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Identifying Conebushes - Leucadendron

The genus Leucadendron is easily identified by the woody cones borne on some plants in each species, and the plants of separate sexes. About half the plants (the females) in any population produce cones which bear seeds, whereas male plants bear cones for the short flowering seasons and never produce seed. Cones consist of spirally arranged floral bracts, which cover and partially hide a floret. After flowering the bracts become hard and woody, forming the conspicuous fruiting cones.

The perianth segments are fused into a tube with only tips free. In male florets the ovaries are reduced. In female florets the style is straight or slightly curved, with a large terminal portion, usually with a slit on the top for receiving pollen.

Male plants are usually more branched than female plants and are often slightly larger with smaller leaves and cones. Male florets are subtended by an inconspicuous floral bract. The perianth segments are free on top half of floret, but fused into a tube below. The style is always present, and serves as a pollen presenter, but other female organs are reduced.

The identification of Conebushes is difficult as it relies very heavily on fruit morphology. Fortunately, many species store their fruit for most of the year in the female cones. In species which do not store their fruit, cones laying on the ground should be collected if it is intended to identify the species. In addition, several other characters are required, from both male and female plants, to identify most species. These include: cone shape and size, the habit of the species and the leaf size, shape and hairyness.

Key

1. Fruit flat with wings, or flat with a prominent ridged edge, never hairy; cones always present on female bushes (EXCEPTION INCLUDED: L. floridum); female floral bracts always encircling the floret base; female ovary usually hairless; male style usually hairless Subgenus ALATOSPERMAE: goto 2

1' Fruit a nut, large and round on both sides, or rounded on one side only (ventricose), occasionally with a pointed end, occasionally with a ridged edge, often hairy; cones not always present year-round on female bushes (find some on ground!); female floral bracts usually free, female ovary usually hairy, male style usually hairy at base Subgenus LEUCADENDRON: goto 5

Subgenus LEUCADENDRON

The subgenus Leucadendron contains a diversity of fruit types and is divided into two major groups:

(A) MEMBRANACEA: The lower part of the mature female perianth is membranous and grows with the fruit to surround it in the cone.

Villosa, Membranacea, Carinata, Uniflora, Aliena, Nervosa, Leucadendron

(B) CRUCIFERAE: The mature female perianth becomes detached from the ovary and splits to form a cross which is pushed up on the top of the growing fruit.

Cuneata, Nucifera, Ventricosa

However, these features are not readily apparent and are not included in the key below. If any cones are available however, search for the perianth: you will find it a useful feature when it is available.

Only a few species store the fruit in cones (serotiny) until a fire kills the plants and allows the fruit to be released. These occur in sections Villosa (4 out of 10 species), Nervosa (1/1) and Leucadendron (4/4). Identification of these species is thus simple as fruit are continuously available.

5. Fruit a biconvex nut or nutlet goto 6

5' Fruit with one side flat, the other rounded, without hairs; cone with elongated stalk and leaf-like involucral bracts goto Section VENTRICOSA

6. Involucral bracts discrete goto 7

6' Involucral bracts tightly stuck together, mature female perianth segments abutting, fruit distinctly pointed below, mottled, with short sparse hairs goto Section CUNEATA

7. Fruit uniform in colour, not mottled, not prominently pointed at the tip goto 8

7' Fruit mottled, with a sharp pointed tip to the fruit, sparsely or densely covered in hairs, never hairless goto Section VILLOSA

8. Fruit not keeled goto 9

8' Fruit elongated, grey, without hairs, with a doubly ridge on the upper surface goto Section CARINATA

9. Female cones containing more than 10 flowers goto 10

9' Female cones containing only one to three flowers or fruit goto Section UNIFLORA

10. Fruit without hairs goto 11

10' Fruit with hairs goto 12

11. Female perianth segments free, segments hairy above, membranous below goto Section MEMBRANACEA

11' Female perianth segments abutting, segments hairy above, spreading and narrow below goto Section NUCIFERA

12. Female perianth plumed, style persisting on fruit to form parachute, fruit without noticeable elaiosome goto Section LEUCADENDRON

12' Female perianth hairless, style persisting in fruit; fruit without noticeable elaiosome, fringed with 8 mm long hairs goto Section NERVOSA

12" Female perianth not plumed, style not persisting in fruit; fruit with conspicuous basal elaiosome goto Section ALIENA

Subgenus ALATOSPERMAE

The subgenus Alatospermae is divided into four sections. More species are concentrated in the southwestern Cape than in Subgenus Leucadendron.

The subgenus is characterized by the fruit being winged, strongly ridged (one species), or triangular in cross-section. The cone bracts are always tightly clasping one another, and separating only in the dried cones. This contrasts with the section Leucadendron which has the cone bracts discrete.

2. Fruit winged, flattened or triangular in shape, never with a rough median ridge, involucral bracts not oily goto 3

2' Fruit flat on one side, with a conspicuous rough median ridge, apex with a small point; involucral bracts dark brown and oily, female perianth segments less than 7 mm long, overlapping like tiles goto Section BRUNNEOBRACTEATA

3. Fruit greater than 5 mm broad, with prominent wing (except some forms of L. xanthoconus), female perianth segments hairless goto 4

3' Fruit less than 5 mm broad, narrowly winged or triangular in section; female perianth segments 2-5 mm long, with short hairs, segments abutting one another without overlap goto Section TRIGONA

4. Fruit compressed, flat and winged; leaves are needle-shaped and round in cross-section in young plants, but may become flat in older plants; bracts of female cones tightly pressed together, not overlapping with bracts above goto Section COMPRESSA

4' Fruit broadly winged, usually triangular in shape; leaves always flat; bracts of female cones tightly pressed together, imbricate (overlapping like tiles with the bracts above) goto Section ALATA

Section MEMBRANACEA: The Arid Conebushes

The Arid Conebushes are characterized by their biconvex, hairless, dark brown, nut-like fruit. The fruit are released after they have ripened. The fruit are thus almost identical to the section Nucifera. They are further distinguished by having the perianth segments of the mature female florets free, and not forming a tube (and therefore fall apart on dissection), with hairs on the upper florets and a membranous texture at the base. All four species are confined to the north-western, arid, mountain ranges. The leaf shape and the size of the female florets are important in distinguishing species.

Leucadendron remotum NIEUWOUDVILLE ARID CONEBUSH
Shrubs up to 1.7 m, single stemmed, but branching from the base. Leaves 29 mm long, 6.5 mm wide in males, 42 mm long, 8 mm wide in females, strap-like (oblanceolate-linear), greyish green, erect, the lower end clasping the stem. Male style hairless. Female inflorescences surrounded by shiny-brown, acute, overlapping involucral bracts. Ovary hairy towards the apex. Style more than 10 mm long.

Leucadendron pubescens GREYMAT ARID CONEBUSH
Erect, dense, 0.6-2.5 m tall shrubs, greyish green with a single stem. Leaves oblanceolate, narrowed to base, twisted, hairless to silvery haired. Involucral bracts acute, soft-haired. Perianth soft haired. Style less than 10 mm long. Cones covered with silvery hairs, 32-40 mm diam., globose.

Leucadendron bonum GIDEONSKOP ARID CONEBUSH
Leaves spathulate, silvery soft-haired. Involucral bracts numerous, acute, with long hairs, open in sunny weather, closed when rainy.

Leucadendron arcuatum RED-EDGED CONEBUSH
Numerous stems arising from a rootstock. Leaves variable, obovate to spathulate, margins red and thickened. Male inflorescences globose. Involucral bracts narrowly triangular, hairless.

Section VILLOSA: The Sandveld Conebushes

The Sandveld Conebushes are characterized by their mottled, densely to sparsely shaggy hairy, nutlike fruit with a pointed apex. Two of the species (dubium and concavum) have fruit much larger than the others and almost hairless. It is helpful to subdivide this group into several dichotomies:

• Fruit shed within two months of flowering (6 spp: brunioides, concavum, coriaceum, dubium, stellare, thymifolium) or
• Fruit retained on the plant (4 spp: cinereum, galpini, levisanus, linifolium).
• Leaves imbricate (clasping the stem and overlapping like tiles on a roof) (5 spp: concavum, dubium, levisanus, stellare, thymifolium) or
• Spreading away from the stem (5 spp: brunioides, cinereum, coriaceum, galpini, linifolium)

Leaf shape and the presence of a rootstock are the most important features used in distinguishing species of sandveld conebushes.

Leucadendron coriaceum ROSETTE CONEBUSH
Thin stems arising from a rootstock. Leaves small, obovate and hairless. Male style with 2 mm long hairs on lower half.

Leucadendron dubium CEDERBERG CONEBUSH VUURSLAANBOS
Single basal stem, and many slender branches. Leaves elliptic, soft haired, closely overlapping and clasped to the stem, 5-9 mm long, 3.5 mm wide. Perianths densely haired. Cones densely haired, fruit released.

Leucadendron concavum PAKHUIS CONEBUSH
Single stem at base. Leaves obovate to circular, soft haired, 9 mm long, 5 mm wide in male, 12 mm long, 7.5 mm wide in females. Cones densely haired.

Leucadendron brunioides brunioides FOETED CONEBUSH
With many thin stems arising from a rootstock. Leaves linear, 23 mm long, 1-2 mm wide, hairless. Male floral bract oblanceolate, shaggy haired.

Leucadendron brunioides flumenlupinum GRAAFWATER FOETED CONEBUSH
With many thin stems arising from a rootstock. Leaves linear to oblong, 23 mm long, 2-4 mm wide in female plants. Male floral bract oblanceolate, shaggy haired.

Leucadendron stellare STAR CONEBUSH
With many stems arising from a rootstock. Leaves, linear-lanceolate with a narrowed tip, sessile, tightly clasping the stem. Fruit 4.5 mm long, 4 mm broad.

Leucadendron thymifolium MALMESBURY CONEBUSH
A few branches arising from a basal stem. Leaves elliptic to oblong. Hypogynous scales present. Cones 18 mm diam., with slightly hairy bracts. Fruit 5.5 mm long, 6 mm broad.

Leucadendron levisanus CAPEFLATS CONEBUSH
Single main stem. Leaves oblanceolate to spathulate, 10 mm long, 4 mm wide. Cones serotinous.

Leucadendron linifolium LINE-LEAF CONEBUSH KRAALTOLBOS
Single main stem. Leaves variable, linear-oblanceolate, twisted. Male inflorescences stalked. Cones serotinous, 13-24 mm diam.

Leucadendron galpinii GLABRESCENT CONEBUSH
A tall, greyish shrub reaching 2-3 m in height. Leaves linear oblanceolate, tapered to base, twisted. Male inflorescences with a 7 mm long stalk. Cones serotinous, 32 mm long, 22 mm diam.

Leucadendron cinereum SCRAGGLY CONEBUSH VAALKNOPBOS
Single main stem. Leaves linear oblanceolate, narrowed to base, with a red apical callus, 27 mm long, 5.5 mm wide in males, 47 mm long, 55 mm wide in females, slightly twisted. Male inflorescences sessile. Perianth segments imbricate.

Section CARINATA: The Ridge-seed Conebushes

The Carinate Conebushes have fruit which are elongated, grey, smooth nuts, with a double ridge down the adaxial (upper) surface, giving the fruit a double keeled appearance. The female perianth segments are free and do not form a tube. The male perianth segments form a tube which is fused to the lower part of the style. Because of this there are no hypogynous scales. Both species in this section release their fruit when ripe.

Leucadendron sericeum WAABOOMS CONEBUSH
Leaves narrowly oblanceolate, soft haired but not silvery, up to 9 mm long, 2 mm wide in the male, females up to 15 mm long, 2.5 mm wide. Involucral leaves same as leaves.

Leucadendron nitidum BOKKEVELD CONEBUSH
Leaves narrowly oblanceolate, densely covered with fine hairs, silvery, 9-15 mm long, 2-4 mm wide in males, 11-20 mm long, 2-5 mm wide in females. Involucral leaves twice as large as stem leaves, yellow, crowded.

Section UNIFLORA: The Pauciflor Conebushes

The section Uniflora is very closely related to section Carinata, especially in the male and female floral structure. However, Pauciflor Conebushes have fruits which are sparsely, short-haired nuts. The female inflorescences bear only one to three flowers at their tips. The female perianth segments are free and do not form a tube. The male inflorescences contain less than 12 terminal flowers. The male perianth segments form a tube which is fused to the lower part of the style. Because of this there are no hypogynous scales and both species are apparently wind pollinated. Both species release their fruit when ripe.

Leucadendron olens YELLOW CONEBUSH
Leaves linear-lanceolate. Female inflorescences bearing 2-3 yellow florets, with large bifid stigmas.

Leucadendron ericifolium ERICOID CONEBUSH HEIDEBLAARGEELBOS
Leaves dimorphic, basal leaves ericoid, upper leaves linear. Male inflorescences very numerous. Female inflorescences bearing a single flower.

Section ALIENA: The Kouga Conebushes

The Kouga Conebushes consist of two species with a sprawling growth habit. Both possess a obovoid, shortly haired nut, which is released when ripe, and are further characterized by shedding the style from the fruit. Both are largely confined to the Uniondale Division.

The section Aliena is closely related to section Leucadendron, as is apparent by the fused female perianth.

Leucadendron singulare KAMMANASIE CONEBUSH
Leaves oblanceolate-linear, narrowing towards the base, with short-dense hairs, silvery.

Leucadendron sorocephaloides WOOLLY CONEBUSH
Leaves ericoid, thick, fleshy. Involucral bracts numerous, lanceolate, acute, brown, conspicuous. Male inflorescences woolly, cream-white. Female inflorescences with woolly cream florets at apex.

Section NERVOSA: The Jonaskop Silver-conebush

The Jonaskop Silver-conebush has a small acutely pointed nutlet, similar to that of the section Leucadendron, but characterized by being fringed with hairs 8 mm long. The female perianth segments are free and hairless, and the style persists in the fruit. This species is most closely related to the silver conebushes of the section Leucadendron.

Leucadendron nervosum JONASKOP CONEBUSH
Branches hairy. Leaves elliptic, leathery. Involucral leaves yellow, smaller towards centre, clothed with long white shaggy hairs. Cones conical, densely soft-haired.

Section LEUCADENDRON: The Silver Conebushes

The Silver Conebushes have fruit consisting of a soft haired nut, with the plumed female perianth segments are fused into a tube which cannot slip off the persistent style and thus forms a parachute for seed dispersal. Fruit are stored for several years in the cones. Leaf morphology is used to distinguish species.

Leucadendron dregei DISH CONEBUSH
Sprawling shrub. Leaves linear oblanceolate, hairless, thick and fleshy. Perianth segments surrounding fruit free, fused only at base.

Leucadendron album LINEAR-LEAF CONETREE
Leaves linear to linear-oblanceolate with a short silvery hairs. The posterior perianth segment has a fleshy base. Perianth segments with short plumes, fused surrounding the fruit, and slide up and down the persistent style.

Leucadendron rubrum SPINNING-TOP CONETREE TOLLETJIESBOS
Leaves oblanceolate, soft-haired aging to hairless, green. Male inflorescences narrow, without involucres. Female inflorescences ovoid with a pointed tip from which the florets' stigmas protrude in a tuft. Fruit with persistent style along which the fused perianth segments with long plumes slide.

Leucadendron argenteum SILVER CONETREE WITTEBOOM
Trees up to 10 m tall. Leaves silver, margins fringed with long hairs. Fruit sparsely long haired, with persistent style along which the fused perianth segments, with long plumes, slide.

Section CUNEATA: The Fusebract Conebushes

The Connate-bract Conebushes are distinguished by their pointed, mottled, shortly and sparsely haired nutlets, by their connate (fused) perianth segments and by their cone bracts tightly pressed together. All three species are found on level ground in the southwestern coastal plain, and shed their fruits annually. Two of the species (corymbosum, laxum) are characterized by a `corymbose' growth habit in which a cluster of short stems is produced at the base of the plant, forming a low round bush, from which slender, long, erect stems emerge.

Leucadendron verticillatum KLAPMUTS CONEBUSH
Leaves narrowly oblanceolate, twisted, densely silvery haired, with a small brown mucron. Perianth segments tightly pressed together.

Leucadendron corymbosum SWARTVELD CONEBUSH
Erect stems and many short branchlets clustered at the base. Leaves needle-like. Male inflorescences 12 mm diam., solitary at the ends of numerous short branches giving corymb appearance. Floral bract 1.5 mm long with 1 mm long bristles. Female inflorescences florets somewhat hidden by curling, spoon-shaped floral bracts.

Leucadendron laxum BREDASDORP CONEBUSH VLEIROSIE
Slender erect stems and many short branchlets clustered at the base. Leaves needle-like. Male inflorescences 6 mm long, 9 mm diam.Floral bract 1.5 mm long with bristles shorter than 1 mm. Cones with points of tightly pressed bracts elevated.

Section NUCIFERA: The Sun Conebushes

The Sun Conebushes are characterized by having large, dark brown, biconvex, hairless nuts. They thus resemble the section Membranacea. The perianth segments in the mature female florets are fused into a tube, narrow below and spreading, with hairs on the upper segments. They do not fall apart into separate segments (as in section Membranacea) All species release their fruit when ripe.

The 16 species are classified on the morphology of the male floret's stigma and the male floret bracts:

Key

1. Abortive stigma in male floret subcylindrical, elongate-club-shaped goto 2

1' Abortive stigma in male floret truncate with a bifid cleft and a pseudostigmatic surface STIGMATIC SUNCONEBUSHES

2. Male floral bract hairy NORTHWESTERN SUNCONEBUSHES

2' Male floral bract hairless SOUTHWESTERN SUNCONEBUSHES

The Stigmatic Sunconebushes comprise 7 species, which can readily be distinguished by stigma of the male flower being truncate and bifid, with a pseudostigmatic pollen presenter. The Stigmatic Sunconebushes occur from Worcester eastwards to Port Elizabeth, and thus comprise the eastern component of the Sun Conebushes. Leucadendron cordatum, L. orientale, L. pubibracteolarum and L. tradouwense may perhaps be considered as distinct forms of L. tinctum, but they appear to differ in sufficient characters to warrant specific status. The formation of the inflorescence bud and the development of the involucral bracts are useful characters in distinguishing between the species.

The Northwestern Sunconebushes comprise 5 species which are characterized by having hairy male floral bracts with subcylindrical male stigmas. The Northwestern Sunconebushes are all centred in the Cederberg and associated Hex River, Gifberg and Nieuwoudville mountains in the north-west of the fynbos. The species in this group are very variable, and are best distinguished by a combination of leaf shape and size, the shape of the involucral bracts and the colour of the flowers.

The Southwestern Sunconebushes comprise 4 species which are characterized by having a subcylindrical pollen presenter and hairless male floral bracts. The Southwestern Sunconebushes are distributed in an arc from the Witteberg southern Cederberg, DuToitskloof and Hottentot Holland mountains, The species in this group are most easily distinguished by plant habit, leaf shape and size and the involucral bracts.

Section NUCIFERA: The Stigmatic Sunconebushes

Leucadendron barkerae SWARTBERG SUNCONEBUSH
Leaves elliptic, sub-glaucous, finely haired, occasionally toothed. Male floral bracts linear with lateral hairs tightly clasping inflorescence base. Female inflorescences bracts overlapping not recurved.

Leucadendron burchellii RIVIERSONDEREND SUNCONEBUSH Leaves elliptic, hairless, dark green. Involucral leaves closely clasping the inflorescence base and overlapping. Male inflorescences flowers spreading widely when open. Male perianth slightly hairy. Female inflorescences with overlapping recurved involucral bracts.

Leucadendron tradouwense TRADOUW SUNCONEBUSH Branches hairless. Leaves erect. Male floral bract hairless. Female inflorescences bracts recurved, not oily. Cones globose or ellipsoid, 1/3-1.6 times as long as broad, Cone bracts broad, obtuse, hairless above, hairy below, not oily. .

Leucadendron orientale VANSTADENS SUNCONEBUSH
Leaves lanceolate to oblanceolate, hairless, erect, grass green. Involucral leaves crowded. Male floral bract with few hairs. Involucral bracts numerous (about 94), oblanceolate, recurved.

Leucadendron pubibracteolatum PURPLE-LEAF SUNCONEBUSH Leaves oblanceolate-elliptic, hairless, deflexed. Male floral bract soft-haired. Female inflorescences involucral bracts (about 89) reddish, recurved, not shiny or oily. Cones bracts recurved. Fruit with ridged perimeter.

Leucadendron tinctum SPICY SUNCONEBUSH
Bushy, spreading shrubs. Branches hairy. Leaves oblong-lanceolate, hairless, glaucous, deflexed. Male floral bract hairless. Female inflorescences strong spicy odour. Cones nearly as broad as long, bracts quadrangular, not recurved, hairless and oily above. Fruit with rounded perimeter.

Leucadendron cordatum DROOPY SUNCONEBUSH BERGKATJIEPIERING Erect or sprawling over rocks. Leaves elliptic or oblong. Involucral leaves larger, exceeding the head in length. Inflorescences pendulous on sharply down-curved branch tips. .

Section NUCIFERA: The Northwestern Sunconebushes

Leucadendron sheilae LOKENBERG SUNCONEBUSH
A small shrub. Branches hairless. Leaves narrowly oblanceolate. Male inflorescences small, greenish, foetid odour. Female inflorescences small, globose, green florets. Floral bract hairy below, ovate obtuse. Involucral bracts hairless, varnished, reddish brown. .

Leucadendron meyerianum VANRHYNSDORP SUNCONEBUSH
Many slender branches. Leaves linear. Male inflorescences small. Female inflorescences small.

Leucadendron glaberrimum OILY SUNCONEBUSH
Erect or spreading shrubs. Leaves oblanceolate, hairless to soft-haired, leathery. Involucre inconspicuous. Male floral bract 4 mm long, linear, oily, hairy below, hairless above. Male flowers yellow. Involucral bracts with a fringe of hairs.

Leucadendron loranthifolium GREENFLOWERED SUNCONEBUSH
Shrub to 2 m tall, single main stem. Leaves glaucous, margin cartilaginous. Male inflorescences green, depressed globose. Involucral bracts ovate. Floral bract linear, densely crispy-haired. Female inflorescences green.

Leucadendron roodii GIFBERG SUNCONEBUSH
Leaves elliptic, hairless. Involucral bracts red, pouch-like.

Section NUCIFERA: The Southwestern Sunconebushes

Leucadendron cadens WITTEBERG CONEBUSH
Prostrate shrub. Leaves narrowly oblanceolate, subsecund. Involucral bract circular in shape. Floral bract 11 mm long, hairless. Perianth with a 9.5 mm tube. Hypogynous scales vestigial. Cones 27 mm long.

Leucadendron gydoense GYDO SUNCONEBUSH
Stems slender, hairless. Leaves narrowly oblanceolate. Involucral leaves similar, not crowded. Floral bract hairless.

Leucadendron sessile SUN CONEBUSH
Leaves narrowly elliptical. Involucral leaves similar, crowded. Brown or purplish brown involucral bracts, not recurved, visible.

Leucadendron daphnoides DUTOITSKLOOF SUNCONEBUSH
Leaves hairless, lanceolate or elliptic. Involucral leaves closely overlapping around the lower portion of the head, hiding the involucral bracts. Male floral bracts with spoon-shaped apices. Female perianth 21 mm long.

Section VENTRICOSA: The Crown Conebushes

The Crown Conebushes are characterized by having a hairless nut, ventricose on one side and keeled on the other, with a narrowly ridged perimeter. The mature female perianth segments are fused together, and exceed 7 mm in length. The cones are unique in having bracts quite free from one another and somewhat leaf-like, and an elongated receptacle. All three species release their fruit after a lengthy (5 month) ripening period. All grow on plains between the mountains.

A possible sixth species, Wynberg Conebush - Leucadendron grandiflorum, known only from a drawing, grew on Wynberg Hill. Distinguished by its pale green, smaller (12-14 mm broad) twisted leaves, a disagreeable odour and smaller male inflorescence, bracts and flowers, it might have been a variant of L. globosum.

Leucadendron globosum GRABOUW CONEBUSH
Stems hairy. Leaves oval, dark green. Female inflorescences with closely overlapping recurved involucral bracts.

Leucadendron chamelaea WITSENBERG CONEBUSH
Erect shrubs to 2.3 m tall, branching immediately from a single main stem. Leaves linear-oblanceolate, 40 mm long, 6 mm wide in males, up to 45 mm long, 6 mm wide in females. Male inflorescences 20 mm diam. Floral bracts in 5 spirals. Cones with numerous obovate, non-overlapping green bracts.

Leucadendron elimense elimense ELIM CONEBUSH
Leaves elliptic, 13-49 mm long, 5-19 mm wide in male, 14-57 mm long, 7-21 mm wide in female, closely spaced. Female inflorescences sides closely overlapped by recurved involucral bracts.

Leucadendron elimense salteri CALEDON CONEBUSH
Leaves 55-67 mm long, 15-25 mm wide in males, 61-78 mm long, 19-26 mm wide in females, elliptic, widely spaced

Leucadendron elimense vyeboomense VYEBOOM CONEBUSH
Leaves 43 mm long, 11 mm wide, similar in both sexes, lanceolate

Section TRIGONA: The Deltaseed Conebushes

The Trigosperm Conebushes have fruit less than 5 mm broad with narrow wings or a sub-triangular cross section. The mature female perianth is always soft-haired and the lobes abut one another without overlapping. With the exception of the Capeflats Conebush L. floridum, the only exception in the subgenus, all species retain their fruit in the cones.

A subgroup comprises the Stream Conebushes (conicum, macowanii, salicifolium) all of which have broad cone bracts, an exposed stigma in the female bud, female perianth forming a tube, and grow in damp places.

Note that some forms of L. xanthoconus (section Alata) have fruit typical of this section.

Leucadendron floridum FLATS CONEBUSH TOLBOS
Branches hairy. Leaves silky-haired, mucronate apex. Perianth fused, tubular.

Leucadendron uliginosum uliginosum OUTENIQUA CONEBUSH
Hairless branches. Leaves linear oblong, flat silvery-silky haired. Male floral bud hairless at tip. Cones bracts silvery-haired. Fruit narrowly winged.

Leucadendron uliginosum glabratum TSITSIKAMMA CONEBUSH
Leaves green, soft-haired when young aging to hairless, 20 mm long, 4 mm wide in males, 26 mm long, 4 mm wide in females

Leucadendron loeriense LOERIE CONEBUSH
Shrubs. Leaves with velvety-erect hairs. Fruit broadly winged.

Leucadendron radiatum LANGEBERG CONEBUSH
Shrubs 0.6 m tall, dense. Leaves sharply twisted, 23 mm long, 5 mm wide.

Leucadendron rourkei UNIONDALE CONEBUSH
Leaves hairless, up to 18 mm long, 3.5 mm wide in males, up to 24 mm long, 45 mm wide in females, oblanceolate. Involucral leaves inconspicuous. Perianth 3.5 mm long.

Leucadendron conicum GARDEN-ROUTE STREAMCONEBUSH
Shrubs or small trees. Leaves narrowly oblanceolate, dark green. Male inflorescences soft-haired. Involucral bracts conspicuous, soft-haired outside.

Leucadendron pondoense PONDOLAND STREAMCONEBUSH
Leaves 35-60 mm long, 2.5-4 mm wide in males, 50-70 mm long, 3-4 mm wide in females. Male inflorescences densely silver-haired. Involucral bracts inconspicuous, linear, 8 in number, soft-haired outside. Fruit shed after 12 months.

Leucadendron salicifolium COMMON STREAMCONEBUSH
Leaves linear, hairless.

Leucadendron macowanii ACACIA-LEAF STREAMCONEBUSH
Leaves narrowly oblanceolate, 53 mm long, 7 mm wide in males, up to 77 mm long, 9 mm wide in females. Involucral leaves absent. Involucral bracts 14 in number, inconspicuous.

Section BRUNNEOBRACTEOTA: The Oilbract Conebush

The Oilbract Conebush is characterized by having a 7-8 mm broad fruit, with the perimeter sharply ridged, with a pointed apex, and a rough median ridge. It also has dark brown involucral bracts, covered with an oily brown, boot polish-like resin, especially in the bud stage.

Leucadendron microcephalum OILBRACT CONEBUSH
Involucral bracts with a dark brown oily stain. Fruit with a double ridge on one side.

Section ALATA: The Clay and Sunshine Conebushes

The Clay and Sunshine Conebushes are characterized by the winged fruits and the female perianth tube with the hairless, free segments overlapping one another. The leaves are never needle-like.

This group of 22 taxa is the largest in the genus and can be divided into two groups:

• The Clay Conebushes with rough (scabrous) leaves, and
• The Sunshine Conebushes with smooth leaves.

The Clay conebushes are the only species in the section Alata to grow on heavy clay soils. There are three species of clay conebushes, distinguished by their leaf sizes and growth habit.

The Sunshine conebushes are typically found on sandy soils, with one on calcareous soils (meridianum) and one on clay gravels (cryptocephalum), although L. salignum is catholic in its requirements. This is a closely knit group, which we will subdivide into three groups purely to aid with their identification. Stem leaves do not include those immediately surrounding the inflorescences.

Key

1. Species resprouting from underground rootstocks
flexuosum, salignum and spissifolium

The latter two species are both widespread, the last with several distinct subspecies. Both of these species are variable and best distinguished from each other by the hairless exposed bracts of the cones of spissifolium and the short-haired bracts of salignum.

1' Species with a single basal stem. Killed by fire. goto 2

2. Longest female stem leaves less than 60 mm long:

diemontianum, foedum, procerum, discolor,& meridianum.

With the exception of L. meridianum, all these species occur in the northwestern and western areas of the Fynbos (west of a line from Stellenbosch to the Hex River, excluding the Cape Peninsula). L. flexuosum fits well in this group. Floret colour and leaf hairyness and size are useful features in distinguishing the species in this group.

2' Longest female stem leaves longer than 60 mm:

cryptocephalum, xanthoconus, eucalyptifolium, coniferum, gandogeri, laureolum & strobilinum.

All these species occur on the eastern portion of the Fynbos, mainly the southern mountains (including the Cape Peninsula) and the coastal flats, with only L. laureolum overlapping with the distribution of the former group. Leaf shape and size and cone size are useful features for distinguishing species in this group.

Leucadendron lanigerum SHALE CONEBUSH Persistent rootstock (single-stemmed at Wolseley). Leaves oblanceolate-linear, hairless, scabrous, slightly twisted.

Leucadendron modestum ROUGH-LEAF CONEBUSH
Single stemmed at base. Leaves narrowly oblanceolate.

Leucadendron stelligerum AGULHAS CONEBUSH
Leaves 23 mm long, 7 mm wide in males, 28 mm long, 8 mm wide in females, not twisted. Involucral leaves partly concealing the head in a star-like cup

Leucadendron xanthoconus SICKLELEAF CONEBUS
Leaves linear oblong, twisted at base, initially silver haired aging to hairless, somewhat sickle-shaped. Cone bracts bilobed, hairy.

Leucadendron diemontianum VISGAT SUNSHINECONEBUSH
Shrubs developing from a single main stem with a cluster of branches at base. Leaves elliptic-linear, clasping stem. Male inflorescences 12 mm diam. Cones 13-14 mm long, 13-18 mm diam.

Leucadendron flexuosum WORCESTER SUNSHINECONEBUSH
Shrubs slender, spindly, with a rootstock. Leaves oblanceolate-linear, hairless. Male inflorescences 14 mm diam. Cones 21 mm long. Flowers: Apr-May

Leucadendron salignum COMMON SUNSHINEBUSH KNOPPIESGEELBOS
Multistemmed shrub from a persistent rootstock. Leaves oblanceolate-linear, hairless, not shining. Floral bracts hairy.

Leucadendron foedum HOPEFIELD SUNSHINECONEBUSH
Shrubs to 2.5 m tall, single stemmed, males rounded. Involucral leaves oblanceolate, pale green. Male florets yellow. Floral bracts almost hairless. Involucral bracts variable, ovate, soft-haired and fringed with hairs.

Leucadendron procerum IVORY SUNSHINECONEBUSH LANGBEENTJIE
Shrubs to 3 m tall, erect. Leaves oblanceolate. Involucral leaves pale green to ivory, crowded. Male florets red. Cone bracts ovate, indented (dimpled) in middle.

Leucadendron discolor PIKETBERG SUNSHINECONEBUSH ROOITOLBOS
Leaves broadly obovate. Male inflorescences 38 mm long, including 10 mm long stalk, 25 mm diam., reddened in bud. Floral bracts hairless. Female inflorescences 26 mm long, 16 mm diam.

Leucadendron eucalyptifolium GUM-LEAF SUNSHINECONEBUSH GROOTGEELBOS
Leaves linear-lanceolate, soft-haired when young aging to hairless, acute apex with a small blunt mucron. Male inflorescences 24 mm long, 16 mm diam.

Leucadendron coniferum MUCRON-LEAF SUNSHINECONEBUSH DUINEGEELBOS
Leaves narrowly oblanceolate, sharp mucronate, hairless when mature. Male stigma hoof-shaped, constricted below tip.

Leucadendron meridianum LIMESTONE SUNSHINECONEBUSH
Densely branched. Leaves silvery haired to hairless. Perianth hairless. Abortive male stigma club-shaped. Cones silvery haired, 28 mm long, 22 mm diam.

Leucadendron gandogeri BROAD-LEAF SUNSHINECONEBUSH GEELBOS
Leaves oblanceolate or elliptic, hairless, large. Involucral leaves crowded, not concealing heads. Involucral bracts about 20, hairless, fringed with hairs, brown. Cones hairless.

Leucadendron cryptocephalum CONCEALED SUNSHINECONEBUSH
Shrubs to 1 m tall. Leaves narrowly lanceolate, pale green. Involucral leaves crowded, almost concealing male head and completely enclosing female head.

Leucadendron laureolum LAURAL-LEAF SUNSHINECONEBUSH GEELBOS
Leaves oblong. Involucral leaves almost concealing head in males, yellow-green and totally concealing heads in females. Involucral bracts brown, hairless. Cones with 8 spiral shallow grooves, 38-46 mm long, 32-37 mm diam. Fruit winged on sides and above.

Leucadendron strobilinum PENINSULA SUNSHINECONEBUSH
Stems stout, purplish, soft-haired. Leaves elliptic, dark green, hairless, with hairy margins especially when young. Involucral leaves cream coloured. Male inflorescences depressed-globose. Cone bracts numerous.

Leucadendron spissifolium spissifolium COMMON SPEAR-LEAF CONEBUSH GEELBOS
Many stems arising from a persistent rootstock. Leaves oblanceolate or linear-oblanceolate, hairless, slightly twisted. Involucral bracts lanceolate, brown, margin hairy.

Leucadendron spissifolium fragrans FRAGRANT SPEAR-LEAF CONEBUSH
Leaves 30-45 mm long, 20-23 mm wide in males, 40-50 mm long, 20-25 mm wide in females, linear, channelled, mucronate, twisted, hairless. Involucral bracts lanceolate acute. Cone bracts hairless.

Leucadendron spissifolium phillipsii KAREEDOUW SPEAR-LEAF CONEBUSH
Leaves up to 50 mm long, 5 mm wide in males, up to 57 mm long, 6.3 mm wide in females, linear to linear-oblanceolate, mucronate, twisted, soft-haired when young becoming hairless. Involucral bracts ovate. Cone bracts hairy across middle.

Leucadendron spissifolium natalensis NATAL SPEAR-LEAF CONEBUSH
Leaves oblanceolate, pointed mucron, twisted below, hairless, 46 mm long, 8 mm wide. Involucral leaves conspicuous, acute apex, bract-like. Cone bracts hairy across middle.

Leucadendron spissifolium oribinum ORIBI SPEAR-LEAF CONEBUSH
Leaves up to 31 mm long, 3 mm wide in males, up to 45 mm long, 4 mm wide in females, linear-oblanceolate, pale apical mucron, sickle-shaped, twisted, hairless. Male inflorescence without involucre.

Section COMPRESSA: The Needle-leaf Conebushes

The Needle-leaf Conebushes all have seedlings with smooth and needle-like (terete) leaves. These may remain terete in the adults or become flattened so that more than one leaf shape (polymorphism) occurs on a plant. The floral bracts of the female head may be partly or completely pressed together, and the fruits are compressed with wings. Identification of the 6 species is best accomplished by comparing leaf shape, sizes of male heads and cones, and the sizes of the retained fruit.

Leucadendron teretifolium NEEDLE-LEAF CONEBUSH WATERBOSSIE
Leaves needle-like, small. Male inflorescences numerous, 14 mm long including stalk. Involucral bracts long acute.

Leucadendron spirale WOLSELEY CONEBUSH
Leaves ericoid, subacute, 4-9 mm long. Male floral bracts rudimentary. Cone bracts bilobed, hairless, overlapping below. Only collected four times (1801,1819, 1820's, 1933)

Leucadendron osbornei KAROO CONEBUSH
Leaves needle-like, short (Intermediate between teretifolium and nobile). Male inflorescences with florets in a short spike. Female inflorescences 28 mm long, 12 mm diam., elongated-ovoid.

Leucadendron nobile KAROO CONEBUSH NAALDBLAARTOLBOS
Leaves needle-like, long. Male inflorescences with florets in a long spike. Female inflorescences 28 mm long, 12 mm diam., elongated-ovoid.

Leucadendron muirii SILVERBALL CONEBUSH KRUIPHOUT
Mature leaves obovate-spathulate, thick and fleshy, rounded apex. Involucral bracts very acute. Cones ashy-grey.

Leucadendron comosum RIDGECONE CONEBUSH
Leaves usually dimorphic. Lower leaves needle-like. Upper leaves broadened. Florets dark red. Male floral bud club-shaped. Cones abutting towards base, up to 100 mm long.

Leucadendron platyspermum PLATE-SEED CONEBUSH KRAALTOLBOS
Male inflorescences with a dense tuft of long white hairs apically which fall off as flowers open. Cones with double-edged bracts abutting in a single spiral. Fruit 15 mm long, 32 mm broad, compressed, flat.

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