Sarracenia Rubra Ssp Jonesii
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''Sarracenia'' ( or ) is a genus comprising 8 to 11 species of North American pitcher plants, commonly called trumpet pitchers. The genus belongs to the family Sarraceniaceae, which also contain the closely allied genera '' Darlingtonia'' and ''
Heliamphora The genus ''Heliamphora'' ( or ; Greek: ''helos'' " marsh" and ''amphoreus'' "amphora") contains 23 species of pitcher plants endemic to South America.McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. '' Sarraceniaceae of South Americ ...
''. ''Sarracenia'' is a genus of
carnivorous plant Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryot ...
s indigenous to the eastern seaboard of the United States, Texas, the Great Lakes area and southeastern Canada, with most species occurring only in the south-east United States (only '' S. purpurea'' occurs in cold-temperate regions). The plant's leaves have evolved into a funnel or pitcher shape in order to trap insects. The plant attracts its insect prey with secretions from extrafloral nectaries on the lip of the pitcher leaves, as well as a combination of the leaves' color and scent. Slippery footing at the pitcher's rim, causes insects to fall inside, where they die and are digested by the plant with proteases and other enzymes.


Description

''Sarracenia'' are
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 ...
perennial plants that grow from a subterranean
rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...
, with many tubular pitcher-shaped leaves radiating out from the growing point, and then turning upwards with their trap openings facing the center of the crown. The trap is a vertical tube with a 'hood' (the operculum) extending over its entrance; and below it the top of the tube usually has a rolled lip (the peristome) which secretes nectar and scents. The hood itself frequently produces nectar too, but in lesser quantities. The inside of the pitcher tube, depending on the species, can be divided into three to five distinguishable zones: zone 1 is the operculum (or hood), zone 2 is the peristome and rest of the trap entrance, while zones 3 and 4 (which in some species are combined) and 5 (only present in ''S. purpurea'') are further divisions of the actual tube. Each of these zones has a specific function, with corresponding morphophysiological characteristics. *Zone 1: ''Operculum''. In most species the operculum covers at least part of the pitcher has an opening, preventing rain from excessively filling the pitcher, which would result in the loss of prey and dilute the digestive fluid. The operculum also serves to guide prey to the pitcher opening, using a combination of color, scent, and downward-pointing hairs to lead insects toward the trap entrance. Some species, specifically '' S. minor'' and '' S. psittacina'', have opercula that hang low over the pitcher entrance. These are also studded with
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...
-free patches, translucent "windows" which confuse prey into attempting to fly through the operculum, thereby causing them to cascade down the pitcher tube. (A similar, better-developed mechanism is found in the closely related ''
Darlingtonia californica ''Darlingtonia californica'' , also called the California pitcher plant, cobra lily, or cobra plant, is a species of carnivorous plant. It is the sole member of the genus ''Darlingtonia'' in the family Sarraceniaceae. This pitcher plant is nativ ...
''). *Zone 2: ''Peristome and trap entrance''. This zone is composed mainly of the peristome, which produces copious amounts of nectar, luring insect prey to land or crawl onto the perilous footing surrounding the pitcher trap. This zone also includes the waxy upper portion of the pitcher tube. Footing on this zone is especially treacherous, as the waxy deposits on surface of this zone cause unwary insects to lose their footing and tumble into the pitcher depths. *Zone 3: Located below Zone 2, this zone features a leaf surface with non-existent footing, as well as a coating of ultra-fine, downward pointing hairs. Insects that have made it this far lose any chance of escape. It is also studded with digestive
gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...
s, which secrete digestive enzymes into the digestive fluid. *Zone 4: This is the final zone in most species. It is filled with digestive fluids, and readily absorbs nutrients released from the insects by the work of the digestive enzymes and bacteria in the pitcher fluid. Along with more digestive glands, this zone features a thick coating of coarse downward pointing hairs, which makes escape from the digestive fluids impossible. *Zone 5: This zone, located below Zone 4 and found only in ''S. purpurea'', is smooth, glabrous, lacks glands, and does not serve as an absorptive zone. Its function is unknown.


Carnivorous mechanism

All ''Sarracenia'' trap insects and other prey without the use of moving parts. Their traps are static and are based on a combination of lures (including color, scent, and nectar) and inescapability – typically the entrances to the traps are one-way by virtue of the highly adapted features listed above. Most species use a combination of scent, waxy deposits (to clog insect feet) and gravity to topple insect prey into their pitcher. Once inside, the insect finds the footing very slippery with a waxy surface covering the walls of the pitcher. Further down the tube, downward-pointing hairs make retreat impossible, and in the lowest region of the tube, a pool of liquid containing digestive enzymes and wetting agents quickly drowns the prey and begins digestion. The exoskeletons are usually not digested, and over the course of the summer fill up the pitcher tube. Only ''S. purpurea'' normally contains significant amounts of rainwater in its tubular pitchers. It is a myth that all species contain water. In fact, the hoods of the other species help to keep out rain water in addition to keeping flying prey from escaping. ''S. psittacina'', the parrot pitcher, uses a lobster-pot style trap that will admit prey (including tadpoles and small fish during floods) but not allow it to find its way out; and sharp inward-pointing hairs force the victim gradually down to the base of the pitcher where it is digested.


Potential narcotic function of coniine

Coniine, a toxic alkaloid also present in poison hemlock, was first detected in 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 mutualists ...
of '' S. flava''. and has since been detected in 7 other species of ''Sarracenia''. While it was demonstrated that concentrated extracts from ''S. flava'' could paralyze ants, it has not been demonstrated that coniine has narcotic effects on insects at the concentrations naturally present in pitchers of ''S. flava''. Other authors hypothesize that coniine may function as an attractant for insects, or may function both as an attractant and a narcotic.


Flowers and seeds

Flowers are produced early in spring, with or slightly ahead of the first pitchers. They are held singly on long stems, generally well above the pitcher traps to avoid the trapping of potential
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
s. The flowers, which depending on species are 3–10 centimeters in diameter, are dramatic and have an elaborate design which prevents self-pollination. It consists of five sepals superintended by three
bract 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 ...
s, numerous anthers, and an umbrella-like five-pointed style, over which five long yellow or red
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 dangle. The whole flower is held upside-down, so that the umbrella-like style catches the
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
dropped by the
anther 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 filam ...
s. The stigmas are located at the tips of the umbrella-like style. The primary pollinators are
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
s. Bees searching for
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 mutualists ...
must force their way past one of the stigmas to enter the chamber formed by the style. Inside, they will inevitably come in contact with a lot of pollen, both from the hanging anthers and from the pollen collected by the style. Upon exiting, the bees must force their way under one of the flap-like petals. This keeps them away from the stigma, avoiding self-pollination. The next flower visited receives on its stigmata some of the first flower's pollen, and the cycle continues. Floral formula: Ca5 Co5 A G(5) The flowers of almost all species are scented. The scent varies, but is often strong and sometimes unpleasant. ''S. flava'' has an especially strong odor resembling cat urine. Flowers generally last about two weeks. At the end of the flowering period, the petals drop and the ovary, if pollinated, begins to swell. The seed forms in five lobes, with one lobe producing significantly smaller numbers of seeds than the other lobes. On average, 300–600 seed are produced, depending on species and pollination success. Seed takes five months to mature, at which point the seed pod turns brown and splits open, scattering seed. The seeds are 1.5–2 mm in length and have a rough, waxy coat which makes it hydrophobic, possibly for seed dispersal by flowing water. ''Sarracenia'' seed requires a stratification period to germinate in large numbers. Plants grown from seed start producing functioning traps almost immediately, although they differ in morphology from adult traps for the first year or so, being simpler in structure. Plants require 3–5 years to reach maturity from seed.


Growth cycle

Pitcher production begins at the end of the flowering period in spring, and lasts until late autumn. At the end of autumn, the pitchers begin to wither and the plants produce non-carnivorous leaves called phyllodia, which play a role in the economics of carnivory in these species. Since the supply of insects during winter is decreased, and the onset of cold weather slows plant metabolism and other processes, putting energy into producing carnivorous leaves would be uneconomical for the plant.


Genetics

The genus has been found to have a Chromosome number of ''2n''=26, though some earlier studies had found that number to be ''2n''=24.


Range and habitat

Seven of the eight species are confined to the south-eastern coastal plain of the United States. One species, ''S. purpurea'', continues north and west well into Canada. The typical habitat is warm-temperate; all ''Sarracenia'' are perennial and require a distinct summer and winter. A few subspecies or varieties (''S. rubra'' subsp. ''alabamensis'', ''S. rubra'' subsp. ''jonesii'', and ''S. purpurea'' var. ''montana'') can be found more inland in mountains (e.g. the Appalachian mountains). ''Sarracenia'' tend to inhabit fens, herb bogs, and seasonally wet grasslands. These habitats tend to be acidic (low pH) with soil made up of sand and '' Sphagnum''
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
. Frequently, the soil will be poor in nutrients, particularly
nitrate Nitrate is a polyatomic ion A polyatomic ion, also known as a molecular ion, is a covalent bonded set of two or more atoms, or of a metal complex, that can be considered to behave as a single unit and that has a net charge that is not zer ...
s, and often continuously leached by moving water or made unavailable to the plant roots by the low pH. The plants gain their advantage from their ability to extract nutrients from insect prey in this mineral-poor environment. The plants prefer strong, direct sunlight with no shade. ''Sarracenia'' habitats in the southeastern Coastal Plain consist primarily of fire-maintained pine savannas, wet prairies, or seepage bogs. Without frequent fire (1–3 years), these habitats undergo ecological succession and are quickly invaded by woody shrubs and trees, which eliminate ''Sarracenia'' by increasing shade and reducing soil moisture. In several cases, carnivorous plant enthusiasts have introduced ''S. purpurea'' into suitable habitats outside of its natural range, where it has naturalized. Some of these populations are decades old; the oldest known occurrence in the Swiss
Jura mountains The Jura Mountains ( , , , ; french: Massif du Jura; german: Juragebirge; it, Massiccio del Giura, rm, Montagnas da Jura) are a sub-alpine mountain range a short distance north of the Western Alps and mainly demarcate a long part of the Frenc ...
is around one hundred years old. Besides
Switzerland ). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...
, such naturalized populations can be found in Ireland, England (Lake District), Germany ( Bavaria, Lusatia) and in Mendocino County along the California coast.


Environmental status

''Sarracenia'' are threatened in the wild by development and the drainage of their habitat. Estimates indicated that 97.5% of ''Sarracenia'' habitat has already been destroyed in the southeastern U.S.,Groves, M., ed. 1993. Horticulture, Trade and Conservation of the Genus ''Sarracenia'' in the Southeastern States of America: Proceedings of a Meeting Held at the Atlanta Botanical Garden, September 22–23, 1993, 17pp. the home of all but one subspecies of ''Sarracenia''. Currently the biggest threats to surviving populations are urban development, drainage of habitat for forestry, runoff of
herbicide Herbicides (, ), also commonly known as weedkillers, are substances used to control undesired plants, also known as weeds.EPA. February 201Pesticides Industry. Sales and Usage 2006 and 2007: Market Estimates. Summary in press releasMain page fo ...
s from agriculture, fire suppression, cut pitcher trade for floristry, and plant trade.Robbins, C. S. 1998. Examination of the U.S. Pitcher-plant Trade With a Focus on the White-topped Pitcher-plant. Traffic Bulletin. Excerpts, Vol. 17, No. 2 (June 1998) The latter two threaten survival of ''Sarracenia'' not only through depletion of healthy population, but also because of the damaging effects (soil compaction and altered moisture levels) of repeated foot and vehicular traffic that comes with harvesting. The Fish and Wildlife Service estimates that approximately 1.6 million pitchers were cut for the domestic market in 1991. Some protective legislation exists. Several southeastern states, such as Florida, Georgia, and South Carolina have conservation laws which protect ''Sarracenia''. However, most of the remaining wetlands in the southeastern U.S. are privately owned. Plants on this land are not protected by state legislation. The key states of Alabama and Mississippi have no such legislation at all, so that even plants on public land have no protection. Three ''Sarracenia'' have been listed as "Federally Endangered" under the USA
Endangered Species Act The Endangered Species Act of 1973 (ESA or "The Act"; 16 U.S.C. § 1531 et seq.) is the primary law in the United States for protecting imperiled species. Designed to protect critically imperiled species from extinction as a "consequence of ec ...
(1973) – '' S. rubra'' subsp. ''alabamensis'' (''S. alabamensis'') in Alabama, ''S. rubra'' subsp. ''jonesii'' (''S. jonesii'') in North and South Carolina, and '' S. oreophila'' in Alabama, Georgia, and North Carolina. These taxa are also on CITES Appendix I, giving them international protection by making export of wild-collected plants illegal. The other species, while appearing on CITES Appendix II, have little federal protection. Some efforts have been made to curb the existing threats to plants. In 2003 the International Carnivorous Plant Society ran a trial distribution program in which young ''S. rubra'' subsp. ''alabamanensis'' plants were grown from seed collected from 3 of the 12 known ''S. alabamanensis'' sites, and were distributed to members in an attempt to increase availability of this plant in cultivation, with the hopes of thereby decreasing the poaching that was endangering the survival of this taxon in the wild. In 1995, the non-profit organization Meadowview Biological Research Station was created to preserve and restore pitcher plant bogs and associated ecosystems in Maryland and Virginia. In 2004, a number of concerned plant enthusiasts founded the North American Sarracenia Conservancy (NASC), which aims to "serve as a living record of the taxonomic, morphological and genetic diversity of the genus Sarracenia for purposes of conservation and cultivation." The NASC is a grassroots Nebraska nonprofit organization working to build a genetic ''Sarracenia'' bank by overseeing the maintenance of genetic strains from all remaining wild populations in cultivation, with the eventual aim of being able to supply these strains for re-introduction in suitable habitats. A similar but centralized collection exists in the UK, with 2000+ clones representing all species (many with location data) and numerous hybrids currently being housed by ''Sarracenia'' expert Mike King. This UK collection is part of the
NCCPG National Plant Collection The National Plant Collection scheme is the main conservation vehicle whereby the Plant Heritage charity (formerly the National Council for the Conservation of Plants and Gardens) can accomplish its mission: to conserve, grow, propagate, docu ...
scheme. While none of these efforts curb the biggest threats – urban development and habitat destruction – they aim to help reduce plant poaching while at the same time making these plants available to future generations. One of the biggest challenges of reintroducing plants back into the wild is the unintended introduction of unwanted species, such as pests, diseases, and invasive weeds. Often, it is human destruction of areas in which the Sarracenia thrive that is a major killer. Aside from determining what genetic material is appropriate for reintroduction (which is up for debate), plants must be semi-aseptic to keep the habitat pristine and sustainable in the long term. Another challenge is maintaining all of the introduced plant material and determining an optimal site to plant them in. A single hurricane or storm event can change the dynamics of a field. Even within a single bog, some areas may be waterlogged, while other areas may become very dry, so identifying the right location is critical. Short term results on private property indicate planting larger specimens into the field have a higher chance of long-term survival compared to planting smaller seedlings.


Taxonomy

The genus ''Sarracenia'' belongs to the family Sarraceniaceae, which also contain the closely allied genera '' Darlingtonia'' and ''
Heliamphora The genus ''Heliamphora'' ( or ; Greek: ''helos'' " marsh" and ''amphoreus'' "amphora") contains 23 species of pitcher plants endemic to South America.McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. '' Sarraceniaceae of South Americ ...
''. Under the Cronquist system, this family was put in the order
Nepenthales Nepenthales (Nepenthales Bercht. & J.Presl) is an order of carnivorous flowering plants in the Cronquist system of plant classification. Cronquist system The order was placed in the subclass Dilleniidae, which in the 1981 version of this sys ...
along with
Nepenthaceae ''Nepenthes'' () is a genus of carnivorous plants, also known as tropical pitcher plants, or monkey cups, in the monotypic family Nepenthaceae. The genus includes about 170 species, and numerous natural and many cultivated hybrids. They are mos ...
and Droseraceae.Cronquist, Arthur. (1981). ''An integrated system of classification of flowering plants''. New York: Columbia University Press. The APG II system, however, assigns Sarraceniaceae to the order Ericales and the other two families to the order Caryophyllales.Angiosperm Phylogeny Group. (2003)
An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II
''Botanical Journal of the Linnean Society'', 141: 399–436.
Typically anywhere from 8 to 11 species of ''Sarracenia'' are generally recognized, depending on individual opinions on the biological species concept and which among many
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
and varieties should be elevated to species status, a common
lumping and splitting Lumpers and splitters are opposing factions in any discipline that has to place individual examples into rigorously defined categories. The lumper–splitter problem occurs when there is the desire to create classifications and assign examples to ...
problem in demarcation.Rice, Barry. (2008)
''Sarracenia'' species lists
Accessed: 10-10-2008.
Some authorities split the described subspecific taxa of ''S. rubra'' into 3 to 5 species. Similarly, ''S. rosea'' is not always recognized as a species distinct from ''S. purpurea''. The most commonly recognized species include: Currently, ''S. rubra'' can be described as having six subspecies, though it is sometimes argued that the subspecies should be elevated to species rank in recognition of the species complex that they are a part of. This division would yield ''S. alabamensis'', ''S. gulfensis'', ''S. jonesii'', ''S. rubra'' ''
sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
'', ''S. viatorum'', and ''S. wherryi''. Others have argued that only some of these demand recognition at the species rank.Barthlott, W., S. Porembski, R. Seine, and I. Theisen. (2007). ''The Curious World of Carnivorous Plants''. Portland: Timber Press.


McPherson & Schnell (2011)

Stewart McPherson and Donald Schnell carried out a comprehensive taxonomic revision of the genus in their 2011
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...
, ''Sarraceniaceae of North America''. They recognized the following taxa:McPherson, S. & D. Schnell 2011. ''Sarraceniaceae of North America''. Redfern Natural History Productions Ltd., Poole. *'' Sarracenia alata'' **''S. alata'' var. ''alata''( autonym) ***''S. alata'' var. ''alata'' f. ''viridescens''S.McPherson & D.E.Schnell **''S. alata'' var. ''atrorubra'' S.McPherson & D.E.Schnell **''S. alata'' var. ''cuprea''S.McPherson & D.E.Schnell **''S. alata'' var. ''nigropurpurea'' P.D'Amato ''ex'' S.McPherson & D.E.Schnell **''S. alata'' var. ''ornata''S.McPherson & D.E.Schnell **''S. alata'' var. ''rubrioperculata''S.McPherson & D.E.Schnell *'' Sarracenia flava'' **''S. flava'' var. ''flava''( autonym) ***''S. flava'' var. ''flava'' f. ''viridescens''S.McPherson & D.E.Schnell **''S. flava'' var. ''atropurpurea''(Hort. W.Bull ''ex'' Mast.) Hort. W.Bull ''ex'' W.Robinson **''S. flava'' var. ''cuprea''D.E.Schnell **''S. flava'' var. ''maxima''Hort. W.Bull ''ex'' Mast. **''S. flava'' var. ''ornata''Hort. Bull ''ex'' W.Robinson **''S. flava'' var. ''rubricorpora''D.E.Schnell **''S. flava'' var. ''rugelii''(Shuttlew. ''ex'' A.DC.) Mast. *'' Sarracenia leucophylla'' **''S. leucophylla'' var. ''leucophylla''( autonym) ***''S. leucophylla'' var. ''leucophylla'' f. ''viridescens''S.McPherson & D.E.Schnell **''S. leucophylla'' var. ''alba''(Hort. T.Baines ''ex'' R.Hogg & T.Moore) J.Pietropaolo & P.Pietropaolo ''ex'' S.McPherson & D.E.Schnell *'' Sarracenia minor'' **''S. minor'' var. ''minor''( autonym) ***''S. minor'' var. ''minor'' f. ''viridescens''S.McPherson & D.E.Schnell **''S. minor'' var. ''okefenokeensis''D.E.Schnell *'' Sarracenia oreophila'' **''S. oreophila'' var. ''oreophila''( autonym) **''S. oreophila'' var. ''ornata''S.McPherson & D.E.Schnell *'' Sarracenia psittacina'' **''S. psittacina'' var. ''psittacina''( autonym) ***''S. psittacina'' var. ''psittacina'' f. ''viridescens''S.McPherson & D.E.Schnell **''S. psittacina'' var. ''okefenokeensis''S.McPherson & D.E.Schnell ***''S. psittacina'' var. ''okefenokeensis'' f. ''luteoviridis''S.McPherson & D.E.Schnell *'' Sarracenia purpurea'' **''S. purpurea'' subsp. ''purpurea''( autonym) ***''S. purpurea'' subsp. ''purpurea'' f. ''heterophylla''(Eaton) Fern. **''S. purpurea'' subsp. ''venosa''(Raf.) Wherry ***''S. purpurea'' subsp. ''venosa'' var. ''venosa''( autonym) ****''S. purpurea'' subsp. ''venosa'' var. ''venosa'' f. ''pallidiflora''S.McPherson & D.E.Schnell ***''S. purpurea'' subsp. ''venosa'' var. ''burkii''D.E.Schnell ****''S. purpurea'' subsp. ''venosa'' var. ''burkii'' f. ''luteola''R.L.Hanrahan & J.Miller ***''S. purpurea'' subsp. ''venosa'' var. ''montana''D.E.Schnell & R.O.Determann *'' Sarracenia rubra'' **''S. rubra'' subsp. ''rubra''( autonym) **''S. rubra'' subsp. ''alabamensis''(Case & R.B.Case) S.McPherson & D.E.Schnell **''S. rubra'' subsp. ''gulfensis''D.E.Schnell ***''S. rubra'' subsp. ''gulfensis'' f. ''luteoviridis''S.McPherson & D.E.Schnell **''S. rubra'' subsp. ''jonesii''(Wherry) Wherry ***''S. rubra'' subsp. ''jonesii'' f. ''viridescens''S.McPherson & D.E.Schnell **''S. rubra'' subsp. ''wherryi''(Case & R.B.Case) D.E.Schnell **''S. rubra'' "Incompletely diagnosed taxon from Georgia and South Carolina"( Undescribed, but see note below) Note: The entity McPherson and Schnell referred to as ''S. rubra'' "Incompletely diagnosed taxon from Georgia and South Carolina" has since been established as ''Sarracenia rubra'' subsp. viatorum B.Rice.


Hybrids

''Sarracenia'' species hybridize and produce fertile offspring freely, making proper classification difficult. ''Sarracenia'' hybrids are able to hybridize further, giving the possibility of hundreds of different hybrids that have multiple species in varying amounts in their ancestry. Since many species ranges overlap, natural hybrids are relatively common. As a result, initial classification included many of these hybrids as separate species. A recent census of the number of hybrids and cultivars of ''Sarracenia'' species revealed about 100 unique hybrids and cultivars in cultivation. Many hybrids of ''Sarracenia'' are still commonly referred to by their obsolete species names, particularly in horticulture. These hybrids are all popularly cultivated by carnivorous plant enthusiasts, and there are consequently a huge number of hybrids and cultivars, most bred for showy pitchers. Some of the more common named hybrids include: * ''Sarracenia'' × ''catesbaei'' = ''S. flava'' × ''S. purpurea'' * ''Sarracenia'' × ''moorei'' = ''S. flava'' × ''S. leucophylla'' * ''Sarracenia'' × ''popei'' = ''S. flava'' × ''S. rubra'' * ''Sarracenia'' × ''harperi'' = ''S. flava'' × ''S. minor'' * ''Sarracenia'' × ''alava'' = ''S. flava'' × ''S. alata'' * ''Sarracenia'' × ''mitchelliana'' = ''S. purpurea'' × ''S. leucophylla'' * ''Sarracenia'' × ''exornata'' = ''S. purpurea'' × ''S. alata'' * ''Sarracenia'' × ''chelsonii'' = ''S. purpurea'' × ''S. rubra'' * ''Sarracenia'' × ''swaniana = ''S. purpurea'' × ''S. minor'' * ''Sarracenia'' × ''courtii'' = ''S. purpurea'' × ''S. psittacina'' * ''Sarracenia'' × ''pureophila'' = ''S. purpurea'' × ''S. oreophila'' * ''Sarracenia'' × ''readii'' = ''S. leucophylla'' × ''S. rubra'' * ''Sarracenia'' × '' farnhamii'' = ''S. leucophylla'' × ''S. rubra'' * ''Sarracenia'' × ''excellens'' = ''S. leucophylla'' × ''S. minor'' * ''Sarracenia'' × ''areolata'' = ''S. leucophylla'' × ''S. alata'' * ''Sarracenia'' × ''wrigleyana'' = ''S. leucophylla'' × ''S. psittacina'' * ''Sarracenia'' × ''ahlesii'' = ''S. alata'' × ''S. rubra'' * ''Sarracenia'' × ''rehderi'' = ''S. rubra'' × ''S. minor'' * ''Sarracenia'' × ''gilpini'' = ''S. rubra'' × ''S. psittacina'' * ''Sarracenia'' × ''formosa'' = ''S. minor'' × ''S. psittacina''


Botanical history

''Sarracenia'' were known to Europeans as early as the 16th century, within a century of Christopher Columbus' discovery of the New World. L'Obel included an illustration of '' S. minor'' in his ''Stirpium Adversaria Nova'' in 1576. The first description and plate of a ''Sarracenia'' to show up in botanical literature was published by Carolus Clusius, who received a partial dried specimen of what was later determined to be ''S. purpurea'' subsp. ''purpurea'', publishing it under the name '' Limonium peregrinum''. The exact origins of this specimen remains unknown, as few explorers are known to have collected plant specimens from the range of this subspecies before that time. Cheek and Young suggest that the most likely source is
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expeditions to what is now Quebec between 1534 and 1541. The fragile flowerless specimen that made its way to Clusius 60 years later was enough to excite his interest, but not enough for him to place it among related plants; his closest guess was the wholly unrelated Sea Lavender genus. The name ''Sarracenia'' was first employed by Michel Sarrazin, the Father of Canadian Botany who in the late 17th century sent live specimens of ''S. purpurea'' to the Parisian botanist Joseph Pitton de Tournefort, who thereupon described the species. Linnaeus adopted this name when he published his ''
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 ...
'' (1753), using it for the two known species at the time: ''S. purpurea'' and ''S. flava''. The first successful flowering in culture occurred in 1773. In 1793
William Bartram William Bartram (April 20, 1739 – July 22, 1823) was an American botanist, ornithologist, natural historian and explorer. Bartram was the author of an acclaimed book, now known by the shortened title ''Bartram's Travels'', which chronicled ...
noted in his book about his travels in the southeast U.S. that numerous insects were caught in the pitchers of these plants, but doubted that any benefit could be derived from them. It was not until 1887 that research by
Joseph H. Mellichamp Joseph is a common male given name, derived from the Hebrew Yosef (יוֹסֵף). "Joseph" is used, along with "Josef", mostly in English, French and partially German languages. This spelling is also found as a variant in the languages of the mo ...
proved the carnivorous nature of this genus. This finding was supported by a study by J.S. Hepburn, E.Q. St. John and F.M. Jones in 1920. Extended field surveys and laboratory studies by Edgar Wherry in the 1930s greatly increased the knowledge of this genus, which has further been extended by the more recent works of C. Ritchie Bell (1949–52),
Donald E. Schnell Donald is a masculine given name derived from the Gaelic name ''Dòmhnall''.. This comes from the Proto-Celtic *''Dumno-ualos'' ("world-ruler" or "world-wielder"). The final -''d'' in ''Donald'' is partly derived from a misinterpretation of the ...
(1970–2002),
Frederick W. Case Frederick may refer to: People * Frederick (given name), the name Nobility Anhalt-Harzgerode *Frederick, Prince of Anhalt-Harzgerode (1613–1670) Austria * Frederick I, Duke of Austria (Babenberg), Duke of Austria from 1195 to 1198 * Frederick ...
(1970–2000s), and
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(1980s-2000s).


Cultivation

''Sarracenia'' are considered easy to grow and are widely propagated and cultivated by gardeners and carnivorous plant enthusiasts. Several hybrids between the very hardy ''S. purpurea'' and showy species like '' S. leucophylla'' are becoming common in garden centers in North America and Europe. ''Sarracenia'' require constantly moist-wet, nutrient free acidic soil. This is most often achieved with a potting mix consisting of peat moss mixed with sand or perlite. As their roots are sensitive to nutrients and minerals, only pure water, such as distilled, rain, or
reverse osmosis Reverse osmosis (RO) is a water purification process that uses a partially permeable membrane to separate ions, unwanted molecules and larger particles from drinking water. In reverse osmosis, an applied pressure is used to overcome osmotic pre ...
water, can be used to water them. ''Sarracenia'' prefer sunny conditions during their growing season but require a dormancy period, with decreased light and temperatures, of a few months in the winter.


Propagation

''Sarracenia'' do not self-pollinate and therefore require hand pollination or access to natural pollinators such as bees. ''Sarracenia'' pollen remains potent for several weeks when refrigerated, and so is stored by cultivators and used to pollinate later-flowering species. Given that all ''Sarracenia'' hybrids are fertile and will hybridize further, this characteristic allows cultivators to produce a limitless number of variants through hybridization. The copious seeds store well if kept dry. In climates or seasons that cannot provide the cold, damp period of stratification required by the seeds for germination, growers mimic this condition by placing the seeds in a refrigerator for 2–6 weeks, depending on species. The seeds are sown on the surface of their substrate and germinate when transferred to warmer, bright conditions. ''Sarracenia'' seedlings all look alike for the first two or three years; the plants reach maturity after four or five years. Regular fertilization (twice a month between April and September) with a balanced fertilizer at the rate of 1 teaspoon per gallon (using a 15-16-17 peat-lite or similar fertilizer) will speed their growth and time to maturity. It is advisable to leach regularly with pure water to prevent the buildup of solutes (fertilizer salts) in the soil. Deep water in a potted plant keeps the soil too waterlogged for proper root functioning. Mature ''Sarracenia'' are commonly propagated by division. Their rhizomes extend and produce new crowns of pitchers over the course of a few growing seasons, and cultivators divide and separate the rhizomes during the plant's winter dormancy or early in the growing season. This technique is also used to separate sections of rhizomes which have no pitchers: when re-potted, the section usually generates a new crown of pitchers. A further technique is employed to encourage new crowns to appear which does not involve division of the rhizome: small notches up to 5 mm deep are cut into the top of the rhizome, whereupon a new crown frequently develops at the site of the notch.


AGM cultivars

The following have won the Royal Horticultural Society's
Award of Garden Merit The Award of Garden Merit (AGM) is a long-established annual award for plants by the British Royal Horticultural Society (RHS). It is based on assessment of the plants' performance under UK growing conditions. History The Award of Garden Merit ...
: *''Sarracenia'' 'Anna Carlisle' *''Sarracenia'' 'Colin Clayton' *''Sarracenia'' 'Dixie Lace' *''Sarracenia'' 'Jenny Helen' *''Sarracenia'' 'Juthatip Soper' *''Sarracenia'' 'Vogel' *''Sarracenia'' × ''catesbaei'' 'Violet' *''Sarracenia'' × ''mitchelliana'' 'Victoria Morley' *''Sarracenia'' × ''moorei'' 'Brooks's Hybrid'


See also

* North American Sarracenia Conservancy * Meadowview Biological Research Station


Notes

:a.Since McPherson & Schnell (2011) did not assign these forms to any particular variety, they are to be placed under the autonymous variety according to the botanical code. :b.This combination had been published previously, but was only validated in McPherson and Schnell's 2011 monograph.


References


Further reading

* Schnell, Donald E. 2002. ''Carnivorous Plants of the United States and Canada''. Portland. * Schnell, Stewart McPherson, Donald E. (2011). ''Sarraceniaceae of North America''. Poole: Redfern Natural History Productions. .


External links


''Sarracenia'' Taxonomy
by Barry Rice
''Sarracenia'' – the Pitcher Plants
by the
Botanical Society of America The Botanical Society of America (BSA) represents professional and amateur botanists, researchers, educators and students in over 80 countries of the world. It functions as a United States nonprofit 501(c)(3) membership society. History The soci ...

''Sarracenia'' Growing Guide and Distribution Map
by Tom's Carnivores
Growing ''Sarracenia''
by the International Carnivorous Plant Society
The Inner World of ''Sarracenia''
by the John Innes Centre {{Taxonbar, from=Q161174 Carnivorous plants of North America Rhizomatous plants Ericales genera