Professional Documents
Culture Documents
a
b Flowering Plants of Africa 65 (2017)
Flowering Plants of Africa
A peer-reviewed journal containing colour plates with descrip-
tions of flowering plants of Africa and neighbouring islands
Edited by
Alicia Grobler
with assistance of
Gillian Condy
Volume 65
Pretoria
2017
ii Flowering Plants of Africa 65 (2017)
Editorial board
R.R. Klopper South African National Biodiversity Institute,
Pretoria, RSA
P.C. Zietsman National Museum, Bloemfontein, RSA
Next volume
Volume 66 is likely to appear in 2019.—The Editor
ISSN 0015-4504
ISBN 978-1-928224-20-4
Flowering Plants of Africa 65 (2017) iii
Contents
Volume 65
Codonorhiza azurea (Eckl. ex Baker) Goldblatt & J.C.Manning in Strelitzia 35: 93 (2015). La-
peirousia corymbosa var. azurea Eckl. ex Baker: 90 (1896). Lapeirousia azurea (Eckl. ex Baker)
Goldblatt in Goldblatt & Manning: 333 (1992). Lapeirousia azurea Eckl.: 31 (1827), nom. nud.
Meristostigma azureum Eckl. ex Steud.: 131 (1841), nom. nud.
Until the recent publication of a monograph of Lapeirousia Pourr. and related genera
by Goldblatt & Manning (2015), which included the new genus Codonorhiza Goldblatt &
J.C.Manning, C. azurea was regarded as a member of Lapeirousia, either as the independ-
ent species L. azurea or included in L. corymbosa var. fastigiata (Lam.) Goldblatt. The new
genus Codonorhiza was erected for an exclusively Western Cape group of species that was
shown by molecular study (Forest et al. 2014) to comprise a monophyletic assemblage
related to Lapeirousia itself. A second group of species, now the genus Afrosolen Goldblatt
& J.C.Manning (Goldblatt & Manning 2016), includes the tropical and eastern southern Afri-
can members of Lapeirousia. The taxonomically isolated relict species, L. neglecta Gold-
blatt, is now treated as the only species of Schizorhiza Goldblatt & J.C.Manning. Nested
between these genera are the Socotran genus Cyanixia Goldblatt & J.C.Manning and the
tropical African Savannosiphon Goldblatt & Marais. The splitting up of Lapeirousia sensu lato
was deemed necessary in order that its constituent genera were natural, or monophyletic
(sharing a single common ancestor) (Goldblatt & Manning 2015). The name Codonorhiza
literally means bell [shaped] root and alludes to the bell-shaped corm.
Codonorhiza is readily recognised by the bell-shaped corms with flat bases that are
covered by dark grey to almost black tunics of compressed fibres, flowers with deeply
forked style branches, and green, somewhat leathery floral bracts, the inner and outer
of which are more or less equal in size (Goldblatt & Manning 2015). Codonorhiza also
stands out from other genera in the Lapeirousia alliance in its distinctive seeds, small in
size and ovoid in shape with a prominent funicular collar, and in having pollen with a
single opercular band. The operculum is two-banded in almost all other members of the
group and is the ancestral condition for subfamily Crocoideae. Divided style branches are
a common, but not unique character in the tribe Watsonieae, to which Codonorhiza and
Lapeirousia belong. Undivided style branches are characteristic of Cyanixia, the tropical
African Zygotritonia Mildbr., and several species of Afrosolen. Flat-based, bell- or cone-
shaped corms also characterise several genera of Crocoideae of the Iridaceae, including
Lapeirousia and Afrosolen. The differences between Codonorhiza and Lapeirousia are also
reflected in their chromosomes. Although both have bimodal karyotypes (chromosome
complements) consisting of one long and many smaller chromosome pairs, Codonorhiza
has a basic chromosome number of n = 10, whereas Lapeirousia has n = 9 or 8 and
Schizorhiza has n = 6.
PLATE 2321.—1, habit, × 1; 2, leaf tip, × 1. Voucher: Goldblatt 6279 in Compton Herbarium, Cape Town.
Artist: John C. Manning.
4 Flowering Plants of Africa 65 (2017)
Codonorhiza azurea is recognised by the dark blue, zygomorphic flowers with the three
abaxial tepals bearing dark red or black markings near the base, and either dark purple or
red-brown pollen. The flowers are tilted backwards so that the unilateral stamens lie almost
horizontally above the adaxial (or dorsal) tepal. The paired green floral bracts are subequal
in length and characteristic of the genus, as is the very large basal foliage leaf. The remain-
ing leaves are much shorter and grade into bract-like structures at the upper nodes of the
stem. The related C. corymbosa (L.) Goldblatt & J.C.Manning has smaller, radially symmetric
flowers with the stamens held erect and surrounding the upright style. Vegetatively, the two
species are similar but the leaves and other organs of C. corymbosa are consistently smaller.
A second related species, C. elandsmontana Goldblatt & J.C.Manning, has flowers similar
to those of C. azurea in floral presentation, thus with tilted flowers with unilateral stamens
arching above the adaxial tepal, but the tepals are paler blue and the abaxial tepals have
white, bracket-shaped markings edged in dark blue.
Pollination of Codonorhiza azurea is somewhat puzzling. The flowers lack scent and the
dark-coloured pollen is unusual for species pollinated by bees, yet the large anthophorine
bee Anthophora diversipes (Anthophoridae) has been recorded visiting the flowers and pre-
sumably foraging for nectar present in the base of the short perianth tube. Hopliine beetles
have also been documented as insect visitors to C. azurea, and the flowers not only provide
a platform for assembly but have the dark tepal markings typical of flowers pollinated by
hopliine beetles. Absence of scent is likewise typical of hopliine-pollinated flowers. At a site
in Malmesbury large, buff-brown hopliines were the only insects that we recorded on the
flowers, using the flowers as sites of assembly and mating. The report of visits to flowers of
C. azurea (as Lapeirousia) by Anthophora diversipes suggests a possible bimodal pollination
system for the species, using both large-bodied bees and hopliine beetles.
Codonorhiza azurea is a narrow endemic of the Western Cape, centred on the Paarde-
berg and surroundings, where it extends from near Paarl north to Malmesbury and Rie-
beek-Kasteel (Figure 1). It is especially prominent in burned or cleared slopes in Elytropap-
pus-dominated renosterveld, on heavier soils, either on clay or granite-derived gravels.
Much of its original habitat has been given over to agriculture and its range is now much
reduced so that the species is regarded as Endangered (Raimondo et al. 2009). In 1992,
Goldblatt & Manning (1992) considered that the species was probably not in immediate
danger of extinction, as it was then still known from sites away from cultivation. Several of
these are now lost to urban expansion and some of its remaining sites are being developed
for vineyards.
Codonorhiza azurea was first figured in a fascicle of N.L. Jacquin’s Icones plantarum
rariorum (Jacquin 1791) under the name Ixia corymbosa L., in a composite painting that
Flowering Plants of Africa 65 (2017) 5
REFERENCES
BAKER, J.G. 1896. Iridaceae. In W.T. Thiselton-Dyer (ed.), Flora capensis 6: 7–171. Reeve,
London.
ECKLON, C.F. 1827. Topographisches Verzeichniss der Pflanzensammlung von C.F. Ecklon.
Esslingen.
FOREST, F., GOLDBLATT, P., MANNING, J.C., BAKER, D., COLVILLE, J., DEVEY, D.S., JOSE,
S., KAYE, M. & BUERKI, S. 2014. Pollinator shifts as triggers of speciation in painted
petal irises (Lapeirousia: Iridaceae). Annals of Botany (London) 113: 357–371.
GOLDBLATT, P. 1972. A revision of the genera Lapeirousia Pourret and Anomatheca Ker in the win-
ter rainfall region of South Africa. Contributions from the Bolus Herbarium 4: 1–111.
GOLDBLATT, P., DAVIES, T.J., MANNING, J.C., VAN DER BANK, M. & SAVOLAINEN, V. 2006.
Phylogeny of Iridaceae subfamily Crocoideae based on combined multigene plastid DNA
analysis. Aliso 22: 399–411.
GOLDBLATT, P. & MANNING, J.C. 1992. Systematics of the southern African Lapeirousia corym-
bosa (Iridaceae–Ixioideae) complex (sect. Fastigiata) and a new species of sect. Paniculata.
South African Journal of Botany 58: 326–336.
GOLDBLATT, P. & MANNING, J.C. 2015. Systematics and biology of Lapeirousia, Codonorhiza,
Psilosiphon and Schizorhiza in southern Africa. Strelitzia 35: 93–96. South African National
Biodiversity Institute, Pretoria.
GOLDBLATT, J.C. & MANNING, J. 2016. Afrosolen Goldblatt & J.C.Manning, a new name for
Psilosiphon Welw. ex Goldblatt & J.C.Manning (Iridaceae), with new combinations. In P.
Goldblatt & J.C. Manning, Nomenclatural adjustments in African plants 2, Bothalia 46(1),
a2024. http://dx.doi.org/10.4102/abc.v46i1.2024.
GOLDBLATT, P., RODRIGUEZ, A., POWELL, M.P., DAVIES, T.J., MANNING, J.C., VAN DER BANK,
M. & SAVOLAINEN, V. 2008. Iridaceae ‘Out of Australasia’? Phylogeny, biogeography, and
divergence time based on plastid DNA sequences. Systematic Botany 33: 495–508.
JACQUIN, N.J. VON. 1791. Ixia corymbosa. Icones Plantarum Rariorum. Wappler, Vienna.
RAIMONDO, D., VON STADEN, L., FODEN, W., VICTOR, J.E., HELME, N.A., TURNER, R.C.,
KAMUNDI, D.A. & MANYAMA, P.A. (eds). 2009. Red List of South African Plants. Strelitzia
25. South African National Biodiversity Institute, Pretoria.
STEUDEL, E.T. 1841. Nomenclator botanicus ed. 2. J.G. Cotta, Stuttgart.
1
3
2
Gladiolus crassifolius Baker in Journal of Botany (London) 14: 334 (1876); Baker: 215 (1892);
Baker: 150 (1896); Lewis et al.: 60 (1972); Goldblatt: 94 (1996); Goldblatt & Manning: 74 (1998).
Gladiolus rachidiflorus Klatt: 339 (1882). Gladiolus thomsonii Baker: 223 (1892). Gladiolus mas-
ukuensis Baker: 283 (1897). Gladiolus tritoniiformis Kuntze: 308 (1898). Gladiolus junodii Baker:
866 (1901). Gladiolus conrathii Baker: 1005 (1904). Gladiolus dieterlenii Phillips: 282 (1917).
Gladiolus L. is one of the two largest genera in Iridaceae, as rich in species as Iris L. and
probably equally as popular in horticulture and the cut flower trade. It is also one of the few
genera of subfamily Crocoideae to have radiated significantly outside the Greater Cape Flo-
ristic Region. Of some 270 species recognised in the genus, 170 occur in southern Africa,
where they are concentrated in the winter-rainfall zone in the extreme southwest of the
subcontinent (Goldblatt & Manning 1998). The genus has undergone a secondary radiation
in the temperate, summer-rainfall grasslands along the eastern highlands, and G. crassifolius
Baker is one of the more common species found here.
Species of Gladiolus are strikingly variable in their vegetative and floral morphology
and the genus is essentially recognised by its disc-like seeds with a papery, circumferential
wing. These are carried in leathery, somewhat inflated capsules that open widely when dry,
each segment ultimately recurving to expose the seeds. A few species have the seed wings
reduced. The expanded stigmatic branches are also characteristic, and most species are
diploid, 2n = 30, with the unusually high basic chromosome number of x = 15.
The genus has no evident close relatives within Crocoideae and a recent phyloge-
netic analysis of DNA sequence data suggested that its closest relative is the monotypic
Melasphaerula Ker Gawl. (Goldblatt et al. 2008). This unusual species has several unique
features that find no parallel in Gladiolus or in any other genus in the family, including its
bell-shaped corms with tunics splitting radially from the centre of the flattened base, a lax,
highly branched inflorescence, and depressed-obcordate capsules dehiscing along their
upper edges.
An analysis of speciation and diversification rates in Gladiolus suggests that the diver-
sity of species in southern Africa stems from an early origin of the genus in the winter-
rainfall region, followed by active speciation in the summer-rainfall parts of the subconti-
nent (Valente et al. 2011). This study proposes a southern African origin of the genus during
the Miocene, followed by radiation into tropical Africa, Madagascar, the Mediterranean
Basin and western Asia. It also supports a relationship between G. crassifolius and other
multi-leaved summer-rainfall species in the genus, including G. densiflorus Baker.
PLATE 2322.—1, plant in flower, much reduced; 2, inflorescence, × 1; 3, flower, × 1.5; 4, fruit, × 1. Voucher
specimen: Condy 255 in National Herbarium, Pretoria. Artist: Gillian Condy.
10 Flowering Plants of Africa 65 (2017)
Gladiolus crassifolius is widely distributed through eastern southern Africa and is still
relatively common in South Africa, Swaziland and Lesotho (Figure 1). It is one of the few
southern African species shared with tropical Africa, where it extends through southern
and eastern Zimbabwe and central Mozambique, northward to Malawi and southwestern
Tanzania, with isolated populations in western Angola. Plants almost always grow in hilly
country, favouring well-drained, rocky grassland habitats, usually in full sun but occasion-
ally in light shade. Typically a late summer-flowering species, G. crassifolius most commonly
blooms from the end of February to late April, but plants are occasionally seen in flower in
late January, even at high elevations. The specimen illustrated here was collected in stony
grassland on the Mpumalanga Escarpment east of Carolina near Lochiel.
The flowers of Gladiolus crassifolius are fairly typical of several smaller-flowered grassland
species in the genus but it is usually readily recognised by the combination of an inclined,
many-flowered spike, and basal fan of several, tough leaves with conspicuously thickened
margins, midribs and secondary veins. The specific epithet crassifolius, meaning thick-leaved,
alludes to the thick, leathery, strongly ribbed leaves. The flowers are most frequently light
pink or salmon with darker pink to purple nectar guides highlighted with yellow. However,
plants with striking orange, red, and coral-pink flowers occur locally in Mpumalanga.
The species was recorded as used traditionally by the Basotho, under the name kxahla-
ya-maloti, with other plants as an enema, and alone as a remedy for headache and lum-
bago (Watt & Breyer-Brandwijk 1932).
Gladiolus crassifolius is most commonly confused with G. densiflorus from the far east-
ern escarpment and coastal lowlands of southern Africa. The critical difference between
the two is in their foliage. Both species have the midrib and margins heavily thickened but
the leaves of G. crassifolius have one or two pairs of secondary veins more conspicuously
thickened than the remaining veins, whereas the leaves of G. densiflorus have numerous,
equally fine, closely spaced minor veins between the midrib and the margins. This distinc-
tion is more easily seen in dried than in living plants. The distribution of the two species
is largely complimentary, with G. crassifolius typically a highland species ranging west and
south of the Mpumalanga Escarpment and G. densiflorus east of the escarpment onto the
KwaZulu-Natal coastal plain and north into lowland Swaziland and Mozambique. Their
ranges overlap along the Mpumalanga Drakensberg but G. crassifolius usually blooms a few
weeks later than G. densiflorus. In the field G. crassifolius can usually also be distinguished
by the pink to salmon flowers strongly marked with purple on the lower tepals. The flowers
of G. densiflorus vary widely in colour but are typically finely speckled with dark spots, and
the lower tepals do not have conspicuous dark markings.
Gladiolus crassifolius is not a particularly variable species but was poorly understood for
many years. The species was based on a collection made by Thomas Cooper in the eastern
Free State in 1862 (Baker 1876), but may have been first collected in 1832 by J.F. Drège in
Flowering Plants of Africa 65 (2017) 11
median 15–20 × 12 mm. Filaments unilateral and arcuate, 12–14 mm long, exserted 6–7 mm
from tube; anthers 7–8 mm long, pale lilac to purple, pollen whitish to pale yellow. Style usu-
ally dividing opposite lower half of anthers, branches ± 3 mm long, seldom reaching past mid-
dle of anthers. Capsules obovoid, 9–12(–14) mm long, 3-lobed above and retuse. Seeds elliptic,
broadly winged, 4–6 × 2.5–4.0 mm. Flowering time: mainly February and March, occasionally
later or earlier at higher elevations; in tropical Africa usually April to June but sometimes August
to November, especially after fires. Plate 2322.
REFERENCES
BAKER, J.G. 1876. New Gladioleae. Journal of Botany, British and Foreign 14: 333–339.
BAKER, J.G. 1892. Handbook of the Irideae. George Bell & Sons, London.
BAKER, J.G. 1896. Iridaceae. In W.T. Thiselton-Dyer (ed.), Flora capensis 6: 7–171. Reeve, London.
BAKER, J.G. 1897. Diagnoses africanae X. Kew Bulletin 1897: 243–300.
BAKER, J.G. 1901. Iridaceae. Bulletin de l’Herbier Boissier, Sér. 2,1: 862–868.
BAKER, J.G. 1904. Iridaceae. Bulletin de l’Herbier Boissier, Sér. 2,4: 1003–1007.
GOLDBLATT, P. 1996. Gladiolus in tropical Africa. Timber Press, Oregon.
GOLDBLATT, P. 2008. Additions to Gladiolus (Iridaceae) in the Flora Zambesiaca Region: the new
species G. metallicola and the reinstatement of G. mosambicensis. Novon 18: 164–167.
GOLDBLATT, P. & J.C. MANNING. 1998. Gladiolus in southern Africa. Fernwood Press, Cape
Town.
GOLDBLATT, P., RODRIGUEZ, A., POWELL, M.P., DAVIES, T.J., MANNING, J.C., VAN DER BANK,
M. & SAVOLAINEN, V. 2008. Iridaceae ‘Out of Australasia’? Phylogeny, biogeography, and
divergence time based on plastid DNA sequences. Systematic Botany 33: 495–508.
KLATT, F.W. 1882. Ergänzungen u, Berichtingungen zu Baker’s Systema Iridacearum. Abhandlun-
gen der Naturforschenden Geselschaft zu Halle 15.
KUNTZE, O. 1898. Revisio generum plantarum, vol. 3. Leipzig.
LEWIS, G.J., OBERMEYER, A.A. & BARNARD, T.T. 1972. A revision of the South African species of
Gladiolus. Journal of South African Botany, Suppl. 10: 1–316.
PHILLIPS, E.P. 1917. A contribution to the Flora of the Leribe Plateau and Environs: with a discus-
sion on the Relationships of the Floras of Basutoland, the Kalahari, and the South-eastern
Regions. Annals of the South African Museum 16: 282.
RENDLE, A.B. 1911. A contribution to the flora of Gazaland: Monocotyledons. Journal of the Lin-
nean Society, Botany 40: 207–235.
VALENTE, L.M., SAVOLAINEN, V., MANNING, J., GOLDBLATT, P. & VARGAS, P. 2011. Explaining
disparities in species richness between Mediterranean floristic regions: a case study in Gladi-
olus (Iridaceae). Global Ecology and Biogeography 20: 881–892.
WATT, J.M. & BREYER-BRANDWIJK, M.G. 1932. The Medicinal and Poisonous Plants of southern
Africa. E. & S. Livingstone, Edinburgh.
Aloe braamvanwykii Gideon F.Sm. & Figueiredo in Bradleya 30: 162–166 (2012); Van Wyk
[B-E.] & Smith: 216–217 (2014).
Within the genus Aloe L. a number of groups are acknowledged as being rather chal-
lenging from a taxonomic point of view. The grass and slender aloes, Aloe sect. Gramini-
aloe Reynolds and Aloe sect. Leptoaloe A.Berger, respectively, are two such groups (Smith
2005). However, no other infrageneric unit in Aloe has defied easy and accurate classifica-
tion more than the maculate aloes, Aloe sect. Pictae Salm-Dyck. These aloes are generally
characterised by having white or whitish H-shaped spots on one or both leaf surfaces, and
their flowers have a perianth tube that is basally swollen. However, even these two charac-
ters show considerable variation. For example, A. immaculata Pillans includes several popu-
lations that have somewhat striated, but generally uniformly green leaves, while the flowers
of A. prinslooi I.Verd. & D.S.Hardy are rather club-shaped and hardly swollen basally. Rep-
resentatives of Aloe sect. Pictae are not the only aloes that have distinct whitish markings on
their leaves. Aloe microstigma Salm-Dyck and A. pictifolia D.S.Hardy, both named for this
character, as well as several other species unrelated to the maculate aloes, also have spotted
leaves.
The adaptive significance of pale or dark spotted leaves, such as those of Aloe braam-
vanwykii (Figure 1), is not well understood and has been the subject of much speculation
and several hypotheses (Farmer 2014). In addition to their presence in some mature aloes,
spotted leaves are a common feature in the juvenile forms of numerous unrelated species
of Aloe, such as A. microstigma (Figure 2). This character is thought to assist young plants
to avoid herbivore attention by enabling them to better blend into seedling establishment
sites in the dappled shade and protection provided by nurse plants. A further explanation
for the spots on the leaves of aloes, and other alooid species, for example of Gasteria Duval
and Haworthia Duval, is that they mimic damage caused by feeding insects. Such mimicry
is observed across diverse plant groups, and is particularly well exemplified by Asystasia
varia N.E.Br. (Acanthaceae), a forest understory herb from KwaZulu-Natal and the Eastern
Cape. This species has irregular silvery patches along the midrib and lateral veins of leaves
which strongly resemble insect leaf-mining activity. It is hypothesised that spotted leaves
may serve as warning signals (albeit by deceit) to would-be leaf eaters. This possibility is
based on the observation that following damage by leaf eaters, many plants respond by
inducing a chemical defence in the same or other leaves (Karban 2015).
As is the case with a number of South Africa’s botanists, Prof. Braam (Abraham Eras-
mus) van Wyk (1952– ) of the Department of Plant and Soil Sciences, University of Pretoria,
was also fascinated by succulents, including species of Aloe, as a child. He grew up on a
PLATE 2323.—1, leaf rosette, × 0.5; 2, cylindrical raceme, × 1; 3, flower development, × 1. Voucher speci-
men: A.E. van Wyk 14132 in H.G.W.J. Schweickerdt Herbarium, Pretoria. Artist: Gillian Condy.
16 Flowering Plants of Africa 65 (2017)
FIGURE 1.—The light green leaves of Aloe braamvanwykii are white-spotted in irregular bands on both
upper and lower surfaces. Photographed on 4 January 2012 next to the Bloemhof road near Wolma-
ransstad, North-West Province, South Africa. Photograph: G.F. Smith.
FIGURE 3.—The red flowers of Aloe braamvanwykii have distinct basal swellings, unlike those of the sym-
patric Aloe grandidentata. Photographed on 5 January 2012 next to the Leeufontein road, near Wol-
maransstad, North-West Province, South Africa. Photograph: G.F. Smith.
erature, the winter- to spring-flowering A. grandidentata Salm-Dyck (Figure 4) was the only
species of maculate Aloe known to grow in the vicinity of Wolmaransstad and Schweizer-
Reneke. In addition to its later flowering time, A. grandidentata is readily identified by a per-
ianth tube that is rather weakly, but still noticeably, constricted above the base (Figure 5).
Braam was familiar with the latter species as it often grows together with the summer-flow-
ering plants, thus reinforcing his suspicion that two different species were involved.
Over a period of many years, Braam regularly mentioned this summer-flowering aloe
to a number of aloe experts, and also took live plants for cultivation to the then Botani-
cal Research Institute in Pretoria (later National Botanical Institute, now the South African
National Biodiversity Institute). However, publications on Aloe, including the treatment of
Aloe in the Flora of South Africa (Glen & Hardy 2000), still failed to account for the presence
of this particular aloe in North-West. When this state of affairs was perpetuated in the first
and second editions of Van Wyk [B-E.] & Smith’s Guide to the aloes of South Africa (Van
Wyk & Smith 1996, 2003), Braam again brought the existence of this particular aloe to the
attention of one of us (GFS).
An opportunity finally arose for two of us (GFS and EF) to more closely examine the
North-West plants just after New Year’s Day in 2012. During this trip the populations, espe-
cially around Wolmaransstad, were studied in detail. This study prompted a more thorough
investigation of other summer-flowering maculate aloes. It soon became apparent that the
18 Flowering Plants of Africa 65 (2017)
Aloe braamvanwykii has a rather brief and concentrated flowering period in mid-
summer, from end-November to February. Plants have consistently been observed to bloom
on Christmas Day (25 December) and New Year’s Day. In years of extreme mid-summer
drought (as has been the case in 2015/2016), many plants in a colony skip flowering. Fruit
set is very quick and mature-sized green fruit are often present while flowers on an inflores-
cence are still opening (Figure 7). This character, however, is not unique to A. braamvanwykii,
and has also been observed in other maculate aloes, such as A. maculata All. (Figure 8).
Flowering Plants of Africa 65 (2017) 19
FIGURE 5.—Flowers of Aloe grandidentata lack prominent basal swellings. Photograph: G.F. Smith.
FIGURE 6.—Plants of Aloe braamvanwykii often form dense clumps consisting of scores of rosettes. Photo-
graph: E. Figueiredo.
20 Flowering Plants of Africa 65 (2017)
FIGURE 8.—As is the case with Aloe braamvanwykii, Aloe maculata also carries mature, bright green fruit
in its clustered, head-shaped inflorescences, along with flowers that are still in bud, and at anthesis.
Photograph: G.F. Smith.
Flowering Plants of Africa 65 (2017) 21
FIGURE 10.—Aloe braamvanwykii flowering on 26 December 2015 in Klerksdorp Thornveld (Mucina et al.
2006) near Wolmaransstad, North-West Province, South Africa. Photograph: A.E. van Wyk.
22 Flowering Plants of Africa 65 (2017)
sometimes confluent yielding milky green surface, texture smooth, linear-attenuate, taper-
ing to apex, 170–260 mm long, 35–55 mm broad at base, basally sheathing; margins very
thin, brown, with triangular marginal teeth, green with light brown tips, ± 4 mm long, same
length throughout, evenly spaced at 10–13 mm apart; exudate pale yellowish, drying pur-
ple. Inflorescence 1–3, successively, 0.65–0.75 m tall, far exceeding height of rosette, cen-
tral raceme longest, 5- to 7-branched from above middle, branches arcuate-erect. Pedun-
cle 270–420 mm long, 8–14 mm broad at base, basally plano-convex, cylindrical above,
light greenish brown with white, powdery bloom; not sterile bracteate; bracts subtending
racemes narrowly triangular, 15–65 mm long, 6–8 mm broad at base, straw-coloured to
light brown, papery, rarely fleshy, many nerved. Racemes cylindrical, 140–170 mm long,
30–50 mm wide; buds erect to suberect, flowers horizontal to drooping when mature.
Floral bracts narrowly triangular, long attenuate, amplexicaul around pedicel, 5–9 mm long,
4–5 mm wide, straw-coloured, papery, 3- or 4-nerved. Pedicels 10–12 mm long, pinkish
brown. Flowers: actinomorphic to slightly zygomorphic, unscented, nectariferous; perianth
greenish tipped in buds, somewhat bicoloured when mature, light pink to mainly orange-
red to bright red with whitish to yellowish longitudinal stripes, tip extremity purplish brown
a b
FIGURE 11.—a, fruit capsules of Aloe braamvanwykii, × 1; b, habit, much reduced. Voucher specimen: A.E.
van Wyk 14132 in H.G.W.J. Schweickerdt Herbarium, Pretoria. Artist: Gillian Condy.
Flowering Plants of Africa 65 (2017) 23
or whitish, lightly pruinose, 20–25 mm long, flattened at base, ± 6 mm across ovary,
distinctly narrowed above ovary to ± 3 mm to form globose basal swelling, enlarging to
6–7 mm towards throat and wide open mouth, tubular-cymbiform; outer segments larger
than inner segments, lorate, free for ± 7 mm, free portion centrally pinkish red, borders
white or light yellowish, acute, segment margins straight, tips slightly recurved; inner seg-
ments narrower than outer, with white or yellowish border and more obtuse apex, free
for upper two-thirds of their length; stamens with cylindrically thread-like to very slightly
flattened, light yellow filaments, 25–28 mm long, all 6 of ± equal length, exserted for
2–5 mm; anthers small, 1–2 mm long, dark brown, versatile; ovary 5–6 mm long, 3 mm
in diameter, light green; style as long as or slightly longer than stamens, minutely capitate,
with small stigma, exserted 1–2 mm. Fruit an erect, bright green, cylindrical, trilocular cap-
sule, 17–22 mm long, 9–11 mm in diameter, apically truncate, dry remains of tepals shed
from around fruit early on, dehiscing loculicidally, chartaceous when dry, apically valves
sigmoidally curved outwards. Seeds dark greyish brown, angled, laterally compressed, 2.5–
3.0 mm long, with up to 1 mm wide off-white wing stretching around periphery of seed.
Chromosome number unknown. Plate 2323; Figure 11.
ACKNOWLEDGEMENTS
We are indebted to: Prof. Braam (A.E.) van Wyk, Department of Plant and Soil Sci-
ences, University of Pretoria, for freely sharing information about Aloe braamvanwykii; Ms
Hester Steyn, National Herbarium, SANBI, for producing the distribution map.
REFERENCES
VAN WYK, B-E. & SMITH, G.F. 2014. Guide to the aloes of South Africa, 3rd ed. Briza, Pretoria.
VON STADEN, L. 2014. Aloe braamvanwykii Gideon F.Sm. & Figueiredo. National Assessment:
Red List of South African Plants version 2015.1. Accessed on 2016/06/07.
11
8
12
7
4
13
14
9
15
5
10
16
Tinospora fragosa (I.Verd.) I.Verd & Troupin subsp. fragosa, in Memoires de l’academie Roy-
ale des Sciences d’Outre-Mer, Classe des sciences naturelles et medicales 13: 1–312 (1962); Van
Jaarsveld: 28 (2016). Desmonema fragosum I.Verd.: 209 (1941).
There are three species of Tinospora indigenous to South Africa, Tinospora caffra (Miers)
Troupin, T. tenera Miers and T. fragosa subsp. fragosa. The genus was recently revised
acknowledging three species in South Africa (De Wet et al. 2016). Tinospora fragosa subsp.
kaokoensis van Jaarsv. was recently named from plants on the Namib margin in Kaokoveld,
Namibia. This subspecies differs from the typical one by its ability to flower on younger
stems, its swollen roots, and its distinctly striated stems that taper at intervals with the thin-
ner portions desiccating spontaneously and acting as a vegetative propagation back-up
(Van Jaarsveld 2016).
PLATE 2324.—1, young branches with leaves, × 1; 2, seed, × 1.5; 3, female fruiting branch, × 1; 4, branch
with shortened, often rounded female inflorescence, × 1; 5 inner sepals, × 4; 6, outer sepals, × 4; 7,
petals, × 4; 8, opened flower from side, × 4; 9, opened flower from top, × 4; 10, flower with sepals
and petals removed, × 4; 11, branch with elongated male inflorescence, × 1; 12, opened flowers, × 4;
13, outer sepals, × 4; 14, inner sepals, × 4; 15, petals, × 4; 16, stamens, × 4. Voucher: Van Jaarsveld
25946 in Compton Herbarium, Cape Town. Artist: Marieta Visagie.
28 Flowering Plants of Africa 65 (2017)
Van der Merwe further noticed the species’ peculiar mode of regeneration from aerial
roots where the plant had been severed, losing contact with the ground. When the local
farmers observed this, they were prompted to name the sprouting Tinospora fragosa subsp.
fragosa Aaron’s rod, in reference to the biblical account: Aaron was chosen leader of the 12
tribes of Israel, but this was disputed by the other tribes. He then asked the tribal leaders to
each bring their walking sticks to be placed in the tent of gathering. The next morning it was
only Aaron’s rod that had sprouted with his authority firmly established.
The plant also came under the attention of the botanist Inez Verdoorn at the Botani-
cal Research Institute in Pretoria, who was in charge of the National Herbarium from
1944–1951 (Gunn & Codd 1981). Realising it was a new species, she named it Desmonema
fragosum I.Verd. in 1941. However, in 1962 Troupin, in his treatment of Tinospora, rightfully
transferred it to this genus.
The savannas of Africa are known for their herbivores, as well as fires, and succulent
plants need specific strategies to survive in this type of environment with its particular dis-
turbance regime. Plants have to either stand their ground where they germinate (defence
mode) or move away somehow. In an attempt to avoid predation, succulents have a vast
number of strategies, consisting mostly of mechanical, chemical and camouflage defensive
modes. Aloes, for example, have a combination of chemical and mechanical defence (bit-
ter sap and thorns) and, in spite of this, are still grazed during severe droughts. Tinospora
fragosa subsp. fragosa grows in dry river valleys, rocky sites and on flats. The species has
the special ability to rapidly twine, enabling it to mobilise and displace itself to a new posi-
tion. It thus follows a path of passive resistance by growing away from danger (i.e. out of
reach of grazing animals). The stems of T. fragosa subsp. fragosa are succulent and juicy,
but with bitter sap. If it is grazed and subsequently loses contact with the soil, it has suf-
ficient reserves in its succulent stems to grow a survival root – a true life line – re-rooting
the plant to ensure its long-term survival, unlike most other creepers that would die upon
loss of soil contact (Van Jaarsveld 2015). The root will only appear during the growing sea-
Flowering Plants of Africa 65 (2017) 29
The ability to sprout from aerial roots is not only confined to Tinospora and has been
observed in other succulent plants, such as Cissus tuberosa DC. from Mexico.
Tinospora fragosa subsp. fragosa is at once distinguished from the other two tinosporas by
its much thicker stems of 30–50 mm in diameter, and abbreviated side branches from where
30 Flowering Plants of Africa 65 (2017)
The vegetation over most of the habitat of Tinospora fragosa subsp. fragosa consists of
mopaneveld (Figure 3), which is part of the Savanna Biome (Mucina & Rutherford 2006).
Plants grow at an altitude of 450 to 1 000 m above sea level. However, near Polokwane
(Limpopo Province) plants have been observed on granite outcrops and also east of Chu-
niespoort in the dry savanna of the Olifants River Valley and near Steelpoort. The region
is dry in winter, with cool evenings, and very high summer temperatures (sometimes up to
40°C) with associated humidity. The average daily summer maximum is 26–29°C and the
average daily winter temperature about 13–17°C. Rainfall is mainly from late spring through
summer, November to May, and ranges between 300 and 500 mm per annum. Tinospora
fragosa subsp. fragosa plants sprout during November and during times of very hot con-
ditions, with temperatures above 35°C, their leaves may change, becoming smaller, more
coriaceous and pointed, and slightly glaucous with the margin undulating and crisped (Fig-
ure 4). The plant becomes deciduous during April or May when the leaves turn yellow,
continuing its growth in November.
At Gonarezhou Nature Reserve (Figure 5) one of us (EJvJ) has observed Tinospora shar-
ing its habitat with various other succulents at the Chivilila Falls on the banks of the Runde
River. Associated succulents includes Adansonia digitata L., Sterculia rogersii N.E.Br., Aloe
globuligemma Pole-Evans, A. chabaudii Schönland, Pachypodium saundersii N.E.Br. and
Flowering Plants of Africa 65 (2017) 31
Sansevieria hallii Chahinian. Our species is not a soil specialist and grows in various types,
including granite-, dolerite- and shale-derived soils. Their arboreal nature also ensures that
they are less affected by occasional fires.
Tinospora fragosa subsp. fragosa grows well as a curiosity plant such as on a protected
window sill in warmer climates. The stems can be hung and when the root appears, it is
best to place a pot below it, allowing the plant to establish. Provide enough moisture dur-
ing the active growing phase, but also withhold water during the dry winters to enable the
plants to rest. One of us (EJvJ) has been growing the plants at the Botanical Society Con-
servatory at Kirstenbosch National Botanical Garden for more than 20 years. Stems were
hung up near the roof of the building and the growth rate of the roots measured at about
10–20 mm per day. The plant rapidly establishes and continuous its growth once it comes
in contact with soil. The initial root gradually thickens (flaking longitudinally) becoming
stem-like. It is best grown in bushveld gardens (Van Jaarsveld 2010). Stems can be placed
in trees or fences and growth is rapid. Once fruits are set it should attract frugivorous birds.
Propagation is easily achieved by pruning the stem into various parts, of which each should
sprout during the growing season.
root free-hanging, 1 mm in diameter at first, fast growing (10–15 mm per day), gradually
thickening to 10 mm in diameter, greyish-green and developing longitudinally peeling bark
with age. Young branches initially green, smooth, becoming grey, later becoming greyish-
brown to dark brown and scaling periderm, covered in lenticels, scars and sometimes
lichens, 30–50 mm in diameter, the distal part with abbreviated ascending to spreading
lateral branches up to 10 mm long from where the flowers appear. Mature branches fragile,
becoming detached with aerial root that grows only during rainy season, free hanging, re-
establishing the plant. Leaves simple, greyish green, alternately arranged, firm, coriaceous,
broadly ovate, often v-shaped and cordate at base, 30–80(100) mm long, 30–80(100) mm
broad, petioles to 100 mm long, pulvinus yellowish green, margin entire, sometimes wavy
(during hot spells), glabrous, nerves palmate. Inflorescence racemose, up to 50 mm long,
cauliflorous. Bracts 1–3, 5 mm long, linear. Flowers small, greenish, inconspicuous, 1–3
per bract. Male inflorescence a simple false raceme 40–50 mm long; sepals 6, ovate, outer
three 1.5 mm long, 1 mm broad, inner three 3.5 mm long, 2.5 mm broad; petals 6, obo-
vate, 2 mm long 1.5 mm broad, often obscurely subcrenate; stamens 6, connate at the
base, 2.5 mm long; filaments 1.5 mm long; anthers longitudinally fissured, dehiscing from
the side. Female flowers on shorter racemes, 20 mm long; sepals 6, outer three 2 mm long,
1.25 mm broad, inner three concave, 4 mm long, 3 mm broad; petals 6, obovate, 2 mm
long, 1.5 mm broad; staminodes 6, opposite petals, 2 mm long; carpels 3, abbreviated stip-
itate; ovary stipitate at base, articulated, obliquely ovoid; stigma subsessile, terminal. Fruit
ripen after leaves appear, reddish, bean-shaped drupe, 8–10 mm long, 5–7 mm in diam-
eter, exocarp a fleshy pulp; endocarp cartilaginous, smooth. Plate 2324.
ACKNOWLEDGEMENTS
We are grateful to various people: Koos Bekker and Karen Roos (owners of Babylons
toren Farm) for allowing one of us (EJvJ) to do research on various succulent and other plant
species; Sally Carney (Harare) for taking one of us (EJvJ) to the Gonarezhou National Park in
July 2015; Sandy Munro, Deana Edwards, Diana Duncan for their company in the field on
the 2015 expedition; and Hester Steyn for providing the distribution map.
REFERENCES
DE WET, H., STRUWIG, M. & VAN WYK, B-E. 2016. Taxonomic notes on the genera Tiliacora
and Tinospora (Menispermaceae) in southern Africa. South African Journal of Botany 103:
283–294.
DE WET, H. & VAN WYK. B-E. 2008. An ethnobotanical survey of southern African Menisper-
maceae. South African Journal of Botany 74: 2–9.
GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern Africa. Balkema, Cape Town.
HEYWOOD, V.H. (ed.) 1978. Flowering Plants of the World. Oxford University Press, Oxford.
JACKSON, W.P.U. 1990. Origins and meanings of names of South African plant genera. UCT Ecolab,
Cape Town.
MIERS, J. 1851. A few remarks on the Menispermaceae. Annals and Magazine of Natural History
7,2: 33–45.
MUCINA, L. & RUTHERFORD, M.C. 2006. The vegetation of South Africa, Lesotho and Swazi-
land. Strelitzia 19. South African National Biodiversity Institute, Pretoria.
RODIN, R.J. 1985. The ethnobotany of the Kwanyama Ovambos. Monographs in Systematic Bot-
any from the Missouri Botany Garden 9: 113–114.
Flowering Plants of Africa 65 (2017) 33
TROUPIN, G. 1962. Monographie des Menispermaceae africaines. Academie Royale des Sciences
d’Otre-Mer, Bruxelles.
VAN JAARSVELD, E.J. 1981. Wonderplant van Suid-Afrika. Veld & Flora, 67,2: 56–59.
VAN JAARSVELD, E.J. 2010. Waterwise Gardening. Tafelberg, Cape Town.
VAN JAARSVELD, E.J. 2015. Succulents and their adaptations to mega-herbivores in the
Gonarezhou National Park. Haseltonia 21: 41–55.
VAN JAARSVELD, E.J. 2016. Tinospora fragosa subsp. kaokoensis (Menispermaceae), a new subspe-
cies from the northern Kaokoveld (Namibia). Aloe 52:1: 28–31.
VERDOORN, I.C. 1941. Desmonema fragosum from the northern Transvaal. Journal of South Afri-
can Botany 7: 209–213.
The Menispermaceae is a very old angiosperm family of about 70 genera and 450
species in the order Ranunculales (APG III 2009; Hoot et al. 2009). Molecular dating has
shown that this family originated 116–105 million years ago (Anderson et al. 2005), and
that the different tribes recognised in the family differentiated either during the Cretaceous
period (145–66 million years ago) or the Palaeocene epoch (66–56 million years ago)
(Jacques & Bertolino 2008).
Members of the Menispermaceae are either herbaceous or woody climbers with dioe-
cious and small, inconspicuous flowers (Kessler 1993). A peculiarity of the anatomy of the
family is the medullary rays that separate the vascular bundles in the stems and rhizomes
(De Wet et al. 2002). The family is distributed throughout the tropical lowland rainforests,
although there are a few species adapted to cooler and drier climates (Hoot et al. 2009). It
is commonly known as the moonseed family, as the shape of the seeds is reminiscent of a
crescent moon (Doria et al. 2008).
Cissampelos hirta is a herbaceous liana with deltate to cordate, subpeltate leaves (Figure 1).
The small, dioecious flowers are subtended by foliaceous bracts, which persist below the red
drupes.
PLATE 2325.—1, flowering stem, × 1; 2, female inflorescence, × 4; 3, fruiting stem, × 1. Voucher specimen:
Struwig 184 in University of Zululand Herbarium, KwaDlangezwa. Artist: Gillian Condy.
1
2
Table 1.—The genera and species of the Menispermaceae in southern Africa and their distribution within the
region. Endemic taxa are indicated by *. N, Namibia; B, Botswana, LIM, Limpopo; NW, North-West; G,
Gauteng; M, Mpumalanga, S, Swaziland; FS, Free State; KZN, KwaZulu-Natal; L, Lesotho; NC, Northern
Cape; WC, Western Cape; EC, Eastern Cape
Cissampelos hirta can easily be confused with C. mucronata A.Rich. and C. torulosa
E.Mey. ex Harv. based on habit and leaf shape. The leaves of C. hirta are, however, coria-
ceous, while those of C. mucronata are puberulent to tomentose and therefore velvety to
the touch. The leaves of C. torulosa have a distinct tuft of hair at the petiolar attachment,
which is absent from C. hirta. Cissampelos capensis L.f. is a woody shrub and not likely to
be confused with the other Cissampelos species. A detailed comparison of the morphology
and anatomy of the four Cissampelos species is given in De Wet et al. (2002). A key to dis-
tinguish the southern African Cissampelos species is given below:
REFERENCES
ANDERSON, C.L., BREMER, K. & FRIIS, E.M. 2005. Dating phylogenetically basal eudicots using
rbcL sequences and multiple fossil reference points. American Journal of Botany 92: 1737–
1748.
APG III. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and
families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: 105–121.
BARBOSA-FILHO, J.M., CA-CUNHA, E.V.L. & GRAY, A.I. 2000. Alkaloids of the Menispermaceae.
In G.A. Cordell (ed.), The Alkaloids, Vol. 54. Academic Press, Illinois.
DE WET, H., TILNEY, P.M. & VAN WYK, B-E. 2002. Vegetative morphology and anatomy of Cis-
sampelos in South Africa. South African Journal of Botany 68: 181–190.
DE WET, H. & VAN WYK, B-E. 2008. An ethnobotanical survey of Southern African Menisper-
maceae. South African Journal of Botany 74: 2–9.
DE WET, H., STRUWIG, M. & VAN WYK, B-E. 2016. Taxonomic notes on Tiliacora and Tinospora
(Menispermaceae) in southern Africa. South African Journal of Botany 103: 283–294.
DORIA, G., JARAMILLO, C.A. & HERRERA, F. 2008. Menispermaceae from the Cerrejón forma-
tion, middle to late Paleocene, Colombia. American Journal of Botany 95: 954–973.
DURAND, T. & SCHINZ, H. 1898. Conspectus Florae Africae, ou énumération des plantes d’Afrique,
Vol. 1. R. Friedländer & Sohn, Berlin.
ENGLER, A. 1899. Beiträge zur Flora von Africa. XVII. Menispermaceae africanae. Botanische Jahr-
bücher für Systematik, Pflanzengeschichte und Pflanzengeographie 26: 393–416.
FOSTER, S. & JOHNSON, R.L. 2008. National Geographic desk reference to nature’s medicine.
National Geographic Society, Washington DC.
GLEN, H.F. 2003. Menispermaceae. In G. Germishuizen & N.L. Meyer (eds), Plants of southern
Africa: an annotated checklist. Strelitzia 14. National Botanical Institute, Pretoria.
40 Flowering Plants of Africa 65 (2017)
HOOT, S.B., ZAUTKE, H., HARRIS, D.J., CRANE, P.R. & NEVES, S.S. 2009. Phylogenetic patterns
in Menispermaceae based on multiple chloroplast data. Systematic Botany 34: 44–56.
JACQUES, F.M.B. & BERTOLINO, P. 2008. Molecular and morphological phylogeny of Menisper-
maceae (Ranunculales). Plant Systematics and Evolution 274: 83–97.
KESSLER, P.J.A. 1993. Menispermaceae. In K. Kubitzki, J.G. Rohwer & V. Bittrich (eds), The families
and genera of vascular plants. Flowering plants — Dicotyledons. Springer-Verlag, Berlin.
KLOPPER, R.R., CHATELAIN, C., BÄNNINGER, V., HABASHI, C., STEYN, H.M., DE WET, B.C.,
ARNOLD, T.H., GAUTIER, L., SMITH, G.F. & SPICHIGER, R. 2006. Checklist of the flower-
ing plants of Sub-Saharan Africa. An index of accepted names and synonyms. South African
Botanical Diversity Network Report No. 42. SABONET, Pretoria.
MIERS, J. 1871. Contributions to botany, iconographic and descriptive, Vol. 3. Williams and Nor-
gate, London.
PETERS, W.C.H. 1862. Naturwissenschaftliche Reise nach Mossambique, auf Befehl Seiner Majestät
des Königs Friedrich Wilhelm IV, in den Jahren 1842 bis 1848 ausgeführt, von Wilhelm C. H.
Peters. Botanik. George Reimer, Berlin.
POOLEY, E. 1998. A field guide to wild flowers: KwaZulu-Natal and the eastern region. Natal Flora
Publication Trust, Durban.
STAFLEU, F.A. & COWAN, R.S. 1981. Taxonomic Literature: A selective guide to botanical publica-
tions and collections with dates, commentaries and types, TL-2. Bohn, Scheltema & Holkema,
Utrecht.
STAFLEU, F.A. & COWAN, R.S. 1983. Taxonomic Literature: A selective guide to botanical publica-
tions and collections with dates, commentaries and types, Tl-2. Bohn, Scheltema & Holkema,
Utrecht.
STEARN, W.T. 1992. Botanical Latin, edn 4. David and Charles, Newton Abbott.
TROUPIN, G. & GONÇALVES, M.L. 1973. Menispermaceae. In A. Fernandes (ed.), Flora de
Moçambique, Vol. 7. Junta de Investigaçáes do Ultramar Centro de Botânica, Lisboa.
4 6 5
Protea foliosa Rourke in Journal of South African Botany 41,2: 53–62 (1975); Rourke: 178
(1980); Vogts: 94 (1982); Rebelo: 64 (1995); Vanderplank: 94 (1999). P. magnoliifolia H.Buek ex
Meisn. in Drège: 118; 127; 213 (1843), nom. nud. P. caulescens E.Mey. ex Meisn. in DC.: 244
(1856), nom. nud. P. tenax (Salisb.) R.Br. var. latifolia Meisn. in DC.: 244 (1856); Hutchinson et al.:
587 (1912).
Protea foliosa is a small, rounded shrub up to 1.5 m tall, but typically shorter than 1 m
tall in populations around Grahamstown. It has many erect, seldom-branched, glabrous
stems arising from a large woody rootstock, giving it a high degree of fire resistance. The
lanceolate, oblanceolate or broadly elliptic leaves are 90–160 mm long, 15–70 mm broad,
narrowed towards the petiole, coriaceous and glabrous, although young leaves are pubes-
cent at first. The upright inflorescences are usually hidden by enfolding leaves. The involu-
cral bracts are greenish yellow, often flushed reddish, and described as rust-coloured (Van-
derplank 1999) or dull carmine (Rourke 1975), in the distal half.
William Burchell first discovered Protea foliosa in the Swartberg Mountains near Rie-
beek East in 1813 and collected specimens again on the Van Stadensberg a year later.
There has been much confusion regarding the correct name for the species. Various invalid
names were applied in the past and at one time it was considered to be a variety of P. tenax
(Salisb.) R.Br.. John Rourke finally resolved the difficulties by describing it anew as P. foliosa
in 1975.
Protea foliosa can be distinguished from related species in the section Crinitae (East-
ern Ground Sugarbushes) (Rebelo 1995) by its shrubby, upright growth habit, erect, rarely
branched, stout (60–80 mm in diameter) stems, and prominent lanceolate to broadly ellip-
tic glabrescent mature leaves that completely surround and obscure the small globose,
multiple-clustered, above-ground inflorescences (Rourke 1975). Protea intonsa Rourke
and P. vogtsiae Rourke are dwarf shrubs with underground, scaled stems. Protea montana
E.Mey. ex Meisn. is a prostrate, many-branched mat-forming shrub. Protea intonsa and
P. montana have needle-like to linear leaves and P. vogtsiae also has longer (120–250 mm)
and narrower (8–30 mm) leaves than P. foliosa. The species was previously much confused
with P. tenax, but that species has decumbent or trailing, rarely branched, stems (3–6 mm
in diameter), secund or subsecund linear-lanceolate to oblanceolate leaves, pointing verti-
cally from the horizontal stem, and wide, shallowly crateriform, seldom clustered inflores-
cences borne at ground level. The species are largely allopatric. Protea foliosa occurs from
near Patensie and Uitenhage in the west to Riebeek East and Grahamstown in the east,
PLATE 2326.—1, foliose, low growing branches bearing terminal inflorescence with carmine-flushed, green
involucral bracts, × 1; 2, florets, × 1; 3, cross section of inflorescence, × 1; 4, mature infructescences
persist on branches, × 1; 5, fire-stimulated infructescence open to expose achenes × 1; 6, empty
involucral receptacles × 1; 7, sterile achenes, × 1; 8, fertile achenes, × 1.5. Voucher specimen: Dold
TD16009 in Selmar Schonland Herbarium, Grahamstown. Artist: Susan Abraham.
44 Flowering Plants of Africa 65 (2017)
Protea foliosa is most frequently encountered along the crests of ridges and southern
slopes in grassy fynbos where it grows in poor sandstone and quartzitic soils. It is common
in its restricted range, sometimes locally abundant on moist sites. This species is distributed
from Patensie in the west to Grahamstown in the east with an interruption of this distribu-
tion range inland of Algoa Bay (Figure 1).
Protea foliosa may begin flowering as early as March and continue until September.
Peak flowering occurs between May and June (Rourke 1980; Melidonis & Peter 2015). Like
other species in the family, pollen is aggregated as tetrads and deposited on the outside of
the stigma, the gynoecium acting as a pollen presenter.
Motion sensing camera traps were used to observe visits to inflorescences of Protea foli-
osa at the farm Upper Gletwyn near Beggars Bush east of Grahamstown. This revealed that
P. foliosa is pollinated primarily by a single species of rodent, Rhabdomys pumilio (Melidonis
& Peter 2015). Unusually, for a rodent pollinated plant (Wiens & Rourke 1978; Biccard &
Midgley 2009; Johnson & Pauw 2014), all recorded visits to the inflorescences occurred
during daylight hours, mainly between 07:00 and 12:00. Rodents clambered on the inflo-
rescences (Figure 2) or surrounding grass tussocks in order to probe inflorescences in search
of nectar, resulting in pollen being loaded onto the rodents’ rostrum and subsequently onto
stigmas with successive visits (Melidonis & Peter 2015).
Other potential pollinators observed at the site include Otomys irroratus and the shrew,
Crocidura cyanea. Both were found to have Protea foliosa pollen tetrads on their rostra and
Flowering Plants of Africa 65 (2017) 45
FIGURE 2.—Motion detecting camera traps reveal that Rhabdomys pumilio (Striped Field Mouse) is the
primary pollinator of Protea foliosa and all recorded visits occurred during the day. Photographs: C.I.
Peter.
in their scat, although to a much lesser extent than Rhabdomys pumilio. Otomys irroratus
and C. cyanea were not captured on camera interacting with P. foliosa inflorescences, fur-
ther supporting the idea that R. pumilio is the primary pollinator of P. foliosa at this site.
The low occurrence of pollen grains in the scat of Otomys irroratus and Crocidura cya-
nea relative to those of Rhabdomys pumilio suggests that pollen-preening is reduced in
these two species, or that absolute pollen loads are lower resulting in less pollen ingestion.
Wester et al. (2009) drew similar conclusions with other rodent species based on their lack
of interaction with the flower and lack of pollen grains found in their scat. The presence of
a large number of pollen grains in the scat of captured R. pumilio individuals provides evi-
dence that this species routinely visit Protea foliosa flowers under natural conditions at this
site.
FIGURE 3.—Cross section of an infructescence FIGURE 4.—A single brief visit by a male Malachite
shows different morphology of fertile (plump Sunbird (Nectarinia famosa) was recorded
and milky-white) vs. infertile (shrivelled and with a motion detecting camera trap. Pho-
brown) achenes. Photograph: C.I. Peter. tograph: C.I. Peter.
However, quantifying seed set in Protea species is difficult, as individual ovules remain
attached to the inflorescence regardless of whether or not they have been fertilized (Wiens
1983; Biccard & Midgley 2009). In addition, sterile and viable achenes do not differ mor-
phologically, which makes them difficult to distinguish externally. Cross-sections of viable
endosperm-containing seeds are swollen and solid white in colour, in contrast to dull-col-
oured and fibrous sterile seeds (Figure 3) (Wiens 1983; Biccard & Midgley 2009; Melidonis
& Peter 2015).
Infructescences appear to persist for a number of years in the plant (serotiny) suggest-
ing the possibility of fire-triggered seed release (pyriscence). To test this possibility a number
of infructescences were harvested randomly from numerous plants. Half of these were left
to dry naturally while the others were placed on a concrete floor and lightly covered with
hay which was then set alight. Within six hours, the infructescences exposed to the fire had
opened completely. Unburnt infructescences took over a week to open to the same degree.
Over the course of the study by Melidonis & Peter (2015), a large number of inflores-
cences were inspected and numerous ants were encountered, presumably attracted by the
nectar. A single unidentified monkey beetle (Scarabaeidae: Hopliini) was also recorded in an
inflorescence. In addition, a camera trap recorded a very brief visit by a male Malachite Sun-
bird (Necatarinia famosa) to a Protea foliosa inflorescence (Figure 4). On more than one occa-
sion, camera traps recorded the presence of small antelope in close proximity to the plants,
but direct feeding was not observed. Many inflorescences and infructescences were found
to contain the frass and larvae of parasitizing insects although these were not identified.
Description (after Rourke 1975, 1980).—Rounded shrub to 1.5 m tall with numerous
straight, erect stems arising from a woody subterranean rootstock. Stems stout, 60–80 mm
Flowering Plants of Africa 65 (2017) 47
REFERENCES
BICCARD, A. & MIDGLEY, J.J. 2009. Rodent pollination in Protea nana. South African Journal of
Botany 75: 720–725.
DOLD, A.P. & COCKS, M.L. 1999. Preliminary list of Xhosa plant names from the Eastern Cape,
South Africa. Bothalia 29,2: 267–292.
DRÈGE, J.F. 1843. Zwei Pflanzengeographische Documente: 118, 127, 213.
HUTCHINSON, J., PHILLIPS, E.P. & STAPF, O. 1912. Proteaceae. In W.T. Thiselton-Dyer (ed.) Flora
capensis 5: 587. Reeve, London.
JOHNSON, C.M. & PAUW, A. 2014. Adaptation for rodent pollination in Leucospermum arena
rium (Proteaceae) despite rapid pollen loss during grooming. Annals of Botany 113: 931–938.
KROPF, A. & GODFREY, R. (eds). 1915. A Kafir-English Dictionary. Lovedale Press, Alice.
MEISNER, C.F. 1856. Proteaceae. In A.P. de Candolle (ed.), Prodromus 14: 244. Victor Masson,
Paris.
MELIDONIS, C. & PETER, C.I. 2015. Diurnal pollination, primarily by a single species of rodent,
documented in Protea foliosa using modified camera traps. South African Journal of Botany
97: 9–15.
PAHL, H.W. (ed.) 1989. The Greater Dictionary of Xhosa. Vol. 3. University of Fort Hare, Alice.
REBELO, T. 1995. Sasol Proteas. A Field Guide to the Proteas of Southern Africa. Fernwood Press,
Vlaeberg.
ROURKE, J.P. 1975. Notes on Protea in South Africa: The Protea tenax tangle. Journal of South
African Botany 41,2: 53–62.
ROURKE, J.P. 1980. The Proteas of Southern Africa. Purnell, Cape Town.
VANDERPLANK, H.J. 1999. Wildflowers of the Port Elizabeth Area. Gamtoos to Swartkops Rivers
(The Coastal Bush and Fynbos Region). Bluecliff Publishing, Hunters Retreat.
VOGTS, M. 1982. South Africa’s Proteaceae. Know Then and Grow Them. Struik, Cape Town.
WESTER, P., STANWAY, R. & PAUW, A. 2009. Mice pollinate the Pagoda Lily, Whiteheadia bifolia
(Hyacinthaceae) — first field observations with photographic documentation of rodent pol-
lination in South Africa. South African Journal of Botany 75: 713–719.
WIENS, D. 1983. Nonflying mammal pollination of southern African Proteas: a non-coevolved
system. Annals of the Missouri Botanical Gardens 70:1–31.
48 Flowering Plants of Africa 65 (2017)
WIENS, D. & ROURKE, J.P. 1978. Rodent pollination in southern African Protea spp. Nature
276:71–73.
Protea namaquana Rourke in South African Journal of Botany 56,2: 261–265 (1990); Rebelo:
73 t.28 (1995).
The Kamiesberg Mountains are by far the most impressive geographical feature in the
landscape of Namaqualand: a great chaotic jumble of huge granite boulders and colossal
granite bornhardts erupting into several peaks over 1 500 m in altitude with Rooiberg in
the south, at 1 706 m, the highest of all. With an average annual winter rainfall of some
350 mm, it is the best watered part of Namaqualand and botanically the most diverse,
known to vegetation ecologists as Kamiesberg Granite Fynbos (Mucina & Rutherford 2006),
while plant geographers have recognised the Kamiesberg as an important centre of endem
ism for over a century (Marloth 1908; Weimarck 1941; Van Wyk & Smith 2001). Hilton-
Taylor (1996) reported that the Kamiesberg is home to 79 endemic plant species, a figure
that is almost certain to rise as more intensive collecting is undertaken.
Here on the upper slopes of Rooiberg, and adjacent peaks at Esels Kop, Rhebokskloof,
Karas and Welkom, Protea namaquana occurs, usually on south- but occasionally on north-
facing slopes, in small fragmented populations of a few dozen plants (Figure 1). Despite the
fact that the upper reaches of the Kamiesberg is largely undisturbed, the increased frequency
of fire due to growing human influence has had the most dramatic effect on this very slow-
maturing species, which may take up to ten or more years from seedling to first flowering.
The species’ assessed Red List status is Critically Endangered (Raimondo et al. 2009). A rough
count by one of us (JPR) in 1980 suggested there were about 300 living adult specimens.
This is not necessarily an accurate assessment, as some populations on Rooiberg had been
burnt prior to that count, leaving a few charred adult specimens as evidence. Due to patchy
burning it was difficult to establish more precise population figures, which could be higher.
Furthermore, plant and population numbers are a dynamic situation, with figures sometimes
changing every few years. Protea namaquana is a serotinous reseeder, but upon investigat-
ing burnt areas in the 1980s, seedling recruitment was observed to be disappointingly low.
A further problem was that some adult populations near Welkom were suffering from very
bad witches broom infestations. This debilitating plant growth disease that affects leaves and
stems is possibly caused by mycoplasma-like organisms, transmitted by mites, and gives rise
to dense bunches of miniature leaves among the normal foliage, weakening the adult plant.
PLATE 2327.—1, flowering branches with open inflorescence and immature inflorescence bud, much reduced;
2, open perianth and curved style, × 2. Voucher specimen, J.P.Rourke 1680 in Compton Herbarium,
Cape Town. Artist: Ellaphie Ward-Hilhorst.
52 Flowering Plants of Africa 65 (2017)
Perhaps the most critical factor in determining the survival of this exceptionally rare
species is the restricted area of suitable habitat above 1 200 m where the microclimate is
favourable for seedling recruitment. Protea namaquana is slow growing and long lived. Fully
mature specimens develop into robust erect shrubs up to two metres in height with a simi-
lar spread, supported by a well-developed, stout, upright trunk, often as much as 200 mm
in diameter at the base. Hoary old specimens like these are well over 50 years old. Some
may even be octogenarians or older.
It was Elsie Esterhuysen, the greatest twentieth-century botanical collector in the Cape
Floristic Region, who made the first recorded collection of this species in 1954. Yet, for a
long time it remained unrecognised, being confused with its vicarious equivalent, Protea
sulphurea E.Phillips, so named for its yellowish involucral bracts. Protea sulphurea is readily
recognised by its procumbent, horizontally sprawling mode of growth and small, elliptic,
glaucous leaves. Protea namaquana, on the other hand, is an upright shrub with considerably
longer (30–60 mm long), narrowly oblanceolate to linear-spathulate leaves with a distinctive
pale, dusty green colour and carmine flushed involucral bracts. Though the distribution areas
of these two taxa are today widely separated by several hundred kilometres, they are clearly
vicariads, almost certainly derived from a common ancestral population which differentiated
into two distinct components as a result of geographical isolation brought about by increas-
ing aridification in the western parts of southern Africa since the Pleistocene.
Flowering peaks in late autumn and winter with a few inflorescences occasionally
opening as late as September. The downwardly facing pendent inflorescences tend to be
partly concealed within the foliage. On opening, each shallowly bowl-shaped inflorescence
produces a strong odour resembling a sweetish, yeasty fermentation, the characteristic syn-
drome of rodent-pollinated proteas (Rourke & Wiens 1977; Wiens & Rourke 1978; Wiens
et al. 1983). Though yet to be confirmed by field observations, rodent pollination in this
species is probably one method of achieving pollen transfer, with the nocturnally active
Namaqua Rock Rat (Aethomys namaquensis) – a widespread and well-known pollinator of
some proteas – being a likely candidate.
Bird pollination is more readily discernible. One of us (JPR) has personally observed
two species of sunbird, the Dusky Sunbird (Nectarinia fusca) and the Malachite Sunbird
(Nectarinia famosa) feeding on open inflorescences. Nevertheless, as in the related Protea
sulphurea where pollination by both birds and rodents is known to occur, the same situa-
tion is likely to prevail in P. namaquana.
Thus far, Protea namaquana has not been successfully cultivated. At Kirstenbosch
National Botanical Garden, Cape Town, seed from the type collection germinated well.
In 1983 140 seedlings were planted out in the protea garden but numbers declined to 120
in 1984 dropping to a mere 25 in February 1986. By 1989 all were dead. The low light
intensity, high rainfall and excessive dampness of Kirstenbosch probably contributed to the
failure of this species which is clearly better suited to sunny, semi-arid inland sites.
REFERENCES
HILTON-TAYLOR, C. 1996. Patterns and characteristics of the flora of the Succulent Karoo Biome,
South Africa. In L.J.G. van der Maesen, X.M. van der Burgt & J.M. Medebach de Rooy (eds),
The Biodiversity of African Plants: 58–72. Kluwer, Dordercht.
MARLOTH, R. 1908. Das Kapland insonderheit das Reich der Kapflora, das Waldgebiet und die
Karroo. Gustav Fischer, Jena.
MUCINA, L. & RUTHERFORD, M.C. 2006. The vegetation of South Africa, Lesotho and Swazi-
land. Strelitzia 19. South African National Biodiversity Institute, Pretoria.
RAIMONDO, D., VON STADEN, L., FODEN, W., VICTOR, J.E., HELME, N.A., TURNER, R.C.,
KAMUNDI, D.A. & MANYAMA, P.A. (eds). 2009. Red list of South African plants. Strelitzia
25. South African National Biodiversity Institute, Pretoria.
REBELO, T. 1995. Sasol Proteas. Fernwood Press, Cape Town.
ROURKE, J.P. 1990. A new species of Protea (Proteaceae) from Namaqualand with comments on
the Kamiesberg as a centre of endemism. South African Journal of Botany 56,2: 261–265.
ROURKE, J.P. & WIENS, D. 1977. Convergent floral evolution in South African and Australian Pro-
teaceae and its possible bearing on pollination by nonflying mammals. Annals of the Missouri
Botanical Garden. 64,1: 1–17.
VAN WYK, A.E. & SMITH, G.F. 2001. Regions of floristic endemism in southern Africa: a review with
emphasis on succulents. Umdaus Press, Hatfield.
WEIMARCK, H. 1941. Phytogeographical groups, centres and intervals within the Cape Flora.
Lunds Universitets Å rs-skrift N.F. Adv. 2 Bd 37. Nr. 5: 1–143.
WIENS, D. & ROURKE, J.P. 1978. Rodent pollination in southern African Protea spp. Nature
276,5683: 71–73.
54 Flowering Plants of Africa 65 (2017)
WIENS, D., ROURKE, J.P., CASPAR, B.B., RICKART, E., LAPINE, T.R., PETERSON, C.J. & CHAN-
NING, A. 1983. Nonflying mammal pollination of southern African Proteas: a non-coevolved
system. Annals of the Missouri Botanical Garden 70,1: 1–31.
The Crassulaceae, a near-cosmopolitan family of about 1 400 species, are well rep-
resented in southern Africa, which, along with Mexico, is recognised as a major centre of
present-day species diversity for the family (ʼt Hart & Eggli 1995; Thiede & Eggli 2007). Five
crassuloid genera are recognised in southern Africa, with Crassula L., with over 150 species,
being the most diverse (Dreyer 1997). The genera Adromischus Lem., Cotyledon L., and
Tylecodon Toelken have similarly diversified extensively in the region. However, this is not
so for the essentially Old World genus Kalanchoe Adans. sensu lato, which with just fewer
than 150 species globally, is the third largest genus in the Crassulaceae (Descoings 2003). In
southern Africa, Kalanchoe is relatively poorly represented, with fewer than 15 indigenous
species, all of which are placed in Kalanchoe sect. Kalanchoe. The greater proportion of
genus members is found naturally occurring in tropical Africa, Madagascar and Asia. Several
species, such as K. delagoensis Eckl. & Zeyh., figured in The Flowering Plants of Africa as
Bryophyllum delagoense (Eckl. & Zeyh.) Schinz (Plate 1938) (Tölken & Leistner 1986), and
hybrids, such as K. ×houghtonii D.B.Ward, are well known internationally as horticultural
subjects. A number of them, especially in Kalanchoe sect. Bryophyllum (Salisb.) Boiteau,
tend to naturalise, becoming weedy and invasive in countries with a mild to temperate
climate (Given 1984; Kapitany 2007; Walters 2011; Smith & Figueiredo 2013; Silva et al.
2015; Smith et al. 2015).
Kalanchoe leblanciae was described by Raymond Hamet just over 100 years ago, based
on material from ‘Delagoa Bay’, in what is today Maputo, Mozambique (Hamet 1912).
[Shortly after this species was described, Raymond Hamet changed his name by hyphenat-
ing his given name and surname to ‘Raymond-Hamet’; therefore his publications are here
cited under either ‘HAMET, R.’, or ‘RAYMOND-HAMET’.] The first gathering of K. leblan-
ciae that we can trace is by Henri-Alexandre Junod, a Swiss Romande missionary, who in
October of 1893 pressed what would become the holotype, noting that it was rare. Despite
this observation, most other Mozambique collections accessioned in South Africa are from
the same general locality as the type. Junod also collected the holotype of K. luciae Raym.-
Hamet, and was commemorated in K. junodii Schinz, which is generally considered con-
specific with K. lanceolata (Forssk.) Pers. (Descoings 2003).
PLATE 2328.—1, inflorescence × 1; 2, stem, middle section × 1. Voucher specimen: Crouch 1288 in H.G.W.J.
Schweickerdt Herbarium, Pretoria. Artist: Gillian Condy.
58 Flowering Plants of Africa 65 (2017)
recorded in Natal [now KwaZulu-Natal] in South Africa by Ross (1972). Evidently, Fernandes
(1983: 65) was alerted to this reference and, in amending the distribution range for the FZ
treatment of the Crassulaceae, indicated that the species occurred also in South Africa.
With some hesitation she gave the distribution in that country as ‘(Natal, Orange Free
State?)’. Apart from noting its presence in Mozambique, Descoings (2003) also reported it
from ‘RSA [Republic of South Africa]’ (Crouch et al. 2016). However, K. leblanciae did not
receive any mention in Tölken’s (1985) Flora of Southern African (FSA) treatment of the Cras-
sulaceae. Consequently it was omitted from all recent catalogues of South and southern
African plants in general (Burgoyne 2003, 2006), and succulent plants specifically (Dreyer
1997), as well as from a preliminary checklist of Mozambican plants (Silva et al. 2004). In
the north, the FZ region is geographically abutted by the Flora of Tropical East Africa region,
an area from which the species has not been reported (Wickens 1987).
Apart from doubts about its geographical distribution range, the taxonomic status of
Kalanchoe leblanciae has also been questioned. In The Plant List (see: www.theplantlist.org)
the name is regarded as being unresolved, based on the World Checklist of Selected Plant
Families in review. When Hamet (1912: 292) described K. leblanciae he noted that in his
previous monograph on the genus (Hamet 1907), the species would key out as the Indian
K. grandiflora Wight & Arn., but that K. leblanciae was quite distinct. Mixed gatherings on
herbarium sheets have further contributed to problems encountered in determining and
identifying specimens of K. leblanciae (see for example Fernandes 1980). In the protologue,
Hamet (1912: 292) also noted that K. leblanciae could be confused with a number of other
Kalanchoe species. These include K. sexangularis N.E.Br., K. paniculata Harv., K. rotundifolia
(Haw.) Haw., K. longiflora Schltr. ex J.M.Wood and K. brachyloba Welw. ex Britten. Kalan-
choe leblanciae may be readily distinguished from particularly K. brachyloba, K. hirta Harv.
and K. longiflora, all of which have clasping leaf bases. He regarded it as similar to K. rotundi-
folia, from which it differs in several leaf and flower characters. Fernandes (1980: 397) further
distinguished K. leblanciae from the polymorphic K. rotundifolia on account of its thicker, ses-
sile leaves that are of a different shape, as well as various flower characters on which she did
not elaborate, but would likely have included at least features provided by Table 1. It is worth
TABLE 1.—Characters on which Kalanchoe leblanciae can be distinguished from K. rotundifolia (see Hamet
1912; Fernandes 1983)
noting here that K. rotundifolia, as currently circumscribed with more than half a dozen syno-
nyms, is exceedingly variable. Although the flower colour of K. leblanciae has been reported
as ranging from ‘…yellow-green, yellow, salmon to bright red…’ (Fernandes 1983; Descoings
2003; Bandeira et al. 2007), we have yet to encounter specimens with flowers that are not
yellow (free portions of petals) and yellowish-green (floral tube). However, in K. rotundifolia
flower colour ranges from salmon through orange to bright red, sometimes even bicoloured,
but all characteristically show pronounced twisting of the corollas (Figure 1). In fact, Fer-
nandes (1983: 42) distinguished K. rotundifolia from the group consisting of K. brachyloba,
K. paniculata and K. leblanciae on the pink to brick flowers of the former species, whereas
the latter three have yellow corollas. Nevertheless, in the same publication, Fernandes
(1983: 65) described K. leblanciae as having a corolla ‘yellow-green to yellow or salmon to
bright red, reddish-brown throughout or with the lobes darker than the tube on drying’. It is
accordingly unclear whether she had ascertained that non-yellow corollas are observed only
once specimens are dry; we note though that one specimen from Inhaca Island in Maputo
Bay was documented as having red flowers (Mogg 28346 in PRE).
Fernandes (1980) further noted that Kalanchoe leblanciae differed from K. paniculata
(Smith et al. 2003) in having smaller leaves that have a different shape, leaf margins that
are not entire, and smaller flowers. In Fernandes (1983: 42) a key to the Kalanchoe spe-
FIGURE 1.—In the extraordinarily variable Kalanchoe rotundifolia flower colour ranges from salmon through
orange to bright red. However, all forms demonstrate twisting of the corollas as the flowers senesce
and start setting fruit, a feature not shared with K. leblanciae. Photograph: N.R. Crouch.
60 Flowering Plants of Africa 65 (2017)
FIGURE 5.—Typical Sand Forest habitat of Kalanchoe leblanciae, Ndumo Game Reserve, northern Maputa-
land, South Africa. Photograph: G.R. Nichols.
Flowering Plants of Africa 65 (2017) 63
ACKNOWLEDGEMENTS
Dr Reto Nyffeler and Franziska Schmid, both of the Herbarium (Z) of the Univer-
sität Zürich, Switzerland; Hermenegildo Matimele (LMA Herbarium, National Institute of
Agronomic Research, Maputo, Mozambique); Alice Massingue (LMU Herbarium, Eduardo
Mondlane University, Maputo, Mozambique); Prof. Fátima Sales, Herbarium Curator (COI),
Department of Life Sciences, University of Coimbra, Portugal, Dr Luis Catarino, Univer-
sity of Lisbon, Portugal; and Laurence Loze (Phanerogamic Herbarium, Conservatoire et
Jardin botaniques, Genève, Switzerland) are thanked for searching their collections for rel-
evant material. Prof. A.E. [Braam] van Wyk, H.G.W.J. Schweickerdt Herbarium, University
of Pretoria, South Africa, kindly allowed reproduction of his Kalanchoe leblanciae photo-
graph taken in the Licuáti Forest Reserve. Geoff Nichols and Michael Hickman generously
provided habitat and habit images, respectively. Curators of CPF, NU, NH, and PRE are
acknowledged for making facilities available during this study. Two anonymous referees are
thanked for useful comments on the manuscript.
REFERENCES
BANDEIRA, S., BOLNICK, D. & BARBOSA, F. 2007. Flores nativas do Sul de Moçambique / Wild
flowers of southern Mozambique. Universidade do Eduardo Mondlane, Departamento de
Ciências Biológicas, Maputo.
BANDEIRA, S., SENKORO, A., BARBOSA, F., MUALASSACE, D. & FIGUEIREDO, E. 2014. The ter-
restrial environment adjacent to Maputo Bay. In S. Bandeira & J. Paula (eds), The Maputo Bay
Ecosystem: 263–278. WIOMSA, Zanzibar Town.
BURGOYNE, P.M. 2003. Kalanchoe. In G. Germishuizen & N.L. Meyer (eds), Plants of southern
Africa: an annotated checklist. Strelitzia 14: 407–408. National Botanical Institute, Pretoria.
BURGOYNE, P.M. 2006. Crassulaceae. In G. Germishuizen, N.L. Meyer, Y. Steenkamp & M. Keith
(eds), A checklist of South African plants. Southern African Botanical Diversity Network
Report No. 41: 336–369. SABONET, Pretoria.
BURGOYNE, P. & VAN WYK, A.E. 1999. Crassula maputensis, a new record for the FSA region.
Bothalia 29,2: 254–255.
Flowering Plants of Africa 65 (2017) 65
CROUCH, N.R., FIGUEIREDO, E. & SMITH, G.F. 2016. Occurrence of the little-known Kalanchoe
leblanciae Raym.-Hamet (Crassulaceae) confirmed in South Africa. Bradleya 34: 70–76.
DESCOINGS, B. 2003. Kalanchoe. In U. Eggli (ed.), Illustrated handbook of succulent plants: Cras-
sulaceae: 143–181. Springer-Verlag, Berlin.
DREYER, L.L. (comp.) 1997. Crassulaceae DC. (Stonecrop or “Plakkie” family). In G.F. Smith, E.J.
van Jaarsveld, T.H. Arnold, F.E. Steffens, R.D. Dixon & J.A. Retief (eds), List of southern Afri-
can succulent plants: 56–63. Umdaus Press, Pretoria.
FERNANDES, R.B. 1980. Notes sur quelques espèces du genre Kalanchoe Adans. Boletim da Socie-
dade Broteriana LIII: 325–442.
FERNANDES, R.[B.] 1983. Crassulaceae. In E. Launert (ed.), Flora zambesiaca 7(1): 3–71. Managing
Committee on behalf of the contributors to Flora zambesiaca, London.
FIGUEIREDO, E. & SMITH, G.F. 2010. What’s in a name: epithets in Aloe L. (Asphodelaceae) and
what to call the next new species. Bradleya 28: 79–102.
FIGUEIREDO, E., SMITH, G.F. & CROUCH, N.R. 2016. The taxonomy and type of Kalanchoe
sexangularis N.E.Br. (Crassulaceae). Bradleya 34: 92–99.
GIVEN, D.R. 1984. Checklist of dicotyledons naturalized in New Zealand 17. Crassulaceae, Escal-
loniaceae, Philadelphaceae, Grossulariaceae, Limnanthaceae. New Zealand Journal of Bot-
any 22: 191–193. [Available at: http://dx.doi.org/10.1080/0028825x.1984.10425250]
HAMET, R. 1907. Monographie du genre Kalanchoe. Bulletin de l’Herbier Boissier, sér. 2,7: 869–
900.
HAMET, R. 1912. Sur un nouveau Kalanchoe de la baie de Delagoa [Kalanchoe leblanciae]. Reper-
torium specierum novarum regni vegetabilis 11: 292–294.
IZIDINE, S.A. 2003. Licuáti Forest Reserve, Mozambique: flora, utilization and conservation. M.Sc.
thesis, University of Pretoria, Pretoria.
KAPITANY, A. 2007. Australian succulent plants. An introduction. Kapitany Concepts, Boronia.
McNEILL, J., BARRIE, F.R., BUCK, W.R., DEMOULIN, V., GREUTER, W., HAWKSWORTH, D.L.,
HERENDEEN, P.S., KNAPP, S., MARHOLD, K., PRADO, J., PRUD’HOMME VAN REINE,
W.F., SMITH, G.F., WIERSEMA, J.H. & TURLAND, N.J. (eds). 2012. International Code of
Nomenclature for algae, fungi, and plants (Melbourne Code) adopted by the Eighteenth Inter-
national Botanical Congress Melbourne, Australia, July 2011. Koeltz Scientific Books, König-
stein.
POOLEY, E.S. 1978. A checklist of plants of Ndumo Game Reserve, north-eastern Zululand. Jour-
nal of South African Botany 44: 1–54.
RAYMOND-HAMET & MARNIER-LAPOSTOLLE, J. 1964. Le genre Kalanchoe au Jardin Botanique
“Les Cèdres”. Archives du Muséum National D’Histoire Naturelle Septième Série, Tome VIII:
1–110; Plates I–XXXVII.
ROSS, J.H. 1972. The flora of Natal. Memoirs of the Botanical Survey of South Africa 39: 1–418.
Botanical Research Institute, Pretoria.
RUTHERFORD, M.C., POWRIE, L.W., LÖTTER, M.C., VON MALTITZ, G.P., EUSTON-BROWN,
D.I.W., MATTHEWS, W.S., DOBSON, L. & McKENZIE, B. 2006. Afrotemperate, Subtropical
and Azonal Forests. In L. Mucina & M.C. Rutherford (eds), The vegetation of South Africa,
Lesotho and Swaziland. Strelitzia 19: 584–614.
SIEBERT, S., BANDEIRA, S., BURROWS, J. & WINTER, P. 2002. SABONET southern Mozambique
expedition 2001. SABONET News 7: 6–18.
SILVA, M.C., IZIDINE, S. & AMUDE, A.B. 2004. A preliminary checklist of the vascular plants of
Mozambique. / Catálogo provisório das plantas superiores de Moçambique. Southern African
Botanical Diversity Network Report No. 30. SABONET, Pretoria.
SILVA, V., FIGUEIREDO, E. & SMITH, G.F. 2015. Alien succulents naturalised and cultivated on the
central west coast of Portugal. Bradleya 33: 58–81.
SMITH, G.F., CROUCH, N.R. & STEYN, E.M.A. 2003. Notes on the distribution and ethnobotany
of Kalanchoe paniculata (Crassulaceae) in southern Africa. Bradleya 21: 21–24.
66 Flowering Plants of Africa 65 (2017)
SMITH, G.F. & FIGUEIREDO, E. 2013. The family Crassulaceae in continental Portugal. Bradleya
31: 76–88.
SMITH, G.F., FIGUEIREDO, E. & SILVA, V. 2015. Kalanchoe ×houghtonii (Crassulaceae) recorded
near Lisbon, Portugal. Bouteloua 20: 97–99.
STEENKAMP, Y., SIEBERT, S.J., SMITH, G.F., HUNTLEY, B.J. & WILLIS, C.K. 2006. Final project
report. Looking back on the SABONET Project: a triumph of regional cooperation. Southern
African Botanical Diversity Network Report No. 43. SABONET, Pretoria.
ʼt HART, H. & EGGLI, U. (eds). 1995. Evolution and systematics of the Crassulaceae. Backhuys
Publishers, Leiden.
THIEDE, J. & EGGLI, U. 2007. Crassulaceae. In K. Kubitzki (ed.), The families and genera of vascular
plants. IX. Flowering Plants. Eudicots: 83–118. Springer-Verlag, Berlin.
TÖLKEN, H.R. 1983. Kalanchoe sexangularis. Natal, Swaziland, Transvaal and Zimbabwe. Cras-
sulaceae. The Flowering Plants of Africa 47: plate 1878.
TÖLKEN, H.R. 1985. Crassulaceae. In O.A. Leistner (ed.), Flora of southern Africa 14: 1–244.
Botanical Research Institute, Pretoria.
TÖLKEN, H.R. & LEISTNER, O.A. 1986. Bryophyllum delagoense. Madagascar and naturalized in
many countries. Crassulaceae. The Flowering Plants of Africa 49: plate 1938.
VAN WYK, A.E. 2003. Dicerocaryum forbesii and D. senecioides. Pedaliaceae. Southeastern and
south-central Africa. Flowering Plants of Africa 58: plates 2198 and 2199.
VAN WYK, A.E. & SMITH, G.F. 2001. Regions of floristic endemism in southern Africa. A review with
emphasis on succulents. Umdaus Press, Pretoria.
WALTERS, M. 2011. Crassulaceae J.St.-Hil. (Stonecrop, Orpine or Houseleek family; Plakkiefami-
lie). In M. Walters, E. Figueiredo, N.R. Crouch, P.J.D. Winter, G.F. Smith, H.G. Zimmermann
& B.K. Mashope, Naturalised and invasive succulents of southern Africa. ABC Taxa 11: 232–
259. The Belgian Development Cooperation, Brussels.
WICKENS, G.E. 1987. Crassulaceae. In R.M. Polhill (ed.), Flora of Tropical East Africa: 1–66. A.A.
Balkema, Rotterdam.
Senna didymobotrya (Fresen.) H.S.Irwin & Barneby in Memoires of the New York Botanical
Garden 35: 467 (1982); Lock: 37 (1989); Pooley: 154 (1993). Cassia didymobotrya Fresen.: 53
(1839); Oliver: 276 (1871); Taubert: 201 (1895); Harms: 498 (1915); Baker: 62 (1911); Brenan
& Greenway: 97 (1949); Steyaert: 504, fig.36 (1952); Mendonça & Torre: 177 (1956); White &
Angus: 120 (1962); Drummond: 243 (1975); Ross: 195 (1972); Gordon-Gray: 79 (1977). Cassia
verdickii De Wild.: 49, t.16 fig.6–11 (1902); Harms: 498 (1915); Baker: 638 (1930).
Senna Mill. is a large genus in the Fabaceae family, subfamily Caesalpinioideae, tribe
Cassieae with approximately 350 species. The subfamily is usually divided into five tribes:
Cercideae, Caesalpineae, Cassieae, Detarieae and Macrolobieae (Tucker 2003; Resende
et al. 2013). Irwin & Turner (1960) included the species of Senna among the approximately
600 species of the genus Cassia sensu lato. In their taxonomic treatment, Irwin & Barneby
(1981 & 1982) subdivided the genus Cassia into Cassia L. emend. Gaertn. sensu stricto,
Chamaecrista Moench and Senna; these three genera were ascribed to subtribe Cassiinae.
Irwin & Barneby (1982) further divided the genus Senna into six sections (Astroiles [1 spe-
cies], Chamaefistula [c. 140 species], Paradictyon [1 species], Peiranisia [c. 55 species], Psilo
rhegma [c. 30 species] and Senna [c. 20 species]) distinguishing them mainly on stem, leaf
and flower characters. This classification has been supported by a number of studies of
these three genera based on morphology, ontogenetic characteristics, molecular systemat-
ics and cytogenetics (Resende et al. 2013). Many species of Senna exhibit enantiostyly floral
symmetry, meaning that the deflection of the style is either to the left or to the right within
the same inflorescence (Marazzi et al. 2006).
The genus Senna is pantropic in distribution but also occurs in most subtropical regions.
Its distribution includes North America, South America, Asia, Australia and Africa (Irwin
& Barneby 1981; Henderson 2001; Du Puy et al. 2002; Marazzii et al. 2006; Orwa et al.
2009). It is paraphyletic and amongst the largest genera in the Caesalpinioideae. In south-
ern Africa Senna is the largest genus in the subtribe Cassiinae, followed by Chamaecrista
and Cassia (Nkonki et al. 2003). Species vary, from trees and shrubs to woody herbs (some
monocarpic) and occur mainly in tropical seasonal vegetation and occasionally in forests,
especially along forest margins. There are about 18 species of Senna present in southern
Africa, of which the majority are naturalised aliens and have been cultivated for their attrac-
tive flowers. Only S. italica subsp. arachoides (Burch.) Lock and S. petersiana (Bolle) Lock
are indigenous (Germishuizen et al. 2006). According to Brummitt et al. (2007), none of the
species in the genus is known to form root nodules. Senna didymobotrya is morphologically
similar to S. alata (L.) Roxb. The former can be distinguished by its oblong-elliptic leaflets
that are shorter than 65 mm, bright yellow flowers with darker veins, and pods that are not
winged; whilst the latter has oblong leaflets that are longer than 60 mm, deep yellow or
orange flowers, and pods that are 4-winged (Brummitt et al. 2007). The genus is character-
PLATE 2329.—1, flowering stem, × 1; 2, fruiting branch, × 1; 3, leaf × 1. Voucher specimen: Jaca & Mkhize
845 in National Herbarium, Pretoria. Artist: Gillian Condy.
70 Flowering Plants of Africa 65 (2017)
The genus name is derived from the Arabic word sana or sanna for species with leaves
and pods with cathartic and laxative properties. Senna didymobotrya is a popular medici-
nal plant within its native range, where it is commonly used as a purgative, an antimalarial
medicine, and used to treat other fevers and jaundice. A decoction of the leaves, stems and
roots is used for the treatment of stomach problems such as diarrhoea, intestinal worms
and general abdominal pains. Higher doses are, however, reported to be toxic and in some
cases may cause vomiting and may be fatal. Root powder mixed in water or infusions of the
fresh parts are known to treat some skin diseases. Sometimes the concoction may cause
the patient to be weak if not taken with milk (Tabuti 2007). When treating children, the
young leaves are cooked in banana leaves to lower toxicity levels and given orally. For live-
stock, a decoction made from leaves is used to treat external parasites such as ticks. The
species is also used to preserve milk for longer periods (up to a year) by coating the inside
of the vessel to be used for storing milk with ash of burnt twigs. In the craft industry the
wood is used to make handicrafts. Outside of its native range (e.g. tropical Asia, America),
Flowering Plants of Africa 65 (2017) 71
S. didymobotrya is reported to be used as a fodder, green manure, cover crop and shade
tree in tea plantations. The species is also know to contain bioactive metabolites, including
anthraquinone, choline, triterpernoids and raffinose (Tabuti 2007; Mining et al. 2014). In
their findings, Mining et al. (2014) concluded that the root bark of S. didymobotrya contains
anthraquinone and triterpenoids and can be used as biopesticides against a bean weevil
beetle (Acanthoscelides obtectus Say) to improve grain production in Kenya.
b c
FIGURE 2.—Senna didymobotrya at Mokopane in Limpopo: a and b, in habitat; c, flowers and fruits; d,
plants form dense thickets. Photographs: T.P. Jaca.
Flowering Plants of Africa 65 (2017) 73
REFERENCES
BAKER, E.G. 1911. Contributions to the Flora of Gazaland. Journal of the proceedings of the Lin-
nean Society Botany 40: 62.
BAKER, E.G. 1930. Leguminosae of Tropical Africa. Erasmus Press, Ghent.
BOTANICAL DATABASE OF SOUTHERN AFRICA (BODATSA). 2016. Senna didymobotrya. South
African National Biodiversity Institute, http://newposa.sanbi.org/sanbi/Explore. [Accessed 17
March 2017].
BRENAN, J.P.M. 1967. Leguminosae: subfamily Caesalpinioideae. In E. Milne-Redhead & R.M.
Polhill (eds), Flora of Tropical East Africa 2: 66–68.
BRENAN, J.P.M. & GREENWAY, P.J. 1949. Checklist of the forest trees and shrubs of the British
Empire 5: Tanganyika Territory (part 2). Imperial Forest Institute, Oxford.
BRUMMITT, R.K., CHIKUNI, A.C., LOCK, J.M. & POLHILL, R.M. 2007. Leguminosae subfamily
Caesalpinioideae. Flora zambesiaca 3(2): 144–152.
DE WILDEMAN, E.M. 1902. Études Sur La Flore du Katanga. Annales du Musee du Congo (Belge).
Botanique 1(4–2): 49, t. 16, fig. 6–11. Brussels.
DRUMMOND, R.B. 1975. A list of trees, shrubs and woody climbers indigenous or naturalised in
Rhodesia. Kirkia 10,1: 243.
DULBERGER, R. 1981. The floral biology of Cassia didymobotrya and C. auriculata (Caesalpini-
aceae). American Journal of Botany 68: 1350–1360.
DU PUY, D.J., LABAT, J.N., RABEVOHITRA, R., VILLIERS, J.F., BOSSER, J. & MOAT, J. 2002. The
Leguminosae of Madagascar. Royal Botanic Gardens, Kew.
ENVIRONMENTAL WEEDS OF AUSTRALIA. 2016. Senna didymobotrya: http://keyserver.lucid-
central.org/weeds/data/media/Html/senna_didymobotrya.htm. [Accessed 10 June 2016].
FOXCROFT, L.C. & RICHARDSON, D.M. 2003. Managing alien plant invasions in the Kruger
National Park, South Africa. In L. Child, J.H. Brock, G. Brundu, K. Prach, P. Pyšek, P.M.
Wade & M. Williamson (eds), Plant Invasions: Ecological Threats and Management Solutions:
385–404. Backhuys Publishers, Kerkwerve.
FRESEN, J.B.G.G.W. 1839. Flora oder Botanische Zeitung: welche Recensionen, Aufsätze, Neuig-
keiten und Nachrichten, die Botanik betreffend, enthält /herausgegeben von der Königl.
Botanischen Gesellschaft in Regensburg 22,1: 53.
GERMISHUIZEN, G., MEYER, N.L., STEENKAMP, Y. & KEITH, M. (eds). 2006. A checklist of South
African plants. South African Botanical Diversity Network Report No. 41. SABONET, Preto-
ria.
74 Flowering Plants of Africa 65 (2017)
The genus Caesalpinia was established by Linnaeus in 1753, represented today by more
or less 200 species, of which most are confined to the New World (Heywood 1978). The
generic name, Caesalpinia, honours the Italian botanist Andreas Caesalpini (1519–1603)
(Jackson 1990). The specific epithet, bracteata, is in reference to the large bracts protecting
the flowers of C. bracteata. Most Caesalpinia species are attractive, scrambling shrubs, with
colourful flowers and many have been taken up in ornamental horticulture.
PLATE 2330.—1, flowering branch, × 1; 2, single flower, × 1.5. Voucher specimen: Van Hoepen 1941 in
National Herbarium, Pretoria. Artist: Gillian Condy.
2
Professor Braam van Wyk of the University of Pretoria brought this species to the atten-
tion of the first author (EJvJ) who decided to visit the site to collect seed for the Botan-
ical Society Conservatory in Kirstenbosch, where he was in charge. On an excursion to
the Kaokoveld in June/July 2001, and after consulting with one of the owners, Sakkie van
Staden, the farm Skroef was visited on 23 July 2001 with Tielman Haumann, a farmer from
the Ghaap Plateau. Caesalpinia bracteata was commonly encountered and often the domi-
nant species in some parts of the farm, and not far from the Orange River. It grew in crev-
ices and among Naros granitoid outcrops of the Namaqua Metamorfic Complex (Miller
& Schalk 1980; Visser 1984; Mendelsohn et al. 2002). Multistemmed shrubs, 1–3 m tall,
grew on a gentle north-facing slope. The vegetation consists of Lower Gariep Broken Veld
(Bushmanland Bioregion of the Nama-Karoo Biome) (Mucina & Rutherford 2006). Here
the plants shared their habitat with species such as Abutilon pycnodon Hochr., Adenolobus
garipensis (E.Mey.) Torre & Hillc., Boscia albitrunca (Burch.) Gilg & Gilg-Ben. and B. foetida
Schinz, Cadaba aphylla (Thunb.) Wild, Commiphora gracilifrondosa Dinter ex J.J.A.van der
Walt, Diopyros acocksii (De Winter) De Winter, Ficus cordata Thunb., Maerua gilgii Schinz,
Gymnosporia tenuispina (Sond.) Szyszyl., Nymania capensis (Thunb.) Lindb., Parkinsonia
africana Sond., Searsia marlothii (Engl.) Moffett, Senegalia mellifera (Vahl) Seigler & Ebinger
subsp. detinens (Burch.) Kyal. & Boatwr., Sisyndite spartea E.Mey. ex Sond., Tribulus terrestris
L., Vachellia erioloba (E.Mey.) P.J.H.Hurter and Ziziphus mucronata Willd. Sufficient seeds
and specimens (E. van Jaarsveld 16805) were collected. This part of southern Africa is a
semi-desert with the climate being hot and dry throughout most of the year. The annual
rainfall has been recorded as less than 100 mm per annum and occurs mainly in spring and
autumn in the form of thundershowers, although the region may occasionally experience
some rain during the winter months. The average daily maximum is 29°C and the average
daily minimum 13°C. During summer, extreme temperatures have been measured and can
sometimes reach 44°C.
80 Flowering Plants of Africa 65 (2017)
The seeds that were collected were sown in sandy soil in the following spring (2001)
in the succulent section of the Kirstenbosch National Botanical Garden. Germination was
rapid and many plants established, some of which were transplanted into the Botanical
Society Conservatory in 2002. These have a slow rate of growth, and the first plants flow-
ered in the Conservatory only in 2008 about six years after being planted out. It was also
from the plants in the Botanical Society Conservatory from which the artist was able to
illustrate the plant in October 2016.
Sakkie van Staden maintained that the species’ distribution extends to the Namibian
side of the Orange River and one of us (EJvJ) decided to inspect this at a later stage. Sub-
sequently an expedition was arranged to the Spelonkberge in Namibia in search of Aloe
dabenorisana Van Jaarsv., which provided the opportunity to investigate for possible Cae-
salpinia bracteata on the farm Naros in Namibia on 25 May 2011. The party consisted of
Wessel Swanepoel, Tielman Haumann, Natanya Mulholland, the farm owner (Nico Strauss)
and myself (EJvJ).
We left for the farm Naros (after which the granite type was named) and headed south
towards the Orange River. As we approached the farm, Naros granitoid outcrops appeared
Flowering Plants of Africa 65 (2017) 81
as expected, and Caesalpinia bracteata was indeed commonly encountered (Figure 2).
The plants were again always confined to crevices of Naros granite and growing in associa-
tion with Sisyndite spartea and Cynanchum (Sarcostemma) pearsonii N.E.Br. Here Caesal-
pinia bracteata grows mainly on south-facing slopes and are slightly smaller than the Skroef
plants. Caesalpinia bracteata also grows here as a dense, multistemmed shrub, up to 1 m
high, occasionally up to 2 m, with the plants just coming into flower (Figure 3). Young stems
are initially green with spines pointing to the ground, turning reddish and blackish as they
mature. Older stems die off, but are replaced by resprouting younger branches from the
plant base (Figure 4). Young leaves are reddish at first, turning olive-green as they mature.
Flowers are an attractive, pinkish colour.
Surprisingly, plants were also recorded by the ecologist Phillip Desmet fairly recently
at Pella to the west. These plants were growing in Eastern Gariep Rocky Desert (Desert
Biome) and extending its range further west into the Desert Biome.
Caesalpinia bracteata was assessed for the Red List of South African plants and its status
assessed as NT D2 (Not Threatened) (Victor et al. 2009).
Caesalpinia bracteata shows horticultural potential for semi-arid to arid gardens and has
been introduced into cultivation. Plants given to Kevin Koen more than a decade ago, and
planted in his garden at Calitzdorp, Western Cape, flower profusely and have here grown
into shrubs of up to 2 m tall. It is best grown in Desert, Succulent Karoo and Nama-Karoo
gardens, where frost is not too severe (Van Jaarsveld 2010).
FIGURE 4.—The lower, multistemmed portion of the plant. Note the white prickles. Photograph: E.J. van
Jaarsveld.
82 Flowering Plants of Africa 65 (2017)
REFERENCES
CRAVEN, P. (ed.). 1999. A checklist of Namibian plant species. Southern African Botanical Diversity
Network Report No. 7. SABONET, Windhoek.
GERMISHUIZEN, G. 1991. Caesalpinia bracteata, a new species from the Onseepkans area of the
Northern Cape Province. Bothalia 21,2: 152–154.
GERMISHUIZEN, G. & CONDY, G. 2001. Caesalpinia rostrata. Flowering Plants of Africa 57: 70–74.
GLEN, H.F. & GERMISHUIZEN, G. 2010. Botanical exploration of southern Africa, edn 2. Strelitzia
26. South African National Biodiversity Institute, Pretoria.
HENDERSON, L. 2001. Alien weeds and invasive plants. Plant Protection Research Institute, Agri-
cultural Research Council, Pretoria.
HEYWOOD, V.H. (ed.). 1978. Flowering plants of the World. Oxford University Press, Oxford.
JACKSON, W.P.U. 1990. Origins and meanings of names of South African plant genera. UCT Ecolab
Botany Department, Cape Town.
LINNAEUS, C. 1753. Species Plantarum. Laurentii Salvii, Stockholm.
MENDELSOHN, J., JARVIS, A., ROBERTS, C. & ROBERTSON, T. 2002. Atlas of Namibia. David
Philip Publishers, Cape Town.
MILLER, R.McG. & SCHALK, K.E.L. 1980. Geological map of South West Africa/Namibia. Geologi-
cal Survey of the Republic of South Africa and South West Africa/Namibia, Pretoria.
MUCINA, L. & RUTHERFORD, M.C. (eds). 2006. The vegetation of South Africa, Lesotho and
Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria.
NKONKI, T. & SWELANKOMO, N. 2003. Fabaceae. In G. Germishuizen & N.L. Meyer (eds), Plants
of southern Africa: an annotated checklist. Strelitzia 14. National Botanical Institute, Pretoria.
VAN JAARSVELD, E.J. 2010. Waterwise gardening in South Africa and Namibia. Struik, Cape Town.
VAN WYK, A.E. & SMITH, G.F. 2001. Regions of floristic endemism in southern Africa. A review
with emphasis on succulents. Umdaus Press, Pretoria.
Flowering Plants of Africa 65 (2017) 83
VICTOR, J.E., DESMET, P. & VAN WYK, A.E. 2009. Caesalpinia bracteata. In D. Raimondo, L. von
Staden, W. Foden, J.E. Victor, N.A. Helme, R.C. Turner, D.A. Kamundi & P.A. Manyama
(eds), Red List of South African plants. Strelitzia 25. South African National Biodiversity Insti-
tute, Pretoria.
VISSER, D.J.L. 1984. Geological map of the Republics of South Africa, Transkei, Bophuthatswana,
Venda and Ciskei and the Kingdoms of Lesotho and Swaziland. Government Printer, Pretoria.
Eriosema distinctum N.E.Br. in Kew Bulletin 4: 103 (1906); Medley Wood: 43 (1907); Bews:
505 (1913); Bews: 113 (1921); Verdcourt: 118 (1971); Ross: 208 (1972). Stirton & Gordon-Gray:
397 (1978).
The genus Eriosema (DC.) G.Don belongs to the tribe Phaseoleae and it consists of
about 150 species worldwide with 100 species in Africa (Leistner 2005; Candido 2016).
The name Eriosema is derived from the Greek words erion in reference to wool, and semeia
in reference to the indumentum on the back of the standard. The species epithet distinc-
tum is of Latin derivation and means separate or different, here probably referring to the
difference in flowers. Eriosema is closely related to Rhynchosia Lour. and differs from it
in seed characteristics; mature seeds have a long, linear hilum flanked by a conspicuous,
whitish, bilobed structure, which is largest near the point of funicular attachment (Grear &
Dengler 1976).
In his treatment of the genus Eriosema, Verdcourt (1971) realised that E. cordatum
E.Mey., E. populifolium Benth. ex Harv. and E. distinctum are very similar and he suggested
that a field study should be carried out to clarify the relationships among these three spe-
cies. Thus Stirton in 1975 did a field study and it revealed that the three species are mor-
phologically different and that they may be distinguished on three main characters, namely
the type of underground system, the vestiture of the plant, and the shape and the size
of the flowers (see the key for more differences between the three species). Eriosema dis-
tinctum is morphologically similar to E. cordatum and E. populifolium of which it can be
distinguished by its large flowers and exsertion of the stigma from the anthers (Stirton &
Gordon-Gray 1978).
Eriosema distinctum was first collected in 1884 by W. Tyson in KwaZulu-Natal and this
old specimen is housed in Compton Herbarium, Cape Town. The first person to describe
E. distinctum was Nicholas Edward Brown in 1906. He closely associated the species to
E. oblongum Harv. (currently known as Eriosema pauciflorum Klotzsch var. pauciflorum) and
E. kraussianum Meisn., but separated it based on its large flowers. Later, in 1978, Stirton &
Gordon-Gray stated that E. distinctum has commonly been confused with E. cordatum. It is
readily distinguished from the latter by its rhizomatous underground system, the presence
of persistent stipels that occur at the apex of the rachis, the general shape of the floral parts,
the length of the fruits and the exserted stigma.
Eriosema distinctum is distributed through southern tropical Africa, i.e. Angola, Zambia,
Zimbabwe, Malawi, Mozambique and southern Africa (Leistner 2005). In South Africa it
occurs mainly in the midlands and uplands of KwaZulu-Natal, extending along the lower
PLATE 2331.—1, flowering stem, × 1; 2, fruiting stem, × 1. Voucher specimens: 1, Condy 246 and 2, Acocks
13877, both in National Herbarium, Pretoria. Artist: Gillian Condy.
1
The common names for Eriosema distinctum are scarlet eriosema and Ubangalala
olukhulu in Zulu. The plant is used as human famine food. The Zulu people use decoctions
from pounded, boiled roots for urinary ailments and for impotence (Hutchings et al. 1996;
Pooley 2003). It is also used as a fish poison.
REFERENCES
BEWS, J.W. 1913. An Ecological Survey of the Midlands of Natal. Annals of the Natal Museum 2:
505.
BEWS, J.W. 1921. Eriosema. The Flora of Natal and Zululand: 113. City Printing works, Pieterma
ritzburg.
BROWN, N.E. 1906. Diagnoses Africanae XVI. Kew Bulletin 4: 103.
CANDIDO, E.S., DE VARGAS, W., VATANPARAST, M., DE FREITAS MANSANO, V., RODRIGUES
MACHADO, S. & FORTUNA-PEREZ, A.P. 2016. A new species of Eriosema (Leguminosae,
Papilionoideae, Phaseoleae) from Mato Grosso do Sul, Brazil, with a secretory structure
novel to the genus. Phytokeys 263,2: 122.
FODEN, W. & POTTER, L. 2005. Eriosema distinctum N.E.Br. National Assessment: Red List of
South African Plants version 2015.1. [Accessed on 9 March 2017].
GREAR, J.W. & DENGLER, N.G. 1976. The Seed Appendage of Eriosema (Fabaceae). Brittonia 28,3:
281–288.
HILLIARD, O.M. & BURTT, B.L. 1987. The Botany of the Southern Natal Drakensberg. Annals of
Kirstenbosch Botanic Gardens: 166. National Botanic Gardens, Cape Town.
HUTCHINGS, A., SCOTT, A.H., LEWIS, G. & CUNNINGHAM, A. 1996. Zulu medicinal plants an
inventory. University of Natal Press, KwaZulu-Natal.
LEISTNER, O.A. 2005. Seed plants of southern tropical Africa: families and genera: 192. Southern
African Botanical Diversity Network Report No. 26. SABONET, Pretoria.
Flowering Plants of Africa 65 (2017) 89
MEDLEY WOOD, J. 1907. Handbook to the Flora of Natal. Bennett & Davis, Durban.
PLANTS OF SOUTHERN AFRICA (POSA). 2016. Eriosema distinctum. http://posa.sanbi.org/search-
spp.php. [Accessed 2016].
POOLEY, E. 2003. Mountain flowers. A field guide to the Flora of the Drakensberg and Lesotho.
Flora Publications Trust, Durban.
ROSS, J.H. 1972. Flora of Natal. Department of Agricultural Technical Services, Pretoria.
SERGIO, A-P. 2012. Zooming in on floral nectar: a first exploration of nectar-associated bacteria in
wild plant communities. Fems Microbiology 80,3: 591–602.
STIRTON, C.H. 1975. A contribution to knowledge of the genus Eriosema (Leguminose, Lotoideae)
in southern Africa (excluding Mozambique & Rhodesia). M.Sc. thesis. University of Natal,
Pietermaritzburg.
STIRTON, C.H. & GORDON-GRAY, K.D. 1978. The Eriosema cordatum Complex. I. The Eriosema
populifolium Group. Bothalia 12,3: 395–404.
VERDCOURT, B. 1971. Studies in the Leguminosae-Papilionoideae for the Flora of Tropical East
Africa. Kew Bulletin 25: 118.
Adenia wilmsii is a rare, erect, little-branched shrublet endemic to the lower foothills of
the Mpumalanga and Limpopo Drakensberg from near Lydenburg in the north to Waterval
Boven in the south (Figure 1). It represents one of 10 species of Adenia Forssk. confined to
South Africa.
The Passifloraceae, to which our species belongs, have a pantropical distribution with
more or less 500 species, most of which are climbers. The largest and best known is the
genus Passiflora L. with its highest diversity of species centered in the Americas and to
which the edible granadilla plant or passion fruit (Passiflora edulis Sims) belongs. Forsskål
named the genus Adenia in 1775 based on A. venenata Forssk. (the type species), which
occurs in the Yemen, and northern eastern Africa (Ethiopia, Kenya, Somalia and Tanzania).
The name Adenia is derived from aden, meaning gland, and pertains to the glands found
on leaves of most of the species (Jackson 1990). Currently the genus Adenia consists of
95 species, of which most are stem or root succulents, distributed in Africa, Madagascar
and Australasia (De Wilde 2002). Most are toxic, occasionally with spines (A. spinosa Burtt
Davy, A. pechuelii (Engl.) Harms), while some have pachypodious caudices (A. fruticosa
Burtt Davy, A. pechuelii, A. spinosa). Adenia is divided into six sections (De Wilde 1971),
based on the position of leaf-glands, the inflorescence architecture and flower construc-
tion. Adenia wilmsii together with its close alies, A. digitata (Harv.) Engl., A. hastata (Harv.)
Schinz and A. huillensis (Welw.) A.Fernald & R.Fernald (all southern Africa), belongs to sec-
tion Blepharanthes (Wight & Arn.) Engl. This section consists of 34 species of small to large
climbers with tubers, of which only two are not African. In section Blepharanthes the glands
are present on the base of the lamina (two or one, or rarely absent), the flowers are uni- or
bisexual, the inflorescences axillary (leaves), and the sepals are also partialy connate into a
calyx tube.
From their close relatives, Adenia wilmsii and A. huillensis (Angola) are immediately
separated by their stems that are without tendrils. However, there the resemblance ends,
as A. huillensis has leaves that are entire and A. wilmsii has leaves that are divided with a
rounded outline.
Adenia wilmsii was named in 1899 by Harms from plants collected by Friedrich Wilms
(1848–1919), a German chemist from Münster, Westphalia, Germany. Wilms came to
South Africa in 1883, arriving at the Cape on 4 July 1883 from where he continued his
journey by sea to Durban. From here he took the train to Pietermaritzburg and then trav-
PLATE 2332.—Plant in fruit, × 1. Voucher specimen: Condy 247, National Herbarium, Pretoria. Artist: Gillian
Condy.
PLATE 2332 Adenia wilmsii
Flowering Plants of Africa 65 (2017) 93
The glands at the bases of leaves act as extrafloral nectaries and secrete a type of nec-
tar. Ants are attracted to the nectar and feed off it, in turn, protecting the plants. Addition-
ally, Adenia plants are inedible to very toxic to humans, as they contain cyanogenic com-
pounds with the possible presence of modeccine.
The genus Adenia, with their attractive grey to green caudices and tapering stems, is
favoured among succulent plant enthusiasts. Popular southern African species with large
caudices include A. glauca Schinz, A. pechuelii and A. spinosa. Adenia fruticosa has an
attractive tapering green and grey caudex. Adenia wilmsii plants, like their close relatives,
have their tubers below or partly above the ground and do not have the same appeal as
most of the larger caudiciform plants; therefore they are mainly grown as a curiosity and
for their rarity. One of us (EJvJ) was privileged to be able to grow A. wilmsii for a number
of years in the Botanical Society Conservatory at Kirstenbosch National Botanical Garden
94 Flowering Plants of Africa 65 (2017)
from plants collected and donated by Fanie Venter, formerly from the University of the
North (University of Limpopo, Turfloop campus). According to Venter (1993), who collected
the plants in the Lydenburg district, A. wilmsii grows between 1 300 and 1 500 m above
sea level. He collected his plants on loam to clay loam, dolerite-derived soil and observed
flowers and fruits from September to December.
Adenia wilmsii is easily grown in a well-drained soil mixture and would be best suited
in bushveld and highveld gardens (Van Jaarsveld 2010). It prefers full sun but will also grow
in light shade.
REFERENCES
DE WILDE, W.J.J.O. 1971. A monograph of the genus Adenia Forssk. (Passifloraceae). Mededelin-
gen Landouwhogeschool Wageninen 71,18: 1–281.
DE WILDE, W.J.J.O. 1976. Adenia. In J.H. Ross (ed.), Flora of southern Africa 22: 106–117.
DE WILDE, W.J.J.O. 2002. Adenia. In U. Eggli (ed.), Illustrated Handbook of Succulent Plants: 336–
350. Springer, Berlin.
FORSSKÅL, P. 1775. Flora Aegyptiaco-Arabica. Haunia, Copenhagen.
GUNN, M. & CODD, L.E. 1980. Botanical exploration of southern Africa. Balkema, Cape Town.
HARMS, H. 1899. Passifloraceae. In A. Engler (ed.), Botanische Jahrbücher fur Systematik, Pflanzen-
geschichte und Pflanzengeographie 26: 238.
JACKSON, W.P.U. 1990. Origins and meanings of names of South African plant genera. UCT Ecolab,
Cape Town.
MUCINA, L. & RUTHERFORD, M.C. (eds). 2006. The vegetation of South Africa, Lesotho and
Swaziland. Strelitzia 19. South African National Biodiversity Institute, Pretoria.
VAN JAARSVELD, E.J. 2010. Waterwise gardening in South Africa and Namibia. Struik, Cape Town.
Flowering Plants of Africa 65 (2017) 95
VENTER, S. 1993. Notes on the genus Adenia with special reference to the Transvaal species. Aloe
30,3 & 4: 83–88.
VON STADEN, L. ARCHER, R.H. & KRYNAUW, S. 2009. Adenia wilmsii. In D. Raimondo, L. von
Staden, W. Foden, J.E. Victor, N.A. Helme, R.C. Turner, D.A. Kamundi & P.A. Manyama
(eds), Red List of South African plants. Strelitzia 25. South African National Biodiversity Insti-
tute, Pretoria.
Esterhuysenia lucilleae Van Jaarsv. sp. nov. differs from E. grahambeckii Van Jaarsv. by its shrub-
by habit and much larger, slightly laterally compressed leaves and floral petals of 35–40 × 1.5 mm,
as well as its flowering time, which is during spring.
TYPE.—South Africa, Western Cape, 3319 (Worcester): Dome Kloof, Hex River Moun-
tains, ledges on south-facing cliffs, (–AD), Van Jaarsveld & Stander 26952 (NBG, holotype).
Cliffs on rugged mountainous areas are of the least botanically explored habitats in the
world for understandable reasons. I have always watched the huge cliffs of the Hex River
Mountains below Matroosberg – at an altitude of 2 249 m the highest peak of the Hex
River Mountains – and for many years wondered about the best way to approach these rock
faces as part of a study of obligatory succulent and bulbous cliff-dwelling plants (cremno-
phytes). I was then fortunate to join the snow protea (Protea cryophila) team, an annual event
arranged by Ernst Hartwig, in January 2013 to the Cederberg where I met Retief Jordaan,
grape farmer and mountaineer from the Hex River Mountains. He told me that for the past
35 years he had been leading mountaineering expeditions down the Groothoekkloof below
the Matroosberg (Figure 1), and that I was welcome to join them on yet another expedi-
tion in February 2013. The Groothoekkloof represents one the most rugged and dangerous
kloofs in the area (Figure 2), with a series of about 12 abseils (some directly into pools or
next to wet waterfalls). The Groothoekkloof starts from near the top of Matroosberg at about
2 000 m above sea level, and can only be explored during the hot, dry season (February)
due to extreme conditions at other times of the year. I joined Jordaan’s team the following
month and was not disappointed, as it was here that I came across an unnamed species of
Esterhuysenia (named E. lucilleae here), as well as a new species of Haemanthus in flower.
This mesemb, an ascending soft-leafed succulent shrublet with glaucous leaves, was
encountered on the middle southern slopes above 1 000 m, growing on a sheer rock wall
(Figure 3). The plant was only found in fruit, and I immediately identified it as a species of
Esterhuysenia by its vegetative growth as well as its capsules, which showed affinity with
E. grahambeckii – a fairly recently named species from the Rooiberg (Van Jaarsveld 2011b).
I realised that it possibly represented a new species. Although I collected cuttings to grow
and observe the flowers, I knew that many of these alpine mesembs refuse to flower in
warm terrain close to sea level and, in spite of the best possible care, these indeed did not
flower at Kirstenbosch National Botanical Garden.
The genus Esterhuysenia (Ruschioideae) is a small mesemb genus of six species, confined
to mountainous peaks along the Western Cape, South Africa. They are characterised by their
PLATE 2333.—1, seedling, much reduced; 2, mature flowering branches, × 1; 3 and 4, developing fruit, × 2
and × 2.5 respectively; 5, mature open capsule, × 2.5. Voucher specimen: Van Jaarsveld & Stander
26952, Compton Herbarium, Cape Town. Artist: Marieta Visagie.
3
1 5
During December 2013 I joined yet another expedition to the Hex River Mountains
with Retief Jordaan and Kallie Pauw, this time to the Buffelshoekkloof where we ascended
and stayed overnight at Perry Refuge (a mountain hut). Early the second morning we con-
tinued to Buffelhoek Peak, past the Shale
Peaks, and descended via Milner Kloof.
Our mission was to investigate the south-
facing cliffs along this portion of the Hex
River Mountains for any cremnophytes.
Again we encountered Esterhuysenia lucil-
leae on cliffs in the Buffelshoekkloof, as well
as Milner Kloof (Figure 4) and now realised
that it was quite common on sheer, south-
facing cliffs along the Hex River Mountains
from the Buffelshoekkloof to Matroosberg
in the northeast.
FIGURE 3.—Esterhuysenia lucilleae at the Groothoekkloof below Matroosberg. Photograph: E.J. van Jaars
veld.
FIGURE 6—Esterhuysenia lucilleae in flower at Stettynskloof where Anso le Roux originally found plants in
2011. Photograph: E.J. van Jaarsveld.
102 Flowering Plants of Africa 65 (2017)
TABLE 1.—Differences in the vegetative, reproductive and habitat characters of Esterhuysenia grahambeckii
and E. lucilleae
Calyx lobes and floral petals Calyx lobes as long as fruit, floral Calyx lobes twice as long as fruit,
petals 25–30 × 1 mm floral petals 35–40 × 1.5 mm
Flowering time and floral diameter Summer (December), 50–65 mm Spring (October), 70–80 mm
Capsule 8–9 mm in diameter, 5 mm deep 7–8 mm in diameter, 5–7 mm deep
Seed 1.3 × 1 mm, reddish-brown 1.5 × 0.9 mm, reddish-brown
Vegetation type Breede Quartz Fynbos South Hex Sandstone Fynbos
Geology (Cape Supergroup) Witteberg Group Table Mountain Group
the plant after pollination when the inner floral parts become visible. Unlike most other
mesembs from non-cliff sites, the cliff-dwelling mesembs’ hygroscopic capsules remain
open after initial opening, with short, erect, covering membranes and slightly flattened
seeds, which are wind dispersed (Van Jaarsveld & Van Wyk 2003). This indicates that wind
is the major agent used by obligatory suc-
culent cremnophytes for dispersing their
seeds. The flattish seeds are deposited in
crevices or among lichens on the cliff where
germination takes place. Seeds of most
other mesembs are dispersed by raindrops,
a mechanism known as ombrohydrochory.
The kinetic energy of falling raindrops eject
the seeds (Hartmann 1990).
our new species thus extends along the Hex River Mountain Range as well at Stettynskloof
near Rawsonville (Figure 1).
At the Groothoekkloof and other sites I have observed that Esterhuysenia lucilleae shares
its habitat with other succulents such as Aloe perfoliata, Crassula atropurpurea var. anomala,
C. dentata, C. nudicaulis, C. obtusa, C. pellucida subsp. spongiosa, Haemanthus sp. and
Haworthia arachnoidea. Plants grow in shallow soil in crevices on cliffs or steep slopes, often
among moss and lichens in full light, and often in areas that cannot be reached by fires.
We have pleasure in naming this species after Lucille Krige, a keen hiker in the moun-
tains of the Western Cape, who regularly notifies me of succulents observed on her expe-
ditions. She brought the flowering plant to my attention, which led to its description here.
FIGURE 8.—Close-up of the rose-pink flowers of Esterhuysenia lucilleae. Note persistent petals of the old
flower. Photograph: E.J. van Jaarsveld.
104 Flowering Plants of Africa 65 (2017)
SPECIMENS EXAMINED
Western Cape: 3319 (Worcester). (–AD): Dome Kloof, sheer cliffs, Van Jaarsveld & Krige
25141, 25563 and Van Jaarsveld & Stander 26952 (NBG); Buffelshoekkloof, sheer cliffs, Van Jaars-
veld, Pauw & Jordaan 25154 (NBG); Milnerkloof, on boulders upper end of Kloof, Van Jaarsveld
& Jordaan 25184 (NBG). (–BC): Groothoekkloof, sheer south-facing cliffs, Van Jaarsveld & Jor-
daan 24703 (NBG). (–CD): Stettynskloof, sheer south-facing cliffs, Van Jaarsveld, Le Roux & Stander
26964 (NBG).
ACKNOWLEDGEMENTS
We are thankful to various people for help. Koos Bekker and Karen Roos, owners of
Babylonstoren Farm, for allowing me to do research on various succulent and other plant
species. We are also thankful for Retief Jordaan, retired wine farmer for arranging the
Groothoekkloof and Buffelskloof expeditions, as well as DeWet Conradie (Kanetvlei) for
allowing us access to Dome Kloof. Anso le Roux (Pokkraal), who brought me photos and
live material from Stettynskloof, and Arne Stander is thanked for assistance in the field near
Dome Kloof. Lucille Krige and Tom Jordan are thanked for taking me to Dome Kloof twice.
REFERENCES
HARTMANN, H.E.K. 1990. Mesembryanthema. In H.P. Linder & A.V. Hall, Systematics, biology
and evolution of some South African taxa. Contributions from the Bolus Herbarium No. 13.
HARTMANN, H.E.K. 2001. Aizoaceae A–E. Illustrated Handbook of Succulent Plants. Springer,
Berlin.
MUCINA, L. & RUTHERFORD, M.C. (eds). 2006. The vegetation of South Africa, Lesotho and
Swaziland. Strelitzia 19. South African National Biodiversity Institute. Pretoria.
VAN JAARSVELD, E.J. 1988. The succulent riches of South Africa and Namibia. Aloe 24, 3 & 4:
11–89.
VAN JAARSVELD, E.J. 2011a. Cremnophilous succulents of southern Africa: diversity, structure and
adaptations. Ph.D. thesis, University of Pretoria, Pretoria.
Flowering Plants of Africa 65 (2017) 105
VAN JAARSVELD, E.J. 2011b. Esterhuysenia grahambeckii, a new rare species in the Aizoaceae
(Mesembryanthema, Ruschioideae) from the Rooiberg, Western Cape South Africa. Brad-
leya 29: 43–50.
VAN JAARSVELD, E.J. & VAN WYK, A.E. 2003. Adaptations without barriers: succulent plants
shaped by cliffs. Aloe 40: 98–103.
Pisonia aculeata L. in Species Plantarum 2: 1026 (1753). Pisonia mitis L.: 1026 (1753); Piso-
nia villosa Poir.: 347 (1804). Pisonia loranthoides Kunth: 197 (1825). Pisonia limonella Blume: 735
(1826). Pisonia sieberi Schltr.: 876 (1849). Pisonia monotaxadenia C.Wright.: 199 (1870). Pisonia
yagua-pinda D.Parodi: 211 (1878).
Pisonia is morphologically distinct from the other Nyctaginaceae genera (namely Boer-
havia L., Commicarpus Standl., Mirabilis L. and Phaeoptilum Radlk.) in southern Africa. Boer-
havia, Commicarpus and Mirabilis are unarmed, annual or perennial herbs, with white, pink,
purple, or variegated flowers, while Pisonia has yellow or greenish flowers. Phaeoptilum is
an armed shrub with greenish-yellow flowers, but differs from Pisonia in having linear leaves
and winged anthocarps (Pisonia has ribbed anthocarps with viscid stipitate glands). A com-
parison of the vegetative and reproductive characters of the Nyctaginaceae genera from
southern Africa is given in Table 1 of Struwig et al. (2015).
A second species of Pisonia, namely P. grandis, occurs in tropical Africa (Klopper et al.
2006). It can be distinguished from P. aculeata in that it is an unarmed and tall tree up to
30 m tall. The bisexual flowers are arranged in terminal cymes and the ribs on the anthocarps
PLATE 2334.—1, flowering branch, × 1; 2, male flower, × 4; 3, female flower, × 1; 4, fruiting branch, × 1; 5,
pair of curved thorns, × 1. Voucher specimen: Struwig 158 in University of Zululand Herbarium. Artist:
Gillian Condy.
1
2 5
a b c
are covered in a single row of viscid, stipitate glands (Stemmerik 1964; Whitehouse 1996).
Pisonia aculeata, however, has unisexual flowers arranged in axillary cymes and the ribs on
the anthocarps are covered in a double row of viscid, stipitate glands (Table 1).
The leaves of Pisonia aculeata are edible (Zomlefer 1994), and the stem and leaves are
used in Jamaica, the Philippines and India to treat various ailments such as rheumatism,
swelling, scabies, jaundice, coughing and venereal diseases (Quisumbing 1951; Ayensu
1978; Nadkarni 2005). In Africa, the juice from the leaves is used as drops against earache
in Côte d’Iviore (De Ruijter 2008), and in Ghana it is considered to be a good indication
of soil that is suitable for growing cacao trees (Hutchinson & Dalziel 1937). There are no
recorded uses for this plant in southern Africa.
TABLE 1.—Comparison of the vegetative and reproductive characters of the Pisonia species present in tropical
Africa
Pisonia aculeata flowers from November to April and produces fruits from January to
May.
REFERENCES
AYENSU, E.S. 1978. Medicinal plants of West Africa. Reference Publications, Michigan.
BLUME, C.L. 1826. Bijdragen tot de flora van Nederlandsch Indie, Vol. 14. Ter Lands Drukkerij,
Batavia.
BOGLE, A.L. 1974. The genera of Nyctaginaceae in the south-eastern United States. Journal of the
Arnold Arboretum 55: 1–37.
BOON, R. 2010. Pooley’s trees of eastern South Africa: A complete guide. Flora and Fauna Publica-
tion Trust, Durban.
BURGER, A.E. 2005. Dispersal and germination of seeds of Pisonia grandis, an Indo-Pacific tropical
tree associated with insular seabird colonies. Journal of Tropical Ecology 21: 263–271.
Flowering Plants of Africa 65 (2017) 111
DE RUIJTER, A. 2008. Pisonia lanceolata (Poir.) Choisy. In G.H. Schmelzer & A. Gurib-Fakim (eds),
Plant resources of tropical Africa 11(1). Medicinal plants 1. PROTA foundation, Wageningen /
Backhuys Publishers, Leiden / CTA, Wageningen.
DOUGLAS, N.A. & SPELLENBERG, R. 2010. A new tribal classification of Nyctaginaceae. Taxon
59: 905–910.
GERMISHUIZEN, G. & MEYER, N.L. (eds). 2003. Plants of southern Africa: An annotated checklist.
Strelitzia 14. National Botanical Institute, Pretoria.
HUTCHINSON, J. & DALZIEL, J.M. 1937. The useful plants of West tropical Africa. Crown Agents
for Overseas Governments and Administrations, London.
KLOPPER, R.R., CHATELAIN, C., BÄNNINGER, V., HABASHI, C., STEYN, H.M., DE WET, B.C.,
ARNOLD, T.H., GAUTIER, L., SMITH, G.F. & SPICHIGER, R. 2006. Checklist of the flower-
ing plants of Sub-Saharan Africa: An index of accepted names and synonyms. South African
Botanical Diversity Network Report No 42. SABONET, Pretoria.
KUNTH, K.S. 1825. Nova Genera et Species Plantarum. Lutetiae Parisiorum: sumtibus. Librariae
Graeco-Latino-Germanico, Paris.
LAMARCK, J.B.A.M. & POIRET, J.L.M. 1804. Encyclopédie méthodique: Botanique. Chez H.
Agasse, Paris.
LINNAEUS, C. 1753. Species Plantarum, Vol. 2. Laurentius Salvius, Stockholm.
NADKARNI, A.K. 2005. Indian Materia Medica, Vol 1. Popular Prakashan, Bombay.
PALGRAVE, K.C. & PALGRAVE, M.C. 2002. Trees of Southern Africa, edn 3. Struik, Cape Town.
PARODI, D. 1878. Contribuciones á la Flora del Paraguay. Anales de la Sociedad Cientifica Argen-
tina 5: 207–214.
PLUMIER, C. 1703. Nova plantarum americanarum genera. Joannem Boudot, Paris.
QUATTROCCHI, U. 2012. CRC World Dictionary of Medicinal and Poisonous Plants: Common
names, scientific names, eponyms, synonyms, and etymology. CRC Press, Boca Raton.
QUISUMBING, E. 1951. Medicinal plants of the Philippines. Bureau of Printing, Manila.
RUMPHIUS, G.E. 1747. Herbarium Amboinense, Vol 5. Apud Fransicum Changuion, Joannem
Catuffe, Hermannum Uytwerf, Amstelaedami.
SAUVALLE, F.A. 1870. Revision catalogi grisebachian vel index plantarum cubensium. Anales de la
Academia de Ciencias Medicas, Fisicas y Naturales de la Habana 7: 194–200.
SCHLECHTENDAL, D.F.L. von 1849. Ueber die gattung Pisonia Plum. Linneae 22: 868–885.
SPELLENBERG, R.W. 2004. Nyctaginaceae. Flora of North America, Vol. 4. Oxford University
Press, New York.
STANNARD, B.L. 1988. Nyctaginaceae. In E. Launert (ed.), Flora zambesiaca, Vol. 9. Halesworth
Press, London.
STEARN, W.T. 1992. Botanical Latin, edn 4. David and Charles, Newton Abbott.
STEMMERIK, J.F. 1964. Nyctaginaceae. In C.G.G.J. van Steenis (ed.), Flora Malesiana, Vol. 6.
P. Noordhoff, Haarlem.
STRUWIG, M., SIEBERT, S.J., ZIETSMAN, L. & CONDY, G. 2015. Phaeoptilum spinosum. Flowering
Plant of Africa 64: 84–89.
WHITEHOUSE, C. 1996. Nyctaginaceae. In R.M. Polhill (ed.), Flora of tropical East Africa. Balkema,
Rotterdam.
ZOMLEFER, W.B. 1994. Guide to flowering plant families. The University of North Carolina Press,
Chapel Hill.
The genus Schizostephanus was established by Bentham and Hooker (1876). It was sunk
under the genus Cynanchum L. by Brown in 1903. Liede (1994) reinstated Schizostephanus
based on the long hairs on the adaxial side of the corolla lobes, stipitate gynostegium and the
attachment of the pollinia along the dorsal rims, as well as the clear sap of its only two mem-
bers, S. gossweileri (S.Moore) Liede and S. alatus. A contribution on S. gossweileri appeared
in a previous volume of this journal (Van Jaarsveld & Visagie 2015) and is only known from
northern Namibia and adjacent southwestern Angola (Bruyns & Klak 2009). The genus Schi
zostephanus is related to Cynanchum, the latter with about 250 species and a cosmopolitan
distribution of which 31 are confined to Africa (Liede 1996). Cynanchum somaliense N.E.Br.
has, for some time, been treated as a close relative of S. alatus, however, the evidence is based
on superficial morphological character expressions (Liede 1996). Species of Cynanchum differ
by their usually evergreen foliage, umbellate inflorescences, distinct milky latex and with most
members not distinctly succulent (Bruyns & Klak 2009). The familiar Cynanchum viminale (L.)
Bassi (Sarcostemma viminale) is probably the best known species recently transferred to Cynan-
chum. Schizostephanus alatus differs from S. gossweileri by its smaller stature, leaves and repro-
ductive features. The stems of S. alatus are 5–7 mm in diameter, with stems of S. gossweileri
10 mm in diameter. Leaves of S. gossweileri are slightly pubescent with blades 60–130 mm
long, whereas they are glabrous with blades 35–80 mm long in S. alatus. Inflorescences are
bostrychoid in S. alatus and elongated in S. gossweileri. The specific epithet, alatus, is in refer-
ence to the species’ follicles which are distinctly winged.
Our depicted plant of Schizostephanus alatus (Van Jaarsveld, Haumann & Swanepoel
23008) was collected on Serra das Neves in southwestern Angola in April 2010 on an
exposed northeastern scree just above Aalwynnek, where we camped at about 1 300 m
PLATE 2335.—1, plant showing mature branches (left), and younger branches with inflorescence (right), × 1;
2, corolla from below, × 3.5; 3, corolla from above, × 3.5; 4, cross section of the young stem, × 1.
Voucher specimen: Van Jaarsveld, Haumann & Swanepoel 23008 in Compton Herbarium, Cape Town.
Artist: Marieta Visagie.
1
3 4
2
FIGURE 2.—Scree habitat of Schizostephanus alatus on Serra das Neves in southwestern Angola at elevation
of about 1 500 m. Photograph: E.J. van Jaarsveld.
116 Flowering Plants of Africa 65 (2017)
FIGURE 3.—Schizostephanus alatus in habitat at the Strydom Tunnel, Mpumalanga. Photograph: E.J. van
Jaarsveld.
Schizostephanus alatus was also collected from screes at the base of cliffs on the Leba
Pass west of Lubango in southwestern Angola (Van Jaarsveld 22609). Here the plants were
growing below steep cliffs on scree in association with Adenia gummifera (Harv.) Harms,
Dombeya rotundifolia (Hochst.) Planch., Euphorbia vallaris L.C.Leach, Kalanchoe lanceolata
(Forssk.) Pers., K. laciniata (L.) DC., Myrothamnus flabellifolius Welw., Obetia carruthersiana,
Sansevieria hyacinthoides and S. pearsonii.
In Mpumalanga plants (Van Jaarsveld 1097) were observed along the Crocodile Gorge
between Mbombela and Malelane growing in granite screes below Homalium dentatum
(Harv.) Warb. trees in Malelane Mountain Bushveld of the Lowveld Bioregion (Mucina &
Rutherford 2006) at an altitude of about 450 m. During the colder winter months the plants
becomes deciduous.
Schizostephanus alatus was also observed on sandstone scree adjacent to the Strydom
Tunnel in Mpumalanga in April 2015 (Van Jaarsveld 25851) (Figure 3). Plants grew in asso-
ciation with Cotyledon barbeyi Schweinf. ex Baker, Crassula ovata (Mill.) Druce, Dombeya
autumnalis I.Verd., Dracaena transvaalensis Baker, Portulacaria afra Jacq. and Sclerochiton
ilicifolius A.Meeuse.
One may very well ask what characteristics give a twining shrub a competitive advan-
tage over typical, woody, savanna plants. Schizostephanus alatus, with its succulent stems, is
Flowering Plants of Africa 65 (2017) 117
able to survive and grow successfully in dry rocky scree where the soil dries out fast. There-
fore, the species has less competition from trees and shrubs in that particular microhabitat,
especially in terms of soil moisture for root growth and light. This ecological advantage ena-
bles S. alatus to grow in soil that remains dry throughout the winter months, where it simply
resprouts new shoots and leaves during the summer rainy season.
Schizostephanus grows well in cultivation, both in containers and, if the climate permits,
in dry, well-drained, hot gardens. It is best grown in bushveld gardens (Van Jaarsveld 2010).
The plant can be propagated from stem cuttings rooted in clean sand. If it is grown outside
of its natural habitat, it fares best when grown in a greenhouse under controlled conditions.
If the climate permits it is best grown on steep embankments, gabions, or rockeries where
frost is either absent or not too severe.
Specimens examined: Angola: Serra das Neves, middle southwestern screes, altitude
1 500 m, 22 April 2010, Van Jaarsveld, Haumann & Swanepoel 23008 (NBG); Leba Pass,
west of Lubango, on screes below cliffs, 1 600 m, 16 January 2009, Van Jaarsveld 22609
(NBG). South Africa: 26 km from Nelspruit [Mbombela] towards Kaap Muiden–Krokodil
Poort, 13 March 1976, Van Jaarsveld 1097 (PRE); Strydom Tunnel, 1 308 m altitude,
12 April 2015, Van Jaarsveld 25851 (NBG).
ACKNOWLEDGEMENTS
We thank Wessel Swanepoel and Tielman Haumann for accompanying one of us (EJvJ)
to Serra das Neves and Professor Brian Huntley for arranging the expedition to southwest-
118 Flowering Plants of Africa 65 (2017)
ern Angola in January 2009. We are further thankful for live plants of Schizostephanus alatus
that was provided by Douglas McMurtry from Witrivier, near Nelspruit to study.
REFERENCES
BENTHAM, G. & HOOKER, J.D. 1876. Genera Plantarum. Part 2. Reeve, London.
BROWN, N.E. 1903. Asclepiadaceae. In W.T. Thiselton-Dyer (ed.), Flora of Tropical Africa 4,1:
231–503. Lovell Reeve, London.
BRUYNS, P.V. & KLAK, C. 2009. The rediscovery of Schizostephanus gossweileri and its phyloge-
netic position. South African Journal of Botany 75: 532–536.
GOYDER, D., HARRIS, T., MASINDE, S., MEVE, U. & VENTER, J. 2012. Apocynaceae. Flora of
Tropical East Africa. Part 2: 115–530.
LIEDE, S. 1994. Myth and reality of the subtribe Astephaniae (Decne.) Schumann (Asclepiadaceae).
Botanical Journal of the Linnean Society 114: 81–98.
LIEDE, S. 1996. A revision of Cynanchum (Asclepiadaceae) in Africa. Annals of the Missouri Botani-
cal Garden 83: 283–345.
LIEDE, S. 2003. Asclepiadaceae (in part). In I. Hedberg, S. Edwards & S. Nemomissa (eds), Flora of
Ethiopia and Eritrea, vol. 4: 99–193.
MUCINA, L. & RUTHERFORD, M.C. 2006. The vegetation of South Africa, Lesotho and Swazi-
land. Strelitzia 19. South African National Biodiversity Institute, Pretoria.
SCHUMANN, K.M. 1893. Asclepiadaceae africanae. Botanische Jahrbücher für Systematik, Pflanz-
engeschichte und Pflanzengeographie 17(1–2): 114–155.
VAN JAARSVELD, E.J. 2010. Waterwise gardening in South Africa and Namibia. Struik, Cape Town.
VAN JAARSVELD, E.J. & VISAGIE, M. 2015. Schizostephanus gossweileri. Flowering Plants of Africa
64: 102–107.
WIGHT, R. & ARNOTT, G.A.W. 1834. Asclepiadeae Indicae. Contributions to the Botany of India:
29–67.
The genus Gomphocarpus R.Br. is a relative small genus from the Apocynaceae (sub-
family Asclepiadoideae) that is naturally found in mostly arid parts of eastern and southern
Africa, extending into the Arabian peninsula, Sinai and north to the Dead Sea (Victor et
al. 2000; Goyder & Nicholas 2001). Due to its robust habit and (in many cases) extensive
populations commonly found in disturbed roadside habitats, Gomphocarpus is one of the
more distinct elements of the Asclepias L. complex in Africa, together with Pachycarpus
E.Mey. and some Xysmalobium R.Br. species. Some species, like Gomphocarpus fruticosus
(L.) Aiton f. and G. physocarpus E.Mey., have become weedy (Forster 1994), but some are
considered to be useful plants (Hutchings et al. 1996; Sobrina et al. 2002; El-Seedi et al.
2013; Hassan-Abdallah et al. 2013; Youssef 2013) in many parts of the world. Surprisingly,
only one Gomphocarpus species (then as Asclepias fruticosa L.) has thus far been covered in
this series (Phillips 1926).
The genus Gomphocarpus was proposed by Robert Brown (1810) and formalised by
Aiton (1811) as a distinct group of the family, characterised by the absence of a tooth in
the corona lobe cavity, and the inflation and ornamentation of the fruit. Brown argued
that the former Asclepias arborescens L., A. fruticosa and A. setosa Forssk. (Linnaeus 1753,
1771; Burmann 1768; Miller 1768; Forsskål 1775; Salisbury 1796) needed to be included
in Gomphocarpus. A long-standing controversy ensued, where taxa of this entity were
either attributed to Asclepias (Schlechter 1896; Brown 1902, 1907–1908; Weimarck 1935)
or to Gomphocarpus (Meyer 1837; Decaisne 1838, 1844). This further coincided with an
alternative view that Asclepias should be retained only for plants occurring in the North-
ern Americas, and that those known as Asclepias from Africa, including Gomphocarpus,
should move to other genera (Decaisne 1838, 1844; Bullock 1952; Smith 1988; Nicho-
las & Goyder 1990; Goyder 1995, 1998a, 1998b, 2001a, 2001b; Fishbein et al. 2011).
An alternate view is that Asclepias in a broad sense should be upheld and include taxa
from both the Old and New Worlds (Weimarck 1935; Woodson 1941, 1954). In the most
recent revision of Gomphocarpus, Goyder and Nicholas (2001) give a detailed overview
of the complexities of the movement of taxa between Gomphocarpus and Asclepias, and
the segregate genera of Asclepias that include Aspidoglossum E.Mey., Cordylogyne E.Mey.,
Glossostelma Schltr., Lagarinthus E.Mey., Pachycarpus, Parapodium E.Mey., Trachycalymma
PLATE 2336.—1, apical portion of stem depicting pendent umbels, × 1; 2, fruit, × 1. Voucher specimen:
Condy 272 in National Herbarium, Pretoria. Artist: Gillian Condy.
122 Flowering Plants of Africa 65 (2017)
The revision by Goyder and Nicholas (2001) of Gomphocarpus, the synopsis of Ascle-
pias for Africa (Goyder 2009), and continued work on the American species of Asclepias
(Fishbein et al. 2011) suggests that understanding the position of Gomphocarpus may be
key to our understanding of how the Old World and New World parts of the Asclepias
complex evolved. This highlights the recurring problem of the circumscription of many
genera of the subtribe Asclepiadinae in Africa, to which the genus Gomphocarpus belong.
Some of the morphological features, for example, are shared between the G. glaucophyl-
lus and Calotropis procera (Aiton) W.T.Aiton and the Xysmalobium undulatum (L.) Aiton f.
groups (Nicholas 1999), indicating a possible common ancestor, although phylogenetic
work shows different relationships.
Characters that delimit the genus Gomphocarpus and distinguish it from Asclepias (at
least in Africa) are presented by Goyder and Nicholas (2001) and summarised in Table 1.
Plants of G. glaucophyllus exhibit the following characteristic features of the genus: numerous
extra-axillary inflorescences with drooping flowers (Figure 1), and the staminal corona lobes
are laterally flattened (Figure 2d, 2e) with a central cavity (Figure 2c) and with teeth on the
upper margins (Figure 2c–e). According to Goyder and Nicholas (2001), and supported here
by one of us (SPB), the morphological structure of the corona lobe is remarkably constant
within the recognised species and this remains one of the single most useful organs for spe-
cies delimitation and recognition (particularly when working with herbarium material).
In the key to the species of Gomphocarpus by Goyder and Nicholas (2001) the genus
is divided into two groups: a group with narrow leaves (the first to be described and bulk
of the recognised taxa) and a group with broad leaves. The first members of the broad-
leaved group were only described some eighty years after the first narrow-leaved members
and are the closely related G. semiamplectens K.Schum. from Angola by Schumann (1893)
and G. glaucophyllus from the mountains around Barberton in South Africa by Schlechter
(1894).
TABLE 1.—Comparison of characters used to distinguish Asclepias from Gomphocarpus (adapted from Goyder
& Nicholas 2001, with additional characters from Brown 1810 indicated with an asterisk[*])
a b
FIGURE 1.—Gomphocarpus glaucophyllus: a, pendent inflorescences axillary along the stem; b, inflorescences
viewed from below; c, plant in habitat showing multiple stems (in this case approximately 35) arising
from underground rhizomatous rootstock. Voucher: Bester 11619 (PRE). Photographs: S.P. Bester.
124 Flowering Plants of Africa 65 (2017)
The only species in the genus with lanceolate or ovate leaves and rounded bases are
Gomphocarpus cancellatus (Burm.f.) Bruyns (with a predominantly winter-rainfall regime in
the southern and western Cape) and the G. glaucophyllus group (that consists of G. glauco-
phyllus, G. swynnertonii (S.Moore) Goyder & Nicholas, G. munonquensis (S.Moore) Goyder
& Nicholas, G. praticola (S.Moore) Goyder & Nicholas and G. semiamplectens). In all of
these species the base is usually cordate. Gomphocarpus glaucophyllus is the only species
that has a distribution as far south as South Africa, while the other species in this group
have a tropical and subtropical distribution. Furthermore, the follicles of G. glaucophyllus
and allies seem particularly variable with respect to the degree of inflation present, which
are invariably smooth. The fruiting pedicels are contorted in almost all species. The disc
of the seed is generally verruculose, but is smooth and glabrous in G. rivularis Schltr. and
pubescent in the G. glaucophyllus group. Seeds of most species have no obvious marginal
rim surrounding the discoid body of the seed, but in taxa of the G. glaucophyllus group the
margins are convoluted.
The plant illustrated here (Condy 272) was collected from Ezemvelo Nature Reserve,
22 km northeast of Bronkhorstspruit, straddling the border between Gauteng and Mpuma-
langa provinces in South Africa. Here plants are occasionally encountered growing on rocky
outcrops, gentle slopes and plains. The reserve falls within the Rand Highveld and Eastern
Highveld Grassland vegetation types of Mucina and Rutherford (2006).
a b c
d f
g h
FIGURE 2.—Gomphocarpus glaucophyllus: a, bracts with marginal hairs; b, flower viewed from under-
neath showing calyx; c, flower viewed from above, note sinus (arrow) of corona lobes; d, gynoste-
gial column showing inner corona lobe (arrow) below anther wings; e, gynostegial column in side
view showing proximal sides of outer corona lobes overtopping the anthers; f, anthers with anther
appendage (arrow) overtopping the style head; g, pollinarium; h, dorsal view of seed, note white
scar on distal end where coma was detached. Scale bars: a, d, e, h, 1 mm; b, c, 2 mm; f, g, 0.5 mm.
Vouchers: a–g, Bester 11619 (PRE); h, MacDonald 13 (PRE). Photographs: S.P. Bester.
126 Flowering Plants of Africa 65 (2017)
The species was assessed and its Red List status established as least concern (LC) for
South Africa due to the species being widespread, common, and not in danger of extinction
(Von Staden 2012). However, the habitat in some parts of South Africa seems to be under
pressure for mining and agricultural development. Concern has been expressed in a TRAFFIC
(the Wildlife Trade Monitoring Network) report by Rukangira (2001) that Gomphocarpus glau-
cophyllus is harvested and traded in Zimbabwe for its medicinal properties.
The plants have various reported uses. The roots are infused in water which is drunk
for asthma or a decoction of it is made to induce vomiting in babies for the relief of various
illnesses (Gelfand et al. 1985; Neuwinger 2000). On the label of a specimen it is indicated
that the plants were subjected to a feeding test and that the seeds (possibly fruit) espe-
cially were eaten by cattle. In Venda, where the plant is known as gunaperi or gunatela
(Fox & Norwood Young 1982; Kuiper 1555 [PRE]), people cook the young leaves and eat it
as a kind of spinach. The leaves and tubers are sources of toxic cardiac glycosides, which
according to Van Wyk et al. (2002), are the most important of all causes of livestock poison-
ing in South Africa. This is in contradiction to Schultz et al. (2005) who reason that although
all Gomphocarpus species contain cardenolides, they are unpalatable and seldom eaten,
Flowering Plants of Africa 65 (2017) 127
usually only incidentally ingested. Two species of Gomphocarpus have been screened for
anticancer properties (Fouche et al. 2008), but due to the well-known toxicity of cardiac
glycosides (Van Wyk et al. 2000), and having only moderate levels of activity, no further
work was carried out. A wide range of active compounds have been isolated from G. glau-
cophyllus (Sawlewicz et al. 1967), mainly from the dried roots, of which the highest per-
centage were glycosides.
Description (adapted from Nicholas (1999) and Goyder and Nicholas (2001) with
additional measurements from specimens at PRE).—Perennial herb arising from a stout rhi-
zomatous rootstock, 20–30 mm in diameter. Stems 1–32(–35), robust and somewhat fleshy,
unbranched, ascending or erect, 0.3–1.0 m tall, 6–14 mm thick, glabrous and commonly
glaucous. Leaves mostly sessile or rarely very shortly petiolate, 40–152 × 13–64 mm, apex
acute, base cordate with stem-clasping lobes, glabrous, pale grey- or blue-green, generally
glaucous with a waxy bloom on both surfaces, primary and secondary veins prominent,
sometimes pink to reddish. Inflorescence in racemose, extra-axillary, one to many nodding
umbels, 4–15-flowered; peduncles 15–60 mm long, mostly half to two-fifths the length of
the subtending leaf, glabrous and generally glaucous; bracts 5.0–11.5 × 0.5–4.2 mm, fili-
form to linear, margins ciliate or not, apex acute; pedicels 17–35 mm long, glabrous, glau-
cous or not. Calyx lobes 5.4–10.0 × 2.1–4.0 mm, oblong to broadly ovate, apex obtuse
to subacute to acute, minutely ciliate marginally, blade glabrous, reflexed, green and glau-
cous or not. Flowers 13.0–17.3 × 11–22 mm, light or apple green, yellowish green to cream,
purplish. Corolla united at base for 1–2 mm, greenish yellow, sometimes purplish on outer
face, ventrally minutely papillate particularly towards base, glabrous dorsally; lobes strongly
or weakly reflexed, 9–13 × 5–8 mm, elliptic to ovate, apex acute, minutely ciliate on one
margin only. Staminal corona lobes attached 1.1–1.8 mm above the base of the staminal
column and as high, erect 4.0–6.2 × 3.0–6.2 × 0.8–1.6 mm (length × depth × width),
compressed-cucullate, subquadrate, 4.2–5.3 mm broad in side view, ridge from middle of
proximal side extending downwards and most pronounced at base extending horizontally
towards outer margin where it fuse with distal end, truncate at top with apical angles of sides
produced into short acute point (± 0.6 mm long) over top of style-apex, rounded at base,
sinus ± two-fifths of height of lobes, yellowish brown, brown with green tips or maroon;
sinus 4.2–5.7 × 1.1–1.5 × 1.3–1.4 mm. Interstaminal corona lobes broadly deltate, 0.62–
0.65 × 0.73–1.20 mm, pseudo-bifid due to base of anther wings pushing down onto middle
of apex of lobe. Staminal column 5–6 mm tall, 3.18 mm in diam. Androecium: quadrate,
± 2.8 × 1.6–1.7 mm; anther wings 2.0–3.2 × 0.5–2.2 mm, triangular, outer margin straight
or weakly concave at base of staminal column; anther appendages 0.6–1.1 × 0.4–1.1 mm,
broadly ovate, obtuse, inflexed over and fully or partly concealing truncate and involute
style-stigma head. Corpuscle 0.4–0.5 × 0.08–0.17 mm, narrowly ovoid to sub-cylindrical,
black with translucent flanges running up sides from attachment of caudicles, these with
acute apices; caudicles 0.47–0.56 × 0.07–0.15 mm, flattened and ± uniform in width but
differentially thickened – inner margin thickened near corpuscle, outer margin thicker near
pollinium, medifixed to corpuscle and fixed apically to pollinia; pollinia 1.18–1.27 × 0.42–
128 Flowering Plants of Africa 65 (2017)
REFERENCES
AITON, W.T. 1811. A Catalogue of the Plants Cultivated in the Royal Botanic Garden at Kew. Hor-
tus Kewensis (2nd ed.), Vol. 3, London.
ASTLE, W.L. 1968–1969. The vegetation and soils of Chishinga, Luapula Province, Zambia. Kirkia
7(1): 73–102.
BAUMANN, G. 2005. Photographic Guide to Wildflowers of Malawi. Wildlife and Environmental
Society of Malawi, Blantyre.
BINNS, B. 1968. A first checklist of the herbaceous flora of Malawi. The Government Printer, Zanba.
BROWN, N.E. 1902. Asclepias. In W.T. Thiselton-Dyer (ed.), Flora of tropical Africa 4(1): 313–353.
Lovell Reeve & Co., London.
BROWN, N.E. 1907–1908. Asclepias. In W.T. Thiselton-Dyer (ed.), Flora capensis 4(1): 663–714.
Reeve, London.
BROWN, R. 1810. On the Asclepiadeae, a natural order of plants separated from the Apocineae
of Jussieu. Memoirs of the Wernerian Natural History Society 1: 12–78.
BULLOCK, A.A. 1952. Notes on African Asclepiadaceae. I. Kew Bulletin 7: 405–426.
BURMAN, N.L. 1768. Florae capensis prodromus. Flora Indica: 1–32. Cornelius Haak, Leiden;
Johannes Schreuder, Amsterdam.
DECAISNE, J. 1838. Etudes sur quelques genres et especes de la famille des Asclepiadees. Annales
des Sciences Naturelles, Botanique, series 2,9: 257–278, 321–348.
DECAISNE, J. 1844. Asclepiadeae. In A. de Candolle (ed.), Prodromus systematis naturalis regni
vegetabilis 8: 490–665. Fortin Masson, Paris.
EL-SEEDI, H.R., BURMAN, R., MANSOUR, A., TURKI, Z., BOULOS, L., GULLBO, J. & GÖRANS-
SON, U. 2013. The traditional medical uses and cytotoxic activities of sixty-one Egyptian
plants: discovery of an active cardiac glycosides from Urginea maritima. Journal of Ethno
pharmacology 145: 746–757.
FISHBEIN, M., CHUBA, D., ELLISON, C., MASON-GAMER, R.J. & LYNCH, S.P. 2011. Phyloge-
netic relationships of Asclepias (Apocynaceae) inferred from non-coding chloroplast DNA
sequences. Systematic Botany 36(4): 1008–1023.
FORSSKÅL, P. 1775. Asclepias. In Flora Aegyptiaco-Arabica: 49–51. Møller, Copenhagen.
FORSTER, P.I. 1994. Diurnal insects associated with the flowers of Gomphocarpus physocarpus
E.Mey. (Asclepiadaceae), an introduced weed in Australia. Biotropica 26(2): 214–217.
FOUCHE, G., CRAGG, G.M., PILLAY, P., KOLESNIKOVA, N., MAHARAJ, V.J. & SENABE, J. 2008.
In vitro anticancer screening of South African plants. Journal of Ethnopharmacology 119(3):
455–461.
FOX, F.W. & NORWOOD YOUNG, M.E. 1982. Food from the Veld: Edible wild plants of Southern
Africa. Delta Books, Johannesburg.
GELFAND, M., MAVI, S., DRUMMOND, R.B. & NDEMERA, B. 1985. The traditional medicinal
practitioner in Zimbabwe. Mambo Press, Gweru.
GOYDER, D.J. 1995. Notes on the African genus Glossostelma (Asclepiadaceae). Kew Bulletin 50:
527–555.
Flowering Plants of Africa 65 (2017) 129
3
2
Ipomoea bolusiana Schinz, Verhandlungen des Botanischen Vereins der Provinz Branden-
burg, Berlin 30: 271 (1888); Hallier f.: 147 (1893); Hallier f.: 53 (1899); Baker & Rendle: 175
(1905–1906); Meeuse: 758 (1957); Roessler: 13 (1967); Adams: t. 122 (1976); Gonçalves: 100, t.
24 (1987); Van Wyk & Malan: 200 (1988); Gonçalves: 106, t. 23 (1992); Lejoly & Lisowski: 106,
t. 15 (1992); Meeuse & Welman: 48 (1996); Fabian & Germishuizen: 334, t. 159c (1997); Roodt:
63 (1998); Meeuse & Welman: 107 (2000). I. simplex Hook.: t. 4206 (1846) nom illegit. I. angusti-
secta Engl.: 245 (1888). I. mesenterioides Hallier f.: 544 (1898). I. bolusiana Schinz var. abbreviata
Hallier f.: 54 (1899). I. bolusiana Schinz var. elongata Hallier f.: 54 (1899). I. praetermissa Rendle:
56 (1901). I. simplex Thunb. var. obtusisepala Rendle: 174 (1905–1906). I. bolusiana Schinz var.
pinnatipartita Verdc.: 118 (1967).
The genus Ipomoea contains about 650 species distributed worldwide in the tropical
and warm temperate countries. Fifty-seven species occur in southern Africa, five of them
are introduced and naturalised. The genus is paraphyletic and can be distinguished from
Convolvulus by its spiny pollen and its stigma which is not linearly divided. It contains annu-
als as well as deciduous or evergreen perennials, climbing or prostrate herbs or shrubs or
even small trees; some species have tubers while a few are caudiciform succulents. Sev-
eral species are ornamental, some are cultivated for their edible roots, flowers, shoots and
leaves, while the seeds or roots of other species provide drugs or medicine.
Mabberley (2008) listed a number of useful species that are either indigenous to Africa
or cultivated on this continent. The tuberous Ipomoea batatas (L.) Lam. (sweet potato), a
hexaploid cultigen with many cultivars, probably originally from Central America, is one
of the world’s most important starch foods and a staple crop in many tropical and warm
areas. Some forms of I. aquatica Forssk. (water spinach), an Old World species, also occur-
ring naturally in Namibia, Botswana and KwaZulu-Natal, are cultivated for their edible
shoots in China and other countries and was introduced into the New World. Ipomoea
obscura (L.) Ker Gawl. from the Old World tropics, also southern Africa, is a pot herb in Sri
Lanka. A root infusion of I. spathulata Hallier f. from East Africa is used against eye disease.
Ipomoea pes-caprae (L.) R.Br. from tropical and subtropical beaches, is a sand binder; the
subsp. brasiliensis (L.) Ooststr., grows in KwaZulu-Natal and the Eastern Cape, South Africa.
Hallucinogens are obtained from the seeds of some species of Ipomoea; the toxin is an
indole alkaloid (Van Wyk et al. 2002). Several indole alkaloids are known from Ipomoea,
such as ergine which is present in many species; large amounts can be fatal. Mabberley
(2008) listed the ornamental I. tricolor Cav. from Mexico and Central America, where the
effective hallucinogenic agent is ergoline. Apparently no South African species have so far
been investigated for these alkaloids.
PLATE 2337.—1, flowering plant, × 1; 2, fruit, × 1; 3, fruit and seed, × 1. Voucher specimens: 1, Condy 279;
2, Bremekamp & Schweickerdt 29552; 3, R. Leendertz 6620 all in National Herbarium, Pretoria. Artist:
Gillian Condy.
134 Flowering Plants of Africa 65 (2017)
With its usually prostrate stems and rather long and very narrow leaves or leaf lobes,
Ipomoea bolusiana, one of the so-called ‘wild morning glories’, is almost inconspicuous
in its natural environment when it is not flowering. However, its fairly large, usually bright
pink, showy flowers attract immediate attention. This species is indigenous to Angola, the
Democratic Republic of the Congo, Zambia, Zimbabwe, Mozambique, Namibia, Botswana,
Swaziland and summer rainfall regions of South Africa (Limpopo, North-West, Gauteng,
Mpumalanga, Free State and KwaZulu-Natal provinces, down to about 30 degrees south in
the latter province) (Figure 1). Gonçalves (1987) noted that it is also found in Tanzania, but
it was not included in the Flora of Tropical East Africa (Verdcourt 1963). Its occurrence in
Madagascar (Meeuse 1957) could not be verified either.
Ipomoea bolusiana has a very wide ecological range and grows in grassland, various
kinds of savanna, woodland or bushveld and shrub- or scrubland. The soil can be deep or
shallow, poorly drained to well-drained sand of various colours, loam or clay which can be
stony, rocky or gravelly. The substrate can be calcrete, dolomite, granite, ironstone, quartz-
ite or sandstone. It can grow in rocky areas, as well as on dune ridges and in pans and
depressions, also on the margins of wetlands. Because of its caudex (succulent and tuber-
ous rootstock), I. bolusiana is very drought resistant; it can survive in disturbed areas like
road- and railway sides, overgrazed and burned land, even on denuded, dry, hard, com-
pacted soil. It can grow in shade, partial shade or full sun, on level to moderate slopes
of all aspects. It has been recorded as solitary, rare, occasional to frequent or common.
The flowering time is from August to April, mostly from October to March. The fruiting
time is mostly from November to April. According to the information on specimen labels
in the National Herbarium, Pretoria (PRE), this species can grow from about 45 to 1 525 m
altitude in southern Africa. Gonçalves (1987) reported that in the Flora zambesiaca area,
I. bolusiana grows in open woodland and savanna, grassland with scattered shrubs, in rocky
and sandy soils at 0 to 1 465 m. According to Raimondo et al. (2009) the Red List status of
I. bolusiana in South Africa is Least Concern.
Many succulent collectors include caudiciform plants in their collections. Rowley (1987)
wrote of the Convolvulaceae: ‘The perennial herbs regenerate from an underground reser-
voir which in a few African species is a solid, smooth-skinned black or brown caudex suited
to grace the glasshouse of any devotee of caudiciforms. Unlike so many caudiciforms, Ipo-
moeas have large, showy bisexual flowers, and because these are borne freely low down
and not just at the top of metres of vine, the plants are suitable for a small glasshouse or
show bench. The caudex, which is sensitive to rot, should be planted above ground on a
grit base and allowed to dry off during the dormant season when the branches die away.’
Among the southern African species credited with having a caudex are Ipomoea bolusiana,
I. simplex Thunb., I. transvaalensis A.Meeuse and I. welwitschii Vatke ex Hallier f. Some rep-
resentatives of the closely related genus Merremia with white or yellow blooms may also
qualify, such as the Namibian endemic M. bipinnatipartita (Engl.) Hallier f. Rowley (1976)
stated, ‘… many South African Convolvulaceae have a massive perennial caudex, and the
fact that it grows underground is no deterrent to the enthusiast who happily pots it above the
soil so that he can admire its distinctive bark and noble contours. And the plants seem none
the worse for such unnatural treatment.’ He wrote the following notes on the cultivation of
I. bolusiana in the northern hemisphere: ‘It offers no special problems in cultivation. I keep it
in the warmest section of the greenhouse and withhold water until the first signs of growth in
Flowering Plants of Africa 65 (2017) 135
The species name bolusiana commemorates the South African botanist and business-
man Harry Bolus (1834–1911) who undertook collecting expeditions to various parts of
South Africa. He described many new species and also produced his own illustrations. His
main interests were the Orchidaceae and Ericaceae; he co-authored the Ericaceae for Flora
capensis. The Bolus Herbarium at the University of Cape Town was named after him after
he had donated his collections and library to that university (Gunn & Codd 1981).
According to Austin & Huáman (1996), Ipomoea bolusiana belongs to section Erpipo-
moea Choisy (section Leiocalyx Hallier f. sensu stricto) which falls under subgenus Eriosper-
mum (Hallier f.) Verdc. ex D.F.Austin. This common and variable species is closely related
to and can be confused with two other species in that section, namely I. simplex and
I. welwitschii. Ipomoea bolusiana differs from the former by its large pink to purple corolla
and from the latter by its leaf morphology and glabrous vegetative parts. Ipomoea simplex
has a white corolla and is endemic to the central and eastern parts of southern Africa,
136 Flowering Plants of Africa 65 (2017)
including Lesotho. The rare and variable I. welwitschii occurs in tropical Africa down to
northern Namibia and Botswana. It is possible that these three species form one large,
variable aggregate species.
The specimen used as voucher for the accompanying plate was cultivated at the Ran-
dom Harvest Nursery in Muldersdrift, Gauteng, from the collection of indigenous plants
built up by the late Charles Craib. It was originally collected in the Rustenburg/Brits area in
the North-West Province.
REFERENCES
ADAMS, J. 1976. Wild flowers of the northern Cape. Department of Nature and Environmental
Conservation, Cape Town.
AUSTIN, D.F. & HUÁMAN, Z. 1996. A synopsis of Ipomoea (Convolvulaceae) in the Americas.
Taxon 45: 3–38.
BAKER, J.G. & RENDLE, A.B. 1905–1906. Convolvulaceae. In W.T. Thiselton-Dyer, Flora of tropical
Africa 4,2: 62–206. Reeve, London.
ENGLER, H.G.A. 1888. Plantae marlothianae. Ein beitrag zur Kenntnis der Flora Südafrikas. Bota-
nische Jahrbücher 10: 242–285.
FABIAN, A. & GERMISHUIZEN, G. 1997. Wild flowers of northern South Africa. Fernwood Press,
Vlaeberg.
GONÇALVES, M.L. 1987. 117, 118. Convolvulaceae–Cuscutaceae. In E. Launert (ed.), Flora zam-
besiaca, Vol. 8,1.
GONÇALVES, M.L. 1992. 113. Convolvulaceae. Flora de Moçambique. Instituto de Investigaçào
Cientifica Tropical. Centro de Botânica, Lisbon.
GUNN, M. & CODD, L.E. 1981. Botanical exploration of southern Africa. A.A. Balkema, Cape Town.
HALLIER, H. 1893. Convolvulaceae africanae. Botanische Jahrbücher 18: 81–160.
HALLIER, H. 1898. Beiträge zur Kenntnis der Afrikanischen Flora. Convolvulaceae. Bulletin de
l’Herbier Boissier 6: 529–548.
Flowering Plants of Africa 65 (2017) 137
Ipomoea cairica (L.) Sweet, Hortus britannicus, 1st edn: 287 (1826); Hallier f.: 148 (1893);
Hutchinson & Dalziel: 216 (1931); Brenan: 8 (1954); Meeuse: 761 (1957); Verdcourt: 125 (1963);
Heine: 351 (1963); Gonçalves: 105, t. 26 (1987); Lejoly & Lisowski: 115, t. 16 (1992); Meeuse
& Welman: 110 (2000). Convolvulus cairicus L.: 922 (1759). Ipomoea palmata Forssk.: 43 (1775).
Convolvulus tuberculatus Desr.: 545 (1791). I. senegalensis Lam.: 464 (1793). I. vesiculosa Beauv.:
73, t. 106 (1819). I. tuberculata (Desr.) Roem. & Schultes: 208 (1819). I. mendesii Welw.: 584
(1859). Iconotype: illustration of Convolvulus aegyptius in Vesling: 75, fig. s.n. (1638).
Ipomoea L. is a very large genus in the family Convolvulaceae (morning glory family)
and has about 650 species distributed worldwide; they are most common in the tropics of
both hemispheres. Ipomoea batatas (L.) Lam., or sweet potato, a food plant with tuberous
roots originally possibly from Central America, is the most important economic species in
the Convolvulaceae. There are 16 genera in this family in southern Africa, with Ipomoea the
biggest genus with 57 representatives. Of these, only five taxa were introduced, and have
escaped from cultivation and became naturalised; they are I. alba L. (moonflower, tropi-
cal Americas), I. carnea Jacq. subsp. fistulosa (Mart. ex Choisy) D.F.Austin (morning glory
bush, tropical Americas), I. indica (Burm.f.) Merr. (pantropical), I. purpurea (L.) Roth (com-
mon morning glory, New World) and the rarely collected I. nil (L.) Roth (Japanese morning
glory, tropical Americas).
Glen (2002) listed 32 species of Ipomoea that are cultivated in especially the warmer
parts of the southern African region and may occasionally occur as culture relicts or gar-
den escapes. Three popular species are often cultivated: I. arborescens (Humb. & Bonpl.)
G.Don (morning glory tree, West Indies), I. quamoclit L. (Cupid flower, cardinal vine, New
World) and I. tricolor Cav. (Japanese morning glory, New World). Verdcourt (1963) listed
about 10 of the more important purely ornamental species that are cultivated in East Africa
of which most are also cultivated in southern Africa.
Henderson (2001) treated the three species of Ipomoea (I. alba, I. indica and I. pur-
purea) that are declared weeds or invaders according to the Conservation of Agricultural
Resources Act (Act No. 43 of the Republic of South Africa 1983, amended in 2001) and
one species (I. carnea subsp. fistulosa) that may soon be declared. Ipomoea alba, an annual
or perennial climber or prostrate creeper that is cultivated for ornamental purposes in the
warmer parts of the world, is a declared weed in KwaZulu-Natal, Mpumalanga and Lim-
popo provinces and a declared invader elsewhere in South Africa. Its invasive status is listed
as ‘transformer’ and it may no longer be traded. Ipomoea indica (perennial) and I. purpurea
(annual) are herbaceous twiners and are both cultivated as ornamentals. Ipomoea indica is
a declared weed and invader in KwaZulu-Natal, Limpopo and Mpumalanga provinces, with
PLATE 2338.—1, flowering branch, × 1; 2, stamens and pistil, × 2; 3, immature fruits, × 1. Voucher speci-
mens: 1, 2, Herman 1594 (flowering) and 3, Herman 1595 (fruiting), all in National Herbarium, Pretoria.
Artist: Gillian Condy.
1
2 3
Verdcourt (1963) described the habitat in tropical East Africa as ‘forest clearings, lake
shores, swampy grassland, hedges, waste and cultivated ground etc.’ at 750 to 1 900 m alti-
tude. Gonçalves (1987) reported that in the Flora zambesiaca area, Ipomoea cairica grows in
‘rainforests, bushland, swampy grassland, river edges, lake shores, roadsides at 0 to 1 676 m
altitude’.
According to Austin & Huáman (1996) this species belongs to section Erpipomoea
Choisy (section Leiocalyx Hallier f. sensu stricto) which falls under subgenus Eriospermum
142 Flowering Plants of Africa 65 (2017)
(Hallier f.) Verdc. ex D.F.Austin. It is closely related to and can be confused with three other
species in that section, namely Ipomoea hochstetteri House (Namibia, Botswana, Swazi-
land, northern South Africa and also tropical Africa), I. tenuipes Verdc. (northern Namibia,
northern Botswana, and Limpopo and Mpumalanga provinces in South Africa, and also
tropical Africa and India) and I. tuberculata Ker Gawl. (northern Namibia, northern Bot-
swana, tropical Africa and also India and Sri Lanka).
These four species all have pseudostipulate and distinctly palmately or pedately 5–9-fid
leaves that are usually quite glabrous. They can be distinguished as follows (see Meeuse &
Welman 2000):
1a Calyx 8–10 mm long; outer sepals saccate at base; corolla yellow with mauve tube;
leaves usually biternately pedate, often with 9 segments . . . . . . . . . . . . . . . . . I. tuberculata
1b Calyx up to ± 7 mm long; outer sepals not saccate; corolla mauve, purple, red or
white; leaves palmately partite, usually with 5 segments. . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2a Corolla 30–60 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. cairica
2b Corolla up to ± 25 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a Peduncles very slender, filiform; corolla often shorter than 16 mm, purplish mauve
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. tenuipes
3b Peduncles rather stout, not filiform; corolla 15–25 mm long, purplish mauve or white
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. hochstetteri
The crushed leaves of Ipomoea cairica are non-aromatic and the cut twigs exude no
sap. Pinheiro & Schlindwein (1998) found that the flowers of this species show a nectar
chamber between the nectariferous disc and the insertion of the filaments at the corolla,
which prevents free access of visitors to the nectar. Nectar can only be reached through
five small openings between the bases of the filaments, which are covered by hairs. Due to
the presence of the nectar chamber, I. cairica liberates nectar only for those bees that have
glossae long enough to reach the nectary.
Watt & Breyer-Brandwijk (1962) reported, inter alia, that the Zulu people drink a mix-
ture of the crushed leaf of Ipomoea cairica for body rashes, especially if accompanied by
fever. The leaf and root contain traces of hydrocyanic acid. The seed is purgative, an action
which is due to muricatin A. The plant has considerable antibiotic action but not in every
Flowering Plants of Africa 65 (2017) 143
case. Gerstner 5112 in PRE recorded in 1944 that this ‘common creeper’ is planted all over
Zululand and sold as a purgative, known as iJalambu. A.M. du Toit s.n. in PRE reported
from Limpopo Province (formerly northern Transvaal) in November 1926 that this plant is
‘extremely palatable and eaten by every class of stock’. However, Watt & Breyer-Brandwijk
(1962) reported that in Australia this plant has been suspected of poisoning horses.
The species name cairica refers to Cairo in Egypt where the species was first collected.
Verdcourt (1963) reported that the taxon discussed here is the rather common and wide-
spread var. cairica, which also occurs in India. The var. indica Hallier f. [Ipomoea palmata
Forssk. var. indica (Hallier f.) Rendle] occurs at about 750 m altitude in thickets near water-
holes in northern Kenya, Somalia and India (Verdcourt 1963). This plant is rather intermedi-
ate between I. cairica var. cairica and I. hochstetteri. The corolla is 23–30 mm long. How-
ever, the present view is that I. cairica has no varieties. According to Mabberley (2008),
I. indica (Burm.) Merr. from tropical South America is a different species that is cultivated
worldwide in the tropics.
REFERENCES
AUSTIN, D.F. & HUÁMAN, Z. 1996. A synopsis of Ipomoea (Convolvulaceae) in the Americas.
Taxon 45: 3–38.
BRENAN, J.P.M. 1954. Plants collected by the Vernay Nyasaland Expedition of 1946 (5). Memoirs
of the New York Botanical Garden, Vol. 9,1. Bronx Park.
CHIARINI, F.E. 2000. Números cromosómicos en dos especies de Ipomoea (Convolvulaceae)
argentinas. Kurtziana 28,2: 309–311.
DESROUSSEAUX, L.A.J. 1791. In J.B. de Lamarck, Encyclopédie méthodique (Dictionnaire encyclo-
pédique de botanique). Vol. 3. Panckoucke, Paris.
144 Flowering Plants of Africa 65 (2017)
Rotheca myricoides (Hochst.) Steane & Mabb. sens. lat., Novon 8: 205 (1998); Fernandes
& Verdcourt: 150 (2000); Fernandes: 136 (2005). Spironema myricoides Hochst. no. 330 (1840);
Hochst. & Steud. (1838). Cyclonema myricoides (Hochst.) Hochst.: 226 (1842); Richard: 171
(1851); Hooker: t. 5838 (1870). Clerodendrum myricoides (Hochst.) Vatke: 535 (1882) [as Clero-
dendron myricoides R.Br.]; Gürke in Engler: 341 (1895) [as Clerodendron myricoides R.Br.]; Bri-
quet: 176 (1895) [as Clerodendron myricoides (Hochst.) Gürke]; Baker: 310 (1900) [as Cleroden-
dron myricoides R.Br.]; Medley Wood: t. 282 (1902) [as Clerodendron myricoides R.Br.]; Pearson:
223 (1912) [as Clerodendron myricoides R.Br.]; Thomas: 86 (1936); Moldenke: 329 (1987) [as
Clerodendrum myricoides (Hochst.) R.Br.]; Verdcourt: 130 (1992); Retief & Herman: 641 (1997);
Fernandes: 47 (1998). Clerodendrum myricoides R.Br. in Salt: lxv (1814), nom. nud.
The genus Rotheca Raf. comprises about 30 species in sub-Saharan Africa (Fernandes
2005) with one species (R. serrata (L.) Steane & Mabb.) native to eastern India, Sri Lanka
and Malaysia, and three species native to Madagascar (R. microphylla (Blume) Callm. &
Phillipson, R. mirabilis (Baker) Callm. & Phillipson and R. nudiflora (Moldenke) Callm. &
Phillipson) (Madagascar Catalogue 2013). About nine species are known from southern
Africa (Namibia, Botswana, Swaziland, Lesotho and South Africa). Members of the genus
Rotheca were previously included under Clerodendrum L., subgenus Cyclonema (Hochst.)
B.Thomas (1936). The genus name Rotheca was reinstated by Steane & Mabberley (1998)
based on differences in morphology and DNA. Rotheca can be distinguished from Clero-
dendrum by the shape of the flower bud: in Rotheca the flower bud is markedly asym-
metrical and the corolla expands abruptly on the lower side so that it looks something
like a golf club; the anterior corolla lobe is usually much larger than the other four lobes.
In Clerodendrum the flower bud is usually symmetrical or, if asymmetrical, it expands
abruptly on the upper side; the anterior corolla lobe is only slightly (if at all) larger than the
other four lobes (Steane & Mabberley 1998). The genera Clerodendrum and Rotheca were
previously treated in the family Verbenaceae, but were transferred to the family Lamiaceae
by Cantino et al. (1992). Subfamilies Ajugoideae and Viticoideae have fruits which either
do not split into 4 distinct mericarps, or rarely do split late in development to produce 4
nutlets (Fernandes 2005).
The genus name Clerodendrum was used by Linnaeus in his Species plantarum (1753)
and Genera plantarum (1754). The spelling was changed to Clerodendron as early as 1763
by Adanson (Moldenke 1985). Since then, there seems to have been considerable confu-
sion about the correct spelling of the name and both names were used randomly by various
authors. Moldenke (1985) pointed out that, according to the rules of botanical nomencla-
ture, there was no reason to change the name and it should remain Clerodendrum L. Rob-
ert Brown (R.Br.) is often given as the author for the species name myricoides, but according
PLATE 2339.—1, flowering branch, × 1; 2, flower, × 2; 3, stamen with ciliate filament, × 2; 4, leaf, × 1.
Voucher specimen: Burrows 6531 in Buffelskloof Nature Reserve & Herbarium, Lydenburg. Artist: Gil-
lian Condy.
148 Flowering Plants of Africa 65 (2017)
to Verdcourt (1992), Brown only mentioned the name in one of his publications without a
description or any references.
Rotheca myricoides is a highly variable species. Thomas (1936), Verdcourt (1992), Fer-
nandes & Verdcourt (2000) and Fernandes (2005) distinguished several subspecies, varie-
ties and formas based on, inter alia, leaf shape, leaf margin and pubescence of the leaves.
Many of the subspecies and varieties are confined to specific small areas. In her account
of Rotheca for Flora zambesiaca, Fernandes (2005) did not mention any occurrence of
R. myricoides representatives in South Africa as she did for instance under R. hirsuta. Nei-
ther did Verdcourt (1992) in his treatment of Clerodendrum for Flora of Tropical East Africa,
as he for instance did for C. glabrum and C. ternatum. The South African material could
perhaps be classified as subsp. myricoides var. myricoides forma myricoides and subsp. myri-
coides var. discolor (Klotzsch) Verdc. forma discolor taxa, but no attempt was made to try
and classify southern African material to subspecific level, as the characters tend to merge
into each other. Fernandes (2005) recognised a separate species, R. reflexa (H.Pearson)
R.Fern., which is confined to the Flora zambesiaca area that approaches some varieties of R.
myricoides. Some specimens in the National Herbarium, Pretoria (PRE) fit the description of
R. reflexa and these two species could probably prove to be conspecific.
The variation in the leaf shape of South African Rotheca myricoides material housed in
PRE is illustrated in Figure 1: the leaves are mostly ovate to rhomboid in outline with the
margins dentate in the upper half to very deeply incised to almost lobed, but can also be
elliptic, or in a few cases, obovate in outline and the entire margin dentate. The indumentum
of the leaves varies as much: usually sparsely hairy on upper surface and sparsely to densely
hairy on lower surface, sometimes almost velvety. In some cases the leaf surfaces seem to
b c d
FIGURE 1.—Line drawings to illustrate different leaf shapes and margins of Rotheca myricoides leaves: a,
ovate with dentate margin (voucher: Joffe 198, PRE); b, rhomboid with coarsely dentate margin
(voucher: Smuts PRE39965, PRE); c, rhomboid with deeply lobed margin (voucher: Van Vuuren 1824,
PRE); d, broadly elliptic, dentate (voucher: Galpin 601, PRE). All drawings × 1. Artist: Gillian Condy.
Flowering Plants of Africa 65 (2017) 149
a b
FIGURE 2.—Scanning electron microscope (SEM) images of peltate hairs: a, upper leaf surface, scale
bar = 100 µm; b, inside of calyx lobes, scale bar = 10 µm. Voucher: Van Wyk 4731, PRE. Micrographs
taken with SEM at Electron Microscopy Unit, University of Pretoria, Pretoria.
be glabrous, but even then some hairs are present, especially along the margins. Most of the
Namibian specimens housed in PRE have narrowly elliptic to elliptic to sometimes obovate,
usually fairly small leaves with the margins entire or shallowly dentate in the upper half.
The leaves of Rotheca myricoides are often described as glandular punctate beneath
(Verdcourt 1992; Fernandes 2005). These ‘glands’ are peltate hairs or peltate glands (Figure
2A). They are not always easy to observe when the leaf surface is densely hairy. Similar
peltate hairs were described for Clerodendrum triphyllum (=R. hirsuta) and C. louwalbertsii
(=R. louwalbertsii) (Herman 1998). They are possibly responsible for the unpleasant scent
when the leaves are crushed. Similar peltate glands were observed on the inside of the
calyx lobes of R. myricoides (Figure 2B) as well as on the skin of the fruit. These glands on
the inside of the calyx lobes were described for Clerodendron myricoides by Medley Wood
(1902), Pearson (1912) and Moldenke (1987), but none of the more recent publications
mention them (Verdcourt 1992; Fernandes & Verdcourt 2000; Fernandes 2005). Peltate
hairs or glands are often present in representatives of the family Lamiaceae (Ascensão et al.
1995; Shan-Shan Huang et al. 2008; Ping Jia et al. 2013; Rusydi et al. 2013).
There seems to be as much variation in the flower colour as in the leaf shape. The
colour of the flowers, as noted on herbarium specimen labels, varies from green and blue
or mauve, white and blue, or blue to mauve. The flower colours of most of the Namib-
ian material housed in PRE are noted on the labels as whitish blue, blue or bluish, purple,
four petals whitish and one blue, or green and blue. Most of the material was identified
by Moldenke as Clerodendrum dekindtii var. dinteri or quoted by Friedrich-Holzhammer
(1967) as C. dekindtii. Fernandes (2005) sank C. dekindtii var. dinteri sensu Moldenke under
R. myricoides subsp. myricoides var. moldenkei. She sank C. dekindtii sensu Moldenke
under R. myricoides subsp. myricoides var. discolor forma discolor. However, she recognised
a separate taxon, subsp. namibiensis, under R. myricoides mainly based on the arrangement
of the flowers in axillary cymes and the flowers being white or with four petals whitish
green and the median petal blue, not all the petals violet.
Rotheca myricoides occurs in the South African provinces of Limpopo, Gauteng, Mpu-
malanga, KwaZulu-Natal (Figure 3) and possibly also North-West. In neighbouring coun-
150 Flowering Plants of Africa 65 (2017)
Rotheca myricoides has long been used in traditional medicine. According to Watt &
Breyer-Brandwijk (1962) and Hutchings (1996) European and African people at Ixopo used
a teaspoonful of the powdered bark for snake bite; the Masai used the root bark as a rem-
edy for East Coast fever in cattle, and for diarrhoea in their calves; the Haya and Shambala
used the plant for dysmenorrhoea, as well as a remedy for cough, furunculosis and swellings
of the body associated with debility; and the root is an East African remedy for the enlarge-
ment of the spleen. The root is also used for pains in the chest, colds, bleeding of the gums,
indigestion and in Zimbabwe for headaches, leaf decoctions for bathing patients with con-
vulsions, while the leaves are used in West Africa for analgesic and antipyretic purposes
(Hutchings 1996).
at each furcation, bracts at first furcation linear, linear-elliptic, obovate to almost leaf-like,
up to 6 mm long; bracts at secondary furcations linear, up to 3 mm long, peduncles and
bracts hairy; pedicles up to 8 mm long, hairy. Calyx with tube cup-shaped, green but some-
times purplish, up to 4 mm long, hairy; calyx lobes 5, semi-circular, up to 4 × 4 mm, apex
roundish to obtuse, hairy on outside, peltate hairs on inside, persistent in fruit. Corolla
zygomorph, tube 7–10 mm long, with 5 free petals, 4 lateral petals (two on each side)
5–12(15) × 4–7 mm, oblong-obovate, often ciliate, greenish, whitish, pale blue or purple,
lower petal pyriform/ob-spathulate, 8–12 × 3.5–10.0 mm, dark blue or purple. Stamens 4,
inserted in corolla throat, opposite lower corolla lip and curved towards it, long exserted,
filaments 23–32 mm long, lower 5–7 mm ciliate-hairy, purple, anthers 2 mm long. Style
long exserted, up to 36 mm long, curved towards lower corolla lip, with 2 uneven stigmatic
lobes, upper lobe 1–3 mm long, lower lobe 1.5–4.0 mm long. Ovary superior, ± globose,
± 2 × 1.5–2.5 mm. Fruit 1–4-lobed drupe, 7–10 × 8–12 mm, purple or black at maturity,
some peltate hairs present on exocarp. Seeds 1–4 pyrenes, reniform. Flowering time: August
to June. Plate 2339.
REFERENCES
ASCENSÃO, L., MARQUES, N. & PAOS, M.S. 1995. Glandular trichomes on vegetative and repro-
ductive organs of Leonotis leonurus (Lamiaceae). Annals of Botany 75: 619–626.
BAKER, J.G. 1900. Verbenaceae. Flora of tropical Africa 5: 273–502.
BRIQUET, J. 1895. Verbenaceae. In A. Engler & K. Prantl (eds), Die Natürlichen Pflanzenfamilien
IV, 3a: 132–182.
BROWN, R. 1814. List of new and rare plants collected in Abyssinia. In H. Salt (ed.), A voyage to
Abyssinia. Append. IV: lxv. F.C. & J. Rivington, London.
CANTINO, P.D., HARLEY, R.M. & WAGSTAFF, S.J. 1992. Genera of Labiatae: status and clas-
sification. In R.M. Harley & T. Reynolds (eds), Advances in Labiate Science: 511–522. Royal
Botanic Gardens, Kew.
FERNANDES, R. 1998. Notes sur les Verbenaceae. XI–Remarques sur quelgues taxa africains du
genre Clerodendrum L. Memórias da Sociedade Broteriana 30: 5–109.
FERNANDES, R. 2005. 129. Lamiaceae. Flora zambesiaca 8,7: 61–153.
FERNANDES, R.B. & VERDCOURT, B. 2000. Rotheca (Labiatae) revived – more new combina-
tions. Kew Bulletin 55: 147–154.
FODEN, W. & POTTER, L. 2005. Rotheca myricoides (Hochst.) Steane & Mabb. National Assess-
ment: Red List of South African Plants version 2015.1. Accessed on 10 June 2016.
FRIEDRICH-HOLZHAMMER, M. 1967. Verbenaceae. Prodromus einer Flora von Südwestafrika
122: 1–10.
GÜRKE, M. 1895. Family Verbenaceae. In A. Engler (ed.), Die Pflanzenwelt Ost-Afrikas und der
Nachbargebiete, Theil C: 337–342.
HERMAN, P.P.J. 1998. The leaf anatomy of two Clerodendrum species (Verbenaceae). South Afri-
can Journal of Botany 64,4: 246–249.
HOCHSTETTER, C.F. 1840. In schaed. Schimperi iter Abyssinicum, sect. 1 & sect. 3. [Printed notes,
distributed to herbaria with Schimper’s specimens, comprising descriptions and type cita-
tions of new taxa.]
HOCHSTETTER, C.F. 1842. Verbenaceae. Flora 25: 225–228.
HOCHSTETTER, C.F. & STEUDEL, E.G. von. 1838. Schimper’s Abyssinische Reise.
HOOKER, J.D. 1870. Cyclonema myricoides. Curtis’s botanical magazine 26: t. 5838.
HUTCHINGS, A. 1996. Zulu medicinal plants: an inventory. University of Natal Press, Pieterma
ritzburg.
152 Flowering Plants of Africa 65 (2017)
4 5 6 1
Ruellia kaokoensis Van Jaarsv. sp. nov. differs from R. bignoniiflora S.Moore by its much small-
er herbaceous habit, glaucous leaves and stems, much shorter diurnal trumpet-shaped maroon
flowers of 60–70 mm long.
Acanthaceae, to which this species belongs, have a cosmopolitan distribution but are
especially abundant in the tropics of Africa, Brazil, Central America and Indo-Malaysia. It
consists of about 2 500 species in 250 genera (Heywood 2007), most of which are shrubs
or herbaceous perennials, with some tending to scramble or climb. Many of the species
have ornamental properties and several have been taken up in the horticultural industry.
Well-known southern African members that are popular in gardens include Anisotes roger-
sii S.Moore, Asystasia gangetica (L.) T.Anderson, Duvernoia adhatodoides E.Mey. ex Nees,
Hypoestes aristata (Vahl) Sol. ex Roem. & Schult., Metarungia longistrobus (C.B.Clarke)
Baden, Petalidium coccineum S.Moore, Salpinctium natalense (C.B.Clarke) T.J.Edwards, vari-
ous Barleria L., Ruellia L. and Crossandra Salisb. species, and the black-eyed Susan (Thun-
bergia alata Bojer ex Sims). Brillantaisia P.Beauv. from tropical Africa, is sometimes cultivated
in suitable areas.
The leaves of Acanthus mollis L. (not indigenous to southern Africa) have been used
as an emblem of the Corinthian capitals in Greek temples. This species is widely grown in
Mediterranean type gardens.
The Acanthaceae family is distinctive and members are characterised by their simple,
ex-stipulate, decussate leaves. Flowers are bisexual, irregular, or two-lipped, arranged in
cymes or racemes with colourful bracts and bracteoles, often subtending the flowers. Petals
and sepals amount to 4–5 each, and 2 or 4 stamens are attached to the petals. The style
has two stigmas dissimilar in length. The fruit is a capsule and the seed borne on hook-
like outgrowths and often released explosively, particularly after rain. Seeds are without
endosperm and with large embryos. Some arid-adapted species are covered in mucous
hairs which, upon wetting, aid in germination (Heywood 2007).
The Acanthaceae are divided into three subfamilies; Ruellia belongs to the subfamily
Acanthoideae, tribe Ruellieae, subtribe Ruelliinae (Balkwill & Welman 2000). Ruellia is dis-
tinguished from close relatives (Crabbea Harv. and Ruelliopsis C.B.Clarke) by its muticous
PLATE 2340.—1, plant in flower, × 1; 2, internal structure of flower, × 1; 3, cross section of ovary, × 1.5; 4,
gynoecium, × 1; 5, gynoecium including sepals, × 1; 6, glandular-puberulous calyx, × 1.5. Voucher
specimen: Van Jaarsveld 16480 in Windhoek Herbarium, Windhoek. Artist: Marieta Visagie.
156 Flowering Plants of Africa 65 (2017)
and smaller upper stigmal lobe, each fruit locule with three seeds, and its distinctive pollen
with large reticulations (Balkwill & Welman 2000). Crabbea has a contorted corolla in bud,
with a cupular disc and a highly congested dichasial cyme surrounded by bracts. Ruelliopis
differs from Ruellia by their anthers which are spurred, the upper lobe of the stigma absent,
with one or two seeds in each locule, and banded pollen (Balkwill & Welman 2000). The
genus name Ruellia was established by Linnaeus in 1753 and commemorates Jean de la
Ruelle of Soissons, physician to Francis I and author of De Natura Stirpium in 1536 (Jackson
1990). The species name kaokoensis is in reference to the region where the species was
first collected. There are 42 species recorded for Ruellia in sub-Saharan Africa (Klopper et
al. 2006). Of these, 13 have been recorded for southern Africa, of which seven for Namibia
(Germishuizen & Meyer 2003). These include R. aspera (Schinz) E.Phillips, R. brandbergensis
Kers, R. currorii T.Anderson, R. diversifolia S.Moore, R. kaokoensis Van Jaarsv., R. otaviensis
P.G.Mey. and R. pilosa L.f. Most are herbaceous perennials growing in various habitats such
as savanna, grassland and semi-desert.
Ruellia kaokoensis is an ascending perennial with glaucous stems and leaves and beau-
tiful maroon to pinkish, trumpet-shaped flowers, from northwestern Namibia and south-
western Angola. Where it occurs it is locally abundant in dry hot subtropical desert con-
ditions. Ruellia kaokoensis is closest to R. bignoniiflora S.Moore, a species widespread in
south tropical Africa (Vollesen 2008). Ruellia bignoniiflora was named by Spencer Le March-
ant Moore in the Journal of Botany (1880) from a plant collected in the Luanda District of
Angola (Welwitsch 5063). Ruellia bignoniiflora is a small shrub, 1.0–1.5 m high, with white
nocturnal flowers that open at dusk and shed the flowers early the next day – clearly a
moth-pollinated species. Its floral tube is 70–130 mm long of which the basal cylindrical
part is 45–110 mm long. Ruellia kaokoensis differs by its much smaller stature (herbaceous
perennial with glaucous leaves and stems), its maroon diurnal flowers (opening during the
morning and falling shortly after dusk) and its corolla tube that is 25–30 mm long.
One of us (EJvJ) first came across Ruellia kaokoensis in the Kaokoveld on an expedition
in June 2000 while studying the obligatory cliff-growing succulents in the lower Cunene val-
ley at Otjiboronbongo. The plant was found in dry and hot semi-desert conditions, on dol-
omitic limestone rocks (Van Jaarsveld 16480). A year later, a specimen was recollected from
the same location (Van Jaarsveld 16823). Plants were found growing scattered on rocky
ridges among mopane (Colophospermum mopane (J.Kirk ex Benth.) J.Kirk ex J.Léonard),
Petalidium coccineum and species of Commiphora (C. virgata Engl., C. multijuga (Hiern)
K.Schum.). Other species observed in the habitat include the annual Thamnosma africana
Engl., Aloe corallina Verd. on the cliffs, Ficus cordata Thunb., F. glumosa Delile, F. ilicina
(Sond.) Miq., Adansonia digitata L., Tinospora fragosa I.Verd. & Troupin and Kalanchoe
laciniata (L.) DC. During an expedition to southwestern Angola, arranged by Brian Huntley
in January 2009 (Van Jaarsveld 2010b), the plant was yet again encountered at Omauha,
Iona National Park among Acacia montis-usti Merxm. & A.Schreib., A. mellifera (M.Vahl.)
Benth. subsp. detinens (Burch.) Brenan, Commiphora glaucescens Engl., C. multijuga, Boscia
albitrunca (Burch.) Gilg & Benedict, Euphorbia virosa Willd., Salvadora persica L., Tamarix
usneoides E.Mey. ex Bunge and Parkinsonia africana Sond. In 2010 Angola was revisited
with Wessel and Hannelie Swanepoel and Tielman Haumann, and R. kaokoensis was found
as far north as Meva Strand near Benguellia along the northern Namib coast (Figure 1 & 2).
Flowering Plants of Africa 65 (2017) 157
FIGURE 2.—Ruellia kaokoensis in flower, growing at Meva Strand near Benguellia (northern Namib) in
southwestern Angola. Photograph: E.J. van Jaarsveld.
158 Flowering Plants of Africa 65 (2017)
cylindrical part is 25 mm long and throat is 27 mm long, surface glandular hairy, lobes
widely spreading, 18–20 × 15 mm, broadly obovate to rounded. Stamens included in
throat, subequal or slightly didynamous; filaments free, 20–25 mm long, glabrous; anthers
5–6 mm long. Ovary slender, glabrous, ± 10 mm long, 4 mm in diameter; stigma 30 mm
long with two subequal lobes 2–5 mm long. Capsule and seed not seen. Flowering time:
November to July. Plate 2340.
REFERENCES
BALKWILL, K. & WELMAN, W.G. 2000. Acanthaceae. In O.A. Leistner (ed.), Seed plants of south-
ern Africa: families and genera. Strelitzia 10: 34–45.
GERMISHUIZEN, G. & MEYER, N.L. (eds). 2003. Plants of southern Africa: an annotated checklist.
Strelitzia 14. National Botanical Institute, Pretoria.
HEYWOOD, V.H. (ed.). 2007. Flowering Plants of the World. Firefly Books, London.
JACKSON, W.P.U. 1990. Origins and meanings of names of South African Plant Genera. UCT Eco-
lab, Cape Town.
KLOPPER, R.R., CHATELAIN, C., BÄNNINGER, V., HABASHI, C., STEYN, H.M., DE WET, B.C.,
ARNOLD, T.H., GAUTIER, L., SMITH, G.F. & SPICHIGER, R. 2006. Checklist of the flower-
ing plants of sub-Saharan Africa. An index of accepted names and synonyms. South African
Botanical Diversity Network Report No. 42. SABONET, Pretoria.
MOORE, S. 1880. Enumeratio Acanthacearum. Journal of Botany, British and Foreign 18: 198.
RAIMONDO, D., VON STADEN, L., FODEN, W., VICTOR, J.E., HELME, N.A., TURNER, R.C.,
KAMUNDI, D.A. & MANYAMA P.A. (eds). 2009. Red List of South African plants. Strelitzia
25. South African National Biodiversity Institute, Pretoria.
VAN JAARSVELD, E.J. 2010a. Waterwise gardening in South Africa and Namibia. Struik, Cape Town.
VAN JAARSVELD, E.J. 2010b. Angolan botanical expedition: Succulent Treasures, January 2009.
Aloe 47,1: 4–35.
VOLLESEN, K. 2008. Acanthaceae (Part 1). In H.J. Beentje & S.A. Ghazanfar (eds), Flora of Tropical
East Africa: 202–214.
INTRODUCTION
Contributions from authors and artists are most welcome. The policy of the editorial
committee of Flowering Plants of Africa (FPA) is to obtain contributions from as wide a
range of authors and artists as possible and to depict a rich and interesting mix of plants
from all over Africa. SANBI reserves the right to purchase original artwork and retain
copyright for FPA. All contributions are assessed by referees.
Plates that have already been published in colour in some other journal or magazine
are generally not acceptable for FPA.
Authors and/or artists are advised to contact the editor (Alicia Grobler) before any
work is undertaken with a view to publication in FPA: a.grobler@sanbi.org.za.
AUTHORS
All plates published in Volumes 1–49, are listed in the Index published in 1988.
There are, however, several completed plates awaiting publication in FPA. Authors
intending to have plants illustrated and written up for FPA are therefore advised to
contact the editor to establish whether a plant has not already been figured for FPA.
Authors are expected to supervise the execution of plates to ensure that they are
botanically correct. As detailed under the section ‘Artists’, a write-up and/or plate will
not be acceptable unless a voucher specimen is made of the figured specimen and
preserved in a recognised herbarium.
Before writing up a text for FPA, the author should ascertain whether the plate
has been approved by the journal’s panel of referees consisting of both botanists and
botanical artists. Only after approval of the plate, should the author produce the text for
submission to the editor. Approved contributions will be published when space allows.
Those in which the names of new taxa are published for the first time usually receive
priority.
1. Synopsis. The correct name of the plant together with its author/s and relevant
literature (name of publication written out in full) is cited. Put a comma after the
author’s name if the publication that follows is his/her own; write the word ‘in’ after
160 Flowering Plants of Africa 65 (2017)
the author’s name if the publication that follows is a journal or other work edited/
compiled by someone else. Following this, are synonyms (listed chronologically) plus
their author/s and relevant literature reference/s reflecting only author, page and year
of publication, e.g. Boris et al.: 14 (1966). The whole synopsis is in one paragraph.
If there are many synonyms and literature references, restrict yourself to the most
important ones. References should be arranged in chronological sequence; where
two or more references by the same author are listed in succession, the author’s
name is repeated with every reference. Author citations of plant names should
follow Brummitt & Powell (1992, Authors of plant names); for other authors, give full
surnames followed by a colon, page number/s and date.
2. Literature references in the text. Should be cited as follows: ‘Jones & Smith (1999)
stated …’ or ‘… (Jones & Smith 1999)’ when giving a reference simply as authority
for a statement. When more than two authors are involved, use the name of the first
author followed by ‘et al.’ Personal communications are given only in the text, not in
the list of references; please include full initials to identify the person more positively.
Referencing websites are generally discouraged unless their content is based on
reviewed scientific research (e.g. databases of plant names, online Floras, specimen
images, Red Lists, etc.) or are otherwise trustworthy and reputable.
3. List of references. All publications referred to in the synopsis and the text, but no
others, are listed at the end of the manuscript under the heading ‘References’. The
references are arranged alphabetically according to authors and chronologically
under each author, with a, b, c, etc. added to the year if the author has published
more than one work in a year. If an author has published both on his/her own and
as senior author with others, the solo publications are listed first and after that, in
strict alphabetical sequence, those published with one or more other authors. Author
names are typed in capitals. Titles of books and journals are written out in full, in
italics. In the case of books, the name of the publisher is followed by a comma and the
place of publication. For websites, provide full website name, address and the date
when it was accessed in square brackets.
4. Text proper. It should be written in language and style acceptable to both the scientist
and informed lay person. The following features should, as far as possible, be described
and discussed in the text:
• Main diagnostic characters for a brief pen picture of the plant.
• Affinities: how the taxon differs from its nearest allies; if necessary keys may be
used to distinguish closely related taxa.
• History of the taxon, where and when first collected and by whom.
• Geographical distribution in Africa: a distribution map, which will be handled as
a figure, is essential; authors should submit a list of Quarter Degree Grid Cells,
or GPS coordinates in decimal degrees in two columns (first column for latitude,
second column for its corresponding longitude), from which the editor’s office
will produce the distribution map.
• Ecology: habitat preferences etc. It is highly recommended that authors submit a
photograph of the plant in habitat or a general habitat image.
• Phenology: time of flowering, fruiting, etc.
Flowering Plants of Africa 65 (2017) 161
5. Description. This is a formal description of the taxon and not merely of the specimen
illustrated. For measurements, use only units of the International System of Units (SI).
Use only mm and/or m.
6. Captions. Supply a caption for the colour plate, indicating the relevant magnifications
and/or reductions, and citing the voucher specimen used for the illustration, i.e.
collector + number + herbarium (full name, not acronym) where the specimen is
housed. The caption ends with the name of the artist. Also supply captions for the
distribution map and any other figures you want to include (please use scale bars
where relevant), making sure all figures are mentioned in the text.
ARTISTS
1. Supervision. All illustrations should be executed under the supervision of the botanist
writing the text to ensure botanical accuracy and that details considered important by
the botanist are adequately depicted.
3. Paper. The paper must be of good quality and as white as possible. Arches or
Saunders Waterford (hot-press, 300 gsm) is recommended. The use of board should
be avoided.
4. Watercolours. The use of good-quality watercolours, e.g. Winsor & Newton (certain
pigments fade with time) or Schmincke (colourfast), is essential. The use of black paint
is not recommended as it is too harsh and tends to kill colour. Similarly, white paint
must be used with caution since it tends to dull adjoining colours and sometimes
reproduces as a bluish colour. Its use should be limited to white hairs and certain
highlights only. To reflect whiteness, endeavour to use the paper colour itself.
5. Subject material, composition etc. For obvious reasons, the subject material should
be representative of the species being illustrated and should be in excellent condition.
Drawing from photographs is not recommended: it is impossible to obtain the same
detail from a photograph as from the living plant.
All parts should be measured by the artist and magnifications indicated on the back
of the plate and the figure(s) where relevant.
162 Flowering Plants of Africa 65 (2017)
The plate should not be overcrowded with too many small dissections. These should
preferably be inserted as separate figures in the text. Dissections or habit sketches
included on the plate should be in pencil or colour, not ink, whereas dissections or
habit sketches to be used as separate text figures should be in ink.
A voucher specimen made of material from the plant(s) illustrated, must be preserved,
given a collector’s number and housed in a recognised herbarium as a permanent
record. This is most important: without a cited voucher specimen, the plate and
write-up will not be accepted.
Show clearly as many features as possible, for example apical buds, leaf axils, hairs,
glands, bracts, stipules, upper and lower surfaces of leaves showing venation, front,
side and back view of flowers, mature fruit, habit and where necessary and feasible,
the underground parts. The supervising botanist will know which features require
accentuation.
The artist’s signature must be unobtrusive but clearly written so that it will stand
reproduction. Numbering of plant parts should be done in light pencil: permanent
numbering will be done by the resident artist in consultation with the editor. No other
annotations should appear on the plate. Information such as species name, collector’s
name and number, date, locality, magnifications etc. should be written on the back of
the plate.
6. Dispatch of plates. Plates should be carefully packed, flat, using a sheet of Masonite
or similar material to prevent bending, and sent by registered or insured post.
Flowering Plants of Africa 65 (2017) 163
Index to Volume 65
Adenia wilmsii (2332) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Aloe braamvanwykii (2323) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Caesalpinia bracteata (2330) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Cissampelos hirta (2325) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Codonorhiza azurea (2321) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Eriosema distinctum (2331) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Esterhuysenia lucilleae (2333) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Gladiolus crassifolius (2322) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Gomphocarpus glaucophyllus (2336) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Ipomoea bolusiana (2337) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
Ipomoea cairica (2338) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Kalanchoe leblanciae (2328) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Pisonia aculeata (2334) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Protea foliosa (2326) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Protea namaquana (2327) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Rotheca myricoides (2339) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Ruellia kaokoensis (2340) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Schizostephanus alatus (2335) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
Senna didymobotrya (2329) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Tinospora fragosa subsp. fragosa (2324) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26