Professional Documents
Culture Documents
2: 115-120 (2005)
Keywords: biogeography, Cyanella alba L.f., Cyanella L„ Eremiolirion amboense (Schinz) J.C.Manning & C.Mannheimer. Eremiolirion
J.C.Manning & F.Forest, gen. nov., Namibia. South Africa. Tecophilaeaceae
ABSTRACT
The generic affiliation o f Cyanella amboensis Schinz has been uncertain since the species w as excluded from the genus
Cyanella L. in 1991. The species has two leaves, a divaricately branching inflorescence, ebracteolate pedicels, and acti-
nomorphic flowers with monomorphic anthers. It is endemic to the western parts of central and northern Namibia. Other
species o f Cyanella have several leaves, racemose inflorescences, bracteolate pedicels, zygomorphic flowers with dimor
phic anthers, and are endemic or near-endemic to the winter rainfall region in southwestern South Africa and southern
Namibia. These differences are consistent with the recognition of the species as a distinct genus within the family.
Phylogenetic analysis of plastid DNA sequences indicates that C. amboensis Schinz is sister to the other species of
Cyanella, a relationship that also supports its independent generic status. The monotvpic genus Eremiolirion is according
ly erected to accommodate the species. Minor differences in flower colour and vegetative morphology in Cyanella alba L.f.
are shown to correlate with the three disjunct groups o f populations in which the species occurs, and these populations are
recognized at the level o f subspecies.
Tecophilaeaceae are a small family of geophytic peren praisal of its relationships within the family. A label
nials well circumscribed by their cormous rootstock, porici- appended to the specimen Bean. VIok & Viviers 1824
dal anthers and semi-inferior ovary (Simpson & Rudall (BOL) indicates that at the time she had considered that
1998). The flowers of all species are attractive, brightly the species might be best transferred to the Chilean genus
coloured and often fragrant (Manning etal. 2002). The fam Conanthera, with which it shares a branched inflores
ily comprises 24 species, currently distributed among eight cence and symmetrical anthers, but the similarity is not
small genera, three of which are monotypic. The genus actually that close. Conanthera has numerous leaves and
Odontostomum Torr. (1 sp.) is endemic to California, a zygomorphic, blue to purple perianth, in contrast to the
Conanthera Ruiz & Pav. (5 spp.). Tecophilaea Bertero ex bifoliate condition and actinomorphic. white perianth of
Colla (2 spp.) and Zephyra D.Don (1 sp.) are restricted to C. amboensis. Currently, therefore, the species is of
Chile, and the remaining four. Cyanastrum Oliv. (3 spp.), uncertain generic affinity. Molecular evidence places C.
Cyanella L. (7 or 8 spp.), Kabuyea Brummitt (1 sp.) and amboensis as sister to a monophyletic clade comprising
Walleria J.Kirk (3 spp.), occur in sub-Saharan Africa the remaining species in the genus (six of the seven
(Simpson & Rudall 1998). An analysis of plastid remaining species are included in this study). This topol
sequence data for the rbcL gene and the trnL-F region for ogy offers two options for resolving the generic position
all genera and most species (18), places the three Chilean of C. amboensis'. the species can either be incorporated
genera as sister to the Califomian-African clade and within an expanded circumscription o f the genus
retrieves all genera as monophyletic (Forest & Manning Cyanella; or a new genus should be erected to accom
2005 in prep.). modate it. The morphological differences between
Cyanella s. str. and C. amboensis are substantial, includ
The African genera have been thoroughly mono ing leaf number, inflorescence structure, floral symme
graphed (Carter 1962; Scott 1991; Brummitt et al. 1998). try, and arrangement of the androecium. Genera in
Among them. Cyanastrum and Kabuyea are found only Tecophilaeaceae as currently circumscribed are homoge
in the tropics, Walleria is widely dispersed through sub neous assemblages of closely related species in which
tropical Africa, and Cyanella is almost restricted to the floral symmetry in particular plays a defining role.
winter rainfall region of South Africa in the extreme Including C. amboensis within Cyanella would not only
southwest of the continent. A single species of Cyanella, enlarge the circumscription of the genus to the extent that
C. amboensis Schinz is found further north, in central it is no longer morphologically coherent but would ren
Namibia. This species is morphologically anomalous der it uniquely heterogeneous in comparison to the other
among the rest of the species of Cyanella and was genera in the family.
excluded from the genus by Scott (1991) pending a reap
Cyanella s. str. is circumscribed on the basis of its
tunicated corm, several to numerous leaves, zygomor
* Compton Herbarium, South African National Biodiversity Institute,
Private Bag X7. 7735 Claremont. Cape Town. phic flowers with dimorphic anthers, and bracteolate
** Leslie Hill Molecular Systematics Laboratory. South African pedicels. All of the species are endemic to or centred in
National Biodiversity Institute. Private Bag X7. 7735 Claremont. Cape the winter rainfall region of southern Africa. Cyanella
Town & Department of Botany. University of Cape Town. Private Bag. amboensis. in contrast, has just two leaves, the flowers
7701 Rondebosch, Cape Town.
*** National Herbarium o f Namibia. National Botanical Research are truly actinomorphic with monomorphic anthers, the
Institute. Private Bag 13184. Windhoek. pedicels are ebracteolate. and the species is distributed in
MS. received: 2005-04-05. summer rainfall Namibia. On the basis of these several
116 Bothalia 35,2 (2005)
FIGURE 1.— Eremiolirion amboense, Mannheimer 2510 (NBG). A, whole plant; B, flower, front view; C, inner tepal; D, outer tepal; E, androe-
cium; F, single anther, adaxial surface; G, half flower detail; H, tip o f anther; I, detail o f mouth o f floral tube showing corona; J, capsule;
K, seed. Scale bar: A -D , J, 10 mm; E-G, 2.5 mm; H, 3.75 mm; I, 1 mm; K, 2 mm. Artist: J.C. Manning.
Bothalia 35,2 (2005) 117
Clanwilliam and the Biedouw Valley. They are separated has never been collected there again. There are other
from the Bokkeveld populations by the arid expanse of instances of mislabelling by Stokoe (Goldblatt & Manning
the Doom River Valley, which supports a succulent 2000: 507).
karoo vegetation in which C. alba does not grow. The
Cederberg populations resemble the Bokkeveld popula Additional material examined
tions in stature but are distinguished from them by their
flower colour. In these plants the tepals (sometimes only NORTHERN CAPE.— 3119 (Calvinia): N o f Nieuwoudtville.
the inner whorl but usually both whorls) are pale yellow Grasberg. (-AC), 16 September 1961. W.F. Barker 9457 (NBG): top of
Van Rhvn's Pass. (-AC). 29 November 1974. W. Wisura 2991 (NBG);
rather than white, and the upper five anthers are invari Van Rhyn's Pass. S side. (-AC). 4 October 1962, G.J. Lewis 6126
ably coherent and marked on their outer (upper) face (NBG); Nieuwoudtville Reserve. (-AC). 8 September 1983. PL. Perry
with a prominent dark maroon or blackish blotch near the & D. Snijman 2351 (NBG); Nieuwoudtville Reserve, (-AC), 12
base. Like the Bokkeveld plants, the pedicels are bracte- October 1983. PL. Pern & D. Snijman 2383 (NBG): Nieuwoudtville,
(-AC), January 1931.' J.W Mathews NBG2270/30 (BOL); near
olate.
Nieuwoudtville. (-AC). September 1941. T.P. Stokoe 8693 (BOL): 11
km from Nieuwoudtville in direction o f Calvinia. Van Wyk’s Farm.
A third group of populations occurs well to the south September 1930, (-AC). L. Bolus 19587 (BOL): ± 15 km S of
at Karoopoort, in small patches of renosterveld that fringe Nieuwoudtville. Matjiesfontein. (-AC), 13 September 1976. M.F.
the arid Tanqua River Basin. These plants are typically Thompson 2902 (NBG); 15 km SSW o f Nieuwoudtville. Oorlogskloof
smaller in stature with fewer, up to three and often just Nature Reserve. Farm Driefontein. (-CA). 10 October 1996. W.A.J.
Pretorius 360 (NBG); Farm Lokenberg. (-CA). 4 September 1985. D.
one flower per plant, borne on pedicels that are mostly
Snijman 905 (NBG); Lokenburg, (-C A ). 29 August 1941. E.
less than 100 mm long, rarely up to 150 mm. The flowers Esterhuysen 5751 (BOL): Menzieskraal. (-CA). 29 September 1933.
are white or pale pink, flushed darker pink on the reverse, Markotters.n. (NBG).
and the upper five anthers are coherent and uniformly yel
low. Superficially the plants resemble small forms of the Doubtful locality
Bokkeveld genotype, and it could be argued that they rep
resent nothing more than depauperate forms from mar WESTERN CAPE.— 3118 (Vanryhnsdorp): Nardouw Pass. (-DD).
September 1941. T.P. Stokoe SAM55591 (SAM).
ginal habitats at the edge of the distribution. However,
closer examination reveals that the bracteoles on the
subsp. flavescens J.C.Manning, subsp. nov.
pedicels have been completely suppressed and are thus
lacking. In this respect the plants are unique in the genus. A ceteris subspeciebus habitu 120-200 mm elato, foliis
lineari-filiformibus et 1-2 mm latis. pedicellis bracteolatis,
The three disjunct groups of populations of Cyanella
floribus 1-4 et dilute flavis vel tepalis extemis albis,
alba are thus morphologically distinct from one another.
antheris superis ad basim maculatis differt.
These morphological distinctions point to long-standing
separation, sufficient to allow genetic drift, suggesting TYPE.—Western Cape. 3218 (Clanwilliam): Biedouw-
that the disjunctions are interpreted to be the result of berg. (-AA), 26 August 1896. R. Schlechter 8686 (NBG,
isolation due to the intervening areas of aridity rather holo.!; BOL. PRE. iso.!).
than the alternative hypothesis of recent fragmentation
through farming activities also offered by Scott (1991). Plants 120-200 mm high. Leaves linear-filiform. 1-2
Since each of the groups of populations is geographical mm wide. Pedicels bracteolate. Flowers 1—4. pale yellow
ly isolated from the other, and is morphologically distin or outer tepals white. Anthers yellow, upper five macu
guishable. it is appropriate to recognize them at the level late. marked with dark blotch near base.
of subspecies. The homogeneous nature of each of the
groups of populations points to genetic continuity within Distribution: Western Cape, northern Cederberg,
each group, while the differences between them indicate between Clanwilliam and Wuppertal, especially common
significant levels of genetic discontinuity. This is highly in the Biedouw River Valley (Figure 3).
significant in the context of the high levels of microgeo
graphic speciation that characterize much of the Cape Additional material examined
flora (Goldblatt & Manning 2000).
WESTERN CAPE.—3218 (Clanwilliam): Clanwilliam. (-BB). 4
Cyanella alha L .f: 201 (1781). Type: Sheet 430/4. August 18%. R Schlechter 8405 (BOL): Clanwilliam. ( BB). no date.
Thunberg collection in herb. Linn. (LINN, lecto.; NBG, C.L. Leipoldt 254 (SAM); Clanw illiam. (-BB). 30 August 1930 (cult ). E
Struts burger s.n. (NBG): 10 km S of Clanwilliam. ( BB). 12 September
microfiche!). 1997. P Goldblatt & J Manning 10741 (MO. NBG). 3219 (Wuppertal):
Biedouwberg. (-AA). 3 September 1933. FM Leighton s.n. (BOL);
subsp. alha Biedouw Hill. ( AA). 1984. G. Scott 2 (NBG); Biedouw Mountain.
( AA). 20 September 1937. G.J. Lewis s.n. (NBG): bottom of pass into
Plants (80 ) 100-200 mm high. Leaves linear. 1-3 Biedouw Valley. (-AA). 24 August 1967. M.F Thompson 352 (NBG);
mm wide. Pedicels bracteolate. Flowers 3-9, white to bottom of hill to Biedouw Valley, (-AA). 9 August 1984. PL. Perry 3145
pale pink. Anthers uniformly yellow. (NBG); WelbedachU AA). 20 September 1937. W.F. Barker 283 (NBG):
Koudeberg near Wuppertal. (-AA). 4 October 1897. H Bolus 9095
Distribution: Northern Cape, Bokkeveld Escarpment (NBG); Citadel Kop. (-AA). 7 September 1953. R.H. Compton 24237
(NBG): Wuppertal. (- AA). 18 August 1938. B Martin NBG1794/37
from just north of Nieuwoudtville southwards to Menzies- (NBG); near Wuppertal. (-AA). 28 August 1951. B E Martin 811 (NBG).
kraal (Figure 3). The collection Stokoe SAM55591 from the
Nardouwsberg north of Clanwilliam, while certainly this subsp m inor J.C.Manning, subsp. nov.
form, is doubtfully correctly localized. The Nardouwsberg
is a sandstone range that provides no suitable habitats for A ceteris subspeciebus habitu 80-150 mm elato. foli
the species and although it is well explored, the species is filiformibus et 0.5-1.5 mm latis. pedicellis ebracteo-
120 Bothalia 35,2 (2005)
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ACKNOWLEDGEMENTS SIMPSON, M.G. & RUDALL, PJ. 1998. Tecophilaeaceae. In K. Kubitzki,
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Dr Peter Bruyns very kindly provided the Latin diagnoses. Hamburg.