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« Armillaria, » 50 , 96 , 12 , 6 ( ).

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– Armillaria.

– .

, Armillaria Armillaria . . , Armillaria .

28 , . . 4 , 18 28 A. borealis. .

: , , ,

...... 4 ...... 6 1 ...... 6 1. 1 ...... 6 1.2 ...... 8 1.3 ...... 10 1. 4 ...... 11 1.5 ...... 13 1.6 ITS ...... 13 1.7 IGS ...... 14 1.8 TEF1α...... 14 1. 9 ...... 15 2 ...... 21 2.1 ...... 21 2.1 ...... 21 2.3 ...... 22 2. 4 ...... 25 3 ...... 26 3.1 ...... 26 3.2 ...... ! . 3.2 ...... ! . 4 ...... 28 ...... 31 C ...... 32

…………………………………………………………………….………………………………………...…………….47

Вв

Armillaria () – . , , . Armillaria . , A. ostoyae A. mellea , , A. sinapina, A. gallica, A. cepistipes – . , Armillaria .

, , . .

Armillaria . , 28 Armillaria ITS, IGS, TEF1α .

:

• ITS, TEF1a, IGS Armillaria • •

, Armillaria .

« », ( №14.Y26.31.0004), № 0356-2019-0024 № 0287-2019-0002.

... . . , ... . . , ... . . , .

ва ча

1

1. 1 А

, , Armillaria [1, 2 , 3 4], [5, 6]. , (ITS) (IGS-1) [7], [8], [9], [10] [11] [12]. , (TEF1α) [13].

Armillaria ( Desarmillaria), 40 [14]. , , :

1. . Armillaria : ( , ) [15].

2. Armillaria , . (, , ) [16]. , ( , ) [17, 18]. Armillaria , , [19]. , , .

3. Armillaria . , . 1-5 , Armillaria . , Armillaria . , , , [20, 21] ( 1). , [22].

1 – Armillaria

1) , (n) ( ) (2n) ( ); 2) . , ; 3) , , ; 4) Armillaria , ; 5) [23].

1.2

, [24, 25]. Armillaria : , . . () , . , in vitro, , [26]. , - , A. cepistipes, A. gallica A. nabsnona [27] A. ostoyae A. gemina [28].

Armillaria ( ), [29]. , () Armillaria 1970- [30, 31]. , Armillaria . . , , A. cepistipes A. sinapina [32, 33].

, , , [34]. , Armillaria. , , , - . [35]. , [36]. . , A. borealis, -, 1- α (TEF1α), [36].

1.3

1990- Armillaria - . 1 (IGS-1), 28S 5S , (ITS), 18S 5.8S [37, 38]. Maphosa et al. [5] TEF1α Armillaria. , TEF1α, -, A. gallica A. cepistipes [39, 40]. [39, 36], .

, , Armillaria , [41]. , 835 , , Guyanagaster Cylindrobasidium Armillaria [42]. [43] [42] , Armillaria Guyanagaster 40-50 .

Armillaria : Desarmillaria, D. ectypa D. tabescens ( A. ectypa A. tabescens) Armillaria, [43].

1. 4

Mycobank (http://www.mycobank.org/) 70 Armillaria, 40 . , Armillaria ( 3), . , .

2 – Armillaria Desarmillaria [23]

* , , § , , # , .

Armillaria [44]. [45, 18]. Armillaria [46, 47]. . (, A. gallica A. cepistipes) , [15]. (, A. mellea, A. ostoyae) , [48].

1.5

, , , Armillaria . , , -, . , A. cepistipes A. sinapina A. ostoyae, A. mellea A. gallica. - [49, 50, 51].

1.6 ITS

ITS (Internal transcribed spacer) – ( 3). ITS . [52]. ITS . ITS (GenBank, EMBL DDBJ) , [53]. , Ascomycota, Penicillium, Aspergillus [54] Fusarium [55], ITS . 1.7 IGS

IGS (Intergrnic spacer) – , [56]. , 18S ( ), 5.8S 26S ( ). : (ITS) (IGS). 5S IGS - IGS1 IGS2 ( 3). IGS [57].

3 - ITS IGS [58].

1.8 TEF1α

TEF1 (translation elongation factor 1α) –

α 1, [59]. , TEF1α [60, 61].

1. 9

, , . , . [62].

1.9.1 (distance matrix)

. , , . , .

– (neighbour joining), . . . , [63].

:

1. . 2. , . 3. .

1 – (neighbour–joining) [64]

, , [65].

1.9.2

, . . , . — . . . , [66].

: 1. , . 2.

1.9.3 ,

GTR (General Time– Reversible) ( 4). : JC69, K80, F81, HKY85, TN93, GTR.

1. Jukes-Cantor (JC)

- (25%) . .

2. Felsenstein 1981 (F81)

–. 25%, .

3. Kimura 2 parameter (K80)

. .

4. Hasegawa-Kishino-Yano (HKY)

, F81 K80. , .

5. Tamura-Nei (TrN) . , .

6. General time reversible (GTR)

.

4 – [11]

, , .

, – (Maximum Likelihood, ML). , . . . , , , [67]. . , . , , . (posterior probablity), , . - [68].

1.9.4 «» «»

. «» , . , , , ( 5). , .

5 – «» [69].

«» , ( 6). .

6 – «» [66]. 2

2.1

28 Armillaria ( ) (-). in situ in vitro .1 .

2.1

in situ 3 :

1 – , ; , , ; Armillaria .

2 – , ( 7 ), , , ; Armillaria , .

3 – , , , ; (˃ 7 ); ( 1 ); Armillaria.

in vitro Abies sibirica 90 . . , Populus tremula, Picea abies . 3 :

1) 90 Abies sibirica:

 1 – 10 % ;  2 – 20-30 % ;  3 – 40-50 % .

1) Populus tremula:

 1 – ;  2 – , ;  3 – , .

2) Picea abies:

 1 – 10-20 % ;  2 – 25-40 %;  3 – 40 %.

2.3

DNeasy Plant Mini Kit (Qiagen Inc. Germantown, MD) ( 1) , 2. 1 –

95 °C 15 94 °C 1 50 °C 1 72 °C 1 2- 4- 35 72 °C 20 16 °C

2 –

№ (5'->3') 1 ITS1 TCCGTAGGTGAACCTGCGG [70] 1 ITS4 TCCTCCGCTTATTGATATGC 2 ITS1-F CTTGGTCATTTAGAGGAAGTAA [71] 2 ITS4-B CAGGAGACTTGTACACGGTCCAG 3 EF595F CGTGACTTCATCAAGAACATG [72] 3 EF1160R CCGATCTTGTAGACGTCCTG 4 Arm EF1-a-FOR GGAACTGGTGAGTTCGAAGCC [73] 4 Arm EF1-a-REV AGACGGAGAGGCTTGTCG GAG 5 IGS1 LR12R CTGAACGCCTCTAAGTCAGAA [74] 5 IGS1 O–1 AGTCCTATGGCCGTGGAT

ITS, TEF-1a IGS 28 ABI Genetic Analyzer 3130xl (Applied Biosystems, Foster City, USA) () , , Armillaria GenBank (www.ncbi.nlm.nih.gov/genbank/). , , , Armillaria, GenBank. .

8 Armillaria 1 Guyanagaster necrorhizus ( 3). , Armillaria . .2 .

3 – TEF1a, ITS, IGS GenBank

К

A. borealis 4 A. gallica 10 A. cepistipes 12 A. mellea 8 A. tabescens 6 A. sinapina 4 A. ostoyae 11 A. nabsnona 4 G. necrorhizus 1 60

2. 4

Ugene v. 34.0 [75] MUSCLE, [76]. - TEF1a Mega7 [77]. , , .. Seaview v. 3.2 [78].

, . , , PartitionFinder2 v. 2.1.1 [79] AIC ( ) [80]. ModelFinder [81]. PartitionFinder2 3 (TEF1a_1, TEF1a_3), (ITS, TEF1a_2) IGS. TRNEF+I+G (TEF1a_1, TEF1a_3), K81UF+I+G (ITS, TEF1_2) TRN+G IGS. : TNe+G4 TEF1α HKY+F+G4 ITS IGS.

c IQ-TREE v 1.6.12, [82]. 1000 .

, , R seqinr [83].

3

3.1

28 Armillaria, ( ) (-). ITS, IGS, TEF1α 28 5 ( 7). 5 A. cepistipes, 2 A. gallica, 3 – A. ostoyae. A. borealis 2 : , 17 , , 1 . 3.

3 – ,

К A. cepistipes 14, 13, 22, 11, 2 A. gallica 7, 8 A. ostoyae 12, 10, 9 20, 18, 3, 4, 21, 6, 28, 5, 23, 27, 26, A. borealis 25, 15, 19, 1, 24, 16 A. botealis 2 17

7. 4

Armillaria . A. borealis, A. gallica, A. ostoyae A. cepistipes, [84].

. , ITS IGS, , , A. ostoyae / A. borealis A. gallica / A. cepistipes / A. sinapina, .. . , A. ostoyae A. borealis . , [85]. , A. cepistipes A. sinapina [86]. TEF1α , . , TEF1α , ITS IGS [86, 44, 40].

TEF1α, , , A. borealis . . , [87, 88]. , . , A. borealis , , A. borealis TEF1α, [89].

Armillaria borealis . , , , . , , - , Armillaria .

A. cepistipes , . . A. borealis A. cepistipes , (mating test) [90].

A. ostoyae, , , , . ( 1 ) , A. borealis.

- A. ostoyae, [91, 92]. , A. borealis . , A. borealis , A. ostoyae, , A. borealis [93, 94]. -, A. borealis , , , , [93].

A. borealis , , A. borealis , A. ostoyae [95], in vitro A. borealis.

Armillaria gallica , . A. cepistipes. , , A. gallica A. cepistipes, , (A. mellea, A. borealis A. ostoyae) [95].

11 10 (A. ostoyae A. cepistipes ) . Armillaria , A. cepistipes A. ostoyae [96]. , , .

Зач

Armillaria. . 4 , 18 28 A. borealis. .

, TEF1α A. borealis. , . , .

C вах чв

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.1 –

in vitro ()

90

Populus Populus

-

Abies sibirica Abies Picea abies Picea situ in ; 3 ;

30 30 ; 2 – ; 1- 30 tremula tremula () () К / , 1 2 3 1 A.borealis 56° 1.873' 92°34.403' Pinus 2 2 3 2 sylvestris L.

2 - A.cepistipes 56°11.292' 92°12.365' Abies sibirica 1 1 1 1 Ledeb. 3 A.borealis N55°59.684' E92°38.156' Populus 0,2 2 2 2 2 tremula L. 4 , A.borealis 55°59.211' 92°37.905' Betula 2 3 2 2 pendula Roth 5 A.borealis N55°59.7' E92°38.2' Populus 0,3 2 2 2 2 tremula L. 6 A.borealis N55°58.3’ E92°37.5’ Populus 0.1 2 3 3 3 tremula L. 7 A.gallica N44° E 135° Abies 2.1 2 2 2 1 , 31,647' 16,980' nephrolepis - (Trautv. ex Maxim.) Maxim. 8 . , A.gallica N52°13.990' E89°39.105' Abies sibirica 1 2 1 1 Ledeb. 9 A.ostoyae N44°38.142' E135°10.465' Abies 4,8 3 3 2 2 , nephrolepis - (Trautv. ex Maxim.) Maxim.

10 A.ostoyae N44°31.723' E135°17.314' Abies 16 3 3 3 2 , nephrolepis - (Trautv. ex Maxim.) Maxim. 11 A.cepistipes N44°31.7' E135°17.3' Abies 1 1 1 0 , nephronlepis - (Trautv. Ex Maxim.) Maxim. 12 A.ostoyae N44°38.439' E135°10.021' Picea 4,8 3 3 3 3 , jezoensis - (Siebold & Zucc.) Carrière 13 A.cepistipes N44°31.637' E135°16.513' Abies 1 1 0 0 , nephronlepis - (Trautv. Ex Maxim.) Maxim. 14 A.cepistipes N54°30.643’ E88°49.912’ Pinus sibirica 1 2 1 2 Du Tour 15 A.borealis N54°38.714' E93°18.074' Picea 16,1 3 3 3 3 obovata Ledeb. 16 - A.borealis N52°26.098’ E88°56.428’ Abies sibirica 20.4 3 3 3 2 Ledeb.

17 A.borealis N55°58.3’ E92°37.5’ Pinus sibirica 3 2 2 2 Du Tour

18 A.borealis 56°0.644' 92°36.342' Pinus 1,1 3 3 3 3 sylvestris L. 19 , A.borealis N54°31.347' E93°23.904' Abies sibirica 20,2 3 3 2 2 Ledeb. 20 A.borealis N54°41.258' E93°11.737' Abies sibirica 14.6 3 3 2 3 Ledeb.

21 A.borealis N55°58.1’ E92°37.1’ Populus 0.1 2 3 3 3 tremula L. 22 A.cepistipes N44°31.637' E135°16.513' Abies 1 1 0 0 , nephronlepis - (Trautv. Ex Maxim.) Maxim. 23 A.borealis N54°38.714' E93°18.074' Picea 16,1 3 3 3 3 obovata Ledeb. 24 A.borealis 55°58.348' 92°36.474' Betula . 2 2 2 2 pendula Roth

25 A.borealis 56° 1.669' 92°34.331' Pinus 0,4 3 3 3 2 sylvestris L. 26 A.borealis 56° 1.727' 92°34.803' Populus 0,4 3 3 3 2 tremula L. 27 A.borealis N54°41.258' E93°11.737' Abies sibirica 31.8 3 3 2 2 Ledeb.

28 A.borealis N54°41.021' E93°12.368' Abies sibirica 25.5 3 3 2 2 Ledeb.

.2 – GenBank

Вид Шта tef1 ITS IGS Источик HAt1S5 HQ285906.1 HQ232292.1 HQ232284.1 Украа HAt5S3 HQ285908.1 HQ232294.1 HQ232286.1 Украа HAt2S5 HQ285907.1 HQ232293.1 HQ232285.1 Украа A. tabescens 96_1_8 AB510804.1 AB510867.1 AB510823.1 Япоя 2006-20-01 AB510807.1 AB510887.1 AB510839.1 Япоя 96_3_3 AB510805.1 AB510868.1 AB510824.1 Япоя

A_12 AB510801.1 AB510864.1 AB510820.1 Япоя 89-07 AB510796 AB510852.1 AB510808 Япоя 94-7 AB510799 AB510863 AB510819 Япоя 94-10-1 AB510798 AB510857 AB510813 Япоя

A. mellea 94-68 AB510803 AB510858 AB510814 Япоя 97-6 AB510797 AB510856 AB510812 Япоя 94_5 AB510802.1 AB510880.1 AB510833.1 Япоя A_10 AB510800.1 AB510860.1 AB510816.1 Япоя NA4 AB510761.1 AB510881.1 AB510834.1 Япоя E5 HQ285905.1 HQ232291.1 HQ232283.1 Фрая Y11DS1 JN657484.1 JN657457.1 JN657430.1 Украа E4 JN657479.1 JN657452.1 JN657425.1 Фрая E6 JN657480.1 JN657453.1 JN657426.1 Фрая A.gallica BRNM 706835 EU251390.1 EU257718.1 EU636240.1 ея HY2a JN657482.1 JN657455.1 JN657428.1 Украа HY1 JN657481.1 JN657454.1 JN657427.1 Украа C1AS JN657483.1 JN657456.1 JN657429.1 Украа NA13 AB510760.1 AB510890.1 AB510842.1 Япоя 90_10_12 AB510790.1 AB510862.1 AB510818.1 Япоя 94_46_01 AB510793.1 AB510898.1 AB510849.1 Япоя BRNM 695717 EU251396.1 EU257716.1 EU257710.1 ея S11AE JN657478.1 JN657451.1 JN657424.1 Украа A. cepistipes BRNM 706814 EU251395.1 EU257715.1 EU257709.1 ея B3 JN657472.1 JN657445.1 JN657418.1 Флядя B2 HQ285902.1 HQ232288.1 HQ232280.1 Флядя Y16AE JN657476.1 JN657449.1 JN657422.1 Украа

C13AE JN657474.1 JN657447.1 JN657420.1 Украа B5 JN657473.1 JN657446.1 JN657419.1 Италя C19AS2 JN657475.1 JN657448.1 JN657421.1 Украа C5CS1 JN657477.1 JN657450.1 JN657423.1 Украа C4 JN657487.1 JN657460.1 JN657433.1 Фрая Y17DS JN657488.1 JN657461.1 JN657434.1 Украа 2002_66_03 AB510781.1 AB510896.1 AB510847.1 Япоя NC8 AB510782.1 AB510897.1 AB510848.1 Япоя C2 JN657486.1 JN657459.1 JN657432.1 Фрая A. ostoyae BRNM 706815 EU251400.1 EU257717.1 EU257711.1 ея HpAg1 JN657489.1 JN657462.1 JN657435.1 Украа C5 HQ285903.1 HQ232289.1 HQ232281.1 Фрая 88-01-19 AB510784 AB510859 AB510815 Япоя 91-01-10 AB510785 AB510865 AB510821 Япоя D20 JN657490.1 JN657463.1 JN657436.1 веаря A618 JN657496.1 JN657469.1 JN657442.1 веаря A1 JN657494.1 JN657467.1 JN657440.1 Флядя A. borealis A5 JN657495.1 JN657468.1 JN657441.1 Герая A722 JN657497.1 JN657470.1 JN657443.1 веаря 00-16-4 AB510765.1 AB510883.1 AB510835.1 Япоя NB4 AB510764 AB510900 AB510851 Япоя A. nabsona 00-4-4 AB510762 AB510876 AB510830 Япоя 00-3-1 AB510766.1 AB510899.1 AB510850.1 Япоя 96-7-1 AB510774.1 AB510873.1 AB510827.1 Япоя 05-7-1 AB510775 AB510886 AB510838 Япоя A. sinapina 05-13-2 AB510776 AB510884 AB510836 Япоя 05-46-1 AB510778 AB510889 AB510841 Япоя G. necrorhizus RAK 31 KU289108.1 KU170948.1 KU254225.1 Гааа