Proteaceae of South Africa

The Proteaceae is one of the most prominent flowering plant families. There are about 1400 species, in more than 60 genera such as Protea, Leucadendron, Leucospermum, Serruria and Paranomus. They occur in southern hemisphere countries. Australia has the most with 45 genera, followed by Africa with 14 genera, of which more than 330 species are found in the southwestern Cape. Other countries where they occur include Central and South America, islands east of New Guinea, New Caledonia, Madagascar, Southeast Asia, New Guinea and New Zealand. The family is an ancient one and existed in the time of the dinosaurs. The species vary from groundcovers to trees. This may be the reason behind its original naming by Carl Linnaeus, the father of taxonomy. Proteus was the mythological Greek god who could see the future and always spoke the truth, but in order to extract a prophecy, one had to grasp him tightly while he changed into numerous animate or inanimate forms.

TAXONOMIC CLASSIFICATION OF PROTEACEAE KINGDOM: Plantae CLADE: Tracheophytes CLADE: Angiosperms ORDER: Proteales FAMILY: Proteaceae SUBFAMILY: Proteoideae SUBFAMILY: Grevilleoideae

Plant taxonomy or classification is the science of naming organisms and placing them in a hierarchical structure, each level being given a name (e.g., kingdom, division (phylum), class, order, family, genus, species).

The family Proteaceae comprises of 83 genera with about 1,660 known species. Genera includes among others Protea, Leucospermum, Leucadendron, Mimetes, Serruria, Aulax, Banksia, Embothrium, Grevillea, Hakea and Macadamia. Species such including Telopea speciosissima), Protea cynaroides, and various species of Banksia, Grevillea, and Leucadendron are sought-after cut flowers, while the nuts of Macadamia integrifolia are widely grown commercially and consumed.

DISTRIBUTION OF PROTEACEAE

Proteaceae are mainly a Southern Hemisphere family, with its main centres of diversity in Australia and South Africa. However members of the Protea family also occur in Central Africa, South and Central America, India, eastern and south eastern Asia, and Oceania. Two species are known from New Zealand, although fossil pollen evidence suggests there were more previously.

Genera endemic to Australia include Banksia and Hakea. Telopea species which resembles the South African pincushion can be found growing in Australia and Tasmania while species from the Grevillea genus occurs in Australia and New Guinea and the Indonesian Islands. Embothrium species are native to South America. A single species is known to occur in Madagascar namely, Malagasia alticola.

The fossil record of some areas, such as New Zealand and Tasmania, show a greater biodiversity for Proteaceae than currently exists, which supports the fact that the distribution of many taxa has changed drastically with the passage of time and that the family has suffered a general decline, including high levels of extinction.

PHYSIOLOGICAL ADAPTATIONS Many of the Proteaceae have specialised proteoid roots, masses of lateral roots and hairs forming an absorptive surface, produced in the leaf litter layer during seasonal growth, and usually shrivelling at the end of the growth season. They are an adaptation to growth in poor, phosphorus-deficient soils, greatly increasing the plants’ access to scarce water and nutrients by exuding carboxylates that mobilise previously unavailable phosphorus. They also increase the root’s absorption surface, but this is a minor feature, as it also increases competition for nutrients against its own root clusters The Protea flower is not a flower, but a flower head or inflorescence, made up of many individual flowers grouped together on a rounded base or receptacle. What looks like the ‘petals’ of the protea ‘flower’ are modified leaves known as floral or involucral bracts. Look inside the cup of involucral bracts and you’ll see many long narrow flowers massed together in the centre. Protea seeds are covered in long hairs. These hairs aid in dispersal and germination: by expanding as the flower head opens, forcing them out of the head; once they are out, the hairs allow the seeds to be more easily moved by the wind; and once they land on the soil the hairs help to anchor the seed to the ground; and finally, the hairs help orientate the seed the right way round and channel water towards it.

PROTEAS OF SOUTH AFRICA

Protea cynaroides / King Protea Surely the best known protea, prized worldwide as a magnificent cut flower and in South Africa honoured as the national flower. The beautiful King protea is a woody shrub with thick stems and large dark green, glossy leaves. With the largest flowers in the family most plants are one metre in height when mature, but may vary according to locality and habitat from 0.35 m to 2 metres in height. The large cream to pink flower heads produce a disappointingly small amount of good seeds, only 1 – 30 percent of flowers result in seed. The seeds are quite large nuts, covered by hairs and stay in the old flower head for a year or more. They are released after a fire and dispersed by wind, rodents and birds. It is visited by nectar feeding birds, such as Sugarbirds and Sunbirds that pollinate the flowers. As they feed on the nectar in the flowers, their heads touch the pollen presenters and transfer pollen from flower to flower. The flowers are also visited by Scarab Beetles and Protea Beetles and many other insects, including bees. The species is not threatened and listed as Least Concerned on the IUCN Red Data Species List.

Protea magnifica / Queen Protea The Queen is one of the most attractive and sought after proteas and has the second largest flower head after P. cynaroides, the king protea. The strikingly beautiful and fluffy flower heads naturally vary in colour. Protea magnifica has a single, stout main stem and is indeed a variable protea, ranging from small erect trees of about 3m in height, to low sprawling shrubs among rocks and some that grow horizontally on the ground. The Queen is a re-seeder–the mother plant will die but the population survives because of the masses of seeds produced by the plant over the years and stored in old flower heads on the plant. Seeds are released after the fire and are dispersed by wind. Currently, this plant is Red Listed as Least Concern (LC), meaning that it is not threatened.

Protea neriifolia / Oleander-leaf protea Protea neriifolia is a very widespread species and occurs from sea-level to 1300 m altitude in the southern coastal mountain ranges from just east of Cape Town to Port Elizabeth. It grows mainly on soils derived from Table Mountain Sandstone, often in large stands. The variability in altitude and locality has led to a wide variation in both flower colour and flowering time. The flowering time is between February and November. Protea neriifolia was first discovered in 1597, was illustrated in 1605, and has the distinction of being the first protea ever to be mentioned in botanical literature. It took quite a while before it was officially recognised as a distinct species by the botanists and it was only described and named in 1810. The flowers are pollinated by scarab beetles, protea beetles and many other insects, as well as by birds. The birds are attracted by both the nectar and the insects visiting the flower. As with most proteas it is adapted to survive the fires by keeping its seeds safely in the old seedheads, which will only be stimulated to open and release the seeds when the plant dies or is killed by fire. These natural fires occur mainly in late summer or autumn and are followed by the first winter rains, which provide the moisture the young seedlings need to grow to a size at which they can survive the long, hot summer.

Protea nitida / Waboom protea It is widely distributed from the Bokkeveld escarpment to the Cape Peninsula and the Winterhoek Mountains and occurs at altitudes of 0-1200 m above sea level. Generally, Protea nitida is slow growing, 5m tall tree with white-grey bark and a trunk diameter of up to 400 mm. Under favourable conditions it may attain a height of up to 10 m and a trunk diameter of 1 m. As with others from the germination of seeds is fire dependant while pollination takes place through birds and insects. Protea nitida has various common names relating to its historical uses. Baboons would climb up the trees to feed on the nectar of the flowers, or baboon sentries would use trees as lookouts, and therefore the plant was given the name bobbejaansuikerbos. Brandhout, the Afrikaans word for firewood, indicates another use. The name waboom originates from the use of the wood for wheel rims and brake blocks of wagons. Interestingly, the name waboom was first recorded in 1720 and has thus been used for far longer than its scientific name. The wood was popular for the manufacture of ornamental furniture. It also made excellent charcoal. The bark was used for tanning leather. The tannin-rich bark was used to prepare an infusion for treating diarrhoea. The leaves were used for making ink. Either dry or fresh leaves were boiled up with a rusty iron nail and a piece of sugar candy. The resulting fluid (a decoction) is a fine blue-black, ideal for dyeing. These days, however, the greatest use for P. nitida is as a garden specimen.

Protea repens / Suikerkan protea Protea repens is a sturdy, dense shrub, 1-4 m tall, with linear hairless leaves and fairly large oblong flowers ranging in colour from cream to deep red. The shape of the flower heads is very distinctive, chalice-shaped, and forms an inverted, brown ‘ice-cream cone’ seedhead. Protea repens occurs in the southern part of South Africa and grows from the South West to the Eastern part of the Cape. Although it mostly occurs on the flats, coastal forelands, lower and middle mountain slopes, it has been found at altitudes up to 1500 metres and can be found scattered in between the other fynbos plants or in dense stands. The flowering period varies from winter flowering in the Western part of the range to summer flowering in the Eastern part. The flowers are pollinated by nectar-feeding birds, such as the Cape Sugarbird and sunbirds, and is also visited by Scarab Beetles and Protea Beetles and many other insects. The birds are attracted by the nectar as well as by the insects visiting the flowers. Protea repens has been exploited for centuries, as a source of firewood as well as for the nectar produced by the flowers and more recently by the cut flower industry. The abundantly produced nectar was collected in the past to be boiled into a sugary syrup, the so-called bossiestroop, meaning ‘bush syrup’, an essential component of 19th century medicine chests in the Cape. Protea repens was grown under glass in the Royal Collections at Kew in 1774 and flowered around 1780, the first protea ever to have been flowered in cultivation away from the Cape. It was also the first protea to have been grown outside in gardens in Australia, New Zealand and California from about 1890. Protea repens was the National Flower of South Africa up to 1976 and has inspired songs such as “Suikerbos ek wil jou he”, which was composed on Lion’s Head near Cape Town.

Protea scolymocephala / Skolliebos protea Protea scolymocephala is a small, neat, upright, well-branched shrub from a single main stem, 0.5-1.5 m tall. The leaves are long and narrow, shaped like an elongated, flattened spoon, 35-90 x 3-6 mm , hairless, with a pointed tip. The shrub bears an abundance of pink-tinged, creamy-green flowerheads from July to November. Protea scolymocephala is serotinous, i.e. it stores its seeds on the plant in the old flowerheads for years, and in the event of a fire, the bush is killed but the seeds survive and germinate en masse with the next rains. This fire-survival strategy works as long as the plants have sufficient time between fires to build up a new seed bank. The hairs on the seeds aid in dispersal and germination: by expanding as the flowerhead dries, forcing them out of the head; once they are out, the hairs allow the seeds to be more easily moved by the wind; and once they land on the soil the hairs help to anchor the seed to the ground; and finally, the hairs help orientate the seed the right way round and channel water towards it. The Skolliebos protea grows on sandy flats and coastal lowlands, often near drainage lines where there is a bit more moisture, from sea level to 400 m. Distribution ranges from the Gifberg and the Sandveld Flats to the Cape Peninsula and the Cape Flats with an outlying population near Hermanus, between Kleinmond and Hawston. It was once abundant on the Cape Flats but it has lost its habitat to urbanization, agriculture and the spread of invasive alien plants like Acacia cyclops (Rookrans) and A. saligna (Port Jackson). The species is currently listed as Vulnerable on the IUCN List of Red Data Plant Species and faces a high risk of extinction in the wild.

Protea odorata / Swartland Sugarbush

Protea odorata is a small, sparsely branched, evergreen, fynbos shrub, that grows about 0.7–1.2 m high. It has a single stem, hairless and brown with age. The leaves are hairless and have a bent spine at the tip that might slightly hurt the hand when touched. The spines are pinkish to reddish when leaves are young and turn black as they age. Protea odorata flowers from late summer to mid-winter (February to June), peaking in abundance in autumn (from March to April). The fruits are serotinous (retained within the seed head on the plant for years) and take about 7 months to ripen.

Protea odorata is endemic to Malmesbury Flats, from Klapmuts to Riverlands. It grows in slightly saline, klipheuwel gravels, mixed with sands in flatlands. Protea odorata almost became extinct (Rebelo 2001). It used to occur in Atlantis and Cape Flats Sand Fynbos, but is extinct in these areas. It survives in a transitional patch of Alluvium Fynbos/Swartland Shale Renosterveld.

Protea odorata flowers have a slight sweet fragrance and are pollinated by wasps. When the proteas die or are killed by fire, the serotinous (retained) fruits open and the seeds fall out and become available food for rodents and birds until the seeds germinate after the autumn and winter rain. The seeds of Protea odorata have a softer coat with hairs that hold the seed into the damp soil after they have been dispersed by wind.

Protea odorata is assessed as Critically Endangered (CR). This is caused by too frequent fire, agricultural expansion, invasive alien plants, overgrazing and road construction. The bulk of the populations are extinct, and only a single marginal population of about 5 plants still survive.

Protea burchellii / Blinksuikerbos Burchell’s sugarbush is an erect, evergreen shrub, up to 2m high. Branches are produced from a single stem. Leaves are olive-green with fine black points, linear to narrowly oblong. Flower are cream-coloured to deep carmine. Fruits with nut seeds, are retained and stored on plants. It is a moderately fast grower, flowering within two years from planting out into correct growing conditions. Flowering time from June to August. Protea burchellii occurs in varied habitats and seems to prefer rich, well-drained soils, in full sun on the lower mountain slopes at altitudes of 100-850 m. It is found in the Hottentots-Holland to Olifants River Mountains, the Cape to Hopefield Flats, Piketberg and the upper Breede River Valley. It also occurs in isolated populations at Witzenbergvlakte. Protea burchellii is pollinated by birds. Seeds are retained on plants in seed heads, to be released once dried out and dispersed by wind. Fynbos is a fire-dependent ecosystem and Burchell’s sugarbush has adapted to this. During a fire, the seeds are safe in their seed heads and survive the blaze to replace the mother plants which are killed in the fire. A root pathogen, Phytophtera, is detrimental to Proteaceae. Symptoms include the plant looking wilted and dry followed by yellowing and death.

LEUCADENDRONS OF SOUTH AFRICA

Leucadendron argenteum / Silver Tree Conservation Status: Rare & Endangered The name Leucadendron is from the Greek, leukos, meaning white, and dendron, tree. This genus is named after and based on the silver tree, from its common name witteboom, meaning white tree, as it was known in the 1690s when the genus was first named. The species name argenteum is Latin and means silver, silvery or ornamented with silver, referring to its striking silver foliage. Leucadendron argenteum is an erect, well-proportioned, ornamental tree, 7 – 10 m tall, with a stout trunk and thick, grey bark. The leaves are silver-grey, covered on both surfaces with thousands of tiny, soft, silvery hairs and fringed with long white hairs. The characteristic silver sheen of the leaves is caused by the hairs. The intensity of the sheen varies with the weather. They are at their most silver in hot, dry weather, when the hairs lie flat to protect the leaves from drying out. In wet weather they are not quite as dazzling, as the hairs stand more erect to allow for free air circulation. Like all leucadendrons, it is dioecious, meaning that the male and female flowers are borne on separate plants. The flowers are in dense heads at the branch tips. The flower heads have a faint, pleasant scent, which attracts the small beetles that do the job of pollination. Leucadendron argenteum occurs on cool, eastern and southerly slopes on granite-derived clay soils, at 100-150 m, and trees are conspicuous on the slopes above Kirstenbosch. There are eight populations spread over an 11 km range with a few outliers in Somerset West, Paarl and Stellenbosch. Only the populations at Rhodes Memorial and Tafelberg Road grow on shale-derived soil. The largest populations are at Kirstenbosch, Wynberg Hill and Lion’s Head. More than half of the wild trees grow in the Kirstenbosch population.

Leucadendron chamelaea / Witzenberg conebush

Conservation Status: Critically Endangered

Leucadendron means ‘white tree’. The species name chamelaea, is derived from the Greek word chamae, which means ‘low, or on the ground’, referring to a creeping habit.

Leucadendron chamelaea is one of the iconic species of the Upper Breede River Valley. This species once occurred commonly in this area, but through the intensification of agricultural practices, it has suffered severe population reduction and is now on the brink of extinction in the wild.

Leucadendron chamelaea is Critically Endangered (CR). A past population reduction of at least 60% is estimated, based on 55% habitat loss because of agriculture and the local extinction of 52% of subpopulations known through herbarium records. Remaining subpopulations are fragmented and largely confined to road verges and adjacent areas. This species is also an indicator of special habitat, and wherever there are populations of this species, numerous other threatened plants at these sites.

This species grows in sandy flats and has rather large flowerheads. It is a single-stemmed species, so is killed by fire and regenerates from seeds. The seeds are held in the cones, but the cones are not tightly closed, so once the seeds are ripe, they can fall to ground out of the cones if they are knocked or blown by a strong wind. Plants flower in spring (September to October), are strongly scented and are pollinated by insects. Seeds ripen within two months.

Leucadendron discolor / Piketberg conebush Conservation Status: Vulnerable The juvenile phase of Leucadendron discolor is a small densely bushy shrub. A year or two after planting out it matures and gradually sends up long erect flowering shoots reaching an eventual height of 2 m and a spread of 1-1.5 m. One of the distinguishing features of this Leucadendron are the grey-green broadly ovate leaves. Another, are the flowerheads which appear in spring. Male flowerheads are bright red and yellow surrounded by creamy yellow involucral leaves. Female plants form a grey-green cone surrounded by pale green involucral leaves that almost enclose the flowerhead. Winged seeds are found inside the cone on the female plant. They ripen a few months after flowering and are retained on the plant. Leucadendron discolor is listed as VU (Vulnerable) and the population is seen to be declining. One of the threats is wild flower harvesting. Male plants are harvested for cut flowers leaving skewed population ratios of male to females. As sub-populations are small, this drastically affects pollination and seed production and therefore future generations. Another threat is habitat loss due to expanding fruit and protea orchards. Coupled with encroaching protea orchards, is the threat of hybridization with other species of Leucadendron planted in the orchards. The Piketberg conebush is pollinated by insects carrying pollen from the male flower onto the female cone. Pollination occurs and seeds form and grow within this cone. The cone remains tightly closed protecting the seeds from the weather and from being eaten by animals. Plants are killed by fire. After a fire all the cones open to release the seed which drops to the ground, germinates and establishes a new population.

Leucadendron macowanii / Wattle-leaf conebush Conservation Status: Critically Endangered Leucadendron macowanii are quite fast growing and should have attained a height of 2 m in three to four years from the date of sowing. The leaves are narrowly oblanceolate, thinly silky at the base, tipped with a fine recurved point. The leaves on female plants are generally longer than those on male plants. The plants flower during winter (July and August), with the male plant looking the most attractive. The male flowerheads are made up of many tiny bright yellow flowers crowded together in small cone-like heads, about 14 mm in diameter and 15 mm long. The female flowerhead is also cone-like, green turning red or reddish brown, 23 mm long. The female flowers can be seen as large yellow stigmas protruding from underneath the green or red scales of the ‘cone’. Neither the male nor the female have any scent. According to the Red List of South African plants, Leucadendron macowanii is Critically Endangered (CR). It is a rare species with a small natural range, and it is endemic to the Cape Peninsula. One small subpopulation remains, the only other known subpopulation is now extinct due to urban expansion.

Leucadendron rubrum / Spinning top conebush Leucadendron rubrum is an erect shrub growing to 2.5 m tall. Plants develop from a single-stemmed base, with male plants being bushier and having smaller leaves than female plants. Young branches in male plants are flexible, sometimes a purplish colour, and covered with fine hairs. As they mature they become smooth. Young female branches are much hairier than the male branches but are fewer in number and fairly stout. Leaves are covered with silver-white hairs when soft and young. On maturing they lose most of the hairs and are a green-grey colour. The female leaves are twisted near the base, which, together with the shape of the female cone imitates the motion and shape of a spinning top. Male and female flower heads are clustered along the ends of branches. Male flower heads are very small, measuring about 11 mm long but are a vibrant bright yellow, and flower in a golden plume. Female flower heads form a cone about 40 mm long. The yellow female stigmas pop out at the top of the cone in a tuft. The bracts forming the cone are a colourful mix of yellow, green, blue and red. Leucadendron rubrum occurs across most of the Fynbos biome from the Bokkeveld escarpment in the north, Table Mountain in the southwest, to the Baviaanskloof Mountains in the east. Although it occurs across a large area, L. rubrum does not show much variation in habit. It grows on dry slopes on Cape Granite or Table Mountain Sandstone at an altitude of between 250 and 1500 m. Male Leucadendron plants produce lots of pollen that is blown by the wind onto the female flowers. Pollination occurs and seeds develop inside the cone on the female plant. Seeds are retained inside the cones for a number of years or until the plant is killed by fire. Each seed has a hairy parachute attached to it, so when the cone opens they float away carried by the wind away from the mother plant.

Leucadendron salignum / Common sunshine conebush

Leucadendrons are dioecious, i.e. separate male and female plants. This is unusual in the protea family. Its long flowering season (May – Dec), coupled with colourful leaves and bracts surrounding the flowers, make this species an attractive garden plant.

There are a range of plants in cultivation, which differ markedly from the usual parent species, most often in growth form, leaf- and bract colour and flowering time. They may be either selections, known as ‘cultivars’ or hybrids, i.e. ‘crosses’ between species. Many hybrids and cultivars have been produced in South Africa, New Zealand, Australia and other countries growing Proteaceae

It is common from Port Elizabeth in the east, to north of Ceres in the west. It occurs on a wide range of soil types, from sea level to an altitude of 2000 m and is quite variable in leaf size as well as leaf- and bract colour.

Leucadendron sessile / Sun conebush Conservation Status: Near Threatened Leucadendron sessile is a 1-2 m tall, rounded, dense, bushy shrub arising from a single stem at ground level, and usually branching low down. Branches are stout and short. Like all leucadendrons this species is dioecious, i.e. male and female flowers are borne on separate plants. The leaves are narrowly elliptical, up to 64 mm long in the male and up to 80 mm long in the female, hairless but not shiny, with red margins and tipped with a blunt, red, recurved point. The leaves increase in size and become more crowded towards the end of the stems. Another characteristic of leucadendrons is the way the leaves that surround the inflorescence, known as the involucral leaves, turn a different colour while the plant is in flower. In Leucadendron sessile the involucral leaves are yellow reddening with age, and the male plants are usually a brighter yellow than the female. Flowering time is mid-winter to early spring (July to August). Both male and female flowers produce nectar from nectaries found inside the perianth tube at the base of the ovary and between each perianth segment, known as hypogynous scales, and thought to be modified petals. Both the male and female flowers contain the same number of floral parts, but in the male flower, the gynoecium (female part) is sterile, and in the female flower, the stamens are sterile. Leucadendron sessile occurs on lower to middle slopes and flats, 10-600 m, in granitic soils, from the Witzenberg, Elandskloof to Slanghoek Mountains and the Hottentots-Holland Mountains from Jonkershoek to Kogelberg. It occurs where the annual rainfall is relatively high and is often supplemented by coastal mist and clouds. It can be seen growing on Sir Lowry’s Pass and along the False Bay coast at Kogel Bay. Leucadendron sessile is Near Threatened (SANBI Interim Red List 2007), meaning that although not at risk it is likely to become threatened in the near future.

Leucadendron strobilinum / Peninsula conebush
Conservation Status: Near Threatened

The word Leucadendron is derived from the Greek word leucos, meaning white, and dendron, meaning a tree, referring to the silver tree, Leucadendron argenteum. Strobilinum comes from the Greek strobilos (cone) and is a reference to the resemblance of this plant’s cones to pine cones.

Leucadendron strobilinum is a large, single-stemmed shrub that branches from the base. It grows up to 2.6 m tall. The leaves are dark green and oval. The leaf tips are red and recurved with a fine point. The fruiting cones are ovoid and hairless and contain flat, winged seeds. It flowers from September-October. It occurs on the Cape Peninsula, and the distribution ranges from Table Mountain to Kommetjie. It grows on damp, rocky slopes.

The plants regenerate from seed. The seeds are stored in the cones on female plants from which they are released after fire. The seeds are dispersed by wind. Leucadendron strobilinum has plants of separate sexes, the male and female flowers occur on separate plants. Flowers are pollinated by small beetles.

Leucadendron teretifolium / Strawberry conebush Conservation Status: Near Threatened The strawberry conebush has a single stem but with dense, bushy branching, creating a compact shrub that grows up to 1 m tall. Leaves are dark green and needle-like. Male plants produce clusters of small, yellow flowerheads at the branch tips. These contain large amounts of pollen, and if you brush past during flowering, a small cloud of pollen will appear. Female plants produce cones that when young, are small and green. Once fertilized and mature, they turn a beautiful rosy red, sometimes dark maroon. Flowering occurs in spring (August–September), and the seeds are held safe on the mother plant within the cones. The strawberry conebush occurs in the central and south-western Cape. Its preferred habitat is mountainous areas and appears along the low foothills all the way up mountain slopes up to 1 350 m altitude. It grows in fynbos and the renosterveld ecotone (the border of vegetation where fynbos meets renosterveld). Soil type can either be shale clay soils (which support renosterveld) or rocky sandstone leached soils (which support fynbos). The Leucadendron genus is dioecious, meaning that males and females occur as separate plants. Male plants produce pollen and female plants produce sticky stigmas. The wind acts as pollinator and blows the pollen from the male plant onto the sticky stigmas on the female plant. Fertilization occurs and many little seeds form and grow within a hard cone on the female plant.

Leucadendron tinctum / Spicy conebush Leucadendron tinctum is a perennial, bushy shrub growing up to 1.3 m tall. The plant grows from a single stem at the base with side branches bearing oblong, elliptical leaves. The leaves are a pale grey-green and during flowering, end in colourful bracts which surround the inflorescence. The male and female inflorescences are borne on separate plants. The flowers are arranged up of a single terminal head. The male flowerhead is yellow, whereas the female flowerhead is a shiny maroon surrounded by 50 oily basal scales. The flowering period for the spicy conebush is in July. Flowers have a pleasant spicy, musk scent, which gives rise to its common name. The female fruiting cones are maroon and contain round nut-like fruits. As this is a widespread species, its distribution covers a rather large area of the Cape, starting on the Agulhas plain and ending in the Langeberg. The plants occur in stony, sandstone soils at an altitude of 20-300 m. Populations have been seen in Potberg, Elim and the Bredasdorp flats to Kleinmond, Groenland, the eastern Hottentots-Holland Mountains and Stettynskloof Mountains. Their higher distribution includes Gysmanshoek, Garcia’s and Robinson’s Passes, the Swartberg and Rooiberg Passes and Matroosberg in the Hex River Mountains. These plants prefer well-drained, sandy soils and seem adaptable to many climatic regions. An unusual characteristic of Leucadendron tinctum is that its anthers act as the pollen-presenter, assuming an erect position for that purpose, instead of the short male pistil that ends at the throat of the perianth tube. The strong odour, conspicuously coloured bracts and nectar-producing hypogynous scales of the inflorescence, indicate insect pollination. The fruit ripens four months after pollination. The seed distribution is rather limited, as it has to rely on animals to disperse its nut-like fruit. An interesting fact about this Leucadendron is that its inflorescence looks different at different times of the year and various forms are present in different distributions due to its limited seed distribution. It has an average speed of growth and can live for many years under correct, well-drained conditions.

LEUCOSPERMUM OF SOUTH AFRICA

Leucospermum bolusii / Gordon’s Bay Pincushion Conservation Status: Rare and Endemic The Gordon’s Bay pincushion is quite unusual for a pincushion in that its flowers are a creamy white colour where they are normally a striking orange or yellow, and fragrant where they normally are not. This pincushion is found in a small area on rocky, west-facing, sandstone slopes of the mountains above Clarence Drive, between Gordon’s Bay and Kogel Bay, overlooking False Bay. Leucospermum bolusii grows into a rounded shrub, about 1.5 m tall. The leaves are hairy when young, and have one apical tooth. The flowerheads are flat and rounded, about 20 mm in diameter, each one made up of 50 – 100 small tubular flowers. The buds are encased by pinkish red bracts, and the flowers are creamy white, with a strong, sweet fragrance that is very noticeable on windless evenings. Flowering time is spring to midsummer (September to December), peaking in late October to early November. The nut-like seeds drop out of the flowers about two months after flowering. The flowers are pollinated by bees, wasps, flies, butterflies and moths. The seeds are dispersed by ants that are attracted to the fleshy skin that covers the seed, called an elaisome. The ants carry the seeds to their underground nests where they consume the elaisome, but the seed remains safe until conditions are right for them to germinate. Leucospermum bolusii is killed by fire, i.e. it is not a resprouter, but its seeds survive. The genus Leucospermum is named from the Greek leukos meaning white, and sperma seed, referring to the white or light-coloured seeds of many species. This species is named after Harry Bolus (1834-1911), a South African botanist and founder of the Bolus Herbarium in Cape Town. It gets its common name from its restricted habitat above Gordon’s Bay, and its Afrikaans name refers to its white flowers and the tick-like appearance of the seeds (wit meaning white, luisies are little ticks and a bos is a bush). Many other pincushions are also called luisiesbos.

Leucospermum conocarpodendron subsp. conocarpodendron / Grey Tree Pincushion Conservation Status: Vulnerable Leucospermum conocarpodendron subsp. conocarpodendron is a tree-like shrub, 3-5 m tall and 3-6 m in diameter, with a dense, rounded habit and often with a gnarled and bent shape. Flowering stems are stout and rigid and are covered with a shaggy layer of fine, silky hairs. Leaves are stalkless (sessile) and have a rounded apex with 3-10 teeth. The leaves are grey-green and covered in a dense layer of fine, short, curly hairs, which require a magnifying lens to be seen properly, and which can be rubbed off. The leaves lose this covering and become hairless after several years (glabrescent). It occurs mainly in the richer, heavy clay soils derived from Cape Granite and shales but also occurs in sandy soils derived from Table Mountain Sandstone in some places. Its habitat is very windy, cool and wet in winter and hot and dry in summer. Leucospermum conocarpodendron subsp. conocarpodendron is endemic to the Cape Peninsula, where it grows on well-drained, north- or west-facing rocky slopes, from sea level to 160 m, over a limited area from the eastern slopes of Devil’s Peak, along the northern and western slopes of Table Mountain and the Twelve Apostles to Llandudno. The brightly coloured and unscented flower heads of Leucospermum conocarpodendron subsp. conocarpodendron are pollinated by birds, sugarbirds and sunbirds being the main pollinators. They perch on the flower heads or the stems and probe into the flower head in search of nectar, and in so doing, their heads and necks come into contact with the pollen presenters. The flower heads are also visited by bees and beetles, in particular the Protea Beetle (Tricosthetha fascicularis) but they do not assist in pollination as they don’t come into contact with the pollen presenter. Leucospermum conocarpodendron subsp. conocarpodendron is fairly fire-resistant and is known to survive some fires, thanks to its height and thick bark. Plants that have survived a fire develop an umbrella-like shape as their lower branches are burned off and new growth is produced from the surviving upper branches. Nevertheless, this species is not a resprouter and plants often die if burned. Leucospermum conocarpodendron seeds have an elaiosome that attracts ants. The ants carry the seeds away to their nests to consume the elaiosome, but disperse the seeds and keep them safe in the process. This is known as myrmecochory. After a fire, the seeds stored in the ants nests germinate.

Leucospermum cordifolium / Common Pincushion Conservation Status: Rare Leucospermum cordifolium is a rounded spreading shrub up to 2 m in diameter and about 1,5 m high, with a single main stem and horizontally spreading stems, hard green leaves and 1 to 3 large inflorescences borne at the end on the stem. The inflorescences consist of a large number of small flowers. It is the stiff protruding styles of the flowers which are the source of the common name “pincushion” for this genus. It grows in acid, nutrient-poor soils in a fairly small area in the Western Cape, from the Kogelberg to the Soetanysberg near Bredasdorp. It is part of the Cape Floral Kingdom and occurs only in the winter rainfall area with its wet winters from May to September and hot, dry summers from December to the end of February. In the early hours of the morning the abundant nectar flow attracts the Cape Sugarbird and three species of Sunbird, and a variety of small insects, which in turn attract the he insectivorous birds that consume the small insects. The flowers are not self-pollinating and depend on the Protea Scarab Beetles and the birds for pollination. The birds are accustomed to the visitors in the Garden and provide great photo opportunities when feeding on the flowers. Only a few large, hard, nut-like seeds are produced by each inflorescence. In their natural environment the seeds are collected by ants, stored in the soil, and germinate only after a fire has killed the mature plants and returned the nutrients back to the soil.

Leucospermum grandiflorum / Rainbow Pincushion Conservation Status: Endangered Leucospermum grandiflorum is an erect, lanky shrub growing 1.5-2.5 m tall. Leaves are grey and hairy and 50-80 mm long. The tips of the leaves have 3 glandular teeth. Flower buds have a hairy appearance and spinning top shape while tightly closed. The perianths (the pins of the pincushion) open in a spiral direction and are bright yellow with a pink pollen presenter on the tip. On maturing the perianth fades to orange and later red. Flowering occurs from July-December and seeds are released two months after flowering. Leucospermum grandiflorum is a fynbos species and is endemic to Boland Granite Fynbos. This vegetation type is characterized by plains and hills with domed granite outcrops. Granite-derived soils are richer than sandstone soils and therefore have been rapidly transformed for agricultural use. In addition to this species being classified as endangered, the entire vegetation type is also classed as Endangered. On the mountain it grows in scattered stands on hot, dry slopes at an altitude of between 80 and 500 m. Its natural distribution is Paarl Mountain, Simonsberg, Paardeberg, Durbanville Hills and Berg River Valley. Endangered! It is estimated that populations have decreased by more than 50% over the past 60 years. Fewer than five remaining locations with plants growing in the wild are known. These populations are not healthy as the habitat quality and number of mature plants is declining. Reasons for this include habitat loss due to agriculture, afforestation and invader alien plants, too frequent wildfires, and illegal wild flower harvesting. Fynbos is a fire-driven system which means the vegetation requires fire once every 10-15 years to regenerate and rejuvenate itself. Fire moves through the fynbos in late summer killing all the mother plants of L. grandiflorum. In early winter the rains begin and new seedlings germinate from the ants’ nests where they were safely protected from the fire. Flowers are pollinated by the Cape sugarbird. After pollination, seeds are produced and released and fall to the ground. The seed has a white waxy coating; this is called an elaiosome and is a favourite food of indigenous ants. Ants disperse the seed by carrying it underground to their nests where they eat off the elaisome leaving the seed in safe storage, hidden away from predators like mice.

Leucospermum oleifolium / Overberg Pincushion Conservation Status: Near Threatened Leucospermum oleifolium is an erect, rounded shrub up to 1 m tall and about 1.5 m in diameter, with a single main stem. The inflorescences are about 4 cm across, in clusters of up to five individual flowerheads at the end of the branches. They open over an extended period of time with the result that one plant provides a colourful spectacle for about four months, from the middle of August to the end of December. The flowerheads open as a pale yellow, which soon turns orange and becomes a brilliant crimson with age. The natural habitat of Leucospermum oleifolium is in the South-Western Cape, from the Du Toit’s Kloof to the Caledon Swartberg. The plants often occur in extensive, dense stands, where they protect one another from prevailing winds. Together with the other fynbos plants a dense cover is established which prevents compaction, keeps the soil cool and reduces the rate of evaporation. During flowering time numerous birds pollinate the flowers and are an added attraction. In the early hours of the morning the abundant nectar flow attracts a variety of small insects, which in their turn attract the Cape Sugar bird as well as Sunbirds. These insectivorous birds make use of the small insects as well as the nectar and in the process transfer pollen from one flower to the other. The flowers are not self-pollinating and depend on the small Scarab beetles and the birds. The birds are accustomed to the visitors in the garden and provide great photo opportunities when feeding on the flowers. Only a few large hard nut-like seeds are produced by each inflorescence and in their natural environment the seeds are collected by ants, stored in the soil and germinate only after a fire has killed the mature plants and returned the nutrients back to the soil.

Leucospermum patersonii / Silver-edge Pincushion Conservation Status: Vulnerable The silver-edge pincushion is a large rounded shrub or small tree with broad green leaves giving it an appearance similar to Leucospermum conocarpodendron. It has beautiful golden orb-like flower heads and is one of the few calcium-loving members of the protea family. Large ornamental rounded shrub or small tree with a single stout main trunk covered with thick corky bark. It bears bright orange pincushion flowers, usually in clusters of three. It is tree-like in stature, up to 4 m tall and between 3 and 6 m in diameter. It is clothed with attractive, light green, broadly oblong, glabrous (hairless) leaves which are toothed at the apex. The leaves are tightly packed and overlapping, giving the plant a bulky, leafy look. The flower heads are bright orange to crimson and very showy. They are similar to, but a bit smaller than those of Leucospermum cordifolium. The flower heads are partly enclosed by the uppermost leaves. Flowers are produced from July to December. Leucospermum patersonii is restricted to alkaline soils on limestone outcrops between Kleinmond and Cape Agulhas. Some of its habitat has been destroyed by urban expansion, development of cut flower farms and alien invasive plants, and it is locally extinct in Hermanus. It is not currently threatened and is listed as Vulnerable (VU), but climate change predictions indicate a significant population reduction by 2025. The brightly coloured, orange flower heads of Leucospermum patersonii are pollinated by sugarbirds and sunbirds. They perch on the flower heads or the stems and probe into the flower head in search of nectar, and in so doing, their heads and necks come into contact with the pollen presenters. The flower heads are also visited by bees and beetles, in particular the green protea beetle (Trichostetha fascicularis), but they do not assist in pollination as they don’t come into contact with the pollen presenter.

Leucospermum reflexum / Rocket Pincushion Conservation Status: Near Threatened These lovely, tall, upright pincushion plants with bright, deep orange to crimson or clear yellow flower heads atop the silvery-grey columns of densely leaved branches are real show-stoppers. Leucospermum reflexum is a rounded, silvery grey-leaved shrub up to 4 m tall with either yellow (var. lutea) or deep orange to crimson (var. reflexum) flower heads which flower from spring to midsummer in the southern hemisphere (August to December). The more or less elliptic, simple leaves are covered with dense grey hairs and have 2-3 glandular teeth at the tip. The rounded flower heads occur at the ends of the branches and are between 80-100 mm across. Each flower is tube-like and made up of velvety floral parts called the perianth segments on which the anthers are placed. These surround the long style which escapes from the tube to form the ‘pin’ and the perianths then curl away from it. Young flowers are curved to start with, and then bend back toward the stem as they mature. The common name of rocket pincushion is very apt because these downward bent styles look like the fiery tail of a rocket trailing behind it. For the same reason, the Afrikaans name perdekop is a good one as the mature flower head looks like a horse’s head with its mane blowing back in the wind. The seeds ripen about 1-2 months after flowering and are then released. L. reflexum has a medium growth rate and usually flowers in its third year. Plants have been known to grow for about 20 years before becoming senescent, but are usually far less long-lived than this. Although the general area in the Cederberg, from Wuppertal to Pakhuis, where the plants occur, tends to be arid, these plants are found on sandstone soils near water sources such as streams. The plants usually occur in groups of a few hundred at altitudes ranging between 1000-2000 m. The yellow variety, L. reflexum var. luteum, comes from the Heuningvlei area in the Cederberg.

Leucospermum rodolentum / Sandveld Pincushion Conservation Status: Vulnerable Leucospermum rodolentum is an upright spreading shrub of up to 3 m (10 ft) high and 4 m (13½ ft) in diameter, that develops from a single trunk at its base of 8–14 cm (3¼–4¾ in) thick, with a smooth, grey bark. The flowering stems are rising up 5–7 mm in diameter and grey due to a dense layer of grey fine crisped hairs. The leaves are elliptic to wedge-shaped, 4–6½ cm long and ¾–1½ cm wide, with a blunt or tapering base, a rounded or bitten-off tip with three to six teeth, and a grey surface due to a dense layer of fine, short, crisped hairs. The flower heads are globe-shaped, 3–3½ cm across, seated or with a short stalk of up to ½ cm long, occurring in groups of two, three or four, rarely individually. The common base of the flowers in the same head is flattened cone-shaped, about 1 cm across. The greyish bracts that subtend the head are oval with a pointy tip, long and wide, tightly overlapping, cartilaginous in consistency. The sandveld pincushion can naturally be found from Darling in the south, through the Hopefield, Piketberg, and Clanwilliam districts, to the Heerenlogementberg and the Nardouw Pass. An isolated population can be found south of the Brandvlei Dam. The species is a prominent constituent of sandveld vegetation that grows on the sandy flats of the west of the Western Cape province, between sea-level and 250 m or rarely 300 m (1000 ft) altitude. It occurs only in loose, very often stabilised Tertiary or Quaternary drift sands. The species is pollinated by insects, such as honey bees, monkey beetle), skippers, and also visited by birds like the Cape sugarbird, orange-breasted sunbird, and Cape weaver. The ripe fruits fall to the ground about two months after flowering, where these are collected by native ants, that carry them to their nests. Here they remain underground, safe for fire, seed-eating rodents and birds, until an overhead fire clears the vegetation and triggers the seeds to germinate.