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-Rojas4, Gina Arancio4
& Raul Moreno4
1
Instituto de Ecología y Biodiversidad (IEB), Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Ñuñoa,
Casilla 653, Santiago, Chile; e-mail: [email protected]
2
Departamento de Ciencias Biológicas Facultad de Ciencias Biológicas y Departamento de Ecología y Biodiversidad Facultad de Ecología y Recursos Naturales, Universidad Andrés Bello, Santiago, Chile
3
Escuela de Ciencias Ambientales, Facultad de Recursos Naturales, Universidad Católica de Temuco, Casilla 15-D, Temuco,
Chile
4
Departamento 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
The study of karyotype morphology among related taxa
is a fundamental aspect to understand the process of genetic variation, genome evolution and speciation, and
the taxonomic circumscriptions as has been broadly described in different plant families (Stebbins 1971; Levin
2002). Regarding Asparagales order, some reports have
discussed the process on chromosome evolution within
the larger families, such as the diverse and worldwidely
distributed Amaryllidaceae family (Sato 1938; Gouss
1949; Flory 1977; Cisternas et al. 2010; Muñoz et al.
2011).
Leucocoryne Lindl. (Alliaceae) is a Chilean endemic genus (Muñoz & Moreira 2000) important as a
component of the desert flower diversity. This genus is
distributed between 20◦ S–37◦ S, 2,500 m above sea level,
with a diversity center between 30◦ S–34◦S in a semiarid
zone (Zöellner 1972; Ravenna 1973; Hoffmannn 1978;
Ravenna 1978; Muñoz & Moreira 2000; Squeo et al.
2001; Mansur et al. 2002). Genus taxa are characterized by the presence of bulbs, linear leaves, spathe of
two lanceolate bracts, umbrella inflorescence at the base
with up to 12 flowers, 6 narrow or wide tepals, 3 or 6
stamens inserted in the tube, 3 cylindrical or flattened
* Corresponding author
c 2012 Institute of Botany, Slovak Academy of Sciences
tepals, superior ovary with several ovules, short style
and capitate stigma (Muñoz & Moreira 2005). These
plants have been described as food source for fossorial rodents, thus playing an important role as primary
producer in trophic chains of semiarid ecosystems (Reig
1970; Contreras & Gutiérrez 1991). Besides, because of
their showy flowers, these plants have been successfully
introduced into cultivation, being used as cut, pot and
garden plants (Hayward 1940; Kroon 1989; Ohkawa et
al. 1996; De la Cuadra et al. 2002; Verdugo & Texeira
2006). However, due to its extraction and habitat degradation by anthropic effect, currently some endangered
taxa are recognized whereas other taxa are insufficiently
known, and scarce information has been compiled on its
conservation (Squeo et al. 2001).
Biosystematics studies within the genus Leucocoryne have been previously documented on the basis of
morphological characteristics (Zöellner 1972; Ravenna
1998; Muñoz & Moreira 2000; Mansur et al. 2002) being currently recognized 14 taxa (Zöellner 1972). However, at present the taxonomic classification of some
populations is controvertial due to the high phenotypic
variation (Muñoz & Moreira 2000). In contrast, karyological data only for ten accurately identified Leucocoryne taxa and other six non-identified taxa are available
P. Jara-Arancio et al.
290
(Zöellner 1972; Crosa 1988; Bahamondes & Labarca
1994; Araneda et al. 2004; Mansur & Cisterna 2005),
but this background information has been scarcely used
in systematic studies. As to this genus, a complex series
of chromosome numbers has been described, including
diploid and polyploid taxa, as well as some metaphases
of hybrid specimens that have been counted (Salas &
Mansur 2004).
In order to increase the karyological antecedent
for Leucocoryne genus, in this work the karyotype
morphology of fifteen taxa is described, and seven
of them are examined here for the first time. Besides, inter-specific cytogenetic relationships are established, including karyological information previously
documented by other authors.
Material and methods
Plants of one or two accessions of fifteen taxa of Leucocoryne were collected from naturally growing populations
and their collection sites are shown in Table 1. Within some
taxa, sub-species or affinities were also examined, including
seventeen taxa in total. In the case of L. talinensis, seeds
previously stored in the laboratory were used in this study.
Nevertheless, adult plants of L. talinensis were obtained
from germinated seeds to corroborate taxonomic indentification. The voucher specimens were deposited at the ULS
herbarium (Universidad de La Serena-CHILE). The taxa
were identified according to taxonomic keys documented by
Zöellner (1972), Ravenna (1998), Muñoz & Moreira (2000),
Mansur et al. (2002) and Zuloaga et al. (2008). In the laboratory, plants were kept with their bulbs submerged in water, with constant aeration, to favor active growth of adventitious roots. After five to ten days, 5 mm-long root tips
were excised from the bulbs. The root tips were pre-treated
with a 0.05% aqueous solution of colchicine at 4 ◦C for 6
hours, fixed in ethanol-glacial acetic acid (3:1) at 4 ◦C for
24 h, and stored in 70% ethanol at 4 ◦C until chromosome
processing was done. The root tips were stained with Feulgen reaction (hydrolyzed for 7 minutes in HCl 1N at 60 ◦C,
stained with Schiff reagent for 60 minutes and washed in
sulfurous water) (Navarrete et al. 1983) and slides were
made by squashing root meristems. The determination of
karyotype parameters was carried out from metaphase images (from ten examined plants per each taxon). Measurements were made on almost five cells, and the short arms
(SA) and long arms (LA) were measured using the software
MicroMeasure 3.3 (Reeves 2001). The total relative length
(LR) of each chromosome pair was calculated and expressed
as a percentage of the total haploid set length. Chromosome
shape was determined based on the centromeric index proposed by Levan et al. (1964) (ratio short arm / total chromosome length) and karyotype was made up in order of
decreasing chromosome length. In addition, arm fundamental number (FN) was estimated for the karyotype of each
taxa where metacentric, submetacentric and subtelocentric
chromosomes showed two arms (short arm and long arm),
and telocentric chromosomes showed one arm (only long
arm) (Spotorno 1985). Intrachromosomal asymmetry index
A1 and interchromosomal asymmetry index A2 proposed by
Romero-Zarco (1986) were estimated for Leucocoryne taxa
and correlated to determine karyotype affinity. Total haploid
set length (in µm) and mean chromosome size (in µm) were
additional characteristics determined here for the karyotype
of each Leucocoryne taxa.
Results
Feulgen stained karyotype of Leucocoryne taxa are
shown in Fig. 1. Karyotypes were described for the
first time for L. appendiculata, L. conferta, L. dimorphopetala, L. macropetala, L. pauciflora, L. vittata, L.
aff. vittata (Table 1). Two plant groups based in the
chromosome numbers 2n = 10 (n = 5) and 2n = 18 (n
= 9) were identified in Leucocoryne taxa studied in this
work.
Chromosome morphology was uniform within of
each studied group, with little variations. Thus in the
taxa group with 2n = 10 three pairs were metacentric
and two pairs were subtelocentric or telocentric (haploid karyotype formula = 3m + 2st or t). In the case
of the taxa group with 2n = 18, seven pairs were metacentric and two pairs were subtelocentric or telocentric
(haploid karyotype formula = 7m + 2st or t). However,
L. ixioides shows seven metacentric, one submetacentric and one subtelocentric pair. Fundamental numbers
(FN) of the haploid set for the group n = 5 were FN =
9 and 10 whereas for the group n = 9 were FN = 16,
17, and 18.
Chromosome morphology displayed for Leucocoryne taxa of suggests the presence of moderately symmetric karyotypes within this genus with predominance
of metacentric chromosomes but with the presence of
telo- or subtelocentric chromosomes, and exceptionally submetacentric chromosomes. The correlation between A1 and A2 asymmetry index are shown in Fig. 2.
Within the group 2n = 18 with more symmetric karyotypes (group A), the A1 values ranged among 0.20 to
0.35 whereas for the group 2n = 10 with less symmetric
karyotype (group B) the A1 values ranged between 0.39
to 0.45. In the case of the A2 values of these, fall within
the range 0.17 to 0.30 for both groups (see A1 and A2
values in Table 1). Thus, the correlation between A1
and A2 indexes plotted for Leucocoryne taxa is consistent with the identification of two different groups
based in chromosome number.
Total chromosome length (THL) and mean chromosome size (MCS) for the studied taxa are shown
in Table 1. Higher total haploid set length and mean
chromosome size within the cytotype 2n = 18 were
described for L. dimorphopetala (THL = 315.63 µm,
MCS = 35.1+2.9 µm), whereas the lower values were
found in L. coquimbensis var. coquimbensis (THL=
98.72 + 0.4 µm, MCS = 11 µm). Similarly, higher total
haploid set length and mean chromosome size within
the cytotype 2n = 10 were described for L. conferta
(THL = 147.58 µm, MCS = 29.5 + 2.0 µm), whereas
lower values were found in L. aff. vitatta (THL = 77.56
+ 2.71 µm, MCS = 15.5 + 0.5 µm). Values of the
other studied Leucocoryne taxa are within the range
described for their respective 2n number.
Secondary constriction (SC) was observed on the
short arms of the chromosome pair 5 in L. macropetala,
L. vittata and L. aff. vittata. In L. alliacea, L. appendiculata, L. coquimbensis, L. dimorphopetala, L. ixioides
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.
THL ± SD
(in µm)
MCS ± SD A1 ± SD A2 ± SD
(in µm)
Index
Index
Taxa
Collection sites
(Latitude – Longitude; Altitude
m a.s.l.)
2n HKF
FN
L. alliacea Miers ex
Lindl.
L. alliacea Miers ex
Lindl.
L. angustipetala Gay
L. angustipetala Gay
L. appendiculata Phil.
L. appendiculata
Phil. *
L. conferta Zoëllner *
Farellones (33◦ 20 S – 70◦ 19 W;
2700)
León muerto (29◦ 20 S – 70◦ 39 W;
2770)
Tofo (28◦ 58 S – 70◦ 56 W; 730)
Domeyko (28◦ 59 S – 70◦ 56 W; 759)
Iquique (20◦ 13 S – 70◦ 09 W; 10)
Pan de Azúcar (26◦ 08 S – 70◦ 35 W;
510)
Cuesta Cavilolén (31◦ 46 S –
71◦ 19 W; 700)
Tongoy (30◦ 15 S – 71◦ 30 W; 40)
18 7m, 2st
18 196.22 ± 13.87 21.8 ± 1.5 0.27 ± 0.0 0.17 ± 0.0
18 7m, 2st
18
10
10
18
18
2st
2st
2st
2st
10 117.01 ± 14.94 23.4 ± 3.0 0.40 ± 0.0 0.18 ± 0.0
10
18 203.58 ± 8.19 22.6 ± 0.9 0.29 ± 0.0 0.23 ± 0.0
18
10 3m, 2st
10 147.58 ± 9.94 29.5 ± 2.0 0.41 ± 0.0 0.30 ± 0.0
18 7m, 1st, 1t
17
Panul (30◦ 00 S – 71◦ 24 W; 100)
18 7m, 1st, 1t
17
Panul (30◦ 00 S – 71◦ 24 W; 100)
18 7m, 1sm, 1st 18 146.56 ± 36.37 16.3 ± 4.1 0.25 ± 0.0 0.17 ± 0.0
Juan Soldado (29◦ 40 S – 71◦ 18 W;
300)
Freirina (28◦ 31 S – 70◦ 04 W; 100)
18 7m, 1sm, 1st 18
18 7m, 2st
18 315.63 ± 26.02 35.1 ± 2.9 0.20 ± 0.0 0.21 ± 0.0
Sauce Pérez (28◦ 40 S – 71◦ 06 W;
607)
El Manzano (33◦ 41 S – 71◦ 09 W;
175)
Quebrada las Vacas (32◦ 42 S –
71◦ 13 W; 200)
Cerro Grande (29◦ 56 S – 71◦ 13 W;
520)
Los Burros (27◦ 54 S – 71◦ 07 W; 50)
Los Burros (27◦ 54 S – 71◦ 07 W; 50)
Paposo (25◦ 00 S – 70◦ 28 W; 50)
Viña del Mar (33◦ 31 S – 71◦ 34 W;
150)
Valparaíso (33◦ 02 S – 71◦ 38 W;
150)
Camino a Rapel (34◦ 05 S –
71◦ 32 W; 70)
Guanaqueros (30◦ 12 S – 71◦ 26 W;
100)
Quebrada las Vacas (32◦ 42 S –
71◦ 13 W; 200)
Colecta planta tipo (30◦ 49 S –
71◦ 33 W; 355)
Amolana (31◦ 12 S – 71◦ 37 W; 200)
Alcones (30◦ 48 S – 71◦ 33 W; 250)
Chigualoco (31◦ 46 S – 71◦ 30 W; 15)
Los Vilos (31◦ 54 S – 71◦ 31 W; 15)
Las Palmas (31◦ 16 S – 71◦ 35 W;
248)
18 7m, 2st
18
L. coquimbensis Phil.
var. coquimbensis
L. coquimbensis Phil.
var. coquimbensis
L. coquimbensis Phil.
var. alba Zoëllner
L. coquimbensis Phil.
var. alba Zoëllner
L. dimorphopetala
(Gay) Rav. *
L. dimorphopetala
(Gay) Rav.
L. ixioides (Hook)
Lindl.
L. ixioides (Hook)
Lindl.
L. macropetala Phil. *
L.
L.
L.
L.
macropetala Phil.
narcissoides Phil.
narcissoides Phil.
foetida Phil. *
L. foetida Phil.
L. pauciflora Phil. *
L. purpurea Gay
L. purpurea Gay
L. talinensis Mansur
L.
L.
L.
L.
L.
violacescens Phil.
violacescens Phil.
vittata Rav. *
vittata Rav.
aff. vittata Rav. *
3m,
3m,
7m,
7m,
98.72 ± 0.40
11.0 ± 0.0 0.30 ± 0.0 0.18 ± 0.0
18 7m, 1sm, 1st 18 274.39 ± 22.18 30.5 ± 2,5 0.28 ± 0.0 0.17 ± 0.0
18 7m, 1sm, 1st 18
97.98 ± 7.05
19.6 ± 1.4 0.40 ± 0.0 0.19 ± 0.0
10 3m, 2st
10
10
18
18
18
10
17 99.29 ± 5.75 11.0 ± 0.6 0.29 ± 0.0 0.23 ± 0.0
17
16 254.37 ± 7.95 28.3 ± 0.9 0.31 ± 0.0 0.19 ± 0.0
3m,
7m,
7m,
7m,
2st
1st, 1t
1st, 1t
2t
18 7m, 2t
16
18 7m, 2st
18 182.23 ± 13.83 20.20 ± 1.5 0.29 ± 0.0 0.18 ± 0.0
10 3m, 1st, 1t
9
10 3m, 1st, 1t
9
18 7m, 1st, 1t
17 240.96 ± 11.69 26.8 ± 1.3 0.35 ± 0.0 0.23 ± 0.0
18
18
10
10
10
arms of the chromosome pair 9 (Fig. 1). SC was not
observed in the remaining taxa.
Discussion
Leucocoryne genus can be characterized as having two
chromosome numbers 2n = 10 and 2n = 18 as has
been described in previous works (Crosa 1988; Baha-
7m,
7m,
3m,
3m,
3m,
88.14 ± 10.71 17.6 ± 2.1 0.45 ± 0.0 0.17 ± 0.0
1sm, 1st 18 283.91 ± 22.23 31.5 ± 2.5 0.28 ± 0.0 0.20 ± 0.0
1sm, 1st 18
2st
10 106.13 ± 8.68 21.2 + 1.7 0.39 ± 0.0 0.17 ± 0.0
2st
10
2st
10 77.56 ± 2.71 15.5 ± 0.5 0.40 ± 0.0 0.21 ± 0.0
mondes & Abarca 1994; Araneda et al. 2004). However, this series of 2n number is extended to include
some taxa described with 2n = 14 such as L. coquimbensis var. alba (Araneda et al. 2004) and the natural
hybrid L. coquimbensis × L. purpurea (Salas & Mansur
2004). Nevertheless, inconsistencies are observed in L.
coquimbensis var. alba due to the fact that two different 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. Scale bar = 10 µm.
Table 2. Cytogenetic data for Leucocoryne species previously studied by other authors.
Taxa
2n
Haploid karyotype formula
Reference
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
L.
18
18
18
18
18
18
18
14
10
10
10
10
10
10
7sm + 2st
7m + 1st + 1t
7m + 1st + 1t
7m + 1st + 1t
7m + 1st + 1t
7m + 2st
7m + 1st + 1t
5m + 1st + 1t
3m + 1st + 1t
3m + 1st + 1t
3m + 1st + 1t
3m + 1sm + 1t
3m + 1sm + 1t
3m + 1sm + 1t
Zoëllner (1972)
Crosa (1988)
Araneda et al. (2004)
Araneda et al. (2004)
Araneda et al. (2004)
Crosa (1988)
Mansur & Cisterna (2005)
Araneda et al. (2004)
Crosa (1988)
Crosa (1988)
Araneda et al. (2004)
Crosa (1988)
Crosa (1988)
Crosa (1988)
ixioides
ixioides
ixioides
coquimbensis var. coquimbensis
narcissoides
violascecens
talinensis
coquimbensis var. alba
odorata (L. foetida)
purpurea
purpurea
alliacea
alliacea
angustipetala
in this work and 2n = 14 in the study of Araneda et al.
(2004). Similarly, L. alliacea has been described previously with a number 2n = 10 (Crosa 1988), whereas in
this work a number 2n = 18 was described (Table 2).
In all cases, taxa identification should be revised using
the voucher specimens deposited in indexed herbarium,
thus corroborating taxonomic identification and clarifying the ambiguous chromosome numbers reported so
far. In our work, collected specimens of all taxa were
checked by comparison with voucher specimens stored
in the indexed ULS herbarium.
Karyotype morphology reported in this work for
Leucocoryne taxa shows resemblances with those described for other taxa of the genus previously examined (Zöellner 1972; Crosa 1988; Bahamondes & Abarca
1994; Araneda et al. 2004; Salas & Mansur 2004). Studied taxa have been adscribed to the groups 2n = 10
and 2n = 18 exhibiting predominance of metacentric
chromosomes (between 60 and 77%) and low quantity
of subtelocentric or telocentric chromosomes (between
22 to 40%) within their respective karyotype formulas
(Fig. 1, Table 1). Most common haploid karyotype formula for the studied taxa belonging to the cytotype
n = 5 was 3m + 2st, while for the cytotype n = 9
was 7m + 2st. It is remarkable that although chromosome number varies between the two mentioned groups,
similarities in gross chromosome morphology among
their karyotypes were observed. Nevertheless, an increment in the number of metacentric chromosomes in the
2n = 18 cytotype is related to lower values of intrachromosomal asymmetry index A1 in these taxa (more
symmetry in group A) regarding to 2n = 10 (less symmetry in group B), although both groups are cohesive
when comparing the interchromosomal asymmetry index A2, whose values are superimposed (between 0.17
and 0.30) (Fig. 2).
All available karyotype data for Leucocoryne are
consistent with Crosa’s (1988) hypothesis, who suggested that cytotype 2n = 10 is diploid and perhaps
ancestral, whereas cytotype 2n = 18 is tetraploid (likely
auto-tetraploid) but with an additional chromosome fusion being probably a derived status. The fundamental
numbers (FN) reported here for both groups of Leucocoryne taxa also support this hypothesis (Table 1).
Then, as an explanation of all this framework, it is possible to suggest that chromosome differentiation in Leu-
P. Jara-Arancio et al.
294
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 increment in number of metacentric chromosomes from 3m
to 7m via genome duplication and by a centric fusion of
telocentric chromosomes from a base cytotype n = 5. In
addition, the number of st-t chromosomes and values of
interchromosomal length are similar among karyotypes
of the taxa (Fig. 2).
Another interesting feature observed in the chromosomes of Leucocoryne taxa studied here are the
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
been described, and only absolute chromosome size
(in µm) for largest and smallest pairs of karyotype
are given (Araneda & Mansur 2004; Salas & Mansur
2004). However, due to different times of pretreatment
with colchicine that affects the chromosome condensation and shorten chromosome metaphases, those values
are not comparable to those described in this current
work. Nevertheless, it is remarkable that the THL estimated here for Leucocoryne taxa are higher than THL
and MCS values previously documented for some Alstroemeria taxa (Liliales; Alstroemeriaceae) (2n = 16,
THL range between 53.9 and 112 µm), which have been
described with larger genome sizes (C-values) within
monocots (Buitendijk & Ramanna 1996; Buitendijk
et al. 1997; Sanso & Hunziker 1998; Sanso 2002). In
this sense, genome size estimation in Leucocoryne taxa
is a pending task and may be an interesting characteristic to study, additional to karyotype morphology.
In the future, additional evidence is neccessary to
explain evolutionary trends within Leucocoryne genus,
including cytogenetic, genomic and genetic methods
which have been broadly useful to study evolution
in polyploid complexes (Soltis et al. 2004). These
data may support a taxonomy based strongly on phylogeny.
Acknowledgements
The authors acknowledge CONAF and ULS, CONC and
SGO herbariums for their valuable help, as well as Mélica
Muñoz’s critical support. We would also thank the Departamento de Biología-ULS, and CONICYT for granting a
fellowship to P. Jara-Arancio. This is a contribution to
the research program of Senda Darwin Biological Station,
Chiloé, Chile. Postdoctoral fellowship from IEB, ICM P05–
002, PFB-23.
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Received May 5, 2011
Accepted September 16, 2011