Crocus
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''Crocus'' (; plural: crocuses or croci) is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of seasonal
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...
s in the family Iridaceae (iris family) comprising about 100 species of perennials growing from corms. They are low growing plants, whose flower stems remain underground, that bear relatively large white, yellow, orange or purple flowers and then become dormant after flowering. Many are cultivated for their flowers, appearing in autumn, winter, or spring. The flowers close at night and in overcast weather conditions. The crocus has been known throughout recorded history, mainly as the source of
saffron Saffron () is a spice derived from the flower of ''Crocus sativus'', commonly known as the "saffron crocus". The vivid crimson stigma (botany), stigma and stigma (botany)#style, styles, called threads, are collected and dried for use mainly ...
. Saffron is obtained from the dried stigma of ''
Crocus sativus ''Crocus sativus'', commonly known as saffron crocus or autumn crocus, is a species of flowering plant in the iris family Iridaceae. A cormous autumn-flowering cultivated perennial, unknown in the wild, it is best known for the culinary use of ...
'', an autumn-blooming
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
. It is valued as a
spice A spice is a seed, fruit, root, bark, or other plant substance primarily used for flavoring or coloring food. Spices are distinguished from herbs, which are the leaves, flowers, or stems of plants used for flavoring or as a garnish. Spices a ...
and
dye A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution and ...
stuff, and is one of the most expensive spices in the world. Iran is the center of saffron production. Crocuses are
native Native may refer to: People * Jus soli, citizenship by right of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Native Americans (disambiguation) In arts and entert ...
to woodland, scrub, and meadows from sea level to alpine tundra from the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on ...
, through North Africa, central and southern
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
, the islands of the Aegean, the
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
and across
Central Asia Central Asia, also known as Middle Asia, is a region of Asia that stretches from the Caspian Sea in the west to western China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north. It includes the fo ...
to
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwes ...
in western
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
. Crocuses may be propagated from seed or from daughter cormels formed on the corm, that eventually produce mature plants. They arrived in Europe from Turkey in the 16th century and became valued as an ornamental flowering plant.


Description

;General ''Crocus'' display the general characteristics of family Iridaceae, which include basal cauline (arising from the aerial stem) leaves that sheath the stem base, hermaphrodite flowers that are relatively large and showy, the perianth petaloid with 2 whorls of 3 tepals each and septal nectaries. The flowers have 3 stamens and a gynoecium of 3 united carpels and an inferior ovary, 3 locules and axile placentation with fruit that is a loculicidal capsule. ''Crocus'' is an acaulescent (lacking a visible lower stem above ground) diminutive seasonal cormous (growing from corms)
herbaceous perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
geophytic genus. The corms are symmetrical and globose or oblate (round in shape with flatted tops and bottoms), and are covered with tunic leaves that are fibrous, membranous or coriaceous (leathery). The corms produce fibrous roots, and contractile roots which adjust the corms depth in the soil, which maybe pulled as deep as into the soil.The roots appear randomly from the lower part of the corm, but in a few species, from a basal ridge. ;Leaves Plants produce several basal linear bifacial green leaves that arise from the corms. These are adaxially (upper surface facing axis) flat or channelled with pale median stripes, while the opposite (abaxial) surface is strongly keeled, with two grooves on either side. The leaves have a distinctive shape in cross-section, being boat-shaped with two lateral arms with margins recurved inwardly towards the central riged keel, forming the sides of the "boat". The keel may be square or rectangular, but is lacking in ''C. carpetanus''. The pale central stripe is caused by parenchymatous cells which lack chloroplasts and may contain air spaces. The leaves are from wide and long. The leaf-like
bracts 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 ...
are membranous, while the smaller bracteoles are either membranous or absent. The leaf bases are surrounded by up to 5 membranous sheaths called cataphylls, a specialised leaf. The bases of the cataphylls form the corm tunic, and their number varies from 3 to 6, and enclose the true leaves (euphylls), bracts, bracteoles and flowering stalk. ;Flowers The number of peduncles (
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 ...
stems) vary from one to several and remain underground, emerging only at the fruiting stage, bearing flowers that are solitary or several, so that a true scape is absent. The flowers are
pedicellate In botany, a pedicel is a stem that attaches a single flower to the inflorescence. Such inflorescences are described as ''pedicellate''. Description Pedicel refers to a structure connecting a single flower to its inflorescence. In the absenc ...
(attached to the peduncle by a short subterranean pedicel stalk). The pedicel is sometimes subtended (below pedicel) by a membranous, sheathing
prophyll 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 ...
(leaf-like structure). The showy, salver to cup-shaped, single or clustered actinomorphic flowers taper off into a narrow tube; the flowers emerge from the ground, and can be white, yellow, lilac to dark purple, or
variegated Variegation is the appearance of differently coloured zones in the leaves and sometimes the stems and fruit of plants. Species with variegated individuals are sometimes found in the understory of tropical rainforests, and this habitat is the ...
in cultivars. The flower tube is long, cylindrical and slender, expanding apically. The floral tube is long and narrow with 6 lobes in 2 whorls. The perianth is 3+3 (3 sepals+3petals) and
gamophyllous 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 o ...
(with fused segments). The tepal whorls are similar, equal or subequal with a smaller inner whorl, and cupped to outspread. The bracts are membranous, but the inner ones are sometimes lacking. The 3 stamens are erect and linear and inserted in the throat of the perianth tube, with anthers shorter than the filaments. Pollen grains are inaperturate (apertures absent) but sometimes spiraperturate (spiral shaped). Each flower has a single style which is
exserted 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 o ...
(projecting beyond the
corolla Corolla may refer to: *Corolla (botany), the petals of a flower, considered as a unit *Toyota Corolla, an automobile model name *Corolla (headgear) A ''corolla'' is an ancient headdress in the form of a small circlet or crown.ovary has 3 carpels with axile placentation. It remains underground, and as the
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 ripen, the pedicel (stem of the flower) grows longer - so 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 ...
is above the soil surface. ;Fruit and seed The fruit is a small membranous capsule, ellipsoid or oblong-ellipsoid in shape and the many
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 globose to ellipsoid. The seed surface is highly variable, including papillate (covered in small protuberances),
digitiform 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 o ...
(finger-like) and other epidermal cell types. In some species the seeds are
arillate An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...
, with fleshy appendages. Crocus seeds have both inner and outer integuments and in some species the outer epidermis may display long papillae.
Embryo-sac A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sex ...
development is Polygonum type. Dehiscence (splitting of the capsule to release the seed) is of the loculicidal type in which it splits through the wall of the locules leaving the septa that separate them intact. ;Karyology ''Crocus'' has extensive aneuploidy (abnormal number of chromosomes), with some uncertainty as to the base number of chromosomes. The chromosome numbers shows extreme variability,ranging from 2n=6 to 2n=70 even within a single species. ;Phytochemistry The Iridaceae contain a wide range of
phenolic compounds In organic chemistry, phenols, sometimes called phenolics, are a class of chemical compounds consisting of one or more hydroxyl groups (— O H) bonded directly to an aromatic hydrocarbon group. The simplest is phenol, . Phenolic compounds are c ...
. However, 6-Hydroxyflavones are found only in ''Crocus'', which is also characterised by the presence of
crocin Crocin is a carotenoid chemical compound that is found in the flowers of crocus and gardenia. Crocin is the chemical primarily responsible for the color of saffron. Chemically, crocin is the diester formed from the disaccharide gentiobiose and ...
s, water-soluble yellow carotenoids, in the floral tissues. Crocin is a
diester In chemistry, an ester is a compound derived from an oxoacid (organic or inorganic) in which at least one hydroxyl group () is replaced by an alkoxy group (), as in the substitution reaction of a carboxylic acid and an alcohol. Glycerides are ...
of
crocetin Crocetin is a natural apocarotenoid dicarboxylic acid that is found in the crocus flower together with its glycoside, crocin, and ''Gardenia jasminoides'' fruits. It forms brick red crystals with a melting point of 285°C. The chemical structur ...
, responsible for the colour of the styles and stigma of ''C. sativus'', and hence saffron. A few species contain
mangiferin Mangiferin is a glucosylxanthone (xanthonoid). This molecule is a glucoside of norathyriol. Natural occurrences Mangiferin was first isolated from the leaves and bark of '' Mangifera indica'' (the mango tree). It can also be extracted from ...
, a glucosylxanthone. While the flowers may vary dramatically between species, there is little variation in the leaves, but sufficient variability in corm tunics that they may be used as an aid in differentiating taxa. :


Taxonomy


History

The crocus was well known to the ancients, being described at least as early as Theophrastus (c. 371c. 287 BC), and was introduced into Britain by the Romans, where the saffron crocus was used as a
dyestuff A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution and ...
. It was reintroduced into Western Europe by the Crusaders. The crocus is mentioned in mediaeval and later herbals, one of the earliest being the 14th century '' Tractatus de Herbis.'' William Turner (1548) states that the crocus is referred to as saffron in English, implying that only ''C. sativus'' was known at that time. However, by 1597 John Gerard writes of "sundry sorts" and uses the term saffron and crocus as interchangeable. He included both spring and autumn flowering crocus, but distinguished Wild Saffron (Crocus) from Meadow Saaffron (Colchicum). He described eleven forms. Some of his specimens were obtained from Clusius. In the following century, John Parkinson in a more detailed account was more careful to include separate chapters for ''Colchicum'', with the common name of meadow saffron, from ''Crocus'' or saffron. Parkinson (1656) states that there are "divers sorts of saffrons" describing 27 spring flowering plants and 4 autumn flowering ones, pointing out that only one of those was the true saffron crocus, which he called ''Crocus verus sativus autumnalis''. Similar accounts are found in continental European herbals, including those of l'Obel in
Flanders Flanders (, ; Dutch: ''Vlaanderen'' ) is the Flemish-speaking northern portion of Belgium and one of the communities, regions and language areas of Belgium. However, there are several overlapping definitions, including ones related to cultu ...
(1576) and Besler's ''
Hortus Eystettensis ''Hortus Eystettensis'' (Garden of Eichstätt) is the short title of a codex produced by Basilius Besler, a Nuremberg apothecary and botanist, in 1613 describing the plants of the garden of the Prince-Bishop of Eichstätt in Bavaria. The Ren ...
'' in
Bavaria Bavaria ( ; ), officially the Free State of Bavaria (german: Freistaat Bayern, link=no ), is a state in the south-east of Germany. With an area of , Bavaria is the largest German state by land area, comprising roughly a fifth of the total l ...
(1613). The genus ''Crocus'' was first formally described by Linnaeus in 1753, with three taxa, and two species, ''C. sativus'' (
type species In 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 species that contains the biological type specim ...
), var. ''officinalis'' (now treated as a synonym of ''C. sativus'') and var. ''vernus'' (now ''C. vernus'') and ''C. bulbocodium'' (now ''
Romulea bulbocodium ''Romulea bulbocodium'' is one of the best-known species from the genus '' Romulea''. The plant, a member of the family Iridaceae, is native to the Mediterranean region (southern Europe, northern Africa, the Middle East) and Sudan.
''). Thus Linnaeus recognised two taxa that are accepted as separate species in modern classifications, one vernal and one autumnal crocus, but incorrectly assumed they were only varieties of a single species, while his second species was actually from a closely related genus that was only recognised later (1772). However a subsequent re-examination of Linnaeus's specimens suggested the presence of several different species that he did not recognise as being separate. Linnaeus' system, based on sexual characteristics, ''Crocus'' was classified as Triandra Monogynia (Three stamens, Single pistil). Linnaeus's system was supplanted by the "natural" system which used a hierarchy of
taxonomic ranks In biological classification, taxonomic rank is the relative level of a group of organisms (a taxon) in an ancestral or hereditary hierarchy. A common system consists of species, genus, family, order, class, phylum, kingdom, domain. While ...
based on weighting of the importance of structural characteristics of the plant. Jussieu (1789) placed the genus Crocus in his ''Ordo'' ( family) Irides or ''Les iris'', as a member of the class ''Stamina epigyna'' (stamens inserted above the ovary) as part of the
monocotyledon Monocotyledons (), commonly referred to as monocots, ( Lilianae '' sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one ...
s, the first level of the division of the
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...
s. One of the first monographs of the genus appeared in 1809, by Haworth, followed in 1829 by that of Sabine, and Herbert in 1847. In 1853, Lindley continued the placement of ''Crocus'' as one of 53 genera in Iridaceae, which he included in a higher order of monocotyledons, the Narcissales. Baker published a monograph on the genus in 1874, adopting a very different schema to that of Herbert. In 1883, Bentham and Hooker described the Irideae (Iridaceae) as having more than 700 species, and divided it into 3 tribes and further into subtribes.
Tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...
Sysyrinchieae as having 2 subtribes, including Ixieae. The latter was circumscribed with four genera, ''Crocus'', '' Syringodea'', ''Galaxia'' ('' Moraea'') and ''
Romulea ''Romulea'' is a genus of flowering plants in the family Iridaceae, first described as a genus in 1772. It is distributed in Europe, the Mediterranean, the Arabian Peninsula, and Africa.Maw (1886), which forms the basis of modern understanding of the genus. Maw built oin the work of Herbert, rejecting Baker's classification. The availability of molecular phylogenetic methods in the late twentieth century has shown that the Iridaceae properly belong within the order Asparagales.


Botanical illustration

The scientific study of the genus in the late eighteenth century was accompanied by detailed descriptions with Botanical illustrations, such as those of William Curtis (1787) and Sims (1803), that appeared in Curtis' Botanical Magazine, with illustrations by Sydenham Edwards. Other illustrations are found in monographs such as those of Haworth (1809) and Sabine (1830), illustrated by Charles John Robertson. The largest collection is found in the most comprehensive monograph, that of Maw (1886). other sources include the portfolios of plates, such as the survey of the plants of France by Masclef (1891). At that time only ''C. sativus'' and ''C. vernus'' were included in the
Flora Flora is all the plant life present in a particular region or time, generally the naturally occurring ( indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or '' skin flora''. ...
of France.


Phylogeny

The genus ''Crocus'' belongs to the
monocot Monocotyledons (), commonly referred to as monocots, (Lilianae '' sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one ...
family Iridaceae (iris family), specifically the large subfamily
Crocoideae Crocoideae is one of the major subfamilies in the family Iridaceae. It contains plants which are widely distributed in the Old World, mainly in Africa, but there are species like some members of the genera '' Romulea'' and ''Gladiolus'' which are ...
. Within that subfamily, crocus is placed on the
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...
, Ixieae (synonym Croceae), one of five. The Ixieae are then subdivided into subtribes, with the genera ''Crocus'', ''
Romulea ''Romulea'' is a genus of flowering plants in the family Iridaceae, first described as a genus in 1772. It is distributed in Europe, the Mediterranean, the Arabian Peninsula, and Africa.Syringodea'' forming subtribe Romuleinae. The Romuleinae have been characterised within the Ixieae by progressively reduced aerial stems. solitary flowers on the stem branches and woody tunics on the corms. They also often have divided style branches. However ''Crocus'' corm tunics are fibrous and membranous rather than woody as in ''Syringodea'', also ''Crocus'' has a ridged and often keeled abaxial leaf surface, while that of ''Syringodea'' is rounded, and the midline adaxial translucency of ''Crocus'' is lacking in ''Syringodea''. ''
Romulea ''Romulea'' is a genus of flowering plants in the family Iridaceae, first described as a genus in 1772. It is distributed in Europe, the Mediterranean, the Arabian Peninsula, and Africa.sister group to ''Syringodea'', the two genera forming a sister group to ''Romulea''.


Subdivision

The genus Crocus consists of about 200 accepted species, which continue to increase, and has undergone a large number of taxonomic classifications. The genus has often been divided into
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 ...
, beginning with that of Haworth (1809) who described two sections based on the presence or absence of hairs in the throat of the flower, while Sabine was the first to realise the importance of the presence or absence of a basal spathe (prophyll) in dividing the genus into two sections, a practice followed by Herbert However Sabine's practice of using trinomials for varieties such as ''C. sulphureus concolor'' is no longer accepted, although Herbert somewhat similarly used varieties and subvarieties, eg ''C. vernus'' var.1 ''Communis'' subvar. 1. ''Obovatus''. Herbert also used geographical distribution as a basis of classification. By the late 19th century Maw (1886), following Herbert, subdivided the genus into two divisions, the ''Involucrati'' and the ''Nudiflori'', and then further divided it into six sections and lastly by flowering times (spring or autumn). Although rejecting the concept of subvarieties, he placed even more emphasis on geography. The most widely accepted system, that proposed by
Brian Mathew Brian Frederick Mathew MBE, VMH is a British botanist, born in the village of Limpsfield, Surrey, England. His particular area of expertise is bulbous plants, particularly ornamental bulbous plants, although he has contributed to other fields ...
in 1982 was based on Maw's system, but with less emphasis on flowering times. This mainly depended on three character states: * the presence or absence of a prophyll (a basal
spathe 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 ...
); * the aspect of the style; * the corm tunic. and included 81 species, however, one of these, ''Crocus medius'' was later recognized as a synonym of ''
Crocus nudiflorus ''Crocus nudiflorus'' is a species of flowering plant in the genus ''Crocus'' of the family Iridaceae. It is an autumn-flowering, dwarf, deciduous perennial found in western Europe from southwestern France to Spain. It has been cultivated since ...
''. The genus, as described by Mathew, consisted of two subgenera, ''Crocirus'' (monotypic for '' Crocus banaticus'') and ''Crocus'' including the remainder of the species, based on whether the anthers were introrse or extrorse ( dehiscence directed towards or away from centre of flower) respectively. Subgenus ''Crocus'' was then divided into two sections, ''Crocus'' and ''Nudiscapus'', based on the presence or absence of the prophyll. Each section was then further divided into six series of ''Crocus'' and nine of ''Nudiscapus''. These series were defined by the division of the style, the corm tunic, flowering time, leaf structure, presence of a bracteole and anther colour. Mathew also introduced the concept of subspecies, including 50 in all, by giving similar but different forms subspecies status if geographically separated, resulting in about 140 distinct taxa. The seven species and ten subspecies discovered since then have been integrated into revisions of this classification, though new species continue to be described, leading to estimates of at least 200 species.


Speciation

Crocus populations have extremely high infra-specific variability with a very diverse spectrum of morphological and phenotypical varieties, while many individual specimens from different species may closely resemble each other. Based on such morphological differences between isolated populations many new species have been named, but without a definition of new species based on molecular and/or karyological information, species can not be confirmed, creating difficulties in determining speciation and hence the exact number of species. The situation is even more complex once hybridisation (combination of taxa) and introgression (transfer of genetic material) are considered.


Molecular phylogeny

The availability of molecular phylogeny methods revealed problems with the traditional systems based on morphology alone. The first analysis of the complete genus was carried out by Mathew and colleagues in 2008 using nucleotide sequences from plastid regions. In particular, the DNA data suggest there are no grounds for isolating ''C. banaticus'' in its own subgenus ''Crociris'', though it is a unique species in the genus. Because it has a prophyll at the base of the pedicel, it therefore would fall within section ''Crocus'', although its exact relationship to the rest of the subgenus remains unclear. Of the 15 series in the Mathew scheme, only seven were monophyletic, and in particular the largest series, ''Biflori'' and ''Reticulati'', which include a third of all species, were non-monophyletic. Another anomalous species, ''C. baytopiorum'', should now be placed in a series of its own, series ''Baytopi''. ''C. gargaricus'' subsp. ''herbertii'' has been raised to species status, as ''C. herbertii''. The autumn-flowering ''C. longiflorus'', the
type species In 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 species that contains the biological type specim ...
of series ''Longiflori'' (long regarded by Mathew as "a disparate assemblage"), appeared to lie within series ''Verni''. In addition, the position of ''C. malyi'' was currently unclear. DNA analysis and morphological studies suggest further that series ''Reticulati'', ''Biflori'' and ''Speciosi'' are "probably inseparable", ''C. adanensis'' and ''C. caspius'' should probably be removed from ''Biflori'', ''C. adanensis'' falls in a clade with ''C. paschei'' as a sister group to the species of series ''Flavi'' and ''C. caspius'' appears to be sister to the species of series ''Orientales''. The study showed "no support for a system of sections as currently defined", although, despite the many inconsistencies between Mathew's 1982 classification and the current hypothesis, "the main assignment of species to the sections and series of that system is actually supported". The authors state, "further studies are required before any firm decisions about a hierarchical system of classification can be considered" and conclude "future re-classification is likely to involve all infrageneric levels, subgenera, sections and series". A further study, using the
internal transcribed spacer Internal transcribed spacer (ITS) is the spacer DNA situated between the small-subunit ribosomal RNA (rRNA) and large-subunit rRNA genes in the chromosome or the corresponding transcribed region in the polycistronic rRNA precursor transcript. ...
region (ITS) of the nuclear ribosomal DNA (rDNA), together with a chloroplast marker, broadly confirmed these findings. ''Crocus'' forms a monophyletic clade, with a basal polytomy of four subclades. The first clade (A) corresponding to section ''Crocus'', but including ''C. sieberi'' and several closely related species (originally included in section ''Nudiscapus'' series ''Reticulati''). The remaining three clades (B-D) include all the remaining species of section ''Nudiscapus''. Of these, B and C are small, corresponding to series ''Orientales'' and ''Carpetani'' respectively, with all remaining series in the large D clade. The exception is ''C. caspius'', originally in series ''Biflori'', which segregates in clade B. Thus, although division of the genus into two sections is well supported, no single morphological character defines these two groups. The ''C. sieberi'' group are assumed to have lost their prophyll secondarily. Of the series, eight could be shown to be monophyletic; ''Crocus'', ''Kotschyani'' and ''Scardici'' (section ''Crocus'') and ''Aleppici'', ''Carpetani'', ''Laevigati'', ''Orientalis'' and ''Speciosi'' (section ''Nudiscapus''). Flowering season did not correspond to molecular groupings and nor did any of the previously used morphological characteristics, indicating a high degree of homoplasy, in which traits are gained or lost independently in different lineages. The remainder of the series could not be supported as natural groupings. Mathew's concept of subspecies status within ''C. biflorus'' could not be supported, each being considered a separate species, resulting in the genus having at least 150 species. A more detailed molecular and morphological study of series ''Verni'' (section ''Crocus'') allowed it to be better characterised and circumscribed, as well as the closely related series ''Longiflori''. Series ''Verni'' ''sensu'' Mathew was found to consist of two groups, the first being ''C. vernus'' ''sensu'' Mathew and the other consisting of ''C. etruscus'', ''ilvensis'', ''kosaninii'' and ''longiflorus''. The taxonomic status of ''C. vernus'' had been uncertain for some time, given the observation that the name was more properly applied to ''C. albiflorus'', requiring a new designation of ''C. neapolitanus'' for those previously known as ''C. vernus''. Subsequently ''C. vernus'' was split into 5 separate species. The incorporation of ''C. longiflorus'' into series ''Verni'' resulted in making series ''Longiflori'' no longer a legitimate taxonomic unit. In section ''Nudiscapus'', series ''Reticulati'' was polyphyletic with species intermingled with series ''Biflori'' and ''Speciosi'', requiring a recircumscription, confining ''Reticulati'' to 8 species, to obtain monophyly. Among the thereby displaced species, are a number of very closely related taxa, referred to as the ''Crocus sieberi'' aggregate, which has been proposed as a new series ''Sieberi''. Other new series, such as ''Isauri'' and ''Lyciotauri'', continue to be created out of the ''Biflori'' series. Mathew's circumscription of ''Crocus'' introduced the rank of subspecies, of which the largest number (14) were those of ''Crocus biflorus'' Miller, the type species of series ''Biflori'', a number which continued to grow. Molecular methods identified these as a polyphyletic assemblage rather than closely related subordinate infraspecific taxa. This necessitated a complete taxonomic revision of series ''Biflori'', elevating each subspecies to species status. A similar issue occurs with ''C. reticulatus'' ''sensu'' Mathew, who created two subspecies, resulting in 9 newly defined species.


Sections and species

The classification of Brian Mathew (1982), as amended in 2009 divides the genus into two sections, further divided by series. The number of series, continues to evolve. * Section ''Crocus'' B.Mathew Species with a basal
prophyll 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 ...
.
Type species In 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 species that contains the biological type specim ...
'' C. sativus'' L. : 6 series * Section ''Nudiscapus'' B.Mathew Species without a basal prophyll.
Type species In 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 species that contains the biological type specim ...
'' C. reticulatus'' Stev. ex Adams : 9 series


Similarly named species

Some crocus species, known as "autumn crocus", flower in late summer and autumn, during (autumnal) rains, after summer's heat and drought, the name autumn crocus is also often used as a common name for ''
Colchicum ''Colchicum'' ( or ) is a genus of perennial flowering plants containing around 160 species which grow from bulb-like corms. It is a member of the botanical family Colchicaceae, and is native to West Asia, Europe, parts of the Mediterranean coa ...
'', which is in its own family ( Colchicaceae) in the lily order Liliales. ''Colchicum'' are also known as meadow saffron, though unlike true
saffron Saffron () is a spice derived from the flower of ''Crocus sativus'', commonly known as the "saffron crocus". The vivid crimson stigma (botany), stigma and stigma (botany)#style, styles, called threads, are collected and dried for use mainly ...
, the plants are toxic. ''Crocus'' species have three stamens while ''Colchicum'' species have six stamens. In addition, crocus have one style, while ''Colchicum'' have three. Some Pulsatilla species are also called "prairie crocus" (previously '' Anemone patens)'' or "wild crocus", but they belong to the buttercup family (
Ranunculaceae Ranunculaceae (buttercup or crowfoot family; Latin "little frog", from "frog") is a family of over 2,000 known species of flowering plants in 43 genera, distributed worldwide. The largest genera are ''Ranunculus'' (600 species), ''Delphinium' ...
). Pulsatilla species, which are commonly called pasqueflowers, in contrast to crocus's, have rhizomes, the foliage is covered with long soft hairs, and the flowers are produced on above-ground stems.


Etymology

"Crocus", the name of the genus is Late Middle English (late 14th century) and also denotes saffron. It is derived via Latin ''crocus ''from the Greek κρόκος (''krokos''). This, in turn, is probably a loan word from a
Semitic language The Semitic languages are a branch of the Afroasiatic language family. They are spoken by more than 330 million people across much of West Asia, the Horn of Africa, and latterly North Africa, Malta, West Africa, Chad, and in large immigrant ...
, related to Hebrew כרכום ''karkōm'',
Aramaic The Aramaic languages, short Aramaic ( syc, ܐܪܡܝܐ, Arāmāyā; oar, 𐤀𐤓𐤌𐤉𐤀; arc, 𐡀𐡓𐡌𐡉𐡀; tmr, אֲרָמִית), are a language family containing many varieties (languages and dialects) that originated i ...
ܟܟܘܪܟܟܡܡܐ ''kurkama'', and
Arabic Arabic (, ' ; , ' or ) is a Semitic language spoken primarily across the Arab world.Semitic languages: an international handbook / edited by Stefan Weninger; in collaboration with Geoffrey Khan, Michael P. Streck, Janet C. E.Watson; Walter ...
''kurkum'', meaning saffron (''Crocus sativus''), "saffron yellow" or turmeric (see '' Curcuma''), another yellow dye. The word ultimately traces back to the
Sanskrit Sanskrit (; attributively , ; nominally , , ) is a classical language belonging to the Indo-Aryan languages, Indo-Aryan branch of the Indo-European languages. It arose in South Asia after its predecessor languages had Trans-cultural diffusion ...
''kunkumam'' () for "saffron". The English name is a learned 16th-century adoption from the Latin, but Old English already had ''croh'' for saffron, introduced by the Romans.


Distribution and habitat

Crocuses are distributed from the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on ...
, from the
Iberian peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, def ...
and North Africa, through central and southern
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
, the islands of the Aegean, the
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
and across
central Central is an adjective usually referring to being in the center of some place or (mathematical) object. Central may also refer to: Directions and generalised locations * Central Africa, a region in the centre of Africa continent, also known a ...
and southwest Asia to
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwes ...
in western
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
, but most species are restricted to Turkey and Asia Minor and the
Balkans The Balkans ( ), also known as the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throughout the who ...
, with the Balkan Peninsula having the largest number of species ( with at least 31), forming the centre of diversity, however they are widely introduced. The distribution of species is described over five contiguous areas from west to east (''see map''). Habitats range from sea level as high as subalpine altitudes, and in a wide range of habitats from woodlands to meadows and deserts, often on stony mountain slopes with good drainage. The majority of species are native to areas with cold winters and hot summers with little rain, and active growth is typically from fall to mid-spring. The natural habitats of crocus species are threatened by human activities, including urbanization, industrialization, and other land disturbances and recreational uses. They are negatively impacted by uncontrolled gathering and heavy grazing by livestock. File:Crocus alatavicus, 2011.jpg, ''Crocus alatavicus'' File:Crocus aleppicus 1.JPG, ''Crocus aleppicus'' File:Crocus ancyrensis002.jpg, ''Crocus ancyrensis'' File:Crocus banaticus.jpg, ''Crocus banaticus'' File:Crocus biflorus 03.jpg, ''Crocus biflorus'' File:Crocus cancellatus04.jpg, ''Crocus cancellatus'' File:Crocus carpetanus.jpg, ''Crocus carpetanus'' File:Crocus cartwrightianus 'Albus'02.jpg, ''Crocus cartwrightianus'' 'Albus' File:Crocus caspius.jpg, ''Crocus caspius'' File:Crocus chrysanthus 'Zwanenburg Bronze'.jpg, ''Crocus chrysanthus''
'Zwanenburg Bronze' File:Albertacce Crocus corsicus.jpg, ''Crocus corsicus'' File:Crocus etruscus02.JPG, ''Crocus etruscus'' 'Zwanenburg' File:Crocus flavus ssp flavus 04.jpg, '' Crocus flavus'' File:Crocus goulimyi3.jpg, ''Crocus goulimyi'' File:Crocus graveolens, Çukurova University Campus, Adana, Turkey - 20060109.jpg, ''Crocus graveolens'' File:Crocus HyemalisTavor1.JPG, ''Crocus hyemalis'' File:Crocus imperati De Jager group 01.JPG, ''Crocus imperati'' 'De Jager' File:Crocus kotschyanus2.jpg, ''Crocus kotschyanus'' File:Crocus laevigatus Fontenayi 03.JPG, ''Crocus laevigatus'' 'Fontenayi' File:Crocus longiflorus5.jpg, ''Crocus longiflorus'' File:Crocus malyi sveti 1.JPG, ''Crocus malyi'' File:Crocus minimus02.jpg, ''Crocus minimus'' File:Crocus nevadensis subsp marcetii FlowerCloseup DehesaBoyaldePuertollano.jpg, ''Crocus nevadensis'' File:Crocus nudiflorus4.jpg, ''Crocus nudiflorus'' File:Crocus olivieri 01.JPG, ''Crocus olivieri'' File:Crocus pallasii 2.jpg, ''Crocus pallasii'' File:Crocus pulchellus02.jpg, ''Crocus pulchellus'' File:Crocus serotinus clusii flower.jpg, ''Crocus serotinus'' subsp. ''clusii'' File:Flors alpines al cim de la muntanya de Parcent.jpg, ''Crocus serotinus'' subsp. ''salzmannii'' File:W 7592.JPG, ''Crocus scharojanii'' File:Crocus tournefortii Flowers.jpg, ''Crocus tournefortii'' File:Crocus at north of the montagne sainte Victoire by JM Rosier 1.JPG, ''Crocus versicolor''


Ecology

The life cycle of ''Crocus'' species begins with the
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 ...
,
germinating Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fern ...
to a
seedling A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (emb ...
, and a mature plant in 3–5 years, however seeds may remain dormant in the soil for several years. The germination stages was first described and illustrated by Maw in his 1886 monograph. In its first year, the crocus produces only a single leaf and creates a corm covered by a thin tunic, about 5–8 mm in size, dependent on the species. In the northern hemisphere, the autumnal crocuses flower between September and November. The vernal (spring) crocuses flowering time depends both on climate and habitat, but is usually mid-winter to spring. Leaves may be synanthous (produced during flowering) or hysteranthous (when the flowers wither away). In the summer, with hot and dry conditions the plant becomes dormant, with all the above ground parts dying back. Colder temperatures in winter then activate the corms. Propagation occurs sexually by seed and asexually by small corms, called cormels or cormlets, produced in the axils of the corms (between tunic scales and body of corm). As the fruit capsule ripens, it emerges from the soil at the base of the flowering stem before
dehiscing Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part; structures that ...
(splitting open) and releasing the seeds. Seed dispersal may be enhanced by ants, at least in species with
arillate An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...
seeds. At night and in overcast weather, the perianth closes. The ovary produces nectar which attracts
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 monophyletic lineage within the superfami ...
s (particularly female bumblebees) and Lepidoptera.


Pests and diseases

Cultivated plants may have their corms consumed by mice and other rodents, including voles, squirrels, and chipmunks; they are also attacked by mildew, gray mold, botrytis, and fusarium rot; root rot may also occur, caused by ''Stromatinia gladioli'' and '' Pythium'' species - the nematode ''
Pratylenchus penetrans ''Pratylenchus penetrans'' is a species of nematode in the genus ''Pratylenchus'', the lesion nematodes. It occurs in temperate regions worldwide, regions between the subtropics and the polar circles. It is an animal that inhabits the roots of ...
'' may also cause root rot. Viruses that are known to infect ''Crocus'' spp include:
Potyvirus ''Potyvirus'' is a genus of positive-strand RNA viruses in the family ''Potyviridae''. Plants serve as natural hosts. The genus is named after member virus ''potato virus Y''. Potyviruses account for about thirty percent of the currently known ...
es, especially bean yellow mosaic virus and also tobacco rattle virus, tobaccos necrosis virus, and
cucumber mosaic virus ''Cucumber mosaic virus'' (CMV) is a plant pathogenic virus in the family ''Bromoviridae''. This virus has a worldwide distribution and a very wide host range, having the reputation of the widest host range of any known plant virus. It can be tra ...
. The foliage may experience rot, rust, and scab diseases and be fed upon by aphids, mites, snails, and slugs. The foliage is eaten by hares, rabbits, and deer; the flowers are sometimes removed by birds, including crows, jackdaws, and magpies.


Cultivation


Saffron

The economic importance of the genus is largely dependent on the single species, ''
Crocus sativus ''Crocus sativus'', commonly known as saffron crocus or autumn crocus, is a species of flowering plant in the iris family Iridaceae. A cormous autumn-flowering cultivated perennial, unknown in the wild, it is best known for the culinary use of ...
'', now known only in cultivation. ''C. sativus'' is grown for the production of
saffron Saffron () is a spice derived from the flower of ''Crocus sativus'', commonly known as the "saffron crocus". The vivid crimson stigma (botany), stigma and stigma (botany)#style, styles, called threads, are collected and dried for use mainly ...
, an orange-red derivative of its dried stigma, and among the most expensive spices in the world. The estimated worldwide production of ''C. sativus'' plants is 205 tons. About 180,000 stigmas from 60,000 flowers are required to produce saffron, which sells for about $10,000US (2018). Modern saffron production is widely cultivated in Kashmir, Iran, Turkey and the Mediterranean from Spain to Asia Minor. An important center is the
eponym An eponym is a person, a place, or a thing after whom or which someone or something is, or is believed to be, named. The adjectives which are derived from the word eponym include ''eponymous'' and ''eponymic''. Usage of the word The term ''epon ...
ous town of Krokos, in the Kozani region of Greece. The saffron product, Krokos Kazanis is a PDO ( Protected Designation of Origin). production is largely indigenous and Iran accounts for 65% of global production, covering 72,162 ha. Saffron is thought to have been used in embalming in Ancient Egypt. it is mentioned in the Old testament, in the Song of Songs as a precious spice and has featured as a dye and fragrance throughout written history, with mention in The Iliad. Cultivation and harvesting of ''C. sativus'' for saffron was first documented in the Mediterranean, notably on the island of
Crete Crete ( el, Κρήτη, translit=, Modern: , Ancient: ) is the largest and most populous of the Greek islands, the 88th largest island in the world and the fifth largest island in the Mediterranean Sea, after Sicily, Sardinia, Cyprus, ...
. Frescos showing them are found there at the
Bronze Age The Bronze Age is a historic period, lasting approximately from 3300 BC to 1200 BC, characterized by the use of bronze, the presence of writing in some areas, and other early features of urban civilization. The Bronze Age is the second pri ...
Minoan site of Knossos, as well as from the comparably aged Akrotiri site on the Aegean island of Santorini, and formed an important part of the Minoan economy and culture and had both a sacred role and use as a
psychoactive drug A psychoactive drug, psychopharmaceutical, psychoactive agent or psychotropic drug is a chemical substance, that changes functions of the nervous system, and results in alterations in perception, mood, consciousness, cognition or behavior. ...
and food additive. Women still gather crocuses in the Akrotiri region.


Horticulture and floriculture

Crocuses were described in Turkish gardens in the early sixteenth century, gathered from the far reaches of the
Ottoman Empire The Ottoman Empire, * ; is an archaic version. The definite article forms and were synonymous * and el, Оθωμανική Αυτοκρατορία, Othōmanikē Avtokratoria, label=none * info page on book at Martin Luther University ...
, where they were seen by visiting European botanists and explorers, among the first of whom was Pierre Belon who arrived in Constantinople in 1547. The first crocus seen in the
Netherlands ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...
, where crocus species were not native, were from corms brought to Vienna in 1562 from Constantinople by the Holy Roman Emperor's ambassador to the Sublime Porte, Ogier Ghiselin de Busbecq. A few corms were forwarded to Carolus Clusius at the
botanical garden A botanical garden or botanic gardenThe terms ''botanic'' and ''botanical'' and ''garden'' or ''gardens'' are used more-or-less interchangeably, although the word ''botanic'' is generally reserved for the earlier, more traditional gardens, an ...
in
Leiden Leiden (; in English and archaic Dutch also Leyden) is a city and municipality in the province of South Holland, Netherlands. The municipality of Leiden has a population of 119,713, but the city forms one densely connected agglomeration w ...
. These were almost certainly cultivated varieties rather than wild species. European visitors to Turkey continued to bring back specimens for gardens in their own country. Prominent among the latter were the gardens at Middelburg in the Netherlands. Jehan Somer, a Middelburg merchant, brought back crocuses among his other specimens in 1592, where they attracted the attention not only of Clusius but of the early Dutch flower painters, notably
Ambrosius Bosschaert Ambrosius Bosschaert the Elder (18 January 1573 – 1621) was a Flemish-born Dutch still life painter and art dealer.
. By 1620, new garden varieties had been developed, and featured in contemporary illustrations, such as that of
Crispijn van de Passe Crispijn van de Passe the Elder, or de Passe (c. 1564, Arnemuiden – buried 6 March 1637, Utrecht) was a Dutch publisher and engraver and founder of a dynasty of engravers comparable to the Wierix family and the Sadelers, though mostly at a ...
in his ''Hortus floridus'' of 1614. and there are accounts of crocus gardens in the seventeenth century, such as the Saffron Garth of Walter Stonehouse at Darfield, Yorkshire. Crocuses are among the most important ornamental geophytes in the global flower industry, ranking sixth in terms of Dutch bulb production (2003–2008) with 463–668  ha under cultivation. The crocus is one of the most popular flowers found in the garden in the late winter and early spring. About 30 of the species are cultivated, among the most popular being ''C. chrysanthus, C. flavus, C. sieberi, C. tommasinianus and C. vernus'', together with hundreds of cultivars derived from them. Both fall and spring blooming crocus's are cultivated for their flowers. Among the first flowers to bloom in spring, their flowering time can vary from fall to the late winter blooming ''C. tommasinianus''; the earliest fall blooming species, ''C. scharojanii'', may flower during the last weeks of July. The varieties cultivated for decoration in gardens and pots mainly represent six species: ''C. vernus'', ''C. chrysanthus'', ''C. flavus'', ''C. sieberi'', ''C. speciosus'' and ''C. tommasinianus''. During the horticulture production year 2009/2010, more than 70 cultivars were grown in Holland, covering an area of 366
hectare The hectare (; SI symbol: ha) is a non-SI metric unit of area equal to a square with 100- metre sides (1 hm2), or 10,000 m2, and is primarily used in the measurement of land. There are 100 hectares in one square kilometre. An acre i ...
s; the most common ones were 'Flower Record and 'King of the Stripes' which accounted for 42 hectares, other species grown included ''C. chrysanthus'', ''C. tommasinianus'', and ''C. flavus'' - all are spring blooming plants. But the most commonly grown plants are the Dutch hybrids with large flowers in a rich palette of colors. Both sexual and asexual means are used to increase the number of plants; seeds and multiplication of corms are the most common means of production, but tissue culture can be used, most commonly for saffron crocus. New corms are formed on top of the older corm which withers away, and cormels are produced from axillary buds. The production of new plants begins with harvested corms in late June to early July, after being graded by corm size the corms are stored around 22 Celsius until early October when they are moved to 17 Celsius until planted later in October and November; flowering occurs in March and the flowers are not removed. Crocuses are also forced to produce flowering plants out of season and the most common species used are ''C. vernus'' and ''C. flavus'', and most of the corms used for forcing come from the Netherlands. Spring flowering types are planted in fall, while fall-blooming types in late summer; typically, the corms are placed 3 to 4 inches deep in well-draining soil in areas with full sun exposure. They do not thrive in heavy clay soils or those that are damp, especially during their summer dormancy period. Commercial crops are produced on raised beds and slopes, to ensure adequate drainage, while horticulturalists often plant on sand beds for the same purpose. Spring flowering types also do well in areas with deciduous trees, where they flower and produce leaves before the trees completely leaf-out. Crocuses are grown in USDA winter zones 3–8. Not all species are hardy in the upper zones; ''C. sativus'' is winter hardy in USDA zones 6 through 8, and ''C. pulchellus'' is hardy in zones 5 through 8. Some are suitable for naturalizing in grass, but mowing off the foliage before it turns yellow produces short lived plants. Some crocuses, especially ''C. tommasinianus'' and its selected forms and hybrids (such as 'Whitewell Purple' and 'Ruby Giant'), seed prolifically and are ideal for naturalizing. They can, however, become weeds in rock gardens, where they will often appear in the middle of choice, mat-forming alpine plants, and can be difficult to remove. Crocus flowers and leaves are protected from frost by a waxy cuticle; in areas where snow and frost occasionally occur in the early spring, it is not uncommon to see early flowering crocuses blooming through a light late snowfall. image:Large flowering of purple crocuses.jpg, field of flowering purple crocuses image:CrocusEABowles.jpg, ''Crocus'' 'E.A. Bowles',
a ''C. chrysanthus'' hybrid Image:Frühlingsblumen Krokus.jpg, Crocus cultivars File:Krokusse violett.jpg, Purple crocuses with closed flowers File:Krokusse im Schnee.jpg, Crocuses appearing through the snow


Autumn crocus

Autumn-flowering species of crocus that are cultivated include: ''C. laevigatus'' has a long flowering period which starts in late autumn or early winter and may continue into February.


Uses

The corms of crocuses have been used as foodstuffs in Syria. The carotenoids found in the styles of ''Crocus'' species, particularly ''C. sativus'' have been shown to inhibit cancer cell proliferation, and have led to interest in potential pharmaceutical applications.


Culture

The crocus or krokos has been known since ancient times, and used in
decorative arts ] The decorative arts are arts or crafts whose object is the design and manufacture of objects that are both beautiful and functional. It includes most of the arts making objects for the interiors of buildings, and interior design, but not usua ...
, such as the Minoan wall paintings in Santorini from ca. 1,600 BC. Representations of the saffron crocus appear frequently in Minoan art nd_pervade_Aegean_art_from_the_Early_Bronze_Age_to_the_Mycenaean_Greece.html" ;"title="Early_Bronze_Age.html" ;"title="nd pervade Aegean art from the Early Bronze Age">nd pervade Aegean art from the Early Bronze Age to the Mycenaean Greece">Mycenaean period. Theophrastos (4th century BC) described the saffron crocus as being valued as a spice and dye, while Homer compares a sunrise to the flower colour. Saffron coloured robes were much admired by women in antiquity and gave the garment Crocota its name. The oil was also valued as a cosmetic. According to Greek legend Crocus or Krokus ( gr, Κρόκος), was a mortal youth the gods turned into a plant bearing his name, the crocus, after his death caused by his great desire and unfulfilled love for the shepherdess Smilax. Other versions state that as he died three tears fell into the flower becoming its three stigmata. Crocuses occur in many flower paintings, one of the earliest being that of
Ambrosius Bosschaert Ambrosius Bosschaert the Elder (18 January 1573 – 1621) was a Flemish-born Dutch still life painter and art dealer.
's ''Composed Bouquet of Spring Flowers'' (1620). In this painting the cream-colored crocus feathered with bronze at the base of the bouquet reflected varieties on the market at that time. Bosschaert, working from a preparatory drawing to paint his composed piece spanning the whole of spring, exaggerated the crocus so that it passes for a tulip, but its narrow, grass-like leaves give it away. The crocus is used in many contexts to symbolically denote spring and new beginnings. For instance, it was used as the emblem of the
2019 FIFA U-20 World Cup The 2019 FIFA U-20 World Cup was the 22nd edition of the FIFA U-20 World Cup, the biennial international men's youth football championship contested by the under-20 national teams of the member associations of FIFA, since its inception in 197 ...
in
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populou ...
to symbolise the emergence of new talent.


Notes


References


Bibliography


Books

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Gardening, horticulture and floriculture

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Dictionaries and encyclopedias

* , (link note, note=''see also'' Online Etymology Dictionary * , ** ** * * * * * * * *


Historical sources (chronological)

;Antiquity * ;14th century * ;16th century * * * (
Internet Archive The Internet Archive is an American digital library with the stated mission of "universal access to all knowledge". It provides free public access to collections of digitized materials, including websites, software applications/games, music, ...
version: also here at
Botanicus The Missouri Botanical Garden is a botanical garden located at 4344 Shaw Boulevard in St. Louis, Missouri. It is also known informally as Shaw's Garden for founder and philanthropist Henry Shaw. Its herbarium, with more than 6.6 million s ...
and here at Biodiversity Heritage Library) ;17th century * * ;18th century * , ''see also'' Species Plantarum * * ;19th century * * * ** *


Chapters

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Articles

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Phylogeny and taxonomy

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Novel taxa

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Historical accounts

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Websites

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Organisations and collections

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Databases and flora

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List of Crocus species
* Flora of China ** , in Flora of China online vol. 24 ** , in Flora of China online vol. 6 * *


External links

{{Authority control Iridaceae genera Garden plants Cormous plants