CARYOLOGIA Vol. 61, no. 1: 92-106, 2008

INTRODUCTION

According to LEVICHEV (1999), the genus Ga-gea Salib. (Liliaceae; tribe Tulipeae, see PATTER-SON and GIVNISH 2002) comprises at least 250 spe-cies, distributed from the Mediterranean region throughout Europe and Asia; many of them being only recently described (LEVICHEV 2001; 2006; ZHAO and ZHAO 2003; 2004; TISON 2004; HENKER 2005; ZARREI and ZARRE 2005; ALI 2006; LEVICHEV and ALI 2006; ZHAO and YANG 2006; PERUZZI et al. 2007). Since several years, this taxonomically extremely difficult genus was the object of several studies on bulb structure (LEVICHEV 1999), mor-phology and ontogeny (HENKER 2005; LEVICHEV 2001; 2006b; TISON 2004; PERUZZI et al. 2007), classical cytotaxonomy (PERUZZI 2003; PERUZZI and AQUARO 2005), embryology (GREILHUBER et al. 2000; CAPARELLI et al. 2006), reproductive biology (GARGANO et al. 2007), pollen morphol-ogy (ZARREI and ZARRE 2005), nomenclature and

taxonomy (BAYER and LÓPEZ GONZÁLEZ 1989; LEVICHEV 1990; CUCCUINI and LUCCIOLI 1995; TISON 2001; 2004b; JOHN et al. 2004; LEVICHEV and TISON 2004; TISON and PERRET 2004; PERUZZI and TISON 2004; 2005; 2006; 2007; PERUZZI 2006; PERUZZI and ZARREI 2007; ZARREI et al. 2007; PE-RUZZI 2008), geographical distribution (TISON 2004c; PERUZZI and GARGANO 2005; PERUZZI and BARTOLUCCI 2006; BARTOLUCCI and PERUZZI 2008; FERRER GALLEGO et al. 2007; PERUZZI and CAPARELLI 2007; ZARREI et al. 2007) and phyloge-netic relationships utilizing cpDNA and nrDNA data (PETERSON et al. 2004; 2008).

The genus Gagea Salisb. consists of at least 13 (PETERSON et al. 2008) sections. Mediterranean species represent about 25% of the existing spe-cies, from all Gagea sections. Representatives (9 species, 15 accessions) of the diverse sec-tion Didymobulbos (K. Koch) Boiss. (except G. tisoniana, from sect. Gagea) are included in this study.

This paper – following two previous contribu-tions on Gagea Salisb. karyology (PERUZZI 2003; PERUZZI and AQUARO 2005) – reports new karyo-logical data from central Mediterranean area and also updates to 2007 the karyological knowledge of that difficult genus (cfr. Table 1).

Contribution to the cytotaxonomical knowledge of the genus Gagea Salisb. (Liliaceae). III. New karyological data from the central Mediterranean areaPeruzzi* Lorenzo

Dipartimento di Biologia, Unità di Botanica Generale e Botanica Sistematica, Università di Pisa, Via Luca Ghini 5, 56126 Pisa, Italy.

Abstract — New karyological data for the genus Gagea Salisb. (Liliaceae) are presented and discussed. First chro-mosome counts for the following taxa are provided: G. chaberti A. Terracc. (2n = 36) from Tunisia; G. sicula Lojac. (2n = 24, 36) and G. chrysantha Schult. et Schult. f. sensu stricto (2n = 36) from Sicily, including their respective loci classici. Chromosome complement of G. mauritanica Durieu (2n = 36) from S France and from a new local-ity from Apulia (S Italy) is reported. Detailed karyotype structure of the recently described C Italian endemic G. tisoniana Peruzzi, Bartolucci, Frignani et Minut. (2n = 24), G. dubia (2n = 48), G. foliosa (J. et C. Presl) Schult. et Schult. f. (2n = 36) and G. granatellii (Parl.) Parl. (2n = 36) are also reported, for the first time. G. bohemica (Za-uschn.) Schult. et Schult. f. resulted tetraploid (2n = 48) in plants from Abruzzo (C Italy) and from locus classicus of G. busambarensis (Tin.) Parl. (Rocca Busambra, NW Sicily), while a new 2n = 72 cytotype is recorded for the first time (Madonie, NW Sicily). Finally, G. villosa (M. Bieb.) Sweet from Abruzzo resulted tetraploid (2n = 48).

Key words: Gagea, karyology, Liliaceae, taxonomy.

* Corresponding author: phone +39 (0)50 2211339; fax +39 (0)50 2211309; e-mail: lperuzzi@biologia.unipi.it

gagea karyology 93

Table 1 — Update of karyological knowledge of the genus Gagea Salisb. Previous data can be found summarized in Peruzzi (2003: table 4).

2n provenance author(s)G. bohemica (Zauschn.) Schult. et Schult. f. 36 Turkey Özhatay 2002

48 C Italy Peruzzi and Bartolucci 2006C Italy present study

[locus classicus of G. busambarensis] Sicily present study72 Sicily present study

G. chaberti A. Terracc. 36 Tunisie present studyG. chrysantha Schult. et Schult. f. s.l. 36 Turkey Özhatay 2002

S Italy Peruzzi and Aquaro 2005SE Sicily Peruzzi and Aquaro 2005

[locus classicus] NW Sicily present studyG. dschungarica Regel 48 Kashmir Koul and Khan 1969 ex Mešicek and Hrouda 1974

60 Kashmir Koul and Khan 1969 ex Mešicek and Hrouda 1974G. dubia A. Terracc. 48 C Italy Peruzzi and Bartolucci 2006

Sicily present studyG. elegans Wall. 72 Kashmir Koul and Khan 1969 ex Mešicek and Hrouda 1974

96 Kashmir Koul and Khan 1969 ex Mešicek and Hrouda 1974132 Kashmir Koul and Khan 1969 ex Mešicek and Hrouda 1974

G. foliosa (J. et C. Presl) Schult. et Schult. f. 36 Sicily Peruzzi and Aquaro 2005Sicily present study

G. fragifera (Vill.) Ehr. Bayer et G. López 84 S Italy Peruzzi and Aquaro 2005Sicily Peruzzi and Aquaro 2005

G. graeca (L.) A. Terracc. 16 Greece Kapasa et al. 200124 Greece Peruzzi et al. unpubl.

G. granatellii (Parl.) Parl. 36 SE Sicily Peruzzi and Aquaro 2005NW Sicily present study

S Italy present study48 C Italy Peruzzi and Bartolucci 2006

G. joannis Grossh. 24 Georgia Davlianidze and Bokeria 2005G. luberonensis J.-M. Tison 36 C Italy Peruzzi and Bartolucci 2006G. lutea (L.) Ker.-Gawl.* 72 Austria Greilhuber et al. 2000

S Italy Peruzzi and Aquaro 2005N Germany Henker 2005

[var. glauca (Blocki) L. Klein] 72 N Germany Henker 2005 [cfr. var. brentae Evers] 72 N Germany Henker 2005

G. mauritanica Durieu 36 S Italy Peruzzi and Aquaro 2005S Italy present study

S France present studyG. megapolitana Henker 72 N Germany Henker 2005

84 N Germany Henker 2005G. minima (L.) Ker-Gawl. 24 S Italy Peruzzi and Aquaro 2005

N Germany Henker 2005G. peduncularis (J. et C. Presl) Pascher 36 S Italy Peruzzi and Caparelli 2007G. polidorii J.-M. Tison 72 S Italy Peruzzi and Aquaro 2005G. pomeranica Ruthe 60 N Germany Henker 2005

72 N Germany Henker 2005G. pratensis (Pers.) Dumort. 36 Turkey Özhatay 2002

60 S Italy Peruzzi and Aquaro 2005C Italy Peruzzi et al. 2007

N Germany Henker 2005Denmark Henker 2005

72 N Germany Henker 2005Sweden Henker 2005

G. pusilla (F. W. Schmidt) Sweet 24 NE Italy Peruzzi et al. 2007G. sicula Lojac. [locus classicus] 24 NW Sicily present study

36 SE Sicily present studyG. sulfurea Miscz. 72 Georgia Davlianidze and Bokeria 2005G. tisoniana Peruzzi et al. 24 C Italy Peruzzi et al. 2007

C Italy present studyG. trinervia (Viv.) Greuter 24 SE Sicily Peruzzi et al. unpubl.G. villosa (M. Bieb.) Sweet var. villosa 36 Turkey Özhatay 2002 48 Turkey Özhatay 2002

S Italy Peruzzi and Aquaro 2005N Germany Henker 2005

C Italy present study

* ERRATA CORRIGE - Dr. H. Henker kindly noticed me an error in my previous publication (Peruzzi 2003: 121): G. lutea is there wrongly reported with 2n = 48 chromosomes from Czech Republic, while actually the authors (Mešicek & Hrouda 1974) refer to it as having a 2n = 72 complement (usual for this species).

peruzzi 94

MATERIALS AND METHODS

Plant material. - Gagea bohemica (Zauschn.) Schult. et Schult. f.: Italy: Abruzzo, Conca di Cap-estrano, province of L’Aquila, 500 m, 18.III.2004, L. Bernardo (cult. Hort. Bot. Pisa University, acc. n. 473/2006; CLU n. 13217); Sicily: Rocca Busambra, province of Palermo, III.2004, J.-M. Tison (cult. Hort. Bot. Pisa University, acc. n. 475/2006); Sicily: Madonie, near the crossroad among Piano Zucchi, Piano Battaglia and Poliz-zi Generosa, eastern slope of Monte dei Cervi, ca. 1500 m, 3.IV.2006, L. Peruzzi, D. Gargano, G. Aquaro, K. F. Caparelli, C. Stefano (cult. Hort. Bot. Pisa University, acc. n. 474/2006; CLU n. 19770); Gagea chaberti A. Terracc.: Tunisia: Tabarka, Barkoucech beach, sandy open grass-lands near the shore, III.2004, J.-M. Tison (cult. Hort. Bot. Pisa University, acc. n. 490/2006); Gagea chrysantha Schult. et Schult. f. s.s.: Sicily: Ficuzza, at the northern foothills of Rocca Busam-bra, province of Palermo, III.2004, J.-M. Tison (cult. Hort. Bot. Pisa University, acc. n. 479/2006; CLU n. 9237); Gagea dubia A. Terracc.: Sicily: Madonie, near the crossroad among Piano Zuc-chi, Piano Battaglia and Polizzi Generosa, eastern slope of Monte dei Cervi, ca. 1500 m, 3.IV.2006, L. Peruzzi, D. Gargano, G. Aquaro, K. F. Caparelli, C. Stefano (cult. Hort. Bot. Pisa University, acc. n. 466/2006; CLU n. 19767, 19768); Gagea fo-liosa (J. et C. Presl) Schult. et Schult. f.: Sicily: Madonie, near the crossroad among Piano Zuc-chi, Piano Battaglia and Polizzi Generosa, eastern slope of Monte dei Cervi, ca. 1500 m, 24.IV.2004, L. Peruzzi et K. F. Caparelli (cult. Hort. Bot. Pi-sa University, acc. n. 489/2006; CLU n. 12689); Gagea granatellii (Parl.) Parl.: Italy: Apulia: Murge: E to S. Simone (province of Taranto), at the mar-gins of a Pinus brutia Ten. artificial wood, karstic grasslands, 250 m, 33 S XE 94.98, 5.III.2006, L. Peruzzi et K. F. Caparelli (CLU n. 18176); Sicily: Madonie, near the crossroad among Piano Zuc-chi, Piano Battaglia and Polizzi Generosa, eastern slope of Monte dei Cervi, ca. 1500 m, 3.IV.2006, L. Peruzzi, D. Gargano, G. Aquaro, K. F. Caparelli, C. Stefano (cult. Hort. Bot. Pisa University, acc. n. 495/2006; CLU n. 19769); Gagea mauritanica Durieu: France: Martigues, pointe de Bonnieu, karstic grasslands near the seashore, 18.II.2004, J.-M. Tison (cult. Hort. Bot. Pisa University, acc. n. 506/2006); Italy: Apulia, Murge: S. Simone (province of Taranto), karstic grasslands, 241 m, 33 S XE 92.98, 5.III.2006, L. Peruzzi et K. F. Ca-parelli (CLU n. 18173); Gagea sicula Lojac.: Sicily: Alpe di Cucco, Ficuzza, near Rocca Busambra,

province of Palermo, III.2004, J.-M. Tison (cult. Hort. Bot. Pisa University, acc. n. 483/2006; CLU n. 9258); Sicily: Iblei, Mount Lauro, III.2004, J.-M. Tison (cult. Hort. Bot. Pisa University, acc. n. 482/2006); Gagea tisoniana Peruzzi, Bartolucci, Frignani et Minut.: Italy: Tuscany, Colline Metal-lifere, Le Cornate di Gerfalco – Montieri (province of Grosseto), 900-1050 m a.s.l., calcareous rocky meadows exposed to S-SW, 01.IV.2005, L. Peruz-zi et F. Frignani (cult. Hort. Bot. Pisa University, acc. n. 501/2006; CLU n. 14922); Gagea villosa (M. Bieb.) Sweet: Italy, Abruzzo: Conca di Cap-estrano, province of L’Aquila, 500 m, 18.III.2004, L. Bernardo (cult. Hort. Bot. Pisa University, acc. n. 468/2006; CLU n. 13216).

Chromosome analysis. - Squash preparations were made from root-tips or young ovules of plants col-lected in situ, according to the following schedule: pretreatment in 0.5% colchicine solution for 4 hours; Carnoy fixing for at least 1 hour; hydrolisis in HCl 1N for 7 minutes at 60°C; staining with leuco-basic fuchsin for 3 hours. Karyotype formulas and terminology are according to LEVAN et al. (1964). At least ten plates were used in order to establish the chromosome numbers, while five plates were meas-ured in order to build the idiograms. Intrachromo-somal (A1) and interchromosomal (A2) asymmetry indexes were calculated according to the method-ology defined by ROMERO ZARCO (1986).

RESULTS

Gagea bohemica (Zauschn.) Schult. et Schult. f. - The chromosome complement of the studied population from Madonie (N Sicily) has revealed to be hexaploid, with 2n = 72 chromosomes (Fig. 1A). Karyotype formula can be expressed as fol-lows: 2n = 72 = 6x = 18st + 6msat + 6sm + 6m + 6msat + 18sm + 6msat + 6msat (Tables 2-3; Fig. 1B). Chromosome size ranges from 1.26 to 3.67 µm in average. The chromosome complement of both the studied populations from Abruzzo and Rocca Busambra (NW Sicily) has revealed to be tetra-ploid, with 2n = 48 chromosomes (Fig. 2A). In the former population, it was possible to establish the karyotype formula (Tables 2-3; Fig. 2B): 2n = 48 = 4x = 12st + 4m + 4sm + 28m. Chromosome size ranges from 1.08 to 2.96 µm in average.

Gagea chaberti A. Terracc. - The chromosome com-plement of the studied population from Tabarka (Tunisia) has revealed to be triploid, with 2n = 36

gagea karyology 95

chromosomes (Fig. 3A). Karyotype formula can be expressed as follows: 2n = 36 = 3x = 9st + 6sm + 3m + 9sm + 9m (Tables 2-3; Fig. 3B). Chromosome size ranges from 1.48 to 4.35 µm in average.

Gagea chrysantha Schult. et Schult. f. s.s. - The chromosome complement of the studied popula-tion from locus classicus of the species (Ficuzza, Sicily) has revealed to be triploid, with 2n = 36

Fig. 1 — Gagea bohemica (Zauschn.) Schult. et Schult. f. A. Metaphasic plate, showing 2n = 72 chromosomes (plants from Madonie); B. haploid idiogram. Scale bars = 5µm.

Fig. 2 — Gagea bohemica (Zauschn.) Schult. et Schult. f. A. Metaphasic plate, showing 2n = 48 chromosomes (plants from Abruzzo); B. haploid idiogram. Scale bars = 5µm.

peruzzi 96

chromosomes (Fig. 4). We were not able to build the idiogram for this species, because we observed only one plate showing the centromeres clearly.

Gagea dubia A. Terracc. - The chromosome com-plement of the studied population from Madonie (N Sicily) has revealed to be tetraploid, with 2n = 48 chromosomes (Fig. 5A). Karyotype formula can be expressed as follows: 2n = 48 = 4x = 12st + 12m + 4sm + 12m + 4sm + 4m (Tables 2-3; Fig. 5B). Chromosome size ranges from 1.56 to 4.93 µm in average.

Gagea foliosa (J. et C. Presl) Schult. et Schult. f. - The chromosome complement of the stud-ied population from Madonie (N Sicily) has re-vealed to be triploid, with 2n = 36 chromosomes (Fig. 6A). Karyotype formula can be expressed as follows: 2n = 36 = 3x = 9st + 3sm + 3st + 3m + 3sm + 3sm + 3msat + 3m + 3st + 3m (Tables 2-3; Fig. 6B). Chromosome size ranges from 1.29 to 4.32 µm in average.

Gagea granatellii (Parl.) Parl. - The chromosome complement of both the studied populations from Apulia and Sicily has revealed to be triploid with 2n = 36 chromosomes (Fig. 7A and 8A, re-spectively). In the former population, karyotype formula can be expressed as follows: 2n = 36 = 3x = 9st + 3sm + 3st + 3m + 3sm + 3m + 3sm + 3m + 3st + 3m (Tables 2-3; Fig. 7B). Chromosome size ranges from 1.56 to 4.49 µm. In the latter popula-tion, karyotype formula is 2n = 36 = 3x = 9st + 3sm + 6m + 3sm + 3m + 6sm + 3m + 3sm (Tables 2-3; Fig. 8B). Chromosome size ranges from 1.34 to 4.14 µm in average.

Gagea mauritanica Durieu - The chromosome complement of both the studied populations from France (Martigues) and Apulia has revealed to be triploid with 2n = 36 chromosomes (Fig. 9A and 10, respectively). In the former population, it was possible to establish the karyotype formula: 2n = 36 = 3x = 9t + 3sm + 6sm + 3m + 6sm + 3m + 3sm + 3m (Tables 2-3; Fig. 9B). Chromosome size ranges from 1.81 to 5.53 µm in average.

Fig. 3 — Gagea chaberti A. Terracc. A. Metaphasic plate, showing 2n = 36 chromosomes; B. haploid idiogram. Scale bars = 5µm.

Fig. 4 — Gagea chrysantha Schult. et Schult. f. s.s. Met-aphasic plate, showing 2n = 36 chromosomes. Scale bars = 5µm.

gagea karyology 97

Fig. 6 — Gagea foliosa (J. et C. Presl) Schult. et Schult. f. A. Metaphasic plate, showing 2n = 36 chromosomes; B. haploid idiogram. Scale bars = 5µm.

Fig. 5 — Gagea dubia A. Terracc. A. Metaphasic plate, showing 2n = 48 chromosomes; B. haploid idiogram. Scale bars = 5µm.

Gagea sicula Lojac. - The chromosome com-plement of the studied population from locus classicus of the species (Ficuzza, Sicily) has re-vealed to be diploid, with 2n = 24 chromosomes (Fig. 11A). Karyotype formula can be expressed as follows: 2n = 24 = 2x = 4st + 2t + 10sm + 8m (Tables 2-3; Fig. 11B). Chromosome size ranges

from 1.84 to 6.18 µm in average. The chromo-some complement of the studied population from Iblei (Sicily) has revealed to be triploid, with 2n = 36 chromosomes (Fig. 12). Because of the average quality of the plates, we were not able to build the idiogram for this cytotype.

peruzzi 98

Tabl

e 2

— M

easu

rem

ents

mad

e on

5 m

etap

hasi

c pl

ates

bel

ongi

ng to

stu

died

Gag

ea s

peci

es. D

ata

wer

e ob

tain

ed fr

om m

icro

phot

ogra

phs,

then

rep

orte

d in

µm

.

III

III

IVV

VI

VII

VII

IIX

XX

IX

IIT

HL

G.b

ohem

ica

6xS

0.62

± 0

.04

0.64

± 0

.18

0.52

± 0

.06

0.83

± 0

.19

0.58

± 0

.01

0.81

± 0

.15

0.71

± 0

.09

0.49

± 0

.02

0.47

± 0

.06

0.44

± 0

.12

0.59

± 0

.12

0.48

± 0

.07

24.2

2 ±

1.52

L3.

04 ±

0.4

62.

40 ±

0.3

22.

25 ±

0.3

61.

36 ±

0.2

61.

32 ±

0.0

41.

09 ±

0.1

60.

95 ±

0.0

21.

14 ±

0.0

11.

01 ±

0.1

00.

91 ±

0.1

30.

75 ±

0.1

30.

78 ±

0.0

7

G.b

ohem

ica

4xS

0.66

± 0

.12

0.53

± 0

.06

0.59

± 0

.09

0.96

± 0

.12

0.69

± 0

.09

0.77

± 0

.10

0.61

± 0

.04

0.70

± 0

.08

0.66

± 0

.05

0.63

± 0

.09

0.55

± 0

.07

0.46

± 0

.08

21.8

0 ±

1.58

L2.

29 ±

0.3

51.

92 ±

0.2

81.

76 ±

0.3

31.

37 ±

0.2

51.

17 ±

0.1

60.

94 ±

0.0

80.

98 ±

0.1

20.

84 ±

0.1

10.

71 ±

0.0

30.

72 ±

0.1

30.

65 ±

0.0

80.

62 ±

0.0

3

G. c

habe

rti 3

xS

0.74

± 0

.16

0.68

± 0

.09

0.62

± 0

.13

0.95

± 0

.09

0.79

± 0

.06

1.12

± 0

.10

0.77

± 0

.09

0.75

± 0

.04

0.73

± 0

.20

0.81

± 0

.11

0.67

± 0

.07

0.68

± 0

.06

31.7

8 ±

0.99

L3.

61 ±

0.1

43.

22 ±

0.2

12.

79 ±

0.2

42.

17 ±

0.1

92.

05 ±

0.1

31.

37 ±

0.1

21.

63 ±

0.1

11.

56 ±

0.0

61.

26 ±

0.0

41.

06 ±

0.2

20.

97 ±

0.0

70.

80 ±

0.0

4

G. d

ubia

4x

S0.

58 ±

0.0

70.

69 ±

0.2

30.

60 ±

0.0

71.

11 ±

0.1

50.

81 ±

0.1

50.

83 ±

0.2

10.

50 ±

0.0

80.

69 ±

0.1

30.

69 ±

0.1

80.

56 ±

0.2

50.

47 ±

0.2

50.

49 ±

0.2

025

.11

± 4.

31L

2.86

± 0

.60

2.52

± 0

.57

1.94

± 0

.49

1.49

± 0

.19

1.38

± 0

.24

1.21

± 0

.26

1.22

± 0

.14

1.01

± 0

.22

0.81

± 0

.21

0.92

± 0

.23

0.96

± 0

.23

0.77

± 0

.20

G. f

olio

sa 3

xS

0.78

± 0

.22

0.57

± 0

.04

0.72

± 0

.11

1.00

± 0

.20

1.16

± 0

.16

0.67

± 0

.14

0.84

± 0

.30

0.85

± 0

.06

0.66

± 0

.15

0.80

± 0

.15

0.53

± 0

.16

0.67

± 0

.14

30.1

4 ±

1.44

L3.

37 ±

0.1

73.

19 ±

0.4

02.

34 ±

0.2

92.

05 ±

0.0

21.

43 ±

0.2

51.

69 ±

0.1

21.

46 ±

0.0

21.

22 ±

0.0

21.

25 ±

0.0

81.

04 ±

0.0

81.

01 ±

0.1

70.

86 ±

0.1

0

G. g

rana

telli

i3x

(Pug

)

S0.

74 ±

0.1

00.

85 ±

0.1

20.

58 ±

0.0

41.

06 ±

0.0

90.

67 ±

0.1

31.

09 ±

0.0

20.

58 ±

0.0

60.

94 ±

0.0

80.

52 ±

0.0

90.

77 ±

0.0

30.

37 ±

0.0

40.

66 ±

0.0

530

.46

± 0.

49L

3.34

± 0

.28

2.63

± 0

.12

2.86

± 0

.34

1.94

± 0

.12

2.00

± 0

.36

1.29

± 0

.09

1.61

± 0

.15

1.03

± 0

.09

1.45

± 0

.03

1.16

± 0

.04

1.27

± 0

.05

1.02

± 0

.09

G. g

rana

telli

i3x

(Mad

)

S0.

78 ±

0.1

60.

63 ±

0.1

40.

63 ±

0.2

10.

81 ±

0.1

9 1.

01 ±

0.1

00.

80 ±

0.1

20.

53 ±

0.0

80.

78 ±

0.0

80.

62 ±

0.1

20.

54 ±

0.1

10.

78 ±

0.1

10.

41 ±

0.1

128

.68

± 2.

62L

3.36

± 0

.58

2.93

± 0

.42

2.56

± 0

.37

1.87

± 0

.22

1.47

± 0

.17

1.30

± 0

.22

1.51

± 0

.07

1.18

± 0

.07

1.18

± 0

.07

1.10

± 0

.14

0.98

± 0

.14

0.93

± 0

.03

G. m

auri

tani

ca

3x

S0.

95 ±

0.1

30.

73 ±

0.1

60.

75 ±

0.1

01.

37 ±

0.2

60.

96 ±

0.1

30.

98 ±

0.2

31.

39 ±

0.1

80.

74 ±

0.1

60.

97 ±

0.2

81.

06 ±

0.2

20.

79 ±

0.1

40.

79 ±

0.0

342

.69

± 4.

80L

4.58

± 0

.67

4.65

± 0

.47

4.31

± 0

.81

2.90

± 0

.35

2.80

± 0

.79

2.48

± 0

.46

1.64

± 0

.23

2.03

± 0

.29

1.78

± 0

.23

1.50

± 0

.13

1.53

± 0

.27

1.02

± 0

.09

G. s

icul

a 2x

S1.

22 ±

0.2

71.

13 ±

0.6

00.

80 ±

0.3

21.

21 ±

0.7

01.

14 ±

0.3

50.

91 ±

0.0

90.

99 ±

0.2

10.

82 ±

0.2

91.

19 ±

0.3

00.

90 ±

0.1

30.

78 ±

0.2

30.

73 ±

0.1

435

.82

± 12

.6L

4.96

± 2

.69

4.94

± 1

.68

4.08

± 1

.02

2.72

± 0

.65

2.51

± 0

.90

2.35

± 0

.83

1.88

± 0

.67

1.95

± 0

.62

1.47

± 0

.46

1.51

± 0

.59

1.31

± 0

.08

1.11

± 0

.34

G. t

ison

iana

2x

S0.

92 ±

0.2

00.

72 ±

0.0

90.

60 ±

0.1

11.

22 ±

0.3

71.

18 ±

0.4

80.

77 ±

0.2

81.

11 ±

0.1

90.

93 ±

0.1

50.

59 ±

0.1

10.

83 ±

0.2

00.

63 ±

0.2

40.

63 ±

0.1

339

.88

± 5.

17L

4.62

± 0

.65

4.32

± 0

.71

3.89

± 0

.57

2.85

± 0

.46

2.33

± 0

.39

2.32

± 0

.43

1.74

± 0

.39

1.64

± 0

.36

1.96

± 0

.24

1.54

± 0

.33

1.53

± 0

.19

1.03

± 0

.30

gagea karyology 99

Fig. 7 — Gagea granatellii (Parl.) Parl. A. Metaphasic plate, showing 2n = 36 chromosomes (plants from Apulia); B. haploid idiogram. Scale bars = 5µm.

Fig. 8 — Gagea granatellii (Parl.) Parl. A. Metaphasic plate, showing 2n = 36 chromosomes (plants from Madonie); B. haploid idiogram. Scale bars = 5µm.

Table 3 — Karyotype features of the studied Gagea species: number of chromosomes of terminal, sub-termi-nal, sub-median and median type (cfr. Levan et al. 1964); A1 and A2 asimmetry indexes (cfr. Romero Zarco 1986).

t st sm m A1 A2

G.bohemica 6x 0 18 24 30 0.49 ± 0.01 0.37 ± 0.05G.bohemica 4x 0 12 4 32 0.34 ± 0.04 0.32 ± 0.05G. chaberti 3x 0 9 15 12 0.49 ± 0.03 0.34 ± 0.02G. dubia 4x 0 12 8 28 0.45 ± 0.03 0.34 ± 0.01G. foliosa 3x 0 15 9 12 0.48 ± 0.04 0.34 ± 0.06G. granatellii 3x (Pug) 0 15 9 12 0.53 ± 0.01 0.32 ± 0.02G. granatellii 3x (Mad) 0 9 15 12 0.53 ± 0.01 0.35 ± 0.07G. mauritanica 3x 9 0 18 9 0.54 ± 0.02 0.35 ± 0.02G. sicula 2x 2 4 10 8 0.53 ± 0.08 0.39 ± 0.07G. tisoniana 2x 6 4 10 4 0.59 ± 0.05 0.37 ± 0.02

peruzzi 100

Gagea tisoniana Peruzzi, Bartolucci, Frignani et Minut. - The chromosome complement of the studied population from locus classicus of the spe-cies (Tuscany) has revealed to be diploid, with 2n = 24 chromosomes (Fig. 13A). Karyotype formula can be expressed as follows: 2n = 24 = 2x = 6st +

Fig. 10 — Gagea mauritanica Durieu. Metaphasic plate, showing 2n = 36 chromosomes (plants from Apulia). Scale bars = 5µm.

Fig. 9 — Gagea mauritanica Durieu. A. Metaphasic plate, showing 2n = 36 chromosomes (plants from France); B. haploid idiogram. Scale bars = 5µm.

4sm + 2st + 2m + 2sm + 2st + 4sm + 2m (Tables 2-3; Fig. 13B). Chromosome size ranges from 1.65 to 5.03 µm in average.

Gagea villosa (M. Bieb.) Sweet. - The chromo-some complement of the studied population from Abruzzo has revealed to be tetraploid, with 2n = 48 chromosomes (Fig. 14). Because of the average quality of the plates, we were not able to build the idiogram for this species.

Karyotype asymmetry - The A1 and A2 asymmetry indexes for the species whose karyotype was stud-ied in detail are reported in Table 3. Fig. 15 shows a comparison of asymmetry indexes among Ga-gea sections whose karyotype was studied in lit-erature, while Fig. 16 points out the relationships within the sect. Didymobulbos (all the investigated species – but for G. tisoniana (sect. Gagea) – be-long to that section).

DISCUSSION

Gagea bohemica (Zauschn,) Schult. et Schult. f. - Euro-Mediterranean species (PERUZZI 2003; PERUZZI and TISON 2004b). Our 2n = 48 count-ing from Abruzzo and Sicily agree with those obtained on plants from Calabria (PERUZZI 2003) and Abruzzo (PERUZZI and BARTOLUCCI 2006). On the other hand, our 2n = 72 counting from Mado-nie (Sicily) results completely new for this species.

gagea karyology 101

Indeed, G. bohemica is known as extremely vari-able in its ploidy level, without any clear relation to geographical distribution (PERUZZI and TISON 2004b). However, until present, only 2n = 24, 36, 48 and 60 cytotypes were known (PERUZZI 2003 and literature cited therein).

Gagea chaberti A. Terracc. - A species likely en-demic to N Africa – Algerie and Tunisia (TISON 2004c). This is the first counting for this triploid unit.

Gagea chrysantha Schult. et Schult. f. s.s. - The typ-ical G. chrysantha is known for a sole certain local-ity: Ficuzza (Sicily), its locus classicus (PERUZZI and TISON 2005). On the other hand, it is the oldest available name for a group of critical and doubtful taxa still under study, named G. chrysantha sensu lato and provisionally including several taxa, such as G. amblyopetala Boiss. & Heldr, G. longifolia Lojac. nom. illeg. (= G. amblyopetala Auct. Fl. Ital.) and G. sicula Lojac. This is the first counting effected on plants to refer to G. chrysantha sensu stricto. Its triploid (2n = 36) level is shared with some population of G. sicula (see beyond) and with all the studied populations of “G. longifolia” (PERUZZI 2003; PERUZZI and AQUARO; 2005).

Gagea dubia A. Terracc. - Mediterranean-W. Asiat-ic species (TISON 2004c). Our result (2n = 48) is the second for Italian plants, and agrees with a previ-ous counting from Abruzzo – C Italy (PERUZZI and BARTOLUCCI 2006) and with data reported from W Asia (Peruzzi, 2003 and literature cited therein). According to the same reference, also diploid 2n = 24 cytotypes are known for this species.

Gagea foliosa (J. et C. Presl) Schult. et Schult. f. - Central Mediterranean species, occurring in Sar-dinia, Sicily and N Algerie (TISON 2004c). This is the first detailed karyotype analysis for this trip-loid unit and it was effected on the same plant material studied by PERUZZI and AQUARO (2005).

Fig. 11 — Gagea sicula Lojac. A. Metaphasic plate, showing 2n = 24 chromosomes (plants from locus classicus); B. haploid idiogram. Scale bars = 5µm.

Fig. 12 — Gagea sicula Lojac. Metaphasic plate, show-ing 2n = 36 chromosomes (plants from Iblei). Scale bars = 5µm.

peruzzi 102

Gagea granatellii (Parl.) Parl. - Central-western Mediterranean species (TISON 2004c). This is the first detailed karyotype analysis for this unit. Our counting (2n = 36) agree with those obtained on plants from Calabria – S Italy (PERUZZI 2003)

and from a different Sicilian locality (PERUZZI and AQUARO 2005). For this species also 2n = 24 (TISON 1998, plants from France) and 2n = 48 (PERUZZI and BARTOLUCCI 2006, plants from Abruzzo) cytotypes are known.

Gagea mauritanica Durieu - Central-western Mediterranean species (TISON 2004c). The stud-ied French population was quoted in MOLINA et al. (1998). Our counting (2n = 36) agree with that obtained on plants from a different locality in Apulia (PERUZZI and AQUARO 2005).

Gagea sicula Lojac. - This taxon – known for Sicily and Calabria (S Italy) – belongs to G. chrysantha group and it is very close to the typical G. chrysantha, but rather distinct from G. longifolia Lojac. nom. illeg. and G. amblyopeta-la Boiss. et Heldr. (PERUZZI and TISON 2005). Our counting from locus classicus evidenced diploid plants, never reported before for G. chrysantha group. On the other hand, triploid plants from Iblei agree with previous reports for Calabrian plants (PERUZZI 2003 sub G. foliosa).

Gagea tisoniana Peruzzi, Bartolucci, Frignani et Minut. - A recently described C Italian endem-ic species, related to G. pusilla (F. W. Schmidt) Sweet (PERUZZI et al. 2007). This is the first de-tailed karyotype analysis for this diploid taxon

Fig. 13 — Gagea tisoniana Peruzzi, Bartolucci, Frignani et Minut. A. Metaphasic plate, showing 2n = 24 chromo-somes; B. haploid idiogram. Scale bars = 5µm.

Fig. 14 — Gagea villosa (M. Bieb.) Sweet. Metapha-sic plate, showing 2n = 48 chromosomes. Scale bars = 5µm.

gagea karyology 103

Fig. 15 — Scatter plot of the karyotype asymmetry indexes according to the different Gagea sections. Data derive from the present study (Table 3) and from literature (cfr. PERUZZI and AQUARO 2005).

Fig. 16 — Scatter plot of the karyotype asymmetry indexes according to the different species of Gagea sect. Didy-mobulbos. Data derive from the present study (Table 3) and from literature (cfr. PERUZZI and AQUARO 2005).

peruzzi 104

and it was effected on plant material from its locus classicus.

Gagea villosa (M. Bieb.) Sweet - Eurasiatic spe-cies (PIGNATTI 1982). Our result (2n = 48) is the second for Italian plants, and agrees with a previ-ous counting from Calabria – S Italy (PERUZZI and AQUARO 2005) and with data reported from sev-eral countries (PERUZZI 2003 and literature cited therein).

Karyotype asymmetry - G. tisoniana, the only representative of G. sect. Gagea investigated, show affinity with a diploid cytotype of G. prat-ensis (Pers.) Dumort. from Balkans (PERUZZI and AQUARO 2005 and literature cited therein). From a general point of view, G. sect. Didymobulbos clearly tends to have less asymmetrical karyotypes than other sections (Fig. 15), as already shown by PERUZZI and AQUARO (2005). As far the relation-ships within G. sect. Didymobulbos are concerned (Fig. 16), G. bohemica appears very variable. G. sicula and G. chrysantha s.l. are very similar to each other, but also to G. villosa and to one G. bohemica accession. G. granatellii and G. mau-ritanica accessions appear very closely related, as well as G. chaberti and G. foliosa ones. Finally, G. dubia occupies a somewhat intermediate po-sition among G. chrysantha group (G. chrysantha s.l. + G. sicula), G. bohemica and G. foliosa.

CONCLUSIONS

This paper evidenced the chromosome com-plements of 10 Gagea representatives, from a total of 15 central Mediterranean populations, involv-ing France, Tunisia and Italy. Among them, are noteworthy the first account on G. chaberti, a new cytotype (2n = 72) for G. bohemica and the first karyotype analysis for G. foliosa, G. granatellii and G. tisoniana.

Our results on karyotype asymmetry confirm – also extending the investigation to an higher number of species and accessions – the trend al-ready evidenced in PERUZZI and AQUARO (2005), particularly concerning the relative low asym-metry of G. sect. Didymobulbos respect to other sections. Within it, there is now delineating some tendency to form species groups, but further in-vestigation is required on this matter.

With the present contribution, for 97 Gagea species it is known something about karyology (in particular for 100 taxa, with a total of 267 dif-

ferent chromosome counting); i.e. ca. 40% of the presumed total number of existing species.

Acknowledgements – I wish to thank: Dr. Jean-Marc Tison (L’Isle d’Abeau, France), for providing bulbs of G. chaberti from Tunisia and G. mauritanica from France; Dr. Gabriella Aquaro and Dr. Katia Francesca Caparelli (University of Calabria) for techni-cal assistance.

REFERENCES

ALI S.I., 2006 — Two new species of Gagea Salisb. (Lil-iaceae) from Pakistan. Pakistan Journal of Botany, 38(1): 43-46.

CAPARELLI K.F., PERUZZI L. and CESCA G., 2006 — A comparative analysis of embryo-sac development in three closely-related Gagea Salisb. species (Liliaceae), with some consideration on their reproductive strate-gies. Plant Biosystems, 140(2): 115-122.

BARTOLUCCI F. and PERUZZI L., 2008 — Distribuzione del genere Gagea Salisb. (Liliaceae) nell’Appennino centro-settentrionale. Biogeographia: in press.

BAYER E. and LÓPEZ GONZÁLEZ G., 1989 — Nomen-clatural notes on some names in Gagea Salisb. (Lil-iaceae). Taxon, 38(4): 643-645.

CUCCUINI P. and LUCCIOLI E., 1995 — Tipificazione di Ornithogalum spathaceum Hayne (Liliaceae) e presenza di Gagea spathacea (Hayne) Salisb. nella flora italiana. Webbia, 49(2): 253-264.

DAVLIANIDZE M.T. and BOKERIA M., 2005 — Contri-bution to the cytotaxonomy of the Georgian flora. XVII International Botanical Congress, Wien 17-23/07/2005; Book of Abstracts: 345.

FERRER GALLEGO P.P., LAGUNA LUMBRERAS E., ALBA VIL-LEGAS S. and TISON J.-M., 2007 — Sobre la presencia de Gagea lacaitae A. Terracc. (Liliaceae) en la flora valenciana. Acta Botanica Malacitana, 32: 1-12.

GARGANO D., PERUZZI L., CAPARELLI K.F. and CESCA G., 2007 — Preliminary observations on the reproduc-tive strategies in five early-flowering species of Gagea Salisb. (Liliaceae). Bocconea, 21: 349-358.

GREILHUBER J., EBERT I., LORENZ A. and VYSKOT B., 2000 — Origin of facultative heterochromatin in the endosperm of Gagea lutea (Liliaceae). Protoplasma, 212: 217-226.

HENKER H., 2005 — Goldsterne und Stinsenpflanzen in Mecklenburg-Vorpommern. Teil 1. Die Goldsterne von Mecklenburg-Vorpommern unter besonderer Berücksichtigung kritisher und neuer Sippen. Bota-nischer Rundbrief für Mecklenburg-Vorpommern, 39: 3-90.

JOHN H., PETERSON A. and PETERSON J., 2004 — Zum taxonomischen Rang zweier kritischer Sippen der Gattung Gagea in Mitteleuropa. Mitteilungen zur floristichen Kartierung in Sachsen-Anhalt (Halle), 9: 15-26.

gagea karyology 105

KAPASA M., NIKOLAIDI T., BAREKA E.P. and KAMARI G., 2001 — Reports (1236-1243). In: Kamari G., Blanché G., Garbari F., Mediterranean chromosome number reports 11. Flora Mediterranea, 11: 448-454.

LEVAN A., FREDGA K. and SANDBERG A.A., 1964 — No-menclature for centromeric position on chromosomes. Hereditas, 52: 201-220.

LEVICHEV I.G., 1990 — The synopsis of the genus Gagea (Liliaceae) from the western Tien-Shan. Botanical Journal (St. Petersburg), 75(2): 225-234.

LEVICHEV I.G., 1999 — The morphology of Gagea Salisb. (Liliaceae) I. Subterranean organs. Flora, 194: 379-392.

LEVICHEV I.G., 2001 — New species of the genus Ga-gea Salisb. (Liliaceae) from western regions of Asia. Turczanimowia, 4(1-2): 5-35.

LEVICHEV I.G., 2006 — Four new species of the genus Gagea salisb. (Liliaceae) from Western Himalayas and the adjoining regions. Pakistan Journal of Bot-any, 38(1): 47-54.

LEVICHEV I.G., 2006b — A review of the Gagea (Lil-iaceae) species in the flora of Caucasus. Botanical Journal (St. Petersburg), 91(6): 917-951.

LEVICHEV I.G. and ALI S.I., 2006 — Seven new species of the genus Gagea Salisb. (Liliaceae) from Western Himalayas and adjoining regions. Pakistan Journal of Botany, 38(1): 55-62.

LEVICHEV I.G. and TISON J.-M., 2004 — Typification of the Caucasian Gagea (Liliaceae) taxa described by Karl Koch. Candollea, 59: 119-133.

MESICEK J. and HROUDA L., 1974 — Chromosome num-bers in Czechoslovak species of Gagea (Liliaceae). Folia Geobotanica and Phytotaxonomica, Praha, 9: 359-368.

MOLINA J., MICHAUD H., ROUX J.-P. and TISON J.-M., 1998 — Gagea mauritanica Durieu (Liliaceae), es-pèce nouvelle puor la flore française. Bulletin Men-suel de la Société Linnéenne de Lyon, 67: 77-78.

ÖZHATAY N., 2002 — Diversity of bulbous monocots in Turkey with special reference to chromosome num-bers. Pure and Applied Chemistry, 74(4): 547-555.

PATTERSON T.B. and GIVNISH T.J., 2002 — Phylogeny, concerted convergence, and phylogenetic niche con-servatism in the core Liliales: Insights from rbcL and ndhF sequence data. Evolution, 56: 233-252.

PERUZZI L., 2003 — Contribution to the cytotaxonomi-cal knowledge of Gagea Salisb. (Liliaceae) sect. Fo-liatae A. Terracc. and synthesis of karyological data. Caryologia, 56(1): 115-128.

PERUZZI L., 2006 — Taxonomic considerations on the nomenclatural types of Gagea rhodiaca A. Terracc. and G. reticulata subsp. africana A. Terracc. (Liliace-ae), kept at Pisa (PI). Atti della Società Toscana di Scienze Naturali, Memorie, serie B, 113: 69-71.

PERUZZI L., 2008 — Hybridity as a main evolutionary force in the genus Gagea Salisb. (Liliaceae). Plant Biosystems, 142(1): 179-183.

PERUZZI L. and AQUARO G., 2005 — Contribution to the cytotaxonomical knowledge of Gagea Salisb.

(Liliaceae). II. Further karyological studies on Italian populations. Candollea, 60(1): 237-253.

PERUZZI L. and BARTOLUCCI F., 2006 — Gagea luberon-ensis J.-M. Tison (Liliaceae) new for the Italian flora. Webbia, 61(1): 1-12.

PERUZZI L., BARTOLUCCI F., FRIGNANI F. and MINUTILLO F., 2007 — G. tisoniana: a new species of Gagea Salisb. sect. Gagea (Liliaceae) from C Italy. Botanical Journal of the Linnean Society, 155(3): 337-347.

PERUZZI L. and CAPARELLI K. F., 2007 — Gagea pedun-cularis (J. & C. Presl) Pascher (Liliaceae) new for the Italian flora. Webbia, 62(2): 261-268.

PERUZZI L. and GARGANO D., 2005 — Distribuzione del genere Gagea Salisb. (Liliaceae) in Calabria. Inform-atore Botanico Italiano, 37(2): 1117-1124.

PERUZZI L. and TISON J.-M., 2004 — Typification and taxonomic status of eleven taxa of Gagea Salisb. (Lil-iaceae) described by Achille and Nicola Terracciano and conserved at Napoli (NAP). Candollea, 59(2): 325-346.

PERUZZI L. and TISON J.-M., 2004b — Verso una revi-sione biosistematica del genere Gagea Salisb. (Lil-iaceae) in Italia. Un nuovo tipo di approccio. Inform-atore Botanico Italiano, 36(2): 470-475.

PERUZZI L. and TISON J.-M., 2005 — Typification and taxonomic status of six taxa of Gagea Salisb. (Liliace-ae) described from Sicily and conserved at Palermo (PAL). Candollea, 60(2): 289-298.

PERUZZI L. and TISON J.-M., 2006 — Typification of the names and taxonomic status of six taxa of Gagea Salisb. (Liliaceae) conserved at Firenze (FI). Candol-lea, 61(2): 293-303.

PERUZZI L. and TISON J.-M., 2007 — Typification of six critical Mediterranean Gagea Salisb. (Liliaceae) spe-cies. Candollea, 62(2): 173-188.

PERUZZI L. and ZARREI M., 2007 — Typification of some critical taxa of Gagea Salisb. (Liliaceae) from W Asia. Candollea 62(2): 237-244.

PETERSON A., JOHN H., KOCH E. and PETERSON J., 2004 — A molecular phylogeny of the genus Gagea (Lil-iaceae) in Germany inferred from non-coding chloro-plast and nuclear DNA sequences. Plant Systematics and Evolution, 245: 145-162.

PETERSON A., LEVICHEV I.G. and PETERSON J., 2008 — Systematics of Gagea and Lloydia (Liliaceae) and infrageneric classification of Gagea based on molecu-lar and morphological data. Molecular Phylogenet-ics and Evolution, 46: 446-465.

PIGNATTI S., 1982 — Flora d’Italia, 3: 352-356. – Edag-ricole, Bologna.

ROMERO ZARCO C., 1986 — A new method for estimat-ing karyotype asymmetry. Taxon, 35(3): 526-530.

TISON J.-M., 1998 — Gagea granatellii (Parl.) Parl. en France. Le Monde des Plantes, 462: 1-6.

TISON J.-M., 2004 — Gagea polidorii J.-M. Tison, espèce méconnue du sud-ouest des Alpes et des Apennins. Acta Botanica Gallica, 151(3): 319-326.

TISON J.-M., 2004b — Identité et situation taxonomique de Gagea polymorpha Boissier. Candollea, 59(1): 109-117.

peruzzi 106

TISON J.-M., 2004c — Contribution à la connaissance du genre Gagea Salisb. (Liliaceae) en Afrique du Nord. Lagascalia, 24: 67-87.

TISON J.-M. and PERRET P., 2004 — Typification d’Ornithogalum pusillum F. W. Schmidt et relations taxonomiques entre Gagea pusilla (F. W. Schmidt) Sweet, Ornithogalum clusii Tausch et G. clusiana Schult. & Schult. f. Candollea, 59(1): 103-108.

ZARREI M. and ZARRE S., 2005 — Pollen morphology of the genus Gagea (Liliaceae) in Iran. Flora, 200: 96-108.

ZARREI M. and ZARRE S., 2005a — A new species of Ga-gea (Liliaceae) from Iran. Nordic Journal of Botany, 23: 269-274.

ZARREI M., ZARRE S., WILKIN P. and RIX M., 2007 — Sys-tematic revision of the genus Gagea Salisb (Liliaceae)

in Iran. Botanical Journal of the Linnean Society, 154(4): 559-588.

ZHAO L.-Q. and YANG J., 2006 — Gagea daqingshan-ensis (Liliaceae), a new species from Inner Mongolia, China. Annales Botanici Fennici, 43: 223-224.

ZHAO Y.-Z. and ZHAO L.-Q., 2003 — A new species of Gagea (Liliaceae) from Nei Mongol, China. Acta Phytotaxonomica Sinica, 41(4): 393-394.

ZHAO Y.-Z. and ZHAO L.-Q., 2004 — Gagea chinen-sis (Liliaceae), a new species from Inner Mongolia, China. Annales Botanici Fennici, 41: 297-298.

Received August 6th 2007; accepted November 9th 2007