**3. Taxonomy and distribution of the genus** *Alstroemeria* **in Chile**

#### **3.1. General morphology**

and comprises about 200 species distributed in the genera *Alstroemeria* L., *Bomarea* Mirb., *Leontochir* Phil. and *Schickendantzia* Pax [4]. Some authors recognized *Taltalia* Ehr. Bayer as an independent genus [5, 6] while others included *Taltalia* and *Schickendantzia* in *Alstroemeria* [7]. Genera have been classified in two subfamilies: (a) Subfamily: Luzuriagoideae with *Drymophila* R. Br. and *Luzuriaga* Ruiz & Pav. and (b) Subfamily: Alstroemerioideae with *Alstroemeria* and *Bomarea* (incl. *Leontochir*) [3, 8] or treated Luzuriagoideae as a separate family Luzuriagaceae [3]. *Alstroemeria* comprises about 80 species endemic to South America (southern South America and Eastern Brazil); *Bomarea* includes 120 species in Central and South America; *Luzuriaga* comprises 4 species (3 species in Chile, 1 in New Zealand) and *Drymophila* 1 species from Australia and Tasmania [8]. Phylogenetic studies using morphology and DNA sequences (*rps16*, *rbcL*) recognized *Alstroemeria* as monophyletic and certainly different from *Leontochir* and *Bomarea*; moreover, three subclades of *Alstroemeria* have been recognized that correspond to northern Chile, central Chile and Brazil [9]. *Alstroemeria* was established by Linnaeus in 1762 in honor to the Swedish botanist Claus von Alströmer [10]. It has its boreal distribution in Venezuela (3°N) and its austral limit in the Patagonia of Chile and Argentina, with two main distribution centers in the continent: Brazil (and adjacent areas of Paraguay and Argentina) and Chile (and the adjacent countries Peru, Bolivia and Argentina) [11–15]. In Chile, *Alstroemeria* represents 1 of the most diverse genera of vascular monocotyledons, comprising more than 50 recognized or accepted taxa (36 species, 11 subspecies and 10 varieties) from which about ca. 82% are endemic to the Mediterranean zone of central Chile [14]. Due to the beauty of their flowers, Chilean species of *Alstroemeria*, locally known as "astromelias or lirios del campo" are appreciated all over the world as ornamental plants [16–18]. Many hybrids and cultivars have been developed in several countries, such as, The Netherlands, England, United States and Japan [14]. The taxonomy of the genus is very complex due to the great variability both in vegetative and floral characters [19]. Moreover, several taxa have been recently described (e.g., *A. werdermannii* var. *flavicans* [20], *A. philippi* var. *albicans* [14], *A. philippi* subsp. *adrianae* [21], *A. hookeri* subsp. *sansebastiana* [22], *A. marticorenae* [19], *A. traudliae* [23]) or nomenclatural changes have been made, affecting the rank status of many taxa. Due to its geographical isolation, Chile contains a unique flora which includes an extraordinary number of endemic plants. Furthermore, the area between the Regions of Atacama and Biobío comprises about 60% of the vascular species of the Chilean flora, with nearly 50% endemic to Chile [24]. In this area, identified as a hotspot of biodiversity [25], lives most of the Chilean species of *Alstroemeria*. This area harbors most of the Chilean population, and is characterized by a strong disturbance triggered mainly by agriculture, industry and forestry. In this chapter, we reviewed the taxonomic literature to make an updated checklist of the species growing in Chile and their synonyms. In addition, we construct distribution maps based on the literature as well as on the database of the Herbarium of the University of Concepción (CONC). Recent studies integrating different source of evidence, such as morphometry, cytogenetic, colorimetry and molecular data for better taxonomic species delimitation are discussed.

The database of the Herbarium of the University of Concepción (CONC) was used to construct preliminary lists of species and the geographic distribution of each taxon. This database contains the following fields: (1) Taxon name; (2) Collector's name; (3) Collector's number; (4) Latitude; (5) Longitude; (6) Elevation; (7) Administrative region; (8) Locality; (9) Collection

**2. Methods**

230 Selected Studies in Biodiversity

*Alstroemeria* comprises mostly perennial species. In 1998, Bayer [5] established the monotypic genus *Taltalia* to separate the annual *A. graminea* Phil., endemic to northern Chile, from *Alstroemeria*. Perennial species have cylindrical rhizomes, from which two kinds of roots born: thin roots and thick roots which contain starch (**Figure 1A**); *A. ligtu* (locally known as "liuto") was used by indigenous people (Mapuches or Araucanos) to produce starch from the thick roots. According to Molina [26], the farmers made from the roots of this plant "a white, light, nutritious and so healthy flour that they usually gave it to the sick persons…" [26]. Aerial stems are erect or decumbent. The leaves are often resupinated, that is, twisted from the petiole or the leaf blade so the lower surface becomes functionally the upper surface; sometimes the leaves form basal rosettes; leaf blades thin or thick, sometimes with papillae; the blade varies in shape from linear to elliptic or ovate; fertile stems usually have reduced leaves but sterile stems have well developed leaves (**Figure 1B**). The flowers are slightly zygomorphic (**Figure 1C**-**D**), with six free tepals in two verticils; the three outer tepals are similar in shape and color; the inner three tepals are differentiated in two upper inner tepals and one lower inner tepal. Upper inner tepals with colored lines (nectar guides) on a lighter background (**Figure 1D**). Stamens 3: the ovary is inferior, 3-carpellate, 3-loculate. The fruit is a six-ribbed loculicidal capsule (**Figure 1C**) with explosive dehiscence, with numerous globose seeds (**Figure 2**).

#### **3.2. Colorimetric studies in Chilean** *Alstroemeria*

In *Alstroemeria*, the flowers varies in color from white to yellow, pink, red, purple and violet according to the species [11, 14]; color is regulated by several pigments including anthocyaninlike 6-hydroxydelphinidine 3-rutinoside, 6-hydroxycyanidin 3-rutinoside, delphinidin 3-malonylglucoside among others, carotenoids and flavonoids [27]. In some groups of plants, as occurs in *Alstroemeria*, it is possible that the taxonomic characters traditionally used do not have sufficient discriminant power to differentiate very close species or varieties within a species complex. In such cases, it may be useful to have characteristics that pose a new perspective on the problem. It has been shown that the color of the corolla, objectively measured, had high taxonomic value when the traditional characters were less informative to distinguish cryptic taxa [28]. In *Alstroemeria*, the color of the flower has often been used in keys and descriptions [11, 14, 15, 22, 23], however, most of the time, the described color corresponds to a subjective perception of the same by the human eye. The color of the flowers is taxonomically significant in *Alstroemeria* [14, 29, 30]; the

**Figure 1.** Morphology of the roots, rhizomes, leaves, flowers and fruits of *Alstroemeria*. (A) Roots and rhizomes of *A. x chrysantha*; (B) sterile leaves in fertile stems and basal rosettes of *A. magenta*; (C) fruits (capsules) of *A. hookeri* subsp. *recumbens*; (D) flower of *A. x chrysantha*. Photos A and D by V.L. Finot; photos B and C by C.M. Baeza.

**Figure 2.** Seeds of *Alstroemeria* as seen with scanning electron microscope (SEM). (A) *Alstroemeria presliana* subsp. *australis*. (B) *Alstroemeria magenta*. (C) *Alstroemeria ligtu* subsp. *simsii*. (D). *Alstroemeria magenta*, surface details. (E) and

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(F) *Alstroemeria x chrysantha*.

Towards an Integrative Taxonomy of the Genus *Alstroemeria* (Alstroemeriaceae) in Chile:… http://dx.doi.org/10.5772/intechopen.71823 233

**Figure 2.** Seeds of *Alstroemeria* as seen with scanning electron microscope (SEM). (A) *Alstroemeria presliana* subsp. *australis*. (B) *Alstroemeria magenta*. (C) *Alstroemeria ligtu* subsp. *simsii*. (D). *Alstroemeria magenta*, surface details. (E) and (F) *Alstroemeria x chrysantha*.

**Figure 1.** Morphology of the roots, rhizomes, leaves, flowers and fruits of *Alstroemeria*. (A) Roots and rhizomes of *A. x chrysantha*; (B) sterile leaves in fertile stems and basal rosettes of *A. magenta*; (C) fruits (capsules) of *A. hookeri* subsp.

*recumbens*; (D) flower of *A. x chrysantha*. Photos A and D by V.L. Finot; photos B and C by C.M. Baeza.

232 Selected Studies in Biodiversity

outer tepals and the lower inner tepals are similar in shape and color but the upper internal tepals are different, showing unique patterns of maculae (actually nectar guides) that are species specific; however, some species showed variability in the background color [30], because it depends on several ecological factors such as temperature and pH [31]. In order to describe objectively the color (mainly the background color) of the different tepals (external, upper internal and lower internal), the CIElab system [32] has been used in some species complexes of Chilean *Alstroemeria* [29, 30] (**Figure 3**). The CIE (Commission Internationale de l'Eclairage) color system uses three coordinates to locate a given color in the color space. The spectrophotometer registers the reflected wavelengths as numerical values (spectral curve) from which the coordinates that place a given color in the color space are calculated. The color expressed in the CIELab scale uses the Cartesian coordinates L\*, a\* and b\*. L\* expresses the luminosity, a\* denotes the green/red value and b\* the blue/yellow value. The degree of luminosity of L\* determines that a color appears lighter or darker and is expressed in a scale from 0 (black or total absorption) to 100 (white). The axis a\* moves from negative values (green) to positive values (red) while axis b\* moves from blue (negative values) to yellow (positive values). The color expressed in the CIELCh scale uses polar coordinates (L\*, C\*, h°), derived from the CIELab scale. C\* denotes chroma (saturation, intensity) and h° denotes hue, expressed as angular measures. Chroma is the distance of the color from the axes a\* and b\* of L\*, calculated as (a\*2 + b\*2 ) 1/2 and represents the color saturation; the hue, h° is calculated as arctg (b\*/a\*). In *A. magnifica* complex, the colorimetric study of the flower helped to elucidate the taxonomic position of *A. pulchra* var. *maxima*. This taxon, originally described by Philippi in 1864 [33], was transferred by Bayer in 1987 to *A. magnifica* with the subspecific rank (*A. magnifica* subsp. *maxima*) [11].

The colorimetric differences between *A. magnifica* and *A. pulchra* as shown in the reflectance spectra were due mainly to the parameters a\* and b\* indicating that *A. magnifica* have tepals comparatively more intense violet than those of *A. pulchra* var. *maxima*. Our results suggest that the color of the flowers can be used as a new taxonomic character in *Alstroemeria* and that var. *maxima* probably belongs to *A. pulchra* as originally proposed and not to *A. magnifica* [29]. Colorimetric studies were carried out also in *A. presliana* [30]. This species comprises two subspecies: subsp. *presliana* and subsp. *australis*. *Alstroemeria presliana* subsp. *presliana* grows in Chile (Regions of Maule and Biobío) and Argentina (Neuqén) [34]; subsp. *australis* in endemic to Chile (Regions of Biobío and Araucanía) [11, 14].

Although the color of the flowers is one of the most important characters to distinguish the subspecies [11], there is a huge variability in color in the flowers of both subspecies. Differences in the spectral reflectance curves were detected between 440 and 540 nm and between 660 and 700 nm in the outer and lower inner tepals. Upper internal tepals differ mainly between 640 and 700 nm. The color measured in the CIELab space is related to the content of anthocyanins so that the flowers containing delphinidin-3-glucosides take on a more blue hue than those containing exclusively cyanidin-3-glucosides [35]. The presence of delphinidin-3-glucosides detected in subsp. *presliana* but not in subsp. *australis* [27] could explain the bluer hue observed in subsp. *presliana* in comparison with subsp. *australis* and the difference observed in the parameter b\*, which takes negative values in subsp. *presliana* and positive values in subsp. *australis* both in outer and lower inner tepals. On the other hand, in the upper inner tepals, b\* was positive (yellow), reaching higher values in subsp. australis and therefore denoting a more intense yellow color in subsp. *australis* than in subsp. *presliana* [30].

**3.3. Geographical distribution**

(Baeza 4376). Photos A and B by C. Baeza; C and D by V. Finot.

In Chile, there are 38 species of *Alstroemeria* and 16 infraspecific taxa (8 subspecies and 8 varieties) and 1 nothospecies (*A. x chrysantha*) [23]. Nevertheless, more than 116 species have been described, most of which are considered synonyms or are names of uncertain application because there is no original material (types) in herbaria (see Checklist below). The description of such high number of taxa can be explained by the extent of morphological variation, especially of the flowers, which harbors most of the characters useful to taxonomy,

**Figure 3.** Flowers of some species of Chilean *Alstroemeria*. (A) *Alstroemeria pulchra*, Maule region, Talca, Río Claro (Baeza 4393). (B) *Alstroemeria ligtu* subsp. *simsii*, Maule region, Talca, Río Claro (Baeza 4395). (C) *Alstroemeria magnifica* var. *sierrae*, Coquimbo region, Caleta Hornos (Baeza 4375). (D) *Alstroemeria x chrysantha*, Coquimbo region, Huanaqueros

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Towards an Integrative Taxonomy of the Genus *Alstroemeria* (Alstroemeriaceae) in Chile:… http://dx.doi.org/10.5772/intechopen.71823 235

**Figure 3.** Flowers of some species of Chilean *Alstroemeria*. (A) *Alstroemeria pulchra*, Maule region, Talca, Río Claro (Baeza 4393). (B) *Alstroemeria ligtu* subsp. *simsii*, Maule region, Talca, Río Claro (Baeza 4395). (C) *Alstroemeria magnifica* var. *sierrae*, Coquimbo region, Caleta Hornos (Baeza 4375). (D) *Alstroemeria x chrysantha*, Coquimbo region, Huanaqueros (Baeza 4376). Photos A and B by C. Baeza; C and D by V. Finot.

#### **3.3. Geographical distribution**

outer tepals and the lower inner tepals are similar in shape and color but the upper internal tepals are different, showing unique patterns of maculae (actually nectar guides) that are species specific; however, some species showed variability in the background color [30], because it depends on several ecological factors such as temperature and pH [31]. In order to describe objectively the color (mainly the background color) of the different tepals (external, upper internal and lower internal), the CIElab system [32] has been used in some species complexes of Chilean *Alstroemeria* [29, 30] (**Figure 3**). The CIE (Commission Internationale de l'Eclairage) color system uses three coordinates to locate a given color in the color space. The spectrophotometer registers the reflected wavelengths as numerical values (spectral curve) from which the coordinates that place a given color in the color space are calculated. The color expressed in the CIELab scale uses the Cartesian coordinates L\*, a\* and b\*. L\* expresses the luminosity, a\* denotes the green/red value and b\* the blue/yellow value. The degree of luminosity of L\* determines that a color appears lighter or darker and is expressed in a scale from 0 (black or total absorption) to 100 (white). The axis a\* moves from negative values (green) to positive values (red) while axis b\* moves from blue (negative values) to yellow (positive values). The color expressed in the CIELCh scale uses polar coordinates (L\*, C\*, h°), derived from the CIELab scale. C\* denotes chroma (saturation, intensity) and h° denotes hue, expressed as angular measures. Chroma is the distance of the color from the axes a\* and b\* of L\*, calculated as

1/2 and represents the color saturation; the hue, h° is calculated as arctg (b\*/a\*). In *A. mag-*

*nifica* complex, the colorimetric study of the flower helped to elucidate the taxonomic position of *A. pulchra* var. *maxima*. This taxon, originally described by Philippi in 1864 [33], was transferred by

The colorimetric differences between *A. magnifica* and *A. pulchra* as shown in the reflectance spectra were due mainly to the parameters a\* and b\* indicating that *A. magnifica* have tepals comparatively more intense violet than those of *A. pulchra* var. *maxima*. Our results suggest that the color of the flowers can be used as a new taxonomic character in *Alstroemeria* and that var. *maxima* probably belongs to *A. pulchra* as originally proposed and not to *A. magnifica* [29]. Colorimetric studies were carried out also in *A. presliana* [30]. This species comprises two subspecies: subsp. *presliana* and subsp. *australis*. *Alstroemeria presliana* subsp. *presliana* grows in Chile (Regions of Maule and Biobío) and Argentina (Neuqén) [34]; subsp. *australis* in endemic

Although the color of the flowers is one of the most important characters to distinguish the subspecies [11], there is a huge variability in color in the flowers of both subspecies. Differences in the spectral reflectance curves were detected between 440 and 540 nm and between 660 and 700 nm in the outer and lower inner tepals. Upper internal tepals differ mainly between 640 and 700 nm. The color measured in the CIELab space is related to the content of anthocyanins so that the flowers containing delphinidin-3-glucosides take on a more blue hue than those containing exclusively cyanidin-3-glucosides [35]. The presence of delphinidin-3-glucosides detected in subsp. *presliana* but not in subsp. *australis* [27] could explain the bluer hue observed in subsp. *presliana* in comparison with subsp. *australis* and the difference observed in the parameter b\*, which takes negative values in subsp. *presliana* and positive values in subsp. *australis* both in outer and lower inner tepals. On the other hand, in the upper inner tepals, b\* was positive (yellow), reaching higher values in subsp. australis and therefore denoting a

more intense yellow color in subsp. *australis* than in subsp. *presliana* [30].

Bayer in 1987 to *A. magnifica* with the subspecific rank (*A. magnifica* subsp. *maxima*) [11].

to Chile (Regions of Biobío and Araucanía) [11, 14].

(a\*2 + b\*2 )

234 Selected Studies in Biodiversity

In Chile, there are 38 species of *Alstroemeria* and 16 infraspecific taxa (8 subspecies and 8 varieties) and 1 nothospecies (*A. x chrysantha*) [23]. Nevertheless, more than 116 species have been described, most of which are considered synonyms or are names of uncertain application because there is no original material (types) in herbaria (see Checklist below). The description of such high number of taxa can be explained by the extent of morphological variation, especially of the flowers, which harbors most of the characters useful to taxonomy, and because microevolutionary processes are still active so that species are not yet completely separated. Chile shares only few taxa with its neighboring countries (Argentina, Bolivia and Peru). In Peru, five species have been mentioned [14, 36, 37], one of which is also present in Chile: *A. violacea*. In Bolivia, three species are found, one of which growth in Chile: *A. aurea* ([38], under *A. aurantiaca*). In Argentina, there are 10 species [34], 5 shared with Chile: *A. andina* var. *venustula*, *A. aurea*, *A. patagonica*, *A. presliana* subsp. *presliana*, *A. pseudospathulata*. Thus, more than 88% of the genus is represented by taxa endemic to Chile (**Figure 4**). In Chile, *Alstroemeria* spreads from 20°S (Tarapacá Region) to 53°S (Magallanes Region) [14, 23]. Most taxa have a very restricted distribution in Chile (**Table 1**). The vast majority of the species are distributed in north (Tarapacá-Coquimbo) and central (Valparaíso-Biobío) Chile (**Figure 5**); only six species growth in southern Chile (Araucanía-Magallanes). The most boreal taxa are *A. lutea* and *A. violacea* that reach the Region of Tarapacá (20°S) in northern Chile. *Alstroemeria lutea* is restricted to the coast of the Tarapacá Region (Iquique) whereas *A. violacea* extends southern to 28°S in the Atacama Region; this species if known also from Peru (Arequipa) [14, 23, 37]. The regions with the largest number of taxa are Atacama (14 taxa), Coquimbo (26), Valparaíso (19) and Metropolitan (14) (**Figure 6**). The number of taxa decreases abruptly southern the Maule Region where 12 taxa are found; in Los Ríos and Los Lagos, only one

**No.**

1 2a 2b 3a 3b

4 5 6 7 8 9 10 11 12a 12b 12c 12d

13 14 15a 15b 15c

16

*A. lutea*

*A. ligtu* subsp. *splendens*

x

*A. ligtu* subsp. *simsii*

*A. ligtu* subsp. *ligtu*

*A. leporina*

*A. kingii*

*A. hookeri* subsp. *sansebastiana*

x

x

x

x

> x

> x

> x

x

x

x

x

x

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237

*A. hookeri* subsp. *recumbens*

*A. hookeri* subsp. *maculata*

x

x

x

x

*A. hookeri* subsp. *hookeri*

*A. graminea*

*A. garaventae*

*A. exerens*

*A. diluta*

*A. cummingiana*

*A. crispata*

*A. citrina*

*A. aurea*

*A. angustifolia* var. *velutina*

*A. angustifolia* var. *angustifolia*

*A. andina* var. *venustula*

*A. andina* var. *andina*

x

x

x

x

> x

x

> x

x

x

x

x

x x

> x

> > x

x

x

x

x

Towards an Integrative Taxonomy of the Genus *Alstroemeria* (Alstroemeriaceae) in Chile:…

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

*A. achirae*

**Taxa**

**AYP**

**TAR**

**ANT**

**ATA**

**COQ**

**VAL**

**RME**

**LBO**

**MAU**

x

**NUB**

**BIO**

**ARA**

**LRI**

**LLA**

**AYS**

**MAG**

**Figure 4.** Species diversity of *Alstroemeria* in South America.

and because microevolutionary processes are still active so that species are not yet completely separated. Chile shares only few taxa with its neighboring countries (Argentina, Bolivia and Peru). In Peru, five species have been mentioned [14, 36, 37], one of which is also present in Chile: *A. violacea*. In Bolivia, three species are found, one of which growth in Chile: *A. aurea* ([38], under *A. aurantiaca*). In Argentina, there are 10 species [34], 5 shared with Chile: *A. andina* var. *venustula*, *A. aurea*, *A. patagonica*, *A. presliana* subsp. *presliana*, *A. pseudospathulata*. Thus, more than 88% of the genus is represented by taxa endemic to Chile (**Figure 4**). In Chile, *Alstroemeria* spreads from 20°S (Tarapacá Region) to 53°S (Magallanes Region) [14, 23]. Most taxa have a very restricted distribution in Chile (**Table 1**). The vast majority of the species are distributed in north (Tarapacá-Coquimbo) and central (Valparaíso-Biobío) Chile (**Figure 5**); only six species growth in southern Chile (Araucanía-Magallanes). The most boreal taxa are *A. lutea* and *A. violacea* that reach the Region of Tarapacá (20°S) in northern Chile. *Alstroemeria lutea* is restricted to the coast of the Tarapacá Region (Iquique) whereas *A. violacea* extends southern to 28°S in the Atacama Region; this species if known also from Peru (Arequipa) [14, 23, 37]. The regions with the largest number of taxa are Atacama (14 taxa), Coquimbo (26), Valparaíso (19) and Metropolitan (14) (**Figure 6**). The number of taxa decreases abruptly southern the Maule Region where 12 taxa are found; in Los Ríos and Los Lagos, only one

236 Selected Studies in Biodiversity

**Figure 4.** Species diversity of *Alstroemeria* in South America.


**No.**

32 33 34 35 36 37a 37b

38 39 AYP =

RME =

LLA =

**Table 1.**

Arica-Parinacota Region; TAR

Metropolitan Region; LBO

Los Lagos Region; AYS

 =

Aysén Region; MAG

 = Presence (x) of the accepted species of *Alstroemeria* in the administrative political regions of Chile.

 =

 =

Tarapacá Region; ANT

O'Higgins Region; MAU

 =

Maule Region; NUB

 = Magallanes and Antártica Chilena Region.

 =

Antofagasta Region; ATA

 = Ñuble Region; BIO

 =

Atacama Region; COQ

 = Biobío Region; ARA

 =

Coquimbo Region; VAL

 = Araucanía Region; LRI

 =

Los Ríos Region;

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Valparaíso Region;

*A. zoellneri*

*A. x chrysantha*

*A. werdermannii* subsp. *werdermannii*

*A. werdermannii* subsp. *flavicans*

*A. violacea*

x

x

x

x x x

x

x

x

x

x

x

*A. versicolor*

*A. umbellata*

*A. traudliae*

*A. spathulata*

**Taxa**

**AYP**

**TAR**

**ANT**

**ATA**

**COQ**

x x

x x

x

x

x

x

x

x

x

x

**VAL**

**RME**

**LBO**

**MAU**

**NUB**

**BIO**

**ARA**

**LRI**

**LLA**

**AYS**

**MAG**


**No.**

17 18a 18b 18c

19 20 21 22 23 24 25a 25b 25c

26 27a 27b

28 29a 29b 29c

30 31a 31b

*A. schizanthoides* var. *schizanthoides*

*A. schizanthoides* var. *alba*

*A. revoluta*

*A. pulchra* var. *maxima*

*A. pulchra* subsp. *pulchra*

x x

x x

x

x

x

x

x

x

x

x

x

x

x

x

x

x

*A. pulchra* subsp. *lavandulacea*

*A. pseudospathulata*

*A. presliana* subsp. *presliana*

*A. presliana* subsp. *australis*

*A. polyphylla*

*A. philippii* var. *philippii*

*A. philippii* var. *albicans*

*A. philippi* subsp. *adrianae*

*A. pelegrina*

*A. patagonica*

*A. parvula*

*A. pallida*

*A. mollensis*

*A. marticorenae*

*A. magnifica* var. *tofoensis*

*A. magnifica* var. *sierrae*

*A. magnifica* var. *magnifica*

*A. magenta*

**Taxa**

**AYP**

**TAR**

**ANT**

**ATA**

**COQ**

x x

x

x

x

x

x x

> x

> > x

x

x x

x

x x

x

x

x

x

x

x

x

x

x

x

x

x

**VAL**

**RME**

**LBO**

**MAU**

**NUB**

**BIO**

**ARA**

**LRI**

**LLA**

**AYS**

**MAG**

238 Selected Studies in Biodiversity

RME Metropolitan Region; LBO O'Higgins Region; MAU Maule Region; NUB Ñuble Region; BIO Biobío Region; ARA Araucanía Region; LRILLA = Los Lagos Region; AYS = Aysén Region; MAG = Magallanes and Antártica Chilena Region.

=

=

=

=

=

=

 =

Los Ríos Region;

**Table 1.** Presence (x) of the accepted species of *Alstroemeria* in the administrative political regions of Chile. species has been collected (*A. aurea*) and *A. patagonica* is found in Aysén and Magallanes being the most austral species of the genus *Alstroemeria* in the world. The latter species grow from 46°30′S to 52°45′S [14] and also in Argentina (Neuquén to Tierra del Fuego) [14, 34]. *Alstroemeria aurea* is the species with the widest distribution in Chile (this species spreads over

10 regions, from the O'Higgins Region, 34°12′S to Torres del Paine National Park, Magallanes Region, 51°21′S). *Alstroemeria revoluta*, the second widely distributed species, spreads from Valparaíso (La Campana National Park, 32°57′S) to Araucanía Region (Traiguén-Galvarino, 38°16′S) and *A. versicolor* ranges from the Metropolitan Region (Rio Clarillo National Reserve, 33°40′S) to Araucanía (Malleco, Renaico, 37°48′S). With these exceptions, most species show very narrow distribution, some of them being confined to a single region, such as *A. lutea* (Tarapacá Region), *A. kingii*, *A. philippi* var. *albicans* and *A. polyphylla* (Atacama Region), *A. andina* var. *venustula*, *A. hookeri* subsp. *maculata*, *A. magnifica*, *A. schizanthoides* var. *alba* and *A. mollensis*, *A. traudliae* (Coquimbo Region), *A. marticorenae* (Valparaíso Region), *A. achirae* (Maule Region), *A. hookeri* subsp. *sansebastiana* (Biobío Region), *A. presliana* subsp. *australis* (Araucanía Region, Nahuelbuta National Park). For the latitudinal distribution of each species, see reference ([14], **Figure 5**). Altitudinally, the genus *Alstroemeria* spreads from the sea level to nearly 4000 m.a.s.l. although most species are found below 2000 m.a.s.l. *Alstroemeria andina*, *A. crispata*, *A. exerens*, *A. pallida*, *A. parvula*, *A. spathulata* and *A. umbellata* can be found

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**Figure 6.** Number of taxa in the 16 administrative regions of Chile. For regions names see **Table 1**.

Cytogenetic studies in *Alstroemeria* have proved to be useful in delimiting species, since each studied taxon has a unique karyotype. These studies have contributed to the delimitation of

above 3000 m of elevation.

**4. Cytogenetic studies in Chilean** *Alstroemeria*

**Figure 5.** Distribution of the genus *Alstroemeria* in Chile. Each point represents at least one collection housed in the herbarium of the University of Concepción, Chile (CONC).

**Figure 6.** Number of taxa in the 16 administrative regions of Chile. For regions names see **Table 1**.

species has been collected (*A. aurea*) and *A. patagonica* is found in Aysén and Magallanes being the most austral species of the genus *Alstroemeria* in the world. The latter species grow from 46°30′S to 52°45′S [14] and also in Argentina (Neuquén to Tierra del Fuego) [14, 34]. *Alstroemeria aurea* is the species with the widest distribution in Chile (this species spreads over

240 Selected Studies in Biodiversity

**Figure 5.** Distribution of the genus *Alstroemeria* in Chile. Each point represents at least one collection housed in the

herbarium of the University of Concepción, Chile (CONC).

10 regions, from the O'Higgins Region, 34°12′S to Torres del Paine National Park, Magallanes Region, 51°21′S). *Alstroemeria revoluta*, the second widely distributed species, spreads from Valparaíso (La Campana National Park, 32°57′S) to Araucanía Region (Traiguén-Galvarino, 38°16′S) and *A. versicolor* ranges from the Metropolitan Region (Rio Clarillo National Reserve, 33°40′S) to Araucanía (Malleco, Renaico, 37°48′S). With these exceptions, most species show very narrow distribution, some of them being confined to a single region, such as *A. lutea* (Tarapacá Region), *A. kingii*, *A. philippi* var. *albicans* and *A. polyphylla* (Atacama Region), *A. andina* var. *venustula*, *A. hookeri* subsp. *maculata*, *A. magnifica*, *A. schizanthoides* var. *alba* and *A. mollensis*, *A. traudliae* (Coquimbo Region), *A. marticorenae* (Valparaíso Region), *A. achirae* (Maule Region), *A. hookeri* subsp. *sansebastiana* (Biobío Region), *A. presliana* subsp. *australis* (Araucanía Region, Nahuelbuta National Park). For the latitudinal distribution of each species, see reference ([14], **Figure 5**). Altitudinally, the genus *Alstroemeria* spreads from the sea level to nearly 4000 m.a.s.l. although most species are found below 2000 m.a.s.l. *Alstroemeria andina*, *A. crispata*, *A. exerens*, *A. pallida*, *A. parvula*, *A. spathulata* and *A. umbellata* can be found above 3000 m of elevation.

#### **4. Cytogenetic studies in Chilean** *Alstroemeria*

Cytogenetic studies in *Alstroemeria* have proved to be useful in delimiting species, since each studied taxon has a unique karyotype. These studies have contributed to the delimitation of the different taxa, as well as to the understanding of the chromosomal processes that determine the divergence among them [39]. Recent studies at the infraspecific level, in taxonomic complexes of the genus, have also been shown to be useful in the recognition of these taxa, either due to differences in the chromosomal architecture or in the asymmetry indexes of the chromosomes [40, 41]. Strasburger [42] was the first researcher to perform chromosome studies in *Alstroemeria* and until 1989 the number of cytological published papers involved no more than 10 different species [43]. In the last 25 years, a wide variety of cytogenetic studies have been carried out in the genus, including physical location of repetitive DNA sequences in *A. aurea* [16, 44, 45], meiosis and mitosis [46], karyology [47–49], variation and size of the genome [50], fluorescent *in situ* hybridization [39, 51, 52] and cytotaxonomy [40, 41, 52–58]. In 15 geographically isolated populations of five species of *Alstroemeria* (*A. aurea*, *A. hookeri*, *A. ligtu*, *A. pelegrina* and *A. presliana*) collected in Chile, karyotypes and variation of RAPD markers have been investigated. Tandemly repeated DNA sequences—5S and 18/25S rDNA genes and the sequence A001-1 were used to characterize karyotypes by fluorescence in situ hybridization (FISH). Ten somatic metaphases per population were used for measurement of chromosome length. Differences in RAPD marker bands were used for characterization of populations, creating a similarity index. FISH with all three DNA probes shows a high degree of polymorphism among and sometimes also within accessions of *A. aurea*, *A. hookeri* and *A. ligtu*. The number of chromosome pairs showing 5S rDNA signals is more different for the investigated species *A. aurea*, *A. hookeri*, *A. ligtu*, *A. pelegrina* and *A. presliana* with 5, 7, 5, 3 and 7, respectively, than the number of 18/25S rDNA signals in this succession with 7, 7, 6, 5 and 7 chromosome pairs, showing a high evolutionary dynamics within the genus. Furthermore, among the four populations of *A. hookeri*, accession 4181 was different in arm length of chromosome 3. RAPD markers (index of similarity) also showed a greater genetic distance of accession 4181 from the other three accessions of *A. hookeri* [39].

with microsatellites. The population of subsp. *simsii* is characterized by having five metacentric chromosomes (chromosome 2 with a microsatellite and 6 with a secondary constriction) and three telocentric chromosomes with satellites. Baeza et al. [39] analyzed four populations of subsp. *ligtu*, defining localization on the chromosomes of the ribosomal genes 5S and 18-45S. Low polymorphic hybridization sites were detected in the populations, and only chromosome 1 presented a polymorphic site of 5S and 18/25S rDNA in the proximal and distal positions, respectively. Three subspecies are recognized within *A. ligtu* complex: subsp. *ligtu*, subsp. *splendens* and subsp. *simsii*. Fourteen populations were collected throughout its distributional range. Chromosome number, karyotype formulae, karyotypes, ideograms, intrachromosomal asymmetry index MCA, and interchromosomal asymmetry index CVCL were calculated [57, 58]. All studied populations showed 2n = 2x = 16 chromosomes. Subspecies *ligtu* and *simsii* are clearly differentiated from each other in MCA and together from subsp. *splendens* with CVCL. Intrachromosomal asymmetry index revealed two population groups within subsp. *splendens*. These populations also differ in karyotype formulae, habitat, soil type and distribution. We concluded that a fourth subspecies should be described from populations located in the lower part of the cordillera de los Andes in the Region of Maule. Populations of higher elevations correspond to those already described as subsp. *splendens* [57]. A comparative karyotype study was carried out among four populations of *A. diluta* subsp. *diluta* and three populations of *A. diluta* subsp. *chrysantha*. The seven populations presented an asymmetric karyotype, with 2n = 2x = 16 chromosomes, and with same karyotype formulae: 3 m + 1sm + 1st + 3 t. The architecture of the karyotype between the subspecies is the same. The scatter plot among MCA versus CVCL shows different groupings between populations of the two subspecies, and the total chromosomes length (TCL) is highest in the populations of subsp. *chrysantha*. According to the results obtained, the populations growing in Valparaíso Region should be considered belong to subsp. *diluta* [58]. We analyzed the karyotypes of 10 populations of *A. magnifica* complex along its natural distribution. All the populations showed an asymmetric karyotype, with 2*n* = 16 chromosomes but with 3 different karyotype formulae. *Alstroemeria magnifica* var. *magnifica* and *A. magnifica* var. *sierrae* presented the same karyotype formula, and *A. magnifica* var. *magenta* and *A. magnifica* var. *tofoensis* each had a different formula. The scatter plot among CVCL versus MCA shows different groupings between populations of the four varieties. Based on these results it is possible to consider raising *Alstroemeria magnifica* var. *magenta* to species rank and *A. magnifica* var. *tofoensis* to subspecies; *A. magnifica* var. *magnifica* and *A. magnifica* var. *sierrae* should each remain as varieties. Nevertheless, these taxonomic changes should be considered tentative, as

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During recent years, an increasing accessibility to molecular data and the development of a vast range of bioinformatics analysis has favored the successful implementation of genetic tools in the identification and conservation of biological diversity [59]. Dominant molecular markers based on random fragment alleles e.g. Inter Simple Sequence Repeat (ISSR), Random Amplified Polymorphic DNA (RAPD) and Amplified Fragment Length Polymorphism (AFLP) have

additional sources of evidence become available.

**5. Molecular studies in** *Alstroemeria*

The study of the chromosomes in *Alstroemeria* has already helped to clarify a number of taxonomic issues within the genus. For example, study of karyotypes in the *A. hookeri* complex permitted change a subspecies to the species rank (*A. cummingiana*), the recognition of a new subspecies (*A. hookeri* subsp. *sansebastiana*) and description of a new species (*A. marticorenae*) [19, 22, 40, 41]. Similar situation occurred in the *A. presliana* complex, where after completing a comparative karyotypic study in 11 populations, it was suggested that *A. presliana* subsp. *australis*, endemic to the cordillera of Nahuelbuta, should be raised to species rank [18].

A number of cytological studies have been completed in the *Alstroemeria ligtu* complex. Buitendijik and Ramanna [16] and Buitendjik et al. [50] found variation in the DNA content and polymorphism of C bands in the chromosomes of subsp. *ligtu*, subsp. *simsii* and subsp. *splendens*. Zhou et al. [51], utilizing FISH, completed the characterization of the genomic DNA of eight highly repetitive sequences in subsp. *ligtu* and *simsii*, showing detailed karyotypes with localization of specific DNA sequences. DAPI staining and acetic orcein, completed a comparative karyotype study of five populations of subsp. *ligtu* from the Region of Biobío and one population of subsp. *simsii* from the Region of Valparaiso [39]. The six populations studied revealed an asymmetric karyotype with 2n = 2x = 16 chromosomes. The populations of subsp. *ligtu* have a haploid formula of four metacentric chromosomes (chromosomes 1 and 2 with microsatellites), one submetacentric with a microsatellite and three telocentric with microsatellites. The population of subsp. *simsii* is characterized by having five metacentric chromosomes (chromosome 2 with a microsatellite and 6 with a secondary constriction) and three telocentric chromosomes with satellites. Baeza et al. [39] analyzed four populations of subsp. *ligtu*, defining localization on the chromosomes of the ribosomal genes 5S and 18-45S. Low polymorphic hybridization sites were detected in the populations, and only chromosome 1 presented a polymorphic site of 5S and 18/25S rDNA in the proximal and distal positions, respectively. Three subspecies are recognized within *A. ligtu* complex: subsp. *ligtu*, subsp. *splendens* and subsp. *simsii*. Fourteen populations were collected throughout its distributional range. Chromosome number, karyotype formulae, karyotypes, ideograms, intrachromosomal asymmetry index MCA, and interchromosomal asymmetry index CVCL were calculated [57, 58]. All studied populations showed 2n = 2x = 16 chromosomes. Subspecies *ligtu* and *simsii* are clearly differentiated from each other in MCA and together from subsp. *splendens* with CVCL. Intrachromosomal asymmetry index revealed two population groups within subsp. *splendens*. These populations also differ in karyotype formulae, habitat, soil type and distribution. We concluded that a fourth subspecies should be described from populations located in the lower part of the cordillera de los Andes in the Region of Maule. Populations of higher elevations correspond to those already described as subsp. *splendens* [57]. A comparative karyotype study was carried out among four populations of *A. diluta* subsp. *diluta* and three populations of *A. diluta* subsp. *chrysantha*. The seven populations presented an asymmetric karyotype, with 2n = 2x = 16 chromosomes, and with same karyotype formulae: 3 m + 1sm + 1st + 3 t. The architecture of the karyotype between the subspecies is the same. The scatter plot among MCA versus CVCL shows different groupings between populations of the two subspecies, and the total chromosomes length (TCL) is highest in the populations of subsp. *chrysantha*. According to the results obtained, the populations growing in Valparaíso Region should be considered belong to subsp. *diluta* [58]. We analyzed the karyotypes of 10 populations of *A. magnifica* complex along its natural distribution. All the populations showed an asymmetric karyotype, with 2*n* = 16 chromosomes but with 3 different karyotype formulae. *Alstroemeria magnifica* var. *magnifica* and *A. magnifica* var. *sierrae* presented the same karyotype formula, and *A. magnifica* var. *magenta* and *A. magnifica* var. *tofoensis* each had a different formula. The scatter plot among CVCL versus MCA shows different groupings between populations of the four varieties. Based on these results it is possible to consider raising *Alstroemeria magnifica* var. *magenta* to species rank and *A. magnifica* var. *tofoensis* to subspecies; *A. magnifica* var. *magnifica* and *A. magnifica* var. *sierrae* should each remain as varieties. Nevertheless, these taxonomic changes should be considered tentative, as additional sources of evidence become available.
