Chromosome numbers of invasive species of the Altai Republic flora. II

Chromosome numbers (2n) in 16 invasive plant species from the families Amaranthaceae, Asteraceae, Caryophyllaceae, Lamiaceae, Onagraceae and Poaceae are reported on the material from the Altai Republic for the first time. Among them, chromosome complements were first examined in Russia for Chaiturus marrubiastrum (L.) Ehrh. ex Reichenb. (2n = 24) and Oenothera villosa Thunb. (2n = 14). Five species: Amaranthus lividus L. (2n = 34), Elisanthe noctiflora (L.) Rupr. (2n = 24), Galeopsis speciosa Mill. (2n = 16), Oenothera biennis L. (2n = 14) and Bromus japonicus Thunb. (2n = 14), were studied first from Siberia. The most part of the species studied are diploids. The general distribution and the history of floristic findings of these species in the Altai Republic are given. The data on chromosome numbers published earlier on the material from Russia are cited.

The Altai Republic is currently one of the regions attracting a large number of tourists from different regions of Russia and other countries. As a result, the vegetation is exposed to significant anthropogenic impact, which contributes to invasion of a large number of alien species in the area and leads to further transformation in biological diversity of the flora. Invasive species, in their turn, may themselves undergo evolutionary changes in the new conditions. That is why, the comprehensive study and biological monitoring of alien species in the natural flora is becoming increasingly important. Given that 64 Lomonosova M. N. et al. Chromosome numbers of invasive species of the Altai Republic flora. II polyploidy is a likely factor in species invasiveness (te Beest et al., 2012, and references therein), we have undertaken a detailed karyological research of all alien species in flora of the Altai Republic. This study is a part of the project on karyological investigation of the invasive flora of South Siberia and is a continuation of previous work (Zykova et al., 2018). We provide an information on the history of floristic studies on examined species at the territory of the Altai Republic, and in some cases -in Siberia. When referring to P. N. Krylov (1909Krylov ( , 1931Krylov ( , 1949 we use the terminology of modern administrativeterritorial division. For all species, the references on chromosome numbers revealed on the material from Russia have been provided as well, as relevant information in the international databases "Еhe Chromosome Counts Database" (Rice et al., 2015) and "Index to Plant Chromosome Numbers" (Goldblatt, Johnson, 1979+) is reflected incompletely.
The ploidy level of samples was determined by direct count of chromosomes in metaphase on squash preparations of root meristem. All species were studied for the first time on the material collected in the Altai Republic. Seeds were germinated at 25 °C (day) and 16 °C (night) on sterile sand. The root tips were pretreated for two hours with 0.2 % colchicine water solution, fixed in 3 : 1 absolute ethanol-glacial acetic acid and stained with 1 % acetic hematoxylin. Metaphase plates were observed under 100× magnification by the Axioscope 40 (Karl Zeiss, Axio Lab) and photographed by the AxiCam MRc 5 digital camera. The herbarium specimens (vouchers) are saved in the Herbarium of the Central Siberian Botanical Garden SB RAS (NS).
For species marked with an asterisk (*), the chromosome number is determined for the first time in Russia, two asterisks (**) -for the first time in Siberia. Annual. This species was found in all developed agricultural areas of the world. In the Altai Republic the first findings were in Gorno-Altaysk (Ilyin, Fedotkina, 2008) and Yaylyu village (Zolotukhin, 1983). Currently, it is a common garden weed occurring on fallow lands and waste places in Gorno-Altaysk, Mayma and Turochak districts (Zykova, 2015).

AMARANTHACEAE
The chromosome number is determined first on Siberian material. The same number was obtained for Primorye Territory (Probatova, 2014, and references therein).  1A).
Annual. North-American species introduced in all continents including polar areas (Nikitin, 1983). It is one of the most aggressive actively spreading adventive species. The first findings in the Altai Republic were made in Ust-Muny village (Krylov, 1949), later it was mentioned for Ust-Kan district (Ilyin, Fedotkina, 2008). According to Zykova (2015), C. canadensis is common now in the northern districts of the Altai Republic.
Our chromosome count matches the number reported on the material from Buryatia Republic and Irkutsk Region (Chepinoga, 2014, and references therein), Kurgan Region (Krasnikov, Korolyuk, 1995), Krasnodar Territory (Probatova et al., 2012) and Primorye Territory (Rostovtseva, 1979;Probatova, 2014, and references therein) Perennial herb. Native area of this species includes West and Central Europe and Middle Asia, from where it is spread as ornamental plant to Eurasia, North America and New Zealand (Kamelin, 1973;Tupitsyna, 2004). Recently P. aurantiaca was found in the Altai Republic (Zykova, An'kova, 2017).
The same chromosome number was known from Leningrad city (Chuksanova et al., 1968), Novosibirsk Region (Krasnikov, Tupitsyna, 2004), Sakhalin Region (Probatova et al., 2007(Probatova et al., , 2017 and Komi Republic (Lavrenko et al., 1990). Annual or biennial. Eurasian species distributed in all continents is one of the most widespread adventive species in Siberia. The first findings on the territory of the Altai Republic were in Kibisen, Yaylyu and Oyrot-Tura villages, and near Karakol Mount (Krylov, 1949). At present, it is common in the northern part of the Altai Republic spreading sometimes to central regions (Zykova, 2015).
Our chromosome count is the first on the material from Siberia. The same chromosome number was reported earlier from Sakhalin Region (Probatova et al., 2007, and references therein). Annual or biennial. Eurasian species introduced elswhere. In the Altai Republic it has been recently found in Turochak village (Zykova, 2015). For several years, the species is actively distributed in disturbed habitats, where it often forms thickets. C. marrubiastrum occasionally enters in light forests.
The chromosome number is counted first in Russia. It confirms the only data previously reported from Romania (Tarnavschi, 1948 1E).
Annual. Euro-Mediterranean species, spreading in Eurasia. In the Altai Republic G. speciosa was first found in the late XX century at the mouth of Sema River, and in Marchela and Anos river basins (Pshenichnaya, 1997). Now this species is actively spreading in Gorno-Altaysk city and Mayma district, less common in Chemal and Shebalino districts (Ilyin, Fedotkina, 2008;Zykova, Erst, 2012;Zykova, 2014bZykova, , 2015. This is the first chromosome count in Siberian population, which agrees with the data reported for Amur Region (Sokolovskaya et al., 1986). Biennial. North American species, settled in Eurasia. In the Altai Republic it is found in Gorno-Altaysk city (Zykova, 2002), Mayma, Chemal (Shaulo et al., 2010) and Turochak ( Zykova, 2017) districts. O. biennis occurs much less often than the following species of Oenothera.
The chromosome number is determined for the first time on Siberian material. The same data were obtained from Stavropol Territory (Magulayev, 1984) and Primorye Territory (Probatova et al., 2014a), as well as Sakhalin Region (Probatova et al., 2007, and references therein). Biennial. North American species spreading in all continents. In the Altai Republic O. villosa was found first in the 1980s in Ust-Koksa and Choya districts (Ebel, 2008), later -in Gorno-Altaysk city and Mayma (Shaulo et al., 2010), Turochak (Zykova, 2015), Chemal (Zykova, 2014a) and Ulagan (Ebel, 2008;Zykova, 2017) districts. The species is quite active in the northern part of the Altai Republic, but in the central part only few locations were found.
The chromosome number is counted from Russia for the first time. Annual. European-W Asian species, as alien elsewhere. In the Altai Republic the first localities were found in 2009 in Gorno-Altaysk city (Shaulo et al., 2010) where this species now is actively propagating, and in Mayma district as well (Zykova, 2015).
The chromosome number is determined on the material from Siberia for the first time. The same data were obtained from the Crimea (Petrova, 1972), Dagestan Republic (Sokolovskaya, Probatova, 1979) and Primorye Territory (Probatova, 2014 and references therein). Annual. The species distribution range covers Central and East Siberia, the Far East, Manchuria. In the Altai Republic it was discovered in late XX century in Choya (Studenikina, 1999) and Ulagan (Pyak, Ebel, 2001) districts. Now this species is quite common in the northern region and is found in central part of the Altai Republic (Shaulo et al., 2010;Seregin, 2012;Zykova, 2014aZykova, , 2015Zykova, , 2017.
Annual or biennial. Euro-Mediterranean species, introduced elsewhere. In the Altai Republic it was imported together with seeds for coating roadside slopes along Chuya Tract in 2011 (Zykova, 2014b). It occurs along the motorway and adjacent meadows; registered only in Mayma district of the Altai Republic (Zykova, 2015).

Conclusion
By now, chromosome numbers of 29 invasive species distributed in the Altai Republic were examined. Among them, the data for 16 species are represented in this study. For the first time Chaiturus marrubiastrum and Oenothera villosa have been studied on the material from Russia. The chromosome numbers of studied species are constant through their distribution range and diploid species prevail between. The exceptions are Tripleurospermum inodorum and Panicum ruderale, for which two chromosome complements are given in the literature. In recent years five polyploid species (Amaranthus lividus, A. retroflexus, Eragrostis amurensis, Hordeum jubatum and Tripleurospermum inodorum) have been introduced most actively in the Altai Republic, as well as lately discovered Pilosella aurantiaca, that is spreading in Turochak district very quickly now. A rarer diploid cytotype (2n = 18) was found in Panicum ruderale which was revealed by previous authors as tetraploid (2n = 36) in the vast majority of studied populations.