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Location: Botanical Garden of Brasilia - Brazil.
Chresta is a genus of South American flowering plants in the family Asteraceae. Close to 14 species are recognized.
Kingdom: Plantae
(unranked): Angiosperms
(unranked): Eudicots
(unranked): Asterids
Order: Asterales
Family: Asteraceae
Subfamily: Cichorioideae
Tribe: Vernonieae
Genus: Chresta Vell. ex DC.
Species: C. sphaerocephala DC.
Binomial name: Chresta sphaerocephala
Centaurea is a genus of over 700 species of herbaceous thistle-like flowering plants in the family Asteraceae. Members of the genus are found only north of the equator, mostly in the Eastern Hemisphere; the Middle East and surrounding regions are particularly species-rich. In the western United States, yellow starthistles are an invasive species. Around the year 1850, seeds from the plant had arrived to the state of California. It is believed that those seeds came from South America.
Common names
Common names for this genus are centaury, centory, starthistles, knapweeds, centaureas and the more ambiguous "bluets"; a vernacular name used for these plants in parts of England is "loggerheads" (common knapweed). The Plectocephalus group – possibly a distinct genus – is known as basketflowers. "Cornflower" is used for a few species, but that term more often specifically means either C. cyanus (the annual cornflower) or Centaurea montana (the perennial cornflower). The common name "centaury" is sometimes used, although this also refers to the unrelated plant genus Centaurium.
The name is said to be in reference to Chiron, the centaur of Greek mythology who discovered medicinal uses of a plant eventually called "centaury".
Description
Knapweeds are robust weedy plants. Their leaves, spiny in some species, are usually deeply divided into elongated lobes at least in the plants' lower part, becoming entire towards the top. The "flowers" (actually pseudanthium inflorescences) are diverse in colour, ranging from intense blues, reds and yellows to any mixture of these and lighter shades towards white. Often, the disk flowers are much darker or lighter than the ray flowers, which also differ in morphology and are sterile. Each pseudanthium sits atop a cup- or basket-like cluster of scaly bracts, hence the name "basketflowers". Many species, in particular those inhabiting more arid regions, have a long and strong taproot.
Certain knapweeds have a tendency to dominate large stretches of landscape together with a few other plants, typically one or two grasses and as many other large herbaceous plants. The common knapweed (C. nigra) for example is plentiful in the mesotrophic grasslands of England and nearby regions. It is most prominently found in pastures or meadows dominated by cock's-foot (Dactylis glomerata) as well as either of crested dog's-tail (Cynosurus cristatus) and false oat-grass (Arrhenatherum elatius). It is also often found in mesotrophic grassland on rendzinas and similar calcareous soils in association with glaucous sedge (Carex flacca), sheep's fescue (Festuca ovina), and either tor-grass (Brachypodium pinnatum) and rough hawkbit (Leontodon hispidus), or upright brome (Bromus erectus). In these grasslands, greater knapweed (C. scabiosa) is found much more rarely by comparison, often in association with red fescue (Festuca rubra) in addition to cock's-foot and false oat-grass.
Due to their habit of dominating ecosystems under good conditions, many Centaurea species can become invasive weeds in regions where they are not native. In parts of North America, diffuse knapweed (C. diffusa), spotted knapweed (C. maculosa) and yellow starthistle (C. solstitialis) cause severe problems in agriculture due to their uncontrolled spread. The seeds are typically transported by human traffic, in particular the tires of all-terrain vehicles. The two knapweeds are harmful mainly because they are strongly allelopathic, producing powerful toxins in their roots that stunt the growth of plants around them not adapted to this. Yellow starthistle, meanwhile, is inedible to most livestock due to its spines and apparently outright poisonous to horses and other equines. However, efficient methods of biological control by insect pests of these weeds have been developed; the knapweeds can also exploited to their detriment by targeted grazing. Controlled burning may also be used, though the timing is important to avoid the plants having seeded already, and neither allowing sufficient time for them to regrow from the rootstock.
Yet other species of Centaurea – mostly ones that occur between Italy and the Caucasus – are endemics of a single island or valley, and some of these are endangered. The Akamas Centaurea (Centaurea akamantis) of Cyprus is almost extinct, while the western Caucasus endemics C. leptophylla and C. straminicephala are at least very rare and C. hedgei and C. pecho from the same region are certainly not abundant either. The last four species would be adversely affected by the proposed Yusufeli Dam, which might actually destroy enough habitat to push the two rarer ones over the brink of extinction.
Centaurea are copious nectar producers, especially on high-lime soils. The high nectar yield of the genus makes it very attractive to insects such as butterflies – including the endangered Karner blue (Plebejus melissa samuelis) which visits introduced spotted knapweed – and day-flying moths – typically Zygaenidae, such as Zygaena loti or the six-spot burnet (Z. filipendulae). The larvae of some other Lepidoptera species use Centaurea species as food plants; see List of Lepidoptera that feed on Centaurea. Several of these are used in biological control of invasive knapweeds and starthistles.
Larvae of several true weevils (Curculionidae) of the subfamily Lixinae also feed on Centaurea. Some genera – such as Larinus whose larval food is flowerheads – have many species especially adapted to particular knapweeds or starthistle and are used in biological control too. These include the yellow starthistle flower weevil (L. curtus) for yellow starthistle, lesser knapweed flower weevil (L. minutus) for diffuse knapweed and blunt knapweed flower weevil (L. obtusus) for spotted knapweed. Broad-nosed seedhead weevil (Bangasternus fausti) larvae eat diffuse, spotted and squarrose knapweed (C. virgata ssp. squarrosa), while those of the yellow starthistle bud weevil (B. orientalis) do not seem to live on anything other than yellow starthistle and occasionally purple starthistle (C. calcitrapa). But perhaps most efficient in destroying developing yellow starthistle seedheads is the larva of the yellow starthistle hairy weevil (Eustenopus villosus). Knapweed root weevil (Cyphocleonus achates) larvae bore into the roots of spotted and to a lesser extentely diffuse knapweed, sometimes killing off the entire plant.
Also used in biological control are Tephritidae (peacock flies) whose larvae feed on Centaurea. Knapweed peacock fly (Chaetorellia acrolophi) larvae eat spotted knapweed and some other species. The yellow starthistle peacock fly (C. australis) has an initial generation each year which often uses cornflower (C. cyanus) as larval food; later generations switch to yellow starthistle. The flies are generally considered less efficient in destroying the growing seedheads than the weevils, but may be superior under certain conditions; employing flies and weevils in combination is expensive and does not noticeably increase their effect.
Use by humans
Although the genus may be considered by a quite significant number of relatively informed individuals to have an overall negative impact on human interests, particularly agricultural interests, the situation is not straightforward enough to simply declare the genus, or, at least, its most aggressively-spreading species, altogether negative. For instance, due to their moderate to high nectar production, which can occur over a comparatively long duration, many species of Centaurea are popular food sources for insects that may otherwise attack certain crops.[citation needed] It may be advisable for some types of farms to allow certain species in this genus, such as cornflower (C. cyanus) in a European setting, to grow adjacent to fields. Although they support and attract many types of beneficial life (not just beetles), these areas are known as beetle banks. When they are present, some pests may be drawn away from crops to them and predatory insects and arachnids that feed upon pest insects will be better-supported by these more naturalized areas. They additionally have the beneficial aspect of supporting pollinators, unlike many field crops such as maize. Moreover, being untreated with pesticides and providing more diversity, plants growing in more wild areas adjacent to farms produce more insects that attract and support birds which can also feed on pests that would harm crops. Insect production is especially high for beetle banks that have enough plants that serve in the role of host plant for immature insects, rather than just in the roles of adult food and/or shelter provision.
Some plants which are considered invasive or problematic in certain areas can have beneficial qualities that outweigh their negative qualities from a human and/or human agricultural point of view, although this sometimes requires some human management – particularly if adequate biological control has not been established for the more aggressive species. An example is wild parsnip, Pastinaca sativa, which produces florets that feed predatory (and other beneficial) insects as well as large tubular stems that provide winter shelter for native bees, wasps, and other organisms that can be beneficial for agriculture. The plant is considered invasive in some areas of the United States and is also often considered undesirable due to its ability to cause contact skin irritation. However, it also serves as a host plant for the black swallowtail butterfly, helps to bring nutrients up from soils with its deep taproot, and possesses evergreen foliage even in climate zones such as US zone 6. This foliage increases soil warmth and moisture which can be beneficial for certain types of life. Perhaps the most dramatic example of a generally disliked plant's beneficial qualities being usually overlooked is the often-despised ragwort, Jacobaea vulgaris, which topped the list by a large amount for nectar production in a UK study, with a production per floral unit of (2921 ± 448μg). This very high nectar production, coupled with its early blooming period, makes the plant helpful for the establishment of bee colonies in spring — a period that is often not well-served by commercial flower meadow seed mixes. It also has the situationally-beneficial quality of being a spring ephemeral, as well as an annual that lacks difficult-to-combat roots. Plants that provide necessary structural supports for invertebrate and small vertebrate predators can help to keep overall pest populations low.
The abundant nectar produced by C. solstitialis flowers attracts many pollinators. This is another reason for the success of the (situationally) highly invasive species. Due to genetic differences related to evolutionary adaption, not all members of Centaurea produce the same amount of nectar. Growing conditions, such as climate and soil, can have a very strong impact, even if the plants grow and flower. For instance, cornflower plants, Centaurea cyanus, produced 33% less seasonal nectar than Centaurea nigra in a UK study. C. nigra also ranked higher than ragwort in another UK study, although ragwort was still in the top 10 for yearly nectar production. The strong nectar production of certain members of the genus can be exploited to the farmer's advantage, possibly in combination with biological control. In particular, the yellow starthistle (C. solstitialis) as well as spotted knapweed (C. maculosa) are major honey plants for beekeepers. Monofloral honey from these plants is light and slightly tangy, and one of the finest honeys produced in the United States – due to its better availability, it is even fraudulently relabeled and sold as the scarce and expensive sourwood honey of the Appalachian Mountains. Placing beehives near stands of Centaurea will cause increased pollination. As most seedheads fail however when biocontrol pests have established themselves, the plants will bloom ever more abundantly in an attempt to replace the destroyed seedheads, to the point where they exhaust their resources in providing food for the pests (seeds), bees (pollen) and humans (honey). Output of allelopathic compounds is also liable to be reduced under such conditions – the plant has to compromise between allocating energy to reproduction and defense. This renders the weeds more likely to be suppressed by native vegetation or crops in the following years, especially if properly timed controlled burning[5] and/or targeted grazing by suitable livestock are also employed. While yellow starthistle and perhaps other species are toxic to equines, some other livestock may eat the non-spiny knapweeds with relish. In Europe, common knapweed (C. nigra) and globe knapweed (C. macrocephala) are locally important pollen sources for honeybees in mid-late summer.
8-Hydroxyquinoline has been identified as a main allelopathic compound produced by diffuse knapweed (C. diffusa); native North American plants are typically sensitive to it, while those of Eastern Europe and Asia Minor usually have coevolved with the knapweed and are little harmed if at all, aided by native microorganisms that break down or even feed on the abundantly secreted compound. Thus, 8-hydroxyquinoline is potentially useful to control American plants that have become invasive weeds in the diffuse knapweed's native range.
Arctiin, found in C. imperialis, has shown anticancer activity in laboratory studies. The roots of the long-lost C. foliosa, an endemic of Hatay Province (Turkey), are used in folk medicine, and other species are presumably too. A South Italian variety[verification needed] of the purple starthistle (C. calcitrapa) is traditionally consumed by ethnic Albanians (Arbëreshë people) in the Vulture area (southern Italy); e.g. in the Arbëreshë communities in Lucania the young whorls of C. calcitrapa are boiled and fried in mixtures with other weedy non-cultivated greens. According to research by the Michael Heinrich group at the Centre for Pharmacognosy and Phytotherapy (School of Pharmacy, University of London) "the antioxidant activity [...] of the young whorls of Centaurea calcitrapa, both in the DPPH and in the lipid peroxidation inhibition assays, [is] very interesting and [the] species should be investigated phytochemically and biochemically focusing on these properties". Extracts from C. calcitrapa were furthermore found to have significant xanthine oxidase (XO)-inhibiting activity.
Spotted knapweed as well as other species are rich in cnicin, a bitter compound found mainly in the leaves and often used to flavor the digestif amaro. In western Crete, Greece a local variety[verification needed] of C. calcitrapa called gourounaki (γουρουνάκι "little pig") also has its leaves eaten boiled by the locals. In the same island an endemic local species, C. idaea called katsoula (κατσούλα), tsita (τσίτα) or aspragatha (ασπραγκάθα), has its leaves eaten boiled by the locals too.
Some species are cultivated as ornamental plants in gardens. As regards other aspects of popular culture, cornflower (C. cyanus) is the floral emblem of Östergötland province (Sweden) – where is it called blåklint, literally "blue mountain" – and of Päijänne Tavastia region in Finland, where it is known as ruiskaunokki ("rye-beaks") or ruiskukka ("rye-flower"). It is also the national flower of Estonia where its local name rukkilill means "rye-lily", Belarus where it is called vałoška (Belarusian: валошка), and one of those of Germany where it is called Kornblume ("cornflower"). The origin of the name "caltrop" for the ancient low-tech area denial weapon is probably in some way connected with C. calcitrapa and its spiny seeds. This plant is attested to by the colloquial name "caltrop" at a time when the weapons were still called by their Roman name tribulus. Lastly, the color cornflower blue is named after C. cyanus. Cornflower is also used as a cut flower.
As namesake member of the subtribe Centaureinae of tribe Cardueae, the knapweeds are probably most closely related to genera such as Carthamus (distaff thistles), Cnicus (blessed thistle), Crupina (crupinas) or Notobasis (Syrian thistle), and somewhat less closely to most other thistles. The monotypic Cnicus seems in fact to properly belong in Centaurea.
Research in the late 20th century shows that Centaurea as traditionally defined is polyphyletic. A number of 19th- and 20th-century efforts to reorganize the genus were not successful, and it is not yet clear what the consequences of the recent research will be for classification of this genus and other related genera. The type species C. centaurium stands somewhat apart from the main lineage of knapweeds and thus the taxonomic consequences of a rearrangement might be severe, with hundreds of species needing to be moved to new genera. It has thus been proposed to change the type species to one of the main lineages to avoid this problem. What seems certain however is that the basketflowers – presently treated as a section Plectocephalus – will be reinstated as a distinct genus in the near future. The rock-centauries (Cheirolophus), formerly usually included in Centaurea, are now already treated as separate genus.
Better-known Centaurea species include:
Centaurea acaulis
Centaurea adpressa
Centaurea aegyptiaca
Centaurea aeolica
Centaurea aggregata
Centaurea akamantis – Akamas centaurea
Centaurea alba
Centaurea albonitens Turrill
Centaurea alpestris
Centaurea alpina
Centaurea ambigua
Centaurea amblyolepis
Centaurea americana – American basketflower, American starthistle
Centaurea ammocyanus
Centaurea antennata Dufour
Centaurea antiochia Boiss.
Centaurea aplolepa
Centaurea aplolepa subsp. carueliana
Centaurea appendicigera C.Koch
Centaurea argentea
Centaurea ascalonica
Centaurea aspera L. – rough starthistle
Centaurea atacamensis (Reiche) I.M.Johnst.
Centaurea atropurpurea
Centaurea ×aurata
Centaurea babylonica L.
Centaurea balsamita
Centaurea behen L. – ak behmen (Turkish)
Centaurea bella
Centaurea benedicta – Cnicus
Centaurea bieberseinii
Centaurea borjae
Centaurea bovina
Centaurea bracteata
Centaurea brevifimbriata Hub.-Mor.
Centaurea bulbosa
Centaurea busambarensis Guss.
Centaurea cachinalensis
Centaurea calcitrapa – purple starthistle, red starthistle, "caltrop"
Centaurea calcitrapoides
Centaurea cariensis Boiss.
Centaurea cariensiformis Hub.-Mor.
Centaurea caroli-henrici Gabrieljan & Dittrich
Centaurea centaurium L.
Centaurea chilensis
Centaurea cineraria – velvet centaurea, dusty miller
Centaurea clementei
Centaurea collina L.
Centaurea corymbosa
Centaurea crithmifolia
Centaurea crocodylium
Centaurea cyanoides J.Berggr. & Wahlenb.
Centaurea cyanus – cornflower, bachelor's button, boutonniere flower, hurtsickle, bluebottle, basketflower
Centaurea damascena
Centaurea debeauxii Gren. & Godr.
Centaurea demirizii Wagenitz
Centaurea depressa – low cornflower
Centaurea deusta
Centaurea diffusa – diffuse knapweed, white knapweed, tumble knapweed
Centaurea diluta – North African knapweed
Centaurea drabifolia Sm.
Centaurea drabifolioides Hub.-Mor.
Centaurea dschungarica
Centaurea emilae Hüseynova et Qaraxani[13]
Centaurea eriophora
Centaurea eryngioides
Centaurea filiformis
Centaurea fischeri Willd.
Centaurea floccosa
Centaurea foliosa Boiss. & Kotschy
Centaurea forojuliensis
Centaurea friderici Vis. – palagruška zečina (Croatian)
Centaurea gayana
Centaurea glaberrima Tausch
Centaurea glastifolia
Centaurea grinensis
Centaurea gymnocarpa
Centaurea haradjianii Wagenitz
Centaurea hedgei
Centaurea helenioides Boiss.
Centaurea hermannii F.Hermann
Centaurea horrida Badarò – fiordaliso spinoso (Italian)
Centaurea hyalolepis
Centaurea hypoleuca
Centaurea iberica – Iberian starthistle, Iberian knapweed
Centaurea idaea – katsoula, tsita (Cretan Greek)
Centaurea imperialis Hausskn. ex Bornm.
Centaurea jabukensis
Centaurea jacea – brown knapweed, brownray knapweed
Centaurea kasakorum
Centaurea kopetaghensis
Centaurea kotschyana Heuff.
Centaurea lanulata
Centaurea leptophylla
Centaurea leucophylla
Centaurea limbata
Centaurea lydia Boiss.
Centaurea macrocephala Puschk. ex Willd. – globe knapweed, Armenian basketflower
Centaurea maculosa – spotted knapweed (might belong in C. stoebe subsp. micranthos)
Centaurea mannagettae
Centaurea margaritalba Klok.
Centaurea marschalliana
Centaurea melitensis – Maltese starthistle; tocalote, tocolote (California)
Centaurea minor
Centaurea moschata – sweet sultan
Centaurea ×moncktonii C.E.Britton – meadow knapweed, protean knapweed (= C. ×pratensis Thuill non Salisb.)
Centaurea monocephala
Centaurea montana – montane knapweed, perennial cornflower, mountain cornflower, mountain bluet
Centaurea napifolia L. – fiordaliso romano (Italian)
Centaurea nervosa Rchb. ex Steud.
Centaurea nigra – common knapweed, black knapweed, lesser knapweed, hardheads
Centaurea nigrescens – Tyrol knapweed, short-fringed knapweed, Tyrol thistle
Centaurea nigrifimbria (C.Koch) Sosn.
Centaurea nivea (Bornm.) Wagenitz
Centaurea onopordifolia
Centaurea orientalis L.
Centaurea ornata Willd.
Centaurea ovina
Centaurea pallescens Delile
Centaurea paniculata L.
Centaurea parlatoris
Centaurea pecho
Centaurea phrygia – wig knapweed
Centaurea pindicola
Centaurea polypodiifolia
Centaurea ×pratensis Salisb. (C. jacea × C. nigra) – meadow knapweed
Centaurea procurrens
Centaurea ×psammogena G.Gayer. (C. diffusa × C. stoebe subsp. micranthos)
Centaurea pseudocaerulescens
Centaurea pseudophrygia C.A.Mey.
Centaurea pulcherrima Willd.
Centaurea pullata L.
Centaurea pumilio
Centaurea ragusina L.
Centaurea rigida
Centaurea rothrockii Greenm. – Mexican basketflower, Rothrock's basketflower, Rothrock's knapweed
Centaurea ruthenica
Centaurea rutifolia Sm.
Centaurea sadleriana – Pannonian knapweed
Centaurea salicifolia Bieb. ex Willd.
Centaurea scabiosa – greater knapweed
Centaurea scannensis
Centaurea scoparia
Centaurea scopulorum Boiss. & Heldr.
Centaurea seguenzae
Centaurea seridis L.
Centaurea sibirica
Centaurea simplicicaulis
Centaurea sinaica
Centaurea solstitialis – yellow starthistle, golden starthistle, yellow cockspur, St. Barnaby's thistle, Barnaby thistle
Centaurea speciosa
Centaurea sphaerocephala L.
Centaurea stenolepis
Centaurea stoebe L.
Centaurea stoebe subsp. micranthos (Gugler) Hayek
Centaurea straminicephala
Centaurea sulphurea – Sicilian starthistle
Centaurea tauromenitana Guss.
Centaurea tenoreana
Centaurea tommasinii
Centaurea transalpina Schleich. ex DC.
Centaurea tchihatcheffii — yanardöner (Turkish)
Centaurea trichocephala Bieb. ex Willd. – featherhead knapweed
Centaurea triniifolia
Centaurea triumfettii All.
Centaurea ucriae Lacaita
Centaurea uniflora Turra
Centaurea verbascifolia Vahl
Centaurea verutum L.
Centaurea virgata
Centaurea virgata subsp. squarrosa – squarrose knapweed
Centaurea wiedemanniana Fisch. & Mey.
Centaurea yozgatensis Wagenitz
Formerly placed here
Plant species placed in Centaurea in former times include:
Acroptilon repens – Russian knapweed (as C. repens)
Cheirolophus crassifolius – Maltese rock-centaury (as C. crassifolia, C. spathulata)
Femeniasia balearica (as C. balearica)
Volutaria muricata (as C. muricata)
Centaurea is a genus of over 700 species of herbaceous thistle-like flowering plants in the family Asteraceae. Members of the genus are found only north of the equator, mostly in the Eastern Hemisphere; the Middle East and surrounding regions are particularly species-rich. In the western United States, yellow starthistles are an invasive species. Around the year 1850, seeds from the plant had arrived to the state of California. It is believed that those seeds came from South America.
Common names
Common names for this genus are centaury, centory, starthistles, knapweeds, centaureas and the more ambiguous "bluets"; a vernacular name used for these plants in parts of England is "loggerheads" (common knapweed). The Plectocephalus group – possibly a distinct genus – is known as basketflowers. "Cornflower" is used for a few species, but that term more often specifically means either C. cyanus (the annual cornflower) or Centaurea montana (the perennial cornflower). The common name "centaury" is sometimes used, although this also refers to the unrelated plant genus Centaurium.
The name is said to be in reference to Chiron, the centaur of Greek mythology who discovered medicinal uses of a plant eventually called "centaury".
Description
Knapweeds are robust weedy plants. Their leaves, spiny in some species, are usually deeply divided into elongated lobes at least in the plants' lower part, becoming entire towards the top. The "flowers" (actually pseudanthium inflorescences) are diverse in colour, ranging from intense blues, reds and yellows to any mixture of these and lighter shades towards white. Often, the disk flowers are much darker or lighter than the ray flowers, which also differ in morphology and are sterile. Each pseudanthium sits atop a cup- or basket-like cluster of scaly bracts, hence the name "basketflowers". Many species, in particular those inhabiting more arid regions, have a long and strong taproot.
Certain knapweeds have a tendency to dominate large stretches of landscape together with a few other plants, typically one or two grasses and as many other large herbaceous plants. The common knapweed (C. nigra) for example is plentiful in the mesotrophic grasslands of England and nearby regions. It is most prominently found in pastures or meadows dominated by cock's-foot (Dactylis glomerata) as well as either of crested dog's-tail (Cynosurus cristatus) and false oat-grass (Arrhenatherum elatius). It is also often found in mesotrophic grassland on rendzinas and similar calcareous soils in association with glaucous sedge (Carex flacca), sheep's fescue (Festuca ovina), and either tor-grass (Brachypodium pinnatum) and rough hawkbit (Leontodon hispidus), or upright brome (Bromus erectus). In these grasslands, greater knapweed (C. scabiosa) is found much more rarely by comparison, often in association with red fescue (Festuca rubra) in addition to cock's-foot and false oat-grass.
Due to their habit of dominating ecosystems under good conditions, many Centaurea species can become invasive weeds in regions where they are not native. In parts of North America, diffuse knapweed (C. diffusa), spotted knapweed (C. maculosa) and yellow starthistle (C. solstitialis) cause severe problems in agriculture due to their uncontrolled spread. The seeds are typically transported by human traffic, in particular the tires of all-terrain vehicles. The two knapweeds are harmful mainly because they are strongly allelopathic, producing powerful toxins in their roots that stunt the growth of plants around them not adapted to this. Yellow starthistle, meanwhile, is inedible to most livestock due to its spines and apparently outright poisonous to horses and other equines. However, efficient methods of biological control by insect pests of these weeds have been developed; the knapweeds can also exploited to their detriment by targeted grazing. Controlled burning may also be used, though the timing is important to avoid the plants having seeded already, and neither allowing sufficient time for them to regrow from the rootstock.
Yet other species of Centaurea – mostly ones that occur between Italy and the Caucasus – are endemics of a single island or valley, and some of these are endangered. The Akamas Centaurea (Centaurea akamantis) of Cyprus is almost extinct, while the western Caucasus endemics C. leptophylla and C. straminicephala are at least very rare and C. hedgei and C. pecho from the same region are certainly not abundant either. The last four species would be adversely affected by the proposed Yusufeli Dam, which might actually destroy enough habitat to push the two rarer ones over the brink of extinction.
Centaurea are copious nectar producers, especially on high-lime soils. The high nectar yield of the genus makes it very attractive to insects such as butterflies – including the endangered Karner blue (Plebejus melissa samuelis) which visits introduced spotted knapweed – and day-flying moths – typically Zygaenidae, such as Zygaena loti or the six-spot burnet (Z. filipendulae). The larvae of some other Lepidoptera species use Centaurea species as food plants; see List of Lepidoptera that feed on Centaurea. Several of these are used in biological control of invasive knapweeds and starthistles.
Larvae of several true weevils (Curculionidae) of the subfamily Lixinae also feed on Centaurea. Some genera – such as Larinus whose larval food is flowerheads – have many species especially adapted to particular knapweeds or starthistle and are used in biological control too. These include the yellow starthistle flower weevil (L. curtus) for yellow starthistle, lesser knapweed flower weevil (L. minutus) for diffuse knapweed and blunt knapweed flower weevil (L. obtusus) for spotted knapweed. Broad-nosed seedhead weevil (Bangasternus fausti) larvae eat diffuse, spotted and squarrose knapweed (C. virgata ssp. squarrosa), while those of the yellow starthistle bud weevil (B. orientalis) do not seem to live on anything other than yellow starthistle and occasionally purple starthistle (C. calcitrapa). But perhaps most efficient in destroying developing yellow starthistle seedheads is the larva of the yellow starthistle hairy weevil (Eustenopus villosus). Knapweed root weevil (Cyphocleonus achates) larvae bore into the roots of spotted and to a lesser extentely diffuse knapweed, sometimes killing off the entire plant.
Also used in biological control are Tephritidae (peacock flies) whose larvae feed on Centaurea. Knapweed peacock fly (Chaetorellia acrolophi) larvae eat spotted knapweed and some other species. The yellow starthistle peacock fly (C. australis) has an initial generation each year which often uses cornflower (C. cyanus) as larval food; later generations switch to yellow starthistle. The flies are generally considered less efficient in destroying the growing seedheads than the weevils, but may be superior under certain conditions; employing flies and weevils in combination is expensive and does not noticeably increase their effect.
Use by humans
Although the genus may be considered by a quite significant number of relatively informed individuals to have an overall negative impact on human interests, particularly agricultural interests, the situation is not straightforward enough to simply declare the genus, or, at least, its most aggressively-spreading species, altogether negative. For instance, due to their moderate to high nectar production, which can occur over a comparatively long duration, many species of Centaurea are popular food sources for insects that may otherwise attack certain crops.[citation needed] It may be advisable for some types of farms to allow certain species in this genus, such as cornflower (C. cyanus) in a European setting, to grow adjacent to fields. Although they support and attract many types of beneficial life (not just beetles), these areas are known as beetle banks. When they are present, some pests may be drawn away from crops to them and predatory insects and arachnids that feed upon pest insects will be better-supported by these more naturalized areas. They additionally have the beneficial aspect of supporting pollinators, unlike many field crops such as maize. Moreover, being untreated with pesticides and providing more diversity, plants growing in more wild areas adjacent to farms produce more insects that attract and support birds which can also feed on pests that would harm crops. Insect production is especially high for beetle banks that have enough plants that serve in the role of host plant for immature insects, rather than just in the roles of adult food and/or shelter provision.
Some plants which are considered invasive or problematic in certain areas can have beneficial qualities that outweigh their negative qualities from a human and/or human agricultural point of view, although this sometimes requires some human management – particularly if adequate biological control has not been established for the more aggressive species. An example is wild parsnip, Pastinaca sativa, which produces florets that feed predatory (and other beneficial) insects as well as large tubular stems that provide winter shelter for native bees, wasps, and other organisms that can be beneficial for agriculture. The plant is considered invasive in some areas of the United States and is also often considered undesirable due to its ability to cause contact skin irritation. However, it also serves as a host plant for the black swallowtail butterfly, helps to bring nutrients up from soils with its deep taproot, and possesses evergreen foliage even in climate zones such as US zone 6. This foliage increases soil warmth and moisture which can be beneficial for certain types of life. Perhaps the most dramatic example of a generally disliked plant's beneficial qualities being usually overlooked is the often-despised ragwort, Jacobaea vulgaris, which topped the list by a large amount for nectar production in a UK study, with a production per floral unit of (2921 ± 448μg). This very high nectar production, coupled with its early blooming period, makes the plant helpful for the establishment of bee colonies in spring — a period that is often not well-served by commercial flower meadow seed mixes. It also has the situationally-beneficial quality of being a spring ephemeral, as well as an annual that lacks difficult-to-combat roots. Plants that provide necessary structural supports for invertebrate and small vertebrate predators can help to keep overall pest populations low.
The abundant nectar produced by C. solstitialis flowers attracts many pollinators. This is another reason for the success of the (situationally) highly invasive species. Due to genetic differences related to evolutionary adaption, not all members of Centaurea produce the same amount of nectar. Growing conditions, such as climate and soil, can have a very strong impact, even if the plants grow and flower. For instance, cornflower plants, Centaurea cyanus, produced 33% less seasonal nectar than Centaurea nigra in a UK study. C. nigra also ranked higher than ragwort in another UK study, although ragwort was still in the top 10 for yearly nectar production. The strong nectar production of certain members of the genus can be exploited to the farmer's advantage, possibly in combination with biological control. In particular, the yellow starthistle (C. solstitialis) as well as spotted knapweed (C. maculosa) are major honey plants for beekeepers. Monofloral honey from these plants is light and slightly tangy, and one of the finest honeys produced in the United States – due to its better availability, it is even fraudulently relabeled and sold as the scarce and expensive sourwood honey of the Appalachian Mountains. Placing beehives near stands of Centaurea will cause increased pollination. As most seedheads fail however when biocontrol pests have established themselves, the plants will bloom ever more abundantly in an attempt to replace the destroyed seedheads, to the point where they exhaust their resources in providing food for the pests (seeds), bees (pollen) and humans (honey). Output of allelopathic compounds is also liable to be reduced under such conditions – the plant has to compromise between allocating energy to reproduction and defense. This renders the weeds more likely to be suppressed by native vegetation or crops in the following years, especially if properly timed controlled burning[5] and/or targeted grazing by suitable livestock are also employed. While yellow starthistle and perhaps other species are toxic to equines, some other livestock may eat the non-spiny knapweeds with relish. In Europe, common knapweed (C. nigra) and globe knapweed (C. macrocephala) are locally important pollen sources for honeybees in mid-late summer.
8-Hydroxyquinoline has been identified as a main allelopathic compound produced by diffuse knapweed (C. diffusa); native North American plants are typically sensitive to it, while those of Eastern Europe and Asia Minor usually have coevolved with the knapweed and are little harmed if at all, aided by native microorganisms that break down or even feed on the abundantly secreted compound. Thus, 8-hydroxyquinoline is potentially useful to control American plants that have become invasive weeds in the diffuse knapweed's native range.
Arctiin, found in C. imperialis, has shown anticancer activity in laboratory studies. The roots of the long-lost C. foliosa, an endemic of Hatay Province (Turkey), are used in folk medicine, and other species are presumably too. A South Italian variety[verification needed] of the purple starthistle (C. calcitrapa) is traditionally consumed by ethnic Albanians (Arbëreshë people) in the Vulture area (southern Italy); e.g. in the Arbëreshë communities in Lucania the young whorls of C. calcitrapa are boiled and fried in mixtures with other weedy non-cultivated greens. According to research by the Michael Heinrich group at the Centre for Pharmacognosy and Phytotherapy (School of Pharmacy, University of London) "the antioxidant activity [...] of the young whorls of Centaurea calcitrapa, both in the DPPH and in the lipid peroxidation inhibition assays, [is] very interesting and [the] species should be investigated phytochemically and biochemically focusing on these properties". Extracts from C. calcitrapa were furthermore found to have significant xanthine oxidase (XO)-inhibiting activity.
Spotted knapweed as well as other species are rich in cnicin, a bitter compound found mainly in the leaves and often used to flavor the digestif amaro. In western Crete, Greece a local variety[verification needed] of C. calcitrapa called gourounaki (γουρουνάκι "little pig") also has its leaves eaten boiled by the locals. In the same island an endemic local species, C. idaea called katsoula (κατσούλα), tsita (τσίτα) or aspragatha (ασπραγκάθα), has its leaves eaten boiled by the locals too.
Some species are cultivated as ornamental plants in gardens. As regards other aspects of popular culture, cornflower (C. cyanus) is the floral emblem of Östergötland province (Sweden) – where is it called blåklint, literally "blue mountain" – and of Päijänne Tavastia region in Finland, where it is known as ruiskaunokki ("rye-beaks") or ruiskukka ("rye-flower"). It is also the national flower of Estonia where its local name rukkilill means "rye-lily", Belarus where it is called vałoška (Belarusian: валошка), and one of those of Germany where it is called Kornblume ("cornflower"). The origin of the name "caltrop" for the ancient low-tech area denial weapon is probably in some way connected with C. calcitrapa and its spiny seeds. This plant is attested to by the colloquial name "caltrop" at a time when the weapons were still called by their Roman name tribulus. Lastly, the color cornflower blue is named after C. cyanus. Cornflower is also used as a cut flower.
As namesake member of the subtribe Centaureinae of tribe Cardueae, the knapweeds are probably most closely related to genera such as Carthamus (distaff thistles), Cnicus (blessed thistle), Crupina (crupinas) or Notobasis (Syrian thistle), and somewhat less closely to most other thistles. The monotypic Cnicus seems in fact to properly belong in Centaurea.
Research in the late 20th century shows that Centaurea as traditionally defined is polyphyletic. A number of 19th- and 20th-century efforts to reorganize the genus were not successful, and it is not yet clear what the consequences of the recent research will be for classification of this genus and other related genera. The type species C. centaurium stands somewhat apart from the main lineage of knapweeds and thus the taxonomic consequences of a rearrangement might be severe, with hundreds of species needing to be moved to new genera. It has thus been proposed to change the type species to one of the main lineages to avoid this problem. What seems certain however is that the basketflowers – presently treated as a section Plectocephalus – will be reinstated as a distinct genus in the near future. The rock-centauries (Cheirolophus), formerly usually included in Centaurea, are now already treated as separate genus.
Better-known Centaurea species include:
Centaurea acaulis
Centaurea adpressa
Centaurea aegyptiaca
Centaurea aeolica
Centaurea aggregata
Centaurea akamantis – Akamas centaurea
Centaurea alba
Centaurea albonitens Turrill
Centaurea alpestris
Centaurea alpina
Centaurea ambigua
Centaurea amblyolepis
Centaurea americana – American basketflower, American starthistle
Centaurea ammocyanus
Centaurea antennata Dufour
Centaurea antiochia Boiss.
Centaurea aplolepa
Centaurea aplolepa subsp. carueliana
Centaurea appendicigera C.Koch
Centaurea argentea
Centaurea ascalonica
Centaurea aspera L. – rough starthistle
Centaurea atacamensis (Reiche) I.M.Johnst.
Centaurea atropurpurea
Centaurea ×aurata
Centaurea babylonica L.
Centaurea balsamita
Centaurea behen L. – ak behmen (Turkish)
Centaurea bella
Centaurea benedicta – Cnicus
Centaurea bieberseinii
Centaurea borjae
Centaurea bovina
Centaurea bracteata
Centaurea brevifimbriata Hub.-Mor.
Centaurea bulbosa
Centaurea busambarensis Guss.
Centaurea cachinalensis
Centaurea calcitrapa – purple starthistle, red starthistle, "caltrop"
Centaurea calcitrapoides
Centaurea cariensis Boiss.
Centaurea cariensiformis Hub.-Mor.
Centaurea caroli-henrici Gabrieljan & Dittrich
Centaurea centaurium L.
Centaurea chilensis
Centaurea cineraria – velvet centaurea, dusty miller
Centaurea clementei
Centaurea collina L.
Centaurea corymbosa
Centaurea crithmifolia
Centaurea crocodylium
Centaurea cyanoides J.Berggr. & Wahlenb.
Centaurea cyanus – cornflower, bachelor's button, boutonniere flower, hurtsickle, bluebottle, basketflower
Centaurea damascena
Centaurea debeauxii Gren. & Godr.
Centaurea demirizii Wagenitz
Centaurea depressa – low cornflower
Centaurea deusta
Centaurea diffusa – diffuse knapweed, white knapweed, tumble knapweed
Centaurea diluta – North African knapweed
Centaurea drabifolia Sm.
Centaurea drabifolioides Hub.-Mor.
Centaurea dschungarica
Centaurea emilae Hüseynova et Qaraxani[13]
Centaurea eriophora
Centaurea eryngioides
Centaurea filiformis
Centaurea fischeri Willd.
Centaurea floccosa
Centaurea foliosa Boiss. & Kotschy
Centaurea forojuliensis
Centaurea friderici Vis. – palagruška zečina (Croatian)
Centaurea gayana
Centaurea glaberrima Tausch
Centaurea glastifolia
Centaurea grinensis
Centaurea gymnocarpa
Centaurea haradjianii Wagenitz
Centaurea hedgei
Centaurea helenioides Boiss.
Centaurea hermannii F.Hermann
Centaurea horrida Badarò – fiordaliso spinoso (Italian)
Centaurea hyalolepis
Centaurea hypoleuca
Centaurea iberica – Iberian starthistle, Iberian knapweed
Centaurea idaea – katsoula, tsita (Cretan Greek)
Centaurea imperialis Hausskn. ex Bornm.
Centaurea jabukensis
Centaurea jacea – brown knapweed, brownray knapweed
Centaurea kasakorum
Centaurea kopetaghensis
Centaurea kotschyana Heuff.
Centaurea lanulata
Centaurea leptophylla
Centaurea leucophylla
Centaurea limbata
Centaurea lydia Boiss.
Centaurea macrocephala Puschk. ex Willd. – globe knapweed, Armenian basketflower
Centaurea maculosa – spotted knapweed (might belong in C. stoebe subsp. micranthos)
Centaurea mannagettae
Centaurea margaritalba Klok.
Centaurea marschalliana
Centaurea melitensis – Maltese starthistle; tocalote, tocolote (California)
Centaurea minor
Centaurea moschata – sweet sultan
Centaurea ×moncktonii C.E.Britton – meadow knapweed, protean knapweed (= C. ×pratensis Thuill non Salisb.)
Centaurea monocephala
Centaurea montana – montane knapweed, perennial cornflower, mountain cornflower, mountain bluet
Centaurea napifolia L. – fiordaliso romano (Italian)
Centaurea nervosa Rchb. ex Steud.
Centaurea nigra – common knapweed, black knapweed, lesser knapweed, hardheads
Centaurea nigrescens – Tyrol knapweed, short-fringed knapweed, Tyrol thistle
Centaurea nigrifimbria (C.Koch) Sosn.
Centaurea nivea (Bornm.) Wagenitz
Centaurea onopordifolia
Centaurea orientalis L.
Centaurea ornata Willd.
Centaurea ovina
Centaurea pallescens Delile
Centaurea paniculata L.
Centaurea parlatoris
Centaurea pecho
Centaurea phrygia – wig knapweed
Centaurea pindicola
Centaurea polypodiifolia
Centaurea ×pratensis Salisb. (C. jacea × C. nigra) – meadow knapweed
Centaurea procurrens
Centaurea ×psammogena G.Gayer. (C. diffusa × C. stoebe subsp. micranthos)
Centaurea pseudocaerulescens
Centaurea pseudophrygia C.A.Mey.
Centaurea pulcherrima Willd.
Centaurea pullata L.
Centaurea pumilio
Centaurea ragusina L.
Centaurea rigida
Centaurea rothrockii Greenm. – Mexican basketflower, Rothrock's basketflower, Rothrock's knapweed
Centaurea ruthenica
Centaurea rutifolia Sm.
Centaurea sadleriana – Pannonian knapweed
Centaurea salicifolia Bieb. ex Willd.
Centaurea scabiosa – greater knapweed
Centaurea scannensis
Centaurea scoparia
Centaurea scopulorum Boiss. & Heldr.
Centaurea seguenzae
Centaurea seridis L.
Centaurea sibirica
Centaurea simplicicaulis
Centaurea sinaica
Centaurea solstitialis – yellow starthistle, golden starthistle, yellow cockspur, St. Barnaby's thistle, Barnaby thistle
Centaurea speciosa
Centaurea sphaerocephala L.
Centaurea stenolepis
Centaurea stoebe L.
Centaurea stoebe subsp. micranthos (Gugler) Hayek
Centaurea straminicephala
Centaurea sulphurea – Sicilian starthistle
Centaurea tauromenitana Guss.
Centaurea tenoreana
Centaurea tommasinii
Centaurea transalpina Schleich. ex DC.
Centaurea tchihatcheffii — yanardöner (Turkish)
Centaurea trichocephala Bieb. ex Willd. – featherhead knapweed
Centaurea triniifolia
Centaurea triumfettii All.
Centaurea ucriae Lacaita
Centaurea uniflora Turra
Centaurea verbascifolia Vahl
Centaurea verutum L.
Centaurea virgata
Centaurea virgata subsp. squarrosa – squarrose knapweed
Centaurea wiedemanniana Fisch. & Mey.
Centaurea yozgatensis Wagenitz
Formerly placed here
Plant species placed in Centaurea in former times include:
Acroptilon repens – Russian knapweed (as C. repens)
Cheirolophus crassifolius – Maltese rock-centaury (as C. crassifolia, C. spathulata)
Femeniasia balearica (as C. balearica)
Volutaria muricata (as C. muricata)
Euphorbia is a very large and diverse genus of flowering plants, commonly called spurge, in the family Euphorbiaceae. "Euphorbia" is sometimes used in ordinary English to collectively refer to all members of Euphorbiaceae (in deference to the type genus), not just to members of the genus.
Euphorbias range from tiny annual plants to large and long-lived trees. with perhaps the tallest being Euphorbia ampliphylla at 30 m (98 ft) or more. The genus has roughly 2,000 members, making it one of the largest genera of flowering plants. It also has one of the largest ranges of chromosome counts, along with Rumex and Senecio. Euphorbia antiquorum is the type species for the genus Euphorbia. It was first described by Carl Linnaeus in 1753 in Species Plantarum.
Some euphorbias are widely available commercially, such as poinsettias at Christmas. Some are commonly cultivated as ornamentals, or collected and highly valued for the aesthetic appearance of their unique floral structures, such as the crown of thorns plant (Euphorbia milii). Succulent euphorbias from the deserts of Southern Africa and Madagascar have evolved physical characteristics and forms similar to cacti of North and South America, so they are often incorrectly referred to as cacti. Some are used as ornamentals in landscaping, because of beautiful or striking overall forms, and drought and heat tolerance.
Euphorbia all share the feature of having a poisonous, latex-like sap and unique floral structures When viewed as a whole, the head of flowers looks like a single flower (a pseudanthium). It has a unique kind of pseudanthium, called a cyathium, where each flower in the head is reduced to its barest essential part needed for sexual reproduction. The individual flowers are either male or female, with the male flowers reduced to only the stamen, and the females to the pistil. These flowers have no sepals, petals, or other parts that are typical of flowers in other kinds of plants. Structures supporting the flower head and other structures underneath have evolved to attract pollinators with nectar, and with shapes and colors that function in a way petals and other flower parts do in other flowers. It is the only genus of plants that has all three kinds of photosynthesis, CAM, C3 and C4.[7]
Etymology
The common name "spurge" derives from the Middle English/Old French espurge ("to purge"), due to the use of the plant's sap as a purgative. The botanical name Euphorbia derives from Euphorbos, the Greek physician of King Juba II of Numidia and Mauretania (52–50 BC – 23 AD), who married the daughter of Anthony and Cleopatra. Juba was a prolific writer on various subjects, including natural history. Euphorbos wrote that one of the cactus-like euphorbias (now called Euphorbia obtusifolia ssp. regis-jubae) was used as a powerful laxative. In 12 BC, Juba named this plant after his physician Euphorbos, as Augustus Caesar had dedicated a statue to the brother of Euphorbos, Antonius Musa, who was the personal physician of Augustus. In 1753, botanist and taxonomist Carl Linnaeus assigned the name Euphorbia to the entire genus in the physician's honor.
Description
The plants are annual, biennial or perennial herbs, woody shrubs, or trees with a caustic, poisonous milky latex. The roots are fine or thick and fleshy or tuberous. Many species are more or less succulent, thorny, or unarmed. The main stem and mostly also the side arms of the succulent species are thick and fleshy, and often winged, 15–91 cm (6–36 in) tall. The succulent trees and large shrubs are mostly confined to southern and eastern Africa, but Euphorbia neutra is native to the Caatinga of Brazil and Euphorbia royleana is found in the Himalayan foothills.[14] The deciduous[citation needed] leaves may be opposite, alternate, or in whorls. In succulent species, the leaves are mostly small and short-lived. The stipules are mostly small, partly transformed into spines or glands, or missing.
Inflorescence and fruit
Like all members of the family Euphorbiaceae, spurges have unisexual flowers.
In Euphorbia, flowers occur in a head, called the cyathium (plural cyathia). Each male or female flower in the cyathium head has only its essential sexual part, in males the stamen, and in females the pistil. The flowers do not have sepals, petals, or nectar to attract pollinators, although other nonflower parts of the plant have an appearance and nectar glands with similar roles. Euphorbias are the only plants known to have this kind of flower head.
Nectar glands and nectar that attract pollinators are held in the involucre, a cup-like part below and supporting the cyathium head. The "involucre" in the genus Euphorbia is not to be confused with the "involucre" in family Asteraceae members, which is a collection of bracts called phyllaries, which surround and encase the unopened flower head, then support the receptacle under it after the flower head opens.
The involucre is above and supported by bract-like modified leaf structures (usually in pairs)[citation needed] called cyathophylls', or cyathial leaves. The cyathophyll often has a superficial appearance of being petals of a flower.
Euphorbia flowers are tiny, and the variation attracting different pollinators, with different forms and colors occurs, in the cyathium, involucre, cyathophyll, or additional parts such as glands that attached to these.
The collection of many flowers may be shaped and arranged to appear collectively as a single individual flower, sometimes called a pseudanthium in the Asteraceae, and also in Euphorbia.
The majority of species are monoecious (bearing male and female flowers on the same plant), although some are dioecious with male and female flowers occurring on different plants. It is not unusual for the central cyathia of a cyme to be purely male, and for lateral cyathia to carry both sexes. Sometimes, young plants or those growing under unfavorable conditions are male only, and only produce female flowers in the cyathia with maturity or as growing conditions improve.
The female flowers reduced to a single pistil usually split into three parts, often with two stigmas at each tip.[citation needed] Male flowers often have anthers in twos.[citation needed] Nectar glands usually occur in fives, may be as few as one,[16] and may be fused into a "U" shape. The cyathophylls often occur in twos, are leaf-like, and may be showy and brightly coloured and attractive to pollinators, or be reduced to barely visible tiny scales.
The fruits are three- or rarely two-compartment capsules, sometimes fleshy, but almost always ripening to a woody container that then splits open, sometimes explosively. The seeds are four-angled, oval, or spherical, and some species have a caruncle.
Xerophytes and succulents
In the genus Euphorbia, succulence in the species has often evolved divergently and to differing degrees. Sometimes, it is difficult to decide, and is a question of interpretation, whether or not a species is really succulent or "only" xerophytic. In some cases, especially with geophytes, plants closely related to the succulents are normal herbs. About 850 species are succulent in the strictest sense. If one includes slightly succulent and xerophytic species, this figure rises to about 1000, representing about 45% of all Euphorbia species.
Irritants
The milky sap of spurges (called "latex") evolved as a deterrent to herbivores. It is white, and transparent when dry, except in E. abdelkuri, where it is yellow. The pressurized sap seeps from the slightest wound and congeals after a few minutes in air. The skin-irritating and caustic effects are largely caused by varying amounts of diterpenes. Triterpenes such as betulin and corresponding esters are other major components of the latex. In contact with mucous membranes (eyes, nose, mouth), the latex can produce extremely painful inflammation. The sap has also been known to cause mild to extreme Keratouveitis, which affects vision. Therefore, spurges should be handled with caution and kept away from children and pets. Wearing eye protection while working in close contact with Euphorbia is advised.[18] Latex on skin should be washed off immediately and thoroughly. Congealed latex is insoluble in water, but can be removed with an emulsifier such as milk or soap. A physician should be consulted if inflammation occurs, as severe eye damage including permanent blindness may result from exposure to the sap.
The poisonous qualities were well known: in the Ethiopian Kebra Nagast, the serpent king Arwe is killed with juice from the Euphorbia.
Uses
Several spurges are grown as garden plants, among them poinsettia (E. pulcherrima) and the succulent E. trigona. E. pekinensis (Chinese: 大戟; pinyin: dàjǐ) is used in traditional Chinese medicine, where it is regarded as one of the 50 fundamental herbs. Several Euphorbia species are used as food plants by the larvae of some Lepidoptera (butterflies and moths), like the spurge hawkmoths (Hyles euphorbiae and Hyles tithymali), as well as the giant leopard moth.
Ingenol mebutate, a drug used to treat actinic keratosis, is a diterpenoid found in Euphorbia peplus.
Euphorbias are often used as hedging plants in many parts of Africa.
Among laypeople, Euphorbia species are among the plant taxa most commonly confused with cacti, especially the stem succulents. Euphorbias secrete a sticky, milky-white fluid with latex, but cacti do not. Individual flowers of euphorbias are usually tiny and nondescript (although structures around the individual flowers may not be), without petals and sepals, unlike cacti, which often have fantastically showy flowers. Euphorbias from desert habitats with growth forms similar to cacti have thorns, which are different from the spines of cacti.
Systematics and taxonomy
The present taxon "Euphorbia" corresponds to its own former subtribe, the Euphorbiinae.[citation needed] It has over 2000 species. Morphological description using the presence of a cyathium (see section above) is consistent with nuclear and chloroplast DNA sequence data in testing of about 10% of its members. This testing supports inclusion of formerly other genera as being best placed in this single genus, including Chamaesyce, Monadenium, Pedilanthus, and poinsettia (E. pulcherrima).
Genetic tests have shown that similar flower head structures or forms within the genus, might not mean close ancestry within the genus. The genetic data show that within the genus, convergent evolution of inflorescence structures may be from ancestral subunits that are not related. So using morphology within the genus becomes problematic for further subgeneric grouping. As stated on the Euphorbia Planetary Biodiversity Inventory project webpage:
Previous morphologically based delimitations of subgenera or sections within the genus should not be taken at face value. The genus is in fact rife with striking examples of morphological convergence in cyathial and vegetative features, which justifies a global approach to studying the genus to obtain a proper phylogenetic understanding of the whole group.... The bottom line is that a number of clades have been placed inside or outside of Euphorbia at different times... few of the subgeneric circumscriptions hold up under DNA sequence analysis.
According to a 2002 publication on studies of DNA sequence data, most of the smaller "satellite genera" around the huge genus Euphorbia nest deep within the latter. Consequently, these taxa, namely the never generally accepted genus Chamaesyce, as well as the smaller genera Cubanthus, Elaeophorbia, Endadenium, Monadenium, Synadenium, and Pedilanthus were transferred to Euphorbia. The entire subtribe Euphorbiinae now consists solely of the genus Euphorbia.
Selected species
See List of Euphorbia species for complete list.
Euphorbia albomarginata – rattlesnake weed, white-margined sandmat
Euphorbia amygdaloides – wood spurge
Euphorbia antisyphilitica – candelilla
Euphorbia balsamifera – sweet tabaiba (Canary Islands)
Euphorbia bulbispina
Euphorbia caducifolia – leafless milk hedge
Euphorbia canariensis – Canary Island spurge, Hercules club (Canary Islands)
Euphorbia candelabrum – candelabra tree of East Africa
Euphorbia caput-medusae – Medusa's head (South Africa)
Euphorbia ceratocarpa – (Sicily and southern Italy)
Euphorbia characias – Mediterranean spurge
Euphorbia coerulescens - blue euphorbia
Euphorbia cotinifolia – copper tree
Euphorbia cyathophora – fire-on-the-mountain
Euphorbia cyparissias – Cypress spurge
Euphorbia decidua
Euphorbia dendroides – tree spurge
Euphorbia epithymoides – cushion spurge
Euphorbia esula – leafy spurge
Euphorbia franckiana
Euphorbia fulgens – scarlet plume
Euphorbia grantii – African milk bush
Euphorbia gregersenii – Gregersen's spurge
Euphorbia griffithii – Griffith's spurge
Euphorbia helioscopia – sun spurge
Euphorbia heterophylla – painted euphorbia, desert poinsettia, fireplant, paint leaf, kaliko
Euphorbia hirta – asthma-plant
Euphorbia hispida
Euphorbia horrida – African milk barrel
Euphorbia ingens – candelabra tree
Euphorbia labatii
Euphorbia lactea – mottled spurge, frilled fan, elkhorn
Euphorbia lathyris – caper spurge, paper spurge, gopher spurge, gopher plant, mole plant
Euphorbia leuconeura – Madagascar jewel
Euphorbia maculata – spotted spurge, prostrate spurge
Euphorbia marginata – snow on the mountain
Euphorbia mammillaris
Euphorbia maritae
Euphorbia milii – crown-of-thorns, Christ plant
Euphorbia misera – cliff spurge, Baja California, Southern California
Euphorbia myrsinites – myrtle spurge, creeping spurge, donkey tail
Euphorbia nivulia – leafy milk hedge
Euphorbia obesa
Euphorbia paralias – sea spurge
Euphorbia pekinensis – Peking spurge
Euphorbia peplis – purple spurge
Euphorbia peplus – petty spurge
Euphorbia polychroma – bonfire
Euphorbia psammogeton – sand spurge
Euphorbia pulcherrima – poinsettia, Mexican flame leaf, Christmas star, winter rose, noche buena, lalupatae, pascua, Atatürk çiçeği (Turkish)
Euphorbia purpurea – Darlington's glade spurge, glade spurge, or purple spurge
Euphorbia resinifera – resin spurge
Euphorbia rigida – gopher spurge, upright myrtle spurge
Euphorbia serrata – serrated spurge, sawtooth spurge
Euphorbia tirucalli – Indian tree spurge, milk bush, pencil tree, firestick
Euphorbia tithymaloides – devil's backbone, redbird cactus, cimora misha (Peru)
Euphorbia trigona – African milk tree, cathedral cactus, Abyssinian euphorbia
Euphorbia tuberosa
Euphorbia virosa – gifboom or poison tree
Hybrids
Euphorbia has been extensively hybridised for garden use, with many cultivars available commercially. Moreover, some hybrid plants have been found growing in the wild, for instance E. × martini Rouy, a cross of E. amygdaloides × E. characias subsp. characias, found in southern France.
Subgenera
The genus Euphorbia is one of the largest and most complex genera of flowering plants, and several botanists have made unsuccessful attempts to subdivide the genus into numerous smaller genera. According to the recent phylogenetic studies, Euphorbia can be divided into four subgenera, each containing several sections and groups. Of these, subgenus Esula is the most basal. The subgenera Chamaesyce and Euphorbia are probably sister taxa, but very closely related to subgenus Rhizanthium. Extensive xeromorph adaptations in all probability evolved several times; it is not known if the common ancestor of the cactus-like Rhizanthium and Euphorbia lineages had been xeromorphic—in which case a more normal morphology would have re-evolved namely in Chamaesyce—or whether extensive xeromorphism is entirely polyphyletic even to the level of the subgenera.
Hike around the Coteau Maigre.
Wanderung um den Coteau Maigre.
Calla (Zantedeschia aethiopica) blossom
Calla (Zantedeschia aethiopica) Blüte
Zantedeschia /ˌzæntɪˈdɛskiə/ is a genus of eight species of herbaceous, perennial, flowering plants in the aroid family, Araceae, native to southern Africa (from South Africa northeast to Malawi). The genus has been introduced, in some form, on every continent (minus Antarctica).
Common names include arum lily for Z. aethiopica, calla and calla lily for Z. elliottiana and Z. rehmannii. However, members of this genus are not true lilies (which belong to the family Liliaceae), and the genera Arum and Calla, although related, are distinct from Zantedeschia, despite visual similarities. The colourful flowers and leaves, of both species and cultivars, are greatly valued and commonly grown as ornamental and garden plants.
Description
Zantedeschia species are rhizomatous, herbaceous, perennial plants with some species, e. g., Zantedeschia aethiopica, growing to 1.2m tall, while Zantedeschia rehmannii does not exceed 60 cm in height, growing in clumps or clusters.
Roots: Contractile, emerging from the top of the tubers in Group II.
Stem: The underground portion is variously described as a thick underground stem, i. e., a rhizome or tuber. While the literature is confusing as to the exact terminology, generally the Zantedeschia aethiopica-Zantedeschia odorata group (Group I) is considered to have rhizomes and the remaining species tubers. The rhizomes are fleshy and branched.
Leaves: Petioles are long, spongy, sheathed at the bases, and of varying lengths, from 15 cm (Zantedeschia rehmannii) to 1.5 m (Zantedeschia aethiopica). The lamina is simple, elongated, and coriaceous with a variety of shapes, including triangular, oval (ovate), with or without a point (elliptic), heart-shaped (cordate), spear-shaped (hastate), lance-shaped (lanceolate), oblong, or circular (orbicular). 15–60 cm in length, 5–25 cm in width. The leaves are dark green in colour, feather-veined (pinnate), and may be erect or spreading with undulate margins. Some species exhibit transparent flecking (maculation), and are therefore described as maculate, while others are immaculate. (see Table I, also New Zealand Calla Council Leaf Shape Images The leaves contain hydathodes that result in guttation.
Inflorescence: Takes the form of a solitary pseudanthium (false flower), with a showy white or yellow spathe (a specialised petal like bract) shaped like a funnel with a yellow, central, finger-like spadix, which carries the true flowers. Both spathe and spadix are carried on or above the leaves on the fleshy flower stem. The shape of the spathe whose overlapping margins form the tubes varies from trumpet shaped (Z. pentlandii) to a tight tube with a tapering tip (Z. rehmannii). The spathe is initially green, but as it unfolds becomes coloured. This may be white as in Z. aethiopica, but other species include yellow and pink. Cultivars have a wide variety of other spathe colours including orange and purple. Inside the spathe, the throat may be darkly coloured. The spathe acts to attract pollinators.
Flowers: Zantedeschia is monoecious, in which separate male (staminate) and female (pistillate) flowers ("imperfect" or "unisexual" flowers) are carried on the spadix. The flowers are small and non-blooming without a perianth. The male flowers contain two to three stamens fused to form a synandrium, and the female flowers have a single, compound pistil with three fused carpels and three locules.
Taxonomy
Zantedeschia is the sole genus in the tribe Zantedeschieae in a 1997 classification of the Araceae.
Species
Eight species are currently recognized:
Zantedeschia aethiopica (L.) Spreng. – giant white arum lily or common arum lily - South Africa, Eswatini, Lesotho
Zantedeschia albomaculata (Hook.) Baill. – spotted arum lily - widespread from South Africa north to Nigeria and Tanzania
Zantedeschia elliottiana (W.Watson) Engl. – yellow or golden arum lily - Mpumalanga Province of South Africa
Zantedeschia jucunda Letty - Leolo Mountains of northern South Africa
Zantedeschia odorata P.L.Perry - Western Cape Province
Zantedeschia pentlandii (R.Whyte ex W.Watson) Wittm. - Mpumalanga Province of South Africa
Zantedeschia rehmannii Engl. – pink arum lily - South Africa, Eswatini, Mozambique
Zantedeschia valida (Letty) Y.Singh - KwaZulu-Natal Province of South Africa
Etymology
The name of the genus was given as a tribute to Italian botanist Giovanni Zantedeschi (1773–1846) by the German botanist Kurt Sprengel (1766–1833).
Distribution and habitat
All species are endemic to central and southern Africa, from Nigeria to Tanzania and South Africa. Z. aethiopica grows naturally in marshy areas and is only deciduous when water becomes scarce. It grows continuously when watered and fed regularly and can survive periods of minor frosts. Z. aethiopica is a very strong and sturdy plant, being able to grow in many soils and habitats, multiplying by rhizome-offsets; it is naturalised and regarded as a weed throughout much of the world. Z. odorata is a rare species, resembling Z. aethiopica, but deciduous and smelling like freesia, endemic to a few localities in South Africa. Z. albomaculata is a widespread and variable species, growing from South Africa north to Kenya, varying in shades of white to cream and pink to orange-shades. Z. jucunda and Z. pentlandii are rare species with large yellow showy flowers. Z. rehmannii is a pink-flowered species with sword shaped leaves. Z. elliotiana is known from horticultural sources only and is probably of hybrid origin.
Introduction
Zantedeschia was introduced to Europe in the seventeenth century as Z. aethiopica, and is now widely naturalised in Europe, North America, Central America, South America, Oceania, and Australasia. In many places it is considered a dangerous invasive species that displaces native vegetation. In the South-West of Western Australia, Z. aethiopica was introduced for horticulture, but has become a widespread and conspicuous weed of watercourses, heath, and wet pastures to the extent that it has been declared a pest in Western Australia and landowners must control it and attempts to sell plants must be reported. Zantedeschia in North America is primarily grown as ornamental cultivars in home gardens.
Habitat
Z. aethiopica grows naturally in marshy areas and is only deciduous when water becomes scarce. It grows continuously when watered and fed regularly and can survive periods of minor frosts. Z. aethiopica is a very strong and sturdy plant, being able to grow in many soils and habitats, multiplying by rhizome-offsets.
Cultivation
All Zantedeschia produce large, showy flowers spathes and are often grown both as ornamental plants and for cut flowers. Zantedeschia are relatively hardy plants, but some are more winter-hardy than others. In this regard there may be considered two groups, a hardy outdoor group with large white flowers (arum lilies) and less hardy group with white-spotted leaves and flowers in many colours (calla lilies), such as yellow, orange, pink and purple.
Hardy forms (arum lilies)
These include Zantedeschia aethiopica and Zantedeschia pentlandii and their cultivars. Zantedeschia aethiopica and some of its relatives can survive at minimum winter temperatures below -23 °C (USDA Zone 6) and many others can be grown in even warmer areas where all the ground does not freeze (USDA Zone 7). Z. pentlandii hybrids include 'Millennium Gold'.
Tender forms (calla lilies)
The more tender specimens are mainly cultivars (hybrids) of Zantedeschia elliotiana and Zantedeschia rehmannii (referred to as elliotiana or rehmannii cultivars or hybrids, or as e.g. Z. × rehmanii), but also Zantedeschia albomaculata and Zantedeschia jucunda. These less hardy forms can only survive winter temperatures to −12 °C (Zones 8). This plant must be grown as tender bulbs or houseplants in cooler areas. Species and hybrids between Z. elliotiana, Z. jucunda, Z. pentlandii and Z. rehmannii appear to have an optimum temperature for growth near 25 °C, with growth being suppressed once daily average temperatures persist at 28 °C.
Seasonal grouping
Other classifications consider two groups based on their seasonal habits. Zantedeschia aethiopica and Zantedeschia odorata form one group (Group I) typified by retaining their leaves in winter, and flowering from late winter to late spring, while the remaining species (Group II) are in leaf from spring to late autumn shedding their leaves in winter (deciduous) and flower during the summer. Zantedeschia aethiopica may retain its leaves all year round (evergreen), otherwise from autumn to late summer, while Zantedeschia odorata retains its leaves from late winter to late spring. The two groups also vary according to the arrangement of the male and female organs. In the first group they are arranged together in the lower part of the spadix, whereas in the latter they are separate, with the female at the base. The Z. aethiopica group also have a fruit that turns soft and orange, whereas the other retains a firm green fruit.
Toxicity
Zantedeschia shares the general properties of the family Araceae in causing contact irritation. Zantedeschia species are also poisonous due to the presence of calcium oxalate crystals in the form of raphides. All parts of the plant are poisonous, typically producing local irritation or a burning sensation in the mouth and occasionally vomiting and diarrhea. However leaves are sometimes cooked and eaten.
Uses
Extensive commercial production of Zantedeschia for cut flowers and/or planting material occurs in California, Colombia, New Zealand and Kenya. Plant breeders in California and New Zealand continue to produce an extensive range of new hybrid cultivars. The so-called white calla derived from Z. aethiopica. All varieties with flowers with shades of yellow, orange, red, purple are mainly derived from Z. albomaculata, Z. pentlandii, Z. elliottiana and Z. rehmanni.
Culture
Zantedeschia has often been used in paintings and is featured in many of Diego Rivera's works of art (see The Flower Vendor, amongst others). It was a favourite subject of the painter Georgia O'Keeffe.
(Wikipedia)
Die Zantedeschien (Zantedeschia) sind die einzige Pflanzengattung der Tribus Zantedeschieae in der Familie der Aronstabgewächse (Araceae).
Bei vier der acht Zantedeschia-Arten verwendet man Calla, Kalla oder selten Calla-Lilien als Trivialnamen, sie sollten jedoch nicht mit Lilien oder Pflanzenarten der Gattung Calla verwechselt werden. Manche Arten und Sorten sind seltene Zimmerpflanzen. Der Name Calla leitet sich vom altgriechischen Wort καλός kalós ab und bedeutet „schön“. Namensverwandte aus der griechischen Mythologie sind die Nymphe Kallisto („die Schönste“) und die Muse der Dichtung und Wissenschaft Kalliope („die Schönstimmige“).
Beschreibung
Zantedeschien sind ausdauernde krautige Pflanzen, die Wuchshöhen bis 2,5 Meter erreichen. Sie bilden ein kriechendes Rhizom oder Knollen als Überdauerungsorgan aus. Manche Arten sind immergrün, andere ziehen zu ungünstigen Jahreszeiten ihr Laub ein. Die grundständigen Laubblätter sind – im Gegensatz zu vielen anderen Gattungen der Einkeimblättrigen Pflanzen – gestielt; die Blattspreiten sind glänzend grün und oft pfeilförmig.
Auf einem unbeblätterten Blütenstandschaft ist der Blütenstand wie bei allen Aronstabgewächsen ausgebildet: Ein einzelnes, oft auffällig gefärbtes Hochblatt, die Spatha, umgibt den Kolben. Die große und deutlich zugespitzte Spatha[2] kann weiß oder gelb sein, selten auch rot, bei Züchtungen gibt es eine große Bandbreite an unterschiedlichen Farben. Zantedeschien sind einhäusig getrenntgeschlechtig (monözisch). Im Kolben (Spadix) stehen viele Blüten zusammen. Im unteren Bereich des Kolbens befindet sich eine Zone nur mit weiblichen Blüten; im längeren oberen Bereich befinden sich nur männliche Blüten. Die kleinen Blüten sind reduziert und besitzen keine Blütenhüllblätter. Männliche Blüten enthalten zwei bis drei Staubblätter. Bei den weiblichen Blüten sind drei Fruchtblätter zu einem synkarpen Fruchtknoten verwachsen.
Sie bilden grüne oder orange Beeren. Die Samen sind kugelig bis eiförmig.
Alle Teile von Zantedeschia aethiopica sind leicht giftig und verursachen starke Schleimhautreizungen. Die Flüssigkeit, die die Zantedeschien über die Blattspitzen absondern, ist ebenfalls giftig und verursacht bei Berührung Hautausschlag und Nesselfieber.
Verbreitung
Alle Arten sind in Afrika heimisch. Am häufigsten sind sie in den Provinzen Südafrikas in der Capensis (Kapprovinz, KwaZulu-Natal und Freistaat) sowie in den Ländern Eswatini und Lesotho. Es gibt die Gattung aber auch weiter nördlich in Angola, Kenia, Malawi, Sambia, Simbabwe und Nigeria. Das natürliche Habitat der sommerblühenden Formen ist ein warmtemperates Klima mit Niederschlägen, die hauptsächlich im Winter fallen. Die anderen Arten, wie beispielsweise Zantedeschia aethiopica und Zantedeschia odorata wachsen in Gebieten mit Sommerregen und blühen im Winter. Oft gedeihen die Pflanzen in sumpfigen Gebieten.
Die Zantedeschienarten benötigen als ursprüngliche Sumpfpflanzen viel Wasser. Gerade in der Wachstumsphase sollte eine Austrocknung der Wurzeln vermieden werden. Nach der Blüte hat die Zantedeschie eine Ruhephase in der die Pflanze keinen Wasserbedarf hat. Bei Zimmercallas handelt es sich meist um die Zantedeschia aethiopica, die von Januar bis April blüht. Auch sie hat ihre Ruhephase im Anschluss an die Blütephase. Zantedeschien brauchen zum optimalen Wachstum leicht sandigen und vor allem äußerst nährstoffreichen Boden.
Systematik
Die Gattung Zantedeschia wurde 1826 durch Kurt Sprengel in Systema Vegetabilium, editio decima sexta, 3, S. 756, 765[4] aufgestellt. Der botanische Gattungsname Zantedeschia ehrt den italienischen Arzt und Botaniker Giovanni Zantedeschi (1773–1846). Typusart ist Zantedeschia aethiopica (L.) Spreng., die von Carl von Linné als Calla aethiopica erstveröffentlicht wurde. Syn. für Zantedeschia Spreng. sind Pseudohomalomena A.D.Hawkes, Richardia Kunth. Zantedeschia ist die einzige Gattung der Tribus Zantedeschieae in der Unterfamilie Aroideae in der Familie der Araceae. Die Tribus Zantedeschieae wurde durch Adolf Engler aufgestellt.
Es gibt acht Zantedeschia-Arten:
Gewöhnliche Calla oder Zimmercalla (Zantedeschia aethiopica (L.) Spreng., Syn.: Calla aethiopica L.): Sie kommt ursprünglich im südlichen Afrika vor.
Gefleckte Calla (Zantedeschia albomaculata (Hook.) Baill.): Sie kommt von Tansania, Demokratischen Republik Kongo, Angola, Malawi, Sambia, Simbabwe bis Südafrika vor.
Mit zwei Unterarten:
Zantedeschia albomaculata (Hook.) Baill. subsp. albomaculata: Sie kommt von Nigeria bis Tansania und im südlichen Afrika vor.
Zantedeschia albomaculata subsp. macrocarpa (englisch) Letty: Sie kommt nur im südlichen Afrika vor.
Goldene Calla oder Elliott-Calla (Zantedeschia elliottiana (W.Watson) Engl.): Sie ist nur aus Kultur bekannt, kommt aber möglicherweise im östlichen Transvaal (Provinz Mpumalanga) wild vor.
Zantedeschia jucunda Letty: Sie kommt nur im nördlichen Südafrika vor.
Zantedeschia odorata P.L.Perry: Sie kommt nur in der Kapprovinz vor.
Zantedeschia pentlandii (W.Watson) Wittm.: Sie kommt nur im östlichen Transvaal in der Provinz Mpumalanga vor.
Rosafarbene Calla oder Rehmann-Calla (Zantedeschia rehmannii Engl.): Sie kommt nur von Mosambik bis Südafrika vor.
Zantedeschia valida (Letty) Y.Singh: Sie kommt nur in KwaZulu-Natal vor.
Nutzung
Viele Sorten werden als Zierpflanzen verwendet.
Über eine weitere Nutzung durch den Menschen ist wenig bekannt. Die Blätter von Zantedeschia aethiopica werden gegart gegessen (alle Pflanzenteile sind roh giftig).
In der Trauerfloristik wird sie als Zeichen der Unsterblichkeit genutzt. In weiß galt sie lange als „Totenblume“. In Gestecken stehen die verschiedenen Sorten heute für Moderne und Eleganz und werden als Symbol für Schönheit und Anerkennung verschenkt. Durch ihre besondere Blütenform, die an einen Kelch erinnert, wurde sie von den Griechen und Römern mit Feierlichkeiten und Freude in Verbindung gebracht. Die Zantedeschie gilt daher in vielen Gegenden noch heute als Glücksbringer.
(Wikipedia)
Bellis perennis is a common European species of daisy, often considered the archetypal species of that name. Many related plants also share the name daisy, so to distinguish this species from other daisies it is sometimes qualified as common daisy, lawn daisy or English daisy. Historically, it has also been commonly known as bruisewort and occasionally woundwort. Bellis perennis is native to western, central and northern Europe, but widely naturalised in most temperate regions including the Americas and Australasia. It is a perennial herbaceous plant with short creeping rhizomes and rosettes of small rounded or spoon-shaped leaves that are from 2–5 cm long and grow flat to the ground. The flowerheads are composite, in the form of a pseudanthium, consisting of many sessile flowers about 2–3 cm in diameter, with white ray florets (often tipped red) and yellow disc florets. 5951
Euphorbia is a very large and diverse genus of flowering plants, commonly called spurge, in the family Euphorbiaceae. "Euphorbia" is sometimes used in ordinary English to collectively refer to all members of Euphorbiaceae (in deference to the type genus), not just to members of the genus.
Euphorbias range from tiny annual plants to large and long-lived trees. with perhaps the tallest being Euphorbia ampliphylla at 30 m (98 ft) or more. The genus has roughly 2,000 members, making it one of the largest genera of flowering plants. It also has one of the largest ranges of chromosome counts, along with Rumex and Senecio. Euphorbia antiquorum is the type species for the genus Euphorbia. It was first described by Carl Linnaeus in 1753 in Species Plantarum.
Some euphorbias are widely available commercially, such as poinsettias at Christmas. Some are commonly cultivated as ornamentals, or collected and highly valued for the aesthetic appearance of their unique floral structures, such as the crown of thorns plant (Euphorbia milii). Succulent euphorbias from the deserts of Southern Africa and Madagascar have evolved physical characteristics and forms similar to cacti of North and South America, so they are often incorrectly referred to as cacti. Some are used as ornamentals in landscaping, because of beautiful or striking overall forms, and drought and heat tolerance.
Euphorbia all share the feature of having a poisonous, latex-like sap and unique floral structures When viewed as a whole, the head of flowers looks like a single flower (a pseudanthium). It has a unique kind of pseudanthium, called a cyathium, where each flower in the head is reduced to its barest essential part needed for sexual reproduction. The individual flowers are either male or female, with the male flowers reduced to only the stamen, and the females to the pistil. These flowers have no sepals, petals, or other parts that are typical of flowers in other kinds of plants. Structures supporting the flower head and other structures underneath have evolved to attract pollinators with nectar, and with shapes and colors that function in a way petals and other flower parts do in other flowers. It is the only genus of plants that has all three kinds of photosynthesis, CAM, C3 and C4.[7]
Etymology
The common name "spurge" derives from the Middle English/Old French espurge ("to purge"), due to the use of the plant's sap as a purgative. The botanical name Euphorbia derives from Euphorbos, the Greek physician of King Juba II of Numidia and Mauretania (52–50 BC – 23 AD), who married the daughter of Anthony and Cleopatra. Juba was a prolific writer on various subjects, including natural history. Euphorbos wrote that one of the cactus-like euphorbias (now called Euphorbia obtusifolia ssp. regis-jubae) was used as a powerful laxative. In 12 BC, Juba named this plant after his physician Euphorbos, as Augustus Caesar had dedicated a statue to the brother of Euphorbos, Antonius Musa, who was the personal physician of Augustus. In 1753, botanist and taxonomist Carl Linnaeus assigned the name Euphorbia to the entire genus in the physician's honor.
Description
The plants are annual, biennial or perennial herbs, woody shrubs, or trees with a caustic, poisonous milky latex. The roots are fine or thick and fleshy or tuberous. Many species are more or less succulent, thorny, or unarmed. The main stem and mostly also the side arms of the succulent species are thick and fleshy, and often winged, 15–91 cm (6–36 in) tall. The succulent trees and large shrubs are mostly confined to southern and eastern Africa, but Euphorbia neutra is native to the Caatinga of Brazil and Euphorbia royleana is found in the Himalayan foothills.[14] The deciduous[citation needed] leaves may be opposite, alternate, or in whorls. In succulent species, the leaves are mostly small and short-lived. The stipules are mostly small, partly transformed into spines or glands, or missing.
Inflorescence and fruit
Like all members of the family Euphorbiaceae, spurges have unisexual flowers.
In Euphorbia, flowers occur in a head, called the cyathium (plural cyathia). Each male or female flower in the cyathium head has only its essential sexual part, in males the stamen, and in females the pistil. The flowers do not have sepals, petals, or nectar to attract pollinators, although other nonflower parts of the plant have an appearance and nectar glands with similar roles. Euphorbias are the only plants known to have this kind of flower head.
Nectar glands and nectar that attract pollinators are held in the involucre, a cup-like part below and supporting the cyathium head. The "involucre" in the genus Euphorbia is not to be confused with the "involucre" in family Asteraceae members, which is a collection of bracts called phyllaries, which surround and encase the unopened flower head, then support the receptacle under it after the flower head opens.
The involucre is above and supported by bract-like modified leaf structures (usually in pairs)[citation needed] called cyathophylls', or cyathial leaves. The cyathophyll often has a superficial appearance of being petals of a flower.
Euphorbia flowers are tiny, and the variation attracting different pollinators, with different forms and colors occurs, in the cyathium, involucre, cyathophyll, or additional parts such as glands that attached to these.
The collection of many flowers may be shaped and arranged to appear collectively as a single individual flower, sometimes called a pseudanthium in the Asteraceae, and also in Euphorbia.
The majority of species are monoecious (bearing male and female flowers on the same plant), although some are dioecious with male and female flowers occurring on different plants. It is not unusual for the central cyathia of a cyme to be purely male, and for lateral cyathia to carry both sexes. Sometimes, young plants or those growing under unfavorable conditions are male only, and only produce female flowers in the cyathia with maturity or as growing conditions improve.
The female flowers reduced to a single pistil usually split into three parts, often with two stigmas at each tip.[citation needed] Male flowers often have anthers in twos.[citation needed] Nectar glands usually occur in fives, may be as few as one,[16] and may be fused into a "U" shape. The cyathophylls often occur in twos, are leaf-like, and may be showy and brightly coloured and attractive to pollinators, or be reduced to barely visible tiny scales.
The fruits are three- or rarely two-compartment capsules, sometimes fleshy, but almost always ripening to a woody container that then splits open, sometimes explosively. The seeds are four-angled, oval, or spherical, and some species have a caruncle.
Xerophytes and succulents
In the genus Euphorbia, succulence in the species has often evolved divergently and to differing degrees. Sometimes, it is difficult to decide, and is a question of interpretation, whether or not a species is really succulent or "only" xerophytic. In some cases, especially with geophytes, plants closely related to the succulents are normal herbs. About 850 species are succulent in the strictest sense. If one includes slightly succulent and xerophytic species, this figure rises to about 1000, representing about 45% of all Euphorbia species.
Irritants
The milky sap of spurges (called "latex") evolved as a deterrent to herbivores. It is white, and transparent when dry, except in E. abdelkuri, where it is yellow. The pressurized sap seeps from the slightest wound and congeals after a few minutes in air. The skin-irritating and caustic effects are largely caused by varying amounts of diterpenes. Triterpenes such as betulin and corresponding esters are other major components of the latex. In contact with mucous membranes (eyes, nose, mouth), the latex can produce extremely painful inflammation. The sap has also been known to cause mild to extreme Keratouveitis, which affects vision. Therefore, spurges should be handled with caution and kept away from children and pets. Wearing eye protection while working in close contact with Euphorbia is advised.[18] Latex on skin should be washed off immediately and thoroughly. Congealed latex is insoluble in water, but can be removed with an emulsifier such as milk or soap. A physician should be consulted if inflammation occurs, as severe eye damage including permanent blindness may result from exposure to the sap.
The poisonous qualities were well known: in the Ethiopian Kebra Nagast, the serpent king Arwe is killed with juice from the Euphorbia.
Uses
Several spurges are grown as garden plants, among them poinsettia (E. pulcherrima) and the succulent E. trigona. E. pekinensis (Chinese: 大戟; pinyin: dàjǐ) is used in traditional Chinese medicine, where it is regarded as one of the 50 fundamental herbs. Several Euphorbia species are used as food plants by the larvae of some Lepidoptera (butterflies and moths), like the spurge hawkmoths (Hyles euphorbiae and Hyles tithymali), as well as the giant leopard moth.
Ingenol mebutate, a drug used to treat actinic keratosis, is a diterpenoid found in Euphorbia peplus.
Euphorbias are often used as hedging plants in many parts of Africa.
Among laypeople, Euphorbia species are among the plant taxa most commonly confused with cacti, especially the stem succulents. Euphorbias secrete a sticky, milky-white fluid with latex, but cacti do not. Individual flowers of euphorbias are usually tiny and nondescript (although structures around the individual flowers may not be), without petals and sepals, unlike cacti, which often have fantastically showy flowers. Euphorbias from desert habitats with growth forms similar to cacti have thorns, which are different from the spines of cacti.
Systematics and taxonomy
The present taxon "Euphorbia" corresponds to its own former subtribe, the Euphorbiinae.[citation needed] It has over 2000 species. Morphological description using the presence of a cyathium (see section above) is consistent with nuclear and chloroplast DNA sequence data in testing of about 10% of its members. This testing supports inclusion of formerly other genera as being best placed in this single genus, including Chamaesyce, Monadenium, Pedilanthus, and poinsettia (E. pulcherrima).
Genetic tests have shown that similar flower head structures or forms within the genus, might not mean close ancestry within the genus. The genetic data show that within the genus, convergent evolution of inflorescence structures may be from ancestral subunits that are not related. So using morphology within the genus becomes problematic for further subgeneric grouping. As stated on the Euphorbia Planetary Biodiversity Inventory project webpage:
Previous morphologically based delimitations of subgenera or sections within the genus should not be taken at face value. The genus is in fact rife with striking examples of morphological convergence in cyathial and vegetative features, which justifies a global approach to studying the genus to obtain a proper phylogenetic understanding of the whole group.... The bottom line is that a number of clades have been placed inside or outside of Euphorbia at different times... few of the subgeneric circumscriptions hold up under DNA sequence analysis.
According to a 2002 publication on studies of DNA sequence data, most of the smaller "satellite genera" around the huge genus Euphorbia nest deep within the latter. Consequently, these taxa, namely the never generally accepted genus Chamaesyce, as well as the smaller genera Cubanthus, Elaeophorbia, Endadenium, Monadenium, Synadenium, and Pedilanthus were transferred to Euphorbia. The entire subtribe Euphorbiinae now consists solely of the genus Euphorbia.
Selected species
See List of Euphorbia species for complete list.
Euphorbia albomarginata – rattlesnake weed, white-margined sandmat
Euphorbia amygdaloides – wood spurge
Euphorbia antisyphilitica – candelilla
Euphorbia balsamifera – sweet tabaiba (Canary Islands)
Euphorbia bulbispina
Euphorbia caducifolia – leafless milk hedge
Euphorbia canariensis – Canary Island spurge, Hercules club (Canary Islands)
Euphorbia candelabrum – candelabra tree of East Africa
Euphorbia caput-medusae – Medusa's head (South Africa)
Euphorbia ceratocarpa – (Sicily and southern Italy)
Euphorbia characias – Mediterranean spurge
Euphorbia coerulescens - blue euphorbia
Euphorbia cotinifolia – copper tree
Euphorbia cyathophora – fire-on-the-mountain
Euphorbia cyparissias – Cypress spurge
Euphorbia decidua
Euphorbia dendroides – tree spurge
Euphorbia epithymoides – cushion spurge
Euphorbia esula – leafy spurge
Euphorbia franckiana
Euphorbia fulgens – scarlet plume
Euphorbia grantii – African milk bush
Euphorbia gregersenii – Gregersen's spurge
Euphorbia griffithii – Griffith's spurge
Euphorbia helioscopia – sun spurge
Euphorbia heterophylla – painted euphorbia, desert poinsettia, fireplant, paint leaf, kaliko
Euphorbia hirta – asthma-plant
Euphorbia hispida
Euphorbia horrida – African milk barrel
Euphorbia ingens – candelabra tree
Euphorbia labatii
Euphorbia lactea – mottled spurge, frilled fan, elkhorn
Euphorbia lathyris – caper spurge, paper spurge, gopher spurge, gopher plant, mole plant
Euphorbia leuconeura – Madagascar jewel
Euphorbia maculata – spotted spurge, prostrate spurge
Euphorbia marginata – snow on the mountain
Euphorbia mammillaris
Euphorbia maritae
Euphorbia milii – crown-of-thorns, Christ plant
Euphorbia misera – cliff spurge, Baja California, Southern California
Euphorbia myrsinites – myrtle spurge, creeping spurge, donkey tail
Euphorbia nivulia – leafy milk hedge
Euphorbia obesa
Euphorbia paralias – sea spurge
Euphorbia pekinensis – Peking spurge
Euphorbia peplis – purple spurge
Euphorbia peplus – petty spurge
Euphorbia polychroma – bonfire
Euphorbia psammogeton – sand spurge
Euphorbia pulcherrima – poinsettia, Mexican flame leaf, Christmas star, winter rose, noche buena, lalupatae, pascua, Atatürk çiçeği (Turkish)
Euphorbia purpurea – Darlington's glade spurge, glade spurge, or purple spurge
Euphorbia resinifera – resin spurge
Euphorbia rigida – gopher spurge, upright myrtle spurge
Euphorbia serrata – serrated spurge, sawtooth spurge
Euphorbia tirucalli – Indian tree spurge, milk bush, pencil tree, firestick
Euphorbia tithymaloides – devil's backbone, redbird cactus, cimora misha (Peru)
Euphorbia trigona – African milk tree, cathedral cactus, Abyssinian euphorbia
Euphorbia tuberosa
Euphorbia virosa – gifboom or poison tree
Hybrids
Euphorbia has been extensively hybridised for garden use, with many cultivars available commercially. Moreover, some hybrid plants have been found growing in the wild, for instance E. × martini Rouy, a cross of E. amygdaloides × E. characias subsp. characias, found in southern France.
Subgenera
The genus Euphorbia is one of the largest and most complex genera of flowering plants, and several botanists have made unsuccessful attempts to subdivide the genus into numerous smaller genera. According to the recent phylogenetic studies, Euphorbia can be divided into four subgenera, each containing several sections and groups. Of these, subgenus Esula is the most basal. The subgenera Chamaesyce and Euphorbia are probably sister taxa, but very closely related to subgenus Rhizanthium. Extensive xeromorph adaptations in all probability evolved several times; it is not known if the common ancestor of the cactus-like Rhizanthium and Euphorbia lineages had been xeromorphic—in which case a more normal morphology would have re-evolved namely in Chamaesyce—or whether extensive xeromorphism is entirely polyphyletic even to the level of the subgenera.
Bellis perennis the daisy, is a European species of the family Asteraceae, often considered the archetypal species of the name daisy. To distinguish this species from other plants known as daisies, it is sometimes qualified as common daisy, lawn daisy or English daisy.
Bellis perennis is a perennial herbaceous plant growing to 20 centimetres (8 inches) in height. It has short creeping rhizomes and rosettes of small rounded or spoon-shaped leaves that are from 2 to 5 cm (3⁄4–2 in) long and grow flat to the ground. The species habitually colonises lawns, and is difficult to eradicate by mowing, hence the term 'lawn daisy'. It blooms from March to September and exhibits the phenomenon of heliotropism, in which the flowers follow the position of the sun in the sky.
The flowerheads are composite, about 2 to 3 cm (3⁄4–1+1⁄4 in) in diameter, in the form of a pseudanthium, consisting of many sessile flowers with white ray florets (often tipped red) and yellow disc florets. Each inflorescence is borne on a single leafless stem 2 to 10 cm (3⁄4–4 in), rarely 15 cm (6 in) tall. The capitulum, or disc of florets, is surrounded by two rows of green bracts known as "phyllaries". The achenes are without pappus.
Bellis may come from bellus, Latin for "pretty", and perennis is Latin for "everlasting".
The name "daisy", possibly originating with this plant, is considered a corruption of "day's eye", because the whole head closes at night and opens in the morning. Geoffrey Chaucer called it "eye of the day". In Medieval times, Bellis perennis or the English Daisy was commonly known as "Mary's Rose". It is also known as bone flower.
Historically, the plant has also been widely known as bruisewort, and occasionally woundwort (although this name is now more closely associated with the genus Stachys).
Bellis perennis is native to western, central and northern Europe, including remote islands such as the Faroe Islands, but has become widely naturalised in most temperate regions, including the Americas and Australasia. It prefers field-like habitats.
The species generally blooms from early to midsummer, although when grown under ideal conditions, it has a very long flowering season and will even produce a few flowers in the middle of mild winters.
It can generally be grown where minimum temperatures are above −35 °C (−30 °F), in full sun to partial shade conditions, and requires little or no maintenance. It has no known serious insect or disease problems and can generally be grown in most well-drained soils. The plant may be propagated either by seed after the last frost, or by division after flowering.
Though not native to the United States, the species is still considered a valuable ground cover in certain garden settings (e.g., as part of English or cottage inspired gardens, as well as spring meadows where low growth and some color is desired in parallel with minimal care and maintenance while helping to crowd out noxious weeds once established and naturalised).
Numerous single- and double-flowered varieties are in cultivation, producing flat or spherical blooms in a range of sizes (1 to 6 cm or 3⁄8 to 2+3⁄8 in) and colours (red, pink and white). They are generally grown from seed as biennial bedding plants. They can also be purchased as plugs in Spring.
It has been reported to be mostly self-fertilizing, but some plants may be self-sterile.
Bellis perennis may be used as a potherb. Young leaves can be eaten raw in salads, or cooked, though the leaves become increasingly astringent with age. Flower buds and petals can be eaten raw in sandwiches, soups and salads. It is also used as a tea and as a vitamin supplement.
B. perennis has astringent properties and has been used in herbal medicine.
Daisies have traditionally been used for making daisy chains in children's games.
Daisy is used as a feminine name, and sometimes as a nickname for people named Margaret, after the French name for the oxeye daisy, marguerite.
The daisy is the national flower of the Netherlands.
ENGLISH
The family Asteraceae or Compositae (known as the aster, daisy, or sunflower family) is the largest family of flowering plants, in terms of number of species.
The name 'Asteraceae' is derived from the type genus Aster, while 'Compositae', an older but still valid name, means composite and refers to the characteristic inflorescence, a special type of pseudanthium found in only a few other angiosperm families. The study of this family is known as synantherology.
The most evident characteristic of Asteraceae is perhaps their inflorescence: a specialised capitulum, technically called a calathid or calathidium, but generally referred to as flower head or, alternatively, simply capitulum. The capitulum is a contracted raceme composed of numerous individual sessile flowers, called the florets, all sharing the same receptacle.
The capitulum of the Asteraceae has evolved many characteristics that make it look superficially like a big single flower. This kind of flower-like inflorescences are quite widespread amongst plants and have been given the name of pseudanthia.
Many bracts form an involucre under the basis of the capitulum; these are called "phyllaries", or "involucral bracts". They may simulate the sepals of the pseudanthium. These are mostly herbaceous but can also be brightly coloured (e.g. Helichrysum) or have a scarious texture. The bracts can be free or fused, and arranged in one to many rows, overlapping like the tiles of a roof (imbricate) or not (this variation is important in identification of tribes and genera).
Each floret may itself be subtended by a bract, called a "palea" or "receptacular bract". These bracts as a group are often called "chaff". The presence or absence of these bracts, their distribution on the receptacle, and their size and shape are all important diagnostic characteristics for genera and tribes.
The florets have five petals fused at the base to form a corolla tube and they may be either actinomorphic or zygomorphic. Disc florets are usually actinomorphic, with five petal lips on the rim of the corolla tube. The petal lips may be either very short, or long, in which case they form deeply lobed petals. The latter is the only kind of floret in the Carduoideae, while the first kind is more widespread. Ray florets are always highly zygomorphic and are characterised by the presence of a ligule, a strap-shaped structure on the edge of the corolla tube consisting of fused petals. In the Asteroideae and other minor subfamilies these are usually borne only on florets at the circumference of the capitulum and have a 3+2 scheme – above the fused corolla tube, three very long fused petals form the ligule, with the other two petals being inconspicuously small. The Cichorioidea has only ray florets, with a 5+0 scheme – all five petals form the ligule. A 4+1 scheme is found in the Barnadesioideae. The tip of the ligule is often divided into teeth, each one representing a petal. Some marginal florets may have no petals at all (filiform floret).
The calyx of the florets may be absent, but when present it is always modified into a pappus of two or more teeth, scales or bristles and this is often involved in the dispersion of the seeds. As with the bracts, the nature of the pappus is an important diagnostic feature.
There are usually five stamens. The filaments are fused to the corolla, while the anthers are generally connate (syngenesious anthers), thus forming a sort of tube around the style (theca). They commonly have basal and/or apical appendages. Pollen is released inside the tube and is collected around the growing style, expelled with a sort of pump mechanism (nüdelspritze) or a brush.
The pistil is made of two connate carpels. The style has two lobes; stigmatic tissue may be located in the interior surface or form two lateral lines. The ovary is inferior and has only one ovule, with basal placentation.
More info: en.wikipedia.org/wiki/Asteraceae
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CASTELLANO
Las Asteráceas (Asteraceae Bercht. & J. Presl), también denominadas Compuestas (Compositae Giseke), reunen mas de 23.000 especies (Jeffrey, 2007; Stevens, 2001) por lo que son la familia de Angiospermas con mayor riqueza y diversidad biológica. La familia está caracterizada por presentar las flores dispuestas en una inflorescencia compuesta denominada capítulo la cual se halla rodeada de una o más filas de brácteas (involucro). El nombre "Asteraceae" deriva del género tipo de la familia Aster, término que -a su vez- proviene del griego ἀστήρ que significa "estrella" y hace alusión a la forma de la inflorescencia. Por otro lado, el nombre "compuestas", más antiguo pero válido, hace referencia al tipo particular de inflorescencia compuesta que caracteriza a la familia y que solo se halla en muy pocas familias de Angiospermas. Las Compuestas presentan una considerable importancia ecológica y económica. Los miembros de esta familia se distribuyen desde las regiones polares hasta los trópicos, conquistando todos los hábitats disponibles, desde los desiertos secos hasta los pantanos y desde las selvas hasta los picos montañosos. En muchas regiones del mundo las Compuestas llegan a integrar hasta el 10% de la flora vernácula.
Las flores son pequeñas, hermafroditas o, en ocasiones, funcionalmente unisexuales o estériles (en este último caso se denominan neutras). Por su simetría pueden ser tanto actinomorfas como zigomorfas, es decir que pueden presentar simetría radial o bilateral, respectivamente. Por el número de piezas que componen cada ciclo, las flores de las compuestas son pentámeras. El cáliz es nulo o bien los sépalos están profundamente modificados como pelos simples, plumosos o setiformes, como escamas o como una pequeña corona membranácea. La corola es gamopétala, los pétalos pueden unirse entre si formando un tubo con 4 ó 5 lóbulos (la flor se denomina tubulosa o flósculo), o dos grupos de pétalos unidos (en el caso de las flores bilabiadas), o pueden presentar un tubo corto y el limbo prolongado lateralmente en una lígula con 3 o 5 dientes (flores liguladas).[a] El androceo presenta 5 estambres, los cuales alternan con los lóbulos de la corola. Los filamentos se hallan insertos en ella. Las anteras se unen entre sí fomando un tubo alrededor del estilo. El gineceo es de ovario ínfero, bicarpelar, con el ovario unilocular. Presentan un solo óvulo anátropo. El pistilo presenta un estilo que está usualmente rodeado en su base por un nectario. Cada estilo se halla dividido distalmente en dos ramas (ramas estilares) que presentan papilas estigmáticas en su cara adaxial dispuestas en dos líneas separadas o bien, en una sola banda continua.
El capítulo es un tipo de inflorescencia racimosa o abierta en la que el pedúnculo se ensancha en la extremidad formando un disco algo grueso, llamado receptáculo común o clinanto. Este receptáculo común se halla rodeado por una o más series de brácteas. Sobre este órgano se disponen las flores sésiles acompañadas o no por sus correspondientes brácteas. Es la inflorescencia típica de las Asteraceae (Compuestas) y de algunos otros géneros como Scabiosa. Inflorescencias en capítulo, rodeado por un involucro de brácteas; en ocasiones existen en el receptáculo escamas de naturaleza bracteal. Los capítulos a su vez pueden agruparse en inflorescencias compuestas diversas.
Más info: es.wikipedia.org/wiki/Asteraceae
Bellis perennis the daisy, is a European species of the family Asteraceae, often considered the archetypal species of the name daisy. To distinguish this species from other plants known as daisies, it is sometimes qualified or known as common daisy, lawn daisy or English daisy.
Bellis perennis is a perennial herbaceous plant growing to 20 centimetres (8 inches) in height. It has short creeping rhizomes and rosettes of small rounded or spoon-shaped leaves that are from 2 to 5 cm (3⁄4–2 in) long and grow flat to the ground. The species habitually colonises lawns, and is difficult to eradicate by mowing, hence the term 'lawn daisy'. It blooms from March to September and exhibits the phenomenon of heliotropism, in which the flowers follow the position of the sun in the sky.
The flowerheads are composite, about 2 to 3 cm (3⁄4–1+1⁄4 in) in diameter, in the form of a pseudanthium, consisting of many sessile flowers with white ray florets (often tipped red) and yellow disc florets. Each inflorescence is borne on a single leafless stem 2 to 10 cm (3⁄4–4 in), rarely 15 cm (6 in) tall. The capitulum, or disc of florets, is surrounded by two rows of green bracts known as "phyllaries". The achenes are without pappus.
*** Arctic Islands 2017 Expedition (polarforskningsportalen.se/en/arctic/expeditions/arktiska...) ***
The flowers here are actually pseudanthia (i.e. false flowers) called cyathia. The yellow mouth-like structures attached to the side of each cyathium are nectar glands.
P1030470
Common sunflower (Helianthus annuus)
Wild Helianthus annuus is a widely branched annual plant with many flower heads. The domestic sunflower, however, often possesses only a single large inflorescence (flower head) atop an unbranched stem.
The term Helianthus annuus is derived from the Greek Helios, meaning sun, and anthos, meaning flower, and from Latin annuus, meaning annual.
What is often called the "flower" of the sunflower is actually a "flower head" (pseudanthium) of numerous small individual five-petaled flowers ("florets"). The outer flowers, which resemble petals, are called ray flowers. Each "petal" consists of a ligule composed of fused petals of an asymmetrical ray flower. They are sexually sterile and may be yellow, red, orange, or other colors. The flowers in the center of the head are called disk flowers. These mature into fruit (sunflower "seeds"). The disk flowers are arranged spirally.
A common misconception is that flowering sunflower heads track the Sun across the sky. Although immature flower buds exhibit this behaviour, the mature flowering heads point in a fixed (and typically easterly) direction throughout the day.
When growth of the flower stalk stops and the flower is mature, the heliotropism also stops and the flower faces east from that moment onward. This eastward orientation allows rapid warming in the morning and, as a result, an increase in pollinator visits.
Common sunflower (Helianthus annuus)
Wild Helianthus annuus is a widely branched annual plant with many flower heads. The domestic sunflower, however, often possesses only a single large inflorescence (flower head) atop an unbranched stem.
The term Helianthus annuus is derived from the Greek Helios, meaning sun, and anthos, meaning flower, and from Latin annuus, meaning annual.
What is often called the "flower" of the sunflower is actually a "flower head" (pseudanthium) of numerous small individual five-petaled flowers ("florets"). The outer flowers, which resemble petals, are called ray flowers. Each "petal" consists of a ligule composed of fused petals of an asymmetrical ray flower. They are sexually sterile and may be yellow, red, orange, or other colors. The flowers in the center of the head are called disk flowers. These mature into fruit (sunflower "seeds"). The disk flowers are arranged spirally.
A common misconception is that flowering sunflower heads track the Sun across the sky. Although immature flower buds exhibit this behaviour, the mature flowering heads point in a fixed (and typically easterly) direction throughout the day.
When growth of the flower stalk stops and the flower is mature, the heliotropism also stops and the flower faces east from that moment onward. This eastward orientation allows rapid warming in the morning and, as a result, an increase in pollinator visits.
Centaurea is a genus of over 700 species of herbaceous thistle-like flowering plants in the family Asteraceae. Members of the genus are found only north of the equator, mostly in the Eastern Hemisphere; the Middle East and surrounding regions are particularly species-rich. In the western United States, yellow starthistles are an invasive species. Around the year 1850, seeds from the plant had arrived to the state of California. It is believed that those seeds came from South America.
Common names
Common names for this genus are centaury, centory, starthistles, knapweeds, centaureas and the more ambiguous "bluets"; a vernacular name used for these plants in parts of England is "loggerheads" (common knapweed). The Plectocephalus group – possibly a distinct genus – is known as basketflowers. "Cornflower" is used for a few species, but that term more often specifically means either C. cyanus (the annual cornflower) or Centaurea montana (the perennial cornflower). The common name "centaury" is sometimes used, although this also refers to the unrelated plant genus Centaurium.
The name is said to be in reference to Chiron, the centaur of Greek mythology who discovered medicinal uses of a plant eventually called "centaury".
Description
Knapweeds are robust weedy plants. Their leaves, spiny in some species, are usually deeply divided into elongated lobes at least in the plants' lower part, becoming entire towards the top. The "flowers" (actually pseudanthium inflorescences) are diverse in colour, ranging from intense blues, reds and yellows to any mixture of these and lighter shades towards white. Often, the disk flowers are much darker or lighter than the ray flowers, which also differ in morphology and are sterile. Each pseudanthium sits atop a cup- or basket-like cluster of scaly bracts, hence the name "basketflowers". Many species, in particular those inhabiting more arid regions, have a long and strong taproot.
Certain knapweeds have a tendency to dominate large stretches of landscape together with a few other plants, typically one or two grasses and as many other large herbaceous plants. The common knapweed (C. nigra) for example is plentiful in the mesotrophic grasslands of England and nearby regions. It is most prominently found in pastures or meadows dominated by cock's-foot (Dactylis glomerata) as well as either of crested dog's-tail (Cynosurus cristatus) and false oat-grass (Arrhenatherum elatius). It is also often found in mesotrophic grassland on rendzinas and similar calcareous soils in association with glaucous sedge (Carex flacca), sheep's fescue (Festuca ovina), and either tor-grass (Brachypodium pinnatum) and rough hawkbit (Leontodon hispidus), or upright brome (Bromus erectus). In these grasslands, greater knapweed (C. scabiosa) is found much more rarely by comparison, often in association with red fescue (Festuca rubra) in addition to cock's-foot and false oat-grass.
Due to their habit of dominating ecosystems under good conditions, many Centaurea species can become invasive weeds in regions where they are not native. In parts of North America, diffuse knapweed (C. diffusa), spotted knapweed (C. maculosa) and yellow starthistle (C. solstitialis) cause severe problems in agriculture due to their uncontrolled spread. The seeds are typically transported by human traffic, in particular the tires of all-terrain vehicles. The two knapweeds are harmful mainly because they are strongly allelopathic, producing powerful toxins in their roots that stunt the growth of plants around them not adapted to this. Yellow starthistle, meanwhile, is inedible to most livestock due to its spines and apparently outright poisonous to horses and other equines. However, efficient methods of biological control by insect pests of these weeds have been developed; the knapweeds can also exploited to their detriment by targeted grazing. Controlled burning may also be used, though the timing is important to avoid the plants having seeded already, and neither allowing sufficient time for them to regrow from the rootstock.
Yet other species of Centaurea – mostly ones that occur between Italy and the Caucasus – are endemics of a single island or valley, and some of these are endangered. The Akamas Centaurea (Centaurea akamantis) of Cyprus is almost extinct, while the western Caucasus endemics C. leptophylla and C. straminicephala are at least very rare and C. hedgei and C. pecho from the same region are certainly not abundant either. The last four species would be adversely affected by the proposed Yusufeli Dam, which might actually destroy enough habitat to push the two rarer ones over the brink of extinction.
Centaurea are copious nectar producers, especially on high-lime soils. The high nectar yield of the genus makes it very attractive to insects such as butterflies – including the endangered Karner blue (Plebejus melissa samuelis) which visits introduced spotted knapweed – and day-flying moths – typically Zygaenidae, such as Zygaena loti or the six-spot burnet (Z. filipendulae). The larvae of some other Lepidoptera species use Centaurea species as food plants; see List of Lepidoptera that feed on Centaurea. Several of these are used in biological control of invasive knapweeds and starthistles.
Larvae of several true weevils (Curculionidae) of the subfamily Lixinae also feed on Centaurea. Some genera – such as Larinus whose larval food is flowerheads – have many species especially adapted to particular knapweeds or starthistle and are used in biological control too. These include the yellow starthistle flower weevil (L. curtus) for yellow starthistle, lesser knapweed flower weevil (L. minutus) for diffuse knapweed and blunt knapweed flower weevil (L. obtusus) for spotted knapweed. Broad-nosed seedhead weevil (Bangasternus fausti) larvae eat diffuse, spotted and squarrose knapweed (C. virgata ssp. squarrosa), while those of the yellow starthistle bud weevil (B. orientalis) do not seem to live on anything other than yellow starthistle and occasionally purple starthistle (C. calcitrapa). But perhaps most efficient in destroying developing yellow starthistle seedheads is the larva of the yellow starthistle hairy weevil (Eustenopus villosus). Knapweed root weevil (Cyphocleonus achates) larvae bore into the roots of spotted and to a lesser extentely diffuse knapweed, sometimes killing off the entire plant.
Also used in biological control are Tephritidae (peacock flies) whose larvae feed on Centaurea. Knapweed peacock fly (Chaetorellia acrolophi) larvae eat spotted knapweed and some other species. The yellow starthistle peacock fly (C. australis) has an initial generation each year which often uses cornflower (C. cyanus) as larval food; later generations switch to yellow starthistle. The flies are generally considered less efficient in destroying the growing seedheads than the weevils, but may be superior under certain conditions; employing flies and weevils in combination is expensive and does not noticeably increase their effect.
Use by humans
Although the genus may be considered by a quite significant number of relatively informed individuals to have an overall negative impact on human interests, particularly agricultural interests, the situation is not straightforward enough to simply declare the genus, or, at least, its most aggressively-spreading species, altogether negative. For instance, due to their moderate to high nectar production, which can occur over a comparatively long duration, many species of Centaurea are popular food sources for insects that may otherwise attack certain crops.[citation needed] It may be advisable for some types of farms to allow certain species in this genus, such as cornflower (C. cyanus) in a European setting, to grow adjacent to fields. Although they support and attract many types of beneficial life (not just beetles), these areas are known as beetle banks. When they are present, some pests may be drawn away from crops to them and predatory insects and arachnids that feed upon pest insects will be better-supported by these more naturalized areas. They additionally have the beneficial aspect of supporting pollinators, unlike many field crops such as maize. Moreover, being untreated with pesticides and providing more diversity, plants growing in more wild areas adjacent to farms produce more insects that attract and support birds which can also feed on pests that would harm crops. Insect production is especially high for beetle banks that have enough plants that serve in the role of host plant for immature insects, rather than just in the roles of adult food and/or shelter provision.
Some plants which are considered invasive or problematic in certain areas can have beneficial qualities that outweigh their negative qualities from a human and/or human agricultural point of view, although this sometimes requires some human management – particularly if adequate biological control has not been established for the more aggressive species. An example is wild parsnip, Pastinaca sativa, which produces florets that feed predatory (and other beneficial) insects as well as large tubular stems that provide winter shelter for native bees, wasps, and other organisms that can be beneficial for agriculture. The plant is considered invasive in some areas of the United States and is also often considered undesirable due to its ability to cause contact skin irritation. However, it also serves as a host plant for the black swallowtail butterfly, helps to bring nutrients up from soils with its deep taproot, and possesses evergreen foliage even in climate zones such as US zone 6. This foliage increases soil warmth and moisture which can be beneficial for certain types of life. Perhaps the most dramatic example of a generally disliked plant's beneficial qualities being usually overlooked is the often-despised ragwort, Jacobaea vulgaris, which topped the list by a large amount for nectar production in a UK study, with a production per floral unit of (2921 ± 448μg). This very high nectar production, coupled with its early blooming period, makes the plant helpful for the establishment of bee colonies in spring — a period that is often not well-served by commercial flower meadow seed mixes. It also has the situationally-beneficial quality of being a spring ephemeral, as well as an annual that lacks difficult-to-combat roots. Plants that provide necessary structural supports for invertebrate and small vertebrate predators can help to keep overall pest populations low.
The abundant nectar produced by C. solstitialis flowers attracts many pollinators. This is another reason for the success of the (situationally) highly invasive species. Due to genetic differences related to evolutionary adaption, not all members of Centaurea produce the same amount of nectar. Growing conditions, such as climate and soil, can have a very strong impact, even if the plants grow and flower. For instance, cornflower plants, Centaurea cyanus, produced 33% less seasonal nectar than Centaurea nigra in a UK study. C. nigra also ranked higher than ragwort in another UK study, although ragwort was still in the top 10 for yearly nectar production. The strong nectar production of certain members of the genus can be exploited to the farmer's advantage, possibly in combination with biological control. In particular, the yellow starthistle (C. solstitialis) as well as spotted knapweed (C. maculosa) are major honey plants for beekeepers. Monofloral honey from these plants is light and slightly tangy, and one of the finest honeys produced in the United States – due to its better availability, it is even fraudulently relabeled and sold as the scarce and expensive sourwood honey of the Appalachian Mountains. Placing beehives near stands of Centaurea will cause increased pollination. As most seedheads fail however when biocontrol pests have established themselves, the plants will bloom ever more abundantly in an attempt to replace the destroyed seedheads, to the point where they exhaust their resources in providing food for the pests (seeds), bees (pollen) and humans (honey). Output of allelopathic compounds is also liable to be reduced under such conditions – the plant has to compromise between allocating energy to reproduction and defense. This renders the weeds more likely to be suppressed by native vegetation or crops in the following years, especially if properly timed controlled burning[5] and/or targeted grazing by suitable livestock are also employed. While yellow starthistle and perhaps other species are toxic to equines, some other livestock may eat the non-spiny knapweeds with relish. In Europe, common knapweed (C. nigra) and globe knapweed (C. macrocephala) are locally important pollen sources for honeybees in mid-late summer.
8-Hydroxyquinoline has been identified as a main allelopathic compound produced by diffuse knapweed (C. diffusa); native North American plants are typically sensitive to it, while those of Eastern Europe and Asia Minor usually have coevolved with the knapweed and are little harmed if at all, aided by native microorganisms that break down or even feed on the abundantly secreted compound. Thus, 8-hydroxyquinoline is potentially useful to control American plants that have become invasive weeds in the diffuse knapweed's native range.
Arctiin, found in C. imperialis, has shown anticancer activity in laboratory studies. The roots of the long-lost C. foliosa, an endemic of Hatay Province (Turkey), are used in folk medicine, and other species are presumably too. A South Italian variety[verification needed] of the purple starthistle (C. calcitrapa) is traditionally consumed by ethnic Albanians (Arbëreshë people) in the Vulture area (southern Italy); e.g. in the Arbëreshë communities in Lucania the young whorls of C. calcitrapa are boiled and fried in mixtures with other weedy non-cultivated greens. According to research by the Michael Heinrich group at the Centre for Pharmacognosy and Phytotherapy (School of Pharmacy, University of London) "the antioxidant activity [...] of the young whorls of Centaurea calcitrapa, both in the DPPH and in the lipid peroxidation inhibition assays, [is] very interesting and [the] species should be investigated phytochemically and biochemically focusing on these properties". Extracts from C. calcitrapa were furthermore found to have significant xanthine oxidase (XO)-inhibiting activity.
Spotted knapweed as well as other species are rich in cnicin, a bitter compound found mainly in the leaves and often used to flavor the digestif amaro. In western Crete, Greece a local variety[verification needed] of C. calcitrapa called gourounaki (γουρουνάκι "little pig") also has its leaves eaten boiled by the locals. In the same island an endemic local species, C. idaea called katsoula (κατσούλα), tsita (τσίτα) or aspragatha (ασπραγκάθα), has its leaves eaten boiled by the locals too.
Some species are cultivated as ornamental plants in gardens. As regards other aspects of popular culture, cornflower (C. cyanus) is the floral emblem of Östergötland province (Sweden) – where is it called blåklint, literally "blue mountain" – and of Päijänne Tavastia region in Finland, where it is known as ruiskaunokki ("rye-beaks") or ruiskukka ("rye-flower"). It is also the national flower of Estonia where its local name rukkilill means "rye-lily", Belarus where it is called vałoška (Belarusian: валошка), and one of those of Germany where it is called Kornblume ("cornflower"). The origin of the name "caltrop" for the ancient low-tech area denial weapon is probably in some way connected with C. calcitrapa and its spiny seeds. This plant is attested to by the colloquial name "caltrop" at a time when the weapons were still called by their Roman name tribulus. Lastly, the color cornflower blue is named after C. cyanus. Cornflower is also used as a cut flower.
As namesake member of the subtribe Centaureinae of tribe Cardueae, the knapweeds are probably most closely related to genera such as Carthamus (distaff thistles), Cnicus (blessed thistle), Crupina (crupinas) or Notobasis (Syrian thistle), and somewhat less closely to most other thistles. The monotypic Cnicus seems in fact to properly belong in Centaurea.
Research in the late 20th century shows that Centaurea as traditionally defined is polyphyletic. A number of 19th- and 20th-century efforts to reorganize the genus were not successful, and it is not yet clear what the consequences of the recent research will be for classification of this genus and other related genera. The type species C. centaurium stands somewhat apart from the main lineage of knapweeds and thus the taxonomic consequences of a rearrangement might be severe, with hundreds of species needing to be moved to new genera. It has thus been proposed to change the type species to one of the main lineages to avoid this problem. What seems certain however is that the basketflowers – presently treated as a section Plectocephalus – will be reinstated as a distinct genus in the near future. The rock-centauries (Cheirolophus), formerly usually included in Centaurea, are now already treated as separate genus.
Better-known Centaurea species include:
Centaurea acaulis
Centaurea adpressa
Centaurea aegyptiaca
Centaurea aeolica
Centaurea aggregata
Centaurea akamantis – Akamas centaurea
Centaurea alba
Centaurea albonitens Turrill
Centaurea alpestris
Centaurea alpina
Centaurea ambigua
Centaurea amblyolepis
Centaurea americana – American basketflower, American starthistle
Centaurea ammocyanus
Centaurea antennata Dufour
Centaurea antiochia Boiss.
Centaurea aplolepa
Centaurea aplolepa subsp. carueliana
Centaurea appendicigera C.Koch
Centaurea argentea
Centaurea ascalonica
Centaurea aspera L. – rough starthistle
Centaurea atacamensis (Reiche) I.M.Johnst.
Centaurea atropurpurea
Centaurea ×aurata
Centaurea babylonica L.
Centaurea balsamita
Centaurea behen L. – ak behmen (Turkish)
Centaurea bella
Centaurea benedicta – Cnicus
Centaurea bieberseinii
Centaurea borjae
Centaurea bovina
Centaurea bracteata
Centaurea brevifimbriata Hub.-Mor.
Centaurea bulbosa
Centaurea busambarensis Guss.
Centaurea cachinalensis
Centaurea calcitrapa – purple starthistle, red starthistle, "caltrop"
Centaurea calcitrapoides
Centaurea cariensis Boiss.
Centaurea cariensiformis Hub.-Mor.
Centaurea caroli-henrici Gabrieljan & Dittrich
Centaurea centaurium L.
Centaurea chilensis
Centaurea cineraria – velvet centaurea, dusty miller
Centaurea clementei
Centaurea collina L.
Centaurea corymbosa
Centaurea crithmifolia
Centaurea crocodylium
Centaurea cyanoides J.Berggr. & Wahlenb.
Centaurea cyanus – cornflower, bachelor's button, boutonniere flower, hurtsickle, bluebottle, basketflower
Centaurea damascena
Centaurea debeauxii Gren. & Godr.
Centaurea demirizii Wagenitz
Centaurea depressa – low cornflower
Centaurea deusta
Centaurea diffusa – diffuse knapweed, white knapweed, tumble knapweed
Centaurea diluta – North African knapweed
Centaurea drabifolia Sm.
Centaurea drabifolioides Hub.-Mor.
Centaurea dschungarica
Centaurea emilae Hüseynova et Qaraxani[13]
Centaurea eriophora
Centaurea eryngioides
Centaurea filiformis
Centaurea fischeri Willd.
Centaurea floccosa
Centaurea foliosa Boiss. & Kotschy
Centaurea forojuliensis
Centaurea friderici Vis. – palagruška zečina (Croatian)
Centaurea gayana
Centaurea glaberrima Tausch
Centaurea glastifolia
Centaurea grinensis
Centaurea gymnocarpa
Centaurea haradjianii Wagenitz
Centaurea hedgei
Centaurea helenioides Boiss.
Centaurea hermannii F.Hermann
Centaurea horrida Badarò – fiordaliso spinoso (Italian)
Centaurea hyalolepis
Centaurea hypoleuca
Centaurea iberica – Iberian starthistle, Iberian knapweed
Centaurea idaea – katsoula, tsita (Cretan Greek)
Centaurea imperialis Hausskn. ex Bornm.
Centaurea jabukensis
Centaurea jacea – brown knapweed, brownray knapweed
Centaurea kasakorum
Centaurea kopetaghensis
Centaurea kotschyana Heuff.
Centaurea lanulata
Centaurea leptophylla
Centaurea leucophylla
Centaurea limbata
Centaurea lydia Boiss.
Centaurea macrocephala Puschk. ex Willd. – globe knapweed, Armenian basketflower
Centaurea maculosa – spotted knapweed (might belong in C. stoebe subsp. micranthos)
Centaurea mannagettae
Centaurea margaritalba Klok.
Centaurea marschalliana
Centaurea melitensis – Maltese starthistle; tocalote, tocolote (California)
Centaurea minor
Centaurea moschata – sweet sultan
Centaurea ×moncktonii C.E.Britton – meadow knapweed, protean knapweed (= C. ×pratensis Thuill non Salisb.)
Centaurea monocephala
Centaurea montana – montane knapweed, perennial cornflower, mountain cornflower, mountain bluet
Centaurea napifolia L. – fiordaliso romano (Italian)
Centaurea nervosa Rchb. ex Steud.
Centaurea nigra – common knapweed, black knapweed, lesser knapweed, hardheads
Centaurea nigrescens – Tyrol knapweed, short-fringed knapweed, Tyrol thistle
Centaurea nigrifimbria (C.Koch) Sosn.
Centaurea nivea (Bornm.) Wagenitz
Centaurea onopordifolia
Centaurea orientalis L.
Centaurea ornata Willd.
Centaurea ovina
Centaurea pallescens Delile
Centaurea paniculata L.
Centaurea parlatoris
Centaurea pecho
Centaurea phrygia – wig knapweed
Centaurea pindicola
Centaurea polypodiifolia
Centaurea ×pratensis Salisb. (C. jacea × C. nigra) – meadow knapweed
Centaurea procurrens
Centaurea ×psammogena G.Gayer. (C. diffusa × C. stoebe subsp. micranthos)
Centaurea pseudocaerulescens
Centaurea pseudophrygia C.A.Mey.
Centaurea pulcherrima Willd.
Centaurea pullata L.
Centaurea pumilio
Centaurea ragusina L.
Centaurea rigida
Centaurea rothrockii Greenm. – Mexican basketflower, Rothrock's basketflower, Rothrock's knapweed
Centaurea ruthenica
Centaurea rutifolia Sm.
Centaurea sadleriana – Pannonian knapweed
Centaurea salicifolia Bieb. ex Willd.
Centaurea scabiosa – greater knapweed
Centaurea scannensis
Centaurea scoparia
Centaurea scopulorum Boiss. & Heldr.
Centaurea seguenzae
Centaurea seridis L.
Centaurea sibirica
Centaurea simplicicaulis
Centaurea sinaica
Centaurea solstitialis – yellow starthistle, golden starthistle, yellow cockspur, St. Barnaby's thistle, Barnaby thistle
Centaurea speciosa
Centaurea sphaerocephala L.
Centaurea stenolepis
Centaurea stoebe L.
Centaurea stoebe subsp. micranthos (Gugler) Hayek
Centaurea straminicephala
Centaurea sulphurea – Sicilian starthistle
Centaurea tauromenitana Guss.
Centaurea tenoreana
Centaurea tommasinii
Centaurea transalpina Schleich. ex DC.
Centaurea tchihatcheffii — yanardöner (Turkish)
Centaurea trichocephala Bieb. ex Willd. – featherhead knapweed
Centaurea triniifolia
Centaurea triumfettii All.
Centaurea ucriae Lacaita
Centaurea uniflora Turra
Centaurea verbascifolia Vahl
Centaurea verutum L.
Centaurea virgata
Centaurea virgata subsp. squarrosa – squarrose knapweed
Centaurea wiedemanniana Fisch. & Mey.
Centaurea yozgatensis Wagenitz
Formerly placed here
Plant species placed in Centaurea in former times include:
Acroptilon repens – Russian knapweed (as C. repens)
Cheirolophus crassifolius – Maltese rock-centaury (as C. crassifolia, C. spathulata)
Femeniasia balearica (as C. balearica)
Volutaria muricata (as C. muricata)
English Daisy - Bellis Perennis
Bellis perennis (/ˈbɛləs pəˈrɛnəs/),[2][3] the daisy, is a European species of the family Asteraceae, often considered the archetypal species of the name daisy. To distinguish this species from other plants known as daisies, it is sometimes qualified or known as common daisy, lawn daisy or English daisy.
Description
Bellis perennis is a perennial herbaceous plant growing to 20 centimetres (8 inches) in height.[4] It has short creeping rhizomes and rosettes of small rounded or spoon-shaped leaves that are from 2 to 5 cm (3⁄4–2 in) long and grow flat to the ground. The species habitually colonises lawns, and is difficult to eradicate by mowing, hence the term 'lawn daisy'. It blooms from March to September[4] and exhibits the phenomenon of heliotropism, in which the flowers follow the position of the sun in the sky.
The flowerheads are composite, about 2 to 3 cm (3⁄4–11⁄4 in) in diameter, in the form of a pseudanthium, consisting of many sessile flowers with white ray florets (often tipped red) and yellow disc florets. Each inflorescence is borne on a single leafless stem 2 to 10 cm (3⁄4–4 in), rarely 15 cm (6 in) tall. The capitulum, or disc of florets, is surrounded by two rows of green bracts known as "phyllaries".[5] The achenes are without pappus.[6]
Etymology
Bellis may come from bellus, Latin for "pretty", and perennis is Latin for "everlasting".[7]
The name "daisy", possibly originating with this plant,[4] is considered a corruption of "day's eye",[8] because the whole head closes at night and opens in the morning. Geoffrey Chaucer called it "eye of the day". In Medieval times, Bellis perennis or the English Daisy was commonly known as "Mary's Rose".[9]
Historically, the plant has also been widely known as bruisewort, and occasionally woundwort (although this name is now more closely associated with the genus Stachys). It is also known as bone flower.[10]
The Spurge family (Euphorbiaceae) is a large family of flowering plants with 240 genera and around 6,000 species. Most are herbs, but some, especially in the tropics, are also shrubs or trees. Some are succulent and resemble cacti.
This family occurs mainly in the tropics, with the majority of the species in the Indo-Malayan region and tropical America a good second. There is a large variety in tropical Africa, but it is not as abundant or varied as in these two other tropical regions. However, Euphorbia also has many species in non-tropical areas such as the Mediterranean, the Middle East, South Africa and southern USA.
The leaves are alternate, seldom opposite, with stipules. They are mainly simple, but where compound, are always palmate, never pinnate. Stipules may be reduced to hairs, glands, or spines, or in succulent species are sometimes absent.
The radially symmetrical flowers are unisexual, with the male and the female flowers usually occurring on the same plant. As can be expected from such a large family, there is a wide variety in the structure of the flowers. They can be monoecious or dioecious. The stamens (the male organs) can number from 1 to 10 (or even more). The female flowers are hypogynous, that is, with a superior ovary.
The genera in tribe Euphorbieae, subtribe Euphorbiinae (Euphorbia and close relatives) show a highly specialized form of pseudanthium ("false flower" made up of several true flowers) called a cyathium. This is usually a small cup-like involucre consisting of fused together bracts and peripheral nectary glands, surrounding a ring of male flowers, each a single stamen. In the middle of the cyathium stands a female flower: a single pistil with branched stigmas. This whole arrangement resembles a single flower.
The fruit is usually a schizocarp, sometimes a drupe. A typical schizocarp is the regma, a capsular fruit with three or more cells, each of which splits open at maturity into separate parts and then breaks away explosively, scattering the small seeds.
The family contains a large variety of phytotoxins (toxic substances produced by plants), mainly diterpene esters, alkaloids, glycosides, and ricin-type toxins.
A milky sap or latex is a characteristic of the subfamilies Euphorbioideae and Crotonoideae. This milky sap is poisonous in the Euphorbioideae, innocuous in the Crotonoideae. White mangrove (Excoecaria agallocha), or Blind-Your-Eye Mangrove sap causes blistering on contact and temporary blindness if it contacts the eyes. Other common names are Milky Mangrove, Buta Buta (Malay), Gewa (Bangladesh). The sap of spurge was used as a purgative.
Recent molecular studies have shown that the enigmatic family Rafflesiaceae, which was only recently recognized to belong to order Malpighiales, is derived from within Euphorbiaceae.
Detail showing structure of one flower--the "petals" are perhaps 6 mm across. The white "petals" are actually bracts, I believe. The female flower is the long-stalked structure jutting out from the center, and there are also clusters of male flowers at the base of the female flower stem--the yellow knobs.
Aha! The Wikipedia article on Euphorbiaceae has a good description of the flowers, and I'll quote it here:
Euphorbia and close relatives show a highly specialized form of pseudanthium ("false flower" made up of several true flowers) called a cyathium. This is usually a small cup-like involucre consisting of fused together bracts and peripheral nectary glands, surrounding a ring of male flowers, each a single stamen. In the middle of the cyathium stands a female flower: a single pistil with branched stigmas. This whole arrangement resembles a single flower.
Thanks, Wikipedia authors!
See also, the article on carpel, explaining the female parts of a flower.
Euphorbia_corollataPCCA20070522-2556A
Walking up to Pico Branco on the North East coast of Porto Santo, Cardoons Cynara cardunculus dot the landscape with their usual rich violet-purple flowers.
This flower had dried out, making it no less spectacular with its tall stems and massively spined capitulum. As a herbaceous perennial it will grow back from its root-stock.
Common on Madeira and Porto Santo, they are a hardy plant, known as Cardos or Pencas, flowering from July through October but often seen off season.
Native, cool season, annual, small, tufted annual herb 2-11 cm tall. Leaves are linear, innermost ones usually reduced to a short membranous hairless sheath. Stems are erect, unbranched and leafless. Flowerheads are terminal cymose heads enclosed by 2 primary bracts with dilated bases; each bract encloses 1–numerous pseudanthia; pseudanthia each consist of 1–30 ± fused female flowers superimposed in 1 or 2 rows, styles usually fused at base, usually with 1 male flower at base; secondary bracts 2 or 3 per pseudanthium. Flowering is in spring. Grows in heath, scrub, mallee, woodland and sclerophyll forest on sand or infertile soils; mostly east and north of the A.C.T., but this one was in Albury.
Superdomain: Neomura
Domain: Eukaryota
(unranked): Diaphoretickes
(unranked): Archaeplastida
Kingdom: Plantae
Clade: Streptophyta
Clade: Embryophytes
Clade: Polysporangiophytes
Subkingdom: Tracheobionta
Superdivision: Spermatophyta
Division: Magnoliophyta
Class: Magnoliopsida
Subclass: Asteridae
Order: Asterales
Family: Asteraceae
Subfamily: Carduoideae
Tribe: Cardueae
Subtribe: Arctiinae
Genus: Arctium
Species: A. minus
Euphorbia Flower in Spring in Bebra Gardens, Knaresborough, North Yorkshire, England.
© Mark Sunderland All Rights Reserved
For licensing enquiries visit www.marksunderland.com
Authors: Chiara Mizzotti, Marco Fambrini, Elisabetta Caporali, Simona Masiero, and Claudio Pugliesi
Citation: Botany, 2015, 93(3): 171-181, 10.1139/cjb-2014-0210
Source: www.nrcresearchpress.com/doi/abs/10.1139/cjb-2014-0210#.U...
Fig. 7. Achenes and embryos of wild type (WT) and tubular ray flower (turf) mutant. (A) A turf inflorescence (side view) showing three fertilized achenes characterized by phytomelanin deposition. Involucral bracts were removed to reveal florets at the periphery of the inflorescence. (B) An empty achene from a WT ray floret (1), a fertile achene from a WT disc floret (3), and a filled achene from a tubular-like ray floret (2). (C) Heteromorphic zygotic embryos isolated from a mature achene at the periphery (tubular-like ray florets) (1) or from a mature achene of the first row of disc florets (2) in a turf inflorescence. Scale bars = 0.48 cm (A), 0.40 cm (B), and 1.28 mm (C).
Botany, NRC Research Press, a division of Canadian Science Publishing. www.nrcresearchpress.com/journal/cjb
For rights and permission, click here [bit.ly/1pL1c5f]
The sunflower (Helianthus annuus) is an annual plant in the family Asteraceae, with a large flower head (capitulum). The stem of the flower can grow up to 3 metres tall, with a flower head that can be 30 cm wide. Other types of sunflowers include the California Royal Sunflower, which has a burgundy (red + purple) flower head.
The flower head is actually an inflorescence made of hundreds or thousands of tiny flowers called florets. The central florets look like the centre of a normal flower, and the outer florets look like yellow petals. All together they make up a "false flower" or pseudanthium. The benefit to the plant is that it is very easily seen by the insects and birds which pollinate it, and it produces thousands of seeds.
Bellis perennis the daisy, is a European species of the family Asteraceae, often considered the archetypal species of the name daisy. To distinguish this species from other plants known as daisies, it is sometimes qualified as common daisy, lawn daisy or English daisy.
Bellis perennis is a perennial herbaceous plant growing to 20 centimetres (8 inches) in height. It has short creeping rhizomes and rosettes of small rounded or spoon-shaped leaves that are from 2 to 5 cm (3⁄4–2 in) long and grow flat to the ground. The species habitually colonises lawns, and is difficult to eradicate by mowing, hence the term 'lawn daisy'. It blooms from March to September and exhibits the phenomenon of heliotropism, in which the flowers follow the position of the sun in the sky.
The flowerheads are composite, about 2 to 3 cm (3⁄4–1+1⁄4 in) in diameter, in the form of a pseudanthium, consisting of many sessile flowers with white ray florets (often tipped red) and yellow disc florets. Each inflorescence is borne on a single leafless stem 2 to 10 cm (3⁄4–4 in), rarely 15 cm (6 in) tall. The capitulum, or disc of florets, is surrounded by two rows of green bracts known as "phyllaries". The achenes are without pappus.
Bellis may come from bellus, Latin for "pretty", and perennis is Latin for "everlasting".
The name "daisy", possibly originating with this plant, is considered a corruption of "day's eye", because the whole head closes at night and opens in the morning. Geoffrey Chaucer called it "eye of the day". In Medieval times, Bellis perennis or the English Daisy was commonly known as "Mary's Rose". It is also known as bone flower.
Historically, the plant has also been widely known as bruisewort, and occasionally woundwort (although this name is now more closely associated with the genus Stachys).
Bellis perennis is native to western, central and northern Europe, including remote islands such as the Faroe Islands, but has become widely naturalised in most temperate regions, including the Americas and Australasia. It prefers field-like habitats.
The species generally blooms from early to midsummer, although when grown under ideal conditions, it has a very long flowering season and will even produce a few flowers in the middle of mild winters.
It can generally be grown where minimum temperatures are above −35 °C (−30 °F), in full sun to partial shade conditions, and requires little or no maintenance. It has no known serious insect or disease problems and can generally be grown in most well-drained soils. The plant may be propagated either by seed after the last frost, or by division after flowering.
Though not native to the United States, the species is still considered a valuable ground cover in certain garden settings (e.g., as part of English or cottage inspired gardens, as well as spring meadows where low growth and some color is desired in parallel with minimal care and maintenance while helping to crowd out noxious weeds once established and naturalised).
Numerous single- and double-flowered varieties are in cultivation, producing flat or spherical blooms in a range of sizes (1 to 6 cm or 3⁄8 to 2+3⁄8 in) and colours (red, pink and white). They are generally grown from seed as biennial bedding plants. They can also be purchased as plugs in Spring.
It has been reported to be mostly self-fertilizing, but some plants may be self-sterile.
Bellis perennis may be used as a potherb. Young leaves can be eaten raw in salads, or cooked, though the leaves become increasingly astringent with age. Flower buds and petals can be eaten raw in sandwiches, soups and salads. It is also used as a tea and as a vitamin supplement.
B. perennis has astringent properties and has been used in herbal medicine.
Daisies have traditionally been used for making daisy chains in children's games.
Daisy is used as a feminine name, and sometimes as a nickname for people named Margaret, after the French name for the oxeye daisy, marguerite.
The daisy is the national flower of the Netherlands.
Common sunflower (Helianthus annuus)
Wild Helianthus annuus is a widely branched annual plant with many flower heads. The domestic sunflower, however, often possesses only a single large inflorescence (flower head) atop an unbranched stem.
The term Helianthus annuus is derived from the Greek Helios, meaning sun, and anthos, meaning flower, and from Latin annuus, meaning annual.
What is often called the "flower" of the sunflower is actually a "flower head" (pseudanthium) of numerous small individual five-petaled flowers ("florets"). The outer flowers, which resemble petals, are called ray flowers. Each "petal" consists of a ligule composed of fused petals of an asymmetrical ray flower. They are sexually sterile and may be yellow, red, orange, or other colors. The flowers in the center of the head are called disk flowers. These mature into fruit (sunflower "seeds"). The disk flowers are arranged spirally.
A common misconception is that flowering sunflower heads track the Sun across the sky. Although immature flower buds exhibit this behaviour, the mature flowering heads point in a fixed (and typically easterly) direction throughout the day.
When growth of the flower stalk stops and the flower is mature, the heliotropism also stops and the flower faces east from that moment onward. This eastward orientation allows rapid warming in the morning and, as a result, an increase in pollinator visits.
*** Arctic Islands 2017 Expedition (polarforskningsportalen.se/en/arctic/expeditions/arktiska...) ***
The common sunflower (Helianthus annuus) is a species of large annual forb of the genus Helianthus. It is commonly grown as a crop for its edible oily seeds. Apart from cooking oil production, it is also used as livestock forage (as a meal or a silage plant), as bird food, in some industrial applications, and as an ornamental in domestic gardens. Wild H. annuus is a widely branched annual plant with many flower heads. The domestic sunflower, however, often possesses only a single large inflorescence (flower head) atop an unbranched stem.
Helianthus annuus (common sunflower) belong to the dicotyledonous category, distinguishing them from monocots. As dicots, sunflowers possess embryos with two veins, known as cotyledons, a characteristic feature that sets them apart. The veins in their leaves exhibit a net-like pattern, in contrast to monocots, which typically display parallel leaf veins.
Etymology
In the binomial name Helianthus annuus the genus name is derived from the Greek ἥλιος : hḗlios 'sun' and ἄνθος : ánthos 'flower'. The species name annuus means 'annual' in Latin.
Domestication
The plant was first domesticated in the Americas. Sunflower seeds were brought to Europe from the Americas in the 16th century, where, along with sunflower oil, they became a widespread cooking ingredient. With time, the bulk of industrial-scale production has shifted to Eastern Europe, and (as of 2020) Russia and Ukraine together produce over half of worldwide seed production.
Description
The plant has an erect rough-hairy stem, reaching typical heights of 3 metres (10 feet). The tallest sunflower on record achieved 9.17 m (30 ft 1 in). Sunflower leaves are broad, coarsely toothed, rough and mostly alternate; those near the bottom are largest and commonly heart-shaped.
Flower
The plant flowers in summer. What is often called the "flower" of the sunflower is actually a "flower head" (pseudanthium), 7.5–12.5 centimetres (3–5 in) wide, of numerous small individual five-petaled flowers ("florets"). The outer flowers, which resemble petals, are called ray flowers. Each "petal" consists of a ligule composed of fused petals of an asymmetrical ray flower. They are sexually sterile and may be yellow, red, orange, or other colors. The spirally arranged flowers in the center of the head are called disk flowers. These mature into fruit (sunflower "seeds").
The prairie sunflower (H. petiolaris) is similar in appearance to the wild common sunflower; the scales in its central disk are tipped by white hairs.
Heliotropism
A common misconception is that flowering sunflower heads track the Sun across the sky. Although immature flower buds exhibit this behaviour, the mature flowering heads point in a fixed (and typically easterly) direction throughout the day. This old misconception was disputed in 1597 by the English botanist John Gerard, who grew sunflowers in his famous herbal garden: "[some] have reported it to turn with the Sun, the which I could never observe, although I have endeavored to find out the truth of it." The uniform alignment of sunflower heads in a field might give some people the false impression that the flowers are tracking the Sun.
This alignment results from heliotropism in an earlier development stage, the young flower stage, before full maturity of flower heads (anthesis). Young sunflowers orient themselves in the direction of the sun. At dawn the head of the flower faces east and moves west throughout the day. When sunflowers reach full maturity they no longer follow the sun, and continuously face east. Young flowers reorient overnight to face east in anticipation of the morning. Their heliotropic motion is a circadian rhythm, synchronized by the sun, which continues if the sun disappears on cloudy days or if plants are moved to constant light. They are able to regulate their circadian rhythm in response to the blue-light emitted by a light source. If a sunflower plant in the bud stage is rotated 180°, the bud will be turning away from the sun for a few days, as resynchronization with the sun takes time.
When growth of the flower stalk stops and the flower is mature, the heliotropism also stops and the flower faces east from that moment onward. This eastward orientation allows rapid warming in the morning and, as a result, an increase in pollinator visits. Sunflowers do not have a pulvinus below their inflorescence. A pulvinus is a flexible segment in the leaf stalks (petiole) of some plant species and functions as a 'joint'. It effectuates leaf motion due to reversible changes in turgor pressure, which occurs without growth. The sensitive plant's closing leaves are a good example of reversible leaf movement via pulvinuli.
Floret arrangement
Generally, each floret is oriented toward the next by approximately the golden angle, 137.5°, producing a pattern of interconnecting spirals, where the number of left spirals and the number of right spirals are successive Fibonacci numbers. Typically, there are 34 spirals in one direction and 55 in the other; however, in a very large sunflower head there could be 89 in one direction and 144 in the other. This pattern produces the most efficient packing of seeds mathematically possible within the flower head.
Genome
The sunflower genome is diploid with a base chromosome number of 17 and an estimated genome size of 2,871–3,189 million base pairs. Some sources claim its true size is around 3.5 billion base pairs (slightly larger than the human genome).
Distribution and habitat
The plant was first domesticated in the Americas. Sunflowers grow best in fertile, moist, well-drained soil with heavy mulch. They often appear on dry open areas and foothills. Outside of cultivation, the common sunflower is found on moist clay based soils in areas with climates similar to Texas. In contrast the related Helianthus debilis and Helianthus petiolaris are found on drier, sandier soils.
The precise native range is difficult to determine. According to Plants of the World Online (POWO) it is native to Arizona, California, and Nevada in the present day United States and to all parts of Mexico except the Gulf Coast and southeast. Though not giving as great of detail, the Missouri Botanical Garden Plant Finder also lists it as native to the Western United States and Canada. The information published by the Biota of North America Program (BONAP) largely agrees with this, showing the common sunflower as native to states west of the Mississippi, though also listed as a noxious weed in Iowa, Minnesota, and Texas. Regardless of its original range it can now be found in almost every part of the world that is not tropical, desert, or tundra.
Ecology
Threats and diseases
Main article: List of sunflower diseases
One of the major threats that sunflowers face today is Fusarium, a filamentous fungus that is found largely in soil and plants. It is a pathogen that over the years has caused an increasing amount of damage and loss of sunflower crops, some as extensive as 80% of damaged crops.
Downy mildew is another disease to which sunflowers are susceptible. Its susceptibility to downy mildew is particular high due to the sunflower's way of growth and development. Sunflower seeds are generally planted only an inch deep in the ground. When such shallow planting is done in moist and soaked earth or soil, it increases the chances of diseases such as downy mildew.
Another major threat to sunflower crops is broomrape, a parasite that attacks the root of the sunflower and causes extensive damage to sunflower crops, as high as 100%.
Cultivation
In commercial planting, seeds are planted 45 cm (1+1⁄2 ft) apart and 2.5 cm (1 in) deep.
History
Common sunflower was one of several plants cultivated by Native Americans in prehistoric North America as part of the Eastern Agricultural Complex. Although it was commonly accepted that the sunflower was first domesticated in what is now the southeastern US, roughly 5,000 years ago, there is evidence that it was first domesticated in Mexico around 2600 BCE. These crops were found in Tabasco, Mexico, at the San Andres dig site. The earliest known examples in the US of a fully domesticated sunflower have been found in Tennessee, and date to around 2300 BCE. Other very early examples come from rockshelter sites in Eastern Kentucky. Many indigenous American peoples used the sunflower as the symbol of their solar deity, including the Aztecs and the Otomi of Mexico and the Incas in South America. In 1510, early Spanish explorers encountered the sunflower in the Americas and carried its seeds back to Europe. Of the four plants known to have been domesticated in eastern North America and to have become important agricultural commodities, the sunflower is currently the most economically important.
Research of phylogeographic relations and population demographic patterns across sunflowers has demonstrated that earlier cultivated sunflowers form a clade from wild populations from the Great Plains, which indicates that there was a single domestication event in central North America. Following the cultivated sunflower's origin, it may have gone through significant bottlenecks dating back to ~5,000 years ago.
In the 16th century the first crop breeds were brought from America to Europe by explorers. Domestic sunflower seeds have been found in Mexico, dating to 2100 BCE. Native American people grew sunflowers as a crop from Mexico to Southern Canada. They then were introduced to the Russian Empire, where oilseed cultivators were located, and the flowers were developed and grown on an industrial scale. The Russian Empire reintroduced this oilseed cultivation process to North America in the mid-20th century; North America began their commercial era of sunflower production and breeding. New breeds of the Helianthus spp. began to become more prominent in new geographical areas. During the 18th century, the use of sunflower oil became very popular in Russia, particularly with members of the Russian Orthodox Church, because only plant-based fats were allowed during Lent, according to fasting traditions. In the early 19th century, it was first commercialized in the village of Alexeyevka in Voronezh Governorate by the merchant named Daniil Bokaryov, who developed a technology suitable for its large-scale extraction, and quickly spread around. The town's coat of arms has included an image of a sunflower ever since.
Production
Sunflower seed production – 2020
Country(Millions of tonnes)
Russia13.3
Ukraine13.1
Argentina3.2
China2.4
Romania2.2
Turkey2.1
United States1.4
World50.2
Source: FAOSTAT of the United Nations
In 2020, world production of sunflower seeds was 50 million tonnes, led by Russia and Ukraine with 53% combined of the total (table).
Fertilizer use
Researchers have analyzed the impact of various nitrogen-based fertilizers on the growth of sunflowers. Ammonium nitrate was found to produce better nitrogen absorption than urea, which performed better in low-temperature areas.
Crop rotation
Sunflower cultivation typically uses crop rotation, often with cereals, soybean, or rapeseed. This reduces idle periods and increases total sunflower production and profitability.
Hybrids and cultivars
In today's market, most of the sunflower seeds provided or grown by farmers are hybrids. Hybrids or hybridized sunflowers are produced by cross-breeding different types and species, for example cultivated sunflowers with wild species. By doing so, new genetic recombinations are obtained ultimately leading to the production of new hybrid species. These hybrid species generally have a higher fitness and carry properties or characteristics that farmers look for, such as resistance to pathogens.
Hybrid, Helianthus annuus dwarf2 does not contain the hormone gibberellin and does not display heliotropic behavior. Plants treated with an external application of the hormone display a temporary restoration of elongation growth patterns. This growth pattern diminished by 35% 7–14 days after final treatment.
Hybrid male sterile and male fertile flowers that display heterogeneity have a low crossover of honeybee visitation. Sensory cues such as pollen odor, diameter of seed head, and height may influence pollinator visitation of pollinators that display constancy behavior patterns.
Sunflowers are grown as ornamentals in a domestic setting. Being easy to grow and producing spectacular results in any good, moist soil in full sun, they are a favourite subject for children. A large number of cultivars, of varying size and color, are now available to grow from seed. The following are cultivars of sunflowers (those marked agm have gained the Royal Horticultural Society's Award of Garden Merit):-
American Giant
Arnika
Autumn Beauty
Aztec Sun
Black Oil
Chianti Hybrid
Claret agm
Dwarf Sunspot
Evening Sun
Florenza
Giant Primrose
Gullick's Variety agm
Incredible
Indian Blanket Hybrid
Irish Eyes
Italian White
Kong Hybrid
Large Grey Stripe
Lemon Queen agm
Loddon Gold agm
Miss Mellish agm
Monarch agm
Mongolian Giant
Moon-Walker
Munchkin
Orange Sun
Pastiche agm
Peach Passion
Peredovik
Prado Red
Red Sun
Ring of Fire
Rostov
Russian Giant
Skyscraper
Solar Eclipse
Soraya
Strawberry Blonde
Sunny Hybrid
Sunsation Yellow
Sunshine
Taiyo
Tarahumara
Teddy Bear agm
Thousand Suns
Titan
Valentine agm
Velvet Queen
Yellow Disk
Uses
Sunflower "whole seed" (fruit) are sold as a snack food, raw or after roasting in ovens, with or without salt and/or seasonings added. Sunflower seeds can be processed into a peanut butter alternative, sunflower butter. It is also sold as food for birds and can be used directly in cooking and salads. Native Americans had multiple uses for sunflowers in the past, such as in bread, medical ointments, dyes and body paints.
Sunflower oil, extracted from the seeds, is used for cooking, as a carrier oil and to produce margarine and biodiesel, as it is cheaper than olive oil. A range of sunflower varieties exist with differing fatty acid compositions; some "high-oleic" types contain a higher level of monounsaturated fats in their oil than even olive oil. The oil is also sometimes used in soap. After World War I, during the Russian Civil War, people in Ukraine used sunflower seed oil in lamps as a substitute for kerosene due to shortages. The light from such a lamp has been described as "miserable" and "smoky."
The cake remaining after the seeds have been processed for oil is used as a livestock feed. The hulls resulting from the dehulling of the seeds before oil extraction can also be fed to domestic animals. Some recently developed cultivars have drooping heads. These cultivars are less attractive to gardeners growing the flowers as ornamental plants, but appeal to farmers, because they reduce bird damage and losses from some plant diseases. Sunflowers also produce latex, and are the subject of experiments to improve their suitability as an alternative crop for producing hypoallergenic rubber.
Traditionally, several Native American groups planted sunflowers on the north edges of their gardens as a "fourth sister" to the better-known three sisters combination of corn, beans, and squash.[61] Annual species are often planted for their allelopathic properties. It was also used by Native Americans to dress hair. Among the Zuni people, the fresh or dried root is chewed by the medicine man before sucking venom from a snakebite and applying a poultice to the wound. This compound poultice of the root is applied with much ceremony to rattlesnake bites.
However, for commercial farmers growing other commodity crops, the wild sunflower is often considered a weed. Especially in the Midwestern US, wild (perennial) species are often found in corn and soybean fields and can decrease yields. The decrease in yield can be attributed to the production of phenolic compounds which are used to reduce competition for nutrients in nutrient-poor growing areas of the common sunflower.
Phytoremediation
Helianthus annuus can be used in phytoremediation to extract pollutants from soil such as lead and other heavy metals, such as cadmium, zinc, cesium, strontium, and uranium, and in rhizofiltration to neutralize radionuclides, such as caesium-137 and strontium-90 from a pond after the Chernobyl disaster. A similar campaign was mounted in response to the Fukushima Daiichi nuclear disaster.[
Culture
Vincent van Gogh – "Lausanne" Sunflowers (1888)
During the 19th century, it was believed that nearby plants of the species would protect a home from malaria.
The Zuni people use the blossoms ceremonially for anthropic worship. Sunflowers were also worshipped by the Incas because they viewed it as a symbol for the Sun.
The flowers are the subject of Vincent van Gogh's Sunflowers series of still-life paintings.
In July 2015, viable seeds were acquired from the field where Malaysia Airlines Flight 17 crashed on a year earlier and were grown in tribute to the 15 Dutch residents of Hilversum who were killed. Earlier that year, Fairfax chief correspondent Paul McGeough and photographer Kate Geraghty had collected 1.5 kg of sunflower seeds from the wreck site for family and friends of the 38 Australian victims, who aimed to give them a poignant symbol of hope.
On May 13, 2021, during the National Costume competition of the Miss Universe 2020 beauty pageant, Miss Dominican Republic Kimberly Jiménez wore a "Goddess of Sunflowers" costume covered in gold and yellow rhinestones that included several real sunflowers sewn onto the fabric.
Modern stories often claim that in Greek mythology, the nymph Clytie transformed into a sunflower when she pined after her former lover Helios, the god of the sun, who spurned her and left her for another. However, sunflowers are not native to Greece or Italy, but to North America. The original story is about another flower, the heliotropium.
National and state symbol
The sunflower is the national flower of Ukraine. Ukrainians used sunflower as a main source of cooking oil instead of butter or lard forbidden by the Orthodox Church when observing Lent. They were planted to clean nuclear waste in Chernobyl. In June 1996, United States, Russia and Ukraine officials planted sunflowers at the Pervomaysk missile base where Soviet nuclear weapons were formerly placed.[69] During the Russo-Ukrainian War, a video widely shared on social media showed a Ukrainian woman confronting a Russian soldier, telling the latter to "take these seeds and put them in your pockets so at least sunflowers will grow when you all lie down here". The sunflower has since become a global symbol of resistance, unity, and hope.
The sunflower is also the state flower of the US state of Kansas, and one of the city flowers of Kitakyūshū, Japan.
Movement symbol
During the late 19th century, the flower was used as the symbol of the Aesthetic Movement.
The sunflower was chosen as the symbol of the Spiritualist Church, for many reasons, but mostly because of the (false) belief that the flowers turn toward the sun as "Spiritualism turns toward the light of truth". Modern Spiritualists often have art or jewelry with sunflower designs.
The sunflower is often used as a symbol of green ideology. The flower is also the symbol of the Vegan Society.
The sunflower is the symbol behind the Sunflower Movement, a 2014 mass protest in Taiwan.
The Hidden Disabilities Sunflower was first used as a visible symbol (typically worn on a lanyard) May 2016 at London Gatwick Airport. It has since come into common usage throughout the UK, and in the Commonwealth more generally.
2019 Photograph, Common Sunflower Patterns in the Capitulum or Pseudanthium (Helianthus annuus, Aster family Asteraceae), McKee-Beshers WMA, Poolesville, Maryland, © 2019.
Centaurea is a genus of over 700 species of herbaceous thistle-like flowering plants in the family Asteraceae. Members of the genus are found only north of the equator, mostly in the Eastern Hemisphere; the Middle East and surrounding regions are particularly species-rich. In the western United States, yellow starthistles are an invasive species. Around the year 1850, seeds from the plant had arrived to the state of California. It is believed that those seeds came from South America.
Common names
Common names for this genus are centaury, centory, starthistles, knapweeds, centaureas and the more ambiguous "bluets"; a vernacular name used for these plants in parts of England is "loggerheads" (common knapweed). The Plectocephalus group – possibly a distinct genus – is known as basketflowers. "Cornflower" is used for a few species, but that term more often specifically means either C. cyanus (the annual cornflower) or Centaurea montana (the perennial cornflower). The common name "centaury" is sometimes used, although this also refers to the unrelated plant genus Centaurium.
The name is said to be in reference to Chiron, the centaur of Greek mythology who discovered medicinal uses of a plant eventually called "centaury".
Description
Knapweeds are robust weedy plants. Their leaves, spiny in some species, are usually deeply divided into elongated lobes at least in the plants' lower part, becoming entire towards the top. The "flowers" (actually pseudanthium inflorescences) are diverse in colour, ranging from intense blues, reds and yellows to any mixture of these and lighter shades towards white. Often, the disk flowers are much darker or lighter than the ray flowers, which also differ in morphology and are sterile. Each pseudanthium sits atop a cup- or basket-like cluster of scaly bracts, hence the name "basketflowers". Many species, in particular those inhabiting more arid regions, have a long and strong taproot.
Certain knapweeds have a tendency to dominate large stretches of landscape together with a few other plants, typically one or two grasses and as many other large herbaceous plants. The common knapweed (C. nigra) for example is plentiful in the mesotrophic grasslands of England and nearby regions. It is most prominently found in pastures or meadows dominated by cock's-foot (Dactylis glomerata) as well as either of crested dog's-tail (Cynosurus cristatus) and false oat-grass (Arrhenatherum elatius). It is also often found in mesotrophic grassland on rendzinas and similar calcareous soils in association with glaucous sedge (Carex flacca), sheep's fescue (Festuca ovina), and either tor-grass (Brachypodium pinnatum) and rough hawkbit (Leontodon hispidus), or upright brome (Bromus erectus). In these grasslands, greater knapweed (C. scabiosa) is found much more rarely by comparison, often in association with red fescue (Festuca rubra) in addition to cock's-foot and false oat-grass.
Due to their habit of dominating ecosystems under good conditions, many Centaurea species can become invasive weeds in regions where they are not native. In parts of North America, diffuse knapweed (C. diffusa), spotted knapweed (C. maculosa) and yellow starthistle (C. solstitialis) cause severe problems in agriculture due to their uncontrolled spread. The seeds are typically transported by human traffic, in particular the tires of all-terrain vehicles. The two knapweeds are harmful mainly because they are strongly allelopathic, producing powerful toxins in their roots that stunt the growth of plants around them not adapted to this. Yellow starthistle, meanwhile, is inedible to most livestock due to its spines and apparently outright poisonous to horses and other equines. However, efficient methods of biological control by insect pests of these weeds have been developed; the knapweeds can also exploited to their detriment by targeted grazing. Controlled burning may also be used, though the timing is important to avoid the plants having seeded already, and neither allowing sufficient time for them to regrow from the rootstock.
Yet other species of Centaurea – mostly ones that occur between Italy and the Caucasus – are endemics of a single island or valley, and some of these are endangered. The Akamas Centaurea (Centaurea akamantis) of Cyprus is almost extinct, while the western Caucasus endemics C. leptophylla and C. straminicephala are at least very rare and C. hedgei and C. pecho from the same region are certainly not abundant either. The last four species would be adversely affected by the proposed Yusufeli Dam, which might actually destroy enough habitat to push the two rarer ones over the brink of extinction.
Centaurea are copious nectar producers, especially on high-lime soils. The high nectar yield of the genus makes it very attractive to insects such as butterflies – including the endangered Karner blue (Plebejus melissa samuelis) which visits introduced spotted knapweed – and day-flying moths – typically Zygaenidae, such as Zygaena loti or the six-spot burnet (Z. filipendulae). The larvae of some other Lepidoptera species use Centaurea species as food plants; see List of Lepidoptera that feed on Centaurea. Several of these are used in biological control of invasive knapweeds and starthistles.
Larvae of several true weevils (Curculionidae) of the subfamily Lixinae also feed on Centaurea. Some genera – such as Larinus whose larval food is flowerheads – have many species especially adapted to particular knapweeds or starthistle and are used in biological control too. These include the yellow starthistle flower weevil (L. curtus) for yellow starthistle, lesser knapweed flower weevil (L. minutus) for diffuse knapweed and blunt knapweed flower weevil (L. obtusus) for spotted knapweed. Broad-nosed seedhead weevil (Bangasternus fausti) larvae eat diffuse, spotted and squarrose knapweed (C. virgata ssp. squarrosa), while those of the yellow starthistle bud weevil (B. orientalis) do not seem to live on anything other than yellow starthistle and occasionally purple starthistle (C. calcitrapa). But perhaps most efficient in destroying developing yellow starthistle seedheads is the larva of the yellow starthistle hairy weevil (Eustenopus villosus). Knapweed root weevil (Cyphocleonus achates) larvae bore into the roots of spotted and to a lesser extentely diffuse knapweed, sometimes killing off the entire plant.
Also used in biological control are Tephritidae (peacock flies) whose larvae feed on Centaurea. Knapweed peacock fly (Chaetorellia acrolophi) larvae eat spotted knapweed and some other species. The yellow starthistle peacock fly (C. australis) has an initial generation each year which often uses cornflower (C. cyanus) as larval food; later generations switch to yellow starthistle. The flies are generally considered less efficient in destroying the growing seedheads than the weevils, but may be superior under certain conditions; employing flies and weevils in combination is expensive and does not noticeably increase their effect.
Use by humans
Although the genus may be considered by a quite significant number of relatively informed individuals to have an overall negative impact on human interests, particularly agricultural interests, the situation is not straightforward enough to simply declare the genus, or, at least, its most aggressively-spreading species, altogether negative. For instance, due to their moderate to high nectar production, which can occur over a comparatively long duration, many species of Centaurea are popular food sources for insects that may otherwise attack certain crops.[citation needed] It may be advisable for some types of farms to allow certain species in this genus, such as cornflower (C. cyanus) in a European setting, to grow adjacent to fields. Although they support and attract many types of beneficial life (not just beetles), these areas are known as beetle banks. When they are present, some pests may be drawn away from crops to them and predatory insects and arachnids that feed upon pest insects will be better-supported by these more naturalized areas. They additionally have the beneficial aspect of supporting pollinators, unlike many field crops such as maize. Moreover, being untreated with pesticides and providing more diversity, plants growing in more wild areas adjacent to farms produce more insects that attract and support birds which can also feed on pests that would harm crops. Insect production is especially high for beetle banks that have enough plants that serve in the role of host plant for immature insects, rather than just in the roles of adult food and/or shelter provision.
Some plants which are considered invasive or problematic in certain areas can have beneficial qualities that outweigh their negative qualities from a human and/or human agricultural point of view, although this sometimes requires some human management – particularly if adequate biological control has not been established for the more aggressive species. An example is wild parsnip, Pastinaca sativa, which produces florets that feed predatory (and other beneficial) insects as well as large tubular stems that provide winter shelter for native bees, wasps, and other organisms that can be beneficial for agriculture. The plant is considered invasive in some areas of the United States and is also often considered undesirable due to its ability to cause contact skin irritation. However, it also serves as a host plant for the black swallowtail butterfly, helps to bring nutrients up from soils with its deep taproot, and possesses evergreen foliage even in climate zones such as US zone 6. This foliage increases soil warmth and moisture which can be beneficial for certain types of life. Perhaps the most dramatic example of a generally disliked plant's beneficial qualities being usually overlooked is the often-despised ragwort, Jacobaea vulgaris, which topped the list by a large amount for nectar production in a UK study, with a production per floral unit of (2921 ± 448μg). This very high nectar production, coupled with its early blooming period, makes the plant helpful for the establishment of bee colonies in spring — a period that is often not well-served by commercial flower meadow seed mixes. It also has the situationally-beneficial quality of being a spring ephemeral, as well as an annual that lacks difficult-to-combat roots. Plants that provide necessary structural supports for invertebrate and small vertebrate predators can help to keep overall pest populations low.
The abundant nectar produced by C. solstitialis flowers attracts many pollinators. This is another reason for the success of the (situationally) highly invasive species. Due to genetic differences related to evolutionary adaption, not all members of Centaurea produce the same amount of nectar. Growing conditions, such as climate and soil, can have a very strong impact, even if the plants grow and flower. For instance, cornflower plants, Centaurea cyanus, produced 33% less seasonal nectar than Centaurea nigra in a UK study. C. nigra also ranked higher than ragwort in another UK study, although ragwort was still in the top 10 for yearly nectar production. The strong nectar production of certain members of the genus can be exploited to the farmer's advantage, possibly in combination with biological control. In particular, the yellow starthistle (C. solstitialis) as well as spotted knapweed (C. maculosa) are major honey plants for beekeepers. Monofloral honey from these plants is light and slightly tangy, and one of the finest honeys produced in the United States – due to its better availability, it is even fraudulently relabeled and sold as the scarce and expensive sourwood honey of the Appalachian Mountains. Placing beehives near stands of Centaurea will cause increased pollination. As most seedheads fail however when biocontrol pests have established themselves, the plants will bloom ever more abundantly in an attempt to replace the destroyed seedheads, to the point where they exhaust their resources in providing food for the pests (seeds), bees (pollen) and humans (honey). Output of allelopathic compounds is also liable to be reduced under such conditions – the plant has to compromise between allocating energy to reproduction and defense. This renders the weeds more likely to be suppressed by native vegetation or crops in the following years, especially if properly timed controlled burning[5] and/or targeted grazing by suitable livestock are also employed. While yellow starthistle and perhaps other species are toxic to equines, some other livestock may eat the non-spiny knapweeds with relish. In Europe, common knapweed (C. nigra) and globe knapweed (C. macrocephala) are locally important pollen sources for honeybees in mid-late summer.
8-Hydroxyquinoline has been identified as a main allelopathic compound produced by diffuse knapweed (C. diffusa); native North American plants are typically sensitive to it, while those of Eastern Europe and Asia Minor usually have coevolved with the knapweed and are little harmed if at all, aided by native microorganisms that break down or even feed on the abundantly secreted compound. Thus, 8-hydroxyquinoline is potentially useful to control American plants that have become invasive weeds in the diffuse knapweed's native range.
Arctiin, found in C. imperialis, has shown anticancer activity in laboratory studies. The roots of the long-lost C. foliosa, an endemic of Hatay Province (Turkey), are used in folk medicine, and other species are presumably too. A South Italian variety[verification needed] of the purple starthistle (C. calcitrapa) is traditionally consumed by ethnic Albanians (Arbëreshë people) in the Vulture area (southern Italy); e.g. in the Arbëreshë communities in Lucania the young whorls of C. calcitrapa are boiled and fried in mixtures with other weedy non-cultivated greens. According to research by the Michael Heinrich group at the Centre for Pharmacognosy and Phytotherapy (School of Pharmacy, University of London) "the antioxidant activity [...] of the young whorls of Centaurea calcitrapa, both in the DPPH and in the lipid peroxidation inhibition assays, [is] very interesting and [the] species should be investigated phytochemically and biochemically focusing on these properties". Extracts from C. calcitrapa were furthermore found to have significant xanthine oxidase (XO)-inhibiting activity.
Spotted knapweed as well as other species are rich in cnicin, a bitter compound found mainly in the leaves and often used to flavor the digestif amaro. In western Crete, Greece a local variety[verification needed] of C. calcitrapa called gourounaki (γουρουνάκι "little pig") also has its leaves eaten boiled by the locals. In the same island an endemic local species, C. idaea called katsoula (κατσούλα), tsita (τσίτα) or aspragatha (ασπραγκάθα), has its leaves eaten boiled by the locals too.
Some species are cultivated as ornamental plants in gardens. As regards other aspects of popular culture, cornflower (C. cyanus) is the floral emblem of Östergötland province (Sweden) – where is it called blåklint, literally "blue mountain" – and of Päijänne Tavastia region in Finland, where it is known as ruiskaunokki ("rye-beaks") or ruiskukka ("rye-flower"). It is also the national flower of Estonia where its local name rukkilill means "rye-lily", Belarus where it is called vałoška (Belarusian: валошка), and one of those of Germany where it is called Kornblume ("cornflower"). The origin of the name "caltrop" for the ancient low-tech area denial weapon is probably in some way connected with C. calcitrapa and its spiny seeds. This plant is attested to by the colloquial name "caltrop" at a time when the weapons were still called by their Roman name tribulus. Lastly, the color cornflower blue is named after C. cyanus. Cornflower is also used as a cut flower.
As namesake member of the subtribe Centaureinae of tribe Cardueae, the knapweeds are probably most closely related to genera such as Carthamus (distaff thistles), Cnicus (blessed thistle), Crupina (crupinas) or Notobasis (Syrian thistle), and somewhat less closely to most other thistles. The monotypic Cnicus seems in fact to properly belong in Centaurea.
Research in the late 20th century shows that Centaurea as traditionally defined is polyphyletic. A number of 19th- and 20th-century efforts to reorganize the genus were not successful, and it is not yet clear what the consequences of the recent research will be for classification of this genus and other related genera. The type species C. centaurium stands somewhat apart from the main lineage of knapweeds and thus the taxonomic consequences of a rearrangement might be severe, with hundreds of species needing to be moved to new genera. It has thus been proposed to change the type species to one of the main lineages to avoid this problem. What seems certain however is that the basketflowers – presently treated as a section Plectocephalus – will be reinstated as a distinct genus in the near future. The rock-centauries (Cheirolophus), formerly usually included in Centaurea, are now already treated as separate genus.
Better-known Centaurea species include:
Centaurea acaulis
Centaurea adpressa
Centaurea aegyptiaca
Centaurea aeolica
Centaurea aggregata
Centaurea akamantis – Akamas centaurea
Centaurea alba
Centaurea albonitens Turrill
Centaurea alpestris
Centaurea alpina
Centaurea ambigua
Centaurea amblyolepis
Centaurea americana – American basketflower, American starthistle
Centaurea ammocyanus
Centaurea antennata Dufour
Centaurea antiochia Boiss.
Centaurea aplolepa
Centaurea aplolepa subsp. carueliana
Centaurea appendicigera C.Koch
Centaurea argentea
Centaurea ascalonica
Centaurea aspera L. – rough starthistle
Centaurea atacamensis (Reiche) I.M.Johnst.
Centaurea atropurpurea
Centaurea ×aurata
Centaurea babylonica L.
Centaurea balsamita
Centaurea behen L. – ak behmen (Turkish)
Centaurea bella
Centaurea benedicta – Cnicus
Centaurea bieberseinii
Centaurea borjae
Centaurea bovina
Centaurea bracteata
Centaurea brevifimbriata Hub.-Mor.
Centaurea bulbosa
Centaurea busambarensis Guss.
Centaurea cachinalensis
Centaurea calcitrapa – purple starthistle, red starthistle, "caltrop"
Centaurea calcitrapoides
Centaurea cariensis Boiss.
Centaurea cariensiformis Hub.-Mor.
Centaurea caroli-henrici Gabrieljan & Dittrich
Centaurea centaurium L.
Centaurea chilensis
Centaurea cineraria – velvet centaurea, dusty miller
Centaurea clementei
Centaurea collina L.
Centaurea corymbosa
Centaurea crithmifolia
Centaurea crocodylium
Centaurea cyanoides J.Berggr. & Wahlenb.
Centaurea cyanus – cornflower, bachelor's button, boutonniere flower, hurtsickle, bluebottle, basketflower
Centaurea damascena
Centaurea debeauxii Gren. & Godr.
Centaurea demirizii Wagenitz
Centaurea depressa – low cornflower
Centaurea deusta
Centaurea diffusa – diffuse knapweed, white knapweed, tumble knapweed
Centaurea diluta – North African knapweed
Centaurea drabifolia Sm.
Centaurea drabifolioides Hub.-Mor.
Centaurea dschungarica
Centaurea emilae Hüseynova et Qaraxani[13]
Centaurea eriophora
Centaurea eryngioides
Centaurea filiformis
Centaurea fischeri Willd.
Centaurea floccosa
Centaurea foliosa Boiss. & Kotschy
Centaurea forojuliensis
Centaurea friderici Vis. – palagruška zečina (Croatian)
Centaurea gayana
Centaurea glaberrima Tausch
Centaurea glastifolia
Centaurea grinensis
Centaurea gymnocarpa
Centaurea haradjianii Wagenitz
Centaurea hedgei
Centaurea helenioides Boiss.
Centaurea hermannii F.Hermann
Centaurea horrida Badarò – fiordaliso spinoso (Italian)
Centaurea hyalolepis
Centaurea hypoleuca
Centaurea iberica – Iberian starthistle, Iberian knapweed
Centaurea idaea – katsoula, tsita (Cretan Greek)
Centaurea imperialis Hausskn. ex Bornm.
Centaurea jabukensis
Centaurea jacea – brown knapweed, brownray knapweed
Centaurea kasakorum
Centaurea kopetaghensis
Centaurea kotschyana Heuff.
Centaurea lanulata
Centaurea leptophylla
Centaurea leucophylla
Centaurea limbata
Centaurea lydia Boiss.
Centaurea macrocephala Puschk. ex Willd. – globe knapweed, Armenian basketflower
Centaurea maculosa – spotted knapweed (might belong in C. stoebe subsp. micranthos)
Centaurea mannagettae
Centaurea margaritalba Klok.
Centaurea marschalliana
Centaurea melitensis – Maltese starthistle; tocalote, tocolote (California)
Centaurea minor
Centaurea moschata – sweet sultan
Centaurea ×moncktonii C.E.Britton – meadow knapweed, protean knapweed (= C. ×pratensis Thuill non Salisb.)
Centaurea monocephala
Centaurea montana – montane knapweed, perennial cornflower, mountain cornflower, mountain bluet
Centaurea napifolia L. – fiordaliso romano (Italian)
Centaurea nervosa Rchb. ex Steud.
Centaurea nigra – common knapweed, black knapweed, lesser knapweed, hardheads
Centaurea nigrescens – Tyrol knapweed, short-fringed knapweed, Tyrol thistle
Centaurea nigrifimbria (C.Koch) Sosn.
Centaurea nivea (Bornm.) Wagenitz
Centaurea onopordifolia
Centaurea orientalis L.
Centaurea ornata Willd.
Centaurea ovina
Centaurea pallescens Delile
Centaurea paniculata L.
Centaurea parlatoris
Centaurea pecho
Centaurea phrygia – wig knapweed
Centaurea pindicola
Centaurea polypodiifolia
Centaurea ×pratensis Salisb. (C. jacea × C. nigra) – meadow knapweed
Centaurea procurrens
Centaurea ×psammogena G.Gayer. (C. diffusa × C. stoebe subsp. micranthos)
Centaurea pseudocaerulescens
Centaurea pseudophrygia C.A.Mey.
Centaurea pulcherrima Willd.
Centaurea pullata L.
Centaurea pumilio
Centaurea ragusina L.
Centaurea rigida
Centaurea rothrockii Greenm. – Mexican basketflower, Rothrock's basketflower, Rothrock's knapweed
Centaurea ruthenica
Centaurea rutifolia Sm.
Centaurea sadleriana – Pannonian knapweed
Centaurea salicifolia Bieb. ex Willd.
Centaurea scabiosa – greater knapweed
Centaurea scannensis
Centaurea scoparia
Centaurea scopulorum Boiss. & Heldr.
Centaurea seguenzae
Centaurea seridis L.
Centaurea sibirica
Centaurea simplicicaulis
Centaurea sinaica
Centaurea solstitialis – yellow starthistle, golden starthistle, yellow cockspur, St. Barnaby's thistle, Barnaby thistle
Centaurea speciosa
Centaurea sphaerocephala L.
Centaurea stenolepis
Centaurea stoebe L.
Centaurea stoebe subsp. micranthos (Gugler) Hayek
Centaurea straminicephala
Centaurea sulphurea – Sicilian starthistle
Centaurea tauromenitana Guss.
Centaurea tenoreana
Centaurea tommasinii
Centaurea transalpina Schleich. ex DC.
Centaurea tchihatcheffii — yanardöner (Turkish)
Centaurea trichocephala Bieb. ex Willd. – featherhead knapweed
Centaurea triniifolia
Centaurea triumfettii All.
Centaurea ucriae Lacaita
Centaurea uniflora Turra
Centaurea verbascifolia Vahl
Centaurea verutum L.
Centaurea virgata
Centaurea virgata subsp. squarrosa – squarrose knapweed
Centaurea wiedemanniana Fisch. & Mey.
Centaurea yozgatensis Wagenitz
Formerly placed here
Plant species placed in Centaurea in former times include:
Acroptilon repens – Russian knapweed (as C. repens)
Cheirolophus crassifolius – Maltese rock-centaury (as C. crassifolia, C. spathulata)
Femeniasia balearica (as C. balearica)
Volutaria muricata (as C. muricata)
A pseudanthium or flower head is a special type of inflorescence, in which anything from a small cluster to hundreds or sometimes thousands of flowers are grouped together to form a single flower-like structure.
Bellis perennis the daisy, is a European species of the family Asteraceae, often considered the archetypal species of the name daisy. To distinguish this species from other plants known as daisies, it is sometimes qualified as common daisy, lawn daisy or English daisy.
Description
Bellis perennis is a perennial herbaceous plant growing to 20 centimetres (8 inches) in height. It has short creeping rhizomes and rosettes of small rounded or spoon-shaped leaves that are from 2 to 5 cm (3⁄4–2 in) long and grow flat to the ground. The species habitually colonises lawns, and is difficult to eradicate by mowing, hence the term 'lawn daisy'. It blooms from March to September and exhibits the phenomenon of heliotropism, in which the flowers follow the position of the sun in the sky.
The flowerheads are composite, about 2 to 3 cm (3⁄4–1+1⁄4 in) in diameter, in the form of a pseudanthium, consisting of many sessile flowers with white ray florets (often tipped red) and yellow disc florets. Each inflorescence is borne on a single leafless stem 2 to 10 cm (3⁄4–4 in), rarely 15 cm (6 in) tall. The capitulum, or disc of florets, is surrounded by two rows of green bracts known as "phyllaries". The achenes are without pappus.
Etymology
Bellis may come from bellus, Latin for "pretty", and perennis is Latin for "everlasting".
The name "daisy", possibly originating with this plant, is considered a corruption of "day's eye", because the whole head closes at night and opens in the morning. Geoffrey Chaucer called it "eye of the day". In Medieval times, Bellis perennis or the English Daisy was commonly known as "Mary's Rose". It is also known as bone flower.
Historically, the plant has also been widely known as bruisewort, and occasionally woundwort (although this name is now more closely associated with the genus Stachys).
Distribution and habitat
Bellis perennis is native to western, central and northern Europe, including remote islands such as the Faroe Islands, but has become widely naturalised in most temperate regions, including the Americas and Australasia. It prefers field-like habitats.
Cultivation
The species generally blooms from early to midsummer, although when grown under ideal conditions, it has a very long flowering season and will even produce a few flowers in the middle of mild winters.
It can generally be grown where minimum temperatures are above −35 °C (−30 °F), in full sun to partial shade conditions, and requires little or no maintenance. It has no known serious insect or disease problems and can generally be grown in most well-drained soils. The plant may be propagated either by seed after the last frost, or by division after flowering.
Though not native to the United States, the species is still considered a valuable ground cover in certain garden settings (e.g., as part of English or cottage inspired gardens, as well as spring meadows where low growth and some color is desired in parallel with minimal care and maintenance while helping to crowd out noxious weeds once established and naturalised).
Numerous single- and double-flowered varieties are in cultivation, producing flat or spherical blooms in a range of sizes (1 to 6 cm or 3⁄8 to 2+3⁄8 in) and colours (red, pink and white). They are generally grown from seed as biennial bedding plants. They can also be purchased as plugs in Spring.
It has been reported to be mostly self-fertilizing, but some plants may be self-sterile.
Uses
Bellis perennis may be used as a potherb. Young leaves can be eaten raw in salads, or cooked, though the leaves become increasingly astringent with age. Flower buds and petals can be eaten raw in sandwiches, soups and salads. It is also used as a tea and as a vitamin supplement.
B. perennis has astringent properties and has been used in herbal medicine.
Daisies have traditionally been used for making daisy chains in children's games.
Culture
Daisy is used as a feminine name, and sometimes as a nickname for people named Margaret, after the French name for the oxeye daisy, marguerite.
The daisy is the national flower of the Netherlands.
The sunflower (Helianthus annuus) is an annual plant in the family Asteraceae, with a large flower head (capitulum). The stem of the flower can grow up to 3 metres tall, with a flower head that can be 30 cm wide. Other types of sunflowers include the California Royal Sunflower, which has a burgundy (red + purple) flower head.
The flower head is actually an inflorescence made of hundreds or thousands of tiny flowers called florets. The central florets look like the centre of a normal flower, and the outer florets look like yellow petals. All together they make up a "false flower" or pseudanthium. The benefit to the plant is that it is very easily seen by the insects and birds which pollinate it, and it produces thousands of seeds.
Authors: Chiara Mizzotti, Marco Fambrini, Elisabetta Caporali, Simona Masiero, and Claudio Pugliesi
Citation: Botany, 2015, 93(3): 171-181, 10.1139/cjb-2014-0210
Source: www.nrcresearchpress.com/doi/abs/10.1139/cjb-2014-0210#.U...
Fig. 1. The tubular ray flower (turf) mutant of sunflower (Helianthus annuus). (A) turf (left) and wild type (WT; right) plants. (B) A ray floret of WT. (C) A tubular-like ray floret of turf. (D) Disc florets of turf (1) and WT (2). Scale bars = 8.6 mm (B), 7.4 mm (C), and 4.8 mm (D).
Botany, NRC Research Press, a division of Canadian Science Publishing. www.nrcresearchpress.com/journal/cjb
For rights and permission, click here [bit.ly/1pL1c5f]
Centaurea is a genus of over 700 species of herbaceous thistle-like flowering plants in the family Asteraceae. Members of the genus are found only north of the equator, mostly in the Eastern Hemisphere; the Middle East and surrounding regions are particularly species-rich. In the western United States, yellow starthistles are an invasive species. Around the year 1850, seeds from the plant had arrived to the state of California. It is believed that those seeds came from South America.
Common names
Common names for this genus are centaury, centory, starthistles, knapweeds, centaureas and the more ambiguous "bluets"; a vernacular name used for these plants in parts of England is "loggerheads" (common knapweed). The Plectocephalus group – possibly a distinct genus – is known as basketflowers. "Cornflower" is used for a few species, but that term more often specifically means either C. cyanus (the annual cornflower) or Centaurea montana (the perennial cornflower). The common name "centaury" is sometimes used, although this also refers to the unrelated plant genus Centaurium.
The name is said to be in reference to Chiron, the centaur of Greek mythology who discovered medicinal uses of a plant eventually called "centaury".
Description
Knapweeds are robust weedy plants. Their leaves, spiny in some species, are usually deeply divided into elongated lobes at least in the plants' lower part, becoming entire towards the top. The "flowers" (actually pseudanthium inflorescences) are diverse in colour, ranging from intense blues, reds and yellows to any mixture of these and lighter shades towards white. Often, the disk flowers are much darker or lighter than the ray flowers, which also differ in morphology and are sterile. Each pseudanthium sits atop a cup- or basket-like cluster of scaly bracts, hence the name "basketflowers". Many species, in particular those inhabiting more arid regions, have a long and strong taproot.
Certain knapweeds have a tendency to dominate large stretches of landscape together with a few other plants, typically one or two grasses and as many other large herbaceous plants. The common knapweed (C. nigra) for example is plentiful in the mesotrophic grasslands of England and nearby regions. It is most prominently found in pastures or meadows dominated by cock's-foot (Dactylis glomerata) as well as either of crested dog's-tail (Cynosurus cristatus) and false oat-grass (Arrhenatherum elatius). It is also often found in mesotrophic grassland on rendzinas and similar calcareous soils in association with glaucous sedge (Carex flacca), sheep's fescue (Festuca ovina), and either tor-grass (Brachypodium pinnatum) and rough hawkbit (Leontodon hispidus), or upright brome (Bromus erectus). In these grasslands, greater knapweed (C. scabiosa) is found much more rarely by comparison, often in association with red fescue (Festuca rubra) in addition to cock's-foot and false oat-grass.
Due to their habit of dominating ecosystems under good conditions, many Centaurea species can become invasive weeds in regions where they are not native. In parts of North America, diffuse knapweed (C. diffusa), spotted knapweed (C. maculosa) and yellow starthistle (C. solstitialis) cause severe problems in agriculture due to their uncontrolled spread. The seeds are typically transported by human traffic, in particular the tires of all-terrain vehicles. The two knapweeds are harmful mainly because they are strongly allelopathic, producing powerful toxins in their roots that stunt the growth of plants around them not adapted to this. Yellow starthistle, meanwhile, is inedible to most livestock due to its spines and apparently outright poisonous to horses and other equines. However, efficient methods of biological control by insect pests of these weeds have been developed; the knapweeds can also exploited to their detriment by targeted grazing. Controlled burning may also be used, though the timing is important to avoid the plants having seeded already, and neither allowing sufficient time for them to regrow from the rootstock.
Yet other species of Centaurea – mostly ones that occur between Italy and the Caucasus – are endemics of a single island or valley, and some of these are endangered. The Akamas Centaurea (Centaurea akamantis) of Cyprus is almost extinct, while the western Caucasus endemics C. leptophylla and C. straminicephala are at least very rare and C. hedgei and C. pecho from the same region are certainly not abundant either. The last four species would be adversely affected by the proposed Yusufeli Dam, which might actually destroy enough habitat to push the two rarer ones over the brink of extinction.
Centaurea are copious nectar producers, especially on high-lime soils. The high nectar yield of the genus makes it very attractive to insects such as butterflies – including the endangered Karner blue (Plebejus melissa samuelis) which visits introduced spotted knapweed – and day-flying moths – typically Zygaenidae, such as Zygaena loti or the six-spot burnet (Z. filipendulae). The larvae of some other Lepidoptera species use Centaurea species as food plants; see List of Lepidoptera that feed on Centaurea. Several of these are used in biological control of invasive knapweeds and starthistles.
Larvae of several true weevils (Curculionidae) of the subfamily Lixinae also feed on Centaurea. Some genera – such as Larinus whose larval food is flowerheads – have many species especially adapted to particular knapweeds or starthistle and are used in biological control too. These include the yellow starthistle flower weevil (L. curtus) for yellow starthistle, lesser knapweed flower weevil (L. minutus) for diffuse knapweed and blunt knapweed flower weevil (L. obtusus) for spotted knapweed. Broad-nosed seedhead weevil (Bangasternus fausti) larvae eat diffuse, spotted and squarrose knapweed (C. virgata ssp. squarrosa), while those of the yellow starthistle bud weevil (B. orientalis) do not seem to live on anything other than yellow starthistle and occasionally purple starthistle (C. calcitrapa). But perhaps most efficient in destroying developing yellow starthistle seedheads is the larva of the yellow starthistle hairy weevil (Eustenopus villosus). Knapweed root weevil (Cyphocleonus achates) larvae bore into the roots of spotted and to a lesser extentely diffuse knapweed, sometimes killing off the entire plant.
Also used in biological control are Tephritidae (peacock flies) whose larvae feed on Centaurea. Knapweed peacock fly (Chaetorellia acrolophi) larvae eat spotted knapweed and some other species. The yellow starthistle peacock fly (C. australis) has an initial generation each year which often uses cornflower (C. cyanus) as larval food; later generations switch to yellow starthistle. The flies are generally considered less efficient in destroying the growing seedheads than the weevils, but may be superior under certain conditions; employing flies and weevils in combination is expensive and does not noticeably increase their effect.
Use by humans
Although the genus may be considered by a quite significant number of relatively informed individuals to have an overall negative impact on human interests, particularly agricultural interests, the situation is not straightforward enough to simply declare the genus, or, at least, its most aggressively-spreading species, altogether negative. For instance, due to their moderate to high nectar production, which can occur over a comparatively long duration, many species of Centaurea are popular food sources for insects that may otherwise attack certain crops.[citation needed] It may be advisable for some types of farms to allow certain species in this genus, such as cornflower (C. cyanus) in a European setting, to grow adjacent to fields. Although they support and attract many types of beneficial life (not just beetles), these areas are known as beetle banks. When they are present, some pests may be drawn away from crops to them and predatory insects and arachnids that feed upon pest insects will be better-supported by these more naturalized areas. They additionally have the beneficial aspect of supporting pollinators, unlike many field crops such as maize. Moreover, being untreated with pesticides and providing more diversity, plants growing in more wild areas adjacent to farms produce more insects that attract and support birds which can also feed on pests that would harm crops. Insect production is especially high for beetle banks that have enough plants that serve in the role of host plant for immature insects, rather than just in the roles of adult food and/or shelter provision.
Some plants which are considered invasive or problematic in certain areas can have beneficial qualities that outweigh their negative qualities from a human and/or human agricultural point of view, although this sometimes requires some human management – particularly if adequate biological control has not been established for the more aggressive species. An example is wild parsnip, Pastinaca sativa, which produces florets that feed predatory (and other beneficial) insects as well as large tubular stems that provide winter shelter for native bees, wasps, and other organisms that can be beneficial for agriculture. The plant is considered invasive in some areas of the United States and is also often considered undesirable due to its ability to cause contact skin irritation. However, it also serves as a host plant for the black swallowtail butterfly, helps to bring nutrients up from soils with its deep taproot, and possesses evergreen foliage even in climate zones such as US zone 6. This foliage increases soil warmth and moisture which can be beneficial for certain types of life. Perhaps the most dramatic example of a generally disliked plant's beneficial qualities being usually overlooked is the often-despised ragwort, Jacobaea vulgaris, which topped the list by a large amount for nectar production in a UK study, with a production per floral unit of (2921 ± 448μg). This very high nectar production, coupled with its early blooming period, makes the plant helpful for the establishment of bee colonies in spring — a period that is often not well-served by commercial flower meadow seed mixes. It also has the situationally-beneficial quality of being a spring ephemeral, as well as an annual that lacks difficult-to-combat roots. Plants that provide necessary structural supports for invertebrate and small vertebrate predators can help to keep overall pest populations low.
The abundant nectar produced by C. solstitialis flowers attracts many pollinators. This is another reason for the success of the (situationally) highly invasive species. Due to genetic differences related to evolutionary adaption, not all members of Centaurea produce the same amount of nectar. Growing conditions, such as climate and soil, can have a very strong impact, even if the plants grow and flower. For instance, cornflower plants, Centaurea cyanus, produced 33% less seasonal nectar than Centaurea nigra in a UK study. C. nigra also ranked higher than ragwort in another UK study, although ragwort was still in the top 10 for yearly nectar production. The strong nectar production of certain members of the genus can be exploited to the farmer's advantage, possibly in combination with biological control. In particular, the yellow starthistle (C. solstitialis) as well as spotted knapweed (C. maculosa) are major honey plants for beekeepers. Monofloral honey from these plants is light and slightly tangy, and one of the finest honeys produced in the United States – due to its better availability, it is even fraudulently relabeled and sold as the scarce and expensive sourwood honey of the Appalachian Mountains. Placing beehives near stands of Centaurea will cause increased pollination. As most seedheads fail however when biocontrol pests have established themselves, the plants will bloom ever more abundantly in an attempt to replace the destroyed seedheads, to the point where they exhaust their resources in providing food for the pests (seeds), bees (pollen) and humans (honey). Output of allelopathic compounds is also liable to be reduced under such conditions – the plant has to compromise between allocating energy to reproduction and defense. This renders the weeds more likely to be suppressed by native vegetation or crops in the following years, especially if properly timed controlled burning[5] and/or targeted grazing by suitable livestock are also employed. While yellow starthistle and perhaps other species are toxic to equines, some other livestock may eat the non-spiny knapweeds with relish. In Europe, common knapweed (C. nigra) and globe knapweed (C. macrocephala) are locally important pollen sources for honeybees in mid-late summer.
8-Hydroxyquinoline has been identified as a main allelopathic compound produced by diffuse knapweed (C. diffusa); native North American plants are typically sensitive to it, while those of Eastern Europe and Asia Minor usually have coevolved with the knapweed and are little harmed if at all, aided by native microorganisms that break down or even feed on the abundantly secreted compound. Thus, 8-hydroxyquinoline is potentially useful to control American plants that have become invasive weeds in the diffuse knapweed's native range.
Arctiin, found in C. imperialis, has shown anticancer activity in laboratory studies. The roots of the long-lost C. foliosa, an endemic of Hatay Province (Turkey), are used in folk medicine, and other species are presumably too. A South Italian variety[verification needed] of the purple starthistle (C. calcitrapa) is traditionally consumed by ethnic Albanians (Arbëreshë people) in the Vulture area (southern Italy); e.g. in the Arbëreshë communities in Lucania the young whorls of C. calcitrapa are boiled and fried in mixtures with other weedy non-cultivated greens. According to research by the Michael Heinrich group at the Centre for Pharmacognosy and Phytotherapy (School of Pharmacy, University of London) "the antioxidant activity [...] of the young whorls of Centaurea calcitrapa, both in the DPPH and in the lipid peroxidation inhibition assays, [is] very interesting and [the] species should be investigated phytochemically and biochemically focusing on these properties". Extracts from C. calcitrapa were furthermore found to have significant xanthine oxidase (XO)-inhibiting activity.
Spotted knapweed as well as other species are rich in cnicin, a bitter compound found mainly in the leaves and often used to flavor the digestif amaro. In western Crete, Greece a local variety[verification needed] of C. calcitrapa called gourounaki (γουρουνάκι "little pig") also has its leaves eaten boiled by the locals. In the same island an endemic local species, C. idaea called katsoula (κατσούλα), tsita (τσίτα) or aspragatha (ασπραγκάθα), has its leaves eaten boiled by the locals too.
Some species are cultivated as ornamental plants in gardens. As regards other aspects of popular culture, cornflower (C. cyanus) is the floral emblem of Östergötland province (Sweden) – where is it called blåklint, literally "blue mountain" – and of Päijänne Tavastia region in Finland, where it is known as ruiskaunokki ("rye-beaks") or ruiskukka ("rye-flower"). It is also the national flower of Estonia where its local name rukkilill means "rye-lily", Belarus where it is called vałoška (Belarusian: валошка), and one of those of Germany where it is called Kornblume ("cornflower"). The origin of the name "caltrop" for the ancient low-tech area denial weapon is probably in some way connected with C. calcitrapa and its spiny seeds. This plant is attested to by the colloquial name "caltrop" at a time when the weapons were still called by their Roman name tribulus. Lastly, the color cornflower blue is named after C. cyanus. Cornflower is also used as a cut flower.
As namesake member of the subtribe Centaureinae of tribe Cardueae, the knapweeds are probably most closely related to genera such as Carthamus (distaff thistles), Cnicus (blessed thistle), Crupina (crupinas) or Notobasis (Syrian thistle), and somewhat less closely to most other thistles. The monotypic Cnicus seems in fact to properly belong in Centaurea.
Research in the late 20th century shows that Centaurea as traditionally defined is polyphyletic. A number of 19th- and 20th-century efforts to reorganize the genus were not successful, and it is not yet clear what the consequences of the recent research will be for classification of this genus and other related genera. The type species C. centaurium stands somewhat apart from the main lineage of knapweeds and thus the taxonomic consequences of a rearrangement might be severe, with hundreds of species needing to be moved to new genera. It has thus been proposed to change the type species to one of the main lineages to avoid this problem. What seems certain however is that the basketflowers – presently treated as a section Plectocephalus – will be reinstated as a distinct genus in the near future. The rock-centauries (Cheirolophus), formerly usually included in Centaurea, are now already treated as separate genus.
Better-known Centaurea species include:
Centaurea acaulis
Centaurea adpressa
Centaurea aegyptiaca
Centaurea aeolica
Centaurea aggregata
Centaurea akamantis – Akamas centaurea
Centaurea alba
Centaurea albonitens Turrill
Centaurea alpestris
Centaurea alpina
Centaurea ambigua
Centaurea amblyolepis
Centaurea americana – American basketflower, American starthistle
Centaurea ammocyanus
Centaurea antennata Dufour
Centaurea antiochia Boiss.
Centaurea aplolepa
Centaurea aplolepa subsp. carueliana
Centaurea appendicigera C.Koch
Centaurea argentea
Centaurea ascalonica
Centaurea aspera L. – rough starthistle
Centaurea atacamensis (Reiche) I.M.Johnst.
Centaurea atropurpurea
Centaurea ×aurata
Centaurea babylonica L.
Centaurea balsamita
Centaurea behen L. – ak behmen (Turkish)
Centaurea bella
Centaurea benedicta – Cnicus
Centaurea bieberseinii
Centaurea borjae
Centaurea bovina
Centaurea bracteata
Centaurea brevifimbriata Hub.-Mor.
Centaurea bulbosa
Centaurea busambarensis Guss.
Centaurea cachinalensis
Centaurea calcitrapa – purple starthistle, red starthistle, "caltrop"
Centaurea calcitrapoides
Centaurea cariensis Boiss.
Centaurea cariensiformis Hub.-Mor.
Centaurea caroli-henrici Gabrieljan & Dittrich
Centaurea centaurium L.
Centaurea chilensis
Centaurea cineraria – velvet centaurea, dusty miller
Centaurea clementei
Centaurea collina L.
Centaurea corymbosa
Centaurea crithmifolia
Centaurea crocodylium
Centaurea cyanoides J.Berggr. & Wahlenb.
Centaurea cyanus – cornflower, bachelor's button, boutonniere flower, hurtsickle, bluebottle, basketflower
Centaurea damascena
Centaurea debeauxii Gren. & Godr.
Centaurea demirizii Wagenitz
Centaurea depressa – low cornflower
Centaurea deusta
Centaurea diffusa – diffuse knapweed, white knapweed, tumble knapweed
Centaurea diluta – North African knapweed
Centaurea drabifolia Sm.
Centaurea drabifolioides Hub.-Mor.
Centaurea dschungarica
Centaurea emilae Hüseynova et Qaraxani[13]
Centaurea eriophora
Centaurea eryngioides
Centaurea filiformis
Centaurea fischeri Willd.
Centaurea floccosa
Centaurea foliosa Boiss. & Kotschy
Centaurea forojuliensis
Centaurea friderici Vis. – palagruška zečina (Croatian)
Centaurea gayana
Centaurea glaberrima Tausch
Centaurea glastifolia
Centaurea grinensis
Centaurea gymnocarpa
Centaurea haradjianii Wagenitz
Centaurea hedgei
Centaurea helenioides Boiss.
Centaurea hermannii F.Hermann
Centaurea horrida Badarò – fiordaliso spinoso (Italian)
Centaurea hyalolepis
Centaurea hypoleuca
Centaurea iberica – Iberian starthistle, Iberian knapweed
Centaurea idaea – katsoula, tsita (Cretan Greek)
Centaurea imperialis Hausskn. ex Bornm.
Centaurea jabukensis
Centaurea jacea – brown knapweed, brownray knapweed
Centaurea kasakorum
Centaurea kopetaghensis
Centaurea kotschyana Heuff.
Centaurea lanulata
Centaurea leptophylla
Centaurea leucophylla
Centaurea limbata
Centaurea lydia Boiss.
Centaurea macrocephala Puschk. ex Willd. – globe knapweed, Armenian basketflower
Centaurea maculosa – spotted knapweed (might belong in C. stoebe subsp. micranthos)
Centaurea mannagettae
Centaurea margaritalba Klok.
Centaurea marschalliana
Centaurea melitensis – Maltese starthistle; tocalote, tocolote (California)
Centaurea minor
Centaurea moschata – sweet sultan
Centaurea ×moncktonii C.E.Britton – meadow knapweed, protean knapweed (= C. ×pratensis Thuill non Salisb.)
Centaurea monocephala
Centaurea montana – montane knapweed, perennial cornflower, mountain cornflower, mountain bluet
Centaurea napifolia L. – fiordaliso romano (Italian)
Centaurea nervosa Rchb. ex Steud.
Centaurea nigra – common knapweed, black knapweed, lesser knapweed, hardheads
Centaurea nigrescens – Tyrol knapweed, short-fringed knapweed, Tyrol thistle
Centaurea nigrifimbria (C.Koch) Sosn.
Centaurea nivea (Bornm.) Wagenitz
Centaurea onopordifolia
Centaurea orientalis L.
Centaurea ornata Willd.
Centaurea ovina
Centaurea pallescens Delile
Centaurea paniculata L.
Centaurea parlatoris
Centaurea pecho
Centaurea phrygia – wig knapweed
Centaurea pindicola
Centaurea polypodiifolia
Centaurea ×pratensis Salisb. (C. jacea × C. nigra) – meadow knapweed
Centaurea procurrens
Centaurea ×psammogena G.Gayer. (C. diffusa × C. stoebe subsp. micranthos)
Centaurea pseudocaerulescens
Centaurea pseudophrygia C.A.Mey.
Centaurea pulcherrima Willd.
Centaurea pullata L.
Centaurea pumilio
Centaurea ragusina L.
Centaurea rigida
Centaurea rothrockii Greenm. – Mexican basketflower, Rothrock's basketflower, Rothrock's knapweed
Centaurea ruthenica
Centaurea rutifolia Sm.
Centaurea sadleriana – Pannonian knapweed
Centaurea salicifolia Bieb. ex Willd.
Centaurea scabiosa – greater knapweed
Centaurea scannensis
Centaurea scoparia
Centaurea scopulorum Boiss. & Heldr.
Centaurea seguenzae
Centaurea seridis L.
Centaurea sibirica
Centaurea simplicicaulis
Centaurea sinaica
Centaurea solstitialis – yellow starthistle, golden starthistle, yellow cockspur, St. Barnaby's thistle, Barnaby thistle
Centaurea speciosa
Centaurea sphaerocephala L.
Centaurea stenolepis
Centaurea stoebe L.
Centaurea stoebe subsp. micranthos (Gugler) Hayek
Centaurea straminicephala
Centaurea sulphurea – Sicilian starthistle
Centaurea tauromenitana Guss.
Centaurea tenoreana
Centaurea tommasinii
Centaurea transalpina Schleich. ex DC.
Centaurea tchihatcheffii — yanardöner (Turkish)
Centaurea trichocephala Bieb. ex Willd. – featherhead knapweed
Centaurea triniifolia
Centaurea triumfettii All.
Centaurea ucriae Lacaita
Centaurea uniflora Turra
Centaurea verbascifolia Vahl
Centaurea verutum L.
Centaurea virgata
Centaurea virgata subsp. squarrosa – squarrose knapweed
Centaurea wiedemanniana Fisch. & Mey.
Centaurea yozgatensis Wagenitz
Formerly placed here
Plant species placed in Centaurea in former times include:
Acroptilon repens – Russian knapweed (as C. repens)
Cheirolophus crassifolius – Maltese rock-centaury (as C. crassifolia, C. spathulata)
Femeniasia balearica (as C. balearica)
Volutaria muricata (as C. muricata)
Common sunflower (Helianthus annuus)
Wild Helianthus annuus is a widely branched annual plant with many flower heads. The domestic sunflower, however, often possesses only a single large inflorescence (flower head) atop an unbranched stem.
The term Helianthus annuus is derived from the Greek Helios, meaning sun, and anthos, meaning flower, and from Latin annuus, meaning annual.
What is often called the "flower" of the sunflower is actually a "flower head" (pseudanthium) of numerous small individual five-petaled flowers ("florets"). The outer flowers, which resemble petals, are called ray flowers. Each "petal" consists of a ligule composed of fused petals of an asymmetrical ray flower. They are sexually sterile and may be yellow, red, orange, or other colors. The flowers in the center of the head are called disk flowers. These mature into fruit (sunflower "seeds"). The disk flowers are arranged spirally.
A common misconception is that flowering sunflower heads track the Sun across the sky. Although immature flower buds exhibit this behaviour, the mature flowering heads point in a fixed (and typically easterly) direction throughout the day.
When growth of the flower stalk stops and the flower is mature, the heliotropism also stops and the flower faces east from that moment onward. This eastward orientation allows rapid warming in the morning and, as a result, an increase in pollinator visits.
Authors: Chiara Mizzotti, Marco Fambrini, Elisabetta Caporali, Simona Masiero, and Claudio Pugliesi
Citation: Botany, 2015, 93(3): 171-181, 10.1139/cjb-2014-0210
Source: www.nrcresearchpress.com/doi/abs/10.1139/cjb-2014-0210#.U...
Fig. 2. Scanning electron micrographs of sunflower floret primordia at various developmental stages in wild type (WT) (A–D, I–L) and tubular ray flower (turf) mutant (E–H, M–P). (A) Stage 1 of WT ray floret buds (white arrows); at the periphery of the inflorescence, note stage 2 of disc floret primordia (black arrowheads). WT ray floret primordium development at stage 3 (B), stage 5 (C), and stage 6 (D). (E) Stage 1 of turf tubular-like ray floret buds (white arrows). turf tubular-like ray floret bud at stage 3 (F), stage 5 (G), and stage 6 (H). (I–L) Progression of WT disc floret primordia development from stage 1 to stage 4. (M–P) Progression of turf disc floret primordia development from stage 1 to stage 5. The stages of floret development are numbered according to Tähtiharju et al. (2012) (see Supplementary Table S21). Scale bars = 100 µm (A–E, G–P) and 50 µm (F).
Botany, NRC Research Press, a division of Canadian Science Publishing. www.nrcresearchpress.com/journal/cjb
For rights and permission, click here [bit.ly/1pL1c5f]
Centaurea is a genus of over 700 species of herbaceous thistle-like flowering plants in the family Asteraceae. Members of the genus are found only north of the equator, mostly in the Eastern Hemisphere; the Middle East and surrounding regions are particularly species-rich. In the western United States, yellow starthistles are an invasive species. Around the year 1850, seeds from the plant had arrived to the state of California. It is believed that those seeds came from South America.
Common names
Common names for this genus are centaury, centory, starthistles, knapweeds, centaureas and the more ambiguous "bluets"; a vernacular name used for these plants in parts of England is "loggerheads" (common knapweed). The Plectocephalus group – possibly a distinct genus – is known as basketflowers. "Cornflower" is used for a few species, but that term more often specifically means either C. cyanus (the annual cornflower) or Centaurea montana (the perennial cornflower). The common name "centaury" is sometimes used, although this also refers to the unrelated plant genus Centaurium.
The name is said to be in reference to Chiron, the centaur of Greek mythology who discovered medicinal uses of a plant eventually called "centaury".
Description
Knapweeds are robust weedy plants. Their leaves, spiny in some species, are usually deeply divided into elongated lobes at least in the plants' lower part, becoming entire towards the top. The "flowers" (actually pseudanthium inflorescences) are diverse in colour, ranging from intense blues, reds and yellows to any mixture of these and lighter shades towards white. Often, the disk flowers are much darker or lighter than the ray flowers, which also differ in morphology and are sterile. Each pseudanthium sits atop a cup- or basket-like cluster of scaly bracts, hence the name "basketflowers". Many species, in particular those inhabiting more arid regions, have a long and strong taproot.
Certain knapweeds have a tendency to dominate large stretches of landscape together with a few other plants, typically one or two grasses and as many other large herbaceous plants. The common knapweed (C. nigra) for example is plentiful in the mesotrophic grasslands of England and nearby regions. It is most prominently found in pastures or meadows dominated by cock's-foot (Dactylis glomerata) as well as either of crested dog's-tail (Cynosurus cristatus) and false oat-grass (Arrhenatherum elatius). It is also often found in mesotrophic grassland on rendzinas and similar calcareous soils in association with glaucous sedge (Carex flacca), sheep's fescue (Festuca ovina), and either tor-grass (Brachypodium pinnatum) and rough hawkbit (Leontodon hispidus), or upright brome (Bromus erectus). In these grasslands, greater knapweed (C. scabiosa) is found much more rarely by comparison, often in association with red fescue (Festuca rubra) in addition to cock's-foot and false oat-grass.
Due to their habit of dominating ecosystems under good conditions, many Centaurea species can become invasive weeds in regions where they are not native. In parts of North America, diffuse knapweed (C. diffusa), spotted knapweed (C. maculosa) and yellow starthistle (C. solstitialis) cause severe problems in agriculture due to their uncontrolled spread. The seeds are typically transported by human traffic, in particular the tires of all-terrain vehicles. The two knapweeds are harmful mainly because they are strongly allelopathic, producing powerful toxins in their roots that stunt the growth of plants around them not adapted to this. Yellow starthistle, meanwhile, is inedible to most livestock due to its spines and apparently outright poisonous to horses and other equines. However, efficient methods of biological control by insect pests of these weeds have been developed; the knapweeds can also exploited to their detriment by targeted grazing. Controlled burning may also be used, though the timing is important to avoid the plants having seeded already, and neither allowing sufficient time for them to regrow from the rootstock.
Yet other species of Centaurea – mostly ones that occur between Italy and the Caucasus – are endemics of a single island or valley, and some of these are endangered. The Akamas Centaurea (Centaurea akamantis) of Cyprus is almost extinct, while the western Caucasus endemics C. leptophylla and C. straminicephala are at least very rare and C. hedgei and C. pecho from the same region are certainly not abundant either. The last four species would be adversely affected by the proposed Yusufeli Dam, which might actually destroy enough habitat to push the two rarer ones over the brink of extinction.
Centaurea are copious nectar producers, especially on high-lime soils. The high nectar yield of the genus makes it very attractive to insects such as butterflies – including the endangered Karner blue (Plebejus melissa samuelis) which visits introduced spotted knapweed – and day-flying moths – typically Zygaenidae, such as Zygaena loti or the six-spot burnet (Z. filipendulae). The larvae of some other Lepidoptera species use Centaurea species as food plants; see List of Lepidoptera that feed on Centaurea. Several of these are used in biological control of invasive knapweeds and starthistles.
Larvae of several true weevils (Curculionidae) of the subfamily Lixinae also feed on Centaurea. Some genera – such as Larinus whose larval food is flowerheads – have many species especially adapted to particular knapweeds or starthistle and are used in biological control too. These include the yellow starthistle flower weevil (L. curtus) for yellow starthistle, lesser knapweed flower weevil (L. minutus) for diffuse knapweed and blunt knapweed flower weevil (L. obtusus) for spotted knapweed. Broad-nosed seedhead weevil (Bangasternus fausti) larvae eat diffuse, spotted and squarrose knapweed (C. virgata ssp. squarrosa), while those of the yellow starthistle bud weevil (B. orientalis) do not seem to live on anything other than yellow starthistle and occasionally purple starthistle (C. calcitrapa). But perhaps most efficient in destroying developing yellow starthistle seedheads is the larva of the yellow starthistle hairy weevil (Eustenopus villosus). Knapweed root weevil (Cyphocleonus achates) larvae bore into the roots of spotted and to a lesser extentely diffuse knapweed, sometimes killing off the entire plant.
Also used in biological control are Tephritidae (peacock flies) whose larvae feed on Centaurea. Knapweed peacock fly (Chaetorellia acrolophi) larvae eat spotted knapweed and some other species. The yellow starthistle peacock fly (C. australis) has an initial generation each year which often uses cornflower (C. cyanus) as larval food; later generations switch to yellow starthistle. The flies are generally considered less efficient in destroying the growing seedheads than the weevils, but may be superior under certain conditions; employing flies and weevils in combination is expensive and does not noticeably increase their effect.
Use by humans
Although the genus may be considered by a quite significant number of relatively informed individuals to have an overall negative impact on human interests, particularly agricultural interests, the situation is not straightforward enough to simply declare the genus, or, at least, its most aggressively-spreading species, altogether negative. For instance, due to their moderate to high nectar production, which can occur over a comparatively long duration, many species of Centaurea are popular food sources for insects that may otherwise attack certain crops.[citation needed] It may be advisable for some types of farms to allow certain species in this genus, such as cornflower (C. cyanus) in a European setting, to grow adjacent to fields. Although they support and attract many types of beneficial life (not just beetles), these areas are known as beetle banks. When they are present, some pests may be drawn away from crops to them and predatory insects and arachnids that feed upon pest insects will be better-supported by these more naturalized areas. They additionally have the beneficial aspect of supporting pollinators, unlike many field crops such as maize. Moreover, being untreated with pesticides and providing more diversity, plants growing in more wild areas adjacent to farms produce more insects that attract and support birds which can also feed on pests that would harm crops. Insect production is especially high for beetle banks that have enough plants that serve in the role of host plant for immature insects, rather than just in the roles of adult food and/or shelter provision.
Some plants which are considered invasive or problematic in certain areas can have beneficial qualities that outweigh their negative qualities from a human and/or human agricultural point of view, although this sometimes requires some human management – particularly if adequate biological control has not been established for the more aggressive species. An example is wild parsnip, Pastinaca sativa, which produces florets that feed predatory (and other beneficial) insects as well as large tubular stems that provide winter shelter for native bees, wasps, and other organisms that can be beneficial for agriculture. The plant is considered invasive in some areas of the United States and is also often considered undesirable due to its ability to cause contact skin irritation. However, it also serves as a host plant for the black swallowtail butterfly, helps to bring nutrients up from soils with its deep taproot, and possesses evergreen foliage even in climate zones such as US zone 6. This foliage increases soil warmth and moisture which can be beneficial for certain types of life. Perhaps the most dramatic example of a generally disliked plant's beneficial qualities being usually overlooked is the often-despised ragwort, Jacobaea vulgaris, which topped the list by a large amount for nectar production in a UK study, with a production per floral unit of (2921 ± 448μg). This very high nectar production, coupled with its early blooming period, makes the plant helpful for the establishment of bee colonies in spring — a period that is often not well-served by commercial flower meadow seed mixes. It also has the situationally-beneficial quality of being a spring ephemeral, as well as an annual that lacks difficult-to-combat roots. Plants that provide necessary structural supports for invertebrate and small vertebrate predators can help to keep overall pest populations low.
The abundant nectar produced by C. solstitialis flowers attracts many pollinators. This is another reason for the success of the (situationally) highly invasive species. Due to genetic differences related to evolutionary adaption, not all members of Centaurea produce the same amount of nectar. Growing conditions, such as climate and soil, can have a very strong impact, even if the plants grow and flower. For instance, cornflower plants, Centaurea cyanus, produced 33% less seasonal nectar than Centaurea nigra in a UK study. C. nigra also ranked higher than ragwort in another UK study, although ragwort was still in the top 10 for yearly nectar production. The strong nectar production of certain members of the genus can be exploited to the farmer's advantage, possibly in combination with biological control. In particular, the yellow starthistle (C. solstitialis) as well as spotted knapweed (C. maculosa) are major honey plants for beekeepers. Monofloral honey from these plants is light and slightly tangy, and one of the finest honeys produced in the United States – due to its better availability, it is even fraudulently relabeled and sold as the scarce and expensive sourwood honey of the Appalachian Mountains. Placing beehives near stands of Centaurea will cause increased pollination. As most seedheads fail however when biocontrol pests have established themselves, the plants will bloom ever more abundantly in an attempt to replace the destroyed seedheads, to the point where they exhaust their resources in providing food for the pests (seeds), bees (pollen) and humans (honey). Output of allelopathic compounds is also liable to be reduced under such conditions – the plant has to compromise between allocating energy to reproduction and defense. This renders the weeds more likely to be suppressed by native vegetation or crops in the following years, especially if properly timed controlled burning[5] and/or targeted grazing by suitable livestock are also employed. While yellow starthistle and perhaps other species are toxic to equines, some other livestock may eat the non-spiny knapweeds with relish. In Europe, common knapweed (C. nigra) and globe knapweed (C. macrocephala) are locally important pollen sources for honeybees in mid-late summer.
8-Hydroxyquinoline has been identified as a main allelopathic compound produced by diffuse knapweed (C. diffusa); native North American plants are typically sensitive to it, while those of Eastern Europe and Asia Minor usually have coevolved with the knapweed and are little harmed if at all, aided by native microorganisms that break down or even feed on the abundantly secreted compound. Thus, 8-hydroxyquinoline is potentially useful to control American plants that have become invasive weeds in the diffuse knapweed's native range.
Arctiin, found in C. imperialis, has shown anticancer activity in laboratory studies. The roots of the long-lost C. foliosa, an endemic of Hatay Province (Turkey), are used in folk medicine, and other species are presumably too. A South Italian variety[verification needed] of the purple starthistle (C. calcitrapa) is traditionally consumed by ethnic Albanians (Arbëreshë people) in the Vulture area (southern Italy); e.g. in the Arbëreshë communities in Lucania the young whorls of C. calcitrapa are boiled and fried in mixtures with other weedy non-cultivated greens. According to research by the Michael Heinrich group at the Centre for Pharmacognosy and Phytotherapy (School of Pharmacy, University of London) "the antioxidant activity [...] of the young whorls of Centaurea calcitrapa, both in the DPPH and in the lipid peroxidation inhibition assays, [is] very interesting and [the] species should be investigated phytochemically and biochemically focusing on these properties". Extracts from C. calcitrapa were furthermore found to have significant xanthine oxidase (XO)-inhibiting activity.
Spotted knapweed as well as other species are rich in cnicin, a bitter compound found mainly in the leaves and often used to flavor the digestif amaro. In western Crete, Greece a local variety[verification needed] of C. calcitrapa called gourounaki (γουρουνάκι "little pig") also has its leaves eaten boiled by the locals. In the same island an endemic local species, C. idaea called katsoula (κατσούλα), tsita (τσίτα) or aspragatha (ασπραγκάθα), has its leaves eaten boiled by the locals too.
Some species are cultivated as ornamental plants in gardens. As regards other aspects of popular culture, cornflower (C. cyanus) is the floral emblem of Östergötland province (Sweden) – where is it called blåklint, literally "blue mountain" – and of Päijänne Tavastia region in Finland, where it is known as ruiskaunokki ("rye-beaks") or ruiskukka ("rye-flower"). It is also the national flower of Estonia where its local name rukkilill means "rye-lily", Belarus where it is called vałoška (Belarusian: валошка), and one of those of Germany where it is called Kornblume ("cornflower"). The origin of the name "caltrop" for the ancient low-tech area denial weapon is probably in some way connected with C. calcitrapa and its spiny seeds. This plant is attested to by the colloquial name "caltrop" at a time when the weapons were still called by their Roman name tribulus. Lastly, the color cornflower blue is named after C. cyanus. Cornflower is also used as a cut flower.
As namesake member of the subtribe Centaureinae of tribe Cardueae, the knapweeds are probably most closely related to genera such as Carthamus (distaff thistles), Cnicus (blessed thistle), Crupina (crupinas) or Notobasis (Syrian thistle), and somewhat less closely to most other thistles. The monotypic Cnicus seems in fact to properly belong in Centaurea.
Research in the late 20th century shows that Centaurea as traditionally defined is polyphyletic. A number of 19th- and 20th-century efforts to reorganize the genus were not successful, and it is not yet clear what the consequences of the recent research will be for classification of this genus and other related genera. The type species C. centaurium stands somewhat apart from the main lineage of knapweeds and thus the taxonomic consequences of a rearrangement might be severe, with hundreds of species needing to be moved to new genera. It has thus been proposed to change the type species to one of the main lineages to avoid this problem. What seems certain however is that the basketflowers – presently treated as a section Plectocephalus – will be reinstated as a distinct genus in the near future. The rock-centauries (Cheirolophus), formerly usually included in Centaurea, are now already treated as separate genus.
Better-known Centaurea species include:
Centaurea acaulis
Centaurea adpressa
Centaurea aegyptiaca
Centaurea aeolica
Centaurea aggregata
Centaurea akamantis – Akamas centaurea
Centaurea alba
Centaurea albonitens Turrill
Centaurea alpestris
Centaurea alpina
Centaurea ambigua
Centaurea amblyolepis
Centaurea americana – American basketflower, American starthistle
Centaurea ammocyanus
Centaurea antennata Dufour
Centaurea antiochia Boiss.
Centaurea aplolepa
Centaurea aplolepa subsp. carueliana
Centaurea appendicigera C.Koch
Centaurea argentea
Centaurea ascalonica
Centaurea aspera L. – rough starthistle
Centaurea atacamensis (Reiche) I.M.Johnst.
Centaurea atropurpurea
Centaurea ×aurata
Centaurea babylonica L.
Centaurea balsamita
Centaurea behen L. – ak behmen (Turkish)
Centaurea bella
Centaurea benedicta – Cnicus
Centaurea bieberseinii
Centaurea borjae
Centaurea bovina
Centaurea bracteata
Centaurea brevifimbriata Hub.-Mor.
Centaurea bulbosa
Centaurea busambarensis Guss.
Centaurea cachinalensis
Centaurea calcitrapa – purple starthistle, red starthistle, "caltrop"
Centaurea calcitrapoides
Centaurea cariensis Boiss.
Centaurea cariensiformis Hub.-Mor.
Centaurea caroli-henrici Gabrieljan & Dittrich
Centaurea centaurium L.
Centaurea chilensis
Centaurea cineraria – velvet centaurea, dusty miller
Centaurea clementei
Centaurea collina L.
Centaurea corymbosa
Centaurea crithmifolia
Centaurea crocodylium
Centaurea cyanoides J.Berggr. & Wahlenb.
Centaurea cyanus – cornflower, bachelor's button, boutonniere flower, hurtsickle, bluebottle, basketflower
Centaurea damascena
Centaurea debeauxii Gren. & Godr.
Centaurea demirizii Wagenitz
Centaurea depressa – low cornflower
Centaurea deusta
Centaurea diffusa – diffuse knapweed, white knapweed, tumble knapweed
Centaurea diluta – North African knapweed
Centaurea drabifolia Sm.
Centaurea drabifolioides Hub.-Mor.
Centaurea dschungarica
Centaurea emilae Hüseynova et Qaraxani[13]
Centaurea eriophora
Centaurea eryngioides
Centaurea filiformis
Centaurea fischeri Willd.
Centaurea floccosa
Centaurea foliosa Boiss. & Kotschy
Centaurea forojuliensis
Centaurea friderici Vis. – palagruška zečina (Croatian)
Centaurea gayana
Centaurea glaberrima Tausch
Centaurea glastifolia
Centaurea grinensis
Centaurea gymnocarpa
Centaurea haradjianii Wagenitz
Centaurea hedgei
Centaurea helenioides Boiss.
Centaurea hermannii F.Hermann
Centaurea horrida Badarò – fiordaliso spinoso (Italian)
Centaurea hyalolepis
Centaurea hypoleuca
Centaurea iberica – Iberian starthistle, Iberian knapweed
Centaurea idaea – katsoula, tsita (Cretan Greek)
Centaurea imperialis Hausskn. ex Bornm.
Centaurea jabukensis
Centaurea jacea – brown knapweed, brownray knapweed
Centaurea kasakorum
Centaurea kopetaghensis
Centaurea kotschyana Heuff.
Centaurea lanulata
Centaurea leptophylla
Centaurea leucophylla
Centaurea limbata
Centaurea lydia Boiss.
Centaurea macrocephala Puschk. ex Willd. – globe knapweed, Armenian basketflower
Centaurea maculosa – spotted knapweed (might belong in C. stoebe subsp. micranthos)
Centaurea mannagettae
Centaurea margaritalba Klok.
Centaurea marschalliana
Centaurea melitensis – Maltese starthistle; tocalote, tocolote (California)
Centaurea minor
Centaurea moschata – sweet sultan
Centaurea ×moncktonii C.E.Britton – meadow knapweed, protean knapweed (= C. ×pratensis Thuill non Salisb.)
Centaurea monocephala
Centaurea montana – montane knapweed, perennial cornflower, mountain cornflower, mountain bluet
Centaurea napifolia L. – fiordaliso romano (Italian)
Centaurea nervosa Rchb. ex Steud.
Centaurea nigra – common knapweed, black knapweed, lesser knapweed, hardheads
Centaurea nigrescens – Tyrol knapweed, short-fringed knapweed, Tyrol thistle
Centaurea nigrifimbria (C.Koch) Sosn.
Centaurea nivea (Bornm.) Wagenitz
Centaurea onopordifolia
Centaurea orientalis L.
Centaurea ornata Willd.
Centaurea ovina
Centaurea pallescens Delile
Centaurea paniculata L.
Centaurea parlatoris
Centaurea pecho
Centaurea phrygia – wig knapweed
Centaurea pindicola
Centaurea polypodiifolia
Centaurea ×pratensis Salisb. (C. jacea × C. nigra) – meadow knapweed
Centaurea procurrens
Centaurea ×psammogena G.Gayer. (C. diffusa × C. stoebe subsp. micranthos)
Centaurea pseudocaerulescens
Centaurea pseudophrygia C.A.Mey.
Centaurea pulcherrima Willd.
Centaurea pullata L.
Centaurea pumilio
Centaurea ragusina L.
Centaurea rigida
Centaurea rothrockii Greenm. – Mexican basketflower, Rothrock's basketflower, Rothrock's knapweed
Centaurea ruthenica
Centaurea rutifolia Sm.
Centaurea sadleriana – Pannonian knapweed
Centaurea salicifolia Bieb. ex Willd.
Centaurea scabiosa – greater knapweed
Centaurea scannensis
Centaurea scoparia
Centaurea scopulorum Boiss. & Heldr.
Centaurea seguenzae
Centaurea seridis L.
Centaurea sibirica
Centaurea simplicicaulis
Centaurea sinaica
Centaurea solstitialis – yellow starthistle, golden starthistle, yellow cockspur, St. Barnaby's thistle, Barnaby thistle
Centaurea speciosa
Centaurea sphaerocephala L.
Centaurea stenolepis
Centaurea stoebe L.
Centaurea stoebe subsp. micranthos (Gugler) Hayek
Centaurea straminicephala
Centaurea sulphurea – Sicilian starthistle
Centaurea tauromenitana Guss.
Centaurea tenoreana
Centaurea tommasinii
Centaurea transalpina Schleich. ex DC.
Centaurea tchihatcheffii — yanardöner (Turkish)
Centaurea trichocephala Bieb. ex Willd. – featherhead knapweed
Centaurea triniifolia
Centaurea triumfettii All.
Centaurea ucriae Lacaita
Centaurea uniflora Turra
Centaurea verbascifolia Vahl
Centaurea verutum L.
Centaurea virgata
Centaurea virgata subsp. squarrosa – squarrose knapweed
Centaurea wiedemanniana Fisch. & Mey.
Centaurea yozgatensis Wagenitz
Formerly placed here
Plant species placed in Centaurea in former times include:
Acroptilon repens – Russian knapweed (as C. repens)
Cheirolophus crassifolius – Maltese rock-centaury (as C. crassifolia, C. spathulata)
Femeniasia balearica (as C. balearica)
Volutaria muricata (as C. muricata)
Euphorbia is a very large and diverse genus of flowering plants, commonly called spurge, in the family Euphorbiaceae. "Euphorbia" is sometimes used in ordinary English to collectively refer to all members of Euphorbiaceae (in deference to the type genus), not just to members of the genus.
Euphorbias range from tiny annual plants to large and long-lived trees. with perhaps the tallest being Euphorbia ampliphylla at 30 m (98 ft) or more. The genus has roughly 2,000 members, making it one of the largest genera of flowering plants. It also has one of the largest ranges of chromosome counts, along with Rumex and Senecio. Euphorbia antiquorum is the type species for the genus Euphorbia. It was first described by Carl Linnaeus in 1753 in Species Plantarum.
Some euphorbias are widely available commercially, such as poinsettias at Christmas. Some are commonly cultivated as ornamentals, or collected and highly valued for the aesthetic appearance of their unique floral structures, such as the crown of thorns plant (Euphorbia milii). Succulent euphorbias from the deserts of Southern Africa and Madagascar have evolved physical characteristics and forms similar to cacti of North and South America, so they are often incorrectly referred to as cacti. Some are used as ornamentals in landscaping, because of beautiful or striking overall forms, and drought and heat tolerance.
Euphorbia all share the feature of having a poisonous, latex-like sap and unique floral structures When viewed as a whole, the head of flowers looks like a single flower (a pseudanthium). It has a unique kind of pseudanthium, called a cyathium, where each flower in the head is reduced to its barest essential part needed for sexual reproduction. The individual flowers are either male or female, with the male flowers reduced to only the stamen, and the females to the pistil. These flowers have no sepals, petals, or other parts that are typical of flowers in other kinds of plants. Structures supporting the flower head and other structures underneath have evolved to attract pollinators with nectar, and with shapes and colors that function in a way petals and other flower parts do in other flowers. It is the only genus of plants that has all three kinds of photosynthesis, CAM, C3 and C4.[7]
Etymology
The common name "spurge" derives from the Middle English/Old French espurge ("to purge"), due to the use of the plant's sap as a purgative. The botanical name Euphorbia derives from Euphorbos, the Greek physician of King Juba II of Numidia and Mauretania (52–50 BC – 23 AD), who married the daughter of Anthony and Cleopatra. Juba was a prolific writer on various subjects, including natural history. Euphorbos wrote that one of the cactus-like euphorbias (now called Euphorbia obtusifolia ssp. regis-jubae) was used as a powerful laxative. In 12 BC, Juba named this plant after his physician Euphorbos, as Augustus Caesar had dedicated a statue to the brother of Euphorbos, Antonius Musa, who was the personal physician of Augustus. In 1753, botanist and taxonomist Carl Linnaeus assigned the name Euphorbia to the entire genus in the physician's honor.
Description
The plants are annual, biennial or perennial herbs, woody shrubs, or trees with a caustic, poisonous milky latex. The roots are fine or thick and fleshy or tuberous. Many species are more or less succulent, thorny, or unarmed. The main stem and mostly also the side arms of the succulent species are thick and fleshy, and often winged, 15–91 cm (6–36 in) tall. The succulent trees and large shrubs are mostly confined to southern and eastern Africa, but Euphorbia neutra is native to the Caatinga of Brazil and Euphorbia royleana is found in the Himalayan foothills.[14] The deciduous[citation needed] leaves may be opposite, alternate, or in whorls. In succulent species, the leaves are mostly small and short-lived. The stipules are mostly small, partly transformed into spines or glands, or missing.
Inflorescence and fruit
Like all members of the family Euphorbiaceae, spurges have unisexual flowers.
In Euphorbia, flowers occur in a head, called the cyathium (plural cyathia). Each male or female flower in the cyathium head has only its essential sexual part, in males the stamen, and in females the pistil. The flowers do not have sepals, petals, or nectar to attract pollinators, although other nonflower parts of the plant have an appearance and nectar glands with similar roles. Euphorbias are the only plants known to have this kind of flower head.
Nectar glands and nectar that attract pollinators are held in the involucre, a cup-like part below and supporting the cyathium head. The "involucre" in the genus Euphorbia is not to be confused with the "involucre" in family Asteraceae members, which is a collection of bracts called phyllaries, which surround and encase the unopened flower head, then support the receptacle under it after the flower head opens.
The involucre is above and supported by bract-like modified leaf structures (usually in pairs)[citation needed] called cyathophylls', or cyathial leaves. The cyathophyll often has a superficial appearance of being petals of a flower.
Euphorbia flowers are tiny, and the variation attracting different pollinators, with different forms and colors occurs, in the cyathium, involucre, cyathophyll, or additional parts such as glands that attached to these.
The collection of many flowers may be shaped and arranged to appear collectively as a single individual flower, sometimes called a pseudanthium in the Asteraceae, and also in Euphorbia.
The majority of species are monoecious (bearing male and female flowers on the same plant), although some are dioecious with male and female flowers occurring on different plants. It is not unusual for the central cyathia of a cyme to be purely male, and for lateral cyathia to carry both sexes. Sometimes, young plants or those growing under unfavorable conditions are male only, and only produce female flowers in the cyathia with maturity or as growing conditions improve.
The female flowers reduced to a single pistil usually split into three parts, often with two stigmas at each tip.[citation needed] Male flowers often have anthers in twos.[citation needed] Nectar glands usually occur in fives, may be as few as one,[16] and may be fused into a "U" shape. The cyathophylls often occur in twos, are leaf-like, and may be showy and brightly coloured and attractive to pollinators, or be reduced to barely visible tiny scales.
The fruits are three- or rarely two-compartment capsules, sometimes fleshy, but almost always ripening to a woody container that then splits open, sometimes explosively. The seeds are four-angled, oval, or spherical, and some species have a caruncle.
Xerophytes and succulents
In the genus Euphorbia, succulence in the species has often evolved divergently and to differing degrees. Sometimes, it is difficult to decide, and is a question of interpretation, whether or not a species is really succulent or "only" xerophytic. In some cases, especially with geophytes, plants closely related to the succulents are normal herbs. About 850 species are succulent in the strictest sense. If one includes slightly succulent and xerophytic species, this figure rises to about 1000, representing about 45% of all Euphorbia species.
Irritants
The milky sap of spurges (called "latex") evolved as a deterrent to herbivores. It is white, and transparent when dry, except in E. abdelkuri, where it is yellow. The pressurized sap seeps from the slightest wound and congeals after a few minutes in air. The skin-irritating and caustic effects are largely caused by varying amounts of diterpenes. Triterpenes such as betulin and corresponding esters are other major components of the latex. In contact with mucous membranes (eyes, nose, mouth), the latex can produce extremely painful inflammation. The sap has also been known to cause mild to extreme Keratouveitis, which affects vision. Therefore, spurges should be handled with caution and kept away from children and pets. Wearing eye protection while working in close contact with Euphorbia is advised.[18] Latex on skin should be washed off immediately and thoroughly. Congealed latex is insoluble in water, but can be removed with an emulsifier such as milk or soap. A physician should be consulted if inflammation occurs, as severe eye damage including permanent blindness may result from exposure to the sap.
The poisonous qualities were well known: in the Ethiopian Kebra Nagast, the serpent king Arwe is killed with juice from the Euphorbia.
Uses
Several spurges are grown as garden plants, among them poinsettia (E. pulcherrima) and the succulent E. trigona. E. pekinensis (Chinese: 大戟; pinyin: dàjǐ) is used in traditional Chinese medicine, where it is regarded as one of the 50 fundamental herbs. Several Euphorbia species are used as food plants by the larvae of some Lepidoptera (butterflies and moths), like the spurge hawkmoths (Hyles euphorbiae and Hyles tithymali), as well as the giant leopard moth.
Ingenol mebutate, a drug used to treat actinic keratosis, is a diterpenoid found in Euphorbia peplus.
Euphorbias are often used as hedging plants in many parts of Africa.
Among laypeople, Euphorbia species are among the plant taxa most commonly confused with cacti, especially the stem succulents. Euphorbias secrete a sticky, milky-white fluid with latex, but cacti do not. Individual flowers of euphorbias are usually tiny and nondescript (although structures around the individual flowers may not be), without petals and sepals, unlike cacti, which often have fantastically showy flowers. Euphorbias from desert habitats with growth forms similar to cacti have thorns, which are different from the spines of cacti.
Systematics and taxonomy
The present taxon "Euphorbia" corresponds to its own former subtribe, the Euphorbiinae.[citation needed] It has over 2000 species. Morphological description using the presence of a cyathium (see section above) is consistent with nuclear and chloroplast DNA sequence data in testing of about 10% of its members. This testing supports inclusion of formerly other genera as being best placed in this single genus, including Chamaesyce, Monadenium, Pedilanthus, and poinsettia (E. pulcherrima).
Genetic tests have shown that similar flower head structures or forms within the genus, might not mean close ancestry within the genus. The genetic data show that within the genus, convergent evolution of inflorescence structures may be from ancestral subunits that are not related. So using morphology within the genus becomes problematic for further subgeneric grouping. As stated on the Euphorbia Planetary Biodiversity Inventory project webpage:
Previous morphologically based delimitations of subgenera or sections within the genus should not be taken at face value. The genus is in fact rife with striking examples of morphological convergence in cyathial and vegetative features, which justifies a global approach to studying the genus to obtain a proper phylogenetic understanding of the whole group.... The bottom line is that a number of clades have been placed inside or outside of Euphorbia at different times... few of the subgeneric circumscriptions hold up under DNA sequence analysis.
According to a 2002 publication on studies of DNA sequence data, most of the smaller "satellite genera" around the huge genus Euphorbia nest deep within the latter. Consequently, these taxa, namely the never generally accepted genus Chamaesyce, as well as the smaller genera Cubanthus, Elaeophorbia, Endadenium, Monadenium, Synadenium, and Pedilanthus were transferred to Euphorbia. The entire subtribe Euphorbiinae now consists solely of the genus Euphorbia.
Selected species
See List of Euphorbia species for complete list.
Euphorbia albomarginata – rattlesnake weed, white-margined sandmat
Euphorbia amygdaloides – wood spurge
Euphorbia antisyphilitica – candelilla
Euphorbia balsamifera – sweet tabaiba (Canary Islands)
Euphorbia bulbispina
Euphorbia caducifolia – leafless milk hedge
Euphorbia canariensis – Canary Island spurge, Hercules club (Canary Islands)
Euphorbia candelabrum – candelabra tree of East Africa
Euphorbia caput-medusae – Medusa's head (South Africa)
Euphorbia ceratocarpa – (Sicily and southern Italy)
Euphorbia characias – Mediterranean spurge
Euphorbia coerulescens - blue euphorbia
Euphorbia cotinifolia – copper tree
Euphorbia cyathophora – fire-on-the-mountain
Euphorbia cyparissias – Cypress spurge
Euphorbia decidua
Euphorbia dendroides – tree spurge
Euphorbia epithymoides – cushion spurge
Euphorbia esula – leafy spurge
Euphorbia franckiana
Euphorbia fulgens – scarlet plume
Euphorbia grantii – African milk bush
Euphorbia gregersenii – Gregersen's spurge
Euphorbia griffithii – Griffith's spurge
Euphorbia helioscopia – sun spurge
Euphorbia heterophylla – painted euphorbia, desert poinsettia, fireplant, paint leaf, kaliko
Euphorbia hirta – asthma-plant
Euphorbia hispida
Euphorbia horrida – African milk barrel
Euphorbia ingens – candelabra tree
Euphorbia labatii
Euphorbia lactea – mottled spurge, frilled fan, elkhorn
Euphorbia lathyris – caper spurge, paper spurge, gopher spurge, gopher plant, mole plant
Euphorbia leuconeura – Madagascar jewel
Euphorbia maculata – spotted spurge, prostrate spurge
Euphorbia marginata – snow on the mountain
Euphorbia mammillaris
Euphorbia maritae
Euphorbia milii – crown-of-thorns, Christ plant
Euphorbia misera – cliff spurge, Baja California, Southern California
Euphorbia myrsinites – myrtle spurge, creeping spurge, donkey tail
Euphorbia nivulia – leafy milk hedge
Euphorbia obesa
Euphorbia paralias – sea spurge
Euphorbia pekinensis – Peking spurge
Euphorbia peplis – purple spurge
Euphorbia peplus – petty spurge
Euphorbia polychroma – bonfire
Euphorbia psammogeton – sand spurge
Euphorbia pulcherrima – poinsettia, Mexican flame leaf, Christmas star, winter rose, noche buena, lalupatae, pascua, Atatürk çiçeği (Turkish)
Euphorbia purpurea – Darlington's glade spurge, glade spurge, or purple spurge
Euphorbia resinifera – resin spurge
Euphorbia rigida – gopher spurge, upright myrtle spurge
Euphorbia serrata – serrated spurge, sawtooth spurge
Euphorbia tirucalli – Indian tree spurge, milk bush, pencil tree, firestick
Euphorbia tithymaloides – devil's backbone, redbird cactus, cimora misha (Peru)
Euphorbia trigona – African milk tree, cathedral cactus, Abyssinian euphorbia
Euphorbia tuberosa
Euphorbia virosa – gifboom or poison tree
Hybrids
Euphorbia has been extensively hybridised for garden use, with many cultivars available commercially. Moreover, some hybrid plants have been found growing in the wild, for instance E. × martini Rouy, a cross of E. amygdaloides × E. characias subsp. characias, found in southern France.
Subgenera
The genus Euphorbia is one of the largest and most complex genera of flowering plants, and several botanists have made unsuccessful attempts to subdivide the genus into numerous smaller genera. According to the recent phylogenetic studies, Euphorbia can be divided into four subgenera, each containing several sections and groups. Of these, subgenus Esula is the most basal. The subgenera Chamaesyce and Euphorbia are probably sister taxa, but very closely related to subgenus Rhizanthium. Extensive xeromorph adaptations in all probability evolved several times; it is not known if the common ancestor of the cactus-like Rhizanthium and Euphorbia lineages had been xeromorphic—in which case a more normal morphology would have re-evolved namely in Chamaesyce—or whether extensive xeromorphism is entirely polyphyletic even to the level of the subgenera.
The "flower" of flowering spurge (Euphorbia sp.), which - botanically speaking - is actually a "false flower" or cyathium. They're tiny - this one was maybe half a centimeter across. The white leaves are not petals but bracts (modified leaves) surrounding the inflorescence in the center. The green disks are actually nectar glands to attract pollinators. The bracts form a cup-like structure (the cyathium) with a female flower at its center and reduced male flowers surrounding it.
When I broke this "pseudo-flower" off the main plant for better placement to take the picture with the Raynox macro lens, the stem was immediately oozing milky white sap, a characteristic of all euphorbs. The sap is a skin irritant and may cause blistering, so if you mess with these plants and tend to be sensitive, be careful!
Euphorbia is a very large and diverse genus of flowering plants, commonly called spurge, in the family Euphorbiaceae. "Euphorbia" is sometimes used in ordinary English to collectively refer to all members of Euphorbiaceae (in deference to the type genus), not just to members of the genus.
Euphorbias range from tiny annual plants to large and long-lived trees. with perhaps the tallest being Euphorbia ampliphylla at 30 m (98 ft) or more. The genus has roughly 2,000 members, making it one of the largest genera of flowering plants. It also has one of the largest ranges of chromosome counts, along with Rumex and Senecio. Euphorbia antiquorum is the type species for the genus Euphorbia. It was first described by Carl Linnaeus in 1753 in Species Plantarum.
Some euphorbias are widely available commercially, such as poinsettias at Christmas. Some are commonly cultivated as ornamentals, or collected and highly valued for the aesthetic appearance of their unique floral structures, such as the crown of thorns plant (Euphorbia milii). Succulent euphorbias from the deserts of Southern Africa and Madagascar have evolved physical characteristics and forms similar to cacti of North and South America, so they are often incorrectly referred to as cacti. Some are used as ornamentals in landscaping, because of beautiful or striking overall forms, and drought and heat tolerance.
Euphorbia all share the feature of having a poisonous, latex-like sap and unique floral structures When viewed as a whole, the head of flowers looks like a single flower (a pseudanthium). It has a unique kind of pseudanthium, called a cyathium, where each flower in the head is reduced to its barest essential part needed for sexual reproduction. The individual flowers are either male or female, with the male flowers reduced to only the stamen, and the females to the pistil. These flowers have no sepals, petals, or other parts that are typical of flowers in other kinds of plants. Structures supporting the flower head and other structures underneath have evolved to attract pollinators with nectar, and with shapes and colors that function in a way petals and other flower parts do in other flowers. It is the only genus of plants that has all three kinds of photosynthesis, CAM, C3 and C4.[7]
Etymology
The common name "spurge" derives from the Middle English/Old French espurge ("to purge"), due to the use of the plant's sap as a purgative. The botanical name Euphorbia derives from Euphorbos, the Greek physician of King Juba II of Numidia and Mauretania (52–50 BC – 23 AD), who married the daughter of Anthony and Cleopatra. Juba was a prolific writer on various subjects, including natural history. Euphorbos wrote that one of the cactus-like euphorbias (now called Euphorbia obtusifolia ssp. regis-jubae) was used as a powerful laxative. In 12 BC, Juba named this plant after his physician Euphorbos, as Augustus Caesar had dedicated a statue to the brother of Euphorbos, Antonius Musa, who was the personal physician of Augustus. In 1753, botanist and taxonomist Carl Linnaeus assigned the name Euphorbia to the entire genus in the physician's honor.
Description
The plants are annual, biennial or perennial herbs, woody shrubs, or trees with a caustic, poisonous milky latex. The roots are fine or thick and fleshy or tuberous. Many species are more or less succulent, thorny, or unarmed. The main stem and mostly also the side arms of the succulent species are thick and fleshy, and often winged, 15–91 cm (6–36 in) tall. The succulent trees and large shrubs are mostly confined to southern and eastern Africa, but Euphorbia neutra is native to the Caatinga of Brazil and Euphorbia royleana is found in the Himalayan foothills.[14] The deciduous[citation needed] leaves may be opposite, alternate, or in whorls. In succulent species, the leaves are mostly small and short-lived. The stipules are mostly small, partly transformed into spines or glands, or missing.
Inflorescence and fruit
Like all members of the family Euphorbiaceae, spurges have unisexual flowers.
In Euphorbia, flowers occur in a head, called the cyathium (plural cyathia). Each male or female flower in the cyathium head has only its essential sexual part, in males the stamen, and in females the pistil. The flowers do not have sepals, petals, or nectar to attract pollinators, although other nonflower parts of the plant have an appearance and nectar glands with similar roles. Euphorbias are the only plants known to have this kind of flower head.
Nectar glands and nectar that attract pollinators are held in the involucre, a cup-like part below and supporting the cyathium head. The "involucre" in the genus Euphorbia is not to be confused with the "involucre" in family Asteraceae members, which is a collection of bracts called phyllaries, which surround and encase the unopened flower head, then support the receptacle under it after the flower head opens.
The involucre is above and supported by bract-like modified leaf structures (usually in pairs)[citation needed] called cyathophylls', or cyathial leaves. The cyathophyll often has a superficial appearance of being petals of a flower.
Euphorbia flowers are tiny, and the variation attracting different pollinators, with different forms and colors occurs, in the cyathium, involucre, cyathophyll, or additional parts such as glands that attached to these.
The collection of many flowers may be shaped and arranged to appear collectively as a single individual flower, sometimes called a pseudanthium in the Asteraceae, and also in Euphorbia.
The majority of species are monoecious (bearing male and female flowers on the same plant), although some are dioecious with male and female flowers occurring on different plants. It is not unusual for the central cyathia of a cyme to be purely male, and for lateral cyathia to carry both sexes. Sometimes, young plants or those growing under unfavorable conditions are male only, and only produce female flowers in the cyathia with maturity or as growing conditions improve.
The female flowers reduced to a single pistil usually split into three parts, often with two stigmas at each tip.[citation needed] Male flowers often have anthers in twos.[citation needed] Nectar glands usually occur in fives, may be as few as one,[16] and may be fused into a "U" shape. The cyathophylls often occur in twos, are leaf-like, and may be showy and brightly coloured and attractive to pollinators, or be reduced to barely visible tiny scales.
The fruits are three- or rarely two-compartment capsules, sometimes fleshy, but almost always ripening to a woody container that then splits open, sometimes explosively. The seeds are four-angled, oval, or spherical, and some species have a caruncle.
Xerophytes and succulents
In the genus Euphorbia, succulence in the species has often evolved divergently and to differing degrees. Sometimes, it is difficult to decide, and is a question of interpretation, whether or not a species is really succulent or "only" xerophytic. In some cases, especially with geophytes, plants closely related to the succulents are normal herbs. About 850 species are succulent in the strictest sense. If one includes slightly succulent and xerophytic species, this figure rises to about 1000, representing about 45% of all Euphorbia species.
Irritants
The milky sap of spurges (called "latex") evolved as a deterrent to herbivores. It is white, and transparent when dry, except in E. abdelkuri, where it is yellow. The pressurized sap seeps from the slightest wound and congeals after a few minutes in air. The skin-irritating and caustic effects are largely caused by varying amounts of diterpenes. Triterpenes such as betulin and corresponding esters are other major components of the latex. In contact with mucous membranes (eyes, nose, mouth), the latex can produce extremely painful inflammation. The sap has also been known to cause mild to extreme Keratouveitis, which affects vision. Therefore, spurges should be handled with caution and kept away from children and pets. Wearing eye protection while working in close contact with Euphorbia is advised.[18] Latex on skin should be washed off immediately and thoroughly. Congealed latex is insoluble in water, but can be removed with an emulsifier such as milk or soap. A physician should be consulted if inflammation occurs, as severe eye damage including permanent blindness may result from exposure to the sap.
The poisonous qualities were well known: in the Ethiopian Kebra Nagast, the serpent king Arwe is killed with juice from the Euphorbia.
Uses
Several spurges are grown as garden plants, among them poinsettia (E. pulcherrima) and the succulent E. trigona. E. pekinensis (Chinese: 大戟; pinyin: dàjǐ) is used in traditional Chinese medicine, where it is regarded as one of the 50 fundamental herbs. Several Euphorbia species are used as food plants by the larvae of some Lepidoptera (butterflies and moths), like the spurge hawkmoths (Hyles euphorbiae and Hyles tithymali), as well as the giant leopard moth.
Ingenol mebutate, a drug used to treat actinic keratosis, is a diterpenoid found in Euphorbia peplus.
Euphorbias are often used as hedging plants in many parts of Africa.
Among laypeople, Euphorbia species are among the plant taxa most commonly confused with cacti, especially the stem succulents. Euphorbias secrete a sticky, milky-white fluid with latex, but cacti do not. Individual flowers of euphorbias are usually tiny and nondescript (although structures around the individual flowers may not be), without petals and sepals, unlike cacti, which often have fantastically showy flowers. Euphorbias from desert habitats with growth forms similar to cacti have thorns, which are different from the spines of cacti.
Systematics and taxonomy
The present taxon "Euphorbia" corresponds to its own former subtribe, the Euphorbiinae.[citation needed] It has over 2000 species. Morphological description using the presence of a cyathium (see section above) is consistent with nuclear and chloroplast DNA sequence data in testing of about 10% of its members. This testing supports inclusion of formerly other genera as being best placed in this single genus, including Chamaesyce, Monadenium, Pedilanthus, and poinsettia (E. pulcherrima).
Genetic tests have shown that similar flower head structures or forms within the genus, might not mean close ancestry within the genus. The genetic data show that within the genus, convergent evolution of inflorescence structures may be from ancestral subunits that are not related. So using morphology within the genus becomes problematic for further subgeneric grouping. As stated on the Euphorbia Planetary Biodiversity Inventory project webpage:
Previous morphologically based delimitations of subgenera or sections within the genus should not be taken at face value. The genus is in fact rife with striking examples of morphological convergence in cyathial and vegetative features, which justifies a global approach to studying the genus to obtain a proper phylogenetic understanding of the whole group.... The bottom line is that a number of clades have been placed inside or outside of Euphorbia at different times... few of the subgeneric circumscriptions hold up under DNA sequence analysis.
According to a 2002 publication on studies of DNA sequence data, most of the smaller "satellite genera" around the huge genus Euphorbia nest deep within the latter. Consequently, these taxa, namely the never generally accepted genus Chamaesyce, as well as the smaller genera Cubanthus, Elaeophorbia, Endadenium, Monadenium, Synadenium, and Pedilanthus were transferred to Euphorbia. The entire subtribe Euphorbiinae now consists solely of the genus Euphorbia.
Selected species
See List of Euphorbia species for complete list.
Euphorbia albomarginata – rattlesnake weed, white-margined sandmat
Euphorbia amygdaloides – wood spurge
Euphorbia antisyphilitica – candelilla
Euphorbia balsamifera – sweet tabaiba (Canary Islands)
Euphorbia bulbispina
Euphorbia caducifolia – leafless milk hedge
Euphorbia canariensis – Canary Island spurge, Hercules club (Canary Islands)
Euphorbia candelabrum – candelabra tree of East Africa
Euphorbia caput-medusae – Medusa's head (South Africa)
Euphorbia ceratocarpa – (Sicily and southern Italy)
Euphorbia characias – Mediterranean spurge
Euphorbia coerulescens - blue euphorbia
Euphorbia cotinifolia – copper tree
Euphorbia cyathophora – fire-on-the-mountain
Euphorbia cyparissias – Cypress spurge
Euphorbia decidua
Euphorbia dendroides – tree spurge
Euphorbia epithymoides – cushion spurge
Euphorbia esula – leafy spurge
Euphorbia franckiana
Euphorbia fulgens – scarlet plume
Euphorbia grantii – African milk bush
Euphorbia gregersenii – Gregersen's spurge
Euphorbia griffithii – Griffith's spurge
Euphorbia helioscopia – sun spurge
Euphorbia heterophylla – painted euphorbia, desert poinsettia, fireplant, paint leaf, kaliko
Euphorbia hirta – asthma-plant
Euphorbia hispida
Euphorbia horrida – African milk barrel
Euphorbia ingens – candelabra tree
Euphorbia labatii
Euphorbia lactea – mottled spurge, frilled fan, elkhorn
Euphorbia lathyris – caper spurge, paper spurge, gopher spurge, gopher plant, mole plant
Euphorbia leuconeura – Madagascar jewel
Euphorbia maculata – spotted spurge, prostrate spurge
Euphorbia marginata – snow on the mountain
Euphorbia mammillaris
Euphorbia maritae
Euphorbia milii – crown-of-thorns, Christ plant
Euphorbia misera – cliff spurge, Baja California, Southern California
Euphorbia myrsinites – myrtle spurge, creeping spurge, donkey tail
Euphorbia nivulia – leafy milk hedge
Euphorbia obesa
Euphorbia paralias – sea spurge
Euphorbia pekinensis – Peking spurge
Euphorbia peplis – purple spurge
Euphorbia peplus – petty spurge
Euphorbia polychroma – bonfire
Euphorbia psammogeton – sand spurge
Euphorbia pulcherrima – poinsettia, Mexican flame leaf, Christmas star, winter rose, noche buena, lalupatae, pascua, Atatürk çiçeği (Turkish)
Euphorbia purpurea – Darlington's glade spurge, glade spurge, or purple spurge
Euphorbia resinifera – resin spurge
Euphorbia rigida – gopher spurge, upright myrtle spurge
Euphorbia serrata – serrated spurge, sawtooth spurge
Euphorbia tirucalli – Indian tree spurge, milk bush, pencil tree, firestick
Euphorbia tithymaloides – devil's backbone, redbird cactus, cimora misha (Peru)
Euphorbia trigona – African milk tree, cathedral cactus, Abyssinian euphorbia
Euphorbia tuberosa
Euphorbia virosa – gifboom or poison tree
Hybrids
Euphorbia has been extensively hybridised for garden use, with many cultivars available commercially. Moreover, some hybrid plants have been found growing in the wild, for instance E. × martini Rouy, a cross of E. amygdaloides × E. characias subsp. characias, found in southern France.
Subgenera
The genus Euphorbia is one of the largest and most complex genera of flowering plants, and several botanists have made unsuccessful attempts to subdivide the genus into numerous smaller genera. According to the recent phylogenetic studies, Euphorbia can be divided into four subgenera, each containing several sections and groups. Of these, subgenus Esula is the most basal. The subgenera Chamaesyce and Euphorbia are probably sister taxa, but very closely related to subgenus Rhizanthium. Extensive xeromorph adaptations in all probability evolved several times; it is not known if the common ancestor of the cactus-like Rhizanthium and Euphorbia lineages had been xeromorphic—in which case a more normal morphology would have re-evolved namely in Chamaesyce—or whether extensive xeromorphism is entirely polyphyletic even to the level of the subgenera.
Asteraceae (Compositae)
Asteridae: Asterales. The Asteraceae are herbs, shrubs, or less commonly trees and are arguably the largest family of flowering plants, comprising about 1,100 genera and 20,000 species that are characterized by having the flowers reduced and organized into an involucrate pseudanthium in the form of a head or capitulum. The leaves are alternate, opposite, or less commonly whorled, and range from simple to pinnately or palmately compound; stipules are absent