Linaria dalmatica
(dalmatian toadflax)

Pictures

Picture	Title	Caption	Copyright
Title Habit Caption Linaria dalmatica (dalmatian toadflax); flowering habit. USA. Copyright ©Eric Coombs/Oregon Department of Agriculture/Bugwood.org - CC BY 3.0 US	Habit	Linaria dalmatica (dalmatian toadflax); flowering habit. USA.	©Eric Coombs/Oregon Department of Agriculture/Bugwood.org - CC BY 3.0 US
Title Infestation Caption Linaria dalmatica (dalmatian toadflax); urban infestation, showing habit. Burns, Oregon, USA. June 2013. Copyright ©Eric Coombs-2013/Oregon Department of Agriculture/Bugwood.org - CC BY 3.0 US	Infestation	Linaria dalmatica (dalmatian toadflax); urban infestation, showing habit. Burns, Oregon, USA. June 2013.	©Eric Coombs-2013/Oregon Department of Agriculture/Bugwood.org - CC BY 3.0 US
Title Flowers Caption Linaria dalmatica (dalmatian toadflax); close view of flowers. USA. Copyright ©Eric Coombs/Oregon Department of Agriculture/Bugwood.org - CC BY 3.0 US	Flowers	Linaria dalmatica (dalmatian toadflax); close view of flowers. USA.	©Eric Coombs/Oregon Department of Agriculture/Bugwood.org - CC BY 3.0 US
Title Leaves Caption Linaria dalmatica (dalmatian toadflax); foliage. USA. Copyright ©Bonnie Million/National Park Service/Bugwood.org - CC BY-NC 3.0 US	Leaves	Linaria dalmatica (dalmatian toadflax); foliage. USA.	©Bonnie Million/National Park Service/Bugwood.org - CC BY-NC 3.0 US
Title Seeds Caption Linaria dalmatica (dalmatian toadflax); seeds. USA. Copyright ©Steve Hurst/USDA NRCS PLANTS Database/Bugwood.org - CC BY-NC 3.0 US	Seeds	Linaria dalmatica (dalmatian toadflax); seeds. USA.	©Steve Hurst/USDA NRCS PLANTS Database/Bugwood.org - CC BY-NC 3.0 US
Title Seedlings Caption Linaria dalmatica (dalmatian toadflax); seedlings. USA. Copyright ©Linda Wilson/University of Idaho/Bugwood.org - CC BY-NC 3.0 US	Seedlings	Linaria dalmatica (dalmatian toadflax); seedlings. USA.	©Linda Wilson/University of Idaho/Bugwood.org - CC BY-NC 3.0 US

Identity

Preferred Scientific Name

• Linaria dalmatica (L.) Mill.

Preferred Common Name

• dalmatian toadflax

Other Scientific Names

• Antirrhinum dalmaticum L.
• Linaria dalmatica var. dalmatica
• Linaria genistifolia subsp. dalmatica (L.) Maire & Petitm.

International Common Names

• English: balkan toadflax; broadleaf toadflax

Local Common Names

• Germany: leinkraut dalmatinische
• Portugal: linaria-dalmática
• Sweden: jättesporre

EPPO code

• LINDA (Linaria genistifolia subsp. dalmatica)

Summary of Invasiveness

L. dalmatica is a herbaceous plant native to western Asia and south east Europe. It is of particular concern in North America where it was introduced in the late nineteenth century. It has since spread across most of the western areas of the USA and Canada (De Clerck-Floate and Turner, 2013). There it has invaded rangelands, rights-of-way and natural habitats. It is classified as noxious in many of the western states of both countries. It is also classed as invasive in South Africa. Its success may be attributed to its high specific leaf area, aggressive root system, prolific seed production and the presence of alkaloids which discourages grazing by livestock, thus allowing it to become dominant (De Clerck-Floate and Turner, 2013).

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Scrophulariales
•                         Family: Scrophulariaceae
•                             Genus: Linaria
•                                 Species: Linaria dalmatica

Notes on Taxonomy and Nomenclature

This species was named Antirrhinum dalmaticum by Linnaeus, but was moved to the genus Linaria as L. dalmatica by Philip Miller in 1768 and that name has been widely accepted. The Royal Botanic Garden Edinburgh (2015) and some other authorities however now use L. genistifolia subsp. dalmatica. The genus Linaria was previously included in the family Scrophulariaceae. Many sources still use this classification but a series of genetic studies resulted in the ‘disintegration’ of the old Scrophulariaceae such that, apart from all the parasitic members of the family being transferred to the Orobanchaceae, Linaria along with Antirrhinum and Scrophularia were transferred to Plantaginaceae (Judd et al., 1999; Olmstead et al., 2001; Judd et al., 2002; APG, 2003). The genus comprises about 100 species.

Two naturally occurring varieties are recognized, L. dalmatica var. dalmatica and L. dalmatica var. macedonica. The latter, sometimes treated as a subspecies, occurs only in Macedonia (Vujnovic and Wein, 2005). There is a further range of horticultural varieties.

A narrow leaved variant mentioned in some publications is now considered to be the related but distinct L. genistifolia, known as narrowleaf or broomleaf toadflax.

Zouhar (2003) reported that hybrids with L. vulgaris had been created in the laboratory. This was later confirmed in the wild in Montana, USA by Ward et al. (2009) with hybrids being viable and fertile.

Description

L. dalmatica is a robust, glaucous, herbaceous perennial with taproots enlarged above, more or less woody, deeply penetrating (1.8 m or more into the ground), spreading by horizontal roots 5–20 cm below the surface, spreading up to 3.6 m from the plant. Adventitious offshoots from these roots are generally sterile, decumbent to weakly ascending, succulent, 3–40 per crown. Stems from the main crown are annual, 40–100 cm high, ascending to erect, more or less woody near the ground. Leaves are alternate, often in two’s or three’s on lower part of the stem, sessile, reflexed to ascending, entire, acute to acuminate at the tip, obtuse to cordate at the base, sometimes concave above, usually amplexicaul, somewhat leathery, glabrous and sometimes rugose, with 3–7 longitudinal veins on lower surface. The lower leaves are 1-5 cm long, up to 1 cm wide, linear to lanceolate; the upper leaves 3–6 cm long, 1-4 cm wide, lanceolate to very broad-ovate. Bracts are similar to upper leaves, or smaller. Flowers are in erect or nodding, simple racemes, usually loose, sometimes compact; 15–55 cm long on pedicels 1–8 mm long. Five sepals, valvate or somewhat imbricate, erect, lanceolate acute to acuminate, 0.7–1.1 cm long, 0.2–0.4 cm wide, margins entire to somewhat undulate. The corolla, including spur 3.3–4.5 cm long, light yellow to yellow, rarely nearly white; tube 6–9 mm long, the upper lip bilobate, often somewhat helmet-shaped, the lower lip trilobate; spur tapered, 1.2–1.8 cm long, straight or slightly curved; four fertile stamens adnate in pairs, didynamous 0.8–1.5 cm long, one reduced staminode; pistil bicarpellate; ovary bilaterally symmetrical ca. 2 mm diameter at anthesis, abruptly narrowed to the terminal style; style filiform, erect, stigma single, terminal, hemispherical; numerous ovules in each locule; capsule long ovoid to nearly spherical, 4–10 mm long, 4–8 mm in diameter, dehiscing terminally on each carpel. Seeds black to purplish-brown, triquetrous or somewhat compressed, muricate-rugose on the surface, 0.7–1.3 mm wide, 30–170 per locule; each seed with narrow wing on the angles. The species is morphologically highly variable (this description simplified from Vujnovic and Wein, 2005, based on original by Alex, 1962).

Plant Type

Distribution

The native distribution of L. dalmatica is restricted to the Balkan Peninsula, Turkey, Iran, Iraq, Syria and Romania. The Royal Botanic Garden Edinburgh (2015) notes that it is locally naturalized in Central Europe therefore, the records from Germany and Hungary presumably refer to introductions. It also occurs as an occasional escape from gardens in Switzerland, UK and elsewhere in Europe. It is now widely naturalized and invasive across Canada and western USA (Leo’s Dictionaries, 2015). L. dalmatica has more recently been spreading in South Africa (Invasive Species South Africa, 2015) and has also naturalized in Australia (Parsons and Cuthbertson, 1992) and in Argentina (Puntieri and Brion, 2003). As it is widely cultivated in gardens, it is likely to occur as an occasional escape in many other countries.

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

L. dalmatica is believed to have been introduced to western USA by the late nineteenth or early twentieth century and had become naturalized in California by 1920. In Canada it was first planted in Ottawa in 1901 and was reported as naturalized in Alberta by 1933 (Vujnovic and Wein, 2005).

Risk of Introduction

The risk of introduction of L. dalmatica continues to be high, though there must be few temperate countries to which it has not already been introduced as an ornamental. Thus the risk of spread is mainly accidental, from careless disposal of material from gardens.

Habitat

In its native range L. dalmatica is a plant of uncultivated fields, vineyards, roadsides and waste places, mainly on coarse textured soils. It may occur on limestone hills and generally favours alkaline soils. In North America, its introduced range, it occurs in similar habitats but has also invaded forest plantations, wood and herbaceous species in the genus Artemisia, pastures, rangelands and to some extent crops such as lucerne/alfalfa (Medicago sativa) (Vujnovic and Wein, 2005).

L. dalmatica has a wide temperature tolerance but on a mountain area in Canada, establishment on disturbed roadsides was limited by the low temperatures experienced at higher elevation (Polinac and Rew, 2014).

Habitat List

Hosts/Species Affected

The crops affected by L. dalmatica are mainly pasture and rangeland species, but alfalfa (Medicago sativa) and forest species such as ponderosa pine (Pinus ponderosa) may also be affected.

Host Plants and Other Plants Affected

Biology and Ecology

Genetics

The chromosome number 2n = 12 has been reported for L. dalmatica (Missouri Botanical Garden, 2015).

Reproductive Biology

L. dalmatica spreads both by seed and by its spreading root system. The shoots arising from the roots are described as ‘sterile’, but presumably will develop fully if separated from the parent plants.

Under laboratory conditions some seeds will germinate the same season in which produced. Germination from dry storage increases for two to three years, with a periodic low in November and a high in April. Over 90% germination is obtained with two to three year old seeds in the laboratory. In the field, germination occurs in both spring and autumn and seedlings can emerge from a soil depth of 2-2.5 cm. Seeds may be dormant for a few weeks after maturation. Thereafter they have no particular requirement (Robocker, 1970).

Seedlings which emerge in the autumn generally do not survive the following year. Most emergences occur in the spring, following stratification over winter. Cyclic alterations of 10 days stratification at 2°C and 10 days drying at room temperature for two cycles induced higher rates of germination than did any other factor. The highest germination of seeds planted at different depths in five soil textures was found to be from a depth of 1.27 cm in loamy sand (Alex, 1962). Seeds germinate from 2.5 cm in clay and loamy soils and from 5 cm in sandy soils. Seedling recruitment can be seriously reduced by plant competition (Grieshop and Nowierski, 2002).

Shoots growing from the crown flower from May and fruit by August, producing large numbers of seeds, up to 250 per capsule and up to 500,000 per plant. The flowers are self-sterile and are insect pollinated (Vujnovic and Wein, 2005). Seventeen species of bumble bee (Bombus spp.), honey bee (Apis mellifera) and the leaf cutting bee Megachile periherta were identified as pollinators, with bumble bees being most common (97% of observations) (De Clerck-Floate et al., 1997).

Physiology and Phenology

A study by Alex (1962) found that roots of seedlings penetrated downward ca. 1 cm/day during the first 31 days in fine sandy loam. Thereafter the rate was slower but roots reached a depth of 56 cm after 105 days. The total length of all roots of one 105 day old plant grown in fine sandy loam exceeded 27 m. Adventitious stem buds were present on the primary axis after 22 days of age and on the second and third order branch roots at 105 days of age. Stem buds on branch roots were mostly more than 16 cm distant from the primary axis and at depths of 2 cm to more than 10 cm below the ground surface (Alex, 1962). Once established, seedlings can flower and set seed within the first year (Vujnovic and Wein, 2005).

L. dalmatica has been shown to benefit greatly from enhanced carbon dioxide levels. In one study, at 600 ppm CO₂, the biomass of L. dalmatica increased by 13 fold, seed production by 32 fold and clonal expansion by seven fold, much larger than the effects observed on the native C₃ grass Pascopyrum smithii [Elymus smithii] (Blumenthal et al., 2013). Increased temperatures of 1.5/3.0°C (day/night) had little effect (Blumenthal et al., 2013). However a study by Dijkstra et al. (2010) found much less pronounced increases and emphasized the potential interactions with nitrogen and water levels.

Longevity

The average life of a plant is about three years. The life of a stand varies and depends on the interactions of the plant's growth habits with environmental factors. A stand may disappear within three years under severe competition. However the development of floral stems from secondary crown points on lateral roots may prolong or sustain the stand beyond the life of the original plants (Robocker, 1974).

Seeds persist in the field for at least 10 years (Robocker, 1974). A study by Bruns (1965) found that seeds could still germinate after 12 months of submergence in canal water.

Nutrition

Nitrogen addition has a strong positive effect on plant performance, with a two fold increase in both biomass and seed production (Jamieson et al., 2010).

Associations

In North America, L. dalmatica is associated with pastures and rangelands dominated by Bromus tectorum, B. japonicus, Stipa comata, Koeleria cristata [Koeleria pyramidata], Festuca idahoensis and Artemisia tridentata (Vujnovic and Wein, 2005).

Environmental Requirements

L. dalmatica is favoured by high water and nitrogen availability, but is relatively unpalatable to cattle. Cover of L. dalmatica was high on steeper slopes, particularly those with southern aspects (Jamieson et al., 2010). The beneficial effects of nitrogen on L. dalmatica can be neutralized by addition of labile carbon to the soil (Blumenthal, 2009).

Wilson et al. (2005) indicate that L. dalmatica is particularly favoured in areas with dry summers. Nevertheless, snow can be highly beneficial to L. dalmatica, apparently in terms of providing high moisture during its establishment phase (Blumenthal et al., 2008).

Post wildfire conditions in high severity burned areas favour increased density, cover, reproduction and spread of L. dalmatica. Whilst native species richness may be reduced, suggesting that the invasive species would persist, at least in the short term, at the expense of natives (Dodge et al., 2008). L. dalmatica was also found to benefit from prescribed fire in a rangeland situation (Jacobs and Sheley, 2003).

Climate

Latitude/Altitude Ranges

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil drainage

• free

Soil reaction

• acid
• alkaline
• neutral

Soil texture

• heavy
• light
• medium

Special soil tolerances

• shallow

Natural enemies

Notes on Natural Enemies

The natural enemies of L. dalmatica have been well studied as a part of efforts to achieve biological control. Potential biocontrol organisms have included Calophasia lunula, Mecinus janthiniformis, Eteobalea intermediaella, Rhinusa neta, R. linariae and R. antirrhini. Recent studies include the stem boring weevils M. laeviceps and M. peterharrisi and the stem-galling weevil R. rara.

Means of Movement and Dispersal

Natural Dispersal

Seeds of L. dalmatica mainly fall near the parent plant but may also be spread to some extent by wind (Robocker, 1974). L. dalmatica can also reproduce vegetatively by the production of new shoots from roots and root fragments.

Vector Transmission

Some dispersal can occur locally via grazing by cows and deer. It has been suggested that seed can remain viable after ingestion and egestion (Robocker, 1974).

Accidental Introduction

Accidental introduction can occur on a local basis by contaminated cultivation equipment spreading seeds or root fragments.

Intentional Introduction

Intentional introductions have readily happened, as L. dalmatica has been imported into a number of countries for ornamental purposes.

Pathway Causes

Pathway Vectors

Impact Summary

Economic Impact

L. dalmatica asserts an economic impact on productive grassland and rangelands in Canada and the USA. This species becomes dominant as a result of vigorous vegetative growth and competition, especially under conditions of overgrazing and drought, but also perhaps as a result of allelopathy via the action of iridoid glycosides under low nitrogen conditions (Jamieson et al., 2013). Jacobs and Sing (2006) indicate that grass production can be 2.5 times lower in dense infestations of L. dalmatica than in similar areas without the weed. No other quantitative estimates of damage have been seen. It is rarely reported in other crops, except in lucerne/alfalfa (Medicago sativa) and in pine plantations (Pinus species) (Vujnovic and Wein, 2005).

De Clerck-Floate and Turner (2013) refer to its potential toxicity to livestock, though other authors generally suggest that they avoid it. Sing and Peterson (2011) discuss in detail the various metabolites in L. dalmatica and their potential for toxicity to animals. This species can also be a reservoir for crop pathogens including cucumber mosaic virus and it can have a tendency to increase soil erosion. 

Environmental Impact

Impact on Biodiversity

Although L. dalmatica has become dominant in a range of ecologies in North America, including natural habitats, there have been no reports of the serious effects on biodiversity or individual endangered species. Sing and Peterson (2011) make detailed attempts to quantify the environmental risks from L. dalmatica and the levels of the weed which would cause significant reductions. They concluded that more than six L. dalmatica plants per 10 m² could cause a 25% reduction in Hedeoma diffusum. A dry weight of 1,163 kg/ha L. dalmatica is the threshold beyond which there could be 25% reduction in cool season grasses.

Risk and Impact Factors

• Proved invasive outside its native range
• Highly adaptable to different environments
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Pioneering in disturbed areas
• Long lived
• Fast growing
• Has high reproductive potential
• Has propagules that can remain viable for more than one year
• Reproduces asexually
• Has high genetic variability

• Ecosystem change/ habitat alteration
• Host damage
• Monoculture formation
• Negatively impacts agriculture
• Negatively impacts forestry
• Reduced native biodiversity
• Threat to/ loss of native species

• Competition - monopolizing resources
• Competition - smothering
• Pest and disease transmission

• Highly likely to be transported internationally accidentally
• Highly likely to be transported internationally deliberately
• Difficult to identify/detect as a commodity contaminant
• Difficult/costly to control

Uses

Economic Value

Due to its showy flowers, L. dalmatica has been widely cultivated as an ornamental in gardens. There is an early record of it being so used in Italy before 1594 and by the nineteenth century it was present in England and by the end of that century in the USA (Vujnovic and Wein, 2005). It is still commercially available as seed or as root pieces in Europe and Asia. Exotic toadflaxes including L. dalmatica also continue to be sold commercially in the USA, although this is illegal in a number of states where they are listed as noxious (Wilson et al., 2005).

Uses List

Environmental

• Amenity

General

• Sociocultural value

Ornamental

• Propagation material
• Seed trade

Similarities to Other Species/Conditions

L. dalmatica is quite distinct from the co-occurring L. vulgaris, which has narrow strap like leaves. L. genistifolia, which also occurs in the USA, also differs from L. dalmatica in its much narrower leaves and relatively small corolla (2-5 cm), while the seeds are without a winged margin.

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Prevention

SPS Measures

L. dalmatica is listed as noxious in Arizona, California, Colorado, Idaho, Minnesota, Montana, Nevada, North Dakota, New Mexico, Oregon, south Dakota, Utah, Washington and Wyoming, USA (USDA-NRCS, 2015). In British Colombia, Canada, it is listed as noxious in Alberta and Manitoba (De Clerck-Floate and Turner, 2013).

Control

Cultural Control and Sanitary Measures

It is well recognized that L. dalmatica is quick to exploit gaps and weaknesses in established grass stands. It can be kept out of areas by maintenance of a dense healthy stand. Grasses may vary in their effectiveness and in one study of grasses to suppress L. dalmatica, the greatest reduction (95%) was found with Elymus lanceolatus (Rose et al., 2001). However, once established it may not be readily suppressed by newly sown grasses. Early detection is vital for the control of new infestations.

Physical/Mechanical Control

Mowing is relatively ineffective for controlling L. dalmatica as the weed can readily recover from the root systems and the removal of competition from other vegetation may allow it to increase. Similarly burning may encourage growth rather than suppress it. In crop land intensive cultivation for at least two years with at 8-10 cultivations in the first year may be required. Hand pulling can prevent seeding but only annual grubbing and pulling over a 10-15 year period can control it fully (Vujnovic and Wein, 2005).

Biological Control

L. dalmatica has been the subject of a number of biological control programmes. According to De Clerck-Floate and Turner (2013), five insect biocontrol agents have been tested and intentionally introduced into Canada for control of L. dalmatica. The first field release took place in 1963.

Of the five insects released, three have established on L. dalmatica. These include the seed capsule weevil Rhinusa antirrhini, the defoliating moth Calophasia lunula and the stem boring weevil Mecinus janthiniformis. The latter had previously been referred to as ‘M. janthinus’ but differing results from different populations of this ‘species’ led to the realisation that two species were involved. Only M. janthiniformis is effective on this L. dalmatica while L. janthinus is effective on L. vulgaris (Toševski et al., 2011). Toševski et al. (2013) have now described a fast and accurate way of distinguishing the two Mecinus species haplotypes using PCR-RFLP diagnostic assay of the mitochondrial cytochrome oxidase subunit II (COII) gene.

R. antirrhini has established well in British Colombia, Canada and populations are increasing. It is however uncertain whether this seed feeding agent can have a significant impact on the weed (De Clerck-Floate and Turner, 2013).

In Canada, Calophasia lunula has only become established in southern British Colombia and not at all in Alberta, apparently due to low temperatures (De Clerck-Floate and Turner, 2013). Jamieson et al. (2010) found that C. lunula has a mechanism to de-activate the iridoid glycosides, defence mechanisms, in L. dalmatica but ‘M. janthinus’ does not.

M. janthiniformis is by far the most successful biocontrol agent released in Canada. Damage is caused by both adults and larvae and conspicuous reductions in the weed have been recorded. Results are less successful in Alberta almost certainly due to severe winter temperatures. Some disappointing results may also be due to the wrong Mecinus species being released. Weed and Schwarzländer (2014) suggest that the impact of M. janthiniformis may vary considerably due to site specific variation in rainfall and density dependent processes. As a result alternative control methods should be prioritized in areas where herbivore impact is expected to be low. Studies have recently started on two additional Mecinus species, M. laeviceps and M. peterharrisi (Toševski et al., 2016). A petition for the introduction of the stem-galling weevil R. rara n North America is pending.

Two species introduced but not proving successful are the root galling weevil R. linariae which proved to have a preference for L. vulgaris and is no longer being exploited and the root moth Eteobalea intermediella which established in 1998 but had died out by 2002 and is no longer being used (De Clerck-Floate and Turner., 2013). The related E. serratella was thought by Quarles (2007) to be having an impact on L. dalmatica, but Shelton (2015) notes that L. dalmatica is a relatively poor host for this species and it appears that it is no longer of interest.

Grubb et al. (2002) suggested that the naturally occurring flower feeding beetle Brachypterolus pulicarius might prove to be an important component in the integrated weed management of L. dalmatica but that the adults show a distinct preference for L. vulgaris and its further distribution in North America was not recommended (MacKinnon et al., 2007). The seed capsule weevil R. neta also occurs widely in British Colombia, Canada but is not being actively distributed (De Clerck-Floate and Turner 2013).

Wilson et al. (2005) provide extremely detailed and valuable information on the biology of the biocontrol organisms and the planning, implementation and monitoring of biological control programmes against both L. dalmatica and L. vulgaris.

Chemical Control

Phenoxy-propionic herbicides are superior to phenoxyacetic herbicides in controlling L. dalmatica. Satisfactory control can be obtained with fenoprop. Rates of fenoprop required to check or to control the plant did not injure perennial grasses. Dicamba has also been effective however it is resistant to triclopyr and fluroxypy (Vujnovic and Wein, 2005).

Picloram can be effective when applied at various stages of growth, but granules applied to the soil in the autumn can be more effective than a foliar application at the same rate in spring (Vujnovic and Wein, 2005). A combination of fenoprop and picloram has also provided control (Robocker, 1968).

Aminocyclopyrachlor (140 and 280 g ae ha^-1) and aminocyclopyrachlor+chlorsulfuron (140 g/ha+53 g/ha) gave second year control; chlorsulfuron at the dormant stage (105 and 158 g/ha) and aminopyralid at the rosette stage (245 g/ha) also gave 2 years of control. The treatments had only minor effects on grass species (Kyser and DiTomaso, 2013).

Herbicides recommended by USDA Forest Service (2012) for use in non-crop areas include imazapic, glyphosate, rimsulfuron, sulfometuron and chlorsulfuron.

IPM

Quarles (2007) suggests that integrated control of L. dalmatica could involve combinations of clean cultivation, pulling, mowing, burning, grazing, competitive planting, using biological control agents and herbicides. Suggestions for integrated control strategies are also made by the USDA Forest Service (2012). Suggestions include checking hay and straw for the presence of seed, feeding certified weed-free hay pellets to horses, combining control methods and ensuring that follow up treatment is carried out to treat plants that are missed.

Ecosystem Restoration

There are indications that the reduction of L. dalmatica in mixed natural plant communities has tended to encourage the invasion of other equally undesirable weedy species such as Bromus tectorum and Centaurea maculosa [Centaurea podospermifolia].

References

Alex JF, 1962. The taxonomy, history, and distribution of Linaria dalmatica. Canadian Journal of Botany, 40:295-307.

APG, 2003. An update of the angiosperm phylogeny group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society, 141(4):399-436.

Bennett BA; Catling PM; Cody WJ; Argus G, 2010. New records of vascular plants in the Yukon Territory VIII. Canadian Field-Naturalist, 124(1):1-27. http://www.ofnc.ca/cfn/124-1/Bennett_etal.pdf

Blumenthal DM, 2009. Carbon addition interacts with water availability to reduce invasive forb establishment in a semi-arid grassland. Biological Invasions, 11(6):1281-1290. http://www.springerlink.com/content/mt735ugn15j4r857/?p=42b4f64f958e451a81e464b81de7a590&pi=6

Blumenthal DM; Resco V; Morgan JA; Williams DG; LeCain DR; Hardy EM; Pendall E; Bladyka E, 2013. Invasive forb benefits from water savings by native plants and carbon fertilization under elevated CO2 and warming. New Phytologist, 200(4):1156-1165. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1469-8137

Bruns VF, 1965. The effects of fresh water storage on the germination of certain weed seeds. Weeds, 13(1):38-40.

Clerck-Floate R de; Richards KW, 1997. Pollination ecology and biocontrol: developing release strategies for seed-feeding insects on Dalmatian toadflax. In: Acta Horticulturae, No. 437 [ed. by Richards, K. W.]. 379-384.

Clerck-Floate RA de; Turner SC, 2013. Linaria dalmatica (L.) Miller, Dalmatian toadflax (Plantaginaceae). In: Biological control programmes in Canada 2001-2012 [ed. by Mason, P. G.\Gillespie, D. R.]. Wallingford, UK: CABI, 342-353. http://www.cabi.org/cabebooks/ebook/20133355782

Dijkstra FA; Blumenthal D; Morgan JA; LeCain DR; Follett RF, 2010. Elevated CO2 effects on semi-arid grassland plants in relation to water availability and competition. Functional Ecology, 24(5):1152-1161.

Dodge RS; Fulé PZ; Sieg CH, 2008. Dalmatian toadflax (Linaria dalmatica) response to wildfire in a southwestern USA forest. Écoscience, 15(2):213-222. http://www.bioone.org/perlserv/?request=get-current-issue

GBIF, 2015. Global Biodiversity Information Facility. http://www.gbif.org/species

Grieshop MJ; Nowierski RM, 2002. Selected factors affecting seedling recruitment of Dalmatian toadflax. Journal of Range Management, 55(6):612-619.

Grubb RT; Nowierski RM; Sheley RL, 2002. Effects of Brachypterolus pulicarius (L.) (Coleoptera: Nitidulidae) on growth and seed production of Dalmatian toadflax, Linaria genistifolia ssp. dalmatica (L.) Maire and Petitmengin (Scrophulariaceae). Biological Control, 23(2):107-114.

Holm LC; Pancho JV; Herberger JP; Plucknett DL, 1979. A Geographical Atlas of World Weeds. New York: Wiley-Interscience.

Invasive Species South Africa, 2015. Plants A-Z: Flora that is invasive in South Africa. South Africa: Invasive Species South Africa. http://www.invasives.org.za/plants/plants-a-z

Jacobs J; Sing S, 2006. Ecology and management of dalmatian toadflax (Linaria dalmatica (L.) Mill.). Invasive Species Technical Note No. Mt-3. Montana, USA: USDA Natural Resources Conservation Service, 9 pp.

Jacobs JS; Sheley RL, 2003. Prescribed fire effects on dalmation toadflax. Journal of Range Management, 56(2):193-197.

Jamieson MA; Bowers MD, 2010. Iridoid glycoside variation in the invasive plant dalmatian toadflax, Linaria dalmatica (Plantaginaceae), and sequestration by the biological control agent, Calophasia lunula. Journal of Chemical Ecology, 36(1):70-79. http://www.springerlink.com/link.asp?id=104273

Jamieson MA; Quintero C; Blumenthal DM, 2013. Interactive effects of simulated nitrogen deposition and altered precipitation patterns on plant allelochemical concentrations. Journal of Chemical Ecology, 39(9):1204-1208. http://rd.springer.com/journal/10886

Judd WS; Campbell CS; Kellogg EA; Stevens PF, 1999. Plant systematics: a phylogenetic approach. Sunderland, USA: Sinauer Associates Incorporated, xvi + 464 pp.

Judd WS; Campbell CS; Kellogg EA; Stevens PF; Donoghue MJ, 2002. Plant Systematics: a Phylogenetic Approach. Second Edition. Sunderland, Massachusetts, USA: Sinauer Associates, 576 pp.

Kyser GB; DiTomaso JM, 2013. Effect of timing on chemical control of Dalmatian toadflax (Linaria dalmatica) in California. Invasive Plant Science and Management, 6(3):362-370. http://wssajournals.org/loi/ipsm

Leo's Dictionaries, 2015. Dalmatian toadflax., Germany: Rechnerbetriebsgruppe der Fakultät für Informatik of Technische Universität München. http://dict.leo.org/forum/viewWrongentry.php?idThread=1141898&idForum=6&lang=en&lp=ende

MacKinnon DK; Hufbauer RA; Norton AP, 2007. Evaluating host use of an accidentally introduced herbivore on two invasive toadflaxes. Biological Control, 41(2):184-189. http://www.sciencedirect.com/science/journal/10499644

Missouri Botanical Garden, 2015. Tropicos database. St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/

Olmstead RG; Pamphilis CWde; Wolfe AD; Young ND; Elisons WJ; Reeves PA, 2001. Disintegration of the Scropulariaceae. American Journal of Botany, 88:348-361.

Parsons WT; Cuthbertson EG, 1992. Noxious Weeds of Australia. Melbourne, Australia: Inkata Press, 692 pp.

Pollnac FW; Rew LJ, 2014. Life after establishment: factors structuring the success of a mountain invader away from disturbed roadsides. Biological Invasions, 16(8):1689-1698. http://rd.springer.com/article/10.1007/s10530-013-0617-6

Puntieri JG; Brion CAM, 2003. Linaria genistifolia subsp. dalmatica and L. purpurea (Scrophulariaceae), new records for Argentina. (Linaria genistifolia subsp. dalmatica y Linaria purpurea (Scrophulariacae), nuevas citas para la Argentina.) Hickenia, 3(38-47):157-159.

Quarles W, 2007. Integrated management of yellow and Dalmation toadflax. IPM Practitioner, 29(5/6):1-7.

Robocker WC, 1968. Control of dalmatian toadflax. Journal of Range Management, 21(2):94-8.

Robocker WC, 1970. Seed characteristics and seedling emergence of Dalmatian toadflax. Weed Science, 18(6):720-5.

Robocker WC, 1974. Life history, ecology and control of Dalmatian toadflax. Technical Bulletin, Washington Agricultural Experiment Station, No. 79. 20 pp.

Rose KK; Hild AL; Whitson TD; Koch DW; Tassell Lvan, 2001. Competitive effects of cool-season grasses on re-establishment of three weed species. Weed Technology, 15(4):885-891.

Royal Botanic Garden Edinburgh, 2015. Flora Europaea. Edinburgh, UK: Royal Botanic Garden Edinburgh. http://rbg-web2.rbge.org.uk/FE/fe.html

Shelton A, 2015. Biological control: a guide to natural enemies in North America. Ithaca, New York, USA: Cornell University College of Agricultural and Life Sciences. http://www.biocontrol.entomology.cornell.edu/weedfeed/EteobaleaSerratella.php

Sing SE; Peterson RKD, 2011. Assessing environmental risks for established invasive weeds: dalmation (Linaria dalmatica) and yellow (L. vulgaris) toadflax in North America. International Journal of Environmental Research in Public Health, 8(7):2828-2853.

Sing SE; Weaver DK; Nowierski RM; Markin GP, 2008. Long-term field evaluation of Mecinus janthinus releases against Dalmatian toadflax in Montana (USA). In: Proceedings of the XII International Symposium on Biological Control of Weeds, La Grande Motte, France, 22-27 April, 2007 [ed. by Julien, M. H.\Sforza, R.\Bon, M. C.\Evans, H. C.\Hatcher, P. E.\Hinz, H. L.\Rector, B. G.]. Wallingford, UK: CAB International, 620-624. http://www.cabi.org/cabebooks/ebook/20093001887

Stace C, 1991. New Flora of the British Isles. Cambridge, UK: Cambridge University Press.

Toševski I; Caldara R; Jovic J; Hernández-Vera G; Baviera C; Gassmann A; Emerson BC, 2011. Morphological, molecular and biological evidence reveal two cryptic species in Mecinus janthinus Germar (Coleoptera, Curculionidae), a successful biological control agent of Dalmatian toadflax, Linaria dalmatica (Lamiales, Plantaginaceae). Systematic Entomology, 36(4):741-753. http://onlinelibrary.wiley.com/doi/10.1111/j.1365-3113.2011.00593.x/abstract

Toševski I; Gassmann A; Krstic O; Jovic J, 2016. Annual Report 2015. Delémont, Switzerland: CABI Europe-Switzerland, 28.

Toševski I; Jovic J; Krstic O; Gassmann A, 2013. PCR-RFLP-based method for reliable discrimination of cryptic species within Mecinus janthinus species complex (Mecinini, Curculionidae) introduced in North America for biological control of invasive toadflaxes. BioControl, 58(4):563-573. http://rd.springer.com/journal/10526

USDA Forest Service, 2012. Invasive Species: field guide for managing dalmatian and yellow toadflaxes in the Southwest. Albuquerque, New Mexico, USA: USDA-Forest Service. http://www.fs.usda.gov/detailfull/r3/forest-grasslandhealth/invasivespecies/?cid=stelprdb5410251&width=full

USDA-ARS, 2015. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

USDA-NRCS, 2015. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Vujnovic K; Wein RW, 2005. The biology of Canadian weeds. 106. Linaria dalmatica (L.) Mill. Canadian Journal of Plant Science, 77:483-491.

Ward SM; Fleischmann CE; Turner MF; Sing SE, 2009. Hybridization between invasive populations of Dalmatian toadflax (Linaria dalmatica) and yellow toadflax (Linaria vulgaris). Invasive Plant Science and Management, 2(4):369-378. http://www.wssa.net

Weed AS; Schwarzländer M, 2014. Density dependence, precipitation and biological control agent herbivory influence landscape-scale dynamics of the invasive Eurasian plant Linaria dalmatica. Journal of Applied Ecology, 51(3):825-834. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1365-2664

Wilson LM; Sing SE; Piper GL; Hansen RW; Clerck-Floate Rde; MacKinnon DK; Randall CB, 2005. Biology and biological control of dalmatian and yellow toadflax. FHTET-05-13., USA: USDA Forest Service. http://www.invasive.org/weeds/ToadflaxBook.pdf

Zouhar K, 2003. Fire Effects Information System (FEIS)., USA: US Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory. http://www.feis-crs.org/beta/

Distribution References

Bennett B A, Catling P M, Cody W J, Argus G, 2010. New records of vascular plants in the Yukon Territory VIII. Canadian Field-Naturalist. 124 (1), 1-27. http://www.ofnc.ca/cfn/124-1/Bennett_etal.pdf

CABI, Undated. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

GBIF, 2015. Global Biodiversity Information Facility. http://www.gbif.org/species

Holm LC, Pancho JV, Herberger JP, Plucknett DL, 1979. A Geographical Atlas of World Weeds., New York, Wiley-Interscience.

Leo's Dictionaries, 2015. Dalmatian toadflax., Germany: Rechnerbetriebsgruppe der Fakultät für Informatik of Technische Universität München. http://dict.leo.org/forum/viewWrongentry.php?idThread=1141898&idForum=6&lang=en&lp=ende

Parsons W T, Cuthbertson E G, 1992. Noxious weeds of Australia. Melbourne, Australia: Inkarta Press. 692 pp.

Puntieri J G, Brion C A M, 2003. Linaria genistifolia subsp. dalmatica and L. purpurea (Scrophulariaceae), new records for Argentina. (Linaria genistifolia subsp. dalmatica y Linaria purpurea (Scrophulariacae), nuevas citas para la Argentina.). In: Hickenia. 3 (38-47) San Isidro, Argentina: Instituto de Botánica Darwinion. 157-159.

Royal Botanic Garden, 2015. Flora Europaea., Edinburgh, UK: Royal Botanic Garden Edinburgh. http://rbg-web2.rbge.org.uk/FE/fe.html

Stace C, 1991. New Flora of the British Isles. Cambridge, UK: Cambridge University Press.

USDA-ARS, 2015. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

USDA-NRCS, 2015. The PLANTS Database. Greensboro, North Carolina, USA: National Plant Data Team. https://plants.sc.egov.usda.gov

Vujnovic K, Wein RW, 2005. The biology of Canadian weeds. 106. Linaria dalmatica (L.) Mill. In: Canadian Journal of Plant Science, 77 483-491.

Links to Websites

Contributors

13/12/2016 Updated by:

André Gassmann, CABI-CH, Switzerland

12/06/2015 Original text by:

Chris Parker, Consultant, UK

Distribution Maps