Taxonomic Diversity

Species diversity in eukaryotes, including fungi, inhabiting permafrost horizons, collected in the Arctic, has been the subject of intense research over the last decade (Dmitriev et al. 1997; Kochkina et al. 2001; Vishnivetskaya et al. 2003). However, the complete list of the fungi detected has not been reported. The published results are summed up in Table 7.1, which lists mycelial fungi detected in permafrost horizons of the Northern hemisphere. To make the picture complete, we also indicate reports on soils of Arctic tundra, in which the same fungal species can be found as in deep horizons of Arctic habitats. Table 7.1 shows that the fungi of Arctic permafrost exhibit considerable taxonomic diversity. Analysis of the occurrence frequencies of fungal species demonstrated that Geomyces pannorum, Cladosporium spp., and Aspergillus spp. were the most common. The genus Penicillium was represented by the greatest number of species. Interestingly, species occurring most frequently in deep horizons were also isolated from modern Arctic soils in the course of long-term studies.

Table 7.1 Fungal biodiversity in Arctic permafrosta

Species

References

Acremonium pteridii W. Gams et

Ozerskaya et al. (2008)

Frankland

A. salmoneum W.Gams et Lodha

Ozerskaya et al. (2008)

Alternaria alternata (Fries) Keissler

Kirtsidely (1999a), Ivanushkina et al. (2005),

Kurek et al. (2007)

Arthrinium sphaerospermum Fuckel

Ivanushkina et al. (2005)

Aspergillus fumigatus Fresen.

Ivanushkina et al. (2005)

A. niger van Tiegh.

Kirtsidely (1999a), Etienne (2002),

Ivanushkina et al. (2005, 2007)

A. oryzae (Ahlb.) E.Cohn

Ivanushkina et al. (2005)

A. sclerotiorum G.A. Huber

Stakhov et al. (2008)

A. sydowii (Bainier et Sattory)

Ivanushkina et al. (2005, 2007)

Thom et Church

A. versicolor (Vuill.) Tirab.

Zabawski (1982), Ivanushkina et al. (2005,

2007), Kurek et al. (2007)

Aureobasidium pullulans (de Bary)

Ivanushkina et al. (2005), Kirtsidely (1999a)

G. Arnaud var. pullulans

A. pullulans (de Bary) G. Arnaud var.

Ozerskaya et al. (2008)

melanogenum Herm.-Nijh.

Bispora antennata (Pers.) E.W. Mason

Ivanushkina et al. (2005)

Botrytis cinerea Pers.

Ivanushkina et al. (2005), Kurek et al. (2007)

Chaetomium globosum Kunze

Zabawski (1982), Ivanushkina et al. (2005)

C.indicum Corda

Ivanushkina et al. (2005)

Chaetophoma sp.

Ivanushkina et al. (2005)

Chrysosporium merdarium (Ehrenb.)

Ozerskaya et al. (2008)

J.W. Carmich

Cladosporium cladosporioides (Fres.)

Cooke and Fournelle (1960), Zabawski (1982),

G.A. de Vries

Kirtsidely (1999a), Ivanushkina et al. (2005),

Kurek et al. (2007)

(continued)

88

S. Ozerskaya et al.

Table 7.1 (continued)

Species

References

C. herbarum (Pers.) Link

Zabawski (1982), Ivanushkina et al. (2005),

Kurek et al. (2007)

C. macrocarpum Preuss

Ivanushkina et al. (2005)

C. sphaerospermum Penz.

Ivanushkina et al. (2005), Stakhov et al. (2008)

Engyodontium album (Limber) de Hoog

Ivanushkina et al. (2005)

Eurotium amstelodami L. Mangin

Ozerskaya et al. (2008)

E. herbariorum (F.H. Wigg.) Link

Ozerskaya et al. (2008)

E. rubrum W. Bremer

Ivanushkina et al. (2005)

Exophiala jeanselmei (Langeron)

Ivanushkina et al. (2007) (Langeron) McGinnis

var. heteromorpha (Nannf.) de Hoog

et A.A. Padhge

Fusarium oxysporum Schltdl.

Cooke and Fournelle (1960), Kirtsidely (1999a),

Ivanushkina et al. (2005)

F solani (Mart.) Sacc.

Ozerskaya et al. (2008)

Geotrichum candidum Link

Ivanushkina et al. (2005)

Geomyces pannorum (Link) Sigler et

Kirtsidely (1999a), Etienne (2002),

J.W. Carmichael

Ivanushkina et al. (2005), Kurek et al.

(2007), Stakhov et al. (2008)

G.vinaceus Dal Vesco

Ozerskaya et al. (2008)

Gliocladium sp.

Ozerskaya et al. (2008)

Lecythophora mutabilis (J.F.H. Beyma)

Ozerskaya et al. (2008)

W.Gams et McGinnis

Malbranchea pulchella Sacc. et Penz.

Ozerskaya et al. (2008)

Monodictys glauca (Cooke et Harkn.)

Ivanushkina et al. (2005)

S. Hughes

Mucor plumbeus Bonord.

Ivanushkina et al. (2005)

Paecilomyces variotii Bainier

Ivanushkina et al. (2005)

Penicillium aurantiogriseum Dierckx

Zabawski (1982), Kirtsidely (1999a, b),

Ivanushkina et al. (2005, 2007), Kurek et al.

(2007), Stakhov et al. (2008)

P brevicompactum Dierckx

Zabawski (1982), Kirtsidely (1999a),

Etienne (2002), Ivanushkina et al. (2005)

P. chrysogenum Thom

Zabawski (1982), Kirtsidely (1999a),

Ivanushkina et al. (2005, 2007),

Kurek et al. (2007)

P. citrinum Thom

Ivanushkina et al. (2005)

P. crustosum Thom

Ivanushkina et al. (2005)

P. decumbens Thom

Ivanushkina et al. (2005)

P. glabrum (Wehmer) Westling

Zabawski (1982), Kirtsidely (1999a, b, 2001),

Ivanushkina et al. (2005)

P. granulatum Bainier

Zabawski (1982), Kirtsidely (2001),

Ozerskaya et al. (2008), Stakhov et al. (2008)

P. griseofulvum Dierckx

Ozerskaya et al. (2008)

P. melinii Thom

Ivanushkina et al. (2007)

P. miczynskii K.M. Zalessky

Kirtsidely (1999a, b), Ivanushkina et al. (2007)

P. minioluteum Dierckx

Ivanushkina et al. (2005, 2007)

P. puberulum Bainier

Ivanushkina et al. (2005)

P. purpurogenum Stoll

Kirtsidely (1999a), Ivanushkina et al. (2005)

(continued)

(continued)

Table 7.1 (continued)

Species

References

P. restrictum J.C. Gilman et E.V. Abbott P rugulosum Thom P simplicissimum (Oudem.) Thom

P variabile Sopp P verrucosum Dierckx P viridicatum Westling Papulaspora sp.

Phialophora fastigiata (Lagerb. et Melin)

Conant P melinii (Nannf.) Conant Phoma crystallifera Gruyter, Noordel.

et Boerema Ph. destructive Plowr. Ph. jolyana Piroz. et Morgan-Jones var.

jolyana Ph. herbarum Westend. Ph. nebulosa (Pers.) Berk. Rhinocladiella atrovirens Nannf. Scopulariopsis candida (Gueg.) Vuill. Stachybotrys chartarum (Ehrenb.) S.Hughes

Sphaeronaemella mougeotii (Fr.) Sacc. Sporotrichum pruinosum J.C. Gilman et E.V. Abbott Thysanophora penicillioides (Roum.)

W.B. Kendr. Trichoderma longibrachiatum Rifai Ulocladium atrum Preuss U. botrytis Preuss Valsa sordida Nitschke Verticillium sp.

Xylohypha nigrescens (Pers.) E.W. Mason

Mycelia sterile (white; dark; dark with sclerotia)

Ivanushkina et al. (2005) Ivanushkina et al. (2005) Zabawski (1982), Kirtsidely (1999a),

Ozerskaya et al. (2008) Ivanushkina et al. (2005) Ivanushkina et al. (2005, 2007) Ivanushkina et al. (2007) Ivanushkina et al. (2005) Zabawski (1982), Kurek et al. (2007),

Stakhov et al. (2008) Ozerskaya et al. (2008) Stakhov et al. (2008)

Ozerskaya et al. (2008) Ozerskaya et al. (2008)

Stakhov et al. (2008)

Ozerskaya et al. (2008), Stakhov et al. (2008) Ivanushkina et al. (2005) Ivanushkina et al. (2007) Ivanushkina et al. (2005)

Ivanushkina et al. (2007) Ivanushkina et al. (2005)

Ozerskaya et al. (2008)

Ivanushkina et al. (2005) Ozerskaya et al. (2008) Ivanushkina et al. (2005) Ozerskaya et al. (2008) Ivanushkina et al. (2005) Ozerskaya et al. (2008)

Zabawski (1982), Kirtsidely (1999a, b);

Kurek et al. (2007), Ozerskaya et al. (2008), Stakhov et al. (2008)

aFungi from superficial horizons of Arctic landscapes of various locations were studied in the following articles: Cooke and Fournelle (1960), Zabawski (1982), Kirtsidely (1999a), Kirtsidely (1999b), Etienne (2002), Kirtsidely (2001), Kurek et al. (2007)

We attempted to compare parameters characterizing the diversity of mycelial fungi from Arctic permafrost, which were determined by (1) the conventional technique of inoculating solid nutritive media with aqueous suspensions of the specimens, and (2) the method of analyzing DNA isolated directly from the same specimens (DNA identification was performed using the database of nucleic acid sequences GenBank; Lydolph et al. 2005). In comparing the two approaches, we used samples (comprising permafrost sediments of Holocene and late Pleistocene) collected in the eastern Arctic. Only at the level of families (or higher taxa) was it possible to compare the taxonomic diversity parameters of the fungi, because the method of direct DNA isolation did not favor precise species assignment (Table 7.2).

The results of our comparative analysis made it possible to assess the concordance in species composition between the isolated complexes of mycelial fungi; for this, we used the Sorensen index of similarity, the values of which were at the level of 40%. The number of higher taxa isolated by plating was slightly lower, which

Table 7.2 Two methods of study of fungal biodiversity (DNA culture-independent method; Plate culture-dependent method)

Class

Subclass

Order

Family

Method

Ascomycetes

Dothideomyc-

Dothideales

Dothioraceae

DNA

Plate

etidae

Incertae sedis

-

Plate

Incertae sedis

Pseudoperis-

DNA

-

poriaceae

Mycosphaer

Mycosphaer-

DNA

Plate

ellales

ellaceae

Pleosporales

Incertae sedis

-

Plate

Leptosphaeriaceae

DNA

-

Pleosporaceae

DNA

Plate

Sporormiaceae

DNA

-

Erysiphomyc-

Erysiphales

Erysiphaceae

DNA

-

etidae

Eurotiales

Trichocomaceae

DNA

Plate

Incertae sedis

Incertae sedis

Incertae sedis

DNA

Plate

Myxotrichaceae

-

Plate

Pseudeurotiaceae

-

Plate

Leotiomyc-

Helotiales

Dermateaceae

-

Plate

etidae

Helotiaceae

DNA

-

Hyaloscyphaceae

DNA

-

Sclerotiniaceae

DNA

Plate

Rhytismatales

Rhytismataceae

DNA

-

Sordariomyc-

Hypocreales

Hypocreaceae

-

Plate

etidae

Incertae sedis

-

Plate

Nectriaceae

-

Plate

Sordariales

Chaetomiaceae

DNA

-

Sordariaceae

DNA

-

Basidiomycetes

Agaricomyc-

Polyporales

Corticiaceae

DNA

-

etidae

Cyphellaceae

DNA

-

Fomitopsidaceae

DNA

-

Phanerochaetaceae

DNA

-

Coelomycetes

Incertae sedis

Incertae sedis

Incertae sedis

-

Plate

Saccharomyc-

Saccharomyc-

Saccharomycetales Incertae sedis

DNA

Plate

etes

etidae

Urediniomycetes

Incertae sedis

Uredinales

Melampsoraceae

DNA

-

Zygomycetes

Incertae sedis

Basidiobolales

Basidiobolaceae

DNA

-

Mortierellales

Mortierellaceae

DNA

Plate

Mucorales

Mucoraceae

-

Plate

may be due to problems of identification of sterile mycelium. Its identification by molecular-biological techniques may well show that higher fungi are much more abundant in permafrost than is considered to be case today.

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