- 03.00.05 - 2008 . : : , - : . «22» 2008 14:00 002.211.01 . : 197376, - , . , . 2. .: (812) 234-12-37, : (812) 234-45-12. «16» 2008 . C 1 . (Poaceae) ~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) , Colpodium versicolor (Poeae) 2n=2 =4. =2 ( 6 x=2, , Hyacinthaceae Cyperaceae 2 Asteraceae). , . (1965) , Zingeria Colpodium sensu stricto. Pooideae , 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, Agrostis, Arctagrostis, Anthoxanthum, Avena, Avenula, Briza, Calamagrostis, Catabrosa, Catabrosella, Colpodium, Glyceria, Helictotrichon, Hierochloe, Hyalopoa, Melica, Paracolpodium, Phalaris, Phippsia, Poa, Sesleria, Triticum, Vahlodea, Zingeria, , . , Colpodium sensu lato ITS- Catabrosa aquatica, 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.... Sclerochloa dura AF532933 POE 14 (7) A.NNNNN.AT ...G.CA... ..A..T..G. ..C..-A... ...C..T... ...T..T.GA A.C.T...GC Puccinellia 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