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Cheiridopsis alba-oculata (Aizoaceae: Ruschioideae, Ruschieae) —Anew
quartz-endemic from southern Namaqualand, South Africa
C. Klak
a,
⁎,N.A.Helme
b
,P.V.Bruyns
a
a
Bolus Herbarium, Department of Biological Sciences, University of Cape Town, 7701 Rondebosch, South Africa
b
P.O. Box 22652, Scarborough 7975, South Africa
abstractarticle info
Article history:
Received 26 August 2014
Received in revised form 29 October 2014
Accepted 3 November 2014
Available online 27 November 2014
Edited by AR Magee
Keywords:
Cheiridopsis alba-oculata
New species
Taxonomy
Cheiridopsis alba-oculata Klak & N.A. Helme is a new species known from two populations in the Garies region of
southern Namaqualand, where it appears to be restricted to patches of quartz-gravel. The species belongs to
the Cheiridopsis–Odontophorus alliance because of its papillate leaves and the 9- or 10-locular fruits with large
closing bodies. The isophylly and globose fruits with raised tops suggest that it is closely related to C. subg.
Odontophoroides and Odontophorus.C. alba-oculata is geographicallyisolated in this group, whose other members
are restricted to northern Namaqualand. C. alba-oculata resembles Cheiridopsis ponderosa and Cheiridopsis
pilosula in its densely papillate leaves but is distinguished from them by its strongly keeled, apically toothed
leaves.
© 2014 SAAB. Published by Elsevier B.V. All rights reserved.
1. Introduction
Cheiridopsis N.E.Br., Ihlenfeldtia H.E.K. Hartmann and Odontophorus
N.E.Br. are closely related genera in the Conophytum clade (Klak et al.,
2013). They are all highly succulent, compact, caespitose perennials,
only occasionally with longer shoots. Monographs exist for all three
genera (Hartmann, 1976, 1992; Hartmann and Dehn, 1987) but several
new species and subspecies have been described subsequently in
Odontophorus and Cheiridopsis.InOdontophorus,Hartmann (1976)
recognised Odontophorus angustifolius L. Bolus, Odontophorus marlothii
N.E.Br. and Odontophorus nanus L. Bolus. Subsequently, O. angustifolius
subsp. protoparcoides S.A. Hammer (Hammer, 1994)andOdontophorus
pusillus S.A. Hammer were described (Hammer, 1996). Cheiridopsis
currently includes 32 species, including the insufficiently known
Cheiridopsis nelii Schwantes (Hartmann, 2001) in three subgenera,
C. subg. Cheiridopsis,C. subg. Aequifoliae H.E.K. Hartmann and C. subg.
Odontophoroides H.E.K. Hartmann (Hartmann and Dehn, 1987). On the
basis of differences in fruit structure, Cheiridopsis excavata L. Bolus and
Cheiridopsis vanzylii L. Bolus were moved into a new genus, Ihlenfeldtia
(Hartmann, 1992). According to Hartmann (1992), the structure of
the fruit of Ihlenfeldtia suggested a close relationship with Aloinopsis
Schwantes, Tanquana H.E.K. Hartmann and Vanheerdia L. Bolus ex
H.E.K. Hartmann. This was not supported by molecular results (Klak
et al., 2013), which re-affirmed the previous position of Ihlenfeldtia
close to Cheiridopsis, corresponding to the similar anatomy of their
leaves (Hartmann, 1992).
Odontophorus shares many characteristics with Cheiridopsis. These
include the papillate epidermis of the leaves and the large, almost glo-
bose fruits. The toothed keels and margins of the leaves are common
to Odontophorus and some species of Cheiridopsis subg. Odontophoroides
(Hartmann, 2001). Odontophorus was distinguished from Cheiridopsis
(Hartmann, 2001) by the much narrower valve wings (as opposed
to the broader valve wings in Cheiridopsis). However, in the key to
the subgenera of Cheiridopsis (Hartmann, 2001), “always narrow
valve wings”distinguish Cheiridopsis subg. Aequifoliae from C.subg.
Odontophoroides (with valve wings as broad as expanding keels or
broader). There is therefore considerable overlap in the width of the
valve wings between Cheiridopsis and Odontophorus, rendering the
current distinction between the two genera unclear. Similarly, the
considerable homoplasy among characters of the fruit revealed for the
first time in Klak et al. (2013) makes it doubtful whether Ihlenfeldtia is
distinct from Cheiridopsis.
Cheiridopsis is fairly widely distributed, from the district of Lüderitz-
South in Namibia to the western and central parts of the Western Cape
in South Africa (Hartmann and Dehn, 1987). The highest diversity of
seven species is found around Steinkopf (Hartmann and Dehn, 1987),
where three of the four species of Odontophorus also occur (Hartmann,
1976). The southernmost record for Odontophorus is 14 km south of
Springbok, along the road to Hondeklipbaai (O. marlothii). Ihlenfeldtia
occurs slightly to the east, at the edge of northern Namaqualand and
Bushmanland as far east as Pofadder. Although Cheiridopsis extends to
the southern Cape, the diversity drops to 2 or 3 species per half degree
South African Journal of Botany 96 (2015) 1–5
⁎Corresponding author.
E-mail address: Cornelia.Klak@uct.ac.za (C. Klak).
http://dx.doi.org/10.1016/j.sajb.2014.11.001
0254-6299/© 2014 SAAB. Published by Elsevier B.V. All rights reserved.
Contents lists available at ScienceDirect
South African Journal of Botany
journal homepage: www.elsevier.com/locate/sajb
square. Most of the “southern species”belong to C. subg. Cheiridopsis
(Hartmann and Dehn, 1987).
With only one species from each of these genera included by Klak
et al. (2013) in their molecular analysis, we were unable to conclude
whether Cheiridopsis,Ihlenfeldtia and Odontophorus were monophyletic
as currently circumscribed and a more detailed molecular study of this
group will provide better insight into generic and subgeneric bound-
aries among them (Powell et al., in prep.).
Recent generic treatments have favoured broader circumscrip-
tions of genera (Klak et al., 2007; Klak and Bruyns, 2012, 2013) due
to the high levels of homoplasy among many morphological charac-
ters (Klak et al., 2013). Below, we show that the new species could
be placed equally well in Cheiridopsis subg. Odontophoroides or in
Odontophorus. In view of the many shared morphological similarities
between Cheiridopsis subg. Odontophoroides and Odontophorus,we
predict that molecular data will lend further support to an enlarged
generic concept of Cheiridopsis, including both Odontophorus and
Ihlenfeldtia. We therefore describe the new species in Cheiridopsis
subg. Odontophoroides.
2. Materials and methods
Morphological data on the new species came from herbarium mate-
rial and observationsin the field. All measurements are basedon mature
leaves, fresh flowering material and mature capsules. Collections at BOL,
NBG and SAM were consulted.
3. Taxonomic treatment
Cheiridopsis alba-oculata Klak & N.A. Helme, sp. nov. Type: South
Africa. Western Cape: Hondeklipbaai (3017), south of Bruinkop farm
towards Spioenkop, (–DA), on loam with superficial quartz-gravel,
212 m alt., 26 Sep 2013, Klak 2308 (BOL, holo.; K, iso.).
Compact, tufted, succulent perennial to 100 × 300 mm, branches
to 150 mm long, internodes 5–15 mm long, greyish to ochre, rooting
at nodes. Leaves all similar in shape, trigonous, widened and thicker
subapically, 30–80 × 11–16 mm, connate basally into sheath
10–15 mm long, dull to brownish green, epidermis velvety, keel
and margins dentate towards tips. Flowers solitary, 60–70 mm
diam., pedicels 35 mm long with leaf-like bracts positioned near
middle; calyx: lobes 5 or 6, subequal; petaloid staminodes yellow
becoming white towards base, 30.0–33.0 × 1.5 mm, in many rows;
filamentous staminodes few, slightly overtopping stamens; stamens
9mmlong,inmanyrows,filaments white, anthers yellow, only inner-
most filaments papillate towards base; ovary slightly raised on top, styles
9 or 10, green, 5–6 mm long, plumose on inside, with coilomorphic
holonectary. Fruits erect, subglobose, 12–16 × 9–11 mm, top rounded
with moderately raised rims but lower than basal part, basal part hairy,
9- to 10-locular, closing bodies large, blocking entire exit of locule,
covering membranes undulate, sloping downwards towards centre of
fruit and dented around middle, rims bent upwards, without additional
closing devices below covering membranes, keels dark brown, parallel
at base and later diverging apically, valve wing narrow and ending in
awns. Seeds pear-shaped, 1.5 × 1.0 mm, light brown, smooth but
colliculate in micropylar region. Flowering time: August–September.
3.1. Distribution and ecology
C. alba-oculata is known from two populations near Garies in
southern Namaqualand, where it appears to be restricted to patches of
quartz-gravel, in Namaqualand Heuweltjieveld (Mucina et al., 2006)
and known colloquially as Hardeveld. This is a widespread vegetation
type in Namaqualand. The habitat would, however, more correctly be
described as an outlier of the Riethuis-Wallekraal Quartz Vygieveld
(Mucina et al., 2006), which occurs only 30 km away, as it is structurally
and floristically most similar to this unit. However, units smaller than
1 km in width, as is the case for most quartz patches, are typically
unmapped in the South African vegetation map due to scale issues.
The underlying rock-type is gneiss, which has weathered into a
sandy loam, overlain by large, white quartz-pebbles. Such patches of
quartz occupy less than 2% of the total Hardeveld. The slopes may be
slightly convex to slightly concave and are north-, west- or south-facing.
Associated common species include Antimima komkansica (L. Bolus)
H.E.K. Hartmann, Dicrocaulon ramulosum (L. Bolus) Ihlenf., Knersia
diversifolia (L.Bolus)H.E.K.Hartmann&LiedeandMonilaria chrysoleuca
Schwantes (Aizoaceae), Didelta carnosa Aiton, Rhynchopsidium pumilum
DC. (Asteraceae) and Euphorbia restituta N.E.Br. (Euphorbiaceae).
C. alba-oculata is locally common and is a conspicuous element of its
habitat. Only two other species of Cheiridopsis,Cheiridopsis namaquensis
(Sonder) H.E.K. Hartmann and Cheiridopsis denticulata (Haw.) N.E.Br.,
have been recorded in the same quarter degree square (3017DA) as
C. alba-oculata (Fig. 1). However, the three species prefer different
soils: C. namaquensis is mostly found in soils derived from shale, with
the northern populations occurring on granites or quartzites, whereas
C. denticulata prefers gently sloping, gravelly, gneissic soils.
A new species closely related to Jacobsenia vaginata was found near-
by on more alkaline soils and the Sandveld to the west has yielded sev-
eral new species, including Lampranthus procumbens Klak (Aizoaceae)
and Ferraria ornata Goldblatt & J.C. Manning (Iridaceae) (Klak, 2003;
Goldblatt and Manning, 2011). The nearest extensive quartz-fields are
near Riethuis and Wallekraal, some 30 km to the north and near
Komkans, some 70 km to the southeast. Othonna lepidocaulis Schltr.
(Asteraceae), previously known only from Komkans and fields of
quartz-gravel and stony hills on the northern Knersvlakte, some
80 km to the southeast, was also noted and the very small, rarely re-
corded Crassula multiceps Harv. (Crassulaceae) also occurs here.
Thus, the smaller and more isolated patches of quartz-gravel where
C. alba-oculata occurs show links to other quartz fields further north
and to those further south as well but they include some endemic spe-
cies such as C. alba-oculata.
The unscented flowers are typically open from 10 am to 5 pm and
were visited by monkey beetles (Hoplinii), horseflies (Tabanidae) and
bee-flies (Bombylidae).
3.2. Diagnostic characters
Cheiridopsis and its sister genera Odontophorus and Ihlenfeldtia
are distinguished from other genera in Ruschieae by papillate leaves
and multi-locular fruits (8 to 18 locules) with large or rarely small
(Ihlenfeldtia) closing bodies (Hartmann and Dehn, 1987).
Similarly, the diagnostic characters to distinguish the three subgenera
of Cheiridopsis from each other as well as from Ihlenfeldtia and
Odontophorus are mainly found in the leaves and fruits (Hartmann
and Dehn, 1987; Hartmann, 1992).
Common to these three genera are the papillate leaves (Hartmann,
2001) which, however, show differences in the shape and density of
the papillae (Hartmann and Dehn, 1987). The leaves in C. alba-oculata
are densely papillate (Fig. 2A), as found in Cheiridopsis pilosula and
Cheiridopsis speciosa of C.subg.Odontophoroides and also in Odontophorus.
C. alba-oculata cannot be confused with any member of C. subg.
Cheiridopsis, since all members of the latter are heterophyllous (the
first pair of leaves is shorter than the second pair). The other two
subgenera of Cheiridopsis,aswellasOdontophorus and Ihlenfeldtia, are
isophyllous (both pairs of leaves are of the same length). In addition,
the leaves are 30–80 mm long in C. alba-oculata, whereas they never
exceed 50 mm in Odontophorus (Hartmann, 2001). Although the leaves
in C. pilosula and Cheiridopsis ponderosa (both C.subg.Odontophoroides)
may also reach 70–120 mm long (Hartmann, 2001), the leaves of
C. pilosula lack teeth (Hartmann and Dehn, 1987) and, although the
leaves of C. ponderosa may have tiny teeth near the apex, the keels are
rounded. This contrasts with the leaves of C. alba-oculata, which are dis-
tinctly keeled with often quite prominent teeth along the keels and
2C. Klak et al. / South African Journal of Botany 96 (2015) 1–5
margins towards their tips (Fig. 3A), a feature also found in C.subg.
Odontophoroides and in Odontophorus.
It is uncommon for species of Cheiridopsis or Odontophorus to have
distinctly two-coloured petaloid staminodes (white in the lower third
and yellow in the two upper thirds). The flowers in both populations
of C. alba-oculata are bi-coloured (Fig. 3A). This is otherwise known in
C. pilosula,C. denticulata and Cheiridopsis caroli-schmidtii (Dinter & A.
Berger) N.E.Br., which all include populations with unicoloured petaloid
staminodes.
All 14 species of Cheiridopsis subg. Cheiridopsis have mostly decum-
bent capsules. The other two subgenera, as well as Odontophorus,have
predominantly erect capsules and this is also true of C. alba-oculata
Fig. 1. Distribution of Cheiridopsis alba-oculata.
3C. Klak et al. / South African Journal of Botany 96 (2015) 1–5
(Fig. 3A). The new species shares with Odontophorus and Cheiridopsis
the large closing bodies (Fig. 3B), whereas they are always small in
Ihlenfeldtia (Hartmann, 1992). In addition, in terms of the shape of the
fruits, the almost globose, rounded tops of C. alba-oculata (Fig. 3B) are
most similar to those of C.subg.Odontophoroides and of Odontophorus.
In addition, the lower parts of the fruits are densely hairy. Although
Hartmann (1976) suggested that the papillate (hairy) fruits of
Odontophorus are one of the main characteristics separating this
genus from Cheiridopsis, similarly hairy fruits are also found in
C. ponderosa S.A. Hammer and in C. pilosula L. Bolus, where the papil-
lae are shorter. Hartmann (2001) also used this character in the key
to separate the genera Cheiridopsis (“base of capsule rough”)from
Odontophorus (“base of capsule velvety”). However, species with
“velvety”fruits such as C. ponderosa are nevertheless still included
in Cheiridopsis. In addition, the terms “velvety”and “rough”are diffi-
cult to quantify and intermediates exist, as in the case of C. pilosula.
The fruits of the new species can only be confused with members
of Odontophorus or C. subg. Odontophoroides, but none of these occurs
so far to the south.
The seeds are smooth and pear-shaped in the new species (Fig. 2B),
characteristics encountered in both Cheiridopsis and Odontophorus,both
of which may also have more papillate seeds (Hartmann and Dehn,
1987; Hartmann, 2001). The papillate seeds in C. pilosula (Hartmann
and Dehn, 1987) distinguish this species from C. alba-oculata.
3.3. Etymology
The epithet “alba-oculata”is derived from the latin albus = white
and oculus = eye and alludes to the white centre of the flowers.
3.4. Conservation status
The species is known only from two discrete locations, 3 km apart.
The two populations cover a total area (area of occupancy; AOO) of
less than 5 ha. The total number of plants is estimated at between 500
and 1000. No significant threats were observed and, although the area
is grazed by small livestock (mainly sheep), the plants did not appear
to be affected by grazing. According to IUCN 3.1 criteria (IUCN, 2001)
the species is classified as VU D1.
3.5. Additional specimens seen
South Africa—Northern Cape: 3017 (Hondeklipbaai): 30 km west of
Garies, farm Soutfontein 436, along the Outeep River, (–DA), 20 Aug.
2013, on quartz-gravel, 135 m alt., N.A. Helme 8058 (BOL, NBG).
Acknowledgements
The authors received funding from the National Research Founda-
tion (NRF) as an incentive grant for rated researchers. The curators of
BOL, NBG and SAM are thanked for permission to examine specimens.
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