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Ботанический журнал, 2022, T. 107, № 8, стр. 794-799

KARYOLOGICAL STUDY OF SOME GERANIUM SPECIES (GERANIACEAE) FROM ARMENIA

A. Ghukasyan 1, *, R. Adamyan 2, **, A. Poghosyan 2, ***

1 Institute of Botany after A. Takhtajyan of National Academy of Science of RA
0063 Yerevan, Acharyan Str., 1, Armenia

2 Yerevan State University
0025 Yerevan, Alek Manukyan Str., 1, Armenia

* E-mail: botinst@sci.am
** E-mail: ruzanna-74@rambler.ru
*** E-mail: astchik@ysu.am

Поступила в редакцию 25.03.2022
После доработки 31.05.2022
Принята к публикации 14.06.2022

Полный текст (PDF)

Аннотация

The genus Geranium is represented by 20 species in Armenian flora. In the present study 9 species of this genus from the territory of Armenia were karyologically investigated for the first time: G. columbinum, 2n = 18 (sect. Columbinum); G. ibericum, 2n = 28, G. sylvaticum, 2n = 28 (sect. Geranium); G. lucidum, 2n = 40 (sect. Robertiana); G. pyrenaicum, 2n = 26; G. pusillum, 2n = 26; G. molle, 2n = 26; G. rotundifolium, 2n = 26; G. divaricatum, 2n = 26 (sect. Rotundifolia).

Keywords: chromosome number, karyotype, cytorace, flora of Armenia, Geranium

The species of the genus Geranium are common in temperate zones of the Earth and in high mountains of tropical zones. The genus Geranium includes about 400 species (Aedo, 2003). Most species are polymorphic. The genus Geranium is distributed in Armenia and is represented by 20 species from 8 sections: Sanguineum Knuth (1 species), Columbinum Koch (2), Geranium (5), Robertiana Boiss. (2), Palustra Knuth (2), Pyrenaica Knuth (1), Rotundifolia Gams. (5) and Tuberosa Boiss. (2) (Table 1). The classification of the genus was carried out according to the system of R. Knuth (Knuth, 1912).

Table 1.

Chromosomes numbers of the genus Geranium L. species (according to original and literature data)

Section Species Chromosomes numbers (2n)
Sanquineum Knuth G. sanguineum L. ≈56, 80, 82, 84
Columbinum Koch G. columbinum L. 18*
G. dissectum L. 22
Geranium G. ibericum Cav. 28, 56
G. montanum Habl. ex Pall. –
G .platypetalum Fisch. et C.A. Mey. 42
G. sylvaticum L. 24, 28
G. ruprechtii (Woronow) Grossh. –
Robertiana Boiss G. robertianum L. 32, 54, 56, 64
G. lucidum L. 20, ≈40, 40–44, 42
Palustra Knuth G. palustre L. 28, 56
G. collinum Steph. ex Willd. 28
Pyrenaica Knuth G. albanum Bieb. 28
Rotundifolia Gams. G. pyrenaicum Burm. f. 20, 22–24, 26, 28
G. pusillum L. 26, 34
G. molle L. 26
G. rotundifolium L. 26
G. divaricatum Ehrh. 26, 28
Tuberosa Boiss. G. tuberosum L. 28
G. linearilobum DC. –

Note. * The chromosomes numbers determined on the Armenian material by the authors are highlighted in bold.

Determination of chromosome numbers is the first step in any karyological study. As karyological data accumulate they can be used in the systematics. The study of chromosome numbers, in some cases, makes it possible to clarify the issues of evolution, speciation and phylogenetic relationships. Available data on chromosome numbers make it possible to better understand a taxon, species, or even family.

Data on the chromosome numbers of species can be found in different books of references, (Fedorov (ed.), 1969; Takhtajan (ed.), 1993; Moore, 1977; Goldblatt, 1981–1988; Goldblatt, Johnson, 1990–1996). Information on the chromosome numbers of species ocurring in Armenia is presented in the book “Chromosome numbers of flowering plants of the Armenian flora” (Nazarova, Ghukasyan, 2004), which summarizes the data on about 900 species from more than 2000 populations, 51 families belonging to 320 genera. Until now, the members of the genus Geranium growing in Armenia have not been studied karyologically.

MATERIAL AND METHODS

The determination of the chromosome numbers and the description of the species karyotypes were carried out by light microscopy. The seeds were germinated on wet filter paper in Petri dishes in the laboratory (19–21°C). The karyological studies were made on the mitotic metaphases of the meristematic cells from root tips. The root tips were pretreated in 0.4% colchicine solution for 2 hours and fixed in fluid 3:1 alcohol and glacial acetic acid for at least 2 hours at room temperature. After hydrolysis in HCl 1N for 10–15 minutes at 60°C, the root tips were stained in Schiff reagent for 1.5 hours. Then the root tips were squashed on a glass slide with 45% acetic acid. For all chromosome counts, at least 10 plates were examined for each taxon.

The stained roots were washed in warm water and placed on a glass slide. The colored tip of the rootlet (0.5–1 mm) was crushed in 45° acetic acid under a coverslip. After a preliminary examination of the smears under a microscope, the preparations were placed in buthyl alcohol for 5 minutes, then in xylene for 5 minutes, and were placed in Canadian balsam.

The karyological study was based on the samples of the genus Geranium (Geraniaceae) species collected by R. Adamyan, K. Tamanyan, G. Fayvush and others, during expeditions in 2003–2010 in different regions of Armenia. Herbarium material of the Institute of Botany of National Academy of Sciences (ERE) and Yerevan State University (EREU) was studied.

RESULTS AND DISCUSSION

There are 20 species of the genus Geranium in the flora of Armenia, and following species were studied by us karyologically: G. columbinum, 2n = 18 (sect. Columbinum); G. ibericum, 2n = 28, G. sylvaticum, 2n = 28 (sect. Geranium); G. lucidum, 2n = 40 (sect. Robertiana); G. pyrenaicum, 2n = 26; G. pusillum, 2n = 26; G. molle, 2n = 26; G. rotundifolium, 2n = 26; G. divaricatum, 2n = 26 (sect. Rotundifolia) (Тable 1).

According to the literature data on the chromosome numbers of the genus Geranium species, presented in Table 1, the different basic chromosome numbers x = 9, 10, 11, 12, 14, 15, 16, 17, 23, 25 are revealed.

G. columbinum L., 2n = 18

Examined samples: Lori Province, near the village of Akhtala, 805 m a.s.l., 41°09'26''N 44°45'56''E, 24.06.2009, leg. G. Fayvush, К. Tamanyan, N 09-1804 ERE, W, MA, PE, det. R. Adamyan; Tavush Province, above Yenokavan, a cave in the valley of the river Tanzout. Lastiver, 1260–1360 m a.s.l., 41°54'57" N 44°4'46" E, 06/23/2008, leg. G. Faivush, K. Tamanyan, N 08-0790 ERE, W, MA, PE, det. R. Adamyan.

The species is well studied karyologically. According to the literature data, only the diploid cytorace is characteristic for G. columbinum with 2n = 18 (Fedorov, 1969; Moore, 1973; Murin, 1974; Alves, Leitao, 1976; Van Loon, 1984b). Our material also revealed a diploid cytorace 2n = 2х = 18, with a basic chromosome number x = 9. The karyotype of G. columbinum is asymmetric, consisting of 2 pairs of submetacentric and 7 pairs of metacentric chromosomes. Karyotype formula: 2n = 18 = 4SM+14M.

G. ibericum Cav., 2n = 28

Examined samples: Ashotsk Province, climb to the top of Mount Kachal Sar, 2600–2900 m a.s.l., 01.08.2009, leg. E. Gabrielyan, E. Nazarova, M. Hovhannisyan, N 13619 ERCB, det. R. Adamyan; Kotayk Province, Tsakhkadzor, Teghenis 2250 m a.s.l., 40°32'08" N 44°41'02", 05.22.2005, leg. Aedo, Fayvush, Tamanyan et al. No. 17696 MA, det. S. Castroviejo; Kotayk Province, Tsaghkadzor, cableway 2300 m a.s.l., 40°32'08" N 44°41'02", 0.07.2010 leg. R. Adamyan, N 13618 ERCB, det. R. Adamyan.

The species is poorly studied karyologically. According to the literature data, for the European species G. ibericum, mainly the tetraploid cytorace was found, 2n = 4x = 56 (Gauger, 1937; Warburg, 1938), in addition, a diploid cytorace is known for this species, 2n = 28 (Van Loon, 1984a). On our material the diploid cytorace was also identified for this species 2n = 2x = 28, with basic chromosome number x = 14 (Fig. 1.1). The karyotype of G. ibericum is asymmetric, consisting of 7 pairs of metacentric, 4 pairs of submetacentric and 3 pairs of chromosomes with satellites: 2 pairs of metacentric and 1 pair of submetacentric chromosomes with satellites. Karyotype formula: 2n = 28 = 14M + 8SM + 4MS + 2SMS.

Fig. 1.

Metaphase plates of the genus Geranium species.

1 – G. ibericum (2n = 2x = 28); 2 – G. sylvaticum (2n = 2x = 28); 3 – G. lucidum (2n = 4x = 40); 4 – G. pyrenaicum (2n = 2x = 26); 5 – G. pusillum (2n = 2x = 26); 6 – G. molle (2n = 2x = 26); 7 – G. rotundifolium (2n = 2x = 26); 8 – G. divaricatum (2n = 2x = 26).

G. sylvaticum L., 2n = 28

Examined samples: Vayots Dzor, Vayk Province, near the village of Kechut, 1200–2050 m a.s.l., 39°49'N 45°38'E, 19.06.2004, leg. К. Tamanyan, G. Fayvush, M. Hovanisyan, N 04-0851, det. R. Adamyan; Kotayk Province, Hankavan, territory of YSU base, 1600 m a.s.l., 20.07.2010, leg. R. Adamyan, N 13617 ERCB, det. R. Adamyan; Kotayk Province, Tsaghkadzor, Teghenis, 2250 m a.s.l., 40°32'08" N 44°41'02", 06.22.2005, leg. K. Aedo et al., No. 17705 MA, det. R. Adamyan.

The species is well studied karyologically. According to the literature data, mainly the diploid cytorace is characteristic for G. sylvaticum, 2n = 28, with basic chromosome number x = 14 (Fedorov, 1969; Moore, 1973; Uhrikova, Majovsky, 1980; Strid, Franzen, 1981; Arohonka, 1982; Van Loon, 1984a; Takhtajan, 1990). Only for the Belarusian material, S.A. Dmitrieva gives the chromosome number 2n = 24 (Dmitrieva, 1986). Our material also revealed a diploid cytorace for this species, 2n = 2х = 28, with basic chromosome number x = 14 (Fig. 1.2 ). The karyotype of G. sylvaticum is asymmetric, consisting of 6 pairs of metacentric, 5 pairs of submetacentric and 3 pairs of chromosomes with satellites: 2 pairs of metacentric and 1 pair of submetacentric chromosomes with satellites. Karyotype formula: 2n = 28 = 12M+10SM+4MS+ 2SMS.

G. lucidum L., 2n = 40

Examined samples: Syunik Province, on the road Goris × Kapan, 1370–1600 m a.s.l. 39°20' N 46°23'–24' E, 05.07.2003, leg. G. Fayvush, K. Tamanyan, N 03-0859 ERE, W, MA, PE, det. R. Adamyan; Syunik Province, Nerkin Ande village, 660 m a.s.l., 39°03'47" N 46°31'58" E, 06.25.2005, leg. K. Aedo et al., N 313 MA, det. R. Adamyan.

The species is well studied karyologically. According to the literature data, di- and tetraploid cytoraces were found in most cases for the species, 2n = 20, 40 (Fedorov, 1969; Alves, Leitao, 1976; Uhrikova, Majovsky, 1980; Strid, Franzen, 1981; Van Loon, 1984b; Galland, 1988; Luque, Díaz Lifante, 1991; Hollingsworth et al., 1992). In addition, a hexaploid cytorace 2n = 60 (6x) is given for Indian populations of G. lucidum (Aryavand, 1983), and for Belarusian populations, an octoploid chromosome number is also known, 2n = 80 (Petrova, Stanimirova, 2003). For European populations of the species, chromosome numbers 2n = 40–44 were exhibited (Van Loon, 1984 a, b). In the studied specimens of G. lucidum, we found tetraploid cytorace 2n = 4х = 40, with basic chromosome number х = 10 (Fig. 1.3 ). The karyotype of this species is asymmetric, consisting of 5 pairs of metacentric, 12 pairs of submetacentric and 3 pairs of submetacentric chromosomes with satellites. Karyotype formula: 2n = 40 = 10M+24SM+6SMS.

G. pyrenaicum Burm. F., 2n = 26

Examined samples: Lori Province, northwest of the road from Stepanavan to Spitak, 1965 m a.s.l., 40°52'27"N 43°59'39"E, 06.23.2009, leg. G. Faivush et al., N 09-1717 ERE, W, MA, PE, det. R. Adamyan; Tavush Province, vicinity of Dilijan, Kurudakh tract, meadow, 1913 m a.s.l., 40°38'47"N 44°29'51"E, 07.10.2005, leg. K. Tamanyan, G. Faivush, M. Hovhannisyan, N 12616 ERCB, det. R. Adamyan; Kotayk Province, Hankavan, territory of YSU base, forest, 1600 m a.s.l., 07.06.2010, leg. R. Adamyan, N 13615 ERCB, det. R. Adamyan; Armavir Province, Etchmiadzin, Tsiatsan village, 1350 m a.s.l., 40°09'42"N 44°17'28"E, 17.08.2010, leg. R. Adamyan, N 13614 ERCB, det. R. Adamyan.

The species is well studied karyologically. According to the literature data, mainly the diploid cytorace is characteristic for G. pyrenaicum, 2n = 26 (Fedorov, 1969; Moore, 1977; Murin, 1974; Alves, Leitao, 1976; Skalinska et al., 1976, 1978; Strid, 1980; Strid, Franzen, 1981; Mizianty et al., 1983; Van Loon, 1984a; Takhtadjan, 1990; Galland, 1988; Baltisberger, 1991), only for Indian populations chromosome number 2n = 2x = 20 is given, with basic chromosome number x = 10 (Sharma, 1970), and for Europian populations – 2n = 28 with basic number x = 14 (Warburg, 1938). In the studied specimens of the species G. pyrenaicum we also revealed a diploid cytorace 2n = 2x = 26, with basic chromosome number x = 13, which confirms the previously presented numerous data (Fig. 1.4 ). Karyotype of this species is asymmetric, consisting of 8 pairs of metacentric, 3 pairs of submetacentric chromosomes and on 1 pair of meta- and 1 pair of submetacentric chromosomes with satellites. Karyotype formula: 2n = = 26 = 16M+6SM+2MS+2SMS.

G. pusillum L., 2n = 26

Examined samples: Tavush Province, northwest of Ijevan, near Yenokavan village, 1365 m a.s.l., 40°24'54"N 45°18'43"E, 11.06.2008, leg. K. Tamanyan, G. Faivush, E. Vitek, N 08-0704 ERE, W, MA, PE, det. R. Adamyan; Tavush Province, Dilijan, the territory of the House of Composers, 1200–1500 m a.s.l., 40°44'27"N 44°51'47"E, 07.20.2010, leg. R. Adamyan, N 13613 ERCB, det. R. Adamyan; Yerevan, Khanjyan street, 1020 m a.s.l., 40°18'24"N 44°52'72"E, 05.23.2012, leg. R. Adamyan, N 13612 ERCB, det. R. Adamyan.

The karyology of this species has been fairly well investigated. According to the literature data, mainly the diploid cytorace is found for G. рusillum, 2n = 26, with basic chromosome number x = 13 (Fedorov, 1969; Moore, 1977; Murin, 1974; Skalinska et al., 1976; Fernandes Cassas, Pascual, 1978; Arohonka, 1982; Van Loon, 1984b; Buttler, 1989). In the studied specimens of G. рusillum, we found the diploid cytorace 2n = 2x = 26, with basic chromosome number x = 13, which confirms numerous previousdata (Fig. 1.5 ). The karyotype of this species is asymmetric. Diploid set of chromosomes consists of 8 pairs of metacentric, 3 pairs of submetacentric chromosomes and 2 pairs of metacentric chromosomes with satellites. Karyotype formula: 2n = 26 = 16M + 6SM + 4MS.

G. molle L., 2n = 26

Examined sample: Syunik Provonce, not far from the Shikahogh State Reserve, m a.s.l., 06.04.2008, leg. E. Gabrielyan, N 13611 ERCB, det. R. Adamyan.

The karyology of this species has been fairly well investigated. According to the literature data, mainly the diploid cytorace is found for G. molle, 2n = 26 (Fedorov, 1969; Moore, 1977; Alves, Leitao, 1976; Murin, 1978; Franzen, Gustafsson, 1983; Van Loon, 1984b; Hill, 1989). In the studied specimens of the species, we found the diploid cytorace 2n = 2x = 26, with basic chromosome number x = 13, which confirms numerous previous data (Fig. 1.6 ). The karyotype of this species is asymmetric, consisting of 3 pairs of metacentric, 8 pairs of submetacentric chromosomes, and 1 pair of metacentric and 1 pair of submetacentric chromosomes with satellites. Karyotype formula: 2n = 26 = 6M+16SM+2MS+2SMS.

G. rotundifolium L., 2n = 26

Examined samples: Lori Province, bank of the river Berd, 1290 m a.s.l., 41°08'0"N 44°26'02"E, 06.28.2003, leg. G. Faivush, K. Tamanyan, A. Ter-Voskanyan, E. Vitek, N 03-0200 ERE, W, MA, det. R. Adamyan; Syunik Province, Meghri district, at the fork of Shvanidzor village, from the side of Nyuvadi village, 500 m a.s.l. 05.09.2004, leg. E. Gabrielyan, M. Hovhannisyan, N 13610 ERCB, det. K. Tamanyan; Syunik Province, eastern part of Zangezur, west of Goris-Kapan road, near village Gurdbulakh, 1370–1600 m a.s.l., 40°20'0"N 46°23'24"E, 05.07.2003, leg. G. Faivush, K. Tamanyan, A. Ter-Voskanyan, E. Vitek, N 03-0919 ERE, W, MA, det. R. Adamyan.

The karyology of this species has been fairly well investigated. According to the literature data mainly the diploid cytorace is characteristic for G. rotundifolium, 2n = 26 (Fedorov, 1969; Moore, 1977; Alves, Leitao, 1976; Strid, Franzen, 1981; Franzen, Gustafsson, 1983; Buttler, 1989; Galland, 1988). In the studied specimens we also found the diploid cytorace 2n = = 2х = 26, with basic chromosome number x = 13 (Fig. 1.7 ). The karyotype of this species is asymmetric, consisting of 8 pairs of metacentric, 3 pairs of submetacentric chromosomes and 1 pair of meta-, 1 pair of submetacentric chromosomes with satellites. Karyotype formula: 2n = 26 = 16M+6SM+2MS+2SMS.

G. divaricatum Ehrh., 2n = 26

Examined samples: Kotayk Province, near Bjni village, 1550 m a.s.l., 40°27'35"N 44°39'34". 06.17.2004, leg. E. Vitek, K. Tamanyan, G. Faivush, M. Hovhannisyan, A. Ter-Voskanyan, N 04-0571 ERE, W, MA, det. R. Adamyan; Kotayk Province, Zovuni village, near the monument, 1200 m a.s.l., 05.20.2010, leg. R. Adamyan, N 13609 ERCB, det. R. Adamyan; Vayots Dzor Province, village Ger-Ger, 1780 m a.s.l., 39°47'09"N 45°32'15", 06.24.2005, leg. K. Aedo, G. Faivush, K. Tamanian, N 2869 MA, det. A. Guerrero.

The species is poorly studied karyologically. According to the literature data, for the Slovakian G. divaricatum specimens the chromosome number 2n = 26 mainly was found, with basic number x = 13 (Murin, 1974), for the European specimens the diploid cytorace 2n = 28 is given, with basic number x = 14 (Dersch, 1974; Van Loon, 1984?). In our material, we also revealed the diploid cytorace 2n = 2х = 26, with basic chromosome number x = 13 (Fig. 1.8 ). The karyotype of this species is asymmetric, consisting of 8 pairs of metacentric, 4 pairs of submetacentric chromosomes and 1 pair of metacentric chromosomes with satellites. Karyotype formula: 2n = 26 = 16M+8SM+2MS.

CONCLUSIONS

The karyological research of the genus Geranium species of the Armenian flora has revealed the following: we discovered diploid cytoraces for 8 species – G. columbinum, 2n = 18 (sect. Columbinum), G. ibericum, 2n = 28, G. sylvaticum, 2n = 28 (sect. Geranium), G. pyrenaicum, 2n = 26, G. pusillum, 2n = 26, G. molle, 2n = 26, G. rotundifolium, 2n = 26, G. divaricatum, 2n = 26 (sect. Rotundifolia), with different basic chromosome numbers, x = 9, 13, 14. For the species G. lucidum (sect. Robertiana) a tetraploid cytorace 2n = 40 was discovered with the basic chromosome number x = 10. The number of chromosomes for all karyologically studied species of the genus Geranium the flora of Armenia is brought up for the first time.

The species of the section Rotundifolia occurring in Armenia (G. pyrenaicum, 2n = 26; G. pusillum, 2n = 26; G. molle, 2n = 26; G. rotundifolium, 2n = 26; G. divaricatum, 2n = 26) are characterized with the basic main chromosome number x = 13.

The main basic chromosome number x = 9 is typical of the Armenian species of the section Columbinum (G. columbinum, 2n = 18).

The main basic chromosome number x = 10 is typical of the Armenian species of the section Robertiana (G. lucidum, 2n = 40).

The main basic chromosome number x = 14 is typical of the Armenia species of the section Geranium (G. ibericum, 2n = 28; G. sylvaticum, 2n = 28).

Asymmetric karyotype is observed in all the karyologically studied species of the genus Geranium.

Список литературы

  1. Aedo C. 2003. Taxonomic revision of Geranium sect. Trygonium (Geraniaceae). – Bot. Zhurn. 88: 124–131.

  2. Alves M., Leitao M. 1976. Contribucialo para o conhecimento citotaxonomico das spematophyta de Portugal. XIII. Geraniaceae. – Bol. Soc. Brot. 50: 231–245.

  3. Arohonka T. 1982. Chromosome counts of vascular plants of the island Seili in Nauvo, SW Finland. – Biologian–Laitoksen Julkaisuja. 3: 1–12.

  4. Aryavand A. 1983. In IOPB Chromosome number reports LXXIV. – Taxon. 32: 321–322.

  5. Baltisberger M. 1991. Cytological investigations of some Greek plants. – Fl. Medit. 1: 157–173.

  6. Buttler K.P. 1989. Chromosomenzahlen von Gefässpflanzen aus Hessen 4. Folge. – Hess. Florist. Briefe. 38: 11–14.

  7. Dersch G. 1974. Über einige Chromosomenzählungen an mitteleuropäischen Biütenpflanze. II. – Philippia. 2: 75–82.

  8. Dmitrieva S.A. 1986. Chromosome numbers of some species of vascular plants from Belorussia. – Bot. Zhurn. 71 (8): 1145–1147 (In Russ.).

  9. Fedorov A.A. (ed.). 1969. Chromosome Numbers of Flowering Plants. Leningrad. 926 p. (In Russ.).

  10. Fernandes Cassas J., Pascual R. 1978. In Númerous chromosómicos para la flora Española. – Lagascalia. 7: 105–125.

  11. Franzen R., Gustafsson L.A. 1983. Chromosome numbers in flowering plants from the high mountains of Sterea Ellas, Grees. – Willdenowia. 13: 101–106.

  12. Galland N. 1988. Recherche sur l’origine de la flore orophile du Maroc étude caryologique et cytogéographique. – Trav. Inst. Sci. Univ. Mohammed V, Sér. Bot. (Rabat). 35: 1–168.

  13. Gauger W. 1937. Ergebnisse einer zytologischen Untersuchung der Familie der Geraniaceae. I. – Planta. 26 (4): 529–531.

  14. Goldblatt P. 1981. Index to plant chromosome numbers. – Monogr. Syst. Bot. Missouri Bot. Gard. 5: 276–277; 1984. 8: 201–202; 1985. 13: 105; 1988. 23: 117.

  15. Goldblatt P., Johnson D. 1990. Index to plant chromosome numbers. – Monogr. Syst. Bot. Missouri Bot. Gard. 30: 98–99; 1991. 40: 113–114; 1994. 51:108–109; 1996. 58: 124–125.

  16. Hill L.M. 1989. IOPB chromosome data I. – Int. Organ. Plant Biosyst. Newslett. 13: 17–19.

  17. Hollingsworth P., Gornall R., Bailey J. 1992. Contribution to a cytological catalogue of the british and Irish flora, 2. – Watsonia. 19: 134–137.

  18. Knuth R. 1912. Geraniaceae Juss. In: Engler A. Das Pflanzenreich regni vegetabilis conspectus. Leipzig, Engelman. 53: 640 p.

  19. Luque T., Díaz Lifante. 1991. Chromosome numbers of plants collected during Iter Mediterraneum I in the SE of Spain. – Bocconea. 1: 303–364.

  20. Mizianty M., Mirek Z., Frey L. 1983. Chromosome numbers of Polish vascular plants. Part 4. – Acta Soc. Bot. Poloniae. 52: 205–214.

  21. Moore R.I. 1977. Index to plant chromosome numbers for 1973–1974. Regnum Vegetabile. Utrecht. 257 p.

  22. Murin A. 1974. In Index of chromosome numbers of Slovakian flora. Part 4. – Acta Fac. Rerum Nat. Univ. Comenianae, Bot. 23: 1–23.

  23. Murin A. 1978. In Index of chromosome numbers of Slovakian flora. Part 6. – Acta Fac. Rerum Nat. Univ. Comenianae, Bot. 26: 1–42.

  24. Nazarova E.A., Gukasyan A.G. 2004. Chromosome numbers of flowering plants of Armenia flora, Yerevan. Institute of Botany NAS RA. 171 p. (In Russ.).

  25. Petrova A., Stanimirova P. 2003. Karyological study of some Geranium (Geraniaceae) species growing in Bulgaria. – Bocconea. 16 (2): 675–682.

  26. Sharma A.K., 1970. Annual report, 1967–1968. – Res. Bull. Univ. Calcutta Cytogenetics Lab. 2: 1–50.

  27. Skalinska M., Pogan E., Wcislo H. 1976. Further studies in chromosome numbers of Polish angiosperm. XI. – Acta. Biol. Cracov. 19: 107–148.

  28. Skalinska M., Pogan E., Czapik R. 1978. Further studies in chromosome numbers of Polish angiosperm. XII. – Acta. Biol. Cracov. 21: 31–63.

  29. Strid A. 1980. In chromosome number reports LXIX. – Taxon. 29: 709–710.

  30. Strid A., Franzen R. 1981. In chromosome number reports LXXIII. – Taxon. 30: 829–842.

  31. Takhtajan A.L. (ed.) 1993. Chromosome Numbers of flowering plants of the Flora of the USSR. St. Petersburg. Vol. 1. 508 p. (In Russ.).

  32. Uhrikova A., Majovsky J. 1980. In chromosome number reports LXIX. – Taxon. 29: 725–725.

  33. Van Loon, J.C. 1984a. Chromosome numbers in Geranium from Europe, I. The perennial species. – Proceedings Koninklijke Nederlandse Akademie van Wetenschappen. Series C, Biological and Medical Sciences. 87: 263–277.

  34. Van Loon, J.C. 1984b. Chromosome numbers in Geranium from Europe, II. The annual species. – Proceedings Koninklijke Nederlandse Akademie van Wetenschappen. Series C, Biological and Medical Sciences. 87: 279–296.

  35. Warburg E.F. 1938. Taxonomy and relationship in the Geraniales in the light of their cytology. – New Phytol. 37 (2): 130–159.

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