Many popular sources and even scientific data aggregators, including Plants of the World Online (PoWo 2021), stated that this species is native to Eurasia with the continuous distribution from Portugal to Japan. Palaeobotanical data definitely show that the species was present in Europe as early as in Pliocene (Särkinen et al. 2013), but this evidence does not indicate its continuous presence in the territory. As evident from the details of its distribution in particular countries, the species is native in two disjunct areas: the Caucasus (Grossheim 1967) and central China (Li 1973) with adjacent territories.

The species was a common vegetable in pre-historic times (Colledge and Conolly 2014). For this reason, it had been transported with people as they settled in new territories since the Neolithic period (e.g. Kohler-Schneider and Caneppele 2007, Reed 2015, Jin et al. 2020). With humans, it expanded as an archaeophyte to Europe, Central Asia and neighbouring mountainous areas (including Xinjiang). Its occurrence in the Russian Far East (Ignatov 1991) originated from the ancient Chinese colonisation (Schischkin 1936). Its non-native status in Central Asia was established by Pojarkova (1954a).

The species is a neophyte outside Eurasia, in North America and northern Africa.

The species is widely distributed in Central Asia and has been recorded from every country of the region (Kovalevskaya 1987). It was commonly cultivated before the Russian colonisation (Fedtschenko and Fedtschenko 1913) and occurred spontaneously in gardens and around populated places.

Due to a technical error, P. viscosa L. was reported as historically occurring in Uzbekistan (Khassanov et al. 2020). This record was based on a misfiled collection of A. officinarum from Tashkent (cultivated or weedy), dated 1919.

Western Tian-Shan, Northern Tian-Shan, Alay-Turkestan.

The species has been commonly observed in and around populated places, along irrigation ditches and field margins. It was commonly cultivated in the whole country (Spota 1960) but went out of fashion and became rare nowadays (Lazkov, pers. obs.). Historical specimens do not provide any reliable data on its former distribution (Fig. 2); we assume that the cultivation was concentrated in climatically favourable, agricultural areas of western and northern Kyrgyzstan.

Figure 2.  

Recorded distribution of Alkekengi officinarum in Kyrgyzstan, according to historical specimens examined (cultivated plants excluded) and recent observations.

Archaeophyte.

The species is an archaeophyte of the Neolithic period, which was introduced from China in pre-historic times. It has been grown in China for at least six thousand years (Jin et al. 2020) for its edible fruits (Li 1973) and is still consumed in some rural territories (e.g. Kang et al. 2013, Wang et al. 2020).

Escape from confinement: Agriculture.

The species was introduced and originally used as a vegetable. When its role as a vegetable had decreased and was largely forgotten, it was still cultivated as an ornamental and traditional plant.

The species colonised the territory around the places of original cultivation by vegetative growth and seemingly by seed dispersal along water streams (cf. Cappers 1993). Whereas the species was frequently noted in walnut forests in the proximity of villages in Uzbekistan (Kovalevskaya 1961), no such wild occurrence is known in Kyrgyzstan, thus indicating that its seed dispersal was very limited or inefficient. Most likely, the main agent of its local dispersal was humans.

China.

Largely casual (persisting in places of original cultivation) or locally established. All recent observations are from the places of former cultivation (Lazkov, pers. obs.), which should be treated as casual. Not invasive.

Agriculture - no impact (the species currently does not occur as a weed, although it was formerly recorded along fields: Spota 1960). Native ecosystems - no impact (not occurring in native habitats). Urban areas - minor impact (may occur as a ruderal in populated places when the cultivation was abandoned).

Strongly decreasing. The species had been very common in agricultural areas and, at that time, was commonly observed around populated places (Spota 1960). When the tradition of the species cultivation had practically ceased, it disappeared or much decreased in many places and can be rarely seen nowadays (Fig. 3); this observation evidences that the species largely relied on cultivation for its persistence.

Figure 3.  

Alkekengi officinarum, a survivor of long-abandoned cultivation in Bishkek (photo by G. Lazkov, 21 September 2021).

North America.

Europe, Asia (southern Siberia, Central Asia, Eastern Asia), Australia and New Zealand; sporadically also elsewhere.

In Europe, this species belongs to the most widely distributed alien vascular plants (Lambdon et al. 2008). It also belongs to the most invasive plants in Russia (Morozova and Vinogradova 2018) and Belarus (Dzhus 2020).

Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan.

In Central Asia, the species was first recorded in a single locality on the south-eastern margin of Tashkent City, Uzbekistan, in 1990 (Alexeev 1991). The species was found naturalised on irrigated grassland in newly developed city districts. By the beginning of the 2010s, the species was found commonly naturalised and invasive in agricultural areas of Uzbekistan (Maltsev 2013).

In Kazakhstan, the species was first recorded in 2001 near Jänıbek in West Kazakhstan Region. This locality is situated immediately next to the Russian border, and the species was known from the southern Volga Region of Russia by that time (Sukhorukov and Berezutsky 2000). Its current distribution seems to be quite wide, especially in agricultural areas of the north (Plantarium 2021).

The species was first recorded in Kyrgyzstan by Lazkov et al. (2011), based on a single observation in Bishkek dated 2008. More data are reported in the present Contribution.

In Tajikistan, the species was first recorded from Dushanbe City and its vicinities in 2009, along roadside ditches (Nobis and Nowak 2011). It was also found on rice fields near Hissar (Nowak et al. 2013).

At present the species is naturalised in all these four countries. It is widely naturalised and invasive in Kazakhstan and Uzbekistan, but sparsely occurring and not yet invasive in Kyrgyzstan and Tajikistan.

Western Tian-Shan, Northern Tian-Shan, Alay-Turkestan (Fig. 4).

Figure 4.  

Currently recorded distribution of Bidens frondosa in Kyrgyzstan.

Bidens frondosa was recorded in Bishkek in 2008, for the first time in Kyrgyzstan. Since then, small groups of the species have been observed in the city centre (Fig. 5). These occurrences are mostly ephemerous, not lasting long, but their regular re-appearance suggests that the invasion is continuous. The species was also observed established in the Botanical Garden, where it has found a suitable agricultural habitat and experiences little pressure from the environment.

Figure 5.  

Bidens frondosa in Bishkek, Kyrgyzstan (photo by G. Lazkov, 19 June 2018).

During 2011-2020, we also observed small groups of B. frondosa in a few localities in the Fergana Depression, along the border with Uzbekistan. These previously unpublished records suggest that the species may be found elsewhere in the Depression because of its common naturalisation in Uzbekistan (Maltsev 2013).

So far, all the localities are from altitudes between 650 and 1000 m above sea level, and the species shows no tendency to spread to the mountains.

Neophyte.

The first record of the species in Kyrgyzstan, dated 2008 (Lazkov et al. 2011), most likely does not reflect its first arrival in the territory. Taking into account its first observation in Uzbekistan, dated 1990 (Alexeev 1991), with its extensive naturalisation subsequently revealed in the beginning of the 2010s (Maltsev 2013), the introduction should have started from the late Soviet period, during the 1980s, if not earlier.

Transport - Contaminant: Contaminated bait. Transport - Contaminant: Seed contaminant. Transport - Contaminant: Contaminant on animals.

The most likely pathway of introduction of Bidens frondosa in Kyrgyzstan was its arrival with contaminated forage, but we also cannot exclude its appearance on corn fields as a seed contaminant. The occurrences along irrigation ditches may be zoochorous.

Further dispersal may occur with water, humans, domestic animals and water birds.

Naturalised.

Although most of the occurrences observed so far have been represented by just a few plants, and some were proven to have disappeared, the species is apparently on the way to its naturalisation in the country. It can be considered naturalised at least in the Botanical Garden in Bishkek.

No large populations or founder localities have been noticed so far.

Agriculture - minor impact (rarely occurring along irrigation ditches, once recorded as a garden weed). Native ecosystems - no impact (restricted to agricultural and urbanised areas). Urban areas - minor impact (rarely occurs in ruderal places).

Increasing (observed).

Central America.

North and South America, Europe, Africa, Southern Asia, Australia.

Kyrgyzstan, Uzbekistan.

The species was recorded as a rare alien in Uzbekistan, observed as ruderal or escaped from cultivation (Kovalevskaya 1961).

Reported from Kyrgyzstan for the first time here.

Northern Tian-Shan (Fig. 6).

Figure 6.  

The subspontaneous occurrence of Datura innoxia in Kyrgyzstan.

So far, the species has been recently recorded from the only locality on the southern margin of Bishkek City, in 2017, by G. Lazkov. A single individual was noticed on a dumping area of the cemetery. This occurrence is apparently casual.

Neophyte.

The species was cultivated in Kyrgyzstan and Uzbekistan for at least 60 years (Nikitina 1960, Kovalevskaya 1961). It was noted to self-seed (Nikitina 1960) but has never been reported as running wild in Kyrgyzstan. Its current subspontaneous occurrence, first recorded in 2017, may be linked with an increasingly common use of the plant in ornamental cultivation, which has been observed in recent years (Lazkov, pers. obs.; Fig. 7).

Figure 7.  

Datura innoxia in ornamental cultivation in Bishkek (photo by G. Lazkov, 21 July 2020).

Escape from confinement: Ornamental purpose other than horticulture.

The species is cultivated for ornamental purposes in private gardens and public areas, and is sometimes found in waste sites. As in Europe (e.g. Gudžinskas 2017), its subspontaneous occurrences originated through garden waste. Further dispersal does not occur.

Casual.

Agriculture - no impact (not recorded in crop production areas). Native ecosystems - no impact (restricted to urbanised areas). Urban areas - minor impact (rarely escapes and occurs in ruderal places).

Increasing (inferred).

Central America.

Archaeophyte in North and South America, Central and Southern Europe, Africa, Southern and Central Asia, Malesia and Australia. Neophyte in Northern Europe. This species is one of the most widely distributed naturalised alien plants in the world, reaching top-10 in the temperate biome (Pyšek et al. 2017).

The species is a striking example of the plants native to the New World but introduced with ancient human-mediated transport in the pre-Columbian era. Plants of Datura sp. were introduced to the Old World possibly by the 4th century; the exact mechanism of such transportations is still uncertain (Geeta and Gharaibeh 2007).

The species occurs as a common alien plant in all the countries of Central Asia (Kovalevskaya 1987).

It was first recorded from Transoxiana (the territories between the Amudarya River and the Syrdarya River) by Avicenna in the first part of the 11th century, although probably as an imported plant (Geeta and Gharaibeh 2007). Nowadays, it is still frequently seen in ruderal places and on field margins (Fig. 8).

Figure 8.  

Datura stramonium along a cotton field in Andijon District, Uzbekistan (photo by T. Tillaev, 23 July 2017). Source: https://www.plantarium.ru/page/image/id/525187.html (Plantarium 2021).

Western Tian-Shan, Northern Tian-Shan, Alay-Turkestan (Fig. 9).

Figure 9.  

Recorded distribution of Datura stramonium in Kyrgyzstan.

The species was found in major agricultural areas (Nikitina 1960) and considered to occur in the whole territory of Kyrgyzstan, although we have not seen collections from the Eastern Tian-Shan. It was collected from populated places and their vicinities, at altitudes up to nearly 2000 m above sea level.

Archaeophyte.

This species was known from the whole of Central Asia from the beginning of its botanical exploration (Fedtschenko and Fedtschenko 1913). The time of its introduction is uncertain, but the species was known from the territory already in the 11th century (Geeta and Gharaibeh 2007). In agreement with the history of the plant introduction described by Geeta and Gharaibeh (2007), we may speculate that the appearance of Datura stramonium in Central Asia followed the Muslim reconquest of Transoxiana, which occurred by the beginning of the 11th century and was connected with the massive cultural influence from the Arabic world.

Escape from confinement: Horticulture.

The species was originally cultivated as a medicinal plant in India (Geeta and Gharaibeh 2007) and was seemingly introduced as such to Central Asia. Its contemporary occurrence was recorded as a ruderal plant (Deza 1983), with further dispersal by wind and local human activities. Its occasional presence on fields (grain and root vegetables) has also been recorded (Nikitina 1960); in other countries, this may also be a rather new phenomenon, linked with the recent cultivation of soybean, bean and maize that are characterised by larger planting seed material (e.g. Weaver and Warwick 1984).

Naturalised. The species is a component of traditional ruderal vegetation, but also occurs along rivers and around springs.

Agriculture - moderate impact (occasional weed of crops, in fields and gardens). Native ecosystems - minor impact (occurring along streams near populated places). Urban areas - moderate impact (ruderal occurrence).

Stable (inferred).

Central and Southern China, Korea, Japan.

North America, New Zealand (neophyte); Europe, Western and Southern Asia (archaeophyte).

In Europe, the ornamental cultivation of the species has a long history, recorded as common in Britain by Gerard (1597) and in Central Europe by L'Obel (1576) and Clusius (1601) already in the 16th century. It is currently known as naturalised in many countries, including Great Britain (Clement and Foster 1994) and Belgium (Verloove 2021).

Hemerocallis fulva was common in the North American ornamental cultivation since the late 19th century; now it became invasive in several states of the USA, occurring along roadsides and river banks (Pennsylvania Flora Database 2021).

Escaped from traditional ornamental cultivation in Kyrgyzstan and Uzbekistan.

The species was known from subspontaneous occurrences in Kyrgyzstan (Nikitina 1951, Pazij 1971). Its presence in Uzbekistan has not been mentioned in literature, but unpublished herbarium specimens (deposited at LE) were collected from the vicinities of Charvak Village, Tashkent Region (in 1899) and Qora-Qo'rg'on Village, Namangan Region (in 1912).

In historical times, a major part of the mountainous Central Asia, with its highly developed culture in populated oases, was known as Transoxiana (in Latin) or Mavarannahr (in Arabic). This territory, subordinated to various major contemporary states but being de facto autonomous, became the Khanate (then Emirate and finally Republic) of Buxoro from the 16th century until 1924. It included two major cities, Buxoro and Samarqand. The Khanate of Buxoro was characterised by extensive cultivation of numerous fruits, vegetables, ornamental and medicinal plants, which were recorded by early European travellers and native writers of Buxoro and Samarqand (e.g. Meyendorff 1826, Abu Tahir Kojo 1899).

The cultivation of Hemerocallis fulva in Samarqand was recorded by Olga Fedtschenko in 1869 (Regel 1876). This species, therefore, belongs to the ornamental cultivation of the Khanate of Buxoro, predating the Russian colonisation of the country. Its feral occurrences were recorded in the lower mountains surrounding the Fergana Depression in the early 1870s (Regel 1876), thus indicating that the species was capable of running wild, long before the beginning of the botanical records.

As evident from herbarium records (collections of A. Regel at LE, dated 1877), the species was cultivated also in Qulja [Yining], Xinjiang, China. This means that its historical cultivation apparently included also the agricultural areas of northern Kyrgyzstan.

Since the Khanate of Buxoro maintained close connections and trade of medicinal and other plants with India (Meyendorff 1826), we assume that Hemerocallis fulva was originally imported from that country; its broad distribution suggests the early period of introduction. It is also possible that the species was first imported as a vegetable, for its edible flowers and fleshy rhizomes (Li 1970), and was subsequently turned into an ornamental.

Currently, the species is very commonly cultivated in Central Asia (Fig. 10), although seemingly from some commercial sources, different from the historical cultivation. Its recent subspontaneous populations are not recorded.

Figure 10.  

Hemerocallis fulva in modern ornamental cultivation in Tashkent Region, Uzbekistan (photo by T. Tillaev, 28 June 2018). Source: https://www.plantarium.ru/page/image/id/582460.html (Plantarium 2021).

Western Tian-Shan, Alay-Turkestan (Fig. 11).

Figure 11.  

Historical records of Hemerocallis fulva in Kyrgyzstan.

The species was found along rivers and irrigation ditches in the lower mountain belt (950-1100 m a.s.l.) near populated places surrounding the Fergana Depression.

Archaeophyte.

The species was common in ornamental cultivation in the Khanate of Buxoro, and found in the territories around the Fergana Depression that belonged to this state. This introduction is at least some centuries old.

Escape from confinement: Ornamental purpose other than horticulture.

Although the plant is edible, its latest historical use was ornamental cultivation in private gardens (Regel 1876).

Locally naturalised, maintained by vegetative reproduction (colonophyte).

In Kyrgyzstan, feral populations of the species were known along rivers and irrigation ditches near populated places, from the area of semi-wild apple and walnut forests at the lower belt in the north-western part of the Fergana Range (Pazij 1971). These populations had been repeatedly sampled from the early 1870s till 1927 (Fig. 12), thus indicating their conspicuousness. Although these territories belong to the most visited and intensely studied areas in the country (e.g. Sukachev 1949), no further collections or observations originated in the latest 95 years; this indicates that the populations had significantly declined or even disappeared. Their current status or even existence have not been verified; the old feral populations may be currently extinct.

Figure 12.  

The last historical specimen of Hemerocallis fulva from Kyrgyzstan (MW0813045).

Agriculture - no impact (not weedy). Native ecosystems - minor impact (colonising riversides near populated places). Urban areas - minor impact (colonising irrigation ditches in populated places).

Declining (inferred).

Mediterranean, Western Asia (Meusel and Jäger 1965).

Archaeophyte in Temperate and Northern Europe and Temperate Asia, neophyte in Australia and North America.

The species occurs in all the countries of Central Asia (Kovalevskaya 1987).

Western Tian-Shan, Northern Tian-Shan, Eastern Tian-Shan, Alay-Turkestan (Fig. 13).

Figure 13.  

Recorded distribution of Hyoscyamus niger in Kyrgyzstan.

The species is distributed in agricultural areas and populated places across the whole territory of Kyrgyzstan (Nikitina 1960, Deza 1983).

Archaeophyte.

The species was first recorded as being in foreign use in medieval China, Tang Dynasty (Li 1977, Li 2012), corresponding to the 7th-9th centuries. This can be firmly linked with the Muslim conquest of Transoxiana, which occurred during AD 673-751 (Nicolle 2009). Besides the narcotic effect, seeds of the plant were used as a tonic that provides strength in walking for long distances (Li 1977, Li 2012); this effect apparently was valued in the contemporary army.

Escape from confinement: Horticulture.

As the plant is an important sedative, anaesthetic and pain-relieving drug of ancient times, we conclude that it was intentionally introduced with medical purposes and subsequently cultivated in Central Asia. This ancient cultivation has been abandoned long ago, and the species largely occurs as a ruderal plant in or around populated places, or as a weed, or on abandoned fields (Nikitina 1960, Kovalevskaya 1987). Many researchers (Nikitina 1960, Deza 1983, Kovalevskaya 1987) also noted its common occurrence on fields, which assumes its secondary dispersal with contaminated seeds and from ruderal habitats. Its ruderal occurrence relies on winds and human activities.

Naturalised, old invasive plant. The species has been very frequently found in populated places (ruderal places, roadsides) and on fields (wheat, alfalfa) and pastures (Nikitina 1960, Kovalevskaya 1987), but may also occur in native habitats (along riversides, on rocky slopes, in grasslands) near populated places. It is still regularly found in the country (Fig. 14).

Figure 14.  

Hyoscyamus niger near Bishkek City, Kyrgyzstan (photo by G. Chulanova, 25 May 2019). Source: https://www.plantarium.ru/page/image/id/645576.html (Plantarium 2021).

Agriculture - major impact (reported as a common weed of crops, on fields and in gardens). Native ecosystems - major impact (occurring in natural habitats near populated places). Urban areas - major impact (ruderal occurrence).

Stable (inferred).

South America.

North and Central America, Southern Europe and Asia, Africa, Australia.

In Europe, this species belongs to the most widely distributed alien vascular plants (Lambdon et al. 2008).

The species has been naturalised in several provinces of China (Zhang et al. 1994), probably as a field weed.

Sporadically found in Kazakhstan, Kyrgyzstan, Turkmenistan, Uzbekistan (Kovalevskaya 1987).

In Central Asia, the species was historically found in gardens and on melon fields (Pojarkova 1954b, Nikitina 1960, Vasilieva 1965). In the early 20th century (1907-1931), it was recorded as cultivated and ruderal in vegetable gardens of Russian-populated places.

The first record of the species from Central Asia comes from Vannovskoe Village of Turkestan Region (now Tūrar Rysqūlov Village, Türkıstan Region, Kazakhstan). This village was founded by Russian and Ukrainian colonists in 1887, which may be close to the earliest possible date of the species' introduction to Central Asia (following the conquest in 1868).

Its recent occurrence seems to be in ornamental cultivation and ruderal.

Western Tian-Shan, Northern Tian-Shan (Fig. 15).

Figure 15.  

Historical and recent records of Nicandra physalodes in Kyrgyzstan.

The species was found as a weed in agricultural areas of the Chü and Talas Depressions (Nikitina 1960). Recently, it was rediscovered as a ruderal plant (Fig. 16) in a village along the northern side of Ysyk-Köl Lake (iNaturalist 2021).

Figure 16.  

Nicandra physalodes in Cholpon-Ata, Kyrgyzstan (photographed 25 July 2020). Source: https://www.inaturalist.org/observations/54240496 (iNaturalist 2021).

Neophyte.

The plant was introduced in the early Soviet period (first record in 1928 from present-day Talas Town), transported by Russian colonists from their native villages in southern Russia. Its latest record from the Botanical Garden in Bishkek is dated 1955 and may constitute the last remnant of the old cultivation.

Its latest subspontaneous record is dated 2020 and may be linked with recent ornamental cultivation.

Escape from confinement: Agriculture. Escape from confinement: Ornamental purpose other than horticulture.

The plant seems to have been historically introduced for its cultivation and subsequent use in home bakery, and then it has become a weed in and around the places of introduction. This pathway is indicated by the contemporary evidence (Khrebtov 1926, Larionov 1931) and by the recorded occurrences of the plants in vegetable gardens.

Modern pathways of introduction of this plant in Europe include grain import (Suominen 1979, Clement and Foster 1994, Verloove 2021), wool contamination and horticulture (Clement and Foster 1994, Verloove 2021). The species is a very common weed of many crops, especially in warmer countries (Holm et al. 1997).

In Russia, in recent years, Nicandra physalodes has been widely cultivated for ornamental purposes in private gardens and city yards and along streets, and occasionally noted as running wild and occurring in ruderal and dumping places (Plantarium 2021).

A recent record from Ysyk-Köl Region, Kyrgyzstan (iNaturalist 2021) indicated that the species may escape from ornamental cultivation and occur in ruderal places as a casual alien.

Further dispersal was not observed.

Casual.

The historical occurrences have seemingly disappeared (the plant was not naturalised and the historical factors of introduction are no longer in place). The recent ruderal occurrence was represented by a single plant and was apparently casual as well.

Agriculture - no impact (no longer occurring as a weed in gardens or on fields). Native ecosystems - no impact (not found outside populated places). Urban areas - minor impact (ruderal occurrences).

Increasing (inferred).

Central and South America.

Africa, Europe, southern Asia, Australia, North America.

Tajikistan, Uzbekistan. Previously reported in error from Kyrgyzstan.

In Tajikistan, the species was frequently found during the period between 1928 and the 1960s on cotton and sesame fields in the large oasis of Boxtar (Korovin 1934, Kovalevskaya 1986), which is a large agricultural centre of the southern part of the country. Recently it was sporadically noted on rice fields (Nowak et al. 2013).

During the same period, it was recorded on cotton fields also in Uzbekistan, near Tashkent and Samarkand (Kovalevskaya 1961). Some older specimens, collected on cotton fields near Tashkent, were misidentified by Kovalevskaya (1961) as Physalis ixocarpa. These specimens (Fig. 19) are characterised by pale whitish flowers with very faint spots in the throat and by long pedicels, thus matching P. angulata. For this reason, the historical record of P. ixocarpa from Uzbekistan (Khassanov et al. 2020) should be rejected.

Figure 19.  

A specimen of Physalis angulata from Uzbekistan, which was misidentified as P. ixocarpa (TASH).

The only historical record of P. angulata from Bishkek, Kyrgyzstan (Kovalevskaya 1987) is based on herbarium specimens collected from experimental cultivation and kept at FRU and LE. On the basis of this record, the species was listed as occurring in the country (Lazkov and Sultanova 2011, Lazkov and Sultanova 2014). Since this occurrence was not spontaneous, the species should be removed from the flora of Kyrgyzstan.

It seems that P. angulata was introduced to Central Asia largely with cotton cultivation (American varieties introduced in the late 1920s). According to herbarium records, the species persisted on and around cotton fields until the 1960s. No recent records are available, and the current status of the species is unknown (presumably historical casual).

No spontaneous occurrence has been recorded.

Native to Central America.

Widely cultivated as a fruit crop. Introduced in many countries of North America, Europe, Africa, Asia, Australia. Archaeophyte in North America (González-Pérez and Guerrero-Beltrán 2021), neophyte elsewhere.

In arid areas of Asia, the species was introduced to Turkey in the 1990s as a weed of irrigated cotton fields (Bükün et al. 2002), where it became invasive and quickly gained the top status among other weeds (Bükün 2004). In other arid territories, this species is not invasive.

Reported from Kazakhstan, Kyrgyzstan and Uzbekistan for the first time here.

In Kazakhstan, the species was observed for the first (and only) time in 2019, in a damp place at Jabağyly Village, Turkestan Region (Plantarium 2021).

In Kyrgyzstan, it was found for the first time in Bishkek, in 2015.

In Uzbekistan, the species was originally recorded in 1929 and 1930 as a weed of cotton cultivation (Kovalevskaya 1961, Khassanov et al. 2020), but this record was erroneously based on misidentified specimens of P. angulata (see discussion under that species). Since 2008, the species was repeatedly collected as a ruderal plant in Tashkent (Plantarium 2021). Besides, recently it was noted (albeit with a low abundance) as a weed in Tashkent Region: on late carrot fields in Oqqoʻrgʻon District (Axmedov and Narbaev 2019), on tomato fields in Qibray District (Axmedov et al. 2020) and along margins of maize fields in Qibray District (Plantarium 2021). All recent records were misidentified as P. angulata.

Northern Tian-Shan (Fig. 23).

Figure 23.  

Distribution of Physalis philadelphica in Kyrgyzstan.

The species was first recorded by Georgy Lazkov in Bishkek City in 2015, as a few scattered individuals along Toktogul Street and a large population (ca. 20 flowering individuals) on the southern margin of the city. The scattered individuals withered quickly, whereas the fate of the large population was not studied. Further on, one individual was observed in flower in Sokuluk Village in 2021. The plants usually occurred in places with regular water supply, along irrigation ditches.

Neophyte.

The first record from Kyrgyzstan was dated 2015, thus falling within the period of the independence.

Transport - Contaminant: Seed contaminant.

In Kyrgyzstan, the species is sometimes cultivated in private gardens and sold privately in marketplaces (Lazkov, pers. obs.), but its direct escape from cultivation is considered highly unlikely. Its occurrences on roadsides and waste ground in populated places suggest the arrival with contaminated grain or fodder. No further dispersal was noticed.

In Uzbekistan, the species with certainly arrived with contaminated seed material, as indicated by its occurrence on fields. A wide variety of contaminated seed material (carrots, tomato, maize) indicates its North American origin and multiple sources of introduction.

In Russia, the species was introduced in the European part with garden seeds, as a weed of flower beds and vegetable gardens, recorded in the Middle Volga Region in the 1990s (Rakov et al. 2011). Its earliest record in Siberia (Novosibirsk, dated 1944) comes from potato fields (Ebel et al. 2015).

In the USA, the species is cultivated for fruits and frequently escapes from cultivation, becoming established along roadsides and field margins (Sullivan 2004). In Kenya, its original introduction was intentional as a green manure crop that caused a local invasion (Cunningham-van Someren 1957). In the British Isles, the species originated with grain, wool and food refuse (Clement and Foster 1994); it was also known as a contaminant of bird seed (Hanson and Mason 1985).

Further dispersal in other countries was registered as occurring with animals (Cunningham-van Someren 1957) and water (Bükün et al. 2002). Its riverbed occurrence in Spain (Gómez-Bellver et al. 2016) seems to be connected with this type of dispersal.

Presumably North America.

Casual; ephemeral or locally persisting. The species may become established in places with regular water supply; so far, no long-term survival has been observed.

Agriculture - no impact (so far, not recorded on fields, although recent surveys are lacking). Native ecosystems - no impact (not found outside populated places). Urban areas - minor impact (casual occurrence as a ruderal plant).

Increasing (observed).

The species has been noticed in Kyrgyzstan only recently, as a newcomer. Its regular recent occurrence as a weed or ruderal plant in Uzbekistan may suggest further spreading also in Kyrgyzstan. Its recent introduction and subsequent expansion in Turkey (Economou et al. 2016) indicates that the species is potentially invasive in regularly irrigated areas.

Southern Europe, Mediterranean, Southern Asia from Asia Minor to China. The species is most genetically diverse in Asia and may have an Asian origin (Särkinen et al. 2018). In Central Asia, the species is considered native in Turkmenistan, where it was recorded more commonly from riversides in the mountains (Pojarkova 1954b).

Archaeophyte in Boreal and Central Europe, Northern and Central Asia; neophyte in North America, South Africa, Malesia and Australia. One of the most common and widely distributed weeds in the world (Holm et al. 1979).

Native in Turkmenistan, alien in Kazakhstan, Kyrgyzstan, Tajikistan and Uzbekistan.

Historically, Solanum nigrum was a common weed of irrigated fields (wheat, cotton, maize, maash) in Central Asia (Fedtschenko 1915).

Most of the major sources (e.g. Kovalevskaya 1987, Särkinen et al. 2018) do not make a distinction between the native and secondary distributions of S. nigrum because of the old age of its naturalisation in many areas. In this case, we considered type of habitat as the main distinguishing feature (Webb 1985) and assigned the alien status to the areas with overwhelmingly predominant ruderal occurrences.

Western Tian-Shan, Northern Tian-Shan, Alay-Turkestan (Fig. 25).

Figure 25.  

Recorded distribution of Solanum nigrum in Kyrgyzstan.

The species was considered occurring in all parts of the country (Spota 1960, Deza 1983), although we have seen no collections from the Eastern Tian-Shan. It was collected at altitudes between 600 and 1350 m above sea level.

Archaeophyte.

This species was known from the whole of Central Asia from the beginning of its botanical exploration (Fedtschenko and Fedtschenko 1913). The time of its introduction is uncertain but is most likely Neolithic, as the species was recorded among the earliest and most common weeds in the early Neolithic of Germany (Rösch 1998).

Transport - Contaminant: Seed contaminant.

The species is a noxious weed of gardens and fields (Deza 1983), also occurring in ruderal places (Spota 1960). It is capable of infesting a large variety of crops, including wheat and melons (e.g. Cherniakovskaya 1935, Ogg et al. 1981). We assume that Solanum nigrum arrived to the territory as a weed of historical crops (possibly of S. villosum).

Further dispersal occurs with water, contaminated seed and soil.

Naturalised, invasive. The species has been a noxious weed of all crops in Kyrgyzstan (Deza 1983) and remains common to date.

Agriculture - major impact (noxious weed of all crops, in fields and gardens). Native ecosystems - minor impact (occurring along streams and water bodies near populated places). Urban areas - major impact (ruderal occurrence).

Stable (observed).

Southern Europe, Mediterranean, Southern Asia from Asia Minor to China, Africa. In Central Asia, the species is considered native in Turkmenistan, where it was recorded abundantly from riversides in the mountains (as Solanum nigrum s.l.: Pojarkova 1954b).

Archaeophyte in Boreal and Central Europe, Northern and Central Asia and South Africa; neophyte in North America and Australia.

Native in Turkmenistan, alien in Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan.

Similarly to Solanum nigrum, we consider S. villosum as alien in Central Asia north of Turkmenistan because of its occurring exclusively on cultivated lands or in ruderal habitats (Spota 1960, Kovalevskaya 1987). Pojarkova (1955a) considered riversides as a native habitat of the species in the southern part of Central Asia.

Western Tian-Shan, Northern Tian-Shan, Alay-Turkestan (Fig. 27).

Figure 27.  

Distribution of Solanum villosum in Kyrgyzstan.

The species occurs in major agricultural territories: the Fergana Depression, the Chü Depression and the Ysyk-Köl Depression (Spota 1960, Lazkov and Sultanova 2014). It was collected at altitudes between 600 and 1800 m above sea level.

Archaeophyte.

This species was known from the whole of Central Asia from the beginning of its botanical exploration (Pojarkova 1955b). The time of its introduction is uncertain.

Escape from confinement: Agriculture.

The species is especially abundant in Southern Europe, Central and Southern Asia and Africa, where it has been traditionally used as a fruit or leaf vegetable (Cherniakovskaya 1935, Pojarkova 1955a, Baranov 1967, Defelice 2003, Njau Mwai et al. 2007, Särkinen et al. 2018). It was known in Chinese cultivation in the Russian Far East during the 19th century (Schischkin 1936). For this reason, we assume that the species was introduced intentionally as a cultivated plant, rather than unintentionally as a weed. Its current weedy status seems to be secondary.

Further dispersal occurs with water, contaminated seed and soil, in the same way as Solanum nigrum.

Naturalised. The species was noted as a weed in gardens and on fields, locally common but seemingly not truly noxious (Spota 1960).

Agriculture - major impact (locally common weed of crops, in fields and gardens). Native ecosystems - minor impact (occurring along streams and water bodies near populated places). Urban areas - moderate impact (ruderal occurrence).

Stable (inferred).