Cliffortia Linearifolia
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''Cliffortia'', or Caperose is a genus of plants that has been assigned to the
rose family Rosaceae (), the rose family, is a family of flowering plants that includes 4,828 known species in 91 genera. The name is derived from the type genus ''Rosa''. The family includes herbs, shrubs, and trees. Most species are deciduous, but some ...
, with currently 132 known species. Its species can be found in southern Africa, particularly in the
Cape Floristic Region The Cape Floral Region is a floristic region located near the southern tip of South Africa. It is the only floristic region of the Cape Floristic Kingdom, and includes only one floristic province, known as the Cape Floristic Province. The Cap ...
where 124 of the species can be found, 109 of which are
endemic Endemism is the state of a species being found only 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 foun ...
to the CFR. Most species are
ericoid The word "ericoid" is used in modern biological terminology for its literal meanings and for extensions. Ericoid could have more than one meaning, but in practice the most common use is in reference to a plant's habit, to describe small, tough (s ...
shrubs, some small trees up to 5 m (16 ft) high, others more or less herbaceous
groundcover Groundcover or ground cover is any plant that grows low over an area of ground, which protects the topsoil from erosion and drought. In a terrestrial ecosystem, the ground cover forms the layer of vegetation below the shrub layer known as the ...
. All are wind pollinated and have separate male and female flowers in the axils of the leaves, mostly individually, sometimes grouped, which may be on the same plant or on separate plants.


Description

''Cliffortia'' species are mostly upright shrubs, but some species develop into small trees of up to 5 m (16 ft) high, are more or less
herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of ...
groundcover Groundcover or ground cover is any plant that grows low over an area of ground, which protects the topsoil from erosion and drought. In a terrestrial ecosystem, the ground cover forms the layer of vegetation below the shrub layer known as the ...
or grow in a dense tangle. The
stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole (botany), petiole). They are primarily found among dicots and rare among monocots. Stipules are considered part ...
s have merged with the base of the leaf and form a sheath around the branch. The leaves are alternately arranged along the stems, and may consist of three, two or only one leaflet with one or several main veins, seated or on a
leaf stalk In botany, the petiole () is the stalk that attaches the leaf blade to the stem. It is able to twist the leaf to face the sun, producing a characteristic foliage arrangement (spacing of blades), and also optimizing its exposure to sunlight. Outg ...
. Leaflets may be thin or leathery, broad to needle-shaped, with the margin serrated or entire, and may have a spiny tip. ''Cliffortia'' has separate male and female flowers in the
leaf axils A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, fl ...
, which are mostly set individually but sometimes in clusters. Some species have both male and female flowers on the same plant (and are
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system comparable with gynomonoecy, andromonoecy and trimonoecy, and contras ...
), while other species have plants of separate gender (or are
dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
). The flowers never have
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s, which in other plant species predominantly function to attract pollinators, but that function is of cause unnecessary in wind-pollinated taxa, such as in the entire genus ''Cliffortia''. 3-merous flowers almost certainly represent the ancestral state, and it has been suggested that 4-merous flowers have arisen at four independent occasions. Trimerous flowers have three
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...
s of variable size, and in male flowers six to many
stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
s. Tetramerous flowers have four sepals of 5 mm long at most and either four or eight stamens. In female flowers the sepals are united at their base to form a calyx tube, and have one or two styles, that are finely divided like an ostridge feather. One or two
achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple fruit, simple dry fruits, dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and Dehi ...
s may develop in each flower, within the inflating calyx.


Taxonomy

The first who mentioned species that are currently assigned to ''Cliffortia'' was
Leonard Plukenet Leonard Plukenet (1641–1706) was an English botanist, Royal Professor of Botany and gardener to Queen Mary. Biography Plukenet published ''Phytographia'' (London, 1691–1696) in four parts in which he described and illustrated rare exotic p ...
in 1696. 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 binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...
proposed by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
, are not valid however. It was Linnaeus himself who provided the first descriptions in the
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 genus, genera. It is the first work to consistently apply binomial nomenclature ...
, his groundbreaking work of 1753, which he named ''C. ilicifolia'', ''C. ruscifolia'', ''C. polygonifolia'' and ''C. trifoliata''. In 1808, Necker subdivided the genus and created ''Morilandia'', while
Carl Borivoj Presl Carl Borivoj Presl (; 17 February 1794 – 2 October 1852) was a Czech botanist. Biography Presl lived his entire life in Prague, and was a professor of botany at the University of Prague (1833–52).monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
subfamily Rosoidea and the tribe Sanguisorbeae. The immediate
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of ''Cliffortia'' may either be the
holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical reg ...
genus ''
Sanguisorba ''Sanguisorba'' is a genus of flowering plants in the family Rosaceae native to the temperate regions of the Northern Hemisphere. The common name is burnet. Description The plants are herbaceous perennials or small shrubs. The stems grow to 50â ...
'' or the
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
n genus ''
Acaena ''Acaena'' is a genus of about 60 species of mainly evergreen, creeping herbaceous perennial plants and subshrubs in the family Rosaceae, native mainly to the Southern Hemisphere, notably New Zealand, Australia and South America, but with a few ...
''. ''Cliffortia'' is clearly
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
, but within it only four monophyletic subgenera can be recognised. These are named ''Arborea'', ''Cliffortia'', ''Eriocephalina'' and ''Graminea''. Since comparison of homologous DNA of different genes (in the
nucleus Nucleus (: nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucleu ...
and
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
respectively) result in other phylogenetic trees, it is assumed that many ''Cliffortia'' species have hybridized. Their off-spring survived and propagated further, forming new
nothospecies #REDIRECT Hybrid name In botanical nomenclature, a hybrid may be given a hybrid name, which is a special kind of botanical name, but there is no requirement that a hybrid name should be created for plants that are believed to be of hybrid origin. ...
. This is called
reticulate evolution Reticulate evolution, or network evolution is the origination of a lineage through the partial merging of two ancestor lineages, leading to relationships better described by a phylogenetic network than a bifurcating tree. Reticulate patterns can ...
, which is assumed to be widespread within ''Cliffortia''. ''Cliffortia'' has been named in honor of
George Clifford III George Clifford III (7 January 1685, Amsterdam – 10 April 1760, Heemstede) was a wealthy Dutch banker and one of the directors of the Dutch East India Company. He is known for his keen interest in plants and gardens. His summer estate Harteka ...
, a wealthy Dutch banker and director of the
Dutch East India Company The United East India Company ( ; VOC ), commonly known as the Dutch East India Company, was a chartered company, chartered trading company and one of the first joint-stock companies in the world. Established on 20 March 1602 by the States Ge ...
, who enabled Linnaeus to write the
Hortus Cliffortianus The ''Hortus Cliffortianus'' is a work of early botanical literature published in 1737. The work was a collaboration between Carl Linnaeus and the illustrator Georg Dionysius Ehret, financed by George Clifford in 1735–1736. Clifford was a we ...
in 1737.


Ecology

''Cliffortia'' species differ in their tolerance for fire, and roughly three strategies may be distinguished. A large proportion of the species resprouts from the underground
rootstock A rootstock is part of a plant, often an underground part, from which new above-ground growth can be produced. It could also be described as a stem with a well developed root system, to which a bud from another plant is grafted. It can refer to ...
. Many other species have seeds that are collected by ants and survive in their underground nests. Fewer species survive the less intense fires by regrowing from the crown of the plant, but those species grow in locations that do not sustain intense fires, such as isolated from the surrounding vegetation by bare rock (''C. neglecta'', ''C. tuberculata'', ''C. complanata'' and ''C. propinqua''), wet areas (''C. aculeata'', ''C. graminea'', ''C. nivenioides'' and ''C. strobilifera''), or montane grasslands (''C. linearifolia'', ''C. nitidula'' subsp. ''pilosa'', ''C. repens'' and ''C. spathulata'').


Distribution and habitat

''Cliffortia'' species can be found throughout the
Cape Floristic Region The Cape Floral Region is a floristic region located near the southern tip of South Africa. It is the only floristic region of the Cape Floristic Kingdom, and includes only one floristic province, known as the Cape Floristic Province. The Cap ...
, where they almost exclusively grow in
fynbos Fynbos (; , ) is a small belt of natural shrubland or heathland vegetation located in the Western Cape and Eastern Cape provinces of South Africa. The area is predominantly coastal and mountainous, with a Mediterranean climate. The fynbos ...
, and in the transition zones with adjoining
biome A biome () is a distinct geographical region with specific climate, vegetation, and animal life. It consists of a biological community that has formed in response to its physical environment and regional climate. In 1935, Tansley added the ...
s like the
Karoo The Karoo ( ; from the Afrikaans borrowing of the South Khoekhoe Khoemana (also known as !Orakobab or Korana) word is a semidesert natural region of South Africa. No exact definition of what constitutes the Karoo is available, so its extent is ...
and
Albany thickets The Albany thickets is an ecoregion of dense woodland in southern South Africa, which is concentrated around the Albany region of the Eastern Cape (whence the region's name originates). Geography The thickets grow on well-drained sandy soils i ...
. Within the Cape Floristic Region, it occurs in the widest range of environments, from the highest mountain peaks to coastal sand flats. Some species are bound to constantly moist soils (such as ''C. strobilifera'' near streams on acid to neutral soils, and C. longifolia near streams on marine sands and limestone deposits) while others live on the arid, mountainous edge of the
Karoo The Karoo ( ; from the Afrikaans borrowing of the South Khoekhoe Khoemana (also known as !Orakobab or Korana) word is a semidesert natural region of South Africa. No exact definition of what constitutes the Karoo is available, so its extent is ...
in rock crevices. Most ''Cliffortia'' species are limited to acidic and nutrient-poor sandy soils, but some also grow on alkaline limestone soils (''C. falcata'', ''C. ferruginea'', ''C. filifolia'' and ''C. obcordata'') or more fertile shales such as in the Bokkeveld (such as ''C. filicaulis'', ''C. juniperina'' and ''C. polygonifolia''). ''C. complanata'' and ''C. propinqua'' are both confined to shady rock crevices, except in the moisture of the south coast. Outside the Cape Floristic Region most of the ''Cliffortia'' species occur the
afromontane The Afromontane regions are subregions of the Afrotropical realm, one of the Earth's eight biogeographic realms, covering the plant and animal species found in the mountains of Africa and the southern Arabian Peninsula. The Afromontane regions o ...
heathlands, between
Mount Kenya Mount Kenya (Meru people, Meru: ''Kĩrĩmaara,'' Kikuyu people, Kikuyu: ''Kĩrĩnyaga'', Kamba language, Kamba: ''Ki nyaa'', Embu language, Embu: ''Kĩ nyaga'') is an extinct volcano in Kenya and the Highest mountain peaks of Africa, second-highe ...
(''C. nitidula'') in the north to the southern
Drakensberg The Drakensberg (Zulu language, Zulu: uKhahlamba, Sotho language, Sotho: Maloti, Afrikaans: Drakensberge) is the eastern portion of the Great Escarpment, Southern Africa, Great Escarpment, which encloses the central South Africa#Geography, Sout ...
in the south. Few grow on dolorites near the
Great Escarpment The Great Escarpment is a major topographical feature in Africa that consists of steep slopes from the high central Southern African plateauAtlas of Southern Africa. (1984). p. 13. Reader's Digest Association, Cape Town downward in the directio ...
and sandstone at low altitude in the
Eastern Cape The Eastern Cape ( ; ) is one of the nine provinces of South Africa. Its capital is Bhisho, and its largest city is Gqeberha (Port Elizabeth). Due to its climate and nineteenth-century towns, it is a common location for tourists. It is also kno ...
and
Kwazulu-Natal KwaZulu-Natal (, also referred to as KZN) is a Provinces of South Africa, province of South Africa that was created in 1994 when the government merged the Zulu people, Zulu bantustan of KwaZulu ("Place of the Zulu" in Zulu language, Zulu) and ...
.


References


External links


Some photos
{{Taxonbar, from=Q8346921 Sanguisorbinae Rosaceae genera Flora of Southern Africa