Bradleya 29/2011
pages 79 – 92
The genus Cyphostemma (Planch.) Alston (Vitaceae) in Angola
Filipe de Sousa1, Estrela Figueiredo2 and Gideon F. Smith3
1
Department of Plant and Environmental Sciences, University of Gothenburg, Box 461, SE-40530 Göteborg,
Sweden (email: filipedeportugal@gmail.com) (corresponding author).
2
Department of Botany, P.O.Box 77000, Nelson Mandela Metropolitan University, Port Elizabeth, 6031
South Africa / Centre for Functional Ecology, Departamento de Ciências da Vida, Universidade de Coimbra,
3001-455 Coimbra, Portugal (email: estrelafigueiredo@hotmail.com).
3
Office of the Chief Director: Biosystematics Research & Biodiversity Collections, South African National
Biodiversity Institute, Private Bag X101, Pretoria, 0001 South Africa / H.G.W.J. Schweickerdt Herbarium,
Department of Plant Science, University of Pretoria, Pretoria, 0002 South Africa and Centre for Functional
Ecology, Departamento de Ciências da Vida, Universidade de Coimbra, 3001-455 Coimbra, Portugal
(email: g.smith@sanbi.org.za).
Summary: An overview of the 22 taxa recorded in
the genus Cyphostemma (Planch.) Alston
(Vitaceae) in Angola is presented.
An
identification key to all taxa recorded is provided,
together with a referenced list of taxa with
synonymy, geographical distribution range,
endemic status and the citation of type specimens
that originated from the country. Distribution
maps are also presented. A list of herbarium
collections consulted is given in an appendix.
Zusammenfassung: Es wird eine Übersicht über
die 22 aus Angola bekannten Taxa der Gattung
Cyphostemma (Planch.) Alston (Vitaceae)
gegeben. Der Beitrag besteht aus einem Schlüssel
zu allen Taxa, sowie einer referenzierten
Taxonliste mit Synonymen, geographischer
Verbreitung (inkl. Verbreitungskarten), Angaben
zum Endemiestatus und Zitaten der Typusbelege,
soweit sie aus dem Land stammen. Ein Anhang
listet die untersuchten Herbarbelege auf.
Introduction
The family Vitaceae, which includes Vitis vinifera
L., the source of table and wine grapes, consists of
just over 700 species and 13 genera, making this
a rather small family on global standards (Mabberley, 2008). Most species are woody vines with
branched tendrils, but large, fat-stemmed species
are also included in some genera. Flowers are
rather small and insignificant, while the leaves
are generally simple and deeply invaginated, or
compound, and known for their striking autumn
colours. The fruit of some representatives of the
Vitaceae can be poisonous. The Vitaceae in An-
Bradleya 29/2011
gola were treated for Conspectus florae angolensis
(Exell & Mendonça, 1954) and more recently enumerated for Plants of Angola by Retief (2008).
These treatments recognised five Vitaceae genera
occurring naturally in the country [Ampelocissus
Planch., Cayratia Juss., Cissus L., Cyphostemma
(Planch.) Alston and Rhoicissus Planch.], while
Vitis vinifera L. was recorded as having escaped
from cultivation.
The genus Cyphostemma
(Planch.) Alston, previously included in Cissus L.
as sect. Cyphostemma Planch. (Planchon, 1887),
was elevated to the rank of genus by Alston
(1931), a classification adopted by Descoings
(1960). Descoings (1960) made several combinations in Cyphostemma that were invalid because
the basionym was not clearly cited (ICBN Art.
33.2). Later, he published again the combinations
(Descoings, 1967) in order to validate them, but
meanwhile Wild & Drummond (1966) had already
unintentionally validated some of the names,
therefore some names in Descoings’ second set of
combinations are also not accepted (being superfluous names). Cyphostemma comprises c. 150
species (Mabberley, 2008) and is widely distributed throughout tropical Africa. In Angola it is
represented by 22 species, of which nine are endemic. Representatives of Cyphostemma differ
from species included in the related genus Cissus,
through having constricted flower buds and a nectary disk of four glands, which are free from each
other (Descoings, 1960). The majority of species
are erect, prostrate or climbing perennial herbs or
shrubs, with or without tendrils (Wild & Drummond, 1966). Several species of southern African
Cyphostemma are thick-stemmed succulents or
79
have succulent leaves, giving them horticultural
interest among collectors (Smith et al., 1997;
Figueiredo & Smith, 2009).
Material and methods
The collections of Cyphostemma from Angola kept
at the Herbaria of the Centro de Botânica (LISC)
and University of Lisbon (LISU) in Lisbon,
Portugal, and the National Herbarium of South
Africa (PRE), in Pretoria, were examined. The
Herbarium of the University of Coimbra (COI),
Coimbra, Portugal, was contacted for permission
to examine its specimens, but according to its
curator it does not hold any collections of
Cyphostemma from Angola. However, many
specimens from COI are cited in the literature and
types of Angolan Cyphostemma names from that
herbarium
are
accessible
in
JSTOR
(http://plants.jstor.org). These and other types
were therefore examined on-line in JSTOR. The
type material of names established in
Cyphostemma for species that occur in Angola is
indicated in the text, and for those specimens the
herbarium barcode is given to facilitate searches
using the JSTOR facility. Distribution maps were
made using ArcGIS 9.3 from ESRI. Specimens
cited in the literature that we did not examine
were included in the maps using different symbols
(open symbols). The full list of specimens used to
prepare these maps is presented in Appendix 1.
Results
Twenty-two taxa of Cyphostemma are currently
known to occur in Angola, following the recent
recording of C. wittei (Staner) Wild &
R.B.Drumm. and C. congestum (Baker) Desc. ex
Wild & R.B.Drumm. as part of the flora of the
country (Sousa et al., 2011). Two further possible
records remain to be investigated when more
material is available.
List of taxa
H = Habit; D = Distribution by province
(provinces: BO–Bengo, BE–Benguela, BI–Bié,
CA–Cabinda, CC–Cuando-Cubango, CN–Cuanza
Norte, CS–Cuanza Sul, CU–Cunene, HA–
Huambo, HI–Huíla, LA–Luanda, LN–Lunda
Norte, LS–Lunda Sul, MA–Malange, MO–Moxico,
NA–Namibe, UI–Uíge, ZA–Zaire).
1. Cyphostemma adenocaule (Steud. ex
A.Rich.) Desc. ex Wild & R.B.Drumm., Fl.
Zamb. 2: 473 (1966) subsp. adenocaule.
Retief in Figueiredo & Smith, Plants of
80
Angola: 169 (2008). Type from Ethiopia.
Cissus adenocaulis Steud. ex A.Rich., Tent. Fl.
Abyss. 1: 111 (1847). Exell & Mendonça,
C.F.A. 2(1): 66 (1954).
Cyphostemma adenocaule (Steud. ex A.Rich.)
Desc., Notul. Syst. (Paris) 16: 120 (1960),
comb. inval.
H: Climbing herb with tuberous rootstock. D:
BE, BO, CA, CN, CS, HI, LA, MA, NA, ZA.
(Figure 1.)
2. Cyphostemma adenopodum (Sprague)
Desc., Nat. Monspel., Sér. Bot. 18: 218 (1967).
Retief in Figueiredo & Smith, Plants of
Angola: 169 (2008). Type from Uganda.
Cissus adenopoda Sprague, Bull. Misc. Inf. Kew
1906: 247 (1906). Exell & Mendonça, C.F.A.
2(1): 62 (1954).
Cyphostemma adenopodum (Sprague) Desc.,
Notul. Syst. (Paris) 16: 120 (1960), comb. inval.
H: Climbing herb to several metres long from a
tuberous rootstock. D: CA. (Figure 1.)
3. Cyphostemma chloroleucum (Welw. ex
Baker) Desc. ex Wild & R.B.Drumm., Fl.
Zamb. 2,2: 466 (1966). Retief in Figueiredo &
Smith, Plants of Angola: 169 (2008).
Figueiredo et al., Bothalia 39(2): 201 (2009).
Type: Angola, entre Lago Ivantala e
Quilengues,
Welwitsch
1453
(BMBM000838246,
LISU-LISU208728
&
LISU208730 & LISU208729).
Vitis chloroleuca Welw. ex Baker in Oliver, Fl.
Trop. Afr. 1: 406 (1868).
Cissus chloroleuca (Welw. ex Baker) Planch. in
A. & C. De Candolle, Mon. Phan. 5(2): 592
(1887); Exell & Mendonça, C.F.A. 2(1): 60
(1954).
Cyphostemma chloroleucum (Welw. ex Baker)
Desc., Notul. Syst. (Paris) 16: 120 (1960),
comb. inval.
Vitis andongensis Welw. ex Baker in Oliver, Fl.
Trop. Afr. 1: 406 (1868).
Cyphostemma andongensis (Welw. ex Baker)
Desc., Nat. Monspel., Sér. Bot. 18: 218 (1967).
Type: Angola, Pungo Andongo, Pedra de Cambondo,
Welwitsch
1463
(B†?,
BMBM000838248 & BM000838247, COI-COI00005280, K, LISU-LISU208726 & LISU208727).
Cissus hypargyrea Gilg in Warburg, KuneneSambesi-Expedition: 293 (1903).
Cyphostemma hypargyreum (Gilg) Desc., Nat.
Monspel., Sér. Bot. 18: 222 (1967).
Bradleya 29/2011
Key to the species of Cyphostemma recorded for Angola
1.
−
2.
−
3.
−
4.
−
5.
−
6.
−
7.
−
8.
−
9.
−
10.
−
11.
−
12.
−
13.
−
14.
−
15.
−
16.
−
17.
−
18.
−
19.
−
20.
−
21.
−
Tendrils absent ...................................................................................................................................2
Tendrils present................................................................................................................................13
Plants with perennial, often massively thickened, succulent stem and papery bark.....................3
Plants without a thickened, succulent stem and papery bark .........................................................5
Stem squat, sub-spherical; leaflets sub-orbicular...................................................................C. uter
Stem elongated, branched; leaflets not as above ..............................................................................4
Leaflets 5; leaf margins serrate ......................................................................................C. mendesii
Leaflets 3; leaf margins crenate .......................................................................................C. currorii
Leaves simple, sessile or sub-sessile ....................................................................................C. wittei
Leaves compound, usually petiolate ..................................................................................................6
Leaves pinnate.........................................................................................C. violaceo-glandulosum
Leaves digitate ....................................................................................................................................7
Leaflets oblong-lanceolate to linear, glabrous, glabrescent or pubescent .......................................8
Leaflets cordate-ovate, obovate-elliptical to oblanceolate, never glabrous....................................11
Stipules large (up to 22 × 16 mm), cordate, amplexicaul .............................C. grandistipulatum
Stipules lanceolate-acuminate ...........................................................................................................9
Stems and leaves glabrous, stipules usually 2 or more cm long; inflorescence 10–20 cm wide ......
.......................................................................................................C. junceum subsp. jatrophoides
Stems and leaves not glabrous, stipules up to 1.5 cm long; inflorescence up to 6 cm wide..........10
Distal leaves sessile; leaflets serrate-crenate, usually with glandular hairs on the nerves abaxially; inflorescence smaller than subtending leaf.......................................................C. stenolobum
Distal leaves petiolate; leaflets entire to serrate-dentate, sub-glabrous; inflorescence up to 5 cm
wide ..............................................................................................................................C. fugosioides
Plant densely pilose-glandulose from the base...............................................................C. pruriens
Plant pubescent but never densely pilose-glandulose ....................................................................12
Leaves densely ferruginous-tomentose below; inflorescence branches glandular ...........................
.................................................................................................................................C. chloroleucum
Leaves never densely ferruginous-tomentose below; inflorescence branches quite eglandular.......
..............................................................................................................................................C. luteum
Leaves pedately arranged ..........................................................................................C. adenocaule
Leaves digitately arranged...............................................................................................................14
Leaves sessile or sub-sessile (petiole up to 0.5 cm long).................................................................15
Leaves distinctly petiolate................................................................................................................17
Inflorescence branches, calyx and corolla glabrous or puberulous; leaflets ovate-elliptic, up to 5.5
× 2 cm .........................................................................................................................C. subciliatum
Inflorescence branches, calyx and corolla densely white pubescent; leaflets elliptic-lanceolate, up
to 14 × 5 cm .......................................................................................................................................16
Leaflets glabrous, up to 10 × 3 cm, undulate to slightly serrate..................................C. huillense
Leaflets puberulous to pubescent, up to 14 × 5 cm, distinctly dentate to serrate ....C. congestum
Stem densely covered with glandular hairs to 5 mm long ..........................................C. pendulum
Stem not as above .............................................................................................................................18
Leaflets broadly elliptic to sub-orbicular, apex rounded ..........................................C. ruacanense
Leaflets ovate, elliptic or lanceolate, apex acute to acuminate......................................................19
Leaflets 3 ..................................................................................................................C. adenopodum
Leaflets 5 ...........................................................................................................................................20
Inflorescence branches eglandular, densely white-pubescent.......................................C. johannis
Inflorescence branches glandular, densely to sparsely pubescent.................................................21
Flower buds glandular; petioles to 15 cm long; leaves pubescent on nerves below; stipules quite
large, suborbicular-acuminate, up to 25 × 20 mm...................................................C. stipulaceum
Flower buds eglandular; petioles to 5 cm long; leaves densely pubescent to tomentose below;
stipules inconspicuous ..........................................................................................C. crassiusculum
Bradleya 29/2011
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Type:Angola, Kubango, Kabindere, 1150 m,
1899/11/2, Baum 361 (B† holo-; BM, COICOI00005281,
G-G00023853,
K,
WW19010006645 iso-).
Cissus decurrens Gilg & M.Brandt, Bot. Jahrb.
Syst. 46: 452, 504 (1911). Type: Angola, Chibemba (Gambos), Newton 230 (B†? holo-).
H: Herb up to 60 cm high. D: BE, CN, CS, CU, HI,
MA. (Figure 2.)
4. Cyphostemma congestum (Baker) Desc. ex
Wild & R.B.Drumm., Fl. Zamb. 2(2): 473
(1966). Sousa et al., Bothalia 41(2): 260
(2011). Type from Nyasaland.
Vitis congesta Baker in Oliver, Fl. Trop. Afr. 1:
412 (1868).
Cyphostemma congestum (Baker) Desc., Notul.
Syst. (Paris) 16: 120 (1960), comb. inval.
H: Climbing herb with tuberous root. D: CU.
(Figure 2.)
5. Cyphostemma crassiusculum (Baker)
Desc., Nat. Monspel., Sér. Bot. 18: 220 (1967).
Retief in Figueiredo & Smith, Plants of Angola: 169 (2008). Type: Angola, Loanda, entre
Penedo e Conceição, Welwitsch 1495 (BMBM000838266, COI-COI00005288, K, LISULISU208737 & LISU208738).
Vitis crassiuscula Baker in Oliver, Fl. Trop. Afr. 1:
406 (1868).
Cissus crassiuscula (Baker) Planch., Monogr.
Phan. 5: 608 (1887). Exell & Mendonça,C.F.A.
2(1): 68 (1954).
Cyphostemma crassiusculum (Baker) Desc.,
Notul. Syst. (Paris) 16: 120 (1960), comb. inval.
Cissus loandensis Gilg & M.Brandt, Bot. Jahrb.
Syst. 46: 537 (1912).
Cyphostemma loandense (Gilg & M.Brandt) Desc.,
Nat. Monspel., Sér. Bot. 18: 224 (1967). Type:
Angola, Luanda, Gossweiler 294 (B†? lectodesignated by Exell & Mendonça (1954: 68).
H: Climber. D: BE, BO, CN, LA, MA, NA.
Endemic. (Figure 3.)
6. Cyphostemma currorii (Hook.f.) Desc., Nat.
Monspel., Sér. Bot. 18: 220 (1967). Retief in
Figueiredo & Smith, Plants of Angola: 169
(2008). Type: Moçâmedes, Baía dos Elefantes,
Curror s.n. (B†? iso-, K holo-).
Cissus currorii Hook.f., Niger Fl.: 265 (1849).
Exell & Mendonça, C.F.A. 2(1): 70 (1954).
Vitis currorii (Hook.f.) Baker in Oliver, Fl. Trop.
Afr. 1: 401 (1868).
Cyphostemma currorii (Hook.f.) Desc., Notul. Syst.
(Paris) 16: 121 (1960), comb. inval.
Cissus macropus Welw., J. Linn. Soc., Bot. 8: 77 (1864).
Figure 1. Geographical distribution map of C. adenocaule (Steud. ex A.Rich.) Desc. ex Wild &
R.B.Drumm., C. adenopodum (Sprague) Desc. and C. huillense (Exell & Mendonça) Desc. in Angola.
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2
3
Figure 2. Geographical distribution map of C. chloroleucum (Welw. ex Baker) Desc. ex Wild &
R.B.Drumm., C. congestum (Baker) Desc. ex Wild & R.B.Drumm., C. johannis (Exell & Mendonça) Desc.
and C. mendesii F.Sousa in Angola. Figure 3. Geographical distribution map of C. crassiusculum (Baker)
Desc., C. currorii (Hook.f.) Desc. and C. fugosioides (Gilg) Desc. ex Wild & R.B.Drumm. in Angola.
Bradleya 29/2011
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Vitis macropus (Welw.) Baker in Oliver, Fl. Trop.
Afr. 1: 405 (1868).
Cyphostemma macropus (Welw.) Desc., Notul. Syst.
(Paris) 16: 122 (1960), comb. inval.
Cyphostemma macropus (Welw.) Desc., Nat.
Monspel., Sér. Bot. 18: 224 (1967). Type: Angola,
Moçâmedes, Welwitsch 1446 (BM, LISULISU208739 & LISU208740 & LISU208741).
H: Pachycaul trees to 6 m tall, along desert
coastline. D: BE, NA. (Figures 3–6.)
7. Cyphostemma fugosioides (Gilg) Desc. ex
Wild & R.B.Drumm., Fl. Zamb. 2(2): 469
(1966). Retief in Figueiredo & Smith, Plants of
Angola: 169 (2008). Figueiredo et al., Bothalia
39(2): 201 (2009). Type: Angola, Chitanda,
1150 m, 1899/9/27, Baum 174 (B† holo-; BM,
COI-COI00005289, G-G00023854, K, MM0109127, W-W19010009447 iso-).
Cissus fugosioides Gilg in Warburg, KuneneSambesi-Expedition: 294 (1903); Exell & Mendonça, C.F.A. 2(1): 57 (1954).
Cyphostemma fugosioides (Gilg) Desc., Notul.
Syst. (Paris) 16: 121 (1960), comb. inval.
H: Herb up to 50 cm tall, with a tuberous rootstock. D: BE, CU, HI. (Figure 3.)
8. Cyphostemma grandistipulatum (Gilg &
M.Brandt) Desc., Nat. Monspel., Sér. Bot. 18:
222 (1967). Retief in Figueiredo & Smith,
Plants of Angola: 169 (2008). Type: Angola,
Huíla, Antunes 110 (B†? lecto- designated by
Exell & Mendonça (1954: 56)).
Cissus grandistipulata Gilg & M.Brandt, Bot.
Jahrb. Syst. 46: 491 (1912). Exell & Mendonça, C.F.A. 2(1): 56 (1954).
Cyphostemma
grandistipulatum
(Gilg
&
M.Brandt) Desc., Notul. Syst. (Paris) 16: 121
(1960), comb. inval.
H: Succulent? herb up to 90 cm high. D: HI.
(Figure 7.)
9. Cyphostemma huillense (Exell & Mendonça)
Desc., Nat. Monspel., Sér. Bot. 18: 222 (1967).
Retief in Figueiredo & Smith, Plants of
Angola: 169 (2008). Type: Angola, Huíla, Morros de Cualeque [Calueque], 1000−1100 m,
1937, Exell & Mendonça 2708 (BM holo-; COICOI00005292 & COI00005291, LISCLISC001323, iso-).
Cissus huillensis Exell & Mendonça, Bol. Soc.
Brot. 26: 229 (1952). Exell & Mendonça,
C.F.A. 2(1): 62 (1954).
H: Climber. D: CU. Endemic. (Figure 1.)
10. Cyphostemma johannis (Exell & Mendonça)
Desc., Nat. Monspel., Sér. Bot. 18: 222 (1967).
84
Retief in Figueiredo & Smith, Plants of
Angola: 169 (2008). Type: Angola, Cabinda,
Maiombe, Buco Zau, 1916/10, Gossweiler 7238
(BM-BM000838236
p.p.
holo-;
LISCLISC001325 & LISC001324, LISU p.p. iso-).
Cissus johannis Exell & Mendonça, Bol. Soc. Brot.
26: 231 (1952). Exell & Mendonça, C.F.A. 2(1):
64 (1954).
Cyphostemma johannis (Exell & Mendonça) Desc.,
Notul. Syst. (Paris) 16: 122 (1960), comb.inval.
H: Climber. D: CA. Endemic. (Figure 2.)
11. Cyphostemma junceum (Webb) Desc. ex
Wild & R.B. Drumm., Fl. Zamb. 2(2): 465
(1966) subsp. jatrophoides (Welw. ex Baker)
Verdc., Fl. Trop. E. Africa Vitac.: 62 (1993).
Retief in Figueiredo & Smith, Plants of Angola: 169 (2008). Type: Angola, Ambaca, entre
os Rios Lutete e Zamba, Welwitsch 1474 (B†?,
BM-BM000838275, COI-COI00005293, LISULISU208718 & LISU208719 & LISU208720 &
LISU208721).
Vitis jatrophoides Welw. ex Baker in Oliver, Fl.
Trop. Afr. 1: 400 (1868) pro parte excl. spec
Barter.
Cissus jatrophoides (Welw. ex Baker) Planch. in
A. & C. De Candolle, Mon. Phan. 5(2): 579
(1887); Exell & Mendonça, C.F.A. 2(1): 57
(1954).
Cyphostemma jatrophoides (Welw. ex Baker)
Desc., Notul. Syst. (Paris) 16: 122 (1960),
comb. inval.
Cyphostemma jatrophoides (Welw. ex Baker)
Desc., Nat. Monspel., Sér. Bot. 18: 226 (1967).
Cissus
chlorantha
Gilg
in
Warburg,
Kunene-Sambesi-Expedition: 295 (1903).
Type: Angola, Kubango, Chirumbu, 1200 m,
1899/10/14, Baum 278 (B† holo-; GG00023856, W-W19010009245 iso-).
H: Herb up to 1.3 m high, with a tuberous rootstock.
D: BE, BI, CN, CU, HA, HI, LS. (Figure 8.)
Note: Welwitsch 1474 (K) is Cissus petiolata
Hook.f.
12. Cyphostemma luteum (Exell & Mendonça)
Desc., Nat. Monspel., Sér. Bot. 18: 224 (1967).
Retief in Figueiredo & Smith, Plants of Angola: 169 (2008). Type: Angola, Bié, Rio
Cubango, Vila da Ponte, Gossweiler 2355 (BMBM000838219 holo-, LISC-LISC001326 iso-).
Cissus lutea Exell & Mendonça, Bol. Soc. Brot. 26:
228 (1952). Exell & Mendonça, C.F.A. 2(1): 58
(1954).
Cyphostemma luteum (Exell & Mendonça) Desc.,
Notul. Syst. (Paris) 16: 122 (1960), comb. inval.
H: Herb with multi-headed rootstock. D: HI.
Endemic. (Figure 9.)
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5
6
Figure 4. C. currorii in leaf. Photo taken in Namibia on the Groot Spitzkoppe on 4 May 2005 by Tim
Harvey. Figure 5. C. currorii in the dry season. Photo taken in Namibia at Baobab Bend, approximately 40 km south of Opuwo on the C43, on 24 July 2007 by Tim Harvey. Figure 6. Close-up of the
bark of C. currorii. Photo taken in Namibia at Baobab Bend, approximately 40 km south of Opuwo on
the C43, on 10 May 2005 by Tim Harvey.
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7
8
Figure 7. Geographical distribution map of C. grandistipulatum (Gilg & M.Brandt) Desc., C. violaceoglandulosum (Gilg) Desc. and C. wittei (Staner) Wild & R.B.Drumm. in Angola. Figure 8. Geographical distribution map of C. junceum (Webb) Desc. ex Wild & R.B. Drumm. subsp. jatrophoides (Welw. ex
Baker) Verdc., C. ruacanense (Exell & Mendonça) Desc. and C. uter (Exell & Mendonça) Desc. in Angola.
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10
Figure 9. Geographical distribution map of C. luteum (Exell & Mendonça) Desc., C. pendulum (Welw.
ex Baker) Desc. and C. pruriens (Welw. ex Baker) Desc. in Angola. Figure 10. Geographical distribution map of C. stenolobum (Welw. ex Baker) Desc. ex Wild & R.B.Drumm., C. stipulaceum (Baker) Desc.
and C. subciliatum (Baker) Desc. ex Wild & R.B.Drumm. in Angola.
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13. Cyphostemma mendesii F.Sousa, Phytotaxa
7: 36 (2010). Type: Angola, Moçâmedes, Dois
Irmãos, ca. Pedra da Providência, 21 December
1955, Mendes 1145 (holo- & iso- LISC).
H: Rhizomatous herb with succulent stems. D:
NA. Endemic. (Figure 2.)
14. Cyphostemma pendulum (Welw. ex Baker)
Desc., Nat. Monspel., Sér. Bot. 18: 226 (1967).
Retief in Figueiredo & Smith, Plants of
Angola: 169 (2008). Type: Angola, Pungo
Andongo, Presídio, Welwitsch 1472 (BMBM000838242, COI-COI00005296, K, LISULISU208731 &, LISU208734 & LISU208733
& LISU208732).
Vitis pendula Welw. ex Baker in Oliver, Fl. Trop.
Afr. 1: 407 (1868).
Cissus pendula (Welw. ex Baker) Planch. Monogr.
Phan. 5: 594 (1887). Exell & Mendonça, C.F.A.
2(1): 62 (1954).
Cyphostemma pendulum (Welw. ex Baker) Desc.,
Notul. Syst. (Paris) 16: 123 (1960), comb. inval.
H: Climbing herb to 90 cm long. D: HI, MA, NA.
Endemic. (Figure 9.)
15. Cyphostemma pruriens (Welw. ex Baker)
Desc., Nat. Monspel., Sér. Bot. 18: 226 (1967).
Retief in Figueiredo & Smith, Plants of
Angola: 169 (2008). Type: Angola, Huíla, pr.
Lopolo, Welwitsch 1454 (BM-BM000838250,
LISU-LISU208717 & LISU208716).
Vitis pruriens Welw. ex Baker in Oliver, Fl. Trop.
Afr. 1: 408 (1868).
Cissus pruriens (Welw. ex Baker) Planch.,
Monogr. Phan. 5: 595 (1887). Exell & Mendonça, C.F.A. 2(1): 57 (1954).
Cyphostemma pruriens (Welw. ex Baker) Desc.,
Notul. Syst. (Paris) 16: 123 (1960), comb. inval.
H: Erect or climbing herb up to 90 cm long. D: BE,
CU, HA, HI, NA. Endemic. (Figure 9.)
16. Cyphostemma ruacanense (Exell & Mendonça) Desc., Nat. Monspel., Sér. Bot. 18: 227
(1967). Retief in Figueiredo & Smith, Plants of
Angola: 169 (2008). Type: Angola, Huíla, Ruacaná, Rio Cunene, 1000 m, 9 June 1937, Exell
& Mendonça 2769 (BM-BM000838245 holo-;
COI-COI00005297, LISC-LISC001329 &
LISC001328, iso-).
Cissus ruacanensis Exell & Mendonça, Bol. Soc.
Brot. 26: 229 (1952). Exell & Mendonça,
C.F.A. 2(1): 60 (1954).
Cyphostemma ruacanense (Exell & Mendonça) Desc.,
Notul. Syst. (Paris) 16: 124 (1960), comb. inval.
H: Climber. D: CU, NA. (Figure 8.)
17. Cyphostemma stenolobum (Welw. ex Baker)
Desc. ex Wild & R.B.Drumm., Fl. Zamb. 2(2):
88
471 (1966). Retief in Figueiredo & Smith,
Plants of Angola: 169 (2008). Type: Angola,
entre Lago Ivantala e Quilengues, Welwitsch
1449 (BM-BM000838265, COI-COI00005299,
K, LISU-LISU208722 & LISU208724 &
LISU208723 & LISU208725).
Vitis stenoloba Welw. ex Baker in Oliver, Fl. Trop.
Afr. 1: 408 (1868) pro parte excl. Welwitsch
1451.
Cissus stenoloba (Welw. ex Baker) Planch. in A. &
C. De Candolle, Mon. Phan. 5(2): 578 (1887);
Exell & Mendonça, C.F.A. 2(1): 58 (1954).
Cyphostemma stenolobum (Welw. ex Baker) Desc.,
Notul. Syst. (Paris) 16: 124 (1960), comb. inval.
H: Herb up to 1 m tall, from a narrowly elongated,
tuberous rootstock. D: BE, CC, CS, HA, HI,
MA, MO. (Figure 10.)
18. Cyphostemma stipulaceum (Baker) Desc.,
Nat. Monspel., Sér. Bot. 18: 228 (1967). Retief
in Figueiredo & Smith, Plants of Angola: 169
(2008). Type: Cuanza Norte, Cazengo, R.
Luinha,
Welwitsch
1490
(B†?,
BMBM000838244, COI-COI00005300, LISULISU208735 & LISU208736).
Vitis stipulacea Baker in Oliver, Fl. Trop. Afr. 1:
408 (1868).
Cissus stipulacea (Baker) Planch., Monogr. Phan.
5: 597 (1887). Exell & Mendonça, C.F.A. 2(1):
64 (1954).
Cyphostemma stipulaceum (Baker) Desc., Notul.
Syst. (Paris) 16: 125 (1960), comb. inval.
H: Climber. D: CN, MA. Endemic. (Figure 10.)
19. Cyphostemma subciliatum (Baker) Desc. ex
Wild & R.B.Drumm., Fl. Zamb. 2: 474, t. 97a
(1966). Retief in Figueiredo & Smith, Plants of
Angola: 169 (2008). Type from Mozambique.
Vitis subciliata Baker in Oliver, Fl. Trop. Afr. 1:
409 (1868).
Cissus subciliata (Baker) Planch. in A. & C. De
Candolle, Monogr. Phan. 5(2): 594 (1887).
Cyphostemma subciliatum (Baker) Desc., Notul.
Syst. (Paris) 16: 125 (1960), comb. inval.
Cissus marionae Exell & Mendonça, Bol. Soc.
Brot. ser. 2, 26: 230 (1952). Exell & Mendonça,
C.F.A. 2(1): 62 (1954).
Cyphostemma marionae (Exell & Mendonça)
Desc., Nat. Monspel., Sér. Bot. 18: 224 (1967).
Type from Zambia.
H: Climbing herb with a tuberous, napiform root.
D: LS. (Figure 10.)
20. Cyphostemma uter (Exell & Mendonça)
Desc., Nat. Monspel., Sér. Bot. 18: 229 (1967).
Retief in Figueiredo & Smith, Plants of Angola: 169 (2008). Type: Angola, Moçâmedes,
Bradleya 29/2011
11
12
Figure 11. C. uter in leaf in the rainy season. Photo taken approximately 15 km east of the port city of
Namibe in southern Angola on 14 January 2009 by Ian Oliver. Figure 12. C. uter in the dry season.
Photo taken along the road from Leba Pass en route to Namibe in southern Angola on 6 August 2010 by
Gideon F. Smith.
Bradleya 29/2011
89
Providência, 24 October 1937, Abreu 51 (BMBM000838241 iso-; COI-COI00005303 holo-).
Cissus uter Exell & Mendonça, Bol. Soc. Brot., ser.
2, 26: 231−232 (1952). Exell & Mendonça,
C.F.A. 2(1): 70 (1954).
Cyphostemma uter (Exell & Mendonça) Desc.,
Notul. Syst. (Paris) 16: 125 (1960), comb. inval.
H: Deciduous, caudiciform succulent to 1.5 m tall,
in desert coastline. D: CU, NA. (Figures 8,
11–12.)
21. Cyphostemma
violaceo-glandulosum
(Gilg) Desc., Nat. Monspel., Sér. Bot. 18: 230
(1967). Figueiredo et al., Bothalia 39(2): 201
(2009). Retief in Figueiredo & Smith, Plants of
Angola: 169 (2008). Type: Angola, Kubango,
Chirumbu, 1899/10/11, Baum 270 (B†, lectodesignated by Exell & Mendonça (1954: 56);
BM, COI-COI00005305, G-G00023926, K, MM0109115, S-S-G-1413 isolecto-).
Cissus violaceo-glandulosa Gilg in Warburg,
Kunene-Sambesi-Expedition: 294 (1903). Exell
& Mendonça, C.F.A. 2(1): 56 (1954).
Cyphostemma violaceo-glandulosum (Gilg) Desc.,
Notul. Syst. (Paris) 16: 125 (1960), comb. inval.
H: Prostrate herb from a fleshy, woody, multiheaded rootstock. D: BE, CC, CU, HI.
Endemic. (Figure 7.)
22. Cyphostemma wittei (Staner) Wild &
R.B.Drumm., Kirkia 2: 141 (1961). Sousa et al.,
Bothalia 41(2): 260 (2011). Type from Congo
(Katanga).
Cissus wittei Staner, Contr. Fl. Katanga, Suppl.
4: 49 (1932).
H: Prostrate or scrambling herb from a thick
fusiform woody tap-root. D: MO. (Figure 7.)
Insufficiently known taxa
Cyphostemma sp. aff. vogelii. Referred to as
Cissus sp. aff. vogelii by Exell & Mendonça (1954:
66).
Cyphostemma sp. aff. fugosioides. A climber up to
3.5 m high, with tendrils, 5-foliolate leaves with
narrow leaflets.
Acknowledgements
Mr Ian Oliver, Curator of the Karoo Desert National Botanical Garden of South Africa, and Dr
Tim Harvey of California, USA, are thanked for
the use of images of C. uter and C. currorii, respectively. We thank Michele Walters for producing the distribution maps. The curators of
LISC, LISU and PRE are thanked for facilitating
access to their collections. Dr Christien Bredenkamp of SANBI is thanked for constructive
criticism on the paper.
90
References
ALSTON , A.H.G. (1931). Supplement. In TRIMEN,
H., A Hand-book to the Flora of Ceylon, part 6.
London, Dulau & Co.
DESCOINGS, B. (1960). Un genre méconnu de Vitacées: compréhension et distinction des genres “Cissus” L. et “Cyphostemma” (Planch.)
Alston. Notul. Syst. (Paris) 16: 113–125.
DESCOINGS, B. (1967). Note rectificative à propos
de la nomenclature des Cyphostemma (Vitacées). Nat. Monspel. Sér. Bot. 18: 217–230.
EXELL, A.W. & MENDONÇA, F.A. (1954). Vitaceae.
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G.F. & FADEN, R.B. (2009). The botany of the
Cunene-Zambezi Expedition with notes on
Hugo Baum (1867–1950). Bothalia 39: 185–
211.
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a portable dictionary of plants, their classifications and uses. 3rd ed. Cambridge University Press, Cambridge.
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Continuato 5(2): 472–472.
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STEFFENS, F.E., DIXON, R.D. & RETIEF J.A.
(1997). List of southern African succulent
plants. Umdaus Press, Pretoria.
SOUSA, F., FIGUEIREDO, E. & SMITH, G.F. (2011).
Vitaceae. A new and an overlooked record of
Cyphostemma (Planch.) Alston in Angola.
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LAUNERT, E. (ed.), Flora Zambesiaca, 2(2):
439–492. Flora Zambesiaca Managing Committee, London.
Appendix 1. List of specimens used to prepare the maps
C. adenocaule subsp. adenocaule: Albano Rocha
52 (LISC!); Albano Rocha 64 (LISC!); Anchieta 44
(B COI LISU!); Araújo 64 (LISC!); Barbosa &
Correia 9121 (LISC!, PRE!); Beatriz 38 (COI);
Büttner 99 (B); Cameira 324 (LISC!); Exell &
Mendonça 3105 (BM COI LISC!); Exell &
Mendonça 3191 (BM COI); M.E.F.A. 726 (LISC!);
Mendes 499 (LISC!); Mendes 933 (LISC!); Mendes
Bradleya 29/2011
1155 (LISC!); Mendes 1238 (LISC!); Monteiro s.n.
(K); Monteiro, Santos & Murta 122 (LISC!);
Monteiro, Santos & Murta 166 (LISC!); Pearson
2345 (B K); Pearson 2749 (K); Raimundo, Matos &
Figueira 209 (LISC!); Santos 1377 (LISC!, PRE!);
Teixeira 3689 (LISC!); Teixeira et al. 10298
(LISC!); Teixeira et al. 11133 (LISC!); Teixeira et
al. 11498 (LISC!); Welwitsch 1452 (BM LISU!);
Welwitsch 1479 (BM LISU!); Welwitsch 1479b
(BM LISU!); Welwitsch 1492 (BM LISU!);
Welwitsch 1494 (BM LISU!); Welwitsch 1493 (BM
COI K LISU!).
C. adenopodum: Gossweiler 7690 (COI K LISC!
LISU!); Gossweiler 8208 (COI LISU!).
C. chloroleucum: Anchieta 67 (COI LISU!);
Anchieta 86 (COI LISU!); Baum 361 (B† BM COI
G K W); Borges 318 (LISC!); Borges 318a (PRE!);
Gossweiler 2217 (BM COI LISC!); Gossweiler 3649
(BM COI LISC); Gossweiler 3650 (COI LISC!);
Gossweiler 8768 (BM); Gossweiler 9247 (BM COI
K); Henriques 792 (LISC! LISU!); Mendes 197
(LISC!); Mendes 766 (LISC!); Mendes 880 (LISC!);
Mendes 980 (LISC!); Mendes 1129 (LISC!);
Menezes 1277 (LISC!); Menezes 1373 (LISC!);
Menezes 2408 (LISC!); Menezes 3299 (LISC!);
Menezes 3531 (LISC!, PRE!); Newton 230 (B†?);
Santos 53 (LISC!); Santos 79 (LISC!); Teixeira
1468 (LISC!); Teixeira 1822 (LISC!); Welwitsch
1453 (BM LISU!); Welwitsch 1463 (B†? BM COI K
LISU!).
C. congestum: Kotze 57 (PRE!); Rycroft 2423
(PRE!).
C. crassiusculum: Carrisso & Sousa 334 (BM COI
LISC!); Gossweiler 294 (B†?); Gossweiler 294c
(BM); Gossweiler 1510 (BM COI); Gossweiler 5172
(LISC! LISU!); Gossweiler 5278 (BM LISC!
LISU!); Gossweiler 8276 (BM K P); Henriques 65
(LISC!); Humbert 16134 (P); Monteiro, Santos &
Murta 121 (LISC!, PRE!); Thompson 20 (PRE!);
Welwitsch 1469 (BM LISU!); Welwitsch 1492b (B
BM COI K LISU!); Welwitsch 1495 (BM COI K
LISU!).
C. currorii: Curror s.n. (B†? K); Mendes 138
(LISC!); Mendes 424 (LISC!); Mendes 1224
(LISC!); Mendes 1733 (LISC!); Mendes 8262?
(LISC!); Mendonça 4530 (LISC!); Menezes 316
(LISC!); Menezes, Henriques & Brites 2892
(LISC!); Welwitsch 1446 (BM LISU!).
C. fugosioides: Antunes vel Dekindt 587 (B LISC!);
Baum 174 (B†? BM COI G K M W); Faulkner 68
Bradleya 29/2011
(BM K PRE); Gossweiler 1971 (BM); Gossweiler
2196 (BM COI LISC!); Mendes 362 (LISC!);
Mendes 730 (LISC!); Mendes 753 (LISC!); Menezes
1238 (LISC!); Menezes 1254 (LISC!); Menezes 4262
(LISC!).
C. grandistipulatum: Antunes 110 (B†?); Bester
8990 (PRE!); Fritzsche 220 (B); Mendonça 4617
(LISC!); Ramalho s.n. (B); Teixeira e Andrade
5133 (LISC!); Welwitsch 1451 (BM LISU!).
C. huillense: Exell & Mendonça 2708 (BM COI
LISC!); Torre 8747 (LISC!).
C. johannis: Gossweiler 7238 (BMpp LISC!
LISUpp!); M.E.F.A. 457 (LISC!); M.E.F.A. 676
(LISC!); Monteiro & Murta 243 (PRE!).
C. junceum subsp. jatrophoides: Anchieta 47
(LISU!); Barbosa 11164 (LISC!); Baum 278 (B†?
G W); Bester 9237 (PRE!); Gossweiler 2094 (BM);
Mendes 769 (LISC!); Mendes 1073 (LISC!);
Mendonça 4597 (LISC!); Menezes 2170 (LISC!);
Menezes 4274 (LISC!); Menezes, Barroso & Sousa
4514 (LISC!); Monteiro e Murta 1957 (LISC!);
Nolde 81 (B); Silva 228 (LISC!); Teixeira 1666
(LISC!); Teixeira &Andrade 6657 (LISC!); Teixeira
et al. 10793 (LISC!); Wellman 1546 (B); Welwitsch
1474 (B†? BM COI LISU!); Young 1006 (BM COI
LISC); Young 1090 (BM); Young 1281 (BM COI
LISC!).
C. luteum: Gossweiler 2355 (BM LISC!); Mendes
1898 (LISC!).
C. mendesii: Mendes 1145 (LISC!); Mendes 1164
(LISC!).
C. pendulum: Exell & Mendonça 167 (BM COI
LISC!); Exell & Mendonça 180 (BM COI); Mendes
1001 (LISC!); Menezes 3657 (LISC!); Welwitsch
1472 (BM COI K LISU!).
C. pruriens: Castro 8 (COI); Hundt 688 (B, BM,
COI); Huntley, Roberts & Ward 88 (PRE!); Mendes
604 (LISC!); Mendes 1012 (LISC!); Mendes 1099
(LISC!); Menezes, Barroso & Sousa 4426 (LISC!);
Menezes, Barroso & Sousa 4511 (LISC!); Menezes,
Barroso & Sousa 4629 (LISC!); Teixeira 1667
(LISC!); Torre 8405 (LISC!); Welwitsch 1454 (BM
LISU!).
C. ruacanense: Bester 9218 (PRE!); Exell &
Mendonça 2769 (BM COI LISC!); Menezes,
Henriques & Brites 2815 (PRE!); Menezes,
Henriques & Brites 2943 (PRE!); Rycroft 2423
(PRE!).
91
C. sp. aff. fugosioides: Menezes 3527 (PRE!).
C. sp. aff. vogelii: Welwitsch 1482 (BM LISU!).
C. stenolobum: Barros Machado ANG.I.55-288
(LISC!); Barros Machado ANG.I.55–357 (LISC!);
Bester 9340 (PRE!); Couto 37 (LISC!); Dekindt 67
(B); Gossweiler 2240 (BM COI LISC!); Gossweiler
2905 (BM COI LISC!); Hundt 367 (B); Jessen 420
(B); Mendes 606 (LISC!); Mendes 1041 (LISC!);
Mendes 1126 (LISC!); Mendes 1868 (LISC!);
Mendes 2230 (LISC!); Menezes 1584 (LISC!);
Teixeira & Figueira 6658 (LISC!); Torre 8606
(LISC!); Torre 8613 (LISC!); Wellman 1775 (K);
Welwitsch 1449 (BM COI K LISU!); Young 938
(BM COI LISC); Young 1349 (BM).
C. stipulaceum: Exell & Mendonça 278 (COI);
Gossweiler 594 (BM K); Gossweiler 6363 (COI
LISU!); Gossweiler 10318 (BM COI LISC!); Nolde
523 (B BM); Welwitsch 1475 (BM LISU!);
Welwitsch 1490 (B†? BM COI LISU!); Welwitsch
1471 (BM LISU!).
92
C. subciliatum: Exell & Mendonça 1407 (COI).
C. uter: Abreu 51 (BM COI); Bester 9116 (PRE!);
Exell & Mendonça 2182 (BM COI LISC!); Gomes
e Sousa s.n. (COI); Mendes 136 (LISC!); Mendes
136a (LISC!); Mendes 145 (LISC!); Mendes 170
(LISC!); Mendes 1309 (LISC!); Menezes, Barroso
& Sousa 4828 (LISC!); Menezes & Henriques 216
(LISC! LISU!); Pearson 2256 (K); Pimentel
Teixeira s.n. (BM K); Teixeira 1716 (LISC!);
Teixeira et al. 12699 (LISC!).
C. violaceo-glandulosum: Anchieta 77 (LISU!);
Baum 270 (B†? BM COI G K M S); Baum 270a
(B); Carrisso & Mendonça 579 (BM COI); Couto
282 (LISC!); Faulkner 66 (BM K PRE!); Gossweiler
2100 (BM); Henriques 816 (LISC!); Henriques 817
(LISU!); Hundt 195 (B); Mendes 607 (LISC!);
Mendes 1113 (LISC!); Mendes 1114 (LISC!);
Menezes 1248 (LISC!); Menezes 4259 (LISC!);
Teixeira 1525 (LISC!); Teixeira & Andrade 5024
(LISC!); Teixeira & Figueira 4910 (LISC!).
C. wittei: Barros Machado ANG.XII.54–51
(LISC!).
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Estrela Figueiredo
Gideon F. Smith