Content uploaded by Pierre Meerts
Author content
All content in this area was uploaded by Pierre Meerts on Mar 09, 2019
Content may be subject to copyright.
Plant Ecology and Evolution 145 (3): 373–409, 2012
http://dx.doi.org/10.5091/plecevo.2012.668
Synopsis of the genus Chlorophytum (Asparagaceae) in Central Africa
(Democratic Republic of the Congo, Rwanda, Burundi)
Pierre Meerts1,* & Charlotte Sletten Bjorå2
1Herbarium et bibliothèque de botanique africaine, Université Libre de Bruxelles, CP 169, Avenue F.D. Roosevelt 50, BE-1050 Brussels,
Belgium
2Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, NO-0318 Oslo, Norway
Author for correspondence: pmeerts@ulb.ac.be
INTRODUCTION
“[… ] Les Chlorophytum paraissent des végétaux
extrêmement variables et […] les caractères sur lesquels on
croirait pouvoir se baser pour les grouper d’une façon sûre,
peuvent être, eux-mêmes, peut-être, sous la dépendance des
conditions extérieures. […] Il ne nous est possible que de
décrire, comme nouveautés, les plantes qui ne peuvent être
rapportées, sans le moindre doute à un type ancien. Ce sera
aux monographes de l’avenir à assigner à toutes ces espèces
provisoires leur véritable place dans la classication."
(De Wildeman 1921b: 11)
The circumscription of the genus Chlorophytum Ker Gawl.
(Asparagaceae in APG III 2009) was revised by Ober meyer
(1962), Marais & Reilly (1978), Nordal et al. (1990) and
Kativu & Nordal (1993) to include all the species formerly
included in Acrospira Baker, Debesia Kuntze, Dasystachys
Baker, Verdickia De Wild. and most of the African species
formerly included in Anthericum L.
The genus Chlorophytum has recently been revised
for the Flora of Tropical East Africa (FTEA) (Nordal et
al. 1997), the Flora Zambesiaca (FZ) (Kativu et al. 2008)
and the Flore du Gabon (Bjorå & Nordal 2010). In order to
prepare the treatment of the genus for the Flore d’Afrique
Centrale (FAC), we have critically revised all the material of
Chlorophytum from the Democratic Republic of the Congo
(D.R.Congo), Rwanda and Burundi. For a number of taxa,
phylogenetic relationships within the genus have been in-
vestigated using molecular markers, the results of which are
presented elsewhere. We here present a key to the species, a
check-list with a revised synonymy, and a brief phytogeo-
graphic sketch of the genus within the study area. When our
taxonomic treatment departs from FZ and FTEA, a concise
justication is provided. New combinations are coined when
necessary and new synonyms are reported. Detailed accounts
of the taxa, including citation of all specimens studied, will
be published in the Flore d’Afrique Centrale.
All rights reserved. © 2012 National Botanic Garden of Belgium and Royal Botanical Society of Belgium – ISSN 2032-3921
REGULAR PAPER
Background and aims – The genus Chlorophytum is revised for the Democratic Republic of the Congo,
Rwanda and Burundi, in order to prepare the treatment of the genus for the Flore d’Afrique Centrale.
Methods – Herbarium taxonomy and SE micrographs of seeds. All the material kept in BR, BRLU, BRVU,
FT, K, B and KIP has been examined, supplemented with recent eld observations in S Katanga. Nuclear
and chloroplast DNA markers have been used to clarify the taxonomic position of some species.
Key results – A total of 65 taxa (D.R.Congo: 62, Burundi: 19, Rwanda: 8) are reported for the study
area, i.e. fty species, two subspecies, twelve varieties and one form. A total of 27 taxa are new to the
D.R.Congo, ve to Burundi and two to Rwanda. Three taxa are endemic to the D.R.Congo. Three new
varieties are described (Chlorophytum africanum var. nordalianum, C. colubrinum var. upembense, C.
cameronii var. ruziziense). Twelve new combinations, 16 new synonyms and one new name (Chlorophytum
sphagnicolum) are proposed. An identication key is included. SEM pictures of seeds are included for
fteen taxa and colour photographs of living plants for fourteen taxa. A brief phytogeographic outline is
provided. Upper Katanga, with 47 taxa, appears as a prominent center of diversity of the genus.
Key words – taxonomy, Chlorophytum, identication key, Central Africa, phytogeography, revision,
Katanga, seed coat, SEM.
374
Pl. Ecol. Evol. 145 (3), 2012
MATERIAL AND METHODS
All the relevant material from D.R.Congo, Rwanda and Bu-
rundi in BR, BRLU, BRVU, B, FT, K, KIP and LSHI has
been examined. This material was supplemented with our
own collections and eld observations made in Upper Ka-
tanga in 2009 and 2010. No material pertaining to Antheri-
cum L. was found in the study area. No less than 69 names
have been found that are typied by specimens collected in
D.R.Congo and Burundi.
Conventional methods of herbarium taxonomy have
been applied. Seed shape and details of seed testa have been
studied with SEM. To reveal sister relations available spe-
cies were analyzed phylogenetically using nuclear ITS and
chloroplast rps16 and trnL-F sequences (for methods see
Bjorå et al. 2008). This information was also used to check
consistency of our taxonomic views with other recent taxo-
nomic publications. Our work has been considerably facili-
tated by the publication of the revision of the genus Chlo-
rophytum in FTEA (Nordal et al. 1997) and FZ (Kativu et
al. 2008), completed by Poulsen & Nordal (2005) for Guin-
eo-Congolian taxa. Important references for the study area
include De Wildeman (1909–1912, 1913c), Troupin (1955),
the Flore des Spermatophytes du Parc National Albert (Ro-
byns & Tournay 1955), the Flore des Spermatophytes du
Parc National de la Garamba (Troupin 1956) and the Flore
du Rwanda (Troupin 1988).
RESULTS AND DISCUSSION
In total, 65 taxa are recognized for the study area, i.e. fty
species, two subspecies, twelve varieties and one form. One
species new to science was published separately (Meerts
2011). Three new varieties, one new name, twelve new com-
binations and sixteen new synonyms are proposed. No less
than 27 taxa are new to D.R.Congo, ve to Burundi and two
to Rwanda (table 1). A key to identify all taxa is included.
Photographs of living plants are included for fourteen taxa
(gs 1–3). SE micrographs of seeds and seed coat structure
are provided for fteen taxa (gs 4–6). Seed shape and testa
anatomy are variable within the genus Chlorophytum and
provide important taxonomic characters (Marais & Reilly
1978, Nordal et al. 1990). The shape of the seed varies from
disc-shaped to deeply cup-shaped, more or less folded. The
periclinal walls of the testa cells vary from at to convex, in
a few cases with a distinct papilla up to 10 µm long, sepa-
rated by more or less deep furrows. In some cases, seed traits
are useful to discriminate species with very similar oral and
vegetative traits in particular in the complex of Guineo-Con-
golian rain forest species (Poulsen & Nordal 2005).
In many cases, variation in the studied material was more
extensive and complex than in adjacent regions (especially
so for C. blepharophyllum, C. cf. brachystachyum, the C.
cameronii complex, C. macrophyllum, C. sparsiorum, C.
rubribracteatum), and occasionally difcult to reconcile
with other recent revisions.
Phytogeographic outline
Table 1 gives the distribution of taxa in Central Africa as
presence/absence in Rwanda, Burundi and the nine phy-
togeographic regions of D.R.Congo as dened by Robyns
(1948). Species richness in D.R.Congo, Rwanda and Burun-
di (50 species, 65 taxa) is very close to that in FTEA (52 spe-
cies, 58 taxa) and FZ (56 species, 65 taxa). Upper Katanga
stands out as an important centre of diversity of the genus
for C Africa, with 47 taxa. Zambezian species are restricted
mostly to that region of D.R.Congo. Only few species oc-
cur both in the Guineo-Congolian region and in Katanga
(e.g. C. blepharophyllum, C. sparsiorum, C. stolzii, C. sub-
petiolatum). The occurrence in Katanga of disjunct popula-
tions of Southern African species is remarkable (C. recurvi-
folium, C. calyptrocarpum, possibly also C. macrosporum).
A few W African species are represented in Bas-Congo (C.
debile, C. staudtii, C. sphagnicolum, C. warneckei).
Endemic taxa are surprisingly few (two species: C. cordi-
folium, C. arcuatoramosum and two varieties: C. ca meronii
var. ruziziense and C. colubrinum var. upembense) consider-
ing the large area covered by the ora. However, fteen taxa
have a restricted area, just extending to adjacent countries
(table 1).
Upper Katanga alone counts 47 taxa i.e. by far the largest
richness of all phytogeographic districts of D.R.Congo. The
region of Lubumbashi, in particular, appears as an Eldorado
with 29 taxa within a 1000 km² area. Although sampling bias
cannot be ruled out, the high species density in that region
ts in well with the high species richness in the adjoining
regions of N Zambia (FZ), pointing to the “horn of Congo”
as a most important centre of diversity for the genus Chlo-
rophytum in tropical Africa. That region is currently under
heavy anthropic pressure due to the rapidly increasing hu-
man population and dramatic deforestation, and therefore
qualies as a hotspot (sensu Myers et al. 2000) for the genus,
with several taxa probably at risk.
Finally, it is signicant that 25 taxa are known from
less than six collections and six taxa by only one collection
(table 1) highlighting the urgent need for new plant collect-
ing efforts in D.R.Congo. Concerning Chlorophytum, future
sampling effort should be directed to the Katangan highlands
(Kundelungu, Mitwaba, Biano, Manika, Marungu), and the
regions adjoining the Congo-Angola borderline.
Key to Chlorophytum species in D.R.Congo, Rwanda and
Burundi
The identication of species requires fully developed mate-
rial comprising underground parts, leaves, owers, and ma-
ture fruits. Underground parts must be collected carefully as
tuber shape, size and arrangement have high taxonomic val-
ue. Flower colour, and stamen arrangement (actinomorphic
or zygomorphic) should be observed on fresh materials. In
some cases, examination of seed surface at high magnica-
tion (20–40×) is required. Plant size and habit often change
dramatically throughout the growing season. In particular,
leaves are often incompletely developed or lacking altogeth-
er (“hysteranthous” plants) at the time of owering and leaf
measurements at that stage may be grossly underestimated.
375
Meerts & Bjorå, Chlorophytum in Central Africa
Key to Chlorophytum species in D.R.Congo,
Rwanda and Burundi
1. A single ower at each inorescence node...............................................................................Group A
1. Flowers 2–5(–9) together, at least in part of the inorescence nodes (look carefully for ower buds or
naked pedicels at the base of owers at lowermost inorescence nodes).............................................2
2. Peduncle very short or lacking (< 3(–5) cm, to be measured from below the lowermost inorescence
branch or ower).......................................................................................................................Group B
2. Peduncle > 5 cm.....................................................................................................................................3
3. Peduncle with bracts or leaves..................................................................................................Group C
3. Peduncle naked, without bracts (except 1–2 immediately below inorescence) (leaves can have a
long sheathing base forming a pseudostem)..........................................................................................4
4. Plants having the following combination of characters: leaves distichous, pedicel articulated under
the middle, mature fruit transversally ridged, seed coat irregularly folded..............................Group D
4. Plants not having that combination of characters..................................................................................5
5. Inorescence: a branched panicle.............................................................................................Group E
5. Inorescence: a raceme.............................................................................................................Group F
Group A: 1 ower at all inorescence nodes
1. Peduncle with (1–)2–12 bract-like leaves..............................................................................................2
1. Peduncle naked......................................................................................................................................4
2. Perianth narrowly campanulate, constricted above the ovary; inorescence generally a dense raceme;
pedicels 1–3 mm; tepals 1-nerved....................................................................................C. colubrinum
2. Perianth widely open, not constricted; inorescence lax, with rachis visible between owers; pedicels
4–14 mm; tepals 3–5-nerved..................................................................................................................3
3. Leaves developed at owering; bracts auriculate at base and glossy; pedicels c. 5 mm long in fruit...
.......................................................................................................................................C. recurvifolium
3. Leaves developing after owering; bracts rounded at base and not glossy; pedicels 9–14 mm long in
fruit........................................................................................................................................C. nubicum
4. Inorescence branched..........................................................................................................................5
4. Inorescence a raceme...........................................................................................................................6
5. Plant < 20 cm high, with a rosette of 2–4 ovate leaves > 15 mm wide............................C. perfoliatum
5. Plant (20–)40–120 cm high, with leaves distichous, graminiform, < 12 mm wide (Fig. 2D)..................
...............................................................................................................................................C. vestitum
6. Leaves with ovate-elliptic lamina, contracted into a canaliculate petiole as long as the lamina and
forming an angle with it; < 7 owers to a plant; rain forest plant........................................C. occultum
6. Leaves with liform to ovate-elliptic lamina, not petiolate; > 7 owers to a plant; grassland and
woodland plants.....................................................................................................................................7
7. Leaves liform to grass-like, 1–4(–6) mm wide....................................................................................8
7. Leaves linear to ovate-elliptic, 5–40 mm wide......................................................................................9
8. Fruit 2.5–4 mm; leaves and peduncles spreading in all directions; rachis smooth.............C. warneckei
8. Fruit 6–10 mm; leaves distichous and peduncles not spreading in all directions; rachis papillate.........
.......................................................................................................................C. afne var. curviscapum
9. Leaves distichous; raceme lax, with rachis visible; perianth 7–9 mm long; roots wiry with big distal
tubers.........................................................................................................................C. afne var. afne
9. Leaves rosulate; raceme dense, with rachis most often not visible; perianth 4–7 mm long; roots more
or less eshy, without distal tubers (but sometimes with subsessile tubers)........................................10
10. Lower (abaxial) leaf surface glabrous; fruit glabrous; peduncle glabrous or rarely pubescent;
stolon lacking; rhizome very short without clavate tubers; plant 10–40 cm high at owering....
............................................................................................................................................C. africanum
10. Lower (abaxial) leaf surface pubescent; fruit papillate-hairy; peduncle pubescent; rhizome horizontal,
elongate, occasionally stolon-like, bearing normal roots and clavate tubers; plant 3–10 cm at
owering...............................................................................................C. leptoneurum var. katangense
Group B: Flowers 2–5 to a node; peduncle < 3(–5) cm
1. Inorescence developing before the leaves; leaves distichous without a petiole; cataphylls with brown
margin (g. 2E)..............................................................................................................C. hysteranthum
1. Inorescence developing with or after the leaves; leaves rosulate, or, when distichous, with a distinct
petiole; cataphylls uniform in color without brown margin..................................................................2
2. Peduncle densely covered with long papillae; leaf margin strongly crispate.............................C. minor
2. Peduncle smooth or scabrid; leaf margin smooth or undulate...............................................................3
376
Pl. Ecol. Evol. 145 (3), 2012
3. Raceme very lax, with internodes longer than owers and rachis clearly visible between them...........4
3. Raceme dense with internodes shorter than owers and rachis not visible...........................................9
4. Leaves liform or linear, without a petiole............................................................................................5
4. Leaves with ovate lamina contracted into a canaliculate petiole...........................................................8
5. Leaves attenuate in lower third, 5–12(–20) mm wide, laxly veined (space between nerves much wider
than nerves themselves); bracts 5–10 mm long, longer than pedicels.......................................C. debile
5. Leaves parallel-sided, not attenuate at base, 1–4(–6) mm wide, closely veined (space between nerves
narrower than nerves); bracts 1–5 mm long, shorter than pedicels.......................................................6
6. Fruit 6–10 mm long; inorescence a raceme, much shorter than leaves; peduncle and rachis papillate..
........................................................................................................................C. afne var. curviscapum
6. Fruit 2.5–4 mm long; inorescence most often branched, equalling or longer than leaves; rachis
smooth....................................................................................................................................................7
7. Roots wiry with big distal tubers; ower 5–8 mm long with anther 1.5–2.5 mm; leaves liform,
1–2 mm wide; leaves erect...........................................................................................C. angustissimum
7. Roots without tubers; ower 2.5–4 mm long with anthers 0.5–1.5 mm; leaves linear, 1 mm wide;
leaves and peduncules spreading in all directions...........................................................C. warneckei
8. Inorescence < 6 cm long (including peduncle); pedicel with joint in upper half; owers fewer than
7 to a plant; lamina < 8 cm long; tepals greenish; seed testa without apicules......................C. occultum
8. Inorescence 5–30 cm long (including peduncle); pedicel with joint in lower half; owers more
than 7 to a plant; lamina > 8 cm long (at least in some leaves); tepals whitish; seed testa apiculate
......................................................................................................................................C. alismatifolium
9. Pedicel articulated at tip; raceme much longer than broad; bracts persisting, prominent, partly
concealing young fruits, smooth and glabrous...............................................................C. stenopetalum
9. Pedicel without a joint; raceme usually no longer than broad; bracts neither persisting nor concealing
young fruits, often nely papillate on margin or abaxial surface and occasionally ciliate.................10
10. Peduncle and rachis virtually zero; inorescence lax, with owers more or less umbellate and
pedicels much longer than owers; leaf margin smooth.......................C. sp. B “near tetraphyllum”
10. Peduncle and rachis distinct, though very short; inorescence more congested, not umbellate,
with pedicels shorter than owers (but sometimes elongating in fruit); leaf margin often ciliate at
places....................................................................................................................................................11
11. Leaves (in herbarium) thin, membranous and translucent, without a pseudopetiole; rhizome with small
subsessile clavate tubers; fruiting pedicels < 10 mm long, straight; bracts thin, membranaceous, with
nerves not prominent; seed testa with periclinal walls verrucose (gs 3A & 6B)..............C. pusillum
11. Leaves (in herbarium) thick, more or less coriaceous, opaque, distinctly contracted into a pseudopeti-
ole as long as inorescence; roots with distal tubers; fruiting pedicels > 10 mm long, often recurved;
bracts opaque, with prominent nerves; seed testa with periclinal walls smooth (g. 6A)........................
............................................................................................................................................C. geophilum
Group C: Flowers 2–5 at a node; peduncle > 5 cm long with bracts or leaves
1. Flowers narrowly campanulate, not widely open, white; outer surface of tepals papillate...................2
1. Flowers more or less open (star shaped), green or white, not papillate.................................................3
2. Inorescence a raceme or rarely with 1–2 branches; rachis densely papillate; leaves < 2.5 cm broad;
plant < 1.2 m high.............................................................................................................C. longifolium
2. Inorescence with 3–8 branches curved upwards; rachis smooth (rarely with sparse papillae); leaves
> 2.5 cm broad; plant 1.7–3 m high (g. 1A & B)...................................................C. arcuatoramosum
3. Leaves < 5 mm wide; plant < 50 cm high..............................................................................................4
3. Leaves > 5 mm wide; plant > 50 cm high..............................................................................................5
4. Inorescence developing after the leaves; tepals tightly pressed against the ovary after anthesis; roots
spongy but not dilated into fusiform tubers; seeds c. 1 mm.......................................C. calyptrocarpum
4. Inorescence developing before the leaves; tepals free from the ovary after anthesis; roots dilated into
spindle-shaped tubers; seeds c. 2 mm....................................................................................C. nubicum
5. Anthers > 8 mm long; inorescence simple or basally with 1–5 short erect branches.........................6
5. Anthers < 8 mm long; inorescence a panicle with long spreading branches.......................................7
6. Leaves glabrous; shoot base abruptly dilated into a corm more than 4 cm broad, with concentric
ridges bearing regularly arranged brous remains < 1 cm long.................................................C. stolzii
6. Leaves with velvety pubescence; shoot base little dilated, less than 3 cm thick, without concentric
ridges but with irregularly arranged, brous remains > 1 cm long (g. 3E)........................C. velutinum
7. All leaves spaced along the stem; fruit < 4.5 mm long.........................................................C. ruahense
7. All leaves in a large rosette; fruit 4–15 mm long...................................................................................8
377
Meerts & Bjorå, Chlorophytum in Central Africa
8. Leaves, peduncle and pedicels pubescent.............................................................................C. hirsutum
8. Leaves, peduncle and pedicels glabrous................................................................................................9
9. Perianth greenish, campanulate with lower part of tepals enclosing ovary; tepals 3-nerved, > 15 mm
long...................................................................................................................................C. andongense
9. Perianth white, stellate; tepals 5–7-nerved, c. 8–12 mm long........................C. sp. A “near comosum”
Group D: Flowers 2–5 to a node; peduncle > 5 cm long, naked, often at; leaves distichous; pedicel
articulated below middle; mature fruit with transversal ridges; seed coat irregularly folded
1. Cataphylls and outer leaf sheaths with purplish spots or stripes................C. cameronii var. cameronii
1. Cataphylls and outer leaf sheaths without spots or stripes, but sometimes uniformly reddish..............2
2. Tip of outer tepals blackish (g. 3B)...............................................................................C. sphacelatum
2. Tip of outer tepals white, or sometimes pink or greenish......................................................................3
3. Inorescence with 1–5 branches............................................................................................................4
3. Inorescence a simple raceme................................................................................................................5
4. Inorescence branches 1–5, spreading, divaricate; tepals 8–15 mm long; lowest bract not foliaceous,
< 15 mm long; leaves 4–15 mm wide; plant not forming dense tufts....................................C. galpinii
4. Inorescence branches (0–)1(–2), erect, not spreading and divaricate; tepals < 9 mm long; lowest
bract foliaceous, 1–4 cm long; leaves 2–4 mm wide; plant forming dense tufts.....................C. staudtii
5. Leaf margin and lower surface of midrib with very long cilia...................................C. pilosicarinatum
5. Leaf margin and lower surface of midrib glabrous or with short hairs..................................................6
6. Cataphylls (and, often, bracts) with reddish tinge; leaves graminiform, < 5(–10) mm broad (g. 2F)...
..................................................................................................................................C. rubribracteatum
6. Cataphylls green; leaf width variable.....................................................................................................7
7. Leaves liform to linear, 0.7–3 mm wide, conduplicate........................................................................8
7. Leaves linear 3–30 mm wide, at to folded...........................................................................................9
8. Stamens not in two groups, much shorter than perianth, with anthers ≤ 2.5 mm; tepals 5–10 mm
long; leaves liform 0.7–2 mm wide, erect; bracts ± scariose with prominent nerves; peduncle
attened to terete, with prominent ribs (in herbarium); plant in dense tufts; in moist soil, W Congo....
......................................................................................................................................C. sphagnicolum
8. Stamens in two groups, slightly shorter than perianth and with anthers > 2.5 mm; tepals 7–15 mm;
leaves 2–3 mm wide, spreading; bracts not scariose with nerves not prominent; peduncle attened;
plant not cespitose; in dry soil, E Congo and Burundi................................C. cameronii var. ruziziense
9. Cataphylls with brown band at margin (g. 2E); inorescence developing before the leaves, not
exceeding 15 cm long including the peduncle...............................................................C. hysteranthum
9. Cataphylls without brown band at margin; inorescence generally > 13 cm including peduncle,
developing before or after the leaves...................................................................................................10
10. Roots thick, carrot-like, without distal tubers ...............................................................C. subpetiolatum
10. Roots wiry, with distal tubers................................................................................................................11
11. Leaves (3–)4–30 mm wide, with margin and lower surface of mid-nerve smooth to papillose, not
denticulate; inorescence unbranched, without a foliaceous bract; tepals 7–15 mm; anther 4–6 mm;
fruit 5–7 mm long.....................................................................................C. cameronii var. pterocaulon
11. Leaves 2–4 mm wide, with margin scabrid-denticulate to ciliate and lower surface of midrib
denticulate-scabrid; inorescence often with one slender basal branch subtended by a foliaceous bract
1–4 cm long; tepals 7–10 mm; anther 3–4 mm; fruit < 5.5 mm long.....................................C. staudtii
Group E: Flowers 2–5 to a node; peduncle > 3 cm long, without bracts; fruit smooth, not transver-
sally ridged; inorescence branched (exceptionally robust specimens of group F, with branched
inorescence, may key out here)
1. Inorescence with sterile plantlets.........................................................................................................2
1. Inorescence without sterile plantlets...................................................................................................3
2. Tepals 8–10 mm long, 5-nerved; anthers 4–5 mm long; inorescence (with peduncle) 50–150 cm high;
leaves 10–20 mm wide, linear, parallel-sided, without a pseudopetiole.....C. sp. A “near comosum”
2. Tepals 5–8 mm long, 3-nerved; anthers 2–3 mm long; inorescence (with peduncle) 10–60 cm high;
leaves 10–55 mm wide, linear or elliptic with a pseudopetiole......................................C. sparsiorum
3. Leaves hairy or ciliate at places............................................................................................................4
3. Leaves glabrous or papillate..................................................................................................................8
4. Leaves < 3 mm wide, liform to linear..................................................................................................5
4. Leaves > 3 mm wide, of variable shape.................................................................................................6
5. Roots wiry with terminal tubers; fruit about as long as broad, emarginate.................C. angustissimum
5. Roots eshy without tubers; fruit longer than broad, not emarginate........................C. calyptrocarpum
378
Pl. Ecol. Evol. 145 (3), 2012
6. Leaf surface glabrous; margin ciliate (cilia sometimes caducous)..........................C. blepharophyllum
6. Leaf surface hairy...................................................................................................................................7
7. Leaves graminiform, < 12 mm wide, distichous....................................................................C. vestitum
7. Leaves elliptic, 40–80 mm wide, rosulate.............................................................................C. hirsutum
8. Flower > 15 mm long, greenish, urn-shaped; tepals with short ligule above perianth constriction
..........................................................................................................................................C. andongense
8. Flower 3–10 mm long, greenish or white, not urn-shaped; tepals without a ligule..............................9
9. Leaves graminiform, < 15 mm wide, neither attenuate in lower half nor petiolate, distichous..........10
9. Leaves narrowly to broadly ovate or elliptic, petiolate or attenuate, or strap-shaped but, then, rosulate
(not distichous).....................................................................................................................................11
10. Pedicels and owers papillate-pubescent...........................................................................C. pubiorum
10. Pedicels and owers glabrous...........................................................................................C. polystachys
11. Seeds 3–5 mm; leaves distichous; lamina angular-ovate, forming an angle with the petiole and more
or less horizontal..........................................................................................................C. alismatifolium
11. Seeds < 3.5 mm; leaves rosulate; lamina elliptic, ovate, or linear, not forming an angle with the
petiole...................................................................................................................................................12
12. Plant with leaves developing after the inorescence, linear, < 10 mm wide...........................................
.............................................................................................................C. gallabatense var. micranthum
12. Plant with leaves developing before the inorescence, linear or elliptic, > 10 mm wide....................13
13. Pedicels and rachis with hairs or cylindric papillae c. 0.4 mm long....C. gallabatense var. oribundum
13. Pedicels glabrous or rough with papillae < 0.2 mm.............................................................................14
14. Tepals 7–10 mm long, 5-nerved, anthers 4–5 mm..........................................C. sp.A “near comosum”
14. Tepals 4–7 mm long, 3-nerved; anthers 1–3.5 mm..............................................................................15
15. Lamina abruptly contracted into a distinct petiole, with base truncate or rounded.............................16
15. Lamina progressively attenuated towards base, or strap-shaped, without a distinct petiole...............19
16. Petioles sheathing peduncule and forming a pseudostem (g. 3F).............................C. zingiberastrum
16. Petioles not sheathing peduncle; pseudostem lacking..........................................................................17
17. Robust plants, with at least one leaf > 45 × 6 cm, drying black....................................C. orchidastrum
17. Medium robust plants, with all leaves < 38 × 6 cm, drying blackish or greenish...............................18
18. Lamina widest in the lower one-third, 7–15 × 2–4.5 cm; petiole longer than lamina; plants most often
drying blackish green to black; rachis smooth; anther 1–1.5 mm....................................C. lancifolium
18. Lamina broadest near the middle, 12–25 × 4.5–6 cm (measurements only for forms with petiolate
leaves); pseudopetiole shorter than lamina; plants drying greenish; rachis most often scabrid; anther
2–3 mm............................................................................................................................C. sparsiorum
19. Tubers on lateral rootlets; plant not blackening in herbarium; inorescence generally with 2 branches
or more................................................................................................C. gallabatense var. gallabatense
19. Tubers on main roots or lacking; plants blackening or not; inorescence generally with 0–1 branch
(when more than 3 branches then leaves petiolate).............................................................................20
20. Robust plants, drying black, with at least one leaf > 38 × 4.5 cm; capsules 7–12 mm long...................
.........................................................................................................................................C. lipendulum
20. Less robust plants, with all leaves < 38 ×4.5 cm; leaves blackening or not; capsules 3.5–8 mm long.....
.............................................................................................................................................................21
21. Lamina widest in the lower one-third, equalling pseudopetiole or shorter than it; plants drying dark
green to blackish; rachis smooth; anthers 1–1.5 mm........................................................C. lancifolium
21. Lamina widest near the middle, longer than the pseudopetiole; plant blackening or not (when
blackening: leaves strap-shaped); rachis most often scabrid; anthers 2–3 mm..............C. sparsiorum
Group F: Flowers 2–5 to a node; peduncle > 3 cm long, without bracts; fruit not transversally
ridged; inorescence a raceme (depauperate specimens of group E with simple raceme may oc-
casionally key out here)
1. Inorescence with sterile plantlets.........................................................................................................2
1. Inorescence without sterile plantlets....................................................................................................3
2. Tepals 8–10 mm long, 5-nerved; anthers 4–5 mm long; inorescence (with peduncle) 50–150 cm
high; leaves linear, 10–20 mm wide, not attenuate in lower half...................C. sp. A “near comosum”
2. Tepals 5–8 mm long, 3-nerved; anthers 2–3 mm long; inorescence (with peduncle) 10–60 cm high;
leaves linear or elliptic with a pseudopetiole..................................................................C. sparsiorum
3. Leaf margin ciliate; leaves not distichous..............................................................................................4
3. Leaf margin not ciliate or rarely ciliate but then leaves distichous........................................................5
379
Meerts & Bjorå, Chlorophytum in Central Africa
4. Flowers pure white; fruits generally verrucose or tuberculate, slightly trigonous; leaves thin and
membranaceous, spreading, rosulate, not sheathing the peduncle at fruiting stage (though sheathing
when owering); leaf margin generally undulate or crispate, neither thickened nor recurved (g. 1D
& E)......................................................................................................................C. cf. brachystachyum
4. Flower colour variable, never pure white: yellowish, greenish, brownish, purplish; fruit not verrucose,
sharply triquetrous; leaves thick and somewhat eshy to brose, tightly sheathing the base of the
peduncle at fruiting stage, thus appearing superposate; leaf margin not undulate, often recurved or
thickened..................................................................................................................C. blepharophyllum
5. One leaf, broadly ovate, abruptly contracted into a canaliculate petiole which tightly clasps the
peduncle............................................................................................................................C. cordifolium
5. Almost always 2 leaves or more; when a single leaf, petiole not clasping the peduncle......................6
6. Rachis and upper part of peduncle shortly pubescent; leaves generally distichous...............................7
6. Rachis and upper part of peduncle glabrous or papillate; leaves rosulate or, more rarely, distichous but
then rachis not pubescent.......................................................................................................................9
7. Roots carrot-like, i.e. thick at base and progressively attenuate, without tubers; leaves narrowly
ovate-elliptic, more or less contracted into a pseudopetiole (sometimes very short < 5 mm); seed coat
irregularly folded..........................................................................................................C. subpetiolatum
7. Roots wiry, with distal tubers; leaves linear, never attenuate near base; seeds disk- or saucer-
shaped....................................................................................................................................................8
8. Cataphylls with brown margin; plant < 15 cm high at owering, with leaves appearing after the
owers (g. 2E).............................................................................................................C. hysteranthum
8. Cataphylls without brown band; plant 15–60 cm high at owering, with leaves appearing before the
owers........................................................................................................................C. afne var. afne
9. Leaves markedly distichous; lamina angular-ovate, broadest under middle, abruptly contracted into a
canaliculated petiole and forming an angle with it; seeds 3–5 mm..............................C. alismatifolium
9. Leaves rosulate; lamina linear to narrowly elliptic, progressively attenuate at base, without a
canaliculated petiole and usually not forming an angle with it; seeds 1.5–2.5 mm............................10
10. Leaves linear to narrowly elliptic, < 6 mm wide; bracts purplish-brown; roots thick at base, distally
attenuate; leaves appearing before the owers (g. 1F)..................................................C. burundiense
10. Leaves linear to elliptic 5–80 mm wide (when 5 mm wide, then leaves appearing after the
inorescence); bract green to whitish; roots not thick at base, generally with distal tubers................11
11. Tepals 5–6-nerved, 7–13 mm long; bracts 10–30(–50) mm long, often whitish and very conspicuous
before anthesis, soon withering and turning brown; anthers 3–5 mm long.........................................12
11. Tepals 3(–5)-nerved, 3.5–8 mm long; bracts 2–10 mm long, not whitish before anthesis; anthers
1.5–3.5 mm..........................................................................................................................................14
12. Inorescence (with peduncle) 50–150 cm high, often pseudoviviparous, very lax with rachis visible;
leaves linear, 10–20 mm wide, not attenuate in lower half; bracts 5–10 mm long, never whitish and
conspicuous....................................................................................................C. sp. A “near comosum”
12. Inorescence (with peduncle) < 60 cm high, never pseudoviviparous, relatively dense with rachis
often concealed by fruits; leaves 25–80 mm broad, attenuate in lower half; bracts 10–30(–50) mm
long, whitish and very conspicuous before anthesis............................................................................13
13. Fruit obtusely trigonous to rounded in cross-section, not emarginate; seeds variable in shape, from
saucer-shaped to strongly plicate (g. 5A); seed testa with raised periclinal walls (g. 5B); raceme
5–20 cm long, dense, with fruits concealing rachis and lowermost nodes not distant; tepals 7–9 mm;
anthers 3–4 mm, equalling lament; tubers lateral; pedicel articulated at tip (g. 2A–C).......C. clarae
13. Fruit trigonous to triquetrous, emarginate; seeds at to slightly saucer-shaped (g. 5E); seed testa
with periclinal walls shallowly convex or attened (g. 5F); raceme 15–45 cm long, lax to moderately
dense, with rachis visible and lowermost nodes often distant; tepals 8–13 mm; anthers 4–5 mm,
longer than lament; tubers on main roots; pedicels articulated in upper half.............C. macrophyllum
14. Plant with inorescence appearing before the leaves; leaves linear, < 1.0 cm wide; rootstock with
bres....................................................................................................C. gallabatense var. micranthum
14. Leaves completely developed at owering, narrowly to broadly elliptic 12–60 mm wide; rootstock
not brous............................................................................................................................................15
15. Pedicel articulated at the tip; raceme densely owered, with rachis not visible; rachis smooth......
........................................................................................................................................C. stenopetalum
15. Pedicel articulated below the tip; raceme less densely owered, generally with rachis visible; rachis
smooth or papillate...............................................................................................................................16
380
Pl. Ecol. Evol. 145 (3), 2012
16. At least one leaf longer than 38 cm or wider than 5 cm; plant blackish in herbarium; peduncle
robust and stiff, generally > 2.5 mm thick below lowermost ower (in herbarium), always rough...
.........................................................................................................................................C. lipendulum
16. All leaves < 38 cm long and with lamina < 5 cm wide; plant blackish or green in herbarium; peduncle
more slender, generally < 2.5 mm thick below the lowermost ower (in herbarium), rough or
smooth.............................................................................................................................................17
17. Lamina ovate, broadest below the middle, abruptly contracted into a petiole as long as or longer than
lamina; plant blackish-green in herbarium........................................................................C. lancifolium
17. Lamina elliptic to sublinear, broadest at the middle, progressively attenuate at base; plant blackish or
green in herbarium (when blackish, leaves linear)...............................................................................18
18. Seed testa with periclinal walls at to convex; pedicel articulated near or above the middle, straight;
fruit (3–)4–7(–8) × (4–)5.5–7(–9) mm; leaves 12–50 mm wide, with more than (17–)21 nerves..........
.........................................................................................................................................C. sparsiorum
18. Seed testa with periclinal walls apiculate; pedicel articulated below the middle, often recurved when
fruiting; fruit c. 3–4 × 4 mm; leaves 5–15(–25) mm wide, with 11–19 nerves.........................C. debile
Chlorophytum species are often polymorphic. The key
allows to some extent for variation of polymorphic species,
which key out at different places. However, it does not allow
for exceptional character states. The key is based mostly on
original observations on material from D.R.Congo, Rwanda
and Burundi, supplemented by literature data when the mate-
rial collected in the study area was too scarce.
For the description of leaf shape, we follow the recom-
mendations of Anonymous (1962) and, in particular, we
avoid the word “lanceolate”. For conduplicate leaves, leaf
width is expressed as twice the apparent width.
Check-list of species with synonyms
In the following check-list, citation of synonyms is limit-
ed to (i) new synonyms, (ii) all the names typied by type
specimens collected in the study area (D.R.Congo, Burundi,
Rwanda), (iii) all the names that were used in one of the fol-
lowing references: Durand & Durand (1909), De Wildeman
(1909–1912, 1921a), Lebrun et al. (1948), Robyns & Tour-
nay (1955), Troupin (1956, 1988), Schmitz (1971), Lewalle
(1972).
In this account, only one representative specimen is cited
for each phytogeographic district of D.R.Congo as dened
by Robyns (1948). Occasionally, more specimens are cited
for taxa of particular interest. Full species descriptions and
extensive lists of specimens studied will be published in the
Flore d’Afrique Centrale.
1. Chlorophytum afne Baker (1875: 160 & tab. 104); Baker
(1876: 327); Baker (1898: 507); Hanid (1974: 586); Kativu
(1994: 49); Nordal et al. (1997: 23); Kativu et al. (2008: 60).
– Type: Tanzania, Tabora Ditrict, Unyamwezi, Rubugwa,
1860, Speke & Grant s.n. (holo-: K, barcode K000257040).
Two varieties can be recognized:
1. Peduncle erect, straight, > 10 cm long; inorescence
about as long as the leaves; leaves > 10 mm wide.......
........................................................C. afne var. afne
1. Peduncle curved at base, more or less exuous, < 6
cm long; inorescence shorter than the leaves; leaves
< 8 mm wide.......................C. afne var. curviscapum
1a. Chlorophytum afne Baker var. afne
Other representative specimen examined – D.R.Congo: XI:
Lubumbashi, 1939, Salésiens 1147 (BR).
Bodenghien 331 (BR) is an extremely robust specimen with
a raceme of up to 22 cm long.
1b. Chlorophytum afne Baker var. curviscapum (Poelln.)
Hanid (Hanid 1974: 588); Nordal et al. (1997: 23); Ka-
tivu et al. (2008: 61). – Chlorophytum curviscapum Poelln.
(Poellnit z 1942: 122). – Type: Tanzania, Dodoma dist., Uy-
ansi, near Chaya, 3 Jan. 1926, Peter 45827 (holo-: B).
Other representative specimens examined – Burundi: Territ. Bu-
banza, Plaine de la Rusizi, 800 m, 30 Oct. 1973, Reekmans 2872
(BR).
Rwanda: Parc Nation. Kagera, Gabiro, savane herbeuse à Theme-
da-Hyparrhenia, alt. 1500 m, 3 Dec. 1944, Germain 2858 (BR, K).
Cultivated: Yangambi, originaire de la Kagera, Germain 7499 (BR,
K).
A dwarf plant, 5–13 cm high, with grass-like leaves 2–5
mm wide, very short papillate peduncle, and more or less
prostrate inorescence. The cited collections are somewhat
atypical in having smooth leaf margins and recurved fruiting
pedicels. They are supercially similar to C. inconspicuum
(Baker) Nordal, but with ovoid distal tubers, not progres-
sively dilated roots.
C. afne var. curviscapum, though widespread in Zam-
bia and Tanzania, has apparently not been collected yet in
D.R.Congo. Duvigneaud 4380 (BRLU) approaches var. cur-
viscapum, but it has erect peduncle, liform leaves, and tu-
bers with a reticulate surface. More material is needed.
2. Chlorophytum africanum (Baker) Engl. (Baker & Eng-
ler 1892: 470); Kativu (1994: 51); Nordal et al. (1997: 21);
Kativu et al. (2008: 66). – Type: Tanzania, Tabora Dist.,
Rubuga (Rubugwa) 1860, Grant s.n. (lecto-: K, barcode
K000365093).
381
Meerts & Bjorå, Chlorophytum in Central Africa
I II III IV V VI VII VIII IX XI Burundi Rwanda Observations
number of
collections
if ≤5
Distribution
afne var. afne xnew to
D.R.Congo 4 Sudano-Zambezian
afne var.
curviscapum x x new to Burundi
and Rwanda 2 Sudano-Zambezian
africanum var.
africanum xZambia, Tanzania,
S Katanga
africanum var.
nordalianum xnew to
D.R.Congo S Katanga, N Zambia
africanum var.
silvaticum x x x new to Burundi Zambezian-E African
alismatifolium x x x Guineo-Congolian
andongense x x Sudano-Zambezian
angustissimum x x new to
D.R.Congo Sudanian – E trop. African
arcuatoramosum x x Endemic D.R.Congo
blepharophyllum
var.
amplexicaule
x x x new to
D.R.Congo
Katanga, N Zambia, W
Tanzania
blepharophyllum
var.
blepharophyllum
x x x x x x x x x x Afro-tropical
cf.
brachystachyum xnew to
D.R.Congo Sudano-Zambezian
burundiense x Burundi, W Tanzania
calyptrocarpum xnew to
D.R.Congo Zambezian-S African
cameronii var.
cameronii x x x x x x Afro-tropical
cameronii var.
grantii xnew to
D.R.Congo 5 E tropical Africa
cameronii var.
pterocaulon x x x x Afro-tropical
cameronii var.
ruziziense x x
new to
D.R.Congo and
Burundi
E D.R.Congo, Burundi
Table 1 – Check-list of Chlorophytum species in the Democratic Republic of the Congo, Rwanda and Burundi, with their geographic distribution.
Numbering of phytogeographic districts of D.R.Congo after Robyns (1948): I: Côtier; II: Mayombe; III: Bas-Congo; IV: Kasaï; V: Bas-Katanga; VI: Forestier central; VII: Ubangi-Uele;
VIII: Lac Albert; IX: Lacs Edouard & Kivu; XI: Haut-Katanga.
382
Pl. Ecol. Evol. 145 (3), 2012
I II III IV V VI VII VIII IX XI Burundi Rwanda Observations
number of
collections
if ≤ 5
Distribution
clarae x x new to
D.R.Congo
Katanga, N Zambia,
SW Tanzania
colubrinum var.
colubrinum x x x x x x Broad Zambezian
colubrinum var.
upembense xnew to
D.R.Congo 3Endemic D.R.Congo
(Upper Katanga)
cordifolium x 5 Endemic D.R.Congo
(Upper Katanga)
debile x x new to
D.R.Congo 3 W African
lipendulum x x x x x x Guineo-Congolian
gallabatense var.
oribundum xnew to
D.R.Congo 2 Zambezian
gallabatense var.
gallabatense x x x x x Sudano-Zambezian
gallabatense var.
micranthum x x x x x E African-Sudanian
galpinii var.
matabelense x 3 Zambezian - S African
geophilum x x new to
D.R.Congo Sudanian, E African
hirsutum x x Albertine rift
hysteranthum x 4 S Katanga, N Zambia
lancifolium
subsp. cordatum x x x C African
lancifolium
subsp.
lancifolium
x x 3 C African
lancifolium
subsp. togoense x W Guineo-Congolian
leptoneurum var.
katangense x 3 Katanga, Zambia, Malawi
longifolium var.
aureum x 3 Sudanian
macrophyllum x x ? x Sudano-Zambezian
minor xnew to
D.R.Congo 1SW Tanzania, N Zambia,
S Katanga
Table 1 (continued) – Check-list of Chlorophytum species in the Democratic Republic of the Congo, Rwanda and Burundi, with their geographic distribution.
383
Meerts & Bjorå, Chlorophytum in Central Africa
I II III IV V VI VII VIII IX XI Burundi Rwanda Observations
number of
collections
if ≤ 5
Distribution
nubicum xnew to
D.R.Congo 2 Sudano-Zambezian
occultum x 1 N D.R.Congo, Uganda
orchidastrum x x x x x x Guineo-Congolian
perfoliatum xnew to
D.R.Congo 1 Zambia, Malawi, Katanga
pilosicarinatum xnew to
D.R.Congo 2 Angola, Katanga
polystachys xnew to
D.R.Congo 1 Sudano-Zambezian
pubiorum xnew to
D.R.Congo 1 Zambezian
pusillum x Sudano-Zambezian
recurvifolium xnew to
D.R.Congo 3 S African
ruahense xnew to
D.R.Congo 4Katanga, W Tanzania,
N Zambia
rubribracteatum x x Zambezian
sparsiorum x x x x x x x x x x x x Afro-tropical
sphacelatum x x x x Zambezian
sphagnicolum x x new to
D.R.Congo Angola-Congo
staudtii x x new to
D.R.Congo 5 W C African
stenopetalum var.
latifolium xnew to
D.R.Congo S Katanga, N Zambia
stenopetalum var.
stenopetalum x x x x Sudano-Zambezian
stolzii x x x x x x x broad Zambezian
subpetiolatum
var. pilosifolium xS Katanga, N Zambia,
SW Tanzania
subpetiolatum
var.
subpetiolatum
x x x x x x x x x Sudano-Zambezian
Table 1 (continued) – Check-list of Chlorophytum species in the Democratic Republic of the Congo, Rwanda and Burundi, with their geographic distribution.
384
Pl. Ecol. Evol. 145 (3), 2012
Table 1 (continued) – Check-list of Chlorophytum species in the Democratic Republic of the Congo, Rwanda and Burundi, with their geographic distribution.
I II III IV V VI VII VIII IX XI Burundi Rwanda Observations
number of
collections
if ≤ 5
Distribution
subpetiolatum
var.
subpetiolatum f.
variegatum
x x 4 D.R.Congo, Congo, Sudan,
Cameroon
velutinum x Zambia, Malawi, S Katanga
vestitum subsp.
pilosissimum x Zambezian
warneckei xnew to
D.R.Congo 1 W African
zingiberastrum x x
new to
D.R.Congo and
Burundi
Zambezian
Sp. A “near
comosum”x x new to Burundi
and Rwanda ?
Sp. B “near
tetraphyllum”xnew to
D.R.Congo 2 ?
The pattern of pubescence is variable and of taxonomic
value. Three varieties can be recognized, of which the rst
two were treated as distinct species by FTEA and FZ.
1. Leaf margin with dense, crispate, reddish hairs...........
........................................C. africanum var. africanum
1. Leaf margin glabrous or, more rarely, with short pallid
cilia.............................................................................2
2. Peduncle and abaxial surface of cataphylls pubescent
throughout...................C. africanum var. nordalianum
2. Cataphylls glabrous; peduncle glabrous or sparsely
pubescent in upper part...C. africanum var. silvaticum
2a. Chlorophytum africanum (Baker) Engl. var. africanum
– Caesia africana Baker (Baker 1875: 160 & tab. 103a). –
Dasystachys africana (Baker) T.Durand & H.Durand (Du-
rand & Durand 1909: 569). – Dasystachys grantii Benth.
(Bentham & Hooker 1883: 789); Baker (1898: 513); De
Wildeman (1921a: 29), nom. illeg.
Acrospira giorgii De Wild. & Ledoux (De Wildeman
1930: 92). – Type: Env. Lubumbashi, 1923, De Giorgi s.n.
(holo-: BR, barcode BR0000008767325), synon. nov.
Other representative specimens examined – D.R.Congo: XI: Pa-
turage de la Karavia, ferme CSK, Dec. 1933, Quarré 3645 (BR).
A rare taxon in the study area apparently restricted to the
region of Lubumbashi.
2b. Chlorophytum africanum (Baker) Engl. var. nordalia
num Meerts, var. nov.
A typo differt pedunculo et rachide omnino pubescente, cata-
phyllis extus pubescentibus et foliis marginibus non ciliatis.
– Type: D.R.Congo, Haut-Katanga, 5 km NW of Lubum-
bashi, dembo de la Kiamalale, 14 Nov. 1961, Schmitz 7498
(holo-: BR). Fig. 4A & B.
Other representative specimens examined – D.R.Congo: XI:
Lubumbashi, Katuba, dalle latéritique, Dec. 1952, Schmitz 4261
(BR, KIP); Keyberg, forêt claire sur sol relativement frais, Dec.
1947, Schmitz 1250 (BR); Lubumbashi, Kasapa, 23 Oct. 1961, Poe-
lman 74 (BR); Lubumbashi, Keyberg, Vallée Katapa, 21 Nov. 1956,
Detilleux 135 (BR, KIP); Kasumbalesa, savane dérivée en contre-
bas de la colline boisée, 3 Dec. 1959, Duvigneaud 4377L (BRLU).
Other collections examined – Zambia: Ndola, 1957, Duvigneaud
s.n. (BRLU); Ndola, 27 Nov. 1953, Fanshawe F518 (BR, K); Kit-
we, gravelly roadsides, 25 Dec. 1966, Mutimushi 1696 (K); Kitwe,
miombo on laterite, 23 Nov. 1968, Mutimushi 2836 (K).
This new variety is well characterized by a pubescent
peduncle, rachis and abaxial side of the cataphylls. The
cataphylls have a network of dark green veins. It also dif-
fers from var. silvaticum in its seed testa structure, having
strongly buldging, hemisphaeric periclinal walls (Fig. 4A &
B) (vs. at to conical periclinal walls in var. silvaticum). C.
africanum var. nordalianum, having a pubescent peduncle,
can be confounded with C. leptoneurum. The latter species is
usually less robust, has a pubescent abaxial leaf surface and
a creeping rhizome.
Plants with the distinctive traits of var. nordalianum had
already been reported from N Zambia by Kativu et al. (2008:
66).
Var. nordalianum appears to be endemic to the extreme
south-east of D.R.Congo and adjoining regions of Zambia.
385
Meerts & Bjorå, Chlorophytum in Central Africa
2c. Chlorophytum africanum (Baker) Engl. var. silvaticum
(Dammer) Meerts, comb. & stat. nov.
Chlorophytum silvaticum Dammer, Botanische Jahrbücher
für Systematik, Panzengeschichte und Pan zengeographie
48: 365. 1912 (Dammer 1912); Kativu (1994: 75); Nordal
et al. (1997: 20); Kativu et al. (2008: 65). – Type: Tanzania,
Kilwa Dist., Donde, near Kwa Mpanda, 22 Dec. 1900, Busse
1310 (holo-: B, iso-: EA).
Other representative specimens examined – D.R.Congo: IX:
Plaine de la Rusizi, Feb. 1950, Germain 6084 (BR); XI: Territ. Mit-
waba, Simama, sur riv. Dikuluwe, plaine alluviale Dikuluwe-Lura,
19 Jan. 1956, Brynaert 436 (BR).
Burundi: Plaine de la Rusizi, Germain 5546 (BR); Territ. Bujum-
bura, Gihungwe, savane, alt. 850 m, 26 Nov. 1972, Reekmans 2182
(BR).
Intermediates between var. africanum and var. silvaticum
(short, pallid marginal hairs) are occasionally found (e.g.
Germain 5524 (BR, K)).
3. Chlorophytum alismatifolium Baker (Baker 1876: 324);
Baker (1898: 496); Engler (1908: 308, g. 208 A, B);
Troupin (1956: 188); Hepper (1968b: 102); Poulsen & Nor-
dal (2005: 13, gs 1H, 4, 5, 28); Bjorå & Nordal (2010: 4).
– Type: Equatorial Guinea, banks of the Gaboon river, Jul.
1861, Mann 1030 (holo-: K).
Other representative specimens examined – D.R.Congo: V:
Upemba, Rivière Kanonga, galerie forestière, De Witte 5588 (BR);
VI: Penghe, forêt ombragée au bord de l’Ituri, 5 Feb. 1914, Be-
quaert 2312 (BR); VII: Parc Nat. Garamba, Galerie à Irvingia, 15
May 1950, Noirfalise 387 (BR).
The material from D.R.Congo is more variable than re-
ported by Poulsen & Nordal (2005). Leaf length often ex-
ceeds 25 cm, pedicels are up to 10 mm (vs. 5–8 mm), bracts
are larger (10 × 3 mm vs. 5 × 1 mm), and tepals are often
longer (up to 7 mm vs. 3–4 mm). Some collections from
Sudanian Congo (Garamba) have lateral tubers [De Saeger
1207, 1342 (BR)], a trait typical of the East African, closely
related, C. holstii Engl. More material is needed.
Troupin 1802 (BR, K) has obovoid fruits which are much
longer than wide (9 × 5 mm) and pedicels articulated above
the middle, but other traits correspond to those of C. alis-
matifolium. Keay 58 (K), from a plant cultivated at Kew, is
atypical in having leaves up to 42 cm long, and an 80 cm
long inorescence with most owers replaced by plantlets.
Louis 11815 (BR, BRLU, K) with very long (up to 20
cm) foliaceous lowermost bracts has sometimes been mis-
identied as C. bracteatum Hua, but other traits are typical
of C. alismatifolium.
4. Chlorophytum andongense Baker (Baker 1878a: 260);
Baker (1898: 506); Troupin (1956: 188); Obermeyer (1962:
698); Hepper (1968b: 102); Kativu (1994: 52); Nordal et al.
(1997: 10); Kativu et al. (2008: 70, g. 13.1.12). – Type: An-
gola, Pungo Andongo, Dec. 1856, Welwitsch 3770 (holo-:
BM, iso-: K). Fig. 4C.
Chlorophytum longipes Baker (Baker 1878b: 325); De
Wildeman (1913c: 15; 1921a: 28). – Types: Sudan, Jur Ghat-
tas, 30 May 1869, Schweinfurth 1801 (syn-: K) & 10 Jul.
1869, Schweinfurth 2045 (syn-: K).
Other representative specimens examined – D.R.Congo: VII:
Parc Nat. Garamba, Bagbele, Noirfalise 521 (BR); XI: Lubum-
bashi, termitière en forêt claire, 13 Jan. 1954, Schmitz 4545 (BR).
C. andongense is closely related to the southern African
species C. macrosporum Baker. The latter is said to have
longer fruits (11–15 mm vs. 5–9 mm in C. andongense) and
narrower leaves (1.4–2.5 cm wide vs. 2.5–9 cm in C. andon-
gense). A few collections from Katanga have long fruits and
relatively narrow leaves (e.g. Billiet & Jadin 4150 (BR)).
Katangan collections are variable in leaf width from nar-
rowly strap-shaped with more or less undulate margin (traits
of C. macrosporum) to elliptic (trait of C. andongense) (e.g.
Quarré 2733 (BR, P). Duvigneaud 5416A&C (BRLU) is an-
other critical specimen with “macrosporum-like” fruits and
“andongense-like” leaves. We feel that C. andongense and
C. macrosporum might actually not be distinct at species
rank. The group denitely needs revision and, after much
hesitation, we provisionally keep all Katangan collections
under C. andongense.
5. Chlorophytum angustissimum (Poelln.) Nordal (Nordal
et al. 1997: 42). – Anthericum angustissimum Poelln. (Po-
ellnitz 1942: 26). – Type: Tanzania, Tabora District, Ngulu,
from Malongwe to Nyahua, Jan. 1926, Peter 34505 (holo-:
B, not seen). Fig. 4E & F.
Anthericum nigericum Hepper (Hepper 1968a: 459, 455
g. 5/4-6); Hepper (1968b: 97). – Type: Nigeria, Zaria Prov.,
Anaria Forest reserve, 1 Jul. 1957, Keay 37081 (holo-: K).
Other representative specimens examined – D.R.Congo: IX:
Vallée de la Rusizi, groupement de Luberidi, Centre zootechnique,
Paddock III, 14 Jan. 1954, J.F. Laurent 1015bis (BR); Plaine de la
Ruzizi, savane à Themeda, alt. 800 m, Jan. 1950, Germain 5541
(K).
Burundi: Prov. Bubanza, Gihungwe, steppe à Bulbine abyssinica, 8
Mar. 1981, Reekmans 9802 (BR, BRVU, K).
This species was erroneously referred to as Anthericum
calyptrocarpum Baker by Lewalle (1972).
Collections from the study area often have peduncle >
5 cm, i.e. longer than reported in FTEA. Lukuesa 941 (K),
from Katanga, is an exceptionally large specimen with ino-
rescence up to 56 cm including 22–34 cm peduncle. It may
represent a distinct taxon, but more material is needed.
6. Chlorophytum arcuatoramosum R.B.Drummond (Drum-
mond 1953: 120). – Type: D.R.Congo, Mutuy, Feb. 1931,
Quarré 2465 (holo-: BR). Fig. 1A & B, g. 4G & H.
Other representative specimens examined – D.R.Congo: V: Ki-
samba, ferme Selemo, Jan. 1931, Quarré 2350 (BR, K); XI: Tenke,
colline Shimbidi, savane steppique sur sol faiblement cuprifère,
Feb. 2010, Meerts 2010/54 (BRLU).
One of the largest Chlorophytum species in tropical Af-
rica. It is obviously related to C. longifolium (perianth pap-
illate) and to C. colubrinum (perianth constricted above
ovary). Seed testa (g. 4G & H) is similar to that of C. longi-
folium, with strongly convex, rugose, periclinal walls and
perforate furrows (though perforations are narrower: com-
386
Pl. Ecol. Evol. 145 (3), 2012
pare with Nordal et al. 1990: 548, g. 6E). Extraordinary
dimensions (1.7–3 m high), inorescence architecture (3–7
branched) and smooth rachis justify species rank.
C. arcuatoramosum forms a well-supported clade with
species that were formely assigned to the genus Dasystachys
Baker (cf. Bjorå 2008).
7. Chlorophytum blepharophyllum Baker (Baker 1876:
327); Baker (1898: 501); Kativu (1994: 53); Troupin (1955:
234); Troupin (1956: 189); Hepper (1968b: 100); Troupin
(1988: 42; 45, g. 15/1); Nordal et al. (1990: 547); Nordal et
al. (1997: 51); Kativu et al. (2008: 74). – Types: Sudan/Ethi-
opian border, Gallabat, around Matamma, 14 Jun. 1865, Sch-
weinfurth 9 (lecto-: K, here designated, isolecto-: P); Zim-
babwe, Lower Gwelo R., 22 May 1870, Baines s.n. (syn-:
BM, n.v.).
The C. blepharophyllum complex appears to be much
more variable in Katanga than in most other regions of tropi-
cal Africa. Kativu et al. (2008) recognized subsp. rubropyg-
maeum (dwarf plants with reddish cataphyll-like leaves) and
subsp. pendulum (pendulous inorescence) and separated C.
amplexicaule (broad leaves clasping peduncle) as a distinct
species. In Katanga, plant size, intensity of purplish colours,
leaf width and inorescence habit are combined in various
ways sometimes within a single population. None of the
aforementioned traits appears to be of high taxonomic value
in the abundant materials we have seen. Pending a phenetic
analysis with a populational sampling, we feel that the best
option at the moment is a very synthetic treatment, recogniz-
ing “amplexicaule” only at varietal rank.
1. Cataphylls prominent with spreading lamina broadly
ovate to cordate, clasping stem; leaves 4–10(–12) cm
broad; plant 15–80 cm high..........................................
.........................C. blepharophyllum var. amplexicaule
1. Cataphylls not prominent, with small lamina not
clasping stem; leaves 0.5–4 cm broad; plant 5–40 cm
high............C. blepharophyllum var. blepharophyllum
7a. Chlorophytum blepharophyllum Baker var. blepharo
phyllum
Chlorophytum brosum Engl. & K.Krause (Engler &
Krause 1910: 132); De Wildeman (1912a: 344). – Types:
Cameroon, between Duka and Dangadji, May 1909, Leder-
mann 3647 (syn-: B); Korrowalplateau, May 1909, Leder-
mann 3890 (syn-: B); near Balda, May 1909, Ledermann
4031 (syn-: B).
Chlorophytum longebracteatum De Wild. (De Wilde-
man 1911a: 275); De Wildeman (1912a: 345). – Type:
D.R.Congo, Vankerckhovenville, Apr. 1906, Seret 539bis
(holo-: BR)
Chlorophytum nigrescens De Wild. (De Wildeman 1911a:
275); De Wildeman (1912a: 346). – Type: D.R.Congo,
Gunbari, 26 Feb. 1906, Seret s.n. (holo-: BR, barcode
BR0000008766328).
Chlorophytum kerstingii Dammer (Dammer 1912: 363);
De Wildeman (1913c: 14; 1921a: 28). – Type: Togo, Sokodé-
Basari, near Aledyo, Feb. 1901, Kersting 314 (holo-: B, not
seen).
? Anthericum ruwense De Wild. (De Wildeman
1913a: 508); De Wildeman (1913c: 9; 1921a: 27). – Type:
D.R.Congo, Katanga, Ruwe, Oct. 1911, Hock s.n. (holo-:
BR, barcode BR0000008769534).
Other representative specimens examined – D.R.Congo: III:
Région de Kisantu, 1907, J. Gillet s.n. (BR); IV. Mabete, savane
guinéenne, 2 Apr. 1955, Devred 1781 (BR); V: 8°S 23°E, groupe-
ment plus ou moins forestier en bordure d’une galerie, 9 Oct. 1957,
Liben 3834 (BR); VII: Parc National de la Garamba, mont Bawe-
si, savane arbustive, alt. 700 m, 11 Mar. 1941, Germain 627 (BR);
VIII: Kerekere (Aru), alt. 1350 m, savane, 4 Apr. 1960, D. Froment
692 (BR); IX: Plaine de la Rusizi, rives Sange, galerie, Jan. 1950,
Germain 5819 (BR); XI: 20 km NE Lubumbashi, forêt claire sur sol
argileux rouge, 23 Nov. 1955, Schmitz 5092 (BR).
Rwanda: Préfect. Kibungo, route Lulama-Gabiro, km 6, savane ar-
borée, 13 Jan. 1972, Bamps 2957 (BR).
Burundi: Territ. Bubanza, Randa, alt. 900 m, savane boisée, 29
Oct. 1968, Lewalle 3061 (BR).
A very variable taxon. Size is more variable than previou-
sly reported, with many collections < 10 cm high and with
leaves < 1 cm wide. Hysteranthous forms are sometimes ob-
served [Reekmans 7091 (BR)]. Plants from Rwanda tend to
have more densely ciliate leaf margin (up to 200 cilia/cm vs.
c. 50 cilia/cm in general).
Bouxin & Radoux 1054 (BR), from Rwanda, with small
(5 mm long), hanging owers probably deserves taxonomic
recognition, but more material is needed.
7b. Chlorophytum blepharophyllum Baker var. amplexi
caule (Baker) Meerts, comb. & stat. nov.
Chlorophytum amplexicaule Baker, Journal of the Lin-
nean Society 15: 325. 1876 (Baker 1876); Baker (1898:
501); P oellnitz (1946: 277). – Type: Tanzania, near Lake
Tanganyika, Feb. 1875, Cameron s.n. (holo-: K, barcode
K000256904). Fig. 1C.
Chlorophytum homblei De Wild. (De Wildeman 1913a:
514); De Wildeman (1913c: 13, Plate III); De Wildeman
(1921a: 28; 1930: 95). – Type: D.R.Congo, Lubumbashi,
Feb. 1912, Homblé 153bis (holo-: BR), synon. nov.
Other representative specimens examined – D.R.Congo: VII:
Près de la rivière Ndondo, savane à Lophira, 9 Jul. 1955, Boutique
201 (BR). IX: Plaine de la Rusizi, Feb. 1950, Germain 266 (BR);
XI: Kiswishi, monastère des sept sources, forêt claire à Marquesia,
Jan. 2010, Meerts 2010/14 (BRLU).
Var. amplexicaule occurs mostly in miombo woodland
and derived savannas in Katanga, Tanzania and Zambia.
We have maintained var. amplexicaule with some scep-
ticism, as intermediates exist with var. blepharophyllum in
Upper Katanga. It also appears to be extremely variable in
marginal cilia density (cilia sometimes almost lacking) and
extent of reddish tinge; the most vigorous forms occasionally
have a pendulous inorescence [Meerts 2010/36 (BRLU);
Duvigneaud 4432L (BRLU)]. It may eventually turn out to
be an aggregate of convergent forms of C. blepharophyllum
having little in common beyond extreme robustness.
Hysteranthous forms with deep purple leaves are occa-
sionally observed [Meerts & Muding 58 (BRLU); Schmitz
4523 (BR)] (g. 1C); such forms approach subsp. rubropyg-
maeum Bjorå & Nordal, but intermediates exist with normal
forms.
387
Meerts & Bjorå, Chlorophytum in Central Africa
In preliminary phylogenetic analyses this complex has
strong support as a clade, but little internal structure. Both
the Congolese [Meerts 2010/36 (BRLU)] and the Zambian
[Hoell & Nordal 134 (O)] specimens of var. amplexicaule
cluster within the clade, as do purple-leaved ones [Meerts &
Muding 58 (BRLU); Nordal & Bjorå 4578 (O)].
8. Chlorophytum cf. brachystachyum Baker (Baker 1893:
710); Baker (1898: 502); Kativu (1994: 54); Nordal et al.
(1997: 52); Kativu et al. (2008: 77, g. 13.1.14). – Type: Ma-
lawi, Shire Highlands, . at Kew 1893, Buchanan s.n. (holo-:
K, barcode K000401063). Fig. 1D & E; g. 4I & J.
Other representative specimens examined – D.R.Congo: XI: Sur
la piste de la ferme de la Kando, sur termitières, Oct. 1988, Schaijes
4139 (BR); Kiubo, savane arborée sur le rebord des grès de Kiubo
dominant les chutes, 15 Jan. 1960, Duvigneaud 5047 (BRLU); Lu-
bumbashi, Campus de la Kasapa, termitière dans l’arboretum, Nov.
2009, Meerts et al. 77 (BRLU).
Specimens from D.R.Congo are undoubtedly members of
the C. brachystachyum-C. pauper complex (fruit verrucose
to tuberculate, leaf margin ciliate), but they are variable and
their status is critical. They are often much more robust than
both of these species, and the rachis is occasionally papillose
(e.g. Duvigneaud 5047Ch), a trait not mentioned in previous
descriptions of those species. Fruit surface is also variable
in roughness, sometimes within the same population. Bracts
are generally not ciliate. Leaf margin is undulate and has
long cilia (traits of C. brachystachyum) but tubers are lateral
(trait of C. pauper). In preliminary phylogenetic analyses
the Congolese C. cf. brachystachyum [Meerts & Muding 77
(BRLU)] clusters with C. pauper, however, weekly support-
ed. It does not form a monophyletic group with the Zambian
specimen of C. brachystachyum [Bjorå 615 (O)] and further
studies are therefore needed to clarify its taxonomic status.
9. Chlorophytum burundiense Meerts (Meerts 2011: 233).
– Type: Burundi, Territ. Muramvya, sommet du Mont Teza,
prairie d’altitude, 8 Dec. 1972, Baudet 304 (holo-: BR; iso-:
K). Fig. 1F.
Other representative specimens examined – Burundi: Territ. Bu-
ruri, Tora, steppe, alt. 2450 m, 7 Feb. 1971, Lewalle 5131 (BR).
10. Chlorophytum calyptrocarpum (Baker) Kativu (Kativu
& Nordal 1993: 62); Kativu (1994: 55); Kativu et al. (2008:
53, Fig. 13.1.9). – Anthericum calyptrocarpum Baker (Bak-
er 1878: 258); Baker (1898: 480); Poellnitz (1943a: 62). –
Type: Angola, Huilla, between Mumpulla and Lopollo, Dec.
1859, Welwitsch 3786 (holo-: BM, iso-: K).
Other representative specimens examined – D.R.Congo: XI:
Lubumbashi, Mine de l’Etoile, Cu-Co mineralized substrate, 2 Apr.
1990, Tropmetex 50 (BR, MO, K, MPN, WAG, C, IBE, P, PRE).
Almost all collections come from the Lubumbashi area,
most often on Cu-mineralised substrate. All records of C. ca-
lyptrocarpum from Burundi (e.g. Lewalle 1972) are errone-
ous and belong in C. angustissimum.
Congolese collections lack the glands on the peduncle re-
ported in the FZ area.
11. Chlorophytum cameronii (Baker) Kativu (Kativu & Nor-
dal 1993: 62); Nordal & Thulin (1993: 262); Kativu (1994:
55); Nordal (1997: 102); Nordal et al. (1997: 30); Kativu et
al. (2008: 44). – Anthericum cameronii Baker (Baker 1876:
314); Troupin (1988: 31, g. 9/2; 32). – Type: Tanzania, Kig-
oma Dist., S of Kawele, Feb. 1875, Cameron s.n. (holo-: K,
barcode K000257044).
A very variable taxon. Four varieties are here recognized.
1. Base of leaves and cataphylls with purple spots or
stripes...............................C. cameronii var. cameronii
1. Base of leaves and cataphylls unicolourous...............2
2. Lower leaf surface, peduncle and rachis pubescent.....
..............................................C. cameronii var. grantii
2. Leaves, peduncle and rachis glabrous (but leaf margin
occasionally shortly ciliate)........................................3
3. Leaves tightly conduplicate, liform to narrowly
graminiform, 2–3 mm wide, with margin generally
ciliate at base; inorescence 2–5 cm long with
1–5 nodes..........................C. cameronii var. ruziziense
3. Leaves at or folded, not tightly conduplicate, linear,
4–25 mm wide, generally not ciliate; inorescence
(4–) 7–12 cm long with 4–20 nodes............................
......................................C. cameronii var. pterocaulon
11a. Chlorophytum cameronii (Baker) Kativu var. camero
nii (Nordal et al. 1997: 31); Kativu et al. (2008: 44).
Anthericum uyuiense Rendle (Rendle 1895: 415); Baker
(1898: 485); Troupin (1956: 185 & 186, g. 30); Hepper
(1968b: 97). – Type: Tanzania, between Uyui and the coast,
1886, W.E. Taylor s.n. (holo-: BM, barcode BM000911753).
Anthericum congolense De Wild. & T.Durand (De Wilde-
man & T. Durand 1899: 60); De Wildeman & T. Durand
(1901: 242); Durand & Durand (1909: 568); De Wildeman
(1921a: 26; 1930: 94). – Type: D.R.Congo, environs de
Lùbunda (Loubounda), Sep. 1896, Dewèvre 1029 (holo-:
BR).
Anthericum congolense De Wild. & T.Durand var. elon-
gatum De Wild. (De Wildeman 1910b: 265); Troupin (1955:
233). – Type: D.R.Congo, Kasaï, Katola, 8 Apr. 1906, Sapin
s.n. (holo-: BR, barcode BR0000005749973).
Other representative specimens examined – D.R.Congo: IV: Ka-
panga, savane boisée, Nov. 1933, Overlaet 1089 (BR); V: La Pas-
torale, section 2, Oct. 1931, Quarré 2739 (BR); VII: Parc nation.
Garamba, Crête Congo-Nil, forêt claire à Isoberlinia doka, 16 Jun.
1952, Troupin 1246 (BR); XI: Upemba, rive gauche de la Lura
vers Mabwe, alt. 920 m, 27 Oct. 1947, de Witte 03024 (BR).
Rwanda: Territ. Kibungu, colline Muhororo, alt. 1400 m, savane à
bosquets, 26 Nov. 1958, Troupin 8860 (BR).
Burundi: Territ. Cibiteke, savane boisée, alt. 1000 m, 26 May
1967, Lewalle 1964 (BR).
Collections from Rwanda generally have leaves narrower
than in other regions (2–6 mm).
11b. Chlorophytum cameronii (Baker) Kativu var. grantii
(Baker) Nordal (Nordal et al. 1997: 32). – Anthericum grantii
Baker (Baker 1875: 160 & tab. 103b); Baker (1898: 488). –
Type: Mozambique, Mgunda Mkali, Bass Rock, Dec. 1860,
Speke & Grant s.n. (holo-: K, barcode K000257053).
388
Pl. Ecol. Evol. 145 (3), 2012
Figure 1 – Photographs of living plants. A, B, Chlorophytum arcuatoramosum (D.R.Congo, Tenke, Shimbidi, Feb. 2010, Meerts 2010/54);
C, C. blepharophyllum var. amplexicaule, hysteranthous purple-leaved phenotype approaching C. blepharophyllum subsp. rubropygmaeum
(D.R.Congo, Kiswishi, Nov. 2009); D, C. cf. brachystachyum (D.R.Congo, Lubumbashi, Kasapa, termite mound, Nov. 2009); E, C. cf.
brachystachyum in fruit (D.R.Congo, Kipopo, Jan. 2010); F, C. burundiense, cultivated, from Mt. Teza (Burundi).
A B
C D
E F
389
Meerts & Bjorå, Chlorophytum in Central Africa
Anthericum durum Poelln. (Poellnitz 1941: 232). – Type:
D.R.Congo, Katanga, Kapiri, Oct. 1911, Hock s.n. (lec-
to-: BR, barcode BR0000008767035, here designated) &
D.R.Congo, Katanga, Lukoni, 6 Oct. 1911, Hock s.n. (syn-:
BR, barcode BR0000008769350), synon. nov.
Anthericum rigidum De Wild. (De Wildeman 1913a: 507;
1913c: 8); De Wildeman (1921a: 27; 1930: 94); nom. illeg.
non Anthericum rigidum Krause.
A. rigidum De Wild. var. breviscapum De Wild. (De Wil-
deman 1915: 5) nom. illeg. – Type: D.R.Congo, Katanga,
Plateau de la Manika, environs de Katentania, Nov. 1912,
Homblé 754 (holo-: BR).
Other representative specimens examined – D.R.Congo: XI:
Lubumbashi, galerie forestière, Nov. 1927, Quarré 903 (BR); s.l.,
s.d., Quarré 1512 (KIP); s.l., s.d., Quarré 3684 (KIP); Territ. Mit-
waba, Vibijo, alt. 1700 m, savane herbeuse, 27 Mar. 1970, Coget
154 (BR); Tilwizembe, steppe sur pente rocheuse, légèrement
métallifère en Cu et Co, 8 May 1957, Duvigneaud 3077L (BRLU);
env. Fungurume, Kavifwafwaulu, sous forêt claire, 24 Nov. 2008,
Malaisse et al. 428 (BR).
Anthericum durum Poelln., described from Congolese
materials, was erroneously reduced to C. galpinii var. gal-
pinii by Kativu et al. (2008). The latter has a branched ino-
rescence (sometimes a raceme, but with a long aristate lower
bract), and 3-veined tepals. In Anthericum durum, the ino-
rescence is always simple, with a short lower bract and the
tepals have 5 widely spaced veins. Collections from Katanga
are variable in size, pubescence and bract shape. Pubescent
forms of C. cameronii might represent a heterogeneous as-
semblage.
Duvigneaud 3082L (BRLU) from Tilwizembe copper hill
with a forked raceme and scabrid leaves may belong here.
11c. Chlorophytum cameronii (Baker) Kativu var. ptero
caulon (Baker) Nordal (Nordal et al. 1997: 32); Kativu et al.
(2008: 45). – Type: Angola, Pungo Andongo, s.d., Welwitsch
3795 (holo-: BM, iso-: K).
Anthericum korrowalense Engl. & K.Krause (Engler
1910: 125); Troupin (1956: 184). – Type: Cameroon, Kor-
rowal plateau, May 1909, Ledermann 3832 (B, holo-, not
seen).
Anthericum speciosum Rendle (Rendle 1895: 413); Baker
(1898: 486); Lebrun et al. (1948: 35). – Type: Tanzania, be-
tween Zanzibar and Uyui, Feb. 1887, W.E. Taylor s.n. (holo-:
BM, barcode BM000911768).
Other representative specimens examined – D.R.Congo: V:
Kuakidi, alt. 730 m, savane arbustive, 7 Dec. 1957, Lukuesa 464
(BR); VII: Parc Nat. de la Garamba, savane herbeuse sur alluvions
de vallée, 1 Jun. 1951, De Saeger 01170 (BR); X: Makamba, 42 km
S de Bururi, savane herbeuse sur argile rouge, 6 Jan. 1938, Robert
20 (BR); XI: Kilata, zone Sakania, 11 Nov. 1986, D’Hose 34b (BR).
Burundi: Territ. Ruyigi, Kigamba, savane arborée, alt. 1500 m, 2
Jan. 1975, Reekmans 4108 (BR).
In its typical form (peduncle at, rachis undulate, ino-
rescence distichous, pedicels shorter than owers), C. cam-
eronii var. pterocaulon is easily recognized. However, many
collections depart from that description in one or several
traits. Var. pterocaulon is more variable than var. cameronii
and is possibly a paraphyletic aggregate. Some collections
have an inorescence with 1–3 branches [De Witte 3088
(BR) and 5557 (BR) from l’Upemba].
A few collections from Upper Katanga (region of Fungu-
rume) [FC 12 (BR)] have red-coloured cataphylls and leaf
margins. They may well deserve taxonomic recognition, but
more material is needed.
11d. Chlorophytum cameronii (Baker) Kativu var. ruzi
ziense Meerts, var. nov.
A typo differt foliis tenuissimis, conduplicatis, 2–3 mm latis,
13–17 nervatis, marginibus saepe ciliatis, cum vaginibus non
purpureo maculatis. – Type: Burundi, Bubanza, Gihungwe,
alt. 800 m, 23 Nov. 1974, Reekmans 3955 (holo-: BR, iso-:
K).
Other representative specimens examined – D.R.Congo: IX:
Plaine de la Ruzizi, piste Kabunambo-rivière Ruzizi, savane her-
beuse, Jan. 1950, Germain 5517 (BR, FT); Plaine de la Ruzizi, env.
Kihombo, savane à Themeda et Bulbine, Feb. 1950, Germain 6008
(BR, K); Plaine de la Rusizi, Jan. 1929, Humbert 7301 (BR, K, B).
Burundi: Bubanza, Plaine de la Ruzizi, km 30, savane à Bulbine,
alt. 800 m, 25 Nov. 1973, Reekmans 2929 (BR).
This new variety has extremely narrow, conduplicate
leaves 2–3 mm wide, with 13–17 nerves, most often ciliate
in lower third (though not in the membranaceous sheath), not
purple-spotted. It is variable in size, from 20 cm in collec-
tions from high altitude in Burundi to 90 cm in collections
from the Ruzizi plain. This variety is close to C. zanguebari-
cum (Baker) Nordal, another narrow-leaved variant of the C.
cameronii complex, but this does not have tightly condupli-
cate leaves and leaf margin is not ciliate. More research is
needed on the narrow-leaved forms within the C. cameronii
complex.
Germain (1952) referred to this plant as “C. gracillimum
V.D.”, a nomen nudum (see under C. warneckei).
In the preliminary phylogenetic analyses the Congolese
C. cameronii var. ruziziense [Reekmans 3955 (K, BR)] re-
solves as sister to the Ethiopian C. cameronii in a well sup-
ported clade of species that previously were included in the
genus Anthericum.
12. Chlorophytum clarae Bjorå & Nordal (Bjorå et al. 2008:
232, g. 5; 233 gs 7–9; 236 g. 21). – Type: Zambia, North-
ern region, Mansa District, 4 Dec. 2002, Nordal & Bjorå
4542 (holo-: O, not seen). Fig. 2A–C; g. 5A & B.
Representative specimens examined – D.R.Congo: V: Kiala,
Feb. 1954, Thiébaud 383 (BR); XI: Kiswishi, monastère ND des
sept sources, base de termitière en forêt claire, Jan. 2010, Meerts
2010/23 (BRLU); rivière Mashinji, 10°23’S 25°17’E, forêt galerie,
19 Feb. 1987, Billiet & Jadin 4191 [BR, with photographs of plant
cultivated in greenhouse (S 1975; 87-0156)]; Tenke, fossé ombragé
dans le dembo d’empoisonnement de la colline cuprifère de Ka-
bwelunono, Feb. 2010, Meerts 2010/37 (BRLU); Galerie de la Lo-
foï, Jul. 1939, Quarré 5724 (BR).
This taxon has long been confounded with C. macrophyl-
lum (see discussion under that species). It is clearly distinct
by its seeds, variable in shape, from more or less saucer-
shaped to very irregularly angulose-plicate (at to slightly
saucer-shaped in C. macrophyllum), and by its seed coat with
raised periclinal walls and wide furrows (shallowly convex
periclinal walls and narrow furrows in C. macrophyllum,
390
Pl. Ecol. Evol. 145 (3), 2012
g. 5C–F). Other differences include lateral tubers (terminal
in C. macrophyllum), more or less rounded fruits (strongly
trigonous in C. macrophyllum), and a denser raceme. There-
fore, collections without mature fruits or well-collected be-
lowground organs cannot be identied. Other characters
are variable, including anther length (3–5 mm), tepal length
(7.5–12 mm), leaf width (3–8 cm), raceme length (4–25 cm)
and fruit surface (smooth to slightly rugose-verrucose). Col-
lections from Katanga have a more variable seed shape than
reported for Zambia (Bjorå et al. 2008).
It is still unclear whether genuine C. macrophyllum oc-
curs in Katanga.
13. Chlorophytum colubrinum (Baker) Engl. (Engler 1892:
162); Poellnitz (1946: 353); Kativu (1994: 56); Nordal et al.
(1997: 19); Kativu et al. (2008: 66, g. 13.1.12). – Dasys-
tachys colubrina Baker (Baker 1878a: 256 & tab. 35, gs
5–10); De Wildeman (1913c: 12; 1921a: 28; 1930: 96). –
Type: Angola, Huilla, Empalanca, Jan. 1860, Welwitsch 3784
(lecto-: BM; isolecto-: K, B, P).
Two varieties can be recognized.
1. Raceme cylindrical, 4–30 cm long, 4–20 times as
long as wide; leaves 3–40 mm wide; bracts not or
only shortly ciliate; cataphylls glabrous.....................
....................................C. colubrinum var. colubrinum
1. Raceme subglobose to ovoid, 1.5–4 cm long, less
than 2 times as long as wide; leaves 2–4 mm wide;
bracts long ciliate; cataphylls pubescent......................
.....................................C. colubrinum var. upembense
13a. Chlorophytum colubrinum (Baker) Engl. var. colubri
num
Dasystachys verdickii De Wild. (De Wildeman 1902:
10); Durand & Durand (1909: 569); De Wildeman (1921a:
29; 1930: 96) – Chlorophytum verdickii (De Wild.) Poelln.
(P oellnitz 1945: 232) – Type: D.R.Congo, Katanga, Lofoi,
1899, Verdick s.n. (holo-: BR, barcode BR0000008768872).
Dasystachys hockii De Wild. (De Wildeman 1911a: 264);
De Wildeman (1913c: 11; 1921a: 29). – Type: D.R.Congo,
Katanga, Vallée de la Luembe, Feb. 1910, Hock s.n. (holo-:
BR, barcode BR0000008764348).
Dasystachys bequaertii De Wild. (De Wildeman 1913b:
294); De Wildeman (1921a: 28). – Type: D.R.Congo, Katan-
ga, Elisabethville [Lubumbashi], 15 Apr. 1912, Bequaert 333
[published as 533] (holo-: BR).
Chlorophytum subpapillosum Poelln. (Poellnitz 1945:
232). – Type: D.R.Congo, Lubembe, 27 Jan. 1908, Kassner
2394 (holo-: B, iso-: BR).
Chlorophytum poggei Engler. ex Poelln. (Poellnitz 1945:
232). – Type: D.R.Congo, Mukenge, 27 Apr. 1882, Pogge
1491 (holo-: B).
Dasystachys pulchella P.A.Duvign. & Dewit (Duvi-
gneaud & Denaeyer-De Smet 1963: 149, g. 11). – Chlo-
rophytum linearifolium Marais & Reilly (Marais & Reilly
1978: 660). – Type: D.R.Congo, Kolwezi, Dikuluwe, 21 Jan.
1960, Duvigneaud 5141D (holo-: BRLU).
Chlorophytum decoratum (Baker) Marais & Reilly
(Marais & Reilly 1978: 659); Troupin (1988: 44). –Type:
Zambia, Fwambo, Sep. 1893, Carson 26 (holo-: K).
Other representative specimens examined – D.R.Congo: III:
Route de Kenge, Vallée de la Black River, 30 Jul. 1974, Pauwels
5142 (BR); IV: Mayi a Kantshia, Kazumba; savane, Oct. 1978,
Mabika 0168 (BR); V: Gandazjika (Kasaï), savane, 24 Feb. 1953,
Risopoulos 37 (BR); XI: Katanga, Plateau de la Manika, riv. Po-
topoto, savane steppique, 24 Dec. 1988, Schaijes 4232; Mine de
Luiswishi, 14 Feb. 1982, Malaisse & Robbrecht 2118 (BR).
Rwanda: Rusumo, route vers Nyarubuye, savane arbustive sur sol
rocailleux, 25 Jan. 1998, Bridson 306 (BR).
Burundi: Prov. Bururi, Gitwe (Rumonge), forêt claire à Brachyste-
gia, dans un petit ravin, 24 Feb. 1980, Reekmans 8622 (BR).
The species is extremely variable in the size of all its
parts. Part of the variation is correlated to habitat and seems
to be genetic. The most vigorous forms with a very dense ra-
ceme, long tepals (up to 11 mm), big fruits (up to 10 mm) and
a compact infrutescence 25–40 mm wide [Dasystachys ver-
dickii De Wild.; Chlorophytum poggei Engl. ex Poelln.; e.g.
Quarré 8045 (BR)] are striking. However, they are linked to
normal forms by a range of intermediates. At the opposite
extreme of the variation range, slender few-owered forms
from copper-rich soil in Upper Katanga were described as a
distinct species (Dasystachys pulchella P.A.Duvign. & De-
wit; Chlorophytum linearifolium Marais & Reilly) [Cu-soil:
e.g. Malaisse & Robbrecht 2398 (BR, WAG, MO, K, P), Du-
vigneaud 4670 (BRLU); non mineralized soil: Duvigneaud
5292D (BRLU)]. However, intermediates exist between
those and more robust forms even on copper-rich soil and it
does not seem possible to maintain that taxon even at varietal
rank (Faucon et al. 2010).
Some collections from Katanga have ovate-elliptic leaves
with length/width ratio unusually low [50–100 × 20–30 mm:
de Witte 65 (BR); Malaisse 4907 (BR)]. Small forms with
hysteranthous owers sometimes occur [Quarré 1990 (BR)].
A variant with a ramied inorescence is rarely encountered
[e.g. env. Kibunda, forêt sèche, 28 Apr. 1953, Callens 4074
(K)] corresponding to C. pleiostachyum (Baker) T.Durand &
Schinz, but this does not seem to warrant taxonomic recogni-
tion.
Many collections have purple-spotted leaf sheaths but the
taxonomic value of this trait is unclear.
13b. Chlorophytum colubrinum (Baker) Engl. var. upem
bense Meerts, var. nov.
A typo differt racemo sugloboso aut ovoideo, 15–30 mm
longo, cataphyllis pubescentibus et bracteis longe ciliatis. –
Type: D.R.Congo, Haut-Katanga, Parc national de l’Upem-
ba, Buyé-Bala, af. Muyé, savane marécageuse, alt. 1750 m,
30 Mar. 1948, de Witte 03596 (holo-: BR).
Other representative specimen examined – D.R.Congo: XI:
Parc nation. Upemba, savane herbeuse d’altitude autour de la tête
de source de la riv. Kalumengongo, alt. 1830 m, 31 Mar. 1949, de
Witte 05970 (BR).
This variety is well-characterized by a subcapitate to
ovoid inorescence, long ciliate bracts and hairy cataphylls.
It has very narrow leaves (2–4 mm), but this trait is occa-
sionally found in var. colubrinum, especialy in populations
on copper-rich soil.
391
Meerts & Bjorå, Chlorophytum in Central Africa
14. Chlorophytum cordifolium De Wild. (De Wildeman
1914: 111); De Wildeman (1921a: 27). – Type: D.R.Congo,
Katanga, env. Katentania (Plateau de la Manika, Biano), Nov.
1912, Homblé 766 (holo-: BR, barcode BR0000006420581;
iso-: BR, barcode BR0000006420314).
Chlorophytum unifolium Malaisse & Bamps (Malaisse
& Bamps 2009: 225-229, gs 1 & 2). – Type: D.R.Congo,
Katanga, Kavifwafwaulu, Nov. 2006, Malaisse, Kisimba &
Saad 598 (holo-: BR), synon. nov.
Other representative specimens examined – D.R.Congo: XI. Bi-
ano Hôtel, Forêt claire à Brachystegia et Monotes sur sol profond et
léger, 8 Dec. 1959, Duvigneaud 4459L (BRLU).
A remarkable, very distinct species collected only on
few occasions since its description in 1914. Rediscovered in
2006 it was then erroneously described as a new species. Ap-
parently a narrow endemic of the Tenke-Biano area.
15. Chlorophytum debile Baker (Baker 1878a: 260). – Type:
Angola, Ambriz, ad rupes, Welwitsch 3769 (holo-: BM; iso-:
K, P). Fig. 5G & H.
Chlorophytum gilletii Compère (Compère 1962: 211). –
Chlorophytum gracile De Wild. (De Wildeman 1911a: 274);
De Wildeman (1912a: 344) nom. illeg. – Type: D.R.Congo,
Bas-Congo, environs de Kisantu, 1906, Gillet 3963 (holo-:
BR), synon. nov.
Chlorophytum laxum auct. W Afr. (e.g. Baker 1898: 503;
Hepper 1968b: 100; Vanden Berghen 1988: 425, g. 349;
426) non Chlorophytum laxum R.Br.
Other representative specimen examined – D.R.Congo: III. Ter-
rit. Songolo, Kiandu, galerie forestière de la Lunionzo, 16 Dec.
1959, Compère 1049 (BR, K).
This species can be confounded with slender forms of
C. sparsiorum. The seed testa however is very different,
with smooth periclinal walls in C. sparsiorum and apiculate
periclinal walls in C. debile (g. 5G & H). The species is
very variable in size (6–40 cm high, with inorescence more
or less prostrate in the smallest forms). The type specimen
originates from NW Angola, i.e. not far from Congolese col-
lections. Specimens from D.R.Congo and the type of C. de-
bile perfectly match collections from W Africa to which the
name “C. laxum” has long been wrongly applied based upon
a type specimen from Australia (Poulsen & Nordal 2005). In
our opinion, C. debile may be the correct name for most if
not all collections of C. laxum auct. from W Tropical Africa.
Xylem vessel anatomy also indicated that Asian and African
collections of “C. laxum auct.” are not conspecic (Naik &
Nirgude 1981).
Donis 177 (BR), cited in Compère (1962), has blacken-
ing leaves, an unusual trait for C. debile. As it lacks owers
and fruits, identication is uncertain.
16. Chlorophytum lipendulum Baker (Baker 1878a: 260);
Baker (1898: 499); Rendle (1899: 54); Nordal et al. (1997:
59); Poulsen & Nordal (2005: 16, gs. 12 & 13); Bjorå &
Nordal (2010: 4). – Type: Angola, Golungo Alto, s.d., Wel-
witsch 3776 (holo-: BM, iso-: K, P).
Chlorophytum macrophyllum sensu Robyns & Tournay
(1955: 354) non C. macrophyllum (A.Rich.) Aschers.
Other representative specimens examined – D.R.Congo: IV:
Kamembele, galerie forestière, 12 Jan. 1957, Liben 2257 (BR);
V: Kamalenge, galerie forestière, 10 Feb. 1948, Mullenders 2039
(BR); VI: Yangambi, km 13 route N’gazi, sous-bois forêt primi-
tive ombrophile, 12 Jan. 1938, Louis 7466 (BR, K); VII: Mapere-
Suronga, forêt, 20 Dec. 1905, Seret 399 (BR); VIII: Kima Kima
(Mahagi), alt. 1700 m, galerie forestière, 27 May 1959, D. Froment
459 (BR); IX: Mutsora, galerie forestière de la Ngokoye, endroit
fort ombragé, 2 Aug. 1952, de Witte 7816 (BR).
C. lipendulum has been much underrecorded until now
due to confusion with C. macrophyllum (see discussion un-
der the latter species).
C. lipendulum is closely related to and sometimes dif-
cult to discriminate from C. sparsiorum. In particular, small
forms with incompletely blackening leaves are puzzling. In-
trogression might explain the existence of more or less inter-
mediate forms.
17. Chlorophytum gallabatense Schweinf. ex Baker (Baker
1876: 325); Baker (1898: 504); Troupin (1955: 235); Troupin
(1956: 190); Hepper (1968b: 99); Troupin (1988: 43, g. 4/2;
44); Kativu (1994: 60); Nordal et al. (1997: 45); Kativu et
al. (2008: 81). – Type: Sudan/Ethiopian border, Gallabat,
Matamma, Jul. 1865, Schweinfurth 10 (lecto-: K; isolecto-:
B, P).
Three varieties, treated as distinct species in FZ and
FTEA.
1. Rachis and pedicels densely covered with hairs or
cylindric papillae 0.3–0.5 mm long; lower leaf surface
often papillate on veins; fruits c. 7 mm long, generally
longer than broad.....C. gallabatense var. oribundum
1. Rachis and pedicels glabrous or scabrid with conical
papillae < 0.3 mm long; lower leaf surface glabrous;
fruits 4–6 mm long, no longer than broad..................2
2. Leaves developing after inorescence, linear, < 1 cm
wide; inorescence: a raceme or a poorly ramied
panicle with 1–2 branches; rootstock with copious
brous remains of leaves; tubers 15–20 mm long,
mostly terminal.........C. gallabatense var. micranthum
2. Leaves well developed at owering, narrowly elliptic
to almost linear, attenuate towards base, 12–40 mm
wide; inorescence with 3–15 branches; rootstock
most often without bers; tubers < 12 mm, mostly on
lateral rootlets..........C. gallabatense var. gallabatense
17a. Chlorophytum gallabatense Schweinf. ex Baker var.
gallabatense. Fig. 5I & J.
Chlorophytum psammophilum Engl. & Gilg (Engler &
Gilg 1903: 188). – Type: Angola, between Kiubango and
Kuito, alt. 1200 m, 9 Dec. 1899, Baum 517 (holo-: B; iso-: Z,
K), synon. nov.
Chlorophytum ramulosum De Wild. (De Wildeman
1911a: 275); De Wildeman (1912a: 346; 1921b: 19). – Type:
D.R.Congo, route Bambili-Amadi, 4 Mar. 1905, Seret 244
(holo-: BR).
Chlorophytum katangense De Wild. (De Wildeman
1911a: 277); De Wildeman (1913c: 14; 1921a: 28; 1930:
95). – Type: D.R.Congo, Katanga, Vallée de la Luembe, Feb.
392
Pl. Ecol. Evol. 145 (3), 2012
Figure 2 – Photographs of living plants (photos P. Meerts except otherwise stated). A, C. clarae, in ower (D.R.Congo, Kiswishi, termite
mound in miombo, Nov. 2009); B, C. clarae, cultivated in greenhouse (D.R.Congo, from Billiet & Jadin 4191; photo F. Billiet); C, C. clarae,
in fruit (D.R.Congo, Fungurume, Feb. 2010); D, C. vestitum subsp. pilosissimum (D.R.Congo, Kipopo, Jan. 2010); E, C. hysteranthum
(D.R.Congo, Kipopo, Nov. 2009; Meerts & Muding 73); F, C. rubribracteatum (D.R.Congo, Kiswishi, Nov. 2009).
A B
C D
EF
393
Meerts & Bjorå, Chlorophytum in Central Africa
1910, Hock s.n. (holo-: BR, barcode BR0000008766618),
synon. nov.
Chlorophytum breviorum De Wild. (De Wildeman
1913a: 513; 1913c: 12); De Wildeman (1921a: 27; 1930: 95).
– Type: D.R.Congo, Lubumbashi, Feb. 1912, Homblé 153
(holo-: BR).
Chlorophytum hockii De Wild. (De Wildeman 1913a:
514); De Wildeman (1913c: 13; 1921a: 27). – Type:
D.R.Congo, Katanga, Lubumbashi, Oct. 1911, Hock s.n.
(holo-: BR, barcode BR0000008768124).
Other representative specimens examined – D.R.Congo: IV:
Topo-Guilu-Kwango, savane guinéenne, 28 Mar. 1955, Devred
1713 (BR); V: Kiala, savane ensoleillée, Dec. 1954, Thiébaud 397
(BR); VII: Faradja, Kibale-Ituri, savane arbustive, Jul. 1931, Leb-
run 3409 (BR); VIII: Kasengi, Nov. 1935, Brédo 1893 (BR); XI:
Plateau de la Manika, Biano, env. Katentania, termitière, Nov. 1912,
Homblé 760 (BR); Kimilolo, 10 Feb. 1927, Quarré 128 (BR); Piste
Kolwezi-Musokantanda, 0.35 km avant la piste du détournement,
10°45’39”S 25°20’59”E, 1459 m, sur haute termitière en zone dé-
boisée près d’un village, 7 Dec. 1986, Schaijes 3228 (BR, MO).
Var. gallabatense is variable. It is often said to have gla-
brous rachis and pedicels (e.g. Kativu et al. 2008: 39, 82),
thus contrasting with the papillate var. oribundum. This is
not true, as many specimens of var. gallabatense have a pap-
illate rachis too (e.g. C. katangense De Wild.), but the size
and shape of papillae is different (see key) and the distal part
of pedicels is usually smooth in var. gallabatense. In FZ, var.
gallabatense is said to have long anthers (4 mm), longer than
laments, but materials from D.R.Congo have variable an-
ther size (1–3.5 mm), often shorter than laments. Articula-
tion of pedicel is also variable, from lower half to upper half.
FZ recognizes Chlorophytum psammophilum Engl. &
Gilg, said to have narrower leaves, shorter pedicels and te-
pals, stamens longer than perianth (vs. shorter in C. galla-
batense) and white petals (vs. greenish in C. gallabatense).
Some collections from Katanga resemble the type specimen
of that taxon [e.g. Duvigneaud 4455Ch (BRLU)]. However,
we have not been able to nd reliable diagnostic characters
and C. psammophilum is here reduced to C. gallabatense var.
gallabatense.
17b. Chlorophytum gallabatense Schweinf. ex Baker var.
oribundum (Baker) Meerts, comb. & stat. nov.
Chlorophytum oribundum Baker, Bulletin of Miscellaneous
Information, Kew 1897: 285. 1897 (Baker 1897b); Baker
(1898: 505); Kativu (1994: 60); Nordal et al. (1997: 47); Ka-
tivu et al. (2008: 82). – Type: Malawi, Zomba, Dec. 1896,
Whyte s.n. (holo-: K, barcode K000256938, iso-: B, barcode
B100168008).
Other representative specimen examined – D.R.Congo: XI:
Vallée de la Lofoi, Lukafu, sol forestier peu ombragé d’éboulis de
pente, 17 Oct. 1948, Schmitz 2082 (BR).
Symoens 12825j (BRVU), from Lubumbashi, may well
belong here but the plant is very young. Var. oribundum has
never been collected in the fruiting state in D.R.Congo and
the character of fruit length mentioned in the key is based on
material from Zambia.
17c. Chlorophytum gallabatense Schweinf. ex Baker var.
micranthum (Baker) Meerts, comb. & stat. nov.
Chlorophytum micranthum Baker, Journal of Botany, British
and foreign 16: 325. 1878 (Baker 1878b); Baker (1898: 507);
Troupin (1956: 191); Nordal & Thulin (1993: 269). – Type:
Sudan, Jur, s.d., Schweinfurth 1745 (holo-: K, iso-: B).
Chlorophytum bequaertii De Wild. (De Wildeman 1921b:
14); Robyns & Tournay (1955: 356, Plate XLIX); Hepper
(1968b: 102). – Type: D.R.Congo, Kabare, 22 Aug. 1914,
Bequaert 5397 (lecto-: BR, isolecto-: K, here designated) &
Bequaert 5410 [syn-: BR (3 sheets)], synon. nov.
Other representative specimens examined – D.R.Congo: VII:
Kurukwata (Aba), savane, endroit pierreux, 20 Feb. 1957, Gérard
3100 (BR); VIII: Parc national de la Garamba, crête Congo-Nil, km
10, forêt à Isoberlinia, 4 Mar. 1952, Troupin 286 (BR); IX: env.
d’Uvira, Plaine de la Rusizi, alt. 800 m, Jan. 1929, Humbert 7307
(BR, P).
Burundi: Territ. Bubanza, Gihungwe, steppe à Bulbine, alt. 800 m,
30 Nov. 1974, Reekmans 3954 (BR, K).
Rwanda: Parc Nation. Kagera, Runyonza, savane à Acacia et Aloe
brûlée, 20 Aug. 1948, Robyns 3434 (BR).
All collections from Rwanda previously referred to as
“C. gallabatense” (e.g. in Troupin 1988) belong here.
This taxon was treated at species rank by FTEA, but it
was indicated to “be a small and hysteranthous (re adapt-
ed?) form, not deserving more than subspecic rank”, and
this view is supported here. Tepal length, often used to dis-
criminate C. micranthum and C. gallabatense, shows broad
overlap between the two taxa. Var. micranthum often has yel-
lowish owers, but this is not a constant character. Collec-
tions from the Rusizi plain show fully developed leaves, c.
9 mm wide and 20–35 cm long (fully developed leaves not
observed in FTEA).
C. bequaertii was reduced to C. micranthum by Marais &
Reilly (1978) and to C. gallabatense by FZ. Narrow leaves,
sparingly branched inorescence, copious leaf remains and
terminal tubers suggest it belongs here.
The taxonomic signicance of tuber position needs fur-
ther investigation, as tubers are rarely collected.
18. Chlorophytum galpinii (Baker) Kativu var. matabelense
(Baker) Kativu (Kativu & Nordal 1993: 63); Kativu (1994:
6); Kativu et al. (2008: 47, g. 13.1.7). – Anthericum mata-
belense Baker (Baker 1898: 484). – Type: Zimbabwe, Bulila-
mangwe Dist., on banks of Matengwe R., May 1883, Holub
s.n. (holo-: K, barcode K000256914).
Anthericum kapiriense De Wild. (De Wildeman 1915:
4); De Wildeman (1921a: 26). – Type: D.R.Congo, Katanga,
Vallée Kapiri, Feb. 1913, Homblé 1092 (holo-: BR).
Other representative specimens examined – D.R.Congo: XI:
Plateau de la Manika, savane steppique au nord de la tête de source
de la rivière Luilu, 1500 m, 8 Feb. 1987, Schaijes 3322 (BR); Piste
Nzilo-Kyamasumba à 10.5 km par la piste du centre urbain de Nzi-
lo, forêt claire, 01 Aug. 1987, Schaijes 3575 (BR).
C. galpinii var. galpinii was reported by error from
D.R.Congo by FZ (Kativu et al. 2008) (see discussion under
C. cameronii var. grantii).
19. Chlorophytum geophilum Peter ex Poelln. (Poellnitz
1943b: 127; 1946: 292); Hepper (1968b: 100); Kativu (1994:
394
Pl. Ecol. Evol. 145 (3), 2012
Figure 3 – Photographs of living plants (photos P. Meerts except otherwise stated). A, C. pusillum (cultivated in greenhouse, from Billiet &
Jadin 4151; photo F. Billiet); B, C. sphacelatum (D.R.Congo, Kiswishi; photo J. Piqueray); C, C. stenopetalum var. latifolium (D.R.Congo,
Kiswishi; photo J. Piqueray); D, C. subpetiolatum var. pilosifolium (D.R.Congo, Lubumbashi, Kasapa, termite mound, Nov. 2009); E, C.
velutinum (D.R.Congo, Fungurume, copper hill, Feb. 2010); F, C. zingiberastrum (cultivated in greenhouse, from Billiet & Jadin 4195; photo
F. Billiet).
A B
C
D
E
F
395
Meerts & Bjorå, Chlorophytum in Central Africa
63); Nordal et al. (1997: 54, g. 11); Kativu et al. (2008: 87,
g. 13.1.15). – Type: Tanzania, Tabora Dist., Unyamwesi,
Kombe, 24 Jan. 1926, Peter 35409 (holo-: B). Fig. 6A.
Chlorophytum pusillum sensu Troupin (Troupin 1956:
191).
Other representative specimens examined – D.R.Congo: VII:
Parc Nation. Garamba, savane arbustive, 5 Jun. 1950, Noirfalise
441 (BR). XI: Bunkeya, plaine, Dec. 1948, Hoffmann 889 (BR).
C. geophilum and C. pusillum are two closely related spe-
cies and intermediate specimens exist. The collections from
the Parc National de la Garamba are critical.
20. Chlorophytum hirsutum Poulsen & Nordal (Poulsen
& Nordal 1999: 941). – Type: Uganda, Bundibugyo Distr.,
Bwamba County, near Sempaya, 22 Jul. 1938, Thomas 2311
(holo-: K; iso-: BR).
Other representative specimens examined – D.R.Congo: IX:
Territ. Uvira, Lubarika, savane à Loudetia, 29 May 1959, A. Léo-
nard 4463 (BR).
Burundi: Territ. Bubanza, alt. 1100 m, bord de forêt, Dec. 1967,
Lewalle 2549 (BR).
21. Chlorophytum hysteranthum Kativu (Kativu & Nor-
dal 1993: 63); Kativu (1994: 64); Kativu et al. (2008: 49).
– Anthericum breviscapum De Wild. (De Wildeman 1913a:
507); De Wildeman (1913c: 7; 1921a: 26), non Chlorophy-
tum breviscapum Dalzell or Chlorophytum breviscapum
Dammer. – Type: D.R.Congo, Katanga, Lubumbashi, Sep.
1911, Hock s.n. (holo-: BR, barcode BR0000008222695).
Fig. 1K.
Other representative specimens examined – D.R.Congo: XI: Ki-
popo, 25 km NO Lubumbashi, dembo d’argile grise alternativement
marécageuse et très sèche, 28 Sep. 1952, Schmitz 4093 (BR); ibid.,
Nov. 2009, Meerts & Muding 73 (BRLU).
A rare species with a restricted distribution range in S Ka-
tanga and N Zambia. The few collections from D.R.Congo
all come from a restricted area in the vicinity of Lubumbashi.
Rogers 10076 (K) has unusually large leaves up to 20 cm
long suggesting that C. hysteranthum is closely related to C.
afne.
22. Chlorophytum lancifolium Baker (Baker 1878a: 260);
Baker (1898: 498); Kativu (1994: 65); Nordal et al. (1997:
60); Poulsen & Nordal (2005: 16-17); Kativu et al. (2008:
83). – Type: Angola, Pungo Andongo, Nov. 1856, Welwitsch
3772 (lecto-: BM, isolecto-: K); Welwitsch 3773 (syn-: BM,
K).
1. Seed testa apiculate....C. lancifolium subsp. cordatum
1. Seed testa not apiculate..............................................2
2. Leaf base most often attenuate, owers greenish to
yellowish....................C. lancifolium subsp. togoense
2. Leaf base most often truncate to cordate, owers
whitish.....................C. lancifolium subsp. lancifolium
22a. Chlorophytum lancifolium Baker subsp. lancifolium
[Poulsen & Nordal (2005: 17, gs 1J; 18, 19, 35)].
Other representative specimens examined – D.R.Congo: XI:
Sakania, forêt claire mélangée sur terre rouge, 31 Jan. 1960, Du-
vigneaud 5357 (BRLU).
Burundi: Ruyigi, colline Nyabitangu, sous-bois de forêt à Julber-
nardia, 8 Feb. 1979, Reekmans 7661 (BR, K).
Subsp. lancifolium has been much less collected than the
other two subspecies.
22b. Chlorophytum lancifolium Baker subsp. cordatum
(Engl.) Poulsen & Nordal (Poulsen & Nordal 2005: 16, g.
1K; 16, 17, 34). – Chlorophytum cordatum Engl. (Baker &
Engler 1892: 468). – Type: Soudan, Jebel Bangenze, Ni-
amniamland, am Baginse, 25 May 1870, Schweinfurth 173
(holo-: B).
Other representative specimens examined – D.R.Congo: VII:
Montagne Kundungu, 560 m, village Masabe, à 40 km au N de
Bondo, forêt dense primaire sur rocher granitique, 3 Jul. 1978, Le-
joly 4118 (BRLU, BR, K, WAG); Parc nation. Garamba, Dedegwa,
galerie à la tête de source de la Dedegwa, 17 May 1952, De Saeger
2606 (BR, K); VIII: Bunia, Mont Hoyo, ravin, sur rochers humides,
5 Aug. 1976, Pauwels 5751 (BR); XI: Route Kolwezi-Kyamasum-
ba, rivière Mushingi, 19 Feb. 1987, Billiet & Jadin 4192 (BR; culti-
vated in greenhouse: S 1945).
Subsp. cordatum mostly occurs in NE Congo (Poulsen
& Nordal 2005); however, the record from Upper-Katanga
[Billiet & Jadin 4192 (BR)] (seed testa checked with SEM)
is widely disjunct from the main area of the subspecies indi-
cating that subsp. lancifolium and subsp. cordatum are not
completely allopatric.
22c. Chlorophytum lancifolium Baker subsp. togoense
(Engl.) Poulsen & Nordal (Poulsen & Nordal 2005: 17, g.
1L; 20, 21, 36). – Chlorophytum togoense Engl. (Engler
1902: 92); Hepper (1968b: 99). – Type: Togo, Jaggebach, bei
Misa Höhe, 5 Sep. 1890, Büttner 172 (lecto-: B, here desig-
nated); Togo, Fasugu, 30 May 1891, Büttner 665 (syn-: B).
Chlorophytum ealaense De Wild. (De Wildeman 1911a:
274); De Wildeman (1912a: 343; 1916: 164; 1921b: 15). –
Type: D.R.Congo, Eala, Oct. 1907, Pynaert 1040 (lecto-:
BR; isolecto-: B); D.R.Congo, Eala, s.d., Pynaert 557 (syn-:
BR); D.R.Congo, Eala, s.d., Pynaert 1777 (syn-: BR).
Other representative specimens examined – D.R.Congo: VI:
Prov. Equateur, près de la rivière Ikilemba, 26 Aug. 1914, Nannan
63 (BR, K).
Subsp. togoense has a western Guineo-Congolian distri-
bution, having its eastern limit in W D.R.Congo in the region
of Eala. Some collections [(e.g. Gombe (Equateur), Aug.
1903, M. Laurent s.n. (BR, barcode BR0000005720651)]
are more or less intermediate between C. lancifolium and C.
sparsiorum, and might be of hybrid origin.
C. lancifolium is often said to have a scabrid leaf margin
(Nordal et al. 1997, Nordal & Poulsen 2005), but in most
collections examined leaf margin is smooth.
23. Chlorophytum leptoneurum C.H.Wright var. katan
gense (De Wild.) Meerts, comb. & stat. nov.
Verdickia katangensis De Wild. Annales du Musée du Congo,
Botanique, série 4,1: 7. 1902 (De Wildeman 1902); Engler
(1908: 309); Durand & Durand (1909: 569); De Wildeman
(1921a: 27). – Type: D.R.Congo, Katanga, Lukafu, terrains
396
Pl. Ecol. Evol. 145 (3), 2012
Figure 4 – Scanning electron
microscope images of seeds and
seed testa. A & B, Chlorophytum
africanum var. nordalianum, seed
and seed testa (Schmitz 4261); C,
C. andongense (short-fruited form),
seed testa (Schmitz 4545); D, C.
andongense (long-fruited form),
seed testa (Billiet & Jadin 4150);
E & F, C. angustissimum, seed and
seed testa (Lewalle 6452); G & H,
C. arcuatoramosum, seed and seed
testa (Mullenders 142); I & J, C. cf.
brachystachyum, seed and seed testa
(Quarré 1950).
A B
C D
E F
G H
I J
397
Meerts & Bjorå, Chlorophytum in Central Africa
Figure 5 – Scanning electron
microscope images of seeds and
seed testa. A & B, C. clarae, seed
and seed testa (Meerts 2010/37). C
& D, C. macrophyllum, seed and
seed testa (Schimper 1837); E & F,
C. macrophyllum, seed (Allard 370)
and seed testa (Vanderyst 38806);
G & H, C. debile, seed and seed
testa (Compère 1049); I & J, C.
gallabatense var. galabatense, seed
and seed testa (Homblé 1150).
A B
C D
E F
GH
I J
398
Pl. Ecol. Evol. 145 (3), 2012
argileux, Dec. 1899, Verdick 329 (holo-: BR); non Chloro-
phytum katangense De Wild. Fig. 6C, D & I.
Other representative specimens examined – D.R.Congo: XI:
Keyberg, Katuba, prairie sur sol superciel à dalle latéritique, 28
Nov. 1947, Schmitz 1052 (BR); pâturage articiel du dembo de la
Katuba, Jan. 1934, Quarré 3741 (BR).
Other specimens examined – Malawi: N. Prov., Mzimba dist., 6
miles NE Mzambazi, edge dambo, very sandy, 22 Jan. 1978, Pawek
13667 (BR, K).
Zambia: dist. Luanshya, 23 Jan. 1956, Fanshawe 2708 (BR, K,
NDO) [cited by Kativu et al. (2008) as C. macrorhizum].
The three collections from D.R.Congo are indisputably
members of the C. leptoneurum-C. macrorhizum complex.
A striking nding is the observation of hairs on the fruits [a
mixture of obtuse attened papillae c. 0.2–0.4 mm high on
faces and elongate acute hairs up to 0.7 mm long on angles
(g. 6I)], apparently a unique condition in African species of
Chlorophytum. This observation prompted us to check fruits
of collections stored under C. leptoneurum and C. macro-
rhizum in BR. Fanshawe 2708 from Zambia, and Pawek
13667 from Malawi proved to have the same fruits and very
similar habit compared to the Congolese collections. These
ve collections share many traits and obviously represent a
distinct variety with an original combination of traits in the
complex. These collections also highlight the narrow afni-
ties between C. leptoneurum and C. macrorhizum. They have
a striking horizontal rhizome, but of variable thickness from
very thin and leptoneurum-like to thick and more or less
macrorhizum-like. Leaves and bracts often have a reddish
tinge, a trait said to be typical of C. macrorhizum. Seed testa
has convex periclinal walls, without spines (g.6 C & D).
See also note under C. africanum var. nordalianum.
24. Chlorophytum longifolium Baker (Baker 1876: 327);
Baker (1898: 507); Kativu (1994: 66); Nordal et al. (1997:
17); Kativu et al. (2008: 63). – Type: Sudan/Ethiopian bor-
der, Gallabat, Matamma, Aug. 1865, Schweinfurth 8 (lecto-:
P, designated here).
Chlorophytum papillosum Rendle (Rendle 1895: 422);
De Wildeman (1921a: 28); Nordal et al. (1990: 547 g. 5e,
548 g. 6E). – Type: Tanzania, Between Zanzibar and Uyui,
1886, Taylor s.n. (holo-: BM, barcode BM000911761).
This species is represented in D.R.Congo by a variety
with golden-yellow pubescence on young ower buds.
24a. Chlorophytum longifolium Baker var. aureum (Engl.)
Meerts, comb. & stat. nov.
Chlorophytum aureum Engl., Botanische Jahrbücher für
Systematik, Panzengeschichte und Pan zengeographie
15: 469. 1892 (Baker & Engler 1892); Troupin (1955: 234;
1956: 189); Hepper (1968b: 99). – Type: Sudan, Niamniam-
land, river Huuh, 21 May 1870, Schweinfurth 3752 [lecto-:
B (2 sheets: B100249010 & B100277295), here designa-
ted; isolecto-: K]; Sudan, Dschurland, bei der grossen Se-
riba Kutschuk Ali, 29 Apr. 1896, Schweinfurth 1504 (syn-:
K); Wando, steppen in der Wälden, s.d., Schweinfurth 3570
(syn-: K).
Other representative specimens examined – D.R.Congo: VII:
Parc Nation. Garamba, savane arbustive, 26 May 1950, Noirfalise
429 (BR); Uele, Guvane, savane arbustive, May 1931, Lebrun 2881
(BR, K).
This variety has a Sudanian distribution, while the typical
variety is mostly Zambezian.
25. Chlorophytum macrophyllum (A.Rich.) Aschers.
(Schwein furth 1867: 294); Baker (1876: 323); Baker (1898:
498); Engler (1908: 308, g. 208 C & D); Kativu (1994: 67);
Nordal et al. (1997: 49, g. 9); Bjorå et al. (2008: 233, g.
6, 10, 11); Kativu et al. (2008: 85). – Anthericum macro-
phyllum A.Rich. (Richard 1851: 334). – Type: Ethiopia,
Djeladjekanne, Jun. 1840, Quartin-Dillon s.n. (lecto-: P, bar-
code P01851321); Ethiopia, Djeladjeranne, 24 Oct. 1840,
Schimper 1837 (syn-: P, isosyn-: BR, B, MPU). Fig.5 E & F.
Chlorophytum fuchsianum De Wild. (De Wildeman 1904:
102 as fuchsianus); De Wildeman (1905: 38 & Plate XVI;
1909: 54 & Plate 1, gs 1 & 2); Durand & Durand (1909:
568). – Type: D.R.Congo, Kisantu, s.d., Gillet 902 (lecto-:
BR).
Other representative specimens examined – D.R.Congo: III:
Bas-Congo, Mbanza Ngungu, entrée de la petite grotte, 3 Mar.
1987, Billiet & Jadin 4264 (BR); Mvuasi, îlot forestier, 17 Dec.
1947, Devred 4 (BR, K); VI: Eala, s.d., Goossens s.n. (BR).
Burundi: Ubimbi, prairie, 8 Dec. 1965, Lewalle 63 (BR).
C. macrophyllum has long been misinterpreted in
D.R.Congo. Our revision shows that actually only few col-
lections belong to that species, nearly all originating from
SW D.R.Congo. Most other records of that species actually
belong in C. lipendulum. At the owering stage, C. macro-
phyllum and C. lipendulum cannot be confounded. At the
fruiting stage, C. lipendulum is recognized by its blackish
colour in herbarium (pale brownish to brownish-green in
C. macrophyllum), papillate rachis (smooth in C. macro-
phyllum), and strongly saucer-shaped seeds (disk-shaped to
slightly saucer-shaped in C. macrophyllum).
The name C. macrophyllum has been applied to plants
with large owers (> 8 mm long), with 5-nerved tepals and
long anthers (> 3 mm) and large whitish bracts. We have
found that two closely related taxa with that combination of
traits have been confounded in D.R.Congo until now. i.e. C.
macrophyllum and C. clarae. See discussion under the latter
species. Fig. 5C–F shows that seed shape and seed coat de-
tails are very similar for a syntype of C. macrophyllum from
Ethiopia and for a number of collections from W D.R.Congo.
26. Chlorophytum minor Kativu (Kativu 1993: 502); Kativu
(1994: 68); Kativu et al. (2008: 81). – Type: Zambia, Mbala
Dist., Itembwe Gap and Gorge, 7 Jan. 1968, Richards 22872
(holo-: K).
Other representative specimens examined – D.R.Congo: XI:
Route Mitwaba-Manono, km 45, riv. Kalumengongo, 3 Feb. 1986,
Bamps & Malaisse 8619 (BR).
A rare species, restricted to SW Tanzania, N Zambia and
Katanga.
27. Chlorophytum nubicum (Baker) Kativu (Kativu & Nor-
dal 1993: 63); Kativu (1994: 69); Nordal et al. (1997: 15);
Kativu et al. (2008: 55). – Anthericum nubicum Baker (1876:
399
Meerts & Bjorå, Chlorophytum in Central Africa
301); Baker (1898: 484). – Type: Sudan, Jur, White Nile,
Nyangara, s.d., Petherick s.n. (holo-: BM, n.v.).
Other representative specimens examined – D.R.Congo: XI: an-
cienne piste Kolwezi-Likasi, 10°42’54”S 25°35’09”E, savane arbo-
rescente en forêt claire, alt. 1375 m, 21 Aug. 1986, Schaijes 3051
(BR).
Two collections from l’Upemba [de Witte 03621 & 07289
(BR)] key out here but are atypical, having a short peduncle
with only 1(–2) bracts and inorescence nodes with only one
ower. They probably represent a distinct taxon, but more
material, especially with fruits, is needed.
C. nubicum is very close to C. nidulans. The former can
be distinguished by spindle-shaped tubers and anthers > 2
mm.
28. Chlorophytum occultum Poulsen & Nordal (Poulsen &
Nordal 1999: 944, gs. 2C–F, 3, 4); Poulsen & Nordal (2005:
17, g. 1G; 22, 23, 37). – Type: Ouganda, Masindi Dist.,
Budongo Forest Reserve, 2 Oct. 1995, Poulsen, Nkuutu &
Dumba 975 (holo-: C, iso-: MHU, EBB; not seen).
Other representative specimens examined – Not seen. Key after
description in Poulsen & Nordal (2005).
29. Chlorophytum orchidastrum Lindl. (Lindley 1824: 79);
Baker (1898: 500); Hepper (1968b: 101, 102 g. 351); Pani-
grahi (1975: 563); Poulsen & Nordal (2005: 18, g. 1M, 24);
Bjorå & Nordal (2010: 5). – Type: Cultivated “at the garden
of the Horticultural Society, from a plant sent from Sierra
Leone, in 1822, by their collector Mr. George Don H.HS.,
1824” (holo-: CGE).
Chlorophytum seretii De Wild. (De Wildeman 1909: 56;
Plate 2 & 3); De Wildeman (1921b: 21). – Type: D.R.Congo,
env. de Gombari, 26 Apr. 1906, Seret 577 (holo-: BR).
Chlorophytum seretii De Wild. var. likimensis De Wild.
(De Wildeman 1911b: 289); De Wildeman (1912b: 526). –
Type: D.R.Congo, env. de Likimi, 21 Apr. 1910, Malchair
262 (holo-: BR).
Other representative specimens examined – D.R.Congo: II: Ter-
rit. Seke-Banza, Gimbi, vieille forêt secondaire à Terminalia su-
perba, 15 Sep. 1959, Compère 384 (BR, K); III: Gimbi, jachères
secondaires humides, 15 Jun. 1948, J. Laurent 691 (BR); V: Région
de Pene-Yumbi, sous-bois de forêt primitive, Jun. 1952, Germain
7810 (BR); VI: Panga, forêt humide sur terre riche en humus, 16
Dec. 1913, Bequaert 1500 (BR); VIII: Irumu, galerie forestière
dans la savane, 13 Mar. 1914, Bequaert 2966 (BR).
30. Chlorophytum perfoliatum Kativu (Kativu 1993: 503);
Kativu (1994: 71); Kativu et al. (2008: 69, Fig. 13.1.10). –
Type: Zambia, Kasama Dist., 95 km E of Kasama, among
rocks, s.d., Robinson 4344 (holo-: K).
Other representative specimen examined – D.R.Congo: XI:
Tilwizembe, végétation d’une termitière dans la plaine au nord du
gisement, 15 Dec. 1959, Duvigneaud 4601T (BRLU).
A rare species restricted to Zambia, Malawi and Katanga,
with only one collection in D.R.Congo.
31. Chlorophytum pilosicarinatum (Poelln.) Meerts, comb.
nov.
Anthericum pilosicarinatum Poelln., Boletim da Sociedade
Broteriana. XVII, 2A serie: 89. 1943 (Poellnitz 1943a). –
Type: Angola, Bié, Ufer des Cubango, Vila da Ponte, 1906,
Gossweiler 4018 (lecto-: K, here designated).
Other representative specimen examined – D.R.Congo: XI:
Tilwizembe, alignement métallifère rocheux, steppe sur schiste co-
baltifère, 15 Dec. 1959, Duvigneaud 4590L1 (BRLU).
The other syntypes, Gossweiler 2039 & 2663, have not
been found.
The cited collection is apparently the rst record of that
taxon since its description. C. pilosicarinatum belongs in the
C. cameronii complex and might best be treated at varietal
rank within this latter species, but more material is needed.
It is easily recognized by its unique pattern of leaf pubes-
cence, with glabrous leaf surface and long ciliate leaf mar-
gins and lower mid nerve; the hairs are retrorsely oriented at
their base; bracts have the same pattern of pubescence in the
examined collection, but less clearly so in the type specimen.
Chlorophytum galpinii var. norlindhii has a more or less sim-
ilar pattern of pubescence, but in the latter taxon the inores-
cence is lax and branched (vs. unbranched, < 5 cm long in C.
pilosicarinatum), bracts are attenuate in a long ne point (vs.
bracts ovate without a long point), tepals are 3-veined (vs.
5-nerved) and pedicels are terete (vs. narrowly winged in the
upper part).
Mortelmans 142 (BR) may belong here: it is a dwarf
plant with three very narrow cataphyll-like leaves with the
characterictic pattern of pubescence of the species.
32. Chlorophytum polystachys Baker (Baker 1878b: 326;
1898: 509); Hepper (1968b: 496); Kativu (1994: 71); Nordal
et al. (1997: 43); Kativu et al. (2008: 61). – Type: Sudan, Jur
Ghattas, 2 Jul. 1869, Schweinfurth 1838 (holo-: K, 2 sheets,
iso-: B, 3 sheets).
Other representative specimens examined – D.R.Congo: XI:
Lubumbashi, dembo de la Kiboko, 20 Nov. 1957, Schmitz 6037
(BR).
Apparently a rare species with a single collection in the
study area. This specimen is remarkable by its big fusiform
distal tubers, a character rarely reported for that species.
33. Chlorophytum pubiorum Baker (Baker 1876: 329);
Baker (1898: 509); Kativu (1994: 73); Nordal et al. (1997:
44). – Type: Mozambique, Delta du Zambèze, Dec. 1862,
Kirk s.n. (holo-: K, barcode K000256948).
Other representative specimens examined – D.R.Congo: XI:
Musosa, 28 Feb. 1940, Brédo 4019 (BR).
The single collection examined originates from the Con-
go-Zambia borderline, possibly on Zambian territory. The
species should be sought for in the oodplain of the Luapula
River.
34. Chlorophytum pusillum Baker (Baker 1878b: 325);
Baker (1898: 502); Hepper (1968b: 100); Kativu (1994:
73); Nordal et al. (1997: 55); Kativu et al. (2008: 87, 88, g.
13.1.15); non C. pusillum sensu Troupin (1956: 191). – Type:
Sudan, Jur, 10 Jul. 1869, Schweinfurth 2043 (holo-: K, iso-:
B (3 sheets), P, PRE). Figs 3A & 6B.
400
Pl. Ecol. Evol. 145 (3), 2012
Other representative specimens examined – D.R.Congo: XI:
Lubumbashi, arboretum de l’Etoile, sur termitière, 17 Jan. 1948,
Schmitz 1241 (BR); Mont Mamuntamba, 1987, Billiet & Jadin 4151
(BR, MO, P; cultivated in greenhouse: S 2072, Photo 87-0116).
Specimens from D.R.Congo have papillate bracts, which
had not been noticed previously. Seed testa has raised peri-
clinal walls which are verrucose (g. 6B) and distinct from
the convex periclinal walls of C. geophilum (g. 6A). See
also under that species.
35. Chlorophytum recurvifolium (Baker) C.Archer & Ka-
tivu (Archer & Kativu 2001: 31); Kativu et al. (2008: 17).
– Anthericum recurvifolium Baker (Baker 1906: 28). – Type:
Zimbabwe, 6-miles spruit, near Harare, s.d., Cecil 143 (holo-:
K).
Anthericum longistylum Baker (Baker 1876: 305); Ober-
meyer (1962: 686, 687: Fig. 2). – Type: South Africa, Gold
elds, 1870, Baines s.n. (holo-: K, barcode K000256975).
Other representative specimens examined – D.R.Congo: XI:
Kisenge, colline de Mn, bosquet d’Uapaca robynsii, 1956, Du-
vigneaud 2342Li (BRLU); s.l. “Mn”, s.d., Joris D30 (BRLU); Bun-
keya, Dec. 1942, s. rec. 457 (BRLU).
A phytogeographically very interesting record, as this
species has a southern African distribution, reaching its
northern limit in Zimbabwe. Katangan populations are wide-
ly disjunct.
36. Chlorophytum ruahense Engl. (Engler 1900: 361);
Poellnitz (1946: 332); Kativu (1994: 75); Nordal et al. (1997:
13); Kativu et al. (2008: 74, g. 13.1.13). – Type: Tanzania,
Iringa Dist., Uhehe, hanging cliff on Ruaha R., 10 Jan. 1899,
Goetze 461 (holo-: B).
Other representative specimens examined – D.R.Congo: XI:
Baya-Pierkat, végétation d’une colline rocheuse de calcaires de la
Mofya, 7 Jan. 1960, Duvigneaud 4953L1 (BRLU); Route Nguba-
Mokabe Kasadi, à 2 km au S de Kate, forêt claire, 7 Nov. 1988,
Pauwels 7211 (BR).
Katangan collections extend the distribution area of this
rare species, otherwise conned to SW Tanzania and N Zam-
bia, somewhat to W. The seeds are echinulate (seed not seen
in FZ). The leaves of the Congolese collections are variable
in width.
37. Chlorophytum rubribracteatum (De Wild.) Kativu
(Kativu & Nordal 1993: 64); Kativu (1994: 75); Nordal
et al. (1997: 36); Kativu et al. (2008: 50). – Anthericum
rubribracteatum De Wild. (De Wildeman 1913a: 508;
1913c: 9); De Wildeman (1921a: 27). – Type: D.R.Congo,
Lubumbashi, Nov. 1911, Hock s.n. (holo-: BR, barcode
BR0000008761583). Fig. 2F.
Anthericum roseum Poelln. (Poellnitz 1942: 114); Lewal-
le (1972: 173). – Type: Tanzania, Songea dist., E of Ungoni,
Mampyuy, s.d., Busse 727 (holo-: B, not seen; iso-: EA).
Other representative specimens examined – D.R.Congo: XI:
Luiswishi, colline cuprifère, 18 Nov. 1980, Malaisse 11220 (BR,
WAG, MO); Kipopo, dembo sur plateau latéritique, 30 Jan. 1962,
Schmitz 7570 (BR).
Burundi: Territ. Bubanza, ferme de la Randa, savane boisée, s.d.,
Lewalle 1254 (BR).
The species is well characterized by the red-coloured cat-
aphylls, narrowly graminiform leaves, and lax inorescence
often with remote lower nodes. Bracts are generally carinate,
erect, and often clasping the peduncle. However, C. rubri-
bracteatum is much more variable than mentioned in FZ and
FTEA, especially in intensity of red coloration (often lack-
ing on bracts), pubescence (from glabrous to shortly pubes-
cent on lower leaf surface and peduncle), height (5–80 cm)
and leaf width (1–10 mm). Size tends to be negatively cor-
related with pubescence. In Katanga, the largest specimens
occur in tall vegetation on moist soil (e.g. Duvigneaud 5055
(BRLU)). All collections from Burundi also belong to the ro-
bust form. More research is needed to decide if robust forms
deserve taxonomic recognition.
Some collections with broad leaves, pale coloration and
dense pubescence on lower leaf surface and peduncle ap-
proach C. cameronii var. grantii [e.g. Van Meel 805 (BR);
Schmitz 3633 (BR); Duvigneaud 4769C & 4772A (BRLU)].
Robyns 1646 (BR) is an extreme dwarf form with pros-
trate peduncle, which may belong here. More material is
needed.
38. Chlorophytum sparsiorum Baker (Baker 1876: 325);
Baker (1898: 498); Hepper (1968b: 100); Troupin (1988: 43,
g. 4/1; 44); Bjorå & Nordal (2010: 6). – Type: Equatorial
Guinea, Fernando Po, Apr. 1860, Mann 388 (holo-: K).
Chlorophytum semlikiense De Wild. (De Wilde-
man 1921b: 20); Robyns & Tournay (1955: 356). – Type:
D.R.Congo, Kasonsero, Vallée de Semliki, s.d., Bequaert
5041 (holo-: BR).
Chlorophytum beniense De Wild. (De Wildeman 1921b:
13); Lebrun et al. (1948: 36); Robyns & Tournay (1955:
354); Troupin (1988: 42; 45, g. 15/2). – Type: D.R.Congo,
Mayolo (Beni), s.d., Bequaert 3981 (holo-: BR).
Chlorophytum elongato-fusiforme De Wild. (De Wilde-
man 1921b: 15). – Type: D.R.Congo, en aval de Panga, 15
Dec. 1913, Bequaert 1485 (holo-: BR).
Chlorophytum ituriense De Wild. (De Wildeman 1921b:
17); Troupin (1956: 191). – Type: D.R.Congo, Penghe, 17
Feb. 1914, Bequaert 2513 (syn-: BR), Irumu, 12 Mar. 1914,
Bequaert 2398 (syn-: BR).
Chlorophytum butaguense De Wild. (De Wilde-
man 1921b: 19); Robyns & Tournay (1955: 354). – Type:
D.R.Congo, Ruwensori (Butagu), 15 Apr. 1914, Bequaert
3686 (lecto-: BR, barcode BR0000006420253, here des-
ignated); Ruwensori, 16 May 1914, Bequaert 4292 (syn-:
BR).
Chlorophytum kirkii sensu Robyns & Tournay (1955:
356) non Chlorophytum kirkii Baker.
Chlorophytum comosum sensu Baker (1876: 329); Baker
(1897a: 400); Troupin (1988: 42; 45, g. 15/3); Nordal et
al. (1997: 55); Poulsen & Nordal (2005: 15), Kativu et al.
(2008) non Chlorophytum comosum (Thunb.) Jacques.
Other representative specimens examined – D.R.Congo: II:
Luki, est du village de Kimbuya, association végétale sur le replat
du somet du rocher de granit, lisière de la forêt à Gilletiodendron,
401
Meerts & Bjorå, Chlorophytum in Central Africa
Figure 6 – Scanning electron microscope images of seeds and seed testa. A, C. geophilum, seed testa (Noirfalise 441); B, C. pusillum,
seed testa (Billiet & Jadin 4151); C & D, C. leptoneurum var. katangense, seed and seed testa (Quarré 3741); E & F, C. stenopetalum var.
stenopetalum, seed and seed testa (Seret 196); G & H, C. stenopetalum var. latifolium, seed and seed testa (Detilleux 570); I, C. leptoneurum
var. katangense, fruit (Quarré 3741).
AB
H
FED
C
G I
2 Aug. 1957, Wagemans 1594 (BR); IV. Kwango, Kisantu, Panzi,
1925, Vanderyst 16195 (BR); VI: Prov. Orientale, Territ. Mambasa,
rivière Lemda, région Ndesa, 700 m, clairière humide dans forêt à
Gilbertiodendron dewevrei, 8 Jun. 1956, Christiaensen 1763 (BR);
Yangambi, au pied du plateau de l’Isalowe, alt. 470 m, forêt primi-
tive ripicole le long du euve Congo, 18 May 1938, Louis 9437
(BR); Rivière Dhi, pont de l’ancienne route Djugu-Nioka, sous-bois
ombragé humide de la forêt galerie, 19 Apr. 1991, Dhtechuvi 1128
(BR); VIII: Mont Aboro, pied du versant W, alt. 2200 m, forêt de
montagne à Chrysophyllum fulvum, 15 Apr. 1958, Bamps 154 (BR);
Ituri, Djugu (Kibali), alt. 1780 m, forêt, Sep. 1931, Lebrun 3942
(BR); IX: Territ. Kabare, Kahusi, le long d’un caniveau, alt. 2000
m, 11 Feb. 1959, A. Léonard 2983 (BR); XI: Parc Nation. Upem-
ba, Katongo, alt. 1750 m, galerie forestière, 16 Apr. 1948, de Witte
3713 (BR).
Rwanda: Gisakura (Cyangugu), alt. 1950 m, plantation de thé, 23
Jul. 1954, Auquier 3330 (BR); Territ. Shangugu, Rugege, alt. 2000
m, forêt de montagne, 21 Mar. 1956, Christiaensen 1488 (BR).
Burundi: Territ. Bururi, Rumonge, alt. 1000 m, talus ombreux, 16
Mar. 1969, Lewalle 3338 (BR).
This taxon has been known as C. comosum auct. A recent
molecular phylogenetic study (Bjorå et al. 2008) has shown
that this taxon is not monophyletic. In particular, accessions
from C Africa and S Africa belong in different clades. Since
the type specimen originates from S Africa, the epithet “co-
mosum” can no longer be applied to C African material.
C sparsiorum is an extremely polymorphic taxon. A
multivariate statistical analysis (Poulsen & Nordal 2005)
concluded that three infraspecifc taxa can be recognized,
i.e. one with strap-shaped, blackening leaves (“Chlorophy-
tum comosum var. comosum”), a second with elliptic, non
blackening, leaves more or less attenuate into a pseudopeti-
ole (“var. sparsiorum”) and a third with elliptic, non black-
ening leaves, abruptly contracted into a petiole (“var. bipin-
dense”). We have not been able to settle the objective limits
of those taxa in the abundant material examined. Leaf width
and shape and colour in herbarium (blackening or not) show
an almost continuous pattern of variation in material from
Central Africa. Therefore, the placing of many collections is
problematic. Molecular markers may hopefully provide new
insight into this taxonomic conundrum.
402
Pl. Ecol. Evol. 145 (3), 2012
Collections from S Katanga [Schmitz 2752 (BR), Van-
denberghe 56 (BR), Bodenghien 89 (BR), all from “muhulu”
gallery forests] have long, obcordate fruits 10 × 7 mm, and
narrow 20-nerved leaves; seeds however are typical.
Bequaert 2081 (BR) has densely papillate bracts, a char-
acter diagnostic of the W African C. inornatum (Poulsen &
Nordal 2005), but it lacks mature seeds. More material is
needed.
Collections from eastern D.R.Congo and Burundi with
narrow, not blackening, leaves are problematic and need re-
vision [e.g. Deville 257 (BR), Froment 492 (BR)].
Some collections in W D.R.Congo (region of Eala) [e.g.
Gombe (Equateur), août 1903, M. Laurent s.n. (BR, barcode
BR0000005720651)] are intermediate between C. lancifo-
lium subsp. togoense and C. sparsiorum, and might be of
hybrid origin.
Lejoly 4402 (BR, BRLU), from Kisangani, has
4–5-nerved tepals, anthers equalling laments and long
bracts, and therefore appears somewhat intermediate be-
tween C. sparsiorum and the C. macrophyllum complex.
The name “Chlorophytum laxum” has sometimes been
improperly used for slender, narrow-leaved C. sparsiorum.
See also notes under C. lipendulum, C. debile, and C.
“sp. A near comosum”.
39. Chlorophytum sphacelatum (Baker) Kativu (Kativu &
Nordal 1993: 64); Kativu (1994: 76); Nordal et al. (1997:
34); Kativu et al. (2008: 42). – Anthericum sphacelatum
Baker (Baker 1876: 303); Baker (1898: 489); De Wildeman
(1905: 39); Durand & Durand (1909: 568). – Type: Angola,
112 km from Ambriz, Jun. 1873, Monteiro s.n. (holo-: K,
barcode K000256908). Fig. 3B.
Anthericum lukiense De Wild. (De Wildeman 1911a:
273); De Wildeman (1912a: 341). – Type: D.R.Congo, Luki,
1909, Brixhe 6 (holo-: BR).
A. lukiense De Wild. var. intermedium De Wild. (De
Wildeman 1911a: 273); De Wildeman (1912a: 342). –
Type: D.R.Congo, env. Kisantu, 1909, Gillet s.n. (lec-
to-: BR, barcode BR0000008551863, here designated),
D.R.Congo, Mayombe, 1909, Deleval s.n. (syn-: BR, bar-
code BR0000008551559).
A. lukiense De Wild. var. kionzoense De Wild. (De
Wildeman 1911a: 273); De Wildeman (1912a: 342). – Type:
D.R.Congo, Région du Kionzo, 1907, Gillet 4001 (holo-:
BR).
Anthericum hockii De Wild. (De Wildeman 1911a:
265); De Wildeman (1913c: 8; 1921a: 26). – Chlorophytum
sphacelatum (Baker) Kativu var. hockii (De Wild.) Nordal
(Nordal et al. 1997: 35); Kativu et al. (2008: 44). – Type:
D.R.Congo, Katanga, vallée de la Luembe, Feb. 1910, Hock
s.n. (holo-: BR, barcode BR0000008552211), synon. nov.
Anthericum velutinum De Wild. (De Wildeman 1913a:
508; 1913c: 10); De Wildeman (1921a: 27). – Type:
D.R.Congo, Lubumbashi, Dec. 1911, Hock s.n. (holo-: BR,
barcode BR0000008551542).
Anthericum homblei De Wild. (De Wildeman 1914: 108);
De Wildeman (1921a: 26; 1930: 94). – Type: D.R.Congo,
Katanga, Région du Lualaba, Vallée Kapanda, Dec. 1912,
Homblé 967 (holo-: BR).
Chlorophytum sphacelatum (Baker) Kativu var. milan-
jianum (Rendle) Nordal (Nordal 1997: 35); Kativu et al.
(2008: 43). – Type: Malawi, Mt. Mulanje, Oct. 1891, White
123 (holo-: BM), synon. nov.
Other representative specimens examined – D.R.Congo: II:
Luki, sentier vers Kiobo, 4 Feb. 1947, Devred 3101 (BR); III: Gum-
bi Plateau, savane à Hymenocardia, 27 Dec. 1948, Toussaint 694
(BR); XI: Lubumbashi, savane sèche boisée et base des termitières,
Feb. 1912, Homblé 129 (BR); Lubumbashi, herbe de plaine, Dec.
1912, De Giorgi 318 (BR); Kipopo, sol argileux rouge, graveleux,
en forêt claire, 28 Nov. 1962, Schmitz 8093 (BR).
The species is very variable, though always easy to rec-
ognize by its black-tipped tepals. FZ and FTEA recognize
three varieties (var. sphacelatum, var. hockii (De Wild.) Nor-
dal, var. milanjianum (Rendle) Nordal), based upon pedun-
cle hairiness and shape (terete or attened) and inorescence
shape (globose or racemose). In material from D.R.Congo,
the aforementioned traits are often uncorrelated and many
specimens are difcult to assign to one of those taxa, which
is why we do no longer distinguish them.
Some forms have yellowish velvety pubescence [Schmitz
8093 (BR)].
Dwarf variants (< 12 cm) occur under poor, dry or oth-
erwise stressful conditions, e.g. de Witte 07112 (BR) with 2
mm-wide leaves and Duvigneaud 2130L (BRLU) from Kas-
ompi copper-hill with short tepals (5–7 mm) and very short
pedicels. Such forms approach C. graniticola Kativu and C.
pygmaeum (Weim.) Kativu.
40. Chlorophytum sphagnicolum Meerts, nom. nov.
Anthericum andongense Baker, Transactions of the Linnean
Society, Ser. 2, 1: 257. 1878 (Baker 1878a); Baker (1898:
482); Poellnitz (1943a: 57). – Type: Angola, Pungo Andon-
go, s.d., Welwitsch 3798 (holo-: BM, iso-: P), non Chloro-
phytum andongense Baker.
Other representative specimens examined – D.R.Congo: III:
Rivière Luvu entre Ngidinga et Kimvula, 12 Apr. 1948, Du-
vigneaud 704 (BRLU); IV: Entre Giugungi et Popokabata, savane à
sable blanc, Dec. 1929, Lebrun 79 (BR).
In D.R.Congo, this species has the narrowest leaves and
the smallest owers in the C. cameronii complex. It is also
characterized by short anthers (2.5 mm) equalling laments
and by thin roots (< 0.7 mm in herbarium). It has quite unu-
sual habitat requirements occurring on moist nutrient-poor
sand or occasionally in Sphagnum bogs.
The W African Chlorophytum immaculatum (Hepper)
Nordal resembles this taxon (see Hepper 1968a: 457, g.
6/1-3, 458; Vanden Berghen 1988: 417; Fig. 343) and fur-
ther research is needed to decide if they are actually distinct.
We have kept them distinct due to differences in the bracts
(bracts membranaceous, hyaline, often lobed in C. immac-
ulatum, vs. scariose, entire, with prominent nerves in C.
sphagnicolum). Should they turn out to be conspecic, the
name C. immaculatum would have precedence.
Anthericum andongense Baker has been reported from
Katanga by De Wildeman (1913c: 6, 1921a: 26), based upon
403
Meerts & Bjorå, Chlorophytum in Central Africa
Kassner 2462 (K); in our opinion this collection is a depau-
perate specimen of C. rubribracteatum.
41. Chlorophytum staudtii Nordal (Kativu & Nordal 1993:
64). – Anthericum zenkeri Engl. (Engler 1902: 91), Hep-
per (1968b: 97). – Type as C. staudtii. – Type: Cameroon,
Yaounde, Mar. 1894, Zenker & Staudt 268 (holo-: B (†?),
iso-: K, S).
Other representative specimens examined – D.R.Congo: III: vers
Kilongo, territ. Popokabaka, savane, 28 Oct. 1958, Pauwels 444
(BR); IV. Gana, Région des Bamfunuka, Jun.–Jul. 1915, Vanderyst
& Lambrette 5653 (BR).
An insufciently known species closely related to C.
cameronii and possibly best treated at varietal rank. More
material is needed. Narrow-leaved forms of the C. cameronii
complex need revision preferably with molecular markers.
Pauwels 2544 (BR), with 9-mm wide leaves with 23
nerves and 3 inorescence branches 6–11 cm long, is inter-
mediate between C. staudtii and the W African C. limosum
(Baker) Nordal.
42. Chlorophytum stenopetalum Baker (Baker 1876: 331);
Baker (1898: 502); Hepper (1968b: 100); Kativu (1994: 78);
Nordal et al. (1997: 54); Kativu et al. (2008: 86). – Type: Ni-
ger, Niger R, Nupe, 1859, Barter s.n. (holo-: K000407093).
The species is variable in peduncle length, raceme densi-
ty, leaf width and bract size. Two varieties can be recognized.
1. Peduncle 0–1 cm; raceme compact and rigid, with
pedicels 0–2 mm; bracts prominent and persisting,
concealing young fruits; leaves 4–9 cm wide, broadly
ovate, broadly elliptic or obovate, more or less
appressed on the ground (g. 3C)..............................
....................................C. stenopetalum var. latifolium
1. Peduncle 1–5(–15) cm; raceme not rigid, with pedicels
2–8 mm; bracts not concealing young fruits; leaves
1–4.5 cm wide, narrowly elliptic or elliptic, erect to
spreading.............C. stenopetalum var. stenopetalum
42a. Chlorophytum stenopetalum Baker var. stenopetalum
– Chlorophytum schweinfurthii Baker (1898: 503); De Wil-
deman (1912a: 347); Troupin (1956: 192; g. 31). – Type:
Sudan, Jur Ghattas, 21 May 1869, Schweinfurth 1968 (lecto-:
K; isolecto-: B); 12 Jul. 1869, Schweinfurth 2068 [syn-: P, K
(2 sheets)]. Fig. 6E & F.
Other representative specimens examined – D.R.Congo: V: La
Pastorale, section Lualu, vallées et endroits humides, Nov. 1931,
Quarré 2757 (BR); VI: Uele, env. Bambesa, 1936, Pittery 532
(BR); VII: Parc Nation. Garamba, piste centrale vers km 30, alt.
700-800 m, savane arbustive, 30 Apr. 1952, Troupin 861 (BR); XI:
Kiubo, à l’ouest de la route vers Bunkheya, 5 km sud du pont, sol
rouge brun sur schistes, 27 Jan. 1954, van Oosten 211 (BR).
Exceptionally robust forms with peduncle up to 15 cm
with one foliaceous bract, forked raceme are occasionnally
found in Katanga [e.g. Pogge 1479 (B), referred to as C.
macrophyllum by Baker (1898: 498) and Durand & Durand
(1909: 568); Quarré 2209 (BR); Bamps & Malaisse 8009
(BR); Duvigneaud 5229 (BRLU)]. They can be confused
with C. macrophyllum. In the latter species, however, anthers
are longer (4–5 mm vs. 2–3 mm), fruiting pedicels are longer
(10–20 mm vs. 3–10 mm), bracts are not persisting and seed
testa is different (raised periclinal walls with acute ridge, vs.
at to very shallowly convex periclinal walls in C. macro-
phyllum). Such robust forms may deserve taxonomic recog-
nition but more material is needed.
42b. Chlorophytum stenopetalum Baker var. latifolium
(Engl. & K.Krause) Meerts, comb. & stat. nov.
Chlorophytum latifolium Engl. & K.Krause, Botanische
Jahrbücher für Systematik, Panzengeschichte und Pan-
zengeographie 45: 134. 1910 (Engler & Krause 1910: 134).
– Type: Zambia, Broken Hill, 17 Dec. 1907, Kassner 2014
(holo-: B; iso-: K). Fig. 3C; g. 6G & H.
Chlorophytum geophilum sensu Kativu et al. (2008: 87)
quoad Hoell & Nordal 20.
Other representative specimens examined – D.R.Congo: XI: En-
tre Likasi et Kambove, forêt claire sur terre rouge profonde, 27 Dec.
1959, Duvigneaud 4786 (BRLU); Lubumbashi, Keyberg, Kisanga,
barrage, terre rouge jaunâtre en sous-bois, 21 Feb. 1957, Detilleux
570 (BR); Sanctuaire Mikembo, forêt claire, 16 Jan. 2010, Meerts
2010/24 (BRLU).
C. stenopetalum var. latifolium is one of the most wide-
spread Chlorophytum species in miombo woodland in the
Lubumbashi plain. It was reported with doubt from N Zam-
bia in FZ (Kativu et al. 2008: 89).
We prefer to treat this taxon at varietal rank because
broad leaves and short peduncle are not always correlated in
the examined materials, with some collections being inter-
mediate between var. latifolium and var. stenopetalum.
43. Chlorophytum stolzii (K.Krause) Kativu (Kativu & Nor-
dal 1993: 64); Kativu (1994: 78); Nordal et al. (1997: 15);
Kativu et al. (2008: 58, g. 13.1.10); Bjorå & Nordal (2010:
9). – Type: Tanzania, Rungwe Dist., Kyimbila, Mulinda, 12
Oct. 1910, Stolz 339 (holo-: B, iso-: K).
Acrospira asphodeloides Baker (Baker 1878a: 255 &
tab. 34 gs. 4-7); Baker (1898: 477); Rendle (1899: 50); De
Wildeman (1905: 38; 1910b: 264, Plates 42 & 43; 1912a:
370; 1916: 163; 1930: 92); Durand & Durand (1909: 567)
non Chlorophytum asphodeloides C.B.Wright – Type: Ango-
la, Pungo Adongo, Mar. 1857, Welwitsch 3777 (lecto-: BM);
Welwitsch 3778, 3779 (syn-: BM).
Anthericum russissiense Poelln. (Poellnitz 1942: 70). –
Type: Deutsch-Ost-Africa [Burundi], Seengebiet, Usumbu-
ra [Bujumbura], steppen der Russissi-Ebene, 31 Mar. 190?,
Keil 278 (holo-: B).
Other representative specimens examined – D.R.Congo: III:
Mvuazi, dans la savane, 19 Aug. 1957, Delhaye 177 (BR); IV: Vers
Kipindi, Territ. Popokabaka, savane arborée, 4 Jul. 1959, Pauwels
3755 (BR); V: Mugonda, route Abville-Kabambara, 12 Jul. 1932,
Luxen 166 (BR); VI: Gombe, moerasje bij den Congo, 11 Apr.
1947, Jans 456 (BR); XI: Route Kolwesi-Likasi, 13 km à l’est du
carrefour avec les pistes Tenke-Kando, rudéral, 16 Jul. 1983, Scha-
ijes 1931 (BR).
Burundi: Territ. Bujumbura, route Bujumbura-Bugarama, savane
sur talus, 9 Apr. 1972, Lewalle 6705 (BR).
See note under C. velutinum.
404
Pl. Ecol. Evol. 145 (3), 2012
44. Chlorophytum subpetiolatum (Baker) Kativu (Ka-
tivu & Nordal 1993: 64); Kativu (1994: 79); Nordal et al.
(1997: 36); Kativu et al. (2008: 50, g. 13.1.8). – Antheri-
cum subpetiolatum Baker (Baker 1876: 302); Baker (1898:
481); Troupin (1955: 232; 1956: 184; 1988: 32, g. 9/1);
Hepper (1968b: 96). – Type: Mozambique, Morrumbala Mt,
lower Zambezi R., 18 Jan. 1863, Kirk s.n. (holo-: K, barcode
K000256915).
An extremely variable species, though generally easy to
recognize by its pubescent peduncle, carrot-like roots, large
owers, and distichous leaves. A robust pubescent form in
Katanga deserves taxonomic recognition.
1. Leaves rosulate, softly pubescent on abaxial side, 4–8
cm wide, up to 60 cm long when fully developed;
rhizome > 3 cm in diameter; on termite mounds in
miombo woodland (g. 3D)........................................
................................C. subpetiolatum var. pilosifolium
1. Leaves distichous or nearly so, glabrous or, rarely,
shortly pubescent on abaxial side on nerves, 0.4–3 cm
wide, 6–40 cm long when fully developed; rhizome <
3 cm in diameter; in open habitats and woodland........
............................C. subpetiolatum var. subpetiolatum
44a1. Chlorophytum subpetiolatum (Baker) Kativu var.
subpetiolatum
Anthericum monophyllum Baker (Baker 1878b: 324);
Lewalle (1972: 173). – Type: Sudan, Djur, Seriba Ghattas, 28
May 1869, Schweinfurth 1793 (holo-: K).
Anthericum laurentii De Wild. (De Wildeman 1905: 39).
– Type: D.R.Congo, Bas Kasaï, 10 Nov. 1903, E. & M. Lau-
rent s.n. (holo-: BR, barcode BR0000008807335; iso-: BR,
barcode BR0000008807304).
Acrospira laurentii De Wild. (De Wildeman 1906: 211);
Durand & Durand (1909: 568). – Type: D.R.Congo, Kasaï,
Gambumi, 31 Mar. 1905, M. Laurent 449 (holo-: BR),
synon. nov.
Anthericum claessensii De Wild. (De Wildeman 1911a:
272); De Wildeman (1912a: 340). – Type: D.R.Congo, env.
de Bakulu, Oct. 1909, Claessens 119 (holo-: BR).
Anthericum hecqii De Wild. (De Wildeman 1911a: 277);
De Wildeman (1913c: 7; 1921a: 26). – Type: D.R.Congo,
1899, Hecq s.n. (holo-: BR, barcode BR0000008552204).
Anthericum malchairii De Wild. (De Wildeman 1911a:
274); De Wildeman (1912a: 343; 1916: 164). – Type:
D.R.Congo, Likimi, 05 May 1910, Malchair 320 (holo-:
BR).
Anthericum tuberosum De Wild. (De Wildeman 1913a:
508); De Wildeman (1913c: 10) nom. illeg., non Roxb. –
Type: D.R.Congo, Kambove, Oct. 1911, Hock s.n. (holo-:
BR, barcode BR0000008807342).
Acrospira breviscapa De Wild. (De Wildeman 1915: 3);
De Wildeman (1921a: 26). – Type: D.R.Congo, env. Katen-
tania (Biano), Nov. 1912, Homblé 746 (holo-: BR, 2 sheets),
synon. nov.
Anthericum laurentii De Wild. var. minor De Wild. (De
Wildeman 1921a: 10). – Debesia minor (De Wild.) Ro-
byns & Tournay (Robyns & Tournay 1955: 352). – Type:
D.R.Congo, Kabare, 31 Aug. 1914, Bequaert 5503 (holo-:
BR).
Acrospira kapiriensis De Wild. & Ledoux (De Wildeman
1930: 91). – Type: D.R.Congo, Kapiri, 6 Nov. 1913, Charli-
er s.n. (holo-: BR, barcode BR0000008767653), synon. nov.
Anthericum tropicum Poelln. (Poellnitz 1941: 232) nom.
nov. for Anthericum tuberosum De Wild.
Anthericum unifolium Poelln. (Poellnitz 1942: 78). –
Type: Deutsch-Ost-Africa [Burundi], Seengebiet, Usumbura
[Bujumbura], 22 Oct. 1905, Keil 201 (holo-: B).
Anthericum urumburense Poelln. (Poellnitz 1942: 135). –
Type: Deutsch-Ost-Africa [Burundi], Seengebiet, Usumbura
[Bujumbura], 10 Jul. 1904, Keil 76 (holo-: B).
Debesia contorta Lebrun & L.Touss., substitute name
(Lebrun 1947: 268).
Other representative specimens examined – D.R.Congo: III:
1899, Hecq s.n. (BR); IV: Territ. Dibaya, Badibanga, savane, 17
Oct. 1956, Liben 1721 (BR); V: Basase, champs sur terre sablon-
neuse, 13 Oct. 1936, Matagne 235 (BR); VI: Mringa-Lopozi, villa-
ge Pumbi, Oct. 1912, Dauvrin 8 (BR); VII: Bili, lambeau fores-
tier, May 1931, Lebrun 2823 (BR); IX: Parc des Virunga, région
du camp de la Rwindi, vers les Monts Mitumba, 1025 m, savane
à succulentes, 27 Aug. 1956, de Witte 13333 (BR); XI: Parc de
l’Upemba, Kalungwe, alt. 1740 m, savane brûlée, 6 Aug. 1947, de
Witte 2752 (BR).
Burundi: Bujumbura, alt. 850 m, savane, 24 Nov. 1969, Lewalle
4116 (BR).
Rwanda: Parc nation. Kagera, région du Lac Kivumba, Kibongo,
alt. 1300 m, savane arborescente sur sol brun foncé, 16 Jan. 1970,
Bouxin & Radoux 1375 (BR).
Part of the variation is correlated to the habitat and se-
ems to be genetic. Populations from burnt steppic savanna
on poor Kalahari sand or copper soil have one or two small
xeromorphic leaves and are generally hysteranthous. Popula-
tions from less stressful habitats have more numerous, softer
and larger leaves.
Plants with glabrous peduncle and rachis are rarely en-
countered [Michel & Reed 571 (BR)].
Plants from Burundi often have only 1 narrow linear leaf
sheathing the peduncle and have been referred to as Antheri-
cum monophyllum Baker. However, intermediates exist with
other forms and this taxon is not recognized here.
An unusual dwarf variant with tufted habit and erect, lin-
ear leaves is found in Upper Katanga in steppe on copper
contaminated soil [Malaisse et al. 215 (BR)].
Plants with yellow-striped leaves have been collected on
several occasions in NE Congo. They were rst described at
varietal rank but this character seems of low taxonomic value
and the rank of form appears more appropriate.
44a2. Chlorophytum subpetiolatum (Baker) Kativu var.
subpetiolatum f. variegatum (De Wild.) Meerts, comb. &
stat. nov.
Acrospira laurentii De Wild. var. variegata De Wild., Enu-
mération des plantes récoltées par Emile Laurent. III: 212.
1906 (De Wildeman 1906); Durand & Durand (1909: 568). –
Type: D.R.Congo, Eala, 10 Jul. 1919, M. Laurent 784 (holo-:
BR).
405
Meerts & Bjorå, Chlorophytum in Central Africa
Chlorophytum variegatum Schweinf. ex Poelln.
(Poellnit z 1945: 227); De Wildeman (1912a: 347). – Type:
Sudan, Land der Monbuttu, bei Munsa, 1870, Schweinfurth
3365 (holo-: B), synon. nov. non Anthericum variegatum
Hort.
Other representative specimens examined – D.R.Congo: VI:
Territ. Bambesa, Senza, village, Apr. 1921, Claessens 524 (BR);
VII: Bas-Uele, dans un champ de maïs, 20 May 1935, Dewulf 855
(BR); Tukpo, savane, s.d., Du Bois 687 (BR); Yangambi, cultivé,
plante récoltée dans la savane arbustive des env. d’Ango (Uele), 19
Dec. 1947, Germain 485 (BR).
Other materials examined – Congo (Brazzaville): Pays des
Bondjos, sur l’Oubangui, 13 Aug. 1902, A. Chevalier 5162 (P).
This striking form with yellow striped leaves is said to
be cultivated in villages for its supposed magic properties.
It is probably a mutant favoured by cultivation. Apart from
D.R.Congo and Congo (Brazzaville), this form has also been
reported from Cameroun and the Sudan (Poellnitz 1945:
228).
Some collections have a long, apparently not mon-
iliform, rhizome unusual for C. subpetiolatum [M. Laurent
784 (BR)]. If this trait turns out to be constant, a higher rank
might be justied. More material is needed.
44b. Chlorophytum subpetiolatum (Baker) Kativu var.
pilosifolium (De Wild.) Meerts, comb. & stat. nov.
Acrospira breviscapa De Wild. var. pilosaefolia De Wild.,
Contribution à la ore du Katanga. Supplément III: 90. 1930
(De Wildeman 1930). – Type: D.R.Congo, Katanga, Kafubu,
ferme Granat, 8 Nov. 1927, Quarré 775 (holo-: BR). Fig. 3D.
Chlorophytum unyikense Engl. (Engler 1901); Poellnitz
(1946: 346). – Type: Tanzania, Mbeya, Dist., Mbosi Mt, 11
Nov. 1899, Goetze 1426 (holo-: B, iso-: P, PRE), synon. nov.
Other representative specimens examined – D.R.Congo: XI: 9
km NE Lubumbashi, savane de la Luiswishi, alt. 1250 m, 19 Oct.
1973, Bulaimu 714 (BR); savane de la Luiswishi, 26 Oct. 1970,
Malaisse 6686 (BR, K); Lubumbashi, campus de la Kasapa, pied de
termitière boisée, Nov. 2009, Meerts & Muding 76 (BRLU).
Other specimens examined – Zambia: Northern region, Nkolem-
fumu, miombo woodland, alt. 1100 m, 3 Dec. 2007, Hoell & Nordal
GH126 (K, O); Solwezi district, Mwilunga-Solwezi road, alt. 1200
m, on ant hill in red clay soil, 24 Nov. 1962, Richards 17494 (K);
Mufulira, ant hill on plateau woodland, 11 Nov. 1953, Fanshawe
F494 (K).
This variety is not rare on termite mounds in the region
of Lubumbashi. Plants having the distinctive traits of this va-
riety had already been reported from N Zambia by Kativu et
al. (2008: 54).
In preliminary phylogenetic analyses the Congolese
C. subpetiolatum var. pilosifolium [Meerts & Muding 77
(BRLU)] forms a highly supported clade with C. subpetiola-
tum from a wide distribution range from Ethiopia [Herrmann
206 (ETH)] in the North to Zambia in the South [Hoell &
Nordal 15 (O)]. C. subpetiolatum shows extensive morpho-
logical variation, but forms a monophyletic group with C.
suffruticosum as the closest sister species, both highly sup-
ported.
45. Chlorophytum velutinum Kativu (Kativu & Nordal
1993: 503); Kativu (1994: 81); Kativu et al. (2008: 58, g.
13.1.10). – Type: Zambia, Luano, in crevices on rock out-
crop, s.d., Fanshawe 9628 (holo-: K, iso-: NDO). Fig. 3E.
Acrospira homblei De Wild. (De Wildeman 1915: 4); De
Wildeman (1921a: 26). – Type: D.R.Congo, Katanga, Vallée
de Kapiri, Feb. 1913, Homblé 1119 (holo-: BR), non Chloro-
phytum homblei De Wild., synon. nov.
Other representative specimens examined – D.R.Congo: XI: 20
km au N de Luambo, route Likasi-Kolwesi, forêt claire, 12 Mar.
1983, Schaijes 1861 (BR); Fungurume, hill V, steppic savanna on
Cu/Co mineralization substrate, 6 Apr. 1990, Tropmetex 160 (BR,
K, MPN, WAG); Katwe, Kundelungu Plateau, rough grassland with
bare patches, 7 Mar. 1975, Hooper & Townsend 607 (P, K).
This species was rst described by De Wildeman in 1915
as Acrospira homblei. In 1993 it was described again as a
new species by Kativu as Chlorophytum velutinum, without
reference to De Wildeman’s name. The epithet “homblei”
should have precedence. However, the combination “Chloro-
phytum homblei” is illegitimate because C. homblei De Wild.
(= C. blepharophyllum) was described in 1913. Therefore C.
velutinum remains the legitimate name of that species.
C. velutinum has long been confounded with C. stolzii.
Collections from D.R.Congo depart from the description
given in FZ (Kativu et al. 2008: 58) in having broader leaves
(up to 15(–22) mm vs. 4–6 mm in FZ), higher shoots (up to 2
m vs. 95 cm), longer stamens (25 mm vs. 12 mm) and bigger
seeds (2.5 mm vs. 1 mm).
The Congolese C. velutinum [Meerts 2010/32 (BRLU)]
forms a strongly supported clade with C. stolzii [Hoell &
Nordal 101 (O)] from Zambia.
46. Chlorophytum vestitum Baker subsp. pilosissimum
(Engl. & K.Krause) Meerts, comb. nov.
Chlorophytum pilosissimum Engler & K.Krause, Botanische
Jahrbücher für Systematik, Panzengeschichte und Pan-
zengeographie 45: 139. 1910. (Engler & Krause 1910); De
Wildeman (1913c: 15; 1921a: 28). – Type: D.R.Congo, Lu-
bembe, 02 Feb. 1908, Kassner 2436 (holo-: B, iso-: BR, K).
Fig. 2D.
Chlorophytum vestitum sensu Nordal et al. (1997), Ka-
tivu et al. (2008).
Other representative specimens examined – D.R.Congo: XI: Ki-
popo, bord de route en forêt claire, 8 Feb. 1961, Schmitz 7010 (BR).
In a preliminary phylogenetic analysis the Congolese C.
vestitum subsp. pilosissimum [Meerts 2010/53 (BRLU)] re-
solves as a sister to the Zambian C. pubiorum Baker [Nor-
dal 4561 (O)] in a clade with species with distichous leaf ar-
rangement.
This taxon is referred to as C. vestitum in FZ (Kativu et
al. 2008) and FTEA (Nordal et al. 1997). However, the type
specimen of C. vestitum Baker from Mozambique (Kirk s.n.,
K, barcode K000256947) differs in having a peduncle < 10
cm and a simple or poorly ramied inorescence. We con-
sider C. pilosissimum to be conspecic (similar leaves, ow-
ers and fruits), though differing by a much longer peduncle
and a much more ramied panicle. Since the two morphs are
apparently not sympatric (the dwarf variant does not occur in
D.R.Congo), a subspecic rank seems appropriate.
406
Pl. Ecol. Evol. 145 (3), 2012
47. Chlorophytum warneckei (Engl.) Marais & Reilly (Ma-
rais & Reilly 1978: 662); Raadts (1984: 658). – Anthericum
warneckei Engl. (Engler 1902: 91); Hepper (1968b: 97);
Lisowski (2009: 421). – Type: Togo, prope Lome, Warnecke
304 (holo-: B (†?), iso-: P, BM) non Anthericum warneckei
sensu Vanden Berghen (1988: 420; g. 345bis).
Chlorophytum gracillimum Dammer (De Wildeman
1912a: 345), nomen nudum.
Other representative specimens examined – D.R.Congo: Savane
sud-occidentale du Congo, mission Cabra, 1899, Tilman s.n. (BR).
A W African species (Ghana, Guinea, Senegal, Togo), ap-
parently with disjunct populations in SW Congo, but the pre-
cise locality is not known. It has sometimes been confounded
with C. inconspicuum (Baker) Nordal, an E African species
which differs by bigger fruits, and a much reduced inores-
cence (Nordal et al. 2007).
48. Chlorophytum zingiberastrum Nordal & Poulsen (Nor-
dal et al. 1997: 48); Nordal & Poulsen (1998: 937; 938:
Fig. 1); Kativu et al. (2008: 73). – Type: Malawi, Mzimba
Dist., Mzuzu, Marymount, 25 Jan. 1974, Pawek 7987 (holo-:
WAG). Fig. 3F.
Other representative specimens examined – D.R.Congo: XI:
Keyberg, forêt claire sur sol rouge et frais, 5 Feb. 1948, Schmitz
1155 (BR); Kipopo, forêt claire, pied de termitière sur sol rouge,
Mar. 2010, Meerts 2010/63 (BRLU); Route Kolwesi-Kyamasumba,
rivière, Mushingi, forêt galerie, alt. 1010 m, Billiet & Jadin 4195
(BR; also cultivated in greenhouse: S 2063). Burundi: Butare, forêt
claire à Brachystegia sur le mont au N de la Nyakende, 6 Feb. 1952,
Michel & Reed 1113 (BR).
49. Chlorophytum sp. A “near comosum”
A most distinctive member of the C. sparsiorum-C. co-
mosum complex. Leaves rosulate, linear, 20–50(–75) cm ×
10–15(–25) mm, not blackening in herbarium. Inorescence
60–90(–150) cm (including peduncle), erect or decumbent;
peduncle with 1–2 foliaceous bracts up to 20 cm long; ow-
ers often replaced by rosettes of linear leaves (pseudovivi-
pary); owers 10 mm long; tepals minutely papillate on in-
ner side with 5–7 equally spaced nerves; anthers 3–4 mm,
shorter than lament (6 mm); lament scabrid distally and
attened at base. In swamps and marshes, generally in herba-
ceous vegetation.
Specimens examined – D.R.Congo: IX: Lac Kivu, forêt à l’W de
Tshibinda, près du marais Kanzibi, alt. 2000–2400 m, s.d., Humbert
7415 (BR). Rwanda: Forêt de Nyungwe, route Butare-Cyangugu,
sentier au km 100, bord de ruisseau en sous-bois, alt. 2000 m, plan-
te de 80–150 cm, eurs verdâtres, 21 Aug. 1969, Bouxin & Radoux
714 (BR); Territ. Shangugu, tourbière Kamiranzovu, km 85 route
Astrida–Bukavu, alt. 1950 m, Juncetum, 22 Aug. 1957, Deuse 1063
(BR); Territ. Shangugu, route Bukavu–Astrida, marais à Cyperus
latifolius, 23 Jul. 1959, A. Leonard 5098 (BR); Territ. Shangugu,
route Bukavu–Astrida, env. Uwinka, alt. 1900 m, marais, 2 Feb.
1959, Troupin 9707 (BR); Territ. Shangugu, Kamiranjovu, alt. 2000
m, fond de vallée, 22 Mar. 1956, Christiaensen 1514 (BR); Territ.
Cyangugu, forêt Nyungwe, env. Gisakura, alt. 2000 m, marais à
Syzygium, 12 Jun. 1971, Bouxin 968 (BR).
Burundi: Territ. Muramwya, Mont Manga, alt. 2300 m, bord de
ruisseau, 19 Nov. 1972, Reekmans 2078 (BR, BJA).
The C. comosum-C. sparsiorum complex is poorly un-
derstood (see comments under C. sparsiorum). Pending a
thorough taxonomic revision based upon molecular markers,
we prefer not to describe a new species.
50. Chlorophytum sp. B “near tetraphyllum”
A very small plant without a peduncle with long pedicel-
late, more or less umbellate owers. It closely ressembles C.
tetraphyllum (L.f.) Baker a species known only from Ethi-
opia, from which it differs by the non ciliate leaf margin.
However, more material is needed to decide on its taxonomic
status.
Specimens examined – D.R.Congo: XI: Kansenia, 8 Nov. 1958,
Schmitz 6220 (BR, KIP); Kansenia gare, dépressions herbeuses
sans géofrutex dans le steppe à géofrutex et Tephrosia et Crotalaria,
8 Dec. 1959, Duvigneaud 4467L (BRLU).
Uncertain species and names rejected
Chlorophytum huyghei De Wild. (De Wildeman 1909: 55,
Plate 1, gs. 3 & 4). – Type: D.R.Congo, Bokele, 1908,
Huyghe s.n. (BR, photograph); cultivated in greenhouse, 17
Mar. 1922, (BR, Herb. Plant Cult.: 314). – The diagnose and
the accompanying photographs suggest it may have been C.
sparsiorum.
Chlorophytum laurentii De Wild. in schedis. – This name
was handwritten by De Wildeman himself on a sheet (bar-
code BR0000008766946) with the indication “coll. E. Lau-
rent, cult. Gembloux 1901”. This sample is C. macrophyl-
lum. The name Chlorophytum laurentii does not seem to
have been published.
“Chlorophytum dewildemanianum Tournay”, cited in
Schmitz (1971). – Neither the type specimen nor the proto-
logue was found and this name is therefore a nomen nudum.
“Chlorophytum luembense Tournay”, cited in Schmitz
(1971). – Neither the type specimen nor the protologue was
found and this name is therefore a nomen nudum.
“Chlorophytum gracillimum Dammer” (De Wildeman
1912a: 345) and “Chlorophytum gracilimum VD.” [sic], cit-
ed in Germain (1952). – Names sometimes found in schedis
for slender, narrow-leaved forms of the C. cameronii com-
plex in E D.R.Congo and Burundi. Neither the protologue
nor the type specimen was found and this name is therefore a
nomen nudum. See note under C. warneckei.
Species excluded
Anthericum malosanum Baker, cited in Lebrun et al. (1948:
34) belongs in Trachyandra.
SUPPLEMENTARY DATA
Supplementary data are available at Plant Ecology and Evo-
lution, Supplementary Data Site (http://www.ingentacon-
nect.com/content/botbel/plecevo/supp-data), and consist of a
check list of taxa names (pdf format).
407
Meerts & Bjorå, Chlorophytum in Central Africa
ACKNOWLEDGEMENTS
This work was completed while P.M. was on sabbatical
leave. The Université Libre de Bruxelles (Fonds Cassel)
and FRS-FNRS (Belgium) are gratefully acknowledged for
nancial support, the Université de Lubumbashi for logis-
tic support, and the Jardin botanique national de Belgique
for hosting P.M. as a visiting scientist in BR. Inger Nordal
(Oslo) is gratefully acknowledged for her seminal contribu-
tion to Chlorophytum taxonomy, without which the present
work would not have been possible. P.M. is grateful to the
curators of B, BR, BRLU, BRVU, FT, K, KIP, LSHI and
especially to Odile Weber (K) and Sarah Bollendorf (B) for
their kind assistance. Frank Vankaekenberghe (BR) is grate-
fully acknowledged for SEM pictures. Frieda Billiet and
Julien Piqueray kindly provided some of the photographs.
Daniel Geerinck is gratefully acknowledged for fruitful dis-
cussion and advice. Ferdinand Nyiongabo kindly provided
living plants of Chlorophytum burundiense from Mt. Teza
(Burundi). Elmar Robbrecht and two anonymous referees
are gratefully acknowledged for their detailed comments.
This work is dedicated to the memory of Karl von Poellnitz
(1896–1945) who saved the Chlorophytum herbarium speci-
mens from destruction by the bombing raid on Berlin on 3d
March, 1943.
REFERENCES
Anonymous (1962) Report of the Systematics Association Com-
mittee for Descriptive Biological Terminology, II and IIa. Ter-
minology of simple symmetrical plane shapes (Charts 1, 1a).
Taxon 11: 145–155, 245–247.
http://dx.doi.org/10.2307/1216718
http://dx.doi.org/10.2307/1217034
APG III [The Angiosperm Phylogeny Group] (2009) An update
of the Angiosperm Phylogeny Group classication for the or-
ders and families of owering plants: AGP III. Botanical Jour-
nal of the Linnean Society 161: 105–121. http://dx.doi.org/
10.1111/j.1095-8339.2009.00996.x
Archer C., Kativu S. (2001) The correct name in Chlorophytum for
Anthericum longistylum. Bothalia 31: 40–41.
Baker J.G. (1875) The botany of the Speke and Grant expedition.
Monocotyledones Petaloideae. Transactions of the Linnean So-
ciety London 29: 151–163.
Baker J.G. (1876) Revision of the genera and species of Anth-
ericeae and Eriospermae. Journal of the Linnean Society, Bot-
any 15: 253–363. http://dx.doi.org/10.1111/j.1095-8339.1876.
tb00247.x
Baker J.G. (1878a) Report on the Liliaceae, Iridaceae, Hypoxy-
daceae and Haemodoraceae of Welwitsch’s Angolan Herbari-
um. Transactions of the Linnean Society, Series 2, 1: 245–273.
http://dx.doi.org/10.1111/j.1095-8339.1878.tb00040.x
Baker J.G. (1878b) Description of new and little known Liliaceae.
Journal of Botany, British and foreign 16: 321–326.
Baker J.G. (1893) New or noteworthy plants. Gardener’s chronicle
ser. 3, 13: 710–711.
Baker J.G. (1897a) Liliaceae. In Thiselton-Dyer W.T. (ed.), Flora
Capensis 6. Ashford, L. Reeve & Co.
Baker J.G. (1897b) 562. Chlorophytum oribundum. Diagnoses af-
ricanae, X. Bulletin of Miscellaneous Information, Royal Gar-
dens, Kew 128–129: 285.
Baker J.G. (1898) Liliaceae. In: Thiselton-Dyer W.T. (ed.) Flora of
Tropical Africa. Vol. 7: 421–568. London, Lovell Reeve.
Baker J.G. (1906) 797. Anthericum (Phalangium) recurvifolium.
Bulletin of Miscellaneous Information, Royal Botanic Gardens,
Kew 1906: 28.
Baker J.G., Engler A. (1892) Liliaceae africanae. Botanische Jahr-
bücher für Systematik, Panzengeschichte und Panzengeogra-
phie 15: 467–479.
Bentham G., Hooker J.D. (1883) Genera Plantarum 3(2). London,
Reeve & Co.
Bjorå C.S., Hoell, G. Kativu, S., Nordal, I. (2008) New taxa of
Chlorophytum (Anthericaceae) from Southern Tropical Africa
with notes on their sister group relationships. Botanical Journal
of the Linnean Society 157: 223–238. http://dx.doi.org/10.1111/
j.1095-8339.2008.00811.x
Bjorå C.S. (2008) Phylogeny, speciation and biogeography - a study
of Crinum and Chlorophytum (Asparagales) with focus on Afri-
can taxa. PhD thesis: University of Oslo, Oslo, Norway.
Bjorå C.S. & Nordal I. (2010) Anthericaceae. In Sosef M.S.M. et
al. (eds) Flore du Gabon 41: 1–10. Weikersheim, Margraf Pu-
blishers.
Compère P. (1962) Observations sur la ore du Bas-Congo. Bulle-
tin du Jardin botanique de l’État à Bruxelles 32(2): 199–211.
http://dx.doi.org/10.2307/3667229
Dammer U. (1912) Liliaceae africanae IV. Botanische Jahrbücher
für Systematik, Panzengeschichte und Panzengeographie 48:
360–366.
De Wildeman E. (1902) Etudes sur la ore du Katanga. Annales du
Musée du Congo Belge, Bot., sér.4, 1: 1–241.
De Wildeman E. (1904) Etudes de systématique et de géographie
botanique sur la ore du Bas- et du Moyen-Congo. Annales du
Musée du Congo, Botanique, sér. 5, vol. 1(2).
De Wildeman E. (1905–1906) Enumération des plantes récoltées
par Emile Laurent pendant sa dernière mission au Congo. I–III.
Bruxelles, Etat Indépendant du Congo.
De Wildeman E. (1909–1912) Etudes sur la ore du Bas- et du
Moyen-Congo. Annales du Musée du Congo, Botanique, série
5, 3: 1–147 (1909), 149–316 (1910a), 317–533 (1912a).
De Wildeman E. (1910b) Compagnie du Kasaï. Résultats de ses re-
cherches botaniques et agronomiques. Bruxelles, Lesigne.
De Wildeman E. (1911a) Decades novarum specierum Florae Con-
golensis. Bulletin du Jardin botanique de l’État à Bruxelles
3(2): 255–280. http://dx.doi.org/10.2307/3666486
De Wildeman E. (1911b) Plantae Thonnerianae congolenses. II.
Etudes sur la ore des districts des Bangala et de l’Ubangi.
Bruxelles, Misch & Thron.
De Wildeman E. (1912b) Plantae Thonnerianae Congolenses no-
vae. II. Repertorium novarum specierum regni vegetabilis 10:
523–529.
De Wildeman E. (1913a) Decades novarum specierum orae katan-
gensis. I–VII. Repertorium novarum specierum regni vegetabi-
lis 11: 501–524.
De Wildeman E. (1913b) Decades novarum specierum orae katan-
gensis. XII–XIV. Repertorium novarum specierum regni veg-
etabilis 12: 289–298.
De Wildeman E. (1913c) Etudes sur la ore du Katanga. Annales du
Musée du Congo, Botanique, Série IV, vol. II, 1: 1–180.
De Wildeman E. (1914) Decades novarum specierum orae katan-
gensis. XV–XVIII. Repertorium novarum specierum regni veg-
etabilis 13(6): 103–117. http://dx.doi.org/10.2307/3666515
408
Pl. Ecol. Evol. 145 (3), 2012
De Wildeman E. (1915) Decades Novarum Specierum Florae Con-
golensis. Bulletin du Jardin botanique de l’État a Bruxelles
5(1): 1–108. http://dx.doi.org/10.2307/3666484
De Wildeman E. (1916) Additions à la Flore du Congo. Bulletin
du Jardin botanique de l’État à Bruxelles 5(2): 117–268. http://
dx.doi.org/10.2307/3666538
De Wildeman E. (1921a) Contribution à l’étude de la ore du Ka-
tanga. Bruxelles, Comité spécial du Katanga.
De Wildeman E. (1921b) Plantae Bequaertianae. Gand, Hoste.
De Wildeman E. (1930) Contribution à la ore du Katanga. Supplé-
ment III. Bruxelles, Comité spécial du Katanga.
De Wildeman E., Durand Th. (1899) Contribution à la ore du
Congo. Annales du Musée du Congo. Botanique. Série 2: 1–95.
De Wildeman E., Durand Th. (1901) Reliquiae Dewevreanae. An-
nales du Musée du Congo. Botanique. Série III, 2: 81–291.
Drummond R.B. (1953) Tropical African Plants XXIII. Chlorophy-
tum arcuatoramosum. Kew Bulletin 8: 120.
Durand Th., Durand H. (1909) Sylloge Florae Congolanae [Pha-
nerogamae]. Bulletin du Jardin botanique de l’État à Bruxelles
2: 1–716. http://dx.doi.org/10.2307/3666516
Duvigneaud P., Denaeyer-De Smet S. (1963) Cuivre et végétation
au Katanga. Bulletin de la Société Royale de Botanique de
Belgique 96: 92–231.
Engler A. (1892) (“1891”) Über die Hochgebirgsora des tropi-
schen Afrika. Abhandlungen der Königlichen Akademie der
Wissenschaften zu Berlin. Phys.-Math. Klasse 1891(2): 1–461.
Engler A. (1895) Die Panzenwelt Ost-Afrikas und der Nachbarge-
biete. Teil C. Berlin, Dietrich Reimer.
Engler A. (1900) Die von W. Goetze und Dr. Stuhlmann im Ulu-
gurugebirge, sowie die von W. Goetze in der Kisaki- und Khu-
tu-Steppe und in Uhehe gesammelten Panzen. Botanische
Jahrbücher für Systematik, Panzengeschichte und Panzen-
geographie 28: 332–510.
Engler A. (1901) Berichte über die botanischen Ergebnisse der
Nyassa-See- und Kinga-Gebirgs-Expedition der Hermann- und
Elise- geb. Heckmann-Wentzel-Stiftung. IV. Botanische Jahr-
bücher für Systematik, Panzengeschichte und Panzengeogra-
phie 30: 39–445.
Engler A. (1902) Liliaceae Africanae II. Botanische Jahrbücher für
Systematik, Panzengeschichte und Panzengeographie 32:
89–126.
Engler A., Gilg E. (1903) Liliaceae. In: Warburg O. (ed.) Kunene-
Sambesi Expedition H. Baum: 185–197. Berlin, Kolonial Wirt-
schaftlichen Komitees.
Engler A. (1908) Die Panzenwelt Afrikas insbesondere seiner tro-
pischen Gebiete. II. Leipzig, Engelmann.
Engler A., Krause K. (1910) Liliaceae africanae 2. Botanische Jahr-
bücher für Systematik, Panzengeschichte und Panzengeogra-
phie 45: 123–155.
Faucon M.-P., Meersseman A., Ngoy Shutcha M., Mahy G., Ngongo
Luhembwe M., Malaisse F., Meerts P. (2010) Copper endemism
in the Congolese ora: a database of copper afnity and conser-
vational value of cuprophytes. Plant Ecology and Evolution 1:
5–18. http://dx.doi.org/10.5091/plecevo.2010.411
Germain R. (1952) Les associations végétales de la plaine de la
Ruzizi (Congo belge) en relation avec le milieu. Bruxelles, In-
stitut National pour l’Etude Agronomique du Congo, Ser. Sci.,
52.
Hanid M.A. (1974) The identity of Chlorophytum afne (Liliaceae)
and its range of variation in Africa. Kew Bulletin 29: 585–591.
http://dx.doi.org/10.2307/4108001
Hepper C.M. (1968a) Notes on Tropical African Mono-
cotyledons: II. Kew Bulletin 22: 449–467. http://dx.doi.
org/10.2307/4108346
Hepper C.M. (1968b) Liliaceae. In: Hepper F.N. (ed.) Flora of West
Tropical Africa, vol. 3: 90–107. London, Crown Agents for
Overseas Governments and Administrations.
Kativu S. (1993) Four new species of Chlorophytum (Antherica-
ceae) from Tropical Africa. Nordic Journal of Botany 13: 501–
505. http://dx.doi.org/ 10.1111/j.1756-1051.1993.tb00090.x
Kativu S., Nordal J. (1993) New combinations of African species in
the genus Chlorophytum. Nordic Journal of Botany 13: 59–65.
http://dx.doi.org/10.1111/j.1756-1051.1993.tb00016.x
Kativu S. (1994) Synopsis of Chlorophytum (Anthericaceae) in the
Flora Zambesiaca area. Kirkia 15: 43–111.
Kativu S., Hoell G., Bjorå C.S., Nordal J. (2008) Anthericaceae. In:
Timberlake J.F. (ed.) Flora Zambesiaca 13(1). Kew, Royal Bo-
tanic Gardens.
Lebrun J. (1947) La végétation de la plaine alluviale au sud du lac
Edouard. Exploration du Parc National Albert, Mission J. Leb-
run (1937–1938), vol. 1. Bruxelles, Institut des Parcs nationaux
du Congo belge.
Lebrun J., Taton A., Toussaint L. (1948) Contribution à l’étude de
la ore du Parc National de la Kagera. Bruxelles, Institut des
Parcs nationaux du Congo belge.
Lewalle J. (1972) Les étages de végétation du Burundi occidental.
Bulletin du Jardin botanique national de Belgique 42: 1–247.
http://dx.doi.org/10.2307/3667406
Lindley J. (1824) Chlorophytum orchidastrum. The Botanical Reg-
ister 10: 813.
Lisowski S. (2009) Flore (Angiospermes) de la République de
Guinée. Meise, Jardin botanique national de Belgique.
Malaisse F., Bamps P. (2009) Chlorophytum unifolium (Antheri-
caceae), espèce nouvelle du Katanga (Rép. Dém. Congo). Sys-
tematics and Geography of Plants 79: 225–228.
Marais W., Reilly J. (1978) Chlorophytum and its related gen-
era (Liliaceae). Kew Bulletin 32: 653–663. http://dx.doi.
org/10.2307/4109671
Meerts P. (2011) Chlorophytum burundiense (Asparagaceae), a new
species from Burundi and Tanzania. Plant Ecology and Evolu-
tion 144: 233–236. http://dx.doi.org/10.5091/plecevo.2011.609
Myers N., Mittermeier R.A., Mittermeier C.G., da Fonseca G.A.B.,
Kent J. (2000) Biodiversity hotspots for conservation priorities.
Nature 403: 853–858. http://dx.doi.org/10.1038/35002501
Naik V.N., Nirgude S.M. (1981) Vessels in Chlorophytum (Li-
liaceae). Adansonia 2: 201–212.
Nordal I., Eriksen T.E., Fosby M. (1990) Studies on the generic de-
limitation of Anthericaceae. Mitteilungen Institut Allgemeine
Botanik Hamburg 23b: 535–559.
Nordal I., Kativu S., Poulsen A.D. (1997) Anthericaceae. In: Polhill
R.M. (ed.) Flora of Tropical East Africa. Rotterdam, A.A. Bal-
kema.
Nordal I., Thulin M. (1993) Synopsis of Anthericum and Chloro-
phytum (Anthericaceae) in the Horn of Africa, including the
description of nine new species. Nordic Journal of Botany
13: 257–280. http://dx.doi.org/10.1111/j.1756-1051.1993.
tb00047.x
Nordal I. (1997) 6. Hydrocharitaceae to Arecaceae. In: Edwards S.,
Demissew S., Hedberg I. (eds) Flora of Ethiopia and Erithrea.
Addis Ababa & Uppsala, the National Herbarium, Addis Ababa
University.
409
Meerts & Bjorå, Chlorophytum in Central Africa
Nordal I., Poulsen A.D. (1998) Chlorophytum zingiberastrum, a
new species from South-Eastern Africa. Kew Bulletin 53: 937–
942. http://dx.doi.org/10.2307/4118881
Obermeyer A.A. (1962) A revision of the South African species of
Anthericum, Chlorophytum and Trachyandra. Bothalia 7: 669–
767.
Panigrahi G. (1975) Chlorophytum orchidastrum sens. lat. (Lilia-
ceae) from Africa & Asia. Kew Bulletin 30: 563–567. http://
dx.doi.org/10.2307/4103087
Poellnitz K. von (1941) Zur Kenntnis der Gattung Anthericum L.
Repertorium novarum specierum regni vegetabilis 50: 232.
Poellnitz K. von (1942) Die Anthericum-Arten Deutsch-Ost-
Afrikas. Repertorium Novarum Specierum Regni Vegetabilis
51: 8–15, 17–32, 66–80.
Poellnitz K. von (1943a) Die Anthericum-Arten Angolas. Boletim
da Sociedade Broteriana. XVII 2A serie: 55–91.
Poellnitz K. von (1943b) Zur Kenntis der Gattung Chlorophytum
Ker. Berichte der Deutschen Botanischen Gesellschaft 61: 126–
131.
Poellnitz K. von (1945) Neue Chlorophytum-Arten Africas. Portu-
galiae Acta Biologica. Série B, 1(1–2): 221–233.
Poellnitz K. von (1946) Die Chlorophytum-Arten Tanganyikas.
Portugaliae Acta Biologica série B, 1(3–4): 255–383.
Poulsen A.D., Nordal I. (1999) Two new species of Chlorophytum
from Central Africa. Kew Bulletin 54: 941–949. http://dx.doi.
org/10.2307/4111172
Poulsen A.D., Nordal I. (2005) A phenetic analysis and revi sion
of Guineo-Congolian rain forest taxa of Chlorophytum (Anth-
ericaceae). Botanical Journal of the Linnean Society 148: 1–20.
http://dx.doi.org/10.1111/j.1095-8339.2005.00386.x
Raadts E. (1984) Liliaceae. In: J.Brunel J.F., Hiepko P., Scholz
H. (eds) Flore Analytique du Togo: Phanérogames. Englera 4:
3–751.
Rendle A.B. (1895) A Contribution to the Flora of Eastern Tropi-
cal Africa. Journal of the Linnean Society of London, Botany
30: 373–435. http://dx.doi.org/10.1111/j.1095-8339.1894.
tb02417.x
Rendle A.B. (1899) Catalogue of the African plants collected by
Dr. Friedrich Welwitsch in 1853–1861. II. Monocotyledons and
Gymnosperms. London, Trustees of the British Museum.
Richard A. (1851) Tentamen Florae Abyssinicae, vol. 2. Paris, A.
Bertrand.
Robyns W. (1948) Atlas Général du Congo: les territoires phytogéo-
graphiques du Congo belge et du Ruanda-Urundi. Bruxelles,
Institut Royal Colonial Belge, No. 410.
Robyns W., Tournay R. (1955) Flore des Spermatophytes du Parc
National Albert. III. Monocotylées. Bruxelles, Institut des Parcs
nationaux du Congo belge.
Schmitz A. (1971) La végétation de la Plaine de Lubumbashi (Haut-
Katanga). Institut national pour l’étude agronomique du Congo,
sér. scientif. 113. Bruxelles, INEAC.
Schweinfurth G. (1867) Beiträge zur Flora Aethiopiens. Berlin,
Reimer.
Troupin G. (1955) Plantae Africanae IV. Monocotyledoneae. Bul-
letin du Jardin botanique de l’État à Bruxelles 25(3): 221–237.
http://dx.doi.org/10.2307/3667056
Troupin G. (1956) Flore des Spermatophytes du Parc National de
la Garamba. I. Gymnospermes et Monocotylédones. Bruxelles,
Institut des Parcs Nationaux du Congo belge.
Troupin G. (ed.) (1988) Flore du Rwanda, vol. 4. Tervuren, Musée
Royal d’Afrique Centrale.
Vanden Berghen C. (1988) Flore illustrée du Sénégal. Monocotylé-
dones et Ptéridophytes. Tome 9. Monocotylédones: Agavacées
à Orchidacées. Dakar, Gouvernement du Sénégal.
Manuscript received 25 Apr. 2011; accepted in revised version 12
Jan. 2012.
Communicating Editor: Elmar Robbrecht.
