Darwinian

printed version ISSN 0011-6793

Vol.46no.2DarwinianSanIsidroAug./Dec.2008

SYSTEMATICS AND TAXONOMY OF VASCULAR PLANTS

District of Rebutia Minuscula (Cactaceae, Cactoideae)

Nora B. Muruaga 1 , Mary R. Romero Figueroa ? & Roberto Kiesling2
Miguel Lillo Foundation, Laboratory of Plant Taxonomy Fanerogmica, Miguel Lillo 251,
4000, Tucumn, Argentina; nmuruaga@yahoo.com.ar (corresponding author).
IADIZA. Argentine Institute for Arid Zones, Mendoza, Argentina.
Original received on 7 April 2006, accepted on April 4, 2008
Summary.
Species Rebutia distributed from northwestern Argentina to southern Bolivia (Cochabamba to
Tarija). Five species growing in Argentina, including the type species of the
genus: R. lowercase. Phenotypic diversity, very common within species Rebutia, has led to
the publication of numerous specific and infraspecific names, producing a real confusion in
the nomenclature and synonymy distinctness of species, particularly R .lowercase, which
overlaps the boundary of R. wessneriana. The purpose of this study was to reevaluate the
definition of R.lowercase. 's exo-micromorphological characteristics of natural populations
inhabiting the upper montane forest and high pastures of the provinces of Catamarca,
Tucumn, Salta and Jujuy were analyzed. The data were documented with illustrations and
photographs, information of geographical distribution, ecological and phenological
observations is provided. Includes synonyms and nomenclatural types for some of them are
designated. Is considered to R. wessneriana as a subspecies of R. lowercase and proposes a
new combination and a new range for naming: Rebutia lowercase subsp. wessneriana.
Keywords . Argentina; Cactaceae; Cactoideae; Rebutia; Taxonomy.
Abstract . circumscription of Rebutia lowercase (Cactaceae, Cactoideae).
Species of Rebutia are distributed from northwestern Argentina to southern Bolivia
(Cochabamba to Tarija). Five species grow in Argentina, including the type species of the
genus: R. lowercase. phenotypic The diversity, very varying Within the species of
Rebutia, Promoted publication of the multitude numerous specific and infraspecific names,
leading to significant confusion in nomenclature, synonymy, and distinctive characters of its
species, Particularly for R. lowercase, Whose delimitation overlaps With That
of R. wessneriana. The aim of this work is to reevaluate the circumscription
of R. lowercase. Exo-micromorphological Characteristics of Natural Populations growing in
the upper mountain forest and highgrasslands of Catamarca, Tucumn, Salta and Jujuy
provinces, Were analyzed. Data Were documented with illustrations and photographs;
geographic distribution of information is provided as well as ecological and phenological
observations. Synonyms are included and the nomenclatural types for some of them are
established. Rebutia wessneriana is Considered as subspecies of R. lowercase and a new
combination and a new rank is Proposed: Rebutia lowercase subsp. wessneriana.
Keywords . Argentina; Cactaceae; Cactoideae; Rebutia; Taxonomy.

INTRODUCTION
Rebutia lowercase K. Schum. is the type of the genus and one of the five
cited by Kiesling (1999) for Argentina species. It presents a great variability in
their habit, shape and size of the stem, dis position of their hubs, number,
length and color of the thorns, so does the color of its flowers between red
and orange and the arrangement of flowers along the stem. The areolas floral
ornamentation, presence or absence of hairs and bristles, and the degree of
welding of style with the floral tube, are also variable traits within the species,
which can be interpreted in different ways. Buxbaum (1953) mentions that
in R. lowercase floral areolas have hairs or bristles and the floral tube is
welded to the base of the style. In contrast, and using that name in a broader
sense, Hjertson (2005) distinguishes R. lowercase , among other characters,
by having floral areolas hairy, with or without bristles and "solid floral tube", ie,
style, partly welded to the tube.
Moreover, Hjertson (2005) recognizes R. wessneriana Bewer. as a synonym
of R. lowercase characters but does not specify that it is based to synonymy,
whereas Kiesling (1999) treated R. wessneriana as a synonym
of R. marsoneri Werderm,. throughout the previous
literature R. wessneriana and R. lowercase are treated as different species,
mainly by the soldier style floral tube flower larger and more intense red
in R. wessnerianaand freestyle in R. lowercase .
The purpose of this research was to circumscribe R. lowercase based on the
exomicromorfolgicas characteristics presented in their natural habitat, and
discuss its relationship with R. wess-neriana . The characteristics studied are
documented with illustrations and photographs, geographic distribution
information is provided both altitudinal and latitudinal, ecological and
phenological observations.
MATERIALS AND METHODS
A comprehensive assessment of the exo-micromorphological characteristics
in natural populations in the upper montane forest and high pastures of the
provinces of Catamarca, Tucumn, Salta and Jujuy was performed. Original
descriptions and herbarium specimens available in BAB, CTES, LIL, SI, SGO
(Holmgren et al., 1990) were analyzed.
Pollen and seed materials come from samples collected from their
habitat. Pollen grains were acetolyzed according technique Erdt-man (1960)
and mounted in glicerinagelatina. The observations were made with an optical
microscope (OM) Wild. For analysis of sculpture exine material not
acetolizado was used metallized do with gold vapor and microfotogra-rays
were obtained with a scanning electron microscope (SEM) Jeol 35 CF Service
Electron Microscopy, National University of Tucumn .
to observe seed SEM, washed repeatedly with distilled water and mounted in
synthetic enamel specimen holder and coated with gold vapor, then
photomicrographs were obtained with the same scanning electron microscope
(SEM). For descriptions of the terminology seeds Barthlott & Hunt (2000) was
followed.
RESULTS AND CONCLUSIONS

Rebutia wessneriana shares with R. lowercase the following morphological

features: simple, globose stem, globosodeprimido to cylindrical, sometimes
clavate, then pendulum always mamelons tetrahexagonal base; numerous
spines (-30), straight and usually weak, lateral flowers, small, 5 ( -6) cm long,
infundibular, long, narrow tube, red to orange,. stamens numerous spirally
arranged along the floral tube, partially welded to the floral tube at its base,
then freestyle; floral areolas almost all glabrous; a Sometimes, some of the
basal pericarpelo have a few hairs and bristles.
Both entities pollen is similar: isopolar, lobed-fosaperturado, tricolpate; exine
(SEM) puntitegilada with scattered spinules and perforations ( Figs 1. ,
and 2 ).

Fig 1. bean pollen R. lowercase subsp. wessne-Saharan Africa . A, polar view. B, equatorial
view. C, detail of exine (of 390 Muruaga, LIL).

Figure 2. Pollen Grain R. lowercase subsp. lowercase . A, equatorial view. B, detail of exine
(of 298 Muruaga , LIL).

The seeds in both taxa are oval, small (0.8-1.2 mm long.), Brilliant black, the
head cells are generally isodiametric, tetragonal to hexagonal, with limit
anticlinal ribbed, straight and well intracellular gap differentiated and
embossed convex; testa cells seed apex shaped projections have acuminata
or are truncated and the side of lower dome shaped cones, gradually become
smaller in the edge region hilomicropilar; which is oblique and large ( Figs
3.and 4 ).

Figure 3. seed R. lowercase subsp. lowercase . A, detail of testa cells of the apex of the
seed. B, testa cells of striated anticlinal walls ( Muruaga 298, LIL).

Figure 4. seed R. lowercase subsp. wessneriana . A, detail of apex cells of the seed. B cells
testa smooth anticlinal walls (Muruaga 390 , LIL).

These data indicate that R. lowercase and R. wessneriana share most

morphological characteristics and is therefore difficult to draw boundaries
between them. We also note that in R. lowercase and R. wessnerianastyle
floral tube and soldiers are always greater or lesser extent, and both flower
size and color are not enough to keep them as distinct species, as these
characters vary in natural populations. Furthermore, analysis of pollen grains
and seeds reinforce the similarities between the two taxa
tudes. However, R. lowercase and R.wessneriana differ in the degree of
welding floral tube, in the ornamentation of the anticlinal cell walls of the seed
coat in the geographical distribution and, hence, we consider that there are
two distinct species, but R.wessneriana is a subspecies of R. lower case , on
the basis of differential characters you can see in the Table 1 .
Table 1. Differences between subspecies of Rebutia lowercase.

Moreover, R. marsoneri differs from R. lowercase for having a depressedglobose stem to 4 cm diam., areolas with 9-15 spines 3-5 mm long. and red or
yellow flowers.
TAXONOMIC TREATMENT
Rebutia lowercase K. Schum., Monatsschr. Kakteenk. 5 (7):
102. 1895. Echinopsis minus culaFAC Weber in Bois, Dict. Hort. 1:
471. 1896. Echinocactus minusculus FAC Weber in Bois, Dict.Hort. 1:
467. 1896. TYPE: sn icon in Schumann, Monatsschr. Kakteenk. 5 (7):
102. 1895, reproduced by Schumann in Gesamtb. Kakteen: 395, fig. 67. 1898
(lectotype been appointed by M. Hjertson, Cact Init Syst 19:.... 20
2005). Fig. Six .

Figure 5. flower detail. A, R. lowercase subsp. lowercase (for Muruaga 298 ,

LIL). B, R. lowercase subsp. wessne-Saharan (of 390 Muruaga , LIL).

Fig 6. Rebutia lowercase subsp. lowercase . A, general

appearance. B, areola. C, flower. D, longitudinal section of flower and detail of the insertion of
the filaments. E, abaxial view of a flake floral pericarpelo where the areola is observed with
some hairs. F, fruit. G,seed (of 298 Muruaga , LIL).

Stems usually solitary, globose-depressed to cylindrical, 1-14 cm long. x 2-8

cm diam., exceptionally clavate, then pendulums up to 20 (-40) cm long.,
olivaceous-green to green. Hubs base tetra arranged in a hexagonal
spiral. Thorns (10 -) 25 (-30) or whitish tip and chestnut, dark yellow or brown
basis up to 25 mm long, straight and generally weak.. Numerous flowers ( 15
simultaneous), 3-5 (-6) cm long. x (2 -). 3-4 cm diam, laterally located in the
stem, never apical infundibular completely red, sometimes red-blood, or the
yellow receptacle basal, middle or upper portion and red perigonio Brown
reddish-violet to orange or pink. Numerous stamens ( 63), with yellowishwhite or pink filaments arranged spirally along the floral tube 14 mm
long. Style white to reddish; whitish stigma. The floral Areolas usually
glabrous receptacle, sometimes some baseline pericarpelo tube and with few
short hairs, with 1-5 (-10) bristles or without. Fruit globose to subglobose,

sometimes pyriform, greenish, yellowish to reddish pink, triangular to broadly

triangular scales of ca. 2mm long. x 1 mm lat., somewhat fleshy at the base
and membranous at the apex, glabrous or glabrescent. Seeds numerous,
generally small 0.8-1.2 mm long, shiny black testa,. Testa cells at the apex of
the seed-truncated conical projections or conical acuminate that you have
roughly grouped into a ridge or are scattered.
Key to the subspecies of Rebutia lowercase
1. Style welded on (1 -) 3 mm long. the floral tube at its base. Seeds with testa cells of striated
anticlinal walls. Dispatched from Jujuy Catamarca up ............
1a. R. lowercase subsp. minuscule
1. Style soldier in 7 (-10) mm long. the floral tube at its base. Seeds with testa cells smooth
anticlinal walls. It lives in the town of Volcan, Jujuy Province .......
1b. R. lowercase subsp. wessneriana

1a. Lowercase R. K. Schum. subsp. lowercase

Rebutia senilis Backeb, Freund Kakteen-1 (11). 124. sn 1932 fig. TYPE: sn
icon in Backeberg, KakteenFreund 1 (11): 124. 1932 (seen lectoti-po
appointed by M. Hjertson, Cact Init Syst 19:... 21 2005.).
Rebutia senilis var. hyalacantha Backeb, Freund Kakte-in-1 (12).
131. 1932. TYPE: Argentina, Salta, Chicoana, Escoipe Gorge, 1600 m sm,
15-IX-2005, NB 404 Muruaga (LIL neotype here designated).
Rebutia senilis var. stuemeri Backeb, Freund Kakteen-1 (12). 131. fig. sn
1932. TYPE: sn icon in Backeberg, Kakteen-Freund. 1 (12): 131. 1932
(lectotype here designated).
Rebutia xanthocarpa Backeb, Freund Kakteen-1 (12). 131. fig. sn
1932. TYPE: sn icon in Backeberg, Freund Kakteen-1 (12): 131. 1932
(lectotype been appointed by M. Hjertson, Cact Syst Init:... 19: 22 2005.).
Rebutia dasyphrissa Werderm., in Fedde, Repert. Spec. Nov. Regni
Veg. Sonderbeih. C. t. 103.1935. Rebutia
xanthocarpa var. dasyphrissa (Werderm.) Backeb.,
Beitr. Sukkulentenk.Sukkulentenpflege: 3. 1939. TYPE: sn icon in
Werdermann, bluh. And kakteen. Sukk. Pflanzen 26: t. 103. 1935 (lectotype
here designated).
Rebutia violaciflora Backeb., Blatt. Kakteenf. (Unnumbered) 8. 1935. fig. sn
1935. Rebutia lowercase K. Schum. . subsp violaciflora (Backeb.) Donald,
Ashingtonia 2 (3): 43. 1975. TYPE: sn icon in Backeberg
Blatt. Kakteenf. (Unnumbered) 8. 1935 (lectotype here designated).
Rebutia chrysacantha Backeb in Backeberg & Knuth, Kaktus-ABC. 276,
416. "1935", 1936.Rebutia senilis Backeb. subsp. chrysacantha (Backeb.)
Donald, Ashingtonia 2 (3): 50. 1975.TYPE: Argentina, Jujuy, El Carmen, Abra
Santa Laura, 1336 m sm 3-XII-2004, NB 391 Muruaga(LIL neotype here
designated).

Rebutia grandiflora Backeb in Backeberg & Knuth, Kaktus-ABC. 277,

416. "1935", 1936. Rebutia lowercase var. grandiflora (Backeb.) Krainz in
Krainz, Die Kakteen, 15.VII.1960. CVC. (No pagination, sub R.
lowercase var. grandiflora ). Rebutia lowercase subsp. grandiflora (Backeb.)
Donald, Ashingtonia 2 (3): 43. 1975. TYPE: Argentina, Salta, Chicoana,
Quebrada Escoipe 12-XI-1995, R. Kiesling et al. 8723 (SI neotype here
designated).
Rebutia knuthiana Backeb in Backeberg & Knuth, Kaktus-ABC. 277,
416. "1935", 1936. Rebutia violaciflora var. knuthiana (Backeb.) Donald, Nat
Cat Succ. J. 12: 11. 1957. TYPE: Argentina, Salta, Rosario de Lerma, Campo
Quijano, 18-XI-1942, A. Burkart 13162 (SI neotype here designated).
Rebutia senilis Backeb. var. aurescens Backeb in Backeberg & Knuth,
Kaktus-ABC. 278, 416."1935", 1936. TYPE: Argentina, Salta, Chicoana, El
Rodeo, before The Trench, 2300 m sm, 16-IX-2004, NB 385 Muruaga (LIL
neotype here designated).
Rebutia senilis Backeb. var. lilacinorosea Backeb in Backeberg & Knuth,
Kaktus-ABC. 416."1935", 1936. TYPE: Argentina, Salta, Chicoana, 22 Km
west of El Carril, 2770 m sm, 10-X-2004,NB 386 Muruaga (LIL neotype here
designated).
Rebutia xanthocarpa Backeb. var. citricicarpa Fric ex Backeb., Cact. Succ. J.
(U.S.) 23 (3): 83.1951. TYPE: Argentina, Salta, La Caldera, from Yacones to
Potrero de Castilla, 1800 m sm, 14-IX-2004, M. Cecotti & NB 383-ga
Murua (LIL neotype here designated).
Rebutia xanthocarpa Backeb. var. elegans Backeb., Cact. Succ. J. (U.S.) 23
(3): 83. 1951.Rebutia chrysacantha var. elegans (Backeb.) Backeb. Die
Cactaceae 3: 1539. 1959. TYPE: Argentina, Tucuman, Trancas, San Pedro de
Colalao, 1500 m sm, XI-1933, CR 9742 Schreiter(LIL neotype here
designated).
Rebutia xanthocarpa Backeb. var. luteirosea Backeb., Cact. Succ. J (U.S.) 23
(3): 83. 1951.TYPE: Argentina, Salta, Guachipas, Torino, 1100 m sm, 14-IX2005, NB 402 Muruaga (LIL neotype here designated).
Rebutia xanthocarpa Backeb. var. salmonea Fric ex Backeb., Cact. Succ. J.
(U.S.) 23 (3): 83.1951. TYPE: Argentina, Tucuman Tafi Viejo, Cumbres del
Cerro Cabra low Horco, 1900 m sm, 17-IX-1999, NB 324 Muruaga (LIL
neotype here designated).
Rebutia senilis Backeb. var. schieliana Bewer., Kakt. And. Sukk. 8 (7):
105. fig. sn 1957. TYPE: sn icon in Bewerunge, Kakt. And. Sukk. 8 (7):
105. 1957 (lectotype here designated).
Style floral tube welded to the base to 3 mm long., Then free ca. 33 mm long.,
Whitish. Seeds with testa cells of striated anticlinal walls.

Geographic distribution and habitat . Schumann (1895) mentioned

that R. lowercase probably came from Argentina, according to information
received from Mr. Pierre Rebut, French time grower who is dedicated the
genus name.
Subsequently Weber (1896) Quote for the province of Tucumn and
Schumann (1898) repeats the town of Weber, who quite possibly got the
plants directly from F. Schickendantz of Tucumn. Spegazzini (1905) Quote
for Tucuman and Salta, like Kiesling (1999).
campaigns conducted in northwestern Argentina ratify its allowed range for
the provinces of Tucuman and Salta, Jujuy expand its presence where the but
under other names known as the province of Catamarca, where there had
been summoned.
grows in rock crevices exposed upper montane forest in rocky or hilly relief
sites in grassland environment, between 1100-2800 m sm is a component of
the stratum herbaceous and grows with Viguiera tucumanensis(Hook. & Arn.)
Griseb. var. oligodonta (Blake) Cabrera, Stevia yaconensis Hieron., Tagetes
terniflora Kunth,Ipomoea rubriflora O'Donnell and Lamprothyrsus
hieronymi (Kuntze) Pilg., among other species.
Phenology . Both cultured and wild stocks of different species of the material,
it was observed that R.lowercase begins to bloom in mid-August and flowers
and fruits simultaneously. The period of greatest production of flowers and
fruits is at the end of the dry season and beginning of the rainy season
(October-November).
Anthropogenic factor . Collection sites generally do not show any significant
alteration produced by man.However, as a result of livestock in some high
mountain sites, dry grass burned in the months of September and
October. This activity significantly affects populations living there.
Observations . In most cacti described based on a specimen grown
herbarium materials or locations not mentioned, but the names are valid
through 1957 were published (Greuter et al, 2000 Art.. 37.1), but several
descriptions are accompanied by an illustration that the only existing original
material observed by the author, have been designated as lectotypes
here. Designate neotypes for taxa described before January 1, 1958 of
Backeberg publications (in Cact. Succ. J. (U.S.) 23 (3), 1951) and Backeberg
& Kunth (in Kaktus-ABC, "1935", 1936) was not designated as type material
by the authors. Also, not to Bckerberg herbarium specimens, so that almost
all species published since 1958 are invalid.
While following synonyms were already established by other authors, we
consider it appropriate to comment on our observations based on wild
material:
In Rebutia senilis specific epithet refers to the white color of the spines, a
quality that is quite variable within a population. To our knowledge, this name
was created for individuals who are depressed-globose stem 7 cm long. x 8
cm diam., completely covered with thorns about 2.5 cm long.
Rebutia senilis var. stuemeri was created by Backeberg (1932) for their

flowers up to 4 cm long., red-tile inner tepals yellow to reddish white throat;

these characters, highly variable within a population, are not sufficient to
accept a variety within the species.
Variety Rebutia senilis var. lilacinorosea is not accepted because the
difference allocated in the color of the flower is not relevant and .'s color
variations between the species having
Backeberg & Knuth (1936) mention the presence of very thin, sharp, long and
dense spines, yellow plants inRebutia senilis var. aurescens , these
characters are highly variable within a population for which the assigned
differences are not relevant.
Rebutia grandiflora is distinguished, among other characters, by the size of
the flower (6.5 cm long.) size we observed in individuals flowers locality El
Rodeo, Apt. Chicoana, Province of Salta, corresponding to R.lowercase .
's name Rebutia xanthocarpa was based on the yellow color of the fruit, but
this color is part of the variation that can be found in the fruits of the species.
Rebutia xanthocarpa var. salmonea as its varietal epithet implies, I think for
individuals with salmon flowers, varying in color in different individuals.
Rebutia xanthocarpa var. elegans is characterized by long and wispy spiral
spines and ribs. The length and consistency of the spines are variable
characters within the group. Sometimes the hubs are arranged vertically
aligned together giving the appearance of ribs. However, this feature is not
constant within the species.
's name Rebutia chrysacantha was created for individuals who have the
yellow gold spines. Observations in natural populations show that the color of
the spines is variable and may be whitish, yellowish or dark brown.
Backeberg (1977) quotes Rebutia violaciflora to northern Argentina, in Salta
and in the upper part of the Gorge Escoipe. The material collected by us in
flowers exhibit the same red-violet shades that this taxon is based, which are
within the variation of colors presented floral species.
Representative material examined
ARGENTINA. Jujuy . Dept.. Dr. Manuel Belgrano, way Tiraxi 3-XI1974, Schinini 10014 (SELV), from Laguna Tiraxi Treasurer, 1517 m sm, 16IX-2005, 405 Muruaga (LIL). Tucuman . Dept..Trancas, Quebrada de la
Hoyada, 1500 m sm, 26-XI-1920, Schreiter 66740 (LIL); Hualinchay,
Quebrada Pozo del Rio Grande, 1500 m sm, 9-IX-1999, 298
Muruaga (LIL). Dept.. Tafi del Valle, uorco base of Cerro Grande, 2800 m
sm, 17-IX-2000, Boero sn (LIL 605758). Catamarca.Department. Andalgal
The Banderita, 2000 m sm, 20-X-2001, 357 Muruaga (LIL).
1b. Rebutia lowercase K. Schum. subsp. wess-neriana (Bewer.) Muruaga,
stat. & Comb. nov. Rebutia wessneriana Bewer. Sukkulentenkunde 2:
24. 1948. Rebutia krainziana Kesselr. var. wessneriana (Bewer.) Krainz &
Haarm., Katalog. Stdt Zrich. Sukk. Samml. Ed 2. 107. 1967. TYPE: sn icon
in Bewerunge, Sukkulentenkunde 2: 24. 1948 (lectotype been appointed by
M. Hjertson, Cact Init Syst 19:.... 21 2005).
Rebutia calliantha Bewer. Sukkulentenkunde 2: 25. 1948. Rebutia
wessneriana Bewer. var. calliantha (Bewer.) Donald, Nat Cact. Succ. J. 14 (1):

5. 1959. TYPE: Argentina, Jujuy, "Volcano unten" F. Ritter 58 . Loc 2 (sa) (SGO-124559
neotype here designated).

Style floral tube welded to the base to 10 mm long., Free in ca. 12 mm long.,
White, red to pink. Seeds with testa cells smooth anticlinal walls.
Geographical distribution and habitat. It lives in the town of Volcano, Province
of Jujuy, and grows rooted in the clefts of the rocks on the rocky hillsides.
Observations . With no existing material on which the name of the original
taxon was based Rebutia calliantha neotype is designated.
Representative material examined
ARGENTINA. Jujuy . Dept.. Tumbaya, Volcano, Laguna, 19-I-1998, Kiesling
8901 (SI), Volcano, 31-X-1974, MC-8 (BAB), Volcano, Laguna, 2092 m sm 11X-2004, 390 Muruaga ( LIL).
Invalid Names
Rebutia hyalacantha (Backeb.) Backeb in Backeberg, Die Cactaceae 3:.
1551. 1959. nom. illeg.as superfluous, by including R. wessneriana between
synonyms and therefore its type (Greuter et al., 2000. Art. 52.1).
Rebutia permutata Heinrich in Backeberg, Descr. Cat Nov. 3:
13.1963. nom. inval. lack of mention of type item (Greuter et al., 2000. Art.
37.1).
Rebutia wessneriana Bewer. . var permutata (Heinrich) Buining & Donald,
succulenta 51 (12): 225, 1972. comb. inval. because its basionym is invalid.
Rebutia xanthocarpa Backeb. var. violaciflora (Backeb.) Backeb in Backeberg,
Descr. Cat Nov. 1: 31. 1956. nom. inval. lack of mention of the basionym
(Greuter et al., 2000. Art. 33.3).
Acknowledgements
The authors thank the Fundacin Miguel Lillo by the total support that has enabled this work
to be carried out. Reviewers for their valuable suggestions, Mr. Alberto Gutierrez conducting
sheets and the curators of the herbaria for the loan of the requested material.
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