Erodium botrys (long-beaked stork's bill)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Threatened Species
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Erodium botrys (Cav.) Bertol.
Preferred Common Name
- long-beaked stork's bill
Other Scientific Names
- Geranium botrys Cav.
International Common Names
- English: broadleaf filaree
Local Common Names
- English: big heron’s bill; corkscrews; long-beak crowfoot; Mediterranean storksbill; wild geranium
- Spanish: agullots (Catalan); alfileres (Castillian); alfileteros (Castillian); cargola d'agullots (Catalan); ciñueloñ relojes (Castillian)
- France: érodium en grappe
- Germany: Reiherschnabel
- Italy: becco di grù botri
- Spain: relojitos
Summary of InvasivenessTop of page
E. botrys is a winter growing annual originally native to the Mediterranean. It is considered invasive in California and parts of Australia. It is thought to present a significant threat to one or more native plant communities in Victoria, Australia. Elsewhere in southern Australia, it is a common weed of native pastures, open woodlands and grazed grasslands (Weeds of Australia, 2013). In California, E. botrys and annual grasses of Mediterranean origin have largely supplanted the perennial grasses of the pristine Californian prairies (McCown and Williams, 1968).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Dicotyledonae
- Order: Geraniales
- Family: Geraniaceae
- Genus: Erodium
- Species: Erodium botrys
Notes on Taxonomy and NomenclatureTop of page
There are about 60-70 species of Erodium.
DescriptionTop of page
Modified slightly from Webb et al. (1988):
Annual or possibly biennial herb, often reddish; stems initially absent, later several, to about 18 cm long, more or less prostrate, not musk-scented, with dense retrorse stiff hairs. Petioles of basal and lower cauline leaves 2.5-5.5 cm long. Lamina 6-10 × 2.5-3 cm, oblong-ovate or triangular-ovate, cordate at base, pinnatisect; lateral lobes extending to over 3/4 way to hairy midrib, more or less ovate, ciliate, otherwise almost glabrous, acutely lobed; terminal lobe usually larger. Upper cauline leaves somewhat smaller, subsessile or with short petiole; lamina with narrow-oblong, sharply acute lobes. Stipules membranous, triangular-ovate, densely ciliate, silvery, often tinged red, obtuse. Umbels 2-3-flowered; bracts similar to stipules but smaller. Peduncles and pedicels glandular-hairy, somewhat reddish, equal to or greater than the calyx at anthesis. Sepals 5-7.5 mm long, ovate or elliptic-ovate, with simple and glandular hairs or glabrate; apical mucro dark. Petals 9-10 mm long; pinkish mauve with dark crimson vein in lower part; claw very short; limb obovate. Stamens 3.5-4.5 mm long, much widened in lower 1/2; anthers dark purple. Staminodes narrower and much smaller than stamens. Fruit beak (7)-7.5-8.5-(9) cm long, with antrorse hairs. Mericarps 7-8 mm long, with dense, antrorse, stiff hairs; apical pits eglandular, with 2 prominent furrows beneath.
Plant TypeTop of page
DistributionTop of page
E. botrys is native to the wider Mediterranean region and much of Europe. It has been introduced to South Africa, USA, Australia and New Zealand. In the USA it is limited to California, Massachusetts, Maine, Oregon, Texas and Vermont (USDA-NRCS, 2013). It has also been recorded as a ‘wool alien’ in Belgium (Alien plants of Belgium, 2013) and Britain (Salisbury, 1961; Shimwell, 2006), but has never become fully naturalized or spread in either country. The species is considered rare in crops in France and is protected in Bretagne (Hyppa, 2007).
Distribution TableTop of page
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.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|South Africa||Present, Few occurrences||Introduced||Elandsberg flats: lower slopes (Western Cape)|
|Belgium||Present, Few occurrences||Introduced||Rare and much decreasing, always ephemeral alien|
|Federal Republic of Yugoslavia||Present||Native|
|Russia||Present||Present based on regional distribution.|
|United Kingdom||Present, Few occurrences||Introduced||Wool alien|
|United States||Present||Present based on regional distribution.|
|-New South Wales||Present||Introduced||Invasive|
|-Northern Territory||Present||Introduced||Invasive||Southern parts|
|-Western Australia||Present||Introduced||Invasive||Southern parts|
|New Zealand||Present, Few occurrences||Introduced|
History of Introduction and SpreadTop of page
Originally from the Mediterranean region, E. botrys has become naturalized in the USA, Australia and New Zealand. In Australia it was first recorded in 1865 (Australia’s Virtual Herbarium, 2013), but it was not found in New Zealand until 1985 (New Zealand Plants Database, 2013). Stamp (1989) suggested the species was introduced to California during Spanish colonization.
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Australia||1865||Yes||No||Australia’s Virtual Herbarium (2013); Royal Botanic Gardens Sydney (2004); Royal Botanic Gardens Sydney (2013)||Victoria|
|Belgium||1826||No||No||Euro+Med Plantbase (2009); Euro+Med Plantbase (2013)||Accidental introduction, probably in wool. Wool alien in the valley of the river Vesdre near Verviers|
|New Zealand||1985||Yes||No||New Zealand Plants Database (2013)||Accidental introduction, probably from Australia. Near lucerne paddocks in the Upper Clutha Valley|
Risk of IntroductionTop of page
E. botrys has been recorded as a ‘wool alien’ in Belgium (Alien plants of Belgium, 2013) and Britain (Salisbury, 1961; Shimwell, 2006). Wool aliens or wool casuals are plant species taken to northern Europe in sheep or goat wool from elsewhere. This wool was sent to Europe to be processed into clothing, and waste from the mills was formerly distributed to agricultural areas as a slow-acting organic manure (Salisbury, 1961) or as bedding for livestock (Shimwell, 2006).
The risk of introduction to new countries in agricultural seed or raw wool may still be high, although patterns of the world wool trade have changed dramatically in the last hundred years. Strict control of wool and seed imports through phytosanitary regulations and inspections ought to restrict such spread.
Changing global climates may increase the chances of the naturalization and spread of species like E. botrys in countries such as Belgium and Britain, where it occurs as a casual plant (Shimwell, 2006).
HabitatTop of page
Fiz et al. (2006) described the habitat of E. botrys (presumably in Spain and other Mediterranean countries) as ‘widespread, cultivated places and disturbed sites, in sandy acid soils, 0–2000 m.’ Flora of Israel Online (2013) described its native habitat as ‘Mediterranean woodlands and shrublands’.
In California, it dominates the valley annual grasslands and exists over a range of soil types and rainfall regimes (Gordon and Rice, 1992).In parts of California complete roadside dominance of E. botrys is probably the result of roadside mowing, especially when the carpels were dehiscing (Frenkel, 1977). By contrast, the adjacent, unmown, mixed annual grassland is free of the species.
In Australia E. botrys is widespread as a weed of native pastures, open woodlands and grasslands in southern Australia (Weeds of Australia, 2013).
Habitat ListTop of page
|Terrestrial||Managed||Managed grasslands (grazing systems)||Principal habitat|
|Terrestrial||Managed||Disturbed areas||Principal habitat|
Biology and EcologyTop of page
2n = 40 (Fiz et al., 2013), although the same authors also report a hexaploid with 2n=60.
E. botrys is a winter growing annual whose seeds germinate in late summer or early autumn. Stamp (1989) described the dispersal of seeds of the four introduced species of Erodium found in Californian grasslands, observing that all the species have ballistic dispersal, followed by hygroscopic activity of the awned diaspore, which facilitates burial. When dry the awn becomes tightly coiled and when wet it uncoils. Advantages of this self-burial include protection from granivorous rodents and birds, ensuring some seeds remain dormant, avoiding fire, and providing better conditions for germination (Stamp, 1989). The diaspores of E. botrys are heavier, longer, contain heavier seeds and have longer uncoiling and recoiling times for the awns than other species, including the closely related E. brachycarpum (Stamp, 1989). E. botrys can throw its seeds an average of 76.2 cm on dispersal, although surrounding vegetation would effectively reduce this distance.
Self-burial of E. botrys disapores was ineffective on sand and only slightly better on gravel. However, under field conditions 51.3 % of carpels buried themselves successfully. Even so, many diaspores remained on the soil surface in June, a month after the end of seed production and two months after the end of the rainy season. One effect of such a mechanism may simply be to better anchor the seed in the soil and thus help penetration of the root into the soil (Stamp, 1989).
In earlier work Stamp (1984) indicated that E. botrys awns reacted to moisture much more slowly than those of E. cicutarium, which may be because E. botrys grows in wetter habitats, where rainfall is more prolonged and soil surfaces are wetter.
In Australia E. botrys is a winter-growing annual which germinates after good late summer or autumn rains. It then flourishes, flowers and sets seed in early spring. After maturing, the plants break up quickly, leaving virtually no feed or cover over summer (Schroder, 1998). Brown and Bettink (2011) claimed that seeds can germinate at any time of the year in Australia, depending on soil moisture conditions, but that germination usually takes place during April to June (autumn). They say that active growth is from June to November (late autumn to late spring) and flowering from August to November (late winter to spring).
In California, Rice (1985) reported that germination begins with the onset of rain in October, although some germination can occur after late summer rain. Plants overwinter as rosettes and initiate flowering in late February and early March; seeds are dispersed from late March to early June and remain dormant over summer. Also in California, when Gordon and Rice (1992) sowed E. botrys outdoors in November, seedlings emerged by mid-December and the plants flowered from late March through to mid-June. In late June, the authors could not collect shoots of the plants ‘because much of the leaf material had senesced during flowering’.
Physiology and Phenology
The seedlings of E. botrys are very tolerant of dry conditions (Schroder, 1998). The rate of root elongation and depth of penetration for E. botrys are certainly much greater than for the associated species Trifolium subterraneum or Bromus mollis (Ozanne et al., 1965; McCown and Williams, 1968). Gerakis et al. (1975) observed that, although E. botrys had the highest relative water content, transpired the least, and had the lowest transpiration ratio of three associated species grown in pots (the others being B. mollis and T. subterraneum), it appeared to be similar to B. mollis in its sensitivity to drought. However, the authors went on to suggest that the pronounced wilting of E. botrys, even when moisture was only slightly deficient, could be advantageous as a survival mechanism in tolerating drought.
After a short after-ripening period, seed dormancy is due to the water-impermeable seed coat (Rice, 1985). After moistening and placing at a constant 25° C, germination was higher for seeds that had been previously exposed, whilst dry, to alternating temperatures (25° and 45° C) for four months, than for similar seeds held at a constant 25°C. Seeds germinated equally well whether exposed to 12 hour photoperiods or kept in the dark.
The plants themselves are short-lived winter annuals, but the seeds, according to Brown and Betterik (2011) can last for three years or more. Some seeds of the related species E. cicutarium also survived for over three years in the soil even under good conditions for germination (Roberts, 1986).
Population Size and Structure
In Californian annual grasslands, Heady (1958) recorded the variation in population size of E. botrys and other species at three sites: 1. ungrazed by domestic livestock for several years beforehand; 2. where sheep grazing was moderate, and 3. in a grazed area near where sheep were fed in winter. Few plants of E. botrys were present in site 1 and most were in site 2. Numbers decreased through the year as young seedlings disappeared. E. botrys numbers during flowering and fruiting in June varied quite significantly from year to year.
Annual rangeland soils in California are often deficient in sulphur, and McKell et al. (1971) tested the addition of sulphur at three temperatures to monocultures and mixtures of E. botrys, Bromus mollis and Trifolium hirtum under controlled environment conditions. In sulphur-deficient soil the yields of all three species were similar, and tended to increase with temperature. Adding sulphur increased yields of all three species; T. hirtum responded most and B. mollis least. At higher sulphur levels the effect of increased temperatures was more marked. In mixtures of the species, T. hirtum gained more advantage than the other species both in growth and sulphur uptake.
McCown and Williams (1968) pointed out that Bromus and other tall grasses dominate the annual grasslands of California when fertilized, but that in the more typical infertile conditions species of Erodium dominate. They grew plants of B. mollis and E. botrys either under controlled conditions or outdoors in pots and compared growth of the two species in monocultures and mixtures at three levels of sulphur and two levels of nitrogen. At low levels of sulphur Erodium grew much better in mixtures than in monoculture, at least partly because of its superior early root growth. However, at high sulphur levels Bromus was more competitive as its population density increased and when the ratio of Bromus to Erodium plants was 192:3, the Erodium was almost entirely eliminated.
Gerakis et al. (1975) examined the water relations and nutrition of E. botrys, B. mollis and T. subterraneum at different plant densities in pots in a glasshouse. Although all three species tended to show increased mineral concentrations (of N, P, K, Ca, Mg) in their tissues in relation to greater moisture stress, this was most marked in E. botrys. The authors speculated that this accumulation of nutrients may afford some competitive advantage to Erodium in less favourable nutrient environments, giving the plants the ability to use the stored nutrients for rapid growth when temperatures rise in the spring. Another possibility is that the stored nutrients give the plants a high osmotic potential for the rapid uptake of water.
In Australia, E. botrys tends to colonise areas that are relatively bare in autumn, so it is often found where hay has been cut, around livestock camps and along ridges where livestock like to graze, and in paddocks that have been heavily grazed over summer (Schroder, 1998). In Californian annual grasslands, E. botrys grows in association with many other annuals of Mediterranean origin, such as the grasses Aira caryophyllea, Bromus spp., Festuca spp., Hordeum hystrix and Poa annua (Heady, 1958). Along with Bromus diandrus it can dominate the valley annual grasslands and exists over a range of soil types and rainfall regimes (Gordon and Rice, 1992).
A major requirement for the germination and establishment of E. botrys is disturbed ground, relatively free of cover from other species (Rice, 1987; Schroder, 1988). Once the seeds germinate, root growth is rapid and the developing broad rosette leaves can suppress other, later-germinating plants. The species is most competitive against associated grasses and legumes at low levels of sulphur (McKell et al., 1971; McCown and Williams, 1968); at high sulphur levels it is more likely to be outcompeted by grasses.
ClimateTop of page
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
Notes on Natural EnemiesTop of page
According to Plant Viruses Online (2013), E. botrys, E. moschatum and E. cicutarium are susceptible to filaree red leaf luteovirus. Symptoms include vein clearing, with immature leaves cupped and chlorotic, and older leaves red and brittle. The plants are stunted, with no flowers.
Ants and rodents show strong preference for seeds of Erodium spp (Stamp, 1989). Rice (1987) suggested that high populations of Californian vole (Microtus californicus) probably help reduce or eliminate populations and growth of E. botrys and E. brachycarpum in annual Californian grasslands ungrazed by domestic stock. Sheep grazing in turn has a strong negative effect on small mammal populations because less litter accumulates. Therefore Erodium spp. persist under sheep grazing but not in its absence.
Means of Movement and DispersalTop of page
Natural Dispersal (Non-Biotic)
E. botrys uses ballistic dispersal of seeds. The distances travelled by seeds this way are small and transport further afield depends on animal or human assistance.
Vector Transmission (Biotic)
The spiral corkscrew of the awn allow the propagules to become attached to the wool or fur of animals and, in the case of sheep, the seeds can sometimes work their way into the skin (Schroder, 1998). Carriage of sheep from Europe to America and Australia probably introduced E. botrys to these continents, although seeds may also have been carried in hay, straw and bedding taken to feed and care for sheep.
Ants and rodents show a strong preference for seeds of Erodium spp. (Stamp, 1989) and may help disperse them.
The early introduction of Erodium species to the Americas and Australia was probably accidentally in the wool of transported sheep or in the hay, straw and bedding used to support the animals on their long sea voyages. In more recent years, seeds may also have been accidentally introduced (to New Zealand, for example) in agricultural seed (New Zealand Plants Database, 2013). This is the most likely route by which the species reaches new localities, although phytosanitary regulations and inspections ought to prevent this.
Intentional introduction of the species is unlikely.
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
|Economic/livelihood||Positive and negative|
Economic ImpactTop of page
E. botrys invades grazed grassland in both California and Australia. Although it produces some fodder in winter and early spring, when it flowers and seeds in the spring its leaves wither and die (Schroder, 1988). Its propagules can become attached to wool and can even penetrate skin and flesh of grazing animals, reducing the value of both wool and skins (Schroder, 1988). However, the Australian Wool Testing Authority (2013) reported that its seeds are easier to remove from harvested wool than those of Aristida spp. or Stipa spp.
Connor (1997) mentioned photosensitization in lambs and calves, believed to be caused by species of Erodium, in both Australia and New Zealand, although no photosensitizing agent was identified. Stroebel (2002) found that the related species E. moschatum can induce photosensitivity in sheep if ingested in large quantities.
E. cicutarium and probably E. botrys have prostrate stems that, according to Howard (1992), help spread ground fires, and dead plants contribute to fuel loads.
Environmental ImpactTop of page
E. botrys is but one of a large number of Mediterranean plant species that have become established in both California and Australia. Together these species dominate Californian grasslands, in both density and cover, having largely displaced but not eliminated the native American plants originally present (Bartolome, 1979).
In Victoria, Australia, E. botrys is widely regarded as an environmental weed, and it is widespread as a weed of native pastures, open woodlands and grasslands across southern Australia. It is thought to pose a serious threat to one or more native plant communities in Victoria (Weeds of Australia, 2013).
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Pseudobahia bahiifolia (Hartweg's golden sunburst)||NatureServe; USA ESA listing as endangered species||California||Competition - smothering||US Fish and Wildlife Service (2007)|
Risk and Impact FactorsTop of page
- Proved invasive outside its native range
- Pioneering in disturbed areas
- Fast growing
- Has high reproductive potential
- Competition - smothering
- Highly likely to be transported internationally accidentally
UsesTop of page
The related E. cicutarium is important forage for cattle, horses and domestic sheep in California, Nevada and Arizona, and the same probably applies to E. botrys (Howard, 1992). Rosiere (1987) reported that E. cicutarium and E. botrys declined in woodland but increased in grassland as grazing intensified. Both species tend to fluctuate widely from year to year in their contribution to pasture production.
Presumably the contribution of E. botrys to grazing means that it has some economic value, but probably less than that of the species it replaces.
Uses ListTop of page
Animal feed, fodder, forage
- Fodder/animal feed
Similarities to Other Species/ConditionsTop of page
Webb et al. (1988) pointed out that E. botrys is close to E. brachycarpum (Godron) Thell., a segregate species which grows with E. botrys in Australia, Chile and California and is sometimes confused with it. E. brachycarpum has shorter fruit beaks and mericarps. The mericarp has only one shallow hairy pit on either side of the awn base, whilst E. botrys has two to three hairless furrows on each side (Richardson et al., 2006). E. botrys can be distinguished from the related E. cicutarium by its fruit beak, which in E. botrys is usually 7.5-8.5 cm long, compared with 3.0-3.5 cm long in E. cicutarium.
The rate of root elongation and depth of penetration for E. botrys are much greater than for the associated species Trifolium subterraneum or Bromus mollis (Ozanne et al., 1965; McCown and Williams, 1968).
Prevention and ControlTop of page
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.
Where E. botrys grows in grazed pastures, grazing management and fertilizer applications can reduce its growth to some extent. Herbicides can also be used, but these are rarely completely selective to desirable species.
Physical control of isolated plants or cultivation of extensive populations would presumably give control, but the cultivation or physical removal would probably need to remove the fairly deep tap root, and buried seeds may be a residual problem.
Apparently no biological control has been used for any Erodium species.
Brown and Bettink (2011) recommend the use of clopyralid for control of E. botrys in grassland in Western Australia.
The Department of Agriculture and Food Western Australia (2013) lists a number of herbicides for the control of E. botrys and other species of Erodium in cereal crops (wheat, barley, oats and triticum), available at: http://archive.agric.wa.gov.au/objtwr/imported_assets/content/pw/weed/erodium20110510_web.pdf.
Especially in grassland or pastures, chemical control of E. botrys is rarely useful because herbicides will also damage useful species as well as weedy ones. Integrated control is much more likely to be successful (see IPM).
Schroder (1998), in Australia, reiterated the old rule that a dense competitive pasture is the best way to deal with this and other species. He suggested sowing early maturing cultivars of subterranean clover, which have a high seed yield and high hard seed content. Such cultivars can flower and set copious seed before the dry weather starts in spring, leading to a dense germination of clover to compete with E. botrys and other undesirable species. Even better is to establish dense perennial-based pastures of species like lucerne (Medicago sativa), phalaris (Phalarisaquatica) or cocksfoot (Dactylis glomerata). With such species, ensuring a dense cover in autumn is important to prevent invasion by E. botrys.
Control by Utilization
E. botrys can be grazed by livestock, but low fertility conditions and grazing are likely to promote the growth of E. botrys at the expense of other grassland species (McCown and Williams, 1968). Broom and Arnold (1986) observed that Merino sheep grazing annual pastures at the start of the autumn growing period completely avoided E. botrys growing in the presence of Lolium rigidum and Trifolium subterrraneum. This gave E. botrys a competitive advantage over the other species.
ReferencesTop of page
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Webb C J, Sykes W R, Garnock-Jones P J, 1988. Flora of New Zealand Volume IV: Naturalized Pteridophytes, Gymnosperms, Dicotyledons. Christchurch, New Zealand: Department of Scientific and Industrial Research. 1365 pp.
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20/05/13: Original text by:
Ian Popay, consultant, New Zealand, with the support of Landcare Research.
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