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.
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
The subgenus Leucadendron contains a diversity of fruit types and is divided into two major groups:
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
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
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
Leucadendron pubescens GREYMAT
Leucadendron bonum GIDEONSKOP ARID CONEBUSH
Leucadendron arcuatum RED-EDGED CONEBUSH
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:
Leaf shape and the presence of a rootstock are the most important features used in distinguishing species of sandveld conebushes.
Leucadendron coriaceum ROSETTE CONEBUSH
Leucadendron dubium CEDERBERG CONEBUSH VUURSLAANBOS
Leucadendron concavum PAKHUIS
Leucadendron brunioides brunioides FOETED CONEBUSH
Leucadendron brunioides flumenlupinum GRAAFWATER FOETED
Leucadendron stellare STAR CONEBUSH
Leucadendron levisanus CAPEFLATS CONEBUSH
Leucadendron linifolium LINE-LEAF CONEBUSH KRAALTOLBOS
Leucadendron galpinii GLABRESCENT CONEBUSH
Leucadendron cinereum SCRAGGLY
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
Leucadendron nitidum BOKKEVELD CONEBUSH
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
Leucadendron ericifolium ERICOID CONEBUSH HEIDEBLAARGEELBOS
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
Leucadendron sorocephaloides WOOLLY 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
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
Leucadendron album LINEAR-LEAF CONETREE
Leucadendron rubrum SPINNING-TOP CONETREE TOLLETJIESBOS
Leucadendron argenteum SILVER CONETREE WITTEBOOM
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
Leucadendron corymbosum SWARTVELD CONEBUSH
Leucadendron laxum BREDASDORP CONEBUSH VLEIROSIE
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:
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
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
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
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
Leucadendron meyerianum VANRHYNSDORP SUNCONEBUSH
Leucadendron glaberrimum OILY SUNCONEBUSH
Leucadendron loranthifolium GREENFLOWERED SUNCONEBUSH
Leucadendron roodii GIFBERG SUNCONEBUSH
Section NUCIFERA: The Southwestern Sunconebushes
Leucadendron cadens WITTEBERG CONEBUSH
Leucadendron gydoense GYDO SUNCONEBUSH
Leucadendron sessile SUN CONEBUSH
Leucadendron daphnoides DUTOITSKLOOF SUNCONEBUSH
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
Leucadendron chamelaea WITSENBERG CONEBUSH
Leucadendron elimense elimense
Leucadendron elimense salteri CALEDON CONEBUSH
Leucadendron elimense vyeboomense VYEBOOM CONEBUSH
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
Leucadendron uliginosum uliginosum OUTENIQUA CONEBUSH
Leucadendron uliginosum glabratum TSITSIKAMMA CONEBUSH
Leucadendron loeriense LOERIE CONEBUSH
Leucadendron radiatum LANGEBERG CONEBUSH
Leucadendron rourkei UNIONDALE CONEBUSH
Leucadendron conicum GARDEN-ROUTE STREAMCONEBUSH
Leucadendron pondoense PONDOLAND STREAMCONEBUSH
Leucadendron salicifolium COMMON STREAMCONEBUSH
Leucadendron macowanii ACACIA-LEAF STREAMCONEBUSH
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
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 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.
1. Species resprouting from underground rootstocks
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
Leucadendron stelligerum AGULHAS
Leucadendron xanthoconus SICKLELEAF CONEBUS
Leucadendron diemontianum VISGAT
Leucadendron flexuosum WORCESTER
Leucadendron salignum COMMON SUNSHINEBUSH KNOPPIESGEELBOS
Leucadendron foedum HOPEFIELD
Leucadendron procerum IVORY SUNSHINECONEBUSH LANGBEENTJIE
Leucadendron discolor PIKETBERG
Leucadendron eucalyptifolium GUM-LEAF SUNSHINECONEBUSH
Leucadendron coniferum MUCRON-LEAF
Leucadendron meridianum LIMESTONE SUNSHINECONEBUSH
Leucadendron gandogeri BROAD-LEAF
Leucadendron laureolum LAURAL-LEAF SUNSHINECONEBUSH GEELBOS
Leucadendron strobilinum PENINSULA SUNSHINECONEBUSH
Leucadendron spissifolium spissifolium COMMON
SPEAR-LEAF CONEBUSH GEELBOS
Leucadendron spissifolium fragrans FRAGRANT SPEAR-LEAF
Leucadendron spissifolium phillipsii KAREEDOUW
Leucadendron spissifolium natalensis NATAL SPEAR-LEAF
Leucadendron spissifolium oribinum ORIBI SPEAR-LEAF
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
Leucadendron spirale WOLSELEY
Leucadendron osbornei KAROO CONEBUSH
Leucadendron nobile KAROO CONEBUSH NAALDBLAARTOLBOS
Leucadendron muirii SILVERBALL CONEBUSH KRUIPHOUT
Leucadendron comosum RIDGECONE CONEBUSH
Leucadendron platyspermum PLATE-SEED CONEBUSH KRAALTOLBOS