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«22» 2008 14:00 002.211.01 . . . : 197376, - , . . , . 2. .: (812) 234-12-37, : (812) 234-45-12.

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«16» 2008

. . C

1

. () ~10 600 700 ( , 1976; Clayton, Renvoize, 1986; Watson, Dallwitz, 1999). 20% (Shantz, 1954). , ( ) , ( , 1993).

( , 1937; , 1976; , 2003; , 2005; Prasad et al., 2005; , 2007). , . / . , . , , ,

60 Poaceae (Devos, Gale, 1998). , ( , )=7, : Bambusoideae, Oryzoideae Arundoideae x=12, Eragrostideae Panicoideae x=9 10 ( , 1931; Hilu, 2004). . . (1969) , 7. 7, / . . . (1969) , « » , , . , x=9, 10, 12 , , , =7, 6, 5, 4 2 ( , 1969, 1972, 1975; Tzvelev, 1989). . . . . (1974) : 2n=2x=4 - Zingeria biebersteiniana (Aveneae). . . . . (1977) , versicolor () 2n=2 =4. =2 ( 6 x=2, , Hyacinthaceae Cyperaceae 2 Asteraceae).

. , . . . . (1965) , Zingeria Colpodium sensu stricto. , Colpodium Poinae Poeae ( , 1976), Zingeria Agrostidinae ( , 1976) Alopecurinae (Clayton, Renvoize, 1986) Aveneae.

2 Poeae s.l. ( , 1987; Tzvelev, 1989). , ,

, , Zingeria Colpodium. .

: , , 2n=2x=4 Zingeria biebersteiniana ( Aveneae) Colpodium versicolor ( Poeae), , Aveneae Poeae, , 7 6 5. . : 1. , ITS1 5.8S ITS2 45S Zingeria Colpodium , , , , Colpodium sensu lato. 2. , ITS1 5.8S ITS2 Aveneae Poeae

=7. 3. , ITS1 ITS2 , , ,

Aveneae Poeae. 4. 5.8S , Aveneae Poeae, , . . ITS1 5.8S ITS2 45S 50- Pooideae, , , Anthoxanthum, Avena, Avenula, Briza, Calamagrostis, , Catabrosella, Colpodium, , Helictotrichon, Hierochloe, Hyalopoa, Melica, Paracolpodium, Phalaris, , , Sesleria, Triticum, Vahlodea, Zingeria, , . , Colpodium sensu lato ITS- , C. apusii, Catabrosella araratica,

3 C. subornata, C. variegata, Hyalopoa lanatiflora, H. pontica, Paracolpodium altaicum Phippsia concinna. ITS , Zingeria Colpodium , (p-distance) ITS 3.1 4.4%. ITS Colpodium sensu lato , Colpodium versicolor, . . (1964), Colpodium (Watson, Dallwitz, 1992 2008 .) Catabrosella subornata, C. variegata, Paracolpodium altaicum, Hyalopoa lanatiflora, H. pontica. , 7 Colpodium Zingeria Catabrosella araratica, Catabrosella. - , Catabrosa (x=5), Catabrosella (x=5), Anthoxanthum ( =5), Trisetum flavescens (x=6), Phalaris truncata Ph. canariensis (x=6), Briza minor (x=5), Zingeria Colpodium, Briza minor, - , B. elatior B. marcowiczii ( 2n=2x=14, x=7). BEP Catabrosa aquatica Hyalopoa lanatiflora « T» 102 5.8S , , BEP PACCAD. . XI ( , 2003), IV - « » ( , 2005), XVII ( , , 2005), V , ( - , 2005), X ( - , 2005), VIII I(IX) - (2004, 2006), , 300- . ( , 2007), ( , , 2007), « » ( , 2007), XV « » ( , 2008). . 16 . . 214 , , 372 , 276 , . 27 20 .

4

=2 =7, Pooideae. Colpodium versicolor (Stev.) Schmalh.

(Poeae). Colpodium s.str. . 2 I .

1. Colpodium versicolor: 1. 2. 3. , DAPI ( Rodionov et al., 2005).

Colpodium - . Colpodium 1820 . C. stevenii Trin. (=Agrostis versicolor Stev., 1812; C. versicolor (Stev.) Schmalh., 1897) C. monandrum Trin. (=Agrostis algida Soland., 1775), 1821 . Vilfa. Vilfa, C. stevenii, Colpodium, C. monandrum Phippsia. 1824 . R. Brown Phippsia , C. monandrum Phippsia algida R. Br., ( , 1934 ; , 1964, 1987; Tzvelev, 1989). . ., Colpodium . versicolor (Stev.) Schmalh. ( , 1934 ). ( .- . . , A.H.R. Grisebach, E. Boissier, O. Stapf, . . , . . , . . , . . , . . , . . . ) ( . , 1934 ; , 1964 .). . . « »

5 Colpodium 7 « » ( , 1934 ). 1964 . , 15 19 . Colpodium 5 , , , Colpodium sensu stricto Tzvel., x=2; Catabrosella Tzvel., x=5, 6, 7, 9; Paracolpodium Tzvel., x=7; Hyalopoa Tzvel., x=7. Colpodium araraticum (Lipsky) Woron. . . Nevskia Catabrosella ( , 1964; , 1967).

2. Zingeria biebersteiniana: 1. 2. , DAPI ( Rodionov et al., 2005). Zingeria biebersteiniana (Claus) P. Smirn. ( Aveneae) - . 2 I . Zngeria =2 Z. trichopoda (2n=8), Z. kochii (2n=12) Z. pisidica (2n=?).

- ITS1-5.8S -ITS2 45S -

ITS1 ITS2 45S -

- , . , (Hillis, Dixon, 1991; Baldwin et al., 1995).

6

Z. biebersteiniana C. versicolor, , Aveneae Poeae, Meliceae Triticeae. , . , ., - ., , . - . : . . , . . . . . , . . , . . , . . , . . , . . ( ), . . , . . ( ) . . ( ).

3. - (45S ). ITS (internal transcribed spacers), ETS (external transcribed spacers), , .

(ITS) - , ~18S ITS1 5.8S ITS2 26S ~ 650 . . CTAB- (Saghai-Maroof et al., 1984; Doyle, Doyle, 1987; ., 2005), . . «DNAEasyPlantMiniKit» (Qiagen, ). (Mullis et al., 1986). ITS1-F 5'-cttggtcatttagaggaagtaa-3' (Gardes, Brunes, 1993), ITS5 5'-ggaagtaaaagtcgtaacaagg-3' (White et al., 1990) ITS1-P 5'-aaccttatcatttagaggaagg-3' (Ridgway et al., 2003). ITS4 5'- tcctccgcttattgatatgc-3' (White et al., 1990). «PCR-Sprint» (Hybaid Inc., ), « » ( . , ), «Techne TC412» (BarloworldScientific,

7 ), «TGradient» (Biometra, ) «2720 Thermal Cycler» (Applied Biosystems, ). . (Sanger et al., 1977) « » « » ( - ),

( ).

SeqScan v.1.2.2 (Applied Biosystems, ). Chromas Lite v.2.23 (Technelysium Pty Ltd, ).

( ) (http://www.ncbi.nlm.nih.gov), 100 , , .

BLAST (www.ncbi.nlm.nih.gov/BLAST/; Altschul et al., 1997). , MEGA v. 3.1 (www.megasoftware.net; Nei, Kumar, 2000; Kumar et al., 2004): ClustalW (Thompson et al., 1995), p-distance Fitch-Margoliash; , ; - 20000 (Felsenstein, 1985). Meliceae Triticeae.

RNA Structure v. 4.5 (Mathews et al., 2004), (Zuker, 1989; Zuker et al., 1999).

ITS1 Aveneae Poeae «TCGWGRCC» «WTWTAA(A)TC» 211 221 . . ( , 217 . .). G+C 28.1+32.8=60.9%, + =20.2+19.0=39.2%. ITS2 «HWAAYACGCTY» «YGYYYHGACS» 209 221 . . ( , 212.7 . .). G+C 31.7+31.7=63.4%, + =18.4+18.1=36.5%, . . GC- ITS2 , ITS1. 5.8S «MMAYGACTYTC» ( «ACACGACTCTC») «GGGHGTCAYRY» ( «GGGCGTCACGC») 162 163 . . 1 ( , ),

8 ITS , Colpodium. , ITS Poa Avena ( 2 ). 2 ,

ITS C. versicolor 0.0 0.2%, Z. biebersteiniana 0%; Z. trichopoda 2.6%, ITS Z. trichopoda Z. biebersteiniana 1.2 3.7%. Colpodium/Zingeria 3.1 4.4%. P- ITS ITS Poa (7.4 17.3%), ITS ITS Avena (13.9 16.3%). , p- ITS Zingeria Colpodium p- Poa.

ITS1 5.8S ITS2 . 4. , C. versicolor ( , . - ) Z. biebersteiniana ( . , .) Z. trichopoda ( : 5 6.07.1916. . . - . - . . .: . ., . .) - . C. versicolor

Colpodium s.l. Catabrosella araratica. Reticulata Colpodium ( , 1934 ), Nevskia Colpodium ( , 1964) / Nevskia Catabrosella ( , 1976; , 1988), , (2n=6x=42), Catabrosella subornata C. variegata, . , Catabrosella araratica

Nevskia. Catabrosella subornata C. variegata, Hyalopoa lanatiflora, H. pontica ( -112), Paracolpodium altaicum, Catabrosa aquatica . capusii , [(Colpodium s.str.+Zingeria spp.) Catabrosella araratica]. Catabrosa Colpodium s.l. (excl. C. versicolor) Catabrosella s.str., Hyalopoa Paracolpodium Catabrosa Colpodium s.l. ( , 1964). H. pontica -112 ( - , . ) Catabrosella variegata C. subornata, H. lanatiflora, Catabrosa aquatica. H. pontica T-76 ( - , . ) Poa, sect. Ochlopoa, , Poa. ITS Hyalopoa pontica -76 -112, - ,

45S .

9 1. ITS1 5.8S ITS2 Zingeria Colpodium.

11 1111111111 1111111111 1111111122 122233344 4445555555 5666667777 7778899900 0001111224 4667777778 8888889900 6207912612 6790123467 8034570124 5693818924 5890135242 7261235670 3456787912 ZINGERIA BIEBERSTEINIANA* DQ910765 AVE 4(2) GGCTCCGCTG TGCTG-TGCG GATTGCTCAT TGAGCCTTCT CAGTGGGCGG GCGCAACCAG CGTGGCTACT Zingeria biebersteiniana AJ428836 AVE 4(2) ...... -...... ZINGERIA TRICHOPODA* GRU AVE 8(2) A...... -...... M.... S...... T....G. ..C...... Zingeria trichopoda ARM AJ428835 AVE 4(2) A...... -...... C.T...... G.T....G. ..C...... COLPODIUM VERSICOLOR* TEB AY497472 POE 4(2) AA.....T...... -.... .T.C...... T..A.G. ..C...... Colpodium versicolor ARM AJ867446 POE 4(2) AA.....T...... -...... C...... T..A.G. ..C...... Colpodium versicolor GRU AJ867445 POE 4(2) AA.....T...... -.... .T.C...... T..A.G. ..C...... CATABROSELLA ARARATICA* R43 POE42(7) A...... AT A..G.-...... C....G...... A...... T....G...... HYALOPOA PONTICA* T-76 POE 28(7) A...A...A. ...A.-C.Y. .TC.....G. ..C..TC...... C...... T..T..T.G. A.C..T...C HYALOPOA PONTICA* T-76-2 POE 28(7) A...A..CA. T..A.-C.TG .TM..TT.C. ..C.CTC...... C...... T..T..T.G. A.C..Y..GC HYALOPOA PONTICA* T-112 POE 28(7) A...... A. ...G.-C... ..A..T..G. ..C.T-C...... C...... T..T.G. A.C.....GC PARACOLPODIUM ALTAICUM* Alt-001 EF432735 POE 42(7) A...... A. ...G.-C... ..A..TC.G. ..C..-C...... C...... T..T.G. A.C.....GC CATABROSELLA VARIEGATA* T-100 AY862811 POE 10(5) A....T.TA. ...G.-C...... C.T..G. ..C..-C...... C.T...... T..T.G. A.CT....GC CATABROSELLA SUBORNATA* POE N A....T.TA. ...G.-C...... C.T..G. ..C..-C...... C.T...... T..T.G. A.CT....AC HYALOPOA LANATIFLORA* POE 42(7) A.G.....A. ...G.-C..A ..A..T..G. ..C.M-C...... C....T. ...T..T.G. A.CAA...GC CATABROSA AQUATICA* KS-21 EF577510 POE 10(5) A.G.....A. ...G.-C..A ..A..T..G. ..C..-C...... C....T. ...T..T.G. A.CAA...GC CATABROSA CAPUSII* Alt-311 POE 20 (5) A.G.....A. ...G.-C..A ..A..T..G. ..CC.-C...... C.T..T. ...T..T.G. A.CAA.C.GC PHIPPSIA CONCINNA* QKS-13 EU093028 POE 28(7) A.T..T..AT .C.G.-C... ATC..T..-. -.C..-G..A ...C..T... ..AT....G. A.C..T.... dura AF532933 POE 14 (7) A.NNNNN.AT ...G.CA... ..A..T..G. ..C..-A...... C..T...... T..T.GA A.C.T...GC distans AF532934 POE 28 42(7) A.AG....AT ...G.-C... ..A.CT..GC ..C..-C.T. ...C...... AT..T.G. A.C.....GC POA CHAIXII* R33 EF433458 POE 14(7) A...A...... A.-C... .CCC....G. ..C..TC...... C...... T....G. A.C...... POA SUPINA* EF165099 POE 14(7) --A.AAC.A. CA.A.-CA.A .CA....TG. ..CT.GCC.. T....T.... A..TT...G. T...... C AVENA DAMASCENA* AY881171 AVE 14(7) A..A...TAT ..TGAG...... CC....G. CAC-.TC... T.T....T.. ...T.GTTG. .AC.A..G.C AVENA PILOSA* AY530162 AVE 14(7) A..C...TAT -.YGAGC... ..CC...TG. C.C..TC... .GA...... A A..T.GT.GT T.C...CGTC

2222222222 2222223333 3333333333 4444444444 4444444444 4444444444 4444444444 0000111222 2222333556 6666678899 0011111122 2222333333 3344445555 5555666667 4579014024 5678275592 3458904967 2601234902 6789012356 7913580123 5789025672 ZINGERIA BIEBERSTEINIANA* DQ910765 AVE 4(2) CCCTCGTTCA TTTACCCCGA CCTCCGCCCC CCCTCCCTAC GGGTAGTG-A TCGATGCTCT GCATGATGGG Zingeria biebersteiniana AJ428836 AVE 4(2) ...... -...... ZINGERIA TRICHOPODA* GRU AVE 8(2) ...... C..N ...... -...... C...... Zingeria trichopoda ARM AJ428835 AVE 4(2) ...... A...C.-...... C...... COLPODIUM VERSICOLOR* TEB AY497472 POE 4(2) ...... A...... T...... A...C.-...... C...... Colpodium versicolor ARM AJ867446 POE 4(2) ...... A...... T...... A...C.-...... C...... Colpodium versicolor GRU AJ867445 POE 4(2) ...... A...... T...... A...C.-...... C...... CATABROSELLA ARARATICA* R43 POE42(7) ...CT..A...... T.N...... C.G...... C...... HYALOPOA PONTICA* T-76 POE 28(7) .M.C...... T..T...... A..-.CT ...... G. ....G.TCTC ..TC..G... HYALOPOA PONTICA* T-76-2 POE 28(7) ...C.T..CA ...... T .T..TG.G.. TT...... CT N....GT.TT N.CTGTTC.C CCTCCTG... HYALOPOA PONTICA* T-112 POE 28(7) ...C.T...... T .T..T..... T..A..A.CT .T...... G. ...TG..C.C ...C...... PARACOLPODIUM ALTAICUM* Alt-001 EF432735 POE 42(7) C...... T.T .T..T....T T.MA..A.CT .T.C....G. ...TG.TC.C ...CC..... CATABROSELLA VARIEGATA* T-100 AY862811 POE 10(5) ..C.T...... T TT..TA.... T..A..A.CT .T.C.A..G. ...TG.TC.C ...C...... CATABROSELLA SUBORNATA* POE N ...C.T...... T TT..TA.... T..A..A.CT .T.C.A..G. ...GG.TCTC ...C...... HYALOPOA LANATIFLORA* POE 42(7) ...C...... TT..T YT.TT..... T..A..A.CT RK...... G. .T.TGC.CY...... CATABROSA AQUATICA* KS-21 EF577510 POE 10(5) ...C...... TT..T .T.TT..... T..A..A.CT AT...... G. .T.TGC.C...... CATABROSA CAPUSII* Alt-311 POE 20 (5) ...C-....G A.CTANNNNN NNNNNNNNNN T..A..N.CT .T...N..N. .T.TGNNC.C ...C...... PHIPPSIA CONCINNA* QKS-13 EU093028 POE 28(7) ...C...... AT TT.TT..... T..A..A-CT ...-....G. ....G..CTC .TGC...... Sclerochloa dura AF532933 POE 14 (7) ...... T .T.TT..... T..A..A.CT ...... AG. ...TG.TC.C ...C...TA. AF532934 POE 28 42(7) ...... T .T.TT..... T..A..A..T ...... AG. ...TG.TCTC ..GC...TA. POA CHAIXII* R33 EF433458 POE 14(7) ...... T...... A..-.CT ....G...G. ....G.TC.C ..TC..G... POA SUPINA* EF165099 POE 14(7) TG.C...... A.....T.. .T....T.T. ...A..A.CT ...A....G. C...G..C.C .TTC..GTA. AVENA DAMASCENA* AY881171 AVE 14(7) .T.C.TCA...... TTC...... T ...AA...T- ...G.ACAG...... TC.C ...CC.G.-. AVENA PILOSA* AY530162 AVE 14(7) .TTC.KCA...... TTC...... T ...A.A.CCT .A.G.ACAG...... C ...C..G..A

4444444444 4455555555 5555555555 5555555555 5555555555 66666666666 7777888899 9902222233 4455555555 6666667778 8889999999 00000001111 5678013456 8932346724 0201234678 0156892560 3562346789 23567890126 ZINGERIA BIEBERSTEINIANA* DQ910765 AVE 4(2) CCGAGAGCGT TCTGGCATTT ACCATCCACT CGCCATCGGT A-GTATAAA- GTCGCATGACC Zingeria biebersteiniana AJ428836 AVE 4(2) ...... -...... - ...... ZINGERIA TRICHOPODA* GRU AVE 8(2) ...... -...... - ...... Zingeria trichopoda ARM AJ428835 AVE 4(2) ..CGAG...... C...... C.... .-...... - ...... COLPODIUM VERSICOLOR* TEB AY497472 POE 4(2) ...... A...... C. T.T...... T....C.... .-...... - ...... Colpodium versicolor ARM AJ867446 POE 4(2) ...... A...... C. T.T...... T....C.... .-...... - ...... Colpodium versicolor GRU AJ867445 POE 4(2) ...... A...... C. T.T...... T....C.... .-...... - ...... CATABROSELLA ARARATICA* R43 POE42(7) ...... C. ...C...GTC .....C.... .-...... - ...... T.. HYALOPOA PONTICA* T-76 POE 28(7) ...... C....AC. T...... C .....CG... .-....G..- .A...... HYALOPOA PONTICA* T-76-2 POE 28(7) ..A.A.N------HYALOPOA PONTICA* T-112 POE 28(7) ...... T...... CC. G.T...... A .....CAT.. .-....G..- .A..A.....T PARACOLPODIUM ALTAICUM* Alt-001 EF432735 POE 42(7) .T.....T...... CC. G.T...... A T...... C-...G..- .A...... T CATABROSELLA VARIEGATA* T-100 AY862811 POE 10(5) ...... T...... CC. G.T...... A T....C.A.. .C-...G..- .A..A.....T CATABROSELLA SUBORNATA* POE N ...... T...... CC. G.T...... A T....C.A.. .C-...G..- .A..A.....T HYALOPOA LANATIFLORA* POE 42(7) TYT....T.. .T.....AC. T.T..TY..C T....C.... .-....G..- .AY...... T CATABROSA AQUATICA* KS-21 EF577510 POE 10(5) T.T....T.. .T.....AC. T.T..TT..C T....C.... .-....G..- .AT...... T CATABROSA CAPUSII* Alt-311 POE 20 (5) T.T....T.. .T....NNC. T.T..TT..C T....C.... .-....G..- .A...... T PHIPPSIA CONCINNA* QKS-13 EU093028 POE 28(7) .T.....T...... T..C. GAT....G.C ..A.GC...A .-T...-.GC AAT...... T Sclerochloa dura AF532933 POE 14 (7) ...... T...... CC. T.T...... C ..T..C...C .AT...G-GA .A.....A.TT Puccinellia distans AF532934 POE 28 42(7) ...... T...... C...C. T.T.....AC .....C..AC .AT-..G-GA .A...... T POA CHAIXII* R33 EF433458 POE 14(7) ...... C....AC. T...... G.C -A...CG... .-....G..- .A...... POA SUPINA* EF165099 POE 14(7) ..T....TC. ..C.....C. T....A.GAC ...... T... .-.A.AG..- .A.A.T...TT AVENA DAMASCENA* AY881171 AVE 14(7) ...... AT.C ...A..TCCA T.TC.....A ....GCG..A .A-.T-GT.A ...... AVENA PILOSA* AY530162 AVE 14(7) ...... AT.C ...A..TCC. T.-CG....G ...T.CG..C T--.T.G..- ...... A....

: . AVE Aveneae, POE Poeae. (2n) , , , N .

10 2. P- ITS Zingeria Colpodium Poa Avena. P- . .

, , , , H. pontica , Colpodium s.l. Poa, (1895, : , 1934). , , H. pontica Poa capillipes Somm. et Lev., 1893, ( .: , 1934; , 1976). Paracolpodium altaicum Arctagrostis, Colpodium s.l. . . (1964) Arctagrostis Paracolpodium Poa. , , Poa, Arctagrostis Paracolpodium Arctopoa ( . . ) Andinae Poa, Poa ( , , 2008). Colpodium s.l. (excl. Colpodium s.str. et Nevskia) [(Puccinellia distans+Sclerochloa dura) Phippsia]. Phippsia Puccinellia (Love, Love, 1975) Phippsia, Puccinellia Sclerochloa (Choo et al., 1994) . ITS , , , , . ======. 4 . 11. , ITS1 ITS2 ( ). , ITS , . 20000 . POE Poeae, AVE Aveneae. ( ). Colpodium s.l. « » (compressed) .

11

Agrostis spp.*+Lachnagrostis spp. AVE (7) 89 CALAMAGROSTIS NEGLECTA* L-7(2) POE 28(7) CALAMAGROSTIS PURPUREA*KS-26-2 POE 28(7) 26 30 C. PHRAGMITOIDES* KS-27-2 POE 56-91(7) 19 CALAMAGROSTIS LAPPONICA* L-13 POE 28 140(7) Gastridium ventricosum POE 14 28(7) Deyeuxia spp.+Dichelachne spp. AVE (7) 34 49 Polypogon monspeliensis AVE 28(7) Lachnagrostis billardierei NZ AVE 99 92 Lachnagrostis littoralis salaria NZ AVE 91 Lachnagrostis littoralis litto NZ AVE Calamagrostis epigeios POE 28 42 56 (7) 66 Ammophila arenaria AVE 14 28 56(7) 50 Calamagrostis spp. POE (7) 99 SESLERIA spp.* POE (7) 99 Helictotrichon pruinosum AVE 94 Helictotrichon sedenense AVE 14(7) 27 99 Helictotrichon filifolium AVE 28(7) Trisetum youngii NZ AVE 21 86 splendens AVE 14(7) 98 wolfii AVE N 42 Arrhenatherum elatius AVE 28(7) 72 AVENA SPP.* AVE (7) 83 BRIZA ELATIOR* T-44 POE 14(7) 93 BRIZA MARCOWICSII* T-56 POE 14(7) 83 BRIZA MINOR* TzH-001 POE 10(5) 26 99 Briza minor POE 10(5) Phalaris truncata AVE 12(6) 99 PHALARIS CANARIENSIS* AVE 12(6) HIEROCHLOE ALPINA* KS-20 AVE 56(7) 18 81 Hierochloe fusca AVE 84(7) 52 Hierochloe equiseta AVE 41(N) 97 Hierochloe novae-zelandiae AVE 28(7) 99 Anthoxanthum amarum AVE 80 90(5) 99 Anthoxanthum aristatum AVE 10(5) 99 ANTHOXANTHUM ALPINUM* T-88 AVE 10(5) 39 95 ANTHOXANTHUM ALPINUM* Alt-254 AVE 10(5) 99 ANTHOXANTHUM ODORATUM* Alt-248 AVE 20(5) 94 spp. AVE Helictotrichon blaui AVE 99 40 HELICTOTRICHON ASIATICUM* T-70 AVE 14(7) caryophyllea AVE 14 28(7) 80 Avenella flexuosa AVE 28(7) 24 VAHLODEA ATROPURPUREA* KS-28 AVE 14(7) 96 Vahlodea atropurpurea AVE 14(7) 85 s.l. POE (7) ARCTAGROSTIS LATIFOLIA* QKS-23 AVE42 56(7) 88 99 Arctagrostis latifolia AVE 42 56(7) ARCTAGROSTIS LATIFOLIA* CY-2 AVE 42 56(7) 94 Festucella eriopoda POE N 33 99 Hookerochloa hookeriana POE N ARCTOPOA SCHISCHKINII* Alt-55 POE 70(7) 23 94 99 LATIFOLIA* POE 28(7) 84 84 Phleum phleoides AVE 14 28(7) 1 POE 44-88(N) 99 POE 42(7) 32 82 Dupontia fisheri 2 POE 44-88(N) CATABROSELLA ARARATICA* R43 POE 42(7) 99 ZINGERIA BIEBERSTEINIANA* RAKH AVE 4(2) 99 99 Zingeria biebersteiniana AVE 4(2) ZINGERIA TRICHOPODA* GRU AVE 8(2) 96 Zingeria trichopoda ARM AVE 4(2) 49 Colpodium versicolor ARM POE 4(2) 99 COLPODIUM VERSICOLOR* TEB T-94 POE 4(2) 79 Colpodium versicolor GRU POE 4(2) CATABROSA AQUATICA* KS-21 POE 10(5) 46 66 99 CATABROSA CAPUSII* Alt-311 POE 20 (5) 37 HYALOPOA LANATIFLORA* TzH-003 POE 42(7) 79 PARACOLPODIUM ALTAICUM* POE 42(7) HYALOPOA PONTICA* T-112 POE 28(7) 30 49 99 CATABROSELLA SUBORNATA* TzH-002 POE N CATABROSELLA VARIEGATA* T-100 POE 10 (5) 66 99 Sclerochloa dura POE 14(7) Puccinellia distans POE 28 42(7) PHIPPSIA CONCINNA* QKS-13 POE 28(7) 56 99 POE 28(7) 99 POA SUPINA* POE 14(7) POE 14(7) 39 34 HYALOPOA PONTICA* T-76 POE 28(7) HYALOPOA PONTICA* T-76-2 POE 28(7) 27 Poa ramosissima POE 28(7) 46 Poa alpina var. alpina POE 28 42(7) 25 POE 14(7) 71 POA PALUSTRIS* R6 POE 28 42(7) 15 42 Poa flexuosa POE 42(7) 99 Poa hartzii hartzii POE 70(7) POA COMPRESSA* POE 42(7) 97 Poa anceps subsp. polyphylla POE 28(7) 58 Poa litorosa POE 253-265 (N) POA CHAIXII* R33 POE 14(7) 57 51 Poa arctica POE 56 70(7) 98 Poa novae-zelandiae POE 28(7) Poa acicularifolia acicularif POE 28(7) 36 Poa pratensis POE 14-84 (7) 32 Poa abbreviata POE 28 42 70(7) 99 Poa pratensis ssp. alpigena POE 61-72(7) 91 Poa pratensis subsp. alpigena Poa pratensis subsp. alpigena(2) TRITICUM AESTIVUM* TRI 42(7) 99 GLYCERIA spp.* MEL 28 56 60 (7 6 5) 4.

12

5.8S

, 5.8S , , , , ( .: Hershkovitz, Lewis, 1996; Peculis, Greer, 1998; Gottschling, Plotner, 2004). , 5.8S , , , , , ( , , 2002). , 5.8S « » , . 54 102 , BEP PACCAD ( , 2006). 102 , Catabrosa aquatica Hyalopoa lanatiflora.

5. 5.8S Colpodium versicolor T-94 - , 26S . I (Ia+Ib), II III . Aveneae Poeae ( . . 1 2). 54 102, BEP PACCAD.

13 3. III 5.8S . 1 Colpodium versicolor

Cinna latifolia

5.8S Arctagrostis latifolia. 2 Zingeria biebersteiniana, Z. trichopoda Catabrosella araratica.

Arctopooa schischkinii, Arctophila fulva, Phleum phleoides, Dupontia fisheri .

1 Catabrosa aquatica, : Hyalopoa lanatiflora, Puccinellia distans, 2 Hyalopoa pontica T-76 Sclerochloa dura. 3 Hyalopoa pontica T-112 1 Hyalopoa lanatiflora 4 Paracolpodium altaicum c 5 Catabrosella variegata, C. subornata «Y=C/T» 6 Colpodium versicolor

. 5 , III. 3 III . Colpodium versicolor , Zingeria , Zingeria spp. Catabrosella araratica (A1). Catabrosa aquatica Hyalopoa lanatiflora, Puccinellia distans Sclerochloa dura ( 1), Hyalopoa pontica T-76 ( 2) T-112 ( 3), , ( ). III .

14 (1) . . - «G:U», , (2) , « » , - ( . . 4).

4. III 5.8S .

Phippsia algida subsp. concinna QKS-13 . 126 «G A», « T» 135.

Arrhenatherum elatius , 126 135, (2006).

, =2 Zingeria C. versicolor. Catabrosella araratica « » =7. - , 7 2 . , , « » - , ( . 5 . . .). =2 . 6 . I Zingeria Colpodium, 7 2. II =2 Zingeria spp. ( Z) Colpodium spp. ( C). , Colpodium spp. . versicolor

15

6. =2.

( V) , Zingeria biebersteiniana Z+C, - 2n=8 ( IV). - , 45S C / ITS Z TR . ZTR 45S (Kotseruba et al., 2003), 45S CTR, , ITS Z. biebersteiniana, , C. versicolor. Z. trichopoda 2n=8 (2ZTR+2CTR) ( V). c

16 Z. biebersteiniana C. versicolor, Z. trichopoda 2n=2x=4, . C. versicolor Cv , CTR Z. trichopoda, GISH (Kotseruba et al., 2005) . , Z. biebersteiniana Zbi c 2n=2x=4 , - , Zingeria , , ZTR ( GISH . Kotseruba et al., 2003). Z. biebersteiniana, ITS ITS1. , , : X 7 2 , C. versicolor.

, Z. trichopoda. , , , C. versicolor, Z. biebersteiniana,

ITS. , ITS Z. trichopoda , . (2005), ,

. ITS - , , . Z. trichopoda ITS , Z. biebersteiniana, ITS , C. versicolor. : (1) ITS ( ITS1 ITS4, . ITS5 ITS4). ITS1P ITS5 ( 6 . 22) 1 . , , ITS, , , ITS Z. trichopoda , ; (2) , , ITS , 45S CTR Z. trichopoda , Zingeria spp., , 45S ZTR .

17 , ITS, Z. trichopoda - ITS . x=7, Aveneae Poeae, Anthoxanthum alpinum (2n=10, x=5), Briza minor (2n=10, x=5), Catabrosa aquatica (2n=20; x=5), C. capusii (2n=20, x=5), Catabrosella variegata (2n=10, x=5), Phalaris canariensis Ph. truncata (2n=12, x=6). Zingeria biebersteiniana Z. trichopoda Colpodium versicolor , . Catabrosa aquatica c C. capusii Catabrosella variegata, , , [(Colpodium s.str.+Zingeria spp.) Catabrosella araratica]. Briza minor (2n=10, x=5), - , B. elatior B. marcowiczii ( 2n=14, x=7), B. minor . Phalaris truncata Ph. canariensis , 7 (2n=12, x=6) , Briza, . ITS Phalaris Briza. , , , . Seslerieae (Tutin et al., 1980; Watson, Dallwitz, 1992; , 1987), Sesleriinae Poeae Poeae s.l. ( , 1987; Tzvelev, 1989; Soreng, 2003). Sesleria Poaceae ( ITS Sesleria coerulea ITS 2- Sesleria, . ( ) (Quintanar et al., 2007), , Sesleria spp. Aveneae, , ITS1- 5.8S -ITS2 . , , trnT-F (Quintanar et al., 2007), Seslerieae Poeae. ,

, .

18

1. , , ITS1 5.8S ITS2 45S Aveneae Poeae, Meliceae Triticeae. , , , Aveneae Poeae. 2. ITS , x=2 Zingeria biebersteiniana (2n=4), Z. trichopoda (2n=8) Colpodium versicolor (2n=4), , , (p-distance) ITS 3.1 4.4%, Poa. 3. - , 150 ITS1 5.8S ITS2, , - , , Catabrosella araratica (2n=42, x=7). , x=2 Zingeria Colpodium s.str. , . 4. ITS , Colpodium sensu lato, , C. versicolor Colpodium s.l. Catabrosella araratica. Paracolpodium altaicum, Hyalopoa lanatiflora, H. pontica ( -112), Catabrosella subornata C. variegata Catabrosa aquatica Catabrosa capusii ,

[(Colpodium s.str.+Zingeria spp.) Catabrosella araratica]. , Catabrosella araratica Nevskia. 5. (Hyalopoa) (Catabrosella), ITS- , . ,

« » - , . 6. , x=7, Aveneae Poeae, Anthoxanthum alpinum (2n=10, x=5), Catabrosella variegata (2n=10, x=5), Catabrosa aquatica (2n=20, x=5), C. capusii (2n=20, x=5), Briza minor (2n=10, x=5) Phalaris truncata P. canariensis (2n=12, x=6)

19 . , 7- 6- Phalaris truncata Ph. canariensis 7- 5- Briza minor . 7. 5.8S . , Poeae Aveneae , , 5.8S . 5.8S , ( BEP) Hyalopoa lanatiflora Catabrosa aquatica.

1. . ., . ., . ., . . ITS- 5.8S Avena // . XI . . 2. . 17 27 2003 . . 278 279. 2. . ., . ., . ., . . (2n=4) Zingeria biebersteiniana Colpodium versicolor // VIII - . 17 21 2004 . - . . 245. 3. . ., . ., . ., . . Avena macrostachya ITS // VIII - . 17 21 2004 . - . . 245. 4. . ., . ., . ., . . - Avena L. : ITS1, ITS2 5.8S // XXI : . . 6 12 2004. . 2. . 269. 5. Rodionov A.V., Kim E.S., Punina E.O., Machs E.M., Tyupa N.B., Dobroradova M.A., Nossov N.N. Molecular phylogenetic study of Colpodium versicolor and some other Aveneae and Poeae species // XVII International Botanical Congress. Vienna, Austria, . Austria Center, Vienna. 17 23, July, 2005. P. 431 432. 6. . ., . ., . ., . ., . . Avena macrostachya, ITS1 ITS2: Avena // . 2005. . 41. 5. . 646 656. 7. . ., . ., . ., . ., . . Paracolpodium altaicum: - // IV - « ». 12 14 2005. . . 53 54.

20

8. . . - Zingeria biebersteiniana Colpodium versicolor // - . - . 2005. . 32 33. 9. . ., . ., . ., . ., . . Zingeria biebersteiniana Colpodium versicolor: - // V , . 12 15 2005 . - . . 48 49. 10. . ., . ., . ., . ., . ., . . Colpodium sensu lato (Poaceae), ITS1 ITS2 45S // I(IX) - . 21 26 2006 . - . . 30 31. 11. . ., . ., . ., . ., . . Poa schischkinii Tzvel. (=Arctopoa schischkinii (Tzvel.) Probat.) Poeae ITS1 ITS2 45S // V - « ». 21 23 2006 . . . 199 200. 12. . ., . ., . ., . ., . ., . . Aveneae Poeae ITS1 ITS2 45S // . 2007. . 92. 1. P. 57 71. 13. . ., . ., . ., . ., . ., . . Zingeria Colpodium // , 300- . . 17 19 2007. . 66 67. 14. . ., . ., . ., . . ITS 5.8S Anthoxanthum, Hierochloe Phalaris // , 300- . . 17 19 2007 . . 77 79. 15. . ., . ., . ., . ., . ., . . - Poa L. s.l. // « 2007». 16 19 2007 . . . 216 219. 16. . ., . . Poa L. (Poaceae) - // (XVI) « C » ( 3- ). . III. XV

21 « ». . 14 18 2008 . . 209 211.

: , GenBank NCBI ( ) http://www.ncbi.nlm.nih.gov. Agrostis capillaris L. (=Agrostis tenuis Sibth.) N-25 AY520820; Agrostis gigantea Roth Alt-014 EF565133; Agrostis stolonifera L. Alt-052 EF541167; Arctagrostis latifolia (R. Br.) Griseb. EU093029, C -2 EU921830; Anthoxanthum alpinum A. et D. Love T-88 EU093027, Alt-254 FJ010629; A. odoratum L. Alt- 248 FJ010630; Arctopoa schischkinii (Tzvel.) Probat. Alt-55 EU935583; Avena damascena Rajh. et Baum N-1 AY881171; A. longiglumis Dur. N-4 AY522436; A. macrostachya Bal. ex Coss. et Dur. N-8-2 AY522433; A. pilosa M.B. N-28 AY530162; A. hirtula Lag. N-10 AY522435; A. ventricosa Bal. N-21 AY522437; Briza elatior Sibth. et Smith QT-44 EU935582; B. marcowiczii Woronow T-56 FJ008711; B. minor L. TzH-001 EU935584; Calamagrostis lapponica (Wahl.) Hartm. L-13 FJ010631; C. neglecta (Ehrh.) Gaertn., Mey. et Schreb. L-7(2) FJ010632; C. phragmitoides s.str. Hartm. KS-27-2 FJ010633; C. purpurea (Trin.) Trin. KS-26-2 FJ013223; Catabrosa aquatica (L.) P. Beauv. KS-21 EF577510; C. capusii Franchet (=Catabrosa aquatica subsp. capusii (Franchet) Tzvel.) Alt- 311=Ma-6 FJ196299; Catabrosella araratica (Lipsky) Tzvel. R-43 FJ196300; C. subornata E.B.Alexeev (Colpodium humilis subsp. subornata) TzH-002 FJ013225; C. variegata (Boiss.) Tzvel. T-100 AY862811; Colpodium versicolor (Stev.) Schmalh. -94 AY497472; Glyceria maxima (Hartm.) Holmb. E-22 FJ013226; Helictotrichon adzharicum (Albov) Grossheim. (=H. asiaticum (Roshevitz). Grossheim., Avenula hookeri (Scribner) Holub) T-70 AY870327; H. pubescens (Huds.) Pilg. ( ITS1) T-98 AY870326; Hierochloe alpina (Sw. ex Willd.) Roemer et J.A. Schultes (=Anthoxanthum monticola (Bigelow) Y. Schouten et Veldkamp) KS-20 EF577511; Hyalopoa lanatiflora (Roshev.) Tzvel. TzH-003 FJ178781; H. pontica (Bal.) Tzvel. T-76 FJ196302; H. pontica (Bal.) Tzvel. T-112 FJ196303; Paracolpodium altaicum (Trin.) Tzvel. Alt-001 EF432735; Phalaris canariensis L. FJ178782; Phippsia concinna (Th. Fries) Lindeb. (=Phippsia algida subsp. concinna (Th.Fr.) A.Love et D.Love) QKS-13 EU093028; Poa chaixii Vill. R- 33 EF433458; Poa palustris L. R-6 EF555592; Poa supina Schrad EF165099; Sesleria coerulea (L.) Ard. EF565132; Triticum aestivum L. Tr-6 FJ196304; Vahlodea atropurpurea (Wahlenb.) Fries KS-28 FJ013224; Zingeria biebersteiniana (Claus.) P. Smirn DQ910765; Z. trichopoda (Boiss.) P. Smirn. TzH-004 FJ196301.