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Research Article
Taxonomic revision of the southern African species of the genus Cynoglossum L. (Boraginaceae)
expand article infoLydia K. Madika‡, Annah Ntsamaeeng Moteetee‡
‡ University of Johannesburg, Johannesburg, South Africa

Corresponding author: Annah Ntsamaeeng Moteetee ( amoteetee@uj.ac.za )

Academic editor: James Cohen
© 2022 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.
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
Open Access

Abstract

The aim of the study is to provide a revision of the genus Cynoglossum in southern Africa. The 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. The 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 misidentified. Diagnostic characters are described, correct nomenclature, synonyms, typification, distribution maps, as well as the key for identifying the studied species, are provided.

Keywords

Boraginaceae, Cynoglossum, Southern Africa, taxonomic revision

Introduction

The 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 the tropical regions of the Old and New Worlds. This genus comprises about 80–90 (Johnston 1924), to ca. 100 (Weigend et al. 2013), or as many as 200 species (Chacón et al. 2016). The 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). The 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’. This genus was established by Linnaeus (1753) to accommodate six species that he described at the time, namely, Cynoglossum officinale L., C. virginianum L., C. cheirifolium L., C. apenninum L., C. linifolium L., and C. omphaloides L. The 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).

The 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 identification difficult. Another unresolved problem concerns the species relationships in Cynoglossum. Morphological characters that were traditionally used to delineate genera proved to be insufficient, especially in defining 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, Lindelofia 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). This 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 misidentification. The 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 flat gynobases, while in Cynoglossum the nutlets are 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 differentiated 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. The fruits in members of this genus and the floral morphological characters, are normally considered to be of diagnostic importance. The 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. enerve Turcz. (now C. hispidum Thunb.) and C. micranthum Desf. He cited two species, namely, C. leptostachyum DC. and C. hispidum Thunb. 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 Africa as C. geometricum Baker & C.H. Wright] for the first 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 Thunb., C. lanceolatum Forssk., C. obtusicalyx Retief & A.E. van Wyk (endemic to South Africa), and C. spelaeum Hilliard & B.L. Burtt. This paper aims to provide a taxonomic revision of the southern African species, to provide a diagnostic 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 Thiers 2019), were examined for distribution and morphological data. The type specimens of relevant species were studied online from the JSTOR website (https://plants.jstor.org/). Voucher specimens and plant material were collected during several field trips conducted throughout South Africa and Lesotho between October 2018 and December 2019. Specimens collected were deposited in JRAU. The 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. Inflorescence 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 inflorescence to study). Hand drawings representing both the vegetative and reproductive characters were made for all the species (except for C. amabile). The 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. The 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. Then the dried samples were directly mounted on aluminium stubs and sputter-coated with a thin layer of gold before viewing under microscope. This was done to prevent charging of specimens due to accumulation of static electric field, 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. The stems are erect, hollow, simple at the base, and usually branched above. The 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. The 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 differ in shape (cylindrical with a pointed tip in C. alticola, vs. flat 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.

Figure 1. 

SEM micrographs of the adaxial leaf surfaces and the midrib of (A, B) C. alticola (C, D) C. obtusicalyx B close-up of C. alticola displaying cylindrical trichomes with pointed end D close-up of C. obtusicalyx displaying flat 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

The inflorescence is a cyme which is often dichotomously branched with spreading panicles. Flowers are either pedicelled or subsessile, with five parted corollas; corolla white with a blue throat, blue, violet, or magenta (C. hispidum), or rarely white (C. spelaeum). The stamens are included and arise from the base of the tube, they have short filaments and elliptic to oblong shaped anthers. The style is short and relatively thick, with a capitate stigma. The fruit is a schizocarp of four nutlets attached apically to a narrowly conical gynobase. The 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. The 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. The structure and shape of glochidia display an important distinguishing character amongst the southern African species (Table 1), with each species portraying a unique character as can be seen in Figure 2.

Table 1.
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Distinguishing characters between the southern African Cynoglossum species.

Characters C. alticola C. amabile C. austroafricanum C. coeruleum 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 ornamentation Glochidia densely packed on nutlet Marginal glochidia are more distinct than the central glochidia Glochidia more spread towards the margins Glochidia more marginal and on the median line Glochidia covering the whole nutlet Glochidia sparsely arranged on nutlet densely echinulate with glochidia glochidia more marginal and acentric, marginal glochidia are longer compared to the acentric glochidia
Figure 2. 

SEM micrographs of the adaxial surface of the nutlets of eight listed species of Cynoglossum A C. 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).

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., Pflanzenr. 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. officinale 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; first year basal leaves form a rosette, lanceolate or obtuse, often long petiolate. Inflorescences usually elongate, rarely bracteate, sparingly branched or loosely paniculate. Flowers pedicelled or subsessile; blue or violet with distinct veins, rarely white. Calyx five-partite, scarcely enlarged in fruit, patent or reflexed. Corolla tube short, throat closed with obtuse or arched scales; five-lobed, imbricate, obtuse, patent. Stamens five, included in the corolla tube, included, with short filaments, anthers ovoid or shortly oblong, obtuse. Ovary with four distinct lobes from an almost flat receptacle; style short or rather long; stigma small, flat or sub capitate; ovules horizontal, fixed to the central angle of the cell. Nutlets four, depressed, scarcely produced at the apex, convex or flat on the dorsal side or surrounded by an elevated margin, glochidiate (hair-like spines or short prickles). Seeds straight or slightly curve

Diagnostic key to the species

1 Soft woolly hairs covering the entire plant; nutlets thickened; glochidia densely arranged on the nutlet 1. C. alticola
– Stiff 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 1 cm long 3
3 Inflorescences clustered at the apex; corolla bluish purple; glochidia dense at the margins and centre of the nutlet 2. C. amabile
– Inflorescences 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 4 mm 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

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, lanceolate, 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. Inflorescence 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 diameter, 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).

Figure 3. 

Vegetative and reproductive morphology of Cynoglossum alticola A line drawings of: 1. a branch 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).

Phenology

February to May.

Conservation status

Least Concern (Raimondo et al. 2009).

Diagnostic characters

Cynoglossum alticola can be distinguished by its thick, convex nutlets. Among the southern African species, it has a unique appearance due to 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. The difference 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

The 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.

Figure 4. 

Distribution of Cynoglossum alticola based on the specimens examined.

Additional specimens examined

South Africa. Eastern Cape: 3028 (Matatiele): 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).

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. Inflorescence 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).

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).

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 flowers. However, the two species are easily distinguished by their flower colour (C. lanceolatum has white corolla with blue throat, whereas C. amabile has bluish purple corolla). This 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, Thailand, Philippines, and India), based on flower and fruit size. The difference can be observed in the inflorescences, whereby C. furcatum is a much larger plant with inflorescences 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.

Figure 6. 

Recorded distribution of Cynoglossum amabile in southern Africa based on the specimens examined.

Additional specimens examined

South Africa. KwaZulu-Natal: 2930 (Pietermaritzburg): 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 (Kӧnig et al. 2015). This 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.

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 45–100×10–21 mm, narrowly lanceolate to linear lanceolate shaped, acute apex, cuneate base, margins undulate, covered with stiff hairs. Trichomes unicellular, with thick round base on the adaxial surface, simple on the abaxial surface. Inflorescences 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).

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 micrograph 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).

Phenology

December to April.

Conservation status

Least Concern (Raimondo et al. 2009).

Diagnostic characters

Amongst the southern African species, Cynoglossum austroafricanum can be confused with either C. lanceolatum or C. coeruleum var. mannii. This species differs from the two by the colour of the corolla (white corolla with pale blue throat vs pale blue corolla throughout in C. austroafricanum). This latter observation was also noted by Hilliard and Burtt (1986).

Distribution and habitat

The species is distributed in South Africa (North-West, Gauteng, Mpumalanga, Free-State, KwaZulu-Natal, and Eastern Cape Provinces), eSwatini and Lesotho (Figure 8), where it occurs in shady, disturbed areas, and sandy, dry riverbanks.

Figure 8. 

Known distribution of Cynoglossum austroafricanum in southern Africa based on the specimens examined.

Additional specimens examined

South Africa. Limpopo: 2430 (Tzaneen): Lekgalameetse Nature Reserve (-AA), 7 Oct 1986, N. Stalmans 1404 (PRE). North-West: 2626 (Lichtenburg): Lichtenburg (-AA), Feb 1918, L. Kretzshmar 17065 (PRE). Gauteng: 2528 (Pretoria): Brooklyn (-CA), 1 Feb 1931, A.O.D. Mogg 16451 (PRE). Mpumalanga: 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), 8 Feb 1904, J. Burtt-Davy 1472 (PRE). 2630 (Carolina): Between Oshoek border post and Carolina (-BA), Jan 1906, H. Bolus 12161 (PRE). Free State: 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). Kwazulu-Natal: 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 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. Thompson 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). Eastern Cape: 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), 14 Feb 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): Thaba Bosiu (-BC), 1 Mar 1978, M. Schmitz 8205 (PRE); Mahlatsa (-BB), 18 Jan 1941, A. JacotGuillarmod 51 (GRA); Sefikeng 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 Guillarmod 1072 (PRE), 25 Feb 1949, W.J. Barker 21515 (NBG).

No locality details: 27 Nov 1888, H. Medley 4576 (NH); Dec 1946, E. Meston 50 (NU).

Taxonomic notes

According to Hilliard and Burtt (1986), this species was described by Dr. H. Weimarck who did not select a type specimen; therefore, in typifying it they retained the specific epithet.

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 s.n. (K-image! lectotype, here designated). Notes: The 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 90–190×28–56 mm, lanceolate-obtuse, softly hairy, deciduous, margins entire. Stem leaves 35–90×7–25 mm, lanceolate, apex acute, base acute to obtuse, covered with moderately stiff hairs margins entire. Trichomes bulbous based on the upper surface of the leaf, sometimes simple on the lower surface. Inflorescences terminal axillary cymes, few branches spreading dichotomously; 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, 3–4×2.5–3.5 mm; glochidia more marginal and on the median line (Figure 9).

Figure 9. 

Vegetative and reproductive morphological features of Cynoglossum coeruleum var. mannii A line 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).

Phenology

December to April.

Conservation status

Least Concern (Raimondo et al. 2009).

Diagnostic characters

This variety can be easily confused with C. lanceolatum due to the dichotomous branching of the inflorescence but can be distinguished from it by the distribution and density of the glochidia in the nutlets. The 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

This 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.

Figure 10. 

Known distribution of Cynoglossum coeruleum var. mannii in southern Africa based on the specimens examined.

Additional specimens examined

South Africa. Kwazulu-Natal: 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). Eastern Cape: 3023 (Britstown): Kamberg (-CC), 21 Mar 1983, O.M. Hilliard 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 first 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. The correct name is Cynoglossum coeruleum subsp. johnstonii var. mannii, as synonimized 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.science.kew.org/taxon/966730-1) and the World Flora Online (http://www.worldfloraonline.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) There 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 inflorescence character and the branching pattern.

Cynoglossum hispidum Thunb., Prodr. Plant. Cap. 1:34 (1794); Roemer 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], isolectotype). [Note: The HAL specimen is chosen as a lectotype because the specimen 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, Thunberg 168 sub THUNB-UPS 3996 (UPS, microfiche! 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. Inflorescence terminal axillary cyme, branches spreading 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 multiangular (Figure 11).

Figure 11. 

Vegetative and reproductive morphological features of Cynoglossum hispidum A line drawing of obtuse shaped, rosette base leaves, terminal branched fruits, terminal flowers 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).

Phenology

October to March.

Conservation status

Least Concern (Raimondo et al. 2009).

Diagnostic characters

The species can be confused with C. lanceolatum with which it shares a similar branching pattern of the inflorescences and upright brittle hairs covering the whole plant. However, the two differ 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

This 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.

Figure 12. 

Known distribution of Cynoglossum hispidum in southern Africa based on the specimens examined.

Additional specimens examined

South Africa. Limpopo: 2229 (Waterpoort): Evelyn valley (-BD), Feb 1944, RUC Biology Expedition 417 (GRA). 2430 (Tzaneen): Lekgalameetse Nature Reserve (-AA), 4 Dec 1985, M. Stalmans 790 (PRE); Sekhukhune District, Leolo Mountains (-CA), 14 Mar 2007, B. Sachse 471 (PRE). North-West: 2523 (Pomfret): Forbes (-DA), 29 Oct 1959, B. Dlamini s.n. (NH). 2627 (Potchefstroom): Goedgedacht (-AA), 1 May 1932, J.D. Sutton 676 (PRE); Losberg, Elandsfontein (-BC), 11 Dec 1934, J.J. Theron 725 (NH); Potchefstroom (-CA), 23 Nov 1946, W.J. Louw 1525 (PRE). Gauteng: 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). Mpumalanga: 2530 (Mashishing): Boschhoek (-AA), 16 Nov 1933, R.S.N. Young A375 (PRE); Verlorenkloof Reserve, Welgedacht (-AD), 28 Nov 2008, S.P. Bester 8679A (PRE); Machadodorp, Farm Grootvlei (-CB), 21 Oct 1988, P. Burgoyne 457 (PRE); Songimvelo Game Reserve (-DD), 10 Dec 1992, M. Jordaan 2490 (PRE). 2629 (Bethal): Bethal (-AD), 14 Dec 2010, R. Leendertz 9386 (PRE). 2630 (Carolina): Ermelo District, Spitskop (-CA), Dec 1915, R. Potts 5007 (PRE). Northern Cape: 2820 (Kakamas): Krantzkop (-DA), Nov 1911, J. Thode 4768 (NBG). Free State. 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). Kwazulu-Natal: 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 (-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 District, Elandslaagte (-CD), 23 Oct 1964, N.E. Shirley s.n. (NU). 2831 (Nkandla): Empangeni (-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 (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); Cliffdale 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. Getliffe 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), 3 Sep 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). Western Cape: 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). Eastern Cape: 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. Theron 1604 (PRE); Elliotdale District (-DC), 11 Jul 1966, J.L. Gordon-Gray 529 (NH). 3129 (Umtata): Baziya, Tembuland (-CC), no date, Baur 257 (K-image); Blesbok flats 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 off 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. Giffen 264 (PRE). 3227 (Stutterheim): Stutterheim (-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 Guillarmod 279 (GRA). 2929 (Underberg): Thaba Ntšo, Sehlabathebe National Park (-CC), 4–14 Jan 1973, Jacot Guillarmod, Getliffe and Mzamane 65 (PRE); 13 Feb 1976, A. Beverly 471 (PRE).

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. lanceolatum. 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).

Cynoglossum lanceolatum Forssk in Fl. Aegypt. Arab: 41 (1775).

Cynoglossum hirsutum Thunb., Prodr. Pl. Cap.: 34 (1794).

Type: South Africa ♀♂, Precise locality unknown: Roggerveld, C.P. Thunberg 168, sub THUNB-UPS 3995 (UPS-microfiche! 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, designated 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. Basal leaves 85–180×15–23 mm, lanceolate-obtuse, blade elliptic, softly hairy, deciduous. Stem leaves 40–65×8–18 mm, lanceolate, apex acute, base cuneate, covered with moderately stiff hairs. Trichomes brittle, simple on both leaf surfaces. Inflorescence 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).

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 multiangular tip. Voucher specimen: S.P. Bester 4653 (PRE). Drawing scale bar: 7.0 mm. SEM scale bar: 1 mm (B); 100 µm (C).

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 inflorescence, flower colour, and nutlet size. The 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.

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.

Figure 14. 

Known distribution of Cynoglossum lanceolatum in southern Africa based on the specimens examined.

Additional specimens examined

South Africa. Limpopo: 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 (-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 Municipality, Leolo Mountain, SE of Tama Kgoshi (-CA), 14 Mar 2007, B. Sachse 466 (PRE). North-West: 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). Gauteng: 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 Dec 1971, G.J. Bredenkamp 594 (PRE). Mpumalanga: 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): Loskop Dam (-AD), 12 Jan 1967, G.K. Theron 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. Theron 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). Free State: 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). Kwazulu-Natal: 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), 1 Nov 1965, N.E. Shirley s.n. (NU). 2831 (Nkandla): Nkandla, Ferncliff Nature Reserve (-CA), 14 Feb 1994, H. Kennedy 533 (NU); University of KwaZulu-Natal (-DD), 27 May 2003, S.J. Siebert 2333 (NH). 2929 (Underberg): Loteni Nature Reserve (-AD), 13 Dec 1978, M.L. Jacobsz 3937 (PRE); Mpendhle 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. Edwards 2731 (NU). 2930 (Pietermaritzburg): Victoria (-AA), 12 Apr 1955, S.M. Johnson 1152 (NH); Tweedie (-AC), 31 Dec 1927, Forbes 293 (NH); 28 Feb 1982, K.L. Immelman 260 (PRE); Loskop District (-AD), 11 Feb 1946, B.L. Howlett 80 (NH); Pietermaritzburg (-CB), 29 Feb 1976, M. Grice s.n. (NU); 14 Oct 1987, Renecken 6 (GRA); Botanic Garden Estate (-CB), 8 Dec 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 Shepstone): Dumisa (-AD), 9 Dec 1992, A.M. Ngwenya 1071 (NH). Western Cape: 3318 (Cape Town): Zondagsfontein (-DC), Dec-Mar 1930–1, J. Thode A2838 (NH); 18 Oct 1943, B.S. Fischer 498 (NU). Eastern Cape: 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): Amatole Mountain (-DB), 1 May 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. Hilliard 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); 24 Jan 1994, S.R. Hobson 2048 (PRE); Hlambanyathi valley (-AC), 27 Nov 1954, R.J. Compton 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 (Marakabei): Loskop (-AB), 11 Apr 1980, B.N. Ubbink 974 (NH); Molika-lika (-AC), 7 Jan 1954, A. Jacot Guillarmod 1674 (PRE). 2929 (Underberg): Thaba Ntšo, Sehlabathebe National Park (-CC), 4–14 Jan 1973, Jacot Guillarmod, Getliffe, and Mzamane 142 (PRE).

Unknown localities: No locality details, 30 Jan 1948, B.S. Fischer 1426 (NU).

Taxonomic notes

The specimen in the National Herbarium Pretoria (PRE) collected 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.

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. Inflorescence 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).

Figure 15. 

Vegetative and reproductive morphological features of C. obtusicalyx A 1. base leaves 2. stem with alternate stem leaves and terminal flowers 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).

Diagnostic characters

The stems and leaves of Cynoglossum obtusicalyx are densely covered with soft, woolly trichomes which makes it similar to Cynoglossum alticola. However, the two species differ in flower colour and size (deep blue corolla, 4×3 mm vs. pale blue corolla, 5×7 mm in C. obtusicalyx). Cynoglossum obtusicalyx has pale blue flowers with corolla 5–7 mm long whereas C. austroafricanum has bright blue flowers with corolla 2.75–4.25 mm long. The two species also show variation in trichome texture and density, i.e., sparsely hairy with short, stiff 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 Northern and Western Cape Provinces (Figure 16), it occurs in mountainous areas, often growing on screes below cliffs.

Figure 16. 

Known distribution map of C. obtusicalyx in southern Africa based on the specimens examined.

Additional specimens examined

South Africa. Northern Cape: 3119 (Calvinia): Calvinia District (-BD), 22 Sep 1955, J.P.H. Acocks 8509 (PRE); 22 Sep 1955, D.A Leistner 394 (PRE; NBG). Western Cape: 3222 (Beaufort West): Mountain View farm (-BD), 17 Apr 1978, G. Gibbs Russell and R. Hermann 4272 (PRE).

Cynoglossum spelaeum Hilliard & B.L.Burtt in Notes Roy. Bot. Gard., Edinburgh 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 surfaces, soft. Inflorescence 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. Nutlets ovoid, ca. 4×5 mm; glochidia more marginal and acentric, marginal glochidia are longer compared to the acentric glochidia (Figure 17).

Figure 17. 

Morphological features of C. spelaeum A line drawing of a fruiting branch showing stem with alternate, 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).

Figure 18. 

Known distribution of Cynoglossum spelaeum in southern Africa based on the specimens examined.

Phenology

December to March.

Conservation status

Least Concern (Raimondo et al. 2009).

Diagnostic characters

This 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. The abaxial leaf surface has a grey-greenish appearance which is also a unique character of this species.

Distribution and habitat

The 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. Kwazulu-Natal: 2729 (Utrecht): 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. Abbott 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. 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). Eastern Cape: 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 flowers. 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 fits the generic description of four glochidiate 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.

Acknowledgements

We thank the curators of the following herbaria for the loans of material and provision of microfische images: PRE, GRA, NH, NU, and NBG including SAM. We thank Ezemvelo KwaZulu-Natal Wildlife for the plant collecting permit. We thank the University of Johannesburg for logistical and financial support. The first author is indebted to the National Research Foundation for financial support in the form of a bursary. The 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.

References

  • Al-Shehbaz IA (1991) The genera of Boraginaceae in the South-eastern United States. Journal of the Arnold Arboretum 1: 1–169. https://doi.org/10.5962/p.315943
  • Brand A (1921) Borraginaceae-Borraginoideae, Cynoglosseae. In: Engler A (Ed.) Das Pflanzenreich: regni vegetablilis conspectus. W. Engelmann, Leipzig, 183 pp.
  • Burrows JE, Willis CK (2005) Plants of the Nyika Plateau. Southern African Botanical Diversity Network Report No. 31 SABONET, Pretoria, 802 pp.
  • Chacón J, Luebert F, Hilger HH, Ovchinnikova S, Selvi F, Cecchi L, Matt-Guilliams C, Hassenstab-Lehman K, Sutorý K, Simpson MG, Weigend M (2016) The borage family (Boraginaceae s.str.): A revised infrafamilial classification based on new phylogenetic evidence, with emphasis on the placement of some enigmatic genera. Taxon 65(3): 523–546. https://doi.org/10.12705/653.6
  • Cohen JI (2011) A phylogenetic analysis of morphological and molecular characters of Lithospermum L. (Boraginaceae) and related taxa: Evolutionary relationships and character evolution. Cladistics 27(6): 559–580. https://doi.org/10.1111/j.1096-0031.2011.00352.x
  • Cohen JI (2013) A phylogenetic analysis of morphological and molecular characters of Boraginaceae: Evolutionary relationships, taxonomy, and patterns of character evolution. Cladistics 30(2): 139–169. https://doi.org/10.1111/cla.12036
  • Edwards D, Leistner OA (1971) A degree reference system for citing biological records in southern Africa. Mitteilungen des Botanische Staatssammlung Munchen 10: 501–509.
  • Ge-Ling Z, Riedl H, Kamelin R (1995) Boraginaceae. In: Wu ZY, Raven PH (Eds) Science Press, Bejing. Flora of China 16: 329–431.
  • Germishuizen G, Meyer NL (2003) Plants of southern Africa: an annotated checklist. National Botanical Institute: Pretoria. Strelitzia 14: 133–136.
  • Hasenstab-Lehman KE, Simpson MG (2012) Cat’s eyes and popcorn flowers: Phylogenetic systematics of the genus Cryptantha s.l. (Boraginaceae). Systematic Botany 37(3): 738–757. https://doi.org/10.1600/036364412X648706
  • Hilger HH, Greuter W, Stier V (2015) Taxa and names in Cynoglossum sensu lato (Boraginaceae, Cynoglosseae): An annotated, synonymic inventory, with links to the protologues and mention of original material. Biodiversity Data Journal 3: 1–23. https://doi.org/10.3897/BDJ.3.e4831
  • Kӧnig J, Siwakoti M, Hilger HH, Weigend M (2015) A revision of the genus Cynoglossum L. (Boraginaceae Juss.) in Nepal and notes on the widespread Asian species. Phytotaxa 224(1): 1–28. https://doi.org/10.11646/phytotaxa.224.1.1
  • Linnaeus C (1753) Species plantarum, 1: 134–135.
  • Madika LK (2020) The systematics and biological activity of southern African species of the genus Cynoglossum L. (Boraginaceae) (Master’s Dissertation). Johannesburg: University of Johannesburg, 34 pp. https://doi.org/10.3897/arphapreprints.e73746
  • Mill RR, Miller AG (1984) Studies in the flora of Arabia: IX. A synopsis of Paracynoglossum (Boraginaceae). Notes from the Royal Botanic Garden Edinburgh 41: e478.
  • Otero A, Jiménez-Mejías P, Valcárcel V, Vargas P (2014) Molecular phylogenetics and morphology support two new genera (Memorema and Nihon) of Boraginaceae s.s. Phytotaxa 173(4): 241–277. https://doi.org/10.11646/phytotaxa.173.4.1
  • Raimondo D, von Staden L, Foden W, Victor JE, Helme NA, Turner RC, Kamundi DA, Manyama PA (2009) Red List of South African Plants. Strelitzia 25. South African National Biodiversity Institute, Pretoria, 86: 23–35.
  • Retief E (2005) Cynoglossum hispidum. http://pza.sanbi.org/cynoglossum-hispidum [Accessed February 2018]
  • Selvi F, Coppi A, Cecchi L (2011) High epizoochorous specialisation and low DNA sequence divergence in Mediterranean Cynoglossum (Boraginaceae): Evidence from fruit traits and ITS region. Taxon 60(4): 969–985. https://doi.org/10.1002/tax.604003
  • Stapf O, Drummond JR (1906) Cynoglossum amabile, Stapf et Drummond. Bulletin of Miscellaneous Information, Royal Botanic Gardens, Kew. 1906: e202. https://doi.org/10.2307/4120226
  • Sutorý K (2010) Oncaglossum, a new genus of the Boraginaceae, tribe Cynoglosseae, from Mexico. A Journal for Botanical Nomenclature 20(4): 463–469. https://doi.org/10.3417/2008059
  • Thiers B (2019) Index Herbariorum, a Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/ih/
  • Verdcourt B (1991) Boraginaceae. In: Polhill RM (Ed.) Flora of Tropical East Africa. Royal Botanic Gardens, Kew, 124 pp.
  • Von Staden L, Raimondo D, Dayaram A (2013) Taxonomic research priorities for the conservation of the South African flora. South African Journal of Science 109(3/4): 1–10. https://doi.org/10.1590/sajs.2013/1182
  • Weigend M, Leubert F, Selvi F, Brokamp G, Hilger HH (2013) Multiple origins for Hound’s tongues (Cynoglossum L.) and Navel seeds (Omphalodes Mills.) –The phylogeny of the Borage family (Boraginaceae s.str.). Molecular Phylogenetics and Evolution 68(3): 604–618. https://doi.org/10.1016/j.ympev.2013.04.009
  • Wright CH (1904) ‘Boraginaceae’. In: Harvey WH, Sonder OW, Thiselton-Dyer WT (v.4 sect.2). Flora capensis: being a systematic description of the plants of the Cape colony, Caffraria and Port Natal (and neighbouring territories). Lovell, Reeve and CO. , Limited, London, 14 pp. https://doi.org/10.5962/bhl.title.821
  • Xu Q-J, Zhang D-Z, Dang L-J, Li K-P (2009) Triterpene acids from Cynoglossum amabile. Chemical Research in Chinese Universities 25(3): 404–406.
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