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TS'EPiSO REITUMETSE NTHAB!SENG TAOANA
THESIS SUBMITTED IN ACCORDANCE WITH THE REQUIREMENTS FOR f.r .
THE DEGREE
Wil.JJ'J GU$il'fE/& $CIlIEMil'U.liJfE
DEPARTMENT OF BOTANY AND GENETICS
IN THE FACULTY OF NATURAL AND AGRICULTURAL SCIENCES
UNIVERSITY OF THE FREE STATE
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Supervisor: Prof. H.J.T. VENTER
Co-Supervisor: Prof. R.L. VERHOEVEN
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TABLE OF CONTENTS:
CHAPTER: !PAGE #.:
1: INTRODUCTION 1
2: MATERIAL AND METHODS 5
2.1 Leaf surface micro-morphology 5
2.2 Pollen and Translator morphology 7
2.3 Seed surface morphology 8
2.4 Taxonomy 9
~&rRS.1J ~
3: lEAF SURFACE MICRO-MORPHOLOGY 12
3.1 Introduction 12
3.2 Results 13
3.3 Discussion and Conclusions 34
4: POLLEN AND TRANSLATOR MORPHOLOGY 36
4.1 Introduction 36
4.2 Results 38
4.3 Discussion and Conclusions 53
5: SEED SURFACE MORPHOLOGY 55
5.1 Introduction 55
5.2 Results 56
5.3 Discussion and Conclusions 65
6: TAXONOMY OF BASEONEMA Schltr, & Rendie 66
6.1 Historical Background 66
6.1.1 Generic Delimitation 66
6.2 Taxonomy - Baseonema gregorii 67
6.3 Distribution and Ecology 69
6.3.1 Voucher Specimens 70
7: TAXONOMY OF BATESANTHUS N.E. ar, 73
7.1 Historical Background 73
7.2 Taxonomy: Batesanthus 74
1. Batesanthus intrusus 76
2. Batesanthus parviflorus 76
3. Batesanthus purpureus 77
4. Batesanthus ta/botii 77
7.3 Distribution and Ecology - Batesanthus 78
7.3.1 Voucher Specimens 79
1. Batesanthus intrusus 79
2. Batesanthus parviflorus 80
3. Batesanthus purpureus 80
4. Batesanthus ta/botii 80
8: TAXONOMY OF MANGENOTIA Pichon 89
8.1 Historical Background 89
8.2 Taxonomy - Mangenotia eburnea 89
8.3 Distribution and Ecology 91
8.3.1 Voucher Specimens 92
9: TAXONOMY OF MONDIA Skeels 95
9.1 Historical Background 95
9.2 Taxonomy: Mondia 96
1. Mondia ecornuta 97
2. Mondia whitei 99
9.3 Distribution and Ecology - Mondia ecornuta 98
9.3.1 Voucher Specimens 98
9.4 Distribution and Ecology - Mondia whitei 100
9.4.1 Voucher Specimens 102
10: TAXONOMY OF SACLEUXIA Sam. 109
10.1 Historical Background 109
10.2 Taxonomy: Sacleuxia 109
1. Sacleuxia newii 110
2. Sacleuxia tuberosa 113
10.3 Distribution and Ecology - Sacleuxia newii 111
10.3.1 Voucher Specimens 112
10.4 Distribution and Ecology - Sacleuxia tuberosa 114
10.4.1 Voucher Specimens 115
11: TAXONOMY OF SARCORRHIZA Bullock 119
11.1 Historical Background 119
11.2 Taxonomy - Sarcorrhiza epiphytica 119
11.3 Distribution and Ecology 121
11.3.1 Voucher Specimens 122
12: TAXONOMY OF ZACA TEZA Bullock 126
12.1 Historical Background 126
12.2 Taxonomy - Zacateza pedicel/ata 126
12.3 Distribution and Ecology 128
12.3.1 Voucher Specimens 129
13: CLASSIFICATION 132
13.1 Classification 132
13.2 Tribal Position 134
14: GENERAL CONCLUSIONS 135
15: REFERENCES 140
SUMMARY 149
OPSOMMING 151
ACKNOWLEDGEM ENTS 153
APPENDIX A: ABBREVIATIONS AND TIERMINOLOGY
APPENDIX B: SPECIMEN LIST iii
INTRODUCTION:
That the Apocynaceae de Jussieu (1789), Asclepiadaceae Brown (1810) and
Periplocaceae Schltr. (1924) are closely related has been a subject of debate
for many years. The Periplocoideae (Kunze 1990, 1996, Judd et al. 1994,
Struwe et al. 1994, Sennblad & Bremer 1996, Endress et al. 1996, Endress
1997, Liede 1997, Sennbald 1997, Venter & Verhoeven 1997, Verhoeven &
Venter 1998a, b, Endress & Bruyns 2000) as presented in this study, is an
"Old World" (i.e. Africa, Asia, Madagascar, northern Australia and southern
Europe) subfamily as a result of amalgamation of the Apocynaceae s.s. and
Asclepiadaceae (incl. Periplocoideae). Historically the Periplocoideae was
classified in the Asclepiadaceae (Schumam 1895a), but was raised to family
level by Schlechter (1924). The family status has been maintained and used
by Verhoeven & Venter (1988, 1994b); Venter et al. (1990a, b, c), Dave &
Kuriachen (1991), Kunze (1993), Liede & Kunze (1993), Nilsson et al. (1993),
Omlor (1996), Swarupanandan et al. (1996), until the recent change of status.
The asclepiads have been included in the family: Apocynaceae s./. in order to
make the resultant taxon monophyletic, thus reflecting phylogeny of the group.
Morphological and molecular data using_the plastid gene matK has shown that
the Apocynaceae\Asclepiadaceae form a monophyletic group (Civeyrel et al.
1998). The resulting family thus consists of 424 genera, making it the seventh
largest angiosperm family.
Taxonomy entails discovery, identification, description, nomenclature as well
as the synthesis of information on diversity in the form of predictive, stable
and concise classification systems. The predictivity of classifications is
already important for locating chemicals, particularly those of medicinal or
economic importance. Most importantly, data accumulation must be done
while enough of the species are still extant.
1
But then a question of interest is that how do we maintain species diversity
when most parts of Africa are war-torn zones? Somehow our species are
preserved in the herbaria although it is difficult to interpret floral structures
from dried or rehydrated specimens. Of importance is the trend "conservation
through cultivation", the conservation of rare and endangered plants by
growing them in protected habitats (e.g. nature reserves) has now become a
generally accepted practice as is the case with Mondia whifei (Hook. f.)
Skeels. And in this study specifically based on African states, scarcity of
herbarium material, not to mention new collections, makes it even more
difficult to circumscribe the genera.
The Periplocoideae, "Pert" meaning around and "Plecein" to twine, has about
44 accepted generic names and c. 190 species and 18 of which are
monotypic and 9 have two species. This study looks into 12 species from the
following seven (7) selected genera of African Periplocoids:
1. Baseonema Schltr. & Rendie - 1 sp.
2. Bafesanfhus N.E. Br. 4 sp.
3. Mangenofia Pichon 1 sp.
4. Mondia (Hook. f.) Skeels 2 sp.
5. Sacleuxia Baill. 2 sp.
6. Sarcorrhiza Bullock 1 sp.
7. Zacafeza Bullock. 1 sp.
In order to have natural classification systems, taxonomic evidence gathered
from a variety of sources is needed and that means character states should
not be used in isolation but rather to complement other characters. As a
result of which, the "traditional" methods of taxonomy: vegetative and floral
morphology has been used (in part II chapters 6-12). These are supported
with additional morphological characters (in part I - chapters 3-5) using light
microscopy (LM), scanning electron microscopy (SEM) and transmission
electron microscopy (TEM) studies so as to evaluate the generic and/or
infrageneric classification, variability, uniformity and/or stability of the
characters within the Periplocoideae.
2
Vegetative features may contribute to generic delimitation. For instance, the
epiphytic growth form as a character (e.g. Sarcorrhiza and Ischno/epis Jum. &
H. Perrier) is an important feature. The rest of the genera in this study co-
exist as climbing lianas (e.g. Mondia) and/or climbing shrubs. All the species
investigated inhabit tropical rain, swamp and very rarely temperate forests of
Africa, sometimes some of the shrubs are found in semi-arid savanna (e.g.
Sacleuxia).
The flowers of the Periplocoideae are borne in a cymose inflorescence; most
flowers tend to be rotate. The composition and size of the inflorescence are
of little taxonomic significance (Venter & Verhoeven 1997). The flower is
uniquely structured and possesses a translator with a sticky viscidium to
promote animal pollination. Kunze (1993), puts forward that clarifying the
evolution of the translator can solve the question of common ancestry. The
depth of the corolla tube, position of the stamens and gynostegium in relation
to the tube can provide important taxonomic features. However, floral
morphology in the periplocoids is made difficult by the fact that most of the
flowers are small and fleshy.
The corona is one of the most frequently used features for taxon identification
at generic level. It is usually double. Genera within the Periplocoideae have a
corolline corona, usually divided into an outer and inner corona of
interstaminal discs forming a ring around the basal staminal column. These
interstaminal discs are nectariferous in nature (Kunze 1990). According to
Liede and Kunze (1993), the corolline corona is situated in the petal sinuses
that may be homologous to the sinal corolline corona in the Apocynaceae 5.5.
(Kunze 1990) and thus regarded as the oldest form of corona representing the
plesiomorphic type of corona formation. An annulus (secondary corolline
corona) forms a conspicuous tube inside the corolla tube; thus an annulus is
included as a second element within the corolline corona type (Kunze 1993).
The objectives of this study are to add to the existing literature and research
on the Periplocoideae (comprehensive palynological investigation and
3
taxonomic revision of the Periplocoideae being undertaken at the University of
the Free State - Botany Department). That is to:
c> draw up a dichotomous identification key to each of the seven genera and
their species.
<=> determine geographical distribution in the form of maps, ecological and
economic importance of each taxon.
<=> determine taxonomic relationships - through classification system (see
chapter 13).
And at the end to determine the significance and \ or importance of each
character state used in this study.
4
MATERIAL AND METHODS
Plant specimens for morphology and taxonomy of Periplocoids studied were
obtained from herbarium sheets (Appendix B). They were randomly selected
from different localities within the distribution range of the family in Africa.
PART I
No fresh material or FM preserved material was available. Dried flowers,
leaves and seeds were rehydrated in 3% phosphate-buffered glutaraldehyde
(GA) for a period of 36-48 hours prior to analysis.
2.1 leaf Surface Micro-morphology:
Due to limited availability of plant specimens, from two to six specimens were
studied per genus, with the genus Sarcorrhiza limited to only two specimens.
Light Microscopy (LM); the epidermis was peeled off, stained in 1% methylene
blue and mounted on slides with glycerine. Photographs (micrographs) were
taken with a Nikon Microphot FXA microscope. Trichomes were measured
using a Zeiss light microscope.
Scanning Electron Microscope (SEM); the rehydrated leaves were cut into
smaller pieces, postfixed in 1% osmium tetroxide (OS04), dehydrated in an
alcohol series (i.e. 30, 50, 70, 95 & 2 x 100%) and then critical point dried
using a Polaron critical point dryer E3000. Material was glued onto stubs;
gold coated with a BID RAD SEM coating system and examined with a Jeol
Winsem 6400 scanning electron microscope at 10 kV.
5
Herbarium specimens studied far leaf surfaces:
Baseanema gregarii Schltr. & Rendie: Goyder, Masinde, Meve &
Whitehouse 4006 (PRE); Faden, R.B. & A.J 74/436 (MO); Bally, P.R. O. 8745
(K); Field & Powys 174 (K); Bally, P.R.O. 8125 (K).
Batesanthus intrusus S. Moore: Tisserant, R.P. 1480 (P); Adam, JG. 39433
(MO); Louis, J 12992 (K).
Batesanthus perviitorus Norman: Po/hill, R.M. & Kirkup, D.W. 5190 (K);
Deighton, F.G. 3265 (K).
Batesanthus purpureus N.E. Br.: Bates, G.L. 383 (G); Le Testu, M. 5493
(P).
Batesanthus ta/batii S. Moore: Ta/bot, TA. 63 (K); Ta/bot, TA. 2021 (K).
Mangenatia ebumea Pichon: Leeuwenberg, A.JM. 4235 (WAG);
Leeuwenberg, A.JM. 8083 (WAG); Adam, JG. 30434 (MO); Thomas, N.W.
1373 (K).
Mandia ecornute (N.E. Br.) Bullock: Bouquet, A. 631 (P); K/aine, R.P. 577
(P); Musyoki, B.M. & Hansen, O.J 956 (K); Allen, G.E.F. 139 (K); Grote 5795
(K); Fau/kner, H.G. 558 (K).
Mandia whitei (Hook. f.) Skeels: Leeuwenberg, A.JM. 10026 (K); O/deman,
R.A.A. 392 (M); Med/ey Wood, J 6180 (PRE); Strey, R.G. 10347 (PRE);
Venter, H.J T 9282 (BLFU); Biege/, H.M., Pope, G. & Simon, B. 4297
(SRGH); Go/dsmith, B. 5/64 (SRGH); Ward, G.J 3626 (NH).
Sac/euxia newl! (Benth.) Bullock: Drummond & Hems/ey 3364 (K); Faulkner,
H.G. 1434 (K); Verdcourt 246 (K).
Sac/euxia tuberasa (E.A. Bruce) Bullock: Kerfoot, O. 3596 (K); Ford, J 847
(K); Tanner, R.E.S. 360 (K).
Sarcarrhiza epiphytica Bullock: Schlieben 2939 (BR); Bequaert 4494 (BR).
Zacateza pedicel/ata (K. Schum.) Bullock: Schweinfurth 3488 (K);
Gossweiler, J 13684 (K); Evrard 3467 (K); Leornard 698 (K); Louis, J 106
(K).
6
2.2 Pollen and Translator MorlPhology:
Flowers were fixed in 3% GA and then dissected in alcohol. No fresh material
was available except for Mondia whitei (Venter 9329, BLFU). For the genus
Mangenotia, pollen was very sparse. There were no flowers for Sarcorrhiza
epiphytica and Sacleuxia tuberosa. However, prepared stubs from the
laboratory of Prof. R. L. Verhoeven were used where available and indicated
by (*). Pollen samples were collected and acetolysed according to the revised
method of Erdtman (1960), for use in both light and scanning electron
microscopy.
Light Microscopy (LM); acetolysed pollen was mounted in glycerine jelly and
sealed with paraffin wax. Samples were examined and measured with a
Zeiss light microscope. Measurements of tetrad sizes were based on a
minimum of 15 and maximum of 30 per slide specimen. Photos (micrographs)
were taken with a Nikon Microphot FXA microscope.
Translators were mounted in glycerine jelly, examined and measured with a
Zeiss light microscope. Measurements were based on a minimum of three
translators per specimen because some were broken during the preparation
process.
Scanning Electron Microscopy (SEM); acetolysed pollen was air dried on
stubs, coated with gold and examined with a Jeol Winsem 6400 microscope at
5 kV. Translators were also mounted on stubs with double-sided tape, coated
with gold and examined with a Jeol Winsem 6400 microscope at 5 and 10 kV.
Transmission Electron Microscopy (TEM); pollen was fixed in 3% GA,
postfixed in 1% OS04, stained in 0.5% uranyl acetate, dehydrated in an ethyl
alcohol series and embedded in Spurr's low viscocity resin. Sections were
stained with uranyl acetate followed by lead citrate. Sections were cut with a
glass knife. For some species like Sarcorrhiza epiphytica (*), acetolysed
pollen was used for sectioning. Observations and micrographs were made
with a Philips CM 100 electron microscope at 60 kV.
7
Herbarium specimens studied for pollen and translator morphology:
Baseonema gregorii: Fie/d, D. V. & Powys, J.G. 174 (K); Faden, R.B. & A.J.
74/436, (PRE); Verdcourl & Po/hill 2695 (K).
Batesanthus intrusus: Louis, J. 12992 (K); Adam, J.G. 30433 (K); Tisserant,
R.p. 1480 (K).
Batesanthus parviflorus: Deighton, F.G. 3265 (K); Po/hill, R.M. & Kirkup.
D.W. 5190 (K); Gosswei/er, J. 8468 (BM).
Batesanthus purpureus: Bates, G.L. 383 (G, iso); Le Testu, M. 5493 (P).
Batesanthus talbotii: Ta/bot, TA. 63 (K); Ta/bot, TA. 2021 (K).
Mangenotia eburnea: Thomas, N.W. 4546 (K); Thomas, N.W. 4628 (K);
Adam, J.G. 30434 (MO); Leeuwenberg, A.J.M. 4235 (P); Leeuwenberg,
A.J.M. 8083 (BR); Thomas, N.W. 1373 (K); Thomas, N.W. 4628 (K).
Mondia ecornuta: Allen, G.E.F. 139 (K); Fau/kner, H.G. 558 (B, K).
Mondia whitei: Strey, R.G. 10347 (PRE); Pienaar, B.J. 170 (NH);
Leeuwenberg, A.J.M. 10026 (K); Med/ey Wood, J. 6180 (PRE); Ward, G.J.
3626 (NH); Venter, H.J. T 9329 (BLFU); Scheepers, J.G. 1058 (PRE); Venter,
H.J. T 9282 (BLFU).
Sacleuxia newii: Greenway & Kanuri 12852 (K); Drummond, R.B. &
Hems/ey, J.H. 3364 (USe); Verdcourl 246 (K).
Sac/euxia tuberose: Tanner, R.E.S. 360 (K)*.
Sarcorrhiza epiphytica: Semsei 2957 (K)*.
Zacateza pedicel/ata: Muusa, J. 3488 (K); Louis, J. 106 (K).
2.3 SeedSurfaceMorphology:
Due to unavailability of plant material, only the seeds of Baseonema gregorii-
van Someren 184 (K), Batesanthus intrusus - K/aine, R.P. 513 (P),
Mangenotia eburnea - Thomas, N.W. 4544 (K), Mondia whitei - O/deman,
R.A.A. 392 (MO), Sac/euxia newii - Verdcourl 246 (K), Sac/euxia tuberosa -
8
Kerfoot, O. 3596 (K) and Zacateza pedicellata - Evrard 3467 (K) were
examined and studied.
Scanning Electron Microscopy (SEM); rehydrated seeds were rinsed in pH 7.0
phosphate buffer and dehydrated in a series of 30, 50, 70, 95 and 2 x 100%
ethyl alcohol at intervals of 20 minutes. Seeds were critical point dried using a
Polaron critical point dryer and coated with SEM coating system. Subsequent
examinations were conducted with a Jeol Winsem 6400 microscope at 10 kV.
Stereo Microscopy; photos were taken with an Olympus SZ40 stereo
microscope. External form and appearance of the seeds as well as
measurements were done under a Nikon SMZ645 stereo microscope.
PART II
2.4 Taxonomy:
External morphology of available plant specimens was studied with a Nikon
SMZ645 stereo microscope. Plant parts were described in accordance with
the Systematics Association (1962) and Porter (1967). Micrographs of corona
lobes (Plate 1 - following pp. 88) were taken with SEM. Plant distribution, that
is, mapping for localities with locality grid were documented with references
from Bamps (1982), PoihilI (1988), The Times Atlas of the World (1985) and
Pope & Pope (1998). On the other hand, ecological importance was done
with the aid of collection cards! herbarium sheets. In appendix A, all plants
indicated by "(!)" were seen personally, otherwise all other specimens were
seen by my study supervisors.
List of herbaria (Holmgren et al. 1990) from where plant material was
obtained:
ALF Maisons-Alfort: Herbier, Départment de Botanique, Institut
d'Elevage et de Médecine Vétérinaire des Pays Tropicaux, 10
rue Pierre-Currie, F-94704 Maisons-Alfort Cedex, France.
BLFU: Bloemfontein: Geo Potts Herbarium, Botany Department,
9
University of the Free State, P.O. Box 339, Bloemfontein 9300,
Free State Province, South Africa.
BM London: Herbarium, Botany Department, The Natural History
Museum, Cromwell Road, London SW?5BD, England.
BOL Cape Town: Bolus Herbarium, Botany Department, University of
Cape Town, Private Bag, Rondebosch 7700, Western Cape
Province, South Africa.
BR Meise: Herbarium, Nationale Plantentuin van België, Jardin
Botanique National de Belgique, Domein van Bouchout, B-1860
Meise, Belgium.
BRLU: Bruxelles: Herbarium, Laboratorie de Botanique Systématique et
de Phytosociologie, C.P. 169, Université Libre de Bruxelles, 28
Avenue Paul Héger, B-1050 Bruxelles, Belgium.
CO Concarneau: Herbier Crouan, Laboratorie de Biologie Marine,
College de France, B.P. 11, F-2911 0 Concarneau, France.
COl Coimbra: Botanical Institute of the University of Coimbra,
Coimbra, Portugal.
G Géneve: Herbarium, Conservatoire et Jardin botaniques de la
Ville de Geneve, Case postale 60, CH-1292
Chambésy/Genêve, Switzerland.
GRA Grahamstown: Herbarium, Department of Agriculture and Water
Supply, Botanical Research Institute, P.O. Box 101,
Grahamstown 6140, Eastern Cape Province, South Africa.
K Kew: Herbarium, Royal Botanic Gardens, Kew, Richmond,
Surrey TWg :AB, England.
LlSC Lisboa: Herbário, Centro de Botanica, Instituto de lnvestiqacáo
Cientifica Tropical, Jardin-Museu Agricola Tropical, Calcada do
Galváo-Belérn, P-1400 Lisboa, Portugal.
M Munchen: Herbarium, Botanische Staatssammlung, Menzinger
Strasse 67, D-8000 Munchen 19, Federal Republic of Germany.
MO Saint Louis: Herbarium, Missouri Botanical Garden, P.O. Box
299, Saint Louis, Missouri 63166 - 0299, USA.
NBY Newbury: Herbarium, Newbury District Council, Newbury District
Museum Whart Street, Newburry, Berkshire, England.
10
NH Durban: Natal Herbarium, Botanical Research Unit, Botanic
Gardens Road, Durban 4001, Kwazulu-Natal Province, SA.
NU Pietermaritzburg: Herbarium, Botany Department, University of
Natal, P.O. Box 375, Pietermaritzburg 3200, KwaZulu-Natal
Province, South Africa.
P Paris: Herbier, Laboratorie de Phanérogamie, Muséum National
d'Histoire Naturelle 16 rue Buffon, F-75005 Paris, France.
PRE Pretoria: National Herbarium, National Botanical Institute, 2
Cussonia Avenue, Private Bag X 101 Pretoria 0001, Gauteng
Province, South Africa.
PRU Pretoria: H.G.W.J. Schweickerdt Herbarium, Botany Department
University of Pretoria, Pretoria 0002, Gauteng Province, South
Africa.
SAM Cape Town: South African Museum Herbarium, Private Bag X 7
Claremont 7735, Western Cape Province, South Africa - now
deposited at NBY but maintained a separate entity.
SRGH: Harare: National Herbarium and Botanie Garden, P.O. Box
8100, Causeway, Harare, Zimbabwe.
W Wien: Herbarium, Department of Botany, Naturhistorisches
Museum Wien, Burgring 7, a-1014 Wien, Austria.
WAG Wageningen: Herbarium Vadense, Department of Plant
Taxonomy, Agricultural University Postbus 8010, 6700 ED
Wageningen, Netherlands.
Z Zurich: Herbarium, Institut fur Systematische Botanik,
Untversitat Zurich, Zollikerstrasse 107, CH-8008 Zurich,
Switzerland.
11
LEAF SURFACE MICRO~MORPHOlOGY
3.1 INTRODUCTION:
The present investigation into leaf surface morphology of selected African
Periplocoideae genera was undertaken in order to complement other sources
of taxonomic evidence. Results of this chapter are presented in the form of
descriptive analysis following Metcalfe (1987). This chapter looks into the leaf
surface features namely; outline of epidermal cells, cuticular ornamentation,
the stomatal complex i.e. stoma, guard cells and subsidiary cells, as well as
foliar trichome types.
Since little or no information is available on leaf micro-morphology of the
Periplocoideae, the aim of this study was to add to the existing information
and to find the significance of all character states used to ultimately determine
their taxonomic value at generic or species level or within the subfamily as a
whole.
12
3.2 RESULTS:
3.2.1 BASEONEMA Schltr. & Rendie
Baseonema gregorii Schltr. & Rendie
Petiole: (Fig. 3.1 A, B) 4-(7-10)-26 mm long, purple with hirsute surface. The
petiole is more or less circular in cross-section, sometimes adaxially flattened
with a groove in the center.
Lamina: Epidermis (Fig. 3.1 C, D); cells are polygonal and cell walls straight
to slightly undulate. Cuticle (Fig. 3.1 E) is similar on abaxial and adaxial
epidermis, densely striate, with parallel wavy striae extending over periclinal
walls. Stomata (Fig. 3.1 C, F) only on abaxial epidermis. The stomata in
surface view are broadly elliptic to narrowly elliptic with broad apertures. The
region between an outer broad rim (indicated by 'r') and peristomatal rim
(indicated by .. ) (i.e., a cuticular ledge) is smooth. Some stomata are
surrounded by very long radiating striae (Fig. 3.1 F) whereas others have
striae radiating perpendicular to the guard cells. Stomata are paracytic.
Trichomes: (Fig. 3.1 G, H) are present on abaxial and adaxial epidermis.
The abaxial surface has a denser indumentum than the adaxial surface.
However, trichome type on both surfaces is the same. Trichomes are
uniseriate, unicellular, non-glandular and are relatively long and narrow (Table
3.1). The surface of the trichomes is covered with linear, warty outgrowths
(Fig. 3.1 G). Trichomes are straight to curved with an acute apex and the
base is covered with a smooth cuticle (Fig. 3.1 H).
Figure 3.1 Leaf surface of Baseonema gregorii: (A-B) Petiole with trichomes. Faden,
R.B. & A.J. 74/436. Scale bar: A - B = 100 JJm
13
... Figure 3.1 B. gregorii: (C) Surface view of abaxial epidermal cells. Bally, PR. 0 8125.
(0) Surface view of adaxial epidermal cells. Faden, R.B. & A.J. 74/436. (E) Adaxial
epidermis showing densely striate cuticle. Faden, R.B. & A.J. 74/436. (F) Abaxial epidermis
showing stomata with long radiating striae. Faden, R.B. & A.J. 74/436. (G) Abaxial epidermis
with trichomes. Field & Powys 174. (H) Adaxial epidermis with trichomes. Bally 8745.
r = rim of stomata. Scale bar: C, 0 = 50 urn; E, F, G = 10 urn: H = 100 urn
14
... Figure 3.1 B. gregorii: (C) Surface view of abaxial epidermal cells. Bal/y, PR. 0 8125.
(0) Surface view of adaxial epidermal cells. Faden, R.B. & A.J. 74/436. (E) Adaxial
epidermis showing densely striate cuticle. Faden, R.B. & A.J. 74/436. (F) Abaxial epidermis
showing stomata with long radiating striae. Faden, R.B. & A.J. 74/436. (G) Abaxial epidermis
with trichomes. Field & Powys 174. (H) Adaxial epidermis with trichomes. Bal/y 8745.
r = rim of stomata. Scale bar: C, D = 50 urn; E, F, G = 10 urn; H = 100 urn
14
3.2.2 BA TESANTHUS N.E. Brown
Trichomes are present in Batesanthus intrusus and B. talbotii, but absent in
B. parviflorus and B. purpureus.
Batesanthus intrusus S. Moore
Lamina: Epidermis (Fig. 3.2 A, B, C); consists of polygonal cells. Cells of the
abaxial epidermis (Fig. 3.2 A) have undulate and non-pitted anticlinal and
smooth periclinal walls. Anticlinal walls of adaxial epidermis are straight to
slightly undulate and periclinal walls are smooth (Fig.3.2 B, C). Cuticle (Fig.
3.2 D) on both abaxial and adaxial epidermis is smooth. Ornamentation on
the adaxial epidermis is generally a reticulum of rounded or curved ridges.
Stomata (Fig. 3.2 E, F) were observed only on the abaxial epidermis. The
stoma in surface view are broadly elliptic (Fig. 3.2 F) to almost rounded (Fig.
3.2 E) with slightly narrow to broad apertures. The rim ('r') is very prominent
and slightly raised above the subsidiary cells. Subsidiary cells are outlined by
a slightly undulate, raised cuticular ridge (Fig. 3.2 F). Trichomes: (Fig. 3.2
G) are present on abaxial epidermis and sparsely distributed. Trichomes are
uniseriate, unicellular and non-glandular. Trichomes are short and stout,
smooth and straight with an acute apex. The base is nearly rounded.
Batesanthus parviflorus Norman
Lamina: Epidermis (Fig. 3.3 A, B); epidermal cells of the abaxial epidermis
are polygonal with straight to undulate walls. Cuticle (Fig. 3.3 C) varies within
the species from smooth to a slightly wavy. Short-radiating striae extending
over the subsidiary cells occur on the lower surface (Fig. 3.3 E). Stomata
(Fig. 3.3 E) only on abaxial epidermis. The stomata are broadly elliptic with a
slightly raised broad rim and narrow apertures. The region between the guard
cells and the peristomatal rim is smooth. Trichomes: The leaf surface is
glabrous.
Batesanthus purpureus N.E. Br.
Lamina: Epidermis (Fig. 3.4 A, B); abaxial and adaxial surfaces have
polygonal cells. Cell walls are straight to slightly undulate, non-pitted anticlinal
15
and smooth periclinal walls. Cuticle (Fig. 3.4 C) on adaxial epidermal surface
is smooth with slightly undulate walls and raised cuticular ridges over anticlinal
walls. Stomata (Fig. 3.4 D) only on the abaxial epidermis. The stoma in
surface view are broadly elliptic with narrow to wide apertures and a very
distinct rim ('r') which is slightly raised. Trichomes: The leaf surface is
glabrous.
Batesanthus telbotii S. Moore
Lamina: Epidermis (Fig. 3.5 A); consists of polygonal cells. Cell walls of the
abaxial epidermis have undulate, non-pitted anticlinal walls and finely striate
periclinal walls. Cuticle (Fig. 3.5 B, C) on abaxial epidermal surface consists
of short parallel striations around the stomatal region, however striations are
not always in the same direction (Fig. 3.5 C). Adaxial epidermis is densely
striae with wavy parallel striations continuos over irregularly raised periclinal
walls and cuticular grooves over anticlinal walls (Fig. 3.5 B). Stomata (Fig. 3.5
C) observed only on abaxial epidermal surface. The stomata in surface view
are broadly elliptic with wide and occasionally narrow apertures (Fig. 3.5 C).
The rim is broad and distinct. The region between an outer broad rim and the
peristomatal rim is smooth. The stomata are regularly surrounded by short
radiating striae. Stomata are paracytic. Trichomes: (Fig. 3.5 D) were only
found on the abaxial epidermal surface and sparsely distributed. They are
uniseriate, unicellular and non-glandular. Trichomes are short and stout.
They are smooth and straight with an acute apex. The round base is broadly
flattened and embedded in a rosette of epidermal cells.
16
Figure 3.2 Leaf surface of Batesanthus intrusus: (A) Surface view of abaxial
epidermis showing cells with undulate walls. Louis, J. 12992. (B) Surface view of
adaxial epidermis showing cells with straight walls. Adam 30433. (C) Surface view of
adaxial epidermis showing slightly undulate walls. Le Testu 8501. (0) Abaxial
epidermis cuticle. Tisserant 1480. (E) Stoma in abaxial epidermis. Adam 30433. (F)
Stoma in abaxial epidermis. Tisserant 1480. (G) Trichomes on abaxial epidermis. Le
Testu 8501.
Scale bar: A-C = 50 urn; D-F = 10 urn; G = 100 urn
17
A
Figure 3.3 Leaf surface of Batesanthus parviflorus: (A) Surface view of
abaxial epidermis. Deighton, F.C. 3265. (B) Surface view of abaxial
epidermis. Po/hill & Kirkup 5190. (C) Adaxial epidermis showing smooth
cuticle. Po/hill & Kirkup 5190. (0) Adaxial epidermis showing cuticle with
radiating striae. Deighton, F.C. 3265. (E) Abaxial epidermis with stomata
showing parallel striae. Po/hill & Kirkup 5190.
Scale bar: A, B = 50 urn: C-E = 10 urn
18
Figure 3.4 Leaf surface of Betesentbus purpureus: (A) Surface view of
abaxial epidermal cells. Bates, G.L. 383. (B) Surface view of adaxial
epidermis. Bates, G.L. 383. (C) Adaxial epidermis cuticle. Le Testu 5493. (0)
Stomata on abaxial epidermis. Le Testu 5493.
Scale bar: A-B = 50 urn; C = 100 urn: D = 10 urn
19
Figure 3.5 Leaf surface of Batesanthus talbotii: (A) Surface view of
abaxial epidermal cells. Ta/bot, TA. 63. (B) Cuticle on adaxial epidermis.
Ta/bot, TA. 63. (C) Stomata on abaxial epidermis. Ta/bot, TA. 63. (D)
Trichomes on abaxial epidermis. Ta/bot, TA. 63.
Scale bar: A, 0 = 50 urn: B-C = 10 urn
20
3.2.3 MANGENOTIA Pichon
Mangenotia eburnea Pichon
Lamina: Epidermis; epidermal cells in surface view have straight to slightly
undulate, non-pitted, anticlinal walls. Cuticle (Fig. 3.6 A, B): Adaxial epidermis
is characterised by irregular bumps with parallel striations that extend over
individual cells (Fig. 3.6 A). The cuticle is sometimes wavy with variously
orientated striae (Fig. 3.6 B). Globules of papilla cover the abaxial epidermal
surface. Stomata (Fig. 3.6 C, D) were observed only on abaxial epidermis.
The stomata in surface view are narrowly elliptic to broadly elliptic; apertures
are mostly narrow with a distinct raised rim (Fig. 3.6 D). The region between
an outer rim and the peristomatal rim is smooth with the stomata depressed in
between the papilla. Papillae: Abaxial epidermis is beset with papilla.
Papillae are globular and connected together by thread-like radiating ridges
(Fig. 3.6 D). There are 4-6 papillose structures surrounding each stoma.
Trichomes (Fig. 3.6 E): appear to be short and stout and sparsely distributed
on adaxial epidermis. They are straight, smooth with an acute apex. At the
base, there occur long radiating striae (Fig. 3.6 E).
Figure 3.6 Leaf surface of Mangenotia eburnea (A) Cuticle on adaxial
epidermis showing irregular bumps and parallel striations. Thomas, N.W
4628. (8) Adaxial epidermis showing wavy striae. Thomas, N.W 1373.
Scale bar: A-B = 10 urn
21
Figure 3.6 Leaf surface of Mangenotia eburnea: (C) Stomata on abaxial
epidermis. Thomas, N.W 1373. (0) Stomata on abaxial epidermis. Thomas,
N.W 4628. (E) Trichomes on adaxial epidermis. Thomas, N.W 1373.
Scale bar: C-E = 10 IJm
22
3.2.4 MONDIA Skeels
Mondia ecornuta (N.E.Br.) Bullock
Lamina: Epidermis; epidermal cells have slightly undulate anticlinal walls.
Cuticle (Fig. 3.7 A, B) is wavy with parallel radiating striations. Most
specimens studied have wavy and parallel striations with cuticular grooves
over anticlinal walls (Fig.3.7 A). Stomata (Fig. 3.7 C, D) occur only on abaxial
epidermis. The stomata are paracytic and in surface view are mostly elliptic.
The apertures are narrow. Trichomes (Fig. 3.7 E, F) are present on both the
abaxial and adaxial epidermis. However, the upper surface is glabrous or
nearly so. Trichomes are long and narrow and are inserted in a rosette of
epidermal cells (Fig. 3.7 F). The surface is covered with linear, warty
outgrowths (Fig. 3.7 E).
Mondia whitei (Hook. f.) Skeels
Lamina: Epidermis; epidermal cells have straight to slightly undulate anticlinal
walls. Cuticle (Fig. 3.8 A) is wavy with parallel to variously oriented striations
over bumpy periclinal walls as well as deep lying cuticular grooves over
anticlinal walls. Stomata (Fig. 3.8 B) occur only on abaxial epidermis. The
stomata in surface view are narrowly elliptical with a distinct rim. Apertures
are narrow with a peristomatal rim present. However rare, there occur short
radiating striae perpendicular to the guard cells (Fig. 3.8 B). Stomata are
paracytic. Trichomes (Fig. 3.8 C, D) occur on both abaxial and adaxial
epidermis with adaxial epidermis glabrous or nearly so, especially in mature
leaves. The lower surface is sparsely hairy with hairs mostly on the midrib
(Fig. 3.8 C) and secondary veins (Fig. 3.8 D). Trichomes are uniseriate,
unicellular and non-glandular. Trichomes are either beset with warty linear
outgrowths (Fig. 3.8 C) in some specimens (e.g. Goldsmith 5/64Ab-, Oldeman
392Ad- & Ward 3626Ab-) or smooth (Fig. 3.8 D) and straight with an acute apex
(e.g. Biegel et al. 4297Ab-, Medley Wood 6180Ad-, Leeuwenberg 10026 Ab-'Ad).
23
Figure 3.7 Leaf surface of Mondia ecornuta: (A) Cuticle on adaxial
epidermis. Faulkner, H.G. 558. (B) Cuticle on adaxial epidermis. Bouquet,
A. 631. (C) Stomata on abaxial epidermis. Faulkner, H.G. 558. (0)
Stomata on abaxial epidermis showing perpendicular running striations.
Faulkner, H.G. 558. (E) Abaxial epidermis showing trichomes on the
midrib. Musyoki & Hansen 956. (F) Abaxial epidermis showing a rosette of
epidermal cells at trichome insertion (trichome broken). Alien, C.E.F. 139.
Scale bar: A - F = 10 urn
24
Figure 3.8 Leaf surface of Mondia whitei: (A) Adaxial epidermis showing
cuticle with variously orientated striae. Goldsmith, B. 5/64. (B) Stomata on
abaxial epidermis. Venter, H.J. T. 9282. (C) Warty trichome on abaxial
epidermal midrib. Goldsmith, B. 5/64. (0) Smooth trichome on adaxial
epidermis. Leeuwenberg, A.J.M. 10026. Scale bar: A-C 10 urn;0 = 100!Jm
25
3.2.5 SACLEUXIA Bail!.
Leaves are simple and petiolate. Leaves are amphistomatie with paracytic
stomata in Sac/euxia tuberosa (E.A. Bruce) Bullock and hypostomatic with
paracytic stomata in Sacleuxia newii Bail!.
Sacleuxia newii (Benth.) Bullock
Lamina: Epidermis (Fig. 3.9 A, B); consists of polygonal cells (Fig. 3.9 A).
Cells on adaxial and abaxial surface have straight to slightly undulate
anticlinal walls (Fig. 3.9 B). Cuticle (Fig. 3.9 C) is wavy with dense striae on
both surfaces. Striae are variously orientated over slightly raised periclinal
walls. Cuticular grooves occur over anticlinal walls (Fig. 3.9 C). Stomata (Fig.
3.9 D) only observed on abaxial epidermis. The stomata in surface view are
broadly elliptic with a distinct and broad outer rim (Fig. 3.9 D). There is a
narrow outline of peristomatal rim bordering the stomata. Trichomes (Fig. 3.9
E, F) occur on both abaxial and adaxial epidermis. Abaxial epidermis surface
is more hairy than the adaxial and more so on the midrib and venous area.
The midrib is hirsute, forming a whitish line along. Adaxial surface with
sparsely distributed hairs. Trichomes are uniseriate, unicellular, non-glandular
and straight to slightly curved at the tip (Fig. 3.9 E). Trichomes are long and
narrow. Trichome surface is covered with linear, warty outgrowths. The base
is rounded and covered with smooth cuticle and embedded in a rosette of 6-8
epidermal cells (Fig. 3.9 F).
Sacleuxia tuberosa (E.A. Bruce) Bullock
Lamina: Epidermis (Fig. 3.10 A, B); consists of straight to slightly undulate,
non-pitted anticlinal walls on both abaxial (Fig. 3.10 A) and adaxial (Fig. 3.10
B) surfaces. Cuticle (Fig. 3.10 C) is wavy on both surfaces with parallel
striations continuous over the walls. Prominent cuticular grooves correspond
to anticlinal walls. Stomata (Fig. 3.10 0, E) occur on both abaxial and adaxial
epidermis. Stomata on adaxial epidermis are only found near the midrib area.
The stomata in surface view are broadly elliptic with a prominent rim and wide
apertures. Trichomes (Fig. 3.10 F) occur on both abaxial and adaxial
epidermis. Trichomes are uniseriate, unicellular and non-glandular. The
26
trichomes are relatively short and stout - compared to those of Sac/euxia
newii. The surface is covered with linear, warty outgrowths. The base is
rounded and slightly raised above a rosette of epidrmal cells. Trichomes are
mostly straight with an acute apex that is slightly curved in some instances.
Figure 3.9 Leaf surface of Sac/euxia newii: (A) Surface view of abaxial
epidermis. Verdeourl 246. (B) Surface view of adaxial epidermis. Verdeourl
188. (C) Aadaxial epidermis showing wavy, densely striated cuticle. Verdeourl
246. (0) Stomata on abaxial epidermis. Verdeourl 188. (E) Unicellular
trichome on abaxial epidermis. Drummond & Hems/ey 3364. (F) Trichome on
adaxial epidermis showing rosette of epidermal cells. Verdeourl 188.
Scale bar: A, B, E = 50 urn; C, D, E = 10 pm
27
Figure 3.10 Leaf surface of Sac/euxia tuberosa: (A) Surface view of
abaxial epidermis showing slightly undulate walls. Tanner, R.E.S. 360. (B)
Surface view of adaxial epidermis. Kerfoot, O. 3596. (C) Adaxial epidermis
cuticle. Tanner 360. (0) Stomata on abaxial epidermis. Tanner 360. (E)
Stomata on adaxial epidermis. Kerfoot, O. 3596.
Scale bar: A-B = 50 urn: C-D = 10 urn: E = 100 urn
28
3.2.6 SARCORRHIZA Bullock
Sarcorrhiza epiphytica Bullock
The petiole is 4-(8-9)-10 mm long, pubescent and dark crimson. The petiole
is more or less rounded to flattened with longitudinal fissures.
Lamina: Epidermis (Fig. 3.11 A, B); epidermal cells are polygonal. The cells
have slightly undulate, non-pitted anticlinal walls and smooth periclinal walls.
Cuticle (Fig. 3.11 C) is smooth. Stomata (Fig. 3.11 0, E) occur only on the
abaxial epidermis. Most of the stomata in surface view are rounded (Fig. 3.11
D), sometimes broadly elliptic (Fig. 3.11 E). The rim is very distinctive, there
is no peristomatal rim. The stomata are slightly raised (Fig. 3.11 D).
Trichomes (Fig. 3.11 F) were observed on the midrib of both abaxial and
adaxial surfaces. Trichomes are rare and the leaves are almost glabrous in
mature state. Trichomes are uniseriate, unicellular, non-glandular, short and
stout. Trichomes are straight to slightly curved with a tapering point.
29
Figure 3.11 Leaf surface of Sarcorrhiza epiphytica: (A) Surface view of
abaxial epidermis. Bequaert 4494. (B) Surface view of adaxial epidermis.
Bequaert 4494. (C) Adaxial epidermis showing smooth cuticle. Schliebenm
2939. (0) Rounded stoma on abaxial epidermis. Bequaert 4494. (E) Broadly
elliptic stomata on abaxial epidermis. Schliebenm 2939. (F) Trichomes on
abaxial midrib. Bequaert 4494.
Scale bar: A-B = 50 pm: C-E = 10 urn: F = 100 urn
30
3.2.7 ZACATEZA Bullock
Zacateza pedicellata (K. Schum.) Bullock
Lamina: Epidermis (Fig. 3.12 A, B); epidermal cells are polygonal. Cells of
the abaxial epidermis are undulate with pitted anticlinal and finely striate
periclinal walls. Adaxial epidermal cells have straight to slightly undulate,
pitted anticlinal walls, sometimes with non-pitted anticlinal walls. Cuticle (Fig.
3.12 C, D) is wavy and densely striate on adaxial epidermis. In some cases,
the cuticle appears to be smooth with parallel striations in sunken areas (Fig.
3.12 C). Cuticle on abaxial epidermal surface is wavy (Fig. 3.12 E) with
parallel striations extending over cells (Fig. 3.12 F). It can however vary to
radiating striae perpendicular to guard cells (Fig. 3.12 E). Stomata (Fig. 3.12
E, F) were observed on the abaxial epidermis. The stomata in surface view
are narrowly elliptic to broadly elliptic with narrow apertures. The region
between the outer rim and the peristomatal rim is smooth. Variations occur on
the lower surface, with stomata having radiating striae perpendicular to the
guard cells and extended as lateral wings (Fig. 3.12 E). In other specimens,
concentric rings of striae followed by variously orientated striae (Fig. 3.12 F)
occur. Trichomes (Fig 3.12 G, H) are uniseriate. Trichomes are unicellular,
non-glandular, relatively short and stout with acute apex. Trichomes are
sparsely distributed on both surfaces.
Figure 3.12 Leaf surface of Zacateza pedicellata: (A) Surface view of abaxial
epidermis. Gossweiler, J. 13684. (B) Surface view of adaxial epidermis.
Schweinfurth 3488.
Scale Bar: A-B = 50 urn
31
... Figure 3.12 Leaf surface of Zacateza pedicel/ata: (C) Adaxial epidermis showing
parallel, sunken striations. Louis, J. 106. (0) Adaxial epidermis showing densely
striated cuticle. Gossweiler, J. 13684. (E) Stomata on abaxial epidermis. Evrard
3467. (F) Stomata on abaxial epidermis showing concentric rings of striae.
Gossweiler, J. 13684. (G) Trichome on abaxial epidermis. Gossweiler, J. 13684. (H)
Trichome on adaxial epidermis. Gossweiler, J. 13684.
Scale bar: C - H = 10 pm
TABLE 3.1 TYPES AND DENSITY OF FOllAR TRICHOMES IN SELECTED PERiPLOCOID SPECIES
SPECIES Ab- Ad- Warty Smooth Indumentum (trichome cover/density) Uni-
Baseonema gregorii ,{ ,{ J Hirsute ./
Very dense
Batesanthus intrusus J J Puberuient ./
Sparse to glabrous in mature leaves
Batesanthus parviflorus - - - - Glabrous
Batesanthus purpureus - - - - Glabrous
Batesanthus ta/botJJ J J Scabrous ./
Sparsely distributed
Mangenotia eburnea * J J Puberuient ./Sparsely distributed
Mondia ecornuta J J J Puberuient J
Sparsely distributed to glabrous on adaxial epidermis
Mondia whitei J J J J Puberuient to hirsute. Glabrous or nealy so in mature leaves (adaxial) J
Sparsely distributed but dense on the midrib
Sac/euxia newii J oef oef Hirsute oef
Sparsely distributed but dense on the midrib
Sac/euxia tube rosa oef oef oef Pubescent J
Sparsely distributed but dense on the midrib
Sarcorrhiza epiphytica oef J Puberuient ./
Rare, becoming glabrate
Zacateza pedicel/ata J J J Puberuient ./
Rare
Ab- = Abaxial * = Papillose J = Present Uni- = Unicellular
(j.)
(j.) Ad- = Adaxial - = Absent
3.3 DISCUSSION AND CONCLUSIONS
The leaves are hypostomatic in all species except Sac/euxia tuberosa with
amphistomatous leaves. However, the stomata in S. tuberosa are found
towards the apex of the lamina and are situated close to the midrib.
An unsubstantiated belief is that hypostomatous leaves are better adapted to
dry conditions than amphistomatous leaves (Willmer & Fricker 1996).
Spacing of the stomata in the epidermis is characteristic of the species
although variations do occur between species (whereby S. tuberosa - Kerfoot
3596 is the only specimen with stomata on both surfaces) and this is likely
modified by leaf morphology and / or genetic composition?
The leaves are characterized by having paracytic stomata, a character
common among members of the Periplocoideae and this is considered
primitive within the angiosperms (Wilkinson 1979).
The epidermal cells and cuticular ornamentation show very little variation
within the taxa and they are usually uniform in structure, hence of no
taxonomic value. However, Mangenotia eburnea is characterized by the
presence of papillae on the lower surface, a character not common within the
subfamily. Unpublished data of Venter & Verhoeven show presence of
papillae on abaxial epidermis in some species of Crypto/epis R. Br. e.g. C.
stefaninii and in Curroria decidua subsp. vo/ubi/is (Balf. f.) Bullock. The cuticle
is characterised by being smooth (e.g. Batesanthus parviflorus, Sarcorrhiza
epiphytica), wavy or with variously orientated striations (in all other taxa),
sometimes parallel running striations are found.
The Periplocoids in this study are all characterised by having simple,
uniseriate, unicellular and non-glandular trichomes (i.e. hairs of similar shapes
are known to occur) with variations in their distribution. For instance,
Baseonema gregorii is densely pubescent whereas all other taxa are minutely
pubescent to nearly glabrous. As a result of which, trichomes are similar in
structure and therefore of little taxonomic value in this regard.
34
However, when present on the stem, bracts and flowers they are of the same
shape/type as those on the leaves. Trichomes have been found to have
variation patterns on their surface, they are either smooth or with linear warts
and they are straight to slightly curved. Both types were observed in
specimens of Mondia whitei.
Nevertheless, pubescence of the leaf is one feature that varies with leaf age.
For the originally pubescent leaf may remain distinctly hairy, become glabrate
or glabrous or retain some trichomes in sheltered places on the abaxial
surface, such as the side of the midrib (e.g. Sarcorrhiza epiphytica) or as
Hardin (1979) states, in the axils of main secondary veins as these are the
last regions to become glabrous as the leaf ages (e.g. Mondia ecornuta and
M. whiteI). According to Theobald et al. (1979), many instances of the
glabrous condition represent cases where trichomes have degenerated at an
early stage in their development or were lost shortly after maturation.
According to Corsi & Bottega (1999), the non-glandular hairs on the
vegetative and reproductive organs are abundant and longer when the organs
are young. Particularly in the early phases of the ontogenetic cycle, they
presumably collaborate with glandular hairs in mechanical defence, creating a
thick downy layer and they are certainly also involved in protecting the plant
from excessive transpiration and insolation (Corsi & Bottega 1999).
Thus, leaf surface micro-morphology has been found to be of little, if any
taxonomic value within the Periplocoids investigated.
35
POLLEN AND TRANSLATOR MORPHOLOGY
4.1 INTRODUCTION:
Palynologically the Periplocoideae are distinguished from Secamonoideae and
Asclepiadoideae by the presence of tetrads, or free pollinia (in 11 genera;
Verhoeven & Venter 1998a, b), which are shed onto translators at anthesis. The
Secamonoideae are distinguished from Asclepiadoideae by having four pollinia
attached to a translator as opposed to two pollinia in Asclepiadoideae. In the
Secamonoideae and Asclepiadoideae, at the last stage of pollinarium
development, when the anther wall dehisces, the pollinium comes into contact
with, and becomes attached to the translator apparatus (Kunze 1994, Omlor
1996, Civeyrel et al. 1998). The pollinium is not released from the anther at this
stage. The pollinium remains in the anther Iocuie until the pollinator removes the
pollinarium. In the Periplocoideae the translator is attached to the pollinator by
means of an adhesive disc, while in the Secamonoideae and Asclepiadoideae it
is a clasping mechanism.
The Periplocoideae are characterised by having tetrads or free pollinia which are
shed onto cone-shaped or spoon-like translators, each of which consists of an
adhesive disc (by which the translator sticks to the pollinator), a stalk and a
spoon (onto which the tetrads or pollinium is shed at anthesis). The African
Periplocoideae are characterised by having pollen in tetrads, with the exception
of Schlechterella abyssinica (Chiov.) Venter & R.L. Verh. and S. africana (Schltr.)
K. Schum. having tetrads cohering together to form a pollinium (Verhoeven &
Venter 1998b).
36
The pollen morphology of genera such as Raphionacme Harv. (Verhoeven &
Venter 1988), Tacazzea Decne. and Petopentia Bullock (Verhoeven et al. 1989),
Ectadium E. Mey. (Venter et al. 1990b), Curroria Planeh., Mondia Skeels,
Stomatostemma N.E. Br. and Socotranthus Kuntze (Verhoeven & Venter 1993),
Periploca (Verhoeven & Venter 1994a) and genera of the Periplocaceae from
Madagascar (Verhoeven & Venter 1994b) have been investigated. The purpose
of this investigation is to provide palynological information on the seven genera
investigated and to find their taxonomic value.
In five vertical grooves situated around the periphery of the pentangular stylar
head, special epithelial cells secrete the translators consisting of three
morphological parts. The spoon, with its adhesive surface receives pollen
tetrads from adjacent anther halves. The adhesive disc functions to stick to a
visiting pollinator. The three translator parts have been described under different
names, for example, the spoon as a shovel (Safwat 1962), translator (Arekal &
Ramakrishna 1980) and 'Pollenschaufel' (Schick 1982). The stalk has also been
called a stipe (Kunze 1993) or stipes (Schick 1982). The adhesive disc has been
termed 'Klebplatte' (Schick 1982) or viscidium (SchilI & Jakel 1978; Venter et al.
1990 a, b). Schick (1982) introduced the term scutellum for the solid part of the
adhesive disc. According to Arekal and Ramakrishna (1980), translators indicate
a continued evolution from the Apocynoideae to Asclepiadoideae through the
Periplocoideae. Kunze (1993), consider an adhesive disc together with the base
of the stalk as the central element of the Periplocoideae translator.
The Apocynaceae s.s. is distinguished from the Periplocoideae and the
Asclepiadoideae by its single-grained pollen and absence of translators.
However, Apocynum L. (Apocynoideae) have band-like translators, which are
homologous to those of the Periplocoideae and thus indicate a connection
between Periplocoideae and Apocynaceae. Apocynum is the only genus in the
Apocynoideae where pollen is released as tetrads (Nilsson et al. 1993).
37
4.2 RESULTS:
Pollen grains are united in calymmate tetrads. The size of the grain with the
mean average is represented in Table 4.1 and Figure 4. The pollen grains are
arranged decussately (Fig. 4.1 A - N), rhomboidally (Fig. 4.2 A - D) or
tetrahedrally (Fig. 4.3 A - C), the most common arrangement being decussate.
The smallest mean decussate tetrad size was observed in Batesanthus
parviflorus (38.5 urn), Sacleuxia newii (38.9 urn) and Mangenotia eburnea (39
urn). The largest mean decussate tetrad was observed in Baseonema gregorii
(60 urn) and Mondia whitei (55.7 urn).
Pores: (Fig. 4.4 E, H). There are 4-6 pores per grain in all the species
examined. The exception is Baseonema gregorii where 8-10 pores have been
observed. The pores are round to oval and are situated at the junction area of
adjacent grains. The pore edge is often uneven and beset with granules, while
the pores themselves are sometimes covered with a thin layer of exine material
(Fig. 4.1 M, N; 4.2 B, C). The pores range in size from 1.3 - 6.3 urn.
In all the genera, the exine is smooth, and consists of a distal stratum (tectum)
subtended by a granular stratum. The granular stratum consists of unequal sized
granules (Fig. 4.4 A, B, D, G). The intine is well developed (Fig. 4.4 A, C, F).
The inner walls separating the individual grains of the tetrad have the same
structure as the exterior wall, consisting of tectum, granular stratum and intine.
The walls are, however, not continuous, but interrupted by wall bridges consisting
of intine and a granular stratum (Fig. 4.4 C, F).
Translators: There are five translators per flower positioned between the
anthers, with the stalk fitting in a groove on the stigmatic head. The translator
consists of a spoon, stalk and adhesive disc (Fig. 4.5 A - P). The stalk forms the
connection between the spoon and adhesive disc. The spoon is the uppermost
part of the translator and the pollen carrying part. It has an adhesive surface,
which carries the pollen tetrads. In the investigated species there is not a clear
distinction between spoon and stalk. There is a gradual transition from spoon to
38
stalk, making demarcation between spoon and the stalk difficult (Fig. 4.5 G, 0,
P). The stalk measurements were taken from the base of the spoon (where it
becomes narrow) to the upper end of adhesive disc. In most cases there is a
median partition. The spoon varies from elliptic-linear in Mangenotia eburnea
(Fig. 4.5 G), broadly elliptic in Zacateza pedicellata (Fig. 4.5 0, P) and ovate to
broadly ovate in the rest of the species. The adhesive disc is positioned more or
less at right angles to the base of the stalk or in an oblique position (Fig. 4.5 N).
The size of the translators is presented in Table 4.2.
65
E 60 -
2:
en I
Cl
<{
a::: 55 -
lw- t
I-
W
I- 50 - I
<{
eenn
o::::>w 45 - I
Cl
LL.
0 I I I
w
N 40 -
(i) I I I
35 ]/ V
01 1I I I I I I I "I "I
:ë:; '" '" :s III0 '"" 2
~ '~ ~
bl S 0 ~
'"" s0 Q) ~E
I!! ~ E" ~
Q) ~
t:: Q)
0 lJ
bl .S ~ " .QQ") lJ c: (Ij Ï3
.; ...; III ~ <ii Q) Q)Cl.
III III
Cl. III c: c: ~ Cl.
III III <ii ...;III ~ N
III III ~
SPECIES
Figure 4: Differences in pollen range (size) of the Periplocoideae species.
39
TABLE 4.1 - DIAMETER OF POLLEN TETRADS (urn}
TAXON COLLECTOR(S) & NUMBER RANGE - DECCUSATE TETRADS Average + Std.Dev.
Baseonema gregorii Field & Powys 174 [55.92 (51.15 - 60.45) + 2.59] x [56.36 (52.08 - 60.45) + 2.15]
Baseonema gregorii Faden, R.B. & A.J. 74/436 [69.69 (66.03 - 71.61) + 1.55] x [69.63 (66.96 - 72.54) + 1.61]
Baseonema gregorii Verdcourt & PoihilI 2695 [53.94 (47.43 - 64.17) + 3.58] x [54.18 (51.15 - 63.24) + 3.01]
[59.85 (47.43 - 71.61) + 2.57] x [60.06 (51.15 - 72.54) + 2.26] 59.96 + 2.42
Batesanthus intrusus Louis, J. 12992 [50.08 (46.50 - 56.73) + 3.11] x [49.10 (43.68 - 54.87) + 3.03]
B.intrusus [46.07 (42.78 - 49.29) + 1.82] x [45.87 (41.85 - 48.36) + 2.03]
[48.08 (42.78 - 56.73) + 2.47] x [47.48 (41.85 - 54.87) + 2.53] 47.78 + 2.50
Batesanthus parviflorus Deighton, F.C. 3265 [35.03 (32.55 - 39.06) + 2.01] x [35.59 (32.55 - 39.06) + 1.91]
B. parviflorus PoihilI, R.M. & Kirkup, O.W. 5190 [42.72 (39.99 - 44.64) + 1.94] x [41.17 (39.06 - 44.64) + 1.88]
B. parviflorus Gossweir, J. 8468 [38.32 (35.34 - 41.85) + 1.66] x [38.01 (35.34 - 41.85) + 1.92]
[38.69 (32.55 - 44.64) + 1.87] x [38.26 (32.55 - 44.64) + 1.90] 38.48 + 1.89
Batesanthus purpureus Bates, G.L. 383 [42.07 (38.13 - 47.43) + 1.80] x [42.35 (39.06 - 45.57) + 1.43]
B. purpureus Le Testu, M. 5493 [46.94 (41.85 - 54.87) + 2.51] x [46.16 (41.85 - 51.15) + 2.26]
[44.51 (38.13 - 54.87) + 2.16] x [44.26 (39.06 - 51.15) + 1.85] 44.39 + 2.01
Batesanthus ta/botii Talbot, TA 63 [41.51 (38.13 - 46.50) + 2.06] x [42.56 (38.13 - 48.36) + 2.03]
B. ta/botii Talbot, T.A 2021 [46.53 (42.78 - 53.94) + 2.47] x [46.72 (42.78 - 53.01) + 2.08]
[44.02 (38.13 - 53.94) + 2.27] x [44.64 (42.78 - 53.94) + 2.28] 44.33 + 2.28
Mangenotia eburnea Thomas, N.W. 4628 [41.85 (40.92 - 42.78) + 1.32] x [40.46 (39.06 - 41.85) + 1.97]
M. eburnea Adam, J. 30434 [36.72 (33.82 - 41.85) + 2.77] x [36.64 (32.55 - 44.64) + 3.65]
[39.29 (33.82 - 42.78) + 2.05] x [38.55 (32.55 - 44.64) + 2.81] 38.98 + 2.43
- - --------- ------ -----
.j>..
o
TABLE 4.1 - DIAMETER OF POLLEN TETRADS (urn)
TAXON COLLECTOR(S) & NUMBER RANGE - DECUSSATE TETRADS Average + Std. Dev.
Mondia ecomuta Allen, C.E.F. 139 [52.05 (48.36 - 55.80) + 1.85] x [50.97 (48.36 - 54.87) + 1.52]
M. ecomuta Faulkner, H.G. 558 [51.93 (48.36 - 55.80) + 1.99] x [51.90 (49.29 - 54.87) + 1.59]
[51.99 (48.36 - 55.80) + 1.92] x [51.44 (48.36 - 54.87) + 1.56] 51.72 +1.74
Mondia whitei Pienaar, B.J. 170 [51.68 (48.36 - 56.73) + 1.73] x [51.31 (54.87 - 64.17) + 2.00]
M. whitei Meedley Wood 6180 [59.71 (53.94 - 65.10) + 3.24] x [59.46 (54.87 - 64.17) + 2.93]
M. whitei Ward 3626 [56.61 (52.08 - 69.75) + 4.94] x [56.54 (51.15 - 69.75) + 4.76]
M. whitei Verhoeven & Venter 9329 [58.22 (52.08 - 64.17) + 3.49] x [59.21 (52.08 - 65.10) + 4.24]
M. whitei Scheepers 1058 [51.77 (43.71 - 60.45) + 4.87] x [50.99 (45.57 - 57.66) + 3.81]
M. whitei Leewenberg, A.J.M.10026 [53.44 (49.29 - 58.59) + 2.03] x [54.19 (51.15 - 55.80) + 2.03]
M. whitei Venter, H.J.T. 9282 [58.75 (53.01 - 64.17) + 3.83] x [57.97 (54.87 - 65.10) + 3.66]
[55.74 (43.71 - 69.75) + 3.48] x [55.67 (45.57 - 69.75) + 3.35] 55.71 + 3.42
Sac/euxia newii Verdcourt 246 [35.57 (35.34 - 41.85) + 1.87] x [36.83 (35.34 - 38.13) + 1.00]
S. newii Drummond & Hernsley 3364 [37.08 (35.34 - 39.06) + 1.10] x [37.45 (35.34 - 41.85) + 1.63]
S. newii Greenway & Kanuri 12852 [42.35 (37.20 - 47.43) + 2.24] x [41.91 (39.06 - 46.50) + 2.24]
[39.00 (35.34 - 47.43) + 1.62] x [38.73 (35.34 - 46.50) + 1.62] 38.87 + 1.77
Sac/euxia tuberosa NO POLLEN
Sarcorrhiza epip/7ytica NO POLLEN
Zacateza pedicellata Louis, J. 106 [43.56 (39.99 - 46.50) + 1.86] x [42.44 (37.20 - 48.36) + 2.11]
Z. pedicellata Muusa, J. 3488 [44.55 (39.06 - 47.43) + 2.12] x [43.93_(39.06 - 47.43) + 2.08]
[44.06 (39.06 - 47.43) + 1.99] x [38.73 (37.20 - 48.36) + 2.10] 43.63 + 2.05
"'" -
TABLE 4.2 - SIZE (l:!m} OF THE TRANSLATORS & SPOON SHAPE
SPECIES TOTAL LENGTH SPOON LENGTH SPOON WIDTH STALK LENGTH SPOON SHAPE
Baseonema gregorii 1678.88 922.63 550.55 756.25 Broadly - ovate
Baseonema gregorii 1881.55 918.09 592.9 947.83 Broadly - ovate
Baseonema gregorii 2109.03 1082.95 650.38 1026.08 Ovate I
Baseonema gregorii Ave.: 1889.82 974.56 597.94 910.05
Batesanthus intrusus 812.72 578.78 252.15 233.93 Ovate
Batesanthus intrusus 1119.25 756.25 338.8 332.75 Ovate to Angular ovate
Batesanthus intrusus Ave.: 965.99 667.52 295.46 283.34
Batesanthus parviflorus 595.93 396.28 257.13 199.65 Broadly ovate with + round apex
Batesanthus parviflorus 662.48 496.1 378.13 166.38 Broadly ovate with acute apex
Batesanthus parviflorus Ave.: 629.21 446.19 317.63 183.02
Batesanthus purpureus 1249.33 913.55 465.85 378.13 Broadly ovate to Angular ovate
Betesenttuis purpureus Broadly ovate
Batesanthus ta/botii 1288.65 681.63 369.05 583.83 Broadly ovate to Angular ovate I
Mangenotia eburnea Elliptic-linear
Mangenotia eburnea Elliptic-linear
Mondia ecornuta 2305.05 1839.2 1119.25 465.85 Ovate to broadly ovate
Mondia whitei 2994.75 847 2032.8 961.95 Broadly ovate
Mondia whitei 2964.5 411.4 1960.2 1004.3 Ovate
Mondia whitei 2432.1 490.05 1530.65 901.45 Ovate
Mondia whitei Ave.: 2797.12 582.82 1841.22 955.9
Sac/euxia newii 543.48 314.6 292.42 96.8 Broadly ovate to Angular ovate
Sacleuxia newii 496.6 290.4 293.93 116.46 Broadly ovate to Cordate
Sac/euxia newl! Ave.: 520.04 302.5 293.18 106.63
Sac/euxia luberosa Ovate
Sarcorhiza epiphylica Ovate
Zacaleza pedicel/ala 1220.08 445.68 Broadly elliptic
Zacaleza pedicel/ala 1382.43 438.63 Elliptic
.l>-
N Zacateza pedicellata Ave.: 1301.26 442.16
- ---- -----
A
Figure 4.1 Decussate pollen tetrads of Baseonema gregorii: (A) LM
micrograph. Field & Powys 174. (8) A group of tetrads showing different
arrangements (R = rhomboidal). Faden, R.B. & A.J. 74/436. (C) A grain showing
adjacent pores. Field & Powys 174. (D) A grain showing multi pores. Faden, R.B.
& A. J. 74/436. Scale bar: A = 50 urn: B-D = 10 pm
43
E
F
...Figure 4.1 Decussate pollen tetrads of Batesanthus sp.: (E) B. intrusus.
Adam, J.G. 30433. (F) B. purpureus. Le Testu, M.G. 5493. (G) B. parviflorus.
Po/hill & Kirkup 5190. (H) Grain showing irregular and rounded pores. B. telbo tii.
Ta/bot, T.A. 2021.
Scale bar: E-F = 50 urn: G-H = 10 urn
44
J
... Figure 4.1 Decussate pollen tetrads: (I) Mangenotia eburnea. Thomas,
N.W 4628. (J) Mondia whitei. Strey, R. G. 10347. (K) Grain with partially
covered pores. Mangenotia eburnea. Adam, J.G. 3043. (L) Mondia ecornuta.
Faulkner H.G. 558. (M) Grain showing partially covered pores. Mondia
ecornuta. Faulkner, H.G. 558. (N) Grain showing partially covered pores.
Zacateza pedicel/ata. Schweinfurlh, G. 3488.
Scale bar: I-J = 50 urn: K-N = 10 urn
45
Figure 4.2 Rhomboidal pollen tetrads: (A) Grain of Baseonema gregorii.
Field & Powys 174. (B) Grain with covered pores. Batesanthus intrusus. Louis,
J. 12992. (C) Grain with partially covered pores. Batesanthus purpureus.
Faulkner, H. G. 558. (0) Grain of Zacateza pedicel/ata. Schweinfurth, G. 3488.
Scale bar: A-D = 10 urn
46
Figure 4.3 Tetrahedral pollen tetrads: (A) Grain of Baseonema gregorii.
veracourt & Po/hill 2695. (B) Grain of Mondia ecomuta. Fau/kner, H.G. 558.
(C) Grain of Zacateza pedidellata. Louis, J. 106.
Scale bar: A-D = 10 urn
47
Figure 4.4 TEM SECTIONS OF SELECTED PERIPLOCOID POLLEN
GRAINS: (A) Outer wall of Batesanthus parviflorus. Po/hill & Kirkup 5190. (B)
Outer wall of B. parviflorus showing granulose substances of the surface.
Oeighton, F.C. 3265. (C) Inner walls of Mondia ecornuta. Fau/kner, H.G. 558.
(0 Outer wall of M. ecornuta. Fau/kner, H.G. 558.
i = intine, g = granular. t = tectum
48
... Figure 4.4 TEM SECTIONS OF SELECTED PERIPLOCOID POLLEN
GRAINS: (E) A grain showing position of a pore in Sacleuxia newii. Verdcourt
246. (F) Inner walls of Zacateza pedicellata. Louis, J. 106. (G) Outer wall of Z.
pedicellata. Muusa, J. 3488. (H) A grain showing position of a pore in Z.
pedicellata. Louis, J. 106.
i =intine, 9 = granular, t= tectum; t = pore
49
Figure 4.5 TRANSLATORS OF SELECTED PERIPLOCOID SPECIES: (A) Ovate spoon of
Baseonema gregorii. Verdcourt & Po/hill 2695. (B) Ovate spoon of B. gregorii. Verdcourt & Po/hil/
2695. (C) Broadly ovate spoon of Batesanthus intrusus. Tisserant 1480. (D) Broadly ovate spoon
of Batesathus parvif/orus. Po/hill & Kirkup 5190. (E) Broadly ovate spoon of B. parvif/orus. Po/hill
& Kirkup 5190. (F) Broadly ovate spoon of Batesanthus ta/botii. Ta/bot 2021.
Scale bar = 100 urn
50
... Figure 4.5 TRANSLATORS OF SELECTED PERIPLOCOID SPECIES: (G)
Elliptic-linear spoon of Mangenotia eburnea. Leeuwenberg 8083. (H) Ovate
spoon of Mondia ecornuta. Faulkner, H.G. 558. (I) Broadly ovate spoon of
Mondia whitei. Strey, R.G. 10347. (J) Ovate spoon of M. whitei. Strey, R.G.
10347. (K) Angular ovate spoon of Sacleuxia newii. Verdeourl 246. (L) Broadly
ovate spoon of S. newii. Greenway & Kanuri 12852.
Scale bar = 100 urn
.\I\.\01E.t 5 I
t
... Figure 4.5 TRANSLATORS OF SELECTED PERIPLOCOID SPECIES: (M)
Ovate spoon of Sacleuxia tuberosa. Tanner, R.E.S. 360. (N) Ovate spoon of
Sarcorrhiza epiphytica. Semsei 2957. (0) Broadly elliptic spoon of Zacateza
pedicel/ata. Louis, J. 106. (P) Broadly elliptic spoon of Z. pedicel/ata. Louis, J.
106.
Scale bar:: 100 urn
52
4.3 DISCUSSION AND CONCLUSIONS:
Palynological characters, namely pollen morphology (size, shape/arrangement
and number of pores), the translator (shape) and the structure of the exine have
been looked into to establish their taxonomic value.
Pollen morphology is uniform within the subfamily although some variations or
exceptions do occur. Raphionacme Harv. is the one genus that can be
distinguished by the 8-16 pores per pollen gain (SchilI & Jakel 1978, Verhoeven
& Venter 1988) as against 4-6 in the other genera. So far, two genera namely
Schlechterella K. Schum. (Verhoeven & Venter (1998b) and Raphionacme
(Verhoeven & Venter 1988, Nilsson et al. 1993) are known to have multiporate
pollen grains. In this study, it was observed that Baseonema has 8-10 pores per
grain and thus can be distinguished from other genera. Having numerous pores
could be regarded as more advanced than genera with 4-6 pores per grain
(Nilsson et al. 1993). Otherwise, genera with pollinia are regarded as more
advanced than other taxa within the Periplocoideae.
The decussate tetrad size of tetrads varies within the species investigated. The
decussate tetrad size of Baseonema gregorii and Mondia whitei may be used to
distinguish them from the other investigated species. To some extent, the size of
the tetrad may be used to distinguish species that are easily confused, e.g.
Baseonema and Batesanthus. Size of the tetrads has been used by Verhoeven
& Venter (1994) to delimit between species of Periploca L.
Three common arrangements of tetrads were recognized, those are decussate,
rhomboidal and tetrahedral grains. In one genus you may find all three kinds of
arrangement.
In the representatives of the Periplocoideae studied so far, the exine structure
consisting of a solid stratum (tectum), shows little variation (Verhoeven & Venter
1988, 1993, 1994a, b). Camptocarpus Decne. differs from other Periplocoideae
53
genera in that the exine consists of a tectum, granular stratum and foot layer
(Verhoeven & Venter 1994b). The exine structure of the five genera investigated
in this study appears to be without variation.
Although small differences may occur, the pollen morphology of the taxa
investigated is rather uniform and thus of little value in the distinction of the
species and genera investigated.
The unique tri-partite translator of the Periplocoideae consists of the spoon, stalk
and an adhesive disc. The adhesive spoon, which carries the pollen tetrads,
varies in shape and size between genera. Arekal & Ramakrishna (1980) refer to
the shovel (spoon) shaped with a median partition as the most specialized. The
spoon of Baseonema gregorii, Batesanthus intrusus. B. parviflorus, B. talbotii,
Mondia whitei and Sacleuxia tuberosa also have a median partition. The same
structural type has been observed in Tacazzea apiculata Oliv., T. rosmarinifolia
(Decne.) N.E. Br. and T. venosa Decne. (Venter et al. 1990a), Hemidesmus
indicus (L.) R. Br. and Oecalepis hamiltonii Wt. & Arn. (Arekal & Ramakrishna
1980), Raphionacme dyeri Retief & Venter (Verhoeven & Venter 1997).
Translators where the spoon gradually narrows toward the base thus making
demarcation between the spoon and the stalk difficult have been observed. This
feature is also referred to as an extended spoon. This kind of translator has been
found in Raphionacme caerulea E.A. Bruce, R. keayii Bullock, R. lanceolata
Schinz., R. longituba E.A. Bruce, R. madiensis S. Moore and R. michelli De
Wilde (Verhoeven & Venter 1997). Also in Cryptolepis oblongifolia (Meisn.)
Schltr. (Venter & Verhoeven 1997) and C. yemenensis Venter & Rl.Verh.
(Venter & Verhoeven 1999). In the Apocynaceae s.l. a few genera are
characterised by the presence of a primitive translator development (Verhoeven
& Venter 1997).
This study shows that translators have variable spoon-shape types and therefore
of taxonomic value, especially at generic level.
54
SEED SURFACE MORPHOLOGY
5.1 INTRODUCTION:
The Asclepiadaceae (incl. Periplocaceae) and Apocynaceae s.s. are derived
from schizocarpous gynoecia with ovaries united only by their styles or
stigmas and the seeds do not remain attached along a marginal placenta
(Spjut 1994). The gynoecium consists of two free semi-inferior ovaries with
numerous ovules. The two styles are free below, but fused towards the apex,
and dilate into a stigmatic head. The ovules are more or less anatropous,
unitemic (i.e. uni-integumented and not provided with a tegmen) and
tenuinucellate (Corner 1976). Frequently, only one fruitlet of a schizocarpous
follicular gynoecium matures (Cronquist 1981), in which case the fruit is a
follicle. The fruit consists of a paired or single follicle(s). The shape and
texture of the follicles may be fairly constant within a genus, as in Cryptolepis
R. Br., or vary greatly as in Tacazzea Decne. and Raphionacme Harv. (Venter
& Verhoeven 1997). According to Dave & Kuriachen (1991), anatomical
features of the follicles may be significant for higher level distinction.
The Periplocoideae seeds are usually narrowly elliptical in outline, numerous,
flattened or compressed and characteristically dispersed by wind. Attached at
the rear micropylar end is a coma of hairs. Although rare, variations do occur,
for instance, there is no obvious coma at the micropylar end, instead seeds
have a dense fringe of hairs around the entire margin.
The possibility that seed epidermal surface varies between genera has been
taken into consideration in this study. Little has been done on the seed
surface morphology of the Periplocoideae, except for "a note on
Raphionacme namibiana" (Bruyns 1994) and "A new species of Cryptolepis
(Periplocoideae, Apocynaceae) from Arabia" (Venter & Verhoeven 1999).
55
5.2 RESULTS:
There is no specific classification system used but terminology used in this
chapter is based on comparison with the works of: Demissew & Harley
(1992), Venter (2000), Venter & Verhoeven (1999) and Venter et al. (1990).
The fruit is a paired follicle drying brown-black and the surface beset with
longitudinal fissures. The seeds are brown-black and numerous. The shape
is elliptic to obovate [Table 5.1], flattened and/or compressed with a raised
longitudinal fissure along the center that is referred to as a funicular or raphe-
groove in all taxa investigated except for Batesanthus intrusus. The tufted
coma of cream-white to brownish hairs is generally attached at the micropylar
end in all species under investigation, except for Batesanthus intrusus with
hairs extending around the entire margin. The coma spreads out when dry
and exposed to air (Fig. 5.6 D); thus showing that the seeds are wind
dispersed.
Baseonema gregorii: The seed surface is somewhat rugged without an
obvious reticulate pattern (Fig. 5.1 A). Epidermal cells are slightly elongated
and rectangular-like (Fig. 5.1 B), they are bordered by raised ridges along the
margins. Seeds (Fig. 5.1 C) are elliptic to obovate and measure 10 x 3 mm
whilst the length of the coma ranges from 8-12 mm.
Batesanthus intrusus: Seeds are hairy, however, when hairs are removed,
the seed surface is smooth and reticulated (Fig. 5.2 A). Epidermal cells in
surface view are polygonal with about six sided cells. Seeds (Fig. 5.2 B) are
elliptic, ranging from 9-12 x 3-4 mm in size and the length of the coma
ranges from 47-50 mm in length. The coma of hairs extends around the
entire margin instead of the usual micropylar coma. Hairs (Fig. 5.2 C) are
attached from the bottom rear end and form a dense mat covering the whole
one surface (adaxial) of the seed. The other (abaxial) surface is devoid of
hairs.
Mangenotia eburnea: Seed surface has a pitted epidermis (pores) with fine
parallel running striations. There are about 3-5 pores at the edges of
individual cells. Epidermal cells are almost polygonal, isodiametric in various
56
directions and irregularly arranged. Anticlinal epidermal walls are mostly
sinuate (Fig. 5.3 A, B). Seeds (Fig. 5.3 C) are elliptic, ranging from 7-8 x 3
mm in size and the length of the coma ranges from 43-45 mm. The coma at
the micropylar end, when compared to other species in this study is relatively
long (Table 5.1).
Table 5.1: Seed Epidermal Characteristics.
TAXON SEED SEED SIZE LENGTH EPIDERMAL TESTA
SHAPE (mm) OF COMA CELLS (Surface)(mm) (Shape)
Elliptic Elongated &
B. gregorii Obovate 10 x 3 8-12 rectangular Rugged
Polygonal,
B. intrusus Elliptic 9-12 x 3-4 47-50 4-6 sided Hairy
Pitted with
M. eburnea Elliptic 7-8 x 3 43-45 Polygonal parallel
striations
Elliptic Polygonal to
M. whitei Obovate 7-9 x 2-3 23-34 elongated Smooth
Polygonal, Slightly
S. newii Elliptic 7x2 29-34 4-6 sided rough
S. tuberosa Elliptic 6-7 x 2 9-13 Polygonal Smooth
Irregularly
Z. pedicel/ata Elliptic 9 x 2-3 20-23 Polygonal Smooth
Mondia whitei: The seed surface is smooth. Epidermal cells are polygonal,
sometimes slightly elongated, with straight to slightly undulate walls (Fig. 5.4
A, B). The seeds (Fig. 5.4 C) are elliptic to obovate, ranging from 7-9 x 2-3
mm in size. The length of the coma ranges from 23-38 mm. The coma is
attached to the micropylar end of the seed.
Sacleuxia newii: The seed surface is slightly rough with small granular
structures (Fig. 5.5 A). The epidermal cells are 4-6 sided (Fig. 5.5 B). The
seeds (Fig. 5.5 C) are narrowly elliptic and range at 7 x 2 mm in size, with the
length of the coma ranging from 23-29 mm.
Sacleuxia tuberosa: The seed surface is smooth. Epidermal cells are
polygonal and elongated to rectangular (Fig. 5.6 A, B). The seeds (Fig. 5.6 C)
57
are elliptic, ranging from 6-7 x 2 mm in size and the length of the coma
ranges from 9-13 mm. When dry (Fig. 5.6 D), the coma of hairs are spread
out.
Zacateza pedicel/ata: The seed surface is smooth. Epidermal cells are
irregularly shaped with heavy ridges along anticlinal walls (Fig. 5.7 A). The
seeds (Fig. 5.7 B, C) are elliptic and they range from 9 x 2-3 mm in size. The
length of the coma ranges from 20-23 mm. The funicular or raphe-groove
(Fig. 5.7 C) at the center is distinct in this species.
c
Figure 5.1 Seed surface of Baseonema gregor;;: (A) rugged epidermal
surface. (B) rectangular-like epidermal cells. (C) seeds with coma of hairs.
All from van Someren 184 (K). Scale bar: A, B = 10 pm
58
Figure 5.2 Seed surface of Batesanthus intrusus: (A) smooth surface with
polygonal epidermal cells. (8 - C) elliptic seed(s) with hairs extending around the
entire margin covering the whole one surface. All from Klaine, R.P. 513 (P).
Scale bar: A = 10 urn
59
c
Figure 5.3 Seed surface of Mangenotia eburnea: (A-B) pitted surface,
irregularly arranged epidermal cells with sinuate anticlinal walls. (C) seeds
with coma of hairs. All from Thomas, N.W 4544 (K).
Scale bar: A, B = 10 pm
60
c
Figure 5.4 Seed surface of Mondia whitei (A-B) smooth surface with
polygonal to slightly elongated epidermal cells. (C) seeds having coma of
hairs broken-off. All from Oldeman, R.A.A. 392 (MO).
Scale bar: A, B = 10 urn
61
c
Figure 5.5 Seed surface of Sac/euxia newii: (A) rough surface beset
with grain-like substances. (8) 4-6 sided epidermal cells. (C) Seeds, with
hairs at the centre. All from Verdeourl 246 (K).
Scale bar: A, 8 = 10 pm
62
c
Figure 5.6 Seed surface of Sac/euxia tuberosa: smooth surface with
polygonal cells elongated to rectangular. (8) straight to slightly undulated anticlinal
walls. (C) seeds. (0) dry seed showing spread-out hairs.
All from Kerfoot, 0. 3596 (K). Scale bar: A, 8 = 10 urn
63
B
Figure 5.7 Seed surface of Zacateza pedicel/ata: (A) smooth surfaced
epidermal cells. (8) elliptic seed. (C) seed showing funicular (").
All from Evrard 3467 (K).
Scale bar: A = 10 urn
64
5.3 DISCUSSION AND CONCLUSIONS:
Seed surface morphology within the Periplocoideae (Apocynaceae) is a
taxonomic feature that has not been extensively studied. However, due to
scarcity of seeds of selected genera, it has not been easy to find a trend to
ascertain the use of this character state in classification systems.
According to Barthlott (1981), structures of particular importance in the
description of the seed coat characteristics are the relief of the anticlinal cell
wall as well at curvature of the outer periclinal walls. He further states that the
most prominent feature is surface sculpturing, especially the outline of the
cells.
Three types of seed surface sculpturing have been observed:
1) Smooth to reticulate surface and polygonal epidermal cells with deep or
shallow grooves between anticlinal walls as in Batesanthus intrusus,
Mondia whitei, Sacleuxia newii, Sacleuxia tuberosa and Zacateza
pedicel/ata. A feature also observed in Cryptolepis yemenensis Venter &
R.L. Verh., Petopentia natalensis (Schltr.) Bullock, Tacazzea apiculata
D.Oliv. and Tacazzea venosa Decne. Venter et al. (1990a, c).
2) Pitted surface and polygonal cells with pores lying at the edges of
individual cells as in Mangenotia ebumea.
3) A rugged surface without an obvious reticulate pattern and polygonal
epidermal cells with raised ridges over anticlinal walls as in Baseonema
gregorii.
The micropylar coma of hairs is characteristic and uniform, but there occurs
variation in length of the coma whereby Mangenotia ebumea has a long coma
compared to other taxa investigated. However, hairs extending around the
entire margin. have been observed in Batesanthus intrusus, a feature not
common, yet not unique since it is known to occur also in seeds of
Raphionacme namibiana Venter & R.L. Verh. (Periplocoideae), Fockea
sinuata (E. Mey.) Druce (Asclepiadoideae - Marsdenieae) (Bruyns 1994) and
Finlaysonia Wall. (Periplocoideae) (Endress & Bruyns 2000).
65
TAXONOMY OF BASEONEMA schttr. & Rend~e
6.1 HISTORICAL BACKGROUND:
An East African genus: Baseonema Schltr. & Rendie was first established by
Schlechter & Rendie in 1896 to accommodate their new species Baseonema
gregorii Schltr. & Rendie. They based their description on a single collection
from Kenya, collected between 1892-1893.
Baseonema is a Greek derived generic name that implies 'base thread-like';
=nerna is a Greek word that means 'thread'. The specific epithet - gregorii
(Rec. 73C) is a commemorative name in honour of the first collector Dr. J.W.
Gregory.
Baseonema gregorii is a shrub or small liana with climber stem twining. It is
characterized by having interpetiolar stipules, hairy velutinous leaves, red to
purple inflorescence stalks and shiny, green reflexed corolla lobes. Bullock
(1954a) states that 'the type specimen has no twining shoots whereas in a
specimen collected by Kassner at Kibwezi, Kenya on 27-04-1902 the shoots
are evidently twining, although habit variability might be due to the immediate
conditions of the environment and that the same plant may be a twiner during
some years and a non-twiner during others'.
Baseonema gregorii is a rare species (Bullock 1954a), although it is evidently
not yet extinct due to a recent collection of 11-02-1996 from Kenya - 02S
37E BB by Goyder et al. 4006 (PRE).
6.1.1 GENERIC DELIMITATION:
Baseonema is a monotypic genus from East and Equatorial Africa. It has
been separated from the then congeneric genus Baroniella Costantin &
66
Gallaud with seven species that are all endemic to the Madaqascan rainforest
(Klackenberg 1997).
According to Klackenberg (1997), Baseonema and Baroniella are ought to be
kept separate due primarily to different flower morphology. B. gregorii is
characterized by its almost ex-appendiculate anthers and a whorl of small
corolline appendages - that is, inner corona lobes alternating with the
stamens (Schlechter & Rendie 1896), although stamens arise down the
corolla-tube. Corolline coronas are situated primarily in the petal sinuses
(Liede & Kunze 1993). In B. gregorii the staminal filaments are much dilated
at their bases, seemingly situated between the interstaminal discs. These
interstaminal lobes are interpreted as inner corona lobes of corolline origin
(Klackenberg 1997). Kunze (1990) describes these interstaminal discs which
are sometimes mere spots or 'cups' as being nectariferous in nature.
Although inconspicuous, the corolla tube instead of the corolla lobes is the
one that reflexes back at anthesis, eventually exposing the gynostegium (Fig.
6.1 C), a character unknown in Baroniella. Choux (1913) had placed under
Baseonema - B. acuminatum, B. linearis and B. multiflorum of which were
later transferred by Bullock (1955) and revised by Klackenberg (1997) to the
genus Baroniella due to different flower form, habit and ecology. Secondary
veins unite in a crenate pattern in the leaf lamina whereas those of Baroniella
species, if present are straight and parallel and united to a straight sub-
marginal vein.
Pollen morpholoqy' can also be used to distinguish between species of
Baseonema and Batesanthus N.E. Br. Pollen grain size in Baseonema are
relatively larger compared to the smaller grains of Bafesanthus species.
e • Refer to Chapter 4 for review on pollen morphology
6.2 TAXONOMY:
BASEONEMA Schltr. & Rendie in Journal of Botany 34: 97, t. 356 (1896).
TYPE: Baseonema gregorii Schltr. & Rendie
67
Twining shrub. Leaves hairy, orbicular, entire and ciliate. Petioles purplish,
stipules interpetiolar. Inflorescence indeterminately cymose. Flowers 5-
merous. Corolla hairy outside, glabrous within, green, lime-green to yellowish-
green, tube reflexed. Outer corona annular; inner corona lobes lobular.
Filaments with broad bases. Species 1, East Equatorial Africa
Baseonema gregorii Schltr. & Rendie in Journal of Botany 34: 97, t. 356
(1896); K. Schum.: 286 (1897); N.E. Br.: 259 (1902); Thonner: 442 (1915);
Bullock: 59 (1954a); Agnew: 369 (1974).
TYPE: Kenya, K4 Machakos District, Kenani & Ongalea Mts. 1892-1893,
Gregory, J.W. 14 (BM, holotype!).
Twiner with white, milky latex. Stems herbaceous to woody, brownish and
hollow; old stems sparsely pubescent, young stems pubescent. leaves
opposite, simple, with interpetiolar stipules; petiole (4-)7-1 0(-26) mm long,
densely pubescent, purple; lamina (20-)24-60(-97) x (13-)18-63(-93) mm,
broadly obovate to broadly elliptic, apex cuspidate, acuminate to acute, base
rounded or cuneate, velutinous, dark green above, pale green to whitish
below, hypostomatous, venation brochidodromous - evidently so abaxial,
margin entire, undulate, ciliate. Inflorescences lax cymose panicles; primary
peduncles (10-)22-29(-70) mm long, puberulous, purple green to shiny red;
secondary peduncles (10-)22-27(-54) mm long, glabrous or nearly so;
pedicels (5-)11-16(-35) mm long, puberulous, purple. Flower 5-merous,
bisexual, actinomorphic, complete, gynostegium exposed. Calyx of 5 sepals,
1-2 x 0.5-1.5 mm, free, elliptic to ovate, apex acute, villose, green. Corolla
of 5 petals; lobes (4-)5-6(-8) x 1-2.5 mm,free towards apex, ovate, elliptic to
narrowly ovate, apex acute to acuminate, villous outside, glabrous within,
reflexed at anthesis, green, lime green or green yellow; tube: lower tube very
short .± 2 mm long, reflexed; upper tube absent. Corona: outer corona from
apex of corolla tube, lobes almost triangular, apex acute; inner corona of
interstaminal discs, arising in-between the stamens. Androectum; stamens
5, adnate to outer corona lobes and interstaminal discs, positioned from just
below outer corona; anthers basifixed, ovate, apex sub-acute, brownish with
68
whitish membranous flaps (microsporangium), fused to and connivent above
stigma head, connective prominent; filaments free, basely with laterally dilated
bases; pollen tetrads adhere to translator surface, most common arrangement
decussate, otherwise rhomboidal or tetrahedral. GYD1loecium: ovaries free, 2,
semi-globose, semi-inferior, ovules numerous; styles 2, fused terminally;
stigmatic head pentangular, deltoid/ovoid, apex almost conical; translators 5,
from grooves in upper surface of stigmatic head, spoon spathulate with
receptacle ovoid, stalk linear, viscidium sub-circular, sticky. Fruit of paired
follicles; follic/es 52-95 x 7-11 mm, linear-oblong, puberulous to sub-
glabrous; fruit stalk 68-92 mm long, puberulous. Seeds 10-11 x 2 mm,
compressed with a raised horizontal fissure towards centre; coma of hairs 24-
27 mm long. (Figure 6.1 ).
6.3 DISTRIBUTION AND ECOLOGY:
Baseonema gregorii is limited in distribution to the East Africa regions of
Kenya and Tanzania. It is found between the lines of longitude 01°30' -
04°16' S and latitude 37°21' - 39°19' E (Figure 6.2).
Flowering plant specimens were seen from January to April and July, fruits in
March to May (Table 6.1).
Table 6.1 Flowering and fruiting periods of Baseonema gregorii
SEP aCT NOV DEC
It seems to grow at a wide range of altitude, from low-lying areas of 600 m to
higher ones at 1,500 m above sea level.
It is known locally by the Kamba - Kenya tribe as "Muongwa".
In Kenya, it has been found in savanna in association with species of Acacia
Mill., Commiphora Jacq. and Sansevieria Thunb. In the dry evergreen forest
69
dominated by species of Euphorbia L., Ehretia P. Browne, Manilkara Adans.,
Suregada Roxb. ex Rottler and Strychnos L., it occurs on very steep slopes
with poor ground cover. It was also found on sandy soil near rocky outcrops
together with species of Grewia L., Euphorbia, Thunbergia Retz., Adansonia
L., Boswellia Roxb. ex Colebr., Tamarindus L. and Bauhinia L.
6.3.1 VOUCHER SPECIMENS:
KENYA: 02S 38E: 240 km from Mombasa on Nairobi road (-CA).
Verdcourt & PoihilI 2695 (K).
02S 38E: Tsavo National Park, East Simba Hill (-BC). Faden,
R.B. & A.J. 74/436 (PRE).
03S 38E: Mt. Kasigau, 4 km S of Rukanga (-DC). Gilbert, M.G.
& C.I. 6122 (K).
TANZANIA: 03S 37E: T3 Pare District, Lembeni (-DC). Bally, PR.O. 8125
(K).
70
/
Figure 6.1 Baseonema gregorii: (A) Habit showing twining stems, leaves and
flowers. (8) Open flower showing corolla lobes (c), pistil (p) at the centre, stamen (5)
and inner corona (i). (C) Complete flower showing reflexed corolla lobes and
exposed gynostegium. Drawn from Field & Powys 174 (K).
71
o
10
? t. .;\..,-_,.,\'..:.:....... I.f'\)' ~v-, _ •••••-. __ ••• ) /l:.·, .... ···I. ., ::: \
II \....\ \ \,.. ,.....\
20 310
Figure 6.2 Known geographical distributionof Baseonema gregorii.
72
TAXONOMY OIF BATESANTHUS N.IE. a-,
7.1 HISTORiCALBACKGROUND:
Batesanthus N.E. Br. as currently described, comprises of four African
species and has been maintained as a genus separate from Baseonerna
Schltr. & Rendie.
Batesanthus commemorates Mr. G.L. Bates who first collected the type
specimen Batesanthus purpureus N.E. Br. in 1895 from Cameroon.
There are four species of Batesanthus N.E. Br. namely:
1. Bafesanthus intrusus N.E. Br.
2. Batesanthus parviflorus Norman
3. Batesanthus purpureus N.E. Br.
4. Batesanthus ta/botii S. Moore
However, of all the four species, Bullock (1961) in his notes on African
Asclepiadaceae believed that there is no combination of characters that can
be used to delimit between species and published the following synonyms of
B. purpureus N.E. Br.:
Perithrix glabra Pierre in Bull. Soc. Linn. Paris, nov. Sér. No. 8 (1898).
Batesanthus ta/botii S. Moore in Cat. Talb. Nig. PI.: 63 (1913).
Batesanthus ta/botii var. grandifolia S. Moore, l.c. 64 (1913).
Batesanthus g/aber Schltr. In Mildbr. Deutschen Zentral-Afrika
Expedition 1907 -1908 2: 541 (1914).
Batesanthus intrusus S. Moore in Journ. Bot. 58: 267 (1920).
Batesanthus mildbraedi Schltr. In Mildbr. , Wiss. Ergebn. Deutsch.
Zent.-Afr. Exped. 1910- 1911, 2: 80 (1922).
Batesanthus telbotli var. parviflora S. Moore, nomen in herb. Kew;
Hutch. & Dalziel, l.c. in syn (1931).
73
Nevertheless, the present study does not agree with Bullock's synonymy.
Species of Batesanthus are sparsely distributed in Central and more so in the
Western Tropical Africa. The genus's poor representation and thus its sparse
collection may be attributed to its restricted distribution. However, slow
accumulation of material enabled Brown to reduce the inadequately described
Perithrix Pierre, hence now an illegitimate synonym.
The Greek derived name 'intrusus' = thrust in \ inserted refers to the
intrusion of the corolla base, especially in bud, Bullock (1961) dispute
that the so-called intrusion is entirely due to the fact that the buds examined
are at a very early stage of development. In B. intrusus, it should be noted
that, when dried, the flowers appear to be mostly unopened buds, this is due
to the short time the corolla remain expanded (Moore 1920).
B. parviflarus is a Greek derived name referring to the small, profusely
flowering flowers, that is, 'parvis' = small and 'florus' = flower.
B. purpureus, 'purpureus' is a Greek derived name meaning purple,
B. ta/batii commemorates Mr. T.A. Talbot who first collected the type
specimen from Oban in Nigeria.
Brown (1896) and Venter & Verhoeven (1997) placed Batesanfhus (=
Perithrix) under the tribe Periploceae, but Endress & Bruyns (2000) have
excluded Batesanthus from their list of genera under Periplocoideae.
7.2 TAXONOMY:
BATESANTHUS N.E. Br. in Hooker's Icones Plantarum: 25, t. 2500 (1896);
K. Schum.: 286, (2-4) (1897); N.E. Br.: 4(1), 253 (1902); Thonner: 443
(1915); Hutch. & Dalziel: 2(1), 50 (1931); Bullock: 15, 203-204 (1961);
Bullock: 2, 82 (1963a).
TYPE: Batesanthus purpureus N.E. Br.
Shrubs. Stems twining, sometimes erect or climbers. Latex milky-white to
absent. Leaves opposite, large/ample, petiolate, stipulate, stipular fringes
74
interpetiolar, reflexed, amphistomatous; lamina: abaxial surface pale-green to
somewhat whitish, glabrous, adaxial surface dark-green, glabrous to
puberuient on midrib; apex acuminate to acute, base cordate, margin entire to
slightly repand. Inflorescences lax, axillary cymes. Flowers 5-merous.
Calyx free. Corolla rotate, usually glabrous, upper tube absent, lower tube
inconspicuous. Corona 5 lobular, outer corona lobes absent to filiform, inner
corona adnate to filament bases, annular. Androeclum: stamens 5, anthers
white to brownish, broadly ovate to oblong, erect, cohering terminally onto
stigmatic head, apex connate, glabrous; filaments c. 1 mm long, fused to
outer corona, fused sideways into an annulus outside interstaminal discs;
pollen in granular tetrads, porate. GynoecDlUlm: ovaries 2, ovoid, ovules
numerous; styles 2, becoming fused into one apically; stigmatic head
pentangular ovoid to deltoid, apex acuminate to conical; translators 5,
adhering to the grooves of stigmatic head, receptacle rhombic, sometimes
broadly ovate, cleft at center, stalk present, linear, viscidium rounded to half-
moon, sticky. Fruit paired follicles. Seeds comose. Species 4.
KEY TO THE SPECIES:
1a Leaves glabrous or nearly so. Corona lobes annular to long filiform lobes
.............................................................................................. 2
1b Leaves scabrid to hispid on both sides or only below. Corona lobes
annular, emarginate .4
2a Bottom of corolla folded collar-like around pedicel, i.e. intruse,
especially in bud; flowers purple t B. intrusus
21b Bottom of corolla normal, without an intrusion at the base, flowers dark
purple or dark reddish purple 3
3a Corolla lobes less than 10 mm long, corona with annular lobes to filiform
lobes. Climbing shrubs in savanna 2. B. parvif/orus
3b Corolla lobes (10-)15-20 mm long, without filiform corona lobes. Woody
climber in riverine forests 3. B. purpureus
40l Base of corolla not intruse 4. B. ta/botii
411:B>ase of corolla intruse 1. B. intrusus
75
1. Batesanthus intrusus S. Moore in Journal of Botany lviii: 269 (1920).
TYPE: Cameraan - Yaoundé, Bitye. 1917. Bates, G.L. 1392 (BM, holotype!).
== Perithrix glabra Pierre in Bulletin Society. Linnaeus.: 65 (1898)
TYPE: Gabon, Libreville on the Gabaan River. October 1896. Klaine, R.P.
513 (P, holotype!, K, isotype!).
Twining liana 4-5 m high. No latex. Stems herbaceous, purple-reddish or
brown, smooth, terete. leaves sparsely hairy to glabrous; interpetiolar
stipules frill-like; petiole (12-)21-32(-57) mm long; lamina (75-)120-140
(-190) x (25-)60--75(-90) mm long, ovate to lanceolate, secondary veins not
raised, reddish purple. Inflorescences .± 2 flowered; peduncle; primary
peduncle (20-)35-45(-120) mm long, secondary peduncle (25-)35-40(-70)
mm long; pedicel (6-)10-15(-25) mm long, minutely bracteate. Flowers:
blackish purple to violet inside, whitish to pale yellowish green outside. Calyx
2 x 1 mm, lanceolate, puberuient microscopic margins to glabrous,
segmented, purple to violet range. Corolla: lobes 3-11 x 1-3 mm, ovate,
apex acuminate to rounded, pale green outside, dark purple inside; lower tube
intruse at the base. Corona: inner corona awl shaped. Androecium:
stamens arise from mouth of corolla tube; anthers ovate .± 3 x 1 mm.
Gynoecium: ovaries 2, style short, c. 1 mm long. Fruit: follicles 112-120 x
3-4 mm elliptic to broadly elliptic, raphe at the center. Seeds 9-12 x 3-4 mm,
elliptic; coma of hairs 47-50 mm long, hairs extend all around one surface of
the seed. - see chapter 5. (Figure 7.1; 7.2).
2. Batesanthus parviflorus Norman in Journal of Botany lxvii Suppl. 2: 91
(1929).
TYPE: Angola, Guanza - Quilela Camabatela. 29-09-1922. Gossweiler, J.
8468 (BM, holotype!).
Shrub. Stems twining, dark purple. leaves glabrous; petiole (10--)17-20
(-30) mm long; lamina (62-)70--85(-115) x (30-)33-38(--60) mm, ovate to
lanceolate, veins reddish purple, interpetiolar stipules inconspicuous, dentate.
76
lntlorescences many flowered cymose panicle; peduncle (20-)23-38(-75)
mm; pedicel (5-)7-13(-15) mm long, moderately thick base bracteolate.
Flowers small, dark reddish purple. Calyx 2 x 1-1.5 mm, lanceolate, margin
ciliate. Corolla: lobes 4-5 x 1-2 mm, narrowly oblong, apex acuminate to
rounded. Corona sometimes double; outer corona filamentous (Plate 1 - after
page 88), lobes inserted at mouth of corolla tube, exposed at anthesis; inner
corona triangular. Gynoeciu...om:styles short, concealed. Fruit unknown.
(Figure 7.3; 7.4).
3. Batesanthus purpureus N.E. Br. in Hooker's Icones Plantarum PI. 25: t.
2500 (1896); F.T.A. 4(1): 253 (1902); Bullock: 15 (2),203-204 (1961).
TYPE: Cameroon, Efulen - Buie country. 20-09-1895. Bates, G.L. 383 (K,
holotype!).
Woody climber, sometimes twiner, glabrous. Stems covered with small warty
outgrowths, rusty red to purplish. leaves glabrous, stipular fringes
interpetiolar, reflexed, subpersistent; lamina (110-)145-150(-160) x (70-)
75-80(-95) mm, ovate to elliptic. lnflorescences lateral, lower part
dichotomous, higher up undivided, simple. Flowers: dark purple or purple.
Corolla: lobes broadly oblong to ovate, comparatively large at 4-6 x 8-11
mm, apex obtuse to rounded, deeply 5-lobed; lower tube .± 2 mm long.
Corona lobes indistinct; inner corona of 5 minute lobules, ovate, apex acute
to blunt. Androeclum: stamens inserted at mouth of corolla tube. Fruit:
follicles c. 120 mm long x 20 mm diameter, more or less woody, suberect,
shortly beaked. Seeds unknown. (Figure 7.5; 7.6).
4. Batesanthus ta/boW S. Moore in Catalogue of the plants collected by Mr.
& Mrs. P.A. Talbot in the Oban District, South Nigeria, British Museum,
Natural History: 36-64 (1913); Hutch. & Dalziel: 2(1), 52 (1931).
TYPE: Nigeria, Oban. 1912. Talbot, T.A. 2021 (BM, holotype!, K, isotype!).
= B. talbotii var. grandifolia S. Moore
TYPE: Nigeria, Oban. Talbot, T.A. 2021 (BM, holotype).
77
Climbing shrub. Stems brownish, warty, striated when dry. leaves
scabrous, interpetiolar stipules sharply toothed, reflexed; petiole (18-)20-25
(-35) mm long; lamina (120-)130-145(-155) x 70-105 mm, broadly obovate
to broadly elliptic, drying scabrid brown, with minute projections \ emergences
on the surface, more so on abaxial surface, lateral veins looped towards the
margin (i.e. brochidodromous), inserted upon the midrib nearly at right angle.
lntlorescences many-flowered; peduncles s, 30 mm long, glabrous; pedice/s
6-12 mm long, glabrous; bracts ovate, small. Flowers: Calyx 2 x 1 mm,
ovate, segmented, glabrous; Corolla: lobes 6-7 x 2 mm, ovate, apex
rounded, pale green and glabrous outside, purple and papillose within; lower
tube c. 1 mm long. Corona: inner corona a whorl of irregular bumped
interstaminal discs, emarginate. Androeclurn: stamens .± 2 mm long,
moderately thick, arise from mouth of corolla tube. Fruit: follicles 80-90 x 20
mm, ovoid oblong, apex narrow, dark brown. Seeds 5 x 2 mm, elliptic to
ovate, unilaterally keeled, glabrous; coma 25 mm long, pale brown. (Figure
7.7).
, 7.3 DISTRIBUTIONAND IECOlOGY:
Batesanthus intrusus has been collected between 00050'S to 07032'N and
09025'E to 08038'W; B. parviflorus between 08020'S to 08027'N and 05002'E to
12°29'W; B. purpureus between 01046'N to 03053'N and 11017'E to 18°01'E
and B. talbotii between 04009'N to 05017'N and 07055'E to 09°13' (Figure 7.8).
Flowering time is shown in Table 7.1. The only fruits seen were those of
Batesanthus intrusus.
Altitude varies from species to species. For instance, B. intrusus has been
collected at.± 550 m - 1,200 m. B. parviflorus at 1,225 - 1,900 m and B.
talbotii at 980 m. There were no records of altitude for B. purpureus.
78
Table 7.1 Flowering and fruiting periods of Batesanthus* species.
AP MAY JUN JUL DEC
R
B. intrusus
FLOWERS
B. intrusus
FRUITS
B. parviflorus
FLOWERS
B. purpureus
FLOWERS
B. ta/botJJ
FLOWERS
'" No fruiting specimens of B. parviflorus, B. purpureus and B. ta/botii were
recorded.
Batesanthus intrusus and B. ta/botii are rambling twiners in secondary forest
and/or forest edge. On the other hand, B. parviflorus is a climber in upland
evergreen bushland growing in association with Myrica L., Rapanea Aubl. and
Hypericum L., also a climber in secondary bush. B. purpureus is a climber in
riverine forests.
People of Yaoundé - Bitye in Cameroon refer to B. intrusus as "Ekótók",
known as "Bosambala" and "Inaolo a Bosambala" in Zaire - Yangambi and
Basoko respectively.
7.3.1 VOUCHER SPECIMENS:
1. Batesanthus intrusus:
CAMEROON: 06N 14E: Lakka, Distri.ct Barmuda
(-AD). Mait/and 1565 (K).
CENTRAL AFRICAN REPUBLIC: 03N 18E: Boukoko and Mbaiki
regions (-CC). Tisserant 1480 (P).
GABON: OONOgE: Libreville on the Gaboon
79
River (-AD). K/aine, R.P. 513 (P).
OOS12E: Region de Lastoursville
(-DC). Le Testu, M. G. 8501 (P).
LIBERIA: OlN 08W: Yéképa village (-DA).
Adam, J.G. 30433 (MO).
2. Batesanthus parv;f/orus:
ANGOLA: 08S 15E: Guanza, Quilela Camabatela (-AB).
Gosswei/er,J. 8468 (BM).
CAMEROON: 05N 09E: Manengouba Lake (-BB). Po/hill, R.M. &
Kirkup, D. W. 5190 (K).
SIERRA LEONE: 08N 12W: Near Rorules (Roruks) (-BC). Deighton, F.G.
3265 (K).
3. Batesanthus purpureus:
CAMEROON: Not traced. Bates, G.L. 383 (G, K).
GABON: 01N 11E: Haute Ngounié (-CD). Le Testu, M. 5493 (P).
4. Batesanthus ta/bot;;:
NIGERIA: 05N 08E: Oban (-BC). Ta/bot, T.A. 63 (BM).
05N 08E: Oban (-BC). Ta/bot, T.A. 2021 (BM, K).
80
Figure 7.1 Bafesanfhus intrusus: (A) Habit. (8) Leaf insert showing hairs.
(C) Single follicle with stalk. Drawn from Klaine 513 (K).
81
A
0I'\SIÓCNE:MCI \N\'fI.U'5.>4
A\lCM
No~.\'1"
Figure 7.2 Batesanthus intrusus: (A) Flower showing an intruse base.
(8) Open flower showing position of gynoecium (g), corona (c) and stamens
(5). Drawn from Klaine 513 (K).
82
A L-IlOr,'u'rI
Figure 7.3 Batesanthus parviflorus: (A) Specimen with flowers having
corona: (8) Open flower showing filiform corona lobes and reflexed corolla
lobes. (C) Cut flower showing position of gynoecium and insertion of corona
lobes.
Drawn from Oeighton 3265 (K).
83
A
t1a1l1AU _\lIll'", Illom'" ,I.
_\ •rft__' ......... ~~: ~.4.. éi.ou--1k1(---r
.-1 ~,
Jf'
Figure 7.4 Bafesanfhus parviflorus: Specimen, flowers without corona:
(A-B) Habit of profusely flowering specimen.
All from Gossweiler 8468 (BM).
Photos: Gabré Kemp
84
Figure 7.5 Batesanthus purpureus: (A) Habit. (8) Flower with exposed
gynostegium. (C) Open flower showing position of translators on the
pentangular stigmatic head (1-), corona (c) and stamens (5).
Drawn from Bates 383 (G iso).
85
M.S.cel.clum
Figure 7.6 Batesanthus purpureus: (A) Sepal seen from within. (B)
Corona and staminal column. (C) Staminal column partly dissected to show
the position of the pollen carriers. (0) Pollen carriers.
Source: Brown 1896, t. 2500.
86
Figure 7.7 Batesanthus ta/botii: (A) Habit. (8) Flower exposing
gynostegium. (C) Open flower showing position of corona (c), gynoecium (t-)
and stamens (s).
Drawn from Ta/bott 2021 (K).
87
( $ i .' !....(.....7.. lYi
\ '<.y"..,..').' (~:;~. ~ '!.....). '
'\ ..... '. ..~..J~""! ~\ ~.
10 ~ ~ i .:i" .....:...r-. \ 10
\ \ \ '( ' .....:/\>1« ') '/ Ir
\ \ \ \
o .. ,. ..0 '000•• \
\ o 20
Figure 7.8 Known geographical distribution of Batesanthus species.
KEY: B. intrusus ( • ); B. parviflorus (.&.); B. purpureus (Ill); B. talbotii ([Bl).
88
PLATE1: CORONA LOBES.
A. Filiform lobe of Batesanthus parviflorus. B. Obcordate lobe of Mondia whitei. C. Filiform lobe of Zacateza pedicel/ata.
Oeighton, F.C. 3265 (K). Pienaar, B.J. 170 (NH). Gossweiler, J. 13584 (K).
Scale bar = 100 urn Scale bar = 1 mm Scale bar = 1 mm
11"AXONOMY OIF MANGENOTUA Poclhoro
8.1 HISTORICAL BACKGROUND:
The genus Mangenotia was established by Marcel Pichon (1954)
(Recommendation 73B.1 (b)) to commemorate its first collector Professor
Mangenot. His description was based on the first collection of the specimen
from Ivory Coast in 1951.
There is one species, namely Mangenotia eburnea. The specific epithet
eburnea is a latin derivation of "eburneus", meaning ivory white or white with
a yellow tinge.
Mangenotia is a small liana or shrub of about 2-5 m tall, mostly in the tropical
forests of Africa. It is characterised by having a narrowly urceolate to
cylindrical tube as in Cryptolepis R. Br. (Periplocoideae). However, the flower
bud is capitate, conical and remarkably twisted to the right (i.e. dextrorsum).
Pichon (1954) indicated that Mangenotia is nearest to Calotropis R. Br.
(Asclepiadeae) but Bullock (1955) states that it is an obvious lapsus calami for
Cryptolepis to which the genus is closely allied.
8.2 TAXONOMY:
MANGENOTIA Pichon in Bulletin de la Société Botanique de France 101:
246-248 (1954).
TYPE: Mangenotia eburnea Pichon
Twining shrub or small liana, erect at first and later climbing with thin terete
stems; anthers sessile and very hairy; leaf surface puberuient adaxially and
glaucous with papillae projections abaxially; flowers creamy white; West
Africa (coastline)
89
Mangenotia eburnea Pichon in Bulletin de la Société Botanique de France
101: 246-248 (1954); Bullock 10: 587 (1955); Bullock: 81, 84 (1963b).
TYPE: Cote d'lvoire, Adiopodoumé, pres d'Abidjan. 07-06-1951. Mangenot,
S.n. (P, holotype!).
Liane, erect at first, later climbing and twining, with terete thin branches, latex
milky. Stems woody, drying brownish, puberulous or nearly so. leaves
decussate to opposite; petiole (3-)4-5(-7) mm long, puberulous; lamina (21-)
30-43(-50) x (9-)13-19(-22) mm, broadly ovate, ovate to elliptic, apex
acuminate to acute, base rounded, margin entire, glabrous, dark green above,
pale-green to glaucous beneath with hypostomatous surface, venation
brochidodromous. lnflorescences lax cymes, dichasial, puberulous;
peduncles (3-)4-5(-7) mm long; pedicels (4-)6-9(-11) mm long, bracts
small, decussate, outer side puberulous, inner side glabrous. IFIowe rs
pentamerous, complete, greenish, yellow, white or white creamy, flower bud
capitate, conical and contorted to the right (dextrorsum), gynostegium hidden
at base of corolla tube. Calyx of 5 sepals, 2 x 1 mm, pale green, free, ovate,
apex acute, margin somewhat ciliate, bilobed colleters at inner base. Corolla
of 5 petals, pale yellow, membranous; lobes free when open; tube 9-14 mm
long, cylindrical, puberulous outside, glabrous within, lower tube shorter at ± 3
mm long, upper tube cylindrical with constricted throat, longitudinal ridges
within; lobes linear oblong, ± 8 mm long, ivory white. Corona lobes arise at
middle of corolla tube and included within; lobes clavate, c. 1 mm long, with
ovate cushion at base. Androecium: stamens 5, inserted near base of tube;
anthers sub-sessile, hairy, connivent around the stylar head, connective very
prominent with flap/wing-like microsporangia; pollen granular in tetrads.
Gynoecium: ovaries 2, free, broadly ovate to cushion-like (clavuncula),
flattened at the base, half-inferior; styles 2, fused towards apex; stigmatic
head pentangular ovoid; translators 5, situated around the periphery of the
stigmatic head, spathulate with linear receptacle, stalk short to absent,
viscidium circular, sticky. Fruit of paired follicles; follicles 100-128 x 9-12
mm long, linear--oblong and tapering into a blunt point, glabrous with
90
longitudinal fissures on dry surface. Seeds 5-7 x 2 mm, numerous, flattened,
elliptic; coma of hairs whitish brown, 37-44 mm long. (Figure 8.1).
8.3 DISTRIBUTION AND ECOLOGY:
Mangenofia eburnea Pichon is restricted to West Tropical Africa, from Nigeria
to Sénégal between lines of latitude 05001'N - 14019'N and longitude 03023'E
-16045'W (Figure 8.2). Most specimens have been collected around the low-
lying coastline. The species grows at an altitude range of 0-500 m.
The flowering season is around May-August. Fruits were seen around
October-November (Table 8.1).
Table 8.1 Flowering and fruiting periods of Mangenofia eburnea
aCT NOV DEC
FLOWERS
FRUITS
In Ghana, the species mostly grows in shrubland along the Cape Coast and
attains a height of about 2 m. In forest habitat at higher altitudes, from 30 -
500 m, the species grows taller to about 5 m.
Ghana is situated along the coastline of the Gulf of Guinea, and consists
mostly of low-lying savanna regions with a central belt of forest.
The Ivory Coast lies in coastal rain forest, and M. eburnea was found growing
in white sandy soil at an altitude of 2 m. At sea level i.e. Island of Aladin, it
has been found in sandy soil of an old secondary forest.
Liberia is characterised by tropical moist evergreen forest, which towards the
interior, changes into a more deciduous type of forest. Here M. eburnea
grows in riverine, low secondary bush and reaches a height of ± 4 m.
Sénégal is a flat country with sandy soil and an altitude that does not exceed
130 m. It is semi-arid in the north and tropical wet-dry in the south. The
species was found in the forest at an altitude of 8 m.
91
Sierra Leone also presents varying degree of altitude, from 0-130 m. In this
country, quite a number of vernacular names exist that differ from region to
region or are tribal based. For instance, it is known as 'Kpokoyangoei"
(Mende) in Gbangbama, a vernacular name that also refers to species of
Periploca L., and its latex is used to treat craw-craw. ln Matotaka it is
"Njapagba", Yonibana people have two different names, "Kowonruia" and
"Jamolal" whereas those in Port Lokoh refer to it as "Lobwe". In Mayoso it is
known as "Ndifabwa" and "Moikwe" and/or "Cenje" in Mamaka.
8.3.1 VOUCHER SPIECIMlENS:
IVORY COAST: 05N 03W: Forêt de L'Abouabou, between Abidjan and
Grand Bassam (-BO). Leeuwenberg, A.J.M. 4235
(WAG).
05N 04W: .± 10 km. W of Jacqueville, old secondary
forest (-BA). Leeuwenberg, A.J.M. 8083 (BR).
SIERRA LEONE: 08N 13W: Mayoso (-CC). Thomas, N.W 1373 (K).
Not traced. Mamaka. Thomas, N.W. 4544 (K).
92
Figure 8.1 Mangenotia eburnea: (A) Habit. (8) Closed flower, bud
dextorsum; (C) Open flower showing gynostegium at base of tube. (0) Corolla
tube. (E) Anther. (F) Pollen tetrads. (G) Reproductive organs / translators on
apices of stylar head. Source: Pichon (1954).
93
\~.y )1.)...' (i::Ctf !. lY
I "". i". ~ .. ...../.
10
? \.).1'. o.•..., / >r~ N"·: ...!.. "'." .( \ f\i M ••••••• ~ r
20 \ \ \ \
~ , ,.,>' \; ) '/ / I
[ \\ \ \
o 400 800 1000 km
rx \ 10 \ ~ \ 0
Figure 8.2 Known geographical distribution of Mangenotia eburnea.
94
TAXONOMY OF MONDIA Skeels
9.1 HiSTORICAL BACKGROUND:
Mondia with its two species, is an African genus originally placed under the
genus Chlorocodon Hook. f. The genus Mondia Skeels (1911), is a new
combination resulting from Skeel's transfer of Chlorocodon Hook. f. (1871),
thus illegitimate due to its being a later homonym of Chlorocodon Fourreau
(1869) (Bullock 1962a).
The generic name Mondia has been derived from the Zulu vernacular name
"uMondi" , "Mundi", which refers to its aromatic tuberous roots with medicinal
properties, especially Mondia whitei (Hook. f.) Skeels. Conservation through
cultivation is an initiative that has been undertaken at Silverglen nursery in
Durban. Crouch et al. (1998) have taken various approaches to propagate (in
vitro culture) so as to conserve and save the endangered (if not facing
extinction) species of M. whitei. They refer to the species as a versatile plant
whose uses range from the source of stem fibers for rope-making, leaves
cooked as a vegetable, to the deliciously aromatic roots employed as a tonic,
anti-flatulent and aphrodisiac.
The specific epithet "ecornuta" is a Greek derived name, whereas "white""
commemorates Mr. AS. White from Natal who referred to the plant as
'uMondi root of Natal' and then sent a bundle of living roots to Kew. M. whitei
is the correct name instead of the species being spelt as whytei or white;; as it
appears in some publications.
95
9.2 TAXONOMY:
MONDIA Skeels in United States Department of Agriculture Bureau Plant
Industrial Bulletin 223: 45 (1911).
== Chlorocodon Hook. f. in Botanical Magazine: t. 5898 (1871).
TYPE: Mondia whitei (Hook. f.) Skeels. (Chlorocodon whitei Hook. f.).
A tall liana with stems up to 20 m long. Stem twining andlor climbing, woody,
drying brown with parallel striations, sometimes warted, terete. Latex milky
white. leaves opposite, large, petiolate with frill-like, toothed interpetiolar
stipules, reflexed; lamina broadly ovate to broadly elliptic oblong, margin
entire, sometimes ciliate, hypostomatous, abaxial and adaxial surfaces green.
Inflorescences axillary paniculate cymes. Flowers 5-merous; Corolla
campanulate to cupulatus with gynostegium exerted from corolla, glabrous or
nearly so, greenish outside, maroon to reddish brown inside; lower tube .± 1
mm long; upper tube absent. Corona 5, double; outer corona lobes
obcordate, with or without dorsal process, lobes arise at apex of lower corolla
tube, basally fused with staminal filaments and nectaries; inner corona of
interstaminal nectaries, fused basally to filament bases; nectaries sub-erect,
foliose, shute-like lobes, situated basally around style. Androeclum:
stamens 5, inserted at mouth of corolla tube, staminal appendages white;
anthers broadly triangular to linear with a connective, adnate to the dilated
part of the style, connivent into a cone on top of capitate stigmatic head;
filaments free basally, .± 2 mm long, adnate to corolla tube; pollen granular,
parate tetrads. Gynoecium: ovaries 2, free, semi-inferior, ovules numerous;
styles 2, fused into one apically, not exceeding anthers; stigmatic head
pentangular ovoid, apex conical; translators 5, linear to narrowly ovate, in
grooves of the stigmatic head, viscidium rounded, almost attached at right
angle to the stalk. Fruit: follicles widely to horizontally divergent, narrowly
ovoid to lanceolate, apex obtuse. Seeds comose. Species 2.
KEY TO THE SPECIES:
ta Outer corona obcordate without corniculate dorsal process .
96
· 1. Mondia ecornuta
1b Outer corona obcordate with middle lobe subulate and 2 dorsal,
corniculate process... 2. Mondia whitei
1. Mondia ecornuta (N.E. Br.) Bullock in Kew Bulletin 15: 203 (1961);
Agnew: 370 (1974). (Chlorocodon ecornuta N.E. Br. in Bulletin of
Miscellaneous Information Kew 97: 111 (1895a); N.E. Br.: IV: 256 (1895b);
Brenan 5(2); 64 (1949)).
TYPE: Kenya, Ribe near Mombasa. May 1870. Rev. T. Wakefield s.n. (K,
holotype!).
Twiner about 2 m high, glabrous to somewhat pubescent. leaves stipulate,
interpetiolar stipules frill-like, slightly pectinate; petiole (10-)20-25(-35) mm
long; lamina (25-)90-115(-145) x (10-)30-50(-100) mm, apex acuminate to
cuspidate, base rounded to cuneate, sometimes auriculate". Inflorescences:
few flowered; peduncles (7-)10-15(-30) mm long; pedicels (5-)10-15(-35)
mm long. Flower: obconic to cupulatus. Calyx 3 x 2 mm, elliptic to obtuse,
free. Corolla: lobes 7-10 x 3-6 mm (including tube). IFnJlot2: follicles paired.
Seeds2 globose. (Figure 9.1, 9.2).
1. Three specimens, from Congo Bouquet A. 631 and from Gabon Klein«,
R.p. 577 and Le Testu, M.G. 8865 have pubescent stems, leaves, petioles
and peduncles. The flowers are also comparatively smaller. Leaf base is
auriculate.
2. Fruit and seeds descriptions cf. icoKlaine, R.P. 577 (P), herbarium sheet.
97
9.3 DISTRIBUTION AND !ECOLOGY:
Mondia ecornuta is restricted in distribution to the eastern coastline from
Mozambique, Tanzania to Kenya between 03042'S - 07057'S and 36031'E to
39042'E and again collected from western coastline from the Democratic
Republic of Congo and Gabon at 00°50' S - 03°57' to 00°30' Nand 09°25' -
14°38' (Figure 9.5).
The flowering season is from January to November and fruits were recorded
in July and November (Table 9.1).
TABLE 9.1 Flowering and fruiting season of Mondia ecornuta
AUG SEP
This twining liana has been collected at an altitude of 50 m and 200 - 700 m.
The species has been found on limestones outcrop in association with
species of Saintpaulia H.A. Wendl., Securinega Comm. ex Juss., May tenus
Molina and Pandanus Parkinson. It is a creeper, in coastal scrub of Kenya in
young bush and also in hill forests of Tanzania.
People of Tanzania refer to the plant as "Mwendo" or "Wendi" (Kihehe tribe)
and also known as "Mwende" (Kipogoro tribe).
9.3.1 VOUCHIER SPECIMENS:
DEM. REP. OF CONGO: 03S 14E: Brazzaville, village de Kimpélé, km 16
route de Mayama-Mouyondzi (-DC). Bouquet, A.
631 (P).
GABON: OOS 12E: Lastoursville (-DC). Le Testu, M.G.
8865 (P).
98
KENYA: 03S 39E: Ribe near Mombasa (-DC). Wakefield,
T. s.n. (K, hola).
MOZAMBIC;UE: Not traced: Misala River, Hynesa. Allen, C.E.F.
139 (K).
TANZANIA: 05S 38E: Pangani Distr., Bushiri estate (-IBA).
Faulkner, H.G. 558 (K) (Sheet 1 & Collection (2)).
07S 36E: Ulenga Distr., lIingera near Ifakara
(-DC). Haerdi, F. 266\0 (K).
2. Mondia whitei (Hook. f.) Skeels in United States Department of
Agriculture Bureau Plant Industrial Bulletin 223: 45 (1911); Bullock 15(2): 203
(1961); Bullock 2(2): 82 (1963a); Ross: 282 (1973); Killick 45: t. 1792 (1978-
79); Van Jaarsveld 66(3): 90-91 (1980); Retief & Herman 6: 554-555 (1997);
Crouch et. ai.: 21 (1998); McCartan & Crouch 64(5): 313--314 (1998); Victor
et al. 10: 71-98 (2000).
TYPE: South Africa, Natal, Mfundisweni. 12 Dec. 1871. White s.n. (K,
holotype!).
== Chlorocodon whitei Hook. f. in Curtis Botanical Magazine 97: t. 5898
(1871); Bentham 2(2): 740-745 (1876a); K. Schum. 4(2): 215, 217, Fig. 64
O-Q (1895b); Masters (Ed.) 3(18): 234,243, Fig. 48 (1895); Schltr. 34: 314
(1896); Hiern 1: 680 (1898); J.M. Wood & M.S. Evans 1(1): 27-28, t. 31
(1898); N.E Br. 4(1): 255 (1902), 4(1) 542: (1907); Marlath, R. 3(1): 72
(1932); Brenan 5(2): 64 (1949); Bullock 9: 351 (1954b).
= Periploca latifolia K. Schum. in Engier Pflanzenfam Ost-Afrika C: 321
(1895c), 23: 232 (1896).
TYPE: Cameroon, Yaoundé. 28 Jan. 1897. Zeuker 589 (K, holotype!,
isotype!).
= Tacazzea amplifolia S. Moore in Journal of Botany 50: 337 (1912).
SPECIMEN:Angola, Cazengo. Gossweiler, J. 616 (BM, holotype, K, P).
99
= Tacazzea viridis A. Chev. (Expl. Bot. Afr. Occid. France: 429 (1920),
nomen) ex Hutch. & Dalziel 2: 52 (1931), :339 (1937), desc. lat.
SPECIMEN: Ivory Coast, Mankoro district, between Dialakoro and Kénégoué.
Chevalier, A. 21975 (K, holotype!).
Twining climber. Roots thick, succulent with aromatic scent. leaves
herbaceous, interpetiolar stipules dentate, forming a band connecting the pair
of petioles; petiole (11-)20-50(-80) mm long, deeply grooved at center,
pubescent; lamina (40- )80-155( -245) x (15- )55-90( -120) mm, broad ly ovate
to suborbicular, apex acuminate to sharply acute, base cordate to cuneate,
green, glabrous or nearly so adaxial, abaxial surface hirsute, more so on the
midrib. Inflorescences many flowered, pubescent; peduncles (10-)15-25
(-40) mm long; pedicel (5-)10-15(-20) mm long, dichotomously branched;
bracts c. 5 x 1 mm, linear oblong. Flowers rotate to broadly campanulate.
Calyx 2 x 1 mm, ovate, acute, free, membranous, green. Corolla: lobes 6-8
x 3-5 mm, oblong-obtuse with round apex, greenish yellow outside, maroon
to violet purple inside. Corona: outer corona obcordate, central lobe subulate
or horn shaped, 5-6 x 2 mm long, 2 obcordate or corniculate dorsal process,
spreading in open flower, fused to the outside of filament bases (Plate 1 -
after pp. 88). Fruit: follicles 30-120 x 15-25 mm long, ovate to lanceolate,
glabrous, very rarely puberulous; fruit stalk 45-80 mm long, warted. Seeds
obliquely ovate, keeled on both surfaces. (Figure 9.3,9.4).
9.4 DISTRIBUTION AND ECOLOGY:
Mondia whitei is widely distributed (Figure 9.5) in Tropical Africa from South
Africa eastern coastline of KwaZulu - Natal (Otherwise, it has been reported to
occur further inland in the Northern Province (northern Transvaal prior to
1994) to Sudan and western coastline from Namibia to Senegal, that is,
05059'N 00040'W - 12048'N 16018'W to 00027'N 29029'E - 04005'N 32043'E
and 01019'S 12015'E - 30043'S 30040'E. Altitude ranges from 80 - 1,900 m
above sea level.
100
The flowers were seen from November to February in the Southern
Hemisphere and June to August in the Northern Hemisphere and fruits from
March to December (Table 9.2).
Table 9.2 Flowering and fruiting period of Mondia whifei
JAN FEB MAR APR MAY JUN JUL AUG SEP
The species Mondia whifei is currently considered threatened \ vulnerable
(Hilton-Taylor 1996), if not extinct in the wild, with its distribution in South
Africa largely restricted to nature reserves and protected coastal swamp
forests of KwaZulu - Natal. Hooker (1871), made a note that the nearer the
plant grew to the sea, the sweeter and better was the flavour of the root. In
South Africa, Mondia whifei has been declared threatened due to over-
harvesting of the popular "liquorice' (as in Glycyrrhiza L.) roots that are
differently used for medicinal purposes throughout Tropical Africa. In other
parts of Africa, most of which are currently war-zones, there are no recent
collections thus the status is not known. This species mostly grows in
swampy conditions.
It is a climber on forest edge and riverine forest as in Mkuze Nature Reserve,
Hlabisa and Dududuku (South Africa) as well as in dense bush of
Brachysfegia Benth. woodland. Nevertheless, it is also common in Bamboo
thicket of Malawi and in decidous thicket of mixed Acacia Mill. woodland.
Otherwise it has been found in mixed forest edge with red sandy soil. The
plant is usually found in dense bush, the lower portion of the stem being
naked and leafless. The leaves only appearing at the tops of supporting trees
(Wood & Evans 1898) such as Ixora L. sp. and Trema onenietis (L.) BI. In
Ivory Coast, Nigeria, Cameroon and Sudan, the species has been collected
mainly from the savanna to savanna woodland. Otherwise, it has been found
in association with species of Anfhocleisfa AfzeI. ex R. Br., Albizia Durazz.,
101
Berlinia Sol. ex Hook. f. on river banks with Barringfonia J.R. Forst. & G.
Forst.
Uses range from the source of stem fibers for rope-making (Angola) and
medicinally popular among native tribes of Tropical Africa as an aphrodiasic
(Zimbabwe to East-Central Africa), easing of flatulence, settling the stomach
and acting as a tonic (South Africa) (McCartan & Crouch 1998).
There is an initiative at present by the Natal Herbarium together with
Silverglen Medicinal Plant Nursery (Durban) to propagate M. whifei for both
conservation and potential industrial intiatives. Wood & Evans way back in
1898 reported a proposal from the Natal colonists to make a ginger beer-like
beverage from the scented roots.
There are quite a number of vernacular names. People of Angola refer to the
plant as "Mudondo" or "Mundondo", although the name also refers to
members of Tylophora R. Br., some bears a close resemblance to M. whifei
(Hiern 1898). In South Africa, the Zulu call it "uMondi" or "Mundi". The
Lissongo tribal group of Central African Republic call it "Mundondo", "Banda"
or "Molabusia". The Fula of Guinea Bissau refer to it as "Lacadje". "Sedando
omutona" by the Luganda tribe of Uganda. It is "Citumbolo" in Malawi. The
different tribes of Zaire refer to the plant as "Nlondo", "Kimbiolangwa",
"Kumba" by the Ngwako, "Ubasangbwandiya" or "Gatimba" by the Lugware,
"Mujimbaye" by the Tshiluba.
9.4.1 VOUCHER SPECIMIENS:
CAMEROON: 09N 13E: Bosum-Buar (-DC). Mildbraed 9731 (K).
IVORY COAST: 07N 04W: Brabo, 25 km ENE of Bouaké gallery forest
(-DD). Oldeman, R.A.A. 392 (K, MO).
MALAWI: 15S 35E: South region slopes of Zomba Plateau (-AD).
Brummitt, R.K. & Seyani, J.H. 14810 (SRGH).
NIGERIA: 07N 03E: Ibadan Distr., near Busogboro (-BO). Onochie,
34943 (K).
102
SOUTH AFRICA: 23S 30E: Northern Province, Letaba, Duiwelskloof,
Westfalia estate (-CA). Scheepers, J.C. 1058 (PRE).
29S 31E: Kwazulu-Natal, Stanger (-AD). Pienaar, B.J.
170 (NH).
29S 31E: Durban (-CC). Medley Wood, J. 6180 (PRE).
30S 29E: Mfundisweni (-DC). White s.n. (K hola, iso).
ZIMBABWE: 20S 32E: Chipenge Distr., Gungungana Forest Reserve
(-BC). Goldsmith, B. 5/64 (SRGH).
103
Figure 9.1 Habit of Mondia ecornuta: (A) Leaves with flowers, taken from
Wakefie/d, T. s.n. (K). (B-C) Leaves and flowers, taken from Fau/kner, H.G.
558 (K). Photos: Gabré Kemp.
[04
Figure 9.2 (A - N): Mondia ecornuta showing floral parts (A-J, L), seeds
(K), floral diagram (M) and fruit (N).
Source: Herbarium sheet - Klaine, R.P. 577 (P).
105
Figure 9.3 Habit of Mondia whitei. (A) Branch with leaves and flowers,
about natural size. (B) Flower. (C) Calyx. (D) Corona, front and side view. (E)
Translator. (F) Stigma. (G) Follicles, about natural size.
Source: Wood & Evans (1898).
106
Figure 9.4 (A) Natural habit of Mondia whifei in Mkuze (S.A). (8) Terete
stem with leaves and flowers.
Photos: Prof. H.J.T. Venter.
107
o o
10 10
..~.. r··· I i:.... "i~\ I~ -"" .
, , 0... (' l'
0, " 0 .:.
\ \ \ .. ; ,.' '''1. ..... ~ ..... ! /1\ ' .' IP }\.II \ \ \ \ I~' ,_ r: I
o "'0 '" 00" 'm \ '(~L/ I I I I ï
~o \ ~ \ o
Figure 9.5 Known geographical distribution ofMondia ecornuta (..t..) and M. whitei (<I).
108
TAXONOMY OIFSACLEUXIA Bam.
10.1 HISTORICAL BACKGROUND:
Sacleuxia is an East Africa genus with two species, namely S. newii (Benth. )
Bullock (Fig. 10.1) and S. tuberosa (E.A. Bruce) Bullock (Fig. 10.2). Baillon
first published the generic name in 1891 in honour of Mr. Sacleux who
collected the specimen from Zanzibar with S. salicina Baill. as the type
specimen.
S. newii commemorates plant collector, the late Rev. C. New who, according
to herbarium records collected the specimen on his expedition up Kilimanjaro
Mountain (Tanzania)). S. tuberosa refers to the striking potato-like tuberous
roots. "Tuberosus' is a Greek derived word meaning "producing tubers or
swollen into a tuber'.
Gymnolaema Benth. as published by Bentham (1876b) is an orthographic
variant. Bullock (1962a) referred to Gymnoleima Decne. (type species
Lithospermum graminifolium) a genus in the Boraginaceae, as a dead
synonym. Nevertheless, Decaisne's name was validly published and it thus
retains its status for purposes of priority and homonymy. Hence, Sacleuxia
(Recommendation 73B.1 (b)) of the ICBN became the correct name for this
taxon.
Gymno- is a Greek derived word meaning 'naked'; the genus was named on
account of the 'naked throat of the corolla'.
10.2 TAXONOMY:
SACLEUXIA Baill. Histoire Plantarum 10: 265 (1891).
TYPE: Sacleuxia salicina Baill.
109
- Gymnolaema Benth. in Bentham & Hooker, Gen. PI. 2: 740 (1876a); N.E.
Br. 4(1): 241 (1902); K. Schum. 4(2): 211 (1895a); Dale & Greenway:
57- 58 (1961); Sharpe & P.1.Forst. 47: 12-14 (1989), non Gymnoleima
Decne. in AP. Candolle (Ed.): 491-500 (1844).
Type: Gymnolaema newii Benth.
= Macropelma K. Schum. in Engier Pflanzen. Ost. Afr. C: 321 (1895b), 23:
232 (1896).
Type: Macropelma angustifofium K. Schum.
Erect shrubs with tuberous roots. Leaves opposite, simple, margin entire,
hypostomatous, venation brochidodromous, puberulous to somewhat
glabrous, elliptic to oblong-linear, pale-green abaxially, dark-green adaxially.
Inflorescences clustered, panicle or raceme of cymes, few flowered.
flowers pentamerous, bisexual, gynostegium exposed from corolla. Corolla
with broadly campanulate tube, indistinct. Corona annular, obscure with
small fleshy lobes fused to the filaments. Andreeeturn of 5 stamens,
epipetalous; anthers sub-sessile, ovate, connective apically acute, connivent
on top of pentangular ovoid stigmatic head; filaments ·free fused to
interstaminal discs, connate; pollen in granular tetrads. Gynoecium: ovaries
2, sub-globular, semi-inferior, pubescent' styles 2, becoming fused apically;
translators 5. Fruit follicles, paired or clustered, divergent horizontally.
Seeds oblong to obovate, comose. Species 2.
KEY TO THE SPIECIES:
1a Leaves sessile or sub-sessile, blade base cordate; glabrous or nearly so,
at least on adaxial surface. Inflorescences with peduncles slender, 30-
130 mm long; flowers yellow white; ovary glabrous 1. S. newli
1IJ Leaves petiolate, blade base cuneate to obtuse; pubescent, more so
abaxial. Inflorescences with peduncles up to 10 mm long; flowers
purple brown; ovary hairy 2. S. tuberosa
110
1. Sacleuxia newii (Benth.) Bullock in Kew Bulletin 15: 393-394 (1962a)
(Gymnolaema newii Benth.: 74, t. 1186 (1876b)).
TYPE: Tanzania, T2 Moshi Distr., Kilimanjaro mountain. New s.n. (K,
holotype!).
= Sacleuxia salicina Baill. 10: 265 (1891).
TYPE: Zanzibar, Nguru. 01-06-1892. Sacleux 758 (P, holotype!).
= Macropelma angustifolium K. Schum.: 321 (1895c).
TYPE: Tanzania, Merue. Fischer 383 (B \jf). Synonymy after Bullock 1962b
and description of species.
Virgate, branched shrub, 1,5 - 2,5 m high. Roots up to 0.15 m in diameter.
Stem purplish, drying brown, becoming glabrescent, latex milky-white, not
copious. Leaves: petiole 0-3 mm long, sessile or sub-sessile; lamina (33-)
64-75(-127) x (8-)12-15(-19) mm, apex acute, base cordate, adaxial
surface almost glabrous but hirsute on the midrib, abaxial surface pubescent,
more so on the midrib. Inflorescences: peduncles 30-130 mm long,
puberulent; pedicels 2-5 mm long, slender, puberulent; bracts ovate,
clustered. Flowers: Calyx .± 1,5 x 1 mm, free, triangular, sub-glabrous.
Corolla: lower tube .± 1 mm long; lobes 2 x 1 mm, ovate, yellow white,
glabrous. Corona on apex of lower tube, cream coloured; outer corona lobes
present as inconspicuous squarish lobules at base of staminal column.
Andlroecium: stamens connate; anthers sub-sessile, arise from lower tube.
GYll1oecium: translators' receptacle angular ovate on sub-circular viscidium.
Fruit: follicles paired, 30-37 x 5-8 mm, green, shiny, more so inside when
dry, elliptic-linear; fruit stalk 7-9 mm long. Seeds 4-7 x 1-2 mm, compressed
with a coma, shiny, glabrous, oblong; coma of hairs 29-34 mm long. (Figure
10.1 ).
10.3 DISTRIBUTION AND ECOLOGY:
Sacleuxia newii is very much restricted to eastern Africa between latitude
00055'S - 06046'S and longitude 36027'E - 39020'E (Fig. 10.3), that is, in
Kenya and Tanzania. Altitude ranges from 600-1,700 m.
III
Flowers were seen more or less throughout the year as the species occur just
South of the Equator (Table 10.1).
Table 10.1 Flowering and fruiting periods of Sacleuxia newii
JAN FEB
This is a shrub with a variety of habitats, from open bush land to closed
woodland. The most common habitat where the species thrives is in the
bushland, rocky outcrops and slopes with shallow soil cover.
It is a frequent shrub in the more open scrub of Markhamia Seem. ex Baill.,
Xerophyta Juss., Commiphora Jacq., and Steganotaenia Hochst. and may
also be co-dominant with Vellozia Vand. sp. In most of the outcrops and
slopes, where the species was collected, it was growing with species of
Vellozia, Entada Adana., Commiphora, Terminalia L. and Aloe L. In
deciduous bushland, more so secondary forest, it is found with species of
Acacia Mill., Commiphora, Dombeya Cav., Stereospermum Cham.,
Markhamia, Annona L., Afzelia Sm. and Euphorbia L.
The woodland, closely associated to thorn woodland where it was collected,
was characterized by the presence of species like Acacia, Combretum LoefI.,
Sterculia L., as well as tall grass, although grasses are relatively a minor part
of the vegetation. On the south facing, steep rocky slopes of Kanga peak
(Tanzania), it grows together with Xerophyta Juss. and Ericaceae heath.
10.3.1 VOUCHER SPIECIMlENS:
TANZANIA: OSS 38E: T3 Handeni Distr., Kideleko Mts. (-AC). Faulkner,
H.G. 1434 (K).
OSS38E. T3 Lushoto Distr., Amani, Mt. Bomale, East
Usambaras (-BA). Verdcourt 188 (MO).
OSS38E: T3 Tanga Distr., Magunga, sisal estates, footpath
112
between Magunga & Ngua estates, west slopes of E.
Usambaras (-BA). Drummond & Hems/ey 3364 (USe).
05S 39E T3 LushotolTanga Distr., lower Sigi valley, East
Usambaras (-AA). Verdcourt 246 (K, MO).
2. Sacleuxia tuberosa (E.A. Bruce) Bullock in Kew Bulletin 15: 393-394
(1962a); Agnew: 369 (1974). (Gymnolaema tuberosa E.A. Bruce: 304
(1934)).
TYPE: Tanzania, T1 Mwanza, coast of Speke Gulf, Lake Victoria, Burlt, B.O.
2475 (K, holotype!, isotype!).
A pubescent shrub up to 4 m high. Stems terete, thin, sparsely branched,
lactiferous, latex colourless. leaves: petiole 2-5 mm long, puberulent;
lamina (31-)41-80(-131) x (6-)10-14(-19) mm, apex acute to acuminate,
base cuneate to obtuse, midrib tinged with magenta. lnflorescences:
peduncles up to 10 mm long, puberulent; pedicels 3-5 mm long, puberuient.
!Flowers: Calyx puberuient outside. Corolla: lobes ovate, 4 x 2 mm,
puberulent, purple brown; lower tube c. 1 mm long. Corona fleshy, lobules
triangular to linear, dull green; outer corona situated on the mouth of the tube,
, fused into an annulus to filament bases; inner corona of interstaminal discs
fused to inner base of filaments. Androeclum: stamens arise from base of
corolla tube. Gynoecium: ovaries 2, hairy; translators' receptacle broadly
ovate to cordate, viscidium almost rounded to spherical. Fwit paired follicles,
sometimes clustered; follicles (30-)37-42(-50) x (5-)8-12(-15) mm, narrowly
ovoid, tapering into a blunt point, glabrous or nearly so, with longitudinal
fissures on surface; fruit stalk 5--6 mm long. Seeds 4-7 x 1-2 mm, flattened,
drying brownish, obovate; coma with hairs 12-13 mm long. (Figure 10.2).
10.4 DISTRIBUTION AND IECOlOGY:
The species S. tuberosa is endemic to Eastern Africa at an altitude ranging
from 800 - 1,900 m. The range of distribution is between lines of longitude
113
00055'S and 06054'S and lines of latitude 31020'E and 38047'E (Fig. 10.3).
The flowers were seen from October to April and the fruit from April to July
(Table 10.2). However, fruits were seen as early as January (Kerfoot, O.
3596 (K)).
Table 10.2 Flowering and fruiting periods of Sac/euxia tuberosa
Jan Feb Mar Apr May
In Kenya, the species was found mostly on rocky outcrops and/or slopes with
reddish soil, in association with species of Dodonaea MilL, A/oe, Co/eus Lour.,
G/eichenia Sm., Lycopodium L., Dissotis Benth., Cressoceonelum Moench,
sometimes with Acacia, Ormocarpum P. Beauv., Tee/ea Delile, Heeria Meisn.,
Commiphora, Rhus L., Tarchonanthus L., Tinnea Kotschy & Peyr., Bar/eria L.,
Combretum and Ruttya Harv.
There are a number of habitats, all have granite rocks in common. One such
is the rock thicket and rocky grassland. Another example is that of rocky hill
slopes with species of Lannea A. Rich., Sterculia, Brachystegia Benth. and
Loudetia Hochst. ex Steud. lts association with Brachystegia has been
reported to be on a shallow soil cover with brownish loam and deposits of
gravel.
Taking into consideration that most of the habitats encountered so far are dry,
among granite rocks near the lakeshore, it is not strange to find it among
species of A/oe. In the dry Miombo woodland, it is exposed on sloping
rockfaces and in association with species of Xerophyta.
Dry forest - Miombo woodland varies in density from an open park-like growth
of trees to relatively dense stands. Trees which form this forest are practically
the same as those which occur in Acacia - tall grass savanna e.g. Strychnos
L., Brachystegia, Acacia, Ziziphus Mill., Bauhinia L., etc. (Shantz & Marbut
1923)
114
In Tanzania, the Wasukuma tribe uses stems of S. fuberosa to prepare arrow
shafts. They also use the tuberous roots as medication for scabies, the roots
are pounded up green and then rubbed into scabies which has been
previously well washed. Its vernacular name is 'Kisukuma' (Matala).
10.4.1 VOUCHER SPECIMENS:
KENYA: OOS 36E: K4 Naivasha Distr., Ol Longonot Estate (-CD).
Kerfoot, O. 3596 (K).
TANZANIA: 02S 33E T1 Mwanza Distr., Bbarika (-DA). Tanner, R.E.S. 360
(K).
115
Figure 10.1 Sac/euxia newii: (A) Habit. (8) Flower. (C) Open flower: s =
stamens, C = outer corona.
Drawn from Drummond & Hems/ey 3364 (USC).
116
B
.... J J<.. , .. t .,It .. ,,
.. , ... -"'._ ...... 1\'-"..11.
__ t __ ....
Figure 10.2 Habit of Sacleuxia tuberosa: (A) Paired follicles (1-) (Tanner 360
(K)). (8) & (C) Clustered follicles (1-) (Kerfoot 3596 (K)).
Photos: Gabré Kemp.
117
"'"" l'--':h , ! r: ··.··.·····'r···········<;.:···········.·.···,····/·.... /
o .' j :::J······C:::!.; .. .;Y / \ / 0
\~" ..r J .)..t ' / :;~~ ~~4lY
I '\ \/..... ~/' ~
10 10
?
\ \ ' r r1._::.....\..r ') '/ /1\ /: .
• .00 ••• iooo •• \
10 _\ ~ \ o 20
Figure 10.3 Known geographical distribution of Sacleuxia newii (..6.) & S. tuberosa (e).
118
TAXONOMY OF SARCORRHIZA Bullock
11.1 HISTORICAL BACKGROUND:
Sarcorrhiza Bullock is a monotypic genus from Central and East Africa (Figure
11.3). The species Sarcorrhiza epiphytica Bullock was first described and
established by AA Bullock in Hooker's, Icon. PI. (1962b). He based his
description on a collection from Tanzania, Verdcourt, B. & L.O. 1725 (K).
Sarcorrhiza is a generic name that refers to the 'fleshy root-like organs';
sarco- is a Greek derived word meaning fleshy and rhiz- also a Greek
derived word pertaining to the roots or root-like organs. The specific epithet -
epiphytica refers to the epiphytic habit of the shrub. Hence, Sarcorrhiza
epiphytica literally means an "epiphyte with fleshy roots."
Sarcorrhiza epiphytica is an epiphytic shrub, a feature rather unique within the
African Periplocoideae. The only other Periplocoideae epiphytic genus is
Epistemma D.V. Field & J.B. Hall with 3 species found in West Tropical Africa.
Sarcorrhiza is characterized by having cylindrical, fleshy and tuberous roots.
With its fleshy tuberous rootstock, it is similar to the genera within the
Periplocoideae namely - Raphionacme Harv., Petopentia Bullock and
Sacleuxia Baill. which also have large perennial rootstocks. According to
Bullock (1962b), the genus is unique in its overlapping bracts and bracteoles
on the peduncles. It has been in the past confused with Petopentia and
Tacazzea Decne. as can be seen on some herbarium sheets.
11.2 TAXONOMY:
SARCORRHIZA Bullock in Hooker's, Icones Plantarum: ad t. 3585 (1962b).
TYPE: Sarcorrhiza epiphytica Bullock
119
An epiphytic shrub with tuberous roots. Stems twining and convolute.
Distinctly bi-coloured corolla lobes, marginal band yellow-green, central area
spotted blood red and median line blood red to brownish. Corona filiform.
Species 1.
Sarcorrhiza epiphylica Bullock in Hooker's, Icones Plantarum: ad t. 3585
(1962b).
TYPE: Tanzania, T3 Lushoto District, path up Mt. Bomole, behind Amani.
Verdcourt, B. & L.O. 1725 (K, holotype!).
Creeping epiphyte. Tuberous roots, 50-90 mm long and around 35 mm in
diameter, cylindrical, oblong to elliptic and purple brown. Stems contorted,
twining, with adventitious rootlets (Fig. 11.2 C), branching internodes
elongated as long as 150 mm but internodes on lateral branches shorter and
frequently 10-30 mm long, lactiferous. leaves opposite, simple, reticulate;
petiole (4-)8-9(-10) mm long, pubescent, dark crimson to deeply blood-red;
lamina (30-)55-75(-115) x (13-)17-20(-30) mm, narrowly elliptic, apex
acuminate to acute, base cuneate to rounded, margin entire, glabrous or
nearly so with hairs on the main veins, glossy on both surfaces, midrib blood-
red, secondary veins looped towards the edge of the lamina with
inconspicuous tertiary veins, abaxial surface pale green, hypostomatous,
adaxial surface dark green. inflorescences lax, cymose with monochasial
branches, 2-4 flowered; peduncles 10-20 mm long, glabrous or nearly so;
pedicels c. 30 mm long, glabrous or somewhat sparsely pubescent. Flower
5-merous, bisexual, actinomorphic, complete. Calyx of 5 sepals, 10-20 x 10
mm, free, broadly triangular to narrowly ovate, apex acute to ·slightly obtuse,
puberulous, green. Corolla: lower tube annular; upper tube absent; lobes
broadly elliptic, ovate, apex acuminate, bi-coloured, contorted, glabrous,
yellow-green outside, faintly flushed salmon inside, spotted reddish-brown
with a deep blood red to brownish median line. Corona: outer corona 5-
lobed, lobes inserted towards base of lower corolla tube, opposite filaments,
fused basally to staminal feet, forming a coronal annulus together with corona
lobes and stamens, lobes filiform with acute apex, cream, glabrous and
120
sinuate; inner corona form a ridge of interstaminal nectaries towards the base
of the lower corolla tube. Androeclum: stamens 5; anthers ovate, connective
apically acute; filaments free, arise from inner base of corona lobes; pollen in
granular tetrads. GynoeciUlm: ovaries 2, unicarpellate, semi-inferior; styles 2,
becoming fused towards the apex, massive; stigmatic head pentangular,
broadly deltoid/conical; translators arise from adaxial surface of stigmatic
head (cf icon), spoon shovel-like with receptacle obovate, stalk linear,
viscidium rounded. Fruit": follic/es paired, pendulous, 140-450 x 45-50 mm,
horizontally divergent, narrowly cylindrical, puberulous; fruit stalk 9-10 mm
long, pubescent. Seeds obliquely ovate to oblong, compressed, with coma of
hairs. (Figures 11.1 and 11.2).
*Described from herbarium sheet notes
11.3 DISTRIBUTION AND IECOlOGY:
Sarcorhhiza epiphytica Bullock is found in West-Central and East Equatorial
Africa. It is found between the lines of longitude 05014'N and 06°55' Sand
latitude 06°1 OW and 3B03B'E (Figure 11.3).
The flowering season for this species is around November to January (Table
11.1). Fruits were found around May.
Table 11.1 Flowering period of Sarcorrhiza epiphytica
FEB MAR APR MAY JU
FLOWERS
FRUITS
The species was reported at altitudes from 1,000 - 1,900 m.
In Tanzania, near Amani, at an altitude c. 1,000 m, S. eoionynce is found in
the crowns of Piptandenia Benth. (P. buchananii), and Cupressus L. sp.
Some collectors refer to it as a climber because of its upward creeping growth
on its support species. In Democratic Republic of the Congo, it was collected
in Kwango south of the equator, also in the north-east region of Ruwenzori in
Uganda at 1,BOOm.
121
According to Coconet (1998), Ivory Coast, with its low lying flat to undulating
terrain, is characterized by a rain season and dry season and it is a
transitional zone between the moist equatorial climate and the dry tropical
climate with temperatures ranging between 28°C and 37°C. Therefore, it
renders the ecological status of this region more of a mansaan forest than a
tropical rain forest. The collection from Ivory Coast was from a lower altitude
than those found in Tanzania and Democratic Republic of the Congo. It was
collected along the course of River Davo.
11.3.1 VOUCHER SPECIMENS:
TANZANIA: 04S 38E: T3 Lushoto Dstrict, path up Mt. Bomale
(-CD). Verdcourt, B. & i.o. 1725 (K, BLFU).
06S 37E: T6 Morogoro District, Uluguru North, north-
west side of mist forest (-DC). Schlieben, N.
2939 (BR).
UGANDA: DON29E: U2/Zaire, Toro District, Ruwenzori, Kanvui
(-BD). Bequaert 4494 (BR).
122
Figure 11.1 Sarcorrhiza epiphytica. A - tuberous roots; B - habit, showing long and
short shoots; C - part of inflorescence showing bracts and flowers; D - flower from above:
E - flower centre from above; F - stamens from inside; G - section of flower bud; H-
corona and gynostegium; I - pollen carrier; J - section of flower to show corona, stamens
and style-apex (carpels removed).
All drawn from Verdcourt 1725 K. Source: Bullock 1962b.
123
A
c
..:.
Figure 11.2 Sarcorrhiza epiphyfica: (A) Cross section of flower, Bequaert
4494. (8) Twining stem and tuberous root, Verdcourt 1725. (C) Habit
showing leaves (a), part of stem with adventitious roots (b) and follicle (c),
Bequaert 4494.
124
o
\",r'''I).' ,)~\f}, )/
'\ ..• ,A. ril 1:\... II·./.
10
2J
\
\ \ \
-~o 400 800 1000 km \ o 20 ~/o 40 I 50 ~
Figure 11.3 Known geographical distribution of Sarcorrhiza epiphyfica.
125
TAXONOMY OIFZACATEZA Bullock
12.1 HISTORICAL eACKGROUND:
Zacateza is a monotypic genus established by Bullock (1954a), to
accommodate his transfer of Karl Schumann's (1893) Tacazzea pedicellata K.
Schum. (Article 55.1). The type specimen, Schweinfurfh 3483 (K), is a
lectotype and 3488 (K) a syntype declared by Bullock (1954a), both
specimens are on the same sheet. Schumann's types were housed in Berlin
Herbarium, which was bombed in World War 2.
Schumann based his description mostly on floral characteristics of which are
common features in other African Periplocoids. Floral similarity, that is, rotate
flowers, filiform corona lobes as well as anthers with pollen tetrads is a
common characteristic that forms a close relationship between species of
Buckollia Venter & R.L. Verh., Periploca L., Petopentia Bullock, and Tacazzea
Decne. Nevertheless, due to different inflorescence types between Tacazzea
and Zacateza, the latter has been created as a separate taxon with
exceptionally large and coriaceous leaf laminae having conspicuous patent
secondary venation and it is characterized by having slender and relatively
long flower pedicels with axillary clusters of monochasia. In contrast,
Tacazzea has leaf laminae with arching secondary veins and inflorescences
of many flowered cymose panicles.
12.2 TAXONOMY:
ZACATEZA Bullock in Kew Bulletin 9: 351 (1954b).
TYPE: Zacateza pedicellata (K. Schum.) Bullock
Slender liana in gallery and swamp forests. Stem softly woody. Leaves
coriaceous, glabrous with numerous parallel lateral patent veins and fine
126
reticulation in between. Inflorescence cymose. Outer corona lobes filiform.
Follicle falcate, reflexed. Species 1.
Zacateza pedicel/ata (K. Schum.) Bullock in Kew Bulletin 9: 351 (1954b);
Bullock: 82 (1963a) (Tacazzea pedicel/ata K. Schum.: 115 (1893); N.E. Br.
4(1): 263 (1902)).
TYPE: Democratic Republic of the Congo - Munsa MonbuUu land. April
1870, Schweinfurlh 3488 (K, lectotype!).
= Tacazzea pedicel/ata var. occidentalis N.E. Br. in Flora of Tropical Africa
4(1): 263 (1902).
TYPE: Nigeria, Igbessa - Lagos Botanical Garden. January 1893. Mil/en, A.
130 (K, holotype).
Slender twining andlor climbing liana about 4-15 m high with milky white,
sticky latex. Stems thin, softly woody, sometimes much divided from the
base, purplish, bark drying brown, glabrous with longitudinal fissures. leaves
opposite, exstipulate, simple, interpetiolar ridges with dentate colleters; petiole
9-14 mm long; lamina (55-)85-105(-200) x (15-)30-40(-60) mm long,
elliptic to broadly elliptic, apex acuminate, base cordate to rounded, margin
entire, lateral veins perpendicular to midrib, looped towards the margin,
tertiary veins reticulate, abaxial surface pale green with red blotches, adaxial
surface dark green, amphistomatous, coriaceous. Inflorescences cymose,
cluster of monochasial branches with .± 6 flowers per node, glabrous;
peduncle 8-10 mm long; pedicel (25-)35-45(-65) mm long: bracts c. 4 mm
long. Flower 5-merous, greenish white or cream. Calyx of 5 sepals, 0.5-1
mm long, red, free, triangular, apex cuspidate, glabrous. Coro/la of 5 petals;
lobes 6-8 x 4 mm, narrowly ovate, apex acute to acuminate, reflexed at
anthesis, rose-red; lower tube .± 1.5 mm long; upper tube absent. Corona
double; outer corona of 5 filiform lobes (Plate 1 - after pp. 88), .± 6-11 mm
long, frequently as long as the corolla lobes, free, filiform, apex tapering,
greenish; inner corona of triangular, 2-lobed interstaminal discs, fused basally
to filament bases and nectaries into a collar around style. Androecium:
stamens 5, inserted at mouth of corolla tube; anthers whitish, abaxially fused
127
to stigmatic head, broadly triangular to ovate with prominent connective
connivent over stigmatic head; filaments.± 1 mm long, free basally, adnate to
corona; pollen granular in parate tetrads. Gynoecium: ovaries 2, semi-
inferior, free, unicarpellate, slightly elongated, ovules numerous; styles 2,
becoming fused into one apically, semi-inferior; stigma tic head pentangular,
broadly ovoid, apex acuminate; translators 5, arise from sides of stigmatic
head, receptacle broadly obovate to oblong, stalk absent, viscidium rounded.
Fruit paired follicles, glabrous; follicles 65-80 x 10-15 mm, oblong,
horizontally divergent to falcate, apex tapering; fruit stalk 70-80 mm long.
Seeds 5-10 x 2-3 mm, angular ovate, flattened; coma of hairs 20-22 mm
long. (Figure 12.1).
12.3 DISTRIBUTION AND IECOlOGY:
Zacateza pedicel/ata has been collected between 06033'N to 07033'S and
03023'E-29049'E (Figure 12.2).
The flowers and fruits were seen as indicated in Table 12.1.
Table 12.1 Flowering and fruiting periods of Zacateza pedicel/ata
FLOWERS
This liana species seems to thrive best at an altitude of 470-700 m above sea
level. Habitat ranges from gallery forests to swampy areas (e.g. along the
river). In the forests, it was found growing in association with species of
Voacanga Thouars, Cyperus L. and also in the forest margin of Raphia
Beauv. (R. hookerii) and Symphonia L. f. However, in the swampy conditions
especially along the riverbanks, it is yet in association with species of Raphia,
Cyperus (thus an overlap with forest species) as well as Mussaenda L. and
Nephrolepis Schott.
128
People of Yalutcha - Democratic Republic of the Congo refer to the species
as "lnaolo a Bosambala" (dialect of the Turumbu tribe) and called "Wilinduku"
by the Ngwaka tribe of Boyasebégo.
This climbing and/or twining liana is most likely cultivated as an ornamental
plant due to its attractive 'rose-like' flowers (herbarium sheets).
12.3.1 VOUCHER SPECIMENS:
NIGERIA: 06N 03E: Igbessa, Lagos Botanical station (-BC). Mil/en, A. 130
(K).
ZAIRE: OON 24E: Yabalanga, Isangi territory (-CC). Leonard, A. 698
(K!).
03N 28E: Belgian Congo, Munsa Monbuttu land (-CB).
Schweinfurfh, G. 3483/8 (K, lectotype).
06N 19E: Congolan, Bumba territory (-CD). Evrard 3467 (K).
129
Figure 12.1 Zacateza pedicel/ata: (A) Habit. (8) Flower with reflexed corolla
lobes exposing gynostegium. (C) Open flower showing corona insertion of
stamens (s) and stigmatic head. Drawn from Evrard 3467 (K).
130
LI '·OD
/
f /r;/A
r
~i.(~..J.
l.;.
r..~I \ >--~ r:~~1 \~[\ \ \\ \~~ 30/ ~ ,
( ./. I 1/··'·'·····..... )............... ~ ~ , \
2,l- ..· · ! "."\, , ' " ..····1··.'.. .. 1 ' \ _l2Q
) JIl", I_,I.!... ' I '!" .NI\ \j ('\ "'./" .'\ .., .......... .,. ../' '-'.1 i L.
~1}\.)\2.,' ;/~;<\~" /-V /\; 1"10 \~l ,<. ' '( ) ~~J- i _~~
fl\ :0,/,'r !.i~' i; I' ,)... ,. ") " .... > ...'. ( ....... ;...j_'r -. ,,"
c: ( \ \ ......,\,.i e -c.. " .,/
r'\. ") )..,., GIl r _ \~I~."~ !~ -,,...(1.:.:........r. .: ~ ....·. v, .l....... J' ..::...:....:...... ,_.,(......... I /• a •. :::tJ·.: +:;, / ~~: ../V 0J
~O l--~-l---f----t~\T-=\' .-'.i-J ---c, ,..;!r;-lo 0 0 U ij :;r·f ·Y lY
f"\.:'" ! 0 \0 ,.~ "., .., l~ ~; '.1....... r .., It_:) }It ""::L
10
\ \ \
I~\ \ \ \ 1\" ,-_/ r>. 0/ I j
o 400 800 1000 km
Figure 12.2 Known geographical distribution of Zacateza pedicel/ata.
131
CLASS!FICATION
13.1 CLASSIFICATION:
Endress and Bruyns (2000), provide a monophyletic classification of the
family Apocynaceae s.l. that consists of 424 genera distributed among five
subfamilies (Table 13.1).
Table 13.1 Updated classification of the Apocynaceae 5./.
PERIPLOCOIDEAE
Secamoneae
Source: Endress & Bruyns 2000.
132
However, they have not incorporated into their classification system Venter &
Verhoeven (1997), three tribes of the Periplocoideae i.e. Periploceae,
Gymnanthereae and Cryptolepideae stating the overlap of generic names
into tribes and that their results conflict with molecular results of Civeyrel et al.
(1998). The Gonolobeae has been included in the Asclepiadeae
(Swarupanandan et al. 1996, Endress & Bruyns 2000). Hence the question of
tribal classification within the Periplocoideae needs further data assessment
and review.
13.2 TRIBAL POSITiON
This study does not support the tribal classification put forward by Venter &
Verhoeven (1997). They have placed more emphasis on the absence or
presence of a well developed corolla tube, a character or an assumption
which has been proven wrong with accumulation of more data. For instance,
Sacleuxia has been placed under Periploceae (previously in the
Cryptolepideae) due to the absence of a developed corolla tube. The present
state is that the tribal classification has been rejected on the basis that, the
corolla is much more variable within a genus, that is, the corolla tube may be
distinct in some species, but indistinct in other species of the same genus.
All of which give way to a new approach to classification in the form of a
phylogenetic analysis (Venter & Verhoeven 2001, in press).
PIERIPlOCOIDIEAE:
Climbers, tall lianas, erect to scrambling shrubs or shrublets, geophytic herbs,
or rarely epiphytes with watery or milky sap; roots often tuberous. Leaves
with interpetiolar lines, ridges or clustered colleters adaxially at juncture of
petiole and blade. Flowers often fragrant; inflorescence axilliary cymes.
Calyx persistent. Corolla: lobe aestivation usually dextrorse. Corona
double; outer corona of 5 free or fused, epipetalous lobes; inner corona of free
or fused interstaminal discs. Stamens: filaments free or fused to outer or
inner corona; anthers 4-locular, fused to style head, nearly horizontal to
ascending, connectives apically pronounced, apiculate or variously shaped,
133
connivent over style head; pollen shed in T-shaped to rhomboidal, decussate
or linear tetrads, sometimes as pollinia. Nectaries in 5 alternisepalous
troughs at base of filaments. Ovary half\semi inferior; carpels 2, free except
towards base and at apex where they are fused and dilated into style head
with 5 embedded, semi-erect to erect translators consisting of spoon- to
cornet-shaped receptacle (onto which pollen or pollinia is shed), with small
sticky viscidium at lower end; style head enclosed in corolla tube or exposed
from it at anthesis. Fruit a pair of cylindrical-ovoid follicles or a solitary follicle
by abortion. Seeds many, compressed, obliquely ovate, elliptic to oblong with
tuft of hairs at the micropylar end, seldom with a ring of hairs extending
around the entire margin or a membranous wing. Basic chromosome number
X = 11 (diploid 2n = 2x = 22).
134
GENERAL CONCLUSIONS:
The Periplocoideae are more derived than the Apocynoideae but less
advanced than the Asclepiadoideae.
Within the Periplocoideae, the usual state is corolline corona lobes situated
outside the stamens (Klackenberg 1997), except for Malagasy sp. -
Baroniella Costantin & Gallaud having the largest lobes between the stamens
and not opposite to them. The corona in Asclepias L. is primarily annular i.e.
the staminal and interstaminal corona in Asclepiadinae is basically annular
(Kunze 1997). Corolline coronas are widespread in the Periplocoideae.
These are situated primarily in the petal sinuses. This kind of corona has
been observed in some Apocynaceae. No staminal corona occurs in the
Periplocoideae, but are found in the Asclepiadoideae. According to Liede &
Kunze (1993), annular corona (and the annulus) is not homologous with
corolline corona. Corolline corona almost certainly represents the
plesiomorphic type of corona formation. The corona is closely associated with
pollination apparatus, thus more close to the androecium than the corolla and
it is optically attractive and serves as a holding device for pollinators (Endress
1994).
The stamens each have a small, more or less cylindrical filament beneath the
anthers (Endress & Bruyns 2000). These filaments are inserted at different
heights on the corolla tube but always arise on the apex of a thickened ridge.
The number of microsporangia in the stamen has been an important feature
employed in the classification of the family s./. The genera in the
Periplocoideae are easily distinguished by their 4-celled anther from the rest
of the Asclepiadoideae except for the Secamoneae. The 4-celled anther is
135
considered to be plesiomorphic and the 2-celled anther in Asclepiadoideae
more derived (Swarupanandan et al. 1996).
Baseonema is closely allied to Schlechterella K. Schum. and Raphionacme
Harv. due to their multi-parate pollen grains. Thus these three genera are
more advanced than any other taxa within the African Periplocoids, most of
which are characterised by presence of 4-6 pores per pollen grain. On the
other hand, Baseonema is separated from the two geophytes with root-tubers,
by having an annular corona formed from the fusion of staminal elements,
quite distinct from the fusion in Schlechterella whereby an outer corona is
basally fused into an annulus at the mouth of corolla tube. Baseonema is the
least developed within the tribe - Cryptolepideae due to its lack of a distinct
upper corolla tube. In order to justify its position (instead of being in the tribe -
Periploceae) the gynostegium is included near the base of the corolla and
reflexed at anthesis thus exposing the gynostegium.
Batesanthus with its rotate flower with or without filiform corona lobe is
closely associated to Baseonema, these two genera have a reflexed corolla
tube. Bintrusus with a fringe of hairs around the entire seed margin is closely
associated to Finlaysonia obovata Wall. Thus B. intrusus is more derived
within the species and other taxa investigated, all with comose seeds (Table
14.1). The presence of outer corona lobes in one specimen of B. pervittorus
is likely due to genetic modifications. Except for the flower with corona lobes
and being geographical isolated, specimens of B. pervittorus are
morphologically similar.
Mangenotia is closely associated with Cryptolepis by having a distinct upper
corolla tube but separated from the latter by presence of hairy anthers, a
character also observed in species of Periploca together with the presence of
papillae globule on the lower leaf surface instead of striated cuticle. Periploca
can be separated from Mangenotia by its trisegmented, filiform outer corona
lobes.
136
The presence of a well developed corolla tube in Mangenotia makes it more
derived than other taxa with an open, shallow upper tube where pollinators
can easily access the nectar - thus more primitive. An advanced tube as in
Mangenotia, Cryptolepis and Raphionacme can only be reached by
pollinators with a long proboscis. The genera in the Periplocoideae are
animal/insect pollinated. Fly-pollination with open flowers occurs in
Apocynum (Apocynoideae) and in most of the Periplocoideae (Endress 1994).
Mondia with its two liana species found in moist tropical and subtropical Africa
has rotate to broadly campanulate, showy flowers and a unique obcordate
outer corona. Nevertheless, this kind of corona is closely linked to the
trisegmented corona lobes of Periploca and Raphionacme sp. with a robust
central lobule immediately behind the base of the filaments inserted on the
apex of the staminal foot and two rather smalilobuies lateral to the central
lobe. These emergences are of corolline origin (Kunze 1993). Mondia whitei
is more advanced within infrageneric classification due to the structure of the
outer corona lobe.
Sacleuxia and Sarcorrhiza with their tuberous roots recalls the genera
Tacazzea, Raphionacme, Petopentia and Schlechterella. The rootstock found
in Mondia is of medicinal importance.
Sacleuxia with an indistinct, broadly campanulate tube without an upper
corolla tube has been moved from the tribe: Cryptolepideae and placed under
the Periploceae. S. tuberosa with a hairy ovary is more advanced within the
genus compared to the glabrous ovary found in S. newii.
The flowers of Sarcorrhiza and Zacateza are similar to those of Buckollia,
Periploca, Petopentia and Schlechterella, in that they have a shallow lower
corolla tube with the corona lobes (usually filiform) arising just above the
stamens at or near the corolla mouth or the corona lobes may be fused to the
filament bases. Therefore less advanced within the subfamily.
137
Table 14.1: Characteristics of Selected African Periplocoideae.
Genera Tribe Distribution Habit Seeds Corona Pollen Translators Rootstock
Baseonema Cryptolepideae Semi-arid, Shrub Comose Annular Tetrads Specialized
Continental -Twiner Corolline 8-10 pores
East Africa corona per grain
Batesanthus Periploceae Central North Shrubs Comose. Mostly Tetrads Specialized
Africa, very -Twiner Except B. annular 4-6 pores per
rarely West - Climber intrusus with lobules, very grain
Africa a fringe of rarely filiform
hairs around
entire margin
Mangenotia Cryptolep ideae West Tropical Liana Comose Clavate Tetrads Less
Africa - Climbing 4-6 pores per specialized
or twining grain i.e. no clear I
dermaction
. between spoon &
stalk I
Mondia Periploceae Tropical and Liana Comose Obcordate Tetrads Specialized Thick, I
Sub- Tropical - Climbing with or 4-6 pores per Succulent
Africa or twining without dorsal grain with aromatic
processes scent;
medicinally
important
Sacleuxia Periploceae East Africa Shrub Comose Annular, Tetrads Specialized Tuberous
-Erect obscure 4-6 pores per
was Cryptolepideae grain
Sarcorrhiza Periploceae West-Central Epiphyte Comose Annular Tetrads Specialized Tuberous
and East - Creeping 4-6 pores per
Equatorial grain
Africa, very
rarely West
Africa
Zacateza Periploceae Central Africa Liana Comose Filiform outer Tetrads Less
....... -climbing corona lobes 4-6 pores per specialized
w
00 or twining grain I
Sarcorrhiza and Epistemma D.V. Field & J.B. Hall are outstanding as
epiphytes. A climbing character is considered plesiomorphic, whereas erect,
epiphytic, herbaceous, geophytic, shrubby and xerophytic characters are
apomorphic (Venter & Verhoeven 1997):
Within the African Periplocoideae, pollen morphology is fairly uniform, with
most genera characterised by the presence of tetrads. However, with
accumulation of data, Verhoeven and Venter (1998a) have published ten
genera in which pollinia was found. In this study, there is no recorded
occurrence of pollinia.
All the genera investigated most probably have starchless pollen grains.
According to Verhoeven & Venter (1988), the starchy pollen is a feature of
wind pollinated flowering plants whereas insect-pollinated plants have
starchless pollen grains. The spoon with its adhesive surface receives pollen
tetrads from adjacent anther halves. The adhesive disc functions to stick to a
visiting pollinator.
139
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148
Seven genera of the African Periplocoideae namely: Baseonema Schltr. &
Rendie, Batesanthus N.E. Br., Mangenotia Pichon, Mondia Skeels, Sacleuxia
Baill., Sarcorrhiza Bullock and Zacateza Bullock are presented in this study,
with special reference to taxonomy.
A complete account of taxonomic literature, keys as well as known
geographical distribution of the seven genera is given. The type specimens
(in red) and all other specimens studied are presented in Appendix B.
The character states and/or features used to delimit genera are
circumscribed. Floral and vegetative morphology have been extensively
studied with the aid of a stereo-microscope and represented in the form of
diagrammatic sketches or photographic figures. Floral (i.e. corolla, corona,
androecium and gynoecium) features are important to delimit between
species. For instance, Sacleuxia tuberosa is characterised by having a hairy
ovary and the anthers in Mangenotia are hairy. Mondia whitei has an
obcordate corona lobe with two dorsal processes. Most of the genera co-exist
as shrubs and very rarely lianas, but Sarcorrhiza is outstanding as an
epiphyte.
To complement "traditional taxonomy", pollen and translator morphology, leaf
and seed morphology have been studied, with the aid of LM, SEM and TEM.
The pollen is uniform in morphology, with all the genera having pollen in
tetrads and little variation between the species. However, pollen grain size
can be used to a certain extent so as to distinguish between species, for
example, the largest tetrads are found in species of Baseonema qreqoti! &
Mondia whitei (Figure 4). One other important feature is the number of pores
per grain. Most of the Periplocoids are characterised by having 4-6 pores per
grain. This, however, does not apply to multi-parate Baseonema having 8-10
pores per grain, a character so far only known in species of Raphionacme
149
and Schlechterella. Individual grains of a tetrad are held together by wall
bridges. The exine is smooth and consists of an outer homogenous stratum
(tectum) subtended by a granular stratum. The intine is well developed. The
layers turn out to be of little taxonomic value.
Translators are similar in structure, with three parts distinguished, the spoon,
the stalk and an adhesive disc (viscidium). The whole translator varies in size
(although marginal difference is small) within the species (Table 4.2).
The character state of taxonomic value in the leaf surface is the presence of
papillae in Mangenotia eburnea. The cuticle on the leaf is variously striated or
smooth.
The paracytic stomata and smooth or warty, unicellular trichomes have been
found to be of little taxonomic value especially at generic level. However, at
species level, the amphistomatous condition of Sacleuxia tuberosa might be
of taxonomic value. Trichomes are present on the leaf surface of all taxa
except for Batesanthus parviflorus and Batesanthus purpureus with glabrous
leaf surface.
Seeds are characterised by having a coma of hairs. The exception is
Batesanthus intrusus with a fringe of hairs around the entire margin.
150
Sewe genera van die Periplocoideae uit Afrika, naamlik Baseonema Schltr. &
Rendie, Batesanthus N.E. Br., Mangenotia Pichon, Mondia Skeels, Sac/euxia
Baill., Sarcorrhiza Bullock and Zacateza Bullock is taksonomies ondersoek.
'n Volledige uiteensetting van die taksonomiese literatuur, sleutels sowel as
die geografiese verspreiding van die sewe genera word gegee. Die tipe-
eksemplare (in rooi) en al die bestudeerde eksemplare is in Aanhangsel B
gelys.
Die kenmerke en/of strukture wat gebruik is om die genera te omgrens, is
beskryf. Blommorfologie en vegetatiewe kenmerke is intensief met 'n
stereomikroskoop bestudeer en die resultate is in die vorm van sketse en
foto's aangebied. Blomkenmerke (bv. kroon, bykroon, androesium en
ginoesium) is belangrik vir die onderskeiding van spesies. Sac/euxia tuberosa
word deur 'n harige vrugbeginsel terwyl Mangenotia deur harige helmknoppe
gekenmerk. Die meeste genera kom as struike voor en selde as liane.
Sarrcorhiza kom egter as 'n epifiet voor.
Om die morfologiese ondersoek uit te brei, is stuifmeel, stuifmeeldraers,
blaaroppervlak en sade ook met die LM, SEM en TEM (stuifmeel) bestudeer.
Die tipiese Periplocoideae tetrades kom in al sewe genera voor met relatief
min variasie tussen die spesies. Die meeste Periplocoideae word gekenmerk
deur 4-6 porieë per stuifmeelkorrel. In Baseonema kom 8-10 porieë per
stuifmeelkorrel voor, 'n kenmerk wat tot dusver net bekend was in spesies van
Raphionacme en Sch/echterella. Individuele stuifmeelkorrels van 'n tetrade
word by mekaar gehou deur wandbrue. Die eksien is glad en bestaan uit 'n
homogene stratum (tektum) wat begrens word deur 'n granulêre stratum. Die
intien is goed ontwikkel. Die wandlae van die stuifmeelkorrel is nie van
taksonomiese waarde nie.
151
Blaaroppervlakkenmerke wat van taksonomiese waarde is, is: die
teenwoordigheid van papille in Mangenotia eburnea. Die kutikula op die blare
is gestrieerd of glad.
Die parasitiese stomata en gladde of vratagtige, eensellige trigome is van min
taksonomiese waarde, veralop genusvlak. Op spesievlak kan die
amfistomatiese blare van Sacleuxia tuberosa taksonomiese waarde besit.
Trigome kom op die blaaroppervlak van al die taksa voor met die
uittsondering van Batesanthus parviflorus en Batesanthus purpureus waar
geen trigome voorkom nie.
Sade word gekenmerk deur 'n koma van hare. By Batesanthus intrusus kom
die trigome egter op die rand van die saad voor.
152
Many thanks to the National Research Foundation (NRF) and the University of
the North - Qwaqwa Campus for their financial support. And to the University
of the Free State for granting me this opportunity to study.
With special thanks; to my supervisor Prof. H.J.T. Venter for his guidance,
insight and inputs in my work, it has been a pleasure to go through the type
specimens and be baffled how old but precious they are; to my co-supervisor,
Prof. R. L. Verhoeven, for his never ending patience and insight to guide me
thorough the microscopic world .
. To Dr. Andor Venter and Lintle Mohase, your words of encouragement and
suggestions are appreciated.
Dr. L. du Preez who put together excellent scans and Mr. Gabre Kemp for
taking photographs of the "is this really this old" specimens. All the herbaria
listed are thanked for loaning their specimens.
Prof. Rodney Moffett, the one person who instilled the love of plants,
appreciation and the complex herbarium world, I will for always cherish what
you taught me, you are my inspiration. Thank you for all the e-mails of
encouragement.
To my family, Leteketoa, Mats'episo, Thato, Limpho, Khauhelo, Thuto,
Puleng, Palesa, Bokang and Phetetso, those hundreds of calls usually made
my days; your support is really appreciated. And all those who in their special
ways supported me.
And most important, better is the end than the beginning, to my creator,
Almighty God for his blessings and I believe he made everything possible.
I thank you all.
153
APPEND~X A: GLOSSARY
Abbreviations:
leBN International Code of Botanical Nomenclature v.118 (1988)
LM Light Microscope
SEM Scanning Electron Microscope
TEM Transmission Electron Microscope
sp. Species
t. Tabula (Latin for 'table')
B. gregorii Baseonema gregor;; Schltr. & Rendie
B. intrusus Batesanthus intrusus S. Moore
B. parviflorus - Batesanthus parviflorus Norman
B. purpureus - Batesanthus purpureus N.E. Br.
B. ta/botii Batesanthus ta/bot;; S. Moore
M. eburnea - Mangenot;a eburnea Pichon
M. ecornuta - Mond;a ecournuta (N.E. Br.) Bullock
M. white; Mond;a whitei (Hook f.) Skeels
S. newii Sac/euxia new;; (Benth.) Bullock
S. tuberosa - Sac/euxia tuberosa (E.A. Bruce) Bullock
S. epiphytica - Sarcorrhiza epiphytica Bullock
Z. pedicel/ata - Zacateza pedicel/ata (K. Schum.) Bullock
Ic. icone ( = illustration)
cf. confer ( = compare)
a/. a/ii ( = other).
i.e. id est (= that is)
pp page
AlUlthorCitation:
Baill. Baillon
Benth. Bentham
Decne. Decaisne
K. Schum. Karl Schumann
N.E. Br. N.E. Brown
Schltr. Schlechter
locality:
Alt. Altitude
Distr. District
N North
S South
E East
W West
Mts. Mountain(s)
Dem. Rep. Congo - Democratic Republic of the Congo
Symbol:
c. circa (Latin for '_!: or about')
km kilometer
mm millimeter
s.n. sine numero (Latin for 'without collection number')
!Jm micrometer
vidi (Latin for 'I have seen it')
FHI Forestry Herbariu~ in Ibadan
= Nomenclatural synonym I Legitimate synonym
- Illegitimate synonym
[Latin descriptions from: Jones & Luchsinger 1986 and Stearn 1983]
Terminology:
Abaxial Lower surface
Adaxial Upper surface
Amphistomatie Leaves with stomata present on both surfaces
Epistomatic Leaves with stomata on the upper surface only
Hypostomatic Leaves with stomata on the lower surface only
Halo Holotype
Iso lsotype
Lecto Lectotype
Il
APPENDIX B: SPECIMENLIST
BASEONEMA Schltr. & Rendie
Baseonema gregorii Schltr. & Rendie
KENYA
015 37E, 01°30'5 37°47'E: K4 Machakos/Kitui, Yatta Plateau, lower
Ukamba (-DB). Gardner, H.M. 3612; 00-01-1937 (K). Alt. 4,000 m.
025 37E, 02°02'5 37°21 'E: K6 Masai Distr., Carabani valley (-AB). van
Someren 184; 00-00-0000 (K!).
025 37E, 02°12'5 37045'E: K4 Machakos Distr., 1 km S of Hunters Lodge
on track to Kenya Agricultural research station (-BB). Goyder, D.J.,
Masinde, P.S. Meve, U. & Whitehouse, C. 4006; 11-02-1996 (PRE!). Alt.
1,000 m.
025 37E, 02°25'5 37°57'E: K4 Machakos Distr., Kibwezi (-BO). Kássner
705; 27-04-1902 (BM).
025 3SE, 02°07'5 3S006'E: K4 Kitui Distr., Machakos between Yatta gap
and Ikutha (-AA). Bally, P.R.O. 12320; 14-03-1961 (K!).
025 3SE, 02°32'5 3S007'E: 240 km from Mombasa on Nairobi road (-CA).
Verdcourt & Polhi1l2695; 15-04-1960 (BR, K!).
025 3SE, 02°57'5 3S020'E: K4/6 Machakos/Masai Distr., Kenani and
Ongalea mts. (-CD). Gregory, J.W. 14; 00-00-1893 (BM notol).
035 3SE, 03°22'5 3S035'E: K4 Machakos Distr., Tsavo National Park,
east Simba Hill (-BC). Faden, R.B. & A.J. 74/436; 17-04-1974 (BR, K, MO!,
PRE!, WAG! (2x)). Alt. 600 - 630 m.
035 3SE, 03°23'5 3S034'E: K7 Teita/Kilifi Distr., Voi River (= Goshi) near
Voi (-BC). Bally, P.R.O. 8745; 05-02-1933 (K!). Alt. 1,500 m.
035 3SE, 03°50'5 3S03S'E: K7 Teita Distr., Mt. Kasigau, 4 km south of
Rukanga at eastern base (-DC). Gilbert, M.G. & C.I. 6122; 02-05-1981
(K!).
III
035 39E, 03°10'5 39°19'E: K7 Kilifi Distr., c. 60 km from Malindi on raad
from Sala gate to Malindi (-AB). Field, D. V. & Powys, J.G. 174; 07-07-
1977 (K!).
TANZANIA
035 37E, 03°48'5 37°36'E: T3 Pare Distr., Lembeni (-DC). Bal/y, p.R.a.
8125; 02-04-1952 (K!).
045 38E, 04°16'5 38°32'E: T3 Pare/Lushoto, Mkomazi game reserve
(-BC). Cox & Abdallah 2028; 00-00-0000 (K).
IV
BATESANTHUS N.E. Brown
Batesanthus intrusus S. Moore
CAMEROON
03N 12E, 03001'N 12°22'E: Bit ye, Yaoundé (-AB). Bates, G.L. 1392; 00-
00-1917 (BM holo).
OSN 10E, OSoS6'N 10010'E: Bamenda, Bafut-Ngemba forest reserve
(-CC). Hepper 2173; 25-02-1958 (K).
06N 14E, 06°20'N 14°21 'E: District Barmuda, Lakka (-AD). Mait/and
1565; 00-06-1931 (K). Alt. 1,300 m.
CENTRAL AFRICAN REPUBLIC
03N 18E, 030S3'N 18001'E: Boukoko & Mbaiki regions (-CC). Tisserent.
R.P. 1480; 1948-1950 (P!).
DEMOCRATIC REPUBLIC OF THE CONGO
OON 24E, 00047'N 24024'E: Yangambi, valley de la Luuvea, Isangi
Province (-CD). Louis, J. 12992; 19-12-1938 (BR, K!, MO).
OON 24E, 00034'N 2S001 'E: Yangambi et Yakusu, secondary forest, Isangi
Province (-CA). Germain 4908; 00-05-1949 (BR, SRGH).
OON 2SE, 000S6'N 2S009'E: Bengamisa, Banalia Province (-CC). Louis, J.
623; 15-11-1935 (BR, K). Alt..:t 550 m.
01 N 23E, 01°1S'N 23036'E: Grande lie Esabo, Basoko (-BC). Germain
4951; 00-00-0000 (BR, SRGH).
028 20E, 02°08'8 21°34'E: Iwama, Monkoto territory (-DA). Evrard 2826;
00-00-0000 (BR).
Not traced: Bakoko, Isle of Isabe. Germain 4901; 00-06-1949 (SRGH).
GABON
OON 09E, 00028'N 09020'E: Libreville, on the Gaboon River (-AD). K/aine,
R.P. 513; 19-10-1896 (P holo!, K iso).
v
OON09E, 00028'N 09025'E: Libreville (-AD). Klaine, R.P. 2363; 25-03-
1904 (K).
OOS 12E; 00°50'5 12°43'E: Region Lastoursville, Liyanga (-DC). Le
Testu, M.G. 7696; 28-11-1929 (P!).
OOS 12E, 00°50'5 12°43'E: Region de Lastoursville (-DC). Le Testu,
M.G. 8501; 08-11-1930 (P!).
LIBERIA
07N 08W, 07°32'N 08°38'W: Yéképa villlage (-DA). Adam, J.G. 30433;
13-05-1965 (MO!).
Batesanthus parviflorus Norman
ANGOLA
085 15E, 08°20'5 15026'E: Guanza, Quilela Camabatela (-AD).
Gossweiler, J. 8468; 29-09-1922 (BM notol). Alt. 1,225 m.
CAMEROON
05N 09E, 05002'N 09050'E: Manengouba Lake (-BB). Po/hill, R.M. &
Kirlwp, D.W. 5190; 12-03-1984 (K!). Alt. 1,900 m.
SIERRA LEONE
08N 12W, 08°27'N 12029'W: Near Roruks, northern Province (-BC).
Deighton, F.G. 3265; 13-07-1936 (K!).
Batesanthus purpureus N.E. Brown
CAMEROON
03N 13E, 03027'N 13011'E: Bitja, near River Dja (-AC). Bates, G.L. 1855;
00-00-1894 (BR, K).
Not traced: Efulen, Buie country. Bates, G.L. 383; 20-09-1895 (BM,
BR, G (2x) iso!, K notol).
vi
CENTRAL AFRICAN REPUBLIC
03N 1SE, 030S3'N 1So01'E: Oubangui, region de Mbaiki et Boukoka
(-CC). Tisseranf, C. 1480; 13-05-1949 (P! (2x)).
GABON
01N 11E, 01°46'N 11°17'E: Haute Ngounié (-CD). Le Tesfu, M.G. 5493;
03-05-1928 (P!).
Batesanthus ta/botii S. Moore
CAMEROON
04N 09E, 04°09'N 09013'E: Bura/Buea (-AA). Moore, T.S. 666; 00-05-
1929 (K). Alt. 1,100 m.
NIGERIA
OSN OSE, OSo17'N OSo33'E: Oban (-BC). Ta/bott, P.A. 63; 00-00-1911
(BM halo, K! (2x»).
OSNOSE,OSo19'NOSo34'E: Oban (-BC). Ta/bott, T.A. 2074; 00-00-1912
(BM).
04N 07E, 04°39'N 07°SS'E: Eket District (-DB). Ta/bott, T.A. & P.A. 2021;
10-05-1912 (K!).
vii
MANGENOTlA Pichon
Mangenotia eburnea Pichon
DEMOCRATIC REPUBLIC OF THE CONGO
Not traced: Moyen Congo, River Congo (= River Zaire). Pooenquin 173;
00-02-1920 (P).
GHANA
OSN 01W, OSo10'N 01°13'W: Cape Coast (-AA). Hall, Ja. 1010; 08-08-
1956 (K). Alt. 50 m.
OSN 01W, OSo06'N 01°15'W: Cape Coast (-AA). Hall, Ja. 409; 26-12-
1957 (K). Alt. 50 m.
05N 01W, 05010'N 01°13'W: Cape Coast (-AA). Hall, Ja. 1511; 07-07-
1959 (K). Alt. 30 m.
05N 01W, c. 05033'N 01°04'W: Lac Baku, Baku reserve (-CA).
Anonymous 209012; 19-08-1963 (G).
07N 01W, 07°12'N 01°26'W: Mampong scarp (-AB). Vigne 1739; 00-05-
1929 (K). Alt. 430 m.
07N 01W, 07°13'N 01°28'W: 55 km Mampong Ash (-AB). Darko, K.O.
689; 28-07-1952 (BR, K photo!, MO, P).
IVORY COAST (COTE d'IVOIRE)
05N 03W, 05016'N 03052'W: Forest L'Abouabou, between Abidjan and
Grand Bassam, coastal rain forest (-BD). Leeuwenberg, A.JM. 2654; 04-
11-1959 (WAG!). Alt. 2 m.
05N 03W, 05016'N 03052'W: Forest L'Abouabou, between Abidjan and
Grand Bassam, coastal rain-forest (-BD). Leeuwenberg, A.J M. 4235; 23-
05-1962 (P, WAG!). Alt. 2 m.
05N 04W, 05019'N 04°01'W: Forêt d'Adiopodoumé (-AC). Aké Assi 6990;
11-10-1960 (P photo!).
viii
OSIN!04W, OS019'1NI04°01'W: Abidjan (-AC). Gremers 358, 519; 00-10-
1966 (BR).
05N 04W, OS019'N 04°01'W: Forêt d'Adiopodoumé, prés d'Abidjan (-AC).
Mangenot s.n.; 07-06-1951 (P).
OSINI04W, OS011'1Nl04°32'W: :!: 10 km W of Jacqueville, Island Aladdin,
old secondary forest (-IBA). Leeuwenberg, A. J.M. 8083; 03-08-1970 (BR,
WAG!).
G71N104W, 07°27'1NI 04°20'W: Baouké north, Nzi Mbahiakro (-AD).
Chevalier, A. 22282; 04-08-1909 (P).
lNIot traced: Bonake, Orumbo-Boka. Garner 72; 10-07-1963 (K).
UIBER~A
07N 09W, 07°47'1NI09026'W: Zorzor, Gbarnga road, west of St. Paul River
(-CD). Bos 2152; 27-07-1966 (K, WAG).
IN!~GER~A
061N!031E,06°28'1NI 03023'E: Lagos, Skoyi plains (-AID). Dalziel, J.M. 1405;
18-08-1919 (K, MO!).
SIÉNIÉGAl
121N116W, 12°33'1NI16°01'W: Region Zinguinchor, Bissine (-CA). Berhaut,
R.P. 6449; 13-11-1963 (BR, P).
121N116W, 12°3S'1NI 16°16'W: Region Zinguinchor, (-CIB). Berhaut, R.P.
6323; 08-08-1963 (P).
121N116W, 12°4S'1NI16°13'W: Region Bignona, Tanguéme (-CC). Berhaut,
R.p. 6436; 11-11-1963 (P).
12N 16W, 12°48'N 16°1l8'W: Region de Bignona, Basse Casamance,
forêt des Kalounayes (-CD). Berhaut, R.P. 5958; 09-05-1963 (P!).
121N11SW, 12°29'1NI 1S036'W: Region d'Oussouye - Dianthéme (-DA).
Berhaut, R.P. 6550; 20-11-1963 (P).
IX
12N 16W, 12°40'N 16°4S'W: Loc Basse, Casamance, Djibonker (-DB).
van den Berghen 5378; 00-05-1982 (BR).
14N 16W, 14°19'N 16°27'W: Village Fatick (-AD). Adam, J.G. 30434; 05-
10-1962 (MO photo!).
SIERRA LEONE
07N 12W, 07°42'N 12°19'W: Between Mattru and Gbangbama (CB).
Deighton, F.C. 2349; 17-11-1931 (K).
07N 12W, 07°47'N 12°18'W: Gbangbama (-CD). Thomas, N.W. 4077;
30-10-1914 (K!).
08N 11W, 08°39'N 11°S1'W: Matotaka (-DB). Thomas, N.W 1238; 29-
07-1914 (K). Alt. 120 m.
08N 12W, 08°30'N 12°14'W: Yonibana (-CB). Thomas, N.W 4119; 30-
10-1914 (K).
08N 12W, 08°30'N 12°14'W: Yonibana (-CB). Thomas, N.W 4202; 30-10-
1914(K!).
08N 12W, 03°46'N 12°47'W: Port Lokoh (-DD). Thomas, N.W 6585; 15-
12-1914 (K). Alt. 15 m.
08N 12W, 03°46'N 12°47'W: Port Lokoh (-DD). Thomas, N. W 6602; 15-
12-1914 (K).
08N 13W, 08°31'N 13°20'W: Sukudu, Konno (-CB). Oawe 543; 00-08-
1923 (K).
08N 13W, 08°S6'N 13°13'W: Mayoso/Mayoro (-CC). Thomas, N.W. 1373;
06-08-1914 (K!).
Not traced: Mamaka. Thomas, N.W. 4377; 00-00-1914 (K).
Not traced: Mamaka. Thomas, N.W. 4427; 02-11-1914 (K!).
Not traced: Mamaka. Thomas, N.W. 4544, 4546; 02-11-1914 (K!).
Not traced: Mamaka. Thomas, N.W. 4628; 02-11-1914 (K!).
x
MONDIA Skeels
Mondia ecomuta (N.E. Br.) Bullock
DEMOCRATIC REPUBLIC OF THE CONGO
035 14E, 03°57'5 14038'E: Galerie forestiére de la Loualou, village de
Kimpélé, km 16 route de Mayama - Mouyondzi (-DC). Bouquet, A. 631;
05-11-1964 (P!).
GABON
005 12E, 00°50'5 12°43'E: Lastoursville (-DC). Le Testu, M.G. 8865; 18-
04-1932 (P!).
OON09E, 00030'N 09° 25'E: Libreville (-CB). Klaine, R.P. 577; 14-11-
1896 (P!).
KENYA
035 39E, 03°44'5 39042'E: K7 Kilifi District, Chasimba (Chonyi) (-DC).
Musyoki, B.M. & Hansen, O.J. 956; 05-06-1973 (BR!). Alt. 200 m.
03S 39E, 03°53'5 39038'E: K7 Kilifi Distr., Ribe near Mombasa (-DC).
Wakefield, T. s.n.; 00-05-1880 (K holo!).
045 39E, 04°37'5 39022'E: K7 Kwale Distr., 4 km west of Shimani (-CB).
Luke & Robertson 2352; 30-05-1990 (MO). Alt. 500 m.
MOZAMBIQUE
Not traced: Portuguese East Africa, Misala River, Hynesa. Alien, C.E.F.
139; 00-01-1912 (K!).
TANZANIA
055 38E, c. 05°06'5 38°39'E: T3 LushotolTanga, Sigi valley (-BA). Grote
5795; 00-07-1913 (K!). Alt. 500 m.
055 38E, 05°08'5 38°39'E: T3 Lushoto, Amani (-BA). Burlt, B.D. 278;
04-04-1926 (K).
XI
055 38E, 05°08'5 38°39'E: T3 Lushoto, Amani (-BA). Saleman, R. 6222;
13-03-1922 (K).
055 38E, 05°20'5 38°57'E: T3 Pangani Distr., Bushiri estate (-BA).
Fa ulkn er, H.G. 558; 06-05-1950 (K! (x2), BR!). Alt. 50 m.
055 38E, 05°20'5 38°57'E: T3 Pangani Distr., Bushiri estate (-BA).
Faulkner, H.G. 558 (2); 22-11-1950 (K!). Alt. 50 m. [NB: 2nd collection
with follicles, from the same plant as the flowers].
055 38E, 05°24'5 38°S7'E: T3 Pangani, Kumbamtoni (-BD). Tanner,
R.E.S. 1990; 23-07-1955 (K!). Alt. sea level
075 36E, 07°57'5 36°31'E: T6 Ulanga Distr., lIingera near Ifakara (-DC).
Haerdi 266/0; 00-06-1959 (K!, WAG!). Alt. 700 m.
075 37E, c. 07°00'5 37°48'E: T6 Morogoro Distr., Kimbaza (-BB). Parda
319; 30-03-1954 (K).
Mondia whitei (Hook. f.) Skeels
ANGOLA
075 13E, 07°50'5 13°09'E: Luanda, Abriz, Camilemba (-CC). Monteiro,
R., Santos, R. & Murta 397; 07-12-1958 (CO).
085 14E, 08°50'5 14°30'E: District Cuanza Norte, location Colungo
(-DC). Da Si/va, M. 2313; 19-01-1968 (CO). Alt. 550 m.
095 14E, 09°10'5 14°45'E: Colungo Alto (-BB). Gossweiler, J. 4211; 00-
00-0000 (P).
095 14E, 09°10'5 14°45'E: Colungo Alto (-BB). Welwitseh, F. 4217; 00-
00-0000 (BM).
095 14E, 09°41'5 14°25'E: Dondo, River Luachima (-CB). Gossweiler, J.
13856; 20-11-1946 (B, CO, K, P).
095 15E, 09°15'5 15°00'E: Cazengo (Casenga) (-AC). Gossweiler, J.
616; 00-00-0000 (BM, K, P).
095 15E, 09°44'5 15°35'E: Pungo Andongo (-DC). Welwitseh, F. 4218,
4219; 00-11-1878 (BM, K).
xii
125 13E, 12°34'5 13°24'E: BenguelIa District, Bumbo (-CB). Welwitseh,
F. 4221; 00-10-1859 (BM).
135 17E, 13°24'5 17°21'E: Bie Distr., Maxito do Chilesso Canata (-AD).
Teixeira 9962; 10-12-1965 (USC). Alt. 1,640 m.
145 19E, 14°54'5 19°07'E: Cababa, Kakonda, Benguela (-CC).
Gossweiler, J. 4288; 01-03-1907 (CO).
Not traced: Venaut. De Ficalho s.n. 00-00-1882 (P).
Not traced: Queta mts., near Sange. Welwitseh, F. 4211,· 00-11-1878
(BM, G, K).
BURUNDI
Not traced: No locality. Lewalle, J. 921; 00-10-1980 (K).
CAMEROON
03N 11E, 03°53'N 11°27'E: N'kolbison, 2: 8 km west of Yaoundé (-CD).
de Wilde & de Wilde Duyfies 2621; 24-05-1964 (BR, K, MO, P, SRGH, Z).
Alt. 650 m.
03N 11E, 03°53'N 11°27'E: N'kolbison, 2: 8 km west of Yaoundé (-CD).
de Wilde & de Wilde Duyfies 2621 b; 24-05-1964 (MO, WAG). Alt. 650 m.
03N 11E, 03°51'N 11°31'E: Yaoundé (-DC). Zenker 589; 2: 1896 (K!).
03N 11E, 03°S1'N 11°31'E: Yaoundé (-DC). Zenker 1397; 00-00-1897
(BM, G, K, P, WAG, Z holo?).
03N 15E, 03°56'N 15°07'E: 41 km off road Carigombo (Garegoumo) to
Soso, east of Mgbatouri (=Batouri) River, near Bayanga Kadei (-CC).
Leeuwenberg, A.J.M. 6244; 26-07-1965 (K, P, WAG). Alt. 540 m.
03N 18E, 03°S3'N 18°01'E: Region de Mbaiki, Boukoko (-CC). Equipe
1547; 25-07-1949 (BM).
03N 18E, 03°S3'N 18°01'E: Region Mbaiki, locality Centrale de Boukoko
(-CC). Le Testu, M.G. 1541; 25-07-1949 (BM).
03N 18E, 03°53'N 18°01'E: Region Mbaiki, locality Centrale de Boukoko
(-CC). Le Testu, M.G. 2102; 25-07-1949 (BR).
XIII
04N 12E, 04°41'N 12°22'E: Menyang, Nanga Eboko (-CB). Letouzey, R.
2077; 21-05-1959 (P).
04N 13E, 04°S4'N 13°S0'E: 29 km off road Bertoua - Betare Oya (-DD).
Leeuwenberg, A.J.M. 5802; 09-06-1965 (P, WAG). Alt. 750 m.
04N 14E, 04°07'N 14°S2'E: Banga, 65 km SE of Batouri (-BB). Letouzey,
R. 4894; 00-00-0000 (BR, K, P).
04N 14E, c 04°S8'N 14°03'E: Yangamo, 65 km NNW of Batouri (-CC).
Letouzey, R. 2494; 03-05-1962 (BR, K, P).
OSN 10E, OS028'N 10000'E: Village Djang, 40 km west of Bertona (-AC).
Brete/er 2947; 15-05-1962 (BR, K, M, P, WAG). Alt. 650 m.
06N 11E, 06°42'N 11°S6'E: 10 km SSE of Banya (-DB). Bihotonq, M.
186; 08-06-1967 (P).
06N 12E, 06°22'N 12°20'E: Amani (-AD). de Ber/in 346; 00-00-1903 (P).
06N 12E, 06°26'N 12°42'E: 10 km Tibati-Mabouka road, grove in
woodland near Djaoro Ndo (-BC). Leeuwenberg, A.J.M. 10026; 29-06-
1972 (K!).
06N 13E, 06°47'N 13°10'E: Tekel, 40 km NNE from Bagodo (-CC).
Letouzey, R. 4417; 17-07-1966 (P).
06N 14E, 06°31'N 14°18'E: Meiganga et Betuie (-CB). Jacques-Fe/ix, H.
4521; 00-07-1939 (P).
08N 13E, 08°20'N 13020'E: Gere (-AD). Letouzey, J. 2294; 29-06-1959
(K, P).
09N 13E, 090S7'N 13°42'E: Bosum-Buar (-DC). Mi/dbraed 9731; 00-06-
1914 (K photo!). Alt. c. 900 m.
CENTRAL AFRICAN REPUBLIC
03N 1SE, 030SS'N 1S024'E: 1 km road Garigombo (on Mopas
Garegoumo) - Soso, east of Mgbatouri (=Batouri) River near Bayanga
Kadei (-CD). Leeuwenberg, A.J.M. 6244; 26-07-1965 (BR, MO). Alt. 540
m.
xiv
03N 17E, 03°S4'N 17°S6'E: Boukoko (-DD). Tisserant, Cl. Tis. 1541; 00-
00-0000 (G).
03N 17E, 03°S4'N 17°S6'E: Boukoko (-DD). Tisserant, Cl. Tis. 1752; 00-
00-0000 (P).
03N 17E, 03°S4N 17°S6'E: Boukoko (-DD). Tisserant, Cl. Tis. 2102; 00-
00-0000 (P).
03N 18E, 03°S3'N 18°01'E: Mbaiki, station centrale de Boukoko (-CC). Le
Testu, M.G. 1488; 20-05-1949 (BR).
04N 21E, 04°31'N 21°31'E: La Mahake (-DA). Badre, F. 51; 24-05-1968
(P).
OSN 20E, OS040'N 20037'E: Region de Bambari (-DA). Tisserant, T.P.
2204; 29-06-1927 (P).
OSN 20E, OS040'N 20037'E: Region de Bambari (-DA). Tisserant, T.P.
2205; 29-06-1927 (P).
OSN 2SE, OSoOS'N2S001'E: Ombella, Porte Boma (Drouma) (-AA).
Chevalier, A. 5943; 28-10-1902 (P).
06N 22E, 06°2S'N 22°02'E: Tchekeni, Tchéné, River Mluyo (-AC). Le
Testu M.G. 2204; 00-00-0000 (BR).
06N 22E, 06°2S'N 22°02'E: Tchekeni, Tchéné, River Mluyo (-AC). Le
Testu M.G. 2205; 00-00-0000 (BR).
06N 23E, 06°33'N 23°14'E: Yalinga (-CB). Le Testu, M.G. 3957; 11-06-
1923 (BR).
06N 23E, 06°33'N 23°14'E: Yalinga (-CB). Le Testu M.G. 4770; 11-06-
1923 (BR).
DEMOCRATIC REPUBLIC OF THE CONGO
OON 29E, 00027'N 29°29'E: Beni (-BC). Bequaert 5114; 29-07-1914
(BR).
OON29E, 00047'N 29°36'E: Cocalan - Jason village (-DC). Leconte, H.
s.n. 31-01-1864 (P).
xv
02N 29E, 02°45'N 29°03'E: Wamba et Gambari (-CC). Lebrun, J. 3307;
00-07-1931 (BR, K).
02N 19E, 03°24'N 19°47'E: Libenge & Gemena (-BO). Lebrun, J. 1855;
00-12-1930 (BR, K).
03N 20E, 03°45'N 20029'E: Bodangabo (-DC). Evrard, C. 308; 20-02-
1955 (BR).
03N 23E, 03°28'N 23°46'E: Brousse, La Kulu, Aketi Province (-BD).
Braudi, F. 570; 04-08-1931 (BR).
03N 23E, 03°28'N 23°46'E: Brousse, La Kulu (-BO). Braudi, F. 602; 04-
08-1931 (BR).
03N 30E, 03°51'N 30018'E: Kurukwata (Aba), Faradje Province (-CD).
Gérard 3170; 02-05-1957 (BR).
04N 25E, 04°26'N 25°51'E: Tukpwo, Ango Province (-BD). Gérard 4041;
03-07-1959 (BR).
03N 25E, 03°28'N 25°43'E: Bambesa (-BC). Gérard 5139; 05-06-1962
(BR).
03N 26E, 03°09'N 26°53'E: Omadi et Poko (-BB). Lebrun, J. 3081; 00-
06-1931 (BR, K).
03N 26E, 03°38'N 26°56'E: Wangaba (-DB). Seref, F. 876; 00-05-1885
(BR).
03N 27E, 03°53'N 27°45'E: Village Aragi, Niangara Province (-DD). de
Graer, P. 842; 15-06-1937 (BR).
03N 30E, 03°10'N 30040'E: Kiliko, route Aru Aba - Concession (-BA).
Bamps218; 11-06-1958 (BR). AIt.1,160m.
025 18E, 02°50'5 18°23'E: Bokoro (-CD). Jans, E. s.n.; 26-11-1947
(BR).
045 19E, 04°07'5 19°37'E: Ipamu, Idiofa Province (-BA). Vanderysf, H.
12825; 00-12-1922 (BR).
055 13E, 05°50'5 13°03'E: Luki (Boma), station forestier du Mayumbe,
Lukuia Province (-CC). Wagemans, J. 781; 15-03-1954 (BR).
xvi
055 13E, 05°52'5 13003'E: Leopoldville, Luki (Boma) (-CC). Wagemans,
J 979; 00-00-0000 (MO).
055 14E, 05°15'5 14°52'E: Bas Zaire, Mbanza-Ngungu (-BO). Lisowski,
S.56025; 08-01-1980 (BR).
055 14E, 05°27'5 14°54'E: Leopoldville Province, territory Thysville,
M'vuazi, forest de Nkolo (-BD). Oevred, R. 415; 08-12-1948 (BR, K).
055 14E, 05°51'5 14°28'E: Locality Kivala, territory Thysville (-DD).
Compere, D. 773; 06-02-1959 (BR).
055 15E, 05°08'5 15°09'E: Kisantu (Inkisi-Kisantu) (-AA). Gillet, J 230;
00-00-1899 (BR).
065 22E, 06°49'5 22°54'E: Route Bena-Kalangala, Luti (-DD). Liben, L.
2425; 07-02-1957 (SRGH).
065 23E, 06°12'5 23°23'E: Locality route Miabi, territory Bakwanga, Mbuji
Mayi Province (-AB). Liben, L. 1942; 20-11-1956 (BR).
,.
075 23E, 07°01'5 23°27'E: Locality Mwene Ditu (-AB). Liben, L. 3936;
04-11-1957 (BR, SRGH).
Not traced: Moungali. Bouquet, A. 1266; 05-03-1965 (P).
GABON
015 12E, 01°19'5 12°15'E: Movila et Nolende (-AD). Le Testu, M.G.
5152; 22-12-1924 (P).
GHANA
05N OOW,05°59'N 00040'W: Atewa Range forest reserve (-DC). Hall &
Loek 46767; 28-06-1977 (K, MO). Alt. 400 m.
06N OOW,06°39'N 00029'W: Kabakaba east forest reserve (-CB). Hall &
Alobin, G.C. 43496; 17-06-1972 (K).
GUINEA
07N 08W, 07°39'N 08°30'W: Yéképa, Nimba (-DA). Adam, JG. 24248;
00-00-0000 (MO).
xvii
07N oaw, 07°52'N oao29'W: Nzerekore - Lola (-DC). Adam, JG. 25921;
13-06-1970 (MO, SRGH).
07N oaw, 07°45'N oao49'W: Nzérékoré, Sibamon (-DD). Adam, JG.
5420; 00-00-0000 (MO).
OaN 09W, Oa031'N 09°32'W: Macenta, Seredou (-DA). Adam, J.G.
12001; 00-00-0000 (MO).
oaN 09W, Oa031'N 09°32'W: Macenta, Tenebadou (-DA). Adam, JG.
4982; 00-00-0000 (MO).
iON 11W, 10036'N 11°S1'W: Timbo (-DB). Pobeguin, M. 1658; 24-03-
1908 (P).
iON 12W, 10004'N 12°S1 'W: Kindia (Benna) (-BB). Jacques-Fe/ix, H.
1632; 00-06-1937 (K, P).
iON 12W, 10004'N 12°S1'W: Kindia (Benna) (-BB). Jacques-Fe/ix, H.
1750; 00-06-1937 (K, P).
11N 09W, 11°26'N 09°24'W: Near Bafing River on Mau-Touba road
(-BD). Co/enette 55; 04-07-1926 (K). Alt. 500 m.
GUINEA BISSAU
11 N 14W, 11°47'N 14°11 'W: Bafata, Madina fe Mamadii Alfa (-CC).
Santo 2977; 15-11-1952 (K).
11 N 14W, 11°36'N 14°SS'W: Fulacunda, entre Balala & Buba (-DB).
Santo 2155; 01-08-1945 (K, P).
11N 15W, 11°43'N 1S003'W: Fulacunda (-CA). Santa 2161; 09-08-1945
(K, P).
11N 15W, 11°56'N 1S04a'W: Bissau, Bijimita (-DD). Santa" 1760; 03-02-
1945 (K).
12N 15W, 12°04'N 1So23'W: Cantubó (-AB). Perreira, J. 3114; 10-08-
1962 (BR).
xviii
IVORY COAST (COTE d'IVOIRE)
06N 04W, 06°30'N 04°37'W: Tiémélékro (-DA). Garner, P.G. & UB. 22;
23-06-1963 (K).
06N 04W, 06°30'N 04°37'W: 1 km south of Tiémélékro (-DA). Garner,
P.G. & UB. 142; 08-09-1963 (K).
06N OSW, 06°34'N OSo01'W: Lamto reserve, 50 km south of Toumodi
(-CA). Bokdam, J. 2767; 12-06-1968 (BR, MO, WAG).
07N 03W, 07°07'N 03°21 'W: Abo-N'guessan kro (-AB). Garner 41; 07-07-
1963 (K).
07N 04W, 07°48'N 04°S0'W: Brobo, 25 km ENE of Bouaké gallery forest
(-DD). Oldeman, R.A.A. 392; 24-09-1963 (BR, K photo!, MO!, P).
07N OSW, c. 07°14'N OS018'W: 12 km route to Sakasso (Sakassou) (-AD).
Garner, P.G. & UB. 8; 09-08-1963 (K).
07N OSW, c. 07°14'N OS018'W: 12 km route to Sakasso (Sakassou) (-AD).
Garner, P.G. & UB. 96; 09-08-1963 (K).
07N OSW, 07°42'N OSoOO'W: Bouaké (-CA). Céser 1667; 25-05-1982
(ALF).
07N 07W, 07°34'N 07°10'W: Man-Seguela, 55 km (-CA). Bamps 1917; 2-
01-1969 (BR).
08N 02W, 08°03'N 02°4S'W: North of Boudoukou (-BB). Gerling, C. &
Bokdam, J. 253; 14-07-1967 (BR, WAG).
08N OSW, 08°06'N OS026'W: Entré Marabadiassa et Gottoro (-AB).
Chevalier, A. 22025; 04-07-1909 (P).
08N 06W, 08°01'N 06°09'W: Mankono, between Oialakoro and Kénégoré
(-AA). Chevalier, A. 21975; 01-07-1909 (K, P holo).
KENYA
OON 32E, 00022'N 32°38'E: Kiwatule, Kyadondo (-BC). Liebenberg,
L. C.C. 840; 00-05-1929 (K).
OON 34E, c. 0001S'N 34°S2'E: K5 North Kavirondo, Kakamega (-BD).
Dale, I.R. 3431; 00-05-1935 (BR, K).
xix
015 36E, 01°17'S 36°49'E: K4 Nairobi (-BD). Be1/1; 26-03-1952 (K).
LIBERIA
07N 08W, 07°35'N 08°32'W: Yeképa village (-DA). Adam, J.G. 21462;
11-06-1965 (K, MO). Alt. 450 m.
07N 08W, 07°35'N 08°32'W: Yéképa village (-DA). Adam, J.G. 25739;
11-06-1965 (MO, SRGH). Alt. 450 m.
07N 08W, 07°35'N 08°32'W: Yékepa village, Granfield (-DA). Adam, J. G.
26800; 00-00-0000 (MO).
07N 08W, 07°39'N 08°30'W: Nimba (-DA). Val/ah 107; 15-05-1965 (K).
MALAWI
105 33E, 10°59'5 33050'E: Rumpi (Rumphi) District, 3,2 km up Lura
escarpment (-DD). Pawek, J. 13910; 26-02-1978 (K). Alt. 840 m.
105 34E, c. 10°40'5 34°04'E: Lura-Chiwe road, 23 km SE of Chilumba
(-CA). Pawek, J. 13590; 08-01-1978 (K, SRGH).
115 34E, c. 11°36'5 34°18'E: Nkhata Bay secondary school (-CB).
Pawek, J. 5849; 01-10-1972 (K).
115 34E, c. 11°36'5 34°18'E: Nkhata region (-CB). Pawek, J. 6417; 04-
02-1973 (K, MO, SRGH).
155 35E, 15°15'5 35°18'E: Malosa mountain, opposite Domasi valley
(-AD). Sa!ubeni, Banda & Masiye 2692; 00-00-0000 (M).
155 35E, 15°21'5 35°18'E: South region lower slopes of Zomba plateau,
near Mlunguzi stream (-AD). Brummitt, R.K. & Seyani, J.H. 14810; 06-
03-1977 (K, SRGH!). Alt. 1,085 m.
155 35E, 15°23'5 35°19'E: Zomba (= Samba) (-AD). Brummitt & Pate!
15501; 19-04-1980 (K). Alt. 975 m.
155 35E, 15°23'5 35019'E: Zomba, Malemia road near Chichewa office
(-AD). Chirambo, P.C. 102; 04-11-1978 (SRGH).
155 35E, 15°23'5 35°17'E: Zomba Botanical Garden (-AD). Sa!ubeni,
A.J. 2399; 05-01-1979 (MO, SRGH, WAG).
xx
155 35E, 15°23'5 35°19'E: Zomba Botanical Garden (-AD). Salubeni,
A.J 2698; 05-01-1979 (MO, WAG).
155 35E, c. 15°23'5 35019'E: Zomba District (-AD). Wiehe, N. 404; 00-
11-1949 (K, SRGH).
155 35E, c. 15°40'5 35°05'E: Shire Highlands (-CA). Buehanan, J 168;
00-12-1881 (K). Alt. ~ 915 m.
155 35E, 15°29'5 35°14'E: Ntondwe (= Thondwe) (-CB). Cameron, K.J
110; 21-10-1905 (K).
MOZAMBIQUE
145 39E, 14°59'5 39°50'E: Niassa District, Meconta - 21 km from
Meconta para Corrane (-DD). Torre & Paiva 10056; 18-01-1964 (USC).
165 39E, 16°11'5 39°55'E: Antonio Enes, Boila ~ 60 km from Nametil
(-BB). Torre & Correira 17836; 25-01-1968 (USC). Alt. 80 m.
195 34E, 19°56'5 34°03'E: District Manica é Sofala, Tsetserra area, raad
to Mavita (-CC). MOller, T. 511; 30-11-1968 (K, SRGH).
085 31E, 08°47'5 31°23'E: District Huamba, location Chianga (=
Chinhanda) (-CD). Teixeira & Andrade 6330; 30-10-1962 (COl).
NAMIBIA
175 23E, 17°46'5 23°10'E: Oos Caprivi, 32 km van Singalamwe op pad
na Katima Mulilo (-CC). Kil/iek, O.JB. & Leistner, O.A. 3271; 03-01-1959
(PRE).
NIGERIA
06N 03E, 06°27'N 03°28'E: South Nigeria, Lagos (-AD). Batton-Poole
s.n.; 00-00-1946 (K).
07N 3E, 07°13'N 03056'E: Ibadan District, Oke Ado on Ibadan to Ijebu.
Ode motor road (-BB). Tamajong FHI 16796; 17-07-1946 (K).
07N 03E, c. 07°13'N 03°36'E: West of Ibadan, road to Akufo (-BC).
Gentry, A. & Pilz, G. 32963; 00-00-0000 (MO).
xxi
07N 03E, 07°26'N 03°37'E: Olokomeji forest reserve (-BC). Gentry, A. &
Pilz, G. 32664; 00-00-0000 (MO).
07N 03E, 07°23'N 03°56'E: Ibadan District (-BD). Binuyoi & Emwiogbon,
FHI 54548; 00-00-0000 (P).
07N 03E, 07°23'N 03°56'E: Ibadan District, Eleyele plantation by Agbaje's
farm (-BD). Keay, R.WJ. FHI 25675; 00-00-0000 (K).
07N 03E, 07°23'N 03°56'E: Ibadan (-BD). Keay, R.WJ. FHI 46310; 01-
07-1962 (K).
07N 03E, 07°23'N 03056'E: Ibadan, garden of # 2, Jericho road (-BD).
Keay, R.WJ. FHI 46320; 01-07-1962 (K photo!).
07N 03E, 07°23'N 03056'E: Ibadan Distr., near Busogboro (-BO).
Onochie 34943; 04-11-1955 (K).
07N 03E, 07°23'N 03°56'E: South Ibadan, Oke Eleyele (-BD). Sutton s.n.;
15-07-1951 (K).
07N 03E, c. 07°45'N 03°55'E: Ibadan Distr., LLLA. compound (-DD).· Wit,
P. 2227; 03-07-1972 (K).
07N 04E, 07°13'N 04°10'E: West Province, Gambari j, 32 km south east
~
of Ibadan (-AA). van Eijnatten, eLM. 1689; 19-07-1966 (WAG x 2).:
07N 04E, 07°05'N 04°55'E: Ondo state, aka District (-BB). Daramola,
B.O. FHI 91209; 00-00-0000 (MO).
07N 05E, 07°24'N 05003'E: Igbara Oke (-AC). Keay & Meikile 508; 11-
11-1949 (K).
07N 05E, 07°24'N 05003'E: Igbara Oke, side of the road to Igbaru ado
(-AC). Meikile 508; 11-11-1949 (P).
07N 06E, 07°50'N 06°07'E: Province Kabba, Igala Distr., Ogbogbo (-CC).
Latilo FHI 47698; 21-06-1963 (K).
07N 07E, 07°22'N 07°38'E: Province Kabba, Ankpa Distr., along road
from Ankpa to Acharane (-BC). Oaramola, B.O. FHI 38043; 22-06-1958
(K).
Not traced: Bamenda Province, Wum Distr., Aba-ajia on right bank of
river Ife. Ujor, E. FHI 30462; 14-07-1951 (K).
xxii
SIÉNÉGAl
12N 1SW, 12°S0'N 1S000'W: Casamance, forest reserve at Koucouck
(-CC). Miege, J. & Doumbia, F. 1492; 10-09-1962 (ALF).
12N 1SW, 12°44'N 1S033'W: Region de Sêchiou, Forte de Boudie (-DC).
Berhaut, R.p. 6068; 31-10-1983 (P).
12N 1SW, 12°44'N 1S033'W: Region de Séohiou, Forte de Boudie (-DC).
Berhaut, R.p. 6351; 31-10-1983 (BR, M, P).
12N 15W, 12°44'N 1S033'W: Region de Sédtuou, Forte de Boudie (-DC).
Berhaut, R.p. 6357; 31-10-1983 (BR, P).
12N 16W, 12°48'N 16°18'W: Region de Bignona, forest Kalounayes
(-CD). Berhaut, R.P. 7365; 28-08-1968 (BR, M, P).
SIERRA lEONE
09N 11W, 09040'N 11°36'W: Kabala, north Province (-DA). Glanville,
D.R. 267; 18-07-1930 (K). Alt. 400 m.
09N 11W, OgO49' N 11°39'W: Musaia (-DC). Deighton, F.G. 4405; 20-08-
1946 (K).
09N 11W, 09049'N 11°39'W: Musaia (-DC). Deighton, F.G. 4805; 20-08-
1946 (K).
09N 11W, 09°49' N 11°39'W: Musaia (-DC). Jordan, H.D. 494; 22-08-
1951 (K).
SOUTH AFRICA (REPUBLIC OF 50UTH AFRICA)
235 30E, 23°42'5 30006'E: Transvaal, Letaba - Duiwelskloof, Westfalia
estate, small kloof below road to Zulu's bull kraal & 0.8 km from new dam
Westfalia (-CA). Scheepers, J.G. 1058; 24-11-1960 (BM, K, M, PRE!,
PRU, SRGH, WAG, Z). Alt. 1,200 m.
235 30E, 23°46'5 30010'E: Northern Transvaal, Magoebaskloof forest
(-CC). Gerstner, J. 5805; 22-01-1946 (PRE).
235 30E, 23°46'5 30010'E: Transvaal, Sendelingshoek - Noordewenke
(-CC). Keef, J.D.N. S.n. 00-12-1967 (BR, K).
xxiii
275 32E, 27°35'5 32°05'E: Zululand, Ubombo (-CA). Collins, E. 15; DO-
DO-DODO(PRE).
275 32E, 27°35'5 32°05'E: Ubombo (Umbombo) (-AC). Stewart, L. s.n.
00-00-0000 (P).
275 32E, 27°37'5 32°03'E: Ubombo Distr., Mkuze Nature Reserve, wild
garden (-CA). Venter, H.J. T. 9329; 16-11-1999 (BlFU!).
275 32E, 27°37'5 32°03'E: Ubombo Distr., Mkuze Nature Reserve,
confluence of Mkuze and Msunduze Rivers (-CA). Ward, J. 3626; 06-11-
1960 (K, M, NU, NH, PRE, SRGH). Alt. 25 m.
285 31E, 28°38'5 31°06'E: Zululand, Nkandla, near white Umfolozi River
(-CA). Gerstner, J. 617; 20-01-1935 (PRE).
285 31E, 28°58'5 31°46'E: Zululand, Mtunzini, Garden lan Garland
(-DD). Venter, H.J. T. 8781; 06-01-1983 (BlFU!).
285 31E, 28°45'5 31°54'E: Zululand, Empangeni, Ann Hutchings garden
(-DD). Venter, H.J. T. 9282; 30-11-1993 (BlFU!).
285 32E, 28°20'5 32°14'E: Natal, Hlabisa - Dukuduku forest reserve
(-AD). Strey, R.G. 5745; 25-01-1965 (NH, PRE).
285 32E, 28°20'5 32°14'E: Natal, Dududuku swamp forest (-AD). Strey,
R.G. 10347; 25-01-1965 (K, PRE!).
295 30E, 29°36'5 30024'E: Natal, Pietermaritzburg (-CB). Stainbank
11900; 17-01-1908 (NH).
295 31E, 29°20'5 31°18'E: Natal, Stanger (-AD). Pienaar, B.J. 170; 22-
02-1980 (NH!).
295 31E, 29°53'5 31°00'E: Natal, Durban Botanical Garden (-CC).
Medley Wood, J. 10; 00-00-0000 (MO, PRE, SRGH).
295 31E, 29°53'5 31°00'E: Natal, Durban Botanical Garden (-CC).
Medley Wood, J. 197; 00-00-0000 (MO, PRE, SRGH).
295 31E, 29°53'5 31°00'E: Natal, Durban - Wentworth (-CC). Medley
Wood, J. 951; 11-01-1892 (BM, BOL, GRA, SAM).
295 31E, 29°53'5 31,°00'E: Natal, Berea (-CC). Medley Wood, J. 5499;
21-12-1894 (MO, PRE!, SRGH). Alt. 50 m.
xxiv
295 31E, 29°53'5 31°00'E: Durban (-CC). Medley Wood, J. 6180; 20-12-
1896 (BM, PRE!, SAM). Alt. 30 m.
295 31E, 29°53'5 31°00': Natal, Berea (-CC). Medley Wood, J. 6499;
21-12-1894 (PRE).
295 31E, 29°53'5 31°00'E: Durban (-CC). Medley Wood, J. 8653; 00-00-
0000 (MO).
295 31E, 29°53'5 31°00'E: Natal, Berea (-CC). Medley Wood, J. 10754;
17-12-1907 (MO).
295 31E, 29°53'5 31°00'E: Durban, Wentworth (-CC). Medley Wood, J.
s.n.; 00-00-0000 (K, Z).
295 31E, 29°53'5 31°00'E: Natal - on verandah of hotel Avoca (-CC).
Olivier, J.C. 138; 00-02-1925 (NH).
295 31E, 29°53'5 31°00'E: Natal, Durban Botanical Garden (-CC).
Venter, H.J. T. 8976; 14-11-1983 (BLFU!).
295 31E, 29°53'5 31°00': Natal, Durban Botanical Garden (-CC). Venter,
H.J. T. 9068; 20-11-1984 (BLFU!).
305 29E, 30°46'5 29040'E: Mfundisweni (-DC). White s.n.; 00-00-1869
(K hola, iso).
305 30E, 30°19'5 30040'E: Natal, Umzinto, Port Shepstone (-BC).
Medley Wood, J. 3271; 00-00-1884 (NH).
305 30E, 30°19'5 30040'E: Natal, Umzinto, Port Shepstone (-BC).
Medley Wood 4110; 00-00-0000 (NH).
305 30E, 30°43'5 30028'E: Natal, Port Shepstone (-CB). Eyles, A. s.n.;
00-00-1932 (K).
Not traced: Natal. Kew, H. 11/74; 12-12-1871 (K halo! Iso!).
Not traced: Natal. Palm, H. s.n.; 03-08-1911 (K).
5UDAN
04N 32E, 04°05'N 32°43'E: Katire, Torit District (-BA). Jackson 3011;
20-06-1953 (K).
xxv
05N 321E, 05000'N 32°30'E: Equatoria Province, Loti rest house, Torit
District (-BA). Andrews, J.W. 1739; 09-06-1969 (BR, K).
SWAZILAND
265 311E,26°10'5 31°50'E: Stezi (-BB). Young, J.G.; 00-00-0000 (PRE).
TANZANIA
055 381E,05°09'5 38°35'E: T3 LushotolTanga District, Ngua estate forest
reserve (-BA). Semsei 3213; 27-05-1961 (BR, K).
. 055 38E, 05°08'5 3S039'E: T3 Lushoto, Amani (-BA). Warneike 346; 00-
03-1903 (W, Z).
05S 38E, 05°08'5 3S039'E: T3 Lushoto, Amani (-BA). Zimmermann
1125; 18-03-1909 (BM).
05S 38E, 05°05'5 3S045'E: T3 Tanga, 4 km NE of Tongwe, lower slopes
of Mlinga Hili, east Usambaras (-BB). Drummond, RB. & Hems/ey, J.H.
1432; 07-03-1953 (BR, K). Alt. 600 m.
065 36E, 06°50'5 3S059'E: T6 Kilosa (-DD). Lwynnerton, C. 944; 00-00-
0000 (BM).
065 37E, c. 06°55'5 37°50'E: T6 Morogoro Distr., Njurn-ya-ndogo (-DD).
Sch/ieben, H.J. 334; 30-01-1938 (B, BM, BR, G, K, MO, P, W, Z). Alt
1,300 m.
085 361E, 08°49'5 36°43'E: T6 Ulanga, Mahenge Distr., Mzelezi forest
reserve, c. 15 km off Mahenge (-DC). Cribb, Grey-Wi/son & Mwasambi
11064; 19-01-1979 (K). Alt. 725 m.
095 33E, 09°17'5 33039~E: T3 Lushoto, Nyassa Hochland, station
Kyimbila (-BC). Siolz, A. 1837; 00-01-1913 (B, G, K, M, W, WAG, Z).
Alt. 700 m.
095 39f, 09°59'5 39024'E: T8 Lindi, Lutamba, Mirola (= Milola) River
valley (-CD). Sch/ieben, H.J. 5904; 23-01-1935 (K, MO, SRGH).
xxvi
105 35E, 10°16'5 35°39'E: -T8 Songea, River Mutandazi, west of
Gumbiro riverine forest on old termite hill (-BC). Mi/ne-Redhead & Tay/or
10138; 09-05-1956 (K). Alt. 780 m.
105 39E, 10°00'5 39°41'E: T8 Lindi, Lindi-Bazirk, Inhamba pass (-BA).
Sch/ieben, H.J. 5909; 23-01-1935 (BR, USe).
UGANDA
OON30E, 00040'N 30010'E: U2 Toro, 16 km from Fort Portal road (-CA).
Luid 2374; 28-02-1959 (K). Alt. 1,270 m.
OON 31E, 00054'N 31°16'E: U4 Mubende, Buganda Province, locality
Kasambya (-CD). Griffiths 30; 16-05-1957 (K).
OON 32E, 00007'N 32°59'E: U4 Mengo, Buganda Province, Kyagwe,
locality Nansagazi (-BB). Oawkins 706; 24-01-1951 (BR, K). Alt. 1,300
m.
OON32E, 00015'N 32°46'lE: U4 Mengo, Kipayo (-BO). Oummer 871; 00-
00-0000 (BM).
OON 32E, 00015'N 32°46'E: U4 Mengo Distr., Kipayo (-BO). Oummer
2467; 00-05-1915 (BOL, BM, K, MO).
OON33E, c. 00045'N 33°30'E: U3 Busoga (-DC). Brown, E. 264; 06-09-
1905 (K). Alt. 1,300 m.
01N 31E, 01°30'N 31°29'E: U2 Bonyoro District, Bulindi (-DA). Thomas,
N. W. 3890; 13-05-1941 (K). Alt. 1,270 m.
01N 31E, c. 01°40'N 31°30'E: U2 Bunyoro (-DA). Brown, E. 394; 25-11-
1907 (K).
01N 31E, c. 01°47'N 31°35'lE: U2 Bunyoro Distr., Budongo forest reserve
(-DC). Egge/ing 2021; 00-05-1935 (K).
01N 31E, c. 01°40'N 31°30'E: U2 Bunyoro (-DA). Egge/ing 4376; 00-05-
1941 (K).
01N 31E, c. 01°47'N 31°35'E: U2 Bunyoro, Bujenje country, locality
Budongo forest (-DC). Synn att, T.J. 670; 16-09-1971 (K photo!). Alt.
1,050 m.
xxvii
01N 31E, c. 01°47'N 31°35'E: U2 Bunyoro, Bujenje country, locality
Budongo forest (-DC). Synnoii, T.J. 881; 16-09-1971 (MO). Alt. 1,050 m.
ZAMBIA
125 28E, 12°48'5 28°14'E: Kitwe (Kitwe-Nkana) (-CC). Fanshawe, O.B.
2699; 08-01-1956 (K).
135 31E, 13°55'5 31°23'E: Sasare (-CD). Robson, N.K.B. 868; 08-12-
1958 (BM, BR, K, SRGH). Alt. 750 m.
Not traced: Fort Jackson. King, A.E.B. 457; 17-01-1959 (K, SRGH).
ZIMBABWE (RHODESIA)
175 30E, 17°30'5 30059'E: Mazoe (= Mazowe) (-DB). Ey/es, F. 204; 00-
12-1905 (BM, BOL, K, SAM, SRGH).
175 30E, 17°30'5 30059'E: Mazoe (-DB). Ey/es, F. 3235; 18-12-1921 (K,
SAM). Alt. 1,400 m.
175 25E, 17°57'5 25°50'E: Victoria falls (-DD). Alien, C.E.F. 259; 00-01-
1956 (K).
175 25E, 17°57'5 25050'E: Victoria falls (-DD). Lewy, B. 1311; 00-12-
1934 (K).
175 25E, 17°57'5 25050'E: Victoria falls (-DD). Hutchinson & Gil/ett 3468;
08-07-1930 (K).
175 30E, 17°00'5 30030'E: lomagundi (-AD). CanneIl, J.C. 574; 01-11-
1973 (SRGH).
175 31E, 17°43'5 31°03'E: Salisbury (= Harare) (-CA). Ey/es, F. 3385;
28-11-1923 (K).
175 31E, 17°43'5 31°03'E: Salisbury (-CA). Ey/es, F. 5522; 28-11-1923
(SRGH).
175 31E, 17°43'5 31°03'E: Salisbury Botanic Gardens (-CA). Ey/es, F.
5666; 02-11-1927 (SRGH).
175 31E, 17°43'5 31°03'E: Salisbury - springs farm (-CA). Greatrex, F.C.
18213; 11-12-1947 (K, SRGH).
xxviii
175 31E, c. 17°50'5 31°05'E: Salisbury Highlands (-CC). Leach, L.C.
13567; 08-11-1966 (K, SRGH).
175 31E, 17°55'5 31°03'E: Harare (= Mbare) (-CC). Eyles, F. 1379; 21-
11-1927 (B, K). Alt. 1,600 m.
175 27E, 17°52'5 31°22'E: District Goramanzi, Binga swamp forest (-CD).
Brodrick 110; 07-12-1971 (BR, K, M).
185 30E, 18°20'5 30006'E: Mazoe, Iron Duke mine (-AC). Rutherford
426; 00-00-0000 (MO).
185 32E, 18°58'5 32°40'E: Umtali (= Mutare), Kukwanisa training farm
(-DC). Biegel, H.M. 1688; 09-01-1966 (SRGH).
185 32E, 18°58'5 32°40'E: Mutare, Black mountain inn (-DC). Chase,
N.C.460; 28-12-1947 (BM).
185 32E, 18°58'5 32°40'E: Umtali District, Park River, north commonage
(-DC). Chase, N.C. 5424; 27-12-1954 (BM, BR, CO, K, SRGH).
195 32E, c. 19°48'5 32°53'E: Melsetter (= Chimanimani), Welgelegen
(-DD). Ball, J.S. 26; 00-00-0000 (MO, SRGH).
195 32E, c. 19°48'5 32°53'E: Melsetter, Umvumvumvu River (-DD).
Chase, u.c. 459; 24-12-1947 (K, SRGH).
205 30E, 20°38'5 30018'E: District Belingwe, mount Buhwa (-CB).
Biegel, H.M., Pope, G.V. & Simon, B.K. 4297; 04-05-1973 (K photo!
SRGH!). Alt. 1,400 m.
205 32E, 20°24'5 32°43'E: District Chipinga (Chipinge), Gungungana
forest reserve (-BC). Goldsmith, B. 5/64; 00-02-1964 (BR, K photo!, MO,
SRGH!). Alt. 2:. 1,200 m.
Not traced: NW Rhodesia. Rogers, F.Q. 13476; 00-06-1920 (SRGH).
XXIX
SACLEUXIA Baill.
Sacleuxia new;; (Benth.) Bullock
KENYA
OOS 3SE, 00°55'5 3S027'E: K3 Naivasha District, Ol Longonot estate
(-CD). Kerfoot, O. 3395; 29-12-1961 (K!).
025 3SE, 02°37'5 3S002'E: K4 Machakos Distr., Kikumbuliyu - Ruwenzori
(-CA). Scott EI/iot 6149: 1893 - 1894 (K). Alt. 800 - 1,500 m.
035 3SE, 03°11'5 3S031'E: K7 Teita Distr., south of Manga mountain, Ndi
(-BA). Verdcourl & Polhil/2712 16-04-1960 (K).
035 3SE, c.03014'5 3S029'E: Teita Distr., Ndi Hili (-BA). Gil/ett, J.B.
16877: 29-08-1965 (K). Alt. 650 - 700 m.
035 3SE, 03°04'5 3So45'E: Tsavo National Park (-BB). Bal/y, P.R.O.
13369: 12-08-1969 (G). Alt. 800 m.
035 3SE, 03°20'5 3S033'E: K7 Teita Distr., Worssera lookout, east Tsavo
Nat. Park (-BC). Greenway & Kanuri 12852: 23-12-1966 (K!). Alt. 670 m.
035 3SE, 03°20'5 3S035'E: K7 Teita Distr., Tsavo Nat. Park, east Mzinga
Hill (-BC). Gil/ett, J.B. 17237: 09-04-1966 (K). Alt. 750 m.
035 3SE, 03°22'5 3S035'E: K7 Teita Distr., Tsavo Nat. Park, east Simba
Hili (-BC). Faden, R.B. &A.J. 74/450: 18-04-1974 (K). Alt. 600-640 m.
035 3SE, 03°23'5 3S034'E: K7 Teita Distr., Voi, Mazinga Hill (-BC). Bal/y,
P.R.O. 8623: 31-01-1953 (K!). Alt. 620 m (Kikumbuliyu at 830-1,000 m).
035 3SE, 03°47'5 3S052'E: K7 Teita Distr., Kivuko Hill, between
Mackinnon road & Kasigau (-DD). Bal/y, P.R.O. 12728; 26-04-1963 (K).
TANZANIA
035 37E, 03°04'5 37°22'E: T2 Moshi Distr., Kilimanjaro mountain (-AB).
New, S.n.; 00-08-1871 (K holo, photo!).
045 37E, 04°0S'5 37°57'E: T3 Pare Distr., Kisiwani, east Usambaras
(-BB). Greenway, P.J. 4723; 04-11-1936 (K).
045 37E, 04°22'5 37°53'E: T3 Pare Distr., Pare Hills, Suji Mission (-BD).
Bal/y, P.R.O. 4237; 14-01-1945 (K). Alt. 1,800 m.
xxx
055 38E, 05°09'5 38°28'E: T3 Lushoto Distr., Korogwe, Lwenga estates
(-AB). Fau/kner, H.G. 1115; 09-01-1953 (K).
055 38E, 05°29'5 38°01'E: T3 Handeni Distr., Kideleko mountains (-AC).
Archbo/d, N.E. 501; 07-07-1965 (K). Alt. 1,000 m.
055 38E, 05°29'5 38°01'E: T3 Handeni Distr., Kideleko mts. (-AC).
Fau/kner, H.G. 1434; 04-07-1954 (K!). Alt. 670 m.
055 38E, 05°21'5 38°14'E: T3 Handeni Distr., Zindeni Hills (-AD). Burtt,
B.D. 4854; 12-09-1933 (K). Alt. 670 m.
055 38E, 05°27'5 38°16'E: T3 Handeni Distr., Kwa Mkono (-AD).
ArchboId, N.E. 889; 24-06-1969 (K).
055 38E, 05°27'5 38°16'E: T3 Handeni Distr., Kwamkono (Kwa Mkono)
(-AD). Archbo/d, N.E. 2842; 05-05-1981 (K). Alt. 600 m.
055 38E, 05°06'5 38°37'E: T3 Lushoto Distr., Bomole (-BA).
Zimmermann, s.n. 00-00-0000 (K).
055 38E, 05°06'5 38°37'E: T3 Lushoto Distr., Amani, Mt. Bomole, east
Usambaras (-BA). Verdcourt 188; 10-05-1950 (K, MO!). Alt. 1,200 m.
055 38E, 05°06'5 38°37'E: T3 Lushoto Distr., Amani, Bomole Hili (-BA).
Burtt, B.D. 450; 00-03-1926 (K).
055 38E, 05°06'5 38°35'E: T3 Lushoto/Tanga Distr., Ngua, Amani (-BA).
Semsei 2296; 00-09-1955 (K).
055 38E, 05°06'5 38°41'E: T3 Lushoto Distr., Korogwe, Sigi River
valley/gorge, Longuza Hill- east Usambaras (-BA). Brenan 8342; 19-11-
1947 (K).
055 38E, 05°08'5 38°34'E: T3 Tanga Distr., Magunga, sisal estates,
footpath between Magunga and Ngua estates, west slopes of east
Usambaras (-BA). Drummond, R.B. & Hems/ey, J.H. 3364; 19-07-1953
(USe!). Alt. c. 800 m.
055 38E, 05°09'5 38°35'E: T3 Lushoto/Tanga, Ngua forest reserve, east
Usambaras (-BA). Semsei 3265; 06-08-1961 (K).
055 39E, 05°03'5 39°04'E: T3 Lushoto/Tanga Distr., lower Sigi valley,
east Usambaras (-AA). Verdcourt 246; 30-05-1950 (K!, MO!). Alt. 400 m.
xxxi
065 37E, 06°00'5 37°43'E: T6 Morogoro Distr., Nguru mts., south face of
main Kanga forest reserve (-BA). Pais, T. & Temu, R.P.C., 87228; 02-12-
1987 (K!). Alt. 1,250 - 1,500 m.
065 37E, 06°46'5 37°05'E: T6 Kilosa Distr., foothills of Uluguru mts., 10
km From Mandege - Mvumi track (-CC). Thulin & Mhoro 2978; 06-06-
1978 (K). Alt. 900 m.
065 39E, 06°10'5 39020'E: Nguru of ngourou (-AB). Sacleux 758; 01-
06-1892 (P). Alt. 1,200 m.
Not traced: Hidira Kjaro. Or. Kirk %2; [In Hook Icon. PI. 12: 74, plate
1186; Flora of Tropical Africa 4(1): 241].
Sacleuxia tuberosa (E.A. Bruce) Bullock
KENYA
OOS 36E, 00°55'5 36°19'E: K3 Naivasha, SW of Lake Naivasha, Hell's
gate, stream jets, Njorowa gorge (-CD). Verdcourt 3289; 21-10-1962 (K).
Alt. 1,900 m.
OOS 36E, 00°58'5 36°27'E: K3 Naivasha, Ol Longonot estate (-CD).
Kerfoot, 0.3596; 28-01-1962 (K!).
015 36E, 01°105 36°10'E: K3 Naivasha, 32 km from Kikuyu on direct raad
to Narok town rift Wanyaga (-AA). Verdcourt 3551; 20-01-1963 (K). Alt.
1,800 - 2,000 m.
015 36E, c. 01°09'5 36°21'E: K3/6 Naivasha/Masai Distr., west ruins of
central motex of Mt. Suswa (-AB). Glover, P.E. & Samuel 3309; 07-10-
1962 (K). Alt. 1,700 m.
015 38E, 03°46'5 38°47'E: K7 Teita Distr., Kivuko Hill, between
Mackinnon road and Kasigau (-DD). Bal/y, P.R. O. 12728; 26-04-1963
(G).
TANZANIA
025 31E, 02°30'5 31°20'E: T1 Biharamulo Distr., Lake Province (-CB).
Ford, J. 847; 13-11-1948 (K!).
xxxii
025 31E, 02°35'5 31°25'E: T1 Biharamulo, Lake Province (-CB).
Greenway, P.J. 7376; 29-04-1945 (K). Alt. 1,100 m.
025 32E, 02°31'5 32°54'E: T1 Biharamulo, Lake Province, Mwanza
(-DB). Tanner, R.E.S. 608; 30-02-1952 (K). Alt. 1,200 m.
025 32E, 02°36'5 32°45'E: T1 Mwanza, Geita Distr., Lake Province near
Mwanza, Juma island (-DB). Proeter 136; 00-01-1953 (K).
025 33E, c. 02°29'5 33020'E: T1 Mwanza/Kwimba/Musoma, coast of
Speke Gulf, Lake Victoria (-AD). Burtt, B.D. 2475; 01-06-1931 (K holo!,
iso!). AIt.1,100m.
02533E, 02°30'5 33030'E: T1 Mwanza Distr., Bbarika (-DA). Tanner,
R.E.S. 360; 19-07-1951 (K!). Alt. 1,100 m.
035 35E, 03°20'5 35005'E: T2 Masai Distr., 80 km west of Enduien (-AC).
Bally, p.R.O. 11605; 07-07-1957 (G, K). Alt. 1,800 m.
065 37E, c. 06°57'5 37°06'E: T6 Kilosa Distr., Magubike village (-CC).
Poes Sua 88012; 05-02-1988 (K). Alt. 800 - 900 m.
065 37E, 06°42'5 37°36'E: T6 Morogoro Distr., north of Liwale River,
north Nguru mountain side above Manyangu forest (-DA). Drummond,
R.B. & Hems/ey, J.H. 1978; 02-04-1953 (K, M). Alt. 900 m.
065 37E, 06°54'5 37°38'E: T6 Morogoro Distr., Uruguru/Uguruga, Mzinga
area (-DC). Burlt, B.D. 6480; 26-06-1937 (K). Alt. 1,300 m.
xxxiii
SARCORRHIZA Bullock
Sarcorrhiza epiphytica Bullock
DEMOCRATIC REPUBLIC OF THE CONGO
OON29E, 00020'N 29050'E: U2/Zaire, Toro Distr., Ruwenzori, Kanvui
(-BO). Bequaerl 4494; 26-05-1914 (BR!). Alt.:!: 1,800 m.
04817E, 04°00'817°00'E: Kwango, Gimbi-Bas-Congo, Jochere (-AA).
Oevred, A. 1580; 16-02-1955 (K, BLFU photo!)
IVORYCOA8T
05N 06W, 05014'N 06°10'W: :!: 60 km north of Sassandra, River Davo,
East of Beyo (-AA). Leeuwenberg, A.J.M., 2586; 27-01-1959 (WAG).
TANZANIA
048 3SE, 04°45'8 3S03S'E: T3 Lushoto Distr., path up Mt. Bomole
behind Amani (-CD). Vedcourt, B. & L.O. 1725; 24-12-1956 (K holo!;
BLFU photo!). Alt. 1,000 m
058 3SE, 05°06'8 38°38'E: Usambaras, Amani (-BA). Greenway 3679;
02-01-1934 (K). Alt. 1,000 m.
0583SE, 05010'83S03S'E: T3 Tanga Distr., Kwamkoro near sawmill
(-BA). Semsei 2957; 15-12-1959 (K).
068 37E, 06°55'8 37°40'E: T6 Morogoro Distr., Uluguru North, north-
west side of mist forest (-DC). Schlieben, H.J. 2939; 08-11-1932 (BM,
BR!, K, G). Alt. 1,900 m.
XXXIV
ZACA TEZA Bullock
Zacateza pedicellata (K. Schum.) Bullock
ANGOLA
075 20E, c. 07°33'5 20050'E: North van Luanda, Dundo near River
Luachimo (-DB). Gossweiler, J. 13684; 14-10-1946 (CO, K!, P). Alt. 700
m.
BURUNDI
025 29E, 02°54'5 29°49'E: Ngozi, Mukara (Rwegura) (-DD). Reckmans,
M.6092; 27-04-1977 (WAG).
CAMEROON
02N 12E, 02°26'N 12°30'E: Oveng, 30 km WNW of Sangmelina (-BC).
Letouzey, J. 11437; 06-07-1972 (P).
03N 11E, 03°12'N 11°50'E: Between Kondebilong and Meyila (-BB).
Asonganyi 87; 17-05-1980 (P).
03N 11E, c. 03°12'N 11°50'E: Kondebilong and Meyila, 53 km S of
Mbalmayo (-BC). Asonganyi 87; 09-07-1980 (P).
04N 32E, c. 04°46'N 12°32'E: Forest along River Sonaga near Goysum,
20 km west of Dug Oemf. (-DC). Breteler, F.J. 941; 27-01-1961 (WAG).
04N 13E, 04°34'N 13°45'E: Bertoua, 6 km along road to Batouri and
Betare Oya gallery forest (-DB). Breteler, F.J. 1196; 14-03-1961 (BR, K,
LlSC, P, WAG). Alt. c. 650 m.
04N 14E, 04°09'N 14°40'E: 50 km SW of Batouri, E of village Mloundou,
old secondary forest along small river (-BA). Breteler, F.J. 2845; 17-04-
1962 (BR, K, P, WAG).
04N 14E, 04°14'N 14°13'E: Nol, 25 km WSW of Batouri (-AC). Letouzey,
J. 4648; 03-04-1962 (P).
Not traced: East Cameroon, Die Bitjoknam, eyen a 12 km e de
Ngeleulem. Letouzey, J. 11618; 00-00-0000 (BR, K, WAG).
xxxv
Not traced: River de la Riviere ABO entre Koki et Moussoko, 20 km north
Donala. Letouzey, J. 14763; 29-04-1976 (BR, K, P, WAG).
CENTRAL AIFRICAN RIEPUBLlC
OSN 231E, OSo33'N 23014'E: Yalinga - Oubangui River (-CB). Le Tesiu,
M.G. 4744; 21-03-1923 (BR, P).
OSN 231E, c. OSo33'N 23014'E: Oubangui, Boukoko (-CB). Tisserant, R.P.
2531; 18-05-1953 (P).
OSN 23E, c. OSo33'N 23014'1E: Boukoko (-CB). Equipe 2531; 00-00-0000
(P).
OSN 23E, c. OSo33'N 23014'E: Boukoko (-CB). Equipe 2532; 00-00-0000
(P).
DEMOCRATIC REPUBLIC OF THE CONGO
OON 20E, 00028'N 20oS7'E: Befale, Ekekeli (-CD). Evrard 2883; 22-09-
1957 (BR, WAG).
OON 24E, 00048'N 24°10'E: Yabalanga, Isangi territory (-CC). Léonard,
A. 698; 19-05-1958 (BR, K!).
OON 24E, OOoSO'N24°1S'E: Yangole, 20 km west of Yangambi, Isangi
(-CD). Louis, J. 11913; 00-00-0000 (BR).
OON 24E, OOoSO'N 24°1S'E: Yangole, Isangi (-CD). Louis, J. 13517; 03-
02-1939 (K, P).
OON 24E, OOo47'N 24°27'E: Yangambi (-CD). Bo/erna 1148; 26-08-1963
(K).
OON 241E, OOo47'N 24°27'E: Yangambi (-CD). Louis, J. 10524; 27-07-
1938 (K).
OON 24E, OOo43'N 24°32'E: Yalutcha, Isangi (-DA). Germain 242; 09-03-
1940 (BR). Alt. 470 m.
01N 231E, 01°14'N 23°31'E: Barumbu (-BC). Laurent 1379; 00-00-0000
(BR).
01 N 2SE, 01°23'N 2So20'E: Kisangani - Banalia 30 km N of Bengamisa
XXX"l
(-AD). Lisowski 18973; 24-06-1973 (BR).
03N 20E, 03030'N 20007'E: Boyasebego (-CA). Evrard 836; 00-00-0000
(BR).
03N 25E, 03025'N 25043'E: Bambesa (-BC). Gerard 2857; 28-07-1961
(BR, P).
03N 25E, 03028'N 25043'E: Bambesa (-BC). Gerard 4935; 28-07-1961
(P).
03N 28E, 03030N 28°15E: Munsa, MonbuUu Land, Niangara Province
(-CB). Schweinfurth, G. 3483; 03-04-1870 (K lecto!).
03N 28E, 03030N 28°15E: Munsa, Monbuttu Land, Niangara Province
(-CB). Schweinfurlh, G. 3488; 03-04-1870 (K iso!).
04N 19E, 04°58'N 19°57'E: Borun (Boruna) - Kusu (-DD). Dubais 428;
00-07-1934 (BR).
005 18E, 00°06'5 18041'E: Bokuma - Equator (-BA). Louis, J. 106; 11-
09-1935 (BR, K!).
005 18E, 00°45'5 18°09'E: Equator province, territory Bikoro, locality
Mabali (-CC). Densa 28; 15-04-1955 (BR).
005 18E, 00°435 18°32'E: Bikatola et Bikoro (-CA). Lebrun 1444; 00-00-
0000 (BR).
005 18E, 00°50'5 18°00'E: Mabali location, Tumba, Equator region (-CC).
Densa 239; 16-04-1956 (BR).
005 20E, 00°02'5 20058'E: Bongoy (=Bongoie), territory Boende (-BB).
Evrard 3202; 14-01-1958 (BR).
015 23E, 01°06'5 23006'E: Territory Yalikungu, Ikela (-AA). Evrard 5420;
02-01-1958 (K).
015 23E, 01°11'5 23016'E: Territory Ikela (-AB). Evrard 5430; 02-01-
1958 (BR).
025 18E, 02°42'5 18°10'E: Kutu (Lae Leopold), galerie forestiere (-CA).
Lebrun 6600; 00-11-1932 (BR, K, P).
045 15E , 04°05'5 15025'E: Gamakala (-AB). Farron 4141; 14-05-1968
(P).
xxxvii
045 15E, 04°19'5 15019'E: Leopoldville (=Kinshasa) (-AD). Bequaert
7529; 05-05-1915 (BR).
045 15E, 04°06'5 15°51'E: Sao-Ndunu, territory Maluku, Kinshasa
Province (-BB). Breyne 2191; 05-05-1971 (BR).
055 14E, 05°12'5 14°56'E: Wengu River, Boko (-BB). Nsimunde/e 420;
02-05-1978 (BR).
055 14E, 05°12'5 14°56'E: Wengu River, Boko (-BB). Nsimunde/e 424;
02-05-1978 (BR).
065 19E, 06°50'5 19°19'E: Congolan, Bumba territory, forest
Maric;agense (-CD). Evrard 3467; 12-02-1958 (BR, K!).
GABON
OON 12E, 00038'N 12°47'E: d'lpassa, 10 km south of Makokou (-DB).
F/orence 2044; 19-05-1979 (P). Alt. 500 m.
NIGERIA
04N 07E, 04°30'N 07°56'E: South Nigeria, Eket District (-DB). Ta/bott,
T.A. 3265; 00-04-1914 (K).
06N 23E, 06°27'N 03°23'E: Lagos (-AD). Detziel, J.M. 1348; 14-01-1919
(K).
06N 03E, 06°25'N 03°32'E: Igbessa, Lagos Botanical station (-BC).
Mil/en, A. 130; 00-01-1893 (K holo!).
06N 03E, 06°36'N 03028'E: West Nigeria, swamp c. 1 km east Ikorodu
road, Ikeja 11 km (-CB). Kil/ick 248; 04-07-1965 (K).
XXXVIII