ORIGINAL ARTICLE

Additions to the cytologically investigated species of Potentilla L.(Rosaceae) from India

Savita Rani • Sanjeev Kumar • Syed Mudassir Jeelani •

Raghbir Chand Gupta • Santosh Kumari

Received: 26 May 2011 / Accepted: 24 October 2011 / Published online: 15 December 2011

� Springer-Verlag 2011

Abstract At present 14 species of Potentilla L. have been

cytologically worked out from different geographical areas

of Kashmir and Himachal Pradesh in the Western Hima-

layas. New chromosome numbers in nine species—

Potentilla argyrophylla (n = 14), P. atrosanguinea

(n = 7, 14), P. desertorum (n = 7), P. gerardiana

(n = 14), P. indica (n = 14), P. micropetala (n = 14),

P. nepalensis (n = 14), P. sibbaldia (n = 14) and

P. thomsonii (n = 7)—have been reported on a worldwide

basis for the first time. Additional chromosomal races of

polyploid cytotypes for P. argyrophylla (n = 28) and

P. desertorum (n = 14) along with a diploid cytotype

for P. micropetala (n = 7) plus diploid cytotypes for the

five species as P. fulgens (n = 7), P. gelida (n = 7),

P. kleiniana (n = 7), P. sibbaldia (n = 7) and P. sundaica

(n = 7) as well as a tetraploid cytotype for P. fruticosa

(n = 14) all have been cytologically worked out from India

for the first time. The course of meiosis varies from normal

to abnormal in different populations of the majority of

the species, such as P. argyrophylla, P. atrosanguinea,

P. desertorum, P. fruticosa, P. fulgens, P. gelida, P. indica,

P. nepalensis, P. sibbaldia and P. sundaica, except for

normal meiosis observed in P. gerardiana, P. kleiniana,

P. micropetala and P. thomsonii. The anomalous taxa are

marked with meiotic abnormalities in the form of cyto-

mixis, chromosomal stickiness, unoriented bivalents, for-

mation of laggards and bridges resulting in abnormal

microsporogenesis, and production of heterogenous-sized

fertile pollen grains along with reduced pollen fertility. All

the taxa with normal meiotic courses show nearly one

hundred percent pollen fertility.

Keywords Cytotypes � Meiotic abnormalities �Pollen fertility � Polyploidy � Potentilla �Himachal Pradesh � Kashmir

Introduction

The genus Potentilla L. includes over 500 species, mainly

distributed in the temperate regions of the Northern

Hemisphere, including 40 species from India (Santapau

and Henry 1973). The species of the genus are character-

ized as perennial or annual herbs or small shrubs usually

with creeping stems rooted at nodes; leaves are pinnate or

palmately compound; the leaflets are usually lobed with

stipules adnate to the petiole. These species prefer open

habitats, especially alpine or sub-alpine, even in xeric

conditions in the mountains. On the basis of leaf mor-

phology and stamen characters, the genus Potentilla is

charecterized in the systematically difficult genera of the

family Rosaceae (Szafer and Pawlowski 1955). Hooker

(1879) classified the genus Potentilla into three sections

while describing the Flora of British India. Because of

ambiguity at the species level, it has been divided into

many subgenera, sections, several subsections, series and

groups of species (Wolf 1908; B all et al. 1968). However,

in the regional Floras of India pertaining to the Western

Himalayas, no such taxonomic segregation is available

(Chaudhary and Wadhwa 1984; Dhaliwal and Sharma

1999; Kaur and Sharma 2004). Recently, molecular studies

have been carried out for analysis of the phylogeny, phy-

logeography, generic circumscription, cytoptype and

genetic diversification in different species of Potentilla

S. Rani � S. Kumar � S. M. Jeelani (&) �R. C. Gupta � S. Kumari

Department of Botany, Punjabi University,

Patiala 147 002, Punjab, India

e-mail: sandrabi555@gmail.com

123

Plant Syst Evol (2012) 298:485–497

DOI 10.1007/s00606-011-0560-x

from different regions of the world (Eriksson et al. 1998;

Dobes and Paule 2010; Paule et al. 2011). These studies have

provided a phylogenetic framework for the classification of

different tribes of the subfamily Rosoideae, especially for the

tribe Potentilleae, and proposed placement of the Potentilla

species in different sequenes. Although sexuality is the means

of reproduction in the majority of species of the genus, some of

the species are also well known to reproduce apomictically

(Muntzing 1958; Asker and Frost 1970; Holm and Ghatnekar

1996a, b; Gregor et al. 2002). The documented proof of aga-

mospermy and hybridization accompanying the polyploidi-

zation of genomes has also been considered to have played a

significant role in the evolution of the genus (Gentscheff 1938;

Muntzing and Muntzing 1941; Gustafsson 1947a; Rutishauser

1948; Kashin 1999; Nylehn et al. 2003). Regarding cytolog-

ical studies on the genus from India, until now a total of

eleven species—P. atrosanguinea, P. fragarioides, P. fulgens,

P. indica, P. micropetala, P. nepalensis, P. sundaica

and P. supina from the Western Himalayas; P. fulgens and

P. mooniana from the Eastern Himalayas; and P. indica,

P. kleiniana and P. leschenaultiana from South India—

present only scanty cytological information for Indian

species. To fill the lacuna and to make an addition to

the chromosomal database of lesser known angiosper-

mic genera, the present meiotic studies on 14 species

of Potentilla L. were carried out on a population

basis from different geographical areas of the Western

Himalayas.

Materials and methods

For meiotic studies, flower buds were collected from selected

localities of the far distant areas of the Western Himalayas

(Table 1). Smears of appropriately sized flower buds were

made after fixing them in Carnoy’s fixative, using the standard

acetocarmine technique. Pollen fertility was estimated by

mounting mature pollen grains in a glycero-acetocarmine (1:1)

mixture. Well-filled pollen grains with stained nuclei were

taken as apparently fertile, while shrivelled and unstained

pollen grains were counted as sterile. Photomicrographs of

pollen mother cells and pollen grains were made from freshly

prepared slides using the Nikon 80i Eclipse Digital Imaging

System. Voucher specimens were deposited in the Herbarium,

Department of Botany, Punjabi University, Patiala (PUN).

Results and discussion

Cytological study

Detailed cytological studies were carried out on 45 popu-

lations belonging to 14 species of the genus Potentilla. The

data regarding locality with altitude, accessions and mei-

otic chromosome numbers along with figure numbers of the

presently worked out species are presented in Table 1. The

results for each species with new/varied chromosome

counts and abnormal meiotic courses are discussed below.

Potentilla argyrophylla Wall. ex Lehm.

Of the six populations of the species, two each from Kashmir

and Himachal Pradesh all depict n = 14, and the other two

populations again from Himachal Pradesh show n = 28. The

present counts make new records of the tetraploid cytotype

(n = 14) on a worldwide basis and octaploid (n = 28) cyto-

types on an Indian basis. The species was earlier known to have

cytotypes with higher ploidy levels such as 6x = 2n = 42

(Zhukova 1967), 8x = 2n = 56 (Shimotomai 1930; Popoff

1935; Gustafsson 1947b; Zhukova 1967), 9x = 2n = 63

(Gentscheff 1938), and 10x = 2n = 70, 11x = 2n = 77,

12x = 2n = 84, 13x = 2n = 91 and 14x = 2n = 98 (Pop-

off 1935) from different parts of the USSR.

P. atrosanguinea Raf.

Of the six populations of the species, one from Kashmir

and two from Himachal Pradesh show n = 7 and exist as

diploid cytotypes. Of the remaining three populations from

Himachal Pradesh, two depict n = 14, and only one pop-

ulation reveals n = 28. The present chromosome counts of

n = 7 and n = 14 add new diploid and tetraploid cyto-

types, respectively, for the species on a worldwide basis,

while the octaploid cytotype (n = 28) is in conformity with

the earlier reports from India and outside India. The species

was previously known to have 8x to 12x levels as 2n = 56

(Shimotomai 1930; Christoff and Papasova 1943; Gosw-

ami and Metfield 1978), 2n = 63 (Zhukova 1967) and

2n = 74, 84 (Goswami and Metfield 1978) from different

parts of the world.

P. desertorum Bunge

Of the two populations of species collected from Himachal

Pradesh, one shows n = 7, and the other reveals n = 14. The

diploid cytotype (n = 7) has been cytologically worked out

for the first time on a worldwide basis, while the tetraploid

cytotype (n = 14) is chromosomally known for the first time

from India and confirms the previous reports from Mongolia

and Tadzhikistan (Mesicek and Sojak 1969, 1993) and the

USSR (Guinochet and Lefranc 1981).

P. fruticosa Gray

The single accession of the species collected from Hima-

chal Pradesh depicts n = 14 (a tetraploid cytotype) and is

486 S. Rani et al.

123

Table 1 Data showing the location, altitude, accession number, present chromosome number, figure number, nature of the meiotic course,

ploidy level and pollen size in the presently worked out species of Potentilla L. from the Western Himalayas

Name of taxa Locality Altitude

(m)

Accession

no. (PUN)

Meiotic

chromosome

number (n) with

respective figure

Nature of

meiotic

course/ploidy

level (x)

Pollen size (lm)

P. argyrophylla Wall.

ex Lehm

Thajwas, Distt.

Ganderbal,

Kashmir

3,000 54268 14 (Fig. 1a) Normal/4x 23.95 9 20.89

Gulmarg, Distt.

Baramullah,

Kashmir

2,600 54910 14 Normal/4x 23.89 9 20.91

Chota-Bhangal,

Distt. Kangra,

H.P.

2,300 54860 14 Abnormal/4x 22.61 9 18.37–24.94 9 24.01a

Baragran, Distt.

Kangra, H.P.

3,000 54896 14 Abnormal/4x 22.45 9 18.26–25.05 9 23.92

Chur peaks, Distt.

Sirmaur, H.P.

3,200 54876 28 (Fig. 1b) Normal/8x 26.95 9 24.37

Bhangyanimata,

Distt. Sirmaur,

H.P.

2,800 54878 28 Abnormal/8x 24.35 9 19.76–27.15 9 24.79

P. atrosanguinea Raf. Aharbal, Distt.

Kulgam, Kashmir

2,400 54905 7 Normal/2x 22.71 9 20.21

Biching, Distt.

Kangra, H.P.

3,000 54889 7 Normal/2x 22.63 9 19.97

Bara-Bhangal,

Distt. Kangra,

H.P.

4,000 54880 7 (Fig. 1c) Abnormal/2x 21.14 9 18.58–23.16 9 20.69

Haripurdhar, Distt.

Sirmaur, H.P.

2,600 54907 14 (Fig. 1d) Abnormal/4x 22.23 9 19.65–25.29 9 23.78

Jamnala, Distt.

Sirmaur, H.P.

2,600 54867 14 Normal/4x 24.28 9 22.07

Tisri, Distt.

Sirmaur, H.P.

3,000 54868 28 (Fig. 1e) Abnormal/8x 24.63 9 21.75–26.98 9 24.07

P. desertorum Bunge Triund,

Distt. Kangra,H.P.

3,000 54895 7 (Fig. 1f) Abnormal/2x 20.30 9 18.01–23.93 9 21.14

Biching, Distt.

Kangra, H.P.

2,200 54301 14 (Fig. 1g) Normal/4x 24.21 9 19.60

P. fruticosa Gray Tisri, Distt.

Sirmaur, H.P.

3,000 52668 14 (Fig. 1h) Abnormal/4x 23.14 9 20.13–25.11 9 22.29

P. fulgens Wall. ex D.

Don (=P. lineataTrevir.)

Multan, Distt.

Kangra, H.P.

2,300 54884 7 (Fig. 1i) Abnormal/2x 21.70 9 18.05–23.93 9 19.53

Triund, Distt.

Kangra, H.P.

3,000 54885 14 (Fig. 1j) Normal/4x 26.77 9 21.89

Dal lake, Distt.

Kangra, H.P.

3,800 54893 7 Normal/2x 22.21 9 18.25

Nauradhar, Distt.

Sirmaur, H.P.

1,800 54873 7 Abnormal/2x 21.12 9 18.16–23.88 9 19.49

Shillai, Distt.

Sirmaur, H.P.

2,000 54874 7 Normal/2x 22.32 9 18.17

P. gelida C. A. Mey. Thajwas, Distt.

Ganderbal,

Kashmir

3,100 54227 7 (Fig. 1k) Normal/2x 22.77 9 19.36

Chumnai, Distt.

Anantnag,

Kashmir

3,500 54228 7 Abnormal/2x 21.05 9 18.32–23.67 9 19.36

Additions to the cytologically investigated species of Potentilla L. 487

123

Table 1 continued

Name of taxa Locality Altitude

(m)

Accession

no. (PUN)

Meiotic

chromosome

number (n) with

respective figure

Nature of

meiotic

course/ploidy

level (x)

Pollen size (lm)

P. gerardiana Lindl.

(=P. fragarioides L.)

Haripurdhar, Distt.

Sirmaur,H.P.

2,800 54907 14 (Fig. 1l) Normal/4x 24.59 9 22.38

P. indica (Andrews)

T. Wolf (=Duchesneaindica (Andrews)

Focke

Dara, Distt.

Srinagar, Kashmir

1,900 54200 7 Normal/2x 22.92 9 21.79

Dharmshala, Distt.

Kangra, H.P.

1,800 54199 7 (Fig. 1m) Abnormal/2x 23.10 9 19.76–25.46 9 22.02

Timbi, Distt.

Sirmaur, H.P.

1,800 54199 7 Normal/2x 22.79 9 21.68

Shillai, Distt.

Sirmaur, H.P.

2,000 54862 14 (Fig. 1n) Abnormal/4x 22.90 9 19.56–25.38 9 22.93

Nauradhar, Distt.

Sirmaur, H.P.

1,800 54861 14 Normal/2x 24.11 9 22.28

P. kleiniana Wight &

Arn.

Telangna, Distt.

Sirmaur,H.P.

2,800 55698 7 (Fig. 1o) Normal/2x 23.01 9 21.92

P. micropetala D. Don

(=Sibbaldiamicropetala (D. Don)

Hand.-Mazz.

Churdhar, Sirmaur

H.P.

3,600 54911 7 (Fig. 1p) Normal/2x 22.76 9 20.12

Churdhar, Distt.

Sirmaur, H.P.

3,600 54912 14 (Fig. 1q) Normal/4x 25.13 9 23.74

P. nepalensis Hook. Pehalgam, Distt.

Anantnag,

Kashmir

2,300 54901 21 (Fig. 1r) Abnormal/6x 22.96 9 19.45–24.88 9 21.93

Yusmarg, Distt.

Budgam, Kashmir

2,400 54902 14 Normal/2x 23.96 9 22.33

Triund, Distt.

Kangra, H.P.

2,300 54858 7 (Fig. 1s) Normal/2x 21.88 9 19.02

Lohardarhi, Distt.

Kangra, H.P.

2,500 54894 14 (Fig. 1t) Abnormal/4x 22.36 9 19.22–24.49 9 21.68

Haripurdhar, Distt.

Sirmaur, H.P

2,600 54864 7 Abnormal/2x 20.18 9 18.62–22.73 9 20.67

Shillai, Distt.

Sirmaur, H.P.

2,000 54865 14 Normal/4x 24.18 9 22.51

P. sibbaldia Kurtz.

(=Sibbaldia Hornem.)

Tisri, Distt.

Sirmaur, H.P.

3,000 54425 7 (Fig. 1u) Abnormal/2x 20.35 9 18.63–24.13 9 21.74

Churdhar, Distt.

Sirmaur, H.P.

3,600 55065 14 (Fig. 1v) Normal/4x 25.13 9 23.74

P. sundaica (Miq.)

W. Theob. (=Fragariasundaica Blume)

Karmul -Tral, Distt.

Pulwama,

Kashmir

2,300 54263 7 (Fig. 1w) Normal/2x 21.88 9 19.31

Triund, Distt.

Kangra, H.P.

3,000 52496 7 Normal/2x 22.08 9 19.41

Triund, Distt.

Kangra, H.P.

3,000 54883 7 Normal/2x 22.23 9 19.52

Dal lake, Distt.

Kangra, H.P.

2,800 52497 14 (Fig. 1x) Abnormal/4x 23.24 9 20.98–25.27 9 22.17

P. thomsonii Hand.-

Mazz.

Triund, Distt.

Kangra, H.P.

2,800 54886 7 (Fig. 1y) Normal/2x 22.18 9 19.10

Biching, Distt.

Kangra, H.P.

3,000 54887 7 Normal/2x 21.96 9 19.03

H.P. Himachal Pradesha In case of abnormal taxa the pollen size was calculated for both smaller and larger fertile pollen grains

488 S. Rani et al.

123

reported for the first time for India; it is in conformity with

the earlier report of 2n = 28 (Leht 1999) from Russia. The

diploid cytotypes (n = 7) are also known for the species

from Russia (Guinochet and Lefranc 1981; Leht 1991).

P. fulgens Wall. ex D. Don

Of the five populations of the species collected from

Himachal Pradesh, four exhibit diploid cytotypes (n = 7),

and only one population shows a tetraploid cytotype

(n = 14). The cytotype with n = 7 from India has been

cytologically worked out for the first time and is in con-

formity with the earlier report of 2n = 14 (Ikeda 1989)

from Japan. The tetraploid cytotype, n = 14, conforms to a

previous report from the Eastern Himalayas (Sharma and

Sarkar 1967–1968). The species is also known to have

another aneuploid cytotype with 2n = 34 from the Western

Himalayas (Mehra and Dhawan 1966) and octaploids

(2n = 56) from Russia (Popoff 1939).

P. gelida C. A. Mey.

Both populations of the species collected from Himachal

Pradesh depict n = 7 and exist as diploid cytotypes. The

Indian species has been chromosomally identified for the

first time, and n = 7 is in line with a previous report of

2n = 14 (Guinochet 1968) from France.

P. gerardiana Lindl.

The single accession of the species collected from Hima-

chal Pradesh exhibits n = 14. The tetraploid cytotype is

reported for the first time on a worldwide basis. Earlier the

species was reported to have diploids (2n = 14) from

different parts of the USSR and Japan (Krogulevich 1978;

Probatova N and Sokolovskaya 1981; Iwatsubo and Nar-

uhashi 1991) and octaploids (2n = 56) from the Western

Himalayas in India (Mehra and Dhawan 1966).

P. indica (Andrews) T. Wolf

Of the five populations of the species, one from Kashmir

and two from Himachal Pradesh exhibit diploid cytotypes

(n = 7). The other two populations from Himachal Pradesh

depict tetraploid cytotypes (n = 14). The present chro-

mosome count of n = 14 adds a new tetraploid cytotype

for the species on a worldwide basis, whereas n = 7 is in

conformity with the earlier reports from all over the world.

The species is already known to exhibit 2n = 14 (Subra-

manian 1979), 2n = 42 (Delay 1947), 2n = 70 (Kapil and

Modgil 1990) and 2n = 84 (Mehra and Dhawan 1966;

Naruhashi and Takano 1987; Hill 1989; Zhao et al. 1990;

Naruhashi and Iwatsuboo 1991; Naruhashi et al. 2005).

P. kleiniana Wight & Arn.

The single accession of the species collected from Hima-

chal Pradesh exhibits n = 7. The diploid cytotype is

reported for the first time from India and is in confirmity

with an earlier report of 2n = 14 (Shimotomai 1929) from

the USSR.

P. micropetala D. Don

Two populations of the species collected from Himachal

Pradesh show n = 7 and n = 14 cytotypes, respectively.

The present chromosome count of n = 14 adds a new

tetraploid cytotype for the species on a worldwide basis.

The diploid cytotype is reported for the first time from

India and is in conformity with the earlier single report of

2n = 14 (Mesicek and Sojak 1993) from China and Yun-

nan. The species is also known to have octaploids with

2n = 56 (Mehra and Dhawan 1966) from the northern

Western Himalayas in India.

P. nepalensis Hook.

Of the total six populations of the species, only one from

Kashmir shows n = 21 and exists as a hexaploid cytoytpe.

Among others, one population each from Kashmir and

Himachal Pradesh exhibit diploid cytotypes with n = 7.

The remaining two populations from Himachal Pradesh

reveal tetraploid cytotypes (n = 14). The present chro-

mosome count of n = 14 adds a new tetraploid cytotype to

the species on a worldwide basis. The diploid cytotypes

conform to a single previous chromosomal report of

2n = 14 (Popoff 1939) from Russia. The hexaploid cyto-

type is in conformity with the earlier reports of 2n = 42

(Mehra and Dhawan 1966) from the Western Himalayas in

India and outside India from Russia and England (Christoff

and Papasova 1943; Zhukova 1967; Goswami and Metfield

1975).

P. sibbaldia Kurtz.

The two populations of the species collected from Hima-

chal Pradesh show n = 7 and n = 14, respectively. The

present diploid cytotype is chromosomally reported for the

first time from India, whereas the tetraploid cytotype is

new on a worldwide basis. The species was previously

known to have a single diploid report of 2n = 14 (Guinochet

and Lefranc 1981) from Russia.

P. sundaica (Miq.) W. Theob.

Of the four populations of the species, one from Kashmir

and two from Himachal Pradesh show diploid cytotypes

Additions to the cytologically investigated species of Potentilla L. 489

123

(n = 7). Only one population from Himachal Pradesh

reveals a tetraploid cytotype (n = 14). The present chro-

mosome count of n = 7 is reported for the first time from

India and is in line with the previous reports of 2n = 14

(Shimotomai 1929) from different parts of Europe, whereas

2n = 28 is in conformity with the earlier reports from the

Western Himalayas in India (Mehra and Dhawan 1966) and

outside India (Ikeda 1989).

P. thomsonii Hand.-Mazz.

Both populations of the species collected from Himachal

Pradesh show n = 7 and exist as diploids based on x = 7.

The species has been cytologically worked out for the first

time on a worldwide basis.

Chromosome numbers and polyploidy

A perusal of the cumulative cytological literature brings to

light that 168 species/292 cytotypes of the genus Potentilla,

including 17 species (after the present study) from India,

have been worked out. The chromosome numbers in the

genus vary considerably from 2n = 14 to 112, with

2n = 28 (22.26%) being the most common followed by

2n = 14 (20.20%), 2n = 42 (15.06%), 2n = 56 (9.93%),

2n = 35 (5.47%), 2n = 49 (2.73%), 2n = 70 (2.73%) and

2n = 21 (1.71%), while 2n = 20, 34, 38, 39, 48, 62, 74,

77, 84, 91, 98 and 112 are rarely encountered in certain

species on a worldwide basis (c.f., Fedorov 1974; Kumar

and Subramaniam 1986; Index to Plant Chromosome

Numbers, Web, etc.). However, from India among the

different chromosome counts of 2n = 28, 34, 42 and 56,

the cytotype with 2n = 28 remains the most common and

is in line with the world data. As seen at present and as

previously reported by Delgado et al. (2000), Tomasz and

Kolodziejek (2008) and Lundberg (2011), irrespective of a

wide range of ploidy levels, x = 7 remains the single base

number for the genus both for India and worldwide.

Otherwise also the ploidy based on x = 7 in the different

species of subfamily Rosoideae is relatively high, and the

same is exhibited in the genus Potentilla (Johri et al. 1992),

where it is associated with gametophytic apomixis (Dick-

inson et al. 2007). According to Asker and Jerling (1992),

and Nylehn et al. (2003), Potentilla is one such genus in

which apomixis, sexuality, polyploidy and taxonomic

complexity are all combined. In the presently investigated

taxa also the same polyploidy is seen in nearly half of the

taxa, with the occurrence of tetraploids being the highest

(Table 2). Earlier, Stebbins (1950), and Mesicek and Sojak

(1993) categorized two groups of species within the genus,

with one group having species with stable chromosome

numbers, i.e., a single chromosomal race all over their

distribution and the other group of species with complex

ploidy whose chromosome numbers vary even within the

same species. The same holds true for the Western

Himalayas and India as well (Table 2). Analysis of the

worldwide data reveals that 123 species form the pre-

dominant group belonging to the stable chromosome

number category, and only 45 species are in the complex

ploidy level category, this too with some aneuploid cyto-

types as well. However, for Indian species, on the contrary,

a single chromosomal race either at 2x, 4x or 8x occurs less

frequently than those showing complex polyploidy levels

(Table 2). Comparing the present data with that worldwide,

it can be concluded that higher ploidy levels occur rela-

tively less often in Indian species.

In the presently studied species of Potentilla, the role of

hybridization followed by doubling of the chromosome

numbers resulting in allopolyploidy seems to be a major

mode of speciation in the genus Potentilla, as has been

described previously in many angiosperms (Stebbins 1950;

Grant 1981; Masterson 1994; Solits and Solits 2009).

Table 2 Ploidy levels in presently and previously cytologically worked out species of Potentilla L. from India

Species with single ploidy level Species with complex ploidy levels

Ploidy 2x 4x 8x 2x, 4x 2x, 4x, 5x 2x, 4x, 6x 4x, 8x 2x, 4x, 8x

Ch. no. (n) 2n = 14 2n = 28 2n = 56 2n = 14, 2n = 14, 2n = 28, 2n = 14, 2n = 28, 2n = 14

2n = 18 2n = 34 2n = 28, 2n = 56 2n = 28,

2n = 42 2n = 56

Species P. gelida P. fruticosa P. supina P. desertorum P. fulgens P. nepalensis P. argyrophylla P. atrosanguinea

P. kleiniana P. leschenaultiana P. indica P. gerardiana

P. thomsonii P. mooniana P. micropetala

P. sibbaldia

P. sundaica

Frequency 17.64 17.64 5.88 29.41 5.88 5.88 11.76 5.88

(%) 41.16 58.81

490 S. Rani et al.

123

According to recent molecular studies, the role of apomixis

has been considered to be an important phenomenon for the

evolutionary diversification of apomictic plants (Paule

et al. 2011). Based on the reproductive aspect of P. ar-

gentea, with different cytotypes of the species distributed

in Europe, it has been clearly indicated that a correlation

exists in the form in which sexuality and autogamy are

means of multiplication for diploids, whereas apomixis is

generally associated with polyploids (Holm 1995; Holm and

Ghatnekar 1996a, b; Holm et al. 1997; Paule et al. 2011).

The Indian materials have polyploids at various levels

of ploidy in the form of both inter- and intra-specific

Fig. 1 a Potentillaargyrophylla (n = 14), PMC at

Diakinesis. b P. argyrophylla(n = 28), PMC at Diakinesis.

c P. atrosanguinea (n = 7),

PMC at M-I. d P. atrosanguinea(n = 14), PMC at Diakinesis.

e P. atrosanguinea (n = 28),

PMC at Diakinesis. f P.desertorum (n = 7), PMC at

M-I. g P. desertorum (n = 14),

PMC at A-I. h P. fruticosa(n = 14), PMC at M-I. i P.fulgens (n = 7), PMC at M-I.

j P. fulgens (n = 14), PMC at

Diakinesis. k P. gelida (n = 7),

PMC at M-I. l P. gerardiana(n = 14), PMC at Diakinesis.

m P.indica (n = 7), PMC at

M-I. n P.indica (n = 14), PMC

at M-I. o P. kleiniana (n = 7),

PMC at M-I. p P. micropetala(n = 7) PMC at A-I.

q P.micropetala (n = 14) PMC

at A-I. r P. nepalensis (n = 21),

PMC at Diakinesis.

s P.nepalensis (n = 7) PMC at

A-I. t P.nepalensis (n = 14)

PMC at Diakinesis. u P.sibbaldia (n = 7) PMC at M-I.

v P. sibbaldia (n = 14) PMC at

A-I. w P.sundaica (n = 7) PMC

at M-I. x P. sundaica (n = 14)

PMC at Diakinesis. y P.thomsonii (n = 7) PMC at A-I.

Scale 10 lm

Additions to the cytologically investigated species of Potentilla L. 491

123

variability. During the present study, intercytotypes for

different species were found to be sympatric in distribution

and seem to be the result of both systems, i.e., sexuality and

apomictic mode operative selectively in different cyto-

types. This is in line with previous postulations that such

mixing up of the phenomena may be due to the repro-

ductive mode on the chosen donor pollen (Muntzing and

Muntzing 1945), methodological pitfalls of the applied

crossing methods including unintended self-pollination

(Schwendener 1969), mentor effects (Horandl and Temsch

2009) or dominance of maternal markers (Nosrati et al.

2010). From the present study, it is observed that although

the number of species worked out for the chromosome

numbers is very low from the Western Himalayas, the

amount of inter- and intraspecific chromosomal number

diversity is quite high, with euploid series marked in some

of the species from the region, and a few species studied

from other parts of India support the earlier hypothesis of

Panigrahi and Dikshit (1987) of India as the principal

center of origin of Potentilla.

Further, there appears to be a significant size difference

in meiotic chromosomes in different cytotypes in some

species such as P. argyrophylla (n = 28), P. atrosanguinea

(n = 28), P. desertorum (n = 7), P. fulgens (n = 7) and

Fig. 2 a–d Transfer of

chromatin (arrows) in pollen

mother cells (PMCs) at different

stages of meiosis. e PMC

showing chromatin stickiness at

M-I. f PMC showing unoriented

bivalents (arrows) at M-I.

g PMC showing bridges

(arrows) at A-I. h PMC

showing laggard (arrows) at

A-I. i PMC showing laggard

(arrows) at T-I. j Diad with

micronuclei. k Diad showing

incomplete division (arrows).

l Triad showing incomplete

division (arrows). m Triad.

n Tetrad with micronuclei

(arrows). o Polyad.

p Heterogenous sized fertile

pollen grains. q Fertile and

sterile pollen grains. Scale10 lm

492 S. Rani et al.

123

P.indica (n = 7), and also at an interspecific level in P.

gelida, whereas for the rest of the intraspecific cytotypes

and at an interspecific level, the chromosomes appear to

be of similar size. Such observations have been seen

earlier in certain species of Potentilla (Delgado et al.

2000).

Table 3 Data on abnormal meiotic course and pollen fertility in different species of Potentilla L. from the Western Himalayas

Taxa/

accessions

Cytomixis Meiotic course showing PMCs with Pollen

fertility

(%)Percentage of PMCs

involved meiosis-I/

meiosis-II

Number of

PMCs

involved

Chromosomal

stickness at M-I

(%)

Unoriented

bivalents at M-I

(%)

Bridges at

meiosis-I/

meiosis-II (%)

Laggards at

meiosis-I/

meiosis-II (%)

P. argyrophylla

54860 23.33 (28/120)/15.38

(20/130)

2–5 8.33 (10/120) 7.69 (10/130) 13.33 (18/135)/

7.69 (10/130)

7.27 (8/110)/5.21

(6/115)

59.79

54896 19.25 (26/135)/15.38

(20/130)

2–3 5.21 (6/115) 5.51 (7/127) 10.71 (15/140)/

6.15 (8/130)

4.44 (6/135)/5.08

(6/118)

60.62

54878 4.95 (6/121)/– 2–3 3.63 (4/110) 4.23 (5/118) 5.69 (7/123)/6.34

(8/126)

7.25 (9/124)/7.87

(10/127)

60.74

P. atrosanguinea

54880 3.63 (4/110)/– 2–3 – – 2.00 (2/100)/2.72

(3/110)

5.71 (6/105)/3.88

(4/103)

55.45

548907 11.66 (14/120)/5.21 (6/

115)

2–4 2.06 (2/97) – 6.95 (8/115)/– 6.03 (7/116)/– 52.47

54868 9.23 (12/130)/5.18 (7/

135)

2–3 3.47 (4/115) 3.44 (4/116) 7.50 (9/120)/6.40

(8/125)

4.23 (5/118)/– 63.90

P. desertorum

54895 11.66 (14/120)/5.21 (6/

115)

2–4 2.06 (2/97) – 1.81 (2/110)/

2.91(3/103)

5.71 (6/105)/3.88

(4/103)

59.98

P. fruticosa

52668 6.49 (10/154)/6.40 (8/

125)

2–3 – 3.34 (4/120) 3.34 (4/120)/– –/5.60 (7/125) 64.87

P. fulgens

54884 –/– – – –- –/– 3.47 (4/115)/

1.81(2/110)

65.00

54873 –/– – – 1.98 (2/101) 2.85 (3/105)/5.83

(7/120)

3.20 (4/125)/2.30

(3/130)

63.48

P. gelida

54228 11.53 (15/130)/– 2–4 – – –/3.70 (5/135) 2.30 (3/130)/4.13

(6/145)

60.70

P. indica

54199 3.20 (4/125)/– 2–3 1.53 (2/130) 0.90 (1/110) –/– –/– 61.62

54862 4.61 (6/130)/3.57 (4/

112)

2–3 – 2.50 (3/120) 3.17 (4/126)/– 3.47 (4/115)/1.66

(2/120)

62.84

P. nepalensis

54901 –/4.80 (6/125) 2–3 4.28 (6/140) – 8.69 (10/115)/

2.40 (3/125)

2.54 (3/118)/– 63.05

54894 –/– 2–3 3.70 (5/135) 4.34 (6/138) 6.15 (8/130)/3.07

(4/130)

–/– 62.45

54864 5.92 (8/135)/4.28 (6/

140)

2–3 – – 5.18 (7/135)/3.67

(5/136)

5.00 (6/120)/3.20

(4/125)

64.07

P. sibbaldia

54425 –/– – – – 5.45(6/110)/

3.57(4/112)

5.08 (6/118)/3.44

(4/116)

61.60

P. sundaica

52497 16.00 (20/125)/8.69 (10/

115)

2–4 3.63 (4/110) 4.95 (5/101) 6.25 (7/112)/3.63

(4/110)

3.36 (4/119)/– 61.58

Figures in parentheses denote the observed number of abnormal PMCs in the numerator and total number of PMCs observed in the denominator

Additions to the cytologically investigated species of Potentilla L. 493

123

Pollen size

The pollen grain size of the presently investigated taxa as given

in Table 1 shows that the polyploids have in general larger

pollen grains than diploids. Furthermore, among polyploids

there is relatively less difference between tetraploid and hexa-

ploid cytotypes, whereas there is relatively more size difference

when tetraploids are compared to octaploids. Such comparable

results have been previously reported for many angiosperms

(Stebbins 1956; Dijkstra and Speckmann 1980; Strivastava and

Strivastava 2002; Zlesak 2009; Omidbaigi et al. 2010).

Meiotic abnormalities

Meiotic abnormalities have been recorded in some popu-

lations of Potentilla argyrophylla, P. atrosanguinea,

P. desertorum, P. fruticosa, P. fulgens, P. gelida, P. indica,

P. nepalensis, P. sibbaldia and P. sundaica. In such pop-

ulations, abnormalities in the form of cytomixis, chromatin

stickiness, unoriented bivalents, bridges and laggards have

been observed at different stages of meiosis (Fig. 2a–i;

Table 3). These meiotic anomalies indicate the existence of

intraspecific genetic diversities. Such genetic differences

have been seen previously in different plant species

(Baptista-Giacomelli et al. 2000; Sheidai et al. 2003). The

phenomenon of chromatin transfer from early prophase to

the pollen formation stage (Fig. 2a–d) has been observed in

most of these populations with the highest percentage

recorded in populations of P. argyrophylla (Table 3).

Cytomixis and chromatin stickiness are considered to be

the result of genetic factors (Bellucci et al. 2003; Ghaffari

2006; Fadaei et al. 2010) and environmental factors

Table 4 Data on abnormal microsporogenesis in different species of Potentilla L. from the Western Himalayas

Taxa/accessions Microsporogenesis

Monads Diads Triads Tetrads

WMN (%) WM (%) WMN (%) WM (%) WMN (%) WM (%) WMN (%) WM (%)

P. argyrophylla

54860 2.19 (2/91) 1.09 (1/91) 3.29 (3/91) – 6.59 (6/91) 4.39 (4/91) 60.43 (55/91) 21.97 (20/91)

54896 2.91 (3/103) 1.94 (2/103) 1.94 (2/103) 0.97 (1/103) 4.85 (5/103) 3.88 (4/103) 63.10 (65/103) 20.38 (21/103)

54878 1.90 (2/105) 2.85 (3/105) 1.90 (2/105) 1.90 (2/105) 2.85 (3/105) 0.95 (1/105) 68.57 (72/105) 19.04 (20/105)

P. atrosanguinea

54880 1.92 (2/104) – 1.92 (2/104) – 3.84 (4/104) 1.92 (2/104) 65.38 (68/104) 25.00 (26/104)

54907 0.90 (1/110) 0.90 (1/110) 0.90 (1/110) 0.90 (1/110) 2.72 (3/110) 3.63 (4/110) 65.45 (72/110) 24.54 (27/110)

54868 – – 0.91 (1/109) – 1.83 (2/109) 0.91 (1/109) 71.55 (78/109) 24.77 (27/109)

P. desertorum

54895 – – – – 0.98 (1/102) 0.98 (1/102) 78.43 (80/102) 19.60 (20/102)

P. fruticosa

52668 1.83 (2/110) 0.91 (1/109) 1.83 (2/109) – 5.50 (6/109) 6.42 (7/109) 64.22 (70/109) 19.26 (21/109)

P. fulgens

54884 1.94 (2/103) 0.97 (1/103) 1.94 (2/103) – 1.94 (2/103 – 73.78 (76/103) 19.41 (20/103)

54873 0.97 (1/103) – 0.97 (1/103) – 1.94 (2/103) – 73.78 (76/103) 19.41 (20/103)

P. gelida

54228 – 1.00 (1/100) – 1.00 (1/100) 1.00 (1/100) 1.00 (1/100) 80.00 (80/100) 16.00 (16/100)

P. indica

54199 0.98 (1/102) 0.98 (1/102) – 0.98 (1/102) 1.96 (2/102) 1.96 (1/102) 76.47 (78/102) 17.64 (18/102)

54862 0.90 (1/111) 0.90 (1/111) 0.90 (1/111) 0.90 (1/111) 3.60 (4/111) 1.80 (2/111) 72.97 (81/111) 18.91 (21/111)

P. nepalensis

54901 – 0.93 (1/107) 0.93 (1/107) – 2.80 (3/107) 0.98 (1/107) 73.83 (79/107) 20.56 (22/107)

54894 0.88 (1/113) – 0.88 (1/113) 0.88 (1/113) – 3.53 (4/113) 73.45 (83/113) 21.23 (24/113)

54864 – – 0.86 (1/115) – 5/115 (4.34) 1.73 (2/115) 73.04 (84/115) 20.00 (23/115)

P. sibbaldia

54425 0.88 (1/113) – 1.76 (2/113) – 3.53 (4/113) 1.76 (2/113) 74.33 (84/113) 17.69 (20/113)

P. sundaica

52497 0.97 (1/103) – 1.76 (2/103) – 1.94 (1/103) 0.97 (1/103) 75.72 (78/103) 20.38 (21/103)

Figures in parentheses denote the observed number of abnormal PMCs in the numerator and total number of PMCs observed in the denominator

WMN without micronuclei; WM with micronuclei

494 S. Rani et al.

123

(Nirmala and Rao 1996) as well as genomic-environmental

interaction (Baptista-Giacomelli et al. 2000), and seems to

be equally applicable to the presently investigated popu-

lations. Cytomixis in these populations leads to the for-

mation of hypo- and hyperploids PMCs (Fig. 2a) and even

empty PMCs in some cases (Fig. 2b). These meiotic

abnormalities lead to the induction of abnormal polarity of

spindles and formation of nuclei of variable numbers and

sizes during microsporogenesis in the form of monads,

diads, triads and polyads along with micronuclei (Fig. 2j–

o; Table 4). In some of the PMCs, diads/triads show

incomplete division (Fig. 2k, l). As a result of these meiotic

abnormalities, large and small-sized pollen grains are

formed along with reduced pollen fertility (52–65%) in

these meiotically abnormal taxa (Fig. 2p–q; Table 3). The

large-sized pollen grains were seen in the range of 5–7% in

each such population. The occurrence of giant pollen

grains, possibly the unreduced 2n pollen grains, has pre-

viously been reported in several species (Vorsa and Bing-

ham1979; Bertagnolle and Thomson 1995; Sheidai et al.

2008; Fadaei et al. 2010). The unreduced gametes are

known to produce higher polyploidy levels through poly-

ploidization (Villeux 1985). According to Bertagnolle and

Thomson (1995) different cytological mechanisms are

responsible for production of 2n gametes. However, the

pollen fertility in the normal populations has been seen to

be quite high (95–99%).

Acknowledgments The authors are grateful to the University

Grants Commission, New Delhi, under the DRS SAP III of UGC,

FIST of the DST programs and the Rajiv Gandhi National Fellowship

Scheme for providing financial assistance. We are very thankful to the

Director of Botanical Surveys of India, DehraDun, for his help in

identification of plant species.

References

Asker S, Frost S (1970) The ‘‘Potentilla collina problem’’ a

chemotaxonomic approach. Hereditas 66:49–70

Asker SE, Jerling L (1992) Apomixis in plants. CRC Press, Boca

Raton

Ball PW, Pawlowski B, Walters SM (1968) Potentilla L. In: Tutin TG

et al (eds) Flora Europea, 2nd edn. Cambridge University Press,

Cambridge, pp 36–47

Baptista-Giacomelli FR, Pagliarini MS, Almeida JL (2000) Meiotic

behavior in several Brazilian oat cultivars (Avena sativa L.).

Cytol 65:371–378

Bellucci M, Roscini C, Mariani A (2003) Cytomixis in pollen mother

cells of Medicago sativa L. J Hered 94:512–516

Bertagnolle F, Thomson JD (1995) Gametes with the somatic

chromosome number: mechanisms of their formation and role

in the evolution of autopolyploid plants. New Phytol 129:1–22

Chaudhary HJ, Wadhwa BM (1984) Flora of Himachal Pradesh

analysis, 1st edn. Botanical Survey of India, Howrah

Christoff M, Papasova G (1943) Die genetischen Grundlagen der

apomicktischen Fortpflanzung in der Gattung Potentilla. Z. Induct.

Abstamm. Vererb-lehre., 8I, I

Delay C (1947) Recherches Sur la structure des noyaux quiescent

chez les Phanerogames. Rev Cytol Cytophysiol Veg 9:169–222;

10:103–229

Delgado L, Gallego F, Rico E (2000) Karyosystematic study of

Potentilla L. subgen. Potentilla (Rosaceae) in the Iberian

Peninsula. Bot J Linn Soc 132:263–280

Dhaliwal DS, Sharma M (1999) Flora of Kullu District (Himachal

Pradesh). Bishen Singh Mahendra Pal Singh, DehraDun

Dickinson TA, Lo E, Talent N (2007) Ploidy, reproductive biology,

and Rosaceae: understanding evolution and making classifica-

tions. Plant Syst Evol 266:59–78

Dijkstra H, Speckmann GI (1980) Autotetrapliody in Caraway

(Carum carvi L.) for the increase of the aetheric oil content of

the seed. Euphytica 29:89–96

Dobes C, Paule J (2010) A comprehensive chloroplast DNA-based

phylogeny of the genus Potentilla (Rosaceae): implications for

its geographic origin, phylogeography and generic circumscrip-

tion. Mol Phylogenet Evol 56:156–175

Eriksson T, Donoghue MJ, Hibbs MS (1998) Phylogenetic analysis of

Potentilla using DNA sequences of nuclear ribosomal internal

transcribed spacers (ITS), and implications for the classification

of Rosoideae (Rosaceae). Plant Syst Evol 211:155–179

Fadaei F, Sheidai M, Asadi M (2010) Cytological study of the genus

Arenaria L. (Caryophyllaceae). Caryol 63(2):149–156

Fedorov ANA (1974) Chromosome number of flowering plants.

Academy of Science of the USSR Komarov Botancial Institute,

Leningard

Gentscheff G (1938) Uber die pseudogame fortpflanzug bei Poten-tilla. Genetica 20:398–408

Ghaffari SM (2006) Occurrence of diploid and polyploid microspores

in Sorghum bicolor (Poaceae) is the result of cytomixis. Afr J

Biotech 5:1450–1453

Goswami DA, Metfield B (1975) Cytogenetic studies in genus

Potentilla L. New Phytol 75:135–146

Goswami DA, Metfield B (1978) Meiotic studies on two Polyploid

Potentillas, P. ambigens Greene and P. atrosanguinea Lodd.

Caryol 31:109–115

Grant V (1981) Plant Speciation 2nd ed. Columbia University Press,

New York, pp 293–297, 324–328

Gregor T, Rolik J, Weising K (2002) RAPD-Untersuchungen und

Chromosomenzahlungen in der Potentilla-collina-Gruppe (Ros-

aceae). Berichte der Bayerischen Gesellschaft zur Er-forschung

der Heimischen Flora 72:159–167

Guinochet M (1968) Nouvelle contribution al’etude cytotaxinomique

des Potentilla. De la section Aureae. Th Wolf CR. Acad Sci

(Paris) 239:1416–1417

Guinochet M, Lefranc M (1981) In Chromosome number reports

LXXIII. Taxon 30:853–854

Gustafsson A (1947a) Apomixis in higher plants. Part III. Biotype andspecies formation. Lunds Univ Arsskr 243:181–320

Gustafsson A (1947) Apomixis in higher plants. Part-II the causal

aspect of apomixis. Acta Univ Lund (Lunds Univ Arsskr) Avd 2,

43(2):71–178

Hill LM (1989) IOPB chromosome data 1. International organization

of plant biosystematists. Newsletter 13:17–19

Holm S (1995) Unexpectedly high levels of genetic variation in

Potentilla argentea (s.1.). in southern Sweden. Hereditas

123:127–139

Holm S, Ghatnekar L (1996a) Apomixis and sexuality in hexaploid

Potentilla argentea. Hereditas 125:53–60

Holm S, Ghatnekar L (1996b) Sexuality and no apomixis found in

crossing experiments with diploid Potentilla argentea. Hereditas

125:77–82

Holm S, Ghatnekar L, Bengtsson BO (1997) Selfing and outcrossing

but no apomixis in two natural populations of diploid Potentillaargentea. J Evol Biol 10:343–352

Additions to the cytologically investigated species of Potentilla L. 495

123

Hooker JD (1879) Flora of British India, vol II. L. Reeve and Co Ltd.,

Ashford

Horandl E, Temsch EM (2009) Introgression of apomixis into sexual

species is inhibited by mentor effects and ploidy barriers in the

Ranunculus auricomus complex. Ann Bot 104:81–89

Ikeda H (1989) Chromosome numbers of Himalayan Potentilla(Rosaceae). J Jap Bot 64:361–367

Iwatsubo Y, Naruhashi N (1991) Karyomorphological and cytoge-

netical studies in Potentilla (Rosaceae) I. Karyotypes of nine

Japanese species. Cytologia 56:1–10

Johri BM, Ambegaokar KB, Srivastava PS (1992) Comparative

embryology of Angiosperms. Springer-Verlag, Berlin

Kapil M, Modgil D (1990) A new report of pentaploid Fragariaindica Andr. from western Himalaya. Cell Chromosom Res

13:24–26

Kashin AS (1999) The structure of agamocomplexes and the problem

of saltatory speciation in angiosperms. Bot Zurn 84:15–29

Kaur H, Sharma M (2004) Flora of Sirmaur. Bishen Singh Mahendra

Pal Singh, Dehra Dun

Krogulevich RE (1978) Kariologicheskij analiz vidov flory Vostoch-

nogo Sajana. V Flora Pribajkal’ja. Nauka, Novosibirsk, pp 19–48

Kumar V, Subramaniam B (1986) Chromosome atlas of flowering

plants of the Indian subcontinent, vol 1 edn. Dicotyledones BSI,

Calcutta

Leht M (1991) Micromorphological characters in the systematic of

Baltic cinquefoils (Potentilla). Botaanilised uurinused. Scripta

Bot Belg 6:150–164

Leht M (1999) Origin, chromosome number and reproduction biology

of Potentilla fruticosa (Rosaceae) in Estonia and Latvia. Ann

Bot Fenn 162:191–196

Lundberg M (2011). Systematics and polyploid evolution in Poten-tilleae (Rosaceae). University dissertation from Stockholm:

Department of Botany, Stockholm University, Sweden

Masterson J (1994) Stomatal size in fossil plants: evidence for

polyploidy in majority of angiosperms. Science 264:421–423

Mehra PN, Dhawan H (1966) Cytological investigation in the N.

W. Himalayan Rosaceae. Proc 53rd Indian Sci Congr 3:276–277

Mesicek J, Sojak J (1969) Chromosome counts of some Mongolian

plants. Folia Gebot Phytotaxon 4:55–86

Mesicek J, Sojak J (1993) Annotated chromosome numbers of

selected Asiatic Potentilla species. Folia Geobot Phytotaxon

28(4):437–446

Muntzing A (1958) Heteroploidy and polymor-phism in some

apomictic species of Potentilla. Hereditas 44:280–329

Muntzing A, Muntzing G (1941) Some new results concerning

apomixis, sexuality and polymorphism in Potentilla. Bot Notiser

94:237–278

Muntzing A, Muntzing G (1945) The mode of reproduction of hybrids

between sexual and apomictic Potentilla argentea. Bot Notiser

98:49–71

Naruhashi N, Iwatsuboo Y (1991) Karyological studies of Duchesnea

(Rosaceae). Cytol 56(3):143–149

Naruhashi N, Takano H (1987) Chromosome numbers and distribu-

tion of Duchesnea (Rosaceae) in Gifu prefecture. Acta Phyto

Geobot 38(8):155–160

Naruhashi N, Nishikawa T, Iwatsubo Y (2005) Taxonomic relation-

ship between Japanese Potentilla anemonefolia and Himalayan

P. sundaica (Rosaceae). J Phytogeogr Taxon 53:1–11

Nirmala A, Rao PN (1996) Genetics of chromosome numerical

mosaism in higher plants. Nucleus 39:151–175

Nosrati H, Price A, Wilcock CC (2010) No evidence of apomixis in

matroclinal progeny from experimental crosses in the genus

Fragaria (strawberry) based on RAPDs. Euphytica 171:193–202

Nylehn J, Hamre E, Nordal I (2003) Facultative apomixis and

hybridization in arctic Potentilla section Niveae (Rosaceae) from

Svalbard. Bot J Linn Soc 142:373–381

Omidbaigi R, Mirzaee M, Hassani ME, Sedghi Moghadam M (2010)

Induction and identification of polyploidy in basil (Ocimumbasilicum L.) medicinal plant by colchicine treatment. Int J Plant

Prod 4(2):87–98

Panigrahi G, Dikshit BK (1987) Systematics of the genus Potentilla L.

(Rosaceae Juss.)—its infrageneric classification and evolutionary

trends. Bull Bot Surv India 27:177–196

Paule J, Sharbal TF, Dobes C (2011) Apomoctic and sexual lineages

of the Potentilla argentea L. group (Rosaceae): cytotype and

molecular genetic differentiation. Taxon 60(3):721–732

Popoff A (1935) Uber die Fortpflanzungsverhaltnisse der Gattung

Potentilla. Planta Berl 24:510

Popoff A (1939) Karyologische Studien iiber die Entwicklungsvorg-

ange in der Gattung Potentilla 9(3):3–58

Probatova NS, Sokolovskaya AP (1981) Kariologicheskoe

issledovanie sosudistykh rastenij ostrovov Dal’nevostocnogo

gosudarstvennogo morskogo sapovednika. Sb. Cvetkaye Ras-

tenija Ostrovov Dalnevostochnogo Morskogo Sapovednika

pp 92–114

Rutishauser A (1948) Pseudogamie und Polymorphie in der Gattung

Potentilla. Arch Julius Klaus-Stiftung Vererbungsf 23(3/4):

267–424

Santapau H, Henry AN (1973) A dictionary of the flowering plants in

India. C.S.I.R, New Delhi

Schwendener J (1969) Experimente zur Evolution von Potentillaprocumbens Sibth. Ber Schweiz Bot Ges 79:49–92

Sharma A, Sarkar AK (eds) (1967–1968) Chromosome number

reports of plants. In: Annual Report, Cytogenet Lab Dept Bot

Univ Calcutta Res Bull 2:38–48

Sheidai M, Koobaz P, Zehzad B (2003) Meiotic studies of some

Avena L. species and populations. Iran J Sci 14:121–131

Sheidai M, Nikoo M, Gholipour A (2008) Cytogenetic variability and

new chromosome number reports in Silene L. species (Sect.

Lasiostemones, Caryophyllaceae). Acta Biol Szedg 52:313–319

Shimotomai N (1929) Uber die Chromosomenzhalen bei einigen

Potentillen. Sci Rep Tohokn Univ Ser 4(4):369–372

Shimotomai N (1930) Chromosomenzahlen u Phylogene bei der

Gattung Potentilla. Y Sci Hirosh Univ, Botany, I, 1

Solits PS, Solits DE (2009) The role of hybridization in plant

speciation. Annu Rev Plant Biol 60:561–588

Stebbins GL (1950) Variation and evolution in plants. New York

Columbia University Press, New York

Stebbins GL (1956) Cytogenetics and evolution of the grass family.

Am J Bot 43:890–905

Strivastava R, Strivastava GK (2002) Autopolyploids of Helianthusannuus L. Var. morden. Cytol 67:213–220

Subramanian D (1979) Cytological studies in South Indian species of

Rosaceae. Soc Adv Bot Annu Conf, Dep Bot Punjab Agric,

Univ, Ludhiana. Abstr 123–124

Szafer W, Pawlowski B (1955) Potentilla L. In: Szafer W, Pawlowski

B (eds) Flora of Poland vol 7. Panstwowe Wydawnictwo

Naukowe, Warszawa, pp 96–43

Tomasz I, Kolodziejek J (2008) Chromosome numbers of Potentillasubsect. Collinae (Rosaceae) from Poland. Caryologia 61(2):

170–175

Villeux R (1985) Diploid and polyploidy gametes in Crop Plants:

mechanisms of formation and utilization in plant breeding. In:

Janick J (ed) Plant Breed Rev 3. AVI, New York, p 442

Vorsa N, Bingham E (1979) Cytology of 2n pollen formation

in diploid alfalfa, Medicago sativa. Can J Genet Cytol 21:525–

530

Web: Missouri Botanical Garden Tropics, Nomenclatural database

http://mobot.mobot.org/cgi-bin/searchvast, accessed on 28

August 2011

Wolf T (1908) Monographie der Gattung Potentilla. Biblliotbeca Bot

71:1–715

496 S. Rani et al.

123

Zhao ZF, Wang YQ, Huang S (1990) Plant chromosome counts (V).

For Res 3:503–508

Zhukova PG (1967) Karyology of some plants cultivated in the

Arctic-Alpine Botanical Garden. In: Avrorin NA (ed) Plantarum

in Zonam Polarem Transportatio II. Leningrad pp 139–149

Zlesak DC (2009) Pollen diameter and guard cell length as predictor

of ploidy in diverse rose cultivars, species, and breeding lines.

In: Zlesak DC (ed) Roses. Floricult ornament biotechnol, vol 3

(special issue 1). Global science books Ltd., UK, pp 53–70

Additions to the cytologically investigated species of Potentilla L. 497

123