''Cliffortia'', or Caperose is a genus of plants that has been assigned to the

rose family Rosaceae (), the rose family, is a medium-sized family of flowering plants that includes 4,828 known species in 91 genera. The name is derived from the type genus ''Rosa''. Among the most species-rich genera are '' Alchemilla'' (270), ''Sorbus ...

, with currently 132 known species. Its species can be found in southern Africa, particularly in the Cape Floristic Region where 124 of the species can be found, 109 of which are

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 ...

to the CFR. Most species are ericoid 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 over an area of ground. Groundcover provides protection of the topsoil from erosion and drought. In an ecosystem, the ground cover forms the layer of vegetation below the shrub layer known as t ...

. All are

wind pollinated Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Almost all gymnosperms are anemophilous, as are many plants in the order Poales, including grasses, sedges, and rushes. Other common anemophilous pla ...

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 t ...

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). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...

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, 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 ...

. 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 any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...

, 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 alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is conne ...

), while other species have plants of separate gender (or are

dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...

). The flowers never have

petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...

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 The term ''sepalum'' was coined b ...

s of variable size, and in male flowers six to many

stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...

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 dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and indehiscent (they do not ope ...

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 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. 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 genera. It is the first work to consistently apply binomial names and was the ...

, 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 ( cs, Karel Bořivoj 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 cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

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 t ...

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 region ...

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 50 ...

'' or the

Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...

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

, 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 Nucle ...

and

chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...

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 ca ...

, which is assumed to be widespread within ''Cliffortia''. ''Cliffortia'' has been named in homor 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 ( nl, Verenigde Oostindische Compagnie, the VOC) was a chartered company established on the 20th March 1602 by the States General of the Netherlands amalgamating existing companies into the first joint-stock ...

, 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, a wealthy A ...

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 a ...

. 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, where they almost exclusively grow in

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 ...

, and in the transition zones with adjoining

biome A biome () is a biogeographical unit consisting of a biological community that has formed in response to the physical environment in which they are found and a shared regional climate. Biomes may span more than one continent. Biome is a broader ...

s like the

Karoo The Karoo ( ; from the Afrikaans borrowing of the South Khoekhoe !Orakobab or Khoemana word ''ǃ’Aukarob'' "Hardveld") is a semi-desert natural region of South Africa. No exact definition of what constitutes the Karoo is available, so its ext ...

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 in ...

. 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

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 ...

heathlands, between

Mount Kenya Mount Kenya (Kikuyu: ''Kĩrĩnyaga'', Kamba, ''Ki Nyaa'') is the highest mountain in Kenya and the second-highest in Africa, after Kilimanjaro. The highest peaks of the mountain are Batian (), Nelion () and Point Lenana (). Mount Kenya is locat ...

(''C. nitidula'') in the north to the southern

Drakensberg The Drakensberg (Afrikaans: Drakensberge, Zulu: uKhahlambha, Sotho: Maluti) is the eastern portion of the Great Escarpment, which encloses the central Southern African plateau. The Great Escarpment reaches its greatest elevation – within th ...

in the south. Few grow on

dolorite Diabase (), also called dolerite () or microgabbro, is a mafic, holocrystalline, subvolcanic rock equivalent to volcanic basalt or plutonic gabbro. Diabase dikes and sills are typically shallow intrusive bodies and often exhibit fine-grained ...

s 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 provinces of South Africa. Its capital is Bhisho, but its two largest cities are East London and Gqeberha. The second largest province in the country (at 168,966 km2) after Northern Cape, it was formed in ...

and

Kwazulu-Natal KwaZulu-Natal (, also referred to as KZN and known as "the garden province") is a province of South Africa that was created in 1994 when the Zulu bantustan of KwaZulu ("Place of the Zulu" in Zulu) and Natal Province were merged. It is locate ...

References

External links

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