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BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Centaurea diffusa | Diffuse Knapweed

GENERAL BOTANICAL CHARACTERISTICS:


Diffuse knapweed is an exotic, annual, biennial, or short-lived perennial forb [22]. During the juvenile stage, diffuse knapweed is a rosette with deeply divided basal leaves borne on short stalks on a central crown with a taproot. At maturity diffuse knapweed produces 1 upright stem, rarely 2. Stems are 1 to 3 feet (0.3-1 m) tall, with numerous, spreading branches that give the plant a ball-shaped appearance and tumble-weed mobility when broken off. Stem leaves on diffuse knapweed are stalkless, getting smaller and less divided higher up the stem. Flowerheads are solitary or borne in clusters of 2 or 3 at the ends of branches. Diffuse knapweed flowerheads are 3-6 mm in diameter and 8-11 mm long, excluding spines and flowers. Bracts are edged with a fringe of spines, with a longer 1.5-4 mm, erect spine at the tip. Seeds are achenes, 2-3 mm long with a plume of bristle-like hairs that vary from scalelike to 1/8 the length of the seed [18,85].

Once it is established, the highly competitive diffuse knapweed can form monotypic stands. The competitive ability of diffuse knapweed has been attributed to its being adept at depleting soil moisture [108], to allelopathy [14,15,26], and to other competitive or interference mechanisms [14,15].

RAUNKIAER [82] LIFE FORM:


Hemicryptophyte

REGENERATION PROCESSES:


Diffuse knapweed reproduces by seed and is generally biennial. The plants begin as low rosettes and may remain in this form for 1 to several years, until they reach a critical size (e.g. root crown diameter [80]), or stage of development (e.g. leaf number). At this point they respond to vernalization (i.e. low temperatures) by bolting, flowering, setting seed, and dying. Thus, they may behave as annuals, biennials, or short-lived perennials, a habit sometimes designated as "semelparous perennial" [116].

Pollination: Diffuse knapweed is primarily insect pollinated [39,129]. Honeybees, bumble bees, and digger bees are frequent diffuse knapweed flower visitors [39]. Fertilization in diffuse knapweed requires cross-pollination between flowers on different plants. This can limit the reproductive success of isolated individuals, but it also promotes genetic diversity and may thereby improve competitive ability. Watson and Renney [128] reported that diffuse knapweed is self-compatible, but the results of Harrod and Taylor [39] refute this assertion.

Seed production: Average seed production by diffuse knapweed is reported by area as follows:

Location Flowerheads per plant Seeds per flowerhead Seeds per plant Seeds per m2 Reference
British Columbia - rangeland 74 12 925 not reported  
British Columbia - irrigated 1404 13 18,248 not reported [128]
Washington state - averaged across diverse sites and years 89 13 1157 26,400 [93]

Considerable variation in seed production was observed between sites and years in northeastern Washington; nonetheless, seed production was estimated to be 1,000 times what would be necessary to maintain observed levels of infestation [93].

Seed dispersal: Dispersal of diffuse knapweed seed is mainly by wind. Seeds usually remain in the flowerheads after they mature and break away from the receptacle. Dispersal in the vicinity of the parent plant is facilitated by horizontally placed seedheads that open at the top and release seeds as dehydration occurs and plants sway in the wind. Dispersal over longer distances occurs when plants are broken off at ground level and tumble in the wind, dispensing seeds individually from the small opening at the top of the seedheads [128]. This technique adapts well to "hitchhiking" on the frames of vehicles and colonizing the bare shoulders of roads. Seeds may also be transported in mud adhering to vehicles or shoes [129]. Plants bearing seeds may also be carried in rivers and irrigation systems, thus colonizing the banks of waterways [85]. In British Columbia, logging trucks, off-road vehicles, and trail bikes have greatly contributed to the spread of both spotted and diffuse knapweed [112].

Seed banking: Evidence suggests that diffuse knapweed seed germination is distributed over time [69]. This may be considered as evidence of seed banking, although the length of time that diffuse knapweed seeds remain viable in the soil seed bank is undetermined.

Viability: Seed viability information for diffuse knapweed is scarce. In British Columbia , diffuse knapweed seed removed from seedheads at maturity exhibited 40% germination; 25-day-old seed stored under dry conditions exhibited 68% germination; and seeds that overwintered in the seedhead under field conditions exhibited 88% germination [128]. Watson [129] reports successful (93-95%) laboratory germination of 20-month-old seed stored at room temperature [138]. Another laboratory germination test showed greater than 95% viability of diffuse knapweed seeds, although germination values observed in the field were <70% [93].

Germination: Diffuse knapweed seeds germinate in spring (May/June) or late summer/early fall (August/September), and develop into low-lying, tap-rooted rosettes given sufficient moisture [11,68,128]. Diffuse knapweed seeds germinated under a wide range of environmental conditions simulated in the laboratory. Germination of over 80% occurred between 55 and 82 degrees Fahrenheit (13-28 °C) at optimum moisture levels [128]. Diffuse knapweed seeds require more than 55% soil moisture to initiate germination, with optimum emergence between 65 and 70% [107]. Diffuse knapweed seeds germinate best on the soil surface, with emergence rate decreasing as seeding depth increases, and little to no emergence below 1 inch (2.5 cm) [107,128]. Spears and others [107] found diffuse knapweed seeds germinated equally well over the range of 0 to 100% canopy cover. Nolan and Upadyaya [69] observed 3 types of germination behavior in diffuse knapweed with respect to light conditions: nondormant seeds that can germinate in the dark; light-sensitive seeds that germinate after exposure to red light; and light-insensitive dormant seeds that do not respond to exposure to red light. They concluded that bunchgrass rangeland and other open canopy conditions provide favorable light conditions for diffuse knapweed seed germination.

Seedling establishment: Stannard [108] reports high seedling vigor in diffuse knapweed, while Myers and Berube [68] indicate that the greatest mortality in diffuse knapweed occurs between the seedling and rosette stages. Seedling mortality is highest during the summer and is largely related to moisture availability [68,80,93,128]. Once diffuse knapweed seedlings establish as rosettes, they become very drought resistant [11]. Crowding of plants is also a factor in seedling and juvenile mortality, with mortality rates highest among the smallest rosettes, and declining with size under crowded conditions. The proportion of diffuse knapweed plants that flower and produce seed each year (and subsequently die) increases with available growing space [80]. Additionally, Powell [80] observed that the majority of diffuse knapweed rosettes that died during the summer at 1 site in British Columbia were infested with the larvae of the introduced biocontrol beetle Sphenoptera jugoslavica, and suggested that the damage caused by the larvae increased the susceptibility of rosettes to interference-related mortality. Amount and pattern of rainfall are important for diffuse knapweed survival. Wetting increases germination, but when followed by drying (e.g. the wet-dry cycle occurring with the summer convective storms in British Columbia ranges) seedling survival is diminished [11]. High spring precipitation appears to favor diffuse knapweed seedling establishment [101].

Asexual regeneration: Diffuse knapweed can sprout from the root crown after top-kill [85,128]. Other methods of asexual regeneration are not known to occur in diffuse knapweed [128].

SITE CHARACTERISTICS:


Semiarid rangeland and dry open forest sites in the northwestern U.S. are subject to invasion by diffuse knapweed, especially with disturbance. In British Columbia, density of diffuse knapweed was significantly (p<0.05) correlated with degree of soil disturbance, but not with any soil chemical properties [129]. Disturbance also allows diffuse knapweed to invade a wide range of habitats [16,85], where it can rapidly establish dense, often monotypic, stands. Disturbances may be as small as rodent activity or a single hailstorm [50] and may not be recent [85]. Typical disturbed sites subject to diffuse knapweed invasion include transportation corridors, water ways, gravel pits, and industrial areas [88]. Diffuse knapweed is also capable of invading well-managed rangeland [11,50,99].

Diffuse knapweed is tolerant of a wide range of total precipitation and temperature conditions but does best in semiarid and arid conditions, and is most competitive in areas receiving between 12 and 17 inches (305-432 mm) of annual precipitation [11,36,87,128]. Diffuse knapweed is susceptible to flooded or waterlogged conditions, and infestations stop abruptly with an increase in soil moisture near temporary and permanent streams. Irrigation can also eliminate diffuse knapweed [68]. Diffuse knapweed is not competitive in moist microsites such as gullies, depressions, and poorly drained soils [11,85]. Diffuse knapweed seeds require conditions near field capacity for at least 4 days to begin germination. Seedling root growth in diffuse knapweed may be sensitive to saline conditions (electrical conductivity of 4 dS m-1). Seedling root growth may also be sensitive to and germination delayed by water stress (osmotic potential of -0.5 mP or less) [45].

In eastern Washington, diffuse knapweed grows on all aspects and slope positions, from flat to over 60% [85,87,115]. In the Gilpin range, British Columbia, knapweed is commonly found on south-facing slopes below 3,000 feet (900 m) [63]. The following table provides some elevation and precipitation ranges as reported by state or province:

Location Elevation range Annual precipitation range Mean annual temperature range References
Arizona up to 7,000 ft (2,134 m)  ----  ---- [138]
Washington 0-5,000 ft (0-1,500 m) 6-35 inches (150-900 mm)  ---- [85,115]
British Columbia 500-3,000 ft (150-900 m)  ---- 45-49 oF (7-9 °C) [128,129]

Diffuse knapweed is commonly found on well-drained soils such as sandy or gravelly loams or loamy fine sands, with coarse fragments from 0 to over 80% [86,87]. It is less competitive on shallow soils (< 15 inches (38 cm) deep) and very coarse textured soils such as sand or loamy coarse sand, although it may thrive on these sites when disturbance removes other vegetation [87]. It grows best on fertile, well-watered Cryoborolls, mesic Argiudolls, and mesic Hapludolls in open and uncultivated sites with summer drought [36,99].

SUCCESSIONAL STATUS:


Diffuse knapweed is an early successional species that establishes best on disturbed ground. Although diffuse knapweed seeds can germinate in the dark [69,107], diffuse knapweed plants do not grow well in dense shade [85]. Diffuse knapweed can dominate a site over time and persist in monotypic stands [100].

SEASONAL DEVELOPMENT:


The phenology of individual diffuse knapweed plants is highly variable and dependent on climatic and site conditions such as moisture, temperature, and plant density. Diffuse knapweed seedlings usually emerge under favorable conditions in spring and/or fall and develop into rosettes. Most root development occurs during this stage [128]. Rosettes respond to a period of cold temperature (vernalization) and bolt when a critical size or stage of development is reached [80,116]. Diffuse knapweed plants that complete their juvenile growth by fall and overwinter as rosettes usually bolt in early May. Plants that do not complete the juvenile stage by the end of fall remain as rosettes through the 2nd year and may bolt during the 3rd year. Flower buds are formed in early June. Flowering occurs July through September [18,128] or later as permitted by adequate moisture and mild temperatures [85]. Mature seeds are usually formed by mid-August, followed by death of the plant. Dead plants break off at ground level in the spring and tumble with the wind, spreading seed as they roll [128].

Powell [80] found that under conditions of severe crowding, a plant might not flower for 5 or more years. Other exceptions include plants that germinate in early spring that may flower the 1st year (<1% observed) [93], and plants that continue to grow after producing seed to flower again the following year, behaving as short-lived perennials [85].

Related categories for SPECIES: Centaurea diffusa | Diffuse Knapweed

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