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4. Marco teórico

4.4 Metacognición

4.4.1 Habilidades Metacognitivas

Species of Pseudotomentella, Thelephora, Tomentella and Tomentellopsis of the family

Thelephoraceae (Thelephorales, Agaricomycetes) have been reported to form

ectomycorrhizae (ECM). Some ECM not identified to species-level but considered as being caused by Thelephoraceae, are described as Afzeliaerhiza, Fagirhiza, Piceirhiza, Pinirhiza, Populirhiza, Quercirhiza, and Uapacaerhiza (Table 1). Only a few ECM studies considered

ECM of these genera show plectenchymatous outer mantle layers with or without ring-like arrangement. Pseudoparenchymatous mantles are unknown yet.

Tomentella species belong to the most frequently studied ECM fungi because they are

common and often dominant on the root system of coniferous and deciduous forests (Jakucs & Erős-Honti 2008, Kõljalg et al. 2000, Tedersoo et al. 2003, Tedersoo et al. 2007a). Some studies have been published to provide features that could generally characterize Tomentella

ECM. Agerer (2006) referred to (a) dark brown ECM with pseudoparenchymatous mantles and clamps or with cystidia, (b) dark brown ECM with pseudoparenchymatous mantles and blue granules that turn green in KOH, irrespective of the presence of clamps or cystidia. Jakucs & Erős-Honti (2008) reviewed brown ECM that have been characterized in detail and either fit the features compiled by Agerer (2006) or have been proven by nuclear rDNA (nrDNA) analysis as representing a Tomentella species. They compiled further distinctive

characteristics, like angular or star-like arranged outer mantle cells, groups of globular cells on the mantle surface, clamped cystidia, and nodally ramified rhizomorphs densely entwined with very thin peripheral hyphae.

More than 30 Tomentella or Tomentella-like ECM (Tab. 1) have been studied in detail

microscopically since 1988. Most of them share common features summarized by Agerer (2006) and by Jakucs & Erős-Honti (2008). Some, however, that have been identified as

Tomentella species using nrDNA sequence analyses, present plectenchymatous outer mantle

layers as known in other ectomycorrhizal genera of Thelephoraceae. How to identify these thelephoroid ECM morpho-anatomically is still an open question. It appears therefore helpful for future diversity studies to provide a traditional key of determination for thelephoroid ECM, in addition to the already existing and continuously updated synoptic key for all ECM described to date that includes thelephoroid ECM as well (Agerer & Rambold 2004–2009).

After the first detailed description of Tomentella-like ECM on Chinese Pine (Wei & Agerer

2008), some additional Tomentella-like ECM have been found. We present three of them in

this study, compare anatomically all ECM of Thelephoraceae (Tab. 1) and present a key to facilitate their identification.

Table 1 Tomentella and Tomentella-like ECM described morpho-anatomically in detail

since 1988

ECM or fungal partner Host Collecting sites References

Afzeliaerhiza beninensisAfzelia africana Smith Benin, Borgou Yorou & Agerer 2008

Fagirhiza asteromustrataFagus sylvatica L. Hungary, Bükk-Őserdő Jakucs et al. 2008

Fagirhiza setiferaFagus sylvatica L. Germany, Baden-

Württemberg

Brand 1991 “Fagirhiza spinulosaFagus sylvatica L. Germany, Baden-

Württemberg

Brand 1991 “Fagirhiza stellataFagus sylvatica L. Italy, Trient Di Marino et al. 2008

Piceirhiza obscuraPicea abies (L.) H. Karst. Germany Gronbach 1988 “Piceirhiza nigraPicea abies (L.) H. Karst. Germany Berg 1989, Gronbach 1988,

Haug & Pritsch 1992 “Pinirhiza ligulataPinus sylvestris L. Poland, Zmiaca Mleczko 2004b “Pinirhiza amyloideaPinus sylvestris L. Poland, Zmiaca Mleczko 2004a “Pinirhiza cyaneoviridisPinus sylvestris L. Germany, Thuringia Golldack et al. 1998 “Pinirhiza dimorphaPinus sylvestris L. Germany, Brandenburg Golldack et al. 1999 “Pinirhiza tomentelloidesPinus tabulaeformis Carr. China, Ningxia Wei & Agerer 2008

Populirhiza asperulaPopulus alba L. Hungary, Tompa Jakucs et al. 2005 Pseudotomentella humicola

M.J. Larsen Picea abies

(L.) H. Karst. Norway, Akershus Di Marino et al. 2007

Pseudotomentella tristis

(P. Karst.) M.J. Larsen

Salix herbacea L. Italy, Südtirol Agerer 1994 “Quercirhiza ateracusrugosaQuercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006a

Quercirhiza auraterocystidiataQuercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006b “Quercirhiza cumulosaQuercus ilex L.

subsp.ballota (Desf.) Samp

Spain, Navarra De Roman et al. 2002a “Quercihiza flavocystidiataQuercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006c

Quercirhiza nodulosomorphaQuercus suber L. Portugal, Grândola Azul et al. 1999

Quercirhiza squamosaQuercus robur Slovenia Palfner & Agerer 1996 “Quercirhiza summatriangularisQuercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006d “Quercirhiza stellataQuercus ilex L.

Subsp ballota (Desf.) Samp

Spain, Navarra De Roman et al. 2002b “Quercirhiza tomentellocumulataQuercus suber L. Portual, Concelho de

Santiago do Cacém

Azul et al. 2008a “Quercirhiza tomentellocystidiataQuercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006e “Quercirhiza tomentelloepidermoideaQuercus suber L. Portugal, Distrito de Setύbal Azul et al. 2008b

Quercirhiza tomentelloflexuosaQuercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006f

Quercirhiza tomentellofuniculosaQuercus suber L. Portugal, Distrito de Évpra Azul et al. 2006g

Quercirhiza tomentelloreticulataQuercus suber L. Portugal, Distrito de Évpra Azul et al. 2008c “Quercihiza tomentellostellataQuercus suber L. Portugal, Distrito de Évpra Azul et al. 2008d

Thelephora terrestris Ehrh. Picea abies L.

Picea sitchensis (Bong.) Carr.

Germany, Bavaria Britain

Agerer & Weiss 1989, Ingleby et al. 1990

Tomentella brunneorufa M.J.Larsen Eucalyptus sp. Australia Agerer & Bougher 2001 T. ferruginea (Pers.)Pat. Fagus sylvatica L. Germany, Bayern Raidl & Müller 1996 T. galzinii Bourdot (sub nom

Quercirhiza fibulocystidiata”)

Quercus cerris L. Hungary Jakucs et al.1997, Kõljalg et al. 2001

T. pilosa (Burt) Bourdot & Galzin Populus alba L. Hungary, Tompa Jakucs & Agerer 1999 T. stuposa (Link) Stalpers Quercus cerris L.,

Picea abies (L.) H. Karst.

Hungary, Püspökladány Jakucs et al. 2005 Germany, Bayern

T. sublilacina (Ellis & Holw.) Wakef. (Sub nom T. albomarginata )

Pinus sylvestris L. Germany, Hessen Agerer 1996a

T. subtestacea Bourdot & Galzin Populus alba L. Hungary, Tompa Jakucs & Agerer 2001

Tomentellopsis submollis (Svrcek) Hjortstam (sub nom “Fagirhiza rosea” )

Fagus sylvatica L. Germany, Bavaria Brand 1991 “Uapacaerhiza wariensisUapaca guineensis Müll.Arg. Benin, Borgou Yorou et al. 2008

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