Horticulture/Centaurea diffusa

Diffuse knapweed (Centaurea diffusa), also known as white knapweed, is a member of the Genus ../Centaurea/ in the Family ../Asteraceae/. It is native to Asia Minor (Turkey, Syria), the Balkans, (Bulgaria, Greece, Romania), Ukraine, and southern Russia.

Description
Diffuse knapweed is an annual or biennial plant, generally growing to between 10 and 60 cm in height. It has a highly branched stem and a large taproot, as well as a basal rosette of leaves with smaller leaves alternating on the upright stems. Flowers are usually white or pink and grow out of urn-shaped heads carried at the tips of the many branches. Diffuse knapweed often assumes a short rosette form for one year, reaching maximum size, then rapidly growing and flowering during the second year. A single plant can produce approximately 18,000 seeds.

Ecology
Diffuse knapweed is considered an invasive species throughout North America, having established itself in many areas of the continent. C. diffusa was first identified from North America in 1907 when it was found in an alfalfa field in Washington state. The seeds had presumably been transported in an impure alfalfa seed shipment coming from somewhere in the species native range. Now present in at least 19 states in the United States, it has invaded all contiguous states west of the Rockies and additionally in Connecticut, Massachusetts, and New Jersey. Portions of western Canada have also been infested by this plant.

The plant is very resistant to drought, spreads quickly, and possesses other competitive advantages that have allowed it to establish and flourish in its introduced range. Areas in which diffuse knapweed has been established generally are plains rangelands or forest benchlands. Land that has been recently disturbed—by human or natural processes—is favored for the establishment of diffuse knapweed. It has the potential to prosper in semi-arid and arid environments and seems to favor light, dry, porous soils. Areas with large amounts of shade or high levels of water discourage diffuse knapweed growth.

Dispersion occurs in the following ways: Wind is the primary means by which diffuse knapweed seeds are spread.
 * Agriculture – alfalfa contaminated with diffuse knapweed seed can promote the spread of diffuse knapweed;
 * Wildlife – wild animals eating the seeds or transporting the seeds on fur;
 * Wind – seeds blown out of their capsules held on the plant are distributed over a short range, but when the plant dries out it may roll for great distances like tumbleweed, releasing seeds along the way;
 * Water – waterways carry seeds in their flow for long distances before depositing them onto a shore where they germinate.

Control
Effective control of diffuse knapweed requires a fusion of well-executed land management, biological control, physical control, chemical control and reestablishment of the native species. Any method of control must ensure that the root is removed or the plant will grow back. Additionally, native plant growth in areas where diffuse knapweed has been removed should be encouraged to prevent reestablishment.

Analysis
The largest impact of humans on diffuse knapweed is certainly due to our efforts in controlling and eradicating its invasive populations. The several methods outlined in the control section represent a small sample of literally hundreds of approaches being tried with varying levels of effectiveness. Besides reducing the spread of diffuse knapweed, we are also providing selective pressure against the individuals that cannot withstand a certain method of control. Selective pressure, given sufficient time, can cause the adaptation or evolution of invasive species such as diffuse knapweed. If an individual diffuse knapweed plant survives control efforts because of a trait it possesses, its progeny will make up a greater portion of the population than the plants that succumbed to the control.

The success of diffuse knapweed must be attributed to a combination of several mechanisms. Its invasiveness is due to a mix of allelopathy, ERH and superior resource competition. However, the most importance must be attributed to the ERH because diffuse knapweed, while a very effective invasive species in its novel environment, is non-invasive and doesn’t establish monocultures in its native range. It is the differences, biotic and abiotic, between its novel and native surroundings that cause it to be invasive.

To demonstrate that the ERH applies to diffuse knapweed, it is essential to show that the absence of natural enemies has a significant positive effect on its success. One way to show this is to observe the effect of introducing some of diffuse knapweed’s natural enemies into its novel environment. If diffuse knapweed, which generally thrives in its invaded environment, is significantly inhibited through the introduction of natural enemies, it can be concluded that diffuse knapweed is more competitive in the absence of its natural enemies. A recent effort at biocontrol of diffuse knapweed in Camas County, Idaho effectively reduced 8,000 ha (20,000 acres) of knapweed to minimal levels through the release of the lesser knapweed flower weevil and the knapweed root weevil. Since both of the insects released are natural competitors of diffuse knapweed, and since this and other similar efforts at biocontrol have been successful, there is significant evidence that diffuse knapweed benefits from the absence of its natural enemies.

Another aspect of diffuse knapweed’s success relies on the effect of its allelopathic chemicals in its novel environment. Although there is still debate concerning the effectiveness of allelopathic chemicals in the field, the evidence of allelopathic effects demonstrated in a laboratory setting and its propensity to establish monocultures support the importance of allelopathy to diffuse knapweed’s success.

Curiously, diffuse knapweed’s allelopathic chemicals were shown to have a deleterious effect on the North American competitors but were beneficial to its native competitors. While diffuse knapweed’s native competitors are able to compete more effectively in the presence of allelopathic chemicals, the novel competitor’s fitness is decreased. This situation provides an example of the effectiveness of the allelopathy mechanism benefiting from the ERH. The increased effectiveness of allelopathic chemicals cause diffuse knapweed to experience less competitive pressure. As a result, diffuse knapweed is able to establish more predominantly in this new area.

Another connection between allelopathy and the ERH is the fact that concentrations of allelopathic chemicals were found to increase when diffuse knapweed was planted in North American soil as opposed to Eurasian soil. This effect is probably due to the absence of unfavorable soil conditions or soil microorganisms that exist in its native environment. As a result, the allelopathic chemicals will be able to reach higher concentrations, spread farther and therefore be more effective. By effecting more neighboring plants, the favorable changes in soil condition contribute to the success of diffuse knapweed.

Besides the advantages that diffuse knapweed gains from the ERH and allelopathy, it also possesses several characteristically invasive traits. One factor leading to the superior resource competition of diffuse knapweed is its ability to exist in drought conditions. This advantage allows diffuse knapweed to devote its resources to competition while its neighbors are conserving resources to survive. The high number of seeds produced by diffuse knapweed is also a common trait of invasive plants. A higher density of knapweed will not only increase the concentration of allelopathic chemicals in the soil but will also restrict the nutrients available to native plants. Unfortunately, very little research has been conducted to determine the relative competitive ability between diffuse knapweed and its novel competitors. However, tests conducted on the effect of diffuse knapweed on North American grasses in the absence on allelopathic chemicals demonstrated that the fitness of these grasses declined in the presence of diffuse knapweed. Regrettably, we cannot decide if diffuse knapweed is, for general purposes, a better competitor from this data alone. Comparisons of the deleterious effects between these and other pairs of competitors to arrive at a conclusion.

Diffuse knapweed is successful in its novel range primarily because the organisms and conditions that prevent it from becoming invasive in its native environment are absent. It follows that the introduction of species from its native habitat would be an effective method of control. However, the introduction of a non-native organism has the potential to result in another invasive species outbreak. Therefore, any method of biological control must be preceded by analysis of possible effects.