1991 2006 . .. *, .. * * . .. , - * vvgor@skylink.spb.ru ,* *Ira@IB6521.spb.edu - ­ , (, ) . ( ) ( ) Pleurozium schreberi, Hylocomium splendens C. Rangiferina, C. mitis Cladina stellaris . - Pinus sylvestris, P. banksiana, Larix sibirica sl. . ( 10­15%) Picea mariana, P. obovata, Pinus sibirica ( ). - . - , . ( ., 1991, 1995­1997). ( ) - . - . ( ­ ­ ) . 383 , . : . . , 20­50 . , . - ( , 1846; , 1964), . : 1) «» ; 2) . , - , , . ­ 14- . - . , , 150 300 . , . , -0.5º (- .., 1988). ­ -13º; ­ 384 +14º. 104 . . 5º ­ 80­100 . 90 ­110 . 465­500 . ( ) 339­350 . 200­250 , 2 (- .., 1988). Al-Fe- , Al-Fe- (, 1987). , ( 1991­1994 .) 5, 60, 90, 140 160 , . 2005­2006 . : 1) ( - > 70%); 2) - ( 30­70%); 3) ( < 30%). , - - . 1­3. 0,1­0,5 . ( ) 1 2 (20­100 ) 4 . : 1) , 2) ; 3) - - . , 4 . , , 50­100 , . 385 386 [3] [4] 1. (1991­1994 .) [5] , ., ., ., .-1 64.0 63.5 64.5 54.5 89.5 .. .. 40.0 52.5 II 95 98 55.0 II 57.5 II 23P77S 77P23B 100P I II I 100P 99P1S 93P7S 2 -1 , % 25 0 15 70 95[2] .. 0 99 17 56 , 65 . . 9.6 0.3 67 80 94 66 13.9 13.5 0.4 1.4 82 66 13.6 0.4 85 86 5 161/~50 [1] 0.4 0.8 0.8 <0.1 I II I I II I II I I 100P 100P 100P 100P 100P 100P 98P2B 100P 90P10B 87 63 64 163 94/~55[1] 94 10.0 24.5 18.3 0.4 0.1 0.5 75 65 10.4 <0.1 76 140 16.1 1.5 P P P P P P P P P B P P P S P S P B P 115-335 55-63 190-225 240-330 55-63 335 45-63 40-120 40-120 .. 250-290 51-83 200-224 150-180 45-60 50-80 40-63 45-64 114-137 18.0 8.0 12.3 15.4 7.5 18.1 9.3 9.5 9.5 .. 13.2 8.4 16.9 16.3 6.0 8.7 9.0 7.1 16.4 35.0 8.5 25.0 28.0 11.7 48.0 7.5 11.5 11.5 .. 23.0 6.9 28.0 23.7 5.3 11.8 11.0 7.6 23.2 72 440 283 195 88 38 1021 39 .. .. 240 6360 229 23 324 219 801 501 373 : ­ , ; 2 ­ ; 3 ­ 37­1430; 4 ­ : I ­ ; II ­ , 1,3 ; 5­ P ­ ; S ­ ; B ­ . 1 2. ( ) 1991 2006 . 387 85 56 80 82 67 86 87 1992 2005 1991 2005 1992 2006 1992 2005 1991 2005 1994 2005 1994 2005 , 5 18 65 79 66 80 66 79 94 108 163 174 163 174 ,% 95 25 15 70 0 .. 0 - 12 32 32 20 20 24 24 8 8 36 36 32 32 16 16 , % -. 8.1 13.2* 6.3 6.2 15.6 17.9 13.0 5.2* 25.1 21.0 5.7 8.9* 5.9 5.4 Vaccinium vitis-idaea 6.5 3.2* 2.6 2.9 2.9 3.4 5.5 2.5 12.3 7.9* 0.5 0.9 4.3 4.1 Vaccinium myrtillus 0.2 0.2 0.0 0.0 1.6 3.7 0.1 0.3 3.7 4.7 0.0 0.1 0.0 0.1 Empetrum hermaphroditum 0.0 0.9 0.2 1.2 0.5 1.8 5.1 1.4 5.6 5.0 0.8 1.0 0.0 0.1 Arctostaphylos uva-ursi 0.6 2.2 3.0 1.4 0.0 0.0 0.8 0.0 3.5 3.2 1.5 2.7 0.8 0.8 Calluna vulgaris 0.8 6.8*** 0.4 0.1 10.5 9.0 0.9 0.9 0.0 0.0 2.8 3.8 0.9 0.3 -. 20.5 70.6*** 89.4 90.6 88.2 84.7 92.6 89.4 78.5 81.6 91.0 92.3 94.3 92.3 46.3 82.9*** 99.1 99.4 99.2 98.0 96.8 99.8* 91.9 89.9 97.4 98.2 99.8 100 , 8.2 57.8*** 88.6 90.1 87.5 82.9 89.7 89.2 72.2 73.2 88.7 90.7 94.2 92.3 Cladina mitis 0.0 2.7*** 47.7 39.4 52.9 16.7*** 50.2 33.9* 16.5 8.2*** 1.8 0.9 1.5 1.6 Cladina rangiferina 0.0 0.7* 20.7 33.4** 21.4 52.8*** 14.0 32.9** 30.7 35.9 1.7 5.0 7.2 5.4 Cladina stellaris 0.0 0.3** 6.2 11.9** 3.5 7.4** 5.6 8.7 18.2 25.9 81.4 82.4 84.0 84.6 Cladonia uncialis 0.0 1.0*** 8.4 3.0*** 7.7 4.4* 13.4 9.9 5.3 2.4*** 0.3 0.1 0.8 0.6 Cladonia deformis 0.0 16.6*** 0.3 0.1* 0.1 0.0 0.2 0.1 0.0 0.0 1.2 0.6 0.1 0.0 Cladonia gracilis 0.0 0.3** 0.9 0.4 0.2 0.2 0.7 0.4 0.2 0.1 0.3 0.1 0.1 0.0 Cladonia crispata 0.0 11.5*** 2.1 1.2 0.9 0.7 2.4 1.8 0.4 0.1 0.8 0.9 0.1 0.0 Cladonia cornuta 0.0 4.4*** 0.8 0.3** 0.1 0.4 1.2 0.4* 0.1 0.1 0.7 0.4* 0.2 0.0 Cladonia coccifera 0.0 0.8*** 0.2 0.0 0.1 0.0 0.8 0.1* 0.0 0.0 0.0 0.0 0.0 0.0 Cladonia macrophylla 0.0 4.5*** 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Trapeliopsis granulosa 0.6 10.1*** 0.0 0.0 0.0 0.0 0.4 0.0* 0.0 0.0 0.1 0.3 0.0 0.0 Stereocaulon paschale 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.9* 0.8 0.3 0.0 0.0 0.0 0.0 , 12.3 12.8 0.8 0.5 0.7 1.8 2.9 0.1* 6.4 8.4 2.2 1.6 0.1 0.0 Pleurozium schreberi 0.0 0.0 0.0 0.0 0.4 1.4 0.0 0.1 5.4 7.2 0.4 0.3 0.0 0.0 Dicranum scoparium 0.0 0.0 0.0 0.0 0.1 0.1 0.0 0.0 0.3 0.5 0.1 0.0 0.1 0.0 Polytrichum juniperinum 4.9 3.9 0.7 0.4 0.1 0.0 0.1 0.0 0.1 0.1 1.3 0.7 0.0 0.0 Polytrichum piliferum 3.3 6.6* 0.1 0.0 0.0 0.0 1.6 0.0* 0.0 0.3 0.0 0.0 0.0 0.0 Pohlia nutans 4.1 1.0*** 0.0 0.1 0.1 0.1 0.1 0.0 0.0 0.0 0.5 0.6 0.0 0.0 : 3 , * ­ = 0.05; ** ­ = 0.01; *** ­ = 0.001. 388 1992 66 24 95 21.9 10.2** 2.5 3.2** 0.0 4.9* 0.3 0.0 0.0 74.6 53.3*** 39.1** 6.5*** 24.9*** 3.4 1.7*** 0.2** 0.3* 0.0 0.4 1.2 35.5*** 32.8*** 1.0 0.0 1.4 0.0 0.0 0.0 36.9 6.0 20.5 7.0 0.3 0.0 0.8 1.2 0.9 81.6 17.9 14.7 1.3 8.1 3.0 0.3 0.4 0.2 1.2 0.1 0.0 66.9 51.6 4.0 0.6 0.0 3.0 5.7 1.8 42.4 6.5 27.3 6.6 0.2 0.0 0.2* 0.9 0.6 84.3 14.1 12.0 1.5 5.3 3.7 0.1 0.8 0.1 0.3 0.0 0.0 72.2 54.3 5.1 0.8 0.0 4.2 7.6 0.0 37.5 6.2 24.0 6.0 0.9 0.0 0.3 0.0 0.0 78.5 26.9 20.4 1.8 10.7 5.0 2.6 0.1 0.0 0.0 0.0 0.1 58.1 55.7 1.7 0.3 0.2 0.1 0.0 0.0 35.0 4.6 26.6 3.0* 0.4 0.0 0.2 0.1 0.0 81.6 5.7*** 4.6*** 0.4** 2.1*** 2.0* 0.1*** 0.0** 0.1 0.0 0.0 0.0 77.0*** 75.7*** 1.0 0.2 0.0** 0.0 0.1 0.0 23.1 3.5 15.9 1.8 0.1 2.0 0.0 0.0 0.0 84.2 31.6 27.0 1.7 9.6 12.1 2.0 0.5 0.4 0.2 0.6 0.0 57.2 39.1 14.6 1.6 0.0 0.9 0.0 0.0 17 99 98 17.2 5.0 0.9 0.3 0.0 10.3 0.3 0.0 0.0 70.9 84.3 59.8 34.4 11.7 2.2 7.1 0.9 0.8 0.1 0.1 1.3 11.0 6.9 0.6 0.1 2.9 0.0 0.0 0.0 75 2005 79 24 1991 94 32 2005 108 32 1991 94 40 2005 108 40 1991 140 20 64 63 76 2005 154 20 27.8 5.7* 17.1 2.0 0.0 1.8 0.0 0.0 0.0 87.8 37.2 33.5 1.2 13.6 16.4 0.2 0.6 0.3 0.1 0.5 0.0 54.4 43.0 6.6** 1.4 0.0 2.7** 0.0 0.0 3. ( ) - 1991 2006 . , - 12 ,% . ,% .-. , Vaccinium vitis-idaea Vaccinium myrtillus Empetrum hermaphrod. Vaccinium uliginosum Calluna vulgaris Avenella flexuosa Linaea borealis Ledum palustre -. , , Cladina mitis Cladina rangiferina Cladina stellaris Cladonia uncialis Cladonia gracilis Cladonia crispata Peltigera aphthosa Peltigera canina Stereocaulon paschale , Pleurozium schreberi Dicranum scoparium Dicranum polysetum Polytrichum juniperinum Barbilophozia sp. Hylocomium splendens Sphagnum spp. 4. (: Ryan, 2002) , , . () (, 1957). , , , ~100 ; 1,8 ( ., 2004) (. 4). , . , - , (, 1993; Gorshkov, Bakkal, 1996; , , 2002; , , 2004; ., 2005; ., 2005,). (. 5): ~ 30 ­ ; : a) . . ) . () , 2 < 10 1.7 ­5.8 101 5.8 101 ­6.3 102 6.3 101 ­2.8 102 a) (: , 1947) , 1 3.3 104 ­ 1.6 102 < 3.0 101 3.0 101­ 8.3 8.3 ­ 5.0 101 a) , 0.0 0.1 ­ 0.5 0.5 ­ 1.5 1.5 ­ 3.0 3.0 ­ 10.0 , - (-) - - 1.5 101 ­ 1.0 102 5.0 ­ 15.0) 1.0 104­ 105 2.0 102 70 ) 1.3 106 389 390 Bazzaz, 1975; Southwood et al., 1979; Morneau, Payette, 1989; De Granopre et al., 1993; Gorshkov, Bakkal, 1996; , , 2002; , , 2004; ., 2005. Whittaker, Woodwell, 1968; 1969; Whittaker, 1975; , 1971; 1974; .,1977; , 1984; Bormann, Likens, 1979; Bormann, Sidle, 1990; , 1992, , 1995; Helli, Niemi, 1996; , , 2004; , , 2005; ., 2005. Crocker, Major, 1955; Siren, 1955; Whittaker, 1975; Covington, 1981; Auclair, 1985; , , 1985; Morneau, Payette, 1989; Bormann, Siedle, 1990; Kellman, Roulet, 1990; Leak, 1991; , 1993; Gorshkov, Bakkal, 1996; Gorshkov et. al., 1996; Bergeron, Harvey, 1997; , , 2004; , , 2005; ., 2005,. Odum, 1971; Whittaker, 1975; , 1984; , 1998; , , 2002. 5. , 30 50-100 100-200 200­500 . , . ( ~ 12). . . . . ( ) . ( ). . ( ). . . . ~60 ­ ; - - ; 120­150 ­ ; 400­500 ­ ( ). 11 60 ( ) 5­160 ( ) 66­140 ( ). - . 1991(1994) 2005 . (2006) - (. 2, 85, 82 86). ­ 6 8 9 13%, ­ 13 5% (, <0,05). . ­ Calluna vulgaris 1 7% (, <0.001) Vaccinium vitis-idaea 6,5 3% 518 ( 85); Vaccinium vitis-idaea 12 8% 94108 ( 67). - 50­60 . 60 150 - . - . (2070%, <0,001) (860%, <0,001) 518 (. 2). - ­ 518 ( 46 83%, <0,001) 6680 ( 97 99,8%, <0,05) . 391 - 5­ 100 . , , 518 , Trapeliopsis granulosa, 1%. , Trapeliopsis granulosa, Polytrichum piliferum, Pohlia nutans (. 2, 85). 6580 (. 2, 56, 80, 82) Cladina rangiferina, , , Cladina stellaris ( 56, 80) ­ Cladina mitis ( 80, 82) Cladonia uncialis ( 56, 80). 94108 ( 67), C. mitis C. uncialis. - 5­110 - - (Ahti, Oksanen, 1990; , 1993). 150 - . - - . - - 66­154 . 6679 ­ Vaccinium vitis-idaea (510%, <0,001) Empetrum hermaphroditum (0.33%, <0,01) ­ Calluna vulgaris (105%, <0,05). : 94108 ­ Empetrum hermaphroditum 6 3% (<0,05); 140154 ­ Vaccinium vitis-idaea 3.5 6% (<0,05). 392 , , - 50­60 . 90 14 8% , , . - . - 66­154 . - (. . 3, 75, 63). 6680 ( 75) 84 53% (, <0,001), ­ 60 40% (<0,001), C. mitis (347%, <0,001) C. uncialis (72%, <0,001) ­ Cladina rangiferina (1225%, <0,001) Pleurozium schreberi (733%, <0,001) (1136%, <0,001). 94108 ~5569 ( 63) 27 6% (<0,001) 20 5% (<0,001). (5877%, <0,001) Pleurozium schreberi (5676%, <0,001). 94108 140154 - , (. . 3, 64, 76). 140154 (. 3, 64) Dicranum scoparium (157%, <0,01) Barbilophozia sp. (13%, <0,01). 1. , 140 . 5­8% . , 393 - 140 , (, 1993; . 2005). 2. , 518 - 70%, Cladonia deformis, C. crispata Trapeliopsis granulosa. 65­160 - 85­90% . 60 Cladina mitis, 80­110 ­ Cladina rangiferina, 120 ­ Cladina stellaris. , - (, 1993). 3. - 60 80 . - , 60­90 (, , 2000; . 2005). 4. , - , , 90 . 120 . 0604-48902. .. . . . 3, .; ., 1964. . 300­447. .., .. - // . , 2005. . 88­102. 394 .., .., .. // . , 2005. . 271­281. . , 1844 , // . , 1846. . 1 2. .. // . 1993. . 333. . 111­114. .., .. Pinus sylvestris L. // . . 2002. .38. . 1. . 3­24. .., .., .. // . 2004. 2. . 1­11. .., .., .. Betula pubescens (Betulaceae) // . . 2005 . . 41. . 2. . 1­17. .., .., .. //. 2005 . 3. . 37­45. . . . ., 1984. 182 . . . . ., 1984. 247 . .., .., .. . -. 1. // . . 1995. . 80, 9. . 61­75. .., .., .. . -. 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