'n Morfologiese studie van die genus Acacia Miller in Suid-Afrika
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Robbertse, Petrus Johannes
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University of the Free State
Abstract
Showing abstract in English
English: The genus Acacia is an extensive one with representatives in Africa,
America, Asia and Australia. When Bentham revised the subfamily Mimosoideae
in 1875 he divided the taxon into six subgenera. All the South African representatives
of the genus were placed in the two subgenera Vulgares and Gummiferae.
Bentham distinguished between the two subgenera by the differences in
morphology of the stipules. Bentham's classification of the subgenera differs
from that of Oliver (1871), whose grouping of the South African Acacia species
is based on the morphology of the inflorescence. Species which are affected
by this difference are A.albida, A.schweinfurthii and A.kraussiana.
Because of this problem and the fact that, judging from the literature,
there is some uncertainty regarding the application of taxonomic criteria, it is
necessary that the whole genus be revised. Due to the extent of such a project
it was decided to limit this investigation to the South African species with the
main emphasis on characters which could be of taxonomic importance. Where it
was deemed necessary a few species from other geographical areas were included
in the investigation.
Various characters were found which were valuable for considerations of
phylozenetic history of the genus and for delimitation of taxa.
On the basis of the presence of starch grains in the cotyledons and the
morphology of the seedlings, A.albida, A.kraussiana and A.schweinfurthii were
placed in a new subgenus, Farinosae. On the basis of these characters the subgenus
Farinosae shows affinity with the subgenera Phyllodinae, Bothryocephalae
and Pulchellae of Australia and certain Vulgares species of America. The subgenus
Farinosae can possibly be phylogenetically regarded as the oldest extant
taxon of the genus Acacia.
With the species A.albida, A.krau8siana and A.schweinfurthii in a separate
subgenus, the remaining South African members of the subgenera Vulgares and Gummiferae
can be separated on the basis of classical features such as the morphology
of the stipules and inflorescences. Other clear differences which were
found between these two subgenera are as follows:
(a)The flower of the Vulgares (and Farinosae) species contains a cup-like disc
to which the filaments are adnate, and the ovary has a prominent pedicel
(gynophore). In the flower of the Gummiferae species the disc is lacking
and the ovary is almost sessile. A disc and pedicelled ovary are also
found in the flower of A.albida and the name Feidherbia albida which
Chevalier proposed for this taxon in 1934 is therefore not justified. The origin
of the cup-like disc on which the stamens are found can probably be regarded
as a relict of an ancient branched system on which stamens were borne and
which later became reduced.
(b) In the pods of the Vulgares (and Farinosae) species the fibre zone consists
of both cross and longtitudinally arranged fibres, while the pods of the
Gummiferae species have only longtitudinally arranged fibres, or none at all.
(c) In the pinnules of the Vulgares (and Farinosae) species a few layers of
spongy parenchymatous cells are found between the abaxial and adaxial palisade
parenchyma cells. In the pinnules of the Gummiferae species the
abaxial and adaxial palisade cells are continuous.
(d) The seedlings of the Vulgares species have membranous or leaf-like stipules
while the stipules of the Gummiferae species are spinescent. Hooked spines
are absent on the Gummiferae seedlings but appear on the Vulgares seedlings
either dispersed on the internodes or arranged in groups of two or three
directly below the nodes.
On the basis of the morphology of the flower, the anatomy of the pod and the
morphology of the seedlings, the Vulgares and Gummiferae species can be divided
into smaller groups and, in certain cases, into definite species. Keys have
been compiled to facilitate the classification.
Interesting differences between the Vulgares and Gummiferae species were
noted with regard to the nocturnal movements of the leaves. These differences
were, however, not very clear in all cases. In the case of certain species such
as A.polyacantha and A.robusta the nocturnal movements are very conspicuous and
specific (see Fig. 125).
A hypothesis was postulated concerning the evolution of the inflorescence
systems and inflorescences of the genus Acacia. The hypothesis includes a
theory concerning the development of the involucel (cupula). Flowering dates
of the different Acacia species are determined to a large extent by the morphology
of the inflorescence system. Date of flowering can therefore be considered
as a genetic character which nevertheless can be influenced to a certain
extent by ecological conditions.
The development of the ovule and the ontogeny of the seed was investigated.
Ideas were put forward as to the possible homology of the ovule. A study of
seed ontogeny produced information concerning the presence of endosperm in
mature seeds of A.kirkii, A.tortilis and A.xanthophloea. This characteristic
cannot be used as a criterium to remove these three species, as well as other
exotic species containing endosperm in the seed, out of the genus Acacia (cf.
Vassal, 1968).
The arrangement of vascular tissue in the seedling, leaves and young shoots
are discussed. Judging from the arrangement of the primary vascular bundles in
the seedling there is probably a relationship between a monolacunar node with two
leaf traces, a trilacunar node and a node which is called the "fifth type" by
Takhtajan (1969).
It may be concluded that the extant South African taxa of the genus Acacia
can be distinguished from each other on a specific level. By using variable
characters such as degree of pubescence and numerical taxonomic characters such
as seed and leaf size it is possible to carry the subdivision of species too far.
It is therefore recommended that the taxa with an extensive distribution,
manifesting considerable variation, such as A.karroo Hayne, A.caffra Willd.,
A.reficiena Wawra (= A.luederitzii Engl.) and A.robusta Burch., should not be divided
into smaller taxa until more information is available concerning the abovementioned
characteristics.
As a final summary, a key has been composed in which as many characters as
possible have been used to distinguish between the South African Acacia species.