Bradleya 31/2013
page 58-75
A new morphological study of the genus Trichodiadema
(Aizoaceae) permits the description of a new subgenus, T. subg.
Gemiclausa.
H.E.K. Hartmann & I.M. Niesler
Biozentrum Klein Flottbek, Ohnhorststr. 18, 22609 Hamburg, Germany
E mails: hartmann.africa@uni-hamburg.de and Ingeborg.Niesler@uni-hamburg.de
Summary: Based on earlier work on the construc-
tion of the diadem, the name-giving feature of the
genus Trichodiadema, the results of fresh inves-
tigations with a focus on plants that lack “true di-
adems” are presented. The possession of stiff
prickly bristles can be understood to form the
probable synapomorphy of all members of the
genus, the main chemical component being
suberin. More extensive studies of flowers and
capsules permit a subdivision of the genus into
two subgenera, and T. subg. Gemiclausa is de-
scribed here as a new subgenus. A key to the six
included species is given, and all species of the
new subgenus are described.
Zusammenfassung: (H.E.K. Hartmann) Auf der
Grundlage von früheren Arbeiten zum Bau der
Diademe, die der Gattung Trichodiadema ihren
Namen gaben, werden hier die Ergebnisse neuer
Untersuchungen vorgestellt, die sich auf Pflanzen
konzentrieren, welche keine typischen Diademe
ausbilden. Der Besitz von steifen stechenden
Borsten kann als Synapomorphie für die Gattung
angesehen werden, wobei Suberin als eine der
Komponenten bestätigt werden kann. Weiter re-
ichende Untersuchungen an Blüten und Früchten
erlauben die Teilung der Gattung in zwei Unter-
gattungen, von denen T. subg. Gemiclausa hier
als neu beschrieben wird. Ein Schlüssel für die in
die neue Untergattung eingeschlossenen Arten
wird gegeben, und diese Arten werden
beschrieben.
Introduction
When Schwantes (1926: 187) created the name
Trichodiadema for a new genus, he based the
taxon on the possession of a “Borstenschopf”, i.e. a
tuft of bristles at the leaf tips. Eight species
names were listed by Schwantes (T. stelligerum,
T. stellatum, T. densum, T. intonsum, T. se-
tuliferum, T. bulbosum, T. barbatum, and T.
58
Photographs by the authors
mirabilis, in this sequence). The genus name was
only validated by Schwantes in 1927 (106) when
he selected T. stelligerum (placed today as a syn-
onym under T. barbatum) as the “Leitart”, i.e. the
type species.
All eight species possess what Ihlenfeldt
(1980) called “Haarapparat” or diadem proper; he
suggested that only those species should be placed
in the genus that develop such a diadem. The typ-
ical elements of this particular diadem, namely
the apical cone derived from a secondary phel-
logen and the formation of cup cells around the
bases of the bristles have only been identified and
listed by Niesler (1997) as essential properties of
a diadem. She was also the first person who in-
cluded the older studies of Oberstein (1913) and
Weingart (1935), who both had described and fig-
ured the peculiar organs at the leaf tips and had
also confirmed that they contained suberin by ap-
plying Sudan III.
Niesler & Hartmann (2004) were able to
demonstrate that the so-called diadems proper de-
velop at different degrees of activity: the low ac-
tivity line results in two different types, simple
diadems and erect diadems, and the high activity
line in two diadem types, both with radiating bris-
tles, one orange with bristles pointing into all di-
rections, the other with brown to black cup-cells
leaving a central area free of bristles (lists of the
species in Niesler & Hartmann 2004: 63).
In completely independent actions, Bolus de-
scribed several species of Trichodiadema from
May 1928 onwards that included statements like
”apice 2-3 setis erectis albis brevissimis onusta,
rarius ebarbata” (T. occidentale L.Bolus 1928:
150, May) [apically adorned with 2-3 white very
short bristles, more rarely without a beard]. In
none of the descriptions, Bolus gave a reason why
she included the taxon in Trichodiadema, and in
a key to subgroups she distinguished six (Bolus
1963: 17) or eight (Bolus 1966: 203) such groups,
Bradleya 31/2013

Figure 1. The branch points to the rhs, the
younger white bundles of apical bristles in-
dicate that they are not yet stiffened; older
leaf tips from the node below on the lhs
possess orange-brown leaf tips prickly to
the touch. Hartmann & Bayer 34639
(HBG!).
Figure 2. A close-up of a tip shows the or-
ange colour at the bases of the bristles
which themselves are still white. Hart-
mann & Bayer 34499 (HBG!).
Figure 3. With age, the group of bristles
appears more clear-cut at its base. Hart-
mann & Bayer 34639 (HBG!).
Figures 1-5. Leaf tips with tufts of bristles in T. occidentale. Images by I. M. Niesler, except the image
for Figure 5 by U. Meve.
Figures 6-9. SEM-graphs of leaf tips. Images by I. M. Niesler.
Bradleya 31/2013
Figure 4. The median longitudinal section
through the leaf tip reveals that the dark Figure 7. On an older leaf, the tips starts
green assimilating tissue leaves a central withering, all bristles begin to shrink.
chimney free in the area below the bristles Hartmann & Bayer 34499 (HBG!).
with thick outer walls coloured orange
from suberin; some bristles have broken
away in the process of preparation, sup-
porting the interpretion that the tufts are
locally delimited. Hartmann & Bayer
34499 (HBG!).
Figure 8. In a typical diadem, the basal
cambium can be recognised as a straight
transverse line, separating the inner tis-
sues from one another. The bristles are
lifted by the central cone. T. subg. Tri-
chodiadema. Hartmann 34413 (HBG!).
Figure 5. Staining with Sudan III permits
the discrimination of the thick outer epi-
dermal walls of the bristles; note the cen-
tral bristle with a broken tip, supporting
the impression of stiffness. Hartmann &
Bayer 34639 (HBG!).
Figure 9. The central cone lacks a cam-
bium but develops cells in longitudinal
Figure 6. The basal boundary of the bris- rows. The bristles insert along the cone
tles can be traced as a wavy line, but the area. Hartmann & Bayer 34499 (HBG!).
differentiation is very clear. Hartmann &
Bayer 34639 (HBG!).
59
one with soft hairs, named Strumosa, three under
the heading “Leaves conspicuously tipped with
bristles” comprising four (1963) or five (1966) sub-
groups, and another single group under “Leaves
not, or very obscurely, tipped with bristles”. No
further arguments to support the groupings were
supplied. In the relevant species descriptions,
however, the bristles are described as being in-
conspicuous, short, and white, leaving the reader
wondering why they were placed in the bristle-
tipped genus Trichodiadema, defined commonly
as being characterized by coloured apical diadems.
In order to investigate this particular group of
species kept in the genus Trichodiadema since
they were described, extensive field work was un-
dertaken 2004-2006, in addition to older field
studies since 1971, making possible a new assess-
ment regarding the composition and delimitation
of Trichodiadema. In this paper, the focus lies on
material that lacks a diadem in its typical con-
struction. The results permit the establishment of
a new subgenus, T. subg. Gemiclausa H.E.K.Hart-
mann & Niesler, for which a key to the species is
given, and the taxonomy of all included species is
presented.
The formal treatment of the remaining species
of the genus will be published in another paper.
Material and methods
All type material of Trichodiadema has been
studied and reconfirmed in JSTOR in 2012. 428
populations have been investigated in the field
since 1971. Material has been collected and also
transplanted to the greenhouse in Hamburg,
where investigations under the light microscope
were undertaken with a ZEISS 473028; binocu-
lars of ZEISS 475052-9901 were used to draw cap-
sules and to photograph leaf tips; flowers were
documented with Leica DFC 295, CanoScan
5000F, and fine structures were studied in SEM
Philips SEM 505, SEM XL 20, and SEM FEI
QUANTA 250. Herbarium material and a collec-
tion of fruits are kept at HBG.
Analyses of critical features
1. Leaf tips
1.1 A historical review
When Schwantes (1926: 187) first described
the genus Trichodiadema (validated only by
Schwantes 1927: 106, when the type species was
named as “Leitart”), he called the terminal stiff
hairs of the leaves a “Borstenschopf”, i.e. tuft of
bristles and also a tuft of hairs, “Haarschopf”.
Petiver (1709: t. 77, catalogue number 484) de-
scribed and figured a plant of this group as” fol.
apicibus hirtis” [lvs with hairy tips]; Bradley
60
(1716: Dec. I: 6, t. 5) said in his English text
“Leaves are adorned at their points with star-like
thorns, very white and tender” (this plate has
been cited by Miller (1768, nr. 14) and has been
designated as the lectotype of T. stellatum), and
Dillenius (1732: 245, t. 190, f 234) referred to
“folia... in extremitate spinulis (quinque, sex
septemve stellatim dispositis) radiata” giving the
name as “Mesembryanthemum radiatum, ramulis
prolixis, recumbentibus”. Linnaeus (1753: 482)
was the first author to apply the epithet “barba-
tum” (bearded), adding “Foliis apice barbatis”
[with leaves apically bearded]. The term “Bar-
bata” has later been used for a Section (e.g. Salm
Dyck 1836: § 52) in the group “Papillosa”, but the
term “diadem” for the peculiar leaf tip was only
created by Schwantes in his choice of the genus
name (1926: 187).
The unusual fact that yellow to orange or
brown or even black prickly papillae form the
apex of the leaves has initiated several studies in
the anatomy and physiology of the “tufts of bris-
tles”. Oberstein (1913) was the first author to
demonstrate that these bristles contain suberin in
their walls by applying Sudan III (p. 300) and to
report that a cork cambium is formed at the base
of the diadem (his Fig. 7), which latter he classi-
fied as an emergence (“Emergenz”). Weingart
(1935) reported that sclerenchyma was not trace-
able with aniline sulphate (p. 46) in his material.
Ihlenfeldt (1980: 149) mentioned the deposition of
tannins in the walls of the papillae and in the cup-
cells without giving evidence but reports a suberin
layer in the inner wall of a bristle.
1.2 New results
Fresh investigations of recently collected ma-
terial permit a new assessment of leaf tips, espe-
cially those of the plants now placed in T. subg.
Gemiclausa. As a rule, leaf tips of these plants
possess a bunch of exceptionally big and elongated
bladder cells at their tips, white at first and turn-
ing orange to brown with age (Figure 1) while the
leaf itself is still fully turgescent and green. The
change in colour through time can be seen even
on one branch (Figure 1), in habitat and in culti-
vation; it remains visible on herbarium specimens
as well, depending on the state the material was
dried in.
In habitat, the plants show regularly orange
to brown apical tips, prickly to the touch when old
(Figure 1). In younger leaves, the tips are bundled
but translucent or white (Figure 1, upper right
hand side top), a feature apparently more often
found in greenhouse cultivation, suggesting that
more water and less sunlight might result in a
slower formation of colouring components. Char-
Bradleya 31/2013

Figure 10. In the freshly opened flower, the filamentous sta-
mens have already bent outwards showing clearly that they are
longer than the stamens. T. mirabile. Hartmann 34519 (HBG!).
Figure 11. In a longitudinal section through the flower, the tu-
bular arrangement indicates that a young flower was cut, yet
the filamentous staminodes are already distinctly longer than
the stamens. T. peersii. Hartmann 34520 (HBG!).
Figure 12. In T. littlewoodii, the numerous long filamentous
staminodes merge from the stamens into the petals, making a
distinction difficult. This combination is typical of the species.
Hartmann 34331 (HBG!).
Figures 10-15. Flowers of members of T. subg. Trichodiadema. All images by I. M. Niesler.
Bradleya 31/2013
Figure 13. The transition between the floral elements is more
evident in the longitudinal section; visible are also the well-de-
veloped diadems at the calyx lobes tips. T. littlewoodii. Hart-
mann 34331 (HBG!).
Figure 14. The possession of filamentous staminodes longer
than the stamens places the flower of T. attonsum definitely in
T. subg. Trichodiadema. Hartmann et al. 25853 (HBG!).
Figure 15. In principle, the longitudinal section confirms the
typical morphology, but here fewer elements can be recognised.
T. attonsum. Hartmann et al. 25853 (HBG!).
61

Figure 16. Flowers of T. calvatum exhibit the darkest pink in
the subgenus, the darker colouring over the central vein can
hardly be distinguished. Hartmann & Bayer 34621 (HBG!).
Figure 17. The stamens form a central low cone, at their bases
the papillae appear as interwoven, few pointing outwards; note
the pink colouring at the tips of the filaments, a common fea-
ture in flowers of T. subg. Gemiclausa. T. calvatum. Hartmann
& Bayer 34621 (HBG!).
Figure 18. Bright pink dominates also the petal colour in T. oc-
cidentale, but it fades out to a white base while the central deep
purple line appears more prominent (vittate flower). Note some
filamentous staminodes on the rhs. Hartmann & Bayer 34639
(HBG!).
Figures 16-21. Flowers of species in T. subg. Gemiclausa. All images by I. M. Niesler.
62
Figure 19. In the longitudinal section of an older flower, the
white spreading papillae on the filaments and filamentous sta-
minodes form a dense velcro-like cover around the actual cen-
tre. T. occidentale. Hartmann & Bayer 34639 (HBG!).
Figure 20. Purple is restricted to the basal veins of the apically
apricot-coloured petals and to the tips of the stamens and fila-
mentous staminodes. T. gracile. Hartmann & Bayer 34637
(HBG!).
Figure 21. In an older flower, the papillae are more promi-
nent again, and the outer elements start spreading a little. Yet,
the constant shape of the low cone flower is pertained. T.
gracile. Hartmann & Bayer 34637 (HBG!).
Bradleya 31/2013

Figure 22. Flowers of T. pygmaeum agree in the colour pattern
with those of T. gracile. Bayer 7282 = Hartmann & Bayer 34636
(HBG!).
Figure 24. T. strumosum shares its flower colour pattern with
T. gracile and T. pygmaeum, although the colours are even
lighter. Hartmann & Bayer 34625 (HBG!).

Figure 25. In longitudinal section, the very long papillae on the

outer surface of the calyx dominate the picture. The central cone
Figure 23. In longitudinal section, these flowers resemble most comprises also some filamentous staminodes and many basal
closely those of T. occidentale. Bayer 7282 = Hartmann & Bayer papillae united into a layer of felt. Hartmann & Bayer 34625
34636 (HBG!). (HBG!).

Figures 22-26. Flowers of species in

T. subg. Gemiclausa. All images by
I. M. Niesler except that for Figure
26 by M.B.Bayer.

Figure 26 An overview in habitat demonstrates the variation in colouring of petals mainly as

shown by the very young flowers near the bottom rhs in an almost orange colour while younger
flowers on a lant at the lhs are lighter from the beginning. Hartmann & Bayer 34625 (HBG!).

Bradleya 31/2013 63
acteristic is the gradual development of the
colouring of the bristles (Figure 2), but never ex-
tending further down than the tip area (Figure 3).
In habitat and in cultivation, these tips can break
off, especially when touched in handling a speci-
men (Figure 4), e.g. for preparation of herbarium
specimens. In a median longitudinal section, the
central tissue of the leaf continues upwards to the
very base of the tissue forming below and amidst
the bristles (Figure 4), while the veins end below
this region. This central apical cone might be only
parenchymatic since no colouring can be detected,
neither in a fresh section (Figure 4) nor in mate-
rial stained with Sudan III. This difference can
also be noticed when sectioning the leaf tip longi-
tudinally by hand: the hard bristle region pres-
ents more resistance to the movement than the
lower portion which cuts “like butter”. In agree-
ment with these observations, staining with
Sudan III reveals differently coloured layers in
the substantially thickened walls of the bristles
(Figure 5). In such sections, loosely lying calcium
oxalate crystals that have fallen out of cells with
crystal bundles can hide any thin-walled areas
(Figure 5).
In the SEM, the basal border of the tip area
can clearly be seen in very fresh leaves already
(Figure 6), and in older ones the tip has started to
shrink and whither in precisely that area (Figure
7).
A comparison of longitudinal sections in the
SEM demonstrates the differences between the
two types of leaf tips. In a diadem proper, the bris-
tles are lifted by a cone probably derived from a
secondary cambium stretching across the leaf tip
area just below the bases of the bristle bases (Fig-
ure 8), as described by Niesler & Hartmann
(2004). Oberstein (1913) reports a positive reac-
tion of this tissue with Sudan III, and the tissue
has been named a corky cone since. In species
without such diadems, the tips of the leaf show a
similar picture (Figure 9), but no transverse cam-
bium can be identified. Instead, somewhat elon-
gated cells in the interior of the leaf tip can
resemble cell arrangements of a corky tissue (i.e.
cells in vertical rows), and it is this region that
bears the coloured bristles described above (Fig-
ures 1-7).
Samples of T. occidentale show the described
features, and examination of the types of the
species T. gracile, including all three varieties,
possess the same arrangements; the type of T. cal-
vatum exhibits shorter and lighter coloured tips.
In the soft-haired species T. fergusoniae, T. pyg-
maeum, and T. strumosum, the tips colour much
more slowly, more often not at all, and the basal
64
boundary appears to be absent.
The least degree of colouring of leaf tips is
found in T. strumosum and T. pygmaeum, both
with soft papillae all over their leaf surfaces.
Anatomically, no distinct tuft of apical papillae is
formed, and the tissue of assimilating cells runs
around the tip as it does all around the leaf. Only
the light orange of three to four tip cells indicates
a similarity with the diadems proper.
All tufts of bristles are very prickly to the
touch and turn orange to black, depending on the
species. The diadems proper fall off more easily,
due to the cambium at their base, than the simple
tufts.
Cork can be shown to occur in the thick outer
walls of the bristles and in the adjacent much
smaller epidermal cells in all investigated sam-
ples, with or without a diadem proper. Tannin has
only rarely been detected. The question of compo-
nents in the elements of the tuft of bristles is still
in need of more studies on much more material.
2. Flowers
Following a survey of flower colours and pat-
terns (Niesler 1997), a re-examination of more
than 100 preparations of flowers was undertaken.
All populations with diadems proper possess long
filamentous staminodes overtopping the centrally
collected stamens (Figures 10-15). In typical sam-
ples of Trichodiadema subg. Trichodiadema, the
numbers of filamentous staminodes are probably
specific for a species: compare fewer elements in
Figures 10 and 11 against many in Figures 12 and
13. In the beginning of the anthesis, the stamens
and filamentous staminodes form narrow tubes
(Figures 11, 13), towards the end of the flowering
period the filamentous staminodes spread widely
(Figures 10, 12).
This type of flower has also been found in T.
attonsum and T. imitans - both until now excluded
from true Trichodiadema (Figures 14, 15) - be-
cause the central elements form a tube, their tips
elongating and spreading soon.
In contrast to this group of species, all other
populations that lack diadems proper develop
short filaments and filamentous staminodes (Fig-
ures 16-26) in an arrangement very similar to that
described as “short cone type” in the genus
Drosanthemum (Rust et al. 2002). The central el-
ements stand in a broad cone with oblique sides
when the flower opens for the first time (Figures
17, 19, 21, 23, 25, and 26). This cone remains in
shape for several days, the outer elements begin
to spread tardily thereafter (Figures 18, 19, 21, 23,
25, 26) while the petals open and close daily. In
younger flowers, the outer elements of the central
low cone stand clearly apart (e.g. Figures 18, 22,
Bradleya 31/2013
Figure 27. The two closing bodies lie so
close to each other that they seem to form
a single body. Bayer 7177 = Hartmann &
Bayer 34621 (HBG!).
Figure 30. The two closing bodies shown
transversely here are separated by clefts
in the top layer of tissue. Hartmann &
Bayer 34717 (HBG!).
Figure 33. On top of the transverse layer
of expanding tissue lies the placenta, the
swollen tissue below represents the closing
body that lifts the layer above. Bayer 7177
= Hartmann & Bayer 34621 (HBG!).

Figure 31. While the closing bodies are

flattish, the dividing cleft is deepened in Figure 34. A relatively thick closing body
this case. Hartmann & Bayer 34621 is covered by a relatively low layer of ex-
Figure 28. The closing body at the lhs pro- (HBG!). panding tissue. Hartmann & Bayer
trudes further than the smaller swelling at 34626 (HBG!).
the rhs, both ending below the funicles
placed above in the picture. Bayer 7177 =
Hartmann & Bayer 34621 (HBG!).

Figure 32. The well-developed closing

bodies are separated by a broad cleft, The Figure 35. The swelling of the closing bod-
thread-shaped small elements are hyphens ies in the centre peters out towards the
of a fungus. Hartmann & Bayer 34626 front of the section. Note also the hyphens,
(HBG!). as in Figure 32, but in a different popula-
tion here. Hartmann & Bayer 34621
(HBG!).
Figure 29. In the lower middle, two
swellings as closing bodies can be seen
below the placenta in the centre. Hart-
mann & Bayer 34621 (HBG!).
Figures 27-32. Different forms of the closing bodies in SEM, all samples from T. clavatum. Images by I.
M. Niesler. Figures 33-35. Internal structures in closing bodies. T. calvatum. Images by I. M. Niesler.

24, 25), especially clear in the filaments; in most
older flowers, however, a rich development of
papillae can be observed in the lower portion of
the outer elements of the cone, visible as protrud-
ing tiny white fingers sticking together like Vel-
cro surfaces, even felt-like in some cases (Figures
17, 19, 21, 25).
This phenomenon of many densely interwoven
papillae on the outer elements of the central mem-
bers of the androecium has also been observed in
urn-shaped flowers of Amphibolia (Hartmann
1996, 1998; 2001d). Care should be taken not to
confuse the situation with the normal equipment
of stamens with some papillae in the same area,
e.g. also in Trichodiadema subg. Trichodiadema
.

Bradleya 31/2013 65
3. Fruit characters
3.1 Closing bodies
Typical members of the genus Trichodiadema
lack closing bodies (Niesler 2001) but in the de-
scriptions of the six species without a diadem
proper, the presence of closing bodies is men-
tioned. (e.g. in T. calvatum, Bolus 1936). Exten-
sive studies in these six species reveal that they
all share certain features that have, until now, not
been described.
All closing bodies investigated in Trichodi-
adema subg. Gemiclausa consist of two portions,
each placed on the interior of the valve at its edge
to the locule. In an early state of fruit ripening,
the two portions are placed very close to each
other, suggesting one uniform body (Figure 27),
but in another capsule of the same population, a
really solid body can be found (Figure 28) show-
ing also two unequal protrusions to the sides of
the centrally placed placenta. In a young fruit, the
two swellings lie close together (Figure 29), and in
an old fruit (Figure 30), the tissue splits along the
middle of the valve, pushing the two portions of
the placenta apart. The described differentiation
in time is demonstrated here to show that this
type of a twin closing body is not as constant as
other types of closing bodies, e.g. in Argyroderma,
Leipoldtia, Glottiphyllum, or Pleiospilos in which
samples the two halves of the valve are solidly
grown together, always forming a single closing
body of constant shape in a fruit, a population,
and a species, or even in a genus. Closing bodies of
this compact type consist clearly of endocarpal
protrusions derived from tissue underlying the
placenta, often lifted by it (e.g. in Pleiospilos and
Tanquana, Hartmann & Liede 1986: 433-485), i.e.
deeper inside the locule reaching upwards.
In some samples of one and the same species of
T. subg. Gemiclausa, a deep fissure exists from
the beginning between the two closing bodies (Fig-
ure 31), but in yet another population, a less
prominent cleft like a cut is visible (Figure 32).
The different states shown here occur in one and
the same population, even in a single plant or,
most impressive, in a single fruit (Figures 37, 39,
40).
Important is the fact that two distinct closing
bodies can be recognised. This seems to indicate
that the closing bodies may in this case be derived
from placental tissue, as has been found in mem-
bers of the genus Malephora. Sections through the
critical area, however, reveal an entirely different
composition of the tissues. The placenta lies on top
of a layer of cells consisting of elongate vertical el-
ements, indicating that this layer is most probably
part of the expanding sheet (Figure 33). Diving
66
below the placenta, these cells get lower suggest-
ing that they lose their expanding properties.
Below the highest cells lies a conglomerate of
thick-walled tissue, the endocarpal portion of one
of the closing bodies lifting the placenta a little in
this place. A similar situation from a different
population permits a view of the differentiation in
internal tissue in such a closing body (Figure 34),
but a section in still another plant reveals that
closing body material can also lie next to the ex-
panding tissue and differ in internal cell wall
thickenings from the other two forms of closing
body (Figure 35).
What is described here for samples from T. cal-
vatum can be seen in all six investigated species of
Trichodiadema with closing bodies, in all its vari-
ation even within a single fruit (Figures 37, 39,
40). Two aspects are of critical importance:
Firstly, the enormous variation in closing body
shapes, and secondly, the fundamental formation
of twin swellings per valve below the endocarpal
epidermis of the valve in the region that usually
forms the proximal portion of the expanding sheet
(Figures 37-41). Typical closing bodies incorporate
the endocarpal layers of the outer locule wall (e.g.
in Pleiospilos, Hartmann & Liede 1986: 433-485),
a feature absent in the newly discovered “hidden
closing bodies”.
In summary, its appears as if the potential to
develop local swellings on the lower inner side of
the valves (and not at the upper edge of the locule
as a distinct superficial body) represents a special
type of closing body development that has not
been found elsewhere in the Ruschieae up to this
date.
3.2 Covering membranes
The development of closing bodies is function-
ally related to the formation of covering mem-
branes (Hartmann 1988): closing bodies can only
block the distal exit of the locule if some other el-
ement covers the remaining surface of the locule.
Two aspects must be remembered, however:
firstly, covering membranes can be formed inde-
pendently from closing bodies (e.g. in the Lam-
pranthus type of fruit in which funicles block the
exit in place of closing bodies, Hartmann 1988)
and, secondly, the expulsion of seeds takes place
through the nozzle-like openings in the centre of
the fruit, not through the distal exit (Hartmann
1988). Fruits with closing bodies can, therefore,
be expected to possess covering membranes of a
solid texture and morphology in order to meet the
functional requirements, i.e. resilience of the cov-
ering membrane and a somewhat firmer distal
edge touching the closing body.
These conditions are fulfilled for samples from
Bradleya 31/2013
Figure 36. The open capsules shows that
closing bodies are lacking and that reduced Figure 38. In T. fergusoniae, the type
covering membranes leave the numerous specimen exhibits prominent twinned clos-
funicles open to the view. The transparent ing bodies of almost equal shapes. Fergu-
covering membranes permit the conclusion son 18822 (BOL!). Figure 40. T. calvatum produces also most
that they are derived from mesocarp, indi-
different shapes of closing bodies in a sin-
cating also that they are not resilient but
gle fruit, as has been demonstrated in fig-
fall into the empty locule when old. Note
ures 27-35. Hartmann & Bayer 34468
further that the expanding keels lie paral-
(HBG!).
lel and in close contact, leaving no endo-
carp free to develop covering membranes.
T. rogersiae. Rogers 2686/29 Type speci-
men (BOL!).

Figure 39. The open capsule of the type

specimen of T. occidentale represents a
sample of great variation in a single fruit
and almost complete covering membranes.
Smith 5140 (BOL!).
Figure 41. Open capsules of T. gracile
vary less than those of the latter two
Figure 37. Like all members of T. subg. species and within a single fruit. Hart-
Gemiclausa, the open fruit of T. strumo- mann & Bayer 34637 (HBG!).
sum possesses twin closing bodies and per-
manent resilient covering membranes,
derived from endocarpal tissue. Note that,
therefore, the expanding keels diverge
from the base leaving space in the middle
of the valve from which the covering mem-
branes have been formed. Hartmann &
Bayer 34692 (HBG!).

Figures 36-41. Open capsules. Drawings by I. M. Niesler.

all six species with closing bodies: the two parts
covering a locule are each more or less rhombic in
contour (Figures 37-41). The outer two sides are
usually a little shorter than the inner, and the
shortest extension is seen over the middle of the
locule as well as over the wall between the locules;
the distal indentations can be v-shaped (Figures
37, 41), almost straight (Figures 39, 40) or
rounded, resulting in a wavy fringe resembling a
star-shape with ten rays (Figure 38).
Such extensive distally almost fimbriate cov-
ering membranes have also been found in samples
from some species of Trichodiadema lacking clos-
ing bodies, namely T. densum and T. marlothii
with 6-7 locules plus T. fourcadei, and in T. burg-
eri with five locules, all showing inconspicuous in-
dentation. The majority of species of
Trichodiadema without closing bodies, however,
possesses narrow covering membranes of thin con-
sistency probably derived from the interior meso-

Bradleya 31/2013 67
carp of the valve because the membranes are
translucent and can change in shape, depending
on the amount of seed below (Figure 35). The pair
of covering membranes derived from two adjacent
halves of a locule can be oval in outline or heart-
shaped, sometimes angular at the outer side. As
Hartmann (1988) was able to show such covering
membranes are ontogenetically correlated with
the development of expanding keels along the
middle line of the valve, placed close to each other
and hence using up the available endocarpal tis-
sue of the valve. In such a case, the endocarp is
not available to form a covering membrane.
A caveat must be expressed: while the corre-
lation between the closing bodies and the cover-
ing membranes is a functional one, the correlation
between the development of covering membranes
and the expanding keels is an ontogenetic one
since both products need material from the endo-
carpal and mesocarpal layers of the valve.
4. Roots
Most of the 428 populations studied in the field
possess some type of storage roots, either as solid
thickened taproots or with swellings of the far-
reaching only somewhat thickened roots, the
swellings resembling potatoes in appearance, also
called “veldaartappels” and eaten as “veldkos”.
Turnip-shaped taproots are found in all members
of T. subg. Gemiclausa, albeit in different shapes.
In T. occidentale, the irregularly formed thick tap-
roots adapt to the space available between stones
and rock (Figure 45); they can also divide and fill
gaps between rocks in this way. T. gracile, how-
ever, seems to grow in looser soils where it can
form spindle-shaped taproots of considerable
length (up to 18 cm) (Figure 42). The dug-up
plants of T. calvatum resemble a tree in shape due
to a taproot with straight sides, sometimes
branched and usually splitting at the base (Fig-
ure 44).
The roots of the three species of T. subg. Gemi-
clausa with soft hairs are much thicker than those
of the other species, but they break more easily
and are usually fleshy. Most remarkable is T. pyg-
maeum with the biggest storage root (e.g. 12 cm
long and 6 cm thick, Figure 46) found in Tri-
chodiadema and with numerous adventitious
roots to 25 cm long from the numerous pla-
giotropous shoots that present the aspect of a
dense mat. Older branches die back, the above-
ground portion of the plant appearing like a thick
felt in which fine soil can be caught, supplying
new material for further growth. The species was
described from such material; the thick taproot
was unknown at the time. T. strumosum, how-
68
ever, is only known with spindle-shaped roots
(Figure 43).
Storage taproots are rare in T. subg. Trichodi-
adema (e.g. in T. densum); much more common
are widespread root systems that grow often to
more than 50 cm depth and also to the sides. All
roots show some degree of thickening, and
swellings are also common on them. In contrast to
the taproots, these swellings are seasonal or tem-
porary since they can be found on older roots as
hollow shells, mostly broken, most often at the be-
ginning of the rainy season. Some species develop
perpendicular thickened adventitious roots which
can form bundles, and among these, some can also
recycle their contents and leave empty shells be-
hind.
5. Discussion
It is easy to feel the stiffness of the apical bris-
tles on the leaves by hand, leaving no doubt about
the position of such material in the genus Tri-
chodiadema. Nevertheless, it has up to now not
been possible to give evidence which components
cause such prickliness. It seems unlikely that only
cork in the outer bristle walls can result in such
distinct differentiation, and it has been impossi-
ble to proof until now that tannins play a major
role. Tannins are well known to occur in tannin
idioblasts in the interior water tissue of many
Mesembryanthema, but so far no deposition in
walls of the tufts of bristles has been documented.
In consequence, and considering the fact that
in no other taxon of the Aizoaceae prickly leaf tips
have been found, it appears adequate to keep all
species described as members of Trichodiadema
in one genus, based on the observation that all
bristles develop very thick outer walls that stain
with Sudan III (Figure 5).
Common to all samples is also that the conical
inner centre of the tuft of bristles is regularly en-
larged, demonstrated best by the “corky cone”, de-
rived from a transverse cambium from which the
descendants grow in vertical rows, as is typical of
cork (Figure 8). In leaf tips without typical di-
adems, cells lifting the bristles lie also in rows re-
sembling cell arrangements derived from a
cambium, but if such cambium has been there, it
has been exhausted quickly, leaving no distinctive
transverse layer behind. The product does not
stain with Sudan III, suggesting that the tissue
remains more or less parenchymatic (Figure 9).
It can be speculated that the simpler con-
struction of the tuft of bristles may be primary in
the evolution, and that the development of a com-
plicated pattern like the true diadem may be de-
rived, as is also suggested by the fact that
Bradleya 31/2013
Figure 42. Long spindle-shaped
taproots bear erect branches at
their tips that elongate and bend
over with continuous growth over
several years in T. gracile. Hart-
mann & Bayer 34637 (HBG!).
Figure 43. The Figure 44. The storage tap- Figure 45. Shapes of the storage roots in
erect branches roots of T. calvatum are T. occidentale are variable, depending on
from a spindle- unique in the subgenus, dug space that is available in the stony-shaly
shaped taproot are up they resemble a tree in ground. While the above-ground parts of
similar to those of shape with their perpendicular the plants are very similar to those of T.
T. gracile. Hart- sides. Hartmann & Bayer calvatum, the taproots differ distinctly in
mann & Bayer 34468 (HBG!). shape and can be used for identification.
34625 (HBG!). Hartmann & Bayer 34633 (HBG!).

Figure 46. The by far biggest storage

root in the subgenus has been found
in T. pygmaeum which is also unique
in forming a mat of fine plagiotropous
roots forming a mat of up to 40 cm in
diameter in which fine material can
be caught, offering new soil for the
mat. Hartmann & Bayer 34636
(HBG!).
Figure 47. Distribution of all studied populations of Trichodiadema. In light green,
T. subg. Trichodiadema, in purple T. subg. Gemiclausa. The latter occurs only in the
south western part of the entire range, overlapping only in some areas with the very
wide-spread subgenus Trichodiadema. Copyright: H.E.K. Hartmann.

Figures 42-46. Thick roots are characteristic of all species of T. subg. Gemiclausa. All images in habitat,
H.E.K. Hartmann.

Bradleya 31/2013 69
different types of diadems from simple to highly
complex forms can be observed (Niesler & Hart-
mann 2004), visible best in the various shapes of
cups around the bristle bases.
In two character sets, clear boundaries can be
described, namely in flowers and fruits, separat-
ing two groups of species that are recognised as
subgenera here.
Flowers correlated with the occurrence of true
diadems show always a narrow tube of filamen-
tous staminodes overtopping the stamens and
spreading during anthesis (Figures 10-15) AND
lack closing bodies in the capsules (Figure 36).
The same flower and fruit types occur also in
three more species: T. attonsum, T. imitans, and
T. fourcadei, which, however, lack diadems
proper.
In contrast, capsules with closing bodies are
always correlated with the presence of flowers of
the “low cone type” (Figures 16-26). Variable in
such flowers are the numbers of filamentous sta-
minodes, but constant is the central cone with nu-
merous papillae near the bases of the elements,
developing a dense felt during anthesis.
Capsules of plants with true diadems never
possess closing bodies and form incomplete cover-
ing membranes from the mesocarp (Figure 36).
The closing bodies develop by forming swellings
at or below the expanding tissue, mostly one per
half valve, but sometimes as a single body (Fig-
ures 27-35). In contrast to other shapes of closing
bodies, e.g. in Glottiphyllum or Pleiospilos, these
“closing twins” vary in details widely, indicating
that any blocking at the distal side of the locule
is, in combination with the complete covering
membrane, most probably an improvement in
seed dispersal.
Whereas the stiff prickly bristles present a
possible synapomorphy for the genus Trichodi-
adema, character expressions of flowers and cap-
sules can be understood to reflect a subdivision of
the genus into two subgenera: T. subg. Trichodi-
adema and T. subg. Gemiclausa, subg. nov., de-
scribed below, named in allusion to the twin
shaped closing bodies.
A survey over the geographical distribution of
the studied populations (Figure 47) reveals that
members of T. subg. Trichodiadema occur over a
wide area in regions of different rainy seasons and
different amounts of precipitation, hardly reach-
ing into the winter rainfall area. Here, however,
at the edge and near the Cape Floral region, the
members of T. subg. Gemiclausa are found, in
winter rain or just further east, expecting rain at
all seasons. In this particular area, the plants
occur in the vicinity of members of Drosanthe-
70
Key to the subgenera of Trichodiadema
1. Capsules always without closing bodies; flow-
ers with long filamentous staminodes overtop-
ping the stamens, both forming a central bundle
T. subg. Trichodiadema
- Capsules with twin or bifid closing bodies;
flowers without filamentous staminodes or of
the low cone type, i.e. wider than high and con-
stant in shape during anthesis
T. subg. Gemiclausa
mum, especially with those of D. subg. Ossicula
characterised by bone-shaped twin closing bodies
(Hartmann 2009), and with populations of
Acrodon and Brianhuntleya as well (Hartmann &
Niesler 2012), most of these with flowers of the
low cone type and all of them with closing bodies,
distinctly bifid in the last two taxa. The concen-
tration of low cone flowers in this region is best
understood as having been promoted by certain
pollinators, although obviously no hybridisation
takes place as all species involved differ in other
fundamental features. Such agglomeration of sim-
ilarities in flower structure has been observed in
other areas as well, e.g. the spirally unfolding
flowers of deep purple petals in members of Enar-
ganthe, Namaquanthus, and some species of
Cheiridopsis in an area in the southern
Richtersveld.
Taxonomy of Trichodiadema and the new
subgenus T. subg. Gemiclausa H.E.K.Hart-
mann & Niesler
Trichodiadema Schwantes ZSK 1927: 106
Typus T. stelligerum (Haw.) Schwantes = T. bar-
batum (L.) Schwantes Etym Gk trix, hair, di-
adema, crown, allusive to the leaf tip consisting of
tufts of bristles
Erect to decumbent or creeping perennial
shrubs, compact, caespitose or with erect or climb-
ing branches, 4-60 cm tall, roots always thickened
to some degree, forming thick taproots or spindle-
shaped storing-roots, more rarely with adventi-
tious roots, single or as a bundle, or long thinner
roots developing storing tubers like potatoes al-
most 50 cm below the soil surface; internodes
papillate when young, the papillae visible as
white dots, denser below the node, older intern-
odes often smooth; leaves trigonous, the epidermis
with densely arranged huge often thick-walled
bladder cells in rows, their shapes roundish with
or without protrusions in central position or point-
ing upwards or downwards at the ends, giving the
leaves a rough touch resembling a snake’s skin,
the margins basally sometimes with hairs, the
Bradleya 31/2013
 Key to the species of T. subg. Gemiclausa
1. Leaves soft from very long and dense hairs equally distributed over the entire leaf, only few orange
coloured apical cells 2
- Leaves with cobble-stone shaped epidermal cells, only in some species also central longer papillae
more or less loosely dispersed and forming a bundle at the tip 4
2. Branches forming finely rooting mats on the ground, the leaves 6-10 mm long and about 3 mm broad
and thick T. pygmaeum
- Branches erect to decumbent, without adventitious roots and well raised above the ground, leaves up
to 12 mm long and 2 mm broad 3
3. Hairs on leaves from a broad base looking densely villose T. fergusoniae
- Hairs on leaves from a small raised round centre, loosely placed T. strumosum
4. Flowers with a low cone-shaped centre, constant in shape during anthesis, few filamentous stamin-
odes, petals pink or apricot coloured in all shades 5
- Flowers without filamentous staminodes, stamens at first collected in the centre, later as a low erect
tube, petals deep pink to purple T. calvatum
5. Branches spindly, decumbent, and with long internodes T. gracile
- Branches erect with shorter internodes T. occidentale

leaf tips always with tufts of elongate bristles, ei-
ther in a prickly bunch or as a prickly diadem
with a basal cambium layer inside and cup-cells
around the bases of the bristles; flowers solitary,
in a monochasium, or in a cyme of 3, rarely 2-5
flowers, pedicels hairy or papillate like the in-
ternodes, or with smaller papillae, flowers either
with filamentous staminodes distinctly longer
than the stamens, all together forming a tube, or
of the low cone type with short filamentous sta-
minodes and stamens, calyx with (4-) 5-6 (-7)
lobes, the bladder cells elongate, especially at the
base of the calyx, or globular, lobe tips coloured
like the diadem or the tuft of bristles, petals in
shades of white, pink, and yellow, white towards
the base, either uniformly coloured or purple vit-
tate, filamentous staminodes rarely absent, sta-
mens in vittate flowers with many long papillae
in the lower half, protruding later prominently,
(4-) 5-6 (-7) lophomorphic dark green nectaries,
stigmas (4-) 5-6 (-7), flowering almost all year, in
most cases fully expanded at noon, sometimes in
the afternoon, fruit stalks often more prominently
papillate than the internodes, closing bodies ab-
sent in plants with diadems proper, correlated
with a pair of covering membranes derived from
one septum heart-shaped, rhombic, twin closing
bodies present in plants with low cone flowers;
seeds flattened roundish to ovate, light to dark
brown, in most cases the dorsal part darker in
colour, the perisperm region may even be trans-
parent, c. 0.8 mm long, 0.5 mm broad; chromo-
somes 2n = 18, 36.
Ecology: in fissures of rocks, in white quartz
fields, in stony to sandy ground, rarely in shady
localities, regularly browsed.
Distribution: SE part of Namibia; NC, NW,
WC, FS, EC, S Africa.
Note: The characteristic feature of the genus
is the development of long prominent prickly cells
- bristles - at the leaf tips, either as a simple bun-
dle of bristles or structured as so-called diadems.
The new circumscription of the genus Trichodi-
adema is based on the observation that the prickly
apical tuft of big elongate epidermal cells on the
leaves agrees in these properties with the diadems
found in T. subg. Trichodiadema. Until now, such
a formation of apical bristles has not been found
in any other genus of the Aizoaceae. The genus
can be subdivided into two subgenera differing in
their flower construction and in their fruit mor-
phology.
Trichodiadema subg. Gemiclausa
H.E.K.Hartmann & Niesler, subgenus novum
Type species: T. occidentale L.Bolus
Plants with very thick storage tubers, devel-
oping erect or decumbent, rarely creeping peren-
nial branches from the rounded underground
top of the hypocotyl, internodes papillate when
young or smooth and shiny from the beginning;
leaves trigonous, with cobble-stone epidermis or
equipped with central long papillae, in some
groups soft to the touch, the tips equipped with
bundled long papillae usually in erect position,
forming a prickly orange coloured tuft that can
eventually break off; flowers solitary, calyx base
with long hairs, petals deep purple vittate, the

Bradleya 31/2013 71
basic colours pink, shades of apricot or lighter
yellow, filamentous staminodes absent or few,
never longer than the stamens, both forming a
central low tube or a cone broader than high,
constant in shape during anthesis, the long
densely placed basal papillae on the filaments
or the filamentous staminodes developing an in-
terwoven felt; fruits with papillate stalks and
calyx base, valves with broad wings, bifid or
twin closing bodies at the mouth of the opening
of each locule, consisting of endocarpal thicken-
ings lying below or next to the expanding sheets.
Ecology: in fallow fields, in crevices of rocks,
expecting winter rainfall or rain at all seasons.
Distribution: only in the SW part of the area of
the genus, only marginally overlapping with the
area of T. subg. Trichodiadema.
Note: The subgenus is named after the pecu-
liar formation of twin swellings below the ex-
panding sheet, functionally closing bodies, but
until now not found in any other genus of the
Aizoaceae.
T. calvatum L.Bolus NM3 1936: 19 T Esterhuy-
sen 2747/35 (BOL!).
Taproot a fat trunk-shaped tuber, widening a
little to the top and splitting into erect branches
from there, each to 3.8 cm ∅, internodes grey and
soft, later orange and shiny, 10-18 mm long;
leaves with cobble-stone shaped bladder cells with
a short apical papilla, these papillae elongating to
the tip, where they form a slender bundle of bris-
tles fixed laterally at their bases, turning orange
with age, leaves 10-20 mm long, 2-3 mm broad
and thick; flowers with stiff papillae pointing up-
wards on their pedicels, 10-15 mm long, calyx base
with papillae longer than those of the pedicel,
curving upwards or spreading sideways, 30 mm
Ø, petals purplish-pink, filamentous staminodes
none, stamens in a central inconspicuous cone
turning into a tube later, short; fruit with nearly
complete covering membranes, the middle of the
open capsule star-shaped as seen from above, clos-
ing bodies bifid, sometimes split from the basal
split of the valve, covered by expanding sheet or
adjacent to it, closed 6.5 mm Ø. Ecology: in shales
of Ecca or Bockeveld, often in open spaces in
Renosterbosveld, rainfall mainly in winter.
Distribution: Caledon, Montagu, Robertson,
Swellendam, WC, S Africa.
Note: The species survives in remains of
Renosterbosveld in rather a wide area but is
threatened by agriculture. In habit, it looks most
similar to T. occidentale, differing from that
species in its trunk-shaped thickened taproot.
Further material seen: Gerbaulet et al. 23927
72
(HBG!); Hartmann & Bayer 34468, 34621, 34626,
34717 (HBG!).
T. fergusoniae L.Bolus NM2 1928: 9 T Ferguson
18822 (BOL!).
Tubers to 12 cm long, up to 2.5 cm Ø, with
erect to decumbent branches with visible intern-
odes; leaves apically densely arranged, bladder
cells with hairs from a rounded base, disorderly
dispersed over the leaf, resembling uncombed hair
up to the tip where some longer hairs protrude, 9
mm long, 2.5 mm broad, < 2 mm thick; flowers
with hairy pedicels, longer towards the calyx base,
calyx base hairy; fruit with twin closing bodies,
covering membranes triangular, those derived
from one septum forming a rhomb, closed 6 mm
Ø, open 9 mm Ø, 3 mm high.
Ecology: unknown,
Distribution: Riversdale, WC, S Africa.
Note: No fresh flowering material has been
available up to now, but the capsules are distinc-
tive. In the soft long hairs on the leaves, the plants
agree with those of T. pygmaeum and T. strumo-
sum.
T. gracile L.Bolus NM2 1929: 126 T Neethling
1998/24 (BOL!) = T. gracile L.Bolus var. piliferum
L.Bolus 1964: 78 T Ferguson 21440 (BOL!) = T.
gracile L.Bolus var. setiferum L.Bolus 1964: 79 T
Ridley 402/14 (BOL!).
Decumbent branches from a fusiform tuber to
10 cm long, 30 mm Ø in the middle, 15 mm near
the ends, smooth, internodes 10-15 mm long, 1-2
mm broad; leaf bladder cells with an apical and a
basal papilla, lateral basal papillae elongated,
laminal papillae reduced, the c. 6 apical bladder
cells with very long apical bristles, the whole long-
drawn tip orange, falling off with age, leaves 10-40
mm long, 3 mm Ø; flowers solitary, pedicels c. 15
mm long, with long spreading papillae, calyx base
covered with spreading stiff hairs, 18 mm Ø,
petals apricot-coloured or yellowish, deep purple
vittate, the centre forming a low cone type, fila-
mentous staminodes and stamens basally with
papillae; fruit apparently with a bulge of thick-
ened endocarpal tissue below or next to the ex-
panding sheet.
Ecology: very stony dry hillsides, limestone,
rain at all seasons or in winter.
Distribution: Bredasdorp, Caledon, Swellen-
dam, WC, S Africa.
Note: The papillae on the leaves seem to vary
within populations depending on age and water
regime. The plants appear more fragile and thin-
ner stemmed than those of T. occidentale, with
which it might otherwise be confused in habitat.
Bradleya 31/2013
 Further material seen: Gerbaulet et al. 25951
(HBG!); Hartmann & Bayer 34637 (HBG!).
T. occidentale L.Bolus NM1 1928: 150 T Smith
5140 (BOL!).
The thick storage tubers in different shapes,
depending on the available space, sometimes
branched, developing 6-10 erect branches from the
hypocotyl, thick at first, thinner when branching,
the branches forming rather dense mats, intern-
odes short to invisible, finely papillate, later
smooth; leaf bladder cells cobble stone-shaped or
elongate and irregular in shape, with or without
an apical papilla, bladder cells at tip forming elon-
gate bristles up to 0.5 mm long, orange, some-
times falling of with age, leaves 11 mm long, 3 mm
broad; flowers with stiffly hairy pedicels 8 mm
long, solitary, hairs on the calyx base spreading
or turning upwards, 15 mm Ø, petals apricot-
coloured with a deep purple central line, the cen-
tre of the flower forming a low cone, filamentous
staminodes and stamens basally with long papil-
lae densely interwoven but visible between the
outer elements; fruit with two separated closing
bodies as endocarpal swellings below the expand-
ing sheets, closed 6 mm Ø, open 9.5 mm Ø, 5.2
mm h.
Ecology: on shale or limestone, at the edge of
Renosterbosveld, 110 m altitude, rain at all sea-
sons or in winter.
Distribution: Bredasdorp, Caledon, Swellen-
dam, WC, S Africa.
Note: The species seems to grow well in open
places, settling in disturbed areas, natural as well
as man-made, e.g. on low koppies in wheat fields.
Stouter than the otherwise similar members of T.
gracile, to which it becomes even more similar in
cultivation by elongating the internodes.
Further material seen: Hartmann & Bayer
34494, 34497, 34499, 34633, 34639, 34640, 34642,
34704 (HBG!)
T. pygmaeum L.Bolus 1966: 335 T Thomas
27823 (BOL!).
From a thick taproot plagiotropous branches,
these forming adventitious roots and secondary
heads, all in a mat often filled up with fine soil, 1-
3 cm high, up to 25 cm Ø, internodes invisible, c.
1 mm Ø; leaves ovate, bladder cells with hairy
papillae, apically in a group of hairs 0.4 mm long,
leaves 6 mm long, 3 mm Ø; flowers with hairy
pedicels, c. 3 mm long or almost sessile, calyx base
with longer papillae, petals apricot coloured with
a purple central stripe, few filamentous stamin-
odes in the low cone flower type, apically deep
Bradleya 31/2013
purple, like the filaments inside, all equipped with
long papillae in their basal parts; fruit with clos-
ing bodies split in their middle or divided towards
the interior.
Ecology: in finely grained soil, rain at all sea-
sons.
Distribution: Bredasdorp, Swellendam, WC, S
Africa.
Note: T. pygmaeum has been misinterpreted
until now because it was described as having fine
plagiotropous roots. Fresh material, however, re-
vealed that the plants possess originally a very
thick tuber and form lateral roots close to the soil
surface in addition from which secondary heads
can develop. The whole old plant looks like a mat,
and pieces can be removed appearing as a single
plant because they can possess secondary bundles
of fine roots.
In its long hairy leaves, the species agrees with
T. fergusoniae and T. strumosum.
Further material seen: Bayer 7282 (HBG!);
Hartmann & Bayer 34636 (HBG!)
T. strumosum (Haw.) L.Bolus SAG 1928: 127 
M. strumosum Haw. 1821: 190 NT, designated
here, Duncanson 1002/999 from material of Bowie
collected in 1823, drawn 27.9.1825 (K)
Root a thick soft, fragile fusiform tuber,
branches erect to decumbent with long visible in-
ternodes, about 2 mm thick, internodes sparsely
hairy, fragile; leaves almost terete, about 11 mm
long, usually less than 2 mm broad and thick,
with long white distantly placed hairs, apically
collected into an even longer bundle of hairs; flow-
ers with long-haired calyx bases, pedicels with
somewhat shorter hairs, petals pale straw-
coloured or apricot, with a dark purple central
stripe from the base, few filamentous staminodes
in the outer ring of the low cone shaped centre,
not changing in shape during anthesis, with some
to very many interwoven papillae in the basal
part; fruit with solid or sharply divided closing
bodies of endocarpal origin, lifting the expanding
sheet considerably.
Ecology: between rocks and stones in loamy
material in Fynbos vegetation, rainfall at all sea-
sons.
Distribution: Swellendam, WC, S Africa.
Note: Haworth (1821: 190) noted “tuberous,
Hedgehog" in his description. This habit is seen in
an undated drawing of Duncanson (1011/222, K!),
which differs from the description by having a
well-developed white diadem of erect hairs. This
drawing may be the holotype, although Brown
noted "not" on that drawing, based on the white
diadem-like structure, which, however, is just vis-
73
ible in the younger drawing of Duncanson as well.
The second drawing (Duncanson 1002/999, K!),
showing leaves without a spectacular diadem, was
based on material collected by Bowie in 1823 and
produced in 1825 only, both dates younger than
the description and therefore perhaps not the orig-
inal plant. Yet, this latter material is chosen as
the neotype here since there are no discrepancies
between the description and the figure, as far as
both go. The species is similar in growth to T.
gracile, but more conspicuously similar to T. fer-
gusoniae and T. pygmaeum in the soft long hairs
of the leaves.
Further material seen: Bayer 7322 (HBG!);
Hartmann & Bayer 34625 (HBG!).
Acknowledgements
Over many years, the Deutsche Forschungs-
gemeinschaft sponsored research in the family
Aizoaceae, mainly fieldwork in Southern Africa.
Additional grants from Hansische Universitätss-
tiftung Hamburg and Prof. W. Hartmann helped
to undertake even more fieldwork in recent
decades. To all donators we express our gratitude.
We also thank the Nature Conservation authori-
ties in the relevant provinces of South Africa, in
Botswana, Namibia, and Lesotho for permission
to work there and to collect and export material.
Bruce and Daphne Bayer supported extensive
field work during recent years, and many farmers
gave their consent for us to work on their proper-
ties. Several colleagues shared their knowledge
with us, and Ulrich Meve undertook staining ex-
periments to help us to find out the special prop-
erties of the miraculous tufts of bristles. Our work
would not have been possible without the care and
attention Katrin Baumgärtner has given to the
living plants in the greenhouse in Hamburg. To
all of them, we should like to express our sincere
thanks. Last, but not least, we are grateful for
good advice for improvement of the text from an
anonymous referee.
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75