Crepidinae
Systematics
The Crepidinae as a subdivision of the Cichorieae go back to Cassini. In his most elaborated third version of the tribe (Cassini 1830), his "Crepidées" formed one of four "sections" of the tribe. The "Crepidées" included the groups with elongated achenes more or less tapering towards the top and with a white pappus (rarely absent) of non-plumose setae, thus a very large portion of the entire tribe. Hoffmann (1890–1894) and Stebbins (1953) provided each to some extend different circumscriptions of a still fairly widely defined Crepidinae. A major change came only with the morphology-based phylogenetic approach by Bremer (1994), who separated the subtribes Lactucinae and Hyoseridinae (as Sonchinae) from the core Crepidinae, a treatment largely corroborated by all later molecular phylogenies. Kilian & al. (2009) and subsequent contributions (see below) still refined the circumscription of the Crepidinae, in particular regarding its delimitation from the Lactucinae.
Kilian & al. (2009) also separated the Chondrilla alliance as a separate subtribe Chondrillinae. Subsequent ITS phylogenies confirmed a well-supported clade comprising the genera Chondrilla, Phitosia, Willemetia (Tremetsberger & al. 2012; Wang & al. 2020; Yin & al. 2021) and Caucasoseris (Güzel & al. 2022). Whereas the Chondrillinae were mostly resolved as sister group to the Crepidinae in these ITS phylogenies, the plastid DNA phylogeny by Güzel & al. (2022) found the Chondrillinae nested within the Crepidinae. The first well-resolved phylogeny for the Crepidinae by Xu & al. (2024), achieved with hybrid capture phylogenomics, finally clearly revealed the Chondrillinae as deeply nested in the Crepidinae in both the nuclear and plastid DNA phylogenies. Separation of the Chondrillinae is therefore to be rejected.
The Crepidinae (including the reunited Chondrillinae) comprises 29 genera and c. 400 species (the c. 2500 Taraxacum (micro)species [Kirschner & al. 2020] not counted) or more than 1/4 of the species of the tribe, making it the largest subtribe of the Cichorieae.
The comprehensive molecular phylogeny of the subtribe by Xu & al. (2024) in combination with a number of previous Sanger sequencing studies focussing on single genera or generic alliances allows to identify a number of well-supported lineages within the subtribe and their relationships. The study by Xu & al. (2024) also corroborates the previous indications for extensive cytonuclear discordance between Lactucinae and Crepidinae as well as within the Crepidinae (Liu & al. 2013; Wang & al. 2013; Liu & Ren 2014; Kilian & al. 2017; Yin & al. 2021). The classification of the subtribe relies on the nuclear phylogenies and we adopt the designations of the terminal clades by Xu & al. (2024).
The Ixeris clade comprises the genera Ixeris, Ixeridium and Askellia (Xu & al. 2024). The sister group relationship of Ixeris and Ixeridium, in their circumscription by Pak & Kawano (1992) which was confirmed by Shih & Kilian (in Shih & al. 2011) and Nakamura & al. (2014), with Askellia in turn as sister to the former two, was already resolved in previous studies (Zhang & al. 2011; Liu & al. 2013; Wang 2020; Yin & al. 2021). In the analyses by Enke & Gemeinholzer (2008) and Peng & al. (2014), however, Askellia instead appeared as sister to Crepis s.l. Askellia, the former Crepis sect. Ixeridopsis, which has been recognised as a separate genus also by Sennikov & Illarionova (2008) on morphological ground, was first shown in the molecular phylogenetic analysis by Enke & Gemeinholzer (2008) to constitute a genus separate from Crepis. Most of the Askellia species (e.g., A. nana, A. flexuosa), however, used to be treated under Youngia, to which they were reassigned by Adylov & Zuckerwanik (1993). However, Askellia has terete achenes and a basic chromosome number of x = 7 (not present in Crepis), while the achenes of Youngia are compressed and angular and the chromosome number is x = 8. The clade is centred in E Asia with Askellia also extending into North America.
The Youngia clade, comprising the genera Youngia, Crepidiastrum and Lapsanastrum, has been resolved by a number of studies (Deng & al. 2014; Kilian & al. 2009; Zhang & al. 2011; Liu & al. 2013; Peng & al. 2014; Wang & al. 2020; Yin & al. 2021) and has been confirmed by Xu & al. (2024), who added the newly described monotypic genus Qineryangia as sister to the clade of the other genera. Crepidiastrum is circumscribed in the wider sense first established by Shih & Kilian (in Shih & al. 2011) and largely confirmed by Peng & al. (2014), thus embracing the former genera Paraixeris and Crepidifolium (Sennikov & Illarionova 2008). Lapsanastrum (Pak & Bremer 1995), as revealed by Deng & al. (2014), is deeply nested within Youngia, consituting some sort of a parallel case to Lapsana and Rhagadiolus being nested in Crepis. The clade is centred in E Asia.
The Chondrilla clade includes Chondrilla and Willemetia (Xu & al. 2024) and as consecutive sister to them Phitosia (Kilian & al. 2009; Tremetsberger & al. 2012; Wang & al. 2020; Yin & al. 2021) and Caucasoseris (Güzel & al. 2022). The clade is centered and probably also originated in the E Mediterranean-SW Asian region.
The Mojiangia clade (Yin & al. 2021; Xu & al. 2024) includes the single relic species from Yunnan, M. oreophila, which is concluded to be a successor of the maternal Crepidinae ancestor of the allotetraploid Chinese endemic Faberia, which appears as its sister in the plastid DNA tree (Yin & al. 2021 and corroborated by Xu & al. 2024). The genus Faberia has evolved from acient hybridisation between a Lactucinae and a Crepidinae member and is placed in the Lactucinae.
The Taraxacum clade includes the genus Taraxacum only and is sister to the Crepis clade (Xu & al. 2024). This is a new finding because previously Taraxacum has been resolved in ITS phylogenies instead as sister to the Ixeris clade (Kilian & al. 2009; Zhang & al. 2011; Yin & al. 2022).
The Crepis clade includes the genera Crepis and Lapsana in the phylogeny by Xu & al. (2024). The ITS phylogenies by Enke & Gemeinholzer (2008), Wang & al. (2020) and Yin & al. (2021) revealed that in addition Lapsana and Rhagadiolus, two genera lacking a pappus and formerly placed in the Hypochaeridinae (Bremer 1994; Lack 2006), also belong to this clade, confirming indications for their affinities to the Crepidinae by Whitton & al (1995). Enke & Gemeinholzer (2008) had shown that in the both nuclear and plastid DNA analyses the species of Babcock’s Crepis sections Intybellia, Lagoseris, Microcephalum, Phaecasium and Pterotheca form a clade sister to Lapsana and Rhagadiolus, which roughly corresponds to, but is still larger than, the Crepis segregate Lagoseris of some authors. The second and major part of Crepis is sister to that clade. Morphologically, however, both parts of Crepis are ill-delimited from each other, which speaks strongly against separating Lagoseris from Crepis (Enke & Gemeinholzer 2008). Lapsana and Rhagadiolus, in contrast, are morphologically well delimited from Crepis s.l., and it therefore appears, at least for the time being, the best solution to maintain Lapsana and Rhagadiolus and thus accept Crepis as a paraphyletic genus; compare the case of Lapsanastrum and Youngia, above. The monospecific afroalpine Dianthoseris was found by Enke & al. (2008) to be nested right into, and to form a congener of, Crepis s.str.
The Garhadiolus-Heteracia clade (Kilian & al. 2009; Zhang & al. 2011; Kilian & al. 2017; Wang & al. 2020; Yin & al. 2021), including four small or monotypic genera, Garhadiolus, Heteracia, Heteroderis and Lagoseriopsis, was not present in the sampling and phylogeny by Xu & al. (2024). The inclusion of Garhadiolus in the Crepidinae, which was formerly placed in the Hypochaeridinae by Bremer (1994) and Lack (2006), was first indicated by Whitton & al. (1995). Confirmation of this clade and establishing of its relationships, probably somehow related to or part of the Dubyaea-Soroseris clade, is still needed. The clade is centered in SW and Middle Asia.
The large, predominantly Asian Dubyaea-Soroseris clade (Kilian & al. 2009, Zhang & al. 2011; Liu & al. 2013; Wang & al. 2020; Yin & al. 2021; Xu & al. 2024) still needs further studies based on a extended sampling. The circumscription of Dubyaea, the name typified by D. hispida, is not fully settled yet. Based on molecular phylogenetics, Liu & al. (2013) transferred D. glaucescens to Faberia in the Lactucinae and Chen & al. (2021) excluded two species as members of a new genus Lihengia, also member of the Lactucinae. The phylogenetic reconstruction by Xu & Chen (2021) with hybrid capture phylogenomics of the nuclear genome resolved a monophylum of seven species; the relationship of the remaining five species still needs confirmation. Soroseris is a high montane and alpine genus distributed in the Himalayas (Kashmir, Himachal Pradesh; Bhutan, N India, Nepal), SW China and N Myanmar (Stebbins 1940; Zhang & al. 2011); Stebbinsia with its only species S. umbrella is nested right within Soroseris (Zhang & al. 2011), as was already assumed by Lack (2006) from morphology. The clade also includes Nabalus (extending into North America) and Syncalathium, genera formerly placed into the Lactucinae (Bremer 1994; Lack 2006). Syncalathium turned out to be diphylectic; the genus in the sense of the type of the name provided by S. sukaczevii (correctly to be named S. kawaguchii) is a member of the Crepidinae, whereas S. souliei is misplaced in the Crepidinae and actually nested in the Lactucinae (Kilian & al. 2009: 348–350 and Zhang & al. 2011, corroborating the observation by Stebbins (1940: 47–50) inferred from achene morphology). The recognition of Nabalus as a genus separate from Prenanthes (Kilian & al. 2009; Zhang & al. 2011; Schilling & al. 2015; Kilian & al. 2017) and including all North American and several Central and East Asian members of the latter genus (Sennikov 2000), confirms the findings by Stebbins (1940: 63). He had concluded from studies of the achene vascularisation that the species of Nabalus are much closer to Dubyaea and Soroseris than to Prenanthes purpurea, which provides the type of the name Prenanthes. Nabalus is, however, not monophyletic: the Asian Nabalus species are nested in a different clade than the North American N. altissimus and N. trifoliolatus, the latter providing the type of the generic name Nabalus, and the northwestern North American N. sagittatus is sister to those two clades (Zhang & al. 2011; Kilian & al. 2017). Further members of the clade are Hololeion, formerly placed in the Hieraciinae (Bremer 1994; Lack 2006), Sonchella, accommodating two Central to E Asian species (Sennikov & Illarionova 2008; Kilian & al. 2017), and Sinoseris, ecologically outstanding for its distribution in the subtropical (warm-temperate) highlands of Sichuan and Yunnan (Wang & al. 2020).
The relationships between the above clades of the Crepidinae has been resolved for the first time by Xu & al. (2024). According to the nuclear DNA phylogeny of these authors, the clades group into either of two major clades. The Ixeris and the Youngia clades form one major clade. The second major clade comprises the Chondrilla clade and the Mojiangia clade as consecutive sisters to a clade including the Taraxacum and Crepis clades as sistergroup and the Dubyaea-Soroseris clade as sister to the former.
Cytonuclear discordance was detected by Xu & al. (2024) with respect to the CrepisChondrilla clades, which are resolved as sistergroup and sister to the remainder of the subtribe in the plastid DNA phylogeny. Taraxacum and Mojiangia (in a sister group with the Lactucinae genus Faberia, see above) are consecutive sisters to the Dubyaea-Soroseris clade in the plastid DNA phylogeny.A,AA,AB,AC,AD,AE,AF,AG,AH,AI,AJ,AK,B,C,D,E,F,G,H,I,J,K,L,M,N,O,P,Q,R,S,T,U,V,W,X,Y,Z
Kilian & al. (2009) also separated the Chondrilla alliance as a separate subtribe Chondrillinae. Subsequent ITS phylogenies confirmed a well-supported clade comprising the genera Chondrilla, Phitosia, Willemetia (Tremetsberger & al. 2012; Wang & al. 2020; Yin & al. 2021) and Caucasoseris (Güzel & al. 2022). Whereas the Chondrillinae were mostly resolved as sister group to the Crepidinae in these ITS phylogenies, the plastid DNA phylogeny by Güzel & al. (2022) found the Chondrillinae nested within the Crepidinae. The first well-resolved phylogeny for the Crepidinae by Xu & al. (2024), achieved with hybrid capture phylogenomics, finally clearly revealed the Chondrillinae as deeply nested in the Crepidinae in both the nuclear and plastid DNA phylogenies. Separation of the Chondrillinae is therefore to be rejected.
The Crepidinae (including the reunited Chondrillinae) comprises 29 genera and c. 400 species (the c. 2500 Taraxacum (micro)species [Kirschner & al. 2020] not counted) or more than 1/4 of the species of the tribe, making it the largest subtribe of the Cichorieae.
The comprehensive molecular phylogeny of the subtribe by Xu & al. (2024) in combination with a number of previous Sanger sequencing studies focussing on single genera or generic alliances allows to identify a number of well-supported lineages within the subtribe and their relationships. The study by Xu & al. (2024) also corroborates the previous indications for extensive cytonuclear discordance between Lactucinae and Crepidinae as well as within the Crepidinae (Liu & al. 2013; Wang & al. 2013; Liu & Ren 2014; Kilian & al. 2017; Yin & al. 2021). The classification of the subtribe relies on the nuclear phylogenies and we adopt the designations of the terminal clades by Xu & al. (2024).
The Ixeris clade comprises the genera Ixeris, Ixeridium and Askellia (Xu & al. 2024). The sister group relationship of Ixeris and Ixeridium, in their circumscription by Pak & Kawano (1992) which was confirmed by Shih & Kilian (in Shih & al. 2011) and Nakamura & al. (2014), with Askellia in turn as sister to the former two, was already resolved in previous studies (Zhang & al. 2011; Liu & al. 2013; Wang 2020; Yin & al. 2021). In the analyses by Enke & Gemeinholzer (2008) and Peng & al. (2014), however, Askellia instead appeared as sister to Crepis s.l. Askellia, the former Crepis sect. Ixeridopsis, which has been recognised as a separate genus also by Sennikov & Illarionova (2008) on morphological ground, was first shown in the molecular phylogenetic analysis by Enke & Gemeinholzer (2008) to constitute a genus separate from Crepis. Most of the Askellia species (e.g., A. nana, A. flexuosa), however, used to be treated under Youngia, to which they were reassigned by Adylov & Zuckerwanik (1993). However, Askellia has terete achenes and a basic chromosome number of x = 7 (not present in Crepis), while the achenes of Youngia are compressed and angular and the chromosome number is x = 8. The clade is centred in E Asia with Askellia also extending into North America.
The Youngia clade, comprising the genera Youngia, Crepidiastrum and Lapsanastrum, has been resolved by a number of studies (Deng & al. 2014; Kilian & al. 2009; Zhang & al. 2011; Liu & al. 2013; Peng & al. 2014; Wang & al. 2020; Yin & al. 2021) and has been confirmed by Xu & al. (2024), who added the newly described monotypic genus Qineryangia as sister to the clade of the other genera. Crepidiastrum is circumscribed in the wider sense first established by Shih & Kilian (in Shih & al. 2011) and largely confirmed by Peng & al. (2014), thus embracing the former genera Paraixeris and Crepidifolium (Sennikov & Illarionova 2008). Lapsanastrum (Pak & Bremer 1995), as revealed by Deng & al. (2014), is deeply nested within Youngia, consituting some sort of a parallel case to Lapsana and Rhagadiolus being nested in Crepis. The clade is centred in E Asia.
The Chondrilla clade includes Chondrilla and Willemetia (Xu & al. 2024) and as consecutive sister to them Phitosia (Kilian & al. 2009; Tremetsberger & al. 2012; Wang & al. 2020; Yin & al. 2021) and Caucasoseris (Güzel & al. 2022). The clade is centered and probably also originated in the E Mediterranean-SW Asian region.
The Mojiangia clade (Yin & al. 2021; Xu & al. 2024) includes the single relic species from Yunnan, M. oreophila, which is concluded to be a successor of the maternal Crepidinae ancestor of the allotetraploid Chinese endemic Faberia, which appears as its sister in the plastid DNA tree (Yin & al. 2021 and corroborated by Xu & al. 2024). The genus Faberia has evolved from acient hybridisation between a Lactucinae and a Crepidinae member and is placed in the Lactucinae.
The Taraxacum clade includes the genus Taraxacum only and is sister to the Crepis clade (Xu & al. 2024). This is a new finding because previously Taraxacum has been resolved in ITS phylogenies instead as sister to the Ixeris clade (Kilian & al. 2009; Zhang & al. 2011; Yin & al. 2022).
The Crepis clade includes the genera Crepis and Lapsana in the phylogeny by Xu & al. (2024). The ITS phylogenies by Enke & Gemeinholzer (2008), Wang & al. (2020) and Yin & al. (2021) revealed that in addition Lapsana and Rhagadiolus, two genera lacking a pappus and formerly placed in the Hypochaeridinae (Bremer 1994; Lack 2006), also belong to this clade, confirming indications for their affinities to the Crepidinae by Whitton & al (1995). Enke & Gemeinholzer (2008) had shown that in the both nuclear and plastid DNA analyses the species of Babcock’s Crepis sections Intybellia, Lagoseris, Microcephalum, Phaecasium and Pterotheca form a clade sister to Lapsana and Rhagadiolus, which roughly corresponds to, but is still larger than, the Crepis segregate Lagoseris of some authors. The second and major part of Crepis is sister to that clade. Morphologically, however, both parts of Crepis are ill-delimited from each other, which speaks strongly against separating Lagoseris from Crepis (Enke & Gemeinholzer 2008). Lapsana and Rhagadiolus, in contrast, are morphologically well delimited from Crepis s.l., and it therefore appears, at least for the time being, the best solution to maintain Lapsana and Rhagadiolus and thus accept Crepis as a paraphyletic genus; compare the case of Lapsanastrum and Youngia, above. The monospecific afroalpine Dianthoseris was found by Enke & al. (2008) to be nested right into, and to form a congener of, Crepis s.str.
The Garhadiolus-Heteracia clade (Kilian & al. 2009; Zhang & al. 2011; Kilian & al. 2017; Wang & al. 2020; Yin & al. 2021), including four small or monotypic genera, Garhadiolus, Heteracia, Heteroderis and Lagoseriopsis, was not present in the sampling and phylogeny by Xu & al. (2024). The inclusion of Garhadiolus in the Crepidinae, which was formerly placed in the Hypochaeridinae by Bremer (1994) and Lack (2006), was first indicated by Whitton & al. (1995). Confirmation of this clade and establishing of its relationships, probably somehow related to or part of the Dubyaea-Soroseris clade, is still needed. The clade is centered in SW and Middle Asia.
The large, predominantly Asian Dubyaea-Soroseris clade (Kilian & al. 2009, Zhang & al. 2011; Liu & al. 2013; Wang & al. 2020; Yin & al. 2021; Xu & al. 2024) still needs further studies based on a extended sampling. The circumscription of Dubyaea, the name typified by D. hispida, is not fully settled yet. Based on molecular phylogenetics, Liu & al. (2013) transferred D. glaucescens to Faberia in the Lactucinae and Chen & al. (2021) excluded two species as members of a new genus Lihengia, also member of the Lactucinae. The phylogenetic reconstruction by Xu & Chen (2021) with hybrid capture phylogenomics of the nuclear genome resolved a monophylum of seven species; the relationship of the remaining five species still needs confirmation. Soroseris is a high montane and alpine genus distributed in the Himalayas (Kashmir, Himachal Pradesh; Bhutan, N India, Nepal), SW China and N Myanmar (Stebbins 1940; Zhang & al. 2011); Stebbinsia with its only species S. umbrella is nested right within Soroseris (Zhang & al. 2011), as was already assumed by Lack (2006) from morphology. The clade also includes Nabalus (extending into North America) and Syncalathium, genera formerly placed into the Lactucinae (Bremer 1994; Lack 2006). Syncalathium turned out to be diphylectic; the genus in the sense of the type of the name provided by S. sukaczevii (correctly to be named S. kawaguchii) is a member of the Crepidinae, whereas S. souliei is misplaced in the Crepidinae and actually nested in the Lactucinae (Kilian & al. 2009: 348–350 and Zhang & al. 2011, corroborating the observation by Stebbins (1940: 47–50) inferred from achene morphology). The recognition of Nabalus as a genus separate from Prenanthes (Kilian & al. 2009; Zhang & al. 2011; Schilling & al. 2015; Kilian & al. 2017) and including all North American and several Central and East Asian members of the latter genus (Sennikov 2000), confirms the findings by Stebbins (1940: 63). He had concluded from studies of the achene vascularisation that the species of Nabalus are much closer to Dubyaea and Soroseris than to Prenanthes purpurea, which provides the type of the name Prenanthes. Nabalus is, however, not monophyletic: the Asian Nabalus species are nested in a different clade than the North American N. altissimus and N. trifoliolatus, the latter providing the type of the generic name Nabalus, and the northwestern North American N. sagittatus is sister to those two clades (Zhang & al. 2011; Kilian & al. 2017). Further members of the clade are Hololeion, formerly placed in the Hieraciinae (Bremer 1994; Lack 2006), Sonchella, accommodating two Central to E Asian species (Sennikov & Illarionova 2008; Kilian & al. 2017), and Sinoseris, ecologically outstanding for its distribution in the subtropical (warm-temperate) highlands of Sichuan and Yunnan (Wang & al. 2020).
The relationships between the above clades of the Crepidinae has been resolved for the first time by Xu & al. (2024). According to the nuclear DNA phylogeny of these authors, the clades group into either of two major clades. The Ixeris and the Youngia clades form one major clade. The second major clade comprises the Chondrilla clade and the Mojiangia clade as consecutive sisters to a clade including the Taraxacum and Crepis clades as sistergroup and the Dubyaea-Soroseris clade as sister to the former.
Cytonuclear discordance was detected by Xu & al. (2024) with respect to the CrepisChondrilla clades, which are resolved as sistergroup and sister to the remainder of the subtribe in the plastid DNA phylogeny. Taraxacum and Mojiangia (in a sister group with the Lactucinae genus Faberia, see above) are consecutive sisters to the Dubyaea-Soroseris clade in the plastid DNA phylogeny.A,AA,AB,AC,AD,AE,AF,AG,AH,AI,AJ,AK,B,C,D,E,F,G,H,I,J,K,L,M,N,O,P,Q,R,S,T,U,V,W,X,Y,Z
Distribution
Africa: Algeria native ‒ native; Angola native; Azores native; Burundi native; Cameroon native; Canary Is. native ‒ native; Cape Provinces (Eastern Cape Province native, Northern Cape Province introduced, Western Cape Province introduced); Comoros (Comoros introduced); Djibouti introduced; Egypt native; Eritrea native; Ethiopia native; Free State native; Kenya native; KwaZulu-Natal native; Lesotho native; Libya native ‒ native; Madagascar introduced; Madeira native; Malawi native; Mauritius introduced; Morocco native ‒ native (Morocco native); Mozambique native; Namibia introduced; Nigeria native; Northern Provinces (Gauteng native, Mpumalanga native, North-West Province native, Northern Province native); Rodrigues introduced; Rwanda native; Réunion introduced; Selvagens native; Seychelles introduced; Somalia native; Sudan native; Swaziland native; Tanzania native; Tunisia native ‒ native; Uganda native; Zaire native; Zambia native; Zimbabwe native. Antarctic: Falkland Is. native. Asia-Temperate: Afghanistan native ‒ native ‒ native ‒ native; Altay native ‒ native ‒ native; Amur native ‒ native; Buryatiya native; China North-Central (Beijing native, Gansu native, Hebei native, Shaanxi native, Shandong native, Shanxi native); China South-Central (Chongqing native, Guizhou native, Hubei native, Sichuan native, Yunnan native); China Southeast (Anhui native, Fujian native, Guangdong native, Guangxi native, Henan native, Hunan native, Jiangsu native, Jiangxi native, Zhejiang native); Chita native; Cyprus native ‒ native ‒ native; East Aegean Is. native ‒ native; Gulf States (United Arab Emirates native); Hainan native; Inner Mongolia (Nei Mongol native, Ningxia native); Iran native ‒ native ‒ native; Iraq native ‒ native ‒ native; Irkutsk native; Japan native (Hokkaido native, Honshu native, Kyushu native, Shikoku native); Kamchatka native ‒ native; Kazakhstan native ‒ native ‒ native; Khabarovsk native ‒ native; Kirgizistan native ‒ native ‒ native; Korea native (North Korea native, South Korea native); Krasnoyarsk native; Kuril Is. native ‒ native; Lebanon-Syria native (Lebanon native ‒ native, Syria native ‒ native); Magadan native ‒ native; Manchuria (Heilongjiang native, Jilin native, Liaoning native); Mongolia native ‒ native ‒ native; Nansei-shoto native; North Caucasus native ‒ native (Chechnya native, Dagestan native ‒ native, Kabardino-Balkariya native, Karacheyevo-Cherkessiya native, Krasnodar native ‒ native, Severo-Osetiya native, Stavropol native ‒ native); Ogasawara-shoto native; Oman native; Palestine (Israel native ‒ native, Jordan native ‒ native); Primorye native ‒ native; Qinghai native; Sakhalin native ‒ native; Saudi Arabia native ‒ native; Sinai native; Tadzhikistan native ‒ native ‒ native ‒ native; Taiwan native; Tibet native ‒ native; Transcaucasus (Abkhaziya native ‒ native, Adzhariya native ‒ native, Armenia native ‒ native, Azerbaijan native ‒ native, Georgia native ‒ native, Nakhichevan native ‒ native); Turkey native ‒ native ‒ native; Turkmenistan native ‒ native ‒ native; Tuva native; Uzbekistan native ‒ native ‒ native ‒ native; West Siberia native ‒ native; Xinjiang native ‒ native ‒ native ‒ native; Yakutskiya native; Yemen introduced (North Yemen native, South Yemen native). Asia-Tropical: Assam (Assam native, Manipur native, Meghalaya native, Nagaland native); Bangladesh native; Borneo (Sabah native); Cambodia native; East Himalaya native (Arunachal Pradesh native, Bhutan native, Darjiling native, Sikkim native); India (Bihar native, Delhi native, Haryana native, Kerala native, Maharashtra native, Punjab native, Tamil Nadu native, Uttar Pradesh native, West Bengal native); Jawa native; Laos native; Malaya (Peninsular Malaysia native); Maluku native; Myanmar native ‒ native; Nepal native; New Guinea (Irian Jaya native, Papua New Guinea native); Pakistan native ‒ native ‒ native ‒ native; Philippines native; Sri Lanka native; Sulawesi native; Thailand native; Vietnam native; West Himalaya native (Himachal Pradesh native, Jammu-Kashmir native ‒ native, Uttaranchal native). Australasia: Antipodean Is. native; Chatham Is. native; Kermadec Is. introduced; New South Wales (Australian Capital Territory native, New South Wales native); New Zealand North native; New Zealand South native; Northern Territory introduced ‒ introduced; Queensland (Queensland native); South Australia native; Tasmania native; Victoria native; Western Australia (Western Australia native). Europe: Albania native ‒ native; Austria native ‒ native (Austria native ‒ native, Liechtenstein native ‒ native); Baleares native ‒ native; Baltic States (Estonia native, Kaliningrad native ‒ native, Latvia native, Lithuania native); Belarus native ‒ native; Belgium native (Belgium native ‒ native, Luxembourg native ‒ native); Bulgaria native ‒ native; Central European Russia native ‒ native; Corse native ‒ native; Czechoslovakia native (Czech Republic native ‒ native, Slovakia native ‒ native); Denmark native; East European Russia native ‒ native; Finland native; France native ‒ native (Channel Is. native, France native); Føroyar native; Germany native ‒ native; Great Britain native ‒ native; Greece native ‒ native; Hungary native ‒ native; Iceland native; Ireland native (Ireland native, Northern Ireland native); Italy native ‒ native (Italy native); Kriti native ‒ native; Krym native ‒ native; Netherlands native; North European Russia native; Northwest European Russia native; Norway native; Poland native ‒ native; Portugal native ‒ native; Romania native ‒ native; Sardegna native ‒ native; Sicily native ‒ native (Malta native, Sicily native ‒ native); South European Russia native ‒ native; Spain native ‒ native (Andorra native ‒ native, Gibraltar native ‒ native, Spain native ‒ native); Svalbard native; Sweden native; Switzerland native ‒ native; Turkey-in-Europe native ‒ native; Ukraine (Moldova native ‒ native, Ukraine native ‒ native); Yugoslavia (Bosnia-Herzegovina native ‒ native, Croatia native ‒ native, Macedonia native ‒ native, Montenegro native ‒ native, Serbia native ‒ native, Slovenia native ‒ native). Northern America: Alabama native; Alaska native; Alberta native; Aleutian Is. native ‒ native; Arizona native; Arkansas native; British Columbia native; California native; Colorado native; Connecticut native; Delaware native; District of Columbia native; Florida native; Georgia, U.S.A. native; Greenland native; Idaho native; Illinois native; Indiana native; Iowa native; Kansas native; Kentucky native; Labrador native; Louisiana native; Maine native; Manitoba native; Maryland native; Massachusetts native; Mexico Central (México Distrito Federal native, México State native, Puebla native); Mexico Gulf (Veracruz native); Mexico Northeast (Chihuahua native, Coahuila native, Durango native, Hidalgo native, San Luis Potosí native, Tamaulipas native); Mexico Northwest (Sonora native); Mexico Southeast (Chiapas native); Mexico Southwest (Michoacán introduced, Oaxaca native); Michigan native; Minnesota native; Mississippi native; Missouri native; Montana native; Nebraska native; Nevada native; New Brunswick native; New Hampshire native; New Jersey native; New Mexico native; New York native; Newfoundland (Newfoundland native, St.Pierre-Miquelon native); North Carolina native; North Dakota native; Northwest Territories native; Nova Scotia native; Nunavut native; Ohio native; Oklahoma native; Ontario native; Oregon native; Pennsylvania native; Prince Edward I. native; Québec native; Rhode I. native; Saskatchewan native; South Carolina native; South Dakota native; Tennessee native; Texas native; Utah native; Vermont native; Virginia native; Washington native; West Virginia native; Wisconsin native; Wyoming native; Yukon native. Pacific: Cook Is. introduced; Fiji introduced; Hawaii (Hawaiian Is. introduced); New Caledonia introduced. South Africa introduced. South Sudan native. Southern America: Argentina Northeast (Argentina Distrito Federal introduced, Buenos Aires introduced ‒ introduced, Corrientes introduced, Córdoba introduced, Entre Ríos native, Formosa introduced, La Pampa introduced, Misiones introduced); Argentina Northwest (Catamarca introduced, Jujuy introduced, La Rioja native, Mendoza native, Salta native, San Juan native, San Luis introduced, Santiago del Estero introduced, Tucuman introduced); Argentina South (Chubut native, Neuquén native, Rio Negro native, Santa Cruz native, Santa Fé introduced, Tierra del Fuego (Argentina) native); Bahamas introduced; Bermuda native; Bolivia introduced; Brazil South (Paraná native, Rio Grande do Sul introduced, Santa Catarina native); Cayman Is. introduced; Chile Central native (Biobío native, Coquimbo introduced, La Araucania introduced, Maule introduced, O'Higgins introduced, Santiago native, Valparaíso introduced); Chile North (Antofagasta introduced, Atacama introduced); Chile South (Aisén introduced, Los Lagos native, Magellanes native); Colombia native; Costa Rica native; Cuba introduced ‒ introduced; Dominican Republic native; Ecuador introduced; El Salvador introduced; Guatemala native; Haiti (Haiti native, Navassa I. introduced); Honduras native; Jamaica native; Juan Fernández Is. native; Leeward Is. (Virgin Is. introduced); Netherlands Antilles introduced; Nicaragua introduced; Panamá introduced; Paraguay introduced; Peru native; Puerto Rico introduced; Uruguay introduced; Venezuela native; Windward Is. (Barbados introduced, Dominica introduced)
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