Datura innoxia
(downy thorn apple)

Pictures

Picture	Title	Caption	Copyright
Title Flowering habit Caption Datura innoxia (downy thorn apple); flowering habit. Tel Aviv University train station, Tel Aviv, Israel. January 2002. Copyright ©RickP (Rick J. Pelleg)/via wikipedia - CC BY-SA 3.0	Flowering habit	Datura innoxia (downy thorn apple); flowering habit. Tel Aviv University train station, Tel Aviv, Israel. January 2002.	©RickP (Rick J. Pelleg)/via wikipedia - CC BY-SA 3.0
Title Flowering habit Caption Datura innoxia (downy thorn apple); flowering habit. nr. Hyderabad, India. March 2009. Copyright ©J.M. Garg-2009/via wikipedia - CC BY-SA 3.0	Flowering habit	Datura innoxia (downy thorn apple); flowering habit. nr. Hyderabad, India. March 2009.	©J.M. Garg-2009/via wikipedia - CC BY-SA 3.0
Title Flower Caption Datura innoxia (downy thorn apple); close-up of flower. nr. Hyderabad, India. March 2009. Copyright ©J.M. Garg-2009/via wikipedia - CC BY-SA 3.0	Flower	Datura innoxia (downy thorn apple); close-up of flower. nr. Hyderabad, India. March 2009.	©J.M. Garg-2009/via wikipedia - CC BY-SA 3.0
Title Fruiting habit Caption Datura innoxia (downy thorn apple); fruiting habit, with split seedpod. Tel Aviv University train station, Tel Aviv, Israel. December 2005. Copyright ©RickP (Rick J. Pelleg)/via wikipedia - CC BY-SA 3.0	Fruiting habit	Datura innoxia (downy thorn apple); fruiting habit, with split seedpod. Tel Aviv University train station, Tel Aviv, Israel. December 2005.	©RickP (Rick J. Pelleg)/via wikipedia - CC BY-SA 3.0
Title Seeds Caption Datura innoxia (downy thorn apple); seeds. Rambla del Charco, Águilas, Murcia, Spain. August 2011. Copyright ©Philmarin/via wikipedia - CC BY-SA 3.0	Seeds	Datura innoxia (downy thorn apple); seeds. Rambla del Charco, Águilas, Murcia, Spain. August 2011.	©Philmarin/via wikipedia - CC BY-SA 3.0

Identity

Preferred Scientific Name

• Datura innoxia Mill.

Preferred Common Name

• downy thorn apple

Other Scientific Names

• Datura guayaquilensis Kunth
• Datura meteloides DC. ex Dunal
• Datura meteloides Dunal
• Datura velutinosa V.R. Fuentes

International Common Names

• English: angel's trumpet; hoary thorn-apple; Indian-apple; moonapple; sacred datura; thorn-apple
• Spanish: cacaito
• French: datura innocente
• Arabic: datura; tatoora

Local Common Names

• Australia: downy thorn-apple; recurved thorn-apple
• Brazil: datura-européia; trombeta-branca; trombeteira-branca
• China: mao man tuo luo
• Korea, Republic of: teoldogmalpul
• Mexico: toloache
• South Africa: harige stinkblaar
• Sweden: mexikansk spikklubba
• USA: pricklyburr

EPPO code

• DATIN (Datura innoxia)

Summary of Invasiveness

D. innoxia is invasive in the Galapagos islands, New Caledonia, mainland China and Taiwan, and in Ethiopia, Kenya, Namibia and South Africa. It is widely naturalized in the tropics and increasingly in temperate Europe and North America. It is one of several invasive Datura species including D. ferox, D. metel and D. stramonium, that achieve high densities and are difficult to control. It is toxic to animals and humans, with all plant parts and seeds containing toxic alkaloids. Several species of Datura have been spread around the world as contaminants of agricultural seed. Seed inspection and certification agencies should be especially vigilant when batches of seed for planting are imported from areas where Datura species are known to occur. D. innoxia also readily escapes from gardens where it is being cultivated. However, timely cultivation or use of herbicides can easily control the species. D. innoxia has been listed as a Declared Weed in South Africa which is prohibited and must be controlled (AGIS, 2007).

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Solanales
•                         Family: Solanaceae
•                             Genus: Datura
•                                 Species: Datura innoxia

Notes on Taxonomy and Nomenclature

Datura innoxia Mill. is the accepted name in The Plant List (2013) and the Royal Botanic Garden Edinburgh (2014), and is the most commonly used. However, the species name with only a single ‘n’ as D. inoxia is also widely used, e.g. by USDA-ARS (2014), Missouri Botanic Garden (2014), PIER (2014). D. innoxia is the spelling used in this datasheet, with more than 600 records in CAB Direct whereas there are less than 100 for D. inoxia, but this difference in spelling should be taken into consideration when searching the literature.

Two subspecies are described by Missouri Botanic Garden (2014), subsp. quinquecuspidata (Torr.) A.S. Barclay and subsp. lanosa F.A. Barkley. Attention is required with synonymy, as D. fastuosa L. is sometimes reported as a synonym for D. innoxia, but this is incorrect; D. fastuosa is a synonym of D. metel L. (USDA-ARS, 2014). Full descriptions are given by Haegi (1976) of the wild species D. innoxia, D. stramonium, D. ferox, D. leichhardtii, and D. wrightii, and of the cultivated species, D. metel.

D. innoxia is the sister taxon of D. metel next to which is D. wrightii, with D. lanosa being the basal taxon of this group. The combination of genetic and cladistic data indicates that D. innoxia is the most probable progenitor of D. metel (Luna-Cavazos et al., 2009). These are all long-lived, tuberous rooted perennials, in contrast to other species that are tap-rooted annuals.

Phylogenetic analyses by Bye and Sosa (2013) divided species of Datura into three sections, Ceratocaulis, Datura, and Dutra, based upon the position and type of the fruit. The monophyly of the Datura clade was confirmed with two genera, Datura containing herbaceous species and Brugmansia with woody species. The Datura clade consists of two distinct lineages; D. ceratocaula corresponding to the monospecific section Ceratocaulis, and a lineage that gives rise to the other species in a large clade with two groups that do not correspond to the conventional classification of the genus. Species with regularly dehiscent capsular fruits are present in both groups, whereas species with irregularly dehiscent capsules are found only in one group. As section Dutra is polyphyletic, realignment of species within the two traditional sections is suggested: section Datura containing D. arenicola, D. discolor, D. ferox, D. quercifolia, D. kymatocarpa, D. leichhardtii, and D. stramonium; and section Dutra containing D. inoxia, D. lanosa, D. metel, D. reburra, and D. wrightii. The seed margin clearly distinguishes these sections, rather than the combination of fruit position and dehiscence which has been used previously (Bye and Sosa, 2013).

Many common names are applied and misapplied in English. Several species of Datura (and Brugmansia) are known as angel’s trumpet, referring to the distinctive flowers, thorn apple, referring to the spiked fruit, or jimsonweed in North America. However, different species tend to take specific epithets, such as downy thorn, moonapple and others. The genus name Datura is from the Bengali name ‘dhatura’.

Description

D. innoxia is a stout, annual to perennial herb to up to 1 m high, with a spreading crown about 2 m in diameter. Roots may be tuberous. Stems and leaves tomentose, covered with short and soft greyish hairs that give the whole plant a greyish appearance. Mature leaves broadly ovate, the lamina up to 20 cm long, almost entire, slightly sinuate, or irregularly lobed towards base, with conspicuous pinnate venation alternately arranged along the stem. D. innoxia bears white tubular (trumpet) flowers that have green veins, 12-19 cm long, with stigmas well above the anthers. Some people find the flowers fragrant at night, when the plant is blooming. Calyx 5-11 cm long, 3-6-lobed; lobes 13-20 mm long, sometimes incompletely separated. Corolla 12-19 cm long, white with green veins; limb undulate, appearing 10-lobed, alternate lobes broadly triangular or ending in a slender point 5-10 mm long. Stamens not exserted; anthers 8-10 mm long. Style 10-14 cm long; stigma well below anthers. The fruit is a globose or ovoid spiny capsule with numerous slender spines, about 3-5 cm in diameter, deflexed, spiny; spines numerous, slender, sharp, all about equal in length, to 10 mm long; persistent base of calyx to 20 mm long, very prominent. Capsule split when ripe, releasing brown seeds, 4-5 mm long (adapted from PIER, 2014).

Plant Type

Distribution

D. innoxia is native to the tropical and subtropical Americas, from southwestern USA and Mexico, through Central America to northern and western South America, to Peru, and east to Bolivia and Paraguay (USDA-ARS, 2015). Whereas USDA-ARS (2015) noted records from the Caribbean as being possibly naturalized, other sources accept that the species is also native to Caribbean islands.

D. innoxia has been widely introduced and is found throughout the tropics. It is invasive in Namibia (Boyer and Boyer, 1989) and also in temperate areas where introduced as an ornamental but has since naturalized. D. innoxia has spread over Croatia since the early 1980s, occurring in large numbers regenerating from seeds and roots, with past and new localities described by Franjic and Trinajstic (1996). Vukovic et al. (2010) recorded it as invasive but of low occurrence. In the USA it is native only to Texas (USDA-ARS, 2014), but is also recorded in the southwestern states, Arkansas and in the northeast USA as far as Ontario and Quebec, Canada (USDA-NRCS, 2014).

D. innoxia was noted as an invasive alien species in Namibia by Boyer and Boyer (1989) but they observed that it offered less of a threat to native vegetation as compared to Prosopis spp., Nicotiana glauca and Ricinus communis, and that control of these latter species should take priority. Two decades later the situation did not appear to have altered greatly, as Shapaka et al. (2008) identified D. innoxia as one of eight invasive species and active management was recommended, but it was of lower priority as compared to the others. See also Katjirua (2008).

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.

Risk of Introduction

Several species of Datura have been spread around the world as contaminants of agricultural seed. Seed inspection and certification agencies should be especially vigilant when batches of seed for planting are imported from areas where Datura species are known to occur. D. stramonium was introduced into parts of Victoria, Australia, in contaminated safflower seed, and contaminated soyabeans have resulted in new infestations in Norway on several occasions. It is particularly difficult to separate from sorghum seed which is of a similar size (Parsons and Cuthbertson, 1992). Legislation in Queensland, Australia, prohibits the sale of crop seed which is contaminated by D. ferox or D. stramonium (Genn, 1987). D. innoxia was also one of five species recommended for eradication from Isabella Island in the Galapagos archipelago along with the closely-related D. metal (Guézou et al., 2007). It is declared as category 1, prohibited weed in South Africa because of its aggressive growth habit (AGIS, 2007). PIER (2014) gives a very high risk assessment score of 21 for this species in the Pacific region.

Habitat

D. innoxia grows naturally in disturbed areas such as eroded areas, uncultivated fields, vacant lots, overgrazed pastures and rangeland, roadsides, abandoned roadbeds, and fencerows. Disturbance and reduced competition appear to be required for D. innoxia to become established (PIER, 2014). In China it is found growing near villages and on roadsides but is also cultivated (PIER, 2014).

Habitat List

Biology and Ecology

Genetics

A single ploidy level of 2n = 24 for D. innoxia in Mexico was reported by Palominl et al. (1988) and many others. However, colchicine-induced tetraploids have been produced which exceeded the diploids in atropine and scopolamine content and showed better adaptability (Dzhurmanski and Yankulov, 1981). Different levels of ploidy (2n, 4n, 6n) was reported by Badea and Raicu (1983), and haploid, diploid, triploid, tetraploid and hexaploid cytotypes by Gupta et al. (1984).

Reproductive Biology

Reproductive biology of D. innoxia is described in Henderson (2001) and Parsons and Cuthbertson (1992). Datura species, including D. innoxia, may flower overnight and be pollinated by moths. Hummingbirds sometimes also visit the flowers but are affected by the alkaloids in the nectar and must limit their consumption, whereas honeybees are apparently unaffected.

Physiology and Phenology

In the tropics, D. innoxia is a perennial species, but it will behave as an annual in temperate regions. D. innoxia can flower and fruit throughout the year in some environments but only from July to September in India (Parrotta 2001) and September through November in Nicaragua (Stevens et al., 2001). Germination in D. innoxia is slow, poor and variable due to the presence of an inhibitor in the seed coat (Singh, 1974).

Environmental Requirements

D. innoxia grows in a wide variety of tropic and temperate climates. It is found on a wide variety of soils but prefers well-drained soils, on both igneous and sedimentary parent materials. In Puerto Rico, the species grows naturally in areas with 750-1000 mm mean annual precipitation, and from near sea level up to 900 m altitude (Stevens et al., 2001). An after-ripening period of 5-11 months may be required (Parsons and Cuthbertson, 1992) In a study of the effects of temperature and light on germination, maximum germination was obtained under continuous light at alternating temperatures of 20 and 30°C. Germination was reduced under continuous darkness (Barakat et al., 2013).

Climate

Rainfall

Rainfall Regime

Soil Tolerances

Soil drainage

• free

Soil reaction

• acid
• alkaline
• neutral

Soil texture

• light
• medium

Notes on Natural Enemies

A number of diseases have been reported on D. innoxia, including Sclerotium root rot (Thakur and Sastri, 1970), P. nicotianae (reported for the first time in India) (Sattar et al., 1982), Alternaria crassa leaf spot (Grzybowska and Kapala, 1976) and other Alternaria blight (Janardhanan and Husain, 1973). Various viruses have been reported (Garga, 1958), including leaf curl disease (Marwal et al., 2012), virus wilt (Kovachevski, 1974), figwort mosaic virus (Shepherd et al., 1987), and cucumber mosaic virus (Raj et al., 1999). D. innoxia is also a host of the nematode, Xanthomonas vesicatoria (Raj and Moniz, 1975).

Means of Movement and Dispersal

Natural Dispersal

Datura species are thought to spread solely by seed which it produces in large numbers. Both seed capsules and seeds float on water, providing an effective means of dispersal. However, spread by broken stem fragments may also be a minor means of dispersal.

Vector Transmission (Biotic)

The spiny capsule can become attached to the fur of animals, which split open and disperse the seed.

Accidental Introduction

In New Zealand, Datura seeds have been reported to be mainly spread by contamination of agricultural seeds, or by machinery and vehicles, and it is believed to have originated from imported birdseed (Webb et al., 1988). D. innoxia readily escapes from gardens where it is being cultivated.

Intentional Introduction

As ornamental plants, many Datura species have been intentionally introduced around the world, and continue to be grown as such, especially in Europe, North America and China.

Pathway Causes

Pathway Vectors

Plant Trade

Impact Summary

Economic Impact

No economic purpose for the plant has been reported. In some countries it is prohibited to buy, sell or cultivate Datura plants (Henderson, 2001). D. innoxia commonly occurs in a range of crops, including maize, groundnuts, sorghum and temperate cereals, where it may have competitive and allelopathic affects; however, no estimates of economic loss are available.

Environmental Impact

Allelopathic effects of D. innoxia have beenidentified (Hussain et al., 1979). D. innoxia can be found in two national parks in Namibia, Namib Naukluft Park and Daan Viljoen Game Park (Boyer and Boyer, 1989; Shapaka et al., 2008) and in Medvednica Nature Park in Croatia (Vukovic et al., 2010).

Social Impact

If ingested by humans and other animals (including livestock and pets) it can cause fatalities, as all parts of Datura plants are poisonous (Henderson, 2001).

D. innoxia belongs to the family of Solanaceae and contains two main toxic alkaloids, atropine and scopolamine. Seven individuals were admitted to hospital in Athens with an anticholinergic syndrome after eating cooked vegetables (blites, members of Chenopodiaceae), but among the vegetables there was also D. innoxia, which has a similar appearance to blites (Papoutsis et al., 2010). Goetz et al. (2003) report the clinical findings in 14 adolescents in Ohio, USA who became ill after intentional ingestion of toxic seeds identified as D. innoxia. Poisoning was also reported (Marciniak and Sikorski, 1972) in a family who had eaten honey from an apiary near a plantation of D. innoxia and D. stramonium, with scopolamine and hyoscine alkaloids detected in the honey and in the urine of the patients.

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly adaptable to different environments
• Is a habitat generalist
• Pioneering in disturbed areas
• Highly mobile locally
• Fast growing
• Has high reproductive potential
• Has propagules that can remain viable for more than one year
• Has high genetic variability

• Monoculture formation
• Negatively impacts human health
• Negatively impacts animal health
• Reduced native biodiversity
• Threat to/ loss of native species

• Allelopathic
• Competition - monopolizing resources
• Competition - shading
• Poisoning
• Rapid growth
• Produces spines, thorns or burrs

• Highly likely to be transported internationally accidentally
• Highly likely to be transported internationally deliberately
• Highly likely to be transported internationally illegally
• Difficult to identify/detect as a commodity contaminant
• Difficult to identify/detect in the field

Uses

Species of Datura have been used in many parts of the world as narcotics, constituents of folk remedies and shamanistic rituals (Schultes and Hofmann. 1992), and as a source of atropine, hyoscamine and hyoscine alkaloids for pharmacological use in both human and veterinary medicine (Watt and Breyer-Brandwijk, 1962). D. innoxia has been used to treat impotence, asthma, and diarrhoea, as an analgesic, to control fever, to kill parasites, and as a drug for criminal purposes (Parrotta 2001). There has also been considerable interest in the use of extracts of Datura species as botanical pesticides (Oduor–Owino, 1993). El-Bazaoui et al. (2012) identified 53 alkaloids in the roots, stems, leaves, flowers and seeds of D. innoxia – 17 reported for the first time for this species. Also, alkaloid contents for D. innoxia were reported to be: leaves 0.39%, roots 0.54%, and seeds 0.44% (Kosova and Chladek, 1957).

Uses List

Drugs, stimulants, social uses

• Hallucinogen
• Narcotic
• Psychoactive

General

• Ornamental

Medicinal, pharmaceutical

• Source of medicine/pharmaceutical
• Traditional/folklore

Similarities to Other Species/Conditions

Datura species are annual, whereas members of the closely related genus Brugmansia are perennials. However, D. innoxia can be confused with species of both genera, bearing similarly distinctive flowers and fruit. Species of both genera have a trumpet shaped flowers, but those of Brugmansia point downwards with a sweet fragrance, with Datura flowers often pointing upward with spicy or lemony fragrance.

D. innoxia is especially misidentified with other invasive Datura species, including D. ferox, common thornapple (D. stramonium L.), downy thornapple (D. inoxia Mill. = D. innoxia Mill.), hoary thornapple (D. metel L.) and hairy thornapple (D. wrightii Rogel). D.stramonium and D.ferox produce erect fruit capsules on straight stalks with black to greyish seeds, whereas fruit capsules of D. leichhardtii, D. metel, D. inoxia, D. wrightii appear on curved stalks with brown to yellow seeds.

Morphological comparisons between D. innoxia and D. metel showed that D. innoxia could be distinguished by much greater plant height, deeper leaf indentations, larger capsules, a wider corolla (when slit on one edge and rolled out flat), a wider calyx, and heavier and sturdier roots (Kosova and Chladek, 1957). D. innoxia is also reported to be similar to D. metel in habit, but D. innoxia is differentiated by its pubescent whereas D. metel is glabrous.

Parsons and Cuthbertson (1992) distinguish D. innoxia from most other weedy Datura species by the combination of curved seed-stalk, brown to yellow seeds, and hairy leaves; and from D. wrightii by its erect, glandular hairs and pure whiter flowers (lavender-tinged in D. wrightii).

Full descriptions are given by Haegi (1976) of the wild species D. inoxia, D. stramonium, D. ferox, D. leichhardtii, and D. wrightii, and of the cultivated species, D. metel.

Prevention and Control

Control

Physical/mechanical control

Isolated plants should be hand-pulled before they set seed, whereas larger areas of infestation are controlled by cultivation when weeds are in the seedling stage. Cultivation becomes less effective as plants mature – stems become woodier and roots may not become completely severed. Seedlings emerge over a long period of time so repeated cultivations may be necessary to reduce the level of infestation (Parsons and Cuthbertson, 1992). However, tillage may promote seed survival as seeds decay more rapidly on the soil surface than when buried, and there is greater loss of seed to predators under no-till than conventional tillage systems.

Chemical control

D. innoxia can be controlled with 2,4-D in the seedling stage and young growth stages, but become resistant as they mature (Parsons and Cuthbertson, 2001). Other non-selective herbicides include atrazine, diquat, paraquat and glyphosate. The following herbicides are used in specific crops: acifluorfen and bentazone in soybeans and peanuts; 2,4-DB in certain varieties of peanuts; dicamba in grain sorghum and maize; metolachlor in maize; and picloram + 2,4-D in summer cereals (Parsons and Cuthbertson, 2001).

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Contributors

12/11/2014 Original text by:

Nick Pasiecznik, Consultant, France

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