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 beninensis” Afzelia africana Smith Benin, Borgou Yorou & Agerer 2008
“Fagirhiza asteromustrata” Fagus sylvatica L. Hungary, Bükk-Őserdő Jakucs et al. 2008
“Fagirhiza setifera” Fagus sylvatica L. Germany, Baden-
Württemberg
Brand 1991 “Fagirhiza spinulosa” Fagus sylvatica L. Germany, Baden-
Württemberg
Brand 1991 “Fagirhiza stellata” Fagus sylvatica L. Italy, Trient Di Marino et al. 2008
“Piceirhiza obscura” Picea abies (L.) H. Karst. Germany Gronbach 1988 “Piceirhiza nigra” Picea abies (L.) H. Karst. Germany Berg 1989, Gronbach 1988,
Haug & Pritsch 1992 “Pinirhiza ligulata” Pinus sylvestris L. Poland, Zmiaca Mleczko 2004b “Pinirhiza amyloidea” Pinus sylvestris L. Poland, Zmiaca Mleczko 2004a “Pinirhiza cyaneoviridis” Pinus sylvestris L. Germany, Thuringia Golldack et al. 1998 “Pinirhiza dimorpha” Pinus sylvestris L. Germany, Brandenburg Golldack et al. 1999 “Pinirhiza tomentelloides” Pinus tabulaeformis Carr. China, Ningxia Wei & Agerer 2008
“Populirhiza asperula” Populus 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 ateracusrugosa” Quercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006a
“Quercirhiza auraterocystidiata” Quercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006b “Quercirhiza cumulosa” Quercus ilex L.
subsp.ballota (Desf.) Samp
Spain, Navarra De Roman et al. 2002a “Quercihiza flavocystidiata” Quercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006c
“Quercirhiza nodulosomorpha” Quercus suber L. Portugal, Grândola Azul et al. 1999
“Quercirhiza squamosa” Quercus robur Slovenia Palfner & Agerer 1996 “Quercirhiza summatriangularis” Quercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006d “Quercirhiza stellata” Quercus ilex L.
Subsp ballota (Desf.) Samp
Spain, Navarra De Roman et al. 2002b “Quercirhiza tomentellocumulata” Quercus suber L. Portual, Concelho de
Santiago do Cacém
Azul et al. 2008a “Quercirhiza tomentellocystidiata” Quercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006e “Quercirhiza tomentelloepidermoidea” Quercus suber L. Portugal, Distrito de Setύbal Azul et al. 2008b
“Quercirhiza tomentelloflexuosa” Quercus suber L. Portugal, Distrito de Setύbal Azul et al. 2006f
“Quercirhiza tomentellofuniculosa” Quercus suber L. Portugal, Distrito de Évpra Azul et al. 2006g
“Quercirhiza tomentelloreticulata” Quercus suber L. Portugal, Distrito de Évpra Azul et al. 2008c “Quercihiza tomentellostellata” Quercus 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 wariensis” Uapaca guineensis Müll.Arg. Benin, Borgou Yorou et al. 2008