Overview
The Iris family or Iridaceae is a family of perennial, herbaceous and bulbous plants included in the monocot order Asparagales, taking its name from the genus Iris. Almost worldwide in distribution and one of the most important families in horticulture, it includes more than 2000 species. Genera such as Crocus and Iris are significant components of the floras of parts of Eurasia, and Iris also is well-represented in North America. Gladiolus and Moraea are large genera and major constituents of the flora of sub-Saharan and Southern Africa. Sisyrinchium, with more than 140 species, is the most diversified Iridaceae genus in the Americas, where several other genera occur, many of them important in tropical horticulture.1]
All members of Iridaceae have petaloid, soft-textured and colorful perianths in which the three tepals of the inner whorl and the three of t he outer whorl are alike in structure, shape, and often in color. This type of corolla, a feature of all families of Asparagales as well as the related order Liliales, distinguishes them from other monocots, such as grasses, palms or reeds, in which the perianth is either reduced or with the members of one or both of the whorls firm-textured and dry and often brown or green. The character that sets Iridaceae apart from other plants of the Asparagales or from the Liliales, is the male part of the flower, the androecium, which has three stamens?in most related families there are six. Another character that distinguish Iridaceae from most other members of the Asparagales is its inferior ovary, the ovary being superior in most families, with the exception of Orchidaceae and Amaryllidaceae. Apart from their flowers, the Iridaceae can usually be recognized by their characteristic leaves, sword-like and oriented edgewise to the stem and with two identical surfaces. Such leaves are termed isobilateral and u nifacial. In contrast to the Irids, typical plant leaves ? termed dorsiventral and bifacial ? have upper and lower surfaces of different appearance and anatomy.
The Iridaceae originated in Antarctica-Australasia in the late Cretaceous, about 82 million years ago, although the family's subsequent radiation occurred elsewhere, notably in southern Africa and temperate and highland South America, at the end of the Eocene or later.
Currently, 66 genera are recognized, which are distributed among 7 subfamilies and occur in a great variety of habitats. Most species are adapted to seasonal climates that have a pronounced dry or cold period unfavourable for plant growth, during which the plants remain dormant. As a result, most species are deciduous, in that their above-ground parts (leaves and stems) die down when the bulb or corm enters dormancy. The plants thus survive periods that are unfavourable for growth by retreating underground. Evergreen species are restricted to subtropical fo rests or savannah, temperate grasslands and perennially moist fynbos.
Description
In the Iridaceae the perianth is formed of two whorls of three tepals, all similar in structure, shape, and often color. Such a corolla differentiates Liliales and Asparagales from other monocots, where the number or size of the tepals are reduced, or where at least one whorl is papyraceous (firm and dry like a papyrus), and usually green or brown. The characters that differentiate Iridaceae, however, are its three stamen (related families such as Alliaceae and Amaryllidaceae have six) and an inferior ovary. Sword-like leaves parallel to the stem and with normally undifferentiated sides (termed "isobilateral" and "unifacial") are another distinctive feature.
Members of Iridaceae are herbs or, in a few cases, shrubs with woody caudex. They are almost all perennial (three Sisyrinchium species are annuals) that may be either evergreen or se asonal. The rootstock is a rhizome, bulb, or corm.[2] The leaves are found both at the base and on the stem, usually alternate, with the blade oriented parallel to the stem and thus sheathing it at the base. This results in the characteristic fan-like arrangement found in genera like Iris. This type of leaf lacks distinct upper and lower leaf surfaces. In many South African species the leaf has a thickened midrib and often variously thickened or winged margins that may also be crisped. In some species the leaves are needle-like with narrow longitudinal grooves. Species of Moraea are unusual in the family in having channeled leaves with a distinct upper and lower surface.[2][3]
Flowers may be either actinomorphic or zygomorphic. Almo st all the parts are in threes, starting with two equal whorls of three usually large and showy petal-like tepals, distinct or fused in a tube. There are three stamens (rarely two), and their filaments are often partly to completely fused. Anthers have two pollen sacs opening toward the outside, or from their side, and usually along their length. The ovary is located below the tepals (except in Isophysis) with axile (rarely parietal) placentation in three locules. There is a single style branching into three at the top. Iridaceae do not present unisexual flowers, and all flowers have both a style and stamen. Most members other than Sisyrinchium produce nectar from nectaries at the base of the tepals, or on the gynoecium. Iridaceae species are usually pollinated by insects or birds.[1][2] The flowers are collectively arranged in two different types of inflorescences. Simple or branched spikes occur in all Crocoideae. In other subfamilies the basic inflorescence unit is a type of zig-zagging cyme called rhipidium, which is enclosed in enlarged, opposed, bracts called spathes.[1][2]
The fruit is a dry capsule, usually splitting along three sides spontaneously at maturity. It is very variable in shape and texture, from firm to cartilaginous, occasionally woody. In most genera they are tetrahedral or variously angled and without obvious adaptations for dispersal.[3] Seeds are also varied in shape.[4] Winged seeds adapted to wind dispersal characterize Gladiolus and Tritoniopsis and also occur in some species of Hesperantha. Globular seeds with shiny coats that are relatively long-lived occur in several genera of Crocoideae. Chasmanthe aethiopica has fleshy seeds adapted to dispersal by birds and several other species that grow in more wooded places ? like Chasmanthe and some freesias ? have reddish or black seeds that mimic fleshy seeds. They have a hard endosperm, with reserves of hemicellulose, oil, and protein, and a small embryo.[2][5][6]
Distribution and habitat
Members of the Iridaceae are nearly worldwide in distribution, but remain rare in tropical lowlands and at high latitudes. The family is best represented in Southern Africa, especially the w inter-rainfall region in the southwest. Other centers of diversity are temperate South and Central America (with several small genera) and the Mediterranean region (Iris and Crocus). About 2000 species are distributed among some 65 genera are recognized worldwide, just over half of them from Southern Africa, where 38 genera are known. In the Cape Floral Region alone, 707 species and 27 genera are recorded.[1][3]
The family prefers open, seasonal habitats. In Africa, the montane grasslands of eastern South Africa, Swaziland and Lesotho, and the Succulent Karoo and fynbos of the Northern and Western Cape have the most species. Fewer species occur in savannas or the semi-arid central karoo, and very few in forests.[3] T he species grows in a variety of soils, derived from basalt, clay, dolerite, granite, limestone and sandstone, as well as rarer rocks like serpentine. Most species favor loamy soils, often among rocks where drainage is good, but some grow in marshes and others in pure sand. Species grow from just above the high tide mark to over 3,000 metres (9,843 ft) above sea level.[3]
Taxonomy
The family name is based on the genus Iris, the largest and best known genus in Europe. Iris dates from 1753, when it was named by Swedish botanist Car l Linnaeus. Its name derives from the Greek goddess, Iris, who carried messages from Olympus to earth along a rainbow, whose colors were seen by Linnaeus in the multi-hued petals of many of the species.[3] The family name is attributed to Antoine Laurent de Jussieu's 1789 Genera Plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam, and is a conserved name, so that even if an earlier name were to be discovered for the family, Iridaceae would remain valid.[7]
The family has been accepted in all major classification systems of the 20th century. The Cronquist system treated it as part of the order Liliales of the subclass Liliidae,[8] the Takhtajan system placed it in an order Iridales, together with Isophysidaceae and Geo siridaceae treated as single-genus families,[9] and the Thorne system treated it as part of the order Orchidales in its own suborder, Iridineae.[10] The Angiosperm Phylogeny Group in 1998 and 2003 (APG and APG II, respectively) system of flowering plant classification organizes flowering plants into a "selected number of monophyletic suprafamilial groups" and placed Iridaceae in the order Asparagales, which was part of a clade called "Non Commelinoid Monocots".[11][12]
Evolution and phylogeny
The Iridaceae differentiated in the late Cretaceous, about 82 million years ago, and diverged from the next most closely related family, the Doryanthaceae, during the Campanian. The Tasmanian Isophysis is the only extant member of the clade sister to the remainder of the Iridaceae, from which it may have diverged 66 million years ago, in the Maastrichtian. The Iridaceae originated in Antarctica-Australasia, although its subsequent radiation occurred elsewhere, notably in southern Africa and temperate and highland South America at the end of the Eocene or later.[13]
Paleopolyploidy was important in the early diversification of Iridaceae since many genera have basic chromosome numbers which are derived from ancestors with fewer chromosomes. The appearance of polyploidy is important in Northern Hemisphere genera, especially Iris and Crocus, but has an unusually low frequency in Africa, the center of diversity for the family. Changes in basic number, frequent in a few genera, are the result of Robertsonian translocations which led to a reduction in the number of chromosomes.< a href="#endnote_Anone">[a] In all but a few possible examples, accompanying morphological specialization suggests that reduction in the number of chromosomes (dysploid reduction) is involved in gradual changes in base number. Major dysploid series are restricted to a few genera, such as Romulea, the related Crocus as well as Gladiolus and Lapeirousia (all members of subfamily Crocoideae), and Iris, Moraea, and Sisyrinchium (Iridoideae).[14] Complex translocation heterozygotes have been discovered in three species of Homeria (currently included in Gladiolus), a genus from South Africa. In these species, almost every chromosome of the plant has exchanged an arm with another chromosome, which results in a chain of translocated chromosomes. These unusual species, H. tenuis (2n = 10, 9), H. flavescens (2n = 9), and H. pallida (2n i> = 8) are the only aneuploids in the genus; H. pallida also has forms with 2n = 12. The basic chromosome number in Homeria is x = 6 and most species are diploid, a few other polyploid. The genus comprises mostly large-flowered outcrossing species, but self-compatibility and autogamy occur in several, including the three with complex heterozygosity.[15]
Several studies based on morphology and DNA sequence information have shown that the Iridaceae are a monophyletic group included in the order Asparagales.[16][17][18][19] Phylogenetic analyses of sequences of five plastid DNA regions, rbcL, rps4, trnL?F, matK, and rps16, confirm most aspects of the traditional classification of the Iridaceae in four subfamilies and the evolutionary patterns that they imply, importantly the sister relationship of Isophysidoideae to the remainder of the family and the monophyly of Iridoideae. However, it was shown that subfamily Nivenioideae as traditionally defined is paraphyletic: Crocoideae was consistently found nested within it, sister to the core Nivenioideae (the woody genera Klattia, Nivenia, and Witsenia).[1] This clade is sister to Aristea, which in turn is sister to the Madagascarian Geosiris, and then to the Australasian Patersonia.[13][20] This has led to a shift in classification.
Subfamilies
Based on the morphology, anatomy, embryology and chromosome numbers, the family traditionally was divided into four subfamilies (Isophysidodeae, Nivenioideae, Iridioideae and Ixioideae)[21][22] but the results from DNA analysis suggest that several more should be recognized. In fact, the genera Aristea, Geosiris, and Patersonia are now considered as separate subfamilies: Aristeoideae, Geosiridoideae and Patersonioideae, respectively, rendering Nivenioideae and Crocoideae monophyletic.[13][23] Only Iridoideae and Crocoideae are currently subdivided, each into several tribes.
- Isophysidoideae contains the single genus Isophysis, from Tasmania. It is the only member of the family with a superior ovary. It has a star-like yellow to brownish flower.
- Nivenioideae contains three genera (Klattia, Nivenia and Witsenia) and 14 species restricted to the Cape region of South Africa. The plants in this subfamily are true shrubs, with secondary thickening. It is distinguished by having sessile flowers in small, paired clusters surrounded by large bracts, slender styles that are divided into three branches, and nectar produced from glands in the ovary walls. The flowers are always radially symmetrical, with unfused tepals and the rootstock is a rhizome. The basic chromosome number is x= 16.
- Iridoideae, includes 30 genera and about 820 species, is distributed throughout the range of the family, and contains the large genera Iris and Moraea. It is the only subfamily that is represented in South America. The species have flowers in solitary clusters among large bracts, styles that are often petal-like or crested, and nectar (when present) is produced from glands on the tepals. Most species have unfused tepals and the rootstock is usually a rhizome or rarely a bulb. The flowers are almost always radially symmetrical. Bobartia, Dietes and Ferraria belong to this subfamily.
- Patersonioideae contains only one genus (Patersonia) with about 20 species distributed in Sumatra, Borneo, New Guinea, and the periphery of Australia. Plants are more or less woody and rhizomatous with secondary growth. Their flowers are blue and last only for one day. The inner tepals in the flowers are reduced to scales, the filaments are more or less fused. The basic chromosome number is x= 11 or 21.
- Geosiridoideae is represented by only one genus with one species: Geosiris, the only myco-heterotrophic genus of the fam ily, from Madagascar and the Comores. The plants are parastic and lack chlorophyll. They have sessile flowers with the tepals fused at the base, without nectary. They produce minute, dust-like seeds.
- Aristeoideae contains only one genus, Aristea, with about 55 species in sub-Saharan Africa and Madagascar. They are rhizomatous plants with blue flowers which usually last one day, nectaries, and a basic chromosome number x= 16.
- Crocoideae (syn.: Ixioideae), which contains nearly half of the species of the family (28 genera and 995 species), is mostly African. This subfamily contains most of the familiar genera (other than Iris and Moraea) such as Ixia, Gladiolus, Crocus, Freesia and Watsonia. It is easily recognised by its spike-like inflorescence (sometimes bearing solitary flowers) and its tepals joined into a short or long tube. Nectar is produced from glands in the ovary wall and is secreted directly into the base of the floral tube . The flowers are either radially symmetrical or more usually bilaterally symmetrical and two-lipped. The rootstock is a rhizome or, more commonly, a corm.
Genera
Up to 66 genera have been recognised in the family, with a total of around 2,000 species worldwide. The Afrotropic ecozone, and in particular South Africa, have the greatest diversity of genera. DNA sequence information coupled with some associated morphological features provided evidence that several previously recognised New World Tigridieae genera could not retain a generic rank. These are Ainea, Colima, Fosteria, Rigidella and Sessilanthera, which are currently included in Tigridia. The Bolivian genus Cardenanthus was subsumed in Mastigostyla, and Onira and Kelissa were included in the temperate South American genus Cypella.[1] For the same reason, Tamia was included in Calydorea and Tucma in Ennealophus.[1][24]
The genera of Iridaceae, their number of species and geographic distribution are given below.[1][21][22][25][26]
- Afrocrocus [b], one species from Africa.
- Alophia (including Eustylis),[27] five species from the Central and Southern U.S.A. to Brazil.
- Aristea, 56 species distributed in tropical and Southern Africa and Madagascar.
- Babiana, 90 species from tropical to Southern Africa.
- Bobartia, 15 species endemic to Cape Province.
- Calydorea (incl.: Cardiostigma, Itysa, Salpingostylis[28] and Tamia), 21 species from tropical and sub-tropical South America.
- Chasmanthe, three species endemic to Cape Province.
- Cipura, 8 species from Mexico to tropical South America.
- Cobana, one species from Central America.
- Crocosmia, 8 species distributed from Sudan to Southern Africa and Madagascar.
- Crocus, 90 species from Mediterranean Europe to Northwestern China.
- Cyanixia, one species from Socotra.
- Cypella (incl. Onira and Kelissa), 22 species distributed from Peru, Brazil to Northern Argentina.
- Devia, one species endemic to Western Cape.
- Dierama, 43 species from Ethiopia to Southern Africa.
- Dietes, 6 species from Ethiopia to Southern Africa and Lord Howe Island.
- Diplarrhena, two species endemic to Southeastern Australia.
- Duthieastrum, a single species from Northern Cape Province to Free State.
- Eleutherine, two species distributed from Mexico to tropical South America.
- Ennealophus (incl. Tucma and Eurynotia), about 7 species from Ecuador to Northern Brazil and Northwestern Argentina.
- Ferraria, 13 species from southern Za?re to South Africa.
- Freesia (syn. Anomatheca, Lapeirousia), about 56 species distributed from Kenya to Southern Africa.
- Geissorhiza, 85 species endemic to Cape Province.
- Gelasine, 7 species from Braz il to Northeastern Argentina.
- Geosiris, a single species endemic from Madagascar.
- Gladiolus (inc. Oenostachys, Homoglossum, Anomalesia, Antholyza and Acidanthera),[29] 272 species distributed from Europe to Iran, Arabian Peninsula and Africa.
- Herbertia (inc. Sympa), 8 species distributed from Texas to Louisiana, Colombia to Venezuela, Bolivia to southern South America.
- Hesperantha (inc. Schizostylis), 83 species from Cameroon and Ethiopia to Southern Africa.
- Hesperoxiphion, 5 species distributed in Western South America.
- Iris (inc. Belamcanda),[30] includes 260 species distributed from temperate Northern Hemisphere to Philippines.
- Isophysis, a single species from Tasmania.
- < i>Ixia, 61 species from Cape Province.
- Klattia, three species from Southwestern Cape Province.
- Larentia, two species from Mexico, Venezuela, Bolivia to Southern Brazil.
- Lethia, a single species from Bolivia to Brazil (Minas Gerais).
- Libertia, 15 species from Andes to southern South America, and Australia and New Zealand.
- Mastigostyla (incl. Cardenanthus), 17 species from Peru to Northern Argentina.
- Melasphaerula, a single species from Namibia to Cape Province.
- Micranthus, three species from Cape Province.
- Moraea (inc. Bernardiella, Galaxia, Gynandiris, Hexaglottis, Homeria, Sessilstigma and Roggeveldia).[31] includes about 200 species distributed from tropical and South Africa, Mediterranean Europe to western Himalaya.
- Nemastylis , 7 species distributed from Central U.S.A. to Honduras.
- Neomarica, 21 species from Mexico to Trinidad, Brazil to Northeastern Argentina.
- Nivenia, 11 species endemic to Southwestern Cape Province.
- Olsynium, 15 species distributed from western Canada and U.S.A., Costa Rica to Southern South America.
- Orthrosanthus, 9 species from Southern Australia, Mexico to Northwestern Argentina.
- Patersonia, 19 species distributed from Malesia to Australia.
- Phalocallis, a single species from Southern Brazil and Northeastern Argentina.
- Pillansia, a single species from Southwestern Cape Province.
- Pseudotrimezia, 18 species from Southeastern Brazil.
- Radinosiphon, two species from Tanzania to Southern Africa.
- Romulea, about 105 species widely distributed from Macaronesia, Mediterranean Europe, tropical African mountains to Southern Africa and Arabian Peninsula.
- Savannosi phon, a single species distributed from Tanzania to southern tropical Africa.
- Sisyrinchium, about 200 species which range from Hawaiian Islands, Template and Subtropical America to Falkland Islands.
- Solenomelus, two species from Chile and Argentina.
- Sparaxis, 15 species from Southern Africa.
- Syringodea, 7 species from Southern Africa.
- Tapeinia, a single species from southern Chile and Argentina.
- Thereianthus, 8 species from Southwestern Cape Province.
- Tigridia (incl. Ainea, Colima, Fosteria, Rigidella, Sessilanthera), about 50 species distributed from Mexico to El Salvador, and Peru to Northern Chile.
- Trimezia, 41 species from Lesser Antilles to southern tropical America
- Tritonia, 26 species distributed from Tanzania to Southern Africa.
- Tritoniopsis (inc. Anapalina), 23 species from Cape Province.
- Watsonia , 52 species from Southern Africa.
- Witsenia, a single species from Southwestern Cape Province.
- Xenoscapa, two species from Namibia to Cape Province.
- Zygotritonia, four species from tropical Africa.
Ecology
Members of the Iridaceae occur in a great variety of habitats. About the only place they do not grow is in the sea itself, although Gladiolus gueinzii occurs on the seashore just above the high tide mark within reach of the spray. Most species are adapted to seasonal climates that have a pronounced dry or cold period unfavorable for plant growth and during which the plants remain dormant. As a result most species are deciduous. Evergreen species are restricted to subtropical forests or savannah, temperate grasslands and perennially moist fynbos. A few species grow in marshes or along streams and some even grow only in the spray of seasonal waterfalls.[3]
The above-ground parts (leaves and stems) of deciduous species die down when the bulb or corm enters dormancy. The plants thus survive periods that are unfavorable for growth by retreating underground. This is particularly useful adaptation for growth in areas like grasslands and fynbos which regularly have fires in the dry seasons? the plants are dormant and their bulbs or corms are able to survive underground. Veld fires clear the soil surface of competing vegetation and fertilize it with ash. With the arrival of the first rains, the dormant corms are ready to burst into growth, sending up flowers and stems before they can be shaded out by other vegetation. Many Iridaceae species which grow in grassland and fynbos flower best after fires and some fynbos species will only flower in the season after a fire.[3]
The family has a very diverse pollination ecology. Although the majority of species of African Iridaceae are pollinated by Hymenoptera (mostly bees), the remaining species are pollinated mainly, or solely, by insects in the orders Coleoptera (beetles), Diptera (short- and long-proboscid flies) and Lepidoptera (butterflies and moths), or by passerine birds (Nectarinidae). It is now known that pollination systems are predominantly specialized: plants rely on a single species or a few ecologically analogous species for pollination. By contrast, generalist species, which are pollinated by a range of pollinators from at least three pollinator groups, are rare among southern African Iridaceae. In consequence, almost all genera of any size exhibit a range of pollination syndromes, with similar patterns of floral variation having developed repeatedly within different genera.[32]
Most significantly, the diversity of pollination systems increases primarily with floral complexity and secondarily with genus size. Thus, Aristea (approximately 56 species), which has radially symmetric, mostly blue flowers, has three different pollination systems, whereas Sparaxis (15 species), with both zygomorphic and secondarily radially symmetric flowers, in a variety of colors, exhibits five different pollination systems, and Gladiolus, with a similar array of floral types but over two hundred species, exploits seven different pollination systems, some of which have evolved multiple times. As is usual in predominantly specialist pollination systems, floral attractants and rewards correlate closely with pollinator profile, resulting in the development of distinct floral syndromes. Attractants are primarily perianth pigmentation, complemented by a range or floral odors in many species, but flower shape and tepal orientation, in particular functional floral symmetry, may be equally important for some pollinators. The reward to visitors in t he majority of species is nectar, but in others it is pollen, and one species offers non-volatile oil. In the case of hopliine beetles (Scarabaeidae, Hopliinae), flowers provide a stable platform on which to congregate, and the value of pollen, which beetles sometimes consume, as a reward is uncertain.[32]
Conservation
Several species of Iridaceae are endangered or threatened by extinction due to habitat degradation or loss and a restricted geographical distribution. According to the IUCN, the following species are endangered: Babiana longicollis, Cyanixia socotrana (as Babiana socotrana), Gladiolus pole-evansii, Moraea garipensis, Moraea graniticola and Moraea hexaglottis.[33]
Uses
Several cultures have used species of Iridaceae as food, ornamental, condiment or medicinal plants. The Navajo, the largest Native American tribe of North America, used decoctions of Iris missouriensis as an emetic.[34] Pieces of the rhizome of the same species were used to relieve toothaches,[35] or earaches.[36] The mashed roots of Iris versicolor were applied to wounds, presumably as an antiseptic,[37] and the infusions of dry roots of the same species were used to calm pain.[38] Sisyrinchium acre was used in Hawaii in different ways. Leaves or leaf-sap were used as a dye, to give the blue color to tattoos. The use of the leaves, macerated with salt, sugar and other spices was recommended to clean the skin and cure skin diseases.[39] Iris ensata was used in India as anthelmintic and diuretic, and, mixed with other species, to treat venereal diseases.[40] Belamcanda chinensis has a long history of use as medicinal plant in China. Apparently, it was very effective in controlling bacterial, viral and fungal diseases, and in reducing fever and inflammations.[41] Another popular member of the family is Crocus sativus; the spice saffron ? obtained from the stigma ? has been used for centuries in folk medicine as an antispasmodic, aphrodisiac, expectorant, narcotic and sedative.[42]
Iridaceae are of considerable economic importance in ornamental horticulture and the cut-flower industry, especially Iris, Gladiolus, and Freesia. Several other genera (e.g., Crocus, Dietes, Sparaxis, Tritonia, Watsonia) are cultivated in gardens in both tropical and temperate areas. Moraea and Homeria are poisonous and pose significant problems in cattle- and sheep-raising areas, notably in southern Africa.
is a family of perennial, herbaceous and bulbous plants included in the monocot order Asparagales, taking its name from the genus Iris. Almost worldwide in distribution and one of the most important families in horticulture, it includes more than 2000 species. Genera such as Crocus and Iris are significant components of the floras of parts of Eurasia, and Iris also is well-represented in North America. Gladiolus and Moraea are large genera and major constituents of the flora of sub-Saharan and Southern Africa. Sisyrinchium, with more than 140 species, is the most diversified Iridaceae genus in the Americas, where several other genera occur, many of them important in tropical horticulture.1]All members of Iridaceae have petaloid, soft-textured and colorful perianths in which the three tepals of the inner whorl and the three of the outer whorl are alike in structure, shape, and often in color. This type of corolla, a feature of all families of Asparagales as well as the related order Liliales, distinguishes them from other monocots, such as grasses, palms or reeds, in which the perianth is either reduced or with the members of one or both of the whorls firm-textured and dry and often brown or green. The char acter that sets Iridaceae apart from other plants of the Asparagales or from the Liliales, is the male part of the flower, the androecium, which has three stamens?in most related families there are six. Another character that distinguish Iridaceae from most other members of the Asparagales is its inferior ovary, the ovary being superior in most families, with the exception of Orchidaceae and Amaryllidaceae. Apart from their flowers, the Iridaceae can usually be recognized by their characteristic leaves, sword-like and oriented edgewise to the stem and with two identical surfaces. Such leaves are termed isobilateral and unifacial. In contrast to the Irids, typical plant leaves ? termed dorsiventral and bifacial ? have upper and lower surfaces of different appearance and anatomy.
The Iridaceae originated in Antarctica-Australasia in the late Cretaceous, about 82 million years ago, although the family's subsequent radiation occurred elsewhere, notably in southern Africa and temp erate and highland South America, at the end of the Eocene or later.
Currently, 66 genera are recognized, which are distributed among 7 subfamilies and occur in a great variety of habitats. Most species are adapted to seasonal climates that have a pronounced dry or cold period unfavourable for plant growth, during which the plants remain dormant. As a result, most species are deciduous, in that their above-ground parts (leaves and stems) die down when the bulb or corm enters dormancy. The plants thus survive periods that are unfavourable for growth by retreating underground. Evergreen species are restricted to subtropical forests or savannah, temperate grasslands and perennially moist fynbos.
Description
In the Iridaceae the perianth is formed of two whorls of three tepals, all similar in structure, shape, and often color. Such a corolla differentiates Liliales and Asparagales from other monocots, where the number or size of the tepals are reduced, or where at least one whorl is papyraceous (firm and dry like a papyrus), and usually green or brown. The characters that differentiate Iridaceae, however, are its three stamen (related families such as Alliaceae and Amaryllidaceae have six) and an inferior ovary. Sword-like leaves parallel to the stem and with normally undifferentiated sides (termed "isobilateral" and "unifacial") are another distinctive feature.
Members of Iridaceae are herbs or, in a few cases, shrubs with woody caudex. They are almost all perennial (three Sisyrinchium species are annuals) that may be either evergreen or seasonal. The rootstock is a rhizome, bulb, or corm.[2] The leaves are found both at the base and on the stem, usually alternate, with the blade oriented parallel to the stem and thus sheathing it at the base. This results in the characteristic fan-like arrangement found in genera like Iris. This type of leaf lacks distinct upper and lower leaf surfaces. In many South African species the leaf has a thickened midrib and often variously thickened or winged margins that may also be crisped. In some species the leaves are needle-like with narrow longitudinal grooves. Species of Moraea are unusual in the family in having channeled leaves with a distinct upper and lower surface.[2][3]
Flowers may be either actinomorphic or zygomorphic. Almost all the parts are in threes, starting with two equal whorls of three usually large and showy petal-like tepals, distinct or fused in a tube. There are three stamens (rarely two), and their filaments are often partly to completely fused. Anthers have two pollen sacs opening toward the outside, or from their side, and usually along their length. The ovary is located below the tepals (except in Isophysis) with axile (rarely parietal) placentation in three locules. There is a single style branching into three at the top. Iridaceae do not present unisexual flowers, and all flowers have both a style and stamen. Most members other than Sisyrinchium produce nectar from nectaries at the base of the tepals, or on the gynoecium. Iridaceae species are usually pollinated by insects or birds.[1][2] The flowers are collectively arranged in two different types of inflorescences. Simple or branched spikes occur in all Crocoideae. In other subfamilies the basic inflorescence unit is a type of zig-zagging cyme called rhipidium, which is enclosed in enlarged, opposed, bracts called spathes.[1][2]
The fruit is a dry capsule, usually splitting along three sides spontaneously at maturity. It is very variable in shape and texture, from firm to cartilaginous, occasionally woody. In most genera they are tetrahedral or variously angled and without obvious adaptations for dispersal.[3] Seeds are also varied in shape.[4] Winged seeds adapted to wind dispersal characterize Gladiolus and Tritoniopsis and also occur in some species of Hesperantha. Globular seeds with shiny coats that are relatively long-lived occur in several genera of Crocoideae. Chasmanthe aethiopica has fleshy seeds adapted to dispersal by birds and several other species that grow in more wooded places ? like Chasmanthe and some freesias ? have reddish or black seeds that mimic fleshy seeds. T hey have a hard endosperm, with reserves of hemicellulose, oil, and protein, and a small embryo.[2][5][6]
Distribution and habitat
Members of the Iridaceae are nearly worldwide in distribution, but remain rare in tropical lowlands and at high latitudes. The family is best represented in Southern Africa, especially the winter-rainfall region in the southwest. Other centers of diversity are temperate South and Central America (with several small genera) and the Mediterranean region (Iris and Crocus). About 2000 species are distributed among some 65 genera are recognized worldwide, just over half of them from Southern Africa, where 38 genera are known. In the Cape Floral Region alone, 7 07 species and 27 genera are recorded.[1][3]
The family prefers open, seasonal habitats. In Africa, the montane grasslands of eastern South Africa, Swaziland and Lesotho, and the Succulent Karoo and fynbos of the Northern and Western Cape have the most species. Fewer species occur in savannas or the semi-arid central karoo, and very few in forests.[3] The species grows in a variety of soils, derived from basalt, clay, dolerite, granite, limestone and sandstone, as well as rarer rocks like serpentine. Most species favor loamy soils, often among rocks where drainage is good, but some grow in marshes and others in pure sand. Species grow from just above the high tide mark to over 3,000 metres (9,843 ft) above sea level.[3]
Taxonomy
The family name is based on the genus Iris, the largest and best known genus in Europe. Iris dates from 1753, when it was named by Swedish botanist Carl Linnaeus. Its name derives from the Greek goddess, Iris, who carried messages from Olympus to earth along a rainbow, whose colors were seen by Linnaeus in the multi-hued petals of many of the species.[3] The family name is attributed to Antoine Laurent de Jussieu's 1789 Genera Plantar um, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam, and is a conserved name, so that even if an earlier name were to be discovered for the family, Iridaceae would remain valid.[7]
The family has been accepted in all major classification systems of the 20th century. The Cronquist system treated it as part of the order Liliales of the subclass Liliidae,[8] the Takhtajan system placed it in an order Iridales, together with Isophysidaceae and Geosiridaceae treated as single-genus families,[9] and the Thorne system treated it as part of the order Orchidales in its own suborder, Iridineae.[10] The Angiosperm Phylogeny Group in 1998 and 2003 (APG and APG II, respectively) system of fl owering plant classification organizes flowering plants into a "selected number of monophyletic suprafamilial groups" and placed Iridaceae in the order Asparagales, which was part of a clade called "Non Commelinoid Monocots".[11][12]
Evolution and phylogeny
The Iridaceae differentiated in the late Cretaceous, about 82 million years ago, and diverged from the next most closely related family, the Doryanthaceae, during the Campanian. The Tasmanian Isophysis is the only extant member of the clade sister to the remainder of the Iridaceae, from which it may have diverged 66 million years ago, in the Maastrichtian. The Iridaceae originated in Antarctica-Australasia, although its subsequent radiation occurred elsewhere, notably in southern Africa and temperate and highland South America at the end of the Eocene or later.[13]
Paleopolyploidy was important in the early diversification of Iridaceae since many genera have basic chromosome numbers which are derived from ancestors with fewer chromosomes. The appearance of polyploidy is important in Northern Hemisphere genera, especially Iris and Crocus, but has an unusually low frequency in Africa, the center of diversity for the family. Changes in basic number, frequent in a few genera, are the result of Robertsonian translocations which led to a reduction in the number of chromosomes.[a] In all but a few possible examples, accompanying morphological specialization suggests that reduction in the number of chromosomes (dysploid reduction) is involved in gradual changes in base number. Major dysploid series are restricted to a few genera, such as Romulea, the related Crocus as well as Gladiolus and Lapeiro usia (all members of subfamily Crocoideae), and Iris, Moraea, and Sisyrinchium (Iridoideae).[14] Complex translocation heterozygotes have been discovered in three species of Homeria (currently included in Gladiolus), a genus from South Africa. In these species, almost every chromosome of the plant has exchanged an arm with another chromosome, which results in a chain of translocated chromosomes. These unusual species, H. tenuis (2n = 10, 9), H. flavescens (2n = 9), and H. pallida (2n = 8) are the only aneuploids in the genus; H. pallida also has forms with 2n = 12. The basic chromosome number in Homeria is x = 6 and most species are diploid, a few other polyploid. The genus comprises mostly large-flowered outcrossing species, but self-compatibility and autogamy occur in several, including the three with complex heterozygosity.[15]
Several studies based on morphology and DNA sequence information have shown that the Iridaceae are a monophyletic group included in the order Asparagales.[16][17][18][19] Phylogenetic analyses of sequences of five plastid DNA regions, rbcL, rps4, trnL?F, matK, and rps16, confirm most aspects of the traditional classification of the Iridaceae in four subfamilies and the evolutionary patterns that they imply, importantly the sister relationship of Isophysidoideae to the remainder of th e family and the monophyly of Iridoideae. However, it was shown that subfamily Nivenioideae as traditionally defined is paraphyletic: Crocoideae was consistently found nested within it, sister to the core Nivenioideae (the woody genera Klattia, Nivenia, and Witsenia).[1] This clade is sister to Aristea, which in turn is sister to the Madagascarian Geosiris, and then to the Australasian Patersonia.[13][20] This has led to a shift in classification.
Subfamilies
Based on the morphology, anatomy, embryology and chromosome numbers, the family traditionally was divided into four subfamilies (Isophysidodeae, Nivenioideae, Iridioideae and Ixioideae)[21][22] but the results from DNA analysis suggest that several more should be recognized. In fact, the genera Aristea, Geosiris, and Patersonia are now considered as separate subfamilies: Aristeoideae, Geosiridoideae and Patersonioideae, respectively, rendering Nivenioideae and Crocoideae monophyletic.[13][23] Only Iridoideae and Crocoideae are currently subdivided, each into several tribes.
- Isophysidoideae contains the single genus Isophysis, from Tasmania. It is the only member of the family with a superior ovary. It has a star-like yellow to brownish flower.
- Nivenioideae contains three genera (Klattia, Nivenia and Witsenia) and 14 species restricted to the Cape region of South Africa. The plants in this subfamily are true shrubs, with secondary thickening. It is distinguished by having sessile flowers in small, paired clusters surrounded by large bracts, slender styles that are divided into three branches, and nectar produced from glands in the ovary walls. The flowers are always radially symmetrical, with unfused tepals and the rootstock is a rhizome. The basic chromosome number is x= 16.
- Iridoideae, includes 30 genera and about 820 species, is distributed throughout the range of the family, and contains the large genera Iris and Moraea. It is the only subfamily that is represented in South America. The species have flowers in solitary clusters among large bracts, styles that are often petal-like or crested, and nectar (when present) is produced from glands on the tepals. Most species have unfused tepals and the rootstock is usually a rhizome or rarely a bulb. The flowers are almost always radially symmetrical. Bobartia, Dietes and Ferraria belong to this subfamily.
- Patersonioideae contains only one genus (Patersonia) with about 20 species distributed in Sumatra, Borneo, New Guinea, and the periphery of Australia. Plants are more or less woody and rhizomatous with secondary growth. Their flowers are blue and last only for one day. The inner tepals in the flowers are reduced to scales, the filaments are more or less fused. The basic chromosome number is x= 11 or 21.
- Geosiridoideae is represented by only one genus with one species: Geosiris, the only myco-heterotrophic genus of the family, from Madagascar and the Comores. The plants are parastic and lack chlorophyll. They have sessile flowers with the tepals fused at the base, without nectary. They produce minute, dust-like seeds.
- Aristeoideae contains only one genus, Aristea, with about 55 species in sub-Saharan Africa and Madagascar. They are rhizomatous plants with blue flowers which usu ally last one day, nectaries, and a basic chromosome number x= 16.
- Crocoideae (syn.: Ixioideae), which contains nearly half of the species of the family (28 genera and 995 species), is mostly African. This subfamily contains most of the familiar genera (other than Iris and Moraea) such as Ixia, Gladiolus, Crocus, Freesia and Watsonia. It is easily recognised by its spike-like inflorescence (sometimes bearing solitary flowers) and its tepals joined into a short or long tube. Nectar is produced from glands in the ovary wall and is secreted directly into the base of the floral tube. The flowers are either radially symmetrical or more usually bilaterally symmetrical and two-lipped. The rootstock is a rhizome or, more commonly, a corm.
Genera
Up to 66 genera have been recognised in the family, with a total of around 2,000 species worldwide. The Afrotropic ecozone, and in particular South Africa, have the greatest diversity of genera. DNA s equence information coupled with some associated morphological features provided evidence that several previously recognised New World Tigridieae genera could not retain a generic rank. These are Ainea, Colima, Fosteria, Rigidella and Sessilanthera, which are currently included in Tigridia. The Bolivian genus Cardenanthus was subsumed in Mastigostyla, and Onira and Kelissa were included in the temperate South American genus Cypella.[1] For the same reason, Tamia was included in Calydorea and Tucma in Ennealophus.[1][24]
The genera of Iridaceae, their number of species and geographic distribution are given below.[1][21][22][25][26]
- Afrocrocus [b], one species from Africa.
- Alophia (including Eustylis),[27] five species from the Central and Southern U.S.A. to Brazil.
- Aristea, 56 species distributed in tropical and Southern Africa and Madagascar.
- Babiana, 90 species from tropical to Southern Africa.
- Bobartia, 15 species endemic to Cape Province.
- Calydorea (incl.: Cardiostigma, Itysa, Sa lpingostylis[28] and Tamia), 21 species from tropical and sub-tropical South America.
- Chasmanthe, three species endemic to Cape Province.
- Cipura, 8 species from Mexico to tropical South America.
- Cobana, one species from Central America.
- Crocosmia, 8 species distributed from Sudan to Southern Africa and Madagascar.
- Crocus, 90 species from Mediterranean Europe to Northwestern China.
- Cyanixia, one species from Socotra.
- Cypella (incl. Onira and Kelissa), 22 species distributed from Peru, Brazil to Northern Argentina.
- Devia, one species endemic to Western Cape.
- Dierama, 43 species from Ethiopia to Southern Africa.
- Dietes, 6 species from Ethiopia to Southern Africa and Lord Howe Island.
- Diplarrhena, two species endemic to Southeastern Austral ia.
- Duthieastrum, a single species from Northern Cape Province to Free State.
- Eleutherine, two species distributed from Mexico to tropical South America.
- Ennealophus (incl. Tucma and Eurynotia), about 7 species from Ecuador to Northern Brazil and Northwestern Argentina.
- Ferraria, 13 species from southern Za?re to South Africa.
- Freesia (syn. Anomatheca, Lapeirousia), about 56 species distributed from Kenya to Southern Africa.
- Geissorhiza, 85 species endemic to Cape Province.
- Gelasine, 7 species from Brazil to Northeastern Argentina.
- Geosiris, a single species endemic from Madagascar.
- Gladiolus (inc. Oenostachys, Homoglossum, Anomalesia, Antholyza and Acidanthera),[29] 272 species distributed from Europe to Iran, Arabian Peninsula a nd Africa.
- Herbertia (inc. Sympa), 8 species distributed from Texas to Louisiana, Colombia to Venezuela, Bolivia to southern South America.
- Hesperantha (inc. Schizostylis), 83 species from Cameroon and Ethiopia to Southern Africa.
- Hesperoxiphion, 5 species distributed in Western South America.
- Iris (inc. Belamcanda),[30] includes 260 species distributed from temperate Northern Hemisphere to Philippines.
- Isophysis, a single species from Tasmania.
- Ixia, 61 species from Cape Province.
- Klattia, three species from Southwestern Cape Province.
- Larentia, two species from Mexico, Venezuela, Bolivia to Southern Brazil.
- Lethia, a single species from Bolivia to Brazil (Minas Gerais).
- Libertia, 15 species from Andes to southern South America, and Australia and New Zeala nd.
- Mastigostyla (incl. Cardenanthus), 17 species from Peru to Northern Argentina.
- Melasphaerula, a single species from Namibia to Cape Province.
- Micranthus, three species from Cape Province.
- Moraea (inc. Bernardiella, Galaxia, Gynandiris, Hexaglottis, Homeria, Sessilstigma and Roggeveldia).[31] includes about 200 species distributed from tropical and South Africa, Mediterranean Europe to western Himalaya.
- Nemastylis, 7 species distributed from Central U.S.A. to Honduras.
- Neomarica, 21 species from Mexico to Trinidad, Brazil to Northeastern Argentina.
- Nivenia, 11 species endemic to Southwestern Cape Province.
- Olsynium, 15 species distributed from western Canada and U.S.A., Costa Rica to Southern South America.
- Orthrosanthus, 9 specie s from Southern Australia, Mexico to Northwestern Argentina.
- Patersonia, 19 species distributed from Malesia to Australia.
- Phalocallis, a single species from Southern Brazil and Northeastern Argentina.
- Pillansia, a single species from Southwestern Cape Province.
- Pseudotrimezia, 18 species from Southeastern Brazil.
- Radinosiphon, two species from Tanzania to Southern Africa.
- Romulea, about 105 species widely distributed from Macaronesia, Mediterranean Europe, tropical African mountains to Southern Africa and Arabian Peninsula.
- Savannosiphon, a single species distributed from Tanzania to southern tropical Africa.
- Sisyrinchium, about 200 species which range from Hawaiian Islands, Template and Subtropical America to Falkland Islands.
- Solenomelus, two species from Chile and Argentina.
- Sparaxis, 15 species from Southern Africa.
- Syringodea, 7 species from Southern Africa.
- Tapeinia, a single species from southern Chile and Argentina.
- Thereianthus, 8 species from Southwestern Cape Province.
- Tigridia (incl. Ainea, Colima, Fosteria, Rigidella, Sessilanthera), about 50 species distributed from Mexico to El Salvador, and Peru to Northern Chile.
- Trimezia, 41 species from Lesser Antilles to southern tropical America
- Tritonia, 26 species distributed from Tanzania to Southern Africa.
- Tritoniopsis (inc. Anapalina), 23 species from Cape Province.
- Watsonia, 52 species from Southern Africa.
- Witsenia, a single species from Southwestern Cape Province.
- Xenoscapa, two species from Namibia to Cape Province.
- Zygotritonia, four species from tropical Africa.
Ecology
Members of the Iridaceae occur in a great v ariety of habitats. About the only place they do not grow is in the sea itself, although Gladiolus gueinzii occurs on the seashore just above the high tide mark within reach of the spray. Most species are adapted to seasonal climates that have a pronounced dry or cold period unfavorable for plant growth and during which the plants remain dormant. As a result most species are deciduous. Evergreen species are restricted to subtropical forests or savannah, temperate grasslands and perennially moist fynbos. A few species grow in marshes or along streams and some even grow only in the spray of seasonal waterfalls.[3]
The above-ground parts (leaves and stems) of deciduous species die down when the bulb or corm enters dormancy. The plants thus survive periods that are unfavorable for growth by retreating underground. This is particularly useful adaptation for growth in areas like grasslands and fynbos which reg ularly have fires in the dry seasons? the plants are dormant and their bulbs or corms are able to survive underground. Veld fires clear the soil surface of competing vegetation and fertilize it with ash. With the arrival of the first rains, the dormant corms are ready to burst into growth, sending up flowers and stems before they can be shaded out by other vegetation. Many Iridaceae species which grow in grassland and fynbos flower best after fires and some fynbos species will only flower in the season after a fire.[3]
The family has a very diverse pollination ecology. Although the majority of species of African Iridaceae are pollinated by Hymenoptera (mostly bees), the remaining species are pollinated mainly, or solely, by insects in the orders Coleoptera (beetles), Diptera (short- and long-proboscid flies) and Lepidoptera (butterflies and moths), or by passerine birds (Nectarinidae). It is now known that poll ination systems are predominantly specialized: plants rely on a single species or a few ecologically analogous species for pollination. By contrast, generalist species, which are pollinated by a range of pollinators from at least three pollinator groups, are rare among southern African Iridaceae. In consequence, almost all genera of any size exhibit a range of pollination syndromes, with similar patterns of floral variation having developed repeatedly within different genera.[32]
Most significantly, the diversity of pollination systems increases primarily with floral complexity and secondarily with genus size. Thus, Aristea (approximately 56 species), which has radially symmetric, mostly blue flowers, has three different pollination systems, whereas Sparaxis (15 species), with both zygomorphic and secondarily radially symmetric flowers, in a variety of colors, exhibits five different pollination sys tems, and Gladiolus, with a similar array of floral types but over two hundred species, exploits seven different pollination systems, some of which have evolved multiple times. As is usual in predominantly specialist pollination systems, floral attractants and rewards correlate closely with pollinator profile, resulting in the development of distinct floral syndromes. Attractants are primarily perianth pigmentation, complemented by a range or floral odors in many species, but flower shape and tepal orientation, in particular functional floral symmetry, may be equally important for some pollinators. The reward to visitors in the majority of species is nectar, but in others it is pollen, and one species offers non-volatile oil. In the case of hopliine beetles (Scarabaeidae, Hopliinae), flowers provide a stable platform on which to congregate, and the value of pollen, which beetles sometimes consume, as a reward is uncertain.[32]
Conservation
Several species of Iridaceae are endangered or threatened by extinction due to habitat degradation or loss and a restricted geographical distribution. According to the IUCN, the following species are endangered: Babiana longicollis, Cyanixia socotrana (as Babiana socotrana), Gladiolus pole-evansii, Moraea garipensis, Moraea graniticola and Moraea hexaglottis.[33]
Uses
Several cultures have used species of Iridaceae as food, ornamental, condiment or medicinal plants. The Navajo, the largest Native American tribe of North America, used decoctions of Iris missouriensis as an emetic.[34] Pieces of the rhizome of the same species were used to relieve toothaches,[35] or earaches.[36] The mashed roots of Iris versicolor were applied to wounds, presumably as an antiseptic,[37] and the infusions of dry roots of the same species were used to calm pain.[38] Sisyrinchium acre was used in Hawaii in different ways. Leaves or leaf-sap were used as a dye, to give the blue color to tattoos. The use of the leaves, macerated with salt, sugar and other spices was recommended to clean the skin and cure skin diseases.[39] Iris ensa ta was used in India as anthelmintic and diuretic, and, mixed with other species, to treat venereal diseases.[40] Belamcanda chinensis has a long history of use as medicinal plant in China. Apparently, it was very effective in controlling bacterial, viral and fungal diseases, and in reducing fever and inflammations.[41] Another popular member of the family is Crocus sativus; the spice saffron ? obtained from the stigma ? has been used for centuries in folk medicine as an antispasmodic, aphrodisiac, expectorant, narcotic and sedative.[42]
Iridaceae are of considerable economic importance in ornamental horticulture and the cut-flower industry, especially Iris, Gladiolus, and Freesia . Several other genera (e.g., Crocus, Dietes, Sparaxis, Tritonia, Watsonia) are cultivated in gardens in both tropical and temperate areas. Moraea and Homeria are poisonous and pose significant problems in cattle- and sheep-raising areas, notably in southern Africa.
Notes
The family has been accepted in all major classification systems of the 20th century. The Cronquist system treated it as part of the order Liliales of the subclass Liliidae,[8] the Takhtajan system placed it in an order Iridales, together with Isophysidaceae and Geosiridaceae treated as single-genus families,[9] and the Thorne system treated it as part of the order Orchidales in its own sub order, Iridineae.[10] The Angiosperm Phylogeny Group in 1998 and 2003 (APG and APG II, respectively) system of flowering plant classification organizes flowering plants into a "selected number of monophyletic suprafamilial groups" and placed Iridaceae in the order Asparagales, which was part of a clade called "Non Commelinoid Monocots".[11][12]
Evolution and phylogeny
The Iridaceae differentiated in the late Cretaceous, about 82 million years ago, and diverged from the next most closely related family, the Doryanthaceae, during the Campanian. The Tasmanian Isophysis is the only extant member of the clade sister to the remainder of the Iridaceae, from which it may have diverged 66 million years ago, in the Maastrichtian. The Iridaceae originated in Antarctica-Australasia, a lthough its subsequent radiation occurred elsewhere, notably in southern Africa and temperate and highland South America at the end of the Eocene or later.[13]
Paleopolyploidy was important in the early diversification of Iridaceae since many genera have basic chromosome numbers which are derived from ancestors with fewer chromosomes. The appearance of polyploidy is important in Northern Hemisphere genera, especially Iris and Crocus, but has an unusually low frequency in Africa, the center of diversity for the family. Changes in basic number, frequent in a few genera, are the result of Robertsonian translocations which led to a reduction in the number of chromosomes.[a] In all but a few possible examples, accompanying morphological specialization suggests that reduction in the number of chromosomes (dysploid reduction) is invol ved in gradual changes in base number. Major dysploid series are restricted to a few genera, such as Romulea, the related Crocus as well as Gladiolus and Lapeirousia (all members of subfamily Crocoideae), and Iris, Moraea, and Sisyrinchium (Iridoideae).[14] Complex translocation heterozygotes have been discovered in three species of Homeria (currently included in Gladiolus), a genus from South Africa. In these species, almost every chromosome of the plant has exchanged an arm with another chromosome, which results in a chain of translocated chromosomes. These unusual species, H. tenuis (2n = 10, 9), H. flavescens (2n = 9), and H. pallida (2n = 8) are the only aneuploids in the genus; H. pallida also has forms with 2n = 12. The basic chromosome number in Homeria is x = 6 and most species are diploid, a few other p olyploid. The genus comprises mostly large-flowered outcrossing species, but self-compatibility and autogamy occur in several, including the three with complex heterozygosity.[15]
Several studies based on morphology and DNA sequence information have shown that the Iridaceae are a monophyletic group included in the order Asparagales.[16][17][18][19] Phylogenetic analyses of sequences of five plastid DNA regions, rbcL, rps4, trnL?F, matK, and rps16, confirm most aspects of the tra ditional classification of the Iridaceae in four subfamilies and the evolutionary patterns that they imply, importantly the sister relationship of Isophysidoideae to the remainder of the family and the monophyly of Iridoideae. However, it was shown that subfamily Nivenioideae as traditionally defined is paraphyletic: Crocoideae was consistently found nested within it, sister to the core Nivenioideae (the woody genera Klattia, Nivenia, and Witsenia).[1] This clade is sister to Aristea, which in turn is sister to the Madagascarian Geosiris, and then to the Australasian Patersonia.[13][20] This has led to a shift in classification.
Subfamilies
Based on the morphology, anatomy, embryology and chromosome numbers, t he family traditionally was divided into four subfamilies (Isophysidodeae, Nivenioideae, Iridioideae and Ixioideae)[21][22] but the results from DNA analysis suggest that several more should be recognized. In fact, the genera Aristea, Geosiris, and Patersonia are now considered as separate subfamilies: Aristeoideae, Geosiridoideae and Patersonioideae, respectively, rendering Nivenioideae and Crocoideae monophyletic.[13][23] Only Iridoideae and Crocoideae are currently subdivided, each into several tribes.
- Isophysidoideae contains the single genus Isophysis, from Tasmania. It is the only member of the family with a superior o vary. It has a star-like yellow to brownish flower.
- Nivenioideae contains three genera (Klattia, Nivenia and Witsenia) and 14 species restricted to the Cape region of South Africa. The plants in this subfamily are true shrubs, with secondary thickening. It is distinguished by having sessile flowers in small, paired clusters surrounded by large bracts, slender styles that are divided into three branches, and nectar produced from glands in the ovary walls. The flowers are always radially symmetrical, with unfused tepals and the rootstock is a rhizome. The basic chromosome number is x= 16.
- Iridoideae, includes 30 genera and about 820 species, is distributed throughout the range of the family, and contains the large genera Iris and Moraea. It is the only subfamily that is represented in South America. The species have flowers in solitary clusters among large bracts, styles that are often petal-like or crested, and nectar (when present) is produ ced from glands on the tepals. Most species have unfused tepals and the rootstock is usually a rhizome or rarely a bulb. The flowers are almost always radially symmetrical. Bobartia, Dietes and Ferraria belong to this subfamily.
- Patersonioideae contains only one genus (Patersonia) with about 20 species distributed in Sumatra, Borneo, New Guinea, and the periphery of Australia. Plants are more or less woody and rhizomatous with secondary growth. Their flowers are blue and last only for one day. The inner tepals in the flowers are reduced to scales, the filaments are more or less fused. The basic chromosome number is x= 11 or 21.
- Geosiridoideae is represented by only one genus with one species: Geosiris, the only myco-heterotrophic genus of the family, from Madagascar and the Comores. The plants are parastic and lack chlorophyll. They have sessile flowers with the tepals fused at the base, without nectary. They produce minute, dust-like seeds .
- Aristeoideae contains only one genus, Aristea, with about 55 species in sub-Saharan Africa and Madagascar. They are rhizomatous plants with blue flowers which usually last one day, nectaries, and a basic chromosome number x= 16.
- Crocoideae (syn.: Ixioideae), which contains nearly half of the species of the family (28 genera and 995 species), is mostly African. This subfamily contains most of the familiar genera (other than Iris and Moraea) such as Ixia, Gladiolus, Crocus, Freesia and Watsonia. It is easily recognised by its spike-like inflorescence (sometimes bearing solitary flowers) and its tepals joined into a short or long tube. Nectar is produced from glands in the ovary wall and is secreted directly into the base of the floral tube. The flowers are either radially symmetrical or more usually bilaterally symmetrical and two-lipped. The rootstock is a rhizome or, more commonly, a corm.
Genera
Up to 66 gener a have been recognised in the family, with a total of around 2,000 species worldwide. The Afrotropic ecozone, and in particular South Africa, have the greatest diversity of genera. DNA sequence information coupled with some associated morphological features provided evidence that several previously recognised New World Tigridieae genera could not retain a generic rank. These are Ainea, Colima, Fosteria, Rigidella and Sessilanthera, which are currently included in Tigridia. The Bolivian genus Cardenanthus was subsumed in Mastigostyla, and Onira and Kelissa were included in the temperate South American genus Cypella.[1] For the same reason, Tamia was included in Calydorea and Tucma in Ennealophus.[1][24]
The genera of Iridaceae, their number of species and geographic distribution are given below.[1][21][22][25][26]
- Afrocrocus [b], one species from Africa.
- Alophia (including Eustylis),[27] five species from the Central and Southern U.S.A. to Brazil.
- Aristea, 56 species distributed in tropical and Southern Africa and Madagascar.
- Babiana , 90 species from tropical to Southern Africa.
- Bobartia, 15 species endemic to Cape Province.
- Calydorea (incl.: Cardiostigma, Itysa, Salpingostylis[28] and Tamia), 21 species from tropical and sub-tropical South America.
- Chasmanthe, three species endemic to Cape Province.
- Cipura, 8 species from Mexico to tropical South America.
- Cobana, one species from Central America.
- Crocosmia, 8 species distributed from Sudan to Southern Africa and Madagascar.
- Crocus, 90 species from Mediterranean Europe to Northwestern China.
- Cyanixia, one species from Socotra.
- Cypella (incl. Onira and Kelissa), 22 species distributed from Peru, Brazil to Northern Argentina.
- Devia, one species endemic to Western Cape.
- Dierama, 43 species from Et hiopia to Southern Africa.
- Dietes, 6 species from Ethiopia to Southern Africa and Lord Howe Island.
- Diplarrhena, two species endemic to Southeastern Australia.
- Duthieastrum, a single species from Northern Cape Province to Free State.
- Eleutherine, two species distributed from Mexico to tropical South America.
- Ennealophus (incl. Tucma and Eurynotia), about 7 species from Ecuador to Northern Brazil and Northwestern Argentina.
- Ferraria, 13 species from southern Za?re to South Africa.
- Freesia (syn. Anomatheca, Lapeirousia), about 56 species distributed from Kenya to Southern Africa.
- Geissorhiza, 85 species endemic to Cape Province.
- Gelasine, 7 species from Brazil to Northeastern Argentina.
- Geosiris, a single species endemic from Madagascar.
- Gladiolus (inc. Oenostachys, Homoglossum, Anomalesia, Antholy za and Acidanthera),[29] 272 species distributed from Europe to Iran, Arabian Peninsula and Africa.
- Herbertia (inc. Sympa), 8 species distributed from Texas to Louisiana, Colombia to Venezuela, Bolivia to southern South America.
- Hesperantha (inc. Schizostylis), 83 species from Cameroon and Ethiopia to Southern Africa.
- Hesperoxiphion, 5 species distributed in Western South America.
- Iris (inc. Belamcanda),[30] includes 260 species distributed from temperate Northern Hemisphere to Philippines.
- Isophysis, a single species from Tasmania.
- Ixia, 61 species from Cape Province.
- Klattia, three species from Southwestern Cape Province.
- Larentia, two species from Mexico, Venezuela, Bolivia to Southern Br azil.
- Lethia, a single species from Bolivia to Brazil (Minas Gerais).
- Libertia, 15 species from Andes to southern South America, and Australia and New Zealand.
- Mastigostyla (incl. Cardenanthus), 17 species from Peru to Northern Argentina.
- Melasphaerula, a single species from Namibia to Cape Province.
- Micranthus, three species from Cape Province.
- Moraea (inc. Bernardiella, Galaxia, Gynandiris, Hexaglottis, Homeria, Sessilstigma and Roggeveldia).[31] includes about 200 species distributed from tropical and South Africa, Mediterranean Europe to western Himalaya.
- Nemastylis, 7 species distributed from Central U.S.A. to Honduras.
- Neomarica, 21 species from Mexico to Trinidad, Brazil to Northeastern Argentina.
- Nivenia, 11 species endemic t o Southwestern Cape Province.
- Olsynium, 15 species distributed from western Canada and U.S.A., Costa Rica to Southern South America.
- Orthrosanthus, 9 species from Southern Australia, Mexico to Northwestern Argentina.
- Patersonia, 19 species distributed from Malesia to Australia.
- Phalocallis, a single species from Southern Brazil and Northeastern Argentina.
- Pillansia, a single species from Southwestern Cape Province.
- Pseudotrimezia, 18 species from Southeastern Brazil.
- Radinosiphon, two species from Tanzania to Southern Africa.
- Romulea, about 105 species widely distributed from Macaronesia, Mediterranean Europe, tropical African mountains to Southern Africa and Arabian Peninsula.
- Savannosiphon, a single species distributed from Tanzania to southern tropical Africa.
- Sisyrinchium, about 200 species which range from Hawaiian Islands, Template and Subtropical America to Falkland Islands.
- Solenomelus, two species from Chile and Argentina.
- Sparaxis, 15 species from Southern Africa.
- Syringodea, 7 species from Southern Africa.
- Tapeinia, a single species from southern Chile and Argentina.
- Thereianthus, 8 species from Southwestern Cape Province.
- Tigridia (incl. Ainea, Colima, Fosteria, Rigidella, Sessilanthera), about 50 species distributed from Mexico to El Salvador, and Peru to Northern Chile.
- Trimezia, 41 species from Lesser Antilles to southern tropical America
- Tritonia, 26 species distributed from Tanzania to Southern Africa.
- Tritoniopsis (inc. Anapalina), 23 species from Cape Province.
- Watsonia, 52 species from Southern Africa.
- Witsenia, a single species from Southwestern Cape Province.
- Xenoscapa, two species from Namibia to Cape Province.
- Zyg otritonia, four species from tropical Africa.
Ecology
Members of the Iridaceae occur in a great variety of habitats. About the only place they do not grow is in the sea itself, although Gladiolus gueinzii occurs on the seashore just above the high tide mark within reach of the spray. Most species are adapted to seasonal climates that have a pronounced dry or cold period unfavorable for plant growth and during which the plants remain dormant. As a result most species are deciduous. Evergreen species are restricted to subtropical forests or savannah, temperate grasslands and perennially moist fynbos. A few species grow in marshes or along streams and some even grow only in the spray of seasonal waterfalls.[3]
The above-ground parts (leaves and stems) of deciduous species die down when the bulb or corm enters dormancy. The plants thus survive periods that are unfavorable for growth by retreating underground. This is particularly useful adaptation for growth in areas like grasslands and fynbos which regularly have fires in the dry seasons? the plants are dormant and their bulbs or corms are able to survive underground. Veld fires clear the soil surface of competing vegetation and fertilize it with ash. With the arrival of the first rains, the dormant corms are ready to burst into growth, sending up flowers and stems before they can be shaded out by other vegetation. Many Iridaceae species which grow in grassland and fynbos flower best after fires and some fynbos species will only flower in the season after a fire.[3]
The family has a very diverse pollination ecology. Although the majority of species of African Iridaceae are pollinated by Hymenoptera (mostly bees), the remaining species are pollinated mainly, or solely, by inse cts in the orders Coleoptera (beetles), Diptera (short- and long-proboscid flies) and Lepidoptera (butterflies and moths), or by passerine birds (Nectarinidae). It is now known that pollination systems are predominantly specialized: plants rely on a single species or a few ecologically analogous species for pollination. By contrast, generalist species, which are pollinated by a range of pollinators from at least three pollinator groups, are rare among southern African Iridaceae. In consequence, almost all genera of any size exhibit a range of pollination syndromes, with similar patterns of floral variation having developed repeatedly within different genera.[32]
Most significantly, the diversity of pollination systems increases primarily with floral complexity and secondarily with genus size. Thus, Aristea (approximately 56 species), which has radially symmetric, mostly blue flowers, has three different po llination systems, whereas Sparaxis (15 species), with both zygomorphic and secondarily radially symmetric flowers, in a variety of colors, exhibits five different pollination systems, and Gladiolus, with a similar array of floral types but over two hundred species, exploits seven different pollination systems, some of which have evolved multiple times. As is usual in predominantly specialist pollination systems, floral attractants and rewards correlate closely with pollinator profile, resulting in the development of distinct floral syndromes. Attractants are primarily perianth pigmentation, complemented by a range or floral odors in many species, but flower shape and tepal orientation, in particular functional floral symmetry, may be equally important for some pollinators. The reward to visitors in the majority of species is nectar, but in others it is pollen, and one species offers non-volatile oil. In the case of hopliine beetles (Scarabaeidae, Hopliinae), flowers provide a stable platform on which to congregate, and the value of pollen, which beetles sometimes consume, as a reward is uncertain.[32]
Conservation
Several species of Iridaceae are endangered or threatened by extinction due to habitat degradation or loss and a restricted geographical distribution. According to the IUCN, the following species are endangered: Babiana longicollis, Cyanixia socotrana (as Babiana socotrana), Gladiolus pole-evansii, Moraea garipensis, Moraea graniticola and Moraea hexaglottis.[33]
Uses
Several cultures have used species of Iridaceae as food, ornamental, condiment or medicinal plants. The Navajo, the largest Native America n tribe of North America, used decoctions of Iris missouriensis as an emetic.[34] Pieces of the rhizome of the same species were used to relieve toothaches,[35] or earaches.[36] The mashed roots of Iris versicolor were applied to wounds, presumably as an antiseptic,[37] and the infusions of dry roots of the same species were used to calm pain.[38] Sisyrinchium acre was used in Hawaii in different ways. Leaves or leaf-sap were used as a dye, to give the blue color to tattoos. The use of the leaves, macerated with salt, s ugar and other spices was recommended to clean the skin and cure skin diseases.[39] Iris ensata was used in India as anthelmintic and diuretic, and, mixed with other species, to treat venereal diseases.[40] Belamcanda chinensis has a long history of use as medicinal plant in China. Apparently, it was very effective in controlling bacterial, viral and fungal diseases, and in reducing fever and inflammations.[41] Another popular member of the family is Crocus sativus; the spice saffron ? obtained from the stigma ? has been used for centuries in folk medicine as an antispasmodic, aphrodisiac, expectorant, narcotic and sedative.[ 42]
Iridaceae are of considerable economic importance in ornamental horticulture and the cut-flower industry, especially Iris, Gladiolus, and Freesia. Several other genera (e.g., Crocus, Dietes, Sparaxis, Tritonia, Watsonia) are cultivated in gardens in both tropical and temperate areas. Moraea and Homeria are poisonous and pose significant problems in cattle- and sheep-raising areas, notably in southern Africa.
is a family of perennial, herbaceous and bulbous plants included in the monocot order Asparagales, taking its name from the genus Iris. Almost worldwide in distribution and one of the most important families in horticulture, it includes more than 2000 species. Genera such as Crocus and Iris are significant components of the floras of parts of Eurasia, and Iris also is well-represented in North America. Gladiolus and Moraea are large genera and major constituents of the flora of sub-Saharan and South ern Africa. Sisyrinchium, with more than 140 species, is the most diversified Iridaceae genus in the Americas, where several other genera occur, many of them important in tropical horticulture.1]All members of Iridaceae have petaloid, soft-textured and colorful perianths in which the three tepals of the inner whorl and the three of the outer whorl are alike in structure, shape, and often in color. This type of corolla, a feature of all families of Asparagales as well as the related order Liliales, distinguishes them from other monocots, such as grasses, palms or reeds, in which the perianth is either reduced or with the members of one or both of the whorls firm-textured and dry and often brown or green. The character that sets Iridaceae apart from other plants of the Asparagales or from the Liliales, is the male part of the flower, the androecium, which has three stamens?in most related families there are s ix. Another character that distinguish Iridaceae from most other members of the Asparagales is its inferior ovary, the ovary being superior in most families, with the exception of Orchidaceae and Amaryllidaceae. Apart from their flowers, the Iridaceae can usually be recognized by their characteristic leaves, sword-like and oriented edgewise to the stem and with two identical surfaces. Such leaves are termed isobilateral and unifacial. In contrast to the Irids, typical plant leaves ? termed dorsiventral and bifacial ? have upper and lower surfaces of different appearance and anatomy.
The Iridaceae originated in Antarctica-Australasia in the late Cretaceous, about 82 million years ago, although the family's subsequent radiation occurred elsewhere, notably in southern Africa and temperate and highland South America, at the end of the Eocene or later.
Currently, 66 genera are recognized, which are distributed among 7 subfamilies and occur in a great variety of habitats. Mos t species are adapted to seasonal climates that have a pronounced dry or cold period unfavourable for plant growth, during which the plants remain dormant. As a result, most species are deciduous, in that their above-ground parts (leaves and stems) die down when the bulb or corm enters dormancy. The plants thus survive periods that are unfavourable for growth by retreating underground. Evergreen species are restricted to subtropical forests or savannah, temperate grasslands and perennially moist fynbos.
Description
In the Iridaceae the perianth is formed of two whorls of three tepals, all similar in structure, shape, and often color. Such a corolla differentiates Liliales and Asparagales from other monocots, where the number or size of the tepals are reduced, or where at least one whorl is papyraceous (firm and dry like a papyrus), and usually green or brown. The characters that differentiate Iridaceae, however, are its three stam en (related families such as Alliaceae and Amaryllidaceae have six) and an inferior ovary. Sword-like leaves parallel to the stem and with normally undifferentiated sides (termed "isobilateral" and "unifacial") are another distinctive feature.
Members of Iridaceae are herbs or, in a few cases, shrubs with woody caudex. They are almost all perennial (three Sisyrinchium species are annuals) that may be either evergreen or seasonal. The rootstock is a rhizome, bulb, or corm.[2] The leaves are found both at the base and on the stem, usually alternate, with the blade oriented parallel to the stem and thus sheathing it at the base. This results in the characteristic fan-like arrangement found in genera like Iris. This type of leaf lacks distinct upper and lower leaf surfaces. In many South African species the leaf has a thickened midrib and often variously thickened or winged margins that may also be crisped. In some species the leaves are needle-like with narrow longitudinal grooves. Species of Moraea are unusual in the family in having channeled leaves with a distinct upper and lower surface.[2][3]
Flowers may be either actinomorphic or zygomorphic. Almost all the parts are in threes, starting with two equal whorls of three usually large and showy petal-like tepals, distinct or fused in a tube. There are three stamens (rarely two), and their filaments are often partly to completely fused. Anthers have two pollen sacs opening toward the outside, or from their side, and usually along their length. The ovary is located below the tepals (except in Isophysis) with axile (rarely parietal) placentation in three locules. There is a single style branching into three at the top. Iridaceae do not present unisexual flowers, and all flowers have both a style and stamen. Most members other than Sisyrinchium produce nectar from nectaries at the base of the tepals, or on the gynoecium. Iridaceae species are usually pollinated by insects or birds.[1][2] The flowers are collectively arranged in two different types of inflorescences. Simple or branched spikes occur in all Crocoideae. In other subfamilies the basic inflorescence unit is a type of zig-zagging cyme called rhipidium, which is enclosed in enlarged, opposed, bracts called spathes.[1][2]
The fruit is a dry capsule, usually splitting along three sides spontaneously at maturity. It is very variable in shape and texture, from firm t o cartilaginous, occasionally woody. In most genera they are tetrahedral or variously angled and without obvious adaptations for dispersal.[3] Seeds are also varied in shape.[4] Winged seeds adapted to wind dispersal characterize Gladiolus and Tritoniopsis and also occur in some species of Hesperantha. Globular seeds with shiny coats that are relatively long-lived occur in several genera of Crocoideae. Chasmanthe aethiopica has fleshy seeds adapted to dispersal by birds and several other species that grow in more wooded places ? like Chasmanthe and some freesias ? have reddish or black seeds that mimic fleshy seeds. They have a hard endosperm, with reserves of hemicellulose, oil, and protein, and a small embryo.[2][5][6]
Distribution and habitat
Members of the Iridaceae are nearly worldwide in distribution, but remain rare in tropical lowlands and at high latitudes. The family is best represented in Southern Africa, especially the winter-rainfall region in the southwest. Other centers of diversity are temperate South and Central America (with several small genera) and the Mediterranean region (Iris and Crocus). About 2000 species are distributed among some 65 genera are recognized worldwide, just over half of them from Southern Africa, where 38 genera are known. In the Cape Floral Region alone, 707 species and 27 genera are recorded.[1][3]
The family prefers open, seasonal habitats. In Africa, the montane grasslands of eastern South Africa, Swaziland and Lesotho, and the Succulent Karoo and fynbos of the Northern and Western Cape have the most species. Fewer species occur in savannas or the semi-arid central karoo, and very few in forests.[3] The species grows in a variety of soils, derived from basalt, clay, dolerite, granite, limestone and sandstone, as well as rarer rocks like serpentine. Most species favor loamy soils, often among rocks where drainage is good, but some grow in marshes and others in pure sand. Species grow from just above the high tide mark to over 3,000 metres (9,843 ft) above sea level.[3]
Taxonomy
The family name is based on the genus Iris, the largest and best known genus in Europe. Iris dates from 1753, when it was named by Swedish botanist Carl Linnaeus. Its name derives from the Greek goddess, Iris, who carried messages from Olympus to earth along a rainbow, whose colors were seen by Linnaeus in the multi-hued petals of many of the species.[3] The family name is attributed to Antoine Laurent de Jussieu's 1789 Genera Plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam, and is a conserved name, so that even if an earlier name were to be discovered for the family, Iridace ae would remain valid.[7]
The family has been accepted in all major classification systems of the 20th century. The Cronquist system treated it as part of the order Liliales of the subclass Liliidae,[8] the Takhtajan system placed it in an order Iridales, together with Isophysidaceae and Geosiridaceae treated as single-genus families,[9] and the Thorne system treated it as part of the order Orchidales in its own suborder, Iridineae.[10] The Angiosperm Phylogeny Group in 1998 and 2003 (APG and APG II, respectively) system of flowering plant classification organizes flowering plants into a "selected number of monophyletic suprafamilial groups" and placed Iridaceae in the order Asparagales, which was part of a clade called "Non Commelinoid Monocots".[11][12]
Evolution and phylogeny
The Iridaceae differentiated in the late Cretaceous, about 82 million years ago, and diverged from the next most closely related family, the Doryanthaceae, during the Campanian. The Tasmanian Isophysis is the only extant member of the clade sister to the remainder of the Iridaceae, from which it may have diverged 66 million years ago, in the Maastrichtian. The Iridaceae originated in Antarctica-Australasia, although its subsequent radiation occurred elsewhere, notably in southern Africa and temperate and highland South America at the end of the Eocene or later.[13]
Paleopolyploidy was important in the early diversification of Iridaceae since many genera have basic chromosome numbers which are derived from ancestors with fewer chromosomes. The appearance of polyploidy is important in Northern Hemisphere genera, especially Iris and Crocus, but has an unusually low frequency in Africa, the center of diversity for the family. Changes in basic number, frequent in a few genera, are the result of Robertsonian translocations which led to a reduction in the number of chromosomes.[a] In all but a few possible examples, accompanying morphological specialization suggests that reduction in the number of chromosomes (dysploid reduction) is involved in gradual changes in base number. Major dysploid series are restricted to a few genera, such as Romulea, the related Crocus as well as Gladiolus and Lapeirousia (all members of subfamily Crocoideae), and Iris, Moraea, and Sisyrinchium (Iridoideae).[14] Com plex translocation heterozygotes have been discovered in three species of Homeria (currently included in Gladiolus), a genus from South Africa. In these species, almost every chromosome of the plant has exchanged an arm with another chromosome, which results in a chain of translocated chromosomes. These unusual species, H. tenuis (2n = 10, 9), H. flavescens (2n = 9), and H. pallida (2n = 8) are the only aneuploids in the genus; H. pallida also has forms with 2n = 12. The basic chromosome number in Homeria is x = 6 and most species are diploid, a few other polyploid. The genus comprises mostly large-flowered outcrossing species, but self-compatibility and autogamy occur in several, including the three with complex heterozygosity.[15]
Several studies based on morphology and DNA sequence information have shown that the Iridaceae are a monophyletic group incl uded in the order Asparagales.[16][17][18][19] Phylogenetic analyses of sequences of five plastid DNA regions, rbcL, rps4, trnL?F, matK, and rps16, confirm most aspects of the traditional classification of the Iridaceae in four subfamilies and the evolutionary patterns that they imply, importantly the sister relationship of Isophysidoideae to the remainder of the family and the monophyly of Iridoideae. However, it was shown that subfamily Nivenioideae as traditionally defined is paraphyletic: Crocoideae was consistently found nested within it, sister to the core Nivenioideae (the woody genera Klattia, Nivenia, and Witsenia).[1] This clade is sister to Aristea, which in turn is sister to the Madagascarian Geosiris, and then to the Australasian Patersonia.[13][20] This has led to a shift in classification.
Subfamilies
Based on the morphology, anatomy, embryology and chromosome numbers, the family traditionally was divided into four subfamilies (Isophysidodeae, Nivenioideae, Iridioideae and Ixioideae)[21][22] but the results from DNA analysis suggest that several more should be recognized. In fact, the genera Aristea, Geosiris, and Patersonia are now considered as separate subfamilies: Aristeoideae, Geosiridoideae and Patersonioideae, respectively, rendering Nivenioideae and Crocoideae monophyletic.[13][23] Only Iridoideae and Crocoideae are currently subdivided, each into several tribes.
- Isophysidoideae contains the single genus Isophysis, from Tasmania. It is the only member of the family with a superior ovary. It has a star-like yellow to brownish flower.
- Nivenioideae contains three genera (Klattia, Nivenia and Witsenia) and 14 species restricted to the Cape region of South Africa. The plants in this subfamily are true shrubs, with secondary thickening. It is distinguished by having sessile flowers in small, paired clusters surrounded by large bracts, sl ender styles that are divided into three branches, and nectar produced from glands in the ovary walls. The flowers are always radially symmetrical, with unfused tepals and the rootstock is a rhizome. The basic chromosome number is x= 16.
- Iridoideae, includes 30 genera and about 820 species, is distributed throughout the range of the family, and contains the large genera Iris and Moraea. It is the only subfamily that is represented in South America. The species have flowers in solitary clusters among large bracts, styles that are often petal-like or crested, and nectar (when present) is produced from glands on the tepals. Most species have unfused tepals and the rootstock is usually a rhizome or rarely a bulb. The flowers are almost always radially symmetrical. Bobartia, Dietes and Ferraria belong to this subfamily.
- Patersonioideae contains only one genus (Patersonia) with about 20 species distributed in Sumatra, Borneo, New Guinea, a nd the periphery of Australia. Plants are more or less woody and rhizomatous with secondary growth. Their flowers are blue and last only for one day. The inner tepals in the flowers are reduced to scales, the filaments are more or less fused. The basic chromosome number is x= 11 or 21.
- Geosiridoideae is represented by only one genus with one species: Geosiris, the only myco-heterotrophic genus of the family, from Madagascar and the Comores. The plants are parastic and lack chlorophyll. They have sessile flowers with the tepals fused at the base, without nectary. They produce minute, dust-like seeds.
- Aristeoideae contains only one genus, Aristea, with about 55 species in sub-Saharan Africa and Madagascar. They are rhizomatous plants with blue flowers which usually last one day, nectaries, and a basic chromosome number x= 16.
- Crocoideae (syn.: Ixioideae), which contains nearly half of the species of the family (28 genera and 995 species), is mostly African. This subfamily contains most of the familiar genera (other than Iris and Moraea) such as Ixia, Gladiolus, Crocus, Freesia and Watsonia. It is easily recognised by its spike-like inflorescence (sometimes bearing solitary flowers) and its tepals joined into a short or long tube. Nectar is produced from glands in the ovary wall and is secreted directly into the base of the floral tube. The flowers are either radially symmetrical or more usually bilaterally symmetrical and two-lipped. The rootstock is a rhizome or, more commonly, a corm.
Genera
Up to 66 genera have been recognised in the family, with a total of around 2,000 species worldwide. The Afrotropic ecozone, and in particular South Africa, have the greatest diversity of genera. DNA sequence information coupled with some associated morphological features provided evidence that several previously recognised New World Tigridieae genera could not retain a generic rank. These are Ainea, Colima, Fosteria, Rigidella and Sessilanthera, which are currently included in Tigridia. The Bolivian genus Cardenanthus was subsumed in Mastigostyla, and Onira and Kelissa were included in the temperate South American genus Cypella.[1] For the same reason, Tamia was included in Calydorea and Tucma in Ennealophus.[1][24]
The genera of Iridaceae, their number of species and geographic distribution are given below.[1][21][22][25][26]
- Afrocrocus [b], one species from Africa.
- Alophia (including Eustylis),[27] five species from the Central and Southern U.S.A. to Brazil.
- Aristea, 56 species distributed in tropical and Southern Africa and Madagascar.
- Babiana, 90 species from tropical to Southern Africa.
- Bobartia, 15 species endemic to Cape Province.
- Calydorea (incl.: Cardiostigma, Itysa, Salpingostylis[28] and Tamia), 21 species from tropical and sub-tropical South America.
- Cha smanthe, three species endemic to Cape Province.
- Cipura, 8 species from Mexico to tropical South America.
- Cobana, one species from Central America.
- Crocosmia, 8 species distributed from Sudan to Southern Africa and Madagascar.
- Crocus, 90 species from Mediterranean Europe to Northwestern China.
- Cyanixia, one species from Socotra.
- Cypella (incl. Onira and Kelissa), 22 species distributed from Peru, Brazil to Northern Argentina.
- Devia, one species endemic to Western Cape.
- Dierama, 43 species from Ethiopia to Southern Africa.
- Dietes, 6 species from Ethiopia to Southern Africa and Lord Howe Island.
- Diplarrhena, two species endemic to Southeastern Australia.
- Duthieastrum, a single species from Northern Cape Province to Free State.
- Eleutherine, two species distributed from Mexico to tropical South America.
- Enne alophus (incl. Tucma and Eurynotia), about 7 species from Ecuador to Northern Brazil and Northwestern Argentina.
- Ferraria, 13 species from southern Za?re to South Africa.
- Freesia (syn. Anomatheca, Lapeirousia), about 56 species distributed from Kenya to Southern Africa.
- Geissorhiza, 85 species endemic to Cape Province.
- Gelasine, 7 species from Brazil to Northeastern Argentina.
- Geosiris, a single species endemic from Madagascar.
- Gladiolus (inc. Oenostachys, Homoglossum, Anomalesia, Antholyza and Acidanthera),[29] 272 species distributed from Europe to Iran, Arabian Peninsula and Africa.
- Herbertia (inc. Sympa), 8 species distributed from Texas to Louisiana, Colombia to Venezuela, Bolivia to southern South America.
- Hesperantha (inc. S chizostylis), 83 species from Cameroon and Ethiopia to Southern Africa.
- Hesperoxiphion, 5 species distributed in Western South America.
- Iris (inc. Belamcanda),[30] includes 260 species distributed from temperate Northern Hemisphere to Philippines.
- Isophysis, a single species from Tasmania.
- Ixia, 61 species from Cape Province.
- Klattia, three species from Southwestern Cape Province.
- Larentia, two species from Mexico, Venezuela, Bolivia to Southern Brazil.
- Lethia, a single species from Bolivia to Brazil (Minas Gerais).
- Libertia, 15 species from Andes to southern South America, and Australia and New Zealand.
- Mastigostyla (incl. Cardenanthus), 17 species from Peru to Northern Argentina.
- Melasphaerula, a single species from Namibia to Cape Province.
- Micra nthus, three species from Cape Province.
- Moraea (inc. Bernardiella, Galaxia, Gynandiris, Hexaglottis, Homeria, Sessilstigma and Roggeveldia).[31] includes about 200 species distributed from tropical and South Africa, Mediterranean Europe to western Himalaya.
- Nemastylis, 7 species distributed from Central U.S.A. to Honduras.
- Neomarica, 21 species from Mexico to Trinidad, Brazil to Northeastern Argentina.
- Nivenia, 11 species endemic to Southwestern Cape Province.
- Olsynium, 15 species distributed from western Canada and U.S.A., Costa Rica to Southern South America.
- Orthrosanthus, 9 species from Southern Australia, Mexico to Northwestern Argentina.
- Patersonia, 19 species distributed from Malesia to Australia.
- Phalocallis, a single species from Southern Br azil and Northeastern Argentina.
- Pillansia, a single species from Southwestern Cape Province.
- Pseudotrimezia, 18 species from Southeastern Brazil.
- Radinosiphon, two species from Tanzania to Southern Africa.
- Romulea, about 105 species widely distributed from Macaronesia, Mediterranean Europe, tropical African mountains to Southern Africa and Arabian Peninsula.
- Savannosiphon, a single species distributed from Tanzania to southern tropical Africa.
- Sisyrinchium, about 200 species which range from Hawaiian Islands, Template and Subtropical America to Falkland Islands.
- Solenomelus, two species from Chile and Argentina.
- Sparaxis, 15 species from Southern Africa.
- Syringodea, 7 species from Southern Africa.
- Tapeinia, a single species from southern Chile and Argentina.
- Thereianthus, 8 species from Southwestern Cape Province.
- Tigridia (incl. < i>Ainea, Colima, Fosteria, Rigidella, Sessilanthera), about 50 species distributed from Mexico to El Salvador, and Peru to Northern Chile.
- Trimezia, 41 species from Lesser Antilles to southern tropical America
- Tritonia, 26 species distributed from Tanzania to Southern Africa.
- Tritoniopsis (inc. Anapalina), 23 species from Cape Province.
- Watsonia, 52 species from Southern Africa.
- Witsenia, a single species from Southwestern Cape Province.
- Xenoscapa, two species from Namibia to Cape Province.
- Zygotritonia, four species from tropical Africa.
Ecology
Members of the Iridaceae occur in a great variety of habitats. About the only place they do not grow is in the sea itself, although Gladiolus gueinzii occurs on the seashore just above the high tide mark within reach of the spray. Mos t species are adapted to seasonal climates that have a pronounced dry or cold period unfavorable for plant growth and during which the plants remain dormant. As a result most species are deciduous. Evergreen species are restricted to subtropical forests or savannah, temperate grasslands and perennially moist fynbos. A few species grow in marshes or along streams and some even grow only in the spray of seasonal waterfalls.[3]
The above-ground parts (leaves and stems) of deciduous species die down when the bulb or corm enters dormancy. The plants thus survive periods that are unfavorable for growth by retreating underground. This is particularly useful adaptation for growth in areas like grasslands and fynbos which regularly have fires in the dry seasons? the plants are dormant and their bulbs or corms are able to survive underground. Veld fires clear the soil surface of competing vegetation and fertilize it with ash. With the arrival of the first rains, the dormant corms are ready to burst into growth, sending up flowers and stems before they can be shaded out by other vegetation. Many Iridaceae species which grow in grassland and fynbos flower best after fires and some fynbos species will only flower in the season after a fire.[3]
The family has a very diverse pollination ecology. Although the majority of species of African Iridaceae are pollinated by Hymenoptera (mostly bees), the remaining species are pollinated mainly, or solely, by insects in the orders Coleoptera (beetles), Diptera (short- and long-proboscid flies) and Lepidoptera (butterflies and moths), or by passerine birds (Nectarinidae). It is now known that pollination systems are predominantly specialized: plants rely on a single species or a few ecologically analogous species for pollination. By contrast, generalist species, which are pollinated by a rang e of pollinators from at least three pollinator groups, are rare among southern African Iridaceae. In consequence, almost all genera of any size exhibit a range of pollination syndromes, with similar patterns of floral variation having developed repeatedly within different genera.[32]
Most significantly, the diversity of pollination systems increases primarily with floral complexity and secondarily with genus size. Thus, Aristea (approximately 56 species), which has radially symmetric, mostly blue flowers, has three different pollination systems, whereas Sparaxis (15 species), with both zygomorphic and secondarily radially symmetric flowers, in a variety of colors, exhibits five different pollination systems, and Gladiolus, with a similar array of floral types but over two hundred species, exploits seven different pollination systems, some of which have evolved multiple times. As is usual in predominantly specialist pollination systems, floral attractants and rewards correlate closely with pollinator profile, resulting in the development of distinct floral syndromes. Attractants are primarily perianth pigmentation, complemented by a range or floral odors in many species, but flower shape and tepal orientation, in particular functional floral symmetry, may be equally important for some pollinators. The reward to visitors in the majority of species is nectar, but in others it is pollen, and one species offers non-volatile oil. In the case of hopliine beetles (Scarabaeidae, Hopliinae), flowers provide a stable platform on which to congregate, and the value of pollen, which beetles sometimes consume, as a reward is uncertain.[32]
Conservation
Several species of Iridaceae are endangered or threatened by extinction due to habitat deg radation or loss and a restricted geographical distribution. According to the IUCN, the following species are endangered: Babiana longicollis, Cyanixia socotrana (as Babiana socotrana), Gladiolus pole-evansii, Moraea garipensis, Moraea graniticola and Moraea hexaglottis.[33]
Uses
Several cultures have used species of Iridaceae as food, ornamental, condiment or medicinal plants. The Navajo, the largest Native American tribe of North America, used decoctions of Iris missouriensis as an emetic.[34] Pieces of the rhizome of the same species were used to relieve toothaches,[35] or earaches.[36] The mashed roots of Iris versicolor were applied to wounds, presumably as an antiseptic,[37] and the infusions of dry roots of the same species were used to calm pain.[38] Sisyrinchium acre was used in Hawaii in different ways. Leaves or leaf-sap were used as a dye, to give the blue color to tattoos. The use of the leaves, macerated with salt, sugar and other spices was recommended to clean the skin and cure skin diseases.[39] Iris ensata was used in India as anthelmintic and diuretic, and, mixed with other species, to treat venereal diseases.[40] Belamcanda chinensis has a long history of use as medicinal plant in China. Apparently, it was very effective in controlling bacterial, viral and fungal diseases, and in reducing fever and inflammations.[41] Another popular member of the family is Crocus sativus; the spice saffron ? obtained from the stigma ? has been used for centuries in folk medicine as an antispasmodic, aphrodisiac, expectorant, narcotic and sedative.[42]
Iridaceae are of considerable economic importance in ornamental horticulture and the cut-flower industry, especially Iris, Gladiolus, and Freesia. Several other genera (e.g., Crocus, Dietes, Sparaxis, Tritonia, Watsonia) are cultivated in gardens in both tropical and temperate areas. Moraea and Homeria are poisonous and pose significant problems in cattle- and sheep-raising areas, notably in southern Africa.
Notes
References
- ^ a b c d e f g h i Goldblatt, Peter; John Manning (2008). The Iris Family: Natural History & Classification. Portland: Timber Press. ISBN 978-0-88192-897-6.
- ^ a b c d e Goldblatt, Peter (2002). Iridaceae "Iridaceae". Flora of North America. Volume. 26: Magnoliophyta: Liliidae: Liliales and Orchidales. New York: Oxford University Press. p. 348. ISBN 0-19-515208-5. http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=1 0452 Iridaceae. Retrieved 2009-05-12.
- ^ a b c d e f g h Manning, J. (2004). "Iridaceae". South African National Biodiversity Institute. http://www.plantzafrica.com/planthij/iridaceae.htm. Retrieved 2009-05-12.
- ^ Zhao, Yu-tang; Henry J. Noltie & Brian F . Mathew (2000). "Iridaceae". In Wu Zheng-yi & Peter Hamilton Raven (eds.). Flora of China. Volume. 24: Flagellariaceae through Marantaceae. St. Louis: Missouri Botanical Garden. p. 297. ISBN 0-915279-83-5. http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=10452. Retrieved 2009-05-12.
- ^ Dimitri, M. 1987. Enciclopedia Argentina de Agricultura y Jardiner?a. Tomo I. Descripci?n de plantas cultivadas. Editorial ACME S.A.C.I., Buenos Aires.
- ^ Watson, Leslie (2008). "Iridaceae". Florabase, The Western Australia Flora. Western Australia Department of Environment and Conservatio n. http://florabase.calm.wa.gov.au/browse/profile/22784. Retrieved 2009-05-12.
- ^ J. Mcneil et al. (2006). "Nomina Familiarum Bryophytorum et Spermatophytorum Conservanda". International Code of Botanical Nomenclature (VIENNA CODE). http://ibot.sav.sk/icbn/main.htm. Retrieved 2009-05-24.
- ^ Arthur Cronquist. 1988. The Evolution and Classification of Flowering Plants
- ^ A. Takhtajan (1980). "Outline of the classification of flowering plants (Magnoliophya)". Botanical Review 46 (3): 225?359. doi:10.1007/BF02861558.
- ^ Robert F. Thorne. 1992. Classification and Geography of Flowering Plants. Botanical Review 58: 225-348
- ^ APG, 1998. An ordinal Classification for the Families of Flowering Plants. Annals of the Missouri Botanical Garden 85: 531-553
- ^ Angiosperm Phylogeny Group (2003). Full text An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 41: 399-436.
- ^ a b c Goldblatt, P., Aaron Rodriguez, M. P. Powell, T. Jonathan Davies, John C. Manning, M. van der Bank, Vincent Sav olainen. 2008. Iridaceae ?Out of Australasia?? Phylogeny, Biogeography, and Divergence Time Based on Plastid DNA Sequences.Systematic Botany 33:3, 495-508
- ^ Peter Goldblatt and Masahiro Takei. 1997. Chromosome Cytology of Iridaceae-Patterns of Variation, Determination of Ancestral Base Numbers, and Modes of Karyotype Change. Annals of the Missouri Botanical Garden, Vol. 84, No. 2, pp. 285-304
- ^ Peter Goldblatt. 1980. Uneven Diploid Chromosome Numbers and Complex Heterozygosity in Homeria (Iridaceae). Systematic Botany, Vol. 5, No. 4, pp. 337-340
- ^ Chase M. W.. Duvall, M. R., Hills, H. G., Conran, J. G., Cox, A. V., Egu iarte, L. E., Hartwell, J., Fay, M. F., Caddick, L. R., Cameron, K. M., y Hoot, S. 1995. Molecular systematics of Lilianae. In: Rudall, P. J., Cribb, P. J., Cutler, D. F. (eds.) Monocotyledons: Systematics and evolution. Royal Botanic Gardens, Kew, pp. 109-137
- ^ Chase, M. W.,Soltis, D. E., Soltis, P. S., Rudall, P. J., Fay, M. F., Hahn, W. H., Sullivan, S., Joseph, J., Molvray, M., Kores, P. J., Givnish, T. J., Sytsma, K. J., y Pires, J. C. 2000. Higher-level systematics of the monocotyledons: An assessment of current knowledge and a new classification. In.: Wilson, K. L. & Morrison, D. A. (eds.) Monocots: Systematics and Evolution.CSIRO Publ., Collingwood, Australia, pp,:3-16
- ^ Stevenson,D. W. & Loconte, H. 1995. Cladistic analysis of monocot families. In.: Rudall, P. J., C ribb, P. J., Cutler, D. F. (Eds.) Monocotyledons: Systematics and evolution, Royal Botanic Gardens, Kew, pp.: 543-578
- ^ Souza-Chies,T. T., Gabriel Bittar, Sophie Nadot, Leigh Carter, Evelyne Besin and Bernard Lejeune. 1997. Phylogenetic analysis ofIridaceae with parsimony and distance methods using the plastid generps4. Plant Systematics and Evolution 204(1): 109-123
- ^ Tatiana T. Souza-Chies, Gabriel Bittar, Sophie Nadot, Leigh Carter, Evelyne Besin and Bernard Lejeune. 1997. Phylogenetic analysis of Iridaceae with parsimony and distance methods using the plastid generps4. Plant Systematics and Evolution 204(1): 109-123
- ^ a b Goldblatt, P. Phylogeny and Classification of Iridaceae. Annals o f the Missouri Botanical Garden, Vol. 77, No. 4 (1990), pp. 607?627.
- ^ a b Goldblatt, P. 1991. An overview of the systematics, phylogeny and biology of the southern African Iridaceae. Contr. Bolus Herb. 13: 1?74.
- ^ Stevens, P. F. 2001 onwards. Angiosperm Phylogeny Website. Downloaded 2009-5-19.
- ^ De Tullio, L., Germ?n Roitman, Gabriel Bernardello. 2008. Tamia (Iridaceae), a Synonym of Calydorea: Cytological and Morphological Evidence. Systematic Botany 2008 33:3, 509?513
- ^ Reeves, G., Chase, M.W., Goldblatt, P., Rudall, P., Fay, M.F., Cox, A.V., LeJeune, B., y Souza-Chies, T., (2001). Molecular systematics of Iridaceae: Evidence from four plastid DNA regions.. Amer. J. Bot. 88:2074-2087.
- ^ Royal Horticultural Society, Kew. A detailed checklist for family Iridaceae.
- ^ R. K. Brummitt. 1980. Proposal to conserve Alophia over Eustylis. Report of the Committee for Spermatophyta, 22. Taxon, Vol. 29, No. 4 (August, 1980), pp. 489?493
- ^ Goldblatt, P. y J.E. Henrich. Calydorea Herbert (Iridaceae-Tigridieae): Notes on this New World Genus and Reduction to Synonymy of Salping ostylis, Cardiostigma, Itysa, and Catila. Annals of the Missouri Botanical Garden, Vol. 78, No. 2 (1991), pp. 504?511
- ^ Goldblatt, P. & De Vos M. P. The reduction of Oenostachys, Homoglossum and Anomalesia, putative sunbird pollinated genera, in Gladiolus L. (Iridaceae-Ixioideae). Bulletin du Mus?um national d'histoire naturelle. Section B, Adansonia 11 (4): 417?428, 1989.
- ^ Goldblatt, P. & David J. Mabberley. Belamcanda included in Iris, and the New Combination I. domestica (Iridaceae: Irideae). Novon 15 (1), pp. 128?132, (2005).
- ^ Goldblatt, P. Reduction of Bernardiella, Galaxia, Gyna ndriris, Hexaglottis, Homeria, and Roggeveldia in Moraea (Iridaceae: Irideae). Novon, Vol. 8, No. 4 (1998), pp. 371?377
- ^ a b Goldblatt, P. & J.C. Manning. 2006. Radiation of Pollination Systems in the Iridaceae of sub-Saharan Africa. Ann Bot 97: 317-344.
- ^ IUCN 2008. [<www.iucnredlist.org> 2008 IUCN Red List of Threatened Species.] Downloaded on 19 May 2009.
- ^ Vestal, Paul A. 1952 The Ethnobotany of the Ramah Navaho. Papers of the Peabody Museum of American Archaeology and Ethnology 40(4):1?94 (p. 21).
- ^ Nickerson, Gifford S. 1966 Some Data on Plains and Great Basin Indian Uses of Certain Native Plants. Tebiwa 9(1):45?51 (p. 47)
- ^ Train, Percy, James R. Henrichs and W. Andrew Archer 1941 Medicinal Uses of Plants by Indian Tribes of Nevada. Washington DC. U.S. Department of Agriculture (p. 89, 90)
- ^ Raymond, Marcel. 1945 Notes Ethnobotaniques Sur Les Tete-De-Boule De Manouan. Contributions de l'Institut botanique l'Universite de Montreal 55:113?134 (p. 129))
- ^ Speck, Frank G., R.B. Hassrick and E.S. Carpenter 1942 Rappahannock Herbals, Folk-Lore and Science of Cures. Proceedings of the Delaware County Institute of Science 10:7?55. (p. 28)
- ^ Sisyrinchium acre - Hawaiian Ethnobotany Online Database
- ^ Chopra. R. N., Nayar. S. L. and Chopra. I. C. Glossary of Indian Medicinal Plants (Including the Supplement). Council of Scientific and Industrial Research, New Delhi. 1986
- ^ Bown. D. Encyclopaedia of Herbs and their Uses. Dorling Kindersley, London. 1995 ISBN 0-7513-0203-1
- ^ Chevallier. A. The Encyclopedia of Medicinal Plants. Dorling Kindersley. London 1996 ISBN 978-0-7513-0314-8
Bibliography
- Goldblatt, P. & John Manning. 2008. The Iris Family: Natural History & Classification. Timber Press, 290 pp. ISBN 9780881928976.
- For a comple te description of the diverse types of fruits and seeds in the Iridaceae, see Kirkbride, J.H., Jr., C.R. Gunn, and M.J. Dallwitz. 2006. Family Guide for Fruits and Seeds, vers. 1.0. Iridaceae. Accessed May 18, 2009.
External links
Taxonomy
The Family Iridaceae is further organized into finer groupings including:
- Subfamily (13): Asteroideae · Crocoideae · Faboideae · Iridoideae · Isophysidoideae · Ixioideae · Nivenioideae · Pyroideae · Ranunculoideae · Rhododendroideae · Ulmoideae · Violoideae · Yuccoideae
- Tribe (19): Anemoneae · Bombini · Coreopsideae · Eupatorieae · Gentianeae · Heliantheae · Irideae · Ixieae · Maleae · Mariceae · Narcisseae · Pillansieae · Rhododendreae · Sisyrinchieae · Tephrosieae · Tigridieae · Verbeneae · Violeae · Watsonieae
- Subtribe (2): Euphorbiinae · Sarcanthinae
- Genus (222): Acaste · Acidanthera · Aegilotriticum · Aglaea · Agretta · Ainea · Alatavia · Alophia · Anaclanthe · Anactorion · Anapalina · Anisanthus · Anomalesia · Anomalostylus · Anomatheca · Anomaza · Antholyza · Aristea · Asarina · Babiana · Ballosporum · Barnardiella · Beatonia · Beilia · Belamcanda · Bermudiana · Biris · Bobartia · Botherbe · Calydorea · Cardenanthus · Cardiostigma · Catila · Chamaeiris · Chamelum · Chasmanthe · Chasmatocallis · Chlamydostylus · Cipura · Cleanthe · Cobana · Colima · Coresantha · Costia · Crociris · Crocosmia · Crocus · Cryptobasis · Curtonus · Cyanixia · Cypella · Devia · Diaphane · Diasia · Dichone · Dierama · Dietes · Diplarrena · Diplarrhena · Dortania · Dugaldia · Duthiastrum · Echthronema · Eleutherine · Engysiphon · Ennealophus · Eremiris · Eriphilema · Eurynotia · Eustylis · Evansia · Eveltria · Exacum · Exohebea · Ferraria · Fosteria · Freesea · Freesia · Freuchenia · Galatea · Galathea · Galaxia · Gattenhofia · Geanthus · Geissorhiza · Geissorrhiza · Gelasine · Gemmingia · Genlisia · Genosiris · Geosiris · Gladiolus · Glumosia · Gordlinia · Grafia · Gynandriris · Hebea · Hecaste · Helixyra · Herbertia · Hermodactylus · Hesperantha · Hesperanthus · Hesperoxiphion · Hewardia · Hexaglottis · Homeria · Homoglossum · Hydastylus · Hydrotaenia · Hymenostigma · Hyptissa · Iridodictyum · Iridopsis · Iris · Isis · Isodon · Isophysis · Itysa · Ixia · Juno · Kelissa · Kentia · Kentrosiphon · Klattia · Lansbergia · Lapeirousia · Lapeyrousia · Larentia · Lemboglossum · Lemonia · Lethia · Libertia · Lomenia · Marica · Mastigostyla · Melasphaerula · Meristostigma · Micranthus · Microloma · Montbretia · Moraea · Morettia · Morphixia · Myosotidium · Naron · Nemastylis · Nemastylus · Neomarica · Neubeckia · Niduregelia · Nivenia · Oenostachys · Olsynium · Oncocyclus · Onira · Orthrosanthus · Ourisia · Ovieda · Pardanthus · Patersonia · Petamenes · Peyrousia · Phaianthes · Phaiophleps · Phalocallis · Pillansia · Pseudotrimezia · Ptilotrichum · Radinosiphon · Remaclea · Rheome · Rigidella · Roggeveldia · Romulea · Safran · Salpingostylis · Sasaella · Savannosiphon · Schizostylis · Sedadia · Sessilanthera · Sessilistigma · Siphonostylis · Sisyrinchium · Solenomelus · Sophronia · Souza · Sparaxis · Spatalanthus · Sphaerospora · Sphenostigma · Streptanthera · Susarium · Synnotia · Syringodea · Tamia · Tapeinia · Tchihatchewia · Tekelia · Thelysia · Thereianthus · Tigridia · Trichonema · Trifurcia · Trimeza · Trimezia · Tritonia · Tritoniopsis · Tritonixia · Vieusseuxia · Vincetoxicum · Watsonia · Winklera · Witsenia · Wuerthia · X Pardancanda · Xenoscapa · Xerodraba · Xiphion · Xyridion · Zygotritonia
- Species: ZipcodeZoo has pages for 33,864 species, subspecies, varieties, forms, and cultivars in the Family Iridaceae.
Genera
Acaste
Acaste was a name attributed to two characters in Greek mythology. [more]
Acidanthera
Gladiolus (from Latin, the diminutive of gladius, a sword) is a genus of perennial bulbous flowering plants in the iris family (Iridaceae). Sometimes called the sword lily, the most widely used English common name for these plants is simply gladiolus (plural gladioli, gladioluses or sometimes gladiolas). [more]
Aegilotriticum
Aglaea
Aglaea or Agla?a (Greek: "splendor, brilliant, shining one") is the name of several figures in Greek mythology. [more]
Agretta
Ainea
Ainea is a genus of perennial, herbaceous and bulbous plants belonging to the iris family (Iridaceae), endemic from Oaxaca in Mexico. They have white flowers with free stamens and a bifid style. The genus include only one species, Ainea conzattii, which grows in coniferous forest up to 2000 m.a.s.l. Currently, this genus is considered a synonym of Tigridia. [more]
Alatavia
Alophia
Alophia is a small genus of perennial, herbaceous and bulbous plants in the iris family (Iridaceae). The genus comprise five species that occur from center and southern North America to Brazil and Argentina. The genus is closely related to Herbertia, Cypella and Tigridia, differentiating from them by some characters of the stamen and the gynoecium. The genus name is derived from the Greek words a-, meaning "without", and lophos, meaning "crest". [more]
Anaclanthe
Anactorion
Anapalina
Anisanthus
Anomalesia
Gladiolus (from Latin, the diminutive of gladius, a sword) is a genus of perennial bulbous flowering plants in the iris family (Iridaceae). Sometimes called the sword lily, the most widely used English common name for these plants is simply gladiolus (plural gladioli, gladioluses or sometimes gladiolas). [more]
Anomalostylus
Anomatheca
Anomaza
Antholyza
Gladiolus (from Latin, the diminutive of gladius, a sword) is a genus of perennial bulbous flowering plants in the iris family (Iridaceae). Sometimes called the sword lily, the most widely used English common name for these plants is simply gladiolus (plural gladioli, gladioluses or sometimes gladiolas). [more]
Aristea
Aristea is a genus of perennial, herbaceous and rhizomatous species of flowering plants in the iris family (Iridaceae). The genus include 56 species which are distributed in tropical and southern Africa, as well as Madagascar. The genus name is derived from the Greek word arista, meaning "awn". [more]
Asarina
Asarina is a genus comprising 16 species of strongly sprawling or twining perennials, native to Mexico, southwestern USA, and southern Europe. Originally placed in the Scrophulariaceae (figwort family), they have more recently been moved to the Plantaginaceae (plantain family). Leaves are often triangular, toothed, downy and hairy with twining flower stalks. Flowers are attractive trumpet-shaped with broad green sepals and pale throat-spotted corolla in varying sizes, resemble snapdragons, and may be white, yellow, pink, purple, and shades in between. Some species are often placed in the genus Maurandya. [more]
Babiana
Babiana Ker Gawler () is a genus of flowering plants in the Family Iridaceae composed of about 80 species. Most of these species (about 49) are found in the southwestern Cape of Africa, with the remainder distributed in Namaqualand and Northern Cape Province. The native range of the genus is from southern Namibia to the Eastern Cape to southern Zimbabwe, with one species from Socotra off the coast of Somalia (although this may not be a species of Babiana). The genus name is derived from the Dutch word baviaan, referring to the African monkey that consumes the corms of plants in the genus. [more]
Ballosporum
Barnardiella
Beatonia
Beilia
Belamcanda
Iris domestica (Blackberry lily, Leopard flower, Leopard lily; syn. Belamcanda chinensis, Belamcanda punctata Moench, Gemmingia chinensis (L.) Kuntze, Iris chinensis Curtis, Ixia chinensis L., Morea chinensis, Pardanthus chinensis (L.) Ker Gawl.) is an ornamental plant in the Iridaceae family. In 2005, based on molecular DNA sequence evidence, Belamcanda chinensis, the sole species in the genus Belamcanda, was transferred to the genus Iris and renamed Iris domestica. [more]
Bermudiana
Biris
Bobartia
Botherbe
Calydorea
Herbs, perennial, from tunicate, ovoid bulbs; tunic brown, dry, brittle, papery. Stems simple or branched. Leaves few, basal larger; blade pleated, linear-lanceolate. Inflorescences rhipidiate, few-flowered; spathes green, unequal, outer shorter than inner, apex usually brown, acute, dry. Flowers short-lived, erect, actinomorphic; tepals spreading from base, distinct, blue to mauve, ± equal [outer whorl considerably larger than inner]; stamens distinct or filaments variously connate; anthers sometimes connate basally; style eccentric [central] when flower fully open, recurving, slender, branching between middle of filaments and anther apices [branching distal to anthers or 3-lobed apically]; branches [lobes] ascending, undivided, short, stigmatic apically. Capsules ovoid to oblong, cartilaginous, apex truncate. Seeds many, prismatic; seed coat brown. x = 7.[1] [more]
Cardenanthus
Cardiostigma
Catila
Chamaeiris
Chamelum
Chasmanthe
Chasmanthe is a genus of flowering plants in the iris family which originate from South Africa. In their native habitat the flowers are pollinated by sunbirds. The genus name is derived from the Greek words chasme, meaning "gaping", and anthos, meaning "flower". [more]
Chasmatocallis
Chlamydostylus
Cipura
Cleanthe
Cobana
Colima
A Genus in the Kingdom Animalia.[2] [more]
Coresantha
Costia
Crociris
Crocosmia
Crocosmia (; J.E. Planchon, 1851) is a small perennial genus in the iris family Iridaceae, native to the grasslands of Cape Floristic Region, South Africa. [more]
Crocus
Crocus (plural: crocuses, croci) is a genus in the iris family comprising about 80 species of perennials growing from corms. Many are cultivated for their flowers appearing in autumn, winter, or spring. Crocuses are native to woodland, scrub and meadows from sea level to alpine tundra in central and southern Europe, North Africa and the Middle East, on the islands of the Aegean, and across Central Asia to western China. [more]
Cryptobasis
Curtonus
Cyanixia
Cypella
Devia
Diaphane
Diasia
Dichone
Dierama
Dietes
Dietes is a genus of rhizomatous plants of the family Iridaceae. Common names include Fortnight lily, African iris, Morea or Moraea iris, Japanese iris and Butterfly iris, each of which may be used differently in different regions for one or more of the four species within the genus. [more]
Diplarrena
Diplarrena is a genus of 2 species in the family Iridaceae and native to south-eastern Australia. The name is from Greek diploos (double) and arren (male), as Diplarrena has only two functional stamens; all other Iridaceae have three. This name is often misspelled "Diplarrhena", an error that began with George Bentham's Flora Australiensis in 1873. The genus name is derived from the Greek words diploos, meaning "double", and arren, meaning "male". [more]
Diplarrhena
Diplarrena is a genus of 2 species in the family Iridaceae and native to south-eastern Australia. The name is from Greek diploos (double) and arren (male), as Diplarrena has only two functional stamens; all other Iridaceae have three. This name is often misspelled "Diplarrhena", an error that began with George Bentham's Flora Australiensis in 1873. The genus name is derived from the Greek words diploos, meaning "double", and arren, meaning "male". [more]
Dortania
Dugaldia
Duthiastrum
Echthronema
Eleutherine
Engysiphon
Ennealophus
Eremiris
Eriphilema
Eurynotia
Eustylis
Evansia
Eveltria
Orthrosanthus is a genus of flowering plants in the family Iridaceae. It can be seen in Australia, Central and South America. The genus name is derived from the Greek words orthros, meaning "morning", and anthos, meaning "flower". [more]
Exacum
Exacum is a genus of plant in family Gentianaceae. It contains the following species (but this list may be incomplete): [more]
Exohebea
Ferraria
Ferraria is a genus of 11 species of monocotyledonous flowering plants in the family Iridaceae, native to tropical and southern Africa. They are herbaceous corm-bearing plants growing to 30?45 cm tall. Some species have an unpleasant scent similar to rotting meat and are pollinated by flies, while others have a pleasant scent. The genus name is a tribute to 15th century artist Giovanni Ferrari. [more]
Fosteria
A Genus in the Kingdom Animalia. [more]
Freesea
Freesia
Freesia Ecklon ex Klatt is a genus of 14?16 species of flowering plants in the family Iridaceae, native to Africa. Of the 14 species, 12 are native to Cape Province, South Africa, the remaining two to tropical Africa, one species extending north of the equator to Sudan. [more]
Freuchenia
Galatea
Galathea
Galathea is one of the largest genera of squat lobsters, containing 70 currently recognised species (17 in the Atlantic Ocean, 22 in the Indian Ocean and 43 in the Pacific Ocean). Most species of Galathea live in shallow waters. [more]
Galaxia
Galaxia can refer to: [more]
Gattenhofia
Geanthus
Geissorhiza
Geissorrhiza
Gelasine
Gemmingia
Genlisia
Genosiris
Geosiris
Geosiris is a genus in the Iridaceae family of flowering plants. A monotypic genus, it contains the single species Geosiris aphylla, sometimes called the "earth-iris" sometimes not. Native to Madagascar and other islands in the Indian Ocean, G. aphylla is a small myco-heterotroph lacking chlorophyll. The genus name is derived from the Greek words geos, meaning "earth", and iris, referring to the Iris family of plants. [more]
Gladiolus
Gladiolus (from Latin, the diminutive of gladius, a sword) is a genus of perennial bulbous flowering plants in the iris family (Iridaceae). Sometimes called the sword lily, the most widely used English common name for these plants is simply gladiolus (plural gladioli, gladioluses or sometimes gladiolas). [more]
Glumosia
Gordlinia
Grafia
Gynandriris
Hebea
Hecaste
Helixyra
Herbertia
Herbs, perennial, from tunicate, ovoid bulbs; tunic brown, dry, brittle, papery. Stems simple or branched. Leaves few, basal larger; blade pleated, linear-lanceolate. Inflorescences rhipidiate, few-flowered; spathes green, unequal, inner spathe exceeding outer, apex brown, acute, usually dry. Flowers short-lived, erect, unscented, actinomorphic; tepals spreading, distinct, blue to mauve with white markings, unequal, outer whorl more than 2 times inner; filaments connate; anthers diverging, appressed to style branches; style slender, branching at apex of filament column; branches diverging from base, flattened, divided apically into 2 slender lobes, apically stigmatic. Capsules ovoid, apex truncate. Seeds many, prismatic; seed coat brown. x = 7.[3] [more]
Hermodactylus
Hesperantha
Hesperanthus
Hesperoxiphion
Hewardia
Hexaglottis
Homeria
Homeria, commonly known as Cape tulips, is a genus of the botanical family Iridaceae, which has 32 species of corm-bearing perennial herbs. In addition, these plants are used as ornamental plants. In the early 19th century, this genus was recognized as a single species of tulip, Tulipa breyniana[1]. Currently, this genus is considered a synonym of Moraea. [more]
Homoglossum
Hydastylus
Hydrotaenia
Hymenostigma
Hyptissa
Iridodictyum
Iridopsis
Iris
Iris has three main meanings, related by their derivation from the Greek word for rainbow: [more]
Isis
Isis (Greek: , original Egyptian pronunciation more likely Aset) is a goddess in Ancient Egyptian religious beliefs, whose worship spread throughout the Greco-Roman world. She was worshipped as the ideal mother and wife as well as the patron of nature and magic. She was the friend of slaves, sinners, artisans, and the downtrodden, and she listened to the prayers of the wealthy, maidens, aristocrats, and rulers. Isis is often depicted as the mother of Horus, the hawk-headed god of war and protection (although in some traditions Horus's mother was Hathor), and she is depicted suckling him in an attitude similar to that of the Virgin Mary with the Child Jesus. Isis is also known as protector of the dead and goddess of children. [more]
Isodon
A Genus in the Kingdom Animalia.[4] [more]
Isophysis
Itysa
Ixia
The genus Ixia consists of a number of cormous plants native to South Africa from the Iridaceae family and Ixioideae subfamily. Some of them are known as the corn lily. Some distinctive traits include: sword-like leaves, and long wiry stems with star-shaped flowers. It usually prefers well-drained soil. The popular corn lily has specific, not very intense fragrance as vegetables. That is why it's often visited by many insects such as bees. The Ixia are also used sometimes as ornamental plants. [more]
Juno
Kelissa
Kentia
Kentrosiphon
Klattia
Lansbergia
Lapeirousia
Lapeyrousia
Larentia
A Genus in the Kingdom Animalia.[5] [more]
Lemboglossum
Lemonia
Lethia
Libertia
Libertia is a genus of monocotyledenous plants in the family Iridaceae containing 15 species in the southern hemisphere. A number of species are endemic to New Zealand. [more]
Lomenia
Marica
Marica is a derived from Miriam. It may also refer to: [more]
Mastigostyla
Melasphaerula
Meristostigma
Micranthus
Microloma
Montbretia
Crocosmia (; J.E. Planchon, 1851) is a small perennial genus in the iris family Iridaceae, native to the grasslands of Cape Floristic Region, South Africa. [more]
Moraea
Moraea is a genus of plant in family Iridaceae. It contains the following species (but this list may be incomplete): [more]
Morettia
Morphixia
Myosotidium
Myosotidium is a genus of plants belonging to the family Boraginaceae. This genus is represented by the single species Myosotidium hortensia, the Chatham Islands forget-me-not, which is endemic to the Chatham Islands, New Zealand. [more]
Naron
Nemastylis
Herbs, perennial, from tunicate, ovoid bulbs; tunic brown, dry, brittle, papery. Stems simple or branched. Leaves few; blade pleated, lanceolate to ± linear. Inflorescences rhipidiate, few-flowered; spathes green, unequal, outer shorter than inner, apex usually brown, acute, dry. Flowers short-lived, actinomorphic; tepals spreading from base, distinct, blue [to mauve, or white], ± equal [outer whorl slightly larger than inner]; filaments distinct or partly to completely connate; anthers initially erect, collapsing in spiral fashion after dehiscing; style branching ± at or shortly below base of anthers; branches divided ± to base, 2-armed; arms filiform, extending horizontally on either side of subtending anther, stigmatic apically. Capsules ovoid to oblong, cartilaginous, apex truncate to turbinate. Seeds many, ± prismatic, or compressed, winged; seed coat brown. x = 7.[6] [more]
Nemastylus
Neomarica
Neomarica (Walking Iris, Apostle's Iris or Apostle Plant) is a genus of 16 species of plants in family Iridaceae, native to tropical regions of western Africa, Central and South America, with the highest diversity (12 species) in Brazil. The genus name is derived from the Greek words neo, meaning "new", and Marica, the Roman nymph. [more]
Neubeckia
Niduregelia
Nivenia
Oenostachys
Olsynium
Olsynium (huilmo in Chile; formerly part of Sisyrinchium) is a genus of 12 species of summer-dormant rhizomatous perennials in the iris family, native to sunny hillsides in South America and western North America. [more]
Oncocyclus
Onira
Orthrosanthus
Orthrosanthus is a genus of flowering plants in the family Iridaceae. It can be seen in Australia, Central and South America. The genus name is derived from the Greek words orthros, meaning "morning", and anthos, meaning "flower". [more]
Ourisia
Ovieda
Pardanthus
Patersonia
Patersonia (or Purple flag) is a genus of the Iridaceae with about 20 species in Australia and several in the Malesian region. The genus name is a tribute to the first Lieutenant Governor of New South Wales in Australia, William Paterson. [more]
Petamenes
Peyrousia
Phaianthes
Phaiophleps
Phalocallis
Pillansia
Pillansia is a genus of flowering plants in the family Iridaceae. The genus name is a tribute to the South African botanist , who brought the species to the attention of Harriet Margaret Louisa Bolus. [more]
Pseudotrimezia
Ptilotrichum
Radinosiphon
Remaclea
Rheome
Rigidella
Roggeveldia
Romulea
Herbs, perennial, from corms. Stems: true stem aerial [subterranean and only peduncles of individual flowers aerial], simple or branched. Leaves 1-several; blade linear, oval to terete in cross section, 4-grooved (2 on each surface, on either side of thickened midrib) [2-grooved]. Inflorescences solitary flowers, interpreted as sessile, thus without pedicels; bracts inserted at ovary base, green or flushed with purple, unequal, outer exceeding inner, firm, inner bract margins membranous to scarious (rarely entirely scarious). Flowers odorless [fragrant], actinomorphic; tepals forming wide cup, connate into tube, subequal, outer whorl often slightly larger than inner; perianth tube funnel-shaped [or cylindric]; stamens symmetrical; filaments distinct [connate]; anthers usually erect, contiguous; style branching opposite or beyond anthers into 3 slender branches divided for ± 1/2 length. Capsules ovoid-oblong, walls firm, cartilaginous. Seeds many, globose; seed coat light to dark brown, often smooth, shiny. x = 13 or 14.[7] [more]
Safran
Safran is a French conglomerate involved in defense, aerospace propulsion and equipment, and security. It is the result of a merger between the propulsion and aerospace equipment group SNECMA and the defense conglomerate SAGEM. Its headquarters are located in Paris. [more]
Salpingostylis
Sasaella
Sasaella is a genus of bamboo. [more]
Savannosiphon
Schizostylis
Sedadia
Sessilanthera
Sessilistigma
Siphonostylis
Sisyrinchium
Sisyrinchium (Blue-eyed Grasses) is a genus of 70-150 species of annual to perennial plants of the iris family, native to the New World. [more]
Solenomelus
Sophronia
Sophronitis, abbreviated Soph in horticultural trade, is a genus of small, epiphytic or lithophytic orchids, growing in the damp montane forest of eastern Brazil, Paraguay and NE Argentina. Currently, 65 species are recognized. [more]
Souza
Sousa may mean a number of different things. [more]
Sparaxis
Sparaxis (Harlequin Flower) is a genus in the family Iridaceae with about 13 species endemic to Cape Province, South Africa. [more]
Spatalanthus
Sphaerospora
Sphenostigma
Streptanthera
Susarium
Synnotia
Syringodea
Tamia
Tamia Marilyn Hill (n?e Washington; born May 8, 1975), and known professionally as Tamia, is a Canadian R&B and soul singer, songwriter, record producer, composer, model, entrepreneur and philanthropist, as well as an occasional actress. [more]
Tapeinia
Tchihatchewia
Tekelia
Thelysia
Thereianthus
Tigridia
Tigridia (tiger-flowers or shell flowers) is a genus of bulbous or cormous plants, belonging to the family Iridaceae. They have large showy flowers and one species, Tigridia pavonia, is often cultivated for this. The approximately thirty five species in this family grow in the Western Hemisphere, from Mexico to Chile. The tigridia flower is short lived, each often blooming for only one day, but often several flowers will bloom from the same stalk. Usually they are dormant during the winter dry-season. Its roots are edible and was eaten by the Aztecs of Mexico who called it cacomitl and its flower ocÄ“lÅxÅchitl "Jaguar flower". [more]
Trichonema
Trifurcia
Trimeza
Trimezia
Tritonia
Tritonia refers to a genus. The name derives from the god Triton. It may refer to: [more]
Tritoniopsis
Tritonixia
Vieusseuxia
Vincetoxicum
Vincetoxicum is a genus of plant in family Apocynaceae. It contains the following species (but this list may be incomplete): [more]
Watsonia
Watsonia can refer to: [more]
Winklera
Witsenia
Witsenia is a genus of flowering plants in the family Iridaceae. The genus name is a tribute to the botany patron Nicolaas Witsen. It is impossible that Witsen helped finance Carl Peter Thunberg's explorations in South Africa and Japan. [more]
Wuerthia
X Pardancanda
Xenoscapa
Xenoscapa is a genus of herbaceous, perennial and bulbous plants in the Iris family (Iridaceae). It consists of only two species distributed in Africa, and closely related to the genera Freesia and . The genus name is derived from the Greek words xenos, meaning "strange", and scapa, meaning "flowering stem". [more]
Xerodraba
Xiphion
Xyridion
Zygotritonia
At least 7 species and subspecies belong to the Genus Zygotritonia.
More info about the Genus Zygotritonia may be found here.
References
- ^ a b c d e f g h i Goldblatt, Peter; John Manning (2008). The Iris Family: Natural History & Classification. Portland: Timber Press. ISBN 978-0-88192-897-6.
- ^ a b c d e Goldblatt, Peter (2002). Iridaceae "Iridaceae". Flora of North America. Volume. 26: Magnoliophyta: Liliidae: Liliales and Orchidales. New York: Oxford University Press. p. 348. ISBN 0-19-515208-5. http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10452 Iridaceae. Retrieved 2009-05-12.
- ^ a b c d e f g h Manning, J. (2004). "Iridaceae". South African National Biodiversity Institute. http://www.plantzafrica.com/planthij/iridaceae.htm. Retrieved 2009-05-12.
- ^ Zhao, Yu-tang; Henry J. Noltie & Brian F. Mathew (2000). "Iridaceae". In Wu Zheng-yi & Peter Hamilton Raven (eds.). Flora of China. Volume. 24: Flagellariaceae through Marantaceae. St. Louis: Missouri Botanical Garden. p. ;297. ISBN 0-915279-83-5. http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=10452. Retrieved 2009-05-12.
- ^ Dimitri, M. 1987. Enciclopedia Argentina de Agricultura y Jardiner?a. Tomo I. Descripci?n de plantas cultivadas. Editorial ACME S.A.C.I., Buenos Aires.
- ^ Watson, Leslie (2008). "Iridaceae". Florabase, The Western Australia Flora. Western Australia Department of Environment and Conservation. http://florabase.calm.wa.gov.au/browse/profile/22784. Retrieved 2009-05-12.
- ^ J. Mcneil et al. (2006). "Nomina Familiarum Bryophytorum et Spermatophytorum Conservanda". International Code of Botanical Nomenclature (VIENNA CODE). http://ibot.sav.sk/icbn/main.htm. Retrieved 2009-05-24.
- ^ Arthur Cronquist. 1988. The Evolution and Classification of Flowering Plants
- ^ A. Takhtajan (1980). "Outline of the classification of flowering plants (Magnoliophya)". Botanical Review 46 (3): 225?359. doi:10.1007/BF02861558.
- ^ Robert F. Thorne. 1992. Classification and Geography of Flowering Plants. Botanical Review 58: 225-348
- ^ APG, 1998. An ordinal Classification for the Families of Flowering Plants. Annals of the Missouri Botanical Garden 85: 531-553
- ^ Angiosperm Phylogeny Group (2003). Full text An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 41: 399-436.
- ^ a b c Goldblatt, P., Aaron Rodriguez, M. P. Powell, T. Jonathan Davies, John C. Manning, M. van der Bank, Vincent Savolainen. 2008. Iridaceae ?Out of Australasia?? Phylogeny, Biogeography, and Divergence Time Based on Plastid DNA Sequences.Systematic Botany 33:3, 495-508
- ^ Peter Goldblatt and Masahiro Takei. 1997. Chromosome Cytology of Iridaceae-Patterns of Variation, Determination of Ancestral Base Numbers, and Modes of Karyotype Change. Annals of the Missouri Botanical Garden, Vol. 84, No. 2, pp. 285-304
- ^ Peter Goldblatt. 1980. Uneven Diploid Chromosome Numbers and Complex Heterozygosity in Homeria (Iridaceae). Systematic Botany, Vol. 5, No. 4, pp. 337-340
- ^ Chase M. W.. Duvall, M. R., Hills, H. G., Conran, J. G., Cox, A. V., Eguiarte, L. E., Hartwell, J., Fay, M. F., Caddick, L. R., Cameron, K. M., y Hoot, S. 1995. Molecular systematics of Lilianae. In: Rudall, P. J., Cribb, P. J., Cutler, D. F. (eds.) Monocotyledons: Systematics and evolution. Royal Botanic Gardens, Kew, pp. 109-137
- ^ Chase, M. W.,Soltis, D. E., Soltis, P. S., Rudall, P. J., Fay, M. F., Hahn, W. H., Sullivan, S., Joseph, J., Molvray, M., Kores, P. J., Givnish, T. J., Sytsma, K. J., y Pires, J. C. 2000. Higher-level systematics of the monocotyledons: An assessment of current knowledge and a new classification. In.: Wilson, K. L. & Morrison, D. A. (eds.) Monocots: Systematics and Evolution.CSIRO Publ., Collingwood, Australia, pp,:3-16
- ^ Stevenson,D. W. & Loconte, H. 1995. Cladistic analysis of monocot families. In.: Rudall, P. J., Cribb, P. J., Cutler, D. F. (Eds.) Monocotyledons: Systematics and evolution, Royal Botanic Gardens, Kew, pp.: 543-578
- ^ Souza-Chies,T. T., Gabriel Bittar, Sophie Nadot, Leigh Carter, Evelyne Besin and Bernard Lejeune. 1997. Phylogenetic analysis ofIridaceae with parsimony and distance methods using the plastid generps4. Plant Systematics and Evolution 204(1): 109-123
- ^ Tatiana T. Souza-Chies, Gabriel Bittar, Sophie Nadot, Leigh Carter, Evelyne Besin and Bernard Lejeune. 1997. Phylogenetic analysis of Iridaceae with parsimony and distance methods using the plastid generps4. Plant Systematics and Evolution 204(1): 109-123
- ^ a b Goldblatt, P. Phylogeny and Classification of Iridaceae. Annals of the Missouri Botanical Garden, Vol. 77, No. 4 (1990), pp. 607?627.
- ^ a b Goldblatt, P. 1991. An overview of the systematics, phylogeny and biology of the southern African Iridaceae. Contr. Bolus Herb. 13: 1?74.
- ^ Stevens, P. F. 2001 onwards. Angiosperm Phylogeny Website. Downloaded 2009-5-19.
- ^ De Tullio, L., Germ?n Roitman, Gabriel Bernardello. 2008. Tamia (Iridaceae), a Synonym of Calydorea: Cytological and Morphological Evidence. Systematic Botany 2008 33:3, 509?513
- ^ Reeves, G., Chase, M.W., Goldblatt, P., Rudall, P., Fay, M.F., Cox, A.V., LeJeune, B., y Souza-Chies, T., (2001). Molecular systematics of Iridaceae: Evidence from four plast id DNA regions.. Amer. J. Bot. 88:2074-2087.
- ^ Royal Horticultural Society, Kew. A detailed checklist for family Iridaceae.
- ^ R. K. Brummitt. 1980. Proposal to conserve Alophia over Eustylis. Report of the Committee for Spermatophyta, 22. Taxon, Vol. 29, No. 4 (August, 1980), pp. 489?493
- ^ Goldblatt, P. y J.E. Henrich. Calydorea Herbert (Iridaceae-Tigridieae): Notes on this New World Genus and Reduction to Synonymy of Salpingostylis, Cardiostigma, Itysa, and Catila. Annals of the Missouri Botanical Garden, Vol. 78, No. 2 (1991), pp. 504?511
- ^ Goldblatt, P. & De Vos M. P. The reductio n of Oenostachys, Homoglossum and Anomalesia, putative sunbird pollinated genera, in Gladiolus L. (Iridaceae-Ixioideae). Bulletin du Mus?um national d'histoire naturelle. Section B, Adansonia 11 (4): 417?428, 1989.
- ^ Goldblatt, P. & David J. Mabberley. Belamcanda included in Iris, and the New Combination I. domestica (Iridaceae: Irideae). Novon 15 (1), pp. 128?132, (2005).
- ^ Goldblatt, P. Reduction of Bernardiella, Galaxia, Gynandriris, Hexaglottis, Homeria, and Roggeveldia in Moraea (Iridaceae: Irideae). Novon, Vol. 8, No. 4 (1998), pp. 371?377
- ^ a b Goldblatt, P. & J.C. Manning. 2006. Radiation of Pollination Systems in the Iridaceae of sub-Saharan Africa. Ann Bot 97: 317-344.
- ^ IUCN 2008. [<www.iucnredlist.org> 2008 IUCN Red List of Threatened Species.] Downloaded on 19 May 2009.
- ^ Vestal, Paul A. 1952 The Ethnobotany of the Ramah Navaho. Papers of the Peabody Museum of American Archaeology and Ethnology 40(4):1?94 (p. 21).
- ^ Nickerson, Gifford S. 1966 Some Data on Plains and Great Basin Indian Uses of Certain Native Plants. Tebiwa 9(1):45?51 (p. 47)
- ^ Train, Percy, James R. Henrichs and W. A ndrew Archer 1941 Medicinal Uses of Plants by Indian Tribes of Nevada. Washington DC. U.S. Department of Agriculture (p. 89, 90)
- ^ Raymond, Marcel. 1945 Notes Ethnobotaniques Sur Les Tete-De-Boule De Manouan. Contributions de l'Institut botanique l'Universite de Montreal 55:113?134 (p. 129))
- ^ Speck, Frank G., R.B. Hassrick and E.S. Carpenter 1942 Rappahannock Herbals, Folk-Lore and Science of Cures. Proceedings of the Delaware County Institute of Science 10:7?55. (p. 28)
- ^ Sisyrinchium acre - Hawaiian Ethnobotany Online Database
- ^ Chopra. R. N., Nayar. S. L. and Chopra. I. C. Glossary of Ind ian Medicinal Plants (Including the Supplement). Council of Scientific and Industrial Research, New Delhi. 1986
- ^ Bown. D. Encyclopaedia of Herbs and their Uses. Dorling Kindersley, London. 1995 ISBN 0-7513-0203-1
- ^ Chevallier. A. The Encyclopedia of Medicinal Plants. Dorling Kindersley. London 1996 ISBN 978-0-7513-0314-8
Bibliography
- De Vos, M. P. 1972. The genus Romulea in South Africa. J. S. African Bot., suppl. 9.
- De Vos, M. P. 1983. Romulea. In: Department of Agricultural Technical Services [South Africa]. 1963+. Flora of Southern Africa.... 19+ vols. (some partial). Pretoria. Vol. 7, part 2, fasc. 2, pp. 1073.
- Foster, R. C. 1945. Studies in the Iridaceae, III. Contr. Gray Herb. 155: 3-54.
- Goldblatt, P. and J. E. Henrich. 1991. Calydorea Herbert (IridoideaeTigridieae): Notes on this New World genus and reduction to synonymy of Salpingostylis, Cardiostigma, Itysa, and Catila. Ann. Missouri Bot. Gard. 78: 504511.
- Manning, J. C. and P. Goldblatt. 2001. A synoptic review of Romulea (Iridaceae: Crocoideae) in sub-Saharan Africa, the Arabian Peninsula and Socotra including new species, biological notes, and a new infrageneric classification. Adansonia, sér. 3, 23: 59108.
- Small, J. K. 1931b. Bartrams Ixia coelestina rediscovered. J. New York Bot. Gard. 32: 155161.
Footnotes
- Peter Goldblatt "Calydorea". in Flora of North America Vol. 26 Page 17, 349, 400, 401. Oxford University Press. Online at EFloras.org.
- http://bugguide.net/index.php?q=search&keys=Colima&search=Search
- Peter Goldblatt "Herbertia". in Flora of North America Vol. 26 Page 17, 349, 396. Oxford University Press. Online at EFloras.org.
- http://www.ubio.org/browser/details.php?namebankID=113831
- http://bugguide.net/index.php?q=search&keys=Larentia&search=Search
- Peter Goldblatt "Nemastylis". in Flora of North America Vol. 26 Page 17, 349, 398. Oxford University Press. Online at EFloras.org.
- Peter Goldblatt "Romulea". in Flora of North America Vol. 26 Page 348, 349, 407. Oxford University Press. Online at EFloras.org.
- ^ a b c d e f g h i Goldblatt, Peter; John Manning (2008). The Iris Family: Natural History & Classification. Portland: Timber Press. ISBN 978-0-88192-897-6.
- ^ a b c d e Goldblatt, Peter (2002). Iridaceae "Iridaceae". Flora of North America. Volume. 26: Magnoliophyta: Liliidae: Liliales and Orchidales. New York: Oxford University Press. p. 348. ISBN 0-19-515208-5. http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10452 Iridaceae. Retrieved 2009-05-12.
- ^ a b c d e f g h Manning, J. (2004). "Iridaceae". South African National Biodiversity Institute. http://www.plantzafrica.com/planthij/iridaceae.htm. Retrieved 2009-05-12.
- ^ Zhao, Yu-tang; Henry J. Noltie & Brian F. Mathew (2000). "Iridaceae". In Wu Zheng-yi & Peter Hamilton Raven (eds.). Flora of China. Volume. 24: Flagellariaceae through Marantaceae. St. Louis: Missouri Botanical Garden. p. 297. ISBN 0-915279-83-5. http://www.efloras.org/florataxon.aspx?flora_id=2&ta xon_id=10452. Retrieved 2009-05-12.
- ^ Dimitri, M. 1987. Enciclopedia Argentina de Agricultura y Jardiner?a. Tomo I. Descripci?n de plantas cultivadas. Editorial ACME S.A.C.I., Buenos Aires.
- ^ Watson, Leslie (2008). "Iridaceae". Florabase, The Western Australia Flora. Western Australia Department of Environment and Conservation. http://florabase.calm.wa.gov.au/browse/profile/22784. Retrieved 2009-05-12.
- ^ J. Mcneil et al. (2006). "Nomina Familiarum Bryophytorum et Spermatophytorum Conservanda". International Code of Botanical Nomenclature (VIENNA CODE). http://ibot.sav.sk/icbn/main.htm. Retrieved 2009-05-24.
- ^ Arthur Cronquist. 1988. The Evolution and Classification of Flowering Plants
- ^ A. Takhtajan (1980). "Outline of the classification of flowering plants (Magnoliophya)". Botanical Review 46 (3): 225?359. doi:10.1007/BF02861558.
- ^ Robert F. Thorne. 1992. Classification and Geography of Flowering Plants. Botanical Review 58: 225-348
- ^ APG, 1998. An ordinal Classification for the Families of Flowering Plants. Annals of the Missouri Botanical Garden 85: 531-553
- ^ Angiosperm Phylogeny Group (2003). Full text An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 41: 399-436.
- ^ a b c Goldblatt, P., Aaron Rodriguez, M. P. Powell, T. Jonathan Davies, John C. Manning, M. van der Bank, Vincent Savolainen. 2008. Iridaceae ?Out of Australasia?? Phylogeny, Biogeography, and Divergence Time Based on Plastid DNA Sequences.Systematic Botany 33:3, 495-508
- ^ Peter Goldblatt and Masahiro Takei. 1997. Chromosome Cytology of Iridaceae-Patterns of Variation, Determination of Ancestral Base Num bers, and Modes of Karyotype Change. Annals of the Missouri Botanical Garden, Vol. 84, No. 2, pp. 285-304
- ^ Peter Goldblatt. 1980. Uneven Diploid Chromosome Numbers and Complex Heterozygosity in Homeria (Iridaceae). Systematic Botany, Vol. 5, No. 4, pp. 337-340
- ^ Chase M. W.. Duvall, M. R., Hills, H. G., Conran, J. G., Cox, A. V., Eguiarte, L. E., Hartwell, J., Fay, M. F., Caddick, L. R., Cameron, K. M., y Hoot, S. 1995. Molecular systematics of Lilianae. In: Rudall, P. J., Cribb, P. J., Cutler, D. F. (eds.) Monocotyledons: Systematics and evolution. Royal Botanic Gardens, Kew, pp. 109-137
- ^ Chase, M. W.,Soltis, D. E., Soltis, P. S., Rudall, P. J., Fay, M. F., Hahn, W. H., Sulliva n, S., Joseph, J., Molvray, M., Kores, P. J., Givnish, T. J., Sytsma, K. J., y Pires, J. C. 2000. Higher-level systematics of the monocotyledons: An assessment of current knowledge and a new classification. In.: Wilson, K. L. & Morrison, D. A. (eds.) Monocots: Systematics and Evolution.CSIRO Publ., Collingwood, Australia, pp,:3-16
- ^ Stevenson,D. W. & Loconte, H. 1995. Cladistic analysis of monocot families. In.: Rudall, P. J., Cribb, P. J., Cutler, D. F. (Eds.) Monocotyledons: Systematics and evolution, Royal Botanic Gardens, Kew, pp.: 543-578
- ^ Souza-Chies,T. T., Gabriel Bittar, Sophie Nadot, Leigh Carter, Evelyne Besin and Bernard Lejeune. 1997. Phylogenetic analysis ofIridaceae with parsimony and distance methods using the plastid generps4. Plant Systematics and Evolution 204(1): 109-123
- ^ Tatiana T. Souza-Chies, Gabriel Bittar, Sophie Nadot, Leigh Carter, Evelyne Besin and Bernard Lejeune. 1997. Phylogenetic analysis of Iridaceae with parsimony and distance methods using the plastid generps4. Plant Systematics and Evolution 204(1): 109-123
- ^ a b Goldblatt, P. Phylogeny and Classification of Iridaceae. Annals of the Missouri Botanical Garden, Vol. 77, No. 4 (1990), pp. 607?627.
- ^ a b Goldblatt, P. 1991. An overview of the systematics, phylogeny and biology of the southern African Iridaceae. Contr. Bolus Herb. 13: 1?74.
- ^ Stevens, P. F. 2001 onwards. Angiosperm Phylogeny Website. Downloaded 2009-5-19.
- ^ De Tullio, L., Germ?n Roitman, Gabriel Bernardello. 2008. Tamia (Iridaceae), a Synonym of Calydorea: Cytological and Morphological Evidence. Systematic Botany 2008 33:3, 509?513
- ^ Reeves, G., Chase, M.W., Goldblatt, P., Rudall, P., Fay, M.F., Cox, A.V., LeJeune, B., y Souza-Chies, T., (2001). Molecular systematics of Iridaceae: Evidence from four plastid DNA regions.. Amer. J. Bot. 88:2074-2087.
- ^ Royal Horticultural Society, Kew. A detailed checklist for family Iridaceae.
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