Karyological Study in Fifteen Leucocoryne Taxa

Biologia 67/2: 289—295, 2012 Section Botany DOI: 10.2478/s11756-012-0001-5

Karyological study in ﬁfteen Leucocoryne taxa (Alliaceae)

Paola Jara-Arancio1,2*, Pedro Jara-Seguel3, Claudio Palma-Rojas 4,GinaArancio4 &RaulMoreno4

1Instituto de Ecología y Biodiversidad (IEB), Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Nu˜˜ noa, Casilla 653, Santiago, Chile; e-mail: [email protected] 2Departamento de Ciencias Biológicas Facultad de Ciencias Biológicas y Departamento de Ecología y Biodiversidad Fac- ultad de Ecología y Recursos Naturales, Universidad Andrés Bello, Santiago, Chile 3Escuela de Ciencias Ambientales, Facultad de Recursos Naturales, Universidad Católica de Temuco, Casilla 15-D, Temuco, Chile 4Departamento de Biología, Facultad de Ciencias, Universidad de La Serena, Casilla 599, La Serena, Chile

Abstract: The karyotype of fifteen Leucocoryne taxa was studied, assessing characteristics such as chromosome morphology and size, secondary constriction location, and asymmetry level. Two groups of Leucocoryne taxa were described based on chromosome number (2n = 10 and 2n = 18) and karyotype asymmetry. The haploid karyotype formula for the group 2n = 10 was 3m + 2st (or 2t), whereas for the group 2n = 18 was 7m + 2st (or 2t). Such results corroborate the karyotype descriptions previously carried out for some taxa of the genus. Leucocoryne taxa showed a high resemblance in chromosome morphology, but inter-specific differences were found in mean chromosome size. These data and previous studies based on gross chromosome morphology support Crosa’s hypothesis, which suggests that the cytotype 2n = 10 is diploid and perhaps ancestral, whereas that the cytotype 2n = 18 is tetraploid but with an additional chromosome fusion being probably a derived status. Key words: Leucocoryne; karyotype morphology; polyploidy

Introduction tepals, superior ovary with several ovules, short style and capitate stigma (Muñoz & Moreira 2005). These The study of karyotype morphology among related taxa plants have been described as food source for fosso- is a fundamental aspect to understand the process of ge- rial rodents, thus playing an important role as primary netic variation, genome evolution and speciation, and producer in trophic chains of semiarid ecosystems (Reig the taxonomic circumscriptions as has been broadly de- 1970; Contreras & Gutiérrez 1991). Besides, because of scribed in different plant families (Stebbins 1971; Levin their showy flowers, these plants have been successfully 2002). Regarding Asparagales order, some reports have introduced into cultivation, being used as cut, pot and discussed the process on chromosome evolution within garden plants (Hayward 1940; Kroon 1989; Ohkawa et the larger families, such as the diverse and worldwidely al. 1996; De la Cuadra et al. 2002; Verdugo & Texeira distributed Amaryllidaceae family (Sato 1938; Gouss 2006). However, due to its extraction and habitat degra- 1949; Flory 1977; Cisternas et al. 2010; Muñoz et al. dation by anthropic effect, currently some endangered 2011). taxa are recognized whereas other taxa are insufficiently Leucocoryne Lindl. (Alliaceae) is a Chilean en- known, and scarce information has been compiled on its demic genus (Muñoz & Moreira 2000) important as a conservation (Squeo et al. 2001). component of the desert flower diversity. This genus is Biosystematics studies within the genus Leucoco- distributed between 20◦S–37◦S, 2,500 m above sea level, ryne have been previously documented on the basis of with a diversity center between 30◦S–34◦Sinasemiarid morphological characteristics (Zöellner 1972; Ravenna zone (Zöellner 1972; Ravenna 1973; Hoffmannn 1978; 1998; Muñoz & Moreira 2000; Mansur et al. 2002) be- Ravenna 1978; Muñoz & Moreira 2000; Squeo et al. ing currently recognized 14 taxa (Zöellner 1972). How- 2001; Mansur et al. 2002). Genus taxa are character- ever, at present the taxonomic classification of some ized by the presence of bulbs, linear leaves, spathe of populations is controvertial due to the high phenotypic two lanceolate bracts, umbrella inflorescence at the base variation (Muñoz & Moreira 2000). In contrast, kary- with up to 12 flowers, 6 narrow or wide tepals, 3 or 6 ological data only for ten accurately identified Leucoco- stamens inserted in the tube, 3 cylindrical or flattened ryne taxa and other six non-identified taxa are available

* Corresponding author

c 2012 Institute of Botany, Slovak Academy of Sciences 290 P. Jara-Arancio et al.

(Zöellner 1972; Crosa 1988; Bahamondes & Labarca Results 1994; Araneda et al. 2004; Mansur & Cisterna 2005), but this background information has been scarcely used Feulgen stained karyotype of Leucocoryne taxa are in systematic studies. As to this genus, a complex series shown in Fig. 1. Karyotypes were described for the of chromosome numbers has been described, including first time for L. appendiculata, L. conferta, L. dimor- diploid and polyploid taxa, as well as some metaphases phopetala, L. macropetala, L. pauciflora, L. vittata, L. of hybrid specimens that have been counted (Salas & aff. vittata (Table 1). Two plant groups based in the Mansur 2004). chromosome numbers 2n = 10 (n = 5) and 2n = 18 (n In order to increase the karyological antecedent = 9) were identified in Leucocoryne taxa studied in this for Leucocoryne genus, in this work the karyotype work. morphology of fifteen taxa is described, and seven Chromosome morphology was uniform within of of them are examined here for the first time. Be- each studied group, with little variations. Thus in the sides, inter-specific cytogenetic relationships are estab- taxa group with 2n = 10 three pairs were metacentric lished, including karyological information previously and two pairs were subtelocentric or telocentric (hap- documented by other authors. loid karyotype formula = 3m + 2st or t). In the case of the taxa group with 2n = 18, seven pairs were meta- Material and methods centric and two pairs were subtelocentric or telocentric (haploid karyotype formula = 7m + 2st or t). However, Plants of one or two accessions of fifteen taxa of Leuco- L. ixioides shows seven metacentric, one submetacen- coryne were collected from naturally growing populations tric and one subtelocentric pair. Fundamental numbers and their collection sites are shown in Table 1. Within some (FN) of the haploid set for the group n = 5 were FN = taxa, sub-species or affinities were also examined, including seventeen taxa in total. In the case of L. talinensis, seeds 9 and 10 whereas for the group n = 9 were FN = 16, previously stored in the laboratory were used in this study. 17, and 18. Nevertheless, adult plants of L. talinensis were obtained Chromosome morphology displayed for Leucoco- from germinated seeds to corroborate taxonomic indenti- ryne taxa of suggests the presence of moderately sym- fication. The voucher specimens were deposited at the ULS metric karyotypes within this genus with predominance herbarium (Universidad de La Serena-CHILE). The taxa of metacentric chromosomes but with the presence of were identified according to taxonomic keys documented by telo- or subtelocentric chromosomes, and exception- Zöellner (1972), Ravenna (1998), Muñoz & Moreira (2000), ally submetacentric chromosomes. The correlation be- Mansur et al. (2002) and Zuloaga et al. (2008). In the laboratory, plants were kept with their bulbs submerged in wa- tween A1 and A2 asymmetry index are shown in Fig. 2. ter, with constant aeration, to favor active growth of ad- Within the group 2n = 18 with more symmetric kary- ventitious roots. After five to ten days, 5 mm-long root tips otypes (group A), the A1 values ranged among 0.20 to were excised from the bulbs. The root tips were pre-treated 0.35 whereas for the group 2n = 10 with less symmetric with a 0.05% aqueous solution of colchicine at 4 ◦Cfor6 karyotype (group B) the A1 values ranged between 0.39 ◦ hours, fixed in ethanol-glacial acetic acid (3:1) at 4 Cfor to 0.45. In the case of the A2 values of these, fall within ◦ 24 h, and stored in 70% ethanol at 4 C until chromosome the range 0.17 to 0.30 for both groups (see A1 and A2 processing was done. The root tips were stained with Feul- ◦ values in Table 1). Thus, the correlation between A1 gen reaction (hydrolyzed for 7 minutes in HCl 1N at 60 C, andA2indexesplottedforLeucocoryne taxa is con- stained with Schiff reagent for 60 minutes and washed in sulfurous water) (Navarrete et al. 1983) and slides were sistent with the identification of two different groups made by squashing root meristems. The determination of based in chromosome number. karyotype parameters was carried out from metaphase im- Total chromosome length (THL) and mean chro- ages (from ten examined plants per each taxon). Measure- mosome size (MCS) for the studied taxa are shown ments were made on almost five cells, and the short arms in Table 1. Higher total haploid set length and mean (SA) and long arms (LA) were measured using the software chromosome size within the cytotype 2n = 18 were MicroMeasure 3.3 (Reeves 2001). The total relative length described for L. dimorphopetala (THL = 315.63 µm, (LR) of each chromosome pair was calculated and expressed MCS = 35.1+2.9 µm), whereas the lower values were as a percentage of the total haploid set length. Chromosome found in L. coquimbensis var. coquimbensis (THL= shape was determined based on the centromeric index pro- µ µ posed by Levan et al. (1964) (ratio short arm / total chro- 98.72 + 0.4 m, MCS = 11 m). Similarly, higher total mosome length) and karyotype was made up in order of haploid set length and mean chromosome size within decreasing chromosome length. In addition, arm fundamen- the cytotype 2n = 10 were described for L. conferta tal number (FN) was estimated for the karyotype of each (THL = 147.58 µm, MCS = 29.5 + 2.0 µm), whereas taxa where metacentric, submetacentric and subtelocentric lower values were found in L. aff. vitatta (THL = 77.56 chromosomes showed two arms (short arm and long arm), +2.71µm, MCS = 15.5 + 0.5 µm). Values of the and telocentric chromosomes showed one arm (only long other studied Leucocoryne taxa are within the range arm) (Spotorno 1985). Intrachromosomal asymmetry index described for their respective 2n number. A1 and interchromosomal asymmetry index A2 proposed by Secondary constriction (SC) was observed on the Romero-Zarco (1986) were estimated for Leucocoryne taxa L. macropetala, and correlated to determine karyotype affinity. Total haploid short arms of the chromosome pair 5 in L. vittata L. vittata. L. alliacea, L. appendic- set length (in µm) and mean chromosome size (in µm) were and aff. In additional characteristics determined here for the karyotype ulata, L. coquimbensis, L. dimorphopetala, L. ixioides of each Leucocoryne taxa. and L. violascescens, SC was observed on the short Karyological study in fifteen Leucocoryne taxa (Alliaceae) 291

Table 1. Cytogenetic characters of Leucocoryne taxa studied in this work. HKF, haploid karyotype formula; FN, arm fundamental number of haploid set; THL, total haploid set length; MCS, mean chromosome size. masl, meters above sea level. (*) Species studied for the ﬁrst time.

Taxa Collection sites 2n HKF FN THL ± SD MCS ± SD A1 ± SD A2 ± SD (Latitude – Longitude; Altitude (in µm) (in µm) Index Index m a.s.l.)

L. alliacea Miers ex Farellones (33◦20S–70◦19W; 18 7m, 2st 18 196.22 ± 13.87 21.8 ± 1.5 0.27 ± 0.0 0.17 ± 0.0 Lindl. 2700) L. alliacea Miers ex León muerto (29◦20S–70◦39W; 18 7m, 2st 18 Lindl. 2770) L. angustipetala Gay Tofo (28◦58S–70◦56W; 730) 10 3m, 2st 10 117.01 ± 14.94 23.4 ± 3.0 0.40 ± 0.0 0.18 ± 0.0 L. angustipetala Gay Domeyko (28◦59S–70◦56W; 759) 10 3m, 2st 10 L. appendiculata Phil. Iquique (20◦13S–70◦09W; 10) 18 7m, 2st 18 203.58 ± 8.19 22.6 ± 0.9 0.29 ± 0.0 0.23 ± 0.0 L. appendiculata Pan de Azúcar (26◦08S–70◦35W; 18 7m, 2st 18 Phil. * 510) L. conferta Zoëllner * Cuesta Cavilolén (31◦46S– 10 3m, 2st 10 147.58 ± 9.94 29.5 ± 2.0 0.41 ± 0.0 0.30 ± 0.0 71◦19W; 700) L. coquimbensis Phil. Tongoy (30◦15S–71◦30W; 40) 18 7m, 1st, 1t 17 98.72 ± 0.40 11.0 ± 0.0 0.30 ± 0.0 0.18 ± 0.0 var. coquimbensis L. coquimbensis Phil. Panul (30◦00S–71◦24W; 100) 18 7m, 1st, 1t 17 var. coquimbensis L. coquimbensis Phil. Panul (30◦00S–71◦24W; 100) 18 7m, 1sm, 1st 18 146.56 ± 36.37 16.3 ± 4.1 0.25 ± 0.0 0.17 ± 0.0 var. alba Zoëllner L. coquimbensis Phil. Juan Soldado (29◦40S–71◦18W; 18 7m, 1sm, 1st 18 var. alba Zoëllner 300) L. dimorphopetala Freirina (28◦31S–70◦04W; 100) 18 7m, 2st 18 315.63 ± 26.02 35.1 ± 2.9 0.20 ± 0.0 0.21 ± 0.0 (Gay) Rav. * L. dimorphopetala Sauce Pérez (28◦40S–71◦06W; 18 7m, 2st 18 (Gay) Rav. 607) L. ixioides (Hook) El Manzano (33◦41S–71◦09W; 18 7m, 1sm, 1st 18 274.39 ± 22.18 30.5 ± 2,5 0.28 ± 0.0 0.17 ± 0.0 Lindl. 175) L. ixioides (Hook) Quebrada las Vacas (32◦42S– 18 7m, 1sm, 1st 18 Lindl. 71◦13W; 200) L. macropetala Phil. * Cerro Grande (29◦56S–71◦13W; 10 3m, 2st 10 97.98 ± 7.05 19.6 ± 1.4 0.40 ± 0.0 0.19 ± 0.0 520) L. macropetala Phil. Los Burros (27◦54S–71◦07W; 50) 10 3m, 2st 10 L. narcissoides Phil. Los Burros (27◦54S–71◦07W; 50) 18 7m, 1st, 1t 17 99.29 ± 5.75 11.0 ± 0.6 0.29 ± 0.0 0.23 ± 0.0 L. narcissoides Phil. Paposo (25◦00S–70◦28W; 50) 18 7m, 1st, 1t 17 L. foetida Phil. * Viña del Mar (33◦31S–71◦34W; 18 7m, 2t 16 254.37 ± 7.95 28.3 ± 0.9 0.31 ± 0.0 0.19 ± 0.0 150) L. foetida Phil. Valparaíso (33◦02S–71◦38W; 18 7m, 2t 16 150) L. pauciflora Phil. * Camino a Rapel (34◦05S– 18 7m, 2st 18 182.23 ± 13.83 20.20 ± 1.5 0.29 ± 0.0 0.18 ± 0.0 71◦32W; 70) L. purpurea Gay Guanaqueros (30◦12S–71◦26W; 10 3m, 1st, 1t 9 88.14 ± 10.71 17.6 ± 2.1 0.45 ± 0.0 0.17 ± 0.0 100) L. purpurea Gay Quebrada las Vacas (32◦42S– 10 3m, 1st, 1t 9 71◦13W; 200) L. talinensis Mansur Colecta planta tipo (30◦49S– 18 7m, 1st, 1t 17 240.96 ± 11.69 26.8 ± 1.3 0.35 ± 0.0 0.23 ± 0.0 71◦33W; 355) L. violacescens Phil. Amolana (31◦12S–71◦37W; 200) 18 7m, 1sm, 1st 18 283.91 ± 22.23 31.5 ± 2.5 0.28 ± 0.0 0.20 ± 0.0 L. violacescens Phil. Alcones (30◦48S–71◦33W; 250) 18 7m, 1sm, 1st 18 L. vittata Rav. * Chigualoco (31◦46S–71◦30W; 15) 10 3m, 2st 10 106.13 ± 8.68 21.2 + 1.7 0.39 ± 0.0 0.17 ± 0.0 L. vittata Rav. Los Vilos (31◦54S–71◦31W; 15) 10 3m, 2st 10 L. aff. vittata Rav. * Las Palmas (31◦16S–71◦35W; 10 3m, 2st 10 77.56 ± 2.71 15.5 ± 0.5 0.40 ± 0.0 0.21 ± 0.0 248)

arms of the chromosome pair 9 (Fig. 1). SC was not mondes & Abarca 1994; Araneda et al. 2004). How- observed in the remaining taxa. ever, this series of 2n number is extended to include some taxa described with 2n = 14 such as L. coquim- Discussion bensis var. alba (Araneda et al. 2004) and the natural hybrid L. coquimbensis × L. purpurea (Salas & Mansur Leucocoryne genus can be characterized as having two 2004). Nevertheless, inconsistencies are observed in L. chromosome numbers 2n = 10 and 2n = 18 as has coquimbensis var.albadue to the fact that two diﬀer- been described in previous works (Crosa 1988; Baha- ent chromosome numbers have been reported, 2n = 18 292 P. Jara-Arancio et al.

Fig. 1a. Karyotypes of: A – Leucocoryne alliacea;B–L. conferta;C–L. appendiculata;D–L. angustipetala;E–L. coquimbensis var. coquimbensis;F–L. coquimbensis var. alba;G–L. dimorphopetala;H–L. ixioides;I–L. macropetala;J–L. narcissoides;K – L. foetida;L–L. pauciﬂora;M–L. purpurea;N–L. talinensis. Karyological study in ﬁfteen Leucocoryne taxa (Alliaceae) 293

Fig. 1b. Karyotypes of: O – L. violacescens;P–L. vittata;Q–L. aﬀ. vittata.Scalebar=10µm.

Table 2. Cytogenetic data for Leucocoryne species previously studied by other authors.

Taxa 2n Haploid karyotype formula Reference

L. ixioides 18 7sm + 2st Zo¨ellner (1972) L. ixioides 18 7m + 1st + 1t Crosa (1988) L. ixioides 18 7m + 1st + 1t Araneda et al. (2004) L. coquimbensis var. coquimbensis 18 7m + 1st + 1t Araneda et al. (2004) L. narcissoides 18 7m + 1st + 1t Araneda et al. (2004) L. violascecens 18 7m + 2st Crosa (1988) L. talinensis 18 7m + 1st + 1t Mansur & Cisterna (2005) L. coquimbensis var. alba 14 5m + 1st + 1t Araneda et al. (2004) L. odorata (L. foetida) 10 3m + 1st + 1t Crosa (1988) L. purpurea 10 3m + 1st + 1t Crosa (1988) L. purpurea 10 3m + 1st + 1t Araneda et al. (2004) L. alliacea 10 3m + 1sm + 1t Crosa (1988) L. alliacea 10 3m + 1sm + 1t Crosa (1988) L. angustipetala 10 3m + 1sm + 1t Crosa (1988)

in this work and 2n = 14 in the study of Araneda et al. n = 5 was 3m + 2st, while for the cytotype n = 9 (2004). Similarly, L. alliacea has been described previ- was 7m + 2st. It is remarkable that although chromo- ously with a number 2n = 10 (Crosa 1988), whereas in some number varies between the two mentioned groups, this work a number 2n = 18 was described (Table 2). similarities in gross chromosome morphology among In all cases, taxa identification should be revised using their karyotypes were observed. Nevertheless, an incre- the voucher specimens deposited in indexed herbarium, ment in the number of metacentric chromosomes in the thus corroborating taxonomic identification and clari- 2n = 18 cytotype is related to lower values of intra- fying the ambiguous chromosome numbers reported so chromosomal asymmetry index A1 in these taxa (more far. In our work, collected specimens of all taxa were symmetry in group A) regarding to 2n = 10 (less sym- checkedbycomparisonwith voucher specimens stored metry in group B), although both groups are cohesive in the indexed ULS herbarium. when comparing the interchromosomal asymmetry in- Karyotype morphology reported in this work for dex A2, whose values are superimposed (between 0.17 Leucocoryne taxa shows resemblances with those de- and 0.30) (Fig. 2). scribed for other taxa of the genus previously exam- All available karyotype data for Leucocoryne are ined (Zöellner 1972; Crosa 1988; Bahamondes & Abarca consistent with Crosa’s (1988) hypothesis, who sug- 1994; Araneda et al. 2004; Salas & Mansur 2004). Stud- gested that cytotype 2n = 10 is diploid and perhaps ied taxa have been adscribed to the groups 2n = 10 ancestral, whereas cytotype 2n = 18 is tetraploid (likely and 2n = 18 exhibiting predominance of metacentric auto-tetraploid) but with an additional chromosome fu- chromosomes (between 60 and 77%) and low quantity sion being probably a derived status. The fundamental of subtelocentric or telocentric chromosomes (between numbers (FN) reported here for both groups of Leu- 22 to 40%) within their respective karyotype formulas cocoryne taxa also support this hypothesis (Table 1). (Fig. 1, Table 1). Most common haploid karyotype for- Then, as an explanation of all this framework, it is pos- mula for the studied taxa belonging to the cytotype sible to suggest that chromosome differentiation in Leu- 294 P. Jara-Arancio et al.

Fig. 2. Correlation between Intrachromosomal Asymmetry Index (A1) and Interchromosomal Asymmetry Index (A2) of Leucocoryne species. The A group represents species with 2n = 18 and B group represents species with 2n = 10. cocoryne taxa may occur basically by both an incre- Acknowledgements ment in number of metacentric chromosomes from 3m to 7m via genome duplication and by a centric fusion of The authors acknowledge CONAF and ULS, CONC and telocentric chromosomes from a base cytotype n = 5. In SGO herbariums for their valuable help, as well as Mélica addition, the number of st-t chromosomes and values of Muñoz’s critical support. We would also thank the Depar- tamento de Biología-ULS, and CONICYT for granting a interchromosomal length are similar among karyotypes fellowship to P. Jara-Arancio. This is a contribution to of the taxa (Fig. 2). the research program of Senda Darwin Biological Station, Another interesting feature observed in the chro- Chiloé, Chile. Postdoctoral fellowship from IEB, ICM P05– mosomes of Leucocoryne taxa studied here are the 002, PFB-23. high values of total haploid set length (THL) and mean chromosome size (MCS) (Table 1). In previous works on Leucocoryne taxa, THL and MCS have not References been described, and only absolute chromosome size µ Araneda L., Salas P. & Mansur L. 2004. Chromosome numbers in (in m) for largest and smallest pairs of karyotype thechileanendemicgenusLeucocoryne (Huilli). J. Am. Soc. are given (Araneda & Mansur 2004; Salas & Mansur Hortic. Sci. 129(1): 77–80. 2004). However, due to different times of pretreatment Bahamondes N. & Labarca V. 1994. Citogenética y probables with colchicine that affects the chromosome condensa- mecanismos de evolución en especies de los géneros Leucoco- tion and shorten chromosome metaphases, those values ryne y Pabellonia (Alliaceae). Tesis de Pedagogía en Biología, Universidad de La Serena, Chile.,85 pp. are not comparable to those described in this current Buitendijk J. & Ramanna M. 1996. Giemsa C-banded karyotypes work. Nevertheless, it is remarkable that the THL esti- of eight species of Alstroemeria L. and some of their hybrids. mated here for Leucocoryne taxa are higher than THL Ann. Bot. 78: 449–457. and MCS values previously documented for some Al- Buitendijk J., Boon F. & Ramanna M. 1997. Nuclear DNA con- stroemeria tents in twelve species of Alstroemeria L. and some of their taxa (Liliales; Alstroemeriaceae) (2n = 16, hybrids. Ann. Bot. 79: 343–353. THL range between 53.9 and 112 µm), which have been Cisternas M., Araneda L., García N. & Baeza C. 2010. Kary- described with larger genome sizes (C-values) within otypic studies in the Chilean genus Placea (Amaryllidaceae). monocots (Buitendijk & Ramanna 1996; Buitendijk Gayana Bot. 67: 198–205 et al. 1997; Sanso & Hunziker 1998; Sanso 2002). In Contreras L.C. & Gutiérrez J.R. 1991. Effects of the subterranean Leucocoryne herbivorous rodent Spalacopus cyanus on herbaceous vegeta- this sense, genome size estimation in taxa tion in arid coastal Chile. Oecologia 87: 106–109. is a pending task and may be an interesting char- Crosa O. 1988. Los cromosomas de nueve especies del género acteristic to study, additional to karyotype morphol- chileno Leucocoryne Lindley (Alliae-Alliavea). Bol. Inv. Agro. ogy. U. La República. Uruguay 17: 1–12. De La Cuadra C., Mansur L., Verdugo G. & Arriagada L. 2002. In the future, additional evidence is neccessary to Deterioro de semillas de Leucocoryne spp. en función del explain evolutionary trends within Leucocoryne genus, tiempo de almacenaje. Agricultura Técnica 62: 46–55. including cytogenetic, genomic and genetic methods Flory W. 1977. Overview of chromosomal evolution in the which have been broadly useful to study evolution Amaryllidaceae. Nucleus 20: 70–88. in polyploid complexes (Soltis et al. 2004). These Gouss J. 1949. Karyology of some South Smerican Amarylli- daceae. Plant Life (Herbertia) 5(4): 54–80. data may support a taxonomy based strongly on phy- Hayward W. 1940. Leucocoryne as a pot plant. Herbertia 7: 205. logeny. Hoffmann A. 1978. Flora silvestre de Chile. Zona central. Edi- ciones Fundación Claudio Gay, Santiago, 225 pp. Karyological study in fifteen Leucocoryne taxa (Alliaceae) 295

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Received May 5, 2011 Accepted September 16, 2011