Адаптация животных к хроническим факторам

³ɛɪɸɲɤɨ´ɜ ɤɨɬɨɪɨɦ ɪɚɫɩɨɥɨɠɟɧɚ ɰɢɬɨɩɥɚɡɦɚ Ʉɪɨɦɟ ɬɨɝɨ ɦɨɠɧɨ ɝɨɜɨɪɢɬɶ ɨɛ ɭɩɥɨɳɟɧɧɨɣ ɛɪɸɲɧɨɣ ɩɨɜɟɪɯɧɨɫɬɢ ɢ ɜɵɩɭɤɥɨɣ ɫɩɢɧɧɨɣ ɉɪɢɭɫɬɶɟɜɚɹ ɱɚɫɬɶ ɞɨɪ ɡɚɥɶɧɨɣ ɩɨɜɟɪɯɧɨɫɬɢ ɦɨɠɟɬ ɨɛɪɚɡɨɜɵɜɚɬɶ ɤɨɡɵɪɟɤ Ʉɨɪɝɚɧɨɜɚ ɍɫɬɶɟ ɦɨɠɟɬ ɛɵɬɶ ɪɚɡɥɢɱɧɨɣ ɜɟɥɢɱɢɧɵ ɢ ɮɨɪɦɵ ɛɨɥɶɲɨɟ Cyclopyxis arcelloides ɦɚɥɨɟ C. ambigua ɨɤɪɭɝɥɨɟ Cyclopyxis ɳɟɥɟɜɢɞɧɨɟ Plagiopyxis ɢ ɪɚɫɩɨɥɚɝɚɬɶɫɹ ɬɟɪɦɢ ɧɚɥɶɧɨ Nebela, Euglypha ɢɥɢ ɷɤɫɰɟɧɬɪɢɱɧɨ Trinema Ɉɧɨ ɦɨɠɟɬ ɛɵɬɶ ɨɤɪɭɠɟɧɨ ɛɨɥɟɟ ɤɪɭɩɧɵɦɢ ɦɢɧɟɪɚɥɶɧɵɦɢ ɱɚɫɬɢɰɚɦɢ ɱɟɦ ɧɚ ɨɫɬɚɥɶɧɨɣ ɱɚɫɬɢ ɪɚɤɨɜɢɧɤɢ

(Centropyxis plagiostoma ɥɢɛɨ ɛɨɥɟɟ ɦɟɥɤɢɦɢ Cyclopyxis kahli ɥɢɛɨ ɩɪɢɭɫɬɶɟ

ɜɵɦɢ ɢɞɢɨɫɨɦɚɦɢ ɫ ɨɞɧɢɦ ɢɥɢ ɧɟɫɤɨɥɶɤɢɦɢ ɡɭɛɱɢɤɚɦɢ Euglypha). Ɇɧɨɝɨɨɛɪɚɡɢɟ ɮɨɪɦ ɪɚɤɨɜɢɧɨɤ ɤɚɤ ɢ ɢɯ ɜɚɪɢɚɛɟɥɶɧɨɫɬɶ ɜɧɭɬɪɢ ɜɢɞɚ ɨɛɭ

ɫɥɨɜɥɟɧɚ ɩɪɢɫɩɨɫɨɛɢɬɟɥɶɧɨɣ ɮɭɧɤɰɢɟɣ ɪɚɤɨɜɢɧɤɚ ɹɜɥɹɟɬɫɹ ɩɨɫɪɟɞɧɢɤɨɦ ɦɟɠɞɭ ɨɪɝɚɧɢɡɦɨɦ ɢ ɦɟɫɬɨɦ ɟɝɨ ɨɛɢɬɚɧɢɹ Sch|nborn Ʉɨɪɝɚɧɨɜɚ

ɋɜɟɞɟɧɢɹ ɨ ɛɢɨɥɨɝɢɢ ɩɨɱɜɨɨɛɢɬɚɸɳɢɯ ɬɟɫɬɚɰɟɣ ɨɬɪɵɜɨɱɧɵ ɢ ɧɨɫɹɬ ɯɚɪɚɤɬɟɪ ɨɬɞɟɥɶɧɵɯ ɧɚɛɥɸɞɟɧɢɣ ɑɬɨ ɫɜɹɡɚɧɨ ɫ ɧɟɨɞɧɨɪɨɞɧɨɫɬɶɸ ɩɨɱɜɟɧɧɵɯ ɭɫɥɨɜɢɣ ɜ ɤɭɥɶɬɭɪɟ ȼ ɧɟɤɨɬɨɪɵɯ ɪɚɛɨɬɚɯ ɨɩɢɫɵɜɚɸɬɫɹ ɚɦɟɛɨɢɞɧɵɟ ɫɬɚɞɢɢ ɢ ɨɪɝɚɧɢɡɦɵ ɫ ɛɨɥɟɟ ɬɨɧɤɢɦɢ ɪɚɤɨɜɢɧɤɚɦɢ ɱɬɨ ɩɨɡɜɨɥɹɟɬ ɩɪɟɞɩɨɥɚɝɚɬɶ ɧɚɥɢɱɢɟ ɭ ɬɟɫɬɚɰɟɣ ɫɥɨɠɧɨɝɨ ɠɢɡɧɟɧɧɨɝɨ ɰɢɤɥɚ ɇɚɛɥɸɞɚɥɢɫɶ ɬɚɤɠɟ ɹɜɥɟɧɢɹ ɚɧɚɥɨɝɢɱɧɵɟ ɥɢɧɶɤɢȽɟɥɶɰɟɪ Ɉɫɧɨɜɧɨɣ ɬɢɩ ɪɚɡɦɧɨɠɟɧɢɹ± ɛɟɫɩɨɥɵɣ ɨɫɭɳɟɫɬɜɥɹɟɦɵɣ ɩɭɬɟɦ ɞɟɥɟɧɢɹ ɧɚɞɜɨɟ Ⱥɦɟɛɵ ɫ ɦɹɝɤɨɣ ɪɚɤɨɜɢɧɤɨɣ ɞɟɥɹɬɫɹ ɜɞɨɥɶ ɫ ɬɜɟɪɞɨɣ± ɩɨɩɟɪɟɤ ɩɪɨɞɨɥɶɧɨɣ ɨɫɢ ɩɨɱɤɨɜɚɧɢɟɦ ȼ ɩɪɨɰɟɫɫɟ ɪɚɡɦɧɨɠɟɧɢɹ ɞɨɱɟɪɧɹɹ ɤɥɟɬɤɚ ɨɬɞɟɥɹ ɟɬɫɹ ɨɬ ɦɚɬɟɪɢɧɫɤɨɣ ɩɪɢ ɞɨɫɬɢɠɟɧɢɢ ɢɦɢ ɨɞɢɧɚɤɨɜɵɯ ɪɚɡɦɟɪɨɜ ɢ ɨɛɪɚɡɨɜɚɧɢɢ ɧɨɜɨɣ ɪɚɤɨɜɢɧɤɢ ɪɢɫ 4. Ⱦɨ ɮɨɪɦɢɪɨɜɚɧɢɹ ɩɟɪɟɬɹɠɤɢ ɜɧɨɜɶ ɫɮɨɪɦɢɪɨɜɚɜɲɟɟ ɫɹ ɹɞɪɨ ɩɟɪɟɯɨɞɢɬ ɜ ɞɨɱɟɪɧɸɸ ɤɥɟɬɤɭ ɉɪɨɞɨɥɠɢɬɟɥɶɧɨɫɬɶ ɩɪɨɰɟɫɫɚ ɞɟɥɟɧɢɹ ɪɚɡ ɥɢɱɧɚ Ɉɧɚ ɦɨɠɟɬ ɞɥɢɬɶɫɹ ɦɢɧɭɬ ɢɥɢ ɜɫɟɝɨ - ɦɢɧɭɬ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɜɢ ɞɚ Ɉɞɧɨɫɥɨɣɧɚɹ ɨɪɝɚɧɢɱɟɫɤɚɹ ɪɚɤɨɜɢɧɤɚ ɨɬɞɟɥɹɟɬɫɹ ɭ ɦɨɥɨɞɵɯ ɨɫɨɛɟɣ ɨɬ ɰɢɬɨɩɥɚɡɦɵ ɰɟɥɢɤɨɦ Arcella ɩɪɨɩɢɬɵɜɚɟɬɫɹ ɫɨɥɹɦɢ ɠɟɥɟɡɚ ɢ ɤɪɟɦɧɟɜɨɣ ɤɢɫɥɨ ɬɨɣ Ⱥɦɟɛɵ ɫ ɤɫɟɧɨɫɨɦɚɦɢ ɢ ɢɞɢɨɫɨɦɚɦɢ ɬɚɤɠɟ ɜɵɞɟɥɹɸɬ ɜɧɚɱɚɥɟ ɨɪɝɚɧɢɱɟɫɤɭɸ ɨɫɧɨɜɭ ɫɥɭɠɚɳɭɸ ɰɟɦɟɧɬɢɪɭɸɳɢɦ ɜɟɳɟɫɬɜɨɦ ɞɥɹ ɤɪɨɸɳɢɯ ɷɥɟɦɟɧɬɨɜ ɗɥɟɦɟɧ ɬɵ ɩɨɤɪɵɬɢɹ ɪɚɤɨɜɢɧɤɢ ɧɚɤɚɩɥɢɜɚɸɬɫɹ ɜ ɰɢɬɨɩɥɚɡɦɟ ɦɚɬɟɪɢɧɫɤɨɣ ɤɥɟɬɤɢ ɟɳɟ ɞɨ ɧɚɱɚɥɚ ɞɟɥɟɧɢɹ ɢ ɩɨɫɥɟ ɨɛɪɚɡɨɜɚɧɢɹ ɩɥɚɡɦɟɧɧɨɣ ɩɨɱɤɢ ɜɵɫɬɢɥɚɸɬ ɟɟ ɩɨɜɟɪɯ ɧɨɫɬɶ

ɍ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɫɬɪɨɹɳɢɯ ɚɝɝɥɸɬɢɧɢɪɨɜɚɧɧɵɟ ɪɚɤɨɜɢɧɵ ɞɟɥɟɧɢɸ ɩɪɟɞɲɟ ɫɬɜɭɟɬ ɩɟɪɢɨɞ ɚɤɬɢɜɧɨɝɨ ɩɨɝɥɨɳɟɧɢɹ ɤɫɟɧɨɫɨɦ ȼ ɧɚɱɚɥɟ ɞɟɥɟɧɢɹ ɢɡ ɭɫɬɶɹ ɦɚɬɟɪɢɧ ɫɤɨɣ ɪɚɤɨɜɢɧɵ ɜɵɩɹɱɢɜɚɟɬɫɹ ɭɱɚɫɬɨɤ ɰɢɬɨɩɥɚɡɦɵ ɪɢɫ ɤɨɬɨɪɵɣ ɭɜɟɥɢɱɢɜɚɟɬɫɹ ɜ ɪɚɡɦɟɪɚɯ ɢ ɩɪɢɧɢɦɚɟɬ ɮɨɪɦɭ ɫɨɨɬɜɟɬɫɬɜɭɸɳɭɸ ɮɨɪɦɟ ɛɭɞɭɳɟɣ ɪɚɤɨɜɢɧɵ Ʉ ɩɨ ɜɟɪɯɧɨɫɬɢ ɜɵɯɨɞɚ ɰɢɬɨɩɥɚɡɦɵ ɩɟɪɟɦɟɳɚɸɬɫɹ ɜɚɤɭɨɥɢ ɫ ɤɫɟɧɨɫɨɦɚɦɢ ɢ ɫ ɨɪɝɚɧɢ ɱɟɫɤɢɦ ɰɟɦɟɧɬɨɦ ɂɧɨɝɞɚ ɚɦɟɛɚ ɨɛɪɚɡɭɟɬɩɚɥɶɰɟɜɢɞɧɵɟ ɜɟɫɶɦɚɩɨɞɜɢɠɧɵɟɩɫɟɜɞɨɩɨ ɞɢɢ ɤɨɬɨɪɵɟ ɦɨɝɭɬ ɡɚɯɜɚɬɵɜɚɬɶ ɱɚɫɬɢɰɵ ɢɡ ɜɧɟɲɧɟɣ ɫɪɟɞɵ ɉɪɨɢɫɯɨɞɢɬ ɷɤɡɨɰɢɬɨɡ ɫɨɞɟɪɠɢɦɨɝɨ ɜɚɤɭɨɥɟɣ ɢ ɤɫɟɧɨɫɨɦɵ ɫɤɪɟɩɥɹɸɬɫɹ ɨɪɝɚɧɢɱɟɫɤɢɦ ɰɟɦɟɧɬɨɦ Ʉɪɭɩɧɵɟ ɱɚɫɬɢɰɵ ɢɡ ɜɧɟɲɧɟɣ ɫɪɟɞɵ ɦɨɝɭɬ ɩɪɨɫɬɨ©ɩɪɢɥɢɩɚɬɶªɤ ɫɬɟɧɤɟ ɞɨɱɟɪɧɟɣ ɪɚɤɨɜɢɧɵ əɞɪɨ ɩɪɟɬɟɪɩɟɜɚɟɬ ɞɟɥɟɧɢɟ ɱɚɫɬɶ ɨɪɝɚɧɟɥɥ ɢ ɨɞɧɨ ɢɡ ɹɞɟɪ ɦɢɝɪɢɪɭɸɬ ɜ ɞɨɱɟɪɧɸɸ ɤɥɟɬɤɭ ɉɪɨɢɫɯɨɞɢɬ ɨɤɨɧɱɚɬɟɥɶɧɨɟ ɪɚɡɞɟɥɟɧɢɟ ɰɢɬɨɩɥɚɡɦɵ ɤɥɟɬɨɤ Ⱦɨɱɟɪɧɹɹ ɤɥɟɬɤɚ ɟɳɟ ɧɟɤɨɬɨɪɨɟ ɜɪɟɦɹ ɞɨɫɬɪɚɢɜɚɟɬ ɪɚɤɨɜɢɧɭ ɮɚɝɨɰɢɬɢɪɭɹ ɤɫɟɧɨɫɨɦɵ ɢ ɜɤɥɸɱɚɹ ɢɯ ɜ ɫɬɟɧɤɭ ɪɚɤɨɜɢɧɵ ɂɡɧɭɬɪɢ ɪɚɤɨɜɢɧɚ ɜɵɫɬɢɥɚɟɬɫɹ ɫɥɨɟɦ ɨɪɝɚɧɢɱɟɫɤɨɝɨ ɰɟɦɟɧɬɚ ɡɚ ɫɱɟɬ ɧɟɝɨ ɮɨɪɦɢɪɭɟɬɫɹ ɨɪɝɚɧɢɱɟɫɤɚɹ ɤɚɟɦɤɚ ɜɨɤɪɭɝ ɭɫɬɶɹ ɢ ɡɚɩɨɥɧɹɸɬɫɹ ɩɪɨɦɟɠɭɬɤɢ ɦɟɠɞɭ ɱɚɫɬɢɰɚɦɢ ɍ ɜɡɪɨɫɥɨɣ ɨɫɨɛɢ ɰɢɬɨɩɥɚɡɦɚ ɨɬɞɟɥɹɟɬɫɹ ɨɬ ɫɬɟɧɨɤ ɪɚɤɨɜɢɧɵ ɢ ɨɛɪɚɡɭɸɬɫɹɷɩɢɩɨɞɢɢ Ȼɟɣɟɪ ɢ ɞɪ ).

ϭϳϲ

Ɋɢɫɭɧɨɤ ɋɯɟɦɚ ɦɨɪɮɨɝɟɧɟɡɚ Arcella spp. Ȼɟɣɟɪ ɢ ɞɪ Ⱥ²ɤɥɟɬɤɚ ɝɨɬɨɜɚɹɤɞɟɥɟɧɢɸ Ȼ ȼ²ɮɨɪɦɢɪɨɜɚɧɢɟɜɵɪɨɫɬɚɰɢɬɨɩɥɚɡɦɵɢɩɟ

ɪɟɦɟɳɟɧɢɟɬɟɤɚɝɟɧɧɵɯɝɪɚɧɭɥ Ƚ²ɨɛɪɚɡɨɜɚɧɢɟɰɢɬɨɩɥɚɡɦɚɬɢɱɟɫɤɨɝɨɤɭɩɨɥɚ ɤ ɢɧɚ ɱɚɥɶɧɨɟɮɨɪɦɢɪɨɜɚɧɢɟɪɚɤɨɜɢɧɵ Ⱦ²ɦɚɬɟɪɢɧɫɤɚɹɢɞɨɱɟɪɧɹɹɪɚɤɨɜɢɧɵ ɫɤɪɟɩɥɟɧ ɧɵɟɡɚɦɤɨɦ ɡ ɜɨɛɥɚɫɬɢ ɭɫɬɶɹ

ȿ ɤɥɟɬɤɢɩɨɫɥɟɞɟɥɟɧɢɹ ɬɝ²ɬɟɤɚɝɟɧɧɵɟɝɪɚɧɭɥɵ ɹ ɹɞɪɚ

Ɋɢɫɭɧɨɤ ɋɯɟɦɚ ɦɨɪɮɨɝɟɧɟɡɚ Difflugia spp. Ȼɟɣɟɪ ɢ ɞɪ

ϭϳϳ

Ⱥ ɤɥɟɬɤɚ ɝɨɬɨɜɚɹɤɞɟɥɟɧɢɸ Ȼ²ɨɛɪɚɡɨɜɚɧɢɟɰɢɬɨɩɥɚɡɦɚɬɢɱɟɫɤɨɝɨɜɵɪɨɫɬɚ ȼ Ƚ²ɧɚɱɚɥɶɧɵɟɷɬɚɩɵɮɨɪɦɢɪɨɜɚɧɢɹɞɨɱɟɪɧɟɣɪɚɤɨɜɢɧɵ Ⱦ²ɡɚɜɟɪɲɟɧɢɟɮɨɪɦɢ ɪɨɜɚɧɢɹɞɨɱɟɪɧɟɣ ɪɚɤɨɜɢɧɵ ɷɩɢɩɨɞɢɢɟɳɟɧɟɨɛɪɚɡɨɜɚɥɢɫɶ ȿ ɤɥɟɬɤɢɩɨɫɥɟɞɟɥɟ ɧɢɹ ɤ²ɤɫɟɧɨɫɨɦɵ ɩ²ɩɫɟɜɞɨɩɨɞɢɢ ɚɝɝɥɸɬɢɧɢɪɭɸɳɢɟɷɤɡɨɝɟɧɧɵɣ ɦɚɬɟɪɢɚɥ ɹ

ɹɞɪɨ

Ⱦɨɫɬɨɜɟɪɧɵɯ ɫɜɟɞɟɧɢɣ ɨ ɫɭɳɟɫɬɜɨɜɚɧɢɢ ɩɨɥɨɜɨɝɨ ɩɪɨɰɟɫɫɚ ɭ ɬɟɫɬɚɰɢɣ ɧɟ ɦɧɨɝɨ ɢɦɟɸɬɫɹ ɥɢɲɶ ɫɥɭɱɚɣɧɵɟ ɢ ɪɚɡɪɨɡɧɟɧɧɵɟ ɧɚɛɥɸɞɟɧɢɹ Ƚɟɥɶɰɟɪ ɉɪɨɞɨɥɠɢɬɟɥɶɧɨɫɬɶ ɠɢɡɧɢ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɡɧɚɱɢɬɟɥɶɧɨ ɜɵɲɟ ɱɟɦ ɭ ɞɪɭɝɢɯ ɝɪɭɩɩ ɩɪɨɫɬɟɣɲɢɯ ɜ ɫɪɟɞɧɟɦ - ɞɧɟɣ ȼ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɜɥɚɠɧɨɫɬɢ ɩɪɨɞɨɥɠɢ ɬɟɥɶɧɨɫɬɶ ɠɢɡɧɢ ɜɚɪɶɢɪɭɟɬ ɨɬ ɨɞɧɨɝɨ ɞɧɹ ɞɨ ɧɟɫɤɨɥɶɤɢɯ ɧɟɞɟɥɶ Ⱦɟɥɟɧɢɟ ɨɬɞɟɥɶ ɧɵɯ ɛɵɫɬɪɨ ɪɚɡɦɧɨɠɚɸɳɢɯɫɹ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɫɟɦɟɣɫɬɜɚ Euglyphidae ɩɪɨɢɫɯɨɞɢɬ ɱɟɪɟɡ - ɞɧɹ ɱɢɫɥɨ ɩɨɤɨɥɟɧɢɣ ɦɨɠɟɬ ɡɚɜɢɫɟɬɶ ɬɨ ɬɢɩɚ ɩɨɱɜɵ ȿɫɬɶ ɞɚɧɧɵɟ ɨ ɫɭɳɟɫɬɜɨɜɚɧɢɢ - ɩɨɤɨɥɟɧɢɣ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɜ ɝɨɞ ɇɚɛɥɸɞɟɧɢɹ ɧɚɞ ɫɟɡɨɧ ɧɨɣ ɞɢɧɚɦɢɤɨɣ ɫɨɨɛɳɟɫɬɜ ɚɦɺɞ ɦɚɥɨɱɢɫɥɟɧɵ ɡɚɪɟɝɢɫɬɪɢɪɨɜɚɧɵ ɥɢɲɶ ɜɟɫɟɧɧɢɟ ɜɫɩɵɲɤɢ ɱɢɫɥɟɧɧɨɫɬɢ ɉɥɨɬɧɨɫɬɶ ɬɟɫɬɚɰɢɣ ɜ ɨɛɴɟɦɟ ɩɨɱɜɵ ɜ ɬɟɱɟɧɢɟ ɝɨɞɚ ɡɧɚɱɢ ɬɟɥɶɧɨ ɢɡɦɟɧɹɟɬɫɹ Ⱦɥɢɬɟɥɶɧɨɫɬɶ ɫɨɯɪɚɧɟɧɢɹ ɪɚɤɨɜɢɧ ɜ ɩɨɱɜɟ ɡɚɜɢɫɢɬ ɨɬ ɜɢɞɚ ɤɨɪɧɟɧɨɠɟɤ ɢ ɤɨɧɤɪɟɬɧɵɯ ɭɫɥɨɜɢɣ ȼ ɯɜɨɣɧɵɯ ɥɟɫɚɯ ɨɧɢ ɦɨɝɭɬ ɪɚɡɪɭɲɚɬɶɫɹ ɜ ɬɟ ɱɟɧɢɟ ɨɞɧɨɝɨ ɝɨɞɚ ȼ ɬɨɠɟ ɜɪɟɦɹ ɧɚɯɨɠɞɟɧɢɟ ɪɚɤɨɜɢɧɨɤ ɜ ɫɭɛɮɨɫɫɢɥɶɧɵɯ ɨɬɥɨ ɠɟɧɢɹɯ ɛɨɥɨɬ ɝɨɜɨɪɢɬ ɨ ɜɨɡɦɨɠɧɨɫɬɢ ɢɯ ɫɨɯɪɚɧɟɧɢɹ ɜ ɚɧɚɷɪɨɛɧɵɯ ɭɫɥɨɜɢɹɯ ɜ ɬɟ ɱɟɧɢɟ ɞɥɢɬɟɥɶɧɨɝɨ ɜɪɟɦɟɧɢ Ƚɟɥɶɰɟɪ

Ɉɫɧɨɜɧɨɣ ɩɢɳɟɣ ɹɜɥɹɸɬɫɹ ɛɚɤɬɟɪɢɢ ɞɢɚɬɨɦɨɜɵɟ ɢ ɧɢɬɱɚɬɵɟ ɜɨɞɨɪɨɫɥɢ ɞɪɭɝɢɟ ɬɟɫɬɚɰɢɣ ɢ ɩɪɨɫɬɟɣɲɢɢ Ʉɪɢɜɨɥɭɰɤɢɣ ȼ ɨɬɥɢɱɢɟ ɨɬ ɝɨɥɵɯ ɮɨɪɦ ɩɪɨɫɬɟɣɲɢɯ ɪɚɤɨɜɢɧɧɵɟ ɚɦɟɛɵ ɩɢɬɚɸɬɫɹ ɧɟɩɨɫɪɟɞɫɬɜɟɧɧɨ ɨɫɬɚɬɤɚɦɢ ɪɚɡɥɚɝɚɸ ɳɢɯɫɹ ɨɪɝɚɧɢɡɦɨɜ ɇɟɤɨɬɨɪɵɟ ɢɡ ɧɢɯ Nebela, Difflugia ɩɨɟɞɚɸɬ ɦɟɥɤɢɯ ɱɥɟɧɢ ɫɬɨɧɨɝɢɯ Yeatus ɉɪɢɧɢɦɚɹ ɜɨ ɜɧɢɦɚɧɢɟ ɛɨɥɶɲɭɸ ɱɢɫɥɟɧɧɨɫɬɶ ɪɚɤɨɜɢɧ ɧɵɯ ɚɦɟɛ ɜ ɩɨɱɜɚɯ ɫ ɜɟɫɶɦɚ ɛɟɞɧɵɦ ɛɚɤɬɟɪɢɚɥɶɧɵɦ ɧɚɫɟɥɟɧɢɟɦ, ɦɨɠɧɨ ɩɪɟɞɩɨɥɚɝɚɬɶ ɱɬɨ ɢɯ ɩɢɳɟɜɵɦɢ ɨɛɴɟɤɬɚɦɢ ɹɜɥɹɸɬɫɹ ɚɫɤɨɦɢɰɟɬɵ ɝɪɢɛɵ ɢ ɪɚɫ ɬɜɨɪɟɧɧɵɟ ɜ ɜɨɞɟ ɩɢɬɚɬɟɥɶɧɵɟ ɜɟɳɟɫɬɜɚ ɍ ɧɟɤɨɬɨɪɵɯ ɜɢɞɨɜ ɩɫɟɜɞɨɩɨɞɢɢ ɧɚɛɥɸ

ɞɚɸɬɫɹ ɢɫɤɥɸɱɢɬɟɥɶɧɨ ɪɟɞɤɨ Plagiopyxis callida, Trigonopyxis arcula ɉɨɷɬɨɦɭ

ɩɢɬɚɧɢɟ ɦɨɠɟɬ ɨɫɭɳɟɫɬɜɥɹɬɶɫɹ ɞɢɮɮɭɡɧɨ ɢɡ ɨɤɪɭɠɚɸɳɟɣ ɫɪɟɞɵ ɉɫɟɜɞɨɫɬɨɦ ɱɚɫɬɨ ɛɵɜɚɟɬ ɩɥɨɬɧɨ ɡɚɤɪɵɬ ³ɩɭɱɤɨɦ´ ɞɟɬɪɢɬɚ ɩɨɷɬɨɦɭ ɞɨɩɭɫɤɚɟɬɫɹ ɜɨɡɦɨɠɧɨɫɬɶ ɩɢɬɚɧɢɹ ɛɟɡ ɜɜɟɞɟɧɢɹ ɩɢɳɟɜɵɯ ɨɛɴɟɤɬɨɜ ɜɧɭɬɪɶ ɰɢɬɨɩɥɚɡɦɵ Ɍɚɤɨɣ ɫɩɨɫɨɛ ɩɢɬɚ ɧɢɹ ɨɩɢɫɚɧ ɞɥɹ Difflugia rubescens ɤɨɪɧɟɧɨɠɤɚ ɩɪɨɤɚɥɵɜɚɟɬ ɨɛɨɥɨɱɤɭ ɜɨɞɨɪɨɫɥɢ (Closterium ɢ³ɜɵɫɚɫɵɜɚɟɬ´ɟɟ ɫɨɞɟɪɠɢɦɨɟ Ƚɟɥɶɰɟɪ ȿɫɬɟɫɬɜɟɧɧɵɦɢ ɜɪɚ ɝɚɦɢ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɹɜɥɹɸɬɫɹ ɯɢɳɧɵɟ ɝɪɢɛɵ Loopagaceae, ɨɛɢɬɚɸɳɢɟ ɜ ɩɨɱɜɟ ɨɥɢɝɨɯɟɬɵ ɬɢɯɨɯɨɞɤɢ ɤɪɭɩɧɵɟ ɢɧɮɭɡɨɪɢɢ ɝɨɥɵɟ ɚɦɟɛɵ ɧɟɦɚɬɨɞɵ ȼɨɡɦɨɠɧɨ

ɩɨɬɪɟɛɥɹɸɬ Testacea ɬɚɤɠɟ ɦɢɤɨɮɚɝɢ Enchytraeidae ɢ Oribatidae Ȼɨɛɪɨɜ

ȼ ɜɚɤɭɨɥɹɯ ɬɟɫɬɚɰɟɣ ɢɦɟɸɬɫɹ ɱɚɫɬɢɰɵ ɝɭɦɭɫɚ ɍɱɢɬɵɜɚɹ ɜɵɫɨɤɭɸ ɱɢɫɥɟɧɧɨɫɬɶ ɬɟɫɬɚɰɢɣ ɜ ɩɨɱɜɚɯ ɫ ɝɪɭɛɵɦ ɝɭɦɭɫɨɦ ɦɨɠɧɨ ɩɪɟɞɩɨɥɚɝɚɬɶ ɢɯ ɭɱɚɫɬɢɟ ɜ ɝɭɦɢɮɢ ɤɚɰɢɢ ɬɪɭɞɧɨ ɪɚɡɥɨɠɢɦɵɯ ɪɚɫɬɢɬɟɥɶɧɵɯ ɨɫɬɚɬɤɨɜ ɥɟɫɧɵɯ ɩɨɱɜ

ɉɪɢ ɧɟɛɥɚɝɨɩɪɢɹɬɧɵɯ ɭɫɥɨɜɢɹɯ ɜ ɱɚɫɬɧɨɫɬɢ ɢɫɫɭɲɟɧɢɢ ɩɨɱɜɵ ɪɚɤɨɜɢɧɧɵɟ ɚɦɟɛɵ ɢɧɰɢɫɬɢɪɭɸɬɫɹ ɪɢɫ ɰɢɬɨɩɥɚɡɦɚ ɨɛɟɡɜɨɠɢɜɚɟɬɫɹ ɫɠɢɦɚɟɬɫɹ ɢ ɨɤɪɭ ɠɚɟɬɫɹ ɡɚɳɢɬɧɵɦɢ ɨɛɨɥɨɱɤɚɦɢ ɉɫɟɜɞɨɫɬɨɦ ɡɚɤɭɩɨɪɢɜɚɟɬɫɹ ɩɪɨɛɨɱɤɨɣ- ɫɥɢɡɢɫɬɵɦ ɫɟɤɪɟɬɨɦ Ɉɤɪɭɝɥɚɹ ɰɢɫɬɚ ɩɨɦɟɳɚɟɬɫɹ ɜ ɛɪɸɲɧɨɣ ɱɚɫɬɢ ɪɚɤɨɜɢɧɤɢ ɉɪɢ ɢɫɫɥɟɞɨɜɚɧɢɢ ɫɭɯɢɯ ɨɛɪɚɡɰɨɜ ɩɨɱɜɵ ɱɚɫɬɨ ɧɚɛɥɸɞɚɸɬ ɢɧɰɢɫɬɢɪɨɜɚɜɲɢɯɫɹ ɚɦɟɛ ɥɢɛɨ ɧɚɯɨɞɹɳɢɯɫɹ ɜ ɫɨɫɬɨɹɧɢɢ ɬɚɤ ɧɚɡɵɜɚɟɦɨɣ ɩɪɟɞɰɢɫɬɵ Ɋɚɤɨɜɢɧɤɢ ɩɟɪɟ ɤɪɵɜɚɸɬɫɹ ɜɪɟɦɟɧɧɨɣ ɡɚɳɢɬɧɨɣ ɞɢɚɮɪɚɝɦɨɣ ɤɨɬɨɪɚɹ ɨɛɪɚɡɭɟɬɫɹ ɜɫɟɝɨ ɡɚ ɧɟ ɫɤɨɥɶɤɨ ɦɢɧɭɬ Ɂɚɳɢɳɟɧɧɨɟ ɞɢɚɮɪɚɝɦɨɣ ɩɪɨɫɬɟɣɲɟɟ ɩɟɪɟɧɨɫɢɬ ɞɥɢɬɟɥɶɧɨɟ ɞɨ - ɯ ɦɟɫɹɰɟɜ ɢɫɫɭɲɟɧɢɟ ɫɪɟɞɵ. ɉɟɪɟɤɪɵɜɚɧɢɹ ɩɫɟɜɞɨɫɬɨɦɚ ɪɚɤɨɜɢɧɤɢ ɜ ɤɚɱɟɫɬɜɟ ɛɵɫɬɪɨɣ ɪɟɚɤɰɢɢ ɧɚ ɤɨɥɟɛɚɧɢɹ ɭɪɨɜɧɹ ɜɥɚɠɧɨɫɬɢ ɯɚɪɚɤɬɟɪɧɵɟ ɞɥɹ ɩɨɱɜɟɧɧɨɣ

ϭϳϴ

ɫɪɟɞɵ ɨɛɢɬɚɧɢɹ ɫɜɨɣɫɬɜɟɧɧɵ ɦɧɨɝɢɦ Euglyphidae ɍ ɪɹɞɚ ɜɢɞɨɜ ɬɟɫɬɚɰɟɣ ɧɚɛɥɸ ɞɚɟɬɫɹ ɨɛɪɚɡɨɜɚɧɢɟ ɭɩɥɨɬɧɟɧɧɨɣ ɡɚɳɢɬɧɨɣ ɩɟɥɥɢɤɭɥɵ ɜɨɤɪɭɝ ɰɢɬɨɩɥɚɡɦɵ ɫɬɚɞɢɹ ɤɚɩɫɭɥɵ ɧɟ ɩɪɟɩɹɬɫɬɜɭɸɳɟɣ ɨɞɧɚɤɨ ɮɨɪɦɢɪɨɜɚɧɢɸ ɩɫɟɜɞɨɩɨɞɢɣ Ʉɨɪɝɚɧɨɜɚ

1997).

Ɋɢɫɭɧɨɤ ɋɯɟɦɚ ɰɢɫɬɵ ɪɚɤɨɜɢɧɧɨɣɚɦɟɛɵ Ȼɟɣɟɪɢɞɪ ɚ²ɚɝɝɥɸɬɢɧɢɪɨɜɚɧɧɚɹ©ɩɪɨɛɤɚªɡɚɤɪɵɜɚɸɳɚɹɭɫɬɶɟɪɚɤɨɜɢɧɵ ɛ²ɪɚɤɨɜɢɧ

ɤɚ ɜ²ɚɝɝɥɸɬɢɧɢɪɨɜɚɧɧɚɹɞɨɩɨɥɧɢɬɟɥɶɧɚɹɫɬɟɧɤɚɰɢɫɬɵ ɝ ɫɟɤɪɟɬɢɪɨɜɚɧɧɚɹ

ɫɬɟɧɤɚɰɢɫɬɵ ɞ²ɰɢɬɨɩɥɚɡɦɚɤɥɟɬɤɢ

Ɉɫɧɨɜɧɵɟ ɮɚɤɬɨɪɵ ɨɩɪɟɞɟɥɹɸɳɢɟ ɫɨɫɬɚɜ ɫɨɨɛɳɟɫɬɜ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɜɤɥɸɱɚɸɬ ɜɨɞɧɵɣ ɪɟɠɢɦ ɫɪɟɞɵ ɪɇ ɫɨɞɟɪɠɚɧɢɟ ɦɢɧɟɪɚɥɶɧɵɯ ɜɟɳɟɫɬɜ ɤɢɫɥɨɪɨ ɞɚ ɬɟɦɩɟɪɚɬɭɪɭ ɨɫɜɟɳɟɧɧɨɫɬɶ ɧɚɥɢɱɢɟ ɫɬɪɨɢɬɟɥɶɧɨɝɨ ɦɚɬɟɪɢɚɥɚ ɞɥɹ ɮɨɪɦɢɪɨ ɜɚɧɢɹ ɪɚɤɨɜɢɧɤɢ ɢ ɩɢɬɚɬɟɥɶɧɵɟ ɜɟɳɟɫɬɜɚ Ɂɚɜɚɪɡɢɧ ɍɫɥɨɜɢɹ ɜɨɞɧɨɝɨ ɪɟ ɠɢɦɚ ɜ ɡɧɚɱɢɬɟɥɶɧɨɣ ɫɬɟɩɟɧɢ ɪɟɝɭɥɢɪɭɸɬ ɨɛɢɥɢɟ ɢ ɫɬɪɭɤɬɭɪɭ ɫɨɨɛɳɟɫɬɜ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɭɪɨɜɟɧɶ ɜɥɚɠɧɨɫɬɢ ɨɬɪɚɠɚɟɬɫɹ ɧɚ ɱɢɫɥɟɧɧɨɫɬɢ ɜɢɞɨɜ ɚ ɬɚɤɠɟ ɧɚ ɢɯ ɫɨɨɬɧɨɲɟɧɢɢ ɢ ɞɨɦɢɧɢɪɨɜɚɧɢɢ Ɋɚɤɨɜɢɧɧɵɟ ɚɦɟɛɵ ɷɤɫɰɢɫɬɢɪɭɸɬɫɹ ɢ ɩɟɪɟ ɯɨɞɹɬ ɜ ɛɨɥɟɟ ɚɤɬɢɜɧɨɟ ɫɨɫɬɨɹɧɢɟ ɩɪɢ ɛɨɥɟɟ ɜɵɫɨɤɨɦ ɭɪɨɜɧɟ ɜɥɚɠɧɨɫɬɢ ɱɟɦ ɞɪɭ ɝɢɟ ɩɪɟɞɫɬɚɜɢɬɟɥɢ ɠɢɜɨɬɧɨɝɨ ɧɚɫɟɥɟɧɢɹ ɩɨɱɜ ɧɚɩɪɢɦɟɪ ɧɟɦɚɬɨɞɵ ɢ ɤɨɥɨɜɪɚɬɤɢ ɩɪɢɱɟɦ ɜɪɟɦɹ ɪɟɚɤɬɢɜɚɰɢɢ ɡɚɜɢɫɢɬ ɨɬ ɩɪɨɞɨɥɠɢɬɟɥɶɧɨɫɬɢ ɩɪɟɞɲɟɫɬɜɨɜɚɜɲɟɝɨ ɩɟɪɢɨɞɚ Ȼɨɛɪɨɜ ɉɪɢ ɚɧɚɥɢɡɟ ɝɪɭɩɩɢɪɨɜɨɤ ɚɦɺɛ ɜ ɩɨɱɜɚɯ ɫ ɪɚɡɥɢɱɧɵɦɢ ɡɧɚɱɟɧɢɹɦɢ ɪɇ ɜɵɞɟɥɹɸɬɫɹ ɜɢɞɵ ɷɜɪɢɢɨɧɧɵɟ Plagiopyxis declivis ɫɬɟɧɨɢɨɧɧɵɟ ɚɰɢɞɨɮɢɥɶɧɵɟ Copythion dubium ɢ ɫɬɟɧɨɢɨɧɧɵɟ ɛɚɡɨɮɢɥɶɧɵɟ Geopyxella sylvicola, C. halophila Ɋɚɤɨɜɢɧɧɵɟ ɚɦɟɛɵ ɦɨɝɭɬ ɩɨɫɟɥɹɬɶɫɹ ɜ ɫɭɛɫɬɪɚɬɚɯ ɝɞɟ ɤɢ ɫɥɨɬɧɨɫɬɶ ɫɢɥɶɧɨ ɨɬɥɢɱɚɟɬɫɹ ɨɬ ɩɨɱɜɵ ɜ ɱɚɫɬɧɨɫɬɢ ɜɨ ɦɯɚɯ ɩɨɤɪɵɜɚɸɳɢɯ ɫɬɜɨ ɥɵ ɞɟɪɟɜɶɟɜ Booth ɂɡ ɞɪɭɝɢɯ ɫɜɨɣɫɬɜ ɜɥɢɹɸɳɢɯ ɧɚ ɫɨɫɬɚɜ ɧɚɫɟɥɟɧɢɹ ɬɟɫ ɬɚɰɟɣ ɫɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ ɫɨɞɟɪɠɚɧɢɟ ɭɝɥɟɤɢɫɥɨɝɨ ɤɚɥɶɰɢɹ ɉɨɱɜɵ ɛɨɝɚɬɵɟ ɤɚɥɶɰɢɟɦ ɯɚɪɚɤɬɟɪɢɡɭɸɬɫɹ ɫɩɟɰɢɮɢɱɟɫɤɨɣ ɮɚɭɧɨɣ ɬɟɫɬɚɰɢɣ ɍɫɥɨɜɢɹ ɡɚɫɨɥɟɧ ɧɵɯ ɩɨɱɜ ɩɟɪɟɧɨɫɢɬ ɥɢɲɶ ɦɚɥɨɟ ɱɢɫɥɨ ɜɢɞɨɜ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ Centropyxis halophila, Plagiopyxis minuta. ɋɜɨɣɫɬɜɚ ɝɭɦɭɫɚ ɜɥɢɹɸɬ ɧɚ ɫɨɫɬɚɜ ɢ ɪɚɫɩɪɨɫɬɪɚɧɟ ɧɢɟ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ Ɍɢɩ ɝɭɦɭɫɚ ɯɚɪɚɤɬɟɪɢɡɭɟɬɫɹ ɝɥɚɜɧɵɦ ɨɛɪɚɡɨɦ ɪɚɡɥɢɱɢɹɦɢ ɜ ɫɬɟɩɟɧɢ ɢ ɫɤɨɪɨɫɬɢ ɦɢɧɟɪɚɥɢɡɚɰɢɢ ɨɪɝɚɧɢɱɟɫɤɢɯ ɨɫɬɚɬɤɨɜ ɬ ɟ ɫɨɨɬɧɨɲɟɧɢɟɦ ɭɝɥɟɪɨɞɚ ɢ ɚɡɨɬɚ C/N ɋɜɨɟɨɛɪɚɡɢɟ ɬɨɝɨ ɢɥɢ ɢɧɨɝɨ ɦɟɫɬɨɨɛɢɬɚɧɢɹ ɜ ɛɨɥɶɲɨɣ ɫɬɟɩɟɧɢ ɨɩɪɟɞɟɥɹɟɬɫɹ ɫɜɨɟɨɛɪɚɡɢɟɦ ɩɨɞɫɬɢɥɤɢ ɡɚɜɢɫɹɳɟɣ ɨɬ ɯɚɪɚɤɬɟɪɚ ɪɚɫɬɢ ɬɟɥɶɧɨɫɬɢ Ʌɟɫɧɚɹ ɩɨɞɫɬɢɥɤɚ ɨɫɨɛɵɣ ɤɨɦɩɨɧɟɧɬ ɛɢɨɝɟɨɰɟɧɨɡɚ ɢɝɪɚɸɳɢɣ ɜɚɠ ɧɭɸ ɪɨɥɶ ɜ ɩɨɱɜɨɨɛɪɚɡɨɜɚɬɟɥɶɧɨɦ ɩɪɨɰɟɫɫɟ ɩɨɷɬɨɦɭ ɜ ɩɨɱɜɟɧɧɨ-ɡɨɨɥɨɝɢɱɟɫɤɢɯ ɢɫɫɥɟɞɨɜɚɧɢɹɯ ɧɟɨɛɯɨɞɢɦ ɬɳɚɬɟɥɶɧɵɣ ɚɧɚɥɢɡ ɟɟ ɧɚɫɟɥɟɧɢɹ ɤɨɬɨɪɵɣ ɦɨɠɟɬ ɞɚɬɶ ɞɨɩɨɥɧɢɬɟɥɶɧɵɟ ɫɜɟɞɟɧɢɹ ɨ ɬɟɱɟɧɢɢ ɝɭɦɭɫɨɨɛɪɚɡɨɜɚɬɟɥɶɧɨɝɨ ɩɪɨɰɟɫɫɚ.

ɇɚɢɛɨɥɟɟ ɛɨɝɚɬɚ ɬɟɫɬɚɰɟɹɦɢ ɮɚɭɧɚ ɜ ɜɢɞɨɜɨɝɨ ɪɚɡɧɨɨɛɪɚɡɢɹ ɛɨɥɨɬɧɵɯ ɩɨɱɜ ɢ ɯɜɨɣɧɵɯ ɥɟɫɨɜ Ⱦɟɧɢɫɟɧɤɨɜ ȼ ɷɬɨɦ ɨɬɧɨɲɟɧɢɢ ɩɨɤɚɡɚɬɟɥɟɧ ɚɧɚɥɢɡ ɞɚɧ ɧɵɯ ɩɨɥɭɱɟɧɧɵɯ ɞɥɹ ɩɨɞɡɨɥɢɫɬɵɯ ɩɨɱɜ Ɇɨɫɤɨɜɫɤɨɣ ɨɛɥɚɫɬɢ ɫɮɨɪɦɢɪɨɜɚɧɧɵɯ ɩɨɞ ɟɥɶɧɢɤɚɦɢ ɢ ɥɢɫɬɜɟɧɧɵɦɢ ɥɟɫɚɦɢ Ȼɨɥɶɲɢɦ ɜɢɞɨɜɵɦ ɪɚɡɧɨɨɛɪɚɡɢɟɦ ɜɢ

ϭϳϵ

ɞɨɜ ɢ ɤɨɧɰɟɧɬɪɚɰɢɟɣ ɪɚɤɨɜɢɧɨɤ ɛɨɥɟɟ ɬɵɫ ɜ ɝ ɚɛɫɨɥɸɬɧɨ ɫɭɯɨɝɨ ɫɭɛɫɬɪɚ ɬɚ ɨɬɥɢɱɚɥɢɫɶ ɤɢɫɥɵɟ ɩɨɱɜɵ ɟɥɶɧɢɤɨɜ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫɨ ɡɧɚɱɢɬɟɥɶɧɨ ɛɨɥɟɟ ɛɟɞ ɧɵɦ ɥɢɩɧɹɤɨɦ ɜɢɞɨɜ ɨɤɨɥɨ ɬɵɫ ɷɤɡ ɋ ɝɥɭɛɢɧɨɣ ɩɨ ɩɪɨɮɢɥɸ ɩɨɱɜɵ ɭɫ ɥɨɜɢɹ ɨɛɢɬɚɧɢɸ ɞɥɹ ɬɟɫɬɚɰɟɣ ɭɯɭɞɲɚɸɬɫɹ ɭɜɟɥɢɱɢɜɚɟɬɫɹ ɩɥɨɬɧɨɫɬɶ ɢ ɭɦɟɧɶɲɚɟɬɫɹ ɩɨɪɢɫɬɨɫɬɶ ɩɨɱɜɵ ɜ ɫɜɹɡɢ ɫ ɱɟɦ ɢɡɦɟɧɹɟɬɫɹ ɟɟ ɜɨɞɧɨ-ɜɨɡɞɭɲɧɵɣ ɪɟɠɢɦ ȼɫɥɟɞɫɬɜɢɟ ɷɬɨɝɨ ɩɪɨɢɫɯɨɞɢɬ ɡɚɤɨɧɨɦɟɪɧɨɟ ɢɡɦɟɧɟɧɢɟ ɮɚɭɧɵ ɬɟɫɬɚɰɟɣ ɱɬɨ ɩɪɨɹɜɥɹɟɬɫɹ ɜ ɨɛɟɞɧɟɧɢɢ ɧɟ ɬɨɥɶɤɨ ɜɢɞɨɜɨɝɨ ɫɨɫɬɚɜɚ ɢ ɱɢɫɥɟɧɧɨɫɬɢ ɧɨ ɢ ɤɨɦɩɥɟɤɫɚ ɷɤɨɦɨɪɮɨɬɢɩɨɜ ɜ ɝɟɧɟɬɢɱɟɫɤɢɯ ɝɨɪɢɡɨɧɬɚɯ. ȼɟɪɬɢɤɚɥɶɧɨɟ ɪɚɫɩɪɟɞɟ ɥɟɧɢɟ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɨɛɭɫɥɨɜɥɟɧɨ ɫɨɞɟɪɠɚɧɢɟɦ ɝɭɦɭɫɚ ɪɚɡɜɢɬɨɫɬɶɸ ɚɤɤɭɦɭ ɥɹɬɢɜɧɵɯ ɝɨɪɢɡɨɧɬɨɜ ɤɚɱɟɫɬɜɨɦ ɢ ɤɨɥɢɱɟɫɬɜɨɦ ɨɪɝɚɧɢɱɟɫɤɨɝɨ ɜɟɳɟɫɬɜɚ ɩɨɫɬɭ ɩɚɸɳɟɝɨ ɜ ɩɨɱɜɭ ɫ ɪɚɫɬɢɬɟɥɶɧɵɦ ɨɩɚɞɨɦ Ɉɩɬɢɦɚɥɶɧɵɣ ɭɪɨɜɟɧɶ ɨɛɢɬɚɧɢɹ ɝɪɚɧɢɰɚ ɦɟɠɞɭ ɝɨɪɢɡɨɧɬɚɦɢ Ⱥ0 ɢ Ⱥ1 ɝɞɟ ɬɟɫɬɚɰɟɣ ɜ ɧɟɫɤɨɥɶɤɨ ɪɚɡ ɛɨɥɶɲɟ ɱɟɦ ɜ ɩɨɜɟɪɯɧɨɫɬɧɨɦ ɫɥɨɟ ɨɩɚɞɚ ȼ ɩɪɟɞɟɥɚɯ ɝɨɪɢɡɨɧɬɨɜ Ⱥ1 ɢ Ⱥ2 ɮɚɭɧɚ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɡɚɦɟɬɧɨ ɛɟɞɧɟɟ ɤɚɤ ɩɨ ɨɛɢɥɢɸ ɬɚɤ ɢ ɩɨ ɜɢɞɨɜɨɦɭ ɪɚɡɧɨɨɛɪɚɡɢɸ ɱɟɦ ɜ ɜɟɪɯ ɧɟɣ ɱɚɫɬɢ ɩɪɨɮɢɥɹ ɋɬɪɚɬɢɮɢɤɚɰɢɹ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɜ ɝɨɪɢɡɨɧɬɚɯ ɩɨɱɜ ɡɚɜɢɫɢɬ ɧɟ ɬɨɥɶɤɨ ɨɬ ɪɚɫɩɪɟɞɟɥɟɧɢɹ ɝɭɦɭɫɚ ɧɨ ɢ ɨɬ ɜɟɥɢɱɢɧɵ ɩɨɱɜɟɧɧɵɯ ɩɨɪ ɬɨɥɳɢɧɵ ɜɨɞɧɵɯ ɩɥɟɧɨɤ ɫɩɨɫɨɛɧɨɫɬɢ ɩɪɨɬɢɜɨɫɬɨɹɬɶ ɜɵɫɵɯɚɧɢɸ ɢ ɧɚɥɢɱɢɹ ɫɬɪɨɢɬɟɥɶɧɨɝɨ ɦɚɬɟɪɢɚɥɚ Ɂɚɤɨɧɨɦɟɪɧɨɫɬɢ ɜɟɪɬɢɤɚɥɶɧɨɝɨ ɪɚɫɩɪɟɞɟɥɟɧɢɹ ɩɪɨɫɬɟɣɲɢɯ ɬɟɫɧɨ ɫɜɹ ɡɚɧɵ ɫ ɬɢɩɨɦ ɩɨɱɜɵ ɢ ɦɨɝɭɬ ɫɥɭɠɢɬɶ ɛɨɥɟɟ ɱɭɬɤɢɦ ɢɧɞɢɤɚɬɨɪɨɦ ɢɯ ɫɜɨɣɫɬɜ ɱɟɦ

ɜɢɞɨɜɨɟ ɪɚɡɧɨɨɛɪɚɡɢɟ Kishaba, Mitchell, 2005).

Ɋɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɨɬɧɨɫɹɬ ɤ ɝɪɭɩɩɚɦ ɩɨɱɜɟɧɧɨɣ ɮɚɭɧɵ ɚɤɬɢɜɧɨ ɭɱɚɫɬ ɜɭɸɳɢɯ ɜ ɬɪɚɧɫɮɨɪɦɚɰɢɢ ɨɪɝɚɧɢɱɟɫɤɨɝɨ ɜɟɳɟɫɬɜɚ ɜ ɨɫɨɛɟɧɧɨɫɬɢ ɜ ɬɚɟɠɧɵɯ ɩɨɱɜɚɯ ɗɤɫɩɟɪɢɦɟɧɬɚɥɶɧɨ ɩɨɤɚɡɚɧɨ ɱɬɨ ɪɹɞ ɮɨɪɦ ɪɚɡɪɭɲɚɟɬ ɥɢɝɧɢɧ ɢ ɰɟɥɥɸ ɥɨɡɭ Schonborn ɹɜɥɹɹɫɶ ɩɟɪɜɢɱɧɵɦɢ ɞɟɫɬɪɭɤɬɨɪɚɦɢ ɨɩɚɞɚ ȼ ɰɢɤɥɟ ɛɢɨ ɝɟɧɧɨɝɨ ɤɪɟɦɧɟɡɟɦɚ ɜ ɤɨɬɨɪɨɦ ɪɨɥɶ ɠɢɜɨɬɧɵɯ ɜ ɰɟɥɨɦ ɜɟɫɶɦɚ ɧɟɜɟɥɢɤɚ ɬɟɫɬɚ ɰɟɢ ɫɢɧɬɟɡɢɪɭɸɬ ɞɢɨɤɫɢɞ ɤɪɟɦɧɢɹ ɧɟɨɛɯɨɞɢɦɵɣ ɞɥɹ ɩɨɫɬɪɨɟɧɢɹ ɪɚɤɨɜɢɧɨɤɊɚɯɥɟɟɜɚ Ɉɞɧɢɦ ɢɡ ɝɥɚɜɧɵɯ ɧɚɩɪɚɜɥɟɧɢɣ ɩɨɱɜɟɧɧɨɣ ɩɪɨɬɢɫɬɨɥɨɝɢɢ ɹɜ ɥɹɟɬɫɹ ɩɨɞɬɜɟɪɠɞɟɧɢɟ ɢ ɪɚɡɜɢɬɢɟ ɩɨɥɨɠɟɧɢɹ ɨ ɬɨɦ ɱɬɨ ɩɪɨɫɬɟɣɲɢɟ ɜɵɞɟɥɹɸɬ ɛɢɨɥɨɝɢɱɟɫɤɢ ɚɤɬɢɜɧɵɟ ɜɟɳɟɫɬɜɚ ɇɢɤɨɥɸɤ Ɋɹɞɨɦ ɢɫɫɥɟɞɨɜɚɧɢɣ ɭɫɬɚ ɧɨɜɥɟɧɨ ɱɬɨ ɜ ɡɨɧɚɯ ɤɨɪɧɟɜɵɯ ɫɢɫɬɟɦ ɪɚɫɬɟɧɢɣ ɩɪɨɢɫɯɨɞɢɬ ɚɤɬɢɜɧɨɟ ɧɚɤɨɩɥɟɧɢɟ ɩɪɨɫɬɟɣɲɢɯ ɝɞɟ ɢɯ ɤɨɥɢɱɟɫɬɜɨ ɜ ɞɟɫɹɬɤɢ ɢ ɫɨɬɧɢ ɪɚɡ ɛɨɥɶɲɟ ɱɟɦ ɜ ɨɤɪɭɠɚɸɳɟɣ ɫɪɟɞɟ Ɉɫɧɨɜɧɚɹ ɩɪɢɱɢɧɚ ɡɧɚɱɢɬɟɥɶɧɨɝɨ ɧɚɤɨɩɥɟɧɢɹ ɩɪɨɫɬɟɣɲɢɯ ɜ ɪɢɡɨɫɮɟɪɟ ɪɚɫ ɬɟɧɢɣ - ɩɨɜɵɲɟɧɧɨɟ ɫɨɫɪɟɞɨɬɨɱɟɧɢɟ ɪɚɡɥɢɱɧɵɯ ɛɚɤɬɟɪɢɣ ɤɨɬɨɪɵɦɢ ɩɪɨɫɬɟɣɲɢɟ ɩɢɬɚɸɬɫɹ. ɉɪɨɫɬɟɣɲɢɟ ɫɨɜɦɟɫɬɧɨ ɫ ɛɚɤɬɟɪɢɹɦɢ ɨɛɪɚɡɭɸɬ ɫɨɨɛɳɟɫɬɜɚ Ɂɚɜɚɪɡɢɧ 2004) ȼ ɛɨɥɶɲɢɧɫɬɜɟ ɫɥɭɱɚɟɜ ɩɪɨɞɭɰɢɪɭɟɬɫɹ ɝɟɬɟɪɨɚɭɤɫɢɧɵ ɤɨɬɨɪɵɟ ɨɤɚɡɵɜɚ ɸɬ ɢɡɛɢɪɚɬɟɥɶɧɨɟ ɪɨɫɬɨɚɤɬɢɜɢɪɭɸɳɟɟ ɞɟɣɫɬɜɢɟ ɧɚ ɜɫɯɨɠɟɫɬɶ ɫɟɦɹɧ ȼɵɞɟɥɟɧɢɟ ɜ ɤɭɥɶɬɭɪɚɯ ɚɦɟɛ ɢ ɚɡɨɬɨɛɚɤɬɟɪɨɜ ɧɚɛɨɪɚ ɪɨɫɬɨɚɤɬɢɜɢɪɭɸɳɢɯ ɢɧɞɨɥɶɧɵɯ ɜɟ ɳɟɫɬɜ, ɫɜɢɞɟɬɟɥɶɫɬɜɭɟɬ ɨ ɬɨɦ ɱɬɨ ɩɨɱɜɟɧɧɵɟ ɦɢɤɪɨɨɪɝɚɧɢɡɦɵ ɨɛɥɚɞɚɸɬ ɲɢɪɨ ɤɢɦɢ ɜɨɡɦɨɠɧɨɫɬɹɦɢ ɞɥɹ ɩɪɨɞɭɰɢɪɨɜɚɧɢɹ ɛɢɨɥɨɝɢɱɟɫɤɢ ɚɤɬɢɜɧɵɯ ɜɟɳɟɫɬɜ ɤɨ ɬɨɪɵɟ ɦɨɝɭɬ ɦɟɧɹɬɶɫɹ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɨɤɪɭɠɚɸɳɢɯ ɭɫɥɨɜɢɣ ɫɪɟɞɵ ɉɪɨɞɭɤɬɵ ɨɛɦɟɧɚ ɜɟɳɟɫɬɜ ɩɪɨɫɬɟɣɲɢɯ ɩɨɥɨɠɢɬɟɥɶɧɨ ɜɥɢɹɸɬ ɧɚ ɩɪɨɪɚɫɬɚɧɢɟ ɫɟɦɹɧ ɯɥɨɩ ɱɚɬɧɢɤɚ.

ɇɟɨɛɯɨɞɢɦɨɫɬɶ ɩɟɪɟɞɜɢɠɟɧɢɹ ɜ ɭɡɤɢɯ ɦɟɠɩɨɱɜɟɧɧɵɯ ɩɪɨɦɟɠɭɬɤɚɯ ɩɨɪɚɯ ɤɚɩɢɥɥɹɪɚɯ ©ɩɪɟɪɵɜɢɫɬɨɫɬɶªɪɚɫɩɪɟɞɟɥɟɧɢɹ ɜɥɚɝɢ ɢ ɪɚɡɧɨɨɛɪɚɡɢɟ ɮɢɡɢɱɟɫɤɢɯ ɮɨɪɦ ɧɟɩɨɫɬɨɹɧɫɬɜɨ ɫɬɟɩɟɧɢ ɭɜɥɚɠɧɟɧɢɹ ɨɩɪɟɞɟɥɹɸɬ ɫɜɨɟɨɛɪɚɡɢɟ ɩɨɱɜɵ ɤɚɤ ɫɪɟɞɵ ɨɛɢɬɚɧɢɹ ɩɪɨɫɬɟɣɲɢɯ Kishaba, Mitchell ɇɚɱɚɥɶɧɵɦ ɷɬɚɩɨɦ ɮɨɪɦɢ ɪɨɜɚɧɢɹ ɤɨɦɩɥɟɤɫɚ ɩɨɱɜɨɨɛɢɬɚɸɳɢɯ ɬɟɫɬɚɰɟɣ ɩɨ-ɜɢɞɢɦɨɦɭ ɛɵɥɨ ɩɨɫɬɟɩɟɧɧɨɟ ɨɫɜɨɟɧɢɟ ɜɨɞɧɵɦɢ ɝɪɭɩɩɚɦɢ ɩɪɢɛɪɟɠɧɨɣ ɪɚɫɬɢɬɟɥɶɧɨɫɬɢ ɢ ɩɪɨɧɢɤɧɨɜɟɧɢɟ ɢɯ ɜɨ ɜɥɚɠɧɵɣ ɦɨɯɨɜɨɣ ɩɨɤɪɨɜ ɩɨɞ ɤɪɨɧɚɦɢ ɞɟɪɟɜɶɟɜ Ⱦɨɝɟɥɶ Ȼɢɨɬɨɩɵ ɜ ɤɨɬɨ ɪɵɯ ɨɛɢɬɚɸɬ ɪɚɤɨɜɢɧɧɵɟ ɚɦɟɛɵ ɒɟɧɛɨɪɧ ɪɚɫɩɨɥɨɠɢɥ ɜ ɜɢɞɟ ɩɨɫɥɟɞɨɜɚ ɬɟɥɶɧɨɝɨ ɪɹɞɚ ɜɨɞɧɚɹ ɪɚɫɬɢɬɟɥɶɧɨɫɬɶ ɫɮɚɝɧɨɜɵɟ ɢ ɥɢɫɬɨɫɬɟɛɟɥɶɧɵɟ ɦɯɢ ɞɨɧɧɵɟ

ϭϴϬ

ɨɬɥɨɠɟɧɢɹ ɜɨɞɨɟɦɨɜ ɢ ɝɭɦɭɫɨɜɵɣ ɝɨɪɢɡɨɧɬ ɩɨɱɜɵ ɗɤɨɥɨɝɢɱɟɫɤɢɣ ɪɹɞ ɯɚɪɚɤɬɟɪɢ ɡɭɟɬɫɹ ɛɨɥɟɟ ɠɟɫɬɤɢɦɢ ɭɫɥɨɜɢɹɦɢ ɨɛɢɬɚɧɢɹ ɤɨɬɨɪɵɦ ɭɞɨɜɥɟɬɜɨɪɹɸɬ ɜɫɟ ɦɟɧɶɲɟɟ ɤɨɥɢɱɟɫɬɜɨ ɠɢɡɧɟɧɧɵɯ ɮɨɪɦ ɑɟɦ ɛɨɥɟɟ ɠɟɫɬɤɢɟ ɬɪɟɛɨɜɚɧɢɹ ɩɪɟɞɴɹɜɥɹɟɬ ɫɪɟɞɚ ɬɟɦ ɨɬɱɟɬɥɢɜɟɟ ɫɬɚɧɨɜɹɬɫɹ ɚɞɚɩɬɚɰɢɢ ɤ ɧɟɣ ɜɵɪɚɠɟɧɧɵɟ ɜ ɝɭɦɭɫɨɜɨɦ ɝɨɪɢɡɨɧɬɟ ɝɞɟ ɨɫɧɨɜɧɵɦ ɮɚɤɬɨɪɨɦ ɹɜɥɹɟɬɫɹ ɤɨɥɢɱɟɫɬɜɨ ɞɨɫɬɭɩɧɨɣ ɜɥɚɝɢ Ⱥɞɚɩɬɚɰɢɢ ɫɜɹɡɚ ɧɵ ɫ ɦɨɪɮɨɥɨɝɢɟɣ ɪɚɤɨɜɢɧɵ ɮɨɪɦɢɪɭɸɬ ɩɪɟɞɜɟɪɢɹ ɢɡɨɥɢɪɭɸɳɟɝɨ ɭɫɬɶɟ ɩɫɟɜ ɞɨɫɬɨɦ ɨɬ ɜɧɟɲɧɟɣ ɫɪɟɞɵ Ƚɟɥɶɰɟɪ Ƚɥɚɜɧɵɟ ɬɟɧɞɟɧɰɢɢ ɚɞɚɩɬɢɜɧɵɯ ɩɪɟ ɨɛɪɚɡɨɜɚɧɢɣ ɭ ɤɨɪɧɟɧɨɠɟɤ ɨɫɜɨɢɜɲɢɯ ɩɨɱɜɟɧɧɭɸ ɫɪɟɞɭ ɨɛɢɬɚɧɢɹ ɤɚɫɚɸɬɫɹ ɦɨɪɮɨɥɨɝɢɢ ɪɚɤɨɜɢɧɤɢ ɢ ɫɜɹɡɚɧɵ ɫ ɨɛɟɫɩɟɱɟɧɢɟɦ ɩɟɪɟɞɜɢɠɟɧɢɹ ɢ ɡɚɳɢɬɵ ɨɬ ɜɵ ɫɵɯɚɧɢɹ ȼ ɩɪɨɫɬɟɣɲɟɦ ɫɥɭɱɚɟ Cochilopodium ɪɚɤɨɜɢɧɤɚ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɜ ɫɭɳɧɨɫɬɢ ɬɨɧɤɭɸ ɩɥɚɫɬɢɱɧɭɸ ɨɛɨɥɨɱɤɭ ɫɩɨɫɨɛɧɭɸ ɞɟɮɨɪɦɢɪɨɜɚɬɶɫɹ Ɋɚɤɨɜɢɧ ɤɚ Arcella ɬɚɤɠɟ ɨɞɧɨɫɥɨɣɧɚ ɢ ɫɨɫɬɨɢɬ ɢɡ ɨɪɝɚɧɢɱɟɫɤɨɝɨ ɜɟɳɟɫɬɜɚ ɛɟɥɤɨɜɨɣ ɩɪɢ ɪɨɞɵ ɢ ɢɦɟɟɬ ɜɫɟ ɯɚɪɚɤɬɟɪɧɵɟ ɱɟɪɬɵ ɧɚɫɬɨɹɳɟɣ ɪɚɤɨɜɢɧɤɢ Ɋɚɤɨɜɢɧɤɚ ɫ ɩɨɤɪɵɬɢ ɟɦ ɢɡ ɦɢɧɟɪɚɥɶɧɵɯ ɱɚɫɬɢɰ ɜ ɜɨɞɧɨɣ ɫɪɟɞɟ ɩɪɚɤɬɢɱɟɫɤɢ ɨɞɧɨɫɥɨɣɧɵ ɢ ɩɪɟɞɫɬɚɜɥɹɸɬ ɫɨɛɨɣ ɨɪɝɚɧɢɱɟɫɤɨɟ ɰɟɦɟɧɬɢɪɭɸɳɟɟ ɜɟɳɟɫɬɜɨ Centropyxis aculeata ɫ ɪɚɡɛɪɨɫɚɧɧɵɦɢ ɧɚ ɧɟɦ ɨɬɞɟɥɶɧɵɦɢ ɤɫɟɧɨɫɨɦɚɦɢ ȼ ɞɨɧɧɵɯ ɨɬɥɨɠɟɧɢ ɹɯ ɢ ɜ ɩɨɱɜɟ ɭ ɚɦɺɛ ɢɦɟɸɬɫɹ ɩɥɨɬɧɨ ɰɟɦɟɧɬɢɪɨɜɚɧɧɵɟ ɤɫɟɧɨɫɨɦɵ ɢ ɩɪɟɞɫɬɚɜɥɹɸɬ ɫɩɥɨɲɧɨɣ ɩɪɨɱɧɵɣ ɩɨɤɪɨɜ ɉɨɤɪɵɜɚɸɳɢɟ ɪɚɤɨɜɢɧɤɢ ɢɞɢɨɫɨɦɵ ɬɚɤɠɟ ɩɥɨɬɧɨ ɩɪɢɥɟɝɚɸɬ ɞɪɭɝ ɤ ɞɪɭɝɭ ɨɛɪɚɡɭɹ ɦɢɧɢɚɬɸɪɧɵɣ ɩɚɧɰɢɪɶ Tracheleuglypha, Euglypha ɢ ɞɪɭɝɢɟ Ȼɟɣɟɪ ɢ ɞɪ ɇɚɥɢɱɢɟ ɜ ɩɨɱɜɟ ɫɢɫɬɟɦɵ ɦɟɥɤɢɯ ɩɨɥɨɫɬɟɣ ɩɪɢɜɟɥɨ ɤ ɡɧɚɱɢɬɟɥɶɧɨɦɭ ɭɦɟɧɶɲɟɧɢɸ ɪɚɡɦɟɪɨɜ ɩɨɱɜɨɨɛɢɬɚɸɳɢɯ ɩɪɨɫɬɟɣɲɢɯ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɨɛɢɬɚɬɟɥɹɦɢ ɜɨɞɧɨɣ ɫɪɟɞɵ ɋɪɟɞɧɢɟ ɪɚɡɦɟɪɵ ɩɨɱɜɨɨɛɢɬɚɸɳɢɯ ɮɨɪɦ

± ɩɨɪɹɞɤɚ ɦɤɦ Ɇɨɪɮɨɥɨɝɢɹ ɪɚɤɨɜɢɧɨɤ ɡɚɜɢɫɢɬ ɨɬ ɫɜɨɣɫɬɜ ɛɢɨɬɨɩɚ ɢ ɡɚɩɚɫɚ ɜɥɚɝɢ Ɋɚɤɨɜɢɧɤɢ ɩɨɱɜɨɨɛɢɬɚɸɳɢɯ ɬɟɫɬɚɰɟɣ ɨɬɥɢɱɚɸɬɫɹ ɩɪɨɫɬɨɬɨɣ ɮɨɪɦɵ ɜɧɟɲɧɢɟ ɜɵɫɬɭɩɵ ɢɝɥɵ ɲɢɩɵ ɫɩɨɫɨɛɫɬɜɭɸɳɢɟ ɩɚɪɟɧɢɸ ɜ ɜɨɞɟ ɢɥɢ ɮɢɤɫɚɰɢɢ ɧɚ ɪɚɫɬɢɬɟɥɶɧɨɫɬɢ ɨɬɫɭɬɫɬɜɭɸɬ ɢɥɢ ɪɚɡɜɢɬɵ ɫɥɚɛɨ ɇɚɢɛɨɥɶɲɭɸ ɪɨɥɶ ɜ ɡɚɫɟɥɟ ɧɢɢ ɪɚɤɨɜɢɧɧɵɦɢ ɚɦɟɛɚɦɢ ɦɢɧɟɪɚɥɶɧɵɯ ɫɥɨɟɜ ɩɨɱɜɵ ɫɵɝɪɚɥɨ ɩɪɢɨɛɪɟɬɟɧɢɟ ɨɤ ɪɭɝɥɨɣ ɮɨɪɦɵ ɢ ɭɩɥɨɳɟɧɧɨɣ ɜɟɧɬɪɚɥɶɧɨɣ ɩɨɜɟɪɯɧɨɫɬɢ

ɂɡɦɟɧɟɧɢɹ ɜ ɦɨɪɮɨɥɨɝɢɢ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɢɡ ɜɨɞɧɨɣ ɢ ɩɨɱɜɟɧɧɨɣ ɫɪɟɞɵ ɨɛɢɬɚɧɢɹ ɫɜɹɡɚɧɵ ɫ ɪɚɫɩɨɥɨɠɟɧɢɟɦ ɢ ɫɬɪɨɟɧɢɟɦ ɭɫɬɶɟɜ ȼ ɩɨɱɜɚɯ ɩɪɟɨɛɥɚɞɚɸɬ ɮɨɪɦɵ ɫ ɬɟɧɞɟɧɰɢɟɣ ɤ ɭɦɟɧɶɲɟɧɢɸ ɭɫɬɶɹ ɢ ɢɡɨɥɹɰɢɢ ɟɝɨ ɨɬ ɜɧɟɲɧɟɣ ɫɪɟɞɵ ɱɬɨ ɫɧɢɠɚɟɬ ɨɩɚɫɧɨɫɬɶ ɜɵɫɵɯɚɧɢɹ ɩɪɨɫɬɟɣɲɢɯ ɋɩɟɰɢɚɥɢɡɚɰɢɹ ɩɨɱɜɟɧɧɵɯ ɮɨɪɦ ɡɚ ɤɥɸɱɚɸɳɚɹɫɹ ɜ ɢɡɨɥɹɰɢɢ ɭɫɬɶɹ ɢ ɫɨɡɞɚɧɢɢ ɛɭɮɟɪɧɨɝɨ ɩɪɨɫɬɪɚɧɫɬɜɚ ɦɟɠɞɭ ɨɪɝɚ ɧɢɡɦɨɦ ɢ ɚɬɦɨɫɮɟɪɨɣ ɢɞɟɬ ɩɨ ɩɭɬɢ ɪɚɡɜɢɬɢɹ ɩɥɚɝɢɨɫɬɨɦɢɢ ɢ ɤɪɢɩɬɨɫɬɨɦɢɢ ɍ ɩɥɚɝɢɨɫɬɨɦɧɵɯ ɮɨɪɦ ɋentropyxidae, Euglyphidae ɭɫɬɶɟ ɫɞɜɢɧɭɬɨ ɩɨ ɜɟɧɬɪɚɥɶɧɨɣ ɩɨɜɟɪɯɧɨɫɬɢ ɤ ɟɟ ɤɪɚɸ ɱɬɨ ɩɪɢɜɨɞɢɬ ɤ ɨɛɪɚɡɨɜɚɧɢɸ ɩɟɪɟɞɧɟɝɨ ɢ ɡɚɞɧɟɝɨ ɤɨɧɰɨɜ ɪɚɤɨɜɢɧɤɢ ɛɢɥɚɬɟɪɚɥɶɧɚɹ ɫɢɦɦɟɬɪɢɹ ɉɟɪɟɞɧɢɣ ɤɨɧɟɰ ɧɟɫɭɳɢɣ ɭɫɬɶɟ ɭɩɥɨɳɚ ɟɬɫɹ ɜ ɡɚɞɧɟɦ ɛɨɥɶɲɟɝɨ ɨɛɴɟɦɚ ³ɛɪɸɲɤɨ´ ɩɨɦɟɳɚɟɬɫɹ ɨɫɧɨɜɧɚɹ ɦɚɫɫɚ ɰɢɬɨ ɩɥɚɡɦɵ ɉɨɞɨɛɧɚɹ ɬɟɧɞɟɧɰɢɹ ɢɥɢ ɟɞɜɚ ɧɚɦɟɱɟɧɚ ɢɥɢ ɞɨɫɬɚɬɨɱɧɨ ɪɚɡɜɢɬɚ ɡɚ ɫɱɟɬ ɭɝɥɭɛɥɟɧɢɹ ɭɫɬɶɹ ɢ ɨɛɪɚɡɨɜɚɧɢɹ ɩɪɢɭɫɬɶɟɜɨɣ ɜɨɪɨɧɤɢ ɩɪɢ ɷɬɨɦ ɫɩɢɧɧɚɹ ɫɬɨɪɨɧɚ ɪɚɤɨɜɢɧɤɢ ɦɨɠɟɬ ɨɤɪɭɠɚɬɶ ɭɫɬɶɟ ɜɚɥɢɤɨɦ ɜ ɜɢɞɟ ɛɨɥɟɟ ɢɥɢ ɦɟɧɟɟ ɜɵɪɚɠɟɧɧɨɝɨ ɤɨɡɵɪɶɤɚ ɩɥɚɝɢɨɫɬɨɦɢɹ ɫ ɤɨɡɵɪɶɤɨɦ ɩɪɢɤɪɵɜɚɹ ɟɝɨ ɨɬ ɡɚɫɨɪɟɧɢɹ ɢ ɭɦɟɧɶɲɚɹ ɢɫɩɚɪɟɧɢɟ Ʉɪɢɩɬɨɫɬɨɦɢɹ ɫɨɡɞɚɟɬɫɹ ɡɚ ɫɱɟɬ³ɩɟɪɟɤɪɵɜɚɧɢɹ´ɳɟɥɟɜɢɞɧɨɝɨ ɭɫɬɶɹ ɪɚɫɩɨɥɨɠɟɧɧɨɝɨ ɧɚ ɛɨɥɟɟ ɢɥɢ ɦɟɧɟɟ ɭɝɥɭɛɥɟɧɧɨɣ ɛɪɸɲɧɨɣ ɩɨɜɟɪɯɧɨɫɬɢ ɧɚɜɢ ɫɚɸɳɢɦ ɤɨɡɵɪɶɤɨɦ ɪɢɫɭɧɨɤ

ϭϴϭ

Ɋɢɫɭɧɨɤ ɋɯɟɦɚ ɦɨɪɮɨɬɢɩɨɜ ɪɚɤɨɜɢɧɨɤ Ƚɟɥɶɰɟɪ 1- ɚɩɥɚɤɚɬɢɡɦ ɚ - ɩɥɚɧ ɛ± ɩɪɨɮɢɥɶ± ɬɪɚɯɟɥɨɫɬɨɦɢɹ± ɚɤɪɨɫɬɨɦɢɹ ɚ - ɛɟɡ ɥɚɬɟɪɚɥɶɧɨɝɨ ɫɠɚɬɢɹ ɛ - ɫ ɥɚɬɟɪɚɥɶɧɵɦ ɫɠɚɬɢɟɦ 4 - ɰɟɧɬɪɨɫɬɨɦɢɹ ɫ ɭɩɥɨɳɟɧɧɨɣ ɜɟɧɬɪɚɥɶɧɨɣ ɩɨɜɟɪɯɧɨɫɬɶɸ ɚ - ɩɥɚɧ ɛ± ɩɪɨɮɢɥɶ -

Ⱥ - ɩɥɚɝɢɨɫɬɨɦɢɹ ɩɪɨɫɬɚɹ Ȼ - ɫ ɤɨɡɵɪɶɤɨɦ ɚ - ɩɥɚɧ ɛ± ɩɪɨɮɢɥɶ -Ⱥ - ɤɪɢɩɬɨ ɫɬɨɦɢɹ ɩɪɨɫɬɚɹ Ȼ - ɫ ɤɨɡɵɪɶɤɨɦ ɚ - ɩɥɚɧ ɛ - ɩɪɨɮɢɥɶ.

ɇɚ ɨɫɧɨɜɚɧɢɢ ɨɩɢɫɚɧɧɵɯ ɨɫɨɛɟɧɧɨɫɬɟɣ ɫɬɪɨɟɧɢɹ ɩɪɟɞɥɨɠɟɧɚ ɤɥɚɫɫɢɮɢɤɚ ɰɢɹ ɦɨɪɮɨɥɨɝɢɱɟɫɤɢɯ ɬɢɩɨɜ ɢ ɭɫɬɚɧɨɜɥɟɧɚ ɜɡɚɢɦɨɫɜɹɡɶ ɦɟɠɞɭ ɦɨɪɮɨɥɨɝɢɟɣ ɪɚ ɤɨɜɢɧ ɢ ɫɜɨɣɫɬɜɚɦɢ ɛɢɨɬɨɩɚ Bonnet Ɍɪɚɯɟɥɨɫɬɨɦɧɵɣ ɬɢɩ - ɪɚɤɨɜɢɧɤɢ ɫ ɞɭɝɨɨɛɪɚɡɧɵɦ ɭɫɬɶɟɦ ɫɨɫɬɨɹɬ ɢɡ ɞɜɭɯ ɱɚɫɬɟɣ± ©ɲɟɣɤɢªɢ©ɛɪɸɲɤɚªɩɪɢɱɟɦ ɨɫɶ ɲɟɣɤɢ ɫɦɟɳɟɧɚ ɩɨ ɨɬɧɨɲɟɧɢɸ ɤ ɨɫɢ ɛɪɸɲɤɚ ɱɬɨ ɧɚɪɹɞɭ ɫ ɛɨɤɨɜɵɦ ɭɩɥɨɳɟɧɢɟɦ ɪɚɤɨɜɢɧɤɢ ɫɨɡɞɚɟɬ ɞɜɭɫɬɨɪɨɧɧɸɸ ɫɢɦɦɟɬɪɢɸ ɒɟɣɤɚ ɨɬɞɟɥɟɧɚ ɨɬ ɨɫɧɨɜɧɨɣ ɱɚɫɬɢ ɪɚɤɨɜɢɧɵ ɞɢɚɮɪɚɝɦɨɣ ɢɥɢ ɩɟɪɟɦɵɱɤɨɣ ɢ ɢɝɪɚɟɬ ɪɨɥɶ ɩɪɟɞɞɜɟɪɢɹ Leariereusia). Ⱥɪɰɟɥɥɨɜɢɞɧɵɣ ɬɢɩ Ɋɚɤɨɜɢɧɤɚ ɭɩɥɨɳɟɧɧɨ-ɞɢɫɤɨɜɢɞɧɨɣ ɮɨɪɦɵ ɩɫɟɜɞɨɫɬɨɦ ɪɚɫ ɩɨɥɨɠɟɧ ɰɟɧɬɪɚɥɶɧɨ ɧɚɛɥɸɞɚɟɬɫɹ ɪɚɞɢɚɥɶɧɚɹ ɫɢɦɦɟɬɪɢɹ Arcella Ⱥɤɪɨɫɬɨɦɧɵɣ ɬɢɩ Ɋɚɤɨɜɢɧɤɚ ɫ ɬɟɪɦɢɧɚɥɶɧɨ ɪɚɫɩɨɥɨɠɟɧɧɵɦ ɭɫɬɶɟɦ ɩɪɢ ɨɫɟɜɨɣ ɫɢɦɦɟɬɪɢɢ ɜɨɡɦɨɠɧɨ ɥɚɬɟɪɚɥɶɧɨɟ ɫɠɚɬɢɟ Difflugia, Nebela, Euglypha ɢ ɞɪɭɝɢɟ ɐɢɤɥɨɫɬɨɦ ɧɵɣ ɬɢɩ ɋɮɟɪɢɱɟɫɤɢɟ ɢɥɢ ɩɨɥɭɫɮɟɪɢɱɟɫɤɢɟ ɮɨɪɦɵ ɫ ɭɩɥɨɳɟɧɧɨɣ ɰɟɧɬɪɚɥɶɧɨɣ ɩɨɜɟɪɯɧɨɫɬɶɸ ©ɩɨɞɨɲɜɨɣª ɢ ɰɟɧɬɪɚɥɶɧɨ ɪɚɫɩɨɥɨɠɟɧɧɵɦ ɭɫɬɶɟɦ ɫɢɦɦɟɬɪɢɹ ɨɫɟɜɚɹ Phryganella, Cyclopyxis ɢ ɞɪ ɉɥɚɝɢɨɫɬɨɦɧɵɣ ɬɢɩ ɚ ɩɪɨɫɬɨɣ± ɪɚɫɩɨɥɨ ɠɟɧ ɷɤɫɰɟɧɬɪɢɱɧɨ ɧɚ ɜɟɧɬɪɚɥɶɧɨɣ ɫɬɨɪɨɧɟ ɩɨɥɨɫɬɶ ɪɚɤɨɜɢɧɤɢ ɧɟ ɪɚɡɞɟɥɟɧɚ ɧɚ

ɛɪɸɲɤɨ ɢ ɤɨɡɵɪɟɤ Centropyxis plagiostoma, Geopyxella sylvicola ɛ ɫ ɤɨɡɵɪɶɤɨɦ

± ɩɫɟɜɞɨɫɬɨɦ ɛɨɥɟɟ ɢɥɢ ɦɟɧɟɟ ɩɪɢɤɪɵɬ ɜɵɫɬɭɩɨɦ ɞɨɪɡɚɥɶɧɨɣ ɫɬɟɧɤɢ Ɂɚ ɫɱɟɬ ɭɝ ɥɭɛɥɟɧɢɹ ɛɪɸɲɧɨɣ ɫɬɨɪɨɧɵ ɩɨɥɨɫɬɶ ɪɚɤɨɜɢɧɤɢ ɪɚɡɞɟɥɟɧɚ ɧɚ ɛɪɸɲɤɨ ɢ ɤɨɡɵɪɟɤ (Centropyxis aerophila, C. sylvatica Ȼɢɥɚɬɟɪɚɥɶɧɚɹ ɫɢɦɦɟɬɪɢɹ Ʉɪɢɩɬɨɫɬɨɦɧɵɣ ɬɢɩ ɚ ɩɪɨɫɬɨɣ± ɳɟɥɟɜɢɞɧɵɣ ɷɤɫɰɟɧɬɪɢɱɧɨ ɪɚɫɩɨɥɨɠɟɧɧɵɣ ɧɚ ɜɟɧɬɪɚɥɶɧɨɣ ɩɨ ɞɨɲɜɟ ɩɫɟɜɞɨɫɬɨɦ ɩɪɢɤɪɵɬ ɜɵɫɬɭɩɨɦ ɞɨɪɡɚɥɶɧɨɣ ɫɬɟɧɤɢ ɉɨɥɨɫɬɶ ɪɚɤɨɜɢɧɤɢ ɧɟ ɪɚɡɞɟɥɟɧɚ Plagiopyxis minuta, P. penardi ɛ ɫ ɤɨɡɵɪɶɤɨɦ ɷɤɫɰɟɧɬɪɢɱɧɵɣ ɳɟɥɟ ɜɢɞɧɵɣ ɩɫɟɜɞɨɫɬɨɦ ɭɝɥɭɛɥɟɧ ɜɧɭɬɪɶ ɪɚɤɨɜɢɧɤɢ ɢ ɫɤɪɵɬ ɤɨɡɵɪɶɤɨɦ ɉɨɥɨɫɬɶ ɫɨ ɟɞɢɧɹɸɳɚɹ ɟɝɨ ɫ ɜɧɟɲɧɟɣ ɫɪɟɞɨɣ ɭɡɤɚɹ Ɋɚɤɨɜɢɧɤɚ ɪɚɡɞɟɥɟɧɚ ɧɚ ɛɪɸɲɤɨ ɢ ɤɨɡɵ

ϭϴϮ

ɪɟɤ Plagiopyxis callida, Paracentropyxis mimetica Ȼɢɥɚɬɟɪɚɥɶɧɚɹ ɫɢɦɦɟɬɪɢɹ

Ʉɪɨɦɟ ɭɤɚɡɚɧɧɵɯ ɜɵɲɟ ɚɞɚɩɬɚɰɢɣ ɬɚɤɠɟ ɫɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ ɢ ɞɪɭɝɢɟ ɦɨɪɮɨɥɨɝɢ ɱɟɫɤɢɟ ɚɞɚɩɬɚɰɢɢ ɮɨɪɦɚ ɪɚɡɦɟɪ ɪɚɤɨɜɢɧɨɤ ɪɚɫɩɨɥɨɠɟɧɢɟ ɫɬɪɨɟɧɢɟ ɢ ɪɚɡɦɟɪ ɩɫɟɜɞɨɫɬɨɦɚ ɜɧɟɲɧɢɟ ɜɵɫɬɭɩɵ ɢɝɥɵ ɲɢɩɵ ɧɚɥɢɱɢɟ ɩɥɨɫɤɨɣ ɜɟɧɬɪɚɥɶɧɨɣ ɩɨ ɞɨɲɜɵ ɨɛɪɚɡɨɜɚɧɢɟ ɜɪɟɦɟɧɧɨɣ ɨɛɨɥɨɱɤɢ ɰɢɫɬɵ Ʉɨɪɝɚɧɨɜɚ Ⱦɥɹ ɭɫɩɟɲɧɨɝɨ ɡɚɫɟɥɟɧɢɹ ɩɨɱɜɵ ɜɨɞɧɵɟ ɠɢɜɨɬɧɵɟ ɞɨɥɠɧɵ ɨɛɥɚɞɚɬɶ ɫɥɟɞɭɸɳɢɦɢ ɮɢɡɢɨɥɨɝɢɱɟɫɤɢɦɢ ɫɜɨɣɫɬɜɚɦɢ ɬɨɥɟɪɚɧɬɧɨɫɬɶɸ ɤ ɢɡɦɟɧɟɧɢɸ ɪɇ ɩɪɨɫɬɨɬɨɣ ɪɚɡ ɦɧɨɠɟɧɢɹ ɢ ɧɟɬɪɟɛɨɜɚɬɟɥɶɧɨɫɬɶɸ ɤ ɩɢɳɟ Stout Ɋɚɤɨɜɢɧɧɵɟ ɚɦɺɛɵ ɫ ɜɵ ɫɨɤɢɦ ɭɪɨɜɧɟɦ ɦɨɪɮɨɥɨɝɢɱɟɫɤɨɣ ɢɡɦɟɧɱɢɜɨɫɬɢ ɢ ɚɞɚɩɬɚɰɢɹɦɢ ɤ ɷɤɫɬɪɟɦɚɥɶɧɵɦ ɭɫɥɨɜɢɹɦ ɨɛɢɬɚɧɢɹ ɨɫɜɨɢɥɢ ɩɨɱɜɟɧɧɭɸ ɫɪɟɞɭ ɨɛɢɬɚɧɢɹ ɩɪɚɤɬɢɱɟɫɤɢ ɩɨɜɫɟɦɟɫɬ ɧɨ.

Ɂɚɝɪɹɡɧɟɧɢɟ ɩɨɱɜɵ ɧɟɮɬɶɸ ɢ ɧɟɮɬɟɩɪɨɞɭɤɬɚɦɢ ɹɜɥɹɟɬɫɹ ɨɞɧɢɦ ɢɡ ɧɚɢɛɨɥɟɟ ɪɚɫɩɪɨɫɬɪɚɧɟɧɧɵɯ ɡɚɝɪɹɡɧɢɬɟɥɟɣ ɋɢɛɢɪɢ ɉɨɫɬɭɩɥɟɧɢɟ ɧɟɮɬɢ ɜ ɩɨɱɜɭ ɜɵɡɵɜɚɟɬ ɭɜɟɥɢɱɟɧɢɟ ɫɨɞɟɪɠɚɧɢɹ ɨɪɝɚɧɢɱɟɫɤɨɝɨ ɭɝɥɟɪɨɞɚ ɢ ɛɢɬɭɦɢɧɨɡɧɵɯ ɜɟɳɟɫɬɜ ȼ ɫɨ ɫɬɚɜɟ ɩɨɱɜɟɧɧɨɝɨ ɜɨɡɞɭɯɚ ɩɪɟɜɚɥɢɪɭɸɬ ɥɟɝɤɢɟ ɮɪɚɤɰɢɢ ɧɟɮɬɢ ɫɧɢɠɚɟɬɫɹ ɜɨɞɨɩɨ ɝɥɨɬɢɬɟɥɶɧɚɹ ɢ ɜɨɞɨɭɞɟɪɠɢɜɚɸɳɚɹ ɫɩɨɫɨɛɧɨɫɬɶ ɩɨɱɜɵ ɩɨɜɵɲɚɟɬɫɹ ɜɨɞɨɩɪɨɧɢ ɰɚɟɦɨɫɬɶ ɭɦɟɧɶɲɚɟɬɫɹ ɫɨɞɟɪɠɚɧɢɟ ɩɨɞɜɢɠɧɵɯ ɮɨɪɦ ɚɡɨɬɚ ɢ ɮɨɫɮɨɪɚ ɩɨɜɵɲɚɟɬɫɹ ɫɨɞɟɪɠɚɧɢɟ ɩɨɞɜɢɠɧɵɯ ɮɨɪɦ ɦɢɤɪɨɷɥɟɦɟɧɬɨɜ Ʉɚɪɬɚɲɟɜ ɋɜɟ ɞɟɧɢɹ ɨ ɜɡɚɢɦɨɞɟɣɫɬɜɢɢ ɩɪɨɫɬɟɣɲɢɯ ɫ ɧɟɮɬɶɸ ɢ ɟɟ ɩɪɨɢɡɜɨɞɧɵɦɢ ɨɝɪɚɧɢɱɟɧɵ ɍɫɬɚɧɨɜɥɟɧɨ ɱɬɨ ɩɪɨɫɬɟɣɲɢɟ ɨɬɧɨɫɢɬɟɥɶɧɨ ɛɨɥɟɟ ɭɫɬɨɣɱɢɜɵ ɤ ɡɚɝɪɹɡɧɟɧɢɸ ɩɨɱ ɜɵ ɧɟɮɬɶɸ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɞɪɭɝɢɦɢ ɝɪɭɩɩɚɦɢ ɛɟɫɩɨɡɜɨɧɨɱɧɵɯ ɂɧɮɭɡɨɪɢɢ ɠɝɭ ɬɢɤɨɧɨɫɰɵ ɢ ɝɨɥɵɟ ɚɦɟɛɵ ɩɨɫɬɨɹɧɧɨ ɩɪɢɫɭɬɫɬɜɭɸɬ ɜ ɡɚɝɪɹɡɧɟɧɧɨɣ ɧɟɮɬɶɸ ɩɨɱɜɟ ɢ ɯɚɪɚɤɬɟɪɢɡɭɸɬɫɹ ɪɚɡɧɨɨɛɪɚɡɧɵɦ ɜɢɞɨɜɵɦ ɫɨɫɬɚɜɨɦ ȼ ɩɟɪɜɵɣ ɝɨɞ ɡɚɝɪɹɡɧɟɧɢɣ ɧɚɛɥɸɞɚɟɬɫɹ ɡɧɚɱɢɬɟɥɶɧɨɟ ɭɜɟɥɢɱɟɧɢɟ ɨɛɢɥɢɹ ɩɪɨɫɬɟɣɲɢɯ Ʌɚɛɨɪɚɬɨɪɧɵɟ ɨɩɵɬɵ ɫ ɪɚɡɥɢɱɧɵɦɢ ɤɨɧɰɟɧɬɪɚɰɢɹɦɢ ɧɟɮɬɢ ɨɬ ɜɟɫɚ ɩɨɱɜɵ ɩɨɤɚɡɚɥɢ ɱɬɨ ɩɪɨɫɬɟɣɲɢɟ ɦɨɝɭɬ ɜɵɞɟɪɠɢɜɚɬɶ ɨɬɧɨɫɢɬɟɥɶɧɨ ɜɵɫɨɤɢɟ ɭɪɨɜɧɢ ɡɚɝɪɹɡɧɟɧɢɣ Ȼɨ ɪɢɫɨɜɢɱ Ⱥɪɬɟɦɶɟɜɚ ɀɟɪɟɛɰɨɜ ) ɑɚɫɬɢɱɧɨ ɪɚɡɥɨɠɢɜɲɚɹɫɹ ɫɵɪɚɹ ɧɟɮɬɶ ɧɟ ɨɤɚɡɵɜɚɟɬ ɫɭɳɟɫɬɜɟɧɧɨɝɨ ɜɥɢɹɧɢɹ ɧɚ ɫɤɨɪɨɫɬɶ ɪɨɫɬɚ ɢɧɮɭɡɨɪɢɣ ɢ ɚɦɟɛ, ɨɬɦɟɱɚɟɬɫɹ ɥɢɲɶ ɫɥɚɛɨ ɜɵɪɚɠɟɧɧɚɹ ɬɟɧɞɟɧɰɢɹ ɤ ɭɜɟɥɢɱɟɧɢɸ ɢɯ ɪɚɡɦɟɪɨɜ

4.2 ȼɥɢɹɧɢɟ ɧɟɮɬɟɡɚɝɪɹɡɧɟɧɢɣ ɧɚ ɫɨɨɛɳɟɫɬɜɚ ɪɚɤɨɜɢɧɧɵɯ ɚɦɺɛ

Ɉɛɴɟɤɬɨɦ ɢɫɫɥɟɞɨɜɚɧɢɣ ɹɜɥɹɥɢɫɶ ɫɨɨɛɳɟɫɬɜɚ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ Ɉɬ©ɝɨɥɵɯª ɚɦɟɛ ɬɟɫɬɚɰɟɢ ɨɬɥɢɱɚɸɬɫɹ ɧɚɥɢɱɢɟɦ ɠɟɫɬɤɨɣ ɪɚɤɨɜɢɧɤɢ± ɫɜɨɟɝɨ ɪɨɞɚ ɷɤɡɨɫɤɟɥɟ ɬɚ Ɋɚɤɨɜɢɧɤɚ ɩɨɫɬɪɨɟɧɧɚɹ ɢɡ ɷɤɡɨɝɟɧɧɨɝɨ ɢɥɢ ɷɧɞɨɝɟɧɧɨɝɨ ɦɚɬɟɪɢɚɥɚ ɜɵɩɨɥɧɹɟɬ ɡɚɳɢɬɧɭɸ ɮɭɧɤɰɢɸ ɢɝɪɚɟɬ ɪɨɥɶ ɛɭɮɟɪɚ ɦɟɠɞɭ ɤɥɟɬɤɨɣ ɢ ɫɪɟɞɨɣ ɋɬɪɨɟɧɢɟ ɪɚɤɨ ɜɢɧɨɤ ɪɚɡɧɨɨɛɪɚɡɧɨ ɢ ɢɫɩɨɥɶɡɭɟɬɫɹ ɞɥɹ ɞɢɮɮɟɪɟɧɰɢɚɰɢɢ ɜɢɞɨɜ ɪɨɞɨɜ ɫɟɦɟɣɫɬɜ Ɇɨɪɮɨɥɨɝɢɱɟɫɤɚɹ ɢɡɦɟɧɱɢɜɨɫɬɶ ɨɛɭɫɥɨɜɥɟɧɚ ɫɩɟɰɢɮɢɤɨɣ ɠɢɡɧɟɧɧɨɣ ɫɪɟɞɵ ɞɥɹ ɨɩɪɟɞɟɥɟɧɧɵɯ ɷɤɨɦɨɪɮ ɮɨɪɦɵ ɪɚɤɨɜɢɧɤɢ ɯɚɪɚɤɬɟɪɚ ɤɪɨɸɳɢɯ ɷɥɟɦɟɧɬɨɜ ɪɚɫɩɨ ɥɨɠɟɧɢɟ ɫɬɪɨɟɧɢɟ ɧɚɪɭɠɧɨɝɨ ɨɬɜɟɪɫɬɢɹ ɢ ɞɪɭɝɢɟ ɷɥɟɦɟɧɬɵ ɦɨɪɮɨɥɨɝɢɢ Ʉɨɦ ɩɥɟɤɫɧɵɟ ɯɚɪɚɤɬɟɪɢɫɬɢɤɢ ɢ ɮɭɧɤɰɢɨɧɚɥɶɧɵɟ ɨɫɨɛɟɧɧɨɫɬɢ ɪɚɤɨɜɢɧɨɤ ɜɵɞɟɥɹɸɬ ɬɟɫɬɚɰɟɣ ɢɡ ɩɪɨɱɢɯ ɝɪɭɩɩ ɩɪɨɫɬɟɣɲɢɯ ɤɚɤ ɭɞɨɛɧɵɣ ɨɛɴɟɤɬ ɷɤɨɥɨɝɢɱɟɫɤɢɯ ɢɫɫɥɟ ɞɨɜɚɧɢɣ ɉɪɢ ɯɪɨɧɢɱɟɫɤɨɦ ɜɥɢɹɧɢɢ ɧɟɮɬɢ ɦɨɠɧɨ ɨɠɢɞɚɬɶ ɢɡɦɟɧɟɧɢɣ ɜ ɫɨɨɛɳɟ ɫɬɜɟ ɪɚɤɨɜɢɧɧɵɯ ɚɦɺɛ ɫɜɹɡɚɧɧɵɯ ɫ ɞɢɮɮɟɪɟɧɰɢɪɨɜɚɧɧɨɣ ɭɫɬɨɣɱɢɜɨɫɬɶɸ ɪɚɡɥɢɱ ɧɵɯ ɜɢɞɨɜ ɐɟɥɶɸ ɩɪɟɞɥɚɝɚɟɦɨɝɨ ɢɫɫɥɟɞɨɜɚɧɢɹ ɹɜɥɹɥɨɫɶ ɢɡɭɱɟɧɢɟ ɚɞɚɩɬɚɰɢɨɧɧɵɯ ɩɪɰɟɫɫɨɜ ɜ ɫɨɨɛɳɟɫɬɜɚɯ ɪɚɤɨɜɢɧɧɵɯ ɚɦɺɛ ɨɬɧɨɫɹɳɢɯɫɹ ɤ ɪɨ

ɞɚɦ Arcella, Centropyxis, Plagiopyxis, Heleopera, Nebela, Euglypha, Trinema,

Cyclopyxis, Assulina, Corytion, Trigonopyxis, Placocista, Amphitrema, Phryganella

ɩɪɢ ɯɪɨɧɢɱɟɫɤɨɦ ɞɟɣɫɬɜɢɢ ɧɟɮɬɟɡɚɝɪɹɡɧɟɧɢɣ Ʉɚɪɬɚɲɟɜ ɋɦɨɥɢɧɚ ȼ ɥɚɛɨɪɚɬɨɪɧɵɯ ɭɫɥɨɜɢɹɯ ɢɫɫɥɟɞɨɜɚɥɚɫɶ ɭɫɬɨɣɱɢɜɨɫɬɶ ɫɨɨɛɳɟɫɬɜ ɪɚɤɨɜɢɧ

ɧɵɯ ɚɦɺɛ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɤɨɧɰɟɧɬɪɚɰɢɢ ɧɟɮɬɢ ȼ ɨɩɵɬɚɯ ɢɫɩɨɥɶɡɨɜɚɥɚɫɶ ɬɨɜɚɪ

ϭϴϯ

ɧɚɹ ɧɟɮɬɶ Ʌɭɝɢɧɟɰɤɨɝɨ ɦɟɫɬɨɪɨɠɞɟɧɢɹ Ⱥɧɚɥɢɡɢɪɭɟɦɵɟ ɩɪɨɛɵ ɩɪɟɞɫɬɚɜɥɹɥɢ ɫɨ ɛɨɣ ɨɛɪɚɡɰɵ ɩɨɱɜɵ, ɜ ɤɨɬɨɪɵɯ ɨɩɪɟɞɟɥɹɥɨɫɶ ɨɬɧɨɫɢɬɟɥɶɧɚɹ ɱɢɫɥɟɧɧɨɫɬɶ ɢ ɜɢɞɨ ɜɨɣ ɫɩɟɤɬɪ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɩɪɨɜɨɞɢɥɢɫɶ ɢɡɦɟɪɟɧɢɹ ɩɨɱɜɟɧɧɨɣ ɜɥɚɠɧɨɫɬɢ Ʉɨ ɥɢɱɟɫɬɜɟɧɧɵɣ ɭɱɟɬ ɬɟɫɬɚɰɢɣ ɩɪɨɜɨɞɢɥɫɹ ɩɪɹɦɵɦ ɦɢɤɪɨɫɤɨɩɢɪɨɜɚɧɢɟɦ ɜɨɞɧɨɣ ɩɨɱɜɟɧɧɨɣ ɫɭɫɩɟɧɡɢɢ ɜ ɱɚɲɤɚɯ ɉɟɬɪɢ ɜ ɨɩɪɟɞɟɥɟɧɧɨɦ ɤɨɥɢɱɟɫɬɜɟ ɩɨɥɟɣ ɡɪɟɧɢɹ Ƚɟɥɶɰɟɪ ɢ ɞɪ 1985). ȼ ɫɩɨɪɧɵɯ ɫɥɭɱɚɹɯ ɪɚɤɨɜɢɧɤɢ ɩɪɢ ɩɨɦɨɳɢ ɩɢɩɟɬɤɢ ɨɬɫɚɠɢ ɜɚɥɢ ɧɚ ɩɪɟɞɦɟɬɧɨɟ ɫɬɟɤɥɨ ɩɨɦɟɳɚɥɢ ɜ ɤɚɩɥɸ ɝɥɢɰɟɪɢɧɚ ɢ ɢɫɫɥɟɞɨɜɚɥɢ ɩɨɞ ɦɢɤ ɪɨɫɤɨɩɨɦ ɉɪɢ ɤɨɥɢɱɟɫɬɜɟɧɧɨɦ ɩɨɞɫɱɟɬɟ ɭɱɢɬɵɜɚɥɢɫɶ ɜɫɟ ɩɨɩɚɞɚɸɳɢɟɫɹ ɪɚɤɨ ɜɢɧɤɢ ɱɢɫɥɨ ɤɨɬɨɪɵɯ ɩɟɪɟɫɱɢɬɵɜɚɥɢ ɧɚ ɝ ɚɛɫɨɥɸɬɧɨ ɫɭɯɨɣ ɩɨɱɜɵ ȼɥɚɠɧɨɫɬɶ ɨɩɪɟɞɟɥɹɥɢ ɜɟɫɨɜɵɦ ɦɟɬɨɞɨɦ Ɉɩɪɟɞɟɥɟɧɢɟ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɩɪɨɜɨɞɢɥɨɫɶ ɩɨ ɫɬɚɧɞɚɪɬɧɵɦ ɦɟɬɨɞɢɤɚɦ Ƚɟɥɶɰɟɪ ɢ ɞɪ ȼ ɪɚɛɨɬɟ ɩɪɢɧɹɬɚ ɫɢɫɬɟɦɚ ɷɭɤɚɪɢ ɨɬ ɩɪɟɞɥɨɠɟɧɧɚɹ ɦɟɠɞɭɧɚɪɨɞɧɵɦ ɤɨɦɢɬɟɬɨɦ Ɉɫɬɚɬɨɱɧɭɸ ɤɨɧɰɟɧɬɪɚɰɢɸ ɧɟɮɬɢ ɨɩɪɟɞɟɥɹɥɢ ɜɟɫɨɜɵɦ ɦɟɬɨɞɨɦ.

Ⱦɥɹ ɨɰɟɧɤɢ ɜɥɢɹɧɢɹ ɧɟɮɬɢ ɧɚ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɜ ɥɚɛɨɪɚɬɨɪɧɵɯ ɭɫɥɨɜɢɹɯ ɢɫɩɨɥɶɡɨɜɚɥɢ ɤɸɜɟɬɵ ɦɚɬɟɪɢɚɥ± ɩɥɚɫɬɦɚɫɫɚ ɪɚɡɦɟɪɵ ɞɥɢɧɚ± ɦ ɜɵɫɨɬɚ±ɦ ɲɢɪɢɧɚ± ɦ ȼ ɥɚɛɨɪɚɬɨɪɧɵɟ ɤɸɜɟɬɵ ɩɨɦɟɳɚɥɚɫɶ ɫɦɟɲɚɧɧɚɹ ɩɪɨɛɚ ɝɭɦɭɫɨɜɨɝɨ ɫɥɨɹ Ⱥ1, 0- ɫɦ ɫɟɪɵɯ ɥɟɫɧɵɯ ɩɨɱɜ ɡɟɪɧɢɫɬɨ-ɤɨɦɤɨɜɚɬɨɣ ɫɬɪɭɤɬɭɪɵ ɜɥɚɠɧɨɫɬɶɸ ɩɨ ɬɪɢ ɤɢɥɨɝɪɚɦɦɚ ɩɨɱɜɵ ɜ ɤɚɠɞɨɣ ɤɸɜɟɬɟ Ⱦɚɧɧɵɣ ɬɢɩ ɩɨɱɜ ɲɢɪɨɤɨ ɪɚɫɩɪɨɫɬɪɚɧɟɧ ɜ Ɂɚɩɚɞɧɨɣ ɋɢɛɢɪɢ ȼ ɤɚɠɞɨɦ ɨɩɵɬɟ ɢɫɩɨɥɶɡɨɜɚɥɨɫɶ ɩɨ ɤɸɜɟɬ Ɉɩɵɬɵ ɩɪɨɜɨɞɢɥɢɫɶ ɩɪɢ ɤɨɦɧɚɬɧɨɣ ɬɟɦɩɟɪɚɬɭɪɟ ɗɤɫɩɟɪɢɦɟɧɬ ɫɨɫɬɨɹɥ ɢɡ ɩɹɬɢ ɨɩɵɬɨɜ ɫ ɨɞɧɨɪɚɡɨɜɵɦɢ ɞɨɡɚɦɢ ɜɧɟɫɟɧɢɹ ɧɟɮɬɢ ɝ ɤɝ ɩɨɱɜɵ ɩɪɢ ɩɚɪɚɥɥɟɥɶɧɨɦ ɤɨɧɬɪɨɥɟ ɇɚɛɥɸɞɟɧɢɹ ɩɪɨɜɨɞɢɥɢɫɶ ɜ ɬɟɱɟɧɢɟ ɬɪɢɞɰɚɬɢ ɫɭɬɨɤ ɉɪɨɛɵ ɩɨɱɜ ɞɥɹ ɚɧɚɥɢɡɚ ɛɪɚɥɢ ɜ ɩɨɜɟɪɯɧɨɫɬɧɨɦ ɝɨɪɢɡɨɧɬɟ ɧɚ ɝɥɭɛɢɧɟ - ɫɦ ȼ ɥɚɛɨɪɚɬɨɪɧɵɯ ɢ ɩɨɥɟɜɵɯ ɢɫɫɥɟɞɨɜɚɧɢɹɯ ɢɫɩɨɥɶɡɨɜɚɥɚɫɶ ɬɨɜɚɪɧɚɹ ɧɟɮɬɶ Ʌɭɝɢɧɟɰɤɨɝɨ ɦɟɫɬɨɪɨɠɞɟɧɢɹ Ɇɟɫɬɨɪɨɠɞɟɧɢɟ Ʌɭɝɢɧɟɰɤɨɟ ɨɬɧɨɫɢɬɫɹ ɤ ɩɚɥɟɨɡɨɣɫɤɨɦɭ ɬɢɩɭ ɧɟɮɬɟɣ ɢ ɡɚɥɟɝɚɟɬ ɜ ɩɪɟɞɟɥɚɯ ɉɭɞɢɧɫɤɨɝɨ ɦɟɝɚɜɚɥɚ ɯɚɪɚɤɬɟɪɢɡɭɸɳɟɝɨɫɹ ɜɵɫɨɤɨɣ ɫɬɟɩɟɧɶɸ ɢɡɜɥɟɱɟɧɢɹ ɜɫɟɯ ɬɢɩɨɜ ɧɢɡɤɨɦɨɥɟɤɭɥɹɪɧɵɯ ɫɢɥɶɧɵɯ ɨɫɧɨɜɚɧɢɣ.

Ɍɚɛɥɢɰɚ Ɉɫɧɨɜɧɵɟ ɮɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɫɜɨɣɫɬɜɚ ɧɟɮɬɢ

ɉɨ ɫɨɞɟɪɠɚɧɢɸ ɫɟɪɵ ɧɟɮɬɶ ɨɬɧɨɫɢɬɫɹ ɤ ɦɚɥɨɫɟɪɧɢɫɬɨɣ, ɫ ɫɨɞɟɪɠɚɧɢɟɦ ɞɨ 0,5 % ɉɨ ɩɥɨɬɧɨɫɬɢ ɢɫɩɨɥɶɡɭɟɦɚɹ ɧɟɮɬɶ ɨɬɧɨɫɢɬɫɹ ɤ ɤɥɚɫɫɭ ɥɟɝɤɢɯ ɧɟɮɬɟɣ±ɝ ɫɦ3), ɩɪɢ ɧɚɢɛɨɥɟɟ ɪɚɫɩɪɨɫɬɪɚɧɟɧɧɵɯ ɜɟɥɢɱɢɧɚɯ± ɝ ɫɦ3. ɇɢɡɤɚɹ ɩɥɨɬɧɨɫɬɶ ɧɟɮɬɢ ɨɛɭɫɥɨɜɥɟɧɚ ɩɪɟɨɛɥɚɞɚɧɢɟɦ ɦɟɬɚɧɨɜɵɯ ɭɝɥɟɜɨɞɨɪɨɞɨɜ ɧɢɡɤɢɦ ɫɨɞɟɪɠɚɧɢɟɦ ɫɦɨɥɢɫɬɨ-ɚɫɮɚɥɶɬɟɧɨɜɵɯ ɤɨɦɩɨɧɟɧɬɨɜ ɜɨ ɮɪɚɤɰɢɨɧɧɨɦ ɨɬɧɨɲɟɧɢɢ

± ɜɵɫɨɤɢɦ ɫɨɞɟɪɠɚɧɢɟɦ ɛɟɧɡɢɧɨɜɵɯ ɢ ɤɟɪɨɫɢɧɨɜɵɯ ɮɪɚɤɰɢɣ Ⱥɤɫɟɧɨɜ ɢ ɞɪɇɟɮɬɶ ɢ ɧɟɮɬɟɩɪɨɞɭɤɬɵ ɨɬɧɨɫɹɬɫɹ ɤ ɬɪɟɬɶɟɦɭ ɤɥɚɫɫɭ ɨɩɚɫɧɨɫɬɢ Ɍɨɤɫɢɱ

ϭϴϰ

ɧɨɫɬɶ ɧɟɮɬɢ ɨɛɴɹɫɧɹɟɬɫɹ ɩɪɢɫɭɬɫɬɜɢɟɦ ɥɟɬɭɱɢɯ ɨɪɝɚɧɢɱɟɫɤɢɯ ɭɝɥɟɜɨɞɨɪɨɞɨɜ ɬɨ ɥɭɨɥ ɤɫɢɥɨɥ ɛɟɧɡɨɥ ɧɚɮɬɚɥɢɧ ɢ ɪɹɞ ɞɪɭɝɢɯ ɮɪɚɤɰɢɣ ɧɟɮɬɢ ȼ ɫɨɫɬɚɜɟ ɧɟɮɬɢ ɫɨ ɞɟɪɠɚɬɫɹ ɦɟɬɚɧ ɢ ɩɪɨɩɚɧ

Ɋɟɡɭɥɶɬɚɬɵ ɥɚɛɨɪɚɬɨɪɧɵɯ ɢɫɫɥɟɞɨɜɚɧɢɣ ɩɨ ɜɥɢɹɧɢɸ ɧɟɮɬɟɡɚɝɪɹɡɧɟɧɢɣ ɧɚ ɫɨɨɛɳɟɫɬɜɚ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɩɪɟɞɫɬɚɜɥɟɧɧɵ ɜ ɬɚɛɥɢɰɟ ɢ ɩɨɡɜɨɥɹɸɬ ɡɚɦɟɬɢɬɶ ɢɡɦɟɧɟɧɢɹ ɱɢɫɥɟɧɧɨɫɬɢ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɩɪɢ ɜɧɟɫɟɧɢɢ ɬɨɜɚɪɧɨɣ ɧɟɮɬɢɝ ɤɝ ɩɨɱɜɵ ɇɚ ɱɢɫɥɟɧɧɨɫɬɶ ɬɟɫɬɚɰɟɣ ɨɤɚɡɵɜɚɸɬ ɜɥɢɹɧɢɟ ɧɟɜɵɫɨɤɢɟ ɤɨɧɰɟɧ ɬɪɚɰɢɢ ɝ ɤɝ ɜɧɟɫɟɧɢɹ ɩɨɥɥɸɬɚɧɬɚ

Ɍɚɛɥɢɰɚ ɂɡɦɟɧɟɧɢɟ ɱɢɫɥɟɧɧɨɫɬɢ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɬɵɫɹɱɢ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɤɨɧ ɰɟɧɬɪɚɰɢɢ ɧɟɮɬɢ ɜ ɩɨɱɜɟ

Ⱥɧɚɥɢɡ ɩɪɟɞɫɬɚɜɥɟɧɧɵɯ ɞɚɧɧɵɯ ɩɨɡɜɨɥɹɟɬ ɡɚɦɟɬɢɬɶ ɱɬɨ ɧɚɢɛɨɥɶɲɟɟ ɜɥɢɹɧɢɟ ɧɚ ɱɢɫɥɟɧɧɨɫɬɶ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɨɤɚɡɵɜɚɟɬ ɧɟɮɬɟɡɚɝɪɹɡɧɟɧɢɹ ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣɝ ɤɝ ɉɪɢ ɧɟɮɬɟɡɚɝɪɹɡɧɟɧɢɹɯ ɩɨɱɜɵ ɧɟɮɬɶɸ ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ ɝ ɤɝ ɩɪɨɢɫɯɨ ɞɢɬ ɩɨɫɬɟɩɟɧɧɨɟ ɫɧɢɠɟɧɢɟ ɱɢɫɥɟɧɧɨɫɬɢ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɞɨ ɬɵɫ ɷɤɡ ɝ ɧɚ ɫɭɬɤɢ ɱɬɨ ɡɧɚɱɢɬɟɥɶɧɨ ɦɟɧɶɲɟ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɧɟɡɚɝɪɹɡɧɟɧɧɨɣ ɩɨɱɜɨɣ ɬɵɫ ɷɤɡ ɝ ɉɪɢ ɤɨɧɰɟɧɬɪɚɰɢɢ ɧɟɮɬɟɡɚɝɪɹɡɧɟɧɢɣ ɢ ɝ ɤɝ ɧɚɛɥɸɞɚɟɬɫɹ ɫɧɢɠɟɧɢɟ ɱɢɫɥɟɧɧɨɫɬɢ ɚɦɺɛ ɧɚ ɫɭɬɤɢ ɧɚ - ɬɵɫ ɷɤɡ ɝ ɢ - ɬɵɫ ɷɤɡ ɝ ɧɚ ɫɭɬɤɢ ɋɧɢɠɟɧɢɟ ɱɢɫɥɟɧɧɨɫɬɢ ɨɛɭɫɥɨɜɥɟɧɨ ɧɟɝɚɬɢɜɧɵɦ ɜɥɢɹɧɢɟɦ ɧɟɮɬɢ ɧɚ ɫɨɨɛɳɟɫɬɜɚ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ȼ ɤɨɧɬɪɨɥɶɧɨɣ ɤɸɜɟɬɟ ɜ ɫɭɬɤɢ ɧɚɛɥɸɞɚɟɬɫɹ ɭɜɟɥɢɱɟɧɢɟ ɱɢɫ ɥɟɧɧɨɫɬɢ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɞɨ ɬɵɫ ɷɤɡ ɝ ɫ ɩɨɫɥɟɞɭɸɳɢɦ ɫɧɢɠɟɧɢɟ ɱɢɫɥɟɧ ɧɨɫɬɢ ɞɨ ɬɵɫ ɷɤɡ ɝ ɍɦɟɧɶɲɟɧɢɟ ɱɢɫɥɟɧɧɨɫɬɢ ɪɚɤɨɜɢɧɧɵɯ ɚɦɟɛ ɜ ɥɚɛɨɪɚ ɬɨɪɧɵɯ ɤɸɜɟɬɚɯ ɩɨɫɥɟ ɜɧɟɫɟɧɢɹ ɧɟɮɬɢ ɨɛɭɫɥɨɜɥɟɧɨ ɢ ɢɡɦɟɧɟɧɢɟɦ ɜɥɚɠɧɨɫɬɢ ɍɫɬɚɧɨɜɥɟɧɨ ɱɬɨ ɜɥɚɠɧɨɫɬɶ ɡɚɝɪɹɡɧɟɧɧɵɯ ɫɥɨɟɜ ɡɧɚɱɢɬɟɥɶɧɨ ɧɢɠɟ ɱɟɦ ɜ©ɱɢɫ ɬɨɣªɩɨɱɜɟ Ɂɚɝɪɹɡɧɟɧɧɚɹ ɧɟɮɬɟɩɪɨɞɭɤɬɚɦɢ ɫɜɟɬɥɨ-ɫɟɪɚɹ ɥɟɫɧɚɹ ɩɨɱɜɚ ɧɟ ɫɩɨɫɨɛ ɧɚ ɩɪɢɧɢɦɚɬɶ ɜɨɞɭ ɢ ɩɪɨɜɨɞɢɬɶ ɟɟ ɜɧɢɡ ɩɨ ɩɨɱɜɟɧɧɨɦɭ ɩɪɨɮɢɥɸ ȼ ɪɚɛɨɬɚɯ ɫ ɜɚɪɶɢɪɨɜɚɧɢɟɦ ɭɫɥɨɜɢɣ ɭɜɥɚɠɧɟɧɢɹ ɨɬ ɜɨɡɞɭɲɧɨ-ɫɭɯɨɝɨ ɫɨɫɬɨɹɧɢɹ ɞɨ ɩɨɥɧɨɝɨ ɨɛɜɨɞɧɟɧɢɹ ɩɨɤɚɡɚɧɨ ɭɜɟɥɢɱɟɧɢɟ ɞɨɥɢ ɠɢɜɵɯ ɨɪɝɚɧɢɡɦɨɜ ɫ ɪɨɫɬɨɦ ɝɢɞɪɨɦɨɪ ɮɢɡɦɚ ɩɨɱɜɵ Ɋɚɤɨɜɢɧɧɵɟ ɚɦɟɛɵ ɷɤɫɰɢɫɬɢɪɭɸɬɫɹ ɢ ɩɟɪɟɯɨɞɹɬ ɜ ɚɤɬɢɜɧɨɟ ɫɨ ɫɬɨɹɧɢɟ ɩɪɢ ɛɨɥɟɟ ɜɵɫɨɤɨɣ ɜɥɚɠɧɨɫɬɢ, ɱɟɦ ɞɪɭɝɢɟ ɩɪɟɞɫɬɚɜɢɬɟɥɢ ɠɢɜɨɬɧɨɝɨ ɧɚ ɫɟɥɟɧɢɹ ɩɨɱɜ ɧɚɩɪɢɦɟɪ ɧɟɦɚɬɨɞɵ ɢ ɤɨɥɨɜɪɚɬɤɢ ɇɚɢɛɨɥɟɟ ɨɩɬɢɦɚɥɶɧɵɦɢ ɞɥɹ

ϭϴϱ