Haworthia Revisited – 60. Haworthia minima

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

minima: small.

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

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

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

a. var. minima

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

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

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

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

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

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

Volume 5, Chapter 7:- Haworthia minima

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

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

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

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Volume 5, Chapter 13:- A February 2009 Miscellany

7807 H.minima.  Swartrug, SE Heidelberg. 002

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

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

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

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

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

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New finds in Haworthia.

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

Map - east of Swellendam

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

1. Haworthia marginata and H. minima

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

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

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

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

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

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

2. Haworthia retusa ‘nigra’

I first allied this element with H. mutica because this is how G.G.Smith referred to his collection from Kransriviermond south of Heidelberg.  Since then there have been many new collections from which can be gathered that this population is a hybrid one between H. retusa ‘turgida’ and H. mirabilis.  There is another collection north of this at Morning Star that appears to have the same parentage but also including H. floribunda.  Then there are populations continuing up the Duiwenhoks and then Klip rivers to northwest of Heidelberg, a population between Heidelberg and Tradouw Pass further west and also a population at Goedverwagting south from there.  Apart from the Morning Star population (February/March) these are all September/October flowering.  There is a population at the southern entrance to Tradouw Pass of the same ilk that is February/March flowering.  Then there is quite a distance between these known populations and a remarkable population at Buffeljags south of Tradouw Pass.  It is remarkable for the fact that it is lauded as a new species viz. H. groenewaldii when I consider it to be generated from the interaction of H. mirabilis, H. mutica and H. floribunda.  I attach no special importance to the fact that it flowers, contrary to H. mutica, in February/March.  This is because I have observed many hybrids between patently different species despite a seasonal difference in flowering time.

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

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

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

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

Perhaps I should close by explaining that I have dropped the use of any rank below that of the species name.  I simply am suggesting that we recognize the need for a trinomial system without the pretension of status, and more greatly honour the binomial as an entity of a greater significance than we may know.   I do this because botany has no proper species definition and consequently species descriptions are just based on wild guesses about possible non-similarity and on the flimsiest of supposed character differences.  The loosely used word “typical” is only truly useful in respect of the one plant dried as an almost unrecognizable specimen that is used to anchor the Latin name.

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

Cross seasonal hybrids observed are-

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

And also Platjieskop, Riversdale.

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

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

Haworthia minima and pumila flowers

6645.1b H. pumila

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

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

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

Haworthia pumila

Haworthia minima

The Haworthia species of the Bontebok National Park

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

Objective:  The significance of the 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 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).

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