Overview

Comprehensive Description

Description

Annual or perennial herbs. Leaves alternate. Capitula solitary, heterogamous; florets tubular; outer neuter; inner bisexual. Phyllaries many-seriate, imbricate, with a pectinate or spiny, apical appendage. Receptacle densely setose; setae twisted. Achenes ± laterally flattened, glabrous when mature. Pappus many-seriate, of persistent, unequal, barbellate, scale-like setae.
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© Mark Hyde, Bart Wursten and Petra Ballings

Source: Flora of Zimbabwe

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Ecology

Associations

Foodplant / gall
Aceria centaurea causes gall of leaf of Centaurea

In Great Britain and/or Ireland:
Foodplant / nest
female of Andrena hattorfiana provisions nest with pollen of Centaurea
Other: minor host/prey

Foodplant / internal feeder
larva of Apion armatum feeds within stem? of Centaurea

Foodplant / internal feeder
larva of Apion onopordi feeds within stem of Centaurea

Foodplant / parasite
sporangium of Bremia lactucae parasitises live leaf of Centaurea
Remarks: season: 9-10
Other: unusual host/prey

Foodplant / open feeder
larva of Cassida vibex grazes on leaf of Centaurea

Plant / resting place / on
adult of Chrysolina marginata may be found on Centaurea
Remarks: season: early 8-mid 11,4-

Foodplant / sap sucker
Dactynotus jaceae sucks sap of stem of Centaurea
Remarks: season: 6-9

Foodplant / saprobe
apothecium of Hyalopeziza millepunctata is saprobic on dead stem of Centaurea
Remarks: season: 10-6

Foodplant / gall
larva of Isocolus fitchi causes gall of leaf (midrib) of Centaurea

Foodplant / gall
larva of Isocolus jaceae causes gall of inflorescence of Centaurea

Foodplant / gall
larva of Isocolus rogenhoferi causes gall of inflorescence of Centaurea

Foodplant / gall
larva of Isocolus scabiosae causes gall of root of Centaurea

Foodplant / saprobe
immersed pseudothecium of Kalmusia clivensis is saprobic on dead stem of Centaurea
Remarks: season: 5-6

Foodplant / internal feeder
larva of Larinus planus feeds within capitulum of Centaurea
Remarks: Other: uncertain

Foodplant / saprobe
erumpent pseudothecium of Leptosphaeria doliolum is saprobic on dead stem of Centaurea
Remarks: season: 1-12

Foodplant / miner
larva of Liriomyza strigata mines leaf of Centaurea

Foodplant / gall
larva of Loewiola centaureae causes gall of leaf (vein) of Centaurea

Foodplant / saprobe
immersed or semi-immersed pseudothecium of Lophiostoma caulium is saprobic on dead stem of Centaurea
Remarks: season: 1-12

Foodplant / saprobe
mostly immersed, becoming partly erumpent to free pseudothecium of Lophiostoma semiliberum is saprobic on dead stem of Centaurea
Remarks: season: 12-4
Other: minor host/prey

Foodplant / saprobe
usually immersed pseudothecium of Lophiostoma vagabundum is saprobic on dead stem of Centaurea
Remarks: season: 1-12
Other: major host/prey

Foodplant / gall
larva of Phanacis centaureae causes gall of stem of Centaurea

Foodplant / saprobe
gregarious or scattered, erumpent apothecium of Pirottaea inopinata is saprobic on dead stem of Centaurea
Remarks: season: 4

Foodplant / saprobe
apothecium of Pyrenopeziza revincta is saprobic on dead stem of Centaurea

Foodplant / feeds on
Selenophoma coelomycetous anamorph of Selenophoma moravica feeds on Centaurea

Foodplant / open feeder
larva of Sphaeroderma testaceum grazes on Centaurea

Foodplant / feeds on
Tanymecus palliatus feeds on Centaurea
Other: major host/prey

Foodplant / internal feeder
larva of Trupanea amoena feeds within capitulum of Centaurea

Foodplant / saprobe
sessile apothecium of Unguicularia incarnatina is saprobic on dead stem of Centaurea
Remarks: season: 4-7

Foodplant / gall
larva of Urophora cuspidata causes gall of capitulum of Centaurea

Foodplant / gall
larva of Urophora jaceana causes gall of capitulum of Centaurea

Foodplant / gall
larva of Urophora quadrifasciata causes gall of capitulum of Centaurea

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
                                        
Specimen Records:121Public Records:52
Specimens with Sequences:98Public Species:24
Specimens with Barcodes:97Public BINs:0
Species:35         
Species With Barcodes:32         
          
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© Barcode of Life Data Systems

Source: Barcode of Life Data Systems (BOLD)

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Locations of barcode samples

Collection Sites: world map showing specimen collection locations for Centaurea

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Source: Barcode of Life Data Systems (BOLD)

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Barcode data

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Wikipedia

Centaurea

For the race horse, see Australasian Oaks or Storm Queen Stakes.
"Basket flower" and spelling variants redirect here. In Australia, this usually means the unrelated plant Adenanthos obovatus.

Centaurea (/ˌsɛntɔːˈrə/)[1] is a genus of between 350 and 600 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. 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. "Cornflowers" is used for a few species, but that term more often specifically means C. cyanus (sometimes also called "Basket Flower"). The common name "centaury" is sometimes used, although this also refers to the unrelated plant genus Centaurium.[2]

Description and ecology[edit]

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.

Common Knapweed (C. nigra), perhaps the single most abundant Centaurea species of England
Centaurea tchihatcheffii (locally known as Yanardöner), a highly distinctive and rare knapweed endemic to Turkey

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 herbs. 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 Festuce (F. rubra) in addition to Cock's-foot, 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.[3] 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.[4]

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.

Mating pair of Knapweed Fritillaries (Melitaea phoebe), male at the left.
These butterflies can spend their entire lives living off a patch of Brown Knapweed (C. jacea).

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 (Lycaeides 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[edit]

Though the overall effect of the entire genus on humans is probably detrimental, this is merely due to the considerable damage done by a handful of species when these are allowed to spread uncontrolled. On a per-species basis, most centaureas are inoffensive and at least pleasant-looking plants. As noted above, many species of Centaurea are popular food for insects that may otherwise attack crops; particular in small-scale farming it is often advisable to allow non-invasive species (like Cornflower, C. cyanus, in Europe) to grow around fields. In such a beetle bank, they will draw pests away from crops, and at the same time by their robust structure provide shelter for invertebrate and small vertebrate predators that will keep overall pest populations low.[5]

The abundant nectar produced by Centaurea flowers attracts many pollinators. This is another reason for the success of the highly invasive species, but as it holds true for the entire genus it is not decisive. Yet it can be exploited to the farmer's advantage in combination with biological control of these weeds: 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 USA – 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[4] 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.[3] Thus, 8-hydroxyquinoline is potentially useful to control American plants that have become invasive weeds in the Diffuse Knapweed's native range.

Arctiin from C. imperialis kills cancer cells in culture

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.[6]

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.[7]

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 – its local name rukkilill meaning "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.[8] Lastly, the color cornflower blue is named after C. cyanus.

Systematics and taxonomy[edit]

As namesake member of the subtribe Centaureinae of tribe Cynareae, 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.[9]

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 lineage 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.[2]

Synonyms[edit]

Selected species[edit]

Species presently placed in Centaurea include:

Formerly placed here[edit]

Plant species placed in Centaurea in former times include:

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