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''Strophocactus wittii'',
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are al ...
''Selenicereus wittii'', known as the Amazon moonflower, is a species of plant in the genus ''
Strophocactus ''Strophocactus'' is a genus of cacti in the subfamily Cactoideae. Its status and circumscription remain somewhat uncertain, with the genus containing one to three species (not always the same ones). Molecular phylogenetic data suggest that it co ...
'' in the cactus family (Cactaceae), and is one of several species commonly called "moonflowers". It was first described in 1900 and is one of three species of cactus found in the central
Amazon basin The Amazon basin is the part of South America drained by the Amazon River and its tributaries. The Amazon drainage basin covers an area of about , or about 35.5 percent of the South American continent. It is located in the countries of Boli ...
. It grows epiphytically on the trunks of trees in seasonally flooded forests of the Amazon basin, which is regularly flooded for a few weeks each year. During this time, the seeds spread through the water, which is unique within the cactus family.


Description


Vegetative characteristics

''Strophocactus wittii'' grows as an
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
, creeping and climbing up trees. The richly branched, leaf-like, flattened stems are
phylloclade Phylloclades and cladodes are flattened, photosynthetic shoots, which are usually considered to be modified branches. The two terms are used either differently or interchangeably by different authors. '' Phyllocladus'', a genus of conifer, is nam ...
s, and are pressed close to the tree trunks of their carriers and form
aerial root Aerial roots are roots above the ground. They are almost always adventitious. They are found in diverse plant species, including epiphytes such as orchids (''Orchidaceae''), tropical coastal swamp trees such as mangroves, banyan figs (''Fic ...
s along their midrib. When exposed to direct sunlight, the elliptical to lanceolate, dark green shoots turn dull red due to strong betalain pigmentation and are therefore clearly visible from a distance. Each segment is up to 60 cm long and 6–14 cm wide, but only 2 to 4 mm thick. Their edges are slightly notched, and are covered with white woolly
areole In botany, areoles are small light- to dark-colored bumps on cacti out of which grow clusters of spines. Areoles are important diagnostic features of cacti, and identify them as a family distinct from other succulent plants. Gordon Rowley - W ...
s about every 8–10 mm. Up to 20 needle-shaped, whitish spines up to 12 mm long arise from the areoles.


Flowers

The salver-shaped
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
s are up to 27 cm long and reach a diameter of 12.5 cm. The long, slender flower tube measures just 9 mm in diameter. The
tepal A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
s then spread out flat. They are pure white and intensely reflect
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiation ...
light. The
hypanthium In angiosperms, a hypanthium or floral cup is a structure where basal portions of the calyx, the corolla, and the stamens form a cup-shaped tube. It is sometimes called a floral tube, a term that is also used for corolla tube and calyx tube. It ...
and the flower tube are scaly and covered with hair-like spines. The
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...
ies at the base of the flower secrete a large amount of clear nectar. The stigma lobes and the lower portion of the style are warty (papillose). The pollen grains of ''Strophocactus wittii'' are hexacolpate, that is, they have six instead of three germ folds on the surface of the pollen grains. In its natural location, ''Strophocactus wittii'' blooms in May. In
greenhouse A greenhouse (also called a glasshouse, or, if with sufficient heating, a hothouse) is a structure with walls and roof made chiefly of transparent material, such as glass, in which plants requiring regulated climatic conditions are grown.These ...
cultivation in Europe it flowers between November and February. The flowers only open for one night. The flower typically begins to open after sunset and is fully open within two hours, closing again at sunrise. Until the flowers are fully open, the flowers initially give off an intense fragrance, which eventually turns into an unpleasant odor. The components responsible for the odor have been identified as
benzyl alcohol Benzyl alcohol is an aromatic alcohol with the formula C6H5CH2OH. The benzyl group is often abbreviated "Bn" (not to be confused with "Bz" which is used for benzoyl), thus benzyl alcohol is denoted as BnOH. Benzyl alcohol is a colorless liquid w ...
,
benzyl benzoate Benzyl benzoate is an organic compound which is used as a medication and insect repellent. As a medication it is used to treat scabies and lice. For scabies either permethrin or malathion is typically preferred. It is applied to the skin as a ...
and benzyl salicylate.


Fruits and seeds

The
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in partic ...
s are elongated, greenish and thorny
berries A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit, although many pips or seeds may be present. Common examples are strawberries, ras ...
, about 3.5 cm long. They mature in their natural location in about a year and then tear open along a longitudinal opening. The pulp contained in the fruit is rather dry. The shell-shaped, glossy black-brown
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
s are about 4 mm long and 2 mm wide. This makes them unusually large for seeds of the cactus family. The hilum and the
micropyle Micropyle may refer to: * Micropyle (botany) a minute opening in the integument of an ovule of a seed plant. * Micropyle (zoology) A micropyle is a pore in the membrane covering the ovum, through which a sperm enters. Micropyles are also found in s ...
of the seeds are united. The
seed coat A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm p ...
is almost smooth. The main part of the seed consists of a layer of greatly enlarged, dead cells that are on the top and are filled with air, which allows the seeds to float in water.


Taxonomy

''Strophocactus wittii'' was discovered in 1899 in the Igapó forests of the Rio Negro near
Manaus Manaus () is the capital and largest city of the Brazilian state of Amazonas. It is the seventh-largest city in Brazil, with an estimated 2020 population of 2,219,580 distributed over a land area of about . Located at the east center of the s ...
by the German businessman and hobby plant collector Nikolaus Heinrich Witt (from whom the second part of the
scientific name 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, bo ...
is derived). He sent a plant to Karl Moritz Schumann in
Berlin-Dahlem Dahlem ( or ) is a locality of the Steglitz-Zehlendorf borough in southwestern Berlin. Until Berlin's 2001 administrative reform it was a part of the former borough of Zehlendorf. It is located between the mansion settlements of Grunewald and ...
, who at first could not classify it taxonomically. Only when Schumann received further specimens from Witt in the autumn of 1900 and discovered fruit on them, was he able to assign them to the genus '' Cereus'', and publish the first
species description A species description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species that have ...
as ''Cereus wittii''. With the choice of the specific epithet, Schumann honored the discoverer of the species. The
botanical illustrator Botanical illustration is the art of depicting the form, color, and details of plant species, frequently in watercolor paintings. They must be scientifically accurate but often also have an artistic component and may be printed with a botanical ...
Margaret Mee (1909–1988) was able to observe and draw the species in its wild habitat. The main study of the biology, ecology and distribution of the species was made by
Wilhelm Barthlott Wilhelm Barthlott (born 1946 in Forst, Germany) is a German botanist and biomimetic materials scientist. His official botanical author citation is Barthlott. Barthlott's areas of specialization are biodiversity (Global distribution, assessment, ...
in 1997. The systematic position of the species has been disputed.
Nathaniel Lord Britton Nathaniel Lord Britton (January 15, 1859 – June 25, 1934) was an American botanist and taxonomist who co-founded the New York Botanical Garden in the Bronx, New York. Early life Britton was born in New Dorp in Staten Island, New York to ...
and
Joseph Nelson Rose Joseph Nelson Rose (January 11, 1862 – May 4, 1928) was an American botanist. He was born in Union County, Indiana. His father died serving during the Civil War when Joseph Rose was a young boy. He later graduated from high school in Libert ...
created the monotypical genus ''Strophocactus'' in 1913 with the only species ''Strophocactus wittii''. They derived the
botanical name A botanical name is a formal scientific name conforming to the ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) and, if it concerns a plant cultigen, the additional cultivar or Group epithets must conform to the ''Interna ...
of the genus from the Greek noun στροφή ''strophe'' for "wind, twist" because the shoots wound or twisted around the host plant. In 1986, Gordon Rowley transferred the species to the genus ''
Selenicereus ''Selenicereus'', sometimes known as moonlight cactus, is a genus of epiphytic, lithophytic, and terrestrial cacti, found in Mexico, Central America, the Caribbean and northern South America. The term night-blooming cereus is also sometimes ...
'' due to its flower structure. In 2003, Ralf Bauer suggested that ''Strophocactus'' should be recognized again and that ''Selenicereus wittii'' should be re-classified in this genus.


Phylogeny

A
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
study of the
Hylocereeae The Hylocereeae are a tribe of cacti. Most are found in the tropical forests of Central and northern South America, and are climbers or epiphytes, unlike most cacti. The tribe includes between six and eight genera in different circumscriptions ...
in 2017 showed that the widely circumscribed genus ''Selenicereus'' was not
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 gr ...
, and neither was ''Strophocactus'': The authors placed ''Selenicereus wittii'' back in ''Strophocactus'', together with the former ''Pseudoacanthocereus sicariguensis'' and ''P. brasiliensis'', and excluded the genus ''Strophocactus'' from the Hylocereeae. The placement as ''Strophocactus wittii'' is accepted by
Plants of the World Online Plants of the World Online (POWO) is an online database published by the Royal Botanic Gardens, Kew. It was launched in March 2017 with the ultimate aim being "to enable users to access information on all the world's known seed-bearing plants by ...
, .


Distribution

''Strophocactus wittii'' is common in the
rainforests Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainforest, ...
of the central
Amazon basin The Amazon basin is the part of South America drained by the Amazon River and its tributaries. The Amazon drainage basin covers an area of about , or about 35.5 percent of the South American continent. It is located in the countries of Boli ...
along
blackwater river A blackwater river is a type of river with a slow-moving channel flowing through forested swamps or wetlands. As vegetation decays, tannins leach into the water, making a transparent, acidic water that is darkly stained, resembling black tea ...
s. There the species grows in abundance in the crown area of the temporarily flooded floodplain forests. The distribution area extends along the Rio Negro and Rio Japurá in
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
over the Río Vaupés, Río Apaporis and Caquetá in
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the ...
to northeast
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = National seal , national_motto = "Firm and Happy f ...
in the
Loreto region Loreto () is Peru's northernmost department and region. Covering almost one-third of Peru's territory, Loreto is by far the nation's largest department; it is also one of the most sparsely populated regions due to its remote location in the Ama ...
to the city of
Iquitos Iquitos (; ) is the capital city of Peru's Maynas Province and Loreto Region. It is the largest metropolis in the Peruvian Amazon, east of the Andes, as well as the ninth-most populous city of Peru. Iquitos is the largest city in the world t ...
and probably includes the southern area of the Amazon basin in
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
. In addition to ''Strophocactus wittii'', only the two cactus species ''
Rhipsalis baccifera ''Rhipsalis baccifera'', commonly known as the mistletoe cactus, is an epiphytic cactus which originates from Central and South America, the Caribbean, and Florida. It is also found throughout the tropics of Africa and into Sri Lanka where it i ...
'' and '' Epiphyllum phyllanthus'' grow there.


Ecology


Pollination

Certain flower characteristics, for example the pure white color, the extreme length of the flower tube, the scent and the nocturnal opening, indicate that the flowers are only pollinated by
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s – in this case sphinx moths. However, pollination has not yet been observed in nature. Due to the length of the flower tube only two species occurring in the natural range of ''Strophocactus wittii'' have a sufficiently long
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
(up to 25 cm): '' Neococytius cluentius'' and '' Amphimoea walkeri''.


Spread

The structure of the seeds with their large, air-filled, outer cells of the seed coat differs from all other types of cactus. The floating seeds disperse through the water (
hydrochory In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...
). The epiphytic orchid species ''
Galeandra devoniana ''Galeandra'', abbreviated as Gal in horticultural trade, is a genus of 37 known species of orchids native to South America, Central America, the West Indies and Florida. Species Species currently accepted as of June 2014: #'' Galeandra arundi ...
,'' which also lives in the Igapó forests of the Rio Negro, spreads in this way, as does the sundew ''
Drosera amazonica ''Drosera'', which is commonly known as the sundews, is one of the largest genera of carnivorous plants, with at least 194 species. 2 volumes. These members of the family Droseraceae lure, capture, and digest insects using stalked mucilaginous ...
''.


Notes


References

{{Taxonbar, from1=Q1292003, from2=Q777440 Echinocereeae Plants described in 1900 Flora of northern South America Flora of western South America