Leucospermum Conocarpodendron
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''Leucospermum conocarpodendron'' is the largest species of the genus Leucospermum, reaching almost tree-like proportions of high with a firm trunk that is covered in a thick layer of cork that protects it from most fires. It has greyish or green narrow or broad inverted egg-shaped leaves with three to ten teeth near the tip and large yellow flowerheads, with firm, bent, yellow styles that stick far beyond the rest of the flower and give the impression of a pincushion. It is commonly known as the tree pincushion in English or goudsboom in
Afrikaans Afrikaans (, ) is a West Germanic language that evolved in the Dutch Cape Colony from the Dutch vernacular of Holland proper (i.e., the Hollandic dialect) used by Dutch, French, and German settlers and their enslaved people. Afrikaans gra ...
. They naturally occur near
Cape Town Cape Town ( af, Kaapstad; , xh, iKapa) is one of South Africa's three capital cities, serving as the seat of the Parliament of South Africa. It is the legislative capital of the country, the oldest city in the country, and the second largest ...
, South Africa. Two subspecies are distinguished. ''L. conocarpodendron'' subsp. ''conocarpodendron'', that has greyish leaves because they have a covering of felty hairs. ''L. conocarpodendron'' subsp. ''viridum'', has green leaves that lack felty hairs. Flowers can be found between August and December.


Description

''L. conocarpodendron'' is an
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...
large shrub of high and in diameter with a rounded crown, which is rigid because of the thick branching at approximately right angles, and with a firm trunk of in diameter that is covered by thick greyish, reddish or blackish bark with a netting of fissures. The flowering branches are rigid, 1–1½ cm (0.4–0.6 in) thick and covered with a dense layer of white or greyish crinkly hairs or long soft silky hairs. The leaves lack a
leaf stalk In botany, the petiole () is the stalk that attaches the leaf blade to the stem, and is able to twist the leaf to face the sun. This gives a characteristic foliage arrangement to the plant. Outgrowths appearing on each side of the petiole in so ...
and stipules, ovate to lance-shaped with the greater width often beyond midlength 6–11½ cm (2.4–4.6 in) long and 2½–5 cm (1–2 in) wide, with a blunt or pointy tip, shallow or deeply incised with three to ten teeth. Dependent on the subspecies, the surface og=f the leaves is either hairless or has a dense covering of soft, greyish, convoluted hairs, occasionally with a fringe of fine hairs. The flower heads sit atop a
stalk Stalk or stalking may refer to: Behaviour * Stalk, the stealthy approach (phase) of a predator towards its prey * Stalking, an act of intrusive behaviour or unwanted attention towards a person * Deer stalking, the pursuit of deer for sport Biol ...
of about 1½ cm (0.6 in) long, are globe- to egg-shaped, and in diameter. They can be found individually or mostly with two or three together near the tip of the branch, regularly partly enclosed by leaves. The common base of the flowers of the same head is narrowly cone-shaped with a pointy tip, 2½–3 cm (1–1.2 in) long and 1–1½ cm (0.4–0.6 in) in diameter. The
bracts In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
that subtend the flower head are oval in shape with a point tip, 1¼–1½ cm (0.5–0.6 in) long and approximately wide, tightly overlapping, with a rubbery consistency and softly hairy. The
bracts In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
supporting the individual flowers enclose them at their base, have a suddenly pointed tip, and are about long, and about wide, rubbery in consistency, woolly at the base and less so near the top. The
perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepals when ...
is 3½–5 cm (1.4–2.0 in) long and yellow in colour. The lower 1 cm (0.4 in) is fused, cylindric, and hairless. The free parts of the four perianth claws curl back when the flower opens, those to the sides and facing the rim of the flower head densely set with long hairs. The one facing the center of the head minutely powdery or very shortly softly hairy. The perianth limbs are lance-shaped with a pointy tip, long, and have long hairs pressed to the surface, except for the one facing the center that is minutely powdery. The style is stout, 1–1½ mm (0.04–0.06 in) thick and 4½–5½ cm (1.8-2.2 in) long, at first bent towards the center of the flower head but getting more straight with age. It is topped by a slight thickening that is called the pollen presenter, which has a broad conical shape with a pointy tip, is long and about wide. Subtending the
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
are four lance-shaped scales with a pointy tip of about long. The subtribe Proteinae, to which the genus ''Leucospermum'' has been assigned, consistently has a basic chromosome number of twelve ( 2n=24).


Differences between the subspecies

The grey tree pincushion or vaalkreupelhout in
Afrikaans Afrikaans (, ) is a West Germanic language that evolved in the Dutch Cape Colony from the Dutch vernacular of Holland proper (i.e., the Hollandic dialect) used by Dutch, French, and German settlers and their enslaved people. Afrikaans gra ...
(subsp. ''conocarpodendron'') has felty hairy leaves due to a dense cover of fine crisped hairs, while the green tree pincushion or groenkreupelhout (subsp. ''viridum'') has green hairless adult leaves, sometimes with a fringe of hairs around the edge. At one location, on the east side of Little Lion's Head near Mount Rhodes, a hybrid swarm between both subspecies is found, where individual plants may have hairiness anywhere between that of both parents. Elsewhere, the populations are uniform and can easily be assigned to either of the subspecies.


Differences with related species

''L. conocarpodendron'' differs from its nearest relatives by its tree-like habit, the narrowly cone-shaped common base of the flower heads, the oval involucral bracts with a pointy tip, and the broad cone-shaped pollen presenter.


Taxonomy

The earliest known description of the species we now know as ''Leucospermum conocarpodendron'' was by Paul Hermann in ''Paradisus Batavus'', a book describing the plants of the
Hortus Botanicus Leiden The Hortus botanicus of Leiden is the oldest botanical garden of the Netherlands, and one of the oldest in the world. It is located in the southwestern part of the historical centre of the city, between the Academy building and the old Leiden Obs ...
(botanical garden of the Leyden university), that was published in 1689, three years after his death. He called it ''Salix conophora Africana'' (African cone-bearing willow), based on his observation of ''Leucospermum conocarpodendron'' on the lower slopes of the Table Mountain. In the following six decades, several other descriptions were published, such as by Leonard Plukenet,
James Petiver James Petiver (c. 1665 – c. 2 April 1718) was a London apothecary, a fellow of the Royal Society as well as London's informal Temple Coffee House Botany Club, famous for his specimen collections in which he traded and study of botany and entomo ...
,
John Ray John Ray FRS (29 November 1627 – 17 January 1705) was a Christian English naturalist widely regarded as one of the earliest of the English parson-naturalists. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after ...
and Herman Boerhaave. Names published before 1753, the year that was chosen as a starting point for the
binominal nomenclature In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
proposed by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
, are not valid however. The tree pincushion was first validly described in the first edition of ''
Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...
'' as ''Leucadendron conocarpodendron'' by Linnaeus in 1753. Johann Jacob Reichard in 1779 reassigned the species to ''Protea'', creating the
new combination ''Combinatio nova'', abbreviated ''comb. nov.'' (sometimes ''n. comb.''), is Latin for "new combination". It is used in taxonomic biology literature when a new name is introduced based on a pre-existing name. The term should not to be confused wi ...
''P. conocarpodendron''. In 1781,
Carl Peter Thunberg Carl Peter Thunberg, also known as Karl Peter von Thunberg, Carl Pehr Thunberg, or Carl Per Thunberg (11 November 1743 – 8 August 1828), was a Swedish naturalist and an "apostle" of Carl Linnaeus. After studying under Linnaeus at Uppsala Un ...
simplified the species name and created ''P. conocarpa'', but because he used the same
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * Ty ...
as Linnaeus, he should have used the unchanged name. Richard Anthony Salisbury created two
superfluous name ''Nomen illegitimum'' (Latin for illegitimate name) is a technical term, used mainly in botany. It is usually abbreviated as ''nom. illeg.'' Although the International Code of Nomenclature for algae, fungi, and plants uses Latin terms for other ki ...
s, ''Protea tortuosa'' in 1796 and ''Leucadendrum crassicaule'' in 1809. In his book ''
On the natural order of plants called Proteaceae ''On the natural order of plants called Proteaceae'', also published as "On the Proteaceae of Jussieu", was a paper written by Robert Brown on the taxonomy of the plant family Proteaceae. It was read to the Linnean Society of London in the first ...
'' that Robert Brown published in 1810, the species was reassigned to the new genus ''Leucospermum'', but he combined it with Brown's invalid simplified species name to ''Leucospermum conocarpum''. In 1874,
Heinrich Wilhelm Buek Heinrich may refer to: People * Heinrich (given name), a given name (including a list of people with the name) * Heinrich (surname), a surname (including a list of people with the name) *Hetty (given name), a given name (including a list of peo ...
made the correct combination ''Leucospermum conocarpodendron''. Another form was described by
Michael Gandoger Abbé Jean Michel Gandoger (10 May 1850 – 4 October 1926), was a French botanist and mycologist. He was born in Arnas, the son of a wealthy vineyard owner in the Beaujolais region. Although he took holy orders at the age of 26, he devoted his ...
in 1901, and he called it ''Leucospermum macowanii''. In 1970,
John Patrick Rourke John Patrick Rourke FMLS (born 26 March 1942, in Cape Town) is a South African botanist, who worked at the Kirstenbosch National Botanical Garden and became curator of the Compton Herbarium. He is a specialist in the flora of the Cape Floristi ...
proposed to distinguish between the typical subspecies (''L. conocarpodendron'' subsp. ''conocarpodendron'') and ''L. conocarpodendron'' subsp. ''viridum''. ''L. conocarpodendron'' is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the section '' conocarpodendron''. The species and subspecies name ''conocarpodendron'' means "tree bearing cone-shaped fruits". The subspecies name ''viridum'' means "green" and is a reference to the leaves' colour. It was called kreupelhout in Dutch (cripple wood) already before 1680, a reference to the twisted branches that together give the tree a "crippled" appearance.


Distribution, habitat and ecology

''L. conocarpodendron'' subsp. ''conocarpodendron'' is an
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
of the
Cape Peninsula The Cape Peninsula ( af, Kaapse Skiereiland) is a generally mountainous peninsula that juts out into the Atlantic Ocean at the south-western extremity of the African continent. At the southern end of the peninsula are Cape Point and the Cape of ...
where it is limited to the eastern slopes of Devils Peak, the northern and western slopes of Table Mountain and the Black Table, to
Llandudno Llandudno (, ) is a seaside resort, town and community in Conwy County Borough, Wales, located on the Creuddyn peninsula, which protrudes into the Irish Sea. In the 2011 UK census, the community – which includes Gogarth, Penrhyn Bay, Craigsi ...
. It grows mainly on heavy clay derived from the weathering of
Cape Granite A cape is a clothing accessory or a sleeveless outer garment which drapes the wearer's back, arms, and chest, and connects at the neck. History Capes were common in medieval Europe, especially when combined with a hood in the chaperon. The ...
but also weathered Table Mountain Sandstone. It prefers north and west exposures that are well drained. ''L. conocarpodendron'' subsp. ''viridum'' has a much wider distribution that borders on that of the typical subspecies. It occupies the remainder of the Cape Peninsula from
Kirstenbosch Kirstenbosch is an important botanical garden nestled at the eastern foot of Table Mountain in Cape Town. The garden is one of 10 National Botanical Gardens covering five of South Africa's six different biomes and administered by the South A ...
to the
Cape of Good Hope The Cape of Good Hope ( af, Kaap die Goeie Hoop ) ;''Kaap'' in isolation: pt, Cabo da Boa Esperança is a rocky headland on the Atlantic coast of the Cape Peninsula in South Africa. A common misconception is that the Cape of Good Hope is t ...
. In addition, it occurs from the upper Berg River Valley, via Pringle Bay and Hermanus to
Stanford Stanford University, officially Leland Stanford Junior University, is a private research university in Stanford, California. The campus occupies , among the largest in the United States, and enrolls over 17,000 students. Stanford is considere ...
. Isolated populations can also be found at
Helderberg Helderberg refers to a planning district of the City of Cape Town metropolitan municipality, the mountain after which it is named, a wine-producing area in the Western Cape province of South Africa, or a small census area in Somerset West. Or ...
,
Simonsberg Simonsberg ( en, Simon's Mountain) is part of the Cape Fold Belt in the Western Cape province of South Africa. It is located between the towns of Stellenbosch, Paarl and Franschhoek, forming a prominent 1399 m high mountain, as it is detached f ...
, and Kogelberg near Durbanville. It occurs on such different soil types as Malmesbury gravel, sand from weathered Table Mountain Sandstone, dune sands, permanently soggy peat, and sometimes on the heavy clay that remains if Cape Granite is decomposed. It mostly grows between
sea level Mean sea level (MSL, often shortened to sea level) is an average surface level of one or more among Earth's coastal bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical datuma standardised g ...
and , sometimes 300 m altitude. At some locations this pincushion is dominant and develops dense stands. Both subspecies have some resistance for the
wildfire A wildfire, forest fire, bushfire, wildland fire or rural fire is an unplanned, uncontrolled and unpredictable fire in an area of Combustibility and flammability, combustible vegetation. Depending on the type of vegetation present, a wildfire ...
s that occur in the
fynbos Fynbos (; meaning fine plants) is a small belt of natural shrubland or heathland vegetation located in the Western Cape and Eastern Cape provinces of South Africa. This area is predominantly coastal and mountainous, with a Mediterranean clim ...
every one or two decades, because the trunk is covered by a thick bark. After the fire has burnt away the soft parts, regrowth takes place from the tip of the higher branches. Repeated mild burning results in an umbrella shaped growth habit. Seed dispersal and survival greatly depends on the symbiosis of many Proteaceae with native ant species, in particular ''
Anoplolepis steingroeveri ''Anoplolepis steingroeveri'' is a species of ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous ...
'' and '' Pheidole capensis'', that carry the fruits to their underground nests, where the
elaiosome Elaiosomes ( grc, ἔλαιον ''élaion'' "oil" + ''sóma'' "body") are fleshy structures that are attached to the seeds of many plant species. The elaiosome is rich in lipids and proteins, and may be variously shaped. Many plants have elaio ...
is eaten, leaving a slick and hard seed underground, safe from consumption by rodents and birds and overhead fires. The seeds would germinate after a fire, due to the larger temperature variations after the overhead vegetation has vanished and chemicals from the charcoal seep with the winter rains and soak the seeds. Seed dispersal is only limited. In an experiment, seeds on average were moved about 2 m (6½ ft) and at most about 10 m. The absence or presence of the elaiosome did not effect the germination rate, but a field trial showed that seeds without elaiosome almost never survive a fire, whereas those with elaiosome all germinated, implying that the burial of the seed by the ants is essential.


Conservation

''Leucospermum conocarpodendron'' subsp. ''conocarpodendron'' is well protected in the
Table Mountain National Park Table Mountain National Park, previously known as the Cape Peninsula National Park, is a national park in Cape Town, South Africa, proclaimed on 29 May 1998, for the purpose of protecting the natural environment of the Table Mountain Chain, and ...
, where it is locally abundant. The subspecies is nonetheless regarded a
vulnerable species A vulnerable species is a species which has been Conservation status, categorized by the International Union for Conservation of Nature as being threatened species, threatened with extinction unless the circumstances that are threatened species, ...
as a significant part of its range was lost due to urbanization and
land conversion Land development is the alteration of landscape in any number of ways such as: * Changing landforms from a natural or semi-natural state for a purpose such as agriculture or housing * Subdividing real estate into lots, typically for the purpose ...
to gum plantations and
invasive plant species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
. Further housing developments of the Cape Town agglomerations could threaten remaining habitat. Alien ant species have driven back native ants. The alien ants eat the elaiosome on the spot, so that the seed is not protected against consumption or fire. Due to the proximity of housing areas, wildfires in its range are suppressed and so allow the
fynbos Fynbos (; meaning fine plants) is a small belt of natural shrubland or heathland vegetation located in the Western Cape and Eastern Cape provinces of South Africa. This area is predominantly coastal and mountainous, with a Mediterranean clim ...
to develop into a thicket less suitable for the grey tree pincushion, causing the fires when they eventually occur to be hotter because of more biomass, which results in more plants dying. Finally, subsp. ''viridum'' is planted in gardens within the range of subsp. ''conocarpodendron'', which will lead to hybridization between the subspies. This occurs even when the plants are quite far apart as both subspecies are bird pollinated. This could eventually lead to the extinction of subsp. ''conocarpodendron''.


References


External links


several photos of subsp. ''conocarpodendron''

several photos of subsp. ''viridum''
{{Taxonbar, from=Q3642255 Trees of South Africa Trees of Mediterranean climate conocarpodendron Endemic flora of South Africa Plants described in 1753 Taxa named by Carl Linnaeus