A New Heterophyllous Spermacoce Species (Rubiaceae) from
the Marungu Highlands, Democratic Republic of the Congo
Author(s): Steven Dessein, Elmar Robbrecht, Erik Smets
Source: Novon: A Journal for Botanical Nomenclature, 16(2):231-234. 2006.
Published By: Missouri Botanical Garden
DOI: 10.3417/1055-3177(2006)16[231:ANHSSR]2.0.CO;2
URL: http://www.bioone.org/doi/
full/10.3417/1055-3177%282006%2916%5B231%3AANHSSR%5D2.0.CO
%3B2
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A New Heterophyllous Spermacoce Species (Rubiaceae) from
the Marungu Highlands, Democratic Republic of the Congo
Steven Dessein and Elmar Robbrecht
National Botanic Garden of Belgium, Domein van Bouchout, B-1860 Meise, Belgium.
steven.dessein@br.fgov.be; elmar.robbrecht@br.fgov.be
Erik Smets
Laboratory of Plant Systematics, Katholieke Universiteit Leuven, Kasteelpark Arenberg 31,
B-3001 Leuven, Belgium. erik.smets@bio.kuleuven.be
ABSTRACT . Spermacoce heteromorpha Dessein (Rubiaceae, Spermacoceae) is newly described and
illustrated. It grows on the Marungu highlands of the
Democratic Republic of the Congo (D.R. Congo) at
altitudes of 1800 to 1900 m. The species resembles
the Tanzanian Spermacoce taylorii Verdcourt, from
which it mainly differs in having terminal inflorescences, only slightly exserted anthers, and densely
pubescent stems. Moreover, Spermacoce heteromorpha
features marked heterophylly, with long leaves at the
plant’s base and much shorter cauline leaves, a type of
heterophylly rarely observed in Spermacoce or in the
Rubiaceae as a whole. The finding of this species
exemplifies the unique floristic richness of this underexplored part of D.R. Congo.
Key words: Democratic Republic of the Congo,
heterophylly, Marungu highlands, Rubiaceae, Spermacoce.
Spermacoce L. is a pantropical genus of the family
Rubiaceae comprising an estimated 275 species
(Dessein et al., 2002; Harwood & Dessein, 2005).
Tropical America, tropical Africa, and the northern
part of Australia are the main centers of diversity. The
genus is characterized by its herbaceous growth form,
the fimbriate stipules connected to the petioles, the 4merous flowers arranged in compact lateral and/or
terminal inflorescences, the bilocular ovaries with
a single ovule per locule, the often pluriaperturate
pollen grains, and the dry dehiscent fruits. Many
botanists know Spermacoce as a genus of annual weeds
of ruderal places. Typical examples are S. latifolia
Aublet and S. verticillata L., two species presumably
native to America, but introduced in Africa, Asia, and
Australia. Narrow endemics, however, are an element
of the genus with a comparable level of importance.
Our study of Spermacoce from the high plateaus of the
Zambezian regional center of endemism, for example,
revealed the existence of endemic or disjunct
Spermacoce species on the Angolan, Zambian, and
Katangan high plateaus (Dessein et al., 2002, 2003).
Within the Lisowski, Malaisse, and Symoens collection of POZG, which was recently put at our disposal,
unicate Spermacoce specimens from the Marungu
highlands (D.R. Congo) were found representing
another narrow endemic as described below.
HETEROPHYLLY
Heterophylly is not uncommon among woody
Rubiaceae (Robbrecht, 1988), but is rarely observed
among the herbaceous representatives of the family.
Within Spermacoce, it has only been recorded for S.
hockii (De Wildeman) Dessein, a perennial from D.R.
Congo (Dessein et al., 2003). In this species, long
linear leaves develop from the woody plant base, while
the cauline leaves are broader and much shorter.
During field work in Zambia, we observed that several
biannual or short-lived perennial Spermacoce species
develop rosettes of basal leaves. This feature was
observed in the field for S. dibrachiata Oliver, S.
perennis Verdcourt, and S. phyteumoides Verdcourt.
Herbarium collections rarely show the basal leaves, as
the basal part is seldom collected. In the three abovementioned species, basal rosette leaves differ only
slightly in size and shape from the cauline leaves. In
S. heteromorpha, on the contrary, the basal leaves are
markedly longer than the cauline ones. The type
specimens of S. taylorii Verdcourt also have basal and
cauline leaves, but the differences between the two
leaf types are minor.
THE MARUNGU HIGH PLATEAU
The Marungu high plateau is situated southwest of
Lake Tanganyika in D.R. Congo. The name of the
plateau is derived from the word ‘‘dilungu,’’ which
signifies ‘‘herbaceous plateau’’ (Verbeken, 1954). The
name refers to a main feature of the environment, the
absence of tree growth in the more elevated areas.
NOVON 16: 231–234. PUBLISHED
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26 JULY 2006.
232
Novon
Very little detail is known, however, about the
vegetation and environmental conditions of the region.
Floristically, too, the area remains underexplored.
Duvigneaud (1952) included the plateau in the
‘‘district des plateaux Katangais,’’ which also comprised the Biano, Kibara, and Kundelungu plateaus.
Later, he placed the Marungu plateau in a district of
its own because the vegetation shows several East
African elements not found on the other Katangan
plateaus (Duvigneaud, 1958). Most subsequent
authors followed his opinion.
The Marungu region reaches altitudes up to 2460 m
(mont Lusale) and is mainly covered by savannahs
(Lisowski et al., 1971). The vegetation of the
savannahs is composed of many species of Cyperaceae
and Poaceae, along with geoxylic herbs. During the
dry season, other plants, such as Helichrysum Miller
species (Asteraceae), may develop (Duvigneaud,
1958; Lisowski et al., 1971). White (1976) pointed
out that the main factor accounting for the absence of
trees is the waterlogged soils of the plateaus, rather
than the presence of sporadic frost periods or temporal
burning. The Marungu plateau is isolated from the
other highland areas and thus forms an island of very
peculiar environmental and edaphic conditions. As
a consequence, the Marungu plateau might house
a large number of endemic taxa, as observed on the
other Katangan plateaus (Lisowski et al., 1971;
Malaisse et al., 1995; Geerinck et al., 1996; Dessein
et al., 2003).
brescent; apex acute; base gradually narrowed; leaf
margins somewhat thickened, beset with short stiff
trichomes or scabrous; midvein somewhat prominent,
densely pubescent toward the base underneath,
sparsely pubescent above, with indistinct secondary
and tertiary veins. Stipules fimbriate; stipule base 4–
18 mm long, beset with short appressed trichomes,
often with stalked colleters in between the fimbriae;
fimbriae 3 to 6, up to 6(212) mm long, ciliate.
Flowers sessile in many-flowered capitula; inflorescences terminal, composed of several condensed
flowering nodes each with a pair of subtending leaves,
up to 3.5 cm wide in flowering stage; bracts stipuliform with numerous setae 3–6 mm long. Calyx tube
ca. 0.5 mm high; lobes 2 or 3 (rarely no distinct lobes
visible), narrowly triangular or leaf-like, often unequal, 2–8 mm long, beset with long hyaline
trichomes, sometimes with numerous setae in between
the lobes. Corolla blue; tube funnel-shaped, 6–
8.5 mm long, glabrous or sparsely pubescent outside,
with a trichome ring around or below its middle
inside; lobes triangular, 4–6 mm long, 2–2.5 mm wide
at the base, pubescent outside, glabrous inside;
anthers narrowly elliptic in outline, 1.5–2 mm long,
exserted; filaments 1–2 mm long. Pollen 14–17zonocolporate, oblate to suboblate; equatorial diameter 67–83 mm; polar outline circular and somewhat lobed; ectocolpi relatively long; endoaperture
a broad endocingulum; tectum perforate to microreticulate; microspines uniformly present; inner
nexine surface granular with endocracks. Ovary
ellipsoid to narrowly ellipsoid, 3–4.5 mm long,
pubescent in the upper part; style 9–12 mm long,
exserted; stigma shortly 2-lobed, ca. 1 mm wide;
nectary disc bipartite. Capsule and mature seeds
unknown; young seeds show the initiation of a basal
elaiosome.
TAXONOMY
Spermacoce heteromorpha Dessein, sp. nov.
TYPE: D.R. Congo. Katanga: plateau des
Marungu, Kibobwa, 15 Feb. 1970, S. Lisowski,
F. Malaisse & J.-J. Symoens 9696 (holotype,
POZG; photos, BR, K, LV, MO, P). Figure 1.
Similis Spermacocae taylorii a qua inflorescentiis terminalibus et compactis, antheris vix exsertis et caulibus
pubescentibus differt. Praeterea bene distincta habitu
heterophyllo manifesto, cum foliis basalibus duplo longioribus quam illis caulinis.
Robust, erect herb with few stems from a woody
perennial base with woody taproot; stems 45–55 cm
tall, strictly erect, with short branches in the upper
part bearing the inflorescences, densely covered with
short, stiff, often ascending trichomes. Plants heterophyllous, with long leaves at the base and shorter
cauline leaves. Leaves decussate, sessile, fused with
the stipule base; blades of basal leaves narrowly
elliptic to linear, (3–)11–22 6 0.7–2.2 cm; cauline
leaves narrowly elliptic, (2–)5–10 6 0.8–1.7 cm,
young leaves often densely covered with short stiff
trichomes above and underneath, older leaves gla-
Distribution, habitat, and phenology. Endemic to
the northern part of the Zambezian regional centre of
endemism: only known from two collections from the
Marungu high plateau (Katanga province, D.R. Congo)
in grassland at 1800–1900 m elevation. Flowering
specimens were collected in February.
Affinities with other species. Spermacoce heteromorpha is most similar to S. taylorii, a narrow endemic
from the Songea district in Tanzania. The habit
resemblance is especially striking: a woody base from
which a few erect stems arise. Furthermore, S. taylorii
shows a tendency toward the same type of heterophylly, much weaker, however, than in S. heteromorpha. Inflorescence and floral morphology clearly
separate the two species. Whereas the flowers of S.
heteromorpha are arranged in many-flowered terminal
capitula, those of S. taylorii are in clusters at the
Volume 16, Number 2
2006
Dessein et al.
Spermacoce heteromorpha (Rubiaceae)
233
Figure 1. Spermacoce heteromorpha Dessein. —A. Habit. —B. Detail of stem. —C. Inflorescence. —D, D’. Flower buds
showing variability of calyx. —E. Corolla, stamens, and style. —F. Open corolla showing stamens and a ring of trichomes. A.
Lisowski, Malaisse & Symoens 9696 (POZG); B–F. Lisowski, Malaisse & Symoens 10061 (POZG).
nodes along the stem, from which the upper 4 or 5 run
together to form a more congested cone-like inflorescence. The corolla tubes are similar, but the
corolla lobes are, based on Verdcourt’s (1976)
measurements, much longer in S. taylorii (10–
11 mm vs. 4–6 mm). The anthers of S. heteromorpha
are only exserted for 1–2 mm, while those of S.
taylorii are exserted for more than 5 mm. Further
evidence for the separation of the two species is found
in the characteristics of the external indumentum:
Spermacoce taylorii is almost entirely glabrous,
whereas S. heteromorpha is densely covered with
trichomes on almost all organs. Spermacoce perennis,
collected from the northeastern part of Zambia, also
has a similar habit. Spermacoce perennis lacks,
however, some very characteristic features of S.
heteromorpha: heterophylly is usually absent or, if
present, the basal leaves are shorter than the cauline
234
Novon
ones; the inflorescences are both terminal and
axillary; the cauline leaves are much narrower; and
secondary nerves are invisible.
———. 1958. Etudes sur la végétation du Katanga et de
ses sols métallifères. Bull. Soc. Roy. Bot. Belgique 90:
127–286.
Geerinck, D., F. Malaisse & M. Schaijes. 1996. Orchidaceae
du district phytogéographique du Haut-Katanga (Zaı̈re):
Biodiversité et endémisme. Pp. 795–801 in L. J. G. van
der Maesen, X. M. van der Burgt & J. M. van Medenbach
de Rooy (editors), The Biodiversity of African Plants:
Proceedings of the XIVth AETFAT Congress. Kluwer
Academic Publishers, Dordrecht.
Harwood, R. & S. Dessein. 2005. Australian Spermacoce
(Rubiaceae: Spermacoceae). I. Northern Territory. Austral.
Syst. Bot. 18: 297–365.
Lisowski, S., F. Malaisse & J. J. Symoens. 1971. Une flore
des hauts plateaux du Katanga. Mitt. Bot. Staatssamml.
München 10: 51–56.
Malaisse, F., J. M. Lecron & M. Schaijes. 1995. Remarques à
propos du genre Monadenium Pax (Euphorbiaceae), en
particulier concernant les espèces de la région zambézienne. Bull. Séances Acad. Roy. Sci. Outre Mer. 40:
389–418.
Robbrecht, E. 1988. Tropical woody Rubiaceae. Opera Bot.
Belg. 1. National Botanic Garden of Belgium, Meise.
Verbeken, A. 1954. Contribution à la géographie historique
du Katanga et de régions voisines. Koninklijk Belgisch
Koloniaal Instituut, Brussel.
Verdcourt, B. 1976. Rubiaceae (part 1): Spermacoce. Pp.
339–374 in R. M. Polhill (editor), Flora of Tropical East
Africa. Crown Agents for Oversea Governments and
Administrations, London.
White, F. 1976. The underground forests of Africa: A
preliminary review. Gard. Bull. Singapore. XXIX: 57–71.
Paratype. D.R. CONGO. Katanga: plateau des Marungu, au pied du mont Kibobwa, au-dessus du village
Kibobwa, steppe au bord d’une petite rivière, 29u429E
7u319S, 1900 m, 19 Feb. 1970, S. Lisowski, F. Malaisse &
J.-J. Symoens 10061 (POZG; photos, BR, K, LV, P).
Acknowledgments. The authors are indebted to
Anja Vandeperre for the illustration and to the curator of POZG for specimens loaned. This study was
supported by grants from the Research Council of the
K.U. Leuven (OT/01/25) and the Research Foundation
Flanders (1.5.069.02). Fieldwork in Zambia was
supported by the ‘‘Stichting tot bevordering van het
wetenschappelijk onderzoek in Afrika.’’
Literature Cited
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species of Spermacoce (Rubiaceae) from the Manika high
plateau (Katanga; R.D. Congo). Nordic J. Bot. 22: 513–523.
———, S. Ntore, E. Robbrecht & E. Smets. 2003. Pollen
and seeds reveal that Spermacoce thymoidea s.l. (African
Rubiaceae, Spermacoceae) represents three endemic or
disjunct species from the Zambezian high plateaus. Syst.
Bot. 28: 130–144.
Duvigneaud, P. 1952. La flore et la végétation du Congo
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