Mycorrhizas in the South American Mediterranean-Type Ecosystem: Chilean Matorral

Abstract

One of the most important microorganisms in the soil are the mycorrhizal fungi;
however, little research exists regarding mycorrhizal symbiosis on the South
American Mediterranean-type ecosystem (MTE) – also commonly known as
Chilean matorral. The aims of this chapter are to highlight and compile the
existing and arising knowledge on mycorrhizal symbiosis of the Chilean MTE,
as well as detect knowledge gaps and propose future directions of research. So
far, the mycorrhizal type of some plant species of the Chilean matorral is known.
Regarding arbuscular mycorrhizal symbiosis, there are few investigations on
mycorrhizal ecology and applied research with agricultural purposes and more
is in development. Some ectomycorrhizal and orchid mycorrhizal symbiosis
research is available on ecological concerns about biodiversity patterns with
applied potential for conservation. The lack of studies on ericoid mycorrhiza
was detected. Finally, in spite of the ecological diversity studies carried on the
mycorrhiza from the Chilean MTE, further studies quantifying the mycorrhizal
contribution should be performed so as to be applied on conservation and
sustainable systems.

Full text

277© Springer Nature Switzerland AG 2019
M. C. Pagano, M. A. Lugo (eds.), Mycorrhizal Fungi in South America,
Fungal Biology, https://doi.org/10.1007/978-3-030-15228-4_14
Chapter 14
Mycorrhizas intheSouth American
Mediterranean-Type Ecosystem: Chilean
Matorral
PatriciaSilva-Flores, AnaAguilar, MaríaJoséDibán,
andMaríaIsabelMujica
14.1 Introduction
The ve Mediterranean-type ecosystems (MTEs, singular: MTE) in the world are
climatically characterized with warm-dry summers and cool-wet winters (Rundel
and Cowling 2013). These ecosystems are located in California, central Chile, the
Mediterranean Basin, the Cape Region of South Africa, and southwestern and south
Australia (Dallman 1998; Rundel and Cowling 2013). A remarkable feature of the
MTEs is the fact that they occupy, in total, less than 3% of the Earth’s surface and
contain almost 50,000 species of vascular plants, which correspond to 20% of the
world’s known species (Cowling etal. 1996; Rundel and Cowling 2013). Also,
many of the plant species are endemic (Cowling etal. 1996) and, at the same time,
P. Silva-Flores (*)
Centro de Estudios Avanzados en Fruticultura (CEAF), Santiago, Chile
Departamento de Botánica, Universidad de Concepción, Concepción, Chile
Micólos ONG, Concepción, Chile
e-mail: psilvaf@ceaf.cl
A. Aguilar
Centro Regional de Innovación Hortofrutícola de Valparaíso (CERES), Quillota, Chile
Ponticia Universidad Católica de Valparaíso, Valparaíso, Chile
M. J. Dibán
Micólos ONG, Concepción, Chile
Departamento de Ciencias Ecológicas, Universidad de Chile, Santiago, Chile
Instituto de Ecología y Biodiversidad (IEB), Ñuñoa, Chile
M. I. Mujica
Instituto de Ecología y Biodiversidad (IEB), Ñuñoa, Chile
Departamento de Ecología, Ponticia Universidad Católica de Chile, Santiago, Chile

278
they are threatened by several human-related factors (Underwood etal. 2009). All
these characteristics have placed the MTEs as biodiversity hotspots (Myers etal.
2000), which means that they are priorities for conservation. Due to this situation,
much research has been done in MTEs, mostly in plants (Dallman 1998), but also
some attention has been paid to animals (Rundel and Cowling 2013). However, the
soil microbiological biodiversity has been overlooked, despite the fact that they are
increasingly being recognized as key players in the restoration of degraded ecosys-
tems (Harris 2009).
One of the most important microorganisms in the soil are the mycorrhizal fungi
(MF) which form symbiotic relations with the roots of approximately 90% of known
plant species (Brundrett and Tedersoo 2018). These mycorrhizal symbiotic relations
occurs in almost all ecosystems (Read 1991; Read and Perez-Moreno 2003; Read
etal. 2004) and as four main types: arbuscular mycorrhiza (AM), ectomycorrhiza
(EcM), orchid mycorrhiza (OrM) and ericoid mycorrhiza (ErM) (Brundrett and
Tedersoo 2018). The MF uptake nutrients from soil and supplies to the plant partner
in return for carbohydrates (Smith and Read 2008) and lipids (Jiang etal. 2017;
Keymer etal. 2017; Luginbuehl etal. 2017). Consequently, the mycorrhizal sym-
biosis, signicantly inuences plant tness, as well as several ecosystem processes
such as carbon, nitrogen and phosphorous cycles, regulation of plant diversity, soil
aggregation and seedling survival (van der Heijden etal. 2015). Thus, because of
the importance of MF to plants, they cannot be ignored in the efforts to preserve
ecosystems as MTEs. However, regarding MF in the MTEs, scarce research has
been performed. In fact, in an ISI Web of Knowledge search (status May 2018), it is
possible to nd only 512 publications in a 10 year span regarding this topic.
Moreover, in the last XIV MEDECOS and XIII AEET Consortium Meeting held in
Spain in February 2017 that gathered 538 participants (Arista etal. 2017), there
were only fteen investigations dealing with mycorrhiza in MTEs– most of them
from the Mediterranean Basin (Álvarez-Garrido etal. 2017; Benito Matías etal.
2017; Parker etal. 2017; Pérez-Izquierdo etal. 2017; Rincón Herranz etal. 2017;
Romero Munar etal. 2017; Verdú 2017; Calviño-Cancela etal. 2017; Dias etal.
2017; Gil-Martínez etal. 2017; Hernández-Rodríguez et al. 2017; López García
etal. 2017; Marañón etal. 2017; Mediavilla etal. 2017; Navarro-Fernández etal.
2017). Recently, researchers have started to focus on MF in the central Chile
MTE.Due to this, in this chapter we aim to highlight and compile the arising and
existing knowledge on mycorrhizas of central Chile MTE.This chapter will focus
on basic and applied research on different mycorrhizal types, as well as on the
detection of knowledge gaps and proposals for future research directions.
14.2 Mycorrhizas inCentral Chile Mediterranean-Type
Ecosystem
The unique South American Mediterranean-type ecosystem is located from 30° to
36° South Latitude in the western portion of the continent (Armesto etal. 2007). It
is surrounded by the Atacama Desert in the north, by the Pacic Ocean in the west,
P. Silva-Flores et al.

279
by the Andes in the east and by the evergreen-deciduous temperate forests in the
south. This topography, which in turn produces antagonistic radiation/moistness
systems and a highly variable soil system, generates environmental gradients that
have produced highly rich plant communities (Armesto etal. 2007). Also, several
types of plant communities occur here, which altogether are known as Chilean
matorral (Armesto etal. 2007).
From a recent study it was possible to extract the proportions of the mycorrhizal
types from up to 1576 native plant species (from a total of 1591) of the Chilean
matorral (Silva-Flores etal. unpublished work). On that study, the proportions of
mycorrhizal types were calculated at three resolution levels (Fig.14.1). The rst one
calculated the proportions using an empirical approach (Bueno et al. 2018), i.e.
using published in peer-reviewed literature in where the mycorrhizal type of a plant
species was assessed through empirical methods (Fig.14.1a). With this approach it
was possible to assess the mycorrhizal type of only 17.3% of the plant species in the
Chilean matorral (Fig.14.1a). The other two levels of resolution used to assess the
mycorrhizal type of the plant species, calculated the proportions using a taxonomic
approach (Bueno etal. 2018), i.e. extrapolating a certain plant species mycorrhizal
type to a taxonomically and phylogenetically related plant species. Extrapolating
information from plant species to plant genus (Fig.14.1b) and subsequently to plant
family (Fig.14.1c), allowed to assess the mycorrhizal type of 78% and 99% (at
genus and family level of resolution respectively) of the plant species of the Chilean
matorral. From the results of that study was possible to extract that the AM type is
in higher proportion relatively to the other mycorrhizal types, independent of the
level of resolution (Fig.14.1). The other mycorrhizal types proportions vary depend-
ing on the level of resolution. In order to learn the exact proportions of all mycor-
rhizal types in the Chilean matorral more empirical data are urgently needed, since
Fig. 14.1 Proportions of mycorrhizal types in the Chilean matorral. Proportions were calculated
from: (a) empirical data of plant species, (b) extrapolation from plant species to genus, and (c)
extrapolation from plant genus to family. AM: arbuscular mycorrhiza, EcM: ectomycorrhiza, ErM:
ericoid mycorrhiza, OrM: orchid mycorrhiza and NM: non-mycorrhiza
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280
it has been recently showed the inaccuracy of the taxonomic approach to describe
plant mycorrhizal types (Bueno etal. 2018). To learn the mycorrhizal type distribu-
tion patterns in plant species can indicate the relationship strength between plant
communities and mycorrhizas (Moora 2014), potentially regulating both the mycor-
rhizal and plant community (Neuenkamp etal. 2018). Consequently, this informa-
tion might be useful in developing tools for restoration of degraded ecosystems as
the Chilean MTE.
It is important to highlight from the above-mentioned research (Silva-Flores
etal. unpublished work) that even with the extrapolation of mycorrhizal type data
from plant species to plant families, there are still 17 plant species in the Chilean
matorral where the mycorrhizal type is absolutely unknown. Two of these species
are in the Chilean national regulation of classication of species according to their
conservation status. The species Berberidopsis corallina Hook. f.
(Berberidopsidaceae) and Gomortega keule (Molina) Baill. (Gomortegaceae) are
both classied as endangered; thus, it would be essential to clarify their mycorrhizal
type, mycorrhizal abundance and other aspects on mycorrhizal biology in order to
assess the relative importance of the MF in the recovery of these plant species.
14.3 Arbuscular Mycorrhiza intheChilean Matorral
Arbuscular mycorrhizal fungi (AMF) are found as root symbionts in 72% of land
plant species (Brundrett and Tedersoo 2018). This symbiosis, known as AM, occurs
between the roots of certain plants and the hyphae of fungi from the Phylum
Glomeromycota (Tedersooetal.2008). With the currently available research, it is
known that the AM symbiosis is the more frequent across the plant species of the
Chilean matorral (Fig.14.1) and consequently probably a key component for this
ecosystem. Despite this, there is scarce information on the topic, probably due the
lack of awareness. In fact, to our knowledge, there are only ve published studies
regarding basic research on AM symbiosis related to the Chilean matorral
(Casanova- Katny et al. 2011; Torres-Mellado et al. 2012; Marín et al. 2017;
Benedetti et al. 2018; Silva-Flores et al. 2019) and two on applied research
(Curaqueo etal. 2010, 2011).
14.3.1 Basic Research onAMSymbiosis intheChilean
Matorral
One study explored the unknown mycorrhizal type of 10 plant species of the
Amaryllidaceae family: Gilliesia curicana, G. graminea, G. montana, Miersia
chilensis, M. leporina, M. myodes, M. tenuiseta, Solaria atropurpurea, S. miersioi-
des and Speea humilis (Torres-Mellado etal. 2012). The study found that all the
plant species had an association with the AM type with a mean colonization
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281
percentage of 45%. The study has an impact for conservation strategies of those
plant species since all of them are either vulnerable or endangered. The authors
concluded that because the plants have a high mean mycorrhization level, they
should be highly AM dependent and thus the associated AMF should be considered
in conservation strategies as plant propagation. The study also suggests that the next
step should be the identication of AMF species associated to the plant species in
order to successfully use it in conservation programs.
In the same line, another investigation also explored the mycorrhizal type of 23
Andean plant species, 18 of which belonged to the AM type (Casanova-Katny etal.
2011). This was done in order to show that cushion-associated plants had a higher
AM colonization in comparison with the same plant species growing in bare soil.
One study explored the factors affecting AMF communities in ten Chilean
Nothofagus forests (Marín etal. 2017), included three sampling plots that according
to Armesto etal. (2007) can be considered as part of the Chilean matorral. One plot
was from Reserva Natural Los Ruiles and was dominated by N. alesandrii (P1). The
other two plots were from Parque Nacional La Campana, one plot dominated by N.
macrocarpa (P2) and the second by Luma apiculata and Peumus boldus (P3). In
that study, AMF communities were studied through the analysis of spores extracted
from soil samples. Only Glomus sp. was found in P1 and P2, whilst in P3 was also
present together with G. diaphanum. Consequently, P3 resulted to have a higher
diversity than P1 and P2, whilst the community composition of P1 and P2 were
similar, but also different from P3. The content in the soil of plant available phos-
phorus and magnesium were the main edaphic variables affecting the AM fungal
community composition in those three plots. The higher spore abundance was in P2,
followed by P3 and P1.
A recent published study, quantied and morphologically identied the AMF
spores associated to nine sites of P. boldus forests (Benedetti etal. 2018). They
reported a total of 23 AM fungal species considering all sites. Funneliformis badium
was present in all sites and with a high abundance relative to the other AM fungal
species. In contrast, Septoglomus constrictum was also present in all sites but with
a low abundance. F. mosseae, Acaulospora spinosa and Rhizophagus irregularis
were also frequent species considering all sites.Finally, in the VI Region of
Libertador Bernardo O’Higgins, the role of biotic and abiotic factors in regulating
soil AMF spore density in two sclerophyllous shrublands were explored. The results
showed a strong regulation of climatic seasons on spore density in both shrublands,
in contrast to plant host species that did not have an effect on soil spore density in
any of the shrublands. Soil factors as clay content, electrical conductivity, soil
organic matter and available phosphorus and nitrogen also affected AMF spore den-
sity (Silva-Flores etal.2019).
There are also several ongoing studies with unpublished data or submitted results
regarding AM symbiosis on the Chilean matorral. In this respect, a study performed
in the Reserva Río Los Cipreses ecosystem (34°27′54″S 70°27′18″W) allowed sci-
entists to have a rst screening of the diversity of AMF in the upper part of the
Cachapoal river (Aguilar etal. unpublished data). Three plant formations were
studied in this ecosystem: the Austrocedrus chilensis, the Matorral and the Espinal.
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282
The main AMF genera found in all three plant formations were Glomus, Acaulospora
and Archeospora (Fig. 14.2). Moreover, the Matorral plant formation showed a
higher species richness relative to the Espinal and Austrocedrus chilensis (Fig.14.2).
The tree plant formations had different species composition (Fig.14.2). Also, a high
level of AMF colonization was observed in the roots of all three plant formations;
Austrocedrus chilensis (90%), the Matorral (75%) and the Espinal (65%). Finally, a
positive correlation was observed between the diversity of AMF and edaphic fac-
tors, such as nitrogen and phosphorus concentration, available and exchangeable
potassium and soil organic matter percentage.
Another study performed in the VI Region of Libertador Bernardo O’Higgins in
central Chile, aimed to assess the mycorrhizal type of the dominant plant species of
the sclerophyllous shrubland plant formation (Silva-Flores etal. submitted). It was
possible to stablish that P. boldus, Kageneckia oblonga, Escallonia pulverulenta,
Quillaja saponaria and Cryptocarya alba were all AM plant species. In Lithrea
caustica it was possible to observe AMF hyphae only in the surface of roots thus
further analyses are recommended to accurately assess whether is an AM plant.
Finally, another study explored the variation of the molecular AMF community
of the sclerophyllous shrubland in relation with host plant species, compartment
(root or soil), physico-chemical soil factors and seasons (Silva-Flores etal. unpub-
lished data). So far, a richness of 153 virtual taxa (VT) has been found. Also, the
main AMF genera found were Glomus, Claroideoglomus and Paraglomus. AMF
richness was regulated by host plant species, while AMF community composition
was regulated by seasons, host plant species, soil compartment (root or soil) and
some physico-chemical soil factors.
All the studies above described indicate that AMF are highly present in the
Chilean matorral and consequently playing an important role on this MTE.However,
all this research is in a descriptive stage and further studies quantifying the AMF
Fig. 14.2 Percentage of AMF spore genera in 100 g of soil in each plant formation studied
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283
contribution to ecosystem processes should be performed, in order to develop ef-
cient conservation strategies for this MTE.
14.3.2 Applied Research onAMF intheChilean Matorral
Two studies related to AMF and applications of it have been performed in the
Chilean Mediterranean agroecosystems. The rst one evaluated the effect of no till-
age and conventional tillage on soil organic matter, arbuscular mycorrhizal hyphae
and soil aggregates (Curaqueo etal. 2010). Tillage affected the quantity and quality
of soil organic matter, as well as AMF activity, glomalin content, and soil aggrega-
tion. No tillage produced higher values of hyphal length and glomalin production in
contrast to conventional tillage. Thus, no tillage favors soil aggregation and conse-
quently contribute to the stability of organic matter of the Mediterranean agroeco-
systems. The second study explored the effect of conventional tillage and no tillage
for 6 and 10 years on AMF propagules (spore density and total and active fungal
hyphae) and glomalin content (Curaqueo etal. 2011). AMF propagules and gloma-
lin content resulted to be higher in a 6 year no tillage system compared with a con-
ventional tillage system and 10 years no tillage system, suggesting that the positive
effects of no tillage system for 6 year vanished after certain time.
Finally, Aguilar etal. (unpublished data) studied the effect of two different agri-
cultural managements (organic vs conventional) on the diversity of AMF present in
the soil of Mediterranean Chilean vineyards. The morphological analysis from
spores of grapevine rhizospheric soil resulted in a total of twelve morphospecies of
AMF (Fig.14.3). Organic management had a higher species richness (11) compared
to conventional management (10). Also, the species composition was different
between managements. The organic management was composed by 2 exclusive
species (Acaulospora sp. and Pacispora scintillans) and 9 shared with conventional
management, while conventional had 1 exclusive (Claroideoglomus etunicatum)
(Fig.14.3). Finally, a molecular analysis showed that the three most common colo-
nizers of grapevine roots, independent of the management, were Funneliformis ver-
ruculosum, Septoglomus constrictum and an unknown Septoglomus sp. This study
provides valuable information since identication of AMF species have the poten-
tial for being used in sustainable management practices to improve grapevine pro-
duction in the Mediterranean region.
The investigation on AMF in Mediterranean agroecosystems shows that AM
symbiosis is also important and contributes to a better performance of the produc-
tive systems (Curaqueo et al. 2010, 2011). However, more detailed studies are
needed in order to assess their role as a provider of ecological services in, for
instance, sustainable agriculture (Johansson etal. 2004).
Finally, the recognition of the crucial role of AMF in the central Chilean MTE is
needed in order to protect the diversity of AMF populations as well as the vegetation
diversity. Additionally, it is also important to considerer the relationships between
AMF and other microorganisms (e.g. PGPR and rhizobia).
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284
14.4 Ectomycorrhiza intheChilean Matorral
In Chile, Nothofagus is the only native plant genus documented as EcM (Garrido
1988). It has 10 species, where six of them can be found in the Chilean matorral
through altitudinal gradient replacement. N. macrocarpa inhabits in small, relictual
and disjunct populations in the top of Coastal Mountain range, forming the northern
limit of genus distribution (Alcaras 2010). In contrast, N. obliqua, N. glauca, N.
alessandri, N. alpina, and N. dombeyi inhabit the altitudinal intermediate zones.
Several studies on Nothofagus forests in central Chile revealed the presence of
43 species of ectomycorrhizal fungi (Fig.14.4), divided in 3 Orders, 9 Families, and
13 Genera (Singer 1969; Moser and Horak 1975; Horak 1980; Garrido 1985, 1988).
Cortinarius is the dominant fungal genus, comprising 56% of the total species rich-
ness (Table14.1). In addition, these forests are characterized by a high degree of
fungal endemism, being 44%of the fungal species endemic to Chile, and 42% are
endemic of the South American Nothofagus forest, comprising central-south of
Chile and southwest of Argentina (Niveiro and Albertó 2012, 2013, 2014; Romano
and Lechner 2013; Romano etal. 2017).Thus, a total of 86% of EcM fungal species
found in the Chilean matorral are endemic of Nothofagus forests in southern South
America (Table14.1).
G11
50 µm
50 µm 50 µm
25 µm
25 µm
25 µm 15 µm 100 µm 25 µm
30 µm
25 µm 25 µm
25 µm
GL1 GL2GL3
GL5AC1GI2
GI3 PA1 PAR1 GL6
GL4
ab cd
hgfe
ij kl
Fig. 14.3 AMF spore morphotypes from soil of ten Chilean grapevine valleys. (a) GI1
Scutellospora sp., (b) GL1 Funeliformis verruculosum, (c) GL2 Uncultured Septoglomus, (d)
GL3, Claroideoglomus etunicatum**, (e) GL4 Uncultured Septoglomus, (f) GI2 Gigaspora sp.,
(g) AC1 Acaulospora sp.*, (h) GL5 Septoglomus constrictum, (i) GI3 Cetrospora gilmorei, (j) PA1
Pacispora scintillans*, (k) PAR1 Paraglomus sp. and (l) GL6 Sclerocystis sp. *species exclusively
from soils with organic management, ** species exclusively from soils with conventional
management
P. Silva-Flores et al.

285
Currently, macromycetes are being studied in forests dominated by N. macro-
carpa in two locations: Cerro El Roble (33°00′S, 71°00′W) and Reserva Natural
Altos de Cantillana (33°52′S, 71°00′W). We have found 17 ectomycorrhizal fungi
corresponding to 10 species of the genus Cortinarius, two species of Inocybe, two
species of Laccaria, one Hebeloma species, one Amanita species and one Paxillus
species (Dibán etal. unpublished work). Although they are in the process of taxo-
nomic determination, none of them coincide with those described in the literature
for N. macrocarpa (Singer 1969; Moser and Horak 1975; Garrido 1985). Thereby,
this study increases documented EcM fungal richness for N. macrocarpa to 28 spe-
cies in total.
Most of the species records previously mentioned are based on the presence of
ectomycorrhizal species through fruiting bodies (Singer 1969; Moser and Horak
1975; Garrido 1985), with few studies conrming the presence of the species in the
roots (Garrido 1988). Thus, in mixed forests with the presence of two or more
Nothofagus species, it makes it difcult to interpret which ectomycorrhizal species
is associated with which host species. In addition, some fungal genera are both,
ectomycorrhizal and saprotrophs (e.g. Ramaria spp.) (Tedersoo etal. 2008). Thus,
in studies based only on fruiting bodies, there is no certainty whether Ramaria spp.
are forming EcM associations or not. Consequently, one of the challenges in the
Fig. 14.4 Some native EcM species: (a) Austropaxillus statuum, (b) Cortinarius austroturmalis,
(c) C. magellanicus, and (d) Descolea antarctica
14 Mycorrhizas intheSouth American Mediterranean-Type Ecosystem: Chilean…

286
Table 14.1 List of ectomycorrhizal fungal species in four localities of the Chilean matorral, and
its distribution. En = endemic, Ch = Chile, Ar = Argentina. S1: Cerro El Roble (33°00′S, 71°00′W),
S2: Altos de Vilches (35°36′S, 71°12′W), S3: Reserva Forestal El Maule (35°50′S, 72°31′W) and
S4: Pilén (35°57′S, 72°25′W)
Species Distribution S1 S2 S3 S4
Amanita diemii Singer Ch, Ar 0 1 1 0
Amanita merxmuelleri Bresinsky & Garrido En Ch 0 0 1 0
Amanita umbrinella Gilb. Et Clel. Gondwanic 0 1 0 0
Austropaxillus boletinoides (sing.) Bresinsky & Jarosch En Ch, Ar 0 0 1 0
Austropaxillus statuum (Speg.) Bresinsky & Jarosch En Ch, Ar 0 1 1 0
Boletus araucarianus Garrido En Ch 0 0 1 0
Boletus putidus Horak En Ch 0 1 0 0
Butyriboletus loyo Philippi En Ch 0 1 0 0
Cortinarius albocanus (Horak & Moser) Peintner & Moser Gondwanic 0 1 1 0
Cortinarius albocinctus Moser En Ch, Ar 1 0 0 0
Cortinarius amoenus (Moser & Horak) Garnier En Ch, Ar 0 0 1 0
Cortinarius argillohygrophanicus Moser & Horak En Ch, Ar 1 0 0 0
Cortinarius aridus Moser En Ch 1 0 0 0
Cortinarius austroturmalis Moser & Horak En Ch, Ar 0 1 1 1
Cortinarius austroturmalis var. austroturmalis En Ch 1 0 0 0
Cortinarius brevisporus Moser En Ch 1 0 0 0
Cortinarius cauquenensis Garrido En Ch 0 0 1 0
Cortinarius coigue Garrido En Ch 0 0 1 0
Cortinarius columbinus Moser & Horak En Ch, Ar 1 0 0 1
Cortinarius darwinii Spegazzini En Ch, Ar 0 1 0 0
Cortinarius elaiotus Moser & Horak En Ch 0 0 0 1
Cortinarius gracilipes Moser En Ch 0 0 0 1
Cortinarius hualo Garrido En Ch 0 0 1 0
Cortinarius magellanicus Spegazzini Native 0 0 1 0
Cortinarius maulensis Moser En Ch, Ar 0 0 0 1
Cortinarius pachynemeus Moser En Ch 1 0 0 0
Cortinarius paguentus Garrido & Horak En Ch 0 0 1 0
Cortinarius roblemaulicola Garrido & Horak En Ch, Ar 0 0 1 0
Cortinarius teraturgus Moser Gondwanic 1 0 0 0
Cortinarius teresae (Garrido) Garnier En Ch 0 0 1 0
Cortinarius tumidipes Moser En Ch, Ar 0 1 0 0
Cortinarius viridurifolius Moser En Ch 0 0 0 1
Descolea antarctica Singer En Ch, Ar 1 1 1 0
Inocybe neuquenensis Singer En Ch, Ar 0 0 1 0
Laccaria ohiensis (Mont.) Singer Broad 0 0 1 0
Paxillus aff involutus (Batsch ex Fr.) Fr. Broad 1 0 0 0
(continued)
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study of EcM in the Chilean matorral is to combine the taxonomy of fruiting bodies
together with direct observation of the roots, and to sequence the described species
to extend the genetic database. Another challenge is to increase sampling locations,
especially in forests of N. alessandri and N. macrocarpa, that are scarcely sampled
and they are both classied as endangered species (Benoit 1989).
14.5 Orchid Mycorrhiza intheChilean Matorral
The Orchidaceae family forms an exclusive type of mycorrhiza, called the orchid
mycorrhiza (OrM). In this association, orchids interact with a polyphyletic group of
life-free saprophytic fungi called Rhizoctonia that includes fungi from three basid-
iomycetes families: Tulasnellaceae, Ceratobasidiaceae and Sebacinaceae (Dearnaley
etal. 2012). In addition to the exchange of nutrients and carbon between fungi and
adult plants (Cameron etal. 2006), OrM are crucial for orchid germination and
seedling development. Orchid seeds are extremely small and lack of energy reserves
(Arditti and Ghani 2000), so they require associating with MF that provide the nutri-
ents and carbon needed to germinate (Rasmussen 2002). This process, known as
symbiotic germination, is one of the dening characteristics of Orchidaceae
(Givnish etal. 2016) and it means that all orchids are mycoheterotrophic (MHT) at
least in one stage of their life. Most orchids are autotrophic at adulthood, but there
are some species that remain fully MHT throughout life (Leake 1994). Furthermore,
some green orchids species present a third nutrition mode called partial MHT or
mixotrophy, in which they obtain carbon from MF and from photosynthesis (Selosse
and Roy 2009; Hynson etal. 2013).
Although Orchidaceae has a tropical center of diversity, it shows a considerable
secondary diversity outside tropical regions (Dressler 1981). This is the case of
Mediterranean ecosystems, where the scarce orchid ora of Southern California is
an exception compared to Mediterranean Australia, Chile, south Africa and south-
ern Europe (Bernhardt 1995).
Table 14.1 (continued)
Species Distribution S1 S2 S3 S4
Russula austrodelica Singer En Ch 0 1 1 0
Russula nothofaginea Singer En Ch, Ar 0 0 1 0
Stephanopus vilchensis Garrido & Horak En Ch 0 1 0 0
Tricholoma cortinatellum Singer En Ch, Ar 0 1 1 0
Tricholoma fagnani Singer En Ch, Ar 0 0 1 0
Tricholoma fusipes Singer En Ch, Ar 0 0 1 0
Zelleromyces alveolatus (sing. & Sm.) Trappe, Lebel &
Castellano
En Ch 1 0 0 0
TO TAL 11 12 20 6
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288
There are 42 orchid species that show continuous or partial distribution through
the Chilean matorral (Novoa etal. 2015), all of them terrestrial and photosynthetic.
Little is known about OrM associations in Chile (Herrera etal. 2019), and even less
in the Chilean matorral, however the evidence available shows these orchids form
associations mainly with fungi form the families Tulasnellaceae and
Ceratobasidiaceae (Herrera etal. 2017). These ndings support observations from
other Mediterranean zones that show that Tulasnellaceae and Ceratobasidiaceae
fungi are the main associates in a high number of orchid species (Girlanda etal.
2011; Jacquemyn et al. 2015). Interestingly, there is no record of Sebacinaceae
forming OrM in Chilean matorral. This result agrees with other studies in Southern
South America (Fracchia et al. 2014a, b) but differs from observations in
Mediterranean Basin (Girlanda etal. 2011). Further studies assessing the distribu-
tion of OrM fungi in soils would help to elucidate the causes of the lack of
Sebacinaceae. Nevertheless, more research on other orchid species including more
populations is needed to conrm this pattern.
Another exciting observation is the variation in the degree of specicity among
orchid species of Chilean matorral. Specicity ranges from generalist associations
like in Chloraea longipetala (Herrera etal. 2017) and Bipinnula mbriata (Steinfort
etal. 2010) to more specialists, as observed in Chloraea gavilu (Herrera etal. 2017).
This agrees with variation on mycorrhizal specicity observed among orchids spe-
cies from Mediterranean Australia (Bonnardeaux etal. 2007; Swarts and Dixon
2009). Additionally, variation in specicity among populations of the same species
was observed in Bipinnula mbriata and B. plumosa, which was related to changes
in soil nutrient availability (Mujica etal. 2016). In the last decade, it has been an
increasing effort to identify OrM in Chilean matorral (Herrera etal. 2019). However,
further studies are required to expand this knowledge and to allow comparisons
between Mediterranean climates. For example, to our knowledge, there is no evalu-
ation of nutritional modes of Chilean matorral orchids, while mixotrophy has been
detected in Mediterranean Basin orchids (Liebel etal. 2010; Girlanda etal. 2011).
This is particularly interesting considering that this nutritional mode might be more
frequent in green orchids than previously thought (Gebauer etal. 2016). There is a
lot to be done in the study of Chilean matorral OrM, especially bearing in mind that
most of Chilean orchids are endemic and insufciently known or in some degree of
threat (Novoa et al. 2015; Herrera et al. 2019); and mainly considering that
knowledge on OrM is crucial for successful strategies in orchid conservation (Batty
etal. 2002; Swarts and Dixon 2009).
14.6 Final Considerations
Mycorrhizal research in the Chilean matorral is evidently scarce. However, an
emerging interest is arising from several researchers– mainly in AM, EcM and
OrM. South America, in general, with their contrasting mycorrhizal patterns in
comparison with the northern hemisphere, climatic conditions and other features
P. Silva-Flores et al.

289
have the potential for new, interesting discoveries (Bueno et al. 2017), and, of
course, the South American MTE is not distant to this option.
AMF research in the Chilean MTE requires the increase of sampling efforts in all
plant communities that constitute the Chilean matorral– with both complementary
morphological and molecular approaches– not only from the soil compartment, but
also from the direct observations of roots. Studies should be done in order to pro-
mote the conservation of AMF with their respective plant hosts. Also, more research
is needed to encourage sustainable agriculture since most of the plants of productive
interest have AM.The Chilean MTE is under high agriculture pressure thus, conser-
vation and production should nd an equilibrium; and, through AM symbiosis
research, this aim could be reached.
Ectomycorrhizal research, is based mainly on fruiting bodies. Thus, here also
direct morphological observation of roots is needed, as well as the use of molecular
approaches in order to increase the knowledge– at least in terms of diversity.
Orchid mycorrhizal research has been focused on the fungal diversity associated
with this symbiosis. However, more studies are needed in this respect since the
orchids of the Chilean matorral are endemic and many are threatened; thus, OrM
research will aid orchid conservation.
It is important to highlight the lack of studies in ErM not only for the Chilean
Matorral, but also at a national level. Thus, an urgent call is made regarding this
mycorrhizal type.
Mycorrhizal research in the Chilean MTE and, in general, in South America is
emerging and lling basic knowledge gaps through ecological diversity studies.
However, in the future, the integration of physiological studies in order to quantify
the contribution of mycorrhizas to ecosystem processes will be necessary as well as
the use of molecular approaches to understand the mechanism of the ecological pat-
terns that we are nding. Lastly, it is worth mentioning that the Chilean matorral is
a biodiversity hotspot; thus, all the mycorrhizal knowledge will be useful for con-
servation purposes, as well as the restoration of already degraded plant communities
of this ecosystem that are constantly submitted to anthropic negative pressure.
Acknowledgements Patricia Silva-Flores was funded by the National Doctorate Grant N°
21140639 of CONICYT and CONICYT Regional/CEAF/R08I1001. P.S.F. also thanks the support
of the Roberto Godoy regular FONDECYT 1190642. Ana Aguilar was funded by the National
Doctorate Grant N° 21120047 and N° 81150505 of CONICYT and VI Scientic Research Fund of
Pacic Hydro SA.A.A. also thanks the support of the regular postdoctoral 2018 grant of the
Ponticia Universidad Católica de Valparaíso. María José Dibán was funded by Luis Felipe
Hinojosa FONDECYT 1150690 and AFB170008. M.J.D also thanks to Dr. Götz Palfner, co-
supervisor of Master Thesis, specically in guiding taxonomic identication of some species.
María Isabel Mujica thanks to CONICYT for the National Doctorate Grant N° 21151009.
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290
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