on plants. Nevertheless we did find a small colony and are confident more would be found if more time was spent searching. The area is quite extensive and we did not consider several north/south valleys that could well harbor plants. The same is true for quite large hinterland to the Kruis River Valley and Etwin Aslander’s plants, similar to the ones we saw, are surely present back there. This small colony reflects further transition to H. retusa ‘turgida’. The leaves are smoother and more translucent. There are some floral differences muted by the small sample and by the fact that these features such as the absence of frilled base to the bract and the presence of a short pedicel, are variations observed within populations elsewhere.
Set 7 MBB8008 H. retusa↔mirabilis, 1km W Bloekombos, Kruisriver (figs 82-83). Also barely, registering as a population, these odd plants support the predicted presence of a direct transition from the Kruisriver mirabiloid-like H. retusa to the known sandstone forms of H. retusa ‘turgida’. Hoof action has severely impacted this area and it sems very probable that there was a larger population at some time or another. The slope militates against the solitary ground-level growth form of the more usual H. retusa but not steep or rocky enough to be home to the turgidoid cliff growing forms.
Set 8 MBB8005 H. retusa ‘turgida’, S Wegwysersrivier (figs 84-98). The use of the equilibrium sign for the preceding is more informative than accurate. It reflects the problem of decision making where it is not clear if H. mirabilis is even in the area. This of course impacts on the situation further east when the reality of H. pygmaea and all its associated variants are brought into the picture. I am sure it is inarguable that we are now confronted with plants that are truly similar to H. retusa ‘turgida’. Curiously the flower is much the same as in MBB8003 and 8004 to the east but there is a difference in that the peduncles were softer and shorter than of plants to the east and suggesting the shorter lax habit of the cliff dwellers. The habitat is unlike the preceding in that the Tertiary deposit is more alluvial and the rock is more boulder-like. Trampling is still a severe problem. Note the two pictures of a H. retusaXfloribunda hybrid.
Set 9 MBB8006 H. floribunda, S Wegwysersrivier (figs 99-107). The presence of this species is no surprise. It was quite abundant although slightly separated from where we found the H. retusa ‘turgida’. It depends on rocks for protection from trampling. This species of course generally flowers in late summer but once again here is a hybrid present with the other parent flowering a good 6 months later. This has been observed in several other places where these same two species occur in close association. [photo file names are mistakenly named MBB8005, they are in fact MBB8006 – ed.]
Conclusion The product of this exploration was based on prediction and I would wish that cognizance was taken of the fact that this could and can be done. Using an aggregate or group category and a host of Latin binomials is not botany and obfuscates things. Perhaps to the benefit of traders, but certainly confusing to the collector who make wish to know more about plants than simply having a label.
Acknowledgement I have already acknowledged the help of Wilhelm and Mandi Zietsman; and Gert and Lynette van Rensburg. We were now also assisted by Wessie Wessels, a retired airline pilot who owns the Bloekomsbos Farm. Kobus Venter came to see my pictures and I really value his interest and useful comment. I wish there was more similar interested and informed comment. ♦
M. B. Bayer (MSc), Kleinbegin Farm, Kuilsriver, South Africa Mrs. Carly Cowell (MSc), Regional Ecologist, Cape Research Centre, Conservation Services, South African National Parks, Cape Town, South Africa.
Objective: The significance of the Bontebo National Park occurrences. The occurrence of members of three aloid “genera” (the three sub-genera of the genus Haworthia could indeed be genera) and the absence of any other member of the Aloids (bar the ubiquitous Aloe ferox) must surely be indicative of the driving forces that determine the flora of the Bonte National Park. This also must surely help establish the significance of the park as a conservancy of considerable merit. The complex interaction of the species enhances even that. The purpose of this report is to examine more closely the variation and nature of a small segment of the Park flora, and demonstrate how much more can be done.
Note: This report has several constraints. Firstly is the situation in which there is no formal general definition and hence understanding of what a plant species is. Secondly there is the generally understood view that there is an evolutionary process at work by which organisms evolve from a common distant origin by genetic mutation and adaptation. Thirdly there are serious flaws in the classification of the Aloid genera. Several essays dealing with these issues by DNA sequencing are weak because they rest on those flaws and consequently do not address some serious questions of relationships that the results pose. Fourthly of course is the reality that the knowledge or intellectual capacity to overcome these deficiencies may be absent. Thus the report is written in the context of all the publications as the original genus revision (Haworthia Revisited, Bayer; Umdaus, 1999) and others available on the internet (HaworthiaUpdates.org).
Introducing the subgenera. There are 6 species of Haworthia that occur in the Bontebok National Park representing each of the three subgenera. These subgenera are not real and have no closer relationship than that between any of the other genera in the Asphodelaceae however these are, or have been recognised.
Subgenus Hexangulares. There are 15 species in this subgenus. Two are in the northwest of South Africa and the others in the Eastern Cape or across the Karoo. In respect of other Aloid genera it seems to be close to Chortolirion. Only one species occurs in the Southern cape viz.H. venosa.
Subgenus Haworthia. Here there are 37 species by my account. Non-botanists raise this figure to as much as 600. By their methodology and ideology it is a great deal more. This subgenus is at odds with all the other Aloid genera and has an unusual internal diversity suggesting very recent speciation. Three species occur in the Park viz. H. retusa (turgida), H. mirabilis and H. floribunda.
Subgenus Robustipedunculares. There are 4 species all in the Southern cape with spill-over into the Little Karoo. One species, H. pumila also occurs in the southern Great Karoo. There is a very close affinity with the genus Astroloba. The four species have a very close affinity and while H. kingiana in the eastern Southern Cape appears to be discrete, the other three definitely are not. The Park occurrences are evidence of this and these seem to represent two species viz. H. marginata and H. minima. See more under the treatment of H. minima below.
Map 1. Bontebok Park
Introducing the species. Illustrations are of a general form for the species. Clonal variability is so high that use of the term “typical” can be misleading. Haworthia venosa.(Fig. 1)
This species occurs primarily along the lower course of the Breede River with its upper river limit just beyond the northwestern corner of the Bontebok National Park. Its lower limit is just north of Malgas. Two populations are known to me off the river and a small way to the west. The species also occurs at Die Eiland in the Gouritz Valley. Its nearest affinities are with H. tessellata that is very widespread across the northern Karoo and Cape; and with H. granulata. The latteris certainly more closely allied to H. tessellata and the separation may just be on the basis of geographic separation rather than on the morphological characters that could be cited. This may be true for H. venosa as well but in my opinion this would conflict with the general evidence of species geography.
Haworthia retusa (turgida).(Fig. 2)
The name is written in this form because in the existing situation where species are confounded by definition, lesser categories are even more so. The bracketing thus suggests that this is a variant of H. retusa. This species is very widespread and abundant east of the Park with one vicariant population just north of Bredasdorp. The (turgida) variant is one I regard as a riverine cliff ecotype and occurs in the park only along the Breede River at the northwest limits of the Park. It is also downstream along the Breede River. H. mutica is a closely related species that in my opinion actually just extends the distribution of H. retusa (retusa) westwards. My opinion further is that an “evolutionary” process is not complete and that H. retusa and H. mirabilis may be the same “species” separated primarily on the basis of a different flowering season. H. mutica does occur just east of the Park (Rotterdam, Mullershof), and my observation is that H. floribunda is involved with H. retusa and H. mirabilis in this occurrence. It is in fact one of the objectives in this report; to ask if this might be true. Flowering time is spring.
This species and its variants appears in various of my publications and often linked to H. mutica and H. pygmaea.
Haworthia mirabilis. (Fig. 3)
This species is very common and widely distributed from Genadendal, Caledon and Napier all the way to east of Albertinia. It is generally summer flowering and extremely diverse. As its interaction with H. retusa in the west generates H. mutica, so I believe the same interaction produces H. pygmaea in the east. H. retusa is spring flowering but hybrids with H. mirabilis are present in the field and there are populations that suggest that these two species are involved. Flowering time is summer.
Extensive records are discussed and illustrated in various publications but primarily in the Update series, specifically Update Vol.3, part 1. The species is incredibly variable and the floral variations are discussed and illustrated in a flower report as Haworthia Update Vol.8 (Alsterworthia 2012).
Haworthia floribunda. (Fig. 4)
This species is also summer flowering and has a curious relationship with both H. retusa and with H. mirabilis also with evidence that such “introgression” is evidenced in populations. It is at its western limit at Swellendam and the variants in this area are extraordinary. Flowering time is summer.
Also incredibly variable with hybridization with both H. mirabilis and H. retusa and it is hypothesized that some Haworthia populations are derived from such introgression (see also Haworthia Updates Vol.5 ,part 1, Chapter 6).
Haworthia marginata. (Fig. 5)
This species occurs from Ashton eastward to east of Riversdale in a rather narrow eas-west band that widens near Bredasdorp. It interacts with both H. pumila and H.minima. The species in this subgenus flower essentially in mid-summer. The Park population together with that of the adjacent Rotterdam population flowers in spring. At Rotterdam there is hybridization with a version of H. minima.
Haworthia minima. (Fig. 6)
This species distribution generally complements that of H. pumila and then follows, but more widely that of H. marginata. In the southwest it extends to near Elim and is common along the near-coastal strip as far east as Hartenbos. The inland distribution is not well known in the area west of the Breede River. It is probable that it was common in the area now extensively cultivated but it is not known to me inland in that area north of Tarentaal. West of Swellendam there are only two records known to me at Sanddrift, Drew and again furher west of Drew. In the one case there is hybridization with H. marginata. In the second case a short distance west of this is the variant ‘poellnitziana’ and I suspect that introgression with H. pumila may have occurred there The essential differences are that H. pumila has a colour range in the brown-green range, the flowers are generally olivaceous, and the tubercles are larger and rougher. The colour range for the smaller H. minima plants is in the blue-green range, the tubercles are smaller and smoother, and the flowers whiter and greener. H. pumila hybridizes naturally with Astroloba corrugata. Also at Sanddrift, H. pumila hybridizes with H. marginata . It is almost inconceivable that it has not hybridized with H. minima and it will be very difficult to superficially judge if plants were indeed hybrids or not without resorting to DNA sequencing (when that technology is advanced enough). H. minima hybridises with H. marginata at Heidelberg (S. Hooikraal) and in the past with H. marginata just northeast of Bredasdorp. Flowering time is summer.
Some variation is presented and discussed in Haworthia Updates Vol.5. part 1, Chapter 7.
The geology of the Park Bontebok National Park (BNP) is home to three main land types listed in the National Land Type Database, the first land type consists of conglomerate, sandstone and mudstone from the Uitenhage Group (Thamm and Johnson, 2006), which are overlain with Tertiary terrace gravel and can be found on the upper slopes of the north-western section of the park. The second land type is mainly tertiary and quaternary terrace gravels and covers the majority of the park, central and eastern sections. Within this there are high-level gravel terraces with silcrete and ferricrete outcrops, alluvial flats of loam and sandy loam. The final land type consists of siltstone, shales and sandstones of the Bokkeveld and Witteberg Group, these are found along the Breede River section in the western corner of the park. Soils comprise mainly of alluvial sand along the low-lying areas and undeveloped podzols in the main portion of the park. (Chief Director of Survey and Land Information 1993).
Map 2 – The geology of the Park
The vegetation and flora of the Park Originally established to protect the threatened Bontebok antelope the Bontebok National Park is situated in the Cape Flora Region, a Global Biodiversity Hotspot (Goldblatt and Manning, 2000). BNP occurs in an ecotone (transition zone) between Fynbos and Renosterveld and the national vegetation map lists it as Swellendam Silcrete Fynbos (Rebelo et al., 2006) having a mix of both Renosterveld and Fynbos species. The conservation status of this vegetation type is Endangered with the largest portion being formally conserved within BNP. Along the Breede River banks Cape Lowland Alluvial vegetation occurs in the park and is listed as Critically Endangered (Driver et al., 2005), only 1% is formally conserved. A total of 650 floral species are listed as occurring in BNP, from 280 genera and 85 families (Kraaij, 2011). Of these twenty-nine species are listed on the Global threatened species list and a further 23 are recognised as species of conservation concern. The families with the largest species occurrence in the park are the Astercaeae, Iridaceae and Fabaceae (Kraaij, 2011), while the families of Asphodelaceae, Crassulaceae, Poaceae and Cyperaceae were well overrepresented in the park. There are also two species endemic to the park (Aspalathus burchelliana and Diosma fallax) and largest of only 2 populations of Erica filamentosa can be found in Bontebok. Succulents only comprised 8% of the total growth forms within the park with geophtyes being the majority (23%) (Kraaij, 2011). BNP flora is one of the richest in the region and conserves flora that are not conserved elsewhere. It is an area of high conservation priority and diversity and forms a major part in the conservation of Renosterveld in the Overberg region.
Map 3 – The vegetation and flora of the Park
The scope and coverage of this report Three of the Haworthia species are known from single populations, and these are indicated on the distribution map (see map 1).
The report therefore covers those species with multiple records, recording their whereabouts and the pictorial plant body and flower variations. Localities are numbered according to M.B. Bayer accession numbers.
A. Haworthia mirabilis. Accessions 6644, 7704, 7744, 7805, 8035, 8043. In Jan 2013 population 7704 was in flower, buds were observed at 8035 but not at 7744. As in my flower report (Update Vol 8) the flowers are not convincingly different from those of H. floribunda. The variation of the plants in especially 7704 and 7805 suggest introgression with H. floribunda.
MBB6644 – Haworthia mirabilis
Flower profiles and faces
2. MBB7704 – Haworthia mirabilis
Flower Profiles, and Faces
3. MBB774 – Haworthia mirabilis
4. MBB7805 – Haworthia mirabilis
Flower profiles, faces, and bud
5. MBB8035 – Haworthia mirabilis
6. MBB8043 – Haworthia mirabilis
B. Haworthia floribunda. Accessions 8017. In flower Jan 2013. A record 3439 exists for the Park, but the precise whereabouts is not recorded. The plant colour and form do not accord with 8017.
7. MBB8017 – Haworthia floribunda
Flower profiles and faces.
Habitat requirements of Haworthia can be so specific as to defy my describing them. Here is a picture looking north of the habitat for 8017 H. cf floribunda. The rocky ridge extends a long way further north and then west, as well as a long way south. Much of it is of course unexplored in enough detail to know if there are Haworthia there or not. At the southern end of the ridge about 1km away H. mirabilis occurs. To the west the ridge continues, rises and then transforms into steep but low cliffs along the Breede River running northwards. On those cliffs one finds H. retusa (turgida). H. mirabilis is again present on north slopes east of the cliffs. H. venosa is on low river cliffs to the south and it may also be where this ridge abuts the river and turns north.
C. Haworthia minima. Accessions 7743, 8036, 8037
In Jan 2013, a single weak inflorescence was observed at 8036. 8037 was flowering well. No flower or buds were observed at 7743. The plants are not the characteristic blue-green colour of regular H. minima, and the tubercles also seem to be rougher. The flower colour is that of H. minima without the olive-green associated with H. pumila. Note has to be taken of a now non-existent population just south of Swellendam on the N2 that flowered in late Spring with an abnormally large flower, and then a population to the near east at Rotterdam that appears to flower in Summer and where there are hybrids with H. marginata. One cannot exclude the possibility that there is a genetic continuity with either or both, H. marginata and H. pumila.
8. MBB7743 – Haworthia minima
9. MBB8035 – Haworthia minima
10. MBB8037 – Haworthia minima
Flower profiles and faces.
D. Haworthia retusa (turgida).
Accession 2420. This is the cliff-dwelling form of H. retusa and it is only known from the steep riverine cliffs on the northwest boundary of the Park. Expected to be in flower in September.
The significance of the population is the similarity of some clones to H. reticulata that is only known eastwards along the Breede River until Drew from Worcester. It also tends to be a clump-forming species favouring steepish rocky slopes as opposed to its geographic sister species H. herbacea that tends to be more solitary and occupying a wider range of habitat.
11. MBB2420 Haworthia retusa ‘turgida’
View downstream from 2420 habitatView up stream from 2420 habitat
References:
DRIVER, A., MAZE, K., ROUGET, M., LOMBARD, A. T., NEL, J., TURPIE, J. K., COWLING, R. M., DESMET, P., GOODMAN, P., HARRIS, J., JONAS, Z., REYERS, B., SINK, K. & STRAUSS, T. (2005) National Spatial Biodiversity Assessment 2004: priorities for biodiversity conservation in South Africa. . Strelitzia 14. South African National Biodiversity Institute.
GOLDBLATT, P. & MANNING, J. (2000) Cape Plants. A conspectus of the Cape Flora of South Africa, Cape Town, National Botanical Institute of South Africa Missouri Botanical Garden Press.
KRAAIJ, T. (2011) The flora of the Bontebok National Park in regional perspective. South African Journal of Botany, 19.
REBELO, A. G., BOUCHER, C., HELME, N., MUCINA, L. & RUTHERFORD, M. C. (2006) Fynbos Biome. IN MUCINA, L. & RUTHERFORD, M. C. (Eds.) The Vegetation of South Africa, Lesotho and Swaziland. Sterlitzia 19. Pretoria, South African National Biodiversity Institute.
THAMM, A. G. & JOHNSON, M. R. (2006) The Cape Supergroup. IN JOHNSON, M. R., ANHAEUSSER, C. R. & THOMAS, J. R. (Eds.) The Geology of South Africa. Johannesburg, The Geological Society of South Africa.
I was at Bontebok Park south of Swellendam this week specifically to get another look at Haworthia floribunda there and why it is so different. On the way I did some exploring at Stormsvlei about 25km west of Swellendam where I know H. mutica Klipport in a shale environment, and a very odd H. mirabilis growing on a small patch of manganic conglomerate. But going south onto the northern slopes of the Bromberg we found three populations of H. mirabilis in sandstone that are again “different” in the sense that “H. groenewaldii” could be different from H. mutica. This is just a local geological phenomenon and fully sequential with H. mirabilis to all compass directions. If you extrapolate this to Swellendam you have to conclude that the Swellendam mix of H. floribunda, H. marginata, H. minima, H. floribunda, H. mutica and H. retusa is the way it is because of the unusual geology of the area. The Bontebok Park is a relatively massive area of tertiary gravel of mostly river origin and derived from Table Mountain Sandstone. Tertiary gravels east and south are derived from silcrete and ferricrete. I do not know the detail of the mineralogy but it most definitely forms the basis of the soils, vegetation and habitat across the Southern Cape. I specifically looked at H. marginata in the park and see that it was in seed i.e. September flowering with massive capsules and seed. Now if Marx can persuade someone that this is a different species, I accept that I am a monkey’s uncle and that the differences in H. marginata elsewhere e.g. Drew, Bredasdorp and Heidelberg or Riversdale, mean there are several similar species. Oh, I forget – that means H. floribunda would also be several species, and so is H. minima. But then H. mutica is of course several species and H. retusa several dozen. H. mirabilis several hundred.
This is 8017 from the Bontebok Park Swellendam. In some circles I am expected to get excited and see this as something new. In the context of the Haworthia in the wider area, this is a variant of H. floribunda. In my flower report it is evident that there is not much difference between the flowers or flowering time of this species and H. mirabilis. It hybridizes with H. retusa, H. retusa (turgida), and H. pygmaea despite differences in flowering time, and also with H. mirabilis. There are populations that I think are an older product of such hybridization (or failure to have ever separated).
Typical – an over-used comment. A white small glass ball is found and named “Marble alba”. Then a red one is found and named “Marble rosea”. Then several other colours turn up and “rosea” is lumped under “Marble alba” as a variety. Automatically all the others become “Marble alba var. alba” if they are not specifically named or not put under “rosea”. If “rosea” had started as “Marble alba var. rosea”, there would automatically have been a “var. alba”. All other glass marbles known and unknown would have been “var. alba” even if no another white marble ever turned up. A type just establishes a name and the best way to determine how that name is used by an author is by ALL the illustrations and pictures he/she notes. A type that establishes a name may be an oddity or a single variant that does not easily establish a use. The advice handed to me by a group of taxonomists in 1972 was that it would be best to scrap all the old Haworthia names and start again from scratch. The group was led by Prof. Schelpe, one of the few professors who as a taxonomist headed the department. He had explained this view in a published article in respect of Gasteria and why he rejected the idea of personally revising the genus. He had no solution for types and names that could not be directly linked to a set of herbarium specimens or similar evidence. Now substitute H. mirabilis for marbles alba and where the contrast of white and red is absent.
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Just what is the type form of H. mirabilis and what constitutes H. magnifica? That is the subject of my question about the way in which the typical variety is recognised. There are 4 populations of H. mirabilis in the Park and many to west, east and south. Imagining that there is also a “maraisii” is just crackpot even more so to say “magnifica”. There is quite definitely one system that you can see from many populations and that system inter-twines with two others also on the basis of many populations. At this stage of the available information on these populations there are a lot less species – see also the article Haworthia flowers – some comments as a character sourcepublished in Haworthia Updates Vol. 8 by Alsterworthia International. I also started off thinking that there was an H. maraisii and an H. magnifica but changed my mind quite early. In the last ten years I have come to see that both they and H. heidelbergensis are nothing but parts of one system viz H. mirabilis. Ask the authors who think otherwise to present some clear evidence based on good random sampling and statistics. See the Haworthia Updates Vol. 4 article Some variation in Haworthia mirabilis var. sublineata by Loucka and Bayer on the stats of H. mirabilis (sublineata).
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Klipport is a farm a bit off the beaten track on the north side of the Bromberg low range. Bromberg is the eastern end of the Riviersonderend Mountains and is cut off from them by the Riviersonderend River that changes direction to the north to do so and then east again to join the Breede River. Bokkeveld shale is strata-wise above the sandstone of this mountain range and a small amount of shale is present here on the south bank of the river. At Klipport it is a narrow sloping stretch of about 500m long and about 75m wide with white gravel and clays. The vegetation it totally different to the grassy fry Fynbos and renosterveld of the Bromberg and has karoid succulent vegetation with the endemic Gibbaeum esterhuysdeniae and a Brianhuntleya species. The site is also home to a smoothish form of Haworthia pumila, and also to a range of H. mutica variants that might have a name attached by now. Another very interesting species there is Drosanthemum micans. This species has an extraordinary range of variation that parallels that of Haworthia and at Klipport is moving to another face as D. lavisii. It would be very instructive if other Haworthia taxonomists could, or would, take note of this kind of parallel and recognize that there are significant clues to how Haworthia species and their variations are part of a larger pattern.
This small shaly area has a very patchy distribution of the species on it with patches of a restioid and Pteronia. Through some weird quirk a German immigrant ploughed up a vast area of very marginal virginal land and planted gums and pines that after 30 years are still far from harvestable. I doubt if they will ever yield anything. This happened under the eyes of Nature Conservation and the Dept of Agriculture that is supposed to manage land use. The area on the river itself was paradoxically a source of timber in the early Cape days and is now severely infested with exotic gum and wattle. It is this riverine growth that sustains the timber enterprise of the adjoining farm Vaandrigsdrift. Not far away between the shale and the sandstone is a manganese deposit that is now being mined. At the end of this deposit is a tiny set of large conglomerate-like rocks with a variant of H. mirabilis. This species also occurs as an interesting ecotype a short way further east where the shale is less transformed along the sandstone interface. In fact I do not know exactly what it it is that has driven this decay of Bokkeveld shale to kaolinic and bentonitic clays where it has been covered with Tertiary marine deposit at some stage in geological history. Whatever it was, is certainly significant with respect to island-like habitats and “conserved” vegetation remnants.
19. 2019.5.23 – On more than one occasion some collector or other has sought retreat from an argument about species to some purported botanical expert (i.e. someone with a botany qualification) to support their wild guesses and assumptions about names. Too often such persons simply assume they know more about classification than any botanists might. Here are two plants of H. floribunda in a population that differ in having these strange white spots in the leaves. According to the H. groenewaldii argument it must surely be a new species. What about all the other plants in the population? Far more significant is that these plants are in a floribunda population not far from where “H. groenewaldii” (= H. mutica) occurs, where occasional plants have significant leaf spotting too. H. floribunda is very closely entangled with the greater retusoid complex where groenewaldii is secondary to the mutica element in that complex. Those spots must have a significant connection. ♦
34. 2019.6.8 – Traveling west to SW Tradouw there is a habitat just like those that house mirabiloids and retusoids. Yet here there is H. floribunda and Aloe brevifolia in separate discrete small areas in an area stretching about 300m. There is Aloe arborescens that likes the creted upper zone of the old terrace and Aloe ferox that hangs around on usually the edges of the exposed and bare clay. Why no mirabil/retus-oids? ♦
40. 2019.6.13 – How does one know anything? Why I ask is because I have so often heard strong opinions based on a great degree of ignorance. The revision process in plant classification is built on the accumulation of information. At one stage all known Haworthia were 5 small Aloe species. It has been pretty much chaos ever since. There are many total imponderables within Haworthia even in its purest form i.e. with Haworthiopsis and Tulista excluded. While it is difficult enough to arrive at a reasonable guess as to what a population may be, it is very much more difficult to explain an own informed guess to someone convinced of something else on the basis of near nothing. Can one explain to a frog in a pond that has not even seen the other side of it, what the ocean is like? At the same time one should not expect the ignorant to take a blind leap of faith. H. groenewaldii as H. mutica, needs to be seen in this context and there is a lot of peripheral stuff that needs to be explained. I did not think H. floribunda was going to be so central to my story, but it is. It occurs across the range of the retusoids from the Gouritz River in the east to west of the Breede River in the south. But it gets lost among the mirabiloids along a east/west line between Riversdale and Swellendam. What it does at Swellendam is extraordinary. H. floribunda is a rather inconspicuous small plant characterised by a flattened leaf and rounded leaf-tip. It is this character that seems to be infused into the mirabiloid/retusoids suggesting that H. floribunda itself may be essential to that group. BUT it has its own complications in that it can be seen to be linked to H. variegata, H. parksiana and H. chlorocantha. How does one know that? It is an opinion probably arising solely from seeing so many things in the field that suggest this. Judging from all my pictures, its chief character seems to be that it is non-photogenic. It does incline to be well drawn into the soil with at least half the leaves below soil surface. My pictures are appalling but here are some to give an idea of what “floribunda” may look like, before I show what it is like at Swellendam.
Some of the weak points emerging in my story…
1. Starting point. Perhaps I should have said a lot more. A great deal more. I had a plant from N Bredasdorp that was H. mutica by my identification, and another from N Mossel Bay as H. pygmaea. They were absolutely identical except for the colour of the floral bracts. And I had B/W pictures from G.G. Smith’s draft “monograph” of plants from Buffeljags River Swellendam, and from Kransriviermond Heidelberg that Smith had labelled H. mutica. These also looked nearly identical.
2. Missing pictures. I have no decent pictorial record of my field activity until I got a digital camera about 12 years ago. So there are gaps in the photographic record.
3. Too much to digest. I have about 300 populations relevant to the story with an average of about 10 pictures/population plus the whole historical record. I can’t throw one out without feeling my story is diminished. How does one transmit the combined picture of all these images to anyone else when some of it is forgotten or incorrectly memorised?
4. Inadequate flowering time data.
5. Inadequate understanding of the geological facts and the time scales involved.
43. 2019.6.17 – Nearer Swellendam to the north west is where there is a bigger version of floribunda similar to this picture. It is unusual because of its pointed leaves. When first found I thought they may be in fact variegata. That seemed very improbable at the time, but since then variegata has appeared at several localities west of the Breede River. It it is otherwise limited to the limestones south of Albertina and west of the Goukou River. So this is an indirect link between the two species that gets very more involved in the south. What is important in the implication of geographical position and physical similarities.
44. 2019.6.18 – Within Swellendam along the highway there is a little patch where floribunda occurs as the more characteristic small plant deeply withdrawn into the soil. Dark coloured and with the leaf tips barely sticking out of the moss and lichen growing on exposed weathered shale.
45. 2019.7.2 – A short way down the road is this monstrous green form of what cannot be anything else but a variant of floribunda. The big question I suppose is “how do you conclude that?” No! You tell me how you can possibly think it’s anything else. It comes from a familiarity with the field, familiarity of what else goes on around Swellendam, familiarity with variability in nature, a bit of academic learning, what haworthias do generally and then just plain common sense.
46. 2019.7.3 – A few kilometers east of Swellendam there is this population of green floribunda while another kilometer east returns the smaller dark more familiar form. This strange behaviour of one species is not peculiar to that species. It simply demonstrates that a tiny shred of circumspection is needed before wild declarations of a new binomial identity.
47. 2019.7.4 – In the Bontebok National park, less than a kilometre away from the green and the ordinary floribunda virtually within Swellendam is this little green form. There are also at least 6 populations of the smaller heidelbergensisoid mirabilis – all different from each other. I surmise that this may be of mixed origin – avoiding the term hybridization that implies pre-existing separate species. I suggest outcome from a common gene pool rather than this simplistic dependence on an even more totally speculative guess.
48. 2019.7.5 – Very many years ago I did see the small floribunda within the Bontebok Park but it was extremely cryptic and I was not able to find it again. But just east of the park is this. What is curious is that it is the second of a series of 5 “species”, almost contiguous populations going south to north, each occupying their own space in a near uniform environment. Two of those populations are Tulista marginata and T. minima that vary in some degree from their own species norm and even flower much earlier than they should be expected to. Added to that is the fact that now there are clearly hybrids. So we have a situation where we have two species that may have never truly separated, interacting again? Similar situations exist throughout the Haworthia domain. And I have yet not told half the known floribunda story either.
2019.7.6 – In carrying on about H. floribunda (that can be further shown to interact with H. variegata, H. chloracantha AND H. parksiana) it may be thought I have lost track of the prime issue. Not so. The gist of the story is that there are two main elements viz retusoid and mirabiloid. This gene pool disgorges elements that can be labelled H. pygmaea, H. retusa and H. emelyae in the east, and H. mutica, H. retusa, and H. mirabilis in the west. H. floribunda simply demands attention by its involvement and by what it does in its own right. This brings me to something else. I have a near polymath friend who does top-level seriously academic work in major plant families. This source mentions the promise of next generation sequencing. In trying to understand the technology all I could really grasp was this end statement … “The only thing slowing us down now is the interpretation of results.” At the same time I was pondering the phylogram for the aloid sequencing I was unhappily involved in (two major attempts). Especially about some peculiar relationships expressed in that phylogram. It is tough having a small brain when such complex issues need scrutiny and question. But those peculiar relationships (I hesitate to point directly at them now because it means many words and browsing slowly through many files for which I lack patience), require to make me say emphatically … ”A two dimensional array can definitely not illustrate what is a minimally three-dimensional space/time product (species)”. So my contribution is, that while the interpretation of the results rests on a two-dimensional phylogram, the result will never be worth what it should be. ♦
112. 2020.03.26 – The trickster – H. floribunda. This is the Draaihoek Albertinia site where I was not sure (ca 1970) if it was floribunda or chloracantha. I went back to check ca 2001. The place was grazed and trampled to a frazzle – sheep and/or ostriches. I found a few specimens right at the far end. They pass for floribunda and especially if you recognise that floribunda has a good many faces. The last picture is from west looking east – so this is the top rocky edge of a south facing drop. The water is the Valse River. Dekenahii is across the way.
So in these very distressing times, lets see where we go next.
113. 2020.03.26 – This is H. chloracantha from east of Tweekuile along the Valse River north Albertinia, so also further east from Draaihoek. A small dense patch of plants under bush at the upper end of a south facing slope – escaping full exposure to sunlight. A few leaves show the round leaf tip of floribunda but the plants are also much more proliferous than I have ever seen in that ‘species’. It is very useful to consider all the variants in this complex to see how and why I arrive at my opinions on their classification. This is not a glamour complex but as systems they work in exactly the same way. I really regret not having good pictures and especially since I first saw this species at Great Brak in 1969.
114. 2020.03.27 – Ouvloere is east of the previous as that is from Tweekuile but the plants are really odd. Unlike the previous these plants are in and under low ground hugging plants at the top of a slope. This must be about 8km west of the Gouritz River.
How would you know this is H. floribunda?
The answer to … How would you know this is H. floribunda is because I say so? So please forgive my apparent arrogance. I would dearly love to show you all the plants and populations that lead to this conclusion because you would surely be able to then make that decision for yourself. These two pictures are also floribunda from very near to ‘splendens’ at Dekriet (Snymanskraal) west of Albertinia. A strange habitat where the rocky ferricrete is exposed but with a collapsed eroded depression in which the plants grow around the edges. When this time of tribulation is over, I hope to go back and get habitat pictures. Reminiscent of parksiana?
115. 2020.03.30 – The Valse Rivier joins the Gourits at Die Eiland where there is now H. chloracantha and eastwards. It supplants floribunda entirely – or is it truly just the same species?
116. 2020.03.30 – The last post was of plants from N Die Eiland (Buisplaas). From the top edge of a west facing shale cliff. These are also Buisplaas but from a low north facing slope of alluvium. I never imagined chloracantha could be so abundant and so resistant to the impact of habitat. But it becomes one of the most remarkable of the species on closer scrutiny.
The great awakening will be when one can truthfully and openly write about, discuss, the reality of basic life forms i.e. species. When we are free of the prejudices, beliefs and misconceptions of science as practiced and religions as preached. The great awakening will be when one can truthfully and openly write about, discuss, the reality of basic life forms i.e. species. When we are free of the prejudices, beliefs and misconceptions of science as practiced and religions as preached. ♦
122. 2020.04.07 – Like with the present pandemic we are coming to something difficult to comprehend. I went to Botlierskop, Rooiheuwel farm, north of Klein Brakrivier hoping to see a slender long-leaved form of Haworthia chloracantha (a JDV record). Instead I came across this population of rather stubby plants. Keep this population in view – two close populations to follow and then an issue of proximities!
Also on a rock slab about 70 meters away. Is this still chloracantha or is it parksiana?
Frantisek Vesely – We saw the similar situation – on slope there was chloracantha (+ hybrids), on flat a few dozens meters away there was parksiana and its hybrids with chloracantha – not difficult to recognize what happened there as they were so close and mixed up but on one spot most of them were chloracantha-like and the other spot were more parksiana-like plants.
600M away there is a population of uniform parksiana. But how can we say hybrids for the previous? How long have these populations been in existence? Is transfer of pollen a problem and over what distance? There seem to me to be a host of unanswerable questions in general about hybridization as an explanation for variability. It is linked to the more general question of co-occurrence – of mixing or non-mixing of species. Ring clines? Also something we should really try to document – i.e. the proximity of the various species to one another. Add to that the occurrence of hybrids.
123. 2020.04.09 – This population, east of Riversdale, is an aspect that really calls for attention – proximity of kinds (species) and hybridization. At this locality there is H. retusa and H. floribunda in contiguous/adjunct/closely neighbouring populations. Among the floribundas there is a single putative hybrid? How often does this happen and why are kinds (species) so seldom growing together? When is an entire population comprised of hybrids? What comes in between? Let me just add that I see very little in the taxonomic literature dealing with the problems of variability that exist in many genera other than Haworthia. It is simply obscured.
I am quite aware that my comments are often cryptic and often deliberately so. As a society we are very misled and never more so than in respect of the present pandemic. I do not see how we can put that aside and think that we can at the same time depend on our collective intelligence to solve and understand anything else e.g. taxonomy of Haworthia. It is a conscious creation and that consciousness has to be lifted to a level that we see things for what they really are. Hybridity is at the root of species definition and as such a critical step to recognising species. At the same time geographical location is also critical. To to think anything can be explained while there is a paranoia about revealing it, is wishful thinking.
2020.04.12 – There is a problem with classification where priority creates difficulty. The name Haworthia retusa has nomenclatural priority over the name turgida, when in actual fact the turgida form is the main one and retusa is a variant of that. It is much easier to then understand, describe and explain the idea that retusa becomes mutica in the west, retusa and mirabilis become pygmaea in the east, while turgida and mirabilis become emelyae in the Karoo. That might be where I should now leave things. There are a hoard of complexities that just get lost in the needs and activities of the majority who have no need of much more than a lot of desiderata. ♦
Introduction. Apart from the vexing question of species delimitation, there is a truly curious phenomenon in how the species interact spatially and also behaviorally. The authors here believe in a systems approach as the only rational approach to Haworthia. With H. floribunda as part of one system (viz. with H. variegata, H. chlorocantha and H. parksiana), H. retusa is another (viz. with H. turgida, H. mirabilis, H. pygmaea, H. mutica and probably H. emelyae). Within these two systems there is secondary interaction and we are trying to unravel the role of the two prime elements with H. mirabilis (as a secondary in the H. retusa system). Bayer has discussed floribunda in HaworthiaUpdates.org and it is not necessary to repeat that here. In this article we are only dealing with single populations of H. floribunda and H. retusa that occur contiguously at a locality SW of Riversdale where they were located by Essie. H. mirabilis does occur 600m to the south-east as a discrete population in a kaolinite exposure site as opposed to the rocky more stable nature of the H. floribunda/H retusa site. It is important to note two things:
This locality is southwest of multiple discrete populations of H. mirabilis as well as H. retusa, while …
It is the only known occurrence for H. floribunda south of the N2 highway from west of Heidelberg to beyond Albertinia.
Preamble. This is neither a taxonomic and nomenclatural debate nor any attempt to explain how these things came about. It is simply the synthesis of the experience of two field enthusiasts and explorers trying to rationalize what they observe in the field. Trying to make sense of many latin names. It is Essie’s suggestion that H. floribunda and H.retusa interact at a deep level and that H. retusa becomes rougher and darker where the two are in close proximity. They do flower at different seasons with a general and approximate 4 month time barrier. Bayer on the other hand suggests that the two species definitely do produce hybrids despite difference in flowering time (observed in several places as isolated individual hybrids). The apparent roughening and darkening observed in H. retusa is more likely to be consequent to a deeper direct connection with H. mirabilis that may have nothing to do with hybridization. As observed elsewhere, hybridization seems to be a convenient excuse to explain phenomenon that have no other obvious origin. What then happens when H. retusa and H. floribunda virtually share habitat here SW Riversdale.
To understand the situation better we have to look at that H. mirabilis population mentioned to the south (MBB6651). I personally have no photographs because I saw the population before I had a digital camera. But I grew them from seed and the accompanying pictures show the incredible variation that suggests an interaction of the retusoid, the mirabiloid and the floribundoid. Essie made a special excursion to obtain 3 field photographs.
The plants. Multiple photographs of the two species accompany this discussion. H. floribunda can be seen to be H. floribunda with perhaps one notable exception that could be dismissed as a hybrid with the greener coloured H. retusa. The H. retusa plants on the other hand do show some variation. The plants are smaller and generally greener than is the case with other nearby populations.There are indeed some plants on the much darker side and this is where an important distinction may be made between the retusoid and the mirabiloid factions (also spring vs late summer flowering) of a single system. The retusoid system is characterized by light coloration with yellows greens and pinks, while the mirabiloid faction is darker green to blackish green with rusty reds and browns. Two plants are shown from an essentially retusoid population about 33m north.
Conclusion. We do not draw any conclusion from this single location that simply adds fuel to our belief in system and that far less formal names are needed to explain the incredible complexity and high degree of variation. (Note to writer … “Keep it simple, stupid”.) We are further suggesting that all these varied populations are elements of just two complex species systems, and not the fragmented cherry-picked elements that the many Latin names suggest.
Footnote by MBB. It is necessary to write something about Essie Esterhuuizen because whether he knows it or not, he has made an incredible contribution to Haworthia by field exploration that is beyond extraordinary and exceptional. I asked him of he could explain his dedication and commitment to Haworthia and this is what he wrote …
“I was fortunate to have grown up in Heidelberg in the Western Cape. As children we spent most of our time playing in the veld around the town. From a very early stage I was thus exposed to Haworthia and needless to say it formed part of the plants on our rockery. Because of their small size I kept 8 different plants in my so called “collection” which I regarded as a collection of notice. Around 1982 I visited the Karoo Botanical Gardens to see if I could find names for the “small aloes” on the rockery when Bruce at that time the Curator of the garden showed me the garden’s collection. Suddenly I realized that if I want to continue with my hobby – the challenge is humongous.”
Now Essie is Afrikaans speaking and Afrikaans is a language very dear to me because it is what I refer to as a heartfelt language – a language of real people and not like English that is used and abused by many cultures and castes. So I asked Essie to rather write in his home language and leave it to me to try to extract the really personal essence of his interest and opinions. So translated by me, this is what he says…
“I am not a scientist, a collector, or a nurseryman – just a plain field observer. In my first field excursion involving the name heidelbergensis, I was shocked to find that what was known and written about these plants was just a drop in a bucket. Just judging and identifying plants on the basis of what is written, or in collections, does not reflect what the situation is in the field. On the other hand, to explain what one observes in the field verbally or in writing is extremely difficult. Field observation becomes like a treasure hunt. When you think you have found some sort of answer, you find another clue to lead you on.”
The crux of the matter is that it is extremely difficult to articulate what is meant in this last sentence. By putting our heads together we agree that there is pattern in this very complex system, as we have just explained in the article “Last Hurrah”. We have planned to collaborate in presenting the evidence for the continuity we say exists within and between two species systems. Essie’s final sentence reads …”For me my observations will never come to an end … the challenges are too great, continuous and wide ranging”.
Fig. 1 H. mirabilis 6651JME SW RiversdaleFig. 2 H. mirabilis 6651JME SW RiversdaleFig. 3 H. mirabilis 6651JME SW RiversdaleFig. 4 H. mirabilis 6651JME SW RiversdaleFig. 5 H. mirabilis 6651 (cult. exSW Riversdale)Fig. 6 H. mirabilis 6651 (cult. exSW Riversdale)Fig. 7 H. mirabilis 6651 (cult. exSW Riversdale)Fig. 8 H. mirabilis 6651 (cult. exSW Riversdale)Fig. 9 H. mirabilis 6651 (cult. exSW Riversdale)Fig. 10 H. mirabilis 6651 (cult. exSW Riversdale)Fig. 11 H. mirabilis 6651 (cult. exSW Riversdale)Fig. 12 H. mirabilis 6651 (cult. exSW Riversdale)
A note on a H. floribunda and H. retusa interaction was written by Bruce Bayer based on the opinions, observations, and exploration of Essie Esterhuizen.