SUMMARY
Haworthia koelmaniorum is a very interesting cryptic species that frequents both a savannah and a grassland biome in western Mpumalanga. The plants have been studied by Charles Craib for about 20 years and this account discusses the findings drawn from this research. Some of the work on Haworthia koelmaniorum variety mcmurtryi has been published and that on Haworthia koelmaniorum variety koelmaniorum is presently in preparation for publication. This latter research is discussed with particular reference to the unique ways in which a fire driven ecology has produced variation in these plants. A recommendation is also made that H. koelmaniorum is treated as a variable plant without any rank below species level.
NEWSLETTER
THE HISTORY, DISTRIBUTION AND HABITAT OF HAWORTHIA KOEL-MANIORUM
H. koelmaniorum was described in 1967 by Mrs. Mauve (Amelia Obermeyer). The plant was brought to the attention of the then Botanical Research Institute by Mrs. T.H.C. Koelman from the Groblersdal area of the central Transvaal (now western Mpumalanga). At the time of her description of the new species Amelia Obermeyer-Mauve remarked that it was with considerable surprise, that this Haworthia was found in the Groblersdal district. It was moreover of great scientific interest that these plants were discovered so far away from the predominantly Cape distribution of the Genus (1967, 1502).
H. koelmaniorum continued after the type description to be found at various places in the Groblersdal area and a map of its currently known distribution is provided by Pieter Bosch (2004: 57). In this map the plants are shown as occurring mostly in a large area south south east and south west of Groblersdal with a narrow tongue of distribution in the hills and mountains in the extreme south west.
Haworthia mcmurtryi now Haworthia koelmaniorum variety mcmurtryi was described by Charles Scott in 1984, (1985: 140 – 141). The plants were originally found by Doughlas Mcmurtryi and the late Mary Bleck in typical highveld hilly grassland with Faurea saligna and Protea caffra two types of trees often found close to the transition between highveld and savannah. H. mcmurtryi was reduced to varietal status under Haworthia koelmaniorum by Bayer in 1999 (1999: 180 - 181). A sophisticated account, upholding the varietal status of H. koelmaniorum variety koelmaniorum and H. koelmaniorum variety mcmurtryi was published by Pieter Bosch in 2004 (2004: 56 – 60). Bosch was the first researcher to look at how the plants live in their environment. He related geological, pedalogical and climatic influence to variety generation with respect to the two varieties of Haworthia koelmaniorum.
Charles Craib questioned the varietal status of H. koelmaniorum variety mcmurtryi in a paper entitled “The Autecology and Status of Haworthia koelmaniorum variety mcmurtryi (Craib 2007: 22 – 27). He argued that the retuse leaved “mcmurtryi” were simply adaptations to a fire driven ecology under circumstances where longer leaved haworthias, the typical “koelmaniorum”, would not be able to survive frequent dry season grass fires.
In a current detailed research paper in preparation (Craib, September 2008) Charles evaluates the status of the two varieties of Haworthia koelmaniorum. Some of the findings are discussed below with specific reference to H. koelmaniorum var. koelmaniorum in the western part of its distribution range and H. koelmaniorum var. mcmurtryi in its typical hilly highveld grassland habitat.
THE CONCEPTS OF SPECIES, SUBSPECIES AND VARIETY IN THE ORGANISATION AND PRESENTATION OF BOTANICAL KNOWLEDGE
In botany species, subspecies, varieties and forms are concepts for organising and presenting knowledge. These concepts cannot be demonstrated empirically. In recent decades attention has been paid to matters such as chemotaxonomy and DNA sequencing which are empirical scientific techniques. Scientific techniques have been used randomly alongside concepts with an absence of necessary theorisation not only at an appropriate philosophical level but also at the level of practical application. At best the techniques of empirical science can provide some level of authenticity for the concepts currently in use.
The study of Haworthia calls into question the usefulness of the concepts themselves. Species in this genus rarely fit neatly into conventional boxes of classification. Students of Haworthia accordingly vary between those who classify virtually each different population of one species as a separate species to those who reduce the same plants to one species with the variations described under species rank. Neither of these situations are particularly satisfactory as both are limited by the prescriptions and limitations of existing concepts. The reductive approach is certainly to be preferred should one’s interest be in trying to understand the development of these plants in space and over time.
The concept of variety with respect to Haworthia koelmaniorum must essentially be based on differences between “koelmaniorum” and “mcmurtryi”. During some 20 years of detailed field research it was found that none of the characters used to distinguish the two varieties of Haworthia koelmaniorum were accurate. This matter is discussed in full detail in the paper currently in preparation. Only one factor that could usually be consistently used to separate the varieties was found. This was the character that H. koelmaniorum var. koelmaniorum often grows more than one flower stem in a given flowering season usually 2 – 3. These are frequently produced over a period of a few weeks. H. koelmaniorum var. mcmurtryi nearly always produces a single flowering stem. These characters however hardly justify any distinction at varietal or any other level.
H. koelmaniorum is best regarded as one variable species, the variation related to the different kinds of niches it frequents in a predominantly bushveld biome and a predominantly grassveld biome. This approach however begs the question as to what we do with the classificatory labels. The logical approach seems to be to abandon them entirely and to simply use the names of places near the colonies to describe the variation. On the other hand if the autecology of the species across its distribution range is adequately understood it is unlikely that detailed descriptions of the variations are either desirable or necessary.
The discussions that follow have been elicited from about 2 decades of field observations. Variations in the plants are entirely related to the kinds of rocky niches they can colonise and grass fire cycles particularly the load of flammable material in different habitats that gets burnt in dry season veld fires.
THE HABITATS OCCUPIED BY HAWORTHIA KOELMANIORUM AND THE ROLE PLAYED BY GRASS FIRES.
H. koelmaniorum var. koelmaniorum occurs in arid savannah at an altitude of about 900 – 1200 metres. The plants are always associated with fixed rocks, usually quartzites tightly wedged into the ground. There is invariably mixed mainly deciduous woodland present at the localities frequented by variety koelmaniorum. The presence of deciduous trees and the leaves that fall from them in the autumn have a notable and hitherto unrecorded role to play in preserving many of the plants from the worst effects of fire. This is discussed below.
The cavities in quartzites tend to be deeper and sometimes also broader than equivalent places in the variety mcmurtryi habitat. In addition the variety mcmurtryi habitat carries much heavier loads of dry grass during the fire season, mostly from May – September. The rocks in the variety mcmurtryi habitat are rhyolite. This rock type forms fissures and cavities that are quite often only slightly elevated above the ground.
The retuse habit of variety mcmurtryi ensures that most of the Haworthias do not get their leaves badly burnt in the grass fires that are such a regular occurrence in their habitat. Dry grass is highly flammable and burns right down to ground level in fires, unless these are driven by high winds. In such instances the burns are more superficial. Occasional variety mcmurtryi plants, with longer leaves, are found in places with deeper cavities amongst large rhyolite rocks.
The smallest variety mcmurtryi plants yet discovered were found on 28 August 2008. They were growing in an area cobbled with rhyolite pebbles and small blocks of these rocks deeply fixed in the ground and protruding just above it. The full rosettes of highly retuse leaves were mostly 2 x 2 cms in size, some even less. These Haworthias appear to represent an extreme adaptation to regular fires. Their small size and very retuse habit render them almost immune to the effects of grass fires. This cobbled habitat is fairly common where variety mcmurtryi occurs but rarely as extensive as at the locality recently found.
The variety koelmaniorum habitat often has thinner loads of dry grass during the winter months and many dry leaves shed by the surrounding trees and shrubs. The deeper cavities between rocks which the Haworthias occupy and scatterings of dry leaves in them, which often only burn partially in fires, permits the establishment of plants with long leaves. In most variety koelmaniorum colonies typical long leaved plants as well as some shorter leaved examples are found. The latter are often indistinguishable in appearance from typical variety mcmurtryi. This data implies that it is neither an advantage nor a disadvantage to have long or retuse leaves in a rocky environment with grass and trees. The considerable variation in variety koelmaniorum colonies, with respect to the size of the plants and length of their leaves, seems to be directly related to the kinds of rocky niches found in quartzite and the composition of the flammable load which consists of grass and leaves.
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