Ectomycorrhizas of Cortinarius helodes and Gyrodon monticola with Alnus acuminata from Argentina.
ABSTRACT Field ectomycorrhizas of Cortinarius helodes Moser, Matheny & Daniele (sp. nov) and Gyrodon monticola Sing. on Alnus acuminata Kunth (Andean alder, aliso del cerro) are described based on morphological and anatomical features. Ectomycorrhizal roots were sampled beneath fruitbodies of C. helodes and G. monticola from two homogeneous A. acuminata forest sites located in Tucuman and Catamarca Provinces in Argentina. C. helodes ectomycorrhizas showed a thick white to beige mantle exuding a milky juice when injured, were bluish toward the apex, and had hyphal strands in the mantle. G. monticola ectomycorrhizas showed some conspicuous features like highly differentiated rhizomorphs, inflated brown cells on the mantle surface, and hyaline and brown emanating hyphae with dolipores. Restriction fragment length polymorphism analysis of the nuclear rDNA internal transcribed spacer provided a distinctive profile for each of the collections of fruitbodies and the mycorrhizal morphotypes.
- SourceAvailable from: Alejandra Becerra[Show abstract] [Hide abstract]
ABSTRACT: The objective of this study was to determine patterns of arbuscular mycorrhizal (AM) colonization of Alnus acuminata Kunth at two natural forests in relation to soil parameters at two different seasons (autumn and spring). The soil parameters studied were field capacity, pH, electrical conductivity, available P, total N and organic matter. The percentage of AM colonization was estimated and correlated to soil properties and to two different seasons. The results indicate that the percentage of AM colonization varied among soil types and was higher in spring than autumn. A significant positive correlation was found between AM colonization and electrical conductivity, organic matter and total Nitrogen. Results of this study provide evidence that AM colonization of A. acuminata can be affected by some soil parameters and seasonality.Ciencia del suelo. 07/2007; 25(1):7-13.
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ABSTRACT: The objective of this study was to determine patterns of arbuscular mycorrhizal (AM) colonization of Alnus acuminata Kunth at two natural forests in relation to soil parameters at two different seasons (autumn and spring). The soil parameters studied were field capacity, pH, electrical conductivity, available P, total N and organic matter. The percentage of AM colonization was estimated and correlated to soil properties and to two different seasons. The results indicate that the percentage of AM colonization varied among soil types and was higher in spring than autumn. A significant positive correlation was found between AM colonization and electrical conductivity, organic matter and total Nitrogen. Results of this study provide evidence that AM colonization of A. acuminata can be affected by some soil parameters and seasonality.Ciencia del Suelo 04/2007; 25(1):7-13.
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ABSTRACT: Two pure Alnus acuminata stands established in a montane forest in central Mexico (Puebla State) were monitored between 2010 and 2013 to confirm and recognize the ectomycorrhizal (EcM) systems of A. acuminata with Lactarius cuspidoaurantiacus and Lactarius herrerae, two recently described species. Through comparison of internal transcribed spacer (ITS) of nuclear ribosomal DNA sequences from basidiomes and ectomycorrhizas sampled in the forest stands, we confirmed their ectomycorrhizal association. The phytobiont was corroborated by comparing ITS sequences obtained from EcM root tips and leaves collected in the study site and from other sequences of A. acuminata available in Genbank. Detailed morphological and anatomical descriptions of the ectomycorrhizal systems are presented and complemented with photographs.Mycorrhiza 01/2015; · 2.99 Impact Factor
DO! 10.1007 /s00572-003-0278-
Ectomycorrhizas 01 Corlinarius helodes and Gyrodon monticola
with Alnus acuminata Irom Argentina
. Laura Domínguez
© Springer -Verlag 2003
28 May 2003 / Accepted: 14 October 2003 / Published online: 26 November 2003
Moser, Matheny & Daniele (sp. nov) and Gyrodon
monticola Sing. on Alnus acuminata Kunth (Andean
alder, aliso del cerro) are described based on morpholog-
ical and anatomical features. Ectomycorrhizal roots were
monticola from two homogeneous A. acuminata forest
sites located in Tucumán and Catamarca Provinces in
Argentina. C. helodes ectomycorrhizas showed a thick
white to beige mantle exuding a milky juice when injured,
were bluish toward the apex, and had hyphal strands in
the mantle. G. monticola ectomycorrhizas showed some
zomorphs, inflated brown cells on the mantle surface,
and hyaline and brown emanating hyphae with dolipores.
Restriction fragment length polymorphism analysis of the
nuclear rDNA internal transcribed spacer provided a
distinctive profile for each of the collections of fruitbod-
ies and the mycorrhizal morphotypes.
Field ectomycorrhizas of Cortinarius helodes
of C. helodes
Gyrodon monticola . Morphological characterization .
Polymerase chain reaction/restriction fragment length
Cortinarius helodes .
A. Becerra (~)
C.C. 495, 5000 Córdoba, Argentina
. G. Daniele . L. Domínguez
de Biología Vegetal (CONICET),
Oregon State University,
Corvallis, OR, 97331, USA
of Forest Science,
USDA Forest Service, Forestry Sciences Laboratory,
Pacific Northwest Research Station,
3200 SW Jefferson Way, Corvallis, OR, 97331, USA
The distribution of Alnus acuminata Kunth ranges from
the highlands of Mexico to the Andes (Dawson 1990),
reaching its southernmost point in the Catamarca province
of Argentina. Alders form ectomycorrhizas,
alder species are used as forage and fuel sources, and
quickly increase the forest biomass over previously
Species identification of mycorrhizal morphotypes is
often required in biodiversity and ecological studies. The
first approach is to characterize mycorrhizas based on
morphological and anatomical features (Miller et al.
1991; Agerer 1991); in many cases, as stated by Agerer
(1986, 1991, 1995) and Brundrett et al. (1996), hyphal
connections can be traced from the fruitbody's stipe base
to the mycorrhizas. Additionally, molecular techniques
such as polymerase chain reaction coupled with restric-
tion fragment length polymorphism analyses (PCR/RFLP)
increase the resolution of identification to the species
level (Pritsch and Buscot 1994; Egger 1995; Baldwin and
Egger 1996; Eberhardt et al. 1999; Horton and Bruns
2001; Mah et al. 2001; Sakakibara et al. 2002).
(Molina 1979, 1981; Gardner and Barrueco 1999). Trappe
(1962) and Horak (1963) list a number of fungi within the
associated with different Alnu s species. Within Cortinar-
iaceae several species have been reported as having a
mycorrhizal association with alders (Stangl 1970; God-
bout and Fortin 1983, 1985; Brunner et al. 1990; Miller et
al. 1991; Moser 2001). Pritsch et al. (1997a, 1997b)
described the ectomycorrhizas of Cortinarius cf. helvel-
loides (Fr.) Fr. and Cortinarius cf. alneus (Mos.) Mos.
associated with A. glutinosa (L.) Gaertn. Within Bole-
taceae, four Gyrodon species have been reported from
North America and Europe associated with Alnus spp.
(Hayward and Thiers 1984). Gyrodon monticola Sing.
was reported growing under A. acuminata in Mexico and
Argentina (Singer and Morello 1960; Hayward and Thiers
1984; Singer and Gomez 1984) and ectomycorrhizas of
is low in alder forest
Moench were described by Agerer et al. (1993).
The aim of this work was to characterize the ectomy-
corrhizas formed by Cortinarius helodes Moser, Matheny
& Daniele and Gyrodon monticola with Alnus acuminata
through morphological and anatomical features, and to
confirm their identity with molecular techniques (PCRJ
lividus (Bull.) Fr. with Alnusincana (L.)
Materials and methods
Two forest sites located in the northwestern
(NOA) were sampled.
Narvaez Range (Catamarca Pravince), elevation 1,820 m; 27°43'S,
65°54'W,average precipitation 1,620 mm. Mean annual temper-
atures range from 5.8 to 24°C for NOA. The vegetation is a nearly
homogeneous A. acuminata forest (height 6-15 m, age 20-30 years)
with a few herbaceous understory
(Aceñolaza 1995). Soil types are Entisols with high organic matter
contentin both locations(Becerra
30x30 m were sampled at each study site.
region of Argentina
Tafí del Valle
800 and 1,200 mm. The
plants such as Duchesnea
et al. 2002). Two plotsof
Sampling and direct identification
through 2001. At every sampling time, soil cores of 15x15 cm to a
depth of about 10 cm were concurrendy
ies. The samples were placed in plastic bags leaving the sample as
undisturbed as possible, and stored at 4°C during transport to the
The sampleswere placed in water and examined
connectionsleading fram fruitbodies
Zeiss stereo microscopeat xlO-40
method of Agerer (1991). Alder roots (which are the only structures
within these forests that present ectomycorrhizas)
within soil cores were easy to identify due to their morphological
present. Within every morphotype,
DNA extraction,while others
anatomical studies. Photographsof mycorrhizas
Leica M420 stereo microscope.
KOH, Melzer's reagent, cotton blue, 70% ethanol, sulpho-vanillin,
Nl1¡OH, and lactic acid) were used for studying
changes of mycorrhizas. Afterwards,
in 70% alcohol and stored at 4°C in the dark.
were visited during summer and fall fram1999
collected below fruitbod-
under a to fungal mandes
magnification according to the
several tips were prepared for
were taken with a
roots were fixed
Agerer (1991, 1999a) and Miller et al. (1991). Mande views were
examinedand photographed with a Zeiss Axiophot
scope at x200-1,000magnifications.
net follows Godbout and Fortin's (1983) nomenc1ature. Fruitbodies
were identified followingSinger
Moser (1978) protocols. Voucher specimens and mycorrhizas were
deposited in the Museo Botánico de Córdoba Herbarium (CORD)
(Holmgren et al. 1990).
of the ectomycorrhizasfollowsthe terminology of
of the HartigCharacterization
and Digilio (1957,1960) and
DNA extraction and amplification
DNA was extracted
lamellae from two fruitbodies
extractions were carried out twice). Species characterization
fungi was based on PCR amplification
spacer (lTS) region of the rDNA gene by using lTS-1F and ITS-4B
primers (Gardes and Bruns 1993). The primer pairs preferentially
amplify specific fragments of basidiomycete
of plant and fungus DNA. We used reagents, protocols, and cyc1ing
parameters as described previously
from one to three raot tips and fram dried
(Gardesand Bruns1993) (both
of the internal transcribed
DNA from mixtures
(Gardes and Bruns 1996).
C. helades were digested with Alu l, Hinf l and Dpn II enzymes,
whereas G. manticala
DNA was digested with Hinf l, Dpn II and
Hae III enzymes loaded side by side for comparison
agarose/2% Nusieve gel and separated by electraphoresis
100 V in a 1% TBE buffer. A 100-base pair DNA ladder (Promega,
Madison, Wis.) was used to determine fragment size. The length of
the completeITS was estimated
praduct run on a 1% agarase/2% Nusieve gel with the 100-base pair
DNA ladder. Gels were stained in ethidium bromide and observed
under ultraviolet light. lmageswere analyzed
Gene Profiler 4.02 software using default parameters. The ITS base
pair lengths are based on scores using a 100-base pair ladder.
weight standards, then plots the log molecular
intersect each standard data point.
weight values of the unknown samples,
each unknownband migrationdistance
linear line equation ( y = mx + b) for each piece (Hook 2002).
the lTS region by RFLP analysis, which was used
mycorrhizaraot samplesand fruitbodies
Aliquots of DNA from mycorrhizas
and fruitbodies of
onto a 1%
for 3 h at
the logarithmof all molecular
weight values that
Various fruitbodies and mycorrhizas were collected in the
field, but we present here two fungi (c. helodes and G.
monticola) associated with A. acuminata according to the
RFLP matches obtained.
The direct attempt to identify ectomycorrhizas of these
fungi by tracing hyphal connections between the stipe
base and mycorrhizal mot was difficult. Within the
collected C. helodes soil samples only a few mycorrhizas
were observed. Gn the other hand, mycorrhizas of G.
monticola were abundant under the fruitbodies. Within
the molecular analysis, the comparison of PCR/RFLP
patterns showed a similar band size between ectomycor-
rhizas and fruitbodies when using three endonuclease
Fig. 1 Light
Quebrada del Portugués (Tucumán Province) and Sierra de Narváez
(Catamarca Province) sites. A Simple (umamified)
root tip of C. helades; bar 0.5 mm. B Crass-section
plectenchymatous to pseudoparenchymatous
(om) (see also Fig. 2 C-D), pseudoparenchymatous
layer (mm) and pseudoparenchymatous
Hartig net (hn); bar 25 ,lim. C Monopodial
ectomycorrhizal raottipof G.
bar 50 ,lim. E Central core of thick hyphae in the
bar 10 ,lim
(A, B) collected
fram the Sierra de Narváez
(C - E) collected
Table 1 Internal
fragment band size for Cortinarius heZodes eolleeted from Sierra de
Narváez(CatamareaProvinee)and Gyrodon monticoZa eolleeted
from Quebradadel Portugués (Tueumán Provinee)
GD19 2 730 455/135/85330/125/a 325/215/195ABOl 1-3 775
455/135/85330/125/a 325/210/190Colleetion no.
ITS length (bp)
GD173 2 895
length and restnetlOn
and Sierra de
enzymes. bp Base pairs
Provine e) obtained
by using, respeetively,
ITS length (bp)
restriction fragment enzymes, thus confirming the iden-
tification of C. helodes and G. monticola as symbionts of
A. acuminata. The ITS was about 750 base pairs for C.
helodes and about 900 base pairs for G. monticola
Description of the morphotypes
Cortinarius helodes ectomycorrhizas
Simple (unramified), straight to bent, 1.2-9.2 mm in
length and 0.4-0.8 mm in diameter, some tips not entirely
colonized (Fig. lA). Mantle, white when young with
bluish to lilac tints toward the apex, pale yellow to beige
when older. Root tips are b1unt, some acute, covered by
the mantle. Mantle surface with soil particles (Fig. lA);
smooth to woolly with few to many emanating hyphae
and hyphal strands. Young mycorrhizas exudate whitish
substance When damaged.
Emanating hyphae (Fig. 2A, B): hyaline, few or
abundant, straight to bent, branched, thin walled, 2.5-
6.5 f1m in diameter, regularly septate with some clamps.
Hyphae with simple anastomoses without septa (Agerer
1991, Type A).
Hyphal strands (Fig. 2G): hyaline to pale yellow,
straight, agglutinated, up to 47 f1m in diameter at the base
with a few thin-walled and clamped emanating hyphae,
3.2-5f1m in diameter.
MycOrrhiza mantle (plane view): mantle continuous
over the root apex. Plectenchymatous
oriented hyphae with clamp connection, 2.4-5
diameter (Fig. 2C), tuming inward into pseudoparenchy-
matous tissue with elongated hyphae (Fig. 2D); middle
outer 1ayer of
Provinee) site. A Emanating hyphae with soil particles and simple
(an) (arrow); bar
bar 10 /lm. C Pleetenehymatous
loosely interwovenhyphae; bar 10 /lm. D Pseudoparenehymatous
transitional mantle layer of elongated hyphae; bar 10 /lm. E mm of
angular to spherieal eells; bar 6.25 /lm. F im of spherieal eells; bar
G Hyphalstrandsof clamped
bar 10 /lm.
epidennal eells (-);
bar 6.25 /lm. For abbreviations,
eolleeted from the Sierra de Narváez (Catamarea
of A. acuminata
and C. heZodes
outer mantle of
hyphaein a parallel
see Fig. 1
of intraeellular at
(Fig. 2E); inner layer with a pseudoparenchymatous
arrangement of small, spherical and thin-walled cells,
3.2-11f1m in diameter (Fig. 2F).
Mycorrhiza (cross section): mantle 130-220 f1m thick
(Fig. lB), differentiated into a plectenchymatous outer
layer of tangentially arranged hyphae 3.2-6.5
angular to globose cells 6.5-27.5 f1m in diameter, and a
pseudoparenchymatous inner layer of small, isodiametric
cells 4-16 f1m in diameter; paraepidermical Hartig net,
hyphae lined up in one row between epidermal cells 1.6-
5 f1m in diameter; epiderma1 cells spherical to elongated
in outline, 8-30.5 f1mx6.5-19.5
are present at the epidermal cells (Fig. 2H).
Color reaction: who1e ectomycorrhizas
with 15% KOH and 70% ethanol; emanating hyphae and
hYPhal strands stain slightly blue with Cotton blue; and
ShOWno reactions with sulpho-vanillin, 1actic acid, and
Melzer' s reagent.
Voucher specimen: ectomycorrhizas under A. acumi-
nata, in herbarium A. Becerra AB 01 (CORD); fruitbod-
ies examined: Argentina: Sierra de Narváez (Catamarca
Province), elevation 1,820 m; 27°43'S, 65°54'W; 27 May
1999, G Daniele GD 187 (CORD), 17 February 2000, GD
19 (CORD), 20 March 2001, GD 211 (CORD).
with a pseudoparenchymatous
to SPherical cells
f1m. Intracellular hyphae
Gyrodon monticola ectomycorrhizas
Monopodial to irregularly pinnate (Fig. 1C), straight,
tortuous, some bent, 0.6-6.3 mm long and 0.2-0.3 mm in
diameter, golden yellow when young, brown when old,
rhizomorphs concolorous with a shiny surface. Root tips
are b1unt, mantle surface has few globose to ellipsoid
brown cystidia and many soil particles.
Emanating hyphae of two types (Fig. 3A, B): hyaline
and brown pigmented, loosely
hyphae are branched, thin walled, 2.4-4 f1m in diameter,
regularly septate, clamped. Brown hYPhae are branched,
thin walled, 3.2-5
f1m in diameter, some complete1y
collapsed; clamp connections are present with conspicu-
ous dolipores (Fig. 3B). Simple anastomoses are present
without septa (Agerer 1991, Type A).
Cystidia (Fig. 3C, D): spherical to clavate with a small
globu1e at the distal end, brownish, 4.8-11.2
Rhizomorphs (Fig. ID, E): usually more or less
perpendicularly oriented,with a restricted
connection to the mantle, orange to brown, frequently
thick (10.1-30.5 /lm in diameter), with a central core of
very thick hyphae (6.4-13 /lm in diameter) (Agerer 1991,
type E), septate, without clamp connections. The remain-
ing hyphae are thin (2.4--4/lm in diameter), clamped, and
thin walled. Few hyphae emerge from the rhizomorphs,
and are thin walled, with or without clamps, hyaline to
brownish, 3.2-5 /lm in diameter.
Mycorrhiza mantle (plane view): usually continuous
over the root apex. Outer layer is plectenchymatous with
patches of roundish to ellipsoidal brown cells (mantle
type F, Agerer 1991), hyphae interwoven, hyaline, thin
walled, with some clamped septa, 3.2-6 /lm in diameter
(Fig. 3E). Toward the Hartig net, hyphae are smaller in
diameter and length, spherical to elongated,
pseudoparenchymatous and tangential arrangement, 2.4-
8 /lm in diameter, thick walled (Fig. 3F).
Mycorrhiza (cross section): mantle 24-57 /lm thick
(Fig. 3G), differentiated into a p1ectenchymatous outer
layer of tangentially orientated hyphae 3.2-6.5 /lm in
diameter, and a pseudoparenchymatous
roundish cells 2.5-9.1
/lm in diameter. Periepidermal
Hartig net has hyphae between epidermal cells in one
row, 1.6-2.5 /lm in diameter (Fig. 3H). Epidermal cells
are spherical to elliptical in outline, thick walled, 11-
17 /lmx6-16 /lm.
Color reaction: whole ectomycorrhizas and emanating
hyphae stain slightly blue with Cotton blue; the root and
the Hartig net stain reddish with sulpho-vanillin; b1each
with NH40H and lactic acid; and show no reaction with
15% KOH, Me1zer's reagent and 70% ethanol.
Voucher specimen: ectomycorrhizas under A. acumi-
nata, in herbarium A. Becerra AB 05 (CORD); fruitbod-
ies examined: Argentina: Quebrada del Portugués, Tafí
del Valle(Tucumán Province),
26°58/S, 65°45/W and Sierra de Narváez (Catamarca
Province),elevation 1,820 m; 27°43/S, 65°54/W;
March 1999, G Daniele GD 173 (CORD), 20 March 2000,
GD 207 (CORD).
inner layer of
cumán Province) and Sierra de Narváez (Catamarca Province) sites.
(h) and brown
surface; bar 6.25 ,lim. B Emanating
Clavate light brown cystidia with a small distal globule (-);
8 ,lim. E Plectenchymatous outer mande of interwoven hyphae; bar
8 ,lim. F im; bar 6.25 ,lim. G Cross-section
matous outer mande (om), im and hn; bar 10 ,lim. H hn surrounding
epidermalcells (tangential section); bar 8 ,lim. For other abbrevi-
ations, see Fig. 1
of A. acuminata
from the Quebrada
and G. monticola
del Portugués (Tu-
b with conspicuous
(b) from mande
bar 10 ,lim. e Spheric brownish cystidia (-);
bar 8 ,lim. D
In this study, the combined approach of morphotyping
ectomycorrhizas of C. helodes, G. monticola with A.
acuminata in the field.
The PCR/RFLP analyses were used successfully for
the identification of the freshly harvested fruitbodies and
the morphotypes. The two mycorrhizas were separated by
their molecular profiles, supporting the morphological
observations. Band size estimates made by using agarose
gels are approximate, with an error rate of about 3%. We
obtained an ITS length of approximately 750 base pairs
for C. helodes, in contrast to a 61O-base pair ITS for
Cortinarius ectomycorrhizas obtained by Pritsch et al.
(1997a). The larger ITS found in our study is directly a
function of using the primers ITS-lF and ITS-4B, which
amplify a larger fragment through the priming sites (ITS-
1 and ITSA)employedby Pritsch
Differences in band size patterns between fruitbodies
and mycorrhizas in the G. monticola RFLP analysis
(around 1%) probably are due to band size estimation
error rateo No other Boletales were observed during the
sampling, nor are there reports of other species from the
Cortinarius and Gyrodon species, like Cortinarius
bibulus Quel.; C. cf. saturninus (Fr.) Fr.; C. cf. alneus
(Mos.) Mos. and Gyrodon lividus have been found
associated with North American and European Alnus
spp. (Horak 1963, 1985; Schmid-Heckel 1985; Miller et
al. 1987; Brunner et al. 1990; Pritsch et al. 1997b).
From this study, C. helodes was identified as a new
species, described by Moser (2001). C. helodes ectomy-
corrhiza showed similar features to C. cf. helvelloides
ectomycorrhiza on A. glutinosa (Pritsch et al. 1997a,
1997b). The most conspicuous
absence of refractive vacuoles
collapsed emanating hyphae in C. helodes. Similarly to
the type A mycorrhiza of A. japonica
Cortinarius sp. (Masui 1926), C. helodes hyphae invade
the epidermal cells (Fig. 2H). Cortinarius cf. alneus
(Pritsch et al. 1997b) and C. bibulus (Miller et al. 1991)
showed comp1etely different
mantle coloration (whitish, silvery to brown when old);
mantle surface (woolly, cottony) and mantle thickness
G. monticola showed mantle coloration, emanating
hyphae with brown pigments and unpigmented hyphae,
cystidia arranged in a nest-like manner on mantle, and
highly differentiated rhizomorphs similar to G. lividus
and A. incana ectomycorrhizas
Nevertheless the latter type is distinguished by sclerotia
lying on ramifications,not present in G. monticola
mycorrhiza. Brunner et al. (1990) found that G. lividus
did not form a Hartig net on A. tenuifolia Nutt. in
synthesis in vitro. G. monticola ectomycorrhizas on A.
acuminata form an obvious Hartig net, although the
mantle morphological features such as color are quite
similar to G. lividus ectomycorrhizas.
us to identifythe
et al. (1997a).
on mantle cells and
(Agerer et al. 1993).
Using direct identification by tracing mycelia fram the
fruitbodies to mycorrhizas (Agerer 1991) [successfully
done by Agerer (1999b, 1999c), Agerer and Beenken
species, among other authors] in a natural mature stand
of A. acuminata was unsuccessful, because more than one
morphotype was found below fruitbodies, as also found
by Sakakibara et al. (2002). Nevertheless, C. helodes and
were identified molecularly
RFLP analysis) and morphologically as symbionts of A.
in native Argentinean forests.
particu1arly thank Dr M. Moser for the identification
helodes sp. nov. Special thanks to Dr Efren Cázares, Oregon State
University and Marcelo Zak for critically reading the manuscript.
A. B. is gratefulto FOMEC and CONICET
provided, E. N. holds a postdoctoral
This work was partially supported by Agencia
(1999, 2001). We thank Tom Horton and Joyce
for their assistance in the mo1ecu1ar analysis. We also
Ciencia 1416/97;1655/99) and
for the fellowship
position from CONICET.
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