Schoepfia
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''Schoepfia'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of small
hemiparasitic A parasitic plant is a plant that derives some or all of its nutritional requirements from another living plant. They make up about 1% of angiosperms and are found in almost every biome. All parasitic plants develop a specialized organ called the ...
trees,
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s belonging to the family
Schoepfiaceae Schoepfiaceae is a family of flowering plants recognized in the APG III system of 2009. The family was previously only recognized by few taxonomists; the plants in question usually being assigned to family Olacaceae and Santalaceae. The genus '' ...
. The genus has long been placed in the
Olacaceae Olacaceae is a family of flowering plants in the order Santalales. They are woody plants, native throughout the tropical regions of the world. , the circumscription of the family varies; some sources maintain a broad family, others split it int ...
family.


Description

Plants in this genus are small trees or
shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
s which exhibit
heterostyly Heterostyly is a unique form of polymorphism and herkogamy in flowers. In a heterostylous species, two or three morphological types of flowers, termed "morphs", exist in the population. On each individual plant, all flowers share the same morph ...
- individuals can have both often cylindrical brachystylous (short styled) flowers and somewhat bell-shaped dolichostylous (longer styled) flowers. In most plants where heterostyly occurs, there is a sexual differentiation between flower types, the brachystylous flowers being functionally male, or one type of flower is
cleistogamous Cleistogamy is a type of automatic self-pollination of certain plants that can propagate by using non-opening, self-pollinating flowers. Especially well known in peanuts, peas, and pansies, this behavior is most widespread in the grass family. H ...
or self-fertile. In ''Schoepfia'' species both flowers are bisexual and can form fruit, the reason for two flower forms is mysterious. The flowers are fragrant and small. They arise from a short peduncle which grows from the leaf axils of a stem. The peduncle is subtended by persistent, imbricate perular
bract In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...
s. The flower is subtended by a three-lobed
epicalyx This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary ...
, it is composed of a bract and two
bracteole In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also look ...
s, which are all united into a single structure. The actual
calyx CALYX, Inc. is a non-profit publisher of art and literature by women founded in 1976 based in Corvallis, Oregon. CALYX publishes both '' CALYX, A Journal of Art and Literature by Women'' twice a year and CALYX Books, which publishes one to three ...
is inconspicuous, it is completely fused to the truncated, cup-shaped base of the flower. From the edges of this base, generally four or five
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 arise, rarely three or six. The lower part of these petals are fused to each other to form a tube of sorts. The leaves of all species are simple, entire, alternate, penninerved and have petioles. The fruits are single-seeded
drupe In botany, a drupe (or stone fruit) is a type of fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pip'' (UK), ''pit'' (US), ''stone'', or ''pyrena'') of hardened endocarp with a seed ...
s. The
epicarp Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Fruitlike structures may develop directly from the seed itself rather than ...
is derived from the swollen base of the flower.


Taxonomy

It is now placed in the
Schoepfiaceae Schoepfiaceae is a family of flowering plants recognized in the APG III system of 2009. The family was previously only recognized by few taxonomists; the plants in question usually being assigned to family Olacaceae and Santalaceae. The genus '' ...
family. The genus has long been placed in the
Olacaceae Olacaceae is a family of flowering plants in the order Santalales. They are woody plants, native throughout the tropical regions of the world. , the circumscription of the family varies; some sources maintain a broad family, others split it int ...
family, although many researchers noted the differences between it and the rest of the family. Recent molecular studies have shown ''Schoepfia'' to be more closely related to the families
Misodendraceae ''Misodendrum'' is a genus of hemiparasites which grow as mistletoes on various species of ''Nothofagus''. Its species are all restricted to South America. The name of the genus is incorrectly spelt in a number of ways, including ''Misodendron'' ...
and
Loranthaceae Loranthaceae, commonly known as the showy mistletoes, is a family of flowering plants. It consists of about 75 genera and 1,000 species of woody plants, many of them hemiparasites. The three terrestrial species are '' Nuytsia floribunda'' (the ...
, and to uphold the criteria of
monophyly 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 comm ...
it must be excluded from Olacaceae. The genus is split into three
sections Section, Sectioning, or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
: two occur exclusively in Asia, with four species altogether, the third section, which includes the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
, only occurs in the Americas and contains all the other species. The type species is ''Schoepfia schreberi''. One possible fossil species, '' S. republicensis'', has been described from
Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
leaves found in the
Klondike Mountain Formation The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington (state), Washington state. The formation is composed of volcanic rocks in the upper unit and volcanic plus lacu ...
of Washington state.


Species

,
Plants of the World Online Plants of the World Online (POWO) is an online taxonomic database published by the Royal Botanic Gardens, Kew. History Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...
accepted the following species: *''
Schoepfia arenaria ''Schoepfia arenaria'' is an extremely rare species of hemiparasitic flowering plant in the Schoepfiaceae family. It grows as a small, multi-trunked tree. It is endemic to Puerto Rico, where it is found growing along the northern coast. A local S ...
'' Britton *'' Schoepfia brasiliensis'' A.DC. *'' Schoepfia californica'' Brandegee *'' Schoepfia chinensis'' Gardner & Champ. *'' Schoepfia clarkii'' Steyerm. *'' Schoepfia cubensis'' Britton & P.Wilson *'' Schoepfia didyma'' C.Wright ex Griseb. *'' Schoepfia flexuosa'' (Ruiz & Pav.) Roem. & Schult. *'' Schoepfia fragrans'' Wall. *'' Schoepfia griffithii'' Tiegh. ex Steenis *'' Schoepfia haitiensis'' Urb. & Britton *'' Schoepfia harrisii'' Urb. *'' Schoepfia jasminodora'' Siebold & Zucc. *'' Schoepfia lucida'' Pulle *'' Schoepfia macrophylla'' Lundell *'' Schoepfia multiflora'' Urb. *'' Schoepfia obovata'' C.Wright - white beefwood *'' Schoepfia paradoxa'' (Bisse & Berazaín) Berazaín ex Acev.-Rodr. *'' Schoepfia parvifolia'' Planch. *'' Schoepfia pringlei'' B.L.Rob. *'' Schoepfia schreberi'' J.F.Gmel. - gulf graytwig *'' Schoepfia scopulorum'' Alain *'' Schoepfia shreveana'' Wiggins *'' Schoepfia stenophylla'' Urb. *''
Schoepfia tepuiensis ''Schoepfia'' is a genus of small hemiparasitic trees, flowering plants belonging to the family Schoepfiaceae. The genus has long been placed in the Olacaceae family. Description Plants in this genus are small trees or shrubs which exhibit het ...
'' Steyerm. *'' Schoepfia tetramera'' Herzog *'' Schoepfia vacciniiflora'' Planch. ex Hemsl. *'' Schoepfia velutina'' Sandwith


Distribution

The genus has a discontinuous distribution, being native to the
Himalaya The Himalayas, or Himalaya ( ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the Earth's highest peaks, including the highest, Mount Everest. More than 100 pea ...
s through China, Japan and Taiwan to
Sumatra Sumatra () is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the list of islands by area, sixth-largest island in the world at 482,286.55 km2 (182,812 mi. ...
, and to tropical and subtropical America. The genus is most diverse in tropical America. Three species occur in territories belonging to the United States: ''Schoepfia arenaria'', ''S. obovata'' and ''S. schreberi''; ''S. schreberi'' occurs in Florida and the Caribbean, the other two are only found in the US Caribbean possessions.


Ecology

Species in this genus are root
hemiparasite A parasitic plant is a plant that derives some or all of its nutritional requirements from another living plant. They make up about 1% of angiosperms and are found in almost every biome. All parasitic plants develop a specialized organ called th ...
s, deriving a portion of their nutrition by invading the roots and stealing the nutrients of neighbouring plants. They do not appear to be very particular in choice of
host plant In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include a ...
s. A 1979 study of ''Schoepfia schreberi'' in Florida and the Bahamas found
haustoria In botany and mycology, a haustorium (plural haustoria) is a rootlike structure that grows into or around another structure to absorb water or nutrients. For example, in mistletoe or members of the broomrape family, the structure penetrates th ...
(specialised organs on the roots used for parasitism) connected to ten different species, belonging to eight different botanical families. Haustoria were also firmly welded to the wrong things: rocks, buried pipes, concrete building foundations, etc. The young leaves of the ''S. schreberi'' are fed upon by the balloon-collared caterpillars of the metalmark butterfly '' Calydna sturnula'', which are only known to feed upon this species.


References

Schoepfiaceae Santalales genera {{Santalales-stub