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Taxonomic revision of the southern African species
of the genus Cynoglossum L. (Boraginaceae)
Lydia K. Madika1, Annah Ntsamaeeng Moteetee1
1Department of Botany & Plant Biotechnology, Faculty of Science, University of Johannesburg, PO Box 524
Auckland Park 2006, Johannesburg, South Africa
Corresponding author: Annah Ntsamaeeng Moteetee (amoteetee@uj.ac.za)
Academic editor: James Cohen|Received 28 July 2021|Accepted 4 December 2021|Published 15 March 2022
Citation: Madika LK, Moteetee AN (2022) Taxonomic revision of the southern African species of the genus
Cynoglossum L. (Boraginaceae). PhytoKeys 193: 9–42. https://doi.org/10.3897/phytokeys.193.72270
Abstract
e aim of the study is to provide a revision of the genus Cynoglossum in southern Africa. e genus is
taxonomically problematic within the family Boraginaceae, due to the morphological similarities it shares
with other closely related genera in the family. Cynoglossum plants are low-growing biennial, perennial,
or rarely annual herbs which are recognizable by their hairy stems and leaves, the latter are usually basal
and long petiolate. Based on the latest checklist, a total of eight species of this genus are listed for the
study region: C. alticola, C. amabile, C. austroafricanum, C. geometricum, C. hispidum, C. lanceolatum,
C. obtusicalyx (endemic to South Africa), and C. spelaeum. e occurrence of C. amabile in the region,
however, requires further investigation since the only existing specimen was collected within a protected
area in the KwaZulu-Natal province. Two specimens collected in the Doornpoort area in Pretoria, Gauteng
province, assigned to this species appear to have been misidentied. Diagnostic characters are described,
correct nomenclature, synonyms, typication, distribution maps, as well as the key for identifying the
studied species, are provided.
Keywords
Boraginaceae, Cynoglossum, Southern Africa, taxonomic revision
Introduction
e type genus Cynoglossum L. of the tribe Cynoglosseae belongs to the angiosperm
family Boraginaceae (forget-me-not or borage family). Members of this genus have
a worldwide distribution with many species occupying the temperate as well as
Copyright Lydia K. Madika & Annah Ntsamaeeng Moteetee. This is an open access article distributed under the terms of the Creative Commons
Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author
and source are credited.
PhytoKeys 193: 9–42 (2022)
doi: 10.3897/phytokeys.193.72270
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Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
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the tropical regions of the Old and New Worlds. is genus comprises about 80–
90 (Johnston 1924), to ca. 100 (Weigend et al. 2013), or as many as 200 species
(Chacón et al. 2016). e highest diversity of species occurs in the Mediterranean
region, harbouring about 20 species (Selvi and Sutorý 2012), and only a few species
are introduced in Australia and one in North America (i.e., C. creticum) (Selvi et al.
2011). e name Cynoglossum is derived from the Greek words ‘cynos’ (of a dog) and
‘glossa’ (tongue), depicting the texture and shape of the leaves in species of the genus
(Retief 2005), hence the common name ‘hound’s tongue’. is genus was established
by Linnaeus (1753) to accommodate six species that he described at the time, namely,
Cynoglossum ocinale L., C. virginianum L., C. cheirifolium L., C. apenninum L.,
C. linifolium L., and C. omphaloides L. e present state of knowledge has renamed
most of these Linnaeus names to the following accepted names, i.e., C. virginianum
L. (=Andersonglossum virginianum (L.) J.I. Cohen), C. cheirifolium L. (=Pardoglossum
cheirifolium (L.) Barbier & Mathez), C. apenninum L. (=Solenanthus apenninum (L.)
Fisch. & C.A. Mey.), C. linifolium L. (=Iberodes linifolium (L.) Serrano, R. Carbajal &
S.Ortiz), and C. omphaloides L.(=Omphalodes verna Moench).
e genus is complex and has been noted by several botanists such as Miller (2005),
Sutorý (2010), Selvi and Sutorý (2012), to be taxonomically challenging within the
family due to the limited morphological variation between the species, which has rendered
identication dicult. Another unresolved problem concerns the species relationships in
Cynoglossum. Morphological characters that were traditionally used to delineate genera
proved to be insucient, especially in dening monophyletic groups within Cynoglossum
as stated by Hilger et al. (2015). Recent phylogenetic studies by Cohen (2011, 2013,
2015), Selvi and Sutorý (2012), and Weigend et al. (2013), revealed that Cynoglossum is
paraphyletic with respect to Cynoglossopsis Brand, Lindeloa Lehm., Paracaryum Boiss.,
Pardoglossum Barbier & Mathez, Rindera Pall., Solenanthus Ledeb., and Trachelanthus
Kunze, suggesting the need for generic realignment. As a contribution to the systematics
of the family, Johnston (1924) grouped half a dozen of genera under a broad concept of
Cynoglossum based on the character of corolla shape and degree of anther exertion. From
this grouping, the genus Rindera (=Mattia) and Paracaryum (=Mattiastrum) were still
recognised as distinct with the only diagnostic feature being the fruit ornamentation.
Cynoglossum portraying zoochory adapted glochidiate dispersal units, Rindera with the
broadly winged diaspores for wind dispersal, and Paracaryum exhibiting the combinatory
features (Greuter 1981). is unresolved relationship of taxa based on morphological
traits led to authors such as Greuter and Burdert’s (Greuter 1981) request a broader
conception of this genus, where Cynoglossum sensu lato forms a single, large genus.
An article by Von Staden et al. (2013), mentioned that taxonomically problematic
genera are most likely to result in poor collection and misidentication. e latter
case has become a reality for the southern African species of Cynoglossum, whereby
members of this genus are also confused with species from other closely related genera
such as Lithospermum L. and Myosotis L., and members of the tribe Eritrichieae
Gϋrke. Lithospermum and Myosotis members are characterized by ovoid nutlets with
broad basal attachment scars and at gynobases, while in Cynoglossum the nutlets are
Taxonomic revision of the genus Cynoglossum in southern Africa 11
depressed-ovoid or subcircular with scars restricted to the apical half of the ventral
surface and have narrowly conical gynobases (Weigend et al. 2013; Otero et al. 2014).
Eritrichieae are dierentiated from Cynoglossum based on the shape of the gynobase,
i.e., narrow pyramidal to subulate gynobase and mostly small nutlets, as opposed to
broadly pyramidal gynobase and mostly larger nutlets for Cynoglossum (Hasenstab-
Lehman and Simpson 2012; Weigend et al. 2013). According to Al-Shehbaz (1991),
Cynoglossum can also be distinguished from the closely related genera Solenanthus Ledeb.
and Trachelanthus Kunze in having included, instead of exerted, stamens and lastly,
from Pardoglossum Barbier & Mathez in having slim, glabrous glochids on the nutlets,
instead of swollen glochids that are densely packed with minute papillae. e fruits in
members of this genus and the oral morphological characters, are normally considered
to be of diagnostic importance. e nutlets of tropical Cynoglossum are generally smaller
than those of species found in the temperate regions (Al-Shehbaz 1991).
In the southern African region, this genus has not yet been taxonomically revised
since the last treatment by Wright (1904), where he recognised two species, C. en-
erve Turcz. (now C. hispidum unb.) and C. micranthum Desf. He cited two species,
namely, C. leptostachyum DC. and C. hispidum unb. as being imperfectly known.
Hilliard and Burtt (1986) recorded the occurrence of Cynoglossum coeruleum subsp.
johnstonii var. mannii (Baker and C.H. Wright) Verdc. [recognised in southern Af-
rica as C. geometricum Baker & C.H. Wright] for the rst time in this region, while
Retief and Van Wyk (1996) recorded C. obtusicalyx Retief & A.E. van Wyk, the only
species endemic to South Africa. While a comprehensive revision is still lacking for
southern African species, revisionary work has been undertaken mainly at regional or
country level, for example East Africa (Verdcourt 1991), China (Shu 1995), Comoro
Islands and Madagascar (Miller 2005), Italy (Selvi and Sutorý 2012), Nepal (Kӧnig
et al. 2015), and Taiwan (Hsiao and Liu 1998). According to the recent checklist by
Germishuizen and Meyer (2003), there are eight species of Cynoglossum occurring in
southern Africa, namely C. alticola Hilliard & B.L. Burtt, C. amabile Stapf. & J.R.
Drumm, C. austroafricanum Hilliard & B.L. Burtt, C. geometricum Baker & C.H.
Wright, C. hispidum unb., C. lanceolatum Forssk., C. obtusicalyx Retief & A.E. van
Wyk (endemic to South Africa), and C. spelaeum Hilliard & B.L. Burtt. is paper
aims to provide a taxonomic revision of the southern African species, to provide a di-
agnostic key, as well as to lay out their distribution maps in the region.
Materials and methods
A total of 316 specimens loaned from the following herbaria: PRE, GRA, NH, NU,
NBG, and SAM (herbarium acronyms following iers 2019), were examined for
distribution and morphological data. e type specimens of relevant species were stud-
ied online from the JSTOR website (https://plants.jstor.org/). Voucher specimens and
plant material were collected during several eld trips conducted throughout South
Africa and Lesotho between October 2018 and December 2019. Specimens collected
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
12
were deposited in JRAU. e system by Edwards and Leistner (1971) was used for
specimen citation under the section ‘additional specimen examined’.
Data on vegetative morphology was obtained by analysing all the specimens provided
per species. Inorescence structures were studied from the freshly collected samples,
herbarium samples, as well as from the original author’s descriptions (in a case where all
the specimens did not contain any inorescence to study). Hand drawings representing
both the vegetative and reproductive characters were made for all the species (except
for C. amabile). e trichomes from the leaf samples and mature nutlets, from as many
specimens as possible, were examined per species, except for C. amabile, where there was
only one specimen available, and C. obtusicalyx, where three specimens were available.
Original descriptions from JSTOR.org were also included in developing the diagnostic
key. e specimens were studied using either the TESCAN VEGA3 scanning electron
microscope (SEM) or the Phenom Desktop SEM. For TESCAN VEGA3 SEM analysis,
samples were immersed in a mixture of 95% ethanol and isoamyl acetate (1:1) for 10
minutes and in pure isoamyl acetate for 15 minutes. After removing isoamyl acetate,
the samples were placed on a holder for critical point drying for an hour. en the
dried samples were directly mounted on aluminium stubs and sputter-coated with a
thin layer of gold before viewing under microscope. is was done to prevent charging
of specimens due to accumulation of static electric eld, and to increase the number
of secondary electrons that can be detected from the surface of the specimen. For the
Phenom Desktop SEM analysis, samples were only air-dried for an hour and were
directly mounted on the aluminium stubs and viewed under microscope. Maps were
plotted using the program CorelDraw Graphics Suite X7 (http://coreldraw.com).
Morphological characters of Cynoglossum species in southern Africa
Vegetative morphology
Members of this genus are either perennial, biennial, or rarely annual herbs which are
recognizable by their hairy stems and leaves. Roots are thickened cream white taproots
with small lateral roots. e stems are erect, hollow, simple at the base, and usually
branched above. e basal leaves are deciduous, long petiolate, lanceolate-obtuse shaped,
cross-venulate, with smooth margins, and are clustered at the lower parts of the stem
forming a rosette, covered with simple trichomes on both the adaxial and abaxial surface.
e stem leaves are alternate, sessile, or petiolate, lanceolate-obtuse shaped, with smooth
margins. Trichomes on the leaves sometimes have a pustulate base typical of Boraginaceae
members (C. austroafricanum, Cynoglossum coeruleum subsp. johnstonii var. mannii and
C. lanceolatum). Vegetative morphology is of limited diagnostic value in distinguishing
between southern African species, however a closer look at the trichomes has shown that
they may be used to distinguish between similar species. For example, both C. alticola and
C. obtusicalyx have a cluster of soft, woolly trichomes which dier in shape (cylindrical with
a pointed tip in C. alticola, vs. at surface and spherical with a blunt tip in C. obtusicalyx)
and density (denser in C. alticola than in C. obtusicalyx) as observed in Figure 1.
Taxonomic revision of the genus Cynoglossum in southern Africa 13
Figure 1. SEM micrographs of the adaxial leaf surfaces and the midrib of (A, B ) C. alticola (C , D ) C. obtu-
sicalyx B close-up of C. alticola displaying cylindrical trichomes with pointed end D close-up of C. obtusicalyx
displaying at trichomes with blunt tip. Vouchers: A , B from L.C.C. Liebenberg 5789 (PRE) C , D from
J.P.H. Acocks 8509 (PRE). SEM images scale bars: 10 mm (A); 50 µm (B); 1 mm (C); 100 µm (D).
Reproductive morphology
e inorescence is a cyme which is often dichotomously branched with spreading pan-
icles. Flowers are either pedicelled or subsessile, with ve parted corollas; corolla white
with a blue throat, blue, violet, or magenta (C. hispidum), or rarely white (C. spelaeum).
e stamens are included and arise from the base of the tube, they have short laments
and elliptic to oblong shaped anthers. e style is short and relatively thick, with a
capitate stigma. e fruit is a schizocarp of four nutlets attached apically to a narrowly
conical gynobase. e nutlets are ovoid with a convex dorsal surface. At maturity, the
nutlets produce glochidia, which are sharp hair-like spines or bristles tipped with barbs.
e glochidia are either swollen at the base or not bulbous-based, they either cover the
whole surface or are well spaced and vary in number. e structure and shape of glo-
chidia display an important distinguishing character amongst the southern African spe-
cies (Table 1), with each species portraying a unique character as can be seen in Figure 2.
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
14
Table 1. Distinguishing characters between the southern African Cynoglossum species.
Charac-
ters
C. alticola C. amabile C. austroafricanum C. coerule-
um subsp.
johnstonii
var. mannii
C. hispidum C. lanceolatum C. obtusicalyx C. spelaeum
Floral
colour
Deep blue Bluish
purple
Pale blue White with
pale blue
throat
Magenta White with pale
blue throat
Pale blue White
Nutlet
shape and
size
Convex,
6–9×5–6
mm
Ovoid to
convex,
2–4×3–4
mm
Ovoid, 3.0–
3.5×2.5–3.0 mm
Ovoid,
3–4×2.5–
3.5 mm
Convex,
5–6×3–5
mm
Ovoid-convex,
3–4×2.5–3.5
mm
Ovoid-convex,
2–4×3–4 mm
Ovoid, ca.
4×5 mm
Nutlet
ornamen-
tation
Glochidia
densely
packed on
nutlet
Marginal
glochidia
are more
distinct
than the
central
glochidia
Glochidia more
spread towards the
margins
Glochidia
more mar-
ginal and
on the
median line
Glochidia
covering
the whole
nutlet
Glochidia
sparsely ar-
ranged on nutlet
densely echi-
nulate with
glochidia
glochidia
more mar-
ginal and
acentric,
marginal
glochidia
are longer
compared to
the acentric
glochidia
Taxonomic treatment
Cynoglossum L., Sp. Pl. 1: 134 (1753) & Gen. Pl.: 5 (1754); Benth. and Hook. f., Gen.
Pl. 2: 848 (1876); C.H. Wright in Fl. Cap. 4: 13 (1904); Baker and C.H. Wright
in Fl. Trop. Afr. 4 (2): 51 (1905); Brand in Engl., Panzenr. 4: 252 (1921); Al-
Shehbaz in J. Arnold Arbor.: 112 (1991); Selvi and Sutorý in Pl. Biosyst. 146 (2):
461–479 (2012); Hilger et al. in Biodivers. Data J. 3: e4831 (2015). Type species:
C. ocinale L.
Paracynoglossum Popov. Fl. URSS xix.: 717 (1953). Type species: P. denticulatum (DC.)
Popov.
Perennial, biennial, or rarely annual herbs, often tall, up to 1,2 m in height, and slightly
branched. Stems and leaves canescent. Indumentum white, simple or tubercled. Leaves
alternate, lanceolate, obtuse or spathulate, entire; rst year basal leaves form a rosette,
lanceolate or obtuse, often long petiolate. Inorescences usually elongate, rarely bracte-
ate, sparingly branched or loosely paniculate. Flowers pedicelled or subsessile; blue or
violet with distinct veins, rarely white. Calyx ve-partite, scarcely enlarged in fruit, pat-
ent or reexed. Corolla tube short, throat closed with obtuse or arched scales; ve-lobed,
imbricate, obtuse, patent. Stamens ve, included in the corolla tube, included, with
short laments, anthers ovoid or shortly oblong, obtuse. Ovary with four distinct lobes
from an almost at receptacle; style short or rather long; stigma small, at or sub capitate;
ovules horizontal, xed to the central angle of the cell. Nutlets four, depressed, scarcely
produced at the apex, convex or at on the dorsal side or surrounded by an elevated
margin, glochidiate (hair-like spines or short prickles). Seeds straight or slightly curve
Taxonomic revision of the genus Cynoglossum in southern Africa 15
Diagnostic key to the species
1 Soft woolly hairs covering the entire plant; nutlets thickened; glochidia
densely arranged on the nutlet ................................................. 1. C. alticola
– Sti bristle hairs covering the entire plant; nutlets slightly swollen; glochidia
sparsely spaced on the nutlet ....................................................................... 2
2 Nutlets 5 mm wide; glochidia thick at the base, fruit stalk up to 2 cm long ..
............................................................................................. 5. C. hispidum
– Nutlets less than 5 mm wide; glochidia uniformly shaped, fruit stalk up to
1cm long ...................................................................................................3
Figure 2. SEM micrographs of the adaxial surface of the nutlets of eight listed species of Cynoglossum
AC.alticola B C. amabile C C. austroafricanum D C. coeruleum var mannii E C. hispidum F C. lanceolatum
G C. obtusicalyx H C. spelaeum. Voucher specimens: A from L.C.C. Liebenberg 5789 (PRE) B from J. Stewart
2021 (NU) C from O.M. Hilliard and B.L. Burtt 11803 (PRE) D from T.B. Sikhakhane 440 (NH) E from
S.P. Bester 12958 (PRE) F from S.P. Bester 4653 (PRE) G from J.P.H. Acocks 8509 (PRE) H from A. Nicholas
and B. Isaacs 1965 (PRE). SEM images scale bars: 1 mm (A, C-E, G-H); 2 mm (B); 500 µm (F).
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
16
3 Inorescences clustered at the apex; corolla bluish purple; glochidia dense at
the margins and centre of the nutlet........................................ 2. C. amabile
– Inorescences not clustered at the apex; corolla blue to white; glochidia dense
at the margins and few at the centre of the nutlet........................................4
4 Spreading long trichomes covering the whole plant; corolla longer than 7 mm
long ................................................................................... 7. C. obtusicalyx
– Sparsely shorter trichomes covering the whole plant; corolla shorter than
4mm long .................................................................................................. 5
5a Corolla white with blue throat:
6 Trichomes not thickened on both leaf surfaces; glochidia evenly distributed
across the nutlet ............................................................... 6. C. lanceolatum
– Trichomes with pustulate base on the abaxial leaf surface; glochidia on the
median line and centre of the nutlet .............................................................
.......................... 4. Cynoglossum coeruleum subsp. johnstonii var. mannii
5b Corolla uniformly coloured:
7 Leaves brightly green coloured on both surfaces, lanceolate-obtuse shaped;
corolla pale blue; length of glochidia uniform throughout the nutlet ............
................................................................................. 3. C. austroafricanum
– Leaves grey green on the abaxial surface, dark green on the adaxial surface,
spathulate-obtuse shaped; corolla white; marginal glochidia longer than
acentric glochidia .................................................................. 8. C. spelaeum
1. Cynoglossum alticola Hilliard & B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh
43(3): 343 (1986).
Type. South Africa ♀♂ Eastern Cape, Barkly East District (3027): Ben McDhui
(-DB), 5 Feb 1983, O.M. Hilliard and B.L. Burtt 16468 (E-image!, holotype; NU-
image!, isotype).
Perennial herbs, 0.2–0.6 m in height. Basal leaves 76–270×8–18 mm, lanceolate,
densely pubescent, persistent; margins entire. Stem leaves 35–120×5–10 mm, lan-
ceolate, apex acute, base cuneate, margins entire, soft woolly hairs. Trichomes spread
equally on both the adaxial and abaxial leaf surfaces, unicellular hair base, not bulbous
on both leaf surfaces. Inorescence racemose, clustered at the apex; pedicel 4–10 mm
long, lengthening considerably in fruit. Calyx ca. 4 mm long, lobes elliptic-oblong,
densely hairy on inner surface, apices obtuse. Corolla deep blue; lobe 4×3 mm diam-
eter, oblong, round apex. Nutlets convex, 6–9×5–6 mm; glochidia short and thick at
the base, densely packed on nutlet, tips multi-angular (Figure 3).
Phenology. February to May.
Conservation status. Least Concern (Raimondo et al. 2009).
Diagnostic characters. Cynoglossum alticola can be distinguished by its thick, con-
vex nutlets. Among the southern African species, it has a unique appearance due to
Taxonomic revision of the genus Cynoglossum in southern Africa 17
the presence of woolly trichomes that cover the whole plant. Furthermore, it has larger
nutlets (6–9×5–6 mm) than other species (less than 6×5 mm). According to Hilliard
and Burtt (1986), this species is related to C. alpinum (Brand) B.L. Burtt from the
highlands of Ethiopia, with which it shares nutlet shape and size, as well as the leaf
texture and colour. e dierence is observed on the fornices (small crests in the corolla
tube of a plant), as in C. alticola they are broad and short while in C. alpinum they are
long and narrow.
Distribution and habitat. e species is restricted to the Eastern Cape Province
in South Africa and Lesotho (Figure 4), where it is found growing on mountainous
terrain and on damp slopes near streams.
Additional specimens examined. South Africa. Eastern Cape: 3028 (Mata-
tiele): Drakensberg, near Barkly East (-CA), 19 Dec 1982, P.B. Phillipson 705
(PRE); between Malpas and Nek (-CA), 13 Dec 1995, T. Dold and M. Cocks 2058
(GRA).
Lesotho. 2929 (Underberg): Mokhotlong District (-AC), Jan 1953, L.C.C.
Liebenberg 5789 (PRE); 28 Feb 1947, A. Jacot Guillarmod 997 (PRE).
Figure 3. Vegetative and reproductive morphology of Cynoglossum alticola A line drawings of: 1.abranch
showing fruiting arrangement at the apex of the stem and stem leaves; 2. long petiolate basal leaves;
3.adaxial surface of nutlet B SEM micrograph showing trichomes on the leaves C SEM micrograph
showing thick and shorter glochidia on the nutlet. Voucher: L.C.C. Liebenberg 5789 (PRE). Drawing scale
bar: 8 mm. SEM images scale bars: 50 µm (B); 50 µm (C).
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
18
2. Cynoglossum amabile Stapf & J.R.Drumm. in Bull. Misc. Inform., Roy. Bot.
Gard., Kew 6:202 (1906).
Type. China ♀♂ Yunnan, Mengtsze, 1894, W. Hancock 133 (K-image! [3 sheets],
lectotype, designated by Verdcourt 1991).
Perennial herb, 0.6 m in height. Basal leaves 50–100×20–35 mm, lanceolate-
elliptic shaped, softly hairy, deciduous, margins entire. Stem leaves 40–100×9–20
mm, lanceolate shaped, apex acute, base cuneate, entire margins, densely covered
with white brittle hairs. Trichomes soft, upright, bulbous based. Inorescence
clustered at the apex, pedicels 5–8 mm long, lengthens considerably in fruit. Calyx
ca. 3 mm long, lobes ovate, grey pubescent, apex subacute. Corolla bluish purple;
lobe 7×9 mm diameter, segments round. Nutlets ovoid, 2–4×3–4 mm, convex
shaped; glochidia short, thick, marginal glochidia are more distinct than the central
glochidia (Figure 5).
Phenology. October to November.
Conservation status. Not evaluated (Raimondo et al. 2009).
Diagnostic characters. Amongst the southern African species, C. amabile can be
confused with C. lanceolatum due to their small-sized nutlets (between 2–4×2.5–4) and
owers. However, the two species are easily distinguished by their ower colour (C.
Figure 4. Distribution of Cynoglossum alticola based on the specimens examined.
Taxonomic revision of the genus Cynoglossum in southern Africa 19
lanceolatum has white corolla with blue throat, whereas C. amabile has bluish purple
corolla). is species was also reported by Stapf and Drummond (1906) and Kӧnig
et al. (2015) to be like C. furcatum Wallich (from Nepal, China, Bhutan, Vietnam,
ailand, Philippines, and India), based on ower and fruit size. e dierence can
be observed in the inorescences, whereby C. furcatum is a much larger plant with
inorescences up to 1 m tall, and C. amabile is up to 0.6 m tall.
Distribution and habitat. Cynoglossum amabile is widely distributed in southern
China where it is usually grown for ornamental purposes and naturalised in many
parts of the world (Xu et al. 2009). According to Germishuizen and Meyer (2003), this
species is only found in KwaZulu-Natal Province (Figure 6), where it grows in open,
disturbed sites, on gravel slopes and sandy, dry riverbanks.
Additional specimens examined. South Africa. KwaZulu-Natal: 2930 (Pietermar-
itzburg): Richmond District, Byrne Village (-CD), 23 Nov 1977, J. Stewart 2021 (NU).
Taxonomic notes. Cynoglossum amabile has been described as a widespread species
which grows in disturbed habitat and can be grown as an ornamental (Kӧnigetal.2015).
Figure 5. Cynoglossum amabile. SEM micrograph of the adaxial surface of A fruit nutlets B Glochidia.
Voucher specimen: J. Stewart 2021 (NU). SEM images scale bar: 2 mm (A); 100 µm (B).
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
20
is species has only been collected once in South Africa by J. Stewart in 1977, since
then there have been no later records or observations of this species in this region.
Attempts to locate this species in the wild were futile. It is questionable whether this
species occurs naturally in the southern African region since its single known locality is
within a protected area in KwaZulu-Natal.
3. Cynoglossum austroafricanum Weim. ex Hilliard & B.L. Burtt in Notes Roy.
Bot. Gard., Edinburgh 43(3):347 (1986).
Cynoglossum austroafricanum Weim., Jacot Guillarmod, Fl. Lesotho: 233 (1971); Gibbs
Russell et al., in Mem. Bot. Surv. S. Afr. 48:109 (1984), nom. nud.
Type. South Africa ♀♂, KwaZulu-Natal, Underberg (2929): Cobham Forest Reserve,
Sipongweni, c.6500ft (-CB), 21 Feb 1981, O.M. Hilliard and B.L. Burtt 14072 (E-
image! [2 sheets], holotype; K-image!; NU-image! [3 sheets], isotype).
Perennial or biennial herbs, 0.3–0.5 m in height. Basal leaves 100–190×15–
30 mm, lanceolate-obtuse, softly hairy, persistent margins entire. Stem leaves
45–100×10–21mm, narrowly lanceolate to linear lanceolate shaped, acute apex,
Figure 6. Recorded distribution of Cynoglossum amabile in southern Africa based on the specimens examined.
Taxonomic revision of the genus Cynoglossum in southern Africa 21
cuneate base, margins undulate, covered with sti hairs. Trichomes unicellu-
lar, with thick round base on the adaxial surface, simple on the abaxial surface.
Inorescences dichotomously branched, loose cymes at the apex, pedicels 4–9 mm
long, and lengthens considerably in fruit. Calyx ca. 2–3 mm long, lobes obtuse,
pubescent on the outer surface, glabrous on the inner surface, apex acute. Corolla
pale blue; lobes 2.75–4.25 mm in diameter, cruciform, apex obtuse. Nutlets ovoid,
3.0–3.5×2.5–3.0 mm; glochidia more spread towards the margins, thin, tip multi-
angular (Figure 7).
Phenology. December to April.
Conservation status. Least Concern (Raimondo et al. 2009).
Diagnostic characters. Amongst the southern African species, Cynoglossum aus-
troafricanum can be confused with either C. lanceolatum or C. coeruleum var. mannii.
is species diers from the two by the colour of the corolla (white corolla with pale
blue throat vs pale blue corolla throughout in C. austroafricanum). is latter observa-
tion was also noted by Hilliard and Burtt (1986).
Figure 7. Vegetative and reproductive morphological features of Cynoglossum austroafricanum A line
drawing of the branching pattern of the fruit stalk, and the alternating stem leaves B SEM micrograph of
the adaxial surface of a fruit nutlet, with the arrangement of glochidia around the nutlet C SEM micro-
graph of the glochidia. Voucher specimen: O.M. Hilliard and B.L. Burtt 11803 (PRE). Drawing scale bar:
7.5 mm. SEM scale bar: 1 mm (B); 100 µm (C).
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
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Distribution and habitat. e species is distributed in South Africa (North-West,
Gauteng, Mpumalanga, Free-State, KwaZulu-Natal, and Eastern Cape Provinces), eS-
watini and Lesotho (Figure 8), where it occurs in shady, disturbed areas, and sandy,
dry riverbanks.
Additional specimens examined. South Africa. L: 2430 (Tzaneen):
Lekgalameetse Nature Reserve (-AA), 7 Oct 1986, N. Stalmans 1404 (PRE). N-
W: 2626 (Lichtenburg): Lichtenburg (-AA), Feb 1918, L. Kretzshmar 17065 (PRE).
G: 2528 (Pretoria): Brooklyn (-CA), 1 Feb 1931, A.O.D. Mogg 16451 (PRE).
M: 2430 (Pilgrim’s rest): Pilgrim’s rest, next to the old railway line (DD),
31 Jan 2019, A.N. Moteetee and L.K. Madika AL06 (JRAU). 2530 (Mashishing):
Mashishing (-AB), Apr 1910, M. Crosby 1989 (PRE); Mashishing District (-CA),
8Feb1904, J. Burtt-Davy 1472 (PRE). 2630 (Carolina): Between Oshoek border post
and Carolina (-BA), Jan 1906, H. Bolus 12161 (PRE). F S: 2828 (Bethlehem):
Golden Gate (-AB), 22 Jan 1951, A. Wiezer 22485 (NBG). (Fouriesburg): Bethlehem
(-CA), 8 Jan 1918, G.Potts 3246 (PRE); Witsieshoek (-DB), 28 Feb 1975, O.M.
Hilliard and B.L. Burtt 8665 (NU). K-N: 2730 (Vryheid): Oshoek
District, Wakkerstroom (-AC), 18 Jan 1961, N.J. Devenish 480 (PRE); Amajoba
District Municipality area, Luiperdkloof farm, Natural Heritage site no.47 (-AD), 25
Figure 8. Known distribution of Cynoglossum austroafricanum in southern Africa based on the specimens
examined.
Taxonomic revision of the genus Cynoglossum in southern Africa 23
Jan 2011, A.M. Ngwenya 3601 (NH). 2829 (Harrismith): Cathedral Peak (-CC), 18
Feb 1983, O.M. Hilliard and B.L. Burtt 16298 (NU); 12 Jan 1984, J. Scott 76 (NH).
2830 (Dundee): Hattingspruit Station (-AA), Dec 1929, D. Johnston 268 (NU).
2929 (Underberg): Mpendhle District, upper Loteni Valley (-AD), 5 Feb 1985, O.M.
Hilliard and B.L. Burtt 18103 (NU, PRE); Loteni Nature Reserve (-BC), 24 Dec 1978,
O.M. Hilliard and B.L. Burtt 11803 (PRE); Polela River (-CB), 21 Apr 1973, M.A.
Rennie 379 (NU); 23 Mar 1977, O.M. Hilliard and B.L. Burtt 9793 (NU); 17 Feb
1982, O.M. Hilliard and B.L. Burtt 15520 (NU). 2930 (Pietermaritzburg): Gate farm
Keerom-Cottingham (-CC), 23 Mar 1969, R.G. Strey 8420 (NH). 2931 (Stanger):
Tugela valley (-AA), 14 Feb 1926, Bayer 47 (NU). 3029 (Kokstad): Hillside (-CA),
Jan 1956, P. ompson 2 (NU); on banks of streams near Kokstad (-CB), Dec 1883,
W. Tyson 1839 (NBG), 17 Jan 1957, L.E. Taylor 5473 (NBG). 3030 (Dumisa): Port
Shepstone (-AD), 22 Oct 1997, J. Arkell 353 (NH). E C: 3128 (Umtata):
Transkei, on the summit of Baziya Mt. (-AD), 23 February 1988, T. Strever 917 (PRE);
Walter Sisulu University, Area 3: East of In-Service centre (-DB), 14Feb 2001, N.
Nombekela 102 (NH). 3126 (Queenstown): Hangklip (-DD), Feb 1960, H. Koepowitz
13147 (GRA). 3129 (Port St. John’s): Ntabankulu Mountain, Gome Forest Station,
Tabankulu (-AB), 11 Nov 1996, T. Dold, E. Cloete and R. White 2940 (GRA). 3227
(Stutterheim): Fort Cunynghame Station (-AD), Nov 1894, T.R. Sim 1860 (NU); no
date 1897, T.R. Sim 20420 (NU).
eSwatini. 2631 (Mbabane): Forbes Reef (-AA), 14 Apr 1960, R.H. Compton 30035
(NBG); Mbabane (-AC), 17 Jan 1951, A. Wuze 22393 (NBG).
Lesotho. 2828 (Bethlehem): Leribe District, LHDA Phase 1A (-AD), 11 Jan 1996,
P.B. Phillipson, C. Mokuku, R. Judd, and C. Hobson 4473 (GRA); Leribe (-AD), no
date, M. Dieterlen 70 (NBG; NH). 2927 (Maseru): aba Bosiu (-BC), 1 Mar 1978,
M. Schmitz 8205 (PRE); Mahlatsa (-BB), 18 Jan 1941, A. JacotGuillarmod 51 (GRA);
Sekeng Ha Fako (-BD), Oct 2018, A.N. Moteetee 56 (JRAU); 30 Dec 2018, A.N.
Moteetee 59 (JRAU). 2929 (Underberg): Mokhotlong (-AC), Mar 1949, A. Jacot Guil-
larmod 1072 (PRE), 25 Feb 1949, W.J. Barker 21515 (NBG).
No locality details: 27 Nov 1888, H. Medley 4576 (NH); Dec 1946, E. Mes-
ton50(NU).
Taxonomic notes. According to Hilliard and Burtt (1986), this species was de-
scribed by Dr. H. Weimarck who did not select a type specimen; therefore, in typifying
it they retained the specic epithet.
4. Cynoglossum coeruleum subsp. johnstonii var. mannii (Baker & C.H. Wright)
Verdc. in Fl.Trop. E. Africa, Boraginac.: 110 (1991).
Cynoglossum mannii Baker & C.H.Wright, in Oliver et al., Fl. Trop. Afr., 4(2.1): 52
(1905) ♀♂, Type as above.
Cynoglossum geometricum Baker & C.H. Wright, in Oliver et al., Fl. Trop. Afr. 4(2.1):
52 (1905). Type: Nyasaland [Malawi] ♀♂ Mount Chiradzulu, no date, A. Whyte
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
24
s.n. (K-image! lectotype, here designated). Notes: e specimen was selected as a
lectotype by B.L. Burtt on the sheet but was never designated formally.
Paracynoglossum geometricum (Baker & C.H. Wright) R.R. Mill. in Notes Royal. Bot.
Gard., Edinb. 41 (3): 478 (1984). ♀♂, Type: Same as above.
Cynoglossum coeruleum subsp. geometricum (Baker & C.H. Wright) S. Edwards in Fl.
Ethiopia & Eritrea 5: 93 (2006), nom. inval.
Type. Cameroon ♀♂, Mount Cameroon, Dec 1862, Mann 2005 (K-image, lectotype!
here designated; K-image [2 sheets] isolectotype!).
Perennial, biennial, or annual herbs, 1.2 m in height. Basal leaves 90–190×28–56mm,
lanceolate-obtuse, softly hairy, deciduous, margins entire. Stem leaves 35–90×7–25 mm,
lanceolate, apex acute, base acute to obtuse, covered with moderately sti hairs margins
entire. Trichomes bulbous based on the upper surface of the leaf, sometimes simple on
the lower surface. Inorescences terminal axillary cymes, few branches spreading dichoto-
mously; pedicel 4–8 mm long, lengthens considerably in fruit. Calyx ca. 21 mm long, lobes
ovate-oblong, adpressed-hairy outside, smooth inside, apex acute. Corolla white with pale
blue throat; lobes ca. 2.1 mm in diameter, campanulate. Nutlets ovoid, 3–4×2.5–3.5mm;
glochidia more marginal and on the median line (Figure 9).
Phenology. December to April.
Conservation status. Least Concern (Raimondo et al. 2009).
Diagnostic characters. is variety can be easily confused with C. lanceolatum
due to the dichotomous branching of the inorescence but can be distinguished from
it by the distribution and density of the glochidia in the nutlets. e glochidia on the
nutlets of Cynoglossum coeruleum var. mannii are more marginal and on the median
line, whereas they are equally distributed around the whole nutlet in C. lanceolatum.
Distribution and habitat. is variety is endemic to South Africa where it is
known only from KwaZulu-Natal and Eastern Cape Provinces (Figure 10). It is also
reported from Malawi, Mozambique, Zambia, and Zimbabwe (Mill and Miller 1984).
It is found in disturbed grassland and sandy areas.
Additional specimens examined. South Africa. K-N: 2731
(Louwsburg): Itala Nature Reserve (-AD), 10 Dec 1987, M. Jordaan 7064 (NH); 9 Dec
1987, A.G. Hutchings 2530 (NU); Zululand District Municipality Area, Abaqulusi
Municipality Area, Tygerskloof Farm (-CD), 24 Jan 2012, A.M. Ngwenya and D.G.A.
Styles 4034 (NH); 7 Mar 2019, A.N. Moteetee and L.K. Madika AL010 (JRAU). 2828
(Bethlehem): Royal Natal National Park (-DB), 17 February 1984, O.M. Hilliard and
B.L. Burtt 17658 (NBG; NU). 2831 (Nkandla): Nhlazatshe farm, (-AA), 4 Mar 1994,
T.B. Sikhakhane 440 (NH). 2929 (Underberg): Mpendhle District, Loteni Nature
Reserve (-AD), 1 Feb 2001, A.M. Ngwenya 1940 (NH), Loteni, upper reaches of river
(-BC), 31 Mar 1984, O.M. Hilliard 8218 (NU, PRE), Sipongweni Mountain (-CD),
20 Mar 1987, O.M. Hilliard and B.L. Burtt 8249 (PRE). 2930 (Pietermaritzburg): On
the Phezulu Game Estate, Botha’s Hill (-DC), 22 Jan 2005, D. Styles 2280 (NH). 3030
(Port Shepstone): M. Stainbank’s farm, mid Illovo (-BB), 23 Dec 2008, A. Young 942
(NU). E C: 3023 (Britstown): Kamberg (-CC), 21 Mar 1983, O.M. Hilliard
Taxonomic revision of the genus Cynoglossum in southern Africa 25
8208 (NU). 3128 (Umtata): Baziya Mission (-CB), 12 Feb 1981, O.M. Hilliard and
B.L. Burtt 13952 (NU); Nenga River, (-DB), 26 Oct 2001, E. Cloete 6342 (GRA).
3226 (Fort Beaufort): Hogsback (-DB), Apr 1956, R. Collett 9775 (GRA); Apr 1955,
A.R.H. Martin 9678 (GRA); Apr 1962, A. Jacot Guillarmod 5544 (GRA); 12 Apr
1955, L.M. Johnson 1152 (GRA); 4 Mar 1973, M. Bradley 55 (GRA).
Taxonomic notes. (i) Cynoglossum geometricum was recorded in the southern African
region (FSA) for the rst time by Hilliard and Burtt (1986). Although the name seems
to be accepted in the Flora of southern African region (FSA), for example, Germishuizen
and Meyer (2003) and Burrows and Willis (2005), it is not the correct name for this
taxon. e correct name is Cynoglossum coeruleum subsp. johnstonii var. mannii, as synoni-
mized by Verdcourt (1991). It is worth noting that the latter name is erroneously listed as
C. coeruleum var. mannii in websites such as Plants of the World Online (http://powo.sci-
Figure 9. Vegetative and reproductive morphological features of Cynoglossum coeruleum var. mannii
Aline drawing of dichotomous branching of the fruit stalk B marginal and median line glochidia on
the adaxial surface of a nutlet C Glochidia evenly sized. Voucher specimen: T.B. Sikhakhane 440 (NH).
Drawing scale bar: 7.5 mm. SEM scale bar: 2 mm (B); 200 µm (C).
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
26
ence.kew.org/taxon/966730-1) and the World Flora Online (http://www.worldoraon-
line.org/). Verdcourt (1991) relegated C. johnstonii to subspecies level under C. coeruleum
and then transferred C. mannii to C. coeruleum as a variety of Cynoglossum coeruleum
subsp. johnstonii. (ii) ere are three sheets of Mann 2005 in Royal Botanic Gardens, Kew
(K), the one with the barcode number K000418935 is chosen as a lectotype because it
displays most of the important characters of the species which can be used to distinguish
this species from the rest, such as the inorescence character and the branching pattern.
5. Cynoglossum hispidum unb., Prodr. Plant. Cap. 1:34 (1794); Roem-
er and Schultes in Syst. Veg., ed. 15 bis 4: 79,761 (1819); C.H. Wright in F.C.
4(2):14(1904); Brand in Engl. Planzenr. 78 [4,252]:146 (1921).
Cynoglossum glomeratum Pursh in Fl. Amer. Sept. 2:729 (1813).
Type. United States of America ♀♂, Louisiana, no date, Bradbury s.n. (PH-image!
holotype). Cynoglossum enerve Turcz. Bull. Soc. Imp. Naturalistes Moscou 1:259
(1840), E.Mey. ex DC., Prodr.10:154 (1846).
Type. South Africa. ♀♂, Eastern Cape, between Omcamwubo and Omcamcaba, no
date, Drѐge d (HAL-image! lectotype, here designated; GDC-image! [3 sheets],
Figure 10. Known distribution of Cynoglossum coeruleum var. mannii in southern Africa based on the
specimens examined.
Taxonomic revision of the genus Cynoglossum in southern Africa 27
isolectotype). [Note: e HAL specimen is chosen as a lectotype because the speci-
men displays the diagnostic characters of the species as described in the protologue].
Echinospermum enerve E.Mey. ex DC. Prodr. 10:154 (1846), nom. nud.
Type. South Africa ♀♂, Precise locality unknown, Lange Kloof, unberg 168 sub
THUNB-UPS 3996 (UPS, microche! holotype).
Perennial or biennial herbs, 0.5–0.76 m in height. Basal leaves 80–250×15–25
mm, lanceolate-obtuse, densely pubescent, deciduous, margins entire. Stem leaves 60–
80×5–12 mm, oblong-lanceolate, apex acute, base cuneate, covered with brittle hairs,
margins entire. Trichomes bulbous based on the upper surface of the leaf, sometimes
simple on the lower surface. Inorescence terminal axillary cyme, branches spread-
ing dichotomously; pedicel up to 20 mm long, lengthens considerably in fruit. Calyx
ca. 5–10 mm long, lobes obtuse, outer surface packed with bulbous-based trichomes,
apex acute. Corolla magenta; lobes ca. 5 mm in diameter, cruciform. Nutlets convex,
5–6×3–5 mm, highly pubescent; glochidia short and thick at the base, tips multian-
gular (Figure 11).
Phenology. October to March.
Conservation status. Least Concern (Raimondo et al. 2009).
Diagnostic characters. e species can be confused with C. lanceolatum with
which it shares a similar branching pattern of the inorescences and upright brittle
hairs covering the whole plant. However, the two dier in the colour of the corolla
(magenta-purplish vs. white with blue throat in C. lanceolatum) and pedicel length (2
cm long opposed to less than 2 cm long in C. lanceolatum).
Distribution and habitat. is species is widely distributed across all provinces
of South Africa. It can also be found in eSwatini and Lesotho (Figure 12). It mostly
occurs in open grasslands, grassy slopes, woodland marshes, and disturbed areas like
abandoned lands.
Additional specimens examined. South Africa. L: 2229 (Waterpoort):
Evelyn valley (-BD), Feb 1944, RUC Biology Expedition 417 (GRA). 2430 (Tza-
neen): Lekgalameetse Nature Reserve (-AA), 4 Dec 1985, M. Stalmans 790 (PRE);
Sekhukhune District, Leolo Mountains (-CA), 14 Mar 2007, B. Sachse 471 (PRE).
N-W: 2523 (Pomfret): Forbes (-DA), 29 Oct 1959, B. Dlamini s.n. (NH).
2627 (Potchefstroom): Goedgedacht (-AA), 1 May 1932, J.D. Sutton 676 (PRE); Los-
berg, Elandsfontein (-BC), 11 Dec 1934, J.J. eron 725 (NH); Potchefstroom (-CA),
23Nov 1946, W.J. Louw 1525 (PRE). G: 2528 (Pretoria): Brooklyn (-CA), 16
Sep 1928, A.O.D. Mogg 15247 (PRE); 6 Apr 2019, A.N. Moteetee and L.K. Madika
AL014 (JRAU); 1931, A.O.D. Mogg 16601 (PRE); 30 Sep 1943, A.O.D. Mogg 17001
(PRE); Doornpoort/ Hartbeesfontein (-CB), 24 Jan 2004, S.P. Bester 4656 (PRE);
Irene District (-CC), Oct 1929, A.A. Obermeyer 27651 (PRE). 2628 (Johannesburg):
Vereeniging District (-AC), 23 Nov 2007, S.P. Bester 8268 (PRE); Suikkerbosrand
Nature Reserve (-AD), 7 Apr 1970, A. Lambrechts 265 (PRE). M: 2530
(Mashishing): Boschhoek (-AA), 16 Nov 1933, R.S.N. Young A375 (PRE); Verloren-
kloof Reserve, Welgedacht (-AD), 28 Nov 2008, S.P. Bester 8679A (PRE); Machado-
dorp, Farm Grootvlei (-CB), 21 Oct 1988, P. Burgoyne 457 (PRE); Songimvelo Game
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
28
Reserve (-DD), 10 Dec 1992, M. Jordaan 2490 (PRE). 2629 (Bethal): Bethal (-AD), 14
Dec2010, R. Leendertz 9386 (PRE). 2630 (Carolina): Ermelo District, Spitskop (-CA),
Dec 1915, R. Potts 5007 (PRE). N C: 2820 (Kakamas): Krantzkop (-DA),
Nov 1911, J. ode 4768 (NBG). F S. 2728 (Frankfort): Farm Reitfontein
(-BC), 28 Jan 1983, E. Retief 1071 (PRE). 2828 (Bethlehem): Qwaqwa National Park
(-BC), 22 Nov 1994, P.C. Zietsman 2558 (NH); Golden Gate National Park (-DA),
12 Dec 1988, Gertenbach and Groenewald 8929 (PRE); Witsieshoek (-DB), 10 Mar
2015, S. Parbhoo 81 (NH). 2829 (Harrismth): Harrismith (-AC), 17 Mar 1981, M.L.
Jacobsz 3092 (PRE); 18 Nov 1978, M.L. Jacobsz 1299 (PRE); Rensburgskop (-AD), 10
Dec 1962, M.L. Jacobsz 199 (NBG; PRE). K-N: 2730 (Vryheid): Oshoek
District, Wakkerstroom (-AD), 19 Nov 1962, N.J. Devenish 954 (PRE); Klipspruit
Dam (-DD), 10 Feb 2005, S.P. Bester 6540 (NH, PRE). 2731 (Louwsburg): Zululand
District municipality (-CD), 29 Oct 2000, T. Edwards and C. Potgieter s.n. (NU); 26
Jan 2012, A.M. Ngwenya and D.G.A. Styles 4085 (NH). 2732 (Ubombo): Ingwavuma
Figure 11. Vegetative and reproductive morphological features of Cynoglossum hispidum A line drawing
of obtuse shaped, rosette base leaves, terminal branched fruits, terminal owers B densely packed with
glochidia nutlets C Glochidia wide at the base, with multiangular tip. Voucher specimen: S.P. Bester
12958 (PRE). Drawing scale bar: 7.5 mm. SEM scale bar: 5 mm; 100 µm (C).
Taxonomic revision of the genus Cynoglossum in southern Africa 29
(-AA), 22 Nov 1969, E.J. Moll 4680 (NH, PRE). 2829 (Harrismith): Cathedral Peak
(-CC), 10 Nov 1956, D.J.B. Killick 1115 (PRE); 13 Oct 1984, J. Scott 251 (NH). 2830
(Dundee): Dundee District (-AA), 26 Nov 1964, N.E. Shirley s.n. (NU); Klipriver Dis-
trict, Elandslaagte (-CD), 23 Oct 1964, N.E. Shirley s.n. (NU). 2831 (Nkandla): Em-
pangeni (-DB), 9 Jul 1965, H.J.T. Venter 1917 (PRE). 2929 (Underberg): Giants Castle
Game Reserve (-AB), 8 Feb 1966, W.R. Trauseld 575 (PRE); (-AD), 8 Nov 2001, T.R.
Green 1217 (NU); Mpendhle District (-BC), 5 Jan 1983, O.M. Hilliard and B.L. Burtt
16218 (PRE); Underberg District (-CB), 3 Feb 1975, O.M. Hilliard and B.L. Burtt
7942 (NU); 3 Feb 1976, O.M. Hilliard and B.L. Burtt 8905 (NU); 24 Mar 1977, O.M.
Hilliard and B.L. Burtt 9818 (NU); 11 Jan 1978, M.A. Rennie 911 (NU); 27 Jan 1982,
M.A. Rennie 1307 (NU); 13 Feb 1983, O.M. Hilliard and B.L. Burtt 17234 (NU); 6
Jan 1984, O.M. Hilliard and B.L. Burtt 17290 (PRE); Underberg District (-CC), 19
Jan 1984, O.M. Hilliard and B.L. Burtt 17346 (NU); 9 Jan 1986, O.M. Hilliard and
B.L. Burtt 18998 (NU); Hlogoma Mountain (-DC), 30 Nov 2015, S.M. Berruti 513
(NH); Donnybrook (-DD), 7 Nov 2013, D.G.A Styles 4579 (NH); 8 Mar 2019, A.N.
Moteetee and L.K. Madika AL011 (JRAU). 2930 (Pietermaritzburg): Lidgetton (-AC),
23 Mar 1920, A.O.D. Mogg 6894 (PRE); 29 Sep 1964, E.J. Moll 1036 (NU, PRE);
13 Jan 1988, B. Grove 98 (NU); Umgeni River (-CA), 20 Oct 1984, J. Manning 538
Figure 12. Known distribution of Cynoglossum hispidum in southern Africa based on the specimens
examined.
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
30
(NU); 8 Mar 2019, A.N. Moteetee and L.K. Madika AL013 (JRAU); Ukulinga farm
(-CB), 8 Mar 1982, J.C. Manning 212 (NU); Nagle Dam (-DA), 15 Sep 1957, M.J.
Wells 1676 (NU); Clidale road (-DC), 17 Aug 2002, D.G.A. Styles 9141 (NU); 30
Jul 2003, P. Wragg 205 (NU),7 Nov 2014, D.G.A. Styles 4925 (NH). 2931 (Stanger):
Groutville (-AD), 14 Oct 1965, E.J. Moll 2500 (NU, PRE). 3029 (Kokstad): Insizwa
(-CA), 24 Feb 1972, R.G. Strey 10827 (NU); Kokstad District (-DA), 1968, C.J. Piek
53 (NH); 25 Feb 1978, T.A. Coleman 985 (NH); no date, F.M. Getlie and T. Edwards
1266 (NU). 3030 (Port Shepstone): M. Stainbank’s farm, mid Illovo (-BB), 24 Sep
2009, A. Young 1155 (NU); Umzinto District (-BC), 3Sep 1983, K. Balkwill and J.C.
Manning 828 (NU); 20 Oct 1997, A.M. Ngwenya 1538 (NH); Port Shepstone (-CB),
3 Oct 1937, A.O.D. Mogg 13873 (PRE); (-CD), 20 Sep 1955, S. McNeil 142 (NU).
3030 (Umzinto): Vernon Crookes Nature Reserve (-BC), 3 Sep 1983, K. Balkwill and
J.C. Manning 808 (NU); 29 Sep 1984, C.F. Kennedy 32 (NU). W C: 3219
(Cape Town): Rivierground in pypsteelbosse (-CC), 18 Dec 1980, W.J. Hanekom 2600
(PRE); Kouebokkeveld Berge, (-CC), 24 Nov 1998, W.J. Hanekom 3120 (NBG). 3322
(Oudtshoorn): Outeniqua Mountains (-CC), 9 Nov 1986, J.H.J Vlok 1693 (PRE).
E C: 3027 (Barkly East): Farm Faskally near England (-DA), 9 Nov 1995,
J.E. Victor 1586 (PRE); Ben McDui (-DC), 6 Jan 1997, T. Dold and M. Cocks 3495
(GRA). 3028 (Umtata): Maclear District (-CA), 16 Jan 2016, S.P. Bester 13207 (PRE);
Maclear (-CC), 7 Nov 1993, S.P. Bester 1535 (NH); Kloof (-DD), 28 Feb 1946, R.
Story 950 (PRE). 3128 (Umtata): Hill above Mhlanfane Forest Station (-BC), 31 Jan
1983, O.M. Hilliard and B.L. Burtt 16341 (NU); Lady Frere, Engcobo (-CC), 25 Nov
1990, E. Cloete 590 (NH); (-DB), 21 Oct 1953, G.C. eron 1604 (PRE); Elliot-
dale District (-DC), 11 Jul 1966, J.L. Gordon-Gray 529 (NH). 3129 (Umtata): Baziya,
Tembuland (-CC), no date, Baur 257 (K-image); Blesbok ats Cathcart Div. Cape,
1838, Drége s.n. (K-image); between St John’s River and Umtsikaba River, Pondoland,
1838, Drége s.n (K-image!). 3225 (Elandsfontein): Elandsfontein (-AA), 13 Dec 2005,
S.P. Bester 6347 (GRA, PRE). 3226 (Fort Beaufort): Along gravel road to Sada o the
R67 (-BD), 8 Feb 1995, J.E. Victor and D.B. Haare 392 (PRE); Mpofu Game Reserve
(-DA), 28 Feb 2006, C.L. Bredenkamp 3338 (PRE); Menzieberg, Amatole Mountains
(-DB), 6 Jan 1986, P.B. Phillipson 1185 (GRA), Fort Beaufort Road, 2 miles (3.22 km)
from Alice (-DD), 22 Oct 1939, M.H. Gien 264 (PRE). 3227 (Stutterheim): Stutter-
heim (-AD), 5 Dec 1942, J.P.H. Acocks 9399 (PRE); Pirie Mission (-CC), exact date not
provided 1888, T.R. Sim 20419 (NU, PRE); 5 Aug 2014, S. Mgcuwa 117 (GRA); King
William’s Town (-DB), Nov 1891, H.G. Flanagan 1193 (PRE). 3326 (Grahamstown):
Faraway (-AD), 4 Dec 1980, A. Jacot Guillarmod 8473 (GRA); Nov 1942, E. Archibald
669 (GRA); Alexandria (-CB), 10 Aug 1953, S. Johnson 691 (GRA); 11 Feb 1953, W.
Marais 185 (PRE). 3327 (Peddie): Peddie District (-AC), 5 Nov 1993, T. Dold and A.
Booi 481 (GRA).
eSwatini. 2631 (Mbabane): Forbes reef (-AA), 29 Oct 1959, B. Dlamini s.n. (NBG).
Lesotho. 2928 (Marakabei): Ntiboho valley (-AC), 2 Jan 1947, A. Jacot Guillar-
mod 279 (GRA). 2929 (Underberg): aba Ntšo, Sehlabathebe National Park (-CC),
4–14 Jan 1973, Jacot Guillarmod, Getlie and Mzamane 65 (PRE); 13 Feb 1976, A.
Beverly 471 (PRE).
Taxonomic revision of the genus Cynoglossum in southern Africa 31
Unknown localities: Feb 1895, Maurice and Evans 474 (NH); Feb 1939, J. Wylie
30120 (NH); Apr 1943, B. Fischer 464 (NU); 15 Dec 1943, W.F. Barker 2796 (NBG);
18 Nov1952, H.B. Gilliland 26862 (PRE); 18 Dec 1966, R.G. Strey 7047 (NH); Mar
2000, T. Edwards 2088 (NU).
Taxonomic notes. Brand (1921) included C. hispidum as a synonym of C. lanceo-
latum. However, the two species have notable variations especially in the colour of the
corolla (magenta-purplish vs. white with blue throat in C. lanceolatum) and pedicel
length (2 cm long opposed to less than 2 cm long in C. lanceolatum).
6 Cynoglossum lanceolatum Forssk in Fl. Aegypt. Arab: 41 (1775).
Cynoglossum hirsutum unb., Prodr. Pl. Cap.: 34 (1794).
Type: South Africa ♀♂, Precise locality unknown: Roggerveld, C.P. unberg 168, sub
THUNB-UPS 3995 (UPS-microche! holotype).
Cynoglossum micranthum Desf.: 220 (1804), nom. nud.
Cynoglossum canescens Willdenow in Enum. Pl.: 180 (1809).
Type: ♀♂, Precise locality unknown, C.L. Willdenow, s.n. (B-W,-image, lectotype, des-
ignated by Kӧnig et al. 2015).
Cynoglossum racemosum Roxb. in Fl. Ind.: 2:6 (1824), nom. illeg.
Echinospermum paniculatum E.Mey. ex. A.DC., Prodr. [A.P.de Candolle] 10: 149,143
(1846). ♀♂, Type: same as above.
Paracynoglossum lanceolatum (Forssk.) R.R.Mill. in Notes Roy. Bot. Gard. Edinb.: 474
(1984). ♀♂Type: same as above.
Type. Yemen, Al Hadiyah, Mar 1763, P. Forsskal 312 (C-image! holotype).
Annual or biennial herbs, 0.5–0.9 m in height, covered with simple hairs. Ba-
sal leaves 85–180×15–23 mm, lanceolate-obtuse, blade elliptic, softly hairy, decidu-
ous. Stem leaves 40–65×8–18 mm, lanceolate, apex acute, base cuneate, covered with
moderately sti hairs. Trichomes brittle, simple on both leaf surfaces. Inorescence
dichotomously branched axillary cyme, pedicel 1–1.5 mm long. Calyx ca. 1–1.5 mm
long, lobe ovate-obtuse, pubescent on the outer surface, inner surface glabrous, apex
acute. Corolla white with pale blue throat; lobes ca. 1×1 mm, campanulate. Nutlets
ovoid-convex, 3–4×2.5–3.5 mm, fully pubescent; glochidia equally thick and long
(Figure 13).
Phenology. August to May.
Conservation status. Least Concern (Raimondo et al. 2009).
Diagnostic characters. Amongst the southern African species, C. lanceolatum is
similar and possibly related to Cynoglossum coeruleum subsp. johnstonii var. mannii
with which it shares the branching pattern of the inorescence, ower colour, and
nutlet size. e two species can be distinguished by the density of the glochidia on the
nutlets and distribution. Cynoglossum lanceolatum nutlets are completely covered with
glochidia, whereas in Cynoglossum coeruleum subsp. johnstonii var. mannii glochidia
tend to be more marginal and acentric.
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
32
Distribution and habitat. Cynoglossum lanceolatum originates from Yemen (Kӧnig
et al. 2015) but reported from Africa (Ge-Ling et al. 1995), Pakistan, India (Joshi
2016), the Mediterranean, and throughout Asia (Verdcourt 1991) and Madagascar
(Kӧnig et al.2015). In South Africa it occurs widely in all provinces, it also occurs in
eSwatini and Lesotho (Figure 14). It is a widespread species that grows in disturbed
habitats throughout parts of Africa.
Additional specimens examined. South Africa. L: 2229 (Waterpoort):
Wylies Poort (-DD), 17 Oct 1938, J.P.M. Acocks 1295 (PRE). 2230 (Messina): Sibasa
District (-CD), 21 Jun 1977, G. Hemm 163 (PRE). 2329 (Polokwane): Polokwane
Nature Reserve (-CD), 12 Jan 1979, Bredenkamp and Van Vuuren 307 (PRE). 2330
(Tzaneen): Letaba (-CA), 2 Apr 1958, J.C. Scheepers 219 (PRE); Woodbush Forest
Reserve (-CC), Jan 1923, H. Wafen 22973 (PRE); 1 Feb 2019, A.N. Moteetee and
L.K. Madika AL07 (JRAU). 2428 (Modimolle): Waterberg, Kwaggasnek-Alma Road
Figure 13. Morphological characters of Cynoglossum lanceolatum A line drawing of the fruiting branch
B Glochidia evenly distributed on the adaxial surface of the nutlet C Glochidia uniform size, with multi-
angular tip. Voucher specimen: S.P. Bester 4653 (PRE). Drawing scale bar: 7.0 mm. SEM scale bar:
1mm(B); 100 µm (C).
Taxonomic revision of the genus Cynoglossum in southern Africa 33
(-CC), 13 Feb 1981, K.L. Immelman 118a (PRE). 2429 (Zebediela): Zebediela (-AA),
5 Feb 1967, B.J. Huntley 1033 (PRE); 2430 (Pilgrim’s Rest): Sekhukhune District Mu-
nicipality, Leolo Mountain, SE of Tama Kgoshi (-CA), 14 Mar 2007, B. Sachse 466
(PRE). N-W: 2526 (Zeerust): Zeerust along road to Koster (-CA), 4 Feb 1976,
F. van der Meulen 597 (PRE). 2527 (Rustenburg): Rustenburg (-CA), 27 Dec 1949,
S.M. Johnson 854 (NBG). 2626 (Lichtenburg): Municipal Caravan Park (-AA), 19 Mar
1978, A. Balsinhas and Harding 3303 (PRE). 2627 (Potchefstroom): Carletonville, A.
Bailey Nature Reserve (-AD), Apr 1983, S. van Wyk 180 (PRE); Mooiriver (-CA), 9
Mar 1984, B. Ubbink 1257 (PRE). 2724 (Taung): West of Harz River near Taung,
120 km north of Kimberley (-DB), 11 Feb 1948, R.J. Robin 3628 (PRE). G:
2528 (Pretoria): Brooklyn (-CA), 20 May 1915, C.A. Smith 191 (PRE); 24 May 1915,
C.A. Smith 194A (PRE); 10 Dec 1915, A.O.D. Mogg 11846 (PRE); Nov 1925, C.A.
Smith 1220 (PRE); Doornpoort, Airport Road (-CA), 26 Oct 2009, S.P. Bester 9734
(PRE); 14 Nov 2018, A.N. Moteetee and L.K. Madika AL01 (JRAU); Doornpoort/
Hartbeesfontein (-CB), 24 Jan 2004, S.P. Bester 4653 (PRE); Brooklyn (-CC), 11 Apr
1920, K.A. Landsdell 864 (PRE). 2628 (Johannesburg): Randburg, Lanseria (-AA), 20
Nov 2018, A.N. Moteetee and L.K. Madika AL02 (JRAU); Suikkersbosrand (-CB), 27
Figure 14. Known distribution of Cynoglossum lanceolatum in southern Africa based on the specimens
examined.
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
34
Dec 1971, G.J. Bredenkamp 594 (PRE). M: 2429 (Zebediela): Mashishing
District (-DD), 8 January 1939, Barnard and Mogg 860 (PRE). 2430 (Pilgrim’s Rest):
Malta near Marinella (-AA), 16 Aug 1984, M. Stalmans 114 (PRE); Ohrigstad Dam
Nature Reserve (-DC), 22 May 1973, N. Jacobsen 2861 (PRE). 2529 (Emalahleni): Lo-
skop Dam (-AD), 12 Jan 1967, G.K. eron 1129 (PRE). 2530 (Mashishing): Witklip
(-BD), 4 Dec 1973, J.P. Kluge 359 (PRE); Lowveld Botanical Gardens (-BD), 18 Dec
1974, E.J. Van Jaarsveld 199 (NBG); Waterval Onder, Ntsinini (-CD), 26 Dec 2008,
K.W. Grieve 250 (PRE); 23 Jan 2011, K.W. Grieve 342 (PRE); Barberton (-DD), 24
Jan 2000, J.J. Meyer 2603 (PRE). 2531 (Beersrust): White River District (-AC), Apr
1983, W. Jacobsen 5363 (PRE). 2630 (Carolina): Chrissiesmeer (-AC), 5 Jan 1972,
G.K. eron 2402 (PRE); Chrissiesmeer, on road to Lochiel (-BA), 6 Mar 1986, M.
Crosby 273 (PRE). 2730 (Vryheid): Farm Doornhoek along Assegaai river (-AA), 15
Dec 2006, S.J. Siebert 3246 (PRE). F S: 2729 (Volksrust): Vrede (-CB), 5 Feb
1987, L. Smook 6455 (PRE). 2828 (Bethlehem): Golden Gate National Park (-DA),
Jan 1963, L.C.C. Liebenberg 6831 (PRE); Witsieshoek (-DB), Mar 1917, H.A. Janod
17317 (PRE). 2829 (Harrismith): Harrismith District (-AC), 8 Mar 1974, M.L. Jacobsz
2036 (PRE). K-N: 2729 (Utrecht): Volksrust (-BD), Jan 1928, C.A. Smith
5730 (PRE). 2730 (Vryheid): Oshoek District, Wakkerstroom (-AB), 26 Dec 2002,
S.J. Siebert and F. Siebert 2279 (PRE); Vryheid Nature Reserve (-DC), 23 Feb 1988,
C.J. Youthed 11 (NH); 7 March 2019, A.N. Moteetee and L.K. Madika AL09 (JRAU).
2731 (Louwsburg): Louwsburg District, Itala Nature Reserve (-CB), 15 Jan 1978, D.J.
Mcdonald 443 (PRE). 2828 (Bethlehem): Crocodile River (-DB), 16 Dec 1893, R.
Schlechter 3980 (NH); Royal Natal National Park (-DB), 17 Feb 1984, O.M. Hilliard
and B.L. Burtt 17662 (NU); Tugela gorge (-DB), 6 Feb 1982, O.M. Hilliard and B.L.
Burtt 15454 (PRE); Tugela valley (-DD), 14 Feb 1926, A.J.W. Bayer 47 (PRE). 2829
(Harrismith): Ladysmith (-DB), 1Nov 1965, N.E. Shirley s.n. (NU). 2831 (Nkandla):
Nkandla, Ferncli Nature Reserve (-CA), 14 Feb 1994, H. Kennedy 533 (NU); Uni-
versity of KwaZulu-Natal (-DD), 27May 2003, S.J. Siebert 2333 (NH). 2929 (Under-
berg): Loteni Nature Reserve (-AD), 13 Dec 1978, M.L. Jacobsz 3937 (PRE); Mpendh-
le District (-BC), 24 Dec 1978, O.M. Hilliard and B.L. Burtt 11804 (NU, PRE); Polela
District (-CB), 6 Apr 1974, M.A. Rennie 545 (NU); Waterfall (-DD), Mar 2002, T. E d -
wards 2731 (NU). 2930 (Pietermaritzburg): Victoria (-AA), 12 Apr 1955, S.M. Johnson
1152 (NH); Tweedie (-AC), 31Dec 1927, Forbes 293 (NH); 28 Feb 1982, K.L. Immel-
man 260 (PRE); Loskop District (-AD), 11 Feb 1946, B.L. Howlett 80 (NH); Pieter-
maritzburg (-CB), 29 Feb 1976, M. Grice s.n. (NU); 14 Oct 1987, Renecken 6 (GRA);
Botanic Garden Estate (-CB), 8Dec 1991, D.G. Stielau 135 (NH, PRE); Cottingham
(-CC), 23 Mar 1969, R.G. Strey 8420 (NU); Durban Road (-CD), 19 Nov 1950, J.G.
Lawn 1820 (NH); Umgeni Water Board, (-DA), 20 Oct 1984, J. Manning 536 (NU);
Inanda (-DB), Dec, J. Medley-Wood 370 (NH); 25 Nov 1983, H.H. Hilger 35 (PRE);
Pinetown District (-DD), 28 Nov 1913, J.M. Wood 12373 (NU). 3029 (Kokstad):
Harding, Victoria East (-CA), 30 Apr 1955, G.L. Lewis 4914 (PRE); Harding, Alfred
District (-DB), 2 Mar 1983, O.M. Hilliard and B.L. Burtt 16755 (NU). 3030 (Port
Taxonomic revision of the genus Cynoglossum in southern Africa 35
Shepstone): Dumisa (-AD), 9Dec1992, A.M. Ngwenya 1071 (NH). W C:
3318 (Cape Town): Zondagsfontein (-DC), Dec-Mar 1930–1, J. ode A2838 (NH);
18 Oct 1943, B.S. Fischer 498 (NU). E C: 3128 (Mjika): Mahlahlane Forest
(-BC), 6 Mar 1985, A. Hutchings 1590 (GRA). 3129 (Port St. John’s): Kloof above Port
St. Johns (-DA), 31 Jan 1936, M.C. Gillett 1257 (PRE). 3226 (Fort Beaufort): Ama-
tole Mountain (-DB), 1May 1986, P.B. Phillipson 1492 (PRE); Fort Beaufort (-DD),
21 Mar 1977, G.E. Gibbs Russell 3735 (GRA). 3326 (Grahamstown): Grahamstown
Nature Reserve (-BC), Feb 1917, J.C. Jane 17131 (PRE); 1 Apr 1952, A.R.H. Martin
9466 (GRA); Featherstone Kloof (-BC), 18 Jul 2001, C.J. Kayombo and A.A. Merti
3719 (GRA). 3227 (Stutterheim): Stutterheim District (-CB), 28 Jan 1979, O.M. Hill-
iard and B.L. Burtt 12427 (NU); Kababu Hills (-CA), 5 Feb 1936, M.C. Gillett 1325
(PRE); East London (-DB), 11 Dec 2001, J.J. Meyer 4067 (PRE).
eSwatini. 2631 (Hhoho): Hhoho District, Masilela area on Maphalaleni road
along Mucucene Hills (-AB), 27 Jan 1994, G. Germishuizen 7173, 7174 (PRE); 24Jan
1994, S.R. Hobson 2048 (PRE); Hlambanyathi valley (-AC), 27 Nov 1954, R.J. Comp-
ton 24864 (NBG); Mbabane (-AC), 14 Jan 1955, R.J. Compton 24834 (PRE).
Lesotho. 2927 (Maseru): Monethi’s, Berea (-BB), 1 Jan 1957, A. Jacot Guillarmod
1910 (PRE); Mountain Road (-BD), Mar 1977, M. Schmitz 7317 (PRE). 2928 (Maraka-
bei): Loskop (-AB), 11 Apr 1980, B.N. Ubbink 974 (NH); Molika-lika (-AC), 7 Jan 1954,
A. Jacot Guillarmod 1674 (PRE). 2929 (Underberg): aba Ntšo, Sehlabathebe National
Park (-CC), 4–14 Jan 1973, Jacot Guillarmod, Getlie, and Mzamane 142 (PRE).
Unknown localities: No locality details, 30 Jan 1948, B.S. Fischer 1426 (NU).
Taxonomic notes. e specimen in the National Herbarium Pretoria (PRE) col-
lected in the Doornpoort area in Gauteng Province (Voucher number: S.P. Bester 9734
(PRE)) belongs to C. lanceolatum and not to C. amabile since it has characters which
are typical of this species, i.e., white corolla with a blue throat instead of bluish-purple
corolla and divaricately branched instead of clustered at the apex.
7. Cynoglossum obtusicalyx Retief & A.E. van Wyk in S. Afr. J. Bot. 62(3): 169–
172 (1996).
Type. South Africa ♀♂, Western Cape, Worcester (3319): 20 km E. of Ceres (-AD),
21 Oct 1958, Acocks 19893 (PRE-image! holotype; NBG, isotype).
Perennial or biennial herbs, ca. 0.45 m in height. Basal leaves 175–250×5–18 mm,
obtuse, densely pubescent, persistent, margins entire. Stem leaves 50–65×5–10 mm,
obtuse-lanceolate, apex acute, base cuneate, soft hairs. Trichomes woolly, soft, non-
bulbous based, unicellular, long, thin. Inorescence dichotomously branched terminal
cymes, pedicel 5–10 mm long, lengthens considerably in fruit. Calyx ca.1.0–1.6 mm
long, lobe oblong, densely pubescent, apices broadly obtuse-truncate shaped. Corolla
pale blue; lobes 5×7 mm diameter, ovate. Nutlets ovoid-convex, 2–4×3–4 mm, densely
echinulate with glochidia (Figure 15).
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
36
Diagnostic characters. e stems and leaves of Cynoglossum obtusicalyx are dense-
ly covered with soft, woolly trichomes which makes it similar to Cynoglossum alticola.
However, the two species dier in ower colour and size (deep blue corolla, 4×3 mm
vs. pale blue corolla, 5×7 mm in C. obtusicalyx). Cynoglossum obtusicalyx has pale blue
owers with corolla 5–7 mm long whereas C. austroafricanum has bright blue owers
with corolla 2.75–4.25 mm long. e two species also show variation in trichome tex-
ture and density, i.e., sparsely hairy with short, sti hairs on both the stem and leaves in
C. austroafricanum vs. densely arranged long, woolly hair on both the stem and leaves
in C. obtusicalyx.
Phenology. October to February.
Conservation status. Least Concern (Raimondo et al. 2009).
Distribution and habitat. Cynoglossum obtusicalyx is endemic to South Africa
where it has been recorded from Calvinia, Worcester, and Beaufort West in the North-
ern and Western Cape Provinces (Figure 16), it occurs in mountainous areas, often
growing on screes below clis.
Additional specimens examined. South Africa. N C: 3119 (Calvin-
ia): Calvinia District (-BD), 22 Sep 1955, J.P.H. Acocks 8509 (PRE); 22 Sep 1955, D.A
Leistner 394 (PRE; NBG). W C: 3222 (Beaufort West): Mountain View
farm (-BD), 17 Apr 1978, G. Gibbs Russell and R. Hermann 4272 (PRE).
Figure 15. Vegetative and reproductive morphological features of C. obtusicalyx A 1. base leaves 2. stem
with alternate stem leaves and terminal owers B SEM micrograph of a nutlet C SEM micrograph of
glochidia around the nutlet. Voucher: J.P.H. Acocks 8509 (PRE). Drawing scale bar: 7.0 mm. SEM images
scale bar: 1 mm (B); 200 µm (C).
Taxonomic revision of the genus Cynoglossum in southern Africa 37
8. Cynoglossum spelaeum Hilliard & B.L.Burtt in Notes Roy. Bot. Gard., Edin-
burgh 37(2): 287 (1979).
Cynoglossum basuticum Weim.ex Guillarmod, Fl. Lesotho 233 (1973), nom. nud.
Type. South Africa ♀♂, KwaZulu-Natal, Underberg (2929): Underberg District,
Cobham Forest Station, Polela valley (-CD), 20 Mar 1977, O.M. Hilliard and B.L.
Burtt 9728 (E, holotype image! K-image!, NU-image! PRE-image! [2 sheets] isotype).
Perennial herbs, ca. 0.4–0.5 m in height. Basal leaves 130–235×25–35 mm,
spathulate-obtuse, soft hairs, deciduous, margins entire. Stem leaves 40–80×10–25
mm, obtuse shaped, dark green adaxial surface, grey green abaxial surface, smooth
margins, soft hairs, sparsely packed. Trichomes unicellular, simple on both leaf sur-
faces, soft. Inorescence corymbose panicle; threadlike pedicel 5 mm long. Calyx ca.
1.5–2.0 mm long, lobes lanceolate-oblong, softly hairy on the outer surface, inner
surface smooth, apex acute. Corolla white; lobes ca. 3×3 mm diameter, obtuse. Nut-
lets ovoid, ca. 4×5mm; glochidia more marginal and acentric, marginal glochidia are
longer compared to the acentric glochidia (Figure 17).
Phenology. December to March.
Figure 16. Known distribution map of C. obtusicalyx in southern Africa based on the specimens examined.
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
38
Conservation status. Least Concern (Raimondo et al. 2009).
Diagnostic characters. is species is characterised by leaves that are sparsely
covered with hairs that have a softer feel, which distinguishes it sharply from all
the other southern African species which have either brittle or woolly hairs. e
abaxial leaf surface has a grey-greenish appearance which is also a unique character
of this species.
Distribution and habitat. e species is distributed in South Africa (Eastern
Cape, Free-State, and KwaZulu-Natal Provinces) and Lesotho (Figure 18). It grows in
loose sandy soil at the edge of an overhang.
Additional specimens examined. South Africa. K-N: 2729 (Utre-
cht): Ncandu State Forest (-DC), 8 May 1984, A. Nicholas and B. Isaacs 1965 (PRE).
2929 (Underberg): Allandale, Lion’s River District (-BC), 24 Jan 1978, O.M. Hilliard
and B.L. Burtt 11258 (NU); Giant’s Castle Nature Reserve (-BC), 07 Jan 1988, A. Ab-
bott 4064 (NH); 3 Feb 1976, O.M. Hilliard and B.L. Burtt 8904 (NU); 2 Jan 1978,
O.M. Hilliard and B.L. Burtt 11172 (NU); 20 Nov 1985, O.M. Hilliard and B.L.
Figure 17. Morphological features of C. spelaeum A line drawing of a fruiting branch showing stem with al-
ternate, sessile, spathulate leaves B SEM micrograph of nutlet showing more marginal and acentric glochidia
C close-up SEM micrograph showing glochidia with multiangular hooks. Voucher specimen: A.Nicholas
and B. Isaacs 1965 (PRE). Drawing scale 8.0 mm. SEM images scale bars: 1 mm (B); 200 µm (C).
Taxonomic revision of the genus Cynoglossum in southern Africa 39
Burtt 18242 (NU, PRE); Bulwer District (-DD), 8 Jan 1974, M.A. Rennie 507 (NU);
8 Mar 2019, A.N. Moteetee and L.K. Madika AL012 (JRAU). E C: 3127
(Lady Frere): Elliot District (-BB), 22 Jan 1979, O.M. Hilliard and B.L. Burtt 123427
(NU). 3226 (Fort Beaufort): Amatole Mountain (-DB), 16 Feb 1986, P.B. Phillipson
1294 (PRE).
Lesotho. 2927 (Maseru): Laikopile Mountain (-CD), Jan 1918, A. Dieterlen 40799
(PRE).
Taxonomic notes. Cynoglossum spelaeum is quite distinctive among the southern
African species having uniquely shaped (i.e., spatulate to obtuse) and coloured (i.e.,
deep green adaxial surface, grey green abaxial surface) leaves, that almost seem leathery
but contain few soft trichomes. It is also the only species among the southern African
species that has completely white owers. Although Hilger et al. (2015) indicated that
the species does not belong to genus Cynoglossum, they did not elaborate the reasons
for the exclusion. Nonetheless, the species ts the generic description of four glo-
chidiate nutlets that are adapted for zoochory. In addition, preliminary molecular data
(Madika 2020), showed a close relationship between C. coeruleum subsp. johnstonii var.
mannii, C. lanceolatum, and C. spelaeum.
Figure 18. Known distribution of Cynoglossum spelaeum in southern Africa based on the specimens
examined.
Lydia K. Madika & Annah Ntsamaeeng Moteetee / PhytoKeys 193: 9–42 (2022)
40
Acknowledgements
We thank the curators of the following herbaria for the loans of material and provision
of microsche images: PRE, GRA, NH, NU, and NBG including SAM. We thank
Ezemvelo KwaZulu-Natal Wildlife for the plant collecting permit. We thank the Uni-
versity of Johannesburg for logistical and nancial support. e rst author is indebted
to the National Research Foundation for nancial support in the form of a bursary.
e Central Analytical Facility of the Faculty of Science, University of Johannesburg
(SPECTRUM) is acknowledged for assistance with scanning electron microscopy. We
thank the three anonymous reviewers for their valuable input.
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