IntroductionI became interested in the summer-rainfall Gerbera species while studying the endangered Gerbera aurantiaca (the Hilton Daisy). Species delineations in this genus in the eastern regions of South Africa are often confusing, as evidenced by the vast array of synonyms, with many specimens difficult to place. Possibly the most difficult and confusing species of the genus to place, due to their variability, are G. ambigua and G. viridifolia.

The genus Gerbera (Asteraceae–Mutisieae) consists of about 30 species found in the Sino-Himalayan region, tropical and southern Africa, with one controversial species (G. hintonii) from South America (Katinas 1998). The southern part of Africa appears to be the centre of diversity for Gerbera in Africa, and the distribution pattern of the genus supports the theory that it originated in southern Gondwanaland, with only three widespread species successfully migrating northwards (Hansen 1985). About 14 species of Gerbera are found in southern Africa, and eight in the eastern Cape, KwaZulu-Natal, Mpumalanga, Swaziland and Limpopo (Germishuizen et al. 2006).

The genus Gerbera in summer-rainfall South AfricaIsabel JohnsonSchool of Biological and Conservation Sciences, University of KwaZulu-Natal Pietermaritzburg,

The genus was named in 1737 by Jan Frederik Gronovius, a student of Carl von Linné (Linnaeus), somewhat puzzlingly after Traugott Gerber, a medical doctor born in Poland in 1710, who created the first botanic garden in Moscow. In spite of exhaustive investigations no obvious connection has been found between the taxon name and Gerber (Ambrosius 2003).

Gerbera is one of the most important South African genera commercially, being amongst the top ten international cut flowers and pot plants. The story of the commercial Gerbera hybrids goes back to the Cambridge Botanical Garden in 1886 when R. Irwin Lynch produced the first hybrids by crossing G. jamesonii, originating from seed from a plant sent to Kew by Robert Jameson via the Durban Botanical Society with G. viridifolia, from some seed sent to Kew by George Thorncroft. Lynch named the hybrid Gerbera × cantebrigiensis. By 1912 an impressive range of colour forms were exhibited at Chelsea. The majority of today’s commercially cultivated forms originate from the crossing of the progenies of these two species (Codd 1979). It is probable that other

A ‘feeding frenzy’ of monkey beetles on Gerbera aurantiaca.

Private Bag X01, Scottsville, 3209, South Africa. E-mail: johnsoni@ukzn.ac.za

Gerbera species have been used in breeding, but no information is available.

Typical Gerbera plants are perennial with spreading, thickened, fleshy roots and stemless rosettes of leathery and usually hairy leaves arising from a crown; daisy-like inflorescences borne singly on an inflorescence stalk, usually with conspicuous ray floret ligules (‘petals’) from white through yellow to deep red and purple. The outer ray florets are usually female with vestigial stamens (staminodes), while the central disc florets are hermaphrodite or functionally male. The florets are characteristically bilabiate (with the outer, laterally fused elongated petals having three small upper lobes or teeth, and the smaller, laterally fused inner petals with two upper lobes or teeth). The fruit or cypselae have an umbrella-like feathery pappus. The bowl shaped inflorescences are visited by a wide range of generalist insect pollinators. As with most summer-rainfall grassland species, gerberas become dormant in the dry winter months and are able to survive fires, periods of drought and very cold conditions due to their fleshy root system.

The only species of conservation concern is G. aurantiaca which is listed as “Endangered” due to habitat specificity, and the continuing threat of transformation and degradation of mistbelt grassland (Raimondo et al. 2009).

Gerbera species in eastern South Africa have been reviewed by Dümmer (1914), by Olive Hilliard in her comprehensive Compositae of Natal (1977) and most recently by Hans Hansen (1985).

Three sections of the genus are found in the Eastern

1. 2. 3.

1. Section Parva Gerbera parva N.E.Br. Common names: none known

This small, dainty Drakensberg endemic is typically found in colonies along streambeds under Leucosidea sericea thickets or on damp cliff faces. The specific epithet is derived from the Latin ‘small’. The species was first collected by Maurice Evans, co-author with Medley Wood of the first volume of Natal Plants, near Bushman’s River in 1894 (Evans 57, K) (Gunn & Codd 1981).

It is a perennial herb with flowering stems up to 200 mm tall, leaves 30 mm in diameter and inflorescences up to 25 mm in diameter. Distinguished by strap-shaped roots, shiny dark green rounded cordate leaves, bracteate scapes (inflorescence stalks bearing bracts), a single series of ray florets lacking staminodes, with ligules white above, white or pink to red below, white pappus and unbeaked cypsela, this species shows very little morphological variation. Flowering occurs from October to December.

A mass of Gerbera aurantiaca plants at Byrne.

Gerbera parva. Photo: Tony Abbott.

Cape, KwaZulu-Natal and Mpumalanga:Section Parva (1 species);Section Piloselloides (1 species);Section Lasiopus (6 species).

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Herbarium specimen of Gerbera parva.

2. Section PiloselloidesGerbera piloselloides (L.) Cass.Common names: Small Yellow Gerbera, Swarttee-bossie, indlebeyempithi, uhlango olimpofu, umoya-wezwe

The most widespread, but the least showy of all the Gerbera species, G. piloselloides occurs from sea level to about 3 500 m in grassland and open woodland throughout tropical and southern Africa, Madagascar and Asia. The specific epithet means ‘with shaggy hairs’.

The species is easily recognized by the hairy, widened peduncle apex below the inflorescence, the pink, white or yellow ray florets which do not, or only slightly exceed the involucral bracts, and the long-beaked (10 mm) fruit with a whitish pappus. The flowering stems are up to 450 mm tall, and the leaves are very variable and sometimes white-felted beneath.

This species is probably self-compatible, an unusual condition for the genus. In spite of the very nondescript inflorescences, fruit set is always impressive, and the conspicuous white seed heads can be seen dotted throughout grasslands in summer. Flowering occurs from July to February.

3. Section Lasiopus This section contains three very showy species, the Barberton Daisy (G. jamesonii), the Hilton Daisy (G. aurantiaca), and G. viridifolia

In total this section has six species (G. ambigua, G. aurantiaca, G. galpinii, G. jamesonii, G. natalensis, and G. viridifolia) (Germishuizen et al. 2006), with its distribution centred in KwaZulu-Natal and Mpumalanga, and is distinguished by ebracteate scapes, biseriate ray florets, beaked cypselae, and a whitish to purple slender pappus (Hansen 1985).

These six species are separated from one another by a few morphological differences and appear to be closely related. They also show a great deal of intraspecific morphological variation, especially G. ambigua and G. viridifolia. This is the group with the greatest taxonomic controversy.

Interestingly, three type specimens from this group were collected by Christian Ferdinand Friedrich Krauss (1812-1890) during his visit to the Pietermaritzburg area of Natal in 1839. These were G. ambigua [= G. kraussii ] “in summitate mont Tafelberg” (Krauss 402, ?P), G. aurantiaca “ad collium latera prope Pietermaritzburg” (Krauss 388, ?P), and G. natalensis “ in collibus prope villum Pietermaritzburg” (Krauss 452, TUB, K). He also collected G. piloselloides from the summit of Table Mountain in KwaZulu-Natal during this expedition, but the specimen (Krauss 49?) cannot be traced.

Gerbera piloselloides at Byrne.

Fruiting head of Gerbera piloselloides.

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The Gerbera kraussii controversyA brief summary of some of the changes to this taxon are necessary since many readers as well as the widely used A fieldguide to the wildflowers of KwaZulu-Natal and the eastern regions (Pooley 1998) use this name.

This species was first described by Schulz Bipontinus in 1844. Closely allied to G. ambigua, but differing in the rusty-white tomentum and distinctly raised nerves on the lower leaf surface, violet pappus colour, and white rays, G. kraussii was accepted as a valid species for the next century.

Many consolidations, separations and recombinations of four of the species in the section Lasiopus (G. ambigua, G. kraussii, G. natalensis and G. viridifolia) have taken place over the last 150 years. To summarise, the most important: Schulz Bipontinus (1844) described G. ambigua, G. kraussii, G. natalensis and G. viridifolia, while Hilliard (1977) sunk G. viridifolia into G. ambigua, but kept G. kraussii and G. natalensis as separate species. In 1985 Hansen sunk G. kraussii into G. ambigua, but kept G viridifolia subsp. viridifolia and G. viridifolia subsp. natalensis as separate taxa.

Hilliard (1994) does not agree with Hansen’s sinking of G. kraussii into G. ambigua. Retief (in Germishuisen et al. 2006) does not recognize G. kraussii and includes it in G. ambigua, but does give G. natalensis and G. viridifolia separate species status. In addition there have been numbers of other species separated and described within these taxa over the years, but this becomes too lengthy to describe here.

For the sake of consistency I have followed the taxonomic arrangement of Retief (in Germishuizen et al. 2006).

Gerbera ambigua (Cass.) Sch. Bip. Common names: Pink and White Gerbera, Botterblom, Griekwateebossie, ucabazane, uhlamvuhloshane, ulimi-lwenkomo

The specific epithet ambigua means ‘doubtful’ – a very apt name for this extremely variable species which occurs throughout the eastern regions of South Africa and tropical Africa to Ethiopia in grassland and open woodland. This species spreads by underground runners, often forming large conspicuous colonies in grassland, very noticeable from afar on a breezy day by the silvery white-felted undersurface of the leaf.

Large numbers of Gerbera ambigua plants in rocky grassland.

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The leaves are extremely variable in size, shape, margin ornamentation, indumentum (hair covering) on upper and lower surfaces and in petiole length. The most consistent character here is that the leaves are white-felted beneath, unlike the other members of the section Lasiopus. The lateral veins are usually prominent on the lower leaf surface. Inflorescences are 14 to 40 mm across, and the ray floret ligules may be white, pink, yellow, mauve or red, and the pappus white to deep purple.

Yellow-rayed forms occur mainly in Mpumalanga and northwards, and also along the KwaZulu-Natal coastal regions. These yellow forms usually have a light-coloured pappus, but specimens collected near Dullstroom had dark centres caused by purple pappus hairs.

White-flowered forms with and without dark centres are commonest in central and southern KwaZulu-Natal, and a purple form has been collected from Umfolozi.

G. ambigua flowers mainly from September to January, but can be found so in any month of the year.

This species grows easily in cultivation, and makes a good edging plant with interesting leaves and a long flowering season.

Natural hybridisation with G. aurantiaca sometimes occurs where the species are found together, producing pink flowered progeny with intermediate leaf characteristics.

Gerbera ambigua leaf showing white-felting and prominent side veins.

Gerbera ambigua inflorescence showing dark centre and white rays.

Gerbera aurantiaca Sch. Bip. Common name: Hilton Daisy

The specific epithet aurantiaca means ‘beautiful orange’, somewhat strange for a characteristically bright red-flowered species. However, on examining the description by Schultz Bipontinus (1844), it seems that the specimen that Krauss sent to him was in fact deep orange, a not unusual but not predominant colour form in most populations. The Swartkops population near Pietermaritzburg where Krauss most likely collected this plant has sadly almost disappeared under urban development and commercial plantations, and less than a dozen plants survive in a tiny fragment of grassland.

G. aurantiaca is a mistbelt grassland endemic found in widely scattered localities from Richmond in the south to Babanango, with a population of yellow-flowered plants in the Ngome area, and another four populations of red and yellow forms in Mpumalanga.

The leathery leaves are approximately 250 × 60 mm in size, elliptical, narrowing to a winged petiole-like base, with the margins entire or obscurely denticulate, lateral veins not prominent, cobwebby underneath when young, but sparsely hairy when mature. The inflorescence may be up to 90 mm in diameter, with ray floret ligules yellow, orange or red, and the purple-violet pappus giving a vivid black centre to young flowers. Flowering takes place mainly from September to December.

This beautiful species is very difficult to cultivate and seldom survives in a garden situation.

Gerbera galpinii Klatt Common name: none known

This species is named for Ernest Edward Galpin (1858–1941), a banker by trade with no formal botanical training who became fascinated with the flora around Barberton in 1889 and kept carefully annotated specimens with many duplicates which were sent to Kew, Bolus and Zurich. G. galpinii was collected during his time as a

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Gerbera aurantiaca from Birnam Wood Hill.

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Magnified photograph of a Gerbera aurantiaca inflorescence showing the bilabiate disc florets.

Gerbera aurantiaca cypsela and pappus.

bank manager in Barberton. Galpin went on to collect over 11 000 specimens (Gunn & Codd 1981).

Unlike other Gerbera species, G. galpinii is confined to moist or swampy ground along the Mpumalanga Drakensberg and south to Wakkerstroom and Utrecht. This species is not well known. Leaves are few, up to 250 × 15 mm, with two-thirds of this petiole-like, narrowly elliptical or lanceolate in shape, glabrous (not hairy), margins entire or denticulate with hardly visible nerves and a long, winged petiole. Inflorescences measure about 18 mm across, and the rays are yellow or white above, red below and the pappus pinkish brown. Flowering is mostly from September to November.

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A yellow form of Gerbera aurantiaca from northern KwaZulu-Natal.

A natural hybrid between Gerbera aurantiaca and G. ambigua.

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Herbarium specimen of Gerbera galpinii.

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Gerbera jamesonii Bolus ex Adlam Common names: Barberton Daisy, Transvaal Daisy, Rooigousblom

The Barberton Daisy was first discovered by A. Rehman between 1875 and 1880, but was named after Robert Jameson (1832–1919) a businessman and politician, who collected seeds of this species near Barberton in about 1885. These were grown by John Medley Wood in the Durban Botanic Gardens and later sent to Kew to become one of the parents of Gerbera hybrids (Codd 1979).

This species is endemic to Mpumalanga, Limpopo and Swaziland and is usually found in some shade under trees.

It is a distinctive species, easily distinguished by its large (up to 680 mm) pinnatifid leaves, large, showy, brick red to yellow inflorescences with rays up to 5 mm long and creamy white pappus.

A very popular garden plant, G. jamesonii grows prolifically throughout southern Africa and beyond. Flowering occurs mainly in spring (September to November), and autumn (March to May).

Gerbera natalensis Sch. Bip. Common name: Groenteebossie

Named for the province where it was first collected, this is often one of the first grassland species to appear after a burn. It is found from the eastern Cape to the Mpumalanga Drakensberg and Swaziland highlands.

The inflorescences, covered with long white silky hairs, emerge before the leaves, which only develop fully after flowering. This feature makes this species easy to recognize. Later in the season it may be confused with G. viridifolia, but the swollen, spindle-shaped or fusiform roots distinguish this species. The leaves are elliptical, up to 8 × 25 mm, with the base tapering into a slender petiole initially covered with long silky hairs which disappear later. The inflorescences are about 20 mm in diameter, rays white above, pink or reddish below, and the pappus is whitish. Flowering time from July to October.

Gerbera jamesonii flower.

Gerbera jamesonii.

Gerbera viridifolia (DC.) Sch. Bip.Common name: Griquateebossie

The specific epithet means ‘green leaves’, possibly describing its characteristic lack of white-felting on the underside of the leaves. The species has an extensive distribution in Africa, and is highly variable, especially in the northern KwaZulu-Natal–Swaziland–Mpumalanga area.

Hilliard (1977) places this taxon in G. ambigua, while Hansen (1984) regards it as a subspecies of G. natalensis.

Gerbera natalensis inflorescence emerging after a grassland burn. Note the absence of leaves.

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Herbarium specimen of Gerbera jamesonii.

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Gerbera viridifolia showing lack of white felting on the underside of the leaves.

In contrast to G. natalensis, the roots are not fusiform, and young leaves and flower stalks are only moderately hairy. The leaves are very variable in shape and size (up to 460 mm long) but usually with crenate, distinct teeth, and lacking white-felting on the underside.

The flower stalks are up to 760 mm long, and the inflorescences variable in size, 13–37 mm wide, with rays in all shades of pink, red, mauve, purple, crimson and magenta and occasionally yellow. The pappus may be purple or whitish.

This species grows fairly easily under cultivation, and the purple forms make very attractive garden subjects. Flowering time is mainly from September to November.

GlossaryModified from Leistner (2000).

bilabiate: 2-lipped, as when two or three lobes of a calyx or corolla stand separate as an upper lip from the others forming a lower lip.

cypsela: a type of achene or fruit formed from a one-loculed, inferior ovary, characteristic of plants in the Compositae/Asteraceae.

ebracteate: without bracts.

felted: (indumentum) matted, with intertwined hairs, resembling felt.

fusiform: thick, but tapering towards both ends, spindle-shaped.

involucre: a series of bracts, usually close together and appressed, below or around a compact head of flowers.

pappus: a series of bristles, hairs or scales round the base of the corolla, and later around the apex of the fruit.

peduncle: the stalk of an inflorescence or a stalk bearing a solitary flower in a 1-flowered inflorescence.

pinnatifid: with the margin pinnately lobed, but not to the midrib.

scape: a leafless flower or inflorescence stalk arising from the ground with radical leaves; naked peduncle.

staminode: an abortive or vestigial stamen.

AcknowledgementsThe following individuals or institutions are thanked as follows: University of KwaZulu-Natal Herbarium, Pietermaritzburg (NU) for images of G. parva and G. natalensis specimens, Tony Abbott and Elsa Pooley for the image of G. parva, Christina Curry (University of KwaZulu-Natal), Trevor Edwards (University of KwaZulu-Natal), Brian Tarr (KwaZulu-Natal National Botanical Garden), and the South African National Biodiversity Institute.

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Gerbera natalensis herbarium specimen showing the characteristic swollen, spindle-shaped or fusiform roots.

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References and further readingAMBROSIUS, P. 2003. How the Barberton Daisy got its name. Veld & Flora 89(1): 30.

CODD, L.E. 1979. The story of Barberton Daisy, Gerbera jamesonii. Veld & Flora December: 114–115.

DÜMMER, R.A. 1914. The South African Gerberas. Journal of the Royal Horticultural Society. Vol. XL. Part II: 236–262.

GERMISHUIZEN, G., MEYER, N.L., STEENKAMP, Y. & KEITH, M. (eds) 2006. A checklist of South African plants. South African National Botanical Diversity Network Report 41. SABONET, Pretoria.

GUNN, M & CODD, L.E. 1981. Botanical exploration of southern Africa. A.A.Balkema, Cape Town.

HANSEN, H.V. 1985. A taxonomic revision of the genus Gerbera (Compositae, Mutisieae) sections Gerbera, Parva, Piloselloides (in Africa), and Lasiopus. Opera Botanica 78: 5–36.

HILLIARD, O.M. 1977. Compositae in Natal. University of Natal Press, Pietermaritzburg.

HILLIARD, O.M. 1994. Taxonomic and nomenclatural changes in Compositae in Natal since publication in 1977. Royal Bot. Gardn. Edinburgh.

JOHNSON, I.M., EDWARDS, T.J. & CONDY, G. 2005. Gerbera aurantiaca. Flowering Plants of Africa 59: t. 2220.

KATINAS, L. 1998. The Mexican Chaptalia hintonii is a Gerbera (Asteraceae, Mutisieae). Novon 8(4): 380–385.

LEISTNER, O. A. 2000. Seed plants of southern Africa: families and genera. Strelitzia 10. National Botanical Institute, Pretoria.

POOLEY, E. 1998. A field guide to the wildflowers of KwaZulu-Natal and the eastern region. Natal Flora Publications Trust, Durban.

POOLEY, E. 2003. Mountain flowers. A field guide to the flora of the Drakensberg and Lesotho. The Flora Publications Trust, Durban.

RAIMONDO, D., VON STADEN, L., FODEN, W., VICTOR, J.E., HELME, N.A., TURNER, R.C., KAMUNDI, D.A. & MANYAMA, P.A. (eds) 2009. Red list of South African plants. Strelitzia 25. South African National Biodiversity Institute, Pretoria.

SCHULTZ BIPONTINUS, C.H. 1844. Enumeratio Compositarum a cl. Dr. Krauss annis 1838–40 in Capitate bona spei et portum Natalensem lectarum. In: Flora oder algemeine botanische Zeitung 27:767–783. Pustet & Manz, Regensburg.

Gerbera aurantiaca, old and new flowers. Photo: David Styles.