Гиперболический рост разнообразия морской и континентальной биот фанерозоя и эволюция сообществ

ÜìêçÄã éÅôÖâ ÅàéãéÉàà, 2008, ÚÓÏ 69, ‹ 3, Ò. 175–194

175

å‡ÚÂÏ‡ÚË˜ÂÒÍÓÂ ÏÓ‰ÂÎËÓ‚‡ÌËÂ „ÎÓ·‡Î¸Ì˚ıËÁÏÂÌÂÌËÈ ·ËÓ‡ÁÌÓÓ·‡ÁËfl, Ì‡·Î˛‰‡ÂÏ˚ı ‚ Ô‡ÎÂ-ÓÌÚÓÎÓ„Ë˜ÂÒÍÓÈ ÎÂÚÓÔËÒË, ÔÓÏÓ„‡ÂÚ ÔÓÌflÚ¸ ÍÛÔ-ÌÓÏ‡Ò¯Ú‡·Ì˚Â Á‡ÍÓÌÓÏÂÌÓÒÚË Ë ÏÂı‡ÌËÁÏ˚ ˝‚Ó-Î˛ˆËË ·ËÓÒÙÂ˚. ÑËÌ‡ÏËÍ‡ ·ËÓ‡ÁÌÓÓ·‡ÁËflÏËÌÛ‚¯Ëı „ÂÓÎÓ„Ë˜ÂÒÍËı ˝ÔÓı Ú‡‰ËˆËÓÌÌÓ ÓÔË-Ò˚‚‡ÂÚÒfl ÔË ÔÓÏÓ˘Ë ÔÓÒÚ˚ı ˝ÍÒÔÓÌÂÌˆË‡Î¸Ì˚ıË ÎÓ„ËÒÚË˜ÂÒÍËı ÏÓ‰ÂÎÂÈ, Á‡ËÏÒÚ‚Ó‚‡ÌÌ˚ı ËÁ ÚÂÓ-ËË ‰ËÌ‡ÏËÍË ÔÓÔÛÎflˆËÈ (Sepkoski, 1991a; Sepkos-ki, Kendrick, 1993; Benton, 1999). ùÍÒÔÓÌÂÌˆË‡Î¸Ì‡flÏÓ‰ÂÎ¸ (

dN

/

dt

=

kN

) ÔÂ‰ÔÓÎ‡„‡ÂÚ ÔÓÒÚÛ˛ ÔÓÎÓÊË-ÚÂÎ¸ÌÛ˛ Ó·‡ÚÌÛ˛ Ò‚flÁ¸ ÏÂÊ‰Û ‡ÁÌÓÓ·‡ÁËÂÏ (

N

,˜ËÒÎÓ Ú‡ÍÒÓÌÓ‚ – ‚Ë‰Ó‚, Ó‰Ó‚ ËÎË ÒÂÏÂÈÒÚ‚) Ë ÒÍÓ-ÓÒÚ¸˛ Â„Ó ÓÒÚ‡ (ÔÓ ÔËÌˆËÔÛ “·ÓÎ¸¯Â Ó‰ËÚÂ-ÎÂÈ – ·ÓÎ¸¯Â ÔÓÚÓÏÍÓ‚”); ‚Á‡ËÏÓ‰ÂÈÒÚ‚ËÂ ÏÂÊ‰ÛÚ‡ÍÒÓÌ‡ÏË ‚ ˝ÚÓÏ ÒÎÛ˜‡Â ÌÂ Û˜ËÚ˚‚‡ÂÚÒfl. ãÓ„ËÒÚË-˜ÂÒÍ‡fl ÏÓ‰ÂÎ¸ (

dN

/

dt

=

k

(

N

max

–

N

)

N

) ‰Ó·‡‚ÎflÂÚ Í˝ÚÓÏÛ ÓÚËˆ‡ÚÂÎ¸ÌÛ˛ Ó·‡ÚÌÛ˛ Ò‚flÁ¸, ÓÒÌÓ‚‡Ì-ÌÛ˛ Ì‡ Ó„‡ÌË˜ÂÌÌÓÒÚË ‰ÓÒÚÛÔÌÓ„Ó ˝ÍÓÎÓ„Ë˜ÂÒÍÓ-„Ó ÔÓÒÚ‡ÌÒÚ‚‡; ‚Á‡ËÏÓ‰ÂÈÒÚ‚ËÂ ÏÂÊ‰Û Ú‡ÍÒÓÌ‡-ÏË Ò‚Ó‰ËÚÒfl Í ÍÓÌÍÛÂÌˆËË Á‡ ÌË¯Ë. äÓÂÎflˆËfl˝ÚËı ÏÓ‰ÂÎÂÈ Ò Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍËÏË ‰‡ÌÌ˚ÏË ÌÂ-‚˚ÒÓÍ‡.

àÒÔÓÎ¸ÁÓ‚‡ÌËÂ “ÔÓÔÛÎflˆËÓÌÌ˚ı” ÏÓ‰ÂÎÂÈ ‰ÎflËÌÚÂÔÂÚ‡ˆËË ‰ËÌ‡ÏËÍË ·ËÓ‡ÁÌÓÓ·‡ÁËfl ‚Â‰ÂÚ ÍË„ÌÓËÓ‚‡ÌË˛ ·ÓÎÂÂ ÒÎÓÊÌ˚ı ‚ÌÛÚËÒËÒÚÂÏÌ˚ıÒ‚flÁÂÈ, ‚ ÚÓÏ ˜ËÒÎÂ Ò‚flÁÂÈ ÍÓÓÔÂ‡ÚË‚ÌÓ„Ó ı‡‡Í-ÚÂ‡, ·Î‡„Ó‰‡fl ÍÓÚÓ˚Ï ·ËÓÒÙÂ‡ ‡Á‚Ë‚‡ÂÚÒflÍ‡Í ˆÂÎÓÒÚÌ˚È Ó·˙ÂÍÚ, ‡ ÓÒÚ ·ËÓ‡ÁÌÓÓ·‡ÁËflÔËÓ·ÂÚ‡ÂÚ ˜ÂÚ˚ Ò‡ÏÓÛÒÍÓfl˛˘Â„ÓÒfl, ˝ÒÍ‡Î‡ˆË-ÓÌÌÓ„Ó ÔÓˆÂÒÒ‡ (å‡ÍÓ‚, 2001). çÂ‰‡‚ÌÓ Ì‡ÏË ·˚-ÎÓ ÔÓÍ‡Á‡ÌÓ, ˜ÚÓ ‰ËÌ‡ÏËÍ‡ ‡ÁÌÓÓ·‡ÁËfl Ù‡ÌÂÓ-ÁÓÈÒÍÓÈ ÏÓÒÍÓÈ ·ËÓÚ˚ (îåÅ) Ì‡ Ó‰Ó‚ÓÏ ÛÓ‚ÌÂÁÌ‡˜ËÚÂÎ¸ÌÓ ÎÛ˜¯Â ÒÓÓÚ‚ÂÚÒÚ‚ÛÂÚ ÏÓ‰ÂÎË „ËÔÂ·Ó-ÎË˜ÂÒÍÓ„Ó ÓÒÚ‡ (

dN

/

dt

=

kN

2

), ÍÓÚÓ‡fl ÔÂ‰ÔÓÎ‡„‡ÂÚÒÛ˘ÂÒÚ‚Ó‚‡ÌËÂ ÍÓÓÔÂ‡ÚË‚Ì˚ı ‚Á‡ËÏÓ‰ÂÈÒÚ‚ËÈÏÂÊ‰Û Ú‡ÍÒÓÌ‡ÏË (å‡ÍÓ‚, äÓÓÚ‡Â‚, 2007; Mar-kov, Korotayev, 2007). ùÚ‡ ÏÓ‰ÂÎ¸ ̄ ËÓÍÓ ÔËÏÂÌfl-ÂÚÒfl ‚ ‰ÂÏÓ„‡ÙËË Ë Ï‡ÍÓÒÓˆËÓÎÓ„ËË ‰Îfl ÓÔËÒ‡-ÌËfl ‰ÓÎ„ÓÒÓ˜ÌÓÈ ‰ËÌ‡ÏËÍË Ì‡Ó‰ÓÌ‡ÒÂÎÂÌËfl(Foerster et al., 1960; ä‡ÔËˆ‡, 1992, 1999; Kremer,1993; Cohen, 1995; Johansen, Sornette, 2001; äÓÓÚ‡-Â‚ Ë ‰., 2007‡,·). ÉËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ı‡‡ÍÚÂ-ÂÌ ‰Îfl ÒÎÓÊÌ˚ı ÌÂÎËÌÂÈÌ˚ı Ò‡ÏÓÓ„‡ÌËÁÛ˛˘ËıÒflÒËÒÚÂÏ, Ò‚flÁ‡ÌÌ˚ı ‚ÌÛÚÂÌÌËÏË ‚Á‡ËÏÓ‰ÂÈÒÚ‚ËflÏË.ÉËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ·ËÓ‡ÁÌÓÓ·‡ÁËfl ÔÓÁ‚ÓÎflÂÚ

ÉàèÖêÅéãàóÖëäàâ êéëí êÄáçééÅêÄáàü åéêëäéâ

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èÓÒÚÛÔËÎ‡ ‚ Â‰‡ÍˆË˛ 13.06.2007 „.

ù‚ÓÎ˛ˆËÓÌÌ‡fl ‰ËÌ‡ÏËÍ‡ ·ËÓ‡ÁÌÓÓ·‡ÁËfl Ú‡‰ËˆËÓÌÌÓ ÓÔËÒ˚‚‡ÂÚÒfl ÔË ÔÓÏÓ˘Ë ÔÓÒÚ˚ı ̋ ÍÒÔÓÌÂÌ-ˆË‡Î¸Ì˚ı Ë ÎÓ„ËÒÚË˜ÂÒÍËı ÏÓ‰ÂÎÂÈ, Á‡ËÏÒÚ‚Ó‚‡ÌÌ˚ı ËÁ ÚÂÓËË ‰ËÌ‡ÏËÍË ÔÓÔÛÎflˆËÈ. çÂ‰‡‚ÌÓ Ì‡ÏË·˚ÎÓ ÔÓÍ‡Á‡ÌÓ, ˜ÚÓ ÓÒÚ Ó‰Ó‚Ó„Ó ‡ÁÌÓÓ·‡ÁËfl Ù‡ÌÂÓÁÓÈÒÍÓÈ ÏÓÒÍÓÈ ·ËÓÚ˚ ÎÛ˜¯Â ÓÔËÒ˚‚‡ÂÚÒfl„ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎ¸˛, ¯ËÓÍÓ ÔËÏÂÌflÂÏÓÈ ‚ Ï‡ÍÓÒÓˆËÓÎÓ„ËË Ë ‰ÂÏÓ„‡ÙËË. Ç Ì‡ÒÚÓfl˘ÂÈ ‡-·ÓÚÂ ÔÓÍ‡Á‡Ì‡ ÔËÎÓÊËÏÓÒÚ¸ ÏÓ‰ÂÎË „ËÔÂ·ÓÎË˜ÂÒÍÓ„Ó ÓÒÚ‡ Ú‡ÍÊÂ Ë Í ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ ·ËÓÚÂ Ù‡-ÌÂÓÁÓfl Ì‡ ÛÓ‚ÌÂ ÒÂÏÂÈÒÚ‚ Ë Ó‰Ó‚, Í ÏÓÒÍÓÈ ·ËÓÚÂ Ì‡ ÛÓ‚ÌÂ ÒÂÏÂÈÒÚ‚, Ó‰Ó‚ Ë ‚Ë‰Ó‚, ‡ Ú‡ÍÊÂ ÍÓ‚ÒÂÈ Ù‡ÌÂÓÁÓÈÒÍÓÈ ·ËÓÚÂ ‚ ˆÂÎÓÏ, ÔË˜ÂÏ ‚ ÔÓÒÎÂ‰ÌÂÏ ÒÎÛ˜‡Â ÒıÓ‰ÒÚ‚Ó Ù‡ÍÚË˜ÂÒÍËı ‰‡ÌÌ˚ı Ò ÏÓ‰Â-Î¸˛ ÓÍ‡Á˚‚‡ÂÚÒfl Ì‡Ë·ÓÎÂÂ ÔÓÎÌ˚Ï. èÓ ‡Ì‡ÎÓ„ËË Ò Ï‡ÍÓÒÓˆËÓÎÓ„Ë˜ÂÒÍËÏË ÏÓ‰ÂÎflÏË, ‡ Ú‡ÍÊÂ Ì‡ ÓÒ-ÌÓ‚Â ËÏÂ˛˘ËıÒfl Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍËı ‰‡ÌÌ˚ı Ï˚ ÔÂ‰ÔÓÎ‡„‡ÂÏ, ̃ ÚÓ „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ·ËÓ‡ÁÌÓ-Ó·‡ÁËfl Ó·˙flÒÌflÂÚÒfl ÌÂÎËÌÂÈÌÓÈ ÔÓÎÓÊËÚÂÎ¸ÌÓÈ Ó·‡ÚÌÓÈ Ò‚flÁ¸˛ ‚ÚÓÓ„Ó ÔÓfl‰Í‡ ÏÂÊ‰Û ÓÒÚÓÏ˜ËÒÎ‡ Ú‡ÍÒÓÌÓ‚ Ë ËÁÏÂÌÂÌËflÏË ÒÚÛÍÚÛ˚ ÒÓÓ·˘ÂÒÚ‚. ùÚ‡ Ò‚flÁ¸ ÏÓÊÂÚ Â‡ÎËÁÓ‚˚‚‡Ú¸Òfl ‰‚ÓflÍÓ: ̃ ÂÂÁÒÌËÊÂÌËÂ ‚ÂÓflÚÌÓÒÚË ‚˚ÏË‡ÌËfl Ú‡ÍÒÓÌÓ‚ (ÓÒÚ ÒÎÓÊÌÓÒÚË Ë ÛÒÚÓÈ˜Ë‚ÓÒÚË ÒÓÓ·˘ÂÒÚ‚ ÒÔÓÒÓ·ÒÚ‚ÛÂÚÓÒÚÛ ÛÒÚÓÈ˜Ë‚ÓÒÚË ‚ıÓ‰fl˘Ëı ‚ Â„Ó ÒÓÒÚ‡‚ Ú‡ÍÒÓÌÓ‚) Ë ˜ÂÂÁ ÔÓ‚˚¯ÂÌËÂ ‚ÂÓflÚÌÓÒÚË Ëı ÔÓfl‚ÎÂÌËfl(ÔÓfl‚ÎÂÌËÂ ÌÓ‚˚ı Ú‡ÍÒÓÌÓ‚ ÒÔÓÒÓ·ÒÚ‚ÛÂÚ ÒÓÁ‰‡ÌË˛ ÌÓ‚˚ı ÌË¯, ÍÓÚÓ˚Â ÏÓ„ÛÚ ·˚Ú¸ ÓÒ‚ÓÂÌ˚ ÒÎÂ‰Û-˛˘ËÏ “ÔÓÍÓÎÂÌËÂÏ” Ú‡ÍÒÓÌÓ‚). ÉËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ‡ÁÌÓÓ·‡ÁËfl Ù‡ÌÂÓÁÓÈÒÍÓÈ ·ËÓÚ˚ Ò‚Ë‰Â-ÚÂÎ¸ÒÚ‚ÛÂÚ Ó ÚÓÏ, ˜ÚÓ ‚‡ÊÌÛ˛ ÓÎ¸ ‚ ˝‚ÓÎ˛ˆËË ÏÓ„ÛÚ Ë„‡Ú¸ ÌÂ ÚÓÎ¸ÍÓ ÍÓÌÍÛÂÌÚÌ˚Â, ÌÓ Ë ÍÓÓÔÂ‡-ÚË‚Ì˚Â ‚Á‡ËÏÓÓÚÌÓ¯ÂÌËfl ÏÂÊ‰Û ˝‚ÓÎ˛ˆËÓÌËÛ˛˘ËÏË ÍÎ‡‰‡ÏË, ·Î‡„Ó‰‡fl ÍÓÚÓ˚Ï ˝‚ÓÎ˛ˆËfl ·ËÓ-‡ÁÌÓÓ·‡ÁËfl ÔËÓ·ÂÚ‡ÂÚ ˜ÂÚ˚ Ò‡ÏÓÛÒÍÓfl˛˘Â„ÓÒfl ÔÓˆÂÒÒ‡.

ìÑä 56:57;573:7

176

ÜìêçÄã éÅôÖâ ÅàéãéÉàà ÚÓÏ 69 ‹ 3 2008

Ï‡ÍÓ‚, ÍÓÓÚ‡Â‚

ÔÂ‰ÔÓÎÓÊËÚ¸, ˜ÚÓ ˝‚ÓÎ˛ˆËfl ·ËÓÚ˚ ÔÂ‰ÒÚ‡‚ÎflÂÚÒÓ·ÓÈ ÔÓˆÂÒÒ Ò ÔÓÎÓÊËÚÂÎ¸ÌÓÈ Ó·‡ÚÌÓÈ Ò‚flÁ¸˛‚ÚÓÓ„Ó ÔÓfl‰Í‡. Ç ‡Á‚ËÚËË ˜ÂÎÓ‚Â˜ÂÒÚ‚‡ ‰Ó Ò‡-ÏÓ„Ó ÔÓÒÎÂ‰ÌÂ„Ó ‚ÂÏÂÌË ÔÓ‰Ó·Ì‡fl Ò‚flÁ¸ Â‡ÎËÁÓ-‚‡Î‡Ò¸ ·Î‡„Ó‰‡fl Ì‡Û˜ÌÓ-ÚÂıÌË˜ÂÒÍÓÏÛ Ë ÍÛÎ¸ÚÛ-ÌÓÏÛ ÔÓ„ÂÒÒÛ (‚ÍÎ˛˜‡fl ÔÓ„ÂÒÒË‚Ì˚Â ËÁÏÂÌÂÌËfl‚ ÒÚÛÍÚÛÂ ˜ÂÎÓ‚Â˜ÂÒÍËı ÒÓÓ·˘ÂÒÚ‚), ÍÓÚÓ˚È‚ÔÎÓÚ¸ ‰Ó 70-ı „Ó‰Ó‚ ÔÓ¯ÎÓ„Ó ‚ÂÍ‡ ÒÔÓÒÓ·ÒÚ‚Ó‚‡ÎÓÒÚÛ Ì‡ÒÂÎÂÌËfl Ë ‚ Ò‚Ó˛ Ó˜ÂÂ‰¸ ÒÚËÏÛÎËÓ‚‡ÎÒfl˝ÚËÏ ÓÒÚÓÏ.

Ç ‡Á‚ËÚËË ·ËÓÚ˚ ‡Ì‡ÎÓ„Ë˜Ì‡fl Ò‚flÁ¸, ÔÓ-‚Ë‰Ë-ÏÓÏÛ, Â‡ÎËÁÛÂÚÒfl ˜ÂÂÁ ÔÓ„ÂÒÒË‚Ì˚Â ËÁÏÂÌÂ-ÌËfl ‚ ÒÚÛÍÚÛÂ ÒÓÓ·˘ÂÒÚ‚. Ç Ì‡ÒÚÓfl˘ÂÈ ‡·ÓÚÂ‡ÒÒÏ‡ÚË‚‡ÂÚÒfl ÔËÎÓÊËÏÓÒÚ¸ „ËÔÂ·ÓÎË˜ÂÒÍÓÈÏÓ‰ÂÎË Í ‰ËÌ‡ÏËÍÂ ‡ÁÌÓÓ·‡ÁËfl îåÅ, Ù‡ÌÂÓ-ÁÓÈÒÍÓÈ ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ ·ËÓÚ˚ (îäÅ) Ë ‚ÒÂÈ Ù‡-ÌÂÓÁÓÈÒÍÓÈ ·ËÓÚ˚ ‚ ˆÂÎÓÏ Ì‡ ‡ÁÌ˚ı Ú‡ÍÒÓÌÓÏË-˜ÂÒÍËı ÛÓ‚Ìflı (ÓÚ ‚Ë‰Ó‚Ó„Ó ‰Ó ÒÂÏÂÈÒÚ‚ÂÌÌÓ„Ó).

åÄíÖêàÄã à åÖíéÑàäÄ

àÒÔÓÎ¸ÁÓ‚‡ÎË ‰‡ÌÌ˚Â ËÁ ÒÎÂ‰Û˛˘Ëı Ò‚Ó‰ÓÍ:1. Å‡Á‡ ‰‡ÌÌ˚ı ëÂÔÍÓÒÍË (http://strata.um-

mp.lsa.umich.edu/jack/), ÒÓ‰ÂÊ‡˘‡fl Ò‚Â‰ÂÌËfl ÓÒÚ‡ÚË„‡ÙË˜ÂÒÍÓÏ ‡ÒÔÓÒÚ‡ÌÂÌËË Ó‰Ó‚ Ù‡ÌÂ-ÓÁÓÈÒÍËı ÏÓÒÍËı ÊË‚ÓÚÌ˚ı. Ç ·‡ÁÂ ‰‡ÌÌ˚ı ËÒ-ÔÓÎ¸ÁÛÂÚÒfl ¯Í‡Î‡, ‰ÂÎfl˘‡fl Ù‡ÌÂÓÁÓÈ Ì‡ 169 ËÌ-ÚÂ‚‡ÎÓ‚, ·ÓÎ¸¯ËÌÒÚ‚Ó ËÁ ÍÓÚÓ˚ı ÒÓÓÚ‚ÂÚÒÚ‚ÛÂÚÔÓ‰˙flÛÒ‡Ï. Ç ‡Ò˜ÂÚ‡ı Û˜ËÚ˚‚‡ÎËÒ¸ Ó‰˚, ‰‡ÚË-Ó‚‡ÌÌ˚Â Ò ÚÓ˜ÌÓÒÚ¸˛ ÌÂ ÏÂÌ¸¯ÂÈ, ˜ÂÏ ‰Ó flÛÒ‡.í‡ÍËı Ó‰Ó‚ ‚ ·‡ÁÂ ‰‡ÌÌ˚ı 30065.

2. ë‚Ó‰Í‡ ëÂÔÍÓÒÍË ÔÓ ÒÚ‡ÚË„‡ÙË˜ÂÒÍÓÏÛ‡ÒÔÓÒÚ‡ÌÂÌË˛ ÒÂÏÂÈÒÚ‚ ÏÓÒÍËı ÊË‚ÓÚÌ˚ı(Sepkoski, 1992).

3. Å‡Á‡ ‰‡ÌÌ˚ı The Fossil Record 2 (Benton, 1993),ÒÓ‰ÂÊ‡˘‡fl Ò‚Â‰ÂÌËfl Ó ÒÚ‡ÚË„‡ÙË˜ÂÒÍÓÏ ‡Ò-ÔÓÒÚ‡ÌÂÌËË ÒÂÏÂÈÒÚ‚ ËÒÍÓÔ‡ÂÏ˚ı Ó„‡ÌËÁÏÓ‚(Í‡Í ÏÓÒÍËı, Ú‡Í Ë ÍÓÌÚËÌÂÌÚ‡Î¸Ì˚ı). Ç ̋ ÚÓÈ ·‡ÁÂ‰‡ÌÌ˚ı ËÒÔÓÎ¸ÁÛÂÚÒfl ¯Í‡Î‡, ‰ÂÎfl˘‡fl Ù‡ÌÂÓÁÓÈÌ‡ 86 ËÌÚÂ‚‡ÎÓ‚, ·ÓÎ¸¯ËÌÒÚ‚Ó ËÁ ÍÓÚÓ˚ı ÒÓÓÚ-‚ÂÚÒÚ‚ÛÂÚ flÛÒ‡Ï. Ç ‡Ò˜ÂÚ‡ı Û˜ËÚ˚‚‡ÎË ÒÂÏÂÈ-ÒÚ‚‡, ÔÓfl‚Ë‚¯ËÂÒfl ÌÂ ‡ÌÂÂ ÍÂÏ·Ëfl (59 ÒÂÏÂÈÒÚ‚ Ò‰ÓÍÂÏ·ËÈÒÍËÏË ÔÂ‰ÒÚ‡‚ËÚÂÎflÏË ·˚ÎË ËÒÍÎ˛˜Â-Ì˚ ‚ ÒËÎÛ ÔÓ·ÎÂÏ‡ÚË˜ÌÓÒÚË ·ÓÎ¸¯ËÌÒÚ‚‡ ̋ ÚËı ÒÂ-ÏÂÈÒÚ‚ Í‡Í ‚ Ú‡ÍÒÓÌÓÏË˜ÂÒÍÓÏ, Ú‡Í Ë ‚ ÒÚ‡ÚË„‡-ÙË˜ÂÒÍÓÏ ÓÚÌÓ¯ÂÌËË, Ë ÚÛ‰ÌÓÒÚË Ëı Ò‡‚ÌÂÌËfl ÒÙ‡ÌÂÓÁÓÈÒÍËÏË ÒÂÏÂÈÒÚ‚‡ÏË); ·˚ÎË ËÒÍÎ˛˜ÂÌ˚Ú‡ÍÊÂ 133 ÒÂÏÂÈÒÚ‚‡ Ò Ì‡ËÏÂÌÂÂ ÚÓ˜Ì˚ÏË ÒÚ‡ÚË-„‡ÙË˜ÂÒÍËÏË ‰‡ÚËÓ‚Í‡ÏË (‚ ÓÒÌÓ‚ÌÓÏ ˝ÚÓ ÒÂ-ÏÂÈÒÚ‚‡ Ì‡ÒÂÍÓÏ˚ı). é·˘ÂÂ ˜ËÒÎÓ Û˜ÚÂÌÌ˚ı ÒÂ-ÏÂÈÒÚ‚ – 6787. äÓÌÚËÌÂÌÚ‡Î¸Ì˚ÏË Ò˜ËÚ‡ÎË ÒÂÏÂÈ-ÒÚ‚‡, ÌÂ ËÏÂ˛˘ËÂ ÏÓÒÍËı ÔÂ‰ÒÚ‡‚ËÚÂÎÂÈ (‡ÚÓÎ¸ÍÓ Ì‡ÁÂÏÌ˚ı Ë ÔÂÒÌÓ‚Ó‰Ì˚ı), ÒÓ„Î‡ÒÌÓ Ò‚Â‰Â-ÌËflÏ, ‚ÌÂÒÂÌÌ˚Ï ‚ ·‡ÁÛ ‰‡ÌÌ˚ı The Fossil Record 2.ëÂÏÂÈÒÚ‚‡, ËÏÂ˛˘ËÂ ÏÓÒÍËı ÔÂ‰ÒÚ‡‚ËÚÂÎÂÈ, Ò˜Ë-Ú‡ÎËÒ¸ ÏÓÒÍËÏË. Ñ‡ÌÌ˚Â ÔÓ ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ ·ËÓ-ÚÂ ‡ÒÒÏ‡ÚË‚‡ÎË Ì‡˜ËÌ‡fl Ò ÒËÎÛ‡, Ú‡Í Í‡Í ·ÓÎÂÂ‡ÌÌËÂ ‰‡ÌÌ˚Â ÓÚ˚‚Ó˜Ì˚ Ë ÒÓÏÌËÚÂÎ¸Ì˚.

ÑËÌ‡ÏËÍ‡ ‚Ë‰Ó‚Ó„Ó ‡ÁÌÓÓ·‡ÁËfl îåÅ ·˚Î‡ÂÍÓÌÒÚÛËÓ‚‡Ì‡ Ì‡ ÓÒÌÓ‚Â ‰‡ÌÌ˚ı ÔÓ ˜ËÒÎÛ Ì‡‰-‚Ë‰Ó‚˚ı Ú‡ÍÒÓÌÓ‚ ÔË ÔÓÏÓ˘Ë ÏÂÚÓ‰ËÍË, ‡Á‡·Ó-Ú‡ÌÌÓÈ ‡ÌÂÂ Ó‰ÌËÏ ËÁ ‡‚ÚÓÓ‚ Ì‡ÒÚÓfl˘ÂÈ ÒÚ‡Ú¸Ë(å‡ÍÓ‚, 2003). óËÒÎÓ ‚Ë‰Ó‚ ‚ Í‡Ê‰ÓÏ ËÌÚÂ‚‡ÎÂ„ÂÓıÓÌÓÎÓ„Ë˜ÂÒÍÓÈ ¯Í‡Î˚ ‚˚˜ËÒÎflÎË ÔÓ ÙÓÏÛ-ÎÂ

s

=

g

×

(

g

/

f

+

f

/

o

+

o

/

c

+

c

/

p

)/4, „‰Â

g

,

f

,

o

,

c

,

p

– ̃ ËÒÎÓÓ‰Ó‚, ÒÂÏÂÈÒÚ‚, ÓÚfl‰Ó‚, ÍÎ‡ÒÒÓ‚ Ë ÚËÔÓ‚ ÒÓÓÚ‚ÂÚ-ÒÚ‚ÂÌÌÓ; ÒÔÂˆË‡Î¸Ì˚Â ÔÓÔ‡‚ÍË ‚ÌÓÒËÎË ‰Îfl ‚Â-ÏÂÌÌ˚ı ÓÍÂÒÚÌÓÒÚÂÈ Ï‡ÒÒÓ‚˚ı ‚˚ÏË‡ÌËÈ Ë ‚Â-ÎËÍÓÈ Ó‰Ó‚ËÍÒÍÓÈ ‡‰Ë‡ˆËË (ÔÓ‰Ó·ÌÂÂ ÒÏ.: å‡-ÍÓ‚, 2003).

àÒÔÓÎ¸ÁÓ‚‡ÎË ‡·ÒÓÎ˛ÚÌ˚Â ‰‡ÚËÓ‚ÍË „‡ÌËˆÒÚ‡ÚË„‡ÙË˜ÂÒÍËı ÔÓ‰‡Á‰ÂÎÂÌËÈ ÔÓ É‡‰ÒÚÂÈÌÛË ‰. (Gradstein et al., 2004). ê‡ÌÂÂ Ì‡ÏË ·˚ÎÓ ÔÓÍ‡-Á‡ÌÓ, ˜ÚÓ ËÁÏÂÌÂÌËfl ‡·ÒÓÎ˛ÚÌ˚ı ‰‡ÚËÓ‚ÓÍ, ÔÓ-ËÁÓ¯Â‰¯ËÂ Á‡ ÔÓÒÎÂ‰ÌËÂ 20 ÎÂÚ (‰Îfl Ò‡‚ÌÂÌËfl ËÒ-ÔÓÎ¸ÁÓ‚‡ÎË ¯Í‡ÎÛ ËÁ ï‡Î‡Ì‰‡ Ë ‰. (Harland et al.,1982)), Ô‡ÍÚË˜ÂÒÍË ÌÂ ‚ÎËfl˛Ú ÌË Ì‡ Ó·˘ËÈ ‚Ë‰ÍË‚˚ı ‡ÁÌÓÓ·‡ÁËfl, ÌË Ì‡ ÒÚÂÔÂÌ¸ Ëı ÒÓÓÚ‚ÂÚ-ÒÚ‚Ëfl Ï‡ÚÂÏ‡ÚË˜ÂÒÍËÏ ÏÓ‰ÂÎflÏ (å‡ÍÓ‚, äÓÓÚ‡-Â‚, 2007).

êÖáìãúíÄíõ

1. ÑËÌ‡ÏËÍ‡ ‡ÁÌÓÓ·‡ÁËfl îåÅ.

ç‡ ÛÓ‚ÌÂ ÒÂ-ÏÂÈÒÚ‚, Ó‰Ó‚ Ë ‚Ë‰Ó‚ ‰ËÌ‡ÏËÍ‡ ‡ÁÌÓÓ·‡ÁËflîåÅ ÍÓÂÎËÛÂÚ Ò „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎ¸˛ÎÛ˜¯Â, ˜ÂÏ Ò ˝ÍÒÔÓÌÂÌˆË‡Î¸ÌÓÈ (ËÒ. 1). ê‡ÌÂÂ ·˚-ÎÓ ÔÓÍ‡Á‡ÌÓ, ˜ÚÓ ÔÓ ÏÂÂ ÒÌËÊÂÌËfl Ú‡ÍÒÓÌÓÏË˜Â-ÒÍÓ„Ó ‡Ì„‡ (‚ fl‰Û ÓÚfl‰˚ – ÒÂÏÂÈÒÚ‚‡ – Ó‰˚)ÓÒÚ ‡ÁÌÓÓ·‡ÁËfl îåÅ ÒÚ‡ÌÓ‚ËÚÒfl ‚ÒÂ ÏÂÌÂÂ ÔÓ-ıÓÊ Ì‡ ÎÓ„ËÒÚË˜ÂÒÍËÈ Ë ‚ÒÂ ·ÓÎÂÂ – Ì‡ ˝ÍÒÔÓÌÂÌˆË-‡Î¸Ì˚È (Lane, Benton, 2003). ùÚÓ Ì‡·Î˛‰ÂÌËÂ ÒÎÂ‰Û-ÂÚ ‰ÓÔÓÎÌËÚ¸ ÚÂÏ, ˜ÚÓ ÔÓ ÏÂÂ ÒÌËÊÂÌËfl ‡Ì„‡ (‚fl‰Û ÒÂÏÂÈÒÚ‚‡ – Ó‰˚ – ‚Ë‰˚) ÁÌ‡˜ËÚÂÎ¸ÌÓ ‚ÓÁ‡Ò-Ú‡ÂÚ ÒıÓ‰ÒÚ‚Ó „‡ÙËÍÓ‚ Ò „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰Â-Î¸˛, ÔË˜ÂÏ ÒıÓ‰ÒÚ‚Ó Ëı Ò ˝ÍÒÔÓÌÂÌÚÓÈ ‚Ó ‚ÒÂıÚÂı ÒÎÛ˜‡flı ÁÌ‡˜ËÚÂÎ¸ÌÓ ÌËÊÂ. äÓÂÎflˆËfl (

R

2

) c„ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎ¸˛ ‰Îfl ÒÂÏÂÈÒÚ‚ – 0.77, Ó-‰Ó‚ – 0.85, ‚Ë‰Ó‚ – 0.91; Ò ˝ÍÒÔÓÌÂÌˆË‡Î¸ÌÓÈ ‰Îfl ÒÂ-ÏÂÈÒÚ‚ – 0.56, Ó‰Ó‚ – 0.46, ‚Ë‰Ó‚ – 0.51.

2. ÑËÌ‡ÏËÍ‡ ‡ÁÌÓÓ·‡ÁËfl îäÅ.

ÑËÌ‡ÏËÍ‡ ̃ ËÒ-Î‡ ÒÂÏÂÈÒÚ‚ îäÅ Ó˜ÂÌ¸ ıÓÓ¯Ó ÓÔËÒ˚‚‡ÂÚÒfl ˝ÍÒ-ÔÓÌÂÌˆË‡Î¸ÌÓÈ ÏÓ‰ÂÎ¸˛ (ËÒ. 2,

‡

,

R

2

= 0.87), Ó‰Ì‡-ÍÓ ÔÓÒÚ‡fl „ËÔÂ·ÓÎË˜ÂÒÍ‡fl ÏÓ‰ÂÎ¸ ÓÔËÒ˚‚‡ÂÚ ÂÂÂ˘Â ÎÛ˜¯Â (ËÒ. 2,

·

, ÍË‚‡fl

2

,

R

2

= 0.95). Ç‚Â‰ÂÌËÂ‚ „ËÔÂ·ÓÎË˜ÂÒÍÛ˛ ÏÓ‰ÂÎ¸ ‰ÓÔÓÎÌËÚÂÎ¸ÌÓÈ ÍÓÌ-ÒÚ‡ÌÚ˚ ÔÓÁ‚ÓÎflÂÚ ÔÓ‚˚ÒËÚ¸ ÍÓÂÎflˆË˛ (ËÒ. 2,

·

,ÍË‚‡fl

3

,

R

2

= 0.97). èË˜ËÌ˚ Á‡ÏÂÚÌÓ„Ó ‡ÒıÓÊ‰Â-ÌËfl ÏÓ‰ÂÎ¸Ì˚ı Ë Ì‡·Î˛‰‡ÂÏ˚ı ÁÌ‡˜ÂÌËÈ ‚ ÎÂ‚ÓÈ˜‡ÒÚË „‡ÙËÍ‡ ‡ÒÒÏ‡ÚË‚‡˛ÚÒfl ‚ ‡Á‰. “é·ÒÛÊ‰Â-ÌËÂ”.

óËÒÎÓ Ú‡ÍÒÓÌÓ‚ Ú‡ÍÓ„Ó ‚˚ÒÓÍÓ„Ó ‡Ì„‡, Í‡Í ÒÂ-ÏÂÈÒÚ‚Ó, fl‚ÎflÂÚÒfl ‰Ó‚ÓÎ¸ÌÓ „Û·ÓÈ ÓˆÂÌÍÓÈ ·ËÓ-‡ÁÌÓÓ·‡ÁËfl. èÓ ‡Ì‡ÎÓ„ËË Ò ÂÁÛÎ¸Ú‡Ú‡ÏË, ÔÓÎÛ-˜ÂÌÌ˚ÏË ÔÓ îåÅ, ÏÓÊÌÓ ÓÊË‰‡Ú¸, ˜ÚÓ ‡Ì‡ÎËÁ ‰Ë-Ì‡ÏËÍË ˜ËÒÎ‡ ÍÓÌÚËÌÂÌÚ‡Î¸Ì˚ı Ó‰Ó‚ ÏÓ„ ·˚


ÉàèÖêÅéãàóÖëäàâ êéëí

177

ÓÍ‡Á‡Ú¸Òfl ·ÓÎÂÂ ÔÓÍ‡Á‡ÚÂÎ¸Ì˚Ï. ä ÒÓÊ‡ÎÂÌË˛,ÔÓÎÌ˚Â Ò‚Ó‰ÍË ÔÓ ÒÚ‡ÚË„‡ÙË˜ÂÒÍÓÏÛ ‡ÒÔÓ-ÒÚ‡ÌÂÌË˛ Ó‰Ó‚ ÍÓÌÚËÌÂÌÚ‡Î¸Ì˚ı Ó„‡ÌËÁÏÓ‚ Ì‡ÒÂ„Ó‰Ìfl¯ÌËÈ ‰ÂÌ¸ ÓÚÒÛÚÒÚ‚Û˛Ú. éÚ˜‡ÒÚË ˝ÚÓÚ ÌÂ-‰ÓÒÚ‡ÚÓÍ ÏÓÊÌÓ ‚ÓÒÔÓÎÌËÚ¸, ÂÍÓÌÒÚÛËÓ‚‡‚ „Ë-ÔÓÚÂÚË˜ÂÒÍÛ˛ ‰ËÌ‡ÏËÍÛ ˜ËÒÎ‡ Ó‰Ó‚ Ì‡ ÓÒÌÓ‚Â˜ËÒÎ‡ ÒÂÏÂÈÒÚ‚. å˚ ÔÓÔ˚Ú‡ÎËÒ¸ ̋ ÚÓ Ò‰ÂÎ‡Ú¸, ËÒıÓ-‰fl ËÁ ÔÂ‰ÔÓÎÓÊÂÌËfl, ˜ÚÓ Ó·˘ËÂ ÍÓÎË˜ÂÒÚ‚ÂÌÌ˚ÂÁ‡ÍÓÌÓÏÂÌÓÒÚË, ÔÓÒÎÂÊË‚‡˛˘ËÂÒfl ‚ ÒÓÓÚÌÓ¯Â-ÌËË ̃ ËÒÎ‡ Ú‡ÍÒÓÌÓ‚ ‡ÁÌÓ„Ó ‡Ì„‡ ‚ îåÅ, ÒÔ‡‚Â‰-ÎË‚˚ Ú‡ÍÊÂ Ë ‰Îfl îäÅ.

Ç îåÅ ˜ËÒÎÓ Ó‰Ó‚, ÒÛ˘ÂÒÚ‚Ó‚‡‚¯Ëı ‚ ‰‡ÌÌÓÏÔÓ‰˙flÛÒÂ, Ò‚flÁ‡ÌÓ Ò ̃ ËÒÎÓÏ ÒÛ˘ÂÒÚ‚Ó‚‡‚¯Ëı ‚ ̋ ÚÓÊÂ ‚ÂÏfl ÒÂÏÂÈÒÚ‚ ÌÂÎËÌÂÈÌÓÈ Á‡‚ËÒËÏÓÒÚ¸˛(ËÒ. 2,

‚

), ÍÓÚÓ‡fl Ò ·ÓÎ¸¯ÓÈ ÚÓ˜ÌÓÒÚ¸˛ (

R

2

= 0.99)ÓÔËÒ˚‚‡ÂÚÒfl Û‡‚ÌÂÌËÂÏ

f

= 0.000004

g

3

– 0.003

g

2

++ 2.9626

g

, „‰Â

f

– ˜ËÒÎÓ ÒÂÏÂÈÒÚ‚,

g

– ˜ËÒÎÓ Ó‰Ó‚.ÖÒÎË ‡ÒÒ˜ËÚ‡Ú¸ ÔË ÔÓÏÓ˘Ë ˝ÚÓÈ ÙÓÏÛÎ˚ ‰ËÌ‡-ÏËÍÛ ˜ËÒÎ‡ Ó‰Ó‚ ‚ îåÅ Ì‡ ÓÒÌÓ‚‡ÌËË ‰‡ÌÌ˚ı ÔÓ˜ËÒÎÛ ÒÂÏÂÈÒÚ‚, ÂÁÛÎ¸Ú‡Ú ÔÓÎÛ˜‡ÂÚÒfl ‚ÂÒ¸Ï‡ ·ÎË-ÁÓÍ Í Ù‡ÍÚË˜ÂÒÍËÏ ‰‡ÌÌ˚Ï ÔÓ Ó‰Ó‚ÓÏÛ ‡ÁÌÓÓ·-‡ÁË˛ (ËÒ. 2,

„

). Ç˚˜ËÒÎÂÌÌ˚È Ì‡ ÓÒÌÓ‚Â

f

„‡ÙËÍ

g

(ËÒ. 2,

„

, ÍË‚‡fl

2

) ÔÓ˜ÚË ÔÓÎÌÓÒÚ¸˛ ÒÓ‚Ô‡‰‡ÂÚ ÒÂ‡Î¸Ì˚Ï (ËÒ. 2,

„

, ÍË‚‡fl

1

). éÚÎË˜Ëfl Ò‚Ó‰flÚÒfl ÍÒ„Î‡ÊË‚‡ÌË˛ Ì‡Ë·ÓÎÂÂ ÂÁÍËı Ï‡ÍÒËÏÛÏÓ‚ Ë ÏË-ÌËÏÛÏÓ‚, ˜ÚÓ ÌÂ ËÏÂÂÚ Ô‡ÍÚË˜ÂÒÍË ÌËÍ‡ÍÓ„Ó ÁÌ‡-˜ÂÌËfl ‰Îfl ÔÓ‰·Ó‡ Ï‡ÚÂÏ‡ÚË˜ÂÒÍËı ÙÛÌÍˆËÈ (˝ÍÒ-ÔÓÌÂÌˆË‡Î¸ÌÓÈ ËÎË „ËÔÂ·ÓÎË˜ÂÒÍÓÈ), ÓÔËÒ˚‚‡˛-˘Ëı „ÂÌÂ‡Î¸Ì˚È ÚÂÌ‰ Ù‡ÌÂÓÁÓÈÒÍÓÈ ‰ËÌ‡ÏËÍË·ËÓ‡ÁÌÓÓ·‡ÁËfl.

ÑÓÔÛÒÚËÏ, ˜ÚÓ ‚ îäÅ Á‡ÍÓÌ ÒÓÓÚÌÓ¯ÂÌËfl

f

Ë

g

·˚Î Ú‡ÍËÏ ÊÂ, Í‡Í ‚ îåÅ. êÂÍÓÌÒÚÛËÓ‚‡ÌÌ‡fl Ì‡ÓÒÌÓ‚Â ˝ÚÓ„Ó ‰ÓÔÛ˘ÂÌËfl ‰ËÌ‡ÏËÍ‡ ˜ËÒÎ‡ Ó‰Ó‚ ‚îäÅ (ËÒ. 2,

‰

) ÓÔËÒ˚‚‡ÂÚÒfl „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ-‰ÂÎ¸˛ Á‡ÏÂÚÌÓ ÎÛ˜¯Â, ˜ÂÏ ˝ÍÒÔÓÌÂÌˆË‡Î¸ÌÓÈ (‰Îfl„ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎË

R

2

= 0.94, ‰Îfl ˝ÍÒÔÓÌÂÌ-ˆË‡Î¸ÌÓÈ

R

2

= 0.86).

3.

ëÓ‚ÓÍÛÔÌ‡fl ‰ËÌ‡ÏËÍ‡ ‡ÁÌÓÓ·‡ÁËfl ‚ÒÂÈ Ù‡-ÌÂÓÁÓÈÒÍÓÈ ·ËÓÚ˚

(ÏÓÒÍÓÈ Ë ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ)ÒÓÓÚ‚ÂÚÒÚ‚ÛÂÚ „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎË Ì‡ÏÌÓ„ÓÎÛ˜¯Â (

R

2

= 0.96), ̃ ÂÏ ̋ ÍÒÔÓÌÂÌˆË‡Î¸ÌÓÈ (

R

2

= 0.80)(ËÒ. 3,

‡

). ÖÒÎË ÒÎÓÊËÚ¸ ‰‡ÌÌ˚Â ÔÓ ‰ËÌ‡ÏËÍÂ Ó‰Ó-‚Ó„Ó ‡ÁÌÓÓ·‡ÁËfl îåÅ (ËÒ. 1,

·

) Ò ÂÍÓÌÒÚÛËÓ-‚‡ÌÌÓÈ ‰ËÌ‡ÏËÍÓÈ Ó‰Ó‚Ó„Ó ‡ÁÌÓÓ·‡ÁËfl îäÅ(ËÒ. 2,

‰

), ÔÓÎÛ˜ËÏ ÔË·ÎËÁËÚÂÎ¸ÌÛ˛ ÓˆÂÌÍÛ ‰ËÌ‡ÏË-ÍË Ó·˘Â„Ó Ó‰Ó‚Ó„Ó ‡ÁÌÓÓ·‡ÁËfl ·ËÓÚ˚ (ËÒ. 3,

·

).Ñ‡ÌÌ˚È „‡ÙËÍ ÚÓÊÂ Ì‡ÏÌÓ„Ó ÎÛ˜¯Â ÍÓÂÎËÛÂÚÒ „ËÔÂ·ÓÎÓÈ (

R

2

= 0.95), ˜ÂÏ Ò ˝ÍÒÔÓÌÂÌÚÓÈ (

R

2

== 0.67).

éÅëìÜÑÖçàÖ

1. ÑËÌ‡ÏËÍ‡ ‡ÁÌÓÓ·‡ÁËfl Ù‡ÌÂÓÁÓÈÒÍÓÈ ·ËÓ-Ú˚ ÒÓÓÚ‚ÂÚÒÚ‚ÛÂÚ ÏÓ‰ÂÎË „ËÔÂ·ÓÎË˜ÂÒÍÓ„Ó Ó-ÒÚ‡.

èÓÎÛ˜ÂÌÌ˚Â ÂÁÛÎ¸Ú‡Ú˚ ÔÓÍ‡Á˚‚‡˛Ú, ̃ ÚÓ „Ë-ÔÂ·ÓÎË˜ÂÒÍ‡fl ÏÓ‰ÂÎ¸ ÔËÎÓÊËÏ‡ ÌÂ ÚÓÎ¸ÍÓ Í ‰Ë-Ì‡ÏËÍÂ ‡ÁÌÓÓ·‡ÁËfl îåÅ Ì‡ ÛÓ‚ÌÂ Ó‰Ó‚, Í‡Í·˚ÎÓ ÓÚÏÂ˜ÂÌÓ Ì‡ÏË ‡ÌÂÂ, ÌÓ Ë Ì‡ ÛÓ‚ÌÂ ÒÂ-

ÏÂÈÒÚ‚ Ë ‚Ë‰Ó‚, ‡ Ú‡ÍÊÂ Í ‰ËÌ‡ÏËÍÂ ‡ÁÌÓÓ·‡ÁËflîäÅ Ë ‚ÒÂÈ ·ËÓÚ˚ ‚ ˆÂÎÓÏ (îåÅ + îäÅ) Ì‡ ÛÓ‚-ÌÂ ÒÂÏÂÈÒÚ‚ Ë Ó‰Ó‚. ÇÓ ‚ÒÂı ˝ÚËı ÒÎÛ˜‡flı ˝ÏÔËË-

1000

200

0–550

óËÒÎÓ ÒÂÏÂÈÒÚ‚

–350 –150

800

600

400

‡

5000

0–550

óËÒÎÓ Ó‰Ó‚

–350 –150

·

45004000350030002500200015001000500

25000

0–550

óËÒÎÓ ‚Ë‰Ó‚

–350 –150

‚

20000

15000

10000

5000

êËÒ. 1.

ëÓÓÚ‚ÂÚÒÚ‚ËÂ ‰ËÌ‡ÏËÍË ‡ÁÌÓÓ·‡ÁËfl îåÅ ÏÓ-‰ÂÎË „ËÔÂ·ÓÎË˜ÂÒÍÓ„Ó ÓÒÚ‡. èÓ „ÓËÁÓÌÚ‡Î¸ÌÓÈ ÓÒË –‚ÂÏfl, ÏÎÌ. ÎÂÚ.

‡

– ̃ ËÒÎÓ ÒÂÏÂÈÒÚ‚ (ÔÓ: Sepkoski, 1992),

·

– Ó‰Ó‚ (ÔÓ ·‡ÁÂ ‰‡ÌÌ˚ı ëÂÔÍÓÒÍË),

‚

– ‚Ë‰Ó‚ (ËÒÔÓÎ¸-ÁÓ‚‡Ì‡ ÛÒÂ‰ÌÂÌÌ‡fl ÓˆÂÌÍ‡ ÔÓ: å‡ÍÓ‚, 2003). íÓÌÍ‡flÎËÌËfl – ˜ËÒÎÓ Ú‡ÍÒÓÌÓ‚, ÚÓÎÒÚ‡fl – „ËÔÂ·ÓÎË˜ÂÒÍËÈÚÂÌ‰, ÔÛÌÍÚË – ˝ÍÒÔÓÌÂÌˆË‡Î¸Ì˚È ÚÂÌ‰. è‡‡ÏÂÚ-˚ ÏÓ‰ÂÎÂÈ: ÒÂÏÂÈÒÚ‚‡, ˝ÍÒÔÓÌÂÌÚ‡:

N

t

= 888

e

0.0026

t

,

R

2

= 0.56, „ËÔÂ·ÓÎ‡:

N

t

= 131660/(100 –

t

),

R

2

= 0.77; Ó-‰˚, ˝ÍÒÔÓÌÂÌÚ‡:

N

t

= 2029

e

0.003

t

;

R

2

= 0.46, „ËÔÂ·ÓÎ‡:

N

t

= 183320/(37 –

t

),

R

2

= 0.85; ‚Ë‰˚, ˝ÍÒÔÓÌÂÌÚ‡:

N

t

== 9443

e0.004t, R2 = 0.51; „ËÔÂ·ÓÎ‡: Nt = 892874/(35 – t),R2 = 0.91.

2

178



1200

200

0–550


–450 –250

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‡

–350 –150 –50

1600

1400

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1200

200

0–550


–450 –250

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600

400

·

–350 –150 –50

1800

1400

800

123

4000

0

óËÒÎÓ Ó‰Ó‚ ‚ ÔÓ‰˙flÛÒÂ

1000

‚

6000

5000

4000

0–550


–450 –250

3000

1000

„

–350 –150 –50

6000

5000

2000

123000

2000

1000

500óËÒÎÓ ÒÂÏÂÈÒÚ‚ ‚ ÔÓ‰˙flÛÒÂ

6000

0–550


–450 –250

4000

‰

–350 –150 –50

10000

8000

2000

1600

êËÒ. 2. ëÓÓÚ‚ÂÚÒÚ‚ËÂ ‰ËÌ‡ÏËÍË ‡ÁÌÓÓ·‡ÁËfl îäÅ ÏÓ‰ÂÎË „ËÔÂ·ÓÎË˜ÂÒÍÓ„Ó ÓÒÚ‡. èÓ „ÓËÁÓÌÚ‡Î¸ÌÓÈ ÓÒË – ‚ÂÏfl,ÏÎÌ. ÎÂÚ (ÍÓÏÂ ÙË„Û˚ ‚). ‡ – ˜ËÒÎÓ ÒÂÏÂÈÒÚ‚; ÔÛÌÍÚËÌ‡fl ÎËÌËfl – ˝ÍÒÔÓÌÂÌˆË‡Î¸Ì˚È ÚÂÌ‰ (Nt = 1334e0.0094t, R2 = 0.87);· – ˜ËÒÎÓ ÒÂÏÂÈÒÚ‚ (1), „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÚÂÌ‰ ·ÂÁ ÍÓÌÒÚ‡ÌÚ˚ (2; Nt = 62089/(38 – t), R2 = 0.95); „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÚÂÌ‰ ÒÍÓÌÒÚ‡ÌÚÓÈ (3; Nt = 124993/(70 – t), –245, R2 = 0.97); ‚ – Á‡‚ËÒËÏÓÒÚ¸ ˜ËÒÎ‡ Ó‰Ó‚ ÓÚ ˜ËÒÎ‡ ÒÂÏÂÈÒÚ‚ ‚ Ù‡ÌÂÓÁÓÈÒÍÓÈ ÏÓ-ÒÍÓÈ ·ËÓÚÂ (Í‡Ê‰‡fl ÚÓ˜Í‡ ÒÓÓÚ‚ÂÚÒÚ‚ÛÂÚ Ó‰ÌÓÏÛ ËÁ ÔÓ‰˙flÛÒÓ‚ Ù‡ÌÂÓÁÓfl, ÎËÌËfl ÚÂÌ‰‡ Ò„ÂÌÂËÓ‚‡Ì‡ Û‡‚ÌÂÌËÂÏ f == 0.000004g3 – 0.003g2 + 2.9626g, „‰Â f – ˜ËÒÎÓ ÒÂÏÂÈÒÚ‚, g – ˜ËÒÎÓ Ó‰Ó‚; R2 = 0.99); „ – ‡ÔÓ·‡ˆËfl ÏÂÚÓ‰‡ ÂÍÓÌÒÚÛÍˆËË˜ËÒÎ‡ Ó‰Ó‚ ÔÓ ‰‡ÌÌ˚Ï Ó ̃ ËÒÎÂ ÒÂÏÂÈÒÚ‚: 1 – Â‡Î¸ÌÓÂ ̃ ËÒÎÓ Ó‰Ó‚ îåÅ, 2 – ‚˚˜ËÒÎÂÌÌÓÂ Ì‡ ÓÒÌÓ‚Â ÔÓÒÎÂ‰ÌÂÈ ÙÓÏÛÎ˚;‰ – ÂÍÓÌÒÚÛËÓ‚‡ÌÌ‡fl ‰ËÌ‡ÏËÍ‡ Ó‰Ó‚Ó„Ó ‡ÁÌÓÓ·‡ÁËfl îäÅ; ÚÓÌÍ‡fl ÎËÌËfl – ˜ËÒÎÓ Ú‡ÍÒÓÌÓ‚, ÚÓÎÒÚ‡fl – „ËÔÂ·ÓÎË˜Â-ÒÍËÈ ÚÂÌ‰, ÔÛÌÍÚË – ˝ÍÒÔÓÌÂÌˆË‡Î¸Ì˚È ÚÂÌ‰; Ô‡‡ÏÂÚ˚ ÏÓ‰ÂÎÂÈ: ˝ÍÒÔÓÌÂÌÚ‡: Nt = 4781e0.0104t, R2 = 0.86; „ËÔÂ·ÓÎ‡:Nt = = 272095/(29 – t), R2 = 0.94.


ÉàèÖêÅéãàóÖëäàâ êéëí 179

˜ÂÒÍËÂ ËÎË ÂÍÓÌÒÚÛËÓ‚‡ÌÌ˚Â ÍË‚˚Â ‰ËÌ‡ÏËÍË‡ÁÌÓÓ·‡ÁËfl ÍÓÂÎËÛ˛Ú Ò „ËÔÂ·ÓÎÓÈ ÎÛ˜¯Â,˜ÂÏ Ò ˝ÍÒÔÓÌÂÌÚÓÈ (Ú‡·ÎËˆ‡).

åÓÊÌÓ Á‡ÏÂÚËÚ¸, ˜ÚÓ Ò ˝ÍÒÔÓÌÂÌˆË‡Î¸ÌÓÈ ÏÓ-‰ÂÎ¸˛ ÎÛ˜¯Â ‚ÒÂ„Ó ÍÓÂÎËÛÂÚ ‰ËÌ‡ÏËÍ‡ ‡ÁÌÓ-Ó·‡ÁËfl îäÅ, ÁÌ‡˜ËÚÂÎ¸ÌÓ ıÛÊÂ – ‚ÒÂÈ ·ËÓÚ˚,Ì‡ËÏÂÌ¸¯‡fl ÍÓÂÎflˆËfl ı‡‡ÍÚÂÌ‡ ‰Îfl îåÅ. ë„ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎ¸˛, Ì‡ÔÓÚË‚, Ï‡ÍÒËÏ‡Î¸-ÌÓ ÍÓÂÎËÛÂÚ ‚Òfl ·ËÓÚ‡ ‚ ˆÂÎÓÏ, ‡ îäÅ Ë îåÅ,‚ÁflÚ˚Â ÔÓ ÓÚ‰ÂÎ¸ÌÓÒÚË, ÍÓÂÎËÛ˛Ú Ò ÌÂÈ ÌÂ-ÒÍÓÎ¸ÍÓ ıÛÊÂ. ç‡ Ì‡¯ ‚Á„Îfl‰, Ï‡ÍÒËÏ‡Î¸Ì‡fl ÍÓ-ÂÎflˆËfl ËÏÂÌÌÓ ÒÓ‚ÓÍÛÔÌÓÈ ‰ËÌ‡ÏËÍË ‡ÁÌÓÓ·‡-ÁËfl ‚ÒÂÈ ·ËÓÚ˚ Ò „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎ¸˛ ÓÚ‡-Ê‡ÂÚ ÚÓÚ Ù‡ÍÚ, ˜ÚÓ ·ËÓÒÙÂ‡ ‡Á‚Ë‚‡Î‡Ò¸ ‚Ù‡ÌÂÓÁÓÂ ‚Ó ÏÌÓ„Ëı ÓÚÌÓ¯ÂÌËflı Í‡Í Â‰ËÌ‡fl ÒË-ÒÚÂÏ‡, ‚ ÍÓÚÓÓÈ ˝‚ÓÎ˛ˆËÓÌÌ˚Â ËÁÏÂÌÂÌËfl ÓÚ-‰ÂÎ¸Ì˚ı ˝ÎÂÏÂÌÚÓ‚ (‚ ÚÓÏ ˜ËÒÎÂ ‰Ë‚ÂÒËÙËÍ‡ˆËflÚÂı ËÎË ËÌ˚ı „ÛÔÔ) ·˚ÎË ‚Á‡ËÏÓÒ‚flÁ‡Ì˚.

ç‡ËÏÂÌ¸¯ËÂ ‡ÁÎË˜Ëfl ÏÂÊ‰Û ‚ÂÎË˜ËÌÓÈ ÍÓ-ÂÎflˆËË Ò ˝ÍÒÔÓÌÂÌˆË‡Î¸ÌÓÈ Ë „ËÔÂ·ÓÎË˜ÂÒÍÓÈÏÓ‰ÂÎ¸˛ ı‡‡ÍÚÂÌ˚ ‰Îfl îäÅ (ËÌ˚ÏË ÒÎÓ‚‡ÏË,‰ËÌ‡ÏËÍ‡ îäÅ ÓÔËÒ˚‚‡ÂÚÒfl ˝ÍÒÔÓÌÂÌÚÓÈ ÔÓ˜ÚËÚ‡Í ÊÂ ıÓÓ¯Ó, Í‡Í Ë „ËÔÂ·ÓÎÓÈ). ùÚÓ Ò‚flÁ‡ÌÓ ‚ÔÂ‚Û˛ Ó˜ÂÂ‰¸ Ò Á‡ÏÂÚÌ˚Ï ‡ÒıÓÊ‰ÂÌËÂÏ ÏÂÊ‰ÛÌ‡·Î˛‰‡ÂÏ˚ÏË Ë ÔÂ‰ÒÍ‡Á‡ÌÌ˚ÏË „ËÔÂ·ÓÎË˜Â-ÒÍÓÈ ÏÓ‰ÂÎ¸˛ ÁÌ‡˜ÂÌËflÏË ‡ÁÌÓÓ·‡ÁËfl ‚ ÎÂ‚ÓÈ˜‡ÒÚË „‡ÙËÍ‡ Ì‡ ËÒ. 2,·. ÉËÔÂ·ÓÎË˜ÂÒÍ‡fl ÏÓ-‰ÂÎ¸ “ÔÂ‰ÒÍ‡Á˚‚‡ÂÚ” Ì‡ ‡ÌÌËı ˝Ú‡Ô‡ı ÒÚ‡ÌÓ‚ÎÂ-ÌËfl ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ ·ËÓÚ˚ (‚ ÒËÎÛÂ, ‰Â‚ÓÌÂ Ë‡ÌÌÂÏ Í‡·ÓÌÂ) ·ÓÎÂÂ ‚˚ÒÓÍÓÂ ‡ÁÌÓÓ·‡ÁËÂ, ̃ ÂÏÚÓ, ÍÓÚÓÓÂ Â‡Î¸ÌÓ Ì‡·Î˛‰‡ÂÚÒfl ‚ ÎÂÚÓÔËÒË (ÍË-‚‡fl 2 Ì‡ ËÒ. 2,· ‚ Ò‚ÓÂÈ ÎÂ‚ÓÈ ˜‡ÒÚË ÔÓıÓ‰ËÚ ‚˚-¯Â, ˜ÂÏ ÍË‚‡fl 1). Ñ‡ÌÌÓÂ ÌÂÒÓÓÚ‚ÂÚÒÚ‚ËÂ ÏÓÊÂÚ·˚Ú¸ ÛÒÚ‡ÌÂÌÓ ‚‚Â‰ÂÌËÂÏ ‚ „ËÔÂ·ÓÎË˜ÂÒÍÛ˛ ÏÓ-‰ÂÎ¸ ‰ÓÔÓÎÌËÚÂÎ¸ÌÓÈ ÍÓÌÒÚ‡ÌÚ˚ (ËÒ. 2,·, ÍË‚‡fl 3).ùÚÓ, Ò Ó‰ÌÓÈ ÒÚÓÓÌ˚, ÔÓ‚˚¯‡ÂÚ ÍÓÂÎflˆË˛ ÏÓ-‰ÂÎË Ò Ù‡ÍÚË˜ÂÒÍËÏË ‰‡ÌÌ˚ÏË (R2 = 0.97), ‡ Ò ‰Û-„ÓÈ – ÎË¯¸ ÔÓ‰˜ÂÍË‚‡ÂÚ ÚÓ ÚÂ·Û˛˘ÂÂ Ó·˙flÒÌÂ-ÌËfl Ó·ÒÚÓflÚÂÎ¸ÒÚ‚Ó, ˜ÚÓ ı‡‡ÍÚÂ „ËÔÂ·ÓÎË˜Â-ÒÍÓ„Ó ÓÒÚ‡ ‡ÁÌÓÓ·‡ÁËfl îäÅ ‚ ÔÓÁ‰ÌÂÈ¯ËÂ ̋ ÔÓıË(Ì‡˜ËÌ‡fl ÒÓ ÒÂ‰ÌÂ„Ó Í‡·ÓÌ‡) ÔÂ‰ÔÓÎ‡„‡ÂÚ ·ÓÎÂÂ‚˚ÒÓÍËÈ ÛÓ‚ÂÌ¸ ‡ÁÌÓÓ·‡ÁËfl Ì‡ Ì‡˜‡Î¸Ì˚ı ˝Ú‡-Ô‡ı ÒÚ‡ÌÓ‚ÎÂÌËfl îäÅ, ̃ ÂÏ Â‡Î¸ÌÓ Ì‡·Î˛‰‡ÂÏ˚È.

ç‡ ÔÂ‚˚È ‚Á„Îfl‰ ̋ ÚÓ ÏÓÊÌÓ ‡ÒˆÂÌËÚ¸ Í‡Í Ò‚Ë-‰ÂÚÂÎ¸ÒÚ‚Ó ÌÂ‰ÓÒÚ‡ÚÓ˜ÌÓ ÔÓÎÌÓ„Ó ÒÓÓÚ‚ÂÚÒÚ‚ËflÓÒÚ‡ ‡ÁÌÓÓ·‡ÁËfl îäÅ „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰Â-ÎË, ÔÓ Í‡ÈÌÂÈ ÏÂÂ Ì‡ Ì‡˜‡Î¸Ì˚ı ˝Ú‡Ô‡ı. Ç ‰ÂÈ-ÒÚ‚ËÚÂÎ¸ÌÓÒÚË, Ì‡ Ì‡¯ ‚Á„Îfl‰, ‰‡ÌÌ˚È ÂÁÛÎ¸Ú‡Úfl‚ÎflÂÚÒfl ÓÊË‰‡ÂÏ˚Ï Ë ÏÓÊÂÚ ·˚Ú¸ ËÌÚÂÔÂÚËÓ-‚‡Ì Í‡Í ‰ÓÔÓÎÌËÚÂÎ¸Ì˚È ‰Ó‚Ó‰ ‚ ÔÓÎ¸ÁÛ ‡‰ÂÍ‚‡Ú-ÌÓÒÚË „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎË. å˚ ‚Ë‰ËÏ ‰‚Â ‚Â-ÓflÚÌ˚Â ÔË˜ËÌ˚ ·ÓÎÂÂ ÌËÁÍÓ„Ó Ì‡·Î˛‰‡ÂÏÓ„Ó‡ÁÌÓÓ·‡ÁËfl îäÅ ‚ ÒËÎÛÂ – ‡ÌÌÂÏ Í‡·ÓÌÂ ÔÓÒ‡‚ÌÂÌË˛ Ò ÏÓ‰ÂÎ¸Ì˚ÏË ÔÂ‰ÒÍ‡Á‡ÌËflÏË.

ÇÓ-ÔÂ‚˚ı, Ì‡ Ì‡˜‡Î¸Ì˚ı ̋ Ú‡Ô‡ı Ò‚ÓÂ„Ó ÒÚ‡ÌÓ‚-ÎÂÌËfl ÍÓÌÚËÌÂÌÚ‡Î¸Ì‡fl ·ËÓÚ‡, Ó˜Â‚Ë‰ÌÓ, Â˘Â ÌÂ ·˚-Î‡ Ò‡ÏÓÒÚÓflÚÂÎ¸ÌÓÈ ÒËÒÚÂÏÓÈ, Í‡ÍÓÈ ÓÌ‡ ÒÚ‡Î‡ ÔÓÁ-ÊÂ, ‡ fl‚ÎflÎ‡Ò¸ ‚Ó ÏÌÓ„Ëı ÓÚÌÓ¯ÂÌËflı ÎË¯¸ “ÔÓ‰ÓÎ-

ëÓÓÚ‚ÂÚÒÚ‚ËÂ ‰ËÌ‡ÏËÍË ·ËÓ‡ÁÌÓÓ·‡ÁËfl ˝ÍÒÔÓÌÂÌˆË-‡Î¸ÌÓÈ Ë „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎflÏ Ì‡ ‡ÁÌ˚ı Ú‡ÍÒÓ-ÌÓÏË˜ÂÒÍËı ÛÓ‚Ìflı

ÅËÓÚ‡í‡ÍÒÓÌÓÏË˜ÂÒÍËÈ ÛÓ‚ÂÌ¸

ÒÂÏÂÈÒÚ‚‡ Ó‰˚ ‚Ë‰˚

åÓÒÍ‡fl 0.77/0.56 0.85/0.46 0.91/0.51äÓÌÚËÌÂÌÚ‡Î¸Ì‡fl 0.95/0.87 0.94/0.86 ?

ÇÒfl 0.96/0.80 0.95/0.67 ?

èËÏÂ˜‡ÌËÂ. èÂ‚ÓÂ ˜ËÒÎÓ ‚ Í‡Ê‰ÓÈ ÍÎÂÚÍÂ ÒÓÓÚ‚ÂÚÒÚ‚ÛÂÚÁÌ‡˜ÂÌË˛ R2 ‰Îfl „ËÔÂ·ÓÎË˜ÂÒÍÓÈ, ‚ÚÓÓÂ – ‰Îfl ˝ÍÒÔÓÌÂÌˆË-‡Î¸ÌÓÈ ÏÓ‰ÂÎË. äÛÒË‚ÓÏ ‚˚‰ÂÎÂÌ˚ ÁÌ‡˜ÂÌËfl, ÔÓÎÛ˜ÂÌÌ˚ÂÌÂ ‰Îfl ˝ÏÔËË˜ÂÒÍËı, ‡ ‰Îfl ÂÍÓÌÒÚÛËÓ‚‡ÌÌ˚ı („ËÔÓÚÂÚË-˜ÂÒÍËı) „‡ÙËÍÓ‚ ‰ËÌ‡ÏËÍË ‡ÁÌÓÓ·‡ÁËfl; ÊËÌ˚Ï ¯ËÙ-ÚÓÏ – ÁÌ‡˜ÂÌËfl, ÓÔÛ·ÎËÍÓ‚‡ÌÌ˚Â Ì‡ÏË ‡ÌÂÂ (å‡ÍÓ‚, äÓÓ-Ú‡Â‚, 2007; Markov, Korotayev, 2007).

1000

0–550


–450 –150

500

‡

14000

0–550


–450 –150

·12000

10000

8000

6000

4000

2000

–250–350 –50

3000

2500

2000

1500

–350 –50–200

êËÒ. 3. ëÓÓÚ‚ÂÚÒÚ‚ËÂ ÒÛÏÏ‡ÌÓÈ ‰ËÌ‡ÏËÍË ‡ÁÌÓÓ·‡-ÁËfl Ù‡ÌÂÓÁÓÈÒÍÓÈ ·ËÓÚ˚ (ÏÓÒÍÓÈ Ë ÍÓÌÚËÌÂÌÚ‡Î¸-ÌÓÈ) ÏÓ‰ÂÎË „ËÔÂ·ÓÎË˜ÂÒÍÓ„Ó ÓÒÚ‡. èÓ „ÓËÁÓÌÚ‡Î¸-ÌÓÈ ÓÒË – ‚ÂÏfl ‚ ÏÎÌ. ÎÂÚ. íÓÌÍ‡fl ÎËÌËfl – ˜ËÒÎÓ Ú‡Í-ÒÓÌÓ‚, ÚÓÎÒÚ‡fl – „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÚÂÌ‰, ÔÛÌÍÚË –˝ÍÒÔÓÌÂÌˆË‡Î¸Ì˚È ÚÂÌ‰. ‡ – ˜ËÒÎÓ ÒÂÏÂÈÒÚ‚ (ÔÓ ‰‡Ì-Ì˚Ï The Fossil Record 2), Ô‡‡ÏÂÚ˚ ÏÓ‰ÂÎÂÈ: ˝ÍÒÔÓ-ÌÂÌÚ‡: Nt = 1783e0.0037t, R2 = 0.80; „ËÔÂ·ÓÎ‡: Nt == 199551/(70 – t), R2 = 0.96; · – ˜ËÒÎÓ Ó‰Ó‚ (ÏÓÒÍËı –Í‡Í Ì‡ ËÒ. 1,·, ÍÓÌÚËÌÂÌÚ‡Î¸Ì˚ı – Í‡Í Ì‡ ËÒ. 2,‰), Ô‡-‡ÏÂÚ˚ ÏÓ‰ÂÎÂÈ: ˝ÍÒÔÓÌÂÌÚ‡: Nt = 5187.6e0.0047t, R2 == 0.67; „ËÔÂ·ÓÎ‡: Nt = 434635/(30 – t), R2 = 0.95.

2*

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ÊÂÌËÂÏ” ËÎË ÔÂËÙÂË˜ÂÒÍÓÈ Ó·Î‡ÒÚ¸˛ ÏÓÒÍÓÈ·ËÓÚ˚. èÂ‚˚Â Ì‡ÁÂÏÌ˚Â ÊË‚ÓÚÌ˚Â Ë ‚˚Ò¯ËÂ‡ÒÚÂÌËfl Á‡ÒÂÎflÎË ÔÂËÏÛ˘ÂÒÚ‚ÂÌÌÓ ÔË·ÂÊÌ˚ÂÓ·Î‡ÒÚË, ÒÓı‡ÌflÎË ÚÂÒÌÛ˛ Ò‚flÁ¸ Ò ‚Ó‰ÌÓÈ ÒÂ‰ÓÈ Ë‚Ó ÏÌÓ„ÓÏ Á‡‚ËÒÂÎË ÓÚ ÏÓÒÍËı ˝ÍÓÒËÒÚÂÏ, ‚ ÚÓÏ˜ËÒÎÂ Ë ‚ ÓÚÌÓ¯ÂÌËË ÚÂÏÔÓ‚ Ò‚ÓÂÈ ‰Ë‚ÂÒËÙËÍ‡-ˆËË. èÓ-‚Ë‰ËÏÓÏÛ, ÚÓÎ¸ÍÓ Í ÍÓÌˆÛ ‰Â‚ÓÌ‡ – Ì‡˜‡ÎÛÍ‡·ÓÌ‡ ÒÎÓÊËÎËÒ¸ ‰ÓÒÚ‡ÚÓ˜ÌÓ ÒÎÓÊÌ˚Â Ë ÛÒÚÓÈ-˜Ë‚˚Â Ì‡ÁÂÏÌ˚Â ÒÓÓ·˘ÂÒÚ‚‡, ‚ ÚÓÏ ˜ËÒÎÂ Ì‡ÒÚÓfl-˘ËÂ ÎÂÒ‡, Ì‡ÒÂÎÂÌÌ˚Â ‡ÁÌÓÓ·‡ÁÌ˚ÏË ˜ÎÂÌËÒÚÓ-ÌÓ„ËÏË Ë ÚÂÚ‡ÔÓ‰‡ÏË. àÏÂÌÌÓ ‚ ̋ ÚÓ ‚ÂÏfl Ò„Î‡ÊË-‚‡ÂÚÒfl Ë ‡ÁÎË˜ËÂ ÏÂÊ‰Û ÏÓ‰ÂÎ¸Ì˚Ï Ë Â‡Î¸Ì˚Ï‡ÁÌÓÓ·‡ÁËÂÏ îäÅ Ì‡ ÛÓ‚ÌÂ ÒÂÏÂÈÒÚ‚. “çÂ‰ÓÒÚ‡-˛˘Û˛” Ù‡ÍˆË˛ ÒÂÏÂÈÒÚ‚ îäÅ ‚ ÒËÎÛÂ – ‡ÌÌÂÏÍ‡·ÓÌÂ, Ó˜Â‚Ë‰ÌÓ, ÒÎÂ‰ÛÂÚ ËÒÍ‡Ú¸ ÒÂ‰Ë ÏÓÒÍËı (‚ÔÂ‚Û˛ Ó˜ÂÂ‰¸ ÔË·ÂÊÌ˚ı Ë ÎËÚÓ‡Î¸Ì˚ı) Ó„‡-ÌËÁÏÓ‚. ëÓ„Î‡ÒÌÓ ÎÓ„ËÍÂ „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎË(ÒÏ. ÌËÊÂ), ÚÂÒÌ˚Â ˝ÍÓÎÓ„Ë˜ÂÒÍËÂ Ë ˝‚ÓÎ˛ˆËÓÌÌ˚ÂÒ‚flÁË ÏÂÊ‰Û ÏÓÒÍÓÈ ·ËÓÚÓÈ Ë ‰Â‚ÌËÏË Ó·ËÚ‡ÚÂ-ÎflÏË ÒÛ¯Ë ‰ÓÎÊÌ˚ ·˚ÎË ÓÍ‡Á˚‚‡Ú¸ ÒÛ˘ÂÒÚ‚ÂÌÌÓÂ‚ÎËflÌËÂ Ì‡ ‰Ë‚ÂÒËÙËÍ‡ˆË˛ îäÅ Ì‡ ‡ÌÌËı ˝Ú‡-Ô‡ı ÂÂ ÒÚ‡ÌÓ‚ÎÂÌËfl.

ÇÓ-‚ÚÓ˚ı, Ì‡ ‡ÌÌËı ˝Ú‡Ô‡ı ‡Á‚ËÚËfl ÍÓÌÚË-ÌÂÌÚ‡Î¸ÌÓÈ ·ËÓÚ˚ ËÒÍÎ˛˜ËÚÂÎ¸ÌÓ ‚‡ÊÌÛ˛ ÓÎ¸‰ÓÎÊÌ˚ ·˚ÎË Ë„‡Ú¸ Ó„‡ÌËÁÏ˚, Ô‡ÍÚË˜ÂÒÍË ÌÂÒÓı‡Ìfl˛˘ËÂÒfl ‚ ËÒÍÓÔ‡ÂÏÓÏ ÒÓÒÚÓflÌËË. êÂ˜¸Ë‰ÂÚ ÔÂÊ‰Â ‚ÒÂ„Ó Ó ÔÓ˜‚ÂÌÌ˚ı ·‡ÍÚÂËflı Ë „Ë-·‡ı, Ó‰ÌÓÍÎÂÚÓ˜Ì˚ı ‚Ó‰ÓÓÒÎflı, ÎË¯‡ÈÌËÍ‡ı, ÌÂ-ÍÓÚÓ˚ı ·ÂÒÔÓÁ‚ÓÌÓ˜Ì˚ı (ÌÂÏ‡ÚÓ‰‡ı, ÓÎË„ÓıÂÚ‡ı,ÏÂÎÍËı ̃ ÎÂÌËÒÚÓÌÓ„Ëı), ÍÓÚÓ˚Â Û˜‡ÒÚ‚Ó‚‡ÎË ‚ ÒÓ-Á‰‡ÌËË ÔÓ˜‚, ÔË„Ó‰Ì˚ı ‰Îfl ÓÒÚ‡ ‚˚Ò¯Ëı ‡ÒÚÂ-ÌËÈ, Ë ÚÂÏ Ò‡Ï˚Ï ÒÓÁ‰‡ÎË ÔÂ‰ÔÓÒ˚ÎÍË ‰Îfl ÛÒÍÓ-ÂÌÌÓ„Ó ÓÒÚ‡ ‡ÁÌÓÓ·‡ÁËfl îäÅ ‚ ÔÓÒÎÂ‰Û˛˘ËÂ˝ÔÓıË.

ç‡·Î˛‰‡ÂÏ˚Â ‡ÒıÓÊ‰ÂÌËfl ÏÂÊ‰Û ÏÓ‰ÂÎ¸ÌÓÈË Ì‡·Î˛‰‡ÂÏÓÈ (ÂÍÓÌÒÚÛËÓ‚‡ÌÌÓÈ) ‰ËÌ‡ÏËÍÓÈÓ‰Ó‚Ó„Ó ‡ÁÌÓÓ·‡ÁËfl îäÅ (ËÒ. 2,‰), ‚ÂÓflÚÌÓ,ÓÚ˜‡ÒÚË Ó·˙flÒÌfl˛ÚÒfl Ó·˙ÂÍÚË‚Ì˚ÏË Ù‡ÍÚÓ‡ÏË,‡ÒÒÏÓÚÂÌ˚ÏË ‚˚¯Â, ÓÚ˜‡ÒÚË – ÌÂÒÓ‚Â¯ÂÌÒÚ‚ÓÏËÒÔÓÎ¸ÁÓ‚‡ÌÌÓÈ ÏÂÚÓ‰ËÍË ÂÍÓÌÒÚÛÍˆËË Ó‰Ó‚Ó-„Ó ‡ÁÌÓÓ·‡ÁËfl îäÅ.

2. ÉËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ·ËÓ‡ÁÌÓÓ·‡ÁËfl –ÂÁÛÎ¸Ú‡Ú ÔÓ„ÂÒÒË‚ÌÓ„Ó ‡Á‚ËÚËfl ÒÓÓ·˘ÂÒÚ‚.çÂÍÓÚÓ˚Â ‚ÂÓflÚÌ˚Â ÏÂı‡ÌËÁÏ˚, Ó·ÂÒÔÂ˜Ë‚‡˛-

˘ËÂ „ËÔÂ·ÓÎË˜ÂÒÍËÈ ı‡‡ÍÚÂ ÓÒÚ‡ ‡ÁÌÓÓ·‡-ÁËfl îåÅ, ‡ÒÒÏÓÚÂÌ˚ Ì‡ÏË ‡ÌÂÂ (å‡ÍÓ‚, äÓ-ÓÚ‡Â‚, 2007; Markov, Korotayev, 2007). çÓ‚˚Â ‰‡Ì-Ì˚Â, ÔË‚Â‰ÂÌÌ˚Â ‚ Ì‡ÒÚÓfl˘ÂÈ ÒÚ‡Ú¸Â, ÔÓÁ‚ÓÎfl˛ÚÔÂ‰ÔÓÎÓÊËÚ¸, ˜ÚÓ ÓÒÚ ‡ÁÌÓÓ·‡ÁËfl îäÅ, ‡ Ú‡Í-ÊÂ ‚ÒÂÈ ·ËÓÚ˚ ‚ ˆÂÎÓÏ ÍÓÌÚÓÎËÓ‚‡ÎÒfl ÒıÓ‰Ì˚-ÏË ÏÂı‡ÌËÁÏ‡ÏË. ä‡Í ·˚ÎÓ ÔÓ‰ÂÏÓÌÒÚËÓ‚‡ÌÓ ‚ÛÍ‡Á‡ÌÌ˚ı ÒÚ‡Ú¸flı, ‰Îfl ËÌÚÂÔÂÚ‡ˆËË „ËÔÂ·ÓÎË-˜ÂÒÍÓ„Ó ÓÒÚ‡ ‡ÁÌÓÓ·‡ÁËfl ·ËÓÚ˚ ÏÓ„ÛÚ ·˚Ú¸ ÔÓ-ÎÂÁÌ˚ ‡Ì‡ÎÓ„ËË Ò ‰Û„ÓÈ ÒÎÓÊÌÓÈ, „ËÔÂ·ÓÎË˜Â-ÒÍË ‡ÒÚÛ˘ÂÈ ÒËÒÚÂÏÓÈ, ‡ ËÏÂÌÌÓ Ò ˜ÂÎÓ‚Â˜Â-ÒÚ‚ÓÏ. ÉËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ Ì‡ÒÂÎÂÌËfl áÂÏÎË ‚1–1958 „„. Ì.˝. ·˚Î ÓÚÍ˚Ú îÂÒÚÂÓÏ Ë Â„Ó ÍÓÎÎÂ-„‡ÏË (Foerster et al., 1960). Ç ‰‡Î¸ÌÂÈ¯ÂÏ ·˚ÎÓ ÔÓ-Í‡Á‡ÌÓ, ˜ÚÓ „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ Ì‡ÒÂÎÂÌËfl ÔÓ-ÒÎÂÊË‚‡ÂÚÒfl Ë ‚ ÚÂ˜ÂÌËÂ ÏÌÓ„Ëı Ú˚Òfl˜ ÎÂÚ ‰Ó Ì.˝.(ä‡ÔËˆ‡, 1992, 1999; Kremer, 1993). Å˚Î ÔÂ‰ÎÓÊÂÌfl‰ Ï‡ÚÂÏ‡ÚË˜ÂÒÍËı ÏÓ‰ÂÎÂÈ, ÓÔËÒ˚‚‡˛˘Ëı „Ë-ÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ Ì‡ÒÂÎÂÌËfl áÂÏÎË (äÓÓÚ‡Â‚ Ë‰., 2007‡,·; Kremer, 1993; Cohen, 1995; èÓ‰Î‡ÁÓ‚,2002; Podlazov, 2004; Tsirel, 2004; Korotayev et al.,2006, Ë ‰.). èÓÍ‡Á‡ÌÓ, ˜ÚÓ „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚfl‚ÎflÂÚÒfl ÒÎÂ‰ÒÚ‚ËÂÏ ÌÂÎËÌÂÈÌÓÈ ÔÓÎÓÊËÚÂÎ¸ÌÓÈÒ‚flÁË ‚ÚÓÓ„Ó ÔÓfl‰Í‡ ÏÂÊ‰Û ‰ÂÏÓ„‡ÙË˜ÂÒÍËÏ ËÚÂıÌÓÎÓ„Ë˜ÂÒÍËÏ ÓÒÚÓÏ: ·ÓÎ¸¯Â Î˛‰ÂÈ – ·ÓÎ¸¯ÂÔÓÚÂÌˆË‡Î¸Ì˚ı ËÁÓ·ÂÚ‡ÚÂÎÂÈ – ÛÒÍÓÂÌËÂ ÚÂıÌÓ-ÎÓ„Ë˜ÂÒÍÓ„Ó ÓÒÚ‡ – ÛÒÍÓÂÌËÂ ÓÒÚ‡ ÌÂÒÛ˘ÂÈ ÒÔÓ-ÒÓ·ÌÓÒÚË áÂÏÎË – ÛÒÍÓÂÌËÂ ÓÒÚ‡ Ì‡ÒÂÎÂÌËfl –·ÓÎ¸¯Â Î˛‰ÂÈ – ·ÓÎ¸¯Â ÔÓÚÂÌˆË‡Î¸Ì˚ı ËÁÓ·ÂÚ‡-ÚÂÎÂÈ Ë Ú.‰. (ËÒ. 4).

å˚ ÔÂ‰ÔÓÎ‡„‡ÂÏ, ˜ÚÓ „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ‡ÁÌÓÓ·‡ÁËfl îåÅ Ú‡ÍÊÂ Ó·˙flÒÌflÂÚÒfl ÌÂÎËÌÂÈ-ÌÓÈ ÔÓÎÓÊËÚÂÎ¸ÌÓÈ Ó·‡ÚÌÓÈ Ò‚flÁ¸˛ ‚ÚÓÓ„Ó ÔÓ-fl‰Í‡ ÏÂÊ‰Û ÓÒÚÓÏ ˜ËÒÎ‡ Ú‡ÍÒÓÌÓ‚ Ë ËÁÏÂÌÂÌËflÏËÒÚÛÍÚÛ˚ ÏÓÒÍËı ÒÓÓ·˘ÂÒÚ‚. ùÚ‡ Ò‚flÁ¸ ÏÓÊÂÚ Â-‡ÎËÁÓ‚˚‚‡Ú¸Òfl ‰‚ÓflÍÓ: ˜ÂÂÁ ÒÌËÊÂÌËÂ ‚ÂÓflÚÌÓ-ÒÚË ‚˚ÏË‡ÌËfl Ú‡ÍÒÓÌÓ‚ Ë ˜ÂÂÁ ÔÓ‚˚¯ÂÌËÂ ‚ÂÓ-flÚÌÓÒÚË Ëı ÔÓfl‚ÎÂÌËfl: ·ÓÎ¸¯Â Ó‰Ó‚ – ‚˚¯Â ‡Î¸Ù‡-‡ÁÌÓÓ·‡ÁËÂ (ÒÂ‰ÌÂÂ ˜ËÒÎÓ Ó‰Ó‚ ‚ ÒÓÓ·˘Â-ÒÚ‚Â) – ÒÓÓ·˘ÂÒÚ‚‡ ÒÚ‡ÌÓ‚flÚÒfl ·ÓÎÂÂ ÒÎÓÊÌ˚ÏË ËÛÒÚÓÈ˜Ë‚˚ÏË – ÒÌËÊ‡ÂÚÒfl ÒÍÓÓÒÚ¸ ‚˚ÏË‡ÌËfl Ë(ËÎË) ‡ÒÚÂÚ ÒÍÓÓÒÚ¸ ÔÓfl‚ÎÂÌËfl – ‡ÁÌÓÓ·‡ÁËÂ‡ÒÚÂÚ ·˚ÒÚÂÂ – ·ÓÎ¸¯Â Ó‰Ó‚ – ‚˚¯Â ‡Î¸Ù‡-‡Á-ÌÓÓ·‡ÁËÂ Ë Ú.‰. àÏÂ˛˘ËÂÒfl ‰‡ÌÌ˚Â ÔÓ ‰ËÌ‡ÏËÍÂ

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êËÒ. 4. ÅÎÓÍ-ÒıÂÏ‡ ÌÂÎËÌÂÈÌÓÈ ÔÓÎÓÊËÚÂÎ¸ÌÓÈ Ó·‡ÚÌÓÈ Ò‚flÁË ÏÂÊ‰Û ÚÂıÌÓÎÓ„Ë˜ÂÒÍËÏ Ë ‰ÂÏÓ„‡ÙË˜ÂÒÍËÏ ÓÒÚÓÏ, „Â-ÌÂËÛ˛˘ÂÈ „ËÔÂ·ÓÎË˜ÂÒÍÓÂ ‡Á‚ËÚËÂ åË-ëËÒÚÂÏ˚.



‡Î¸Ù‡-‡ÁÌÓÓ·‡ÁËfl ÏÓÒÍËı ÒÓÓ·˘ÂÒÚ‚ ‚ Ù‡ÌÂÓ-ÁÓÂ (Bambach, 1977; Sepkoski, 1988) ÒÓ„Î‡ÒÛ˛ÚÒfl Ò˝ÚËÏ ÔÂ‰ÔÓÎÓÊÂÌËÂÏ (å‡ÍÓ‚, äÓÓÚ‡Â‚, 2007).

êÓÒÚ Ó‰Ó‚Ó„Ó ‡ÁÌÓÓ·‡ÁËfl îåÅ ÔÓËÒıÓ‰ËÎ‚ ÁÌ‡˜ËÚÂÎ¸ÌÓÈ ÒÚÂÔÂÌË Á‡ Ò˜ÂÚ ÓÒÚ‡ ÒÂ‰ÌÂÈ ÔÓ-‰ÓÎÊËÚÂÎ¸ÌÓÒÚË ÒÛ˘ÂÒÚ‚Ó‚‡ÌËfl (èë) Ó‰Ó‚ Ë Ì‡-ÍÓÔÎÂÌËfl ‰ÓÎ„ÓÊË‚Û˘Ëı (ÛÒÚÓÈ˜Ë‚˚ı) Ó‰Ó‚ ‚·ËÓÚÂ (å‡ÍÓ‚, 2001, 2002). ëÂ‰Ìflfl èë ÒÂÏÂÈÒÚ‚ÚÓÊÂ ÌÂÛÍÎÓÌÌÓ ÓÒÎ‡ ‚ ÚÂ˜ÂÌËÂ Ù‡ÌÂÓÁÓfl Í‡Í ‚îåÅ, Ú‡Í Ë ‚ îäÅ (ËÒ. 5,‡). Ç ËÒÚÓËË îåÅ ·˚ÎÓÚË ÍÛÔÌÂÈ¯Ëı ÔÂÂÎÓÏÌ˚ı ˝Ú‡Ô‡ (Ó‰Ó‚ËÍÒÍ‡fl‡‰Ë‡ˆËfl Ë ‚˚ÏË‡ÌËfl Ì‡ Û·ÂÊ‡ı ÔÂÏË/ÚË‡Ò‡ ËÏÂÎ‡/Ô‡ÎÂÓ„ÂÌ‡), ÒÓÔflÊÂÌÌ˚ı ÒÓ ÒÚÛÔÂÌ˜‡Ú˚Ï Ó-ÒÚÓÏ ‡Î¸Ù‡-‡ÁÌÓÓ·‡ÁËfl Ë ÛÒÚÓÈ˜Ë‚ÓÒÚË ÏÓÒÍËıÒÓÓ·˘ÂÒÚ‚, ‡ Ú‡ÍÊÂ Ò ÛÒÎÓÊÌÂÌËÂÏ ÌË¯Â‚ÓÈ ÒÚÛÍ-ÚÛ˚ (Bambach, 1977; Sepkoski, 1988, 1994; Bambachet al., 2002; Bush, Bambach, 2004). êÓÒÚ ÒÎÓÊÌÓÒÚË ËÛÒÚÓÈ˜Ë‚ÓÒÚË ÒÓÓ·˘ÂÒÚ‚ ÏÓÊÂÚ ÒÔÓÒÓ·ÒÚ‚Ó‚‡Ú¸ÒÌËÊÂÌË˛ ‚ÂÓflÚÌÓÒÚË ‚˚ÏË‡ÌËfl ‚ıÓ‰fl˘Ëı ‚ ËıÒÓÒÚ‡‚ Ú‡ÍÒÓÌÓ‚ (å‡ÍÓ‚, äÓÓÚ‡Â‚, 2007).

ç‡ÍÓÔÎÂÌËÂ ‚ ·ËÓÚÂ ‰ÓÎ„ÓÊË‚Û˘Ëı, ÛÒÚÓÈ˜Ë-‚˚ı Ú‡ÍÒÓÌÓ‚ ÔÓËÒıÓ‰ËÚ ·Î‡„Ó‰‡fl ËÁ·Ë‡ÚÂÎ¸-ÌÓÏÛ ‚˚ÏË‡ÌË˛ (Sepkoski, 1991a), ‡ Ú‡ÍÊÂ ËÁ·Ë-‡ÚÂÎ¸ÌÓÏÛ ‚ÓÒÒÚ‡ÌÓ‚ÎÂÌË˛ ÔÓÒÎÂ ÍËÁËÒÓ‚ (Bam-bach et al., 2002). ïÓÚfl Ï‡ÒÒÓ‚˚Â ‚˚ÏË‡ÌËfl ÏÓ„ÛÚ·˚Ú¸ ÌÂËÁ·Ë‡ÚÂÎ¸Ì˚ÏË ‚ ÓÔÂ‰ÂÎÂÌÌ˚ı ‡ÒÔÂÍÚ‡ı(Jablonski, 2005), ÓÌË, ÌÂÒÓÏÌÂÌÌÓ, ‚˚ÒÓÍÓ ËÁ·Ë‡-ÚÂÎ¸Ì˚ ‚ ÓÚÌÓ¯ÂÌËË ÒÔÓÒÓ·ÌÓÒÚË Ú‡ÍÒÓÌÓ‚ ÔÂÂ-ÌÓÒËÚ¸ ÂÁÍËÂ ÌÂÔÂ‰ÒÍ‡ÁÛÂÏ˚Â ËÁÏÂÌÂÌËfl ÒÂ‰˚.ùÚÓ ‚Ë‰ÌÓ, ‚ ˜‡ÒÚÌÓÒÚË, ÔÓ ı‡‡ÍÚÂÛ Á‡‚ËÒËÏÓÒÚËÛÓ‚Ìfl ‚˚ÏË‡ÌËfl ÓÚ ‚ÓÁ‡ÒÚ‡ ÍÓ„ÓÚ˚ ‚Ó ‚ÂÏflÏ‡ÒÒÓ‚˚ı ‚˚ÏË‡ÌËÈ (ËÒ. 5,·,‚). óÂÏ ÒÚ‡¯Â ÍÓ-„ÓÚ‡ Í ÏÓÏÂÌÚÛ ÍËÁËÒ‡, ÚÂÏ ÏÂÌ¸¯ËÈ ÔÓˆÂÌÚÚ‡ÍÒÓÌÓ‚ ËÁ ˝ÚÓÈ ÍÓ„ÓÚ˚ ‚˚ÏË‡ÂÚ Ì‡ ÍËÁËÒÌÓÏÛ·ÂÊÂ. Ñ‡Ì‡fl Á‡‚ËÒËÏÓÒÚ¸ ÒÔ‡‚Â‰ÎË‚‡ Í‡Í ‰Îfl

îåÅ (ËÒ. 5,·), Ú‡Í Ë ‰Îfl îäÅ (ËÒ. 5,‚). íÓ˜ÌÓ Ú‡-Í‡fl ÊÂ Í‡ÚËÌ‡ Ì‡·Î˛‰‡ÂÚÒfl Ë ‚Ó ‚ÂÏfl ÔÂËÓ‰Ó‚“ÙÓÌÓ‚Ó„Ó” ‚˚ÏË‡ÌËfl (å‡ÍÓ‚, 2000). ùÚÓ ÓÁÌ‡-˜‡ÂÚ, ˜ÚÓ Ú‡ÍÒÓÌ˚ ‡ÁÎË˜‡˛ÚÒfl ÔÓ Ò‚ÓÂÈ ÛÒÚÓÈ˜Ë-‚ÓÒÚË (ÒÔÓÒÓ·ÌÓÒÚË ÔÓÚË‚ÓÒÚÓflÚ¸ ‚˚ÏË‡ÌË˛), ËÒ ÚÂ˜ÂÌËÂÏ ‚ÂÏÂÌË ‚ Í‡Ê‰ÓÈ ÍÓ„ÓÚÂ ÌÂÛÒÚÓÈ˜Ë-‚˚Â Ú‡ÍÒÓÌ˚ ÓÚÒÂË‚‡˛ÚÒfl, ‡ ÛÒÚÓÈ˜Ë‚˚Â Ì‡Í‡ÔÎË-‚‡˛ÚÒfl. ëÓÓ·˘ÂÒÚ‚‡, ÒÓÒÚ‡‚ÎÂÌÌ˚Â ËÁ ·ÓÎÂÂ ÛÒÚÓÈ-˜Ë‚˚ı Ú‡ÍÒÓÌÓ‚, ‚ Ò‚Ó˛ Ó˜ÂÂ‰¸ ÚÓÊÂ ÔËÓ·ÂÚ‡˛ÚÔÓ‚˚¯ÂÌÌÛ˛ ÛÒÚÓÈ˜Ë‚ÓÒÚ¸. ì·Â‰ËÚÂÎ¸ÌÓÂ ÚÂÓÂ-ÚË˜ÂÒÍÓÂ Ó·ÓÒÌÓ‚‡ÌËÂ ÚÓ„Ó, ˜ÚÓ ÌÂËÁ·ÂÊÌ˚Ï Â-ÁÛÎ¸Ú‡ÚÓÏ ˝‚ÓÎ˛ˆËË ‰ÓÎÊÌÓ ·˚Ú¸ ÂÂ Á‡ÏÂ‰ÎÂÌËÂ

êËÒ. 5. ç‡ÍÓÔÎÂÌËÂ ‰ÓÎ„ÓÊË‚Û˘Ëı ÒÂÏÂÈÒÚ‚ Ë Ó‰Ó‚ ‚Ù‡ÌÂÓÁÓÈÒÍÓÈ ·ËÓÚÂ. ‡ – ÒÂ‰Ìflfl ÔÓ‰ÓÎÊËÚÂÎ¸ÌÓÒÚ¸ÒÛ˘ÂÒÚ‚Ó‚‡ÌËfl (èë) ÒÂÏÂÈÒÚ‚ (ÔÓ ‰‡ÌÌ˚Ï The FossilRecord 2). èÓ ‚ÂÚËÍ‡Î¸ÌÓÈ ÓÒË – ÒÂ‰Ìflfl èë ÒÂÏÂÈÒÚ‚,ÒÛ˘ÂÒÚ‚Ó‚‡‚¯Ëı ‚ ‰‡ÌÌÓÏ flÛÒÂ. èÓ „ÓËÁÓÌÚ‡Î¸ÌÓÈÓÒË – ‚ÂÏfl ‚ ÏÎÌ. ÎÂÚ. íÓÌÍ‡fl ÎËÌËfl – ÏÓÒÍ‡fl ·ËÓÚ‡,ÚÓÎÒÚ‡fl – ÍÓÌÚËÌÂÌÚ‡Î¸Ì‡fl ·ËÓÚ‡. ÑÎfl ÌÂ‚˚ÏÂ¯Ëı ÒÂ-ÏÂÈÒÚ‚ ‚ÏÂÒÚÓ Â‡Î¸ÌÓÈ èë ËÒÔÓÎ¸ÁÓ‚‡Î‡Ò¸ ÓÊË‰‡ÂÏ‡fl(life expectancy), ‡ÒÒ˜ËÚ‡ÌÌ‡fl Ì‡ ÓÒÌÓ‚Â ËÁ‚ÂÒÚÌ˚ı ÚÂÏ-ÔÓ‚ ‚˚ÏË‡ÌËfl ‚ Í‡ÈÌÓÁÓÂ, ËÒıÓ‰fl ËÁ ‰ÓÔÛ˘ÂÌËfl, ̃ ÚÓ Ú‡-ÍËÂ ÊÂ ÚÂÏÔ˚ ‚˚ÏË‡ÌËfl ÒÓı‡ÌflÚÒfl Ë ‚ÔÂ‰¸ (ÔÓ‰Ó·-ÌÂÂ Ó ÏÂÚÓ‰ËÍÂ ‚˚˜ËÒÎÂÌËfl èë ÒÏ.: å‡ÍÓ‚, 2002). · –ËÁ·Ë‡ÚÂÎ¸ÌÓÒÚ¸ ‚˚ÏË‡ÌËfl Ó‰Ó‚ ÏÓÒÍËı Ó„‡ÌËÁÏÓ‚Ì‡ Û·ÂÊÂ ÏÂÁÓÁÓfl Ë Í‡ÈÌÓÁÓfl ‚ Á‡‚ËÒËÏÓÒÚË ÓÚ ‚ÓÁ‡Ò-Ú‡ Ó‰Ó‚ (îåÅ, ÔÓ ·‡ÁÂ ‰‡ÌÌ˚ı ëÂÔÍÓÒÍË). ì˜ÚÂÌ˚ Ó-‰˚, ‰ÓÊË‚¯ËÂ ‰Ó Û·ÂÊ‡ Ë ÎË·Ó ‚˚ÏÂ¯ËÂ Ì‡ ÌÂÏ, ÎË·ÓÔÂÂ¯Â‰¯ËÂ ‚ Í‡ÈÌÓÁÓÈ. èÓ „ÓËÁÓÌÚ‡Î¸ÌÓÈ ÓÒË – ‚Â-Ïfl ÔÓfl‚ÎÂÌËfl Ó‰Ó‚. 1 – ˜ËÒÎÓ Ó‰Ó‚ ‰‡ÌÌÓÈ ÍÓ„ÓÚ˚,ÔÂÂ¯Â‰¯Ëı ‚ Í‡ÈÌÓÁÓÈ, 2 – ˜ËÒÎÓ Ó‰Ó‚ ‰‡ÌÌÓÈ ÍÓ„Ó-Ú˚, ‚˚ÏÂ¯Ëı Ì‡ Û·ÂÊÂ ÏÂÁÓÁÓfl Ë Í‡ÈÌÓÁÓfl, 3 – ÛÓ-‚ÂÌ¸ ‚˚ÏË‡ÌËfl ‚ ÍÓ„ÓÚÂ, % (å‡ÍÓ‚, 2000). ‚ – ÚÓ ÊÂ‰Îfl ÒÂÏÂÈÒÚ‚ ÍÓÌÚËÌÂÌÚ‡Î¸Ì˚ı Ó„‡ÌËÁÏÓ‚ Ì‡ Û·ÂÊÂÔ‡ÎÂÓÁÓfl Ë ÏÂÁÓÁÓfl (îäÅ, ÔÓ The Fossil Record 2). 1 –˜ËÒÎÓ ÒÂÏÂÈÒÚ‚ ‰‡ÌÌÓÈ ÍÓ„ÓÚ˚, ÔÂÂ¯Â‰¯Ëı ‚ ÏÂÁÓ-ÁÓÈ, 2 – ˜ËÒÎÓ ÒÂÏÂÈÒÚ‚ ‰‡ÌÌÓÈ ÍÓ„ÓÚ˚, ‚˚ÏÂ¯Ëı Ì‡Û·ÂÊÂ ËÎË ‚ Ò‡ÏÓÏ Ì‡˜‡ÎÂ ÚË‡Ò‡ (‚ Ú‡Ú‡ÒÍÓÏ ËÎËÒÍËÙÒÍÓÏ ‚ÂÍÂ), 3 – ÛÓ‚ÂÌ¸ ‚˚ÏË‡ÌËfl ‚ ÍÓ„ÓÚÂ, %.

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ìÓ‚ÂÌ¸ ‚˚ÏË‡ÌËfl, %70

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14012010080604020

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–250 –50

600

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500

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200

100

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182



(Ú.Â. Û‚ÂÎË˜ÂÌËÂ ÒÂ‰ÌÂÈ ÔÓ‰ÓÎÊËÚÂÎ¸ÌÓÒÚË ÒÛ˘Â-ÒÚ‚Ó‚‡ÌËfl Ú‡ÍÒÓÌÓ‚), ·˚ÎÓ ‰‡ÌÓ ÌÂ‰‡‚ÌÓ Ç.è. ôÂ-·‡ÍÓ‚˚Ï (2005).

Ç‡ÊÌÛ˛ ÓÎ¸ ‚ ÔÓˆÂÒÒÂ ÓÒÚ‡ ÛÒÚÓÈ˜Ë‚ÓÒÚËÚ‡ÍÒÓÌÓ‚ Ë ÒÓÓ·˘ÂÒÚ‚ Ë„‡ÂÚ ÔÓ˝Ú‡ÔÌ‡fl ÒÏÂÌ‡ ‰Ó-ÏËÌËÛ˛˘Ëı „ÛÔÔ Ó„‡ÌËÁÏÓ‚, ÓÚ‡ÊÂÌÌ‡fl ‚ ÒÎÛ-˜‡Â îåÅ ‚ ÍÓÌˆÂÔˆËË ÚÂı ÔÓÒÎÂ‰Ó‚‡ÚÂÎ¸Ì˚ı ̋ ‚Ó-Î˛ˆËÓÌÌ˚ı Ù‡ÛÌ (ÍÂÏ·ËÈÒÍÓÈ, Ô‡ÎÂÓÁÓÈÒÍÓÈ ËÒÓ‚ÂÏÂÌÌÓÈ, ËÎË ÏÂÁÓ-Í‡ÈÌÓÁÓÈÒÍÓÈ) (Sepkoski,1981, 1991b) (ËÒ. 6,‡). èÓÍ‡Á‡ÌÓ, ˜ÚÓ ‰Îfl Í‡Ê‰ÓÈÔÓÒÎÂ‰Û˛˘ÂÈ Ù‡ÛÌ˚ ı‡‡ÍÚÂÂÌ ·ÓÎ¸¯ËÈ Ó·˙ÂÏ

Á‡ÌËÏ‡ÂÏÓ„Ó ˝ÍÓÎÓ„Ë˜ÂÒÍÓ„Ó ÔÓÒÚ‡ÌÒÚ‚‡, ·ÓÎ¸-¯ÂÂ ˜ËÒÎÓ Ë ‡ÁÌÓÓ·‡ÁËÂ ÓÒ‚ÓÂÌÌ˚ı ÌË¯ (Bam-bach, 1983, 1985; Sepkoski, 1991b).

Ç ıÓ‰Â ˝ÚÓÈ ÒÏÂÌ˚ ‚ ‚˚Ë„˚¯Â Ô‡ÍÚË˜ÂÒÍË‚ÒÂ„‰‡ ÓÍ‡Á˚‚‡ÎËÒ¸ „ÛÔÔ˚ Ò ·ÓÎÂÂ ÌËÁÍËÏ ÚÂÏÔÓÏÓ·ÌÓ‚ÎÂÌËfl Ó‰Ó‚Ó„Ó Ë ÒÂÏÂÈÒÚ‚ÂÌÌÓ„Ó ÒÓÒÚ‡‚‡,Ú.Â. Ò ·ÓÎÂÂ ÛÒÚÓÈ˜Ë‚˚ÏË Ó‰‡ÏË Ë ÒÂÏÂÈÒÚ‚‡ÏË(Valentine et al., 1991; å‡ÍÓ‚, 2002). íËÔË˜Ì˚ÂÔËÏÂ˚ – ÔÓ˝Ú‡ÔÌ‡fl ÒÏÂÌ‡ ‰ÓÏËÌËÓ‚‡ÌËfl ‚ Ú‡-ÍËı Ô‡‡ı ̋ ÍÓÎÓ„Ë˜ÂÒÍË ÒıÓ‰Ì˚ı „ÛÔÔ, Í‡Í ·‡ıËÓ-ÔÓ‰˚ Ë ‰‚ÛÒÚ‚ÓÍË, Ì‡ÛÊÌÓ‡ÍÓ‚ËÌÌ˚Â „ÓÎÓ‚ÓÌÓ-

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3500

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400

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PteridophytaGymnospermophytaMagnoliophyta

0

800‚ 700

600

500

400

300

200

100

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BrachiopodaBivalvia


0–200

‰

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200

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50

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Amphibia 250

ReptiliaMammalia + Aves

êËÒ. 6. ëÏÂÌ‡ ‰ÓÏËÌËÛ˛˘Ëı „ÛÔÔ ‚ ËÒÚÓËË Ù‡ÌÂÓÁÓÈÒÍÓÈ ·ËÓÚ˚. èÓ „ÓËÁÓÌÚ‡Î¸ÌÓÈ ÓÒË – ‚ÂÏfl ‚ ÏÎÌ. ÎÂÚ. ‡ – ‰Ë-Ì‡ÏËÍ‡ ˜ËÒÎ‡ Ó‰Ó‚ ‚ “ÚÂı ˝‚ÓÎ˛ˆËÓÌÌ˚ı Ù‡ÛÌ‡ı” îåÅ (Sepkoski, 1981, 1991b). É‡ÙËÍ ÔÓÒÚÓÂÌ ÔË ÔÓÏÓ˘Ë Sepkos-ki’s Online Genus Database (http://strata.ummp.lsa.umich.edu/jack/) Ì‡ ÓÒÌÓ‚Â flÛÒÌÓÈ ̄ Í‡Î˚. · – ÒÏÂÌ‡ ‰ÓÏËÌËÛ˛˘Ëı „ÛÔÔ‡ÍÚË‚ÌÓ ÔÎ‡‚‡˛˘Ëı ÍÛÔÌ˚ı ıË˘ÌËÍÓ‚ ‚ îåÅ: „ÓÎÓ‚ÓÌÓ„ËÂ Ë ıÓ‰Ó‚˚Â. ‚ – ÒÏÂÌ‡ ‰ÓÏËÌËÛ˛˘Ëı „ÛÔÔ ÌÂÔÓ‰‚ËÊÌ˚ıË Ï‡ÎÓÔÓ‰‚ËÊÌ˚ı ÙËÎ¸Ú‡ÚÓÓ‚ Ò ‰‚ÛÒÚ‚Ó˜‡ÚÓÈ ‡ÍÓ‚ËÌÓÈ ‚ îåÅ: ·‡ıËÓÔÓ‰˚ Ë ‰‚ÛÒÚ‚ÓÍË. „ – ÒÏÂÌ‡ ‰ÓÏËÌËÛ˛˘Ëı„ÛÔÔ Ì‡ÁÂÏÌ˚ı ‡ÒÚÂÌËÈ: ÒÔÓÓ‚˚Â, „ÓÎÓÒÂÏÂÌÌ˚Â, ÔÓÍ˚ÚÓÒÂÏÂÌÌ˚Â. ‰ – ÒÏÂÌ‡ ‰ÓÏËÌËÛ˛˘Ëı „ÛÔÔ ÚÂÚ‡ÔÓ‰: ‡ÏÙË-·ËË, ÂÔÚËÎËË, ÏÎÂÍÓÔËÚ‡˛˘ËÂ + ÔÚËˆ˚.



„ËÂ Ë ÏÓÒÍËÂ ıÓ‰Ó‚˚Â (ËÒ. 6,·,‚). ÑÎfl ·‡ıËÓÔÓ‰Ë Ì‡ÛÊÌÓ‡ÍÓ‚ËÌÌ˚ı „ÓÎÓ‚ÓÌÓ„Ëı ı‡‡ÍÚÂÂÌÁÌ‡˜ËÚÂÎ¸ÌÓ ·ÓÎÂÂ ‚˚ÒÓÍËÈ ÚÂÏÔ Ó·ÌÓ‚ÎÂÌËfl Ó‰Ó-‚Ó„Ó ÒÓÒÚ‡‚‡, ˜ÂÏ ‰Îfl ‰‚ÛÒÚ‚ÓÓÍ Ë ˚·; ‚ ÚÂ˜ÂÌËÂÙ‡ÌÂÓÁÓfl ÒÚÂÔÂÌ¸ ‰ÓÏËÌËÓ‚‡ÌËfl ÔÂ‚˚ı ‰‚Ûı„ÛÔÔ ÒÌËÊ‡Î‡Ò¸, ‚ÚÓ˚ı – ÓÒÎ‡.

ÑÎfl îåÅ ÔÓÍ‡Á‡Ì Ú‡ÍÊÂ ÒÚÛÔÂÌ˜‡Ú˚È ÓÒÚ ÓÚ-ÌÓ¯ÂÌËfl ̃ ËÒÎ‡ Ó‰Ó‚ ÔÓ‰‚ËÊÌ˚ı Ó„‡ÌËÁÏÓ‚ Í ÌÂ-ÔÓ‰‚ËÊÌ˚Ï, “ÙËÁËÓÎÓ„Ë˜ÂÒÍË Á‡·ÛÙÂÂÌÌ˚ı” ÍÌÂÁ‡·ÛÙÂÂÌÌ˚Ï Ë ıË˘ÌËÍÓ‚ Í ÊÂÚ‚‡Ï (Bambachet al., 2002). ùÚË ËÁÏÂÌÂÌËfl Ú‡ÍÊÂ ‰ÓÎÊÌ˚ ·˚ÎËÒÔÓÒÓ·ÒÚ‚Ó‚‡Ú¸ ÓÒÚÛ ÛÒÚÓÈ˜Ë‚ÓÒÚË ÏÓÒÍËı ÒÓÓ·-˘ÂÒÚ‚ ‰‡ÊÂ ·ÂÁ ËÁÏÂÌÂÌËÈ ‡Î¸Ù‡-‡ÁÌÓÓ·‡ÁËfl.ç‡ÔËÏÂ, ‰Ë‚ÂÒËÙËÍ‡ˆËfl ıË˘ÌËÍÓ‚ ÔÂ‰ÔÓÎ‡-„‡ÂÚ, ̃ ÚÓ ıË˘ÌËÍË ÒÚ‡ÌÓ‚flÚÒfl ·ÓÎÂÂ ÒÔÂˆË‡ÎËÁËÓ-‚‡ÌÌ˚ÏË ‚ ÓÚÌÓ¯ÂÌËË ‚˚·Ó‡ ÊÂÚ‚; Ú‡ÍËÂ ıË˘-ÌËÍË fl‚Îfl˛ÚÒfl ·ÓÎÂÂ ˝ÙÙÂÍÚË‚Ì˚ÏË Â„ÛÎflÚÓ‡-ÏË ˜ËÒÎÂÌÌÓÒÚË ÊÂÚ‚.

èÓˆÂÒÒ˚, ‡Ì‡ÎÓ„Ë˜Ì˚Â ÒÏÂÌÂ ˝‚ÓÎ˛ˆËÓÌÌ˚ıÙ‡ÛÌ ‚ îåÅ (ËÒ. 6,‡), ̃ ÂÚÍÓ ÔÓÒÎÂÊË‚‡˛ÚÒfl Ú‡Í-ÊÂ Ë ‚ îäÅ. ëÏÂÌ‡ ‰ÓÏËÌËÛ˛˘Ëı „ÛÔÔ Û Ì‡ÁÂÏ-Ì˚ı ‡ÒÚÂÌËÈ Ë ÚÂÚ‡ÔÓ‰ ÔÓËÒıÓ‰ËÎ‡ ÚÓÊÂ ‚ ÚË˝Ú‡Ô‡: 1) ÒÔÓÓ‚˚Â, ‡ÏÙË·ËË; 2) „ÓÎÓÒÂÏÂÌÌ˚Â,ÂÔÚËÎËË; 3) ÔÓÍ˚ÚÓÒÂÏÂÌÌ˚Â, ÏÎÂÍÓÔËÚ‡˛˘ËÂË ÔÚËˆ˚ (ËÒ. 6,„,‰). çÂÚÛ‰ÌÓ Á‡ÏÂÚËÚ¸, ˜ÚÓ Ë Û‡ÒÚÂÌËÈ, Ë Û ÚÂÚ‡ÔÓ‰ ‚ fl‰Û ÒÏÂÌfl˛˘Ëı ‰Û„ ‰Û-„‡ ‰ÓÏËÌËÛ˛˘Ëı Ú‡ÍÒÓÌÓ‚ Ì‡·Î˛‰‡ÂÚÒfl ÓÒÚ “Á‡-·ÛÙÂÂÌÌÓÒÚË” (ÌÂÁ‡‚ËÒËÏÓÒÚ¸ ‡ÌÌËı ÒÚ‡‰ËÈ ÓÚ‚Ó‰ÌÓÈ ÒÂ‰˚ Ë Ëı Á‡˘Ë˘ÂÌÌÓÒÚ¸, ÚÂÔÎÓÍÓ‚ÌÓÒÚ¸Ë ‰.) Ë “ÔÓ‰‚ËÊÌÓÒÚË” (·ÓÎÂÂ ˝ÙÙÂÍÚË‚Ì˚Â ÏÂı‡-ÌËÁÏ˚ ‡ÒÒÂÎÂÌËfl, ‡ÍÚË‚ÌÓÒÚ¸ ÔÂÂ‰‚ËÊÂÌËfl Ë‰.). çÂÒÓÏÌÂÌÌÓ Ú‡ÍÊÂ, ˜ÚÓ ‚ ˝ÚÓÏ fl‰Û, Í‡Í Ë ‚îåÅ, ÔÓËÒıÓ‰ËÎ ÓÒÚ Ó·˙ÂÏ‡ Á‡ÌËÏ‡ÂÏÓ„Ó ˝ÍÓ-ÎÓ„Ë˜ÂÒÍÓ„Ó ÔÓÒÚ‡ÌÒÚ‚‡, ‡ÁÌÓÓ·‡ÁËfl ˝ÍÓÎÓ„Ë-˜ÂÒÍËı ÒÚ‡ÚÂ„ËÈ Ë Ó·‡ÁÓ‚ ÊËÁÌË. è‡‡ÎÎÂÎËÁÏ ‚˝‚ÓÎ˛ˆËË Ì‡ÁÂÏÌ˚ı ‡ÒÚÂÌËÈ Ë ÚÂÚ‡ÔÓ‰ ÔÓfl‚Îfl-ÂÚÒfl ÌÂ ÚÓÎ¸ÍÓ ‚ ‰ËÌ‡ÏËÍÂ ‡ÁÌÓÓ·‡ÁËfl (ÒıÓ‰ÒÚ‚Ó„‡ÙËÍÓ‚ Ì‡ ËÒ. 6,„ Ë 6,‰), ÌÓ Ë Ì‡ ÏÓÙÓÙËÁËÓÎÓ-„Ë˜ÂÒÍÓÏ ÛÓ‚ÌÂ (‡Á‚ËÚËÂ ‡ÏÌËÓÌ‡ Ë ÔÎ‡ˆÂÌÚ˚ ÛÚÂÚ‡ÔÓ‰, ˝Ì‰ÓÒÔÂÏ‡ Ë Á‡‚flÁË Û ‡ÒÚÂÌËÈ) Ë ‰‡ÊÂÌ‡ ÏÓÎÂÍÛÎflÌÓ-„ÂÌÂÚË˜ÂÒÍÓÏ (Ô‡‡ÎÎÂÎ¸ÌÓÂ ‡Á-‚ËÚËÂ ÒıÓ‰Ì˚ı ÏÂı‡ÌËÁÏÓ‚ „ÂÌÓÏÌÓ„Ó ËÏÔËÌÚËÌ-„‡ Ì‡ ·‡ÁÂ ‰ËÙÙÂÂÌˆËÓ‚‡ÌÌÓ„Ó ÏÂÚËÎËÓ‚‡ÌËflÓ‰ËÚÂÎ¸ÒÍËı „ÂÌÓ‚ Û ÏÎÂÍÓÔËÚ‡˛˘Ëı Ë ÔÓÍ˚ÚÓ-ÒÂÏÂÌÌ˚ı).

êÓÒÚ ÛÒÚÓÈ˜Ë‚ÓÒÚË Ú‡ÍÒÓÌÓ‚ ÌËÁ¯Â„Ó ‡Ì„‡ (‚Ë-‰Ó‚, Ó‰Ó‚, ÒÂÏÂÈÒÚ‚) ‚ fl‰Û ÒÏÂÌfl˛˘Ëı ‰Û„ ‰Û„‡‰ÓÏËÌËÛ˛˘Ëı „ÛÔÔ Ì‡ÁÂÏÌ˚ı Ó„‡ÌËÁÏÓ‚ ‚îäÅ ÚÓÊÂ ÔÓÒÎÂÊË‚‡ÂÚÒfl, ıÓÚfl Ë ÌÂ ÒÚÓÎ¸ ˜ÂÚÍÓ,Í‡Í ‚ ÏÓÒÍÓÈ ·ËÓÚÂ: Ì‡ÔËÏÂ, Û Ì‡ÁÂÏÌ˚ı ‡ÒÚÂ-ÌËÈ Ì‡ ‚Ë‰Ó‚ÓÏ ÛÓ‚ÌÂ ‚˚fl‚ÎÂÌ˚ ‰‡ÊÂ ÔÓÚË‚ÓÔÓ-ÎÓÊÌ˚Â ÚÂÌ‰ÂÌˆËË (Valentine et al., 1991). çÂÛÍÎÓÌ-Ì˚È ÓÒÚ ÒÂ‰ÌÂÈ èë ÒÂÏÂÈÒÚ‚ (ËÒ. 5,‡), Ì‡·Î˛‰‡-ÂÏ˚È Í‡Í Û ÏÓÒÍËı, Ú‡Í Ë Û ÍÓÌÚËÌÂÌÚ‡Î¸Ì˚ıÓ„‡ÌËÁÏÓ‚, Ò‚Ë‰ÂÚÂÎ¸ÒÚ‚ÛÂÚ Ó ÚÓÏ, ˜ÚÓ ÓÒÚÛÒÚÓÈ˜Ë‚ÓÒÚË Ú‡ÍÒÓÌÓ‚ ‚ ıÓ‰Â ˝‚ÓÎ˛ˆËË fl‚ÎflÂÚÒflÓ·˘ËÏ Ô‡‚ËÎÓÏ.

ëÎÂ‰ÛÂÚ ÓÚÏÂÚËÚ¸, ˜ÚÓ ‚Ó ‚ÒÂı ÛÔÓÏflÌÛÚ˚ı ÒÎÛ-˜‡flı ÔÓ˝Ú‡ÔÌÓÈ ÒÏÂÌ˚ ‰ÓÏËÌËÛ˛˘Ëı „ÛÔÔ ÂÁ-ÍËÈ ÓÒÚ ‚ Í‡ÈÌÓÁÓÂ Ë ‚ ÍÓÌˆÂ ÏÂÁÓÁÓfl Ì‡·Î˛‰‡ÂÚÒflÚÓÎ¸ÍÓ Û Ó‰ÌÓÈ ÔÓÒÎÂ‰ÌÂÈ, Ì‡Ë·ÓÎÂÂ “ÔÓ‰‚ËÌÛÚÓÈ”(‚ ÙËÎÓ„ÂÌÂÚË˜ÂÒÍÓÏ ËÎË ˝ÍÓÎÓ„Ë˜ÂÒÍÓÏ ÓÚÌÓ¯Â-ÌËË) „ÛÔÔËÓ‚ÍË, ‡ ËÏÂÌÌÓ: ‚ îåÅ – Û ÏÂÁÓ-Í‡ÈÌÓ-ÁÓÈÒÍÓÈ ̋ ‚ÓÎ˛ˆËÓÌÌÓÈ Ù‡ÛÌ˚, Û ÔÓ‰‚ËÊÌ˚ı Ë “ÙË-ÁËÓÎÓ„Ë˜ÂÒÍË Á‡·ÛÙÂÂÌÌ˚ı” ÏÓÒÍËı Ó„‡ÌËÁÏÓ‚(‚ ÓÚÎË˜ËÂ ÓÚ ÌÂÔÓ‰‚ËÊÌ˚ı Ë ÌÂÁ‡·ÛÙÂÂÌÌ˚ı),ÒÂ‰Ë ÙËÎ¸Ú‡ÚÓÓ‚ Ò ‰‚ÛÒÚ‚Ó˜‡ÚÓÈ ‡ÍÓ‚ËÌÓÈ –Û ‰‚ÛÒÚ‚ÓÓÍ (ÌÓ ÌÂ Û ·‡ıËÓÔÓ‰), ÒÂ‰Ë ÍÛÔÌ˚ı‡ÍÚË‚ÌÓ ÔÎ‡‚‡˛˘Ëı ÏÓÒÍËı ıË˘ÌËÍÓ‚ – Û ıÓ‰Ó-‚˚ı (ÌÓ ÌÂ Û „ÓÎÓ‚ÓÌÓ„Ëı), ÒÂ‰Ë Ì‡ÁÂÏÌ˚ı ‡ÒÚÂ-ÌËÈ – Û ÔÓÍ˚ÚÓÒÂÏÂÌÌ˚ı (ÌÓ ÌÂ Û „ÓÎÓÒÂÏÂÌÌ˚ı ËÒÔÓÓ‚˚ı), ÒÂ‰Ë ÚÂÚ‡ÔÓ‰ – Û ÔÚËˆ Ë ÏÎÂÍÓÔËÚ‡˛-˘Ëı (ÌÓ ÌÂ Û ÂÔÚËÎËÈ Ë ‡ÏÙË·ËÈ).

èÂÂ˜ËÒÎÂÌÌ˚Â ÚÂÌ‰ÂÌˆËË ËÎÎ˛ÒÚËÛ˛Ú ÔÂ-‚˚È ËÁ ‰‚Ûı ‚ÓÁÏÓÊÌ˚ı ÏÂı‡ÌËÁÏÓ‚ ÔÓÎÓÊËÚÂÎ¸-ÌÓÈ Ó·‡ÚÌÓÈ Ò‚flÁË ‚ÚÓÓ„Ó ÔÓfl‰Í‡, ÓÒÌÓ‚‡ÌÌ˚ÈÌ‡ ÒÌËÊÂÌËË ÚÂÏÔÓ‚ ‚˚ÏË‡ÌËfl. ÇÚÓÓÈ ‚ÓÁÏÓÊ-Ì˚È ÏÂı‡ÌËÁÏ ÓÒÌÓ‚‡Ì Ì‡ ÚÓÏ, ˜ÚÓ ÔÓ„ÂÒÒË‚ÌÓÂ‡Á‚ËÚËÂ ÏÓÒÍËı ÒÓÓ·˘ÂÒÚ‚ ÒÔÓÒÓ·ÒÚ‚ÛÂÚ Ú‡ÍÊÂ ËÓÒÚÛ ‚ÂÓflÚÌÓÒÚË ÔÓfl‚ÎÂÌËfl ÌÓ‚˚ı Ú‡ÍÒÓÌÓ‚. àÁ-‚ÂÒÚÌÓ, ˜ÚÓ Ó‰ÌËÏ ËÁ ‚‡ÊÌÂÈ¯Ëı Ù‡ÍÚÓÓ‚, ÒÔÓ-ÒÓ·ÒÚ‚Û˛˘Ëı ‚˚ÒÓÍÓÏÛ ·ËÓ‡ÁÌÓÓ·‡ÁË˛, fl‚Îfl-ÂÚÒfl „ÂÚÂÓ„ÂÌÌÓÒÚ¸ ÛÒÎÓ‚ËÈ Ó·ËÚ‡ÌËfl, ÔÓfl‚Îfl˛-˘‡flÒfl Ì‡ ‡ÁÌ˚ı ÛÓ‚Ìflı Ë ‚ ‡ÁÌÓÏ Ï‡Ò¯Ú‡·Â ‚ÔÓÒÚ‡ÌÒÚ‚Â Ë ‚ÂÏÂÌË (Huston, 1994). ç‡ÔËÏÂ,ÛÒËÎÂÌËÂ ̄ ËÓÚÌÓ„Ó ÍÎËÏ‡ÚË˜ÂÒÍÓ„Ó „‡‰ËÂÌÚ‡ ‚ ÚÂ-˜ÂÌËÂ Í‡ÈÌÓÁÓfl Ò˜ËÚ‡ÂÚÒfl Ó‰ÌÓÈ ËÁ ‚‡ÊÌ˚ı ÔË˜ËÌÒÚÂÏËÚÂÎ¸ÌÓ„Ó Í‡ÈÌÓÁÓÈÒÍÓ„Ó ÓÒÚ‡ ·ËÓ‡ÁÌÓÓ·‡-ÁËfl (Clarke, Crame, 2003). é˜Â‚Ë‰ÌÓ, ˜ÚÓ ÓÒÚÛ ·ËÓ-‡ÁÌÓÓ·‡ÁËfl ‰ÓÎÊÌ‡ ÒÔÓÒÓ·ÒÚ‚Ó‚‡Ú¸ „ÂÚÂÓ„ÂÌ-ÌÓÒÚ¸ ÌÂ ÚÓÎ¸ÍÓ ‡·ËÓÚË˜ÂÒÍÓÈ, ÌÓ Ë ·ËÓÚË˜ÂÒÍÓÈ ÒÂ-‰˚ Ó·ËÚ‡ÌËfl. í‡ÍËÏ Ó·‡ÁÓÏ, ÏÓÊÌÓ ÓÊË‰‡Ú¸, ˜ÚÓ·ËÓ‡ÁÌÓÓ·‡ÁËÂ ‚ÓÓ·˘Â Ë ‡Î¸Ù‡-‡ÁÌÓÓ·‡ÁËÂ ‚˜‡ÒÚÌÓÒÚË Ò‡ÏÓ ÔÓ ÒÂ·Â ‰ÓÎÊÌÓ ·˚Ú¸ Ù‡ÍÚÓÓÏ,ÒÔÓÒÓ·ÒÚ‚Û˛˘ËÏ ‰‡Î¸ÌÂÈ¯ÂÏÛ ÓÒÚÛ ·ËÓ‡ÁÌÓÓ·-‡ÁËfl. èË ˝ÚÓÏ ÌÂÓ·ıÓ‰ËÏÓ Û˜ËÚ˚‚‡Ú¸, ˜ÚÓ ‰‡ÊÂ‚ ÒÓ‚ÂÏÂÌÌÛ˛ ˝ÔÓıÛ, ÍÓ„‰‡ ‡ÁÌÓÓ·‡ÁËÂ ÊËÁÌËÌ‡ ÔÎ‡ÌÂÚÂ, ÔÓ-‚Ë‰ËÏÓÏÛ, ·ÂÒÔÂˆÂ‰ÂÌÚÌÓ ‚ÂÎËÍÓ,ÏÌÓ„ËÂ ÒÓÓ·˘ÂÒÚ‚‡, ÓÒÓ·ÂÌÌÓ ‚ ÛÏÂÂÌÌ˚ı Ë ‚˚ÒÓ-ÍËı ¯ËÓÚ‡ı, Â˘Â Ó˜ÂÌ¸ ‰‡ÎÂÍË ÓÚ Ì‡Ò˚˘ÂÌËfl Ë‡‚ÌÓ‚ÂÒËfl, Ë ÓÒÚ ‡ÁÌÓÓ·‡ÁËfl ‚ ÌËı ÏÓ„ ·˚ ÔÓ-‰ÓÎÊ‡Ú¸Òfl Â˘Â ‰ÓÎ„Ó, ÂÒÎË ‡Á‚ËÚËÂ ˜ÂÎÓ‚Â˜ÂÒÚ‚‡ÌÂ ‚ÓÒÔÂÔflÚÒÚ‚ÛÂÚ ˝ÚÓÏÛ (Clarke, Crame, 2003).

êÂÁÛÎ¸Ú‡Ú˚ ‡Ì‡ÎËÁ‡ ‰ËÌ‡ÏËÍË ÍÓÎË˜ÂÒÚ‚ÂÌÌÓ-„Ó ‡ÒÔÂ‰ÂÎÂÌËfl ‚Ë‰Ó‚˚ı Ó·ËÎËÈ ‚ ÏÓÒÍËı ÒÓÓ·-˘ÂÒÚ‚‡ı ‚ ÚÂ˜ÂÌËÂ Ù‡ÌÂÓÁÓfl (Wagner et al., 2006)„Ó‚ÓflÚ Ó ÚÓÏ, ˜ÚÓ ‚ ÏÂÁÓ-Í‡ÈÌÓÁÓÈÒÍËı ÒÓÓ·˘Â-ÒÚ‚‡ı ÔÓ Ò‡‚ÌÂÌË˛ Ò Ô‡ÎÂÓÁÓÈÒÍËÏË ÔÓfl‚ÎÂÌËÂÌÓ‚˚ı ‚Ë‰Ó‚, ÔÓ-‚Ë‰ËÏÓÏÛ, ÔË‚Ó‰ËÎÓ Í ·ÓÎÂÂ ÒÛ-˘ÂÒÚ‚ÂÌÌÓÏÛ ‡Ò¯ËÂÌË˛ ‰ÓÒÚÛÔÌÓ„Ó ˝ÍÓÎÓ„Ë˜Â-ÒÍÓ„Ó ÔÓÒÚ‡ÌÒÚ‚‡ Ë ÔÓ‚˚¯‡ÎÓ ‚ÂÓflÚÌÓÒÚ¸ ‚ÌÂ‰-ÂÌËfl ‚ ÒÓÓ·˘ÂÒÚ‚Ó ‰ÓÔÓÎÌËÚÂÎ¸Ì˚ı ‚Ë‰Ó‚ (Wag-ner et al., 2006; Solé et al., 2002; Laland et al., 1999). Ç˝ÚÓÏ ÒÎÛ˜‡Â ÒıÓ‰ÒÚ‚Ó ÏÂı‡ÌËÁÏÓ‚, ÓÚ‚ÂÚÒÚ‚ÂÌÌ˚ıÁ‡ „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ‡ÁÌÓÓ·‡ÁËfl îåÅ Ë Ì‡-ÒÂÎÂÌËfl ÏË‡, ÒÚ‡ÌÓ‚ËÚÒfl Â˘Â ·ÓÎÂÂ ÔÓÎÌ˚Ï, ÔÓ-

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ÒÍÓÎ¸ÍÛ ÒÛÏÏ‡ÌÓÂ ˝ÍÓÎÓ„Ë˜ÂÒÍÓÂ ÔÓÒÚ‡ÌÒÚ‚ÓîåÅ ‡Ì‡ÎÓ„Ë˜ÌÓ ÌÂÒÛ˘ÂÈ ÒÔÓÒÓ·ÌÓÒÚË áÂÏÎË ‚‰ÂÏÓ„‡ÙËË. ÖÒÎË ‚Ë‰˚ ÒÔÓÒÓ·Ì˚ ‡Ò¯ËflÚ¸ ‰Ó-ÒÚÛÔÌÓÂ ˝ÍÓÎÓ„Ë˜ÂÒÍÓÂ ÔÓÒÚ‡ÌÒÚ‚Ó ÒÓÓ·˘ÂÒÚ‚‡,ÒÓÁ‰‡‚‡fl ÌÓ‚˚Â ÌË¯Ë, ÚÓ Ëı ÏÓÊÌÓ ÛÔÓ‰Ó·ËÚ¸“ËÁÓ·ÂÚ‡ÚÂÎflÏ” ËÁ ÒÓˆË‡Î¸ÌÓ-‰ÂÏÓ„‡ÙË˜ÂÒÍËıÏÓ‰ÂÎÂÈ, ˜¸Ë ËÁÓ·ÂÚÂÌËfl Û‚ÂÎË˜Ë‚‡˛Ú ÌÂÒÛ˘Û˛ÒÔÓÒÓ·ÌÓÒÚ¸ áÂÏÎË.

ëÎÂ‰ÛÂÚ ÓÚÏÂÚËÚ¸, ˜ÚÓ ÔÂ‰Î‡„‡ÂÏ˚Â Ì‡ÏË ËÌ-ÚÂÔÂÚ‡ˆËË ÓÒÚ‡ ·ËÓ‡ÁÌÓÓ·‡ÁËfl Ë Ó·˙flÒÌÂÌËfl„ËÔÂ·ÓÎË˜ÂÒÍÓ„Ó ı‡‡ÍÚÂ‡ ˝ÚÓ„Ó ÓÒÚ‡, ÌÂÒÓ-ÏÌÂÌÌÓ, ÌÂ fl‚Îfl˛ÚÒfl ÌË Â‰ËÌÒÚ‚ÂÌÌÓ ‚ÓÁÏÓÊÌ˚ÏË,ÌË ËÒ˜ÂÔ˚‚‡˛˘ËÏË. í‡Í, ÌÂ‰‡‚ÌÓ û.É. èÛÁ‡˜ÂÌ-ÍÓ (2006) Ó·‡ÚËÎ ‚ÌËÏ‡ÌËÂ Ì‡ ÔËÒÛÚÒÚ‚ËÂ ‚ Ù‡ÌÂ-ÓÁÓÈÒÍÓÈ ‰ËÌ‡ÏËÍÂ ‡ÁÌÓÓ·‡ÁËfl (Ì‡ ÛÓ‚ÌÂ ÒÂ-ÏÂÈÒÚ‚, ÔÓ ‰‡ÌÌ˚Ï The Fossil Record 2) “„ËÔÂ„ÂÓ-ÏÂÚË˜ÂÒÍÓ„Ó ÚÂÌ‰‡”, ‰Îfl ÙÓÏ‡ÎËÁ‡ˆËË ÍÓÚÓÓ„ÓÓÌ ÔÂ‰ÎÓÊËÎ ÏÓ‰ÂÎË, ÓÒÌÓ‚‡ÌÌ˚Â Ì‡ ÚÂÓËË ËÌ-ÙÓÏ‡ˆËË, „‰Â ‡ÁÌÓÓ·‡ÁËÂ ‡ÒÒÏ‡ÚË‚‡ÂÚÒfl Í‡ÍÙÛÌÍˆËfl Ì‡ÍÓÔÎÂÌËfl ËÌÙÓÏ‡ˆËË ˝‚ÓÎ˛ˆËÓÌËÛ-˛˘ÂÈ ÒËÒÚÂÏÓÈ. ÑÎfl ·ÓÎÂÂ „ÎÛ·ÓÍÓ„Ó ÔÓÌËÏ‡ÌËflÏÂı‡ÌËÁÏÓ‚ ˝‚ÓÎ˛ˆËË ·ËÓ‡ÁÌÓÓ·‡ÁËfl ÔÓÚÂ·Û-ÂÚÒfl ÒËÌÚÂÁ ‡ÁÎË˜Ì˚ı ÔÓ‰ıÓ‰Ó‚, ÒÂ‰Ë ÍÓÚÓ˚ıÔÓÏËÏÓ ‚˚¯ÂÛÔÓÏflÌÛÚ˚ı ËÌÙÓÏ‡ˆËÓÌÌ˚ı ÓÚÏÂ-ÚËÏ Ú‡ÍÊÂ ÔÓ‰ıÓ‰˚, ‡Á‚Ë‚‡˛˘ËÂÒfl ‚ ÔÓÒÎÂ‰ÌËÂ„Ó‰˚ ‚ ˝ÍÓÎÓ„ËË Ë ÒÓˆËÓÎÓ„ËË ‚ Ò‚flÁË Ò ÍÓÌˆÂÔˆË-flÏË “ÛÔÛ„ÓÒÚË” (resilience), “‡‰‡ÔÚË‚ÌÓ„Ó ˆËÍÎ‡” Ë“Ô‡Ì‡ıËË” (Holling, Gunderson, 2002). èÓ‰Ó·Ì˚ÂÔÓ‰ıÓ‰˚, ‡ÒÒÏ‡ÚË‚‡˛˘ËÂÒfl ÌÂÍÓÚÓ˚ÏË ‡‚ÚÓ-‡ÏË Í‡Í ÔÓÎÂÁÌÓÂ ‰ÓÔÓÎÌÂÌËÂ Í ËÒÔÓÎ¸ÁÛÂÏÓÏÛÌ‡ÏË ‚ ‰‡ÌÌÓÈ ‡·ÓÚÂ “ÏË-ÒËÒÚÂÏÌÓÏÛ” ÔÓ‰ıÓ‰Û(Gotts, 2007), ÔÓÁ‚ÓÎfl˛Ú ‡ÒÒÏ‡ÚË‚‡Ú¸ „ÎÓ·‡Î¸-ÌÛ˛ ˝‚ÓÎ˛ˆËÓÌÌÛ˛ Ï‡ÍÓ‰ËÌ‡ÏËÍÛ Í‡Í ËÌÚÂ-„‡Î¸Ì˚È ÂÁÛÎ¸Ú‡Ú ÒÓÔflÊÂÌÌÓÈ ˝‚ÓÎ˛ˆËË ÏÌÓ-ÊÂÒÚ‚‡ ‚ÎÓÊÂÌÌ˚ı ‰Û„ ‚ ‰Û„‡ ÒËÒÚÂÏ ‡ÁÌ˚ıÛÓ‚ÌÂÈ (‚Ë‰Ó‚ Ë ·ÓÎÂÂ ÍÛÔÌ˚ı Ú‡ÍÒÓÌÓ‚, ˝ÍÓÒË-ÒÚÂÏ, ·ËÓÏÓ‚, Â„ËÓÌ‡Î¸Ì˚ı ·ËÓÚË˜ÂÒÍËı ÍÓÏÔÎÂÍ-ÒÓ‚ Ë Ú.‰.). ùÚÓ ÏÓÊÂÚ ÓÍ‡Á‡Ú¸Òfl ÓÒÓ·ÂÌÌÓ ÔÓÎÂÁÌ˚ÏÔË ‡ÒÒÏÓÚÂÌËË ˆËÍÎË˜ÂÒÍËı Ë ÒÚÓı‡ÒÚË˜ÂÒÍËıÒÓÒÚ‡‚Îfl˛˘Ëı ‰ËÌ‡ÏËÍË ·ËÓ‡ÁÌÓÓ·‡ÁËfl.

3. èÓ·ÎÂÏ‡ ‚˚ıÓ‰‡ ÒËÒÚÂÏ ËÁ ÂÊËÏÓ‚ ÒÓ·ÓÒÚÂÌËÂÏ. ÉËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ‡ÁÌÓÓ·‡-ÁËfl ËÎË Ì‡Ó‰ÓÌ‡ÒÂÎÂÌËfl ÌÂ ÏÓÊÂÚ ÔÓ‰ÓÎÊ‡Ú¸Òfl‚Â˜ÌÓ, Ú‡Í Í‡Í „ËÔÂ·ÓÎË˜ÂÒÍË ‡ÒÚÛ˘ËÈ ÔÓÍ‡Á‡-ÚÂÎ¸ ‚ ÓÔÂ‰ÂÎÂÌÌ˚È ÏÓÏÂÌÚ ‚ÂÏÂÌË ‰ÓÎÊÂÌÒÚ‡Ú¸ ·ÂÒÍÓÌÂ˜ÌÓ ·ÓÎ¸¯ËÏ. äÓ„‰‡ îÂÒÚÂ Ë Â„ÓÍÓÎÎÂ„Ë ‰‡‚‡ÎË Ò‚ÓÂÈ ÒÚ‡Ú¸Â (Foerster et al., 1960)ÂÂ ÁÌ‡ÏÂÌËÚÓÂ Ì‡Á‚‡ÌËÂ – “ëÛ‰Ì˚È ‰ÂÌ¸: ÔflÚÌËˆ‡,13 ÌÓfl·fl 2026 „.”, ÓÌË ‚Ó‚ÒÂ ÌÂ ËÏÂÎË ‚ ‚Ë‰Û, ˜ÚÓÌ‡ÒÂÎÂÌËÂ áÂÏÎË ‚ ̋ ÚÓÚ ‰ÂÌ¸ ‰ÂÈÒÚ‚ËÚÂÎ¸ÌÓ ÏÓÊÂÚÒÚ‡Ú¸ ·ÂÒÍÓÌÂ˜Ì˚Ï, ıÓÚfl “˝Òı‡ÚÓÎÓ„Ë˜ÂÒÍËÂ” ‚˚-‚Ó‰˚ ËÁ „ËÔÂ·ÓÎË˜ÂÒÍËı ÏÓ‰ÂÎÂÈ ÓÒÚ‡ åË-ëË-ÒÚÂÏ˚ ·ÂÁ Í‡ÍËı-ÎË·Ó ‰ÓÒÚ‡ÚÓ˜Ì˚ı Ì‡ ÚÓ ÓÒÌÓ‚‡-ÌËÈ ‰Ó ÒËı ÔÓ ‰ÂÎ‡˛ÚÒfl (ÒÏ., Ì‡ÔËÏÂ, Johansen,Sornette, 2001). àÁ ̋ ÚÓÈ ÒÚ‡Ú¸Ë ÒÍÓÂÂ ‚˚ÚÂÍ‡Î Ôfl-ÏÓ ÔÓÚË‚ÓÔÓÎÓÊÌ˚È ÔÓ„ÌÓÁ – Ì‡·Î˛‰‡‚¯ËÈÒfl‚ÔÎÓÚ¸ ‰Ó 1960 „. Ì‡ ÔÓÚflÊÂÌËË ÏÌÓ„Ëı ‚ÂÍÓ‚ „Ë-ÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ÏËÓ‚Ó„Ó Ì‡ÒÂÎÂÌËfl ‰ÓÎÊÂÌËÒÔ˚Ú‡Ú¸ ‚ Ò‡Ï˚Â ·ÎËÊ‡È¯ËÂ „Ó‰˚ ‡‰ËÍ‡Î¸ÌÛ˛

Ú‡ÌÒÙÓÏ‡ˆË˛ Ë ÒÏÂÌËÚ¸Òfl Ì‡ ÔËÌˆËÔË‡Î¸ÌÓËÌÓÈ ÚËÔ ‰ÂÏÓ„‡ÙË˜ÂÒÍÓÈ Ï‡ÍÓ‰ËÌ‡ÏËÍË. ùÚÓÚÔÓ„ÌÓÁ ÒÚ‡Î ·ÎÂÒÚfl˘Â ÓÔ‡‚‰˚‚‡Ú¸Òfl ÛÊÂ ˜ÂÂÁÌÂÒÍÓÎ¸ÍÓ ÎÂÚ ÔÓÒÎÂ ÔÛ·ÎËÍ‡ˆËË ÒÚ‡Ú¸Ë îÂÒÚÂ‡Ë Â„Ó ÍÓÎÎÂ„ (ÒÏ., Ì‡ÔËÏÂ, äÓÓÚ‡Â‚ Ë ‰., 2007·,Ò. 14–22). åË-ëËÒÚÂÏ‡ Ì‡˜‡Î‡ Ò‚ÓÈ ‚˚ıÓ‰ ËÁ Â-ÊËÏ‡ Ò Ó·ÓÒÚÂÌËÂÏ. èË ˝ÚÓÏ ‚˚‚Ó‰ËÚ¸ ÂÂ ËÁ Â-ÊËÏ‡ Ò Ó·ÓÒÚÂÌËÂÏ ÒÚ‡Î‡ Ú‡ ÊÂ Ò‡Ï‡fl ÒËÎ‡, ÍÓÚÓ-‡fl ÂÂ ‚ ˝ÚÓÚ ÂÊËÏ Ë ‚‚ÂÎ‡ – ÛÒÍÓfl˛˘ËÈÒfl ÚÂı-ÌÓÎÓ„Ë˜ÂÒÍËÈ ÓÒÚ. ùÚÓÚ ÓÒÚ (‚ÍÎ˛˜‡˛˘ËÈ ‚ÒÂ·fl Ë ‡Á‚ËÚËÂ ÏÂ‰ËˆËÌÒÍËı ÚÂıÌÓÎÓ„ËÈ) ‚ÔÎÓÚ¸‰Ó ‰ÓÒÚËÊÂÌËfl ÚÓ˜ÍË ÔÂÂ„Ë·‡ ‚ ÍË‚ÓÈ ‰ÂÏÓ„‡-ÙË˜ÂÒÍÓ„Ó ÔÂÂıÓ‰‡ ÓÍ‡Á˚‚‡Î Ò‚ÓÂ Â¯‡˛˘ÂÂ‚ÓÁ‰ÂÈÒÚ‚ËÂ Ì‡ Ï‡ÍÓ‰ÂÏÓ„‡ÙË˜ÂÒÍÛ˛ ‰ËÌ‡ÏËÍÛ˜ÂÂÁ ÒÌËÊÂÌËÂ ÒÏÂÚÌÓÒÚË. ÇÏÂÒÚÂ Ò ÚÂÏ ‰‡Î¸ÌÂÈ-¯ËÈ ÚÂıÌÓÎÓ„Ë˜ÂÒÍËÈ (Ë Ó·ÛÒÎÓ‚ÎÂÌÌ˚È ËÏ ̋ ÍÓÌÓ-ÏË˜ÂÒÍËÈ) ÓÒÚ ‚Â‰ÂÚ Í ÛÒËÎÂÌË˛ ‰ÂÈÒÚ‚Ëfl Ú‡ÍËıÏÓ˘Ì˚ı Ù‡ÍÚÓÓ‚ ÒÌËÊÂÌËfl ÓÊ‰‡ÂÏÓÒÚË, Í‡Í„‡ÏÓÚÌÓÒÚ¸ (‚ ÓÒÓ·ÂÌÌÓÒÚË ÊÂÌÒÍ‡fl), ‡Á‚ËÚËÂ ÒË-ÒÚÂÏ˚ ÒÓˆË‡Î¸ÌÓ„Ó Ó·ÂÒÔÂ˜ÂÌËfl, ÒÓ‚Â¯ÂÌÒÚ‚Ó‚‡-ÌËÂ ÒÂ‰ÒÚ‚ Ë ÏÂÚÓ‰Ó‚ ÔÎ‡ÌËÓ‚‡ÌËfl ÒÂÏ¸Ë, Ëı ‚ÒÂ·ÓÎ¸¯‡fl ‰ÓÒÚÛÔÌÓÒÚ¸ Ë Ú.Ô. ëÌËÊÂÌËÂ ÒÏÂÚÌÓÒÚË‚ÒÍÓÂ ‰ÓÒÚË„‡ÂÚ ÔÓÓ„Ó‚˚ı ÁÌ‡˜ÂÌËÈ, ÔÓÒÎÂ ˜Â„ÓÒÏÂÚÌÓÒÚ¸ ‰‡ÊÂ Ì‡˜ËÌ‡ÂÚ ÌÂÒÍÓÎ¸ÍÓ ‡ÒÚË (ËÁ-Á‡ÒÚ‡ÂÌËfl Ì‡ÒÂÎÂÌËfl), Ë ‚ ‰‡Î¸ÌÂÈ¯ÂÏ Ò‚ÓÂ „Î‡‚ÌÓÂ‚ÓÁ‰ÂÈÒÚ‚ËÂ Ì‡ ÏËÓ‚Û˛ ‰ÂÏÓ„‡ÙË˜ÂÒÍÛ˛ ‰ËÌ‡-ÏËÍÛ ÚÂıÌÓÎÓ„Ë˜ÂÒÍËÈ ÓÒÚ ÓÍ‡Á˚‚‡ÂÚ ÛÊÂ ËÏÂÌÌÓ˜ÂÂÁ ÒÌËÊÂÌËÂ ÓÊ‰‡ÂÏÓÒÚË, ˜ÚÓ ‚Â‰ÂÚ ÍÓ ‚ÒÂ·ÓÎ¸¯ÂÏÛ Á‡ÏÂ‰ÎÂÌË˛ ÚÂÏÔÓ‚ ÓÒÚ‡ ˜ËÒÎÂÌÌÓÒÚËÌ‡ÒÂÎÂÌËfl ÏË‡ Ë, Ú‡ÍËÏ Ó·‡ÁÓÏ, ‚˚‚Ó‰ËÚ åË-ëËÒÚÂÏÛ ËÁ ÂÊËÏ‡ Ò Ó·ÓÒÚÂÌËÂÏ. ë ‰Û„ÓÈ ÒÚÓÓ-Ì˚, ÓÒÚ Ú‡ÍËı ÔÓÍ‡Á‡ÚÂÎÂÈ ÛÓ‚Ìfl ‡Á‚ËÚËfl åË-ëËÒÚÂÏ˚ Í‡Í ÏËÓ‚‡fl „‡ÏÓÚÌÓÒÚ¸ ËÎË ÏËÓ‚‡flÛ·‡ÌËÁ‡ˆËfl Ì‡˜ËÌ‡ÂÚ Á‡ÏÂ‰ÎflÚ¸Òfl ÔÓÒÚÓ ‚ Ò‚flÁËÒ ÔË·ÎËÊÂÌËÂÏ Í ÛÓ‚Ì˛ Ì‡Ò˚˘ÂÌËfl (‚Â‰¸, ÒÍ‡-ÊÂÏ, ÔÓÔÓˆËfl „‡ÏÓÚÌ˚ı ‚ Ó·˘ÂÈ ˜ËÒÎÂÌÌÓÒÚË‚ÁÓÒÎÓ„Ó Ì‡ÒÂÎÂÌËfl ÏË‡ ÔÓÒÚÓ ÔÓ ÓÔÂ‰ÂÎÂÌË˛ÌÂ ÏÓÊÂÚ ÔÂ‚˚¯‡Ú¸ 100%). ÅÓÎÂÂ ÔÓ‰Ó·ÌÓÂ ÓÔË-Ò‡ÌËÂ ˝ÚËı ÔÓˆÂÒÒÓ‚ Ò Ó·ÓÒÌÓ‚‡ÌËÂÏ Ï‡ÚÂÏ‡ÚË˜Â-ÒÍÓÈ ÏÓ‰ÂÎË ËÁ ÚÂı ‰ËÙÙÂÂÌˆË‡Î¸Ì˚ı Û‡‚ÌÂ-ÌËÈ, ÓÔËÒ˚‚‡˛˘ÂÈ Í‡Í ‡Á‚ËÚËÂ åË-ëËÒÚÂÏ˚ ‚ÂÊËÏÂ Ò Ó·ÓÒÚÂÌËÂÏ, Ú‡Í Ë ÂÂ ‚˚ıÓ‰ ËÁ ˝ÚÓ„Ó Â-ÊËÏ‡, ‰‡ÂÚÒfl ‚ ÏÓÌÓ„‡ÙËflı (äÓÓÚ‡Â‚ Ë ‰.,2007‡,·).

çÂÓ·ıÓ‰ËÏÓ ÔÓ‰˜ÂÍÌÛÚ¸, ˜ÚÓ Ì˚ÌÂ¯ÌÂÂ Ô‡‰Â-ÌËÂ ÚÂÏÔÓ‚ ÏË-ÒËÒÚÂÏÌÓ„Ó ÓÒÚ‡ ÍÓÂÌÌ˚Ï Ó·‡-ÁÓÏ ÓÚÎË˜‡ÂÚÒfl ÓÚ ÒÔ‡‰Ó‚ Ë ÍÓÎÂ·‡ÌËÈ ÔÓ¯ÎÓ„Ó.ùÚÓ ÌÂ Ó˜ÂÂ‰ÌÓÂ ÍÓÎÂ·‡ÌËÂ, ̋ ÚÓ Ù‡ÁÓ‚˚È ÔÂÂıÓ‰Ì‡ ÌÓ‚˚È, ÌÂ ÚËÔË˜Ì˚È ‰Îfl ‚ÒÂÈ ÔÂÊÌÂÈ ËÒÚÓËË,ÂÊËÏ ‡Á‚ËÚËfl. ÖÒÎË ‚ÒÂ ÔÂ‰˚‰Û˘ËÂ ÒÔ‡‰˚ ÚÂÏ-ÔÓ‚ ÓÒÚ‡ ˜ËÒÎÂÌÌÓÒÚË Ì‡ÒÂÎÂÌËfl ÏË‡ ÔÓËÒıÓ‰Ë-ÎË Ì‡ ÙÓÌÂ Í‡Ú‡ÒÚÓÙË˜ÂÒÍÓ„Ó Ô‡‰ÂÌËfl ÛÓ‚ÌflÊËÁÌË Ì‡ÒÂÎÂÌËfl Ë ·˚ÎË ‚˚Á‚‡Ì˚ ÔÂÊ‰Â ‚ÒÂ„ÓÛ‚ÂÎË˜ÂÌËÂÏ ÒÏÂÚÌÓÒÚË ‚ÒÎÂ‰ÒÚ‚ËÂ ‡ÁÎË˜Ì˚ıÍ‡Ú‡ÍÎËÁÏÓ‚ (‚ÓÈÌ˚, „ÓÎÓ‰, ̋ ÔË‰ÂÏËË), ÔÓ ÏÂÂ Á‡-‚Â¯ÂÌËfl ̋ ÚËı ·Â‰ÒÚ‚ËÈ ̃ ÂÎÓ‚Â˜ÂÒÚ‚Ó ÓÚÌÓÒËÚÂÎ¸-ÌÓ ·˚ÒÚÓ ‚ÓÒÒÚ‡Ì‡‚ÎË‚‡ÎÓÒ¸ Ë ‚˚ıÓ‰ËÎÓ Ì‡ ÔÂÊ-Ì˛˛ Ú‡ÂÍÚÓË˛, ÚÓ Ì˚ÌÂ¯ÌËÈ ÒÔ‡‰ ÔÓËÒıÓ‰ËÚÌ‡ ÙÓÌÂ ˝ÍÓÌÓÏË˜ÂÒÍÓ„Ó ÔÓ‰˙ÂÏ‡ Ë ‚˚Á‚‡Ì Í‡˜Â-



ÒÚ‚ÂÌÌÓ ÓÚÎË˜Ì˚ÏË ÔË˜ËÌ‡ÏË: ÂÁÍËÏ ÒÌËÊÂÌËÂÏÓÊ‰‡ÂÏÓÒÚË, ÔÓËÒıÓ‰fl˘ËÏ Í‡Í ‡Á ËÁ-Á‡ ÓÒÚ‡ÛÓ‚Ìfl ÊËÁÌË ÓÒÌÓ‚ÌÓÈ Ï‡ÒÒ˚ Ì‡ÒÂÎÂÌËfl åË-ëË-ÒÚÂÏ˚ Ë ‚˚Á‚‡ÌÌÓ„Ó ˝ÚËÏ ÓÒÚÓÏ ÛÓ‚Ìfl Ó·‡ÁÓ-‚‡ÌÌÓÒÚË, Ó·ÂÒÔÂ˜ÂÌÌÓÒÚË ÏÂ‰ËˆËÌÒÍËÏ Ó·ÒÎÛÊË-‚‡ÌËÂÏ (‚ÍÎ˛˜‡fl ÏÂÚÓ‰˚ Ë ÒÂ‰ÒÚ‚‡ ÔÎ‡ÌËÓ‚‡ÌËflÒÂÏ¸Ë), ÒÓˆË‡Î¸Ì˚Ï ÒÚ‡ıÓ‚‡ÌËÂÏ Ë Ú.Ô. ëÌËÊÂÌËÂÚÂÏÔÓ‚ ÓÒÚ‡ „‡ÏÓÚÌÓÒÚË Ë Û·‡ÌËÁËÓ‚‡ÌÌÓÒÚËÚ‡ÍÊÂ ÌÂÂ‰ÍÓ Ì‡·Î˛‰‡ÎÓÒ¸ ‚ ÔÂ‰¯ÂÒÚ‚Û˛˘ËÂ˝ÔÓıË, ÌÓ ÚÓ„‰‡ ÓÌÓ ·˚ÎÓ Ò‚flÁ‡ÌÓ Ò ÌÂı‚‡ÚÍÓÈ ˝ÍÓ-ÌÓÏË˜ÂÒÍËı ÂÒÛÒÓ‚, ‡ ÒÂÈ˜‡Ò ˝ÚÓ Ì‡·Î˛‰‡ÂÚÒfl Ì‡ÙÓÌÂ ‚˚ÒÓ˜‡È¯Ëı ÚÂÏÔÓ‚ ˝ÍÓÌÓÏË˜ÂÒÍÓ„Ó ÓÒÚ‡ ËÒ‚flÁ‡ÌÓ Ò ‚˚ıÓ‰ÓÏ Ì‡ ÛÓ‚ÂÌ¸ Ì‡Ò˚˘ÂÌËfl.

ç‡ ·ÓÎ¸¯ËÌÒÚ‚Â ÔË‚Â‰ÂÌÌ˚ı „‡ÙËÍÓ‚ ‰ËÌ‡-ÏËÍË ‡ÁÌÓÓ·‡ÁËfl ‚ Ô‡‚ÓÈ ˜‡ÒÚË, ÒÓÓÚ‚ÂÚÒÚ‚Û˛-˘ÂÈ Í‡ÈÌÓÁÓ˛ Ë ÓÒÓ·ÂÌÌÓ Â„Ó ‚ÚÓÓÈ ÔÓÎÓ‚ËÌÂ(ÔÓÒÎÂ‰ÌËÂ 30 ÏÎÌ. ÎÂÚ) Á‡ÏÂÚÌÓ ÌÂÍÓÚÓÓÂ ‚˚ÔÓ-Î‡ÊË‚‡ÌËÂ, ÓÚ‡Ê‡˛˘ÂÂ Á‡ÏÂ‰ÎÂÌËÂ ÓÒÚ‡ ‡ÁÌÓ-Ó·‡ÁËfl. ùÚÓ Á‡ÏÂ‰ÎÂÌËÂ Ì‡Ë·ÓÎÂÂ Á‡ÏÂÚÌÓ Ì‡ÛÓ‚ÌÂ ÒÂÏÂÈÒÚ‚ (ËÒ. 1,‡, 2,‡,·, 3,‡). Ö„Ó ÏÓÊÌÓ ËÌ-ÚÂÔÂÚËÓ‚‡Ú¸ ‰‚ÓflÍÓ: ÎË·Ó Í‡Í ‚ÂÏÂÌÌÓÂ Á‡-ÏÂ‰ÎÂÌËÂ, Á‡ ÍÓÚÓ˚Ï ÏÓÊÂÚ ÔÓÒÎÂ‰Ó‚‡Ú¸ ÌÓ‚ÓÂÛÒÍÓÂÌËÂ ÓÒÚ‡, ÎË·Ó Í‡Í Ì‡˜‡ÎÓ ÌÂÓ·‡ÚËÏÓ„Ó‚˚ıÓ‰‡ ·ËÓÚ˚ ËÁ ÂÊËÏ‡ Ò Ó·ÓÒÚÂÌËÂÏ. Ç ÚÂ˜ÂÌËÂ‚ÒÂ„Ó Ù‡ÌÂÓÁÓfl Ë ‚ ÏÓÂ, Ë Ì‡ ÒÛ¯Â “ÓÊ‰‡Â-ÏÓÒÚ¸” (Û‰ÂÎ¸Ì‡fl ÒÍÓÓÒÚ¸ ÔÓfl‚ÎÂÌËfl) Ë “ÒÏÂÚ-ÌÓÒÚ¸” (Û‰ÂÎ¸Ì‡fl ÒÍÓÓÒÚ¸ ‚˚ÏË‡ÌËfl) Ú‡ÍÒÓÌÓ‚ ‚ˆÂÎÓÏ ÒÌËÊ‡ÎËÒ¸ (ËÒ. 7). Ç ÏÓÒÍÓÈ ·ËÓÚÂ ‚ ˝ÚÓÏÒÌËÊÂÌËË ÏÓÊÌÓ Á‡ÏÂÚËÚ¸ ‰‚‡ ˝Ú‡Ô‡, ‡Á‰ÂÎÂÌÌ˚Â

‚ÂÏÂÌÌ˚Ï ÂÁÍËÏ ÔÓ‚˚¯ÂÌËÂÏ ÓÊ‰‡ÂÏÓÒÚË ËÒÏÂÚÌÓÒÚË ÔÓÒÎÂ ‚ÂÎËÍÓ„Ó ‚˚ÏË‡ÌËfl Ì‡ Û·ÂÊÂÔ‡ÎÂÓÁÓfl Ë ÏÂÁÓÁÓfl (ÓÍ. 250 ÏÎÌ. ÎÂÚ Ì‡Á‡‰), Á‡ ÍÓ-ÚÓ˚Ï ÔÓÒÎÂ‰Ó‚‡Î ÌÓ‚˚È ˝Ú‡Ô ÔÓÒÚÂÔÂÌÌÓ„Ó ÒÌË-ÊÂÌËfl (ËÒ. 7,‡,·). Ç‡Ì Ç‡ÎÂÌ, ‚ÔÂ‚˚Â Ó·‡ÚË‚-¯ËÈ ‚ÌËÏ‡ÌËÂ Ì‡ ̋ ÚÓ Ó·ÒÚÓflÚÂÎ¸ÒÚ‚Ó, ÓÚÏÂ˜‡Î, ̃ ÚÓ‚ÂÎËÍÓÂ ‚˚ÏË‡ÌËÂ Í‡Í ·˚ “ÔÂÂ‚ÂÎÓ Ì‡Á‡‰ ˝‚Ó-Î˛ˆËÓÌÌ˚Â ˜‡Ò˚”; ‚ Í‡˜ÂÒÚ‚Â ‚ÓÁÏÓÊÌ˚ı ÔË˜ËÌÒÌËÊÂÌËfl ÒÏÂÚÌÓÒÚË ÓÌ ÛÍ‡Á˚‚‡Î ÓÒÚ ÔËÒÔÓÒÓ·-ÎÂÌÌÓÒÚË, ÒÌËÊÂÌËÂ ÒÚÂÔÂÌË ÔÂÂÍ˚‚‡ÌËfl ‡‰‡Ô-ÚË‚Ì˚ı ÁÓÌ Ë ÍÓÌÍÛÂÌˆËË, ÓÒÚ ÔÓÁËÚË‚ÌÓ„Ó ‚Á‡Ë-ÏÓ‰ÂÈÒÚ‚Ëfl ÏÂÊ‰Û Ú‡ÍÒÓÌ‡ÏË (Van Valen, 1984).

Ç ˆÂÎÓÏ Ì‡ ÓÒÌÓ‚‡ÌËË ‰ËÌ‡ÏËÍË ÓÊ‰‡ÂÏÓÒÚË ËÒÏÂÚÌÓÒÚË ÚÛ‰ÌÓ Ò‰ÂÎ‡Ú¸ Ó‰ÌÓÁÌ‡˜Ì˚È ‚˚·ÓÏÂÊ‰Û ‰‚ÛÏfl ‚˚¯ÂÛÍ‡Á‡ÌÌ˚ÏË ‚‡Ë‡ÌÚ‡ÏË ËÌÚÂ-ÔÂÚ‡ˆËË Í‡ÈÌÓÁÓÈÒÍÓ„Ó Á‡ÏÂ‰ÎÂÌËfl ÓÒÚ‡ ‡ÁÌÓ-Ó·‡ÁËfl. ë Ó‰ÌÓÈ ÒÚÓÓÌ˚, ÓÊ‰‡ÂÏÓÒÚ¸ Ë ÒÏÂÚ-ÌÓÒÚ¸ ‚ ÍÓÌˆÂ Í‡ÈÌÓÁÓfl ÛÔ‡ÎË ‰Ó ‚ÂÒ¸Ï‡ ÌËÁÍËı ÁÌ‡-˜ÂÌËÈ (Ò‚Ë‰ÂÚÂÎ¸ÒÚ‚Ó ‚ ÔÓÎ¸ÁÛ „ËÔÓÚÂÁ˚ Ó·ÓÍÓÌ˜‡ÚÂÎ¸ÌÓÏ ‚˚ıÓ‰Â ËÁ ÂÊËÏ‡ Ò Ó·ÓÒÚÂÌËÂÏ), Ò‰Û„ÓÈ – ‚ ÏÓÒÍÓÈ ·ËÓÚÂ ‡Ì‡ÎÓ„Ë˜Ì‡fl ÒËÚÛ‡ˆËfl Ì‡-·Î˛‰‡Î‡Ò¸ Ë ‚ ÍÓÌˆÂ Ô‡ÎÂÓÁÓfl, ÔÓÒÎÂ ̃ Â„Ó ÔÓÒÎÂ‰Ó-‚‡ÎÓ ‚ÂÎËÍÓÂ ‚˚ÏË‡ÌËÂ Ë ÌÓ‚ÓÂ ÂÁÍÓÂ ÛÒÍÓÂÌËÂÓÒÚ‡ ‡ÁÌÓÓ·‡ÁËfl. ç‡·Î˛‰‡ÂÏÓÂ ‚Ó ‚ÚÓÓÈ ÔÓ-ÎÓ‚ËÌÂ Í‡ÈÌÓÁÓfl Á‡ÏÂ‰ÎÂÌËÂ ÓÒÚ‡ ‡ÁÌÓÓ·‡ÁËflÏÓÊÂÚ ·˚Ú¸ Ò‚flÁ‡ÌÓ Ë Ò ÍÎËÏ‡ÚË˜ÂÒÍËÏË Ù‡ÍÚÓ‡-ÏË („ÎÓ·‡Î¸ÌÓÂ ÔÓıÓÎÓ‰‡ÌËÂ Ì‡ Û·ÂÊÂ ˝ÓˆÂÌ‡ ËÓÎË„ÓˆÂÌ‡ (Dupont-Nivet et al., 2007; Zanazzi et al.,2007)), Ë Ò “Ú‡ÍÒÓÌÓÏË˜ÂÒÍËÏ Ì‡Ò˚˘ÂÌËÂÏ” ÒÂ‰˚,

0–450

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–350 –250 –150 –50

0.070.060.050.040.030.020.01

0–450

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ì‰ÂÎ¸Ì‡fl ÒÍÓÓÒÚ¸ ‚˚ÏË‡ÌËfl ÒÂÏÂÈÒÚ‚

–350 –250 –150 –50

0.04

0.03

0.02

0.01

0–450

‚

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ì‰ÂÎ¸Ì‡fl ÒÍÓÓÒÚ¸ ÔÓfl‚ÎÂÌËfl ÒÂÏÂÈÒÚ‚

–350 –250 –150 –500

–450

0.14„

–550

ì‰ÂÎ¸Ì‡fl ÒÍÓÓÒÚ¸ ‚˚ÏË‡ÌËfl ÒÂÏÂÈÒÚ‚

–350 –250 –150 –50

0.12

0.10

0.08

0.06

0.04

0.02

0.14

0.12

0.10

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êËÒ. 7. ëÌËÊÂÌËÂ “ÓÊ‰‡ÂÏÓÒÚË” Ë “ÒÏÂÚÌÓÒÚË” ÒÂÏÂÈÒÚ‚ ÏÓÒÍËı Ë ÍÓÌÚËÌÂÌÚ‡Î¸Ì˚ı Ó„‡ÌËÁÏÓ‚ ‚ Ù‡ÌÂÓÁÓÂ (ÔÓ ‰‡Ì-Ì˚Ï The Fossil Record 2). èÓ ‚ÂÚËÍ‡Î¸ÌÓÈ ÓÒË – Û‰ÂÎ¸Ì‡fl ÒÍÓÓÒÚ¸ ÔÓfl‚ÎÂÌËfl/‚˚ÏË‡ÌËfl (ÓÚÌÓ¯ÂÌËÂ ˜ËÒÎ‡ ÒÂÏÂÈÒÚ‚,ÔÓfl‚Ë‚¯ËıÒfl/‚˚ÏÂ¯Ëı ‚ ‰‡ÌÌÓÏ flÛÒÂ, Í ˜ËÒÎÛ ÒÂÏÂÈÒÚ‚, ÒÛ˘ÂÒÚ‚Ó‚‡‚¯Ëı ‚ ‰‡ÌÌÓÏ flÛÒÂ, ÛÏÌÓÊÂÌÌÓÏÛ Ì‡ ‰ÎËÚÂÎ¸-ÌÓÒÚ¸ flÛÒ‡ ‚ ÏÎÌ. ÎÂÚ). ‡, · – îåÅ, ‚, „ – îäÅ; ‡, ‚ – ÓÊ‰‡ÂÏÓÒÚ¸, ·, „ – ÒÏÂÚÌÓÒÚ¸.

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ÓÒÓ·ÂÌÌÓ Ì‡ ‚˚Ò¯Ëı Ú‡ÍÒÓÌÓÏË˜ÂÒÍËı ÛÓ‚Ìflı, Ë ÒÔÓ„ÂÒÒËÛ˛˘ËÏ ÓÒÚÓÏ ˝‚ÓÎ˛ˆËÓÌÌÓÈ ÛÒÚÓÈ-˜Ë‚ÓÒÚË Ú‡ÍÒÓÌÓ‚ Ë ÒÓÓ·˘ÂÒÚ‚ (Van Valen, 1984;ôÂ·‡ÍÓ‚, 2005). Ñ‡ÊÂ ‚ ÚÓÏ ÒÎÛ˜‡Â, ÂÒÎË Í‡ÈÌÓ-ÁÓÈÒÍÓÂ Á‡ÏÂ‰ÎÂÌËÂ fl‚ÎflÂÚÒfl ‚ÂÏÂÌÌ˚Ï, ‚ÔÓÎÌÂÓ˜Â‚Ë‰ÌÓ, ˜ÚÓ ‰‡Î¸ÌÂÈ¯ËÈ „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ‡ÁÌÓÓ·‡ÁËfl ÒÏÓ„ ·˚ ÔÓ‰ÓÎÊ‡Ú¸Òfl ÌÂ ·ÓÎÂÂ ÌÂ-ÒÍÓÎ¸ÍËı ‰ÂÒflÚÍÓ‚ ÏÎÌ. ÎÂÚ. èÂ‚ÓÈ ËÁ ÂÊËÏ‡ ÒÓ·ÓÒÚÂÌËÂÏ ‰ÓÎÊÌ‡ ‚˚ÈÚË ‰ËÌ‡ÏËÍ‡ ˜ËÒÎ‡ ÒÂ-ÏÂÈÒÚ‚, Á‡ÚÂÏ Ó‰Ó‚, Ë, Ì‡ÍÓÌÂˆ, ‚Ë‰Ó‚. èÓÁÊÂ Ì‡-˜‡‚¯‡flÒfl ˝‚ÓÎ˛ˆËfl ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ ·ËÓÚ˚, ‚Â-ÓflÚÌÓ, ‰ÓÎÊÌ‡ ·˚Î‡ ·˚ ‚˚ÈÚË ËÁ ÂÊËÏ‡ ÒÓ·ÓÒÚÂÌËÂÏ Ò ÌÂÍÓÚÓ˚Ï Á‡ÔÓÁ‰‡ÌËÂÏ ÓÚÌÓÒË-ÚÂÎ¸ÌÓ ÏÓÒÍÓÈ ·ËÓÚ˚.

4. “ÅËÓÎÓ„Ë˜ÂÒÍ‡fl åË-ëËÒÚÂÏ‡”? é·Ì‡ÛÊÂ-ÌËÂ „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ÚÂÌ‰ÂÌˆËË ÓÒÚ‡ Ú‡ÍÒÓÌÓÏË-˜ÂÒÍÓ„Ó ‡ÁÌÓÓ·‡ÁËfl ÒÛıÓÔÛÚÌÓÈ ·ËÓÚ˚ ‰ÓÎÊÌÓ‚˚Á‚‡Ú¸ Û ‚‰ÛÏ˜Ë‚Ó„Ó ˜ËÚ‡ÚÂÎfl fl‰ ÌÂ‰ÓÛÏÂÌÌ˚ı‚ÓÔÓÒÓ‚, ÌÂ ÒÚÓÎ¸ Ó˜Â‚Ë‰Ì˚ı ÔËÏÂÌËÚÂÎ¸ÌÓ Í‰ËÌ‡ÏËÍÂ ‡ÁÌÓÓ·‡ÁËfl ÏÓÒÍÓÈ ·ËÓÚ˚. ÑÂÈÒÚ‚Ë-ÚÂÎ¸ÌÓ, ÔÂ‰ÎÓÊÂÌÌ˚Â Ì‡ÏË ÏÓ‰ÂÎË „ËÔÂ·ÓÎË˜Â-ÒÍÓ„Ó ÓÒÚ‡ (ÔËÏÂÌËÚÂÎ¸ÌÓ Í‡Í Í ÒÓˆË‡Î¸Ì˚Ï,Ú‡Í Ë ·ËÓÎÓ„Ë˜ÂÒÍËÏ ÒËÒÚÂÏ‡Ï) ËÒıÓ‰flÚ ËÁ Ì‡ÎË-˜Ëfl ‚ ÒÓÓÚ‚ÂÚÒÚ‚Û˛˘Ëı ÒËÒÚÂÏ‡ı ÏÂı‡ÌËÁÏÓ‚ ÔÓ-ÎÓÊËÚÂÎ¸ÌÓÈ Ó·‡ÚÌÓÈ Ò‚flÁË ‚ÚÓÓ„Ó ÔÓfl‰Í‡. í‡-ÍËÏ Ó·‡ÁÓÏ, ÏÓ‰ÂÎË ÓÒÌÓ‚˚‚‡˛ÚÒfl Ì‡ ‰ÓÔÛ˘ÂÌËËÓ ÚÓÏ, ˜ÚÓ ÒÓÓ‚ÂÚÒÚ‚Û˛˘‡fl ˝‚ÓÎ˛ˆËÓÌËÛ˛˘‡flÓ·˘ÌÓÒÚ¸ ÔÂ‰ÒÚ‡‚ÎflÎ‡ ÒÓ·ÓÈ Â‡Î¸ÌÓ Ò‚flÁÌÓÂ ˆÂ-ÎÓÂ, ÒËÒÚÂÏÛ ‚ ÒÚÓ„ÓÏ ÒÏ˚ÒÎÂ ̋ ÚÓ„Ó ÒÎÓ‚‡. ÇÏÂÒÚÂÒ ÚÂÏ ‚ÔÓÎÌÂ Ó˜Â‚Ë‰ÌÓ, ˜ÚÓ ÔÓÒÎÂ ‡ÒÔ‡‰‡ è‡Ì„ÂË,Ì‡˜‡‚¯Â„ÓÒfl ‚ ÚË‡ÒÓ‚ÓÏ ÔÂËÓ‰Â, ÒÛıÓÔÛÚÌ‡fl·ËÓÚ‡, ‚ÁflÚ‡fl ‚ ˆÂÎÓÏ, ÌÂ ÔÂ‰ÒÚ‡‚ÎflÎ‡ ÒÓ·ÓÈ Â-‡Î¸ÌÓ„Ó ÒËÒÚÂÏÌÓ„Ó Â‰ËÌÒÚ‚‡, Ë·Ó Ò‚flÁË ËÁÓÎËÓ-‚‡ÌÌ˚ı Ï‡ÚÂËÍÓ‚ Ò ·ËÓÚÓÈ ÓÒÚ‡Î¸ÌÓÈ ÒÛ¯Ë ‚ ÚÂ-˜ÂÌËÂ ‰ÎËÚÂÎ¸ÌÓ„Ó ‚ÂÏÂÌË ·˚ÎË ÂÁÍÓ ÓÒÎ‡·ÎÂ-Ì˚. åÓÒÍËÂ ÔÓÎË‚˚ ÔÂ‰ÒÚ‡‚Îfl˛Ú ÒÓ·ÓÈ ÔÓ˜ÚËÌÂÔÂÓ‰ÓÎËÏÛ˛ ÔÂ„‡‰Û ‰Îfl fl‰‡ ÍÓÌÚËÌÂÌÚ‡Î¸-Ì˚ı ·ÂÒÔÓÁ‚ÓÌÓ˜Ì˚ı (ËÒÍÎ˛˜ÂÌËÂ ÒÓÒÚ‡‚Îfl˛Ú ÎÂ-Ú‡˛˘ËÂ Ì‡ÒÂÍÓÏ˚Â, ÓÚ˜‡ÒÚË Ô‡ÛÍË, ÏÂÎÍËÂ ·ÂÒÔÓ-Á‚ÓÌÓ˜Ì˚Â Ò ÛÒÚÓÈ˜Ë‚˚ÏË ÔÓÍÓfl˘ËÏËÒfl ÒÚ‡‰ËflÏË,ÍÓÚÓ˚Â ÏÓ„ÛÚ ‡ÁÌÓÒËÚ¸Òfl ‚ÂÚÓÏ, Ë Ú.‰.), ÏÌÓ„Ëı‚˚Ò¯Ëı ‡ÒÚÂÌËÈ, ·ÓÎ¸¯ËÌÒÚ‚‡ ÌÂÎÂÚ‡˛˘Ëı ÚÂÚ-‡ÔÓ‰ Ë ÔÂÒÌÓ‚Ó‰Ì˚ı ˚·. èÓ˝ÚÓÏÛ ÏÌÓ„ËÂ ‚‡Ê-Ì˚Â “·ËÓÎÓ„Ë˜ÂÒÍËÂ ËÌÌÓ‚‡ˆËË” ÌÂ ËÏÂÎË ¯‡ÌÒÓ‚‡ÒÔÓÒÚ‡ÌËÚ¸Òfl Ì‡ ˝ÚË ËÁÓÎËÓ‚‡ÌÌ˚Â Û˜‡ÒÚÍËÒÛ¯Ë (Ì‡ÔËÏÂ, ·ÓÎ¸¯ËÌÒÚ‚Ó „ÛÔÔ ÔÎ‡ˆÂÌÚ‡-Ì˚ı ÏÎÂÍÓÔËÚ‡˛˘Ëı ÌÂ ÔÓÌËÍÎÓ ‚ Ä‚ÒÚ‡ÎË˛).

ëıÓ‰ÌÓÂ ‚ÓÁ‡ÊÂÌËÂ ÔËÏÂÌËÏÓ Ë Í „ËÔÂ·ÓÎË-˜ÂÒÍËÏ ÏÓ‰ÂÎflÏ ‰ËÌ‡ÏËÍË ÏË‡ Î˛‰ÂÈ. éÚÏÂÚËÏ,˜ÚÓ, ‚ÔÂ‚˚Â ÔÓÁÌ‡ÍÓÏË‚¯ËÒ¸ Ò Ï‡ÚÂÏ‡ÚË˜ÂÒÍËÏËÏÓ‰ÂÎflÏË ÓÒÚ‡ Ì‡ÒÂÎÂÌËfl ÏË‡, Ï˚ Ò‡ÏË ËÒÔ˚Ú‡-ÎË ÓÔÂ‰ÂÎÂÌÌÓÂ ÌÂ‰Ó‚ÂËÂ Í ÌËÏ. ÑÂÈÒÚ‚ËÚÂÎ¸ÌÓ,ÓÌË ÔÓ‰‡ÁÛÏÂ‚‡˛Ú, ˜ÚÓ Ì‡ÒÂÎÂÌËÂ ÏË‡ ÏÓ„ÎÓ‡ÒÒÏ‡ÚË‚‡Ú¸Òfl ‚ Í‡˜ÂÒÚ‚Â Â‰ËÌÓÈ ÒËÒÚÂÏ˚ Ì‡ÔÓÚflÊÂÌËÂ ÏÌÓ„Ëı Ú˚Òfl˜ ÎÂÚ, Ë ÛÊÂ îÂÒÚÂ, åÓ-‡ Ë ÄÏËÓÚ ÔflÏÓ ‰ÂÎ‡ÎË ˝ÚÓ ‰ÓÔÛ˘ÂÌËÂ: “é‰Ì‡ÍÓÚÓ, ˜ÚÓ ÏÓÊÂÚ ·˚Ú¸ Ô‡‚ËÎ¸Ì˚Ï ÔÓ ÓÚÌÓ¯ÂÌË˛ Í˝ÎÂÏÂÌÚ‡Ï, ÍÓÚÓ˚Â ËÁ-Á‡ ÓÚÒÛÚÒÚ‚Ëfl ÏÂÊ‰Û ÌËÏË‡‰ÂÍ‚‡ÚÌÓÈ ÍÓÏÏÛÌËÍ‡ˆËË ‰ÓÎÊÌ˚ ÔËÌËÏ‡Ú¸

Û˜‡ÒÚËÂ ‚ ÒÓÂ‚ÌÓ‚‡ÚÂÎ¸ÌÓÈ Ë„Â Ò (ÔÓ˜ÚË) ÌÛÎÂ‚ÓÈÒÛÏÏÓÈ ‚˚Ë„˚¯‡, ÏÓÊÂÚ ·˚Ú¸ ÌÂÔ‡‚ËÎ¸Ì˚Ï ‰Îfl˝ÎÂÏÂÌÚÓ‚, Ó·Î‡‰‡˛˘Ëı ÒËÒÚÂÏÓÈ ÍÓÏÏÛÌËÍ‡ˆËË,ÍÓÚÓ‡fl ‰‡ÂÚ ËÏ ‚ÓÁÏÓÊÌÓÒÚ¸ Ó·‡ÁÓ‚˚‚‡Ú¸ ÍÓ‡-ÎËˆËË, ÔÓÍ‡ ‚ÒÂ ˝ÎÂÏÂÌÚ˚ ÌÂ ÓÍ‡Á˚‚‡˛ÚÒfl ÒÚÓÎ¸ÒËÎ¸ÌÓ Ò‚flÁ‡Ì˚ ÏÂÊ‰Û ÒÓ·ÓÈ, ˜ÚÓ ‚ÒÂ Ì‡ÒÂÎÂÌËÂ ÒÚÓ˜ÍË ÁÂÌËfl ÚÂÓËË Ë„ ÏÓÊÂÚ ‡ÒÒÏ‡ÚË‚‡Ú¸Òfl ‚Í‡˜ÂÒÚ‚Â Â‰ËÌÓ„Ó Ë„ÓÍ‡, ‚Â‰Û˘Â„Ó Ë„Û, ‚ ÍÓÚÓÓÈ‚ ÓÎË ‚ÚÓÓ„Ó Ë„ÓÍ‡-ÓÔÔÓÌÂÌÚ‡ ‚˚ÒÚÛÔ‡ÂÚ ÔË-Ó‰‡” (Foerster et al., 1960, p. 1292).

é‰Ì‡ÍÓ ËÏÂ˛ÚÒfl ÒÂ¸ÂÁÌ˚Â ÓÒÌÓ‚‡ÌËfl ÛÒÓ-ÏÌËÚ¸Òfl ‚ Ó·ÓÒÌÓ‚‡ÌÌÓÒÚË ÔÓ‰Ó·ÌÓ„Ó ‰ÓÔÛ˘ÂÌËfl.ÇÔÎÓÚ¸ ‰Ó Ò‡ÏÓ„Ó ÌÂ‰‡‚ÌÂ„Ó ‚ÂÏÂÌË (‡ ‚ ÓÒÓ·ÂÌ-ÌÓÒÚË ‰Ó 1492 „.) ˜ÂÎÓ‚Â˜ÂÒÚ‚Ó ÌÂ ÔÂ‰ÒÚ‡‚ÎflÎÓ ÒÓ-·ÓÈ ÒËÒÚÂÏ˚ ÌË ‚ Í‡ÍÓÏ Â‡Î¸ÌÓÏ ÒÏ˚ÒÎÂ, Ë·Ó, Ì‡-ÔËÏÂ, ÓÒÚ Ì‡ÒÂÎÂÌËfl Ú‡ÍËı Â„ËÓÌÓ‚, Í‡Í ëÚ‡-˚È ë‚ÂÚ, çÓ‚˚È ë‚ÂÚ, Ä‚ÒÚ‡ÎËfl Ë í‡ÒÏ‡ÌËfl ËÎËÉ‡‚‡ÈÒÍËÂ ÓÒÚÓ‚‡, ÔÓËÒıÓ‰ËÎ ÔÓ˜ÚË ÔÓÎÌÓÒÚ¸˛ÌÂÁ‡‚ËÒËÏÓ ‰Û„ ÓÚ ‰Û„‡. í‡Í, ÔÂ‰ÒÚ‡‚ÎflÂÚÒfl‚ÔÓÎÌÂ Ó˜Â‚Ë‰Ì˚Ï, ˜ÚÓ ·ÛÌ˚Â ‰ÂÏÓ„‡ÙË˜ÂÒÍËÂÔÓˆÂÒÒ˚, ÔÓËÒıÓ‰Ë‚¯ËÂ ‚ I Ú˚Ò. Ì.˝. ‚ Ö‚‡ÁËË,ÌÂ ÓÍ‡Á‡ÎË ‡·ÒÓÎ˛ÚÌÓ ÌËÍ‡ÍÓ„Ó ‚ÎËflÌËfl Ì‡ ÒËÌ-ıÓÌÌÛ˛ ‰ÂÏÓ„‡ÙË˜ÂÒÍÛ˛ ‰ËÌ‡ÏËÍÛ, ÒÍ‡ÊÂÏ,Ó·ËÚ‡ÚÂÎÂÈ í‡ÒÏ‡ÌËË (‰‡ Ë Ó·‡ÚÌÓÂ ‚ÎËflÌËÂ Ú‡Í-ÊÂ ·˚ÎÓ ÌÛÎÂ‚˚Ï).

Ñ‡ÌÌÓÂ ‚ÓÁ‡ÊÂÌËÂ ÔË‚Ó‰ËÚ, Ì‡ÔËÏÂ,û.Ç. òË¯ÍÓ‚ ‚ ÒÚ‡Ú¸Â ÔÓ‰ ÒËÏÔÚÓÏ‡ÚË˜ÂÒÍËÏ Ì‡-Á‚‡ÌËÂÏ “ÑÂÏÓ„‡ÙË˜ÂÒÍËÂ ÔÓıÓÊ‰ÂÌËfl ÙËÁËÍ‡”:“à ÔÓÒÎÂ Ó‚Î‡‰ÂÌËfl Â˜¸˛ Ë ÔËÒ¸ÏÂÌÌÓÒÚ¸˛ ˜ÂÎÓ-‚Â˜ÂÒÚ‚Ó ÏÌÓ„ËÂ Ú˚Òfl˜Ë ÎÂÚ ÓÒÚ‡‚‡ÎÓÒ¸ ÒÚÓÎ¸ ÌÂ-ÏÌÓ„Ó˜ËÒÎÂÌÌ˚Ï Ë ‡ÁÓ·˘ÂÌÌ˚Ï, ˜ÚÓ Ó‰ÌË Â„Ó„ÛÔÔ˚ ÌÂ ËÏÂÎË ÔÓÌflÚËfl Ó ÒÛ˘ÂÒÚ‚Ó‚‡ÌËË ‰Û„Ëı.Ñ‡‚ÌÓ ÎË ÔÓ ËÒÚÓË˜ÂÒÍËÏ ÏÂÍ‡Ï Â‚ÓÔÂÈˆ˚ Ë‡ÁË‡Ú˚ ÛÁÌ‡ÎË Ó ÊËÚÂÎflı á‡Ô‡‰ÌÓ„Ó ÔÓÎÛ¯‡Ëfl?ä‡Í Ú‡ÍÓÂ ˜ÂÎÓ‚Â˜ÂÒÚ‚Ó ÏÓ„ÎÓ ·˚Ú¸ Â‰ËÌ˚Ï ËÌ-ÙÓÏ‡ˆËÓÌÌ˚Ï ÔÓÎÂÏ? Çfl‰ ÎË ä‡ÔËˆ‡ ‰ÓÔÛÒÍ‡ÂÚ,˜ÚÓ ÔÂÒÌË ·‡‰Ó‚ Ë ‡ÒÒÍ‡Á˚ ÒÚ‡ËÍÓ‚ Û ÒÂÏÂÈÌÓ„ÓÓ˜‡„‡ ÔË ÓÚÒÛÚÒÚ‚ËË ÒÔÛÚÌËÍÓ‚˚ı ÂÚ‡ÌÒÎflÚÓ-Ó‚ Á‚Û˜‡ÎË Ì‡ ‚Ò˛ ÓÈÍÛÏÂÌÛ. Ä ÂÒÎË ·˚ Ë Á‚Û˜‡ÎË,ÚÓ Ì‡ flÁ˚ÍÂ, ÌÂÔÓÌflÚÌÓÏ ‰Îfl ÔÓ‰‡‚Îfl˛˘ÂÈ ˜‡ÒÚËÓÈÍÛÏÂÌ˚” (òË¯ÍÓ‚, 2005, Ò. 160). å˚ Ò˜ËÚ‡ÂÏ˝ÚÓ ‚ÓÁ‡ÊÂÌËÂ ÒÓ‚Â¯ÂÌÌÓ Ó·ÓÒÌÓ‚‡ÌÌ˚Ï, Ë·Ó‡‚ÚÓ˚ „ËÔÂ·ÓÎË˜ÂÒÍËı ÏÓ‰ÂÎÂÈ ÓÒÚ‡ Ì‡ÒÂÎÂÌËfláÂÏÎË ‰Ó ÒËı ÔÓ ÌÂ ‰‡ÎË Ì‡ ÌÂ„Ó ‡„ÛÏÂÌÚËÓ‚‡Ì-ÌÓ„Ó ÓÚ‚ÂÚ‡. à Â˜¸ Á‰ÂÒ¸ Ë‰ÂÚ ÌÂ ÚÓÎ¸ÍÓ Óë.è. ä‡ÔËˆÂ, ÌÓ Ë Ó ï. ÙÓÌ îÂÒÚÂÂ Ë Â„Ó ÍÓÎÎÂ-„‡ı, å. äÂÏÂÂ Ë ‰Û„Ëı ‡‚ÚÓ‡ı Ï‡ÚÂÏ‡ÚË˜ÂÒÍËıÏÓ‰ÂÎÂÈ „ËÔÂ·ÓÎË˜ÂÒÍÓ„Ó ÓÒÚ‡ Ì‡Ó‰ÓÌ‡ÒÂÎÂ-ÌËfl áÂÏÎË (èÓ‰Î‡ÁÓ‚, 2002; Cohen, 1995; Hoerner,1975; Johansen, Sornette, 2001; Tsirel, 2004).

íÂÏ ÌÂ ÏÂÌÂÂ Ï˚ ÔÓÎ‡„‡ÂÏ, ˜ÚÓ Í‡ÚËÌ‡ ‚˚ÒÓ-ÍÓ‰ÂÚÂÏËÌËÓ‚‡ÌÌÓÈ ÚÂıÌËÍÓ-˝ÍÓÌÓÏË˜ÂÒÍÓÈ,ÍÛÎ¸ÚÛÌÓÈ Ë ‰ÂÏÓ„‡ÙË˜ÂÒÍÓÈ ‰ËÌ‡ÏËÍË ÏË‡ Á‡ÔÓÒÎÂ‰ÌËÂ ÌÂÒÍÓÎ¸ÍÓ Ú˚Òfl˜ ÎÂÚ ÌË ‚ ÍÓÂÈ ÒÚÂÔÂÌËÌÂ fl‚ÎflÂÚÒfl ÒÎÛ˜‡ÈÌÓÈ. ëÓ·ÒÚ‚ÂÌÌÓ „Ó‚Ófl, ÓÌ‡ ÓÚ-‡Ê‡ÂÚ ‰ËÌ‡ÏËÍÛ ÒÓ‚Â¯ÂÌÌÓ Â‡Î¸ÌÓÈ ÒËÒÚÂÏ˚,Á‡Ó‰Ë‚¯ÂÈÒfl ‚ Ì‡˜‡ÎÂ „ÓÎÓˆÂÌ‡ Ì‡ ÅÎËÊÌÂÏ ÇÓ-ÒÚÓÍÂ ‚ ÌÂÔÓÒÂ‰ÒÚ‚ÂÌÌÓÈ Ò‚flÁË Ò Ì‡˜‡‚¯ÂÈÒfl Ú‡Ï



‡„‡ÌÓÈ (“ÌÂÓÎËÚË˜ÂÒÍÓÈ”) Â‚ÓÎ˛ˆËÂÈ Ë ÔÓÒÚÂ-ÔÂÌÌÓ Óı‚‡ÚË‚¯ÂÈ ÒÓ·ÓÈ ‚ÂÒ¸ ÏË. ÇÒÎÂ‰ Á‡ î‡Ì-ÍÓÏ (Frank, 1990, 1993; Frank, Gills, 1993) Ï˚ Ì‡Á˚-‚‡ÂÏ ̋ ÚÛ ÒËÒÚÂÏÛ åË-ëËÒÚÂÏÓÈ, Ë ‚ÒÎÂ‰ Á‡ ÌËÏ Ï˚ıÓÚËÏ ÔÓ‰˜ÂÍÌÛÚ¸, ̃ ÚÓ Ì‡ ÚÓ, ̃ ÚÓ·˚ åË-ëËÒÚÂÏ‡Óı‚‡ÚËÎ‡ ÒÓ·ÓÈ ‚ÂÒ¸ ÏË, Û¯ÎÓ ÏÌÓ„Ó Ú˚Òfl˜ ÎÂÚ; ËÔÓ˝ÚÓÏÛ Ì‡ ÔÓÚflÊÂÌËË ‡·ÒÓÎ˛ÚÌÓ ·ÓÎ¸¯ÂÈ ̃ ‡ÒÚËÒ‚ÓÂ„Ó ÒÛ˘ÂÒÚ‚Ó‚‡ÌËfl ËÒÚÓËfl åË-ëËÒÚÂÏ˚ ÌË ‚ÍÓÂÏ ÒÎÛ˜‡Â ÌÂ ·˚Î‡ ÚÓÊ‰ÂÒÚ‚ÂÌÌ‡ “‚ÒÂÏËÌÓÈ ËÒ-ÚÓËË”.

éÚÏÂÚËÏ, ˜ÚÓ, Í‡Í ·˚ÎÓ ÔÓÍ‡Á‡ÌÓ Ì‡ÏË ‡ÌÂÂ(äÓÓÚ‡Â‚ Ë ‰., 2007‡,·), ËÏÂÌÌÓ Ò ‡Á‚ËÚËÂÏ åË-ëËÒÚÂÏ˚ Ò‚flÁ‡ÌÓ Ì‡ÎË˜ËÂ „ËÔÂ·ÓÎË˜ÂÒÍÓ„Ó ÚÂÌ-‰‡ ÓÒÚ‡ ̃ ËÒÎÂÌÌÓÒÚË Ì‡ÒÂÎÂÌËfl ÏË‡. ç‡ÎË˜ËÂ „Ë-ÔÂ·ÓÎË˜ÂÒÍÓ„Ó ÚÂÌ‰‡ Ò‚Ë‰ÂÚÂÎ¸ÒÚ‚ÛÂÚ Ó ÚÓÏ, ̃ ÚÓ·óÎ¸¯‡fl ˜‡ÒÚ¸ ÒÓÓÚ‚ÂÚÒÚ‚Û˛˘ÂÈ Ó·˘ÌÓÒÚË (‡ ‚ ÔÓ-ÒÎÂ‰ÌÂÏ ÒÎÛ˜‡Â, Ì‡ÔÓÏÌËÏ, Â˜¸ Ë‰ÂÚ Ó Ì‡Ó‰ÓÌ‡ÒÂ-ÎÂÌËË ÏË‡) ËÏÂÎ‡ ÓÔÂ‰ÂÎÂÌÌÓÂ ÒËÒÚÂÏÌÓÂ Â‰ËÌ-ÒÚ‚Ó, Ë Ì‡Ï ÔÂ‰ÒÚ‡‚ÎflÂÚÒfl, ̃ ÚÓ ‚ Ì‡¯ÂÏ ‡ÒÔÓflÊÂ-ÌËË ËÏÂÂÚÒfl ‰ÓÒÚ‡ÚÓ˜ÌÓ ‰‡ÌÌ˚ı ‰Îfl ÚÓ„Ó, ˜ÚÓ·˚ÛÚ‚ÂÊ‰‡Ú¸, ˜ÚÓ ÔÓ‰Ó·ÌÓÂ ÒËÒÚÂÏÌÓÂ Â‰ËÌÒÚ‚Ó ‚‡ÒÒÏ‡ÚË‚‡ÂÏÛ˛ ˝ÔÓıÛ Â‡Î¸ÌÓ Ì‡·Î˛‰‡ÎÓÒ¸.ÑÂÈÒÚ‚ËÚÂÎ¸ÌÓ, ‚ Ì‡¯ÂÏ ‡ÒÔÓflÊÂÌËË ËÏÂÂÚÒfl ‰Ó-ÒÚ‡ÚÓ˜ÌÓ ‰‡ÌÌ˚ı Ó ÒËÒÚÂÏ‡ÚË˜ÂÒÍÓÏ ‡ÒÔÓÒÚ‡ÌÂ-ÌËË ‚‡ÊÌÂÈ¯Ëı ËÌÌÓ‚‡ˆËÈ (‰ÓÏÂÒÚËˆËÓ‚‡ÌÌ˚ıÁÎ‡ÍÓ‚, ÍÛÔÌÓ„Ó Ë ÏÂÎÍÓ„Ó Ó„‡ÚÓ„Ó ÒÍÓÚ‡, ÎÓ¯‡-‰Ë, ÔÎÛ„‡, ÍÓÎÂÒ‡, ÏÂÚ‡ÎÎÛ„ËË ÏÂ‰Ë, ·ÓÌÁ˚, ‡ ‚‰‡Î¸ÌÂÈ¯ÂÏ Ë ÊÂÎÂÁ‡, Ë Ú.‰.) Ò ÅÎËÊÌÂ„Ó ÇÓÒÚÓÍ‡ÔÓ ‚ÒÂÈ ÒÂ‚ÂÓ‡ÙËÍ‡ÌÒÍÓ-Â‚‡ÁËÈÒÍÓÈ éÈÍÛÏÂ-ÌÂ, Ì‡˜‡‚¯ÂÏÒfl Á‡ ÏÌÓ„Ó Ú˚Òfl˜ ÎÂÚ ‰Ó Ì. ˝. (ÒÏ., Ì‡-ÔËÏÂ, óÛ·‡Ó‚, 1991). Ç ÂÁÛÎ¸Ú‡ÚÂ ‰‡ÌÌ˚ı ÔÓ-ˆÂÒÒÓ‚ ˝‚ÓÎ˛ˆËfl Ó·˘ÂÒÚ‚ ‰‡ÌÌÓ„Ó Ï‡ÍÓÂ„ËÓÌ‡ÛÊÂ ‚ ˝ÚÓ ‚ÂÏfl ÌÂ ÏÓÊÂÚ ‡ÒÒÏ‡ÚË‚‡Ú¸Òfl Í‡ÍÔÓÎÌÓÒÚ¸˛ ÌÂÁ‡‚ËÒËÏ‡fl.

á‰ÂÒ¸ ÔÂ‰ÒÚ‡‚ÎflÂÚÒfl ÌÂÓ·ıÓ‰ËÏ˚Ï ÒÎÂ‰Û˛-˘ËÈ ÍÓÏÏÂÌÚ‡ËÈ. äÓÌÂ˜ÌÓ, Û Ì‡Ò ÌÂ ·˚ÎÓ ·˚ ÓÒ-ÌÓ‚‡ÌËÈ „Ó‚ÓËÚ¸ Ó åË-ëËÒÚÂÏÂ, ÔÓÒÚË‡˛˘ÂÈ-Òfl ÓÚ ÄÚÎ‡ÌÚËÍË ‰Ó íËıÓ„Ó ÓÍÂ‡Ì‡, ‰‡ÊÂ ‰Îfl Ì‡˜‡-Î‡ I Ú˚Ò. Ì.˝., ÂÒÎË ·˚ Ï˚ ÔËÏÂÌflÎË ÍËÚÂËÈ“Ï‡ÒÒÓ‚˚ı ÚÓ‚‡Ó‚” (“bulk-good” criterion), ÔÂ‰ÎÓ-ÊÂÌÌ˚È Ç‡ÎÎÂÒÚ‡ÈÌÓÏ (Wallerstein, 1974, 1987,2004), ÔÓÚÓÏÛ ˜ÚÓ ‚ ˝ÚÓ ‚ÂÏfl Í‡ÍÓÂ-ÎË·Ó ‰‚ËÊÂ-ÌËÂ Ï‡ÒÒÓ‚˚ı ÚÓ‚‡Ó‚, ÒÍ‡ÊÂÏ, ÏÂÊ‰Û äËÚ‡ÂÏ ËÖ‚ÓÔÓÈ ÔÓÎÌÓÒÚ¸˛ ÓÚÒÛÚÒÚ‚Ó‚‡ÎÓ (Ë Ï˚ ÌÂ ËÏÂÂÏÌËÍ‡ÍËı ÓÒÌÓ‚‡ÌËÈ ÌÂ ÒÓ„Î‡ÒËÚ¸Òfl Ò à. Ç‡ÎÎÂ-ÒÚ‡ÈÌÓÏ ‚ Â„Ó ÍÎ‡ÒÒËÙËÍ‡ˆËË ÔÓÔ‡‰‡‚¯Â„Ó ‚ ‰‡Ì-ÌÓÂ ‚ÂÏfl ‚ Ö‚ÓÔÛ ÍËÚ‡ÈÒÍÓ„Ó ¯ÂÎÍ‡ Í‡Í ÔÂ‰ÏÂ-Ú‡ ÓÒÍÓ¯Ë, ÌÓ ÌËÍ‡Í ÌÂ Ï‡ÒÒÓ‚Ó„Ó ÚÓ‚‡‡). é‰Ì‡-ÍÓ åË-ëËÒÚÂÏ‡ I ‚. Ì.˝. (Ë ‰‡ÊÂ I Ú˚Òfl˜ÂÎÂÚËfl ‰ÓÌ.˝.) ÏÓÊÂÚ ‚ÔÓÎÌÂ ·˚Ú¸ ÍÎ‡ÒÒËÙËˆËÓ‚‡Ì‡ ËÏÂÌ-ÌÓ Í‡Í åË-ëËÒÚÂÏ‡, ÂÒÎË Ï˚ ÔËÏÂÌËÏ Á‰ÂÒ¸ ·Ó-ÎÂÂ Ïfl„ÍËÈ ÍËÚÂËÈ “ËÌÙÓÏ‡ˆËÓÌÌÓÈ ÒÂÚË”,ÔÂ‰ÎÓÊÂÌÌ˚È óÂÈÁ-Ñ‡ÌÌÓÏ Ë ïÓÎÎÓÏ (Chase-Dunn, Hall, 1997; ÒÏ. Ú‡ÍÊÂ, Ì‡ÔËÏÂ, óÂ¯ÍÓ‚,1999). èÓ‰˜ÂÍÌÂÏ, ˜ÚÓ, Í‡Í ·˚ÎÓ ÔÓÍ‡Á‡ÌÓ Ì‡ÏË‡ÌÂÂ (äÓÓÚ‡Â‚ Ë ‰., 2005a,·, 2007·), Ì‡ÎË˜ËÂ ËÌ-ÙÓÏ‡ˆËÓÌÌÓÈ ÒÂÚË, Óı‚‡Ú˚‚‡˛˘ÂÈ ‚Ò˛ åË-ëË-ÒÚÂÏÛ, fl‚ÎflÂÚÒfl ÒÓ‚Â¯ÂÌÌÓ ‰ÓÒÚ‡ÚÓ˜Ì˚Ï ÛÒÎÓ‚Ë-ÂÏ, ÍÓÚÓÓÂ ‰ÂÎ‡ÂÚ ‚ÓÁÏÓÊÌ˚Ï ‡ÒÒÏ‡ÚË‚‡Ú¸ ‚Ò˛

åË-ëËÒÚÂÏÛ Í‡Í Â‰ËÌÓÂ ‡Á‚Ë‚‡˛˘ÂÂÒfl ˆÂÎÓÂ.Ñ‡, ‚ I Ú˚Ò. ‰Ó Ì. ˝. Í‡ÍËÂ-ÎË·Ó Ï‡ÒÒË‚Ì˚Â ÚÓ‚‡Ó-ÔÓÚÓÍË ÏÂÊ‰Û íËıÓÓÍÂ‡ÌÒÍËÏ Ë ÄÚÎ‡ÌÚË˜ÂÒÍËÏÔÓ·ÂÂÊ¸flÏË Ö‚‡ÁËË ·˚ÎË ÔËÌˆËÔË‡Î¸ÌÓ ÌÂ-‚ÓÁÏÓÊÌ˚. é‰Ì‡ÍÓ åË-ëËÒÚÂÏ‡ ‰ÓÒÚË„Î‡ Í ̋ ÚÓÏÛ‚ÂÏÂÌË Ú‡ÍÓ„Ó ÛÓ‚Ìfl ËÌÚÂ„‡ˆËË, ÍÓÚÓ˚È ÛÊÂ‰ÂÎ‡Î ‚ÓÁÏÓÊÌ˚Ï ‡ÒÔÓÒÚ‡ÌÂÌËÂ ÔÓ ‚ÒÂÈ åË-ëËÒÚÂÏÂ ÔËÌˆËÔË‡Î¸ÌÓ ‚‡ÊÌ˚ı ÚÂıÌÓÎÓ„ËÈ Á‡ÔÓÏÂÊÛÚÍË ‚ÂÏÂÌË, Á‡ÏÂÚÌÓ ÏÂÌ¸¯ËÂ Ú˚Òfl˜ÂÎÂ-ÚËfl.

ÑÛ„ËÏ ‚‡ÊÌ˚Ï ÏÓÏÂÌÚÓÏ ÏÓÊÂÚ ÔÂ‰ÒÚ‡‚-ÎflÚ¸Òfl ÚÓ Ó·ÒÚÓflÚÂÎ¸ÒÚ‚Ó, ˜ÚÓ ‰‡ÊÂ ‚ I ‚. Ì.˝. åË-ëËÒÚÂÏ‡ Óı‚‡Ú˚‚‡Î‡ Á‡ÏÂÚÌÓ ÏÂÌÂÂ ÔÓÎÓ‚ËÌ˚‚ÒÂÈ Ó·ËÚ‡ÂÏÓÈ ÁÂÏÌÓÈ ÒÛ¯Ë. é‰Ì‡ÍÓ „Ó‡Á‰Ó ·Ó-ÎÂÂ ‚‡ÊÌ˚Ï Á‰ÂÒ¸ ÔÂ‰ÒÚ‡‚ÎflÂÚÒfl ‰Û„ÓÂ Ó·ÒÚÓfl-ÚÂÎ¸ÒÚ‚Ó: ÛÊÂ Í Ì‡˜‡ÎÛ I ‚. Ì.˝. ·ÓÎÂÂ 90% Ì‡ÒÂÎÂ-ÌËfl ÏË‡ ÊËÎÓ ËÏÂÌÌÓ ‚ ÚÂı Â„ËÓÌ‡ı áÂÏÎË, ÍÓÚÓ-˚Â ·˚ÎË ËÌÚÂ„‡Î¸Ì˚ÏË ˜‡ÒÚflÏË åË-ëËÒÚÂÏ˚(ëÂ‰ËÁÂÏÌÓÏÓ¸Â, ëÂ‰ÌËÈ ÇÓÒÚÓÍ, ûÊÌ‡fl, ñÂÌ-Ú‡Î¸Ì‡fl Ë ÇÓÒÚÓ˜Ì‡fl ÄÁËfl) (ÒÏ., Ì‡ÔËÏÂ, Du-rand, 1977). á‡ ÌÂÒÍÓÎ¸ÍÓ Ú˚Òfl˜ÂÎÂÚËÈ ÔÂÂ‰ ˝ÚËÏÏ˚ ËÏÂÂÏ ‰ÂÎÓ Ò ÔÓflÒÓÏ ÍÛÎ¸ÚÛ, Ú‡ÍÊÂ ı‡‡ÍÚÂ-ËÁÓ‚‡‚¯ËÏÒfl Û‰Ë‚ËÚÂÎ¸ÌÓ ÒıÓ‰Ì˚Ï ÛÓ‚ÌÂÏ Ë ı‡-‡ÍÚÂÓÏ ÍÛÎ¸ÚÛÌÓÈ ÒÎÓÊÌÓÒÚË, ÔÓÚflÌÛ‚¯ËÏÒflÓÚ Å‡ÎÍ‡Ì ‚ÔÎÓÚ¸ ‰Ó ‰ÓÎËÌ˚ àÌ‰‡ (ÒÏ., Ì‡ÔËÏÂ:Peregrine, Ember, 2001a,b,c, 2003; Peregrine, 2003).èÓ‰˜ÂÍÌÂÏ, ̃ ÚÓ Ë ‚ ÚÓÏ, Ë ‚ ‰Û„ÓÏ ÒÎÛ˜‡Â Ì‡ÒÂÎÂ-ÌËÂ ÒÓÓÚ‚ÂÚÒÚ‚Û˛˘Ëı ÔÓflÒÓ‚ ‚ÍÎ˛˜‡ÎÓ ‚ ÒÂ·fl·óÎ¸¯Û˛ ˜‡ÒÚ¸ Ì‡ÒÂÎÂÌËfl ÏË‡ ÒÓÓÚ‚ÂÚÒÚ‚Û˛˘ÂÈ˝ÔÓıË. í‡ÍËÏ Ó·‡ÁÓÏ, ÛÊÂ ÌÂÒÍÓÎ¸ÍÓ Ú˚Òfl˜ ÎÂÚ‰ËÌ‡ÏËÍ‡ Ì‡ÒÂÎÂÌËfl ÏË‡ ÓÚ‡Ê‡ÂÚ ÔÂÊ‰Â ‚ÒÂ„ÓËÏÂÌÌÓ ‰ËÌ‡ÏËÍÛ Ì‡ÒÂÎÂÌËfl åË-ëËÒÚÂÏ˚, ˜ÚÓ Ë‰ÂÎ‡ÂÚ ‚ÓÁÏÓÊÌ˚Ï ÂÂ ÓÔËÒ‡ÌËÂ ÔË ÔÓÏÓ˘Ë Ï‡ÚÂ-Ï‡ÚË˜ÂÒÍËı Ï‡ÍÓÏÓ‰ÂÎÂÈ. äÓÌÂ˜ÌÓ, ·ÛÌÓÂ ‡Á-‚ËÚËÂ ÚÂıÌÓÎÓ„ËË ‚ åË-ëËÒÚÂÏÂ ‚ÔÎÓÚ¸ ‰Ó XIX ‚.ÌËÍ‡Í ÌÂ ÒÍ‡Á˚‚‡ÎÓÒ¸, ÒÍ‡ÊÂÏ, Ì‡ ÔÓÔÛÎflˆËÓÌÌÓÈ‰ËÌ‡ÏËÍÂ Ì‡ÒÂÎÂÌËfl í‡ÒÏ‡ÌËË, ÏÌÓ„Ó Ú˚Òfl˜ ÎÂÚ ÍÓ-ÎÂ·‡‚¯Â„ÓÒfl „‰Â-ÚÓ Ì‡ ÛÓ‚ÌÂ 4000 ˜ÂÎÓ‚ÂÍ (Dia-mond, 1999). çÓ ÍË‚‡fl ‰ËÌ‡ÏËÍË Ó·˘ÂÈ ˜ËÒÎÂÌÌÓ-ÒÚË Ì‡ÒÂÎÂÌËfl ÏË‡ ÓÚ‡Ê‡ÂÚ ÔÂÊ‰Â ‚ÒÂ„Ó ÔÓÔÛ-ÎflˆËÓÌÌÛ˛ ‰ËÌ‡ÏËÍÛ åË-ëËÒÚÂÏ˚, ‡ ÌÂ ÚÂı ̃ ‡ÒÚÂÈ˜ÂÎÓ‚Â˜ÂÒÚ‚‡, ÍÓÚÓ˚Â ‚ ÌÂÂ ÌÂ ‚ıÓ‰ËÎË.

èÓ ‚ÒÂÈ ‚Ë‰ËÏÓÒÚË, Ë „ËÔÂ·ÓÎË˜ÂÒÍ‡fl ‰ËÌ‡ÏË-Í‡ ÓÒÚ‡ ‡ÁÌÓÓ·‡ÁËfl ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ ·ËÓÚ˚ ÓÚ-‡Ê‡Î‡ ‰ËÌ‡ÏËÍÛ ËÏÂÌÌÓ ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ “·ËÓÎÓ-„Ë˜ÂÒÍÓÈ åË-ëËÒÚÂÏ˚”, Ú.Â. ÙÎÓ Ë Ù‡ÛÌ ÚÂı Ï‡Ò-ÒË‚Ó‚ ÒÛ¯Ë, ÏÂÊ‰Û ÍÓÚÓ˚ÏË ËÏÂÎËÒ¸ ıÓÚfl ·˚˝ÔËÁÓ‰Ë˜ÂÒÍËÂ ÍÓÌÚ‡ÍÚ˚. è‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍËÂ‰‡ÌÌ˚Â ÔÓ ÚÂÏ ÍÛÔÌ˚Ï Ï‡ÒÒË‚‡Ï ÒÛ¯Ë, ÍÓÚÓ˚ÂÌ‡ıÓ‰ËÎËÒ¸ ‚ ËÁÓÎflˆËË ‰ÎËÚÂÎ¸ÌÓÂ ‚ÂÏfl, ÎË·Ó ÌÂ-‰ÓÒÚ‡ÚÓ˜ÌÓ ÔÓÎÌ˚, ˜ÚÓ·˚ ÒÛ‰ËÚ¸ Ó ÒÓÓÚ‚ÂÚÒÚ‚ËËÎÓÍ‡Î¸Ì˚ı ËÁÏÂÌÂÌËÈ ‡ÁÌÓÓ·‡ÁËfl „ËÔÂ·ÓÎË˜Â-ÒÍÓÈ ÏÓ‰ÂÎË (Ä‚ÒÚ‡ÎËfl, ÄÌÚ‡ÍÚË‰‡), ÎË·Ó ÌÂ ÔÓ-Í‡Á˚‚‡˛Ú ‚˚‡ÊÂÌÌÓÈ „ËÔÂ·ÓÎË˜ÂÒÍÓÈ ‰ËÌ‡ÏË-ÍË (ûÊÌ‡fl ÄÏÂËÍ‡) (Simpson, 1980).

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åÓ‰ÂÎ¸ „ËÔÂ·ÓÎË˜ÂÒÍÓ„Ó ÓÒÚ‡ ıÓÓ¯Ó ÓÔË-Ò˚‚‡ÂÚ Ù‡ÌÂÓÁÓÈÒÍÛ˛ ‰ËÌ‡ÏËÍÛ ‡ÁÌÓÓ·‡ÁËfl ÌÂÚÓÎ¸ÍÓ ÏÓÒÍÓÈ ·ËÓÚ˚ Ì‡ ÛÓ‚ÌÂ Ó‰Ó‚, Í‡Í ·˚ÎÓÔÓÍ‡Á‡ÌÓ ‡ÌÂÂ, ÌÓ Ë ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ ·ËÓÚ˚, ‡Ú‡ÍÊÂ ‚ÒÂÈ ·ËÓÚ˚ ‚ ˆÂÎÓÏ Ì‡ ÛÓ‚ÌÂ Ó‰Ó‚ Ë ÒÂ-ÏÂÈÒÚ‚ (‡ ‰Îfl ÏÓÒÍÓÈ ·ËÓÚ˚ – Ú‡ÍÊÂ Ì‡ ÛÓ‚ÌÂ ‚Ë-‰Ó‚). èÓ ‡Ì‡ÎÓ„ËË Ò Ï‡ÍÓÒÓˆËÓÎÓ„Ë˜ÂÒÍËÏË ÏÓ‰Â-ÎflÏË, ÓÔËÒ˚‚‡˛˘ËÏË „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ Ì‡ÒÂ-ÎÂÌËfl áÂÏÎË, ‡ Ú‡ÍÊÂ Ì‡ ÓÒÌÓ‚Â ËÏÂ˛˘ËıÒflÔ‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍËı ‰‡ÌÌ˚ı Ï˚ ÔÂ‰ÔÓÎ‡„‡ÂÏ, ˜ÚÓ„ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ·ËÓ‡ÁÌÓÓ·‡ÁËfl ‚ Ù‡ÌÂÓ-ÁÓÂ Ó·˙flÒÌflÂÚÒfl ÌÂÎËÌÂÈÌÓÈ ÔÓÎÓÊËÚÂÎ¸ÌÓÈ Ó·‡Ú-ÌÓÈ Ò‚flÁ¸˛ ‚ÚÓÓ„Ó ÔÓfl‰Í‡ ÏÂÊ‰Û ÓÒÚÓÏ ˜ËÒÎ‡Ú‡ÍÒÓÌÓ‚ Ë ËÁÏÂÌÂÌËflÏË ÒÚÛÍÚÛ˚ ÒÓÓ·˘ÂÒÚ‚.ùÚ‡ Ò‚flÁ¸ ÏÓÊÂÚ Â‡ÎËÁÓ‚˚‚‡Ú¸Òfl ‰‚ÓflÍÓ: ÓÒÚ Ó·-˘Â„Ó ‡ÁÌÓÓ·‡ÁËfl ·ËÓÚ˚ Ë ÓÚ‰ÂÎ¸Ì˚ı ÒÓÓ·˘ÂÒÚ‚(‡Î¸Ù‡-‡ÁÌÓÓ·‡ÁËfl) ÏÓÊÂÚ ÒÔÓÒÓ·ÒÚ‚Ó‚‡Ú¸ Í‡ÍÒÌËÊÂÌË˛ ‚ÂÓflÚÌÓÒÚË ‚˚ÏË‡ÌËfl Ú‡ÍÒÓÌÓ‚, Ú‡Í ËÔÓ‚˚¯ÂÌË˛ ‚ÂÓflÚÌÓÒÚË Ëı ÔÓfl‚ÎÂÌËfl: ·ÓÎ¸¯ÂÚ‡ÍÒÓÌÓ‚ – ‚˚¯Â ‡Î¸Ù‡-‡ÁÌÓÓ·‡ÁËÂ (ÒÂ‰ÌÂÂ ˜ËÒ-ÎÓ Ú‡ÍÒÓÌÓ‚ ‚ ÒÓÓ·˘ÂÒÚ‚Â) – ÒÓÓ·˘ÂÒÚ‚‡ ÒÚ‡ÌÓ‚flÚÒfl·ÓÎÂÂ ÒÎÓÊÌ˚ÏË Ë ÛÒÚÓÈ˜Ë‚˚ÏË – ÒÌËÊ‡ÂÚÒfl ÒÍÓ-ÓÒÚ¸ ‚˚ÏË‡ÌËfl Ë (ËÎË) ‡ÒÚÂÚ ÒÍÓÓÒÚ¸ ÔÓfl‚ÎÂ-ÌËfl – ‡ÁÌÓÓ·‡ÁËÂ ‡ÒÚÂÚ ·˚ÒÚÂÂ – ·ÓÎ¸¯Â Ú‡Í-ÒÓÌÓ‚ – ‚˚¯Â ‡Î¸Ù‡-‡ÁÌÓÓ·‡ÁËÂ Ë Ú.‰.

Ç ˝‚ÓÎ˛ˆËÓÌÌÓÈ ‰ËÌ‡ÏËÍÂ ÏÓÒÍÓÈ Ë ÍÓÌÚË-ÌÂÌÚ‡Î¸ÌÓÈ ·ËÓÚ ‚˚fl‚ÎflÂÚÒfl ˆÂÎ˚È fl‰ ÒıÓ‰Ì˚ı˜ÂÚ, Ò‚Ë‰ÂÚÂÎ¸ÒÚ‚Û˛˘Ëı Ó Â‰ËÌÒÚ‚Â ÏÂı‡ÌËÁÏÓ‚ ËÙ‡ÍÚÓÓ‚, ÓÔÂ‰ÂÎfl˛˘Ëı „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ‡ÁÌÓÓ·‡ÁËfl. ä‡Í ‚ îåÅ, Ú‡Í Ë ‚ îäÅ Ì‡·Î˛‰‡-ÂÚÒfl ÓÒÚ ÒÂ‰ÌÂÈ ÔÓ‰ÓÎÊËÚÂÎ¸ÌÓÒÚË ÒÛ˘ÂÒÚ‚Ó-‚‡ÌËfl Ú‡ÍÒÓÌÓ‚ ‚Ó ‚ÂÏÂÌË, Ô‡‡ÎÎÂÎ¸ÌÓÂ ÒÌËÊÂ-ÌËÂ “ÒÏÂÚÌÓÒÚË” Ë “ÓÊ‰‡ÂÏÓÒÚË”, ÒıÓ‰Ì˚È ËÒÛ-ÌÓÍ ÒÏÂÌ˚ ‰ÓÏËÌËÛ˛˘Ëı „ÛÔÔ, ÔÓÒÚÂÔÂÌÌÓÂÌ‡ÍÓÔÎÂÌËÂ ˝‚ÓÎ˛ˆËÓÌÌÓ ÛÒÚÓÈ˜Ë‚˚ı, ÔËÒÔÓÒÓ·-ÎflÂÏ˚ı, “ÙËÁËÓÎÓ„Ë˜ÂÒÍË Á‡·ÛÙÂÂÌÌ˚ı” ÙÓÏ Ò˝ÙÙÂÍÚË‚Ì˚ÏË ÒÔÓÒÓ·‡ÏË Á‡·ÓÚ˚ Ó ÔÓÚÓÏÒÚ‚Â ËÁ‡˘ËÚ˚ ‡ÌÌËı ÒÚ‡‰ËÈ ËÌ‰Ë‚Ë‰Û‡Î¸ÌÓ„Ó ‡Á‚ËÚËflË Ú.‰. ç‡ ‡ÌÌËı ˝Ú‡Ô‡ı ‡Á‚ËÚËfl îäÅ ‚˚fl‚ÎflÂÚÒflÒÛ˘ÂÒÚ‚ÂÌÌÓÂ ‡ÒıÓÊ‰ÂÌËÂ Ò ÔÂ‰ÒÍ‡Á‡ÌËflÏË „Ë-ÔÂ·ÓÎË˜ÂÒÍÓÈ ÏÓ‰ÂÎË (Â‡Î¸ÌÓ Ì‡·Î˛‰‡ÂÏÓÂ ˜ËÒ-ÎÓ Ú‡ÍÒÓÌÓ‚ ÌËÊÂ “ÔÂ‰ÒÍ‡Á‡ÌÌÓ„Ó”), ˜ÚÓ Ó·˙flÒÌfl-ÂÚÒfl, ‚Ó-ÔÂ‚˚ı, ÍÎ˛˜Â‚ÓÈ ÓÎ¸˛ ÌÂ ÒÓı‡Ìfl˛-˘ËıÒfl ‚ Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍÓÈ ÎÂÚÓÔËÒË „ÛÔÔ ‚Ì‡˜‡Î¸Ì˚È ÔÂËÓ‰ ÒÚ‡ÌÓ‚ÎÂÌËfl îäÅ, ‚Ó-‚ÚÓ˚ı,ÚÂÏ Ó·ÒÚÓflÚÂÎ¸ÒÚ‚ÓÏ, ˜ÚÓ îäÅ ‚ Ì‡˜‡ÎÂ Ò‚ÓÂ„Ó‡Á‚ËÚËfl ÔÂ‰ÒÚ‡‚ÎflÎ‡ ÒÓ·ÓÈ ÌÂ Ò‡ÏÓÒÚÓflÚÂÎ¸ÌÛ˛ÒËÒÚÂÏÛ, ‡ “ÓÚ‚ÂÚ‚ÎÂÌËÂ” ËÎË Ï‡„ËÌ‡Î¸ÌÛ˛ ˜‡ÒÚ¸îåÅ. ä ÒıÓ‰ÌÓÏÛ ‚˚‚Ó‰Û ÌÂÁ‡‚ËÒËÏÓ ÓÚ Ì‡Ò ÔË-¯ÂÎ Ä.Ä. îÓÏËÌ Ì‡ ÓÒÌÓ‚Â ‡Ì‡ÎËÁ‡ ËÏÂ˛˘ËıÒfl ‚ÎËÚÂ‡ÚÛÂ „‡ÙËÍÓ‚ ‰ËÌ‡ÏËÍË ‡ÁÌÓÓ·‡ÁËfl îåÅË îäÅ (Ä.Ä. îÓÏËÌ, ÎË˜ÌÓÂ ÒÓÓ·˘.).

ç‡Ë·ÓÎÂÂ ˜ÂÚÍÓ ‚˚‡ÊÂÌÌ‡fl „ËÔÂ·ÓÎË˜ÂÒÍ‡fl‰ËÌ‡ÏËÍ‡ ı‡‡ÍÚÂÌ‡ ‰Îfl ‚ÒÂÈ ·ËÓÚ˚ ‚ ˆÂÎÓÏ. ùÚÓÏÓÊÌÓ ‡ÒÒÏ‡ÚË‚‡Ú¸ Í‡Í ÔÓ‰Ú‚ÂÊ‰ÂÌËÂ Â‰ËÌ-ÒÚ‚‡ ·ËÓÒÙÂ˚, ÍÓÚÓ‡fl ‚ Ù‡ÌÂÓÁÓÂ ‡Á‚Ë‚‡Î‡Ò¸‚Ó ÏÌÓ„Ëı ÓÚÌÓ¯ÂÌËflı Í‡Í ˆÂÎÓÒÚÌ‡fl ÒËÒÚÂÏ‡.

ÉËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ·ËÓ‡ÁÌÓÓ·‡ÁËfl Ò‚Ë‰Â-ÚÂÎ¸ÒÚ‚ÛÂÚ Ó ÚÓÏ, ̃ ÚÓ ‚‡ÊÌÛ˛ ÓÎ¸ ‚ ̋ ‚ÓÎ˛ˆËË ÏÓ-„ÛÚ Ë„‡Ú¸ ÌÂ ÚÓÎ¸ÍÓ ÍÓÌÍÛÂÌÚÌ˚Â, ÌÓ Ë ÍÓÓÔÂ‡-ÚË‚Ì˚Â ‚Á‡ËÏÓÓÚÌÓ¯ÂÌËfl ÏÂÊ‰Û ̋ ‚ÓÎ˛ˆËÓÌËÛ˛-˘ËÏË ÍÎ‡‰‡ÏË. å‡ÍÓ˝‚ÓÎ˛ˆËÓÌÌ‡fl ÍÓÓÔÂ‡ˆËflÏÓÊÂÚ ÔÓfl‚ÎflÚ¸Òfl Í‡Í ‚ ÔÓ‚˚¯ÂÌËË ÛÒÚÓÈ˜Ë‚Ó-ÒÚË ÒÓÓ·˘ÂÒÚ‚, Ú‡Í Ë ‚ ÚÓÏ, ˜ÚÓ ‚ÓÁÌËÍÌÓ‚ÂÌËÂ ÌÓ-‚˚ı ‚Ë‰Ó‚ ÒÔÓÒÓ·ÒÚ‚ÛÂÚ ÔÓfl‚ÎÂÌË˛ ÌÓ‚˚ı ÌË¯, ÍÓ-ÚÓ˚Â ÏÓ„ÛÚ ·˚Ú¸ Á‡ÔÓÎÌÂÌ˚ ÒÎÂ‰Û˛˘ËÏ “ÔÓÍÓÎÂ-ÌËÂÏ” ÌÓ‚˚ı ‚Ë‰Ó‚. ù‚ÓÎ˛ˆËfl ·ËÓ‡ÁÌÓÓ·‡ÁËfl,Ú‡ÍËÏ Ó·‡ÁÓÏ, ÏÓÊÂÚ ‡ÒÒÏ‡ÚË‚‡Ú¸Òfl Í‡Í Ò‡ÏÓ-ÛÒÍÓfl˛˘ËÈÒfl, ˝ÒÍ‡Î‡ˆËÓÌÌ˚È ÔÓˆÂÒÒ.

ê‡·ÓÚ‡ ÔÓ‰‰ÂÊ‡Ì‡ èÓ„‡ÏÏÓÈ 25 èÂÁË‰ËÛ-Ï‡ êÄç “èÓËÒıÓÊ‰ÂÌËÂ Ë ˝‚ÓÎ˛ˆËfl ·ËÓÒÙÂ˚”,ÙÓÌ‰ÓÏ ÒÓ‰ÂÈÒÚ‚Ëfl ÓÚÂ˜ÂÒÚ‚ÂÌÌÓÈ Ì‡ÛÍÂ Ë êîîà(„‡ÌÚ˚ 06-05-39015, 06-05-64641 Ë 06-06-80503).

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åÓÊÂÚ ÎË „ËÔÂ·ÓÎË˜ÂÒÍËÈ ÓÒÚ ·ËÓ‡ÁÌÓÓ·‡ÁËfl ·˚Ú¸ ‡ÚÂÙ‡ÍÚÓÏ ÌÂÔÓÎÌÓÚ˚

Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍÓÈ ÎÂÚÓÔËÒË?

èÓ·ÎÂÏ‡ ‰ÓÒÚÓ‚ÂÌÓÒÚË ÍÓÎË˜ÂÒÚ‚ÂÌÌ˚ı ÓˆÂ-ÌÓÍ ·ËÓ‡ÁÌÓÓ·‡ÁËfl ÏËÌÛ‚¯Ëı „ÂÓÎÓ„Ë˜ÂÒÍËı˝ÔÓı ÔÓ‰Ó·ÌÓ Ó·ÒÛÊ‰‡Î‡Ò¸ Ì‡ÏË ‡ÌÂÂ (å‡ÍÓ‚,äÓÓÚ‡Â‚, 2007; Markov, Korotayev, 2007). Ç ˆÂÎÓÏÌ‡ ÒÂ„Ó‰Ìfl¯ÌËÈ ‰ÂÌ¸ „ËÔÓÚÂÁ‡, ÒÓ„Î‡ÒÌÓ ÍÓÚÓÓÈ‰‡ÌÌ˚Â Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍÓÈ ÎÂÚÓÔËÒË ‡‰ÂÍ‚‡ÚÌÓÓÚ‡Ê‡˛Ú Â‡Î¸Ì˚Â ËÁÏÂÌÂÌËfl ·ËÓ‡ÁÌÓÓ·‡ÁËfl ‚ÔÓ¯ÎÓÏ, ÓÒÚ‡ÂÚÒfl ÌÂ ÓÔÓ‚Â„ÌÛÚÓÈ Ë ÔÓ-ÔÂÊÌÂ-ÏÛ fl‚ÎflÂÚÒfl Ì‡Ë·ÓÎÂÂ Ì‡‰ÂÊÌÓÈ ÓÒÌÓ‚ÓÈ ‰Îfl ÒÓ-‰ÂÊ‡ÚÂÎ¸Ì˚ı ·ËÓÎÓ„Ë˜ÂÒÍËı ËÌÚÂÔÂÚ‡ˆËÈ.

íÂÏ ÌÂ ÏÂÌÂÂ ‚ÓÔÓÒ Ó ÚÓÏ, ÌÂ fl‚ÎflÂÚÒfl ÎË ‡ÚÂ-Ù‡ÍÚÓÏ ËÏÂÌÌÓ „ËÔÂ·ÓÎË˜ÂÒÍËÈ ı‡‡ÍÚÂ ÓÒÚ‡·ËÓ‡ÁÌÓÓ·‡ÁËfl, Á‡ÒÎÛÊË‚‡ÂÚ ÓÚ‰ÂÎ¸ÌÓ„Ó ‡Ò-ÒÏÓÚÂÌËfl. ëıÓ‰ÒÚ‚Ó ÍË‚˚ı ·ËÓ‡ÁÌÓÓ·‡ÁËfl Ò „Ë-ÔÂ·ÓÎÓÈ ‚Ó ÏÌÓ„ÓÏ Ó·ÛÒÎÓ‚ÎÂÌÓ Ëı ÂÁÍËÏ ÔÓ‰˙-ÂÏÓÏ ‚ ÍÓÌˆÂ ÏÂÁÓÁÓfl Ë ‚ Í‡ÈÌÓÁÓÂ. åÓÊÌÓ ÔÂ‰ÔÓ-ÎÓÊËÚ¸, ˜ÚÓ ˝ÚÓÚ ÂÁÍËÈ ÔÓ‰˙ÂÏ, ıÓÌÓÎÓ„Ë˜ÂÒÍËÔËÛÓ˜ÂÌÌ˚È Í Ò‡‚ÌËÚÂÎ¸ÌÓ ÌÂ‰‡‚ÌËÏ ˝ÔÓı‡Ï,Ó·ÛÒÎÓ‚ÎÂÌ ÌÂÔÓÎÌÓÚÓÈ ÎÂÚÓÔËÒË ‚ ÒÓ˜ÂÚ‡ÌËË Ò ˝Ù-ÙÂÍÚÓÏ “ÔËÚflÊÂÌËfl ÒÓ‚ÂÏÂÌÌÓÒÚË” (Raup, 1979).å˚ ÔÓÔ˚Ú‡ÎËÒ¸ ‚ÂËÙËˆËÓ‚‡Ú¸ ˝ÚÓ ÔÂ‰ÔÓÎÓÊÂ-ÌËÂ ÔË ÔÓÏÓ˘Ë ËÏËÚ‡ˆËÓÌÌÓ„Ó ÍÓÏÔ¸˛ÚÂÌÓ„ÓÏÓ‰ÂÎËÓ‚‡ÌËfl.

éÔËÒ‡ÌËÂ ÏÓ‰ÂÎË. àÏËÚËÓ‚‡ÎÓÒ¸ ‡Á‚ËÚËÂ·ËÓÚ˚ ‚ ÚÂ˜ÂÌËÂ 100 ÔÓÒÎÂ‰Ó‚‡ÚÂÎ¸Ì˚ı ‚ÂÏÂÌ-Ì˚ı ËÌÚÂ‚‡ÎÓ‚ (“‚ÂÍÓ‚”). Ç ÏÓÏÂÌÚ ‚ÂÏÂÌË 1 ÔÓ-fl‚ÎflÂÚÒfl 100 ÔÂ‚˚ı Ú‡ÍÒÓÌÓ‚ (“Ó‰Ó‚”); ‚ ‰‡Î¸-ÌÂÈ¯ÂÏ ÔÓfl‚ÎÂÌËÂ Ë ‚˚ÏË‡ÌËÂ Ó‰Ó‚ ÔÓËÒıÓ‰flÚÔÓ ÓÔÂ‰ÂÎÂÌÌÓÏÛ ‡Î„ÓËÚÏÛ (ÒÏ. ÌËÊÂ). ä‡Ê‰ÓÏÛÓ‰Û ËÁÌ‡˜‡Î¸ÌÓ ÔËÒ‚‡Ë‚‡ÂÚÒfl ‚ÂÎË˜ËÌ‡ pi, ÍÓÚÓ-‡fl ÓÔÂ‰ÂÎflÂÚÒfl Í‡Í ÒÎÛ˜‡ÈÌÓÂ ˜ËÒÎÓ ‚ ‰Ë‡Ô‡ÁÓÌÂÓÚ 0 ‰Ó 1. ÇÂÎË˜ËÌ‡ p ı‡‡ÍÚÂËÁÛÂÚ ÒÓ‚ÓÍÛÔÌÓÒÚ¸‚ÒÂı Ô‡‡ÏÂÚÓ‚, ÓÚ ÍÓÚÓ˚ı Á‡‚ËÒËÚ ‚ÂÓflÚÌÓÒÚ¸ÔÓÔ‡‰‡ÌËfl ‰‡ÌÌÓ„Ó Ó‰‡ ‚ Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍÛ˛ ÎÂ-ÚÓÔËÒ¸ (˜ËÒÎÂÌÌÓÒÚ¸ Ó‰‡, Ì‡ÎË˜ËÂ ıÓÓ¯Ó ÒÓı‡-Ìfl˛˘ËıÒfl ÒÍÂÎÂÚÌ˚ı ˜‡ÒÚÂÈ, ÔÎÓ˘‡‰¸ ‡Â‡Î‡ Ë



Ú.‰.). êÂ‡Î¸Ì˚Â Ú‡ÍÒÓÌ˚ ÒËÎ¸ÌÓ ÓÚÎË˜‡˛ÚÒfl ‰Û„ÓÚ ‰Û„‡ ÔÓ ˝ÚËÏ ı‡‡ÍÚÂËÒÚËÍ‡Ï, ÔÓ˝ÚÓÏÛ ‚ÂÓ-flÚÌÓÒÚ¸ Ëı ÔÓÔ‡‰‡ÌËfl ‚ ÎÂÚÓÔËÒ¸ ‰‡ÎÂÍÓ ÌÂ Ó‰ËÌ‡-ÍÓ‚‡. àÏÂÌÌÓ ˝ÚË ‡ÁÎË˜Ëfl Ë ÔËÁ‚‡Ì‡ ÓÚÓ·‡ÁËÚ¸‚ÂÎË˜ËÌ‡ pi, ‡ÁÌ‡fl Û ‡ÁÌ˚ı Ó‰Ó‚. èÓÏËÏÓ ÏÂÊ-Ó‰Ó‚˚ı ‡ÁÎË˜ËÈ ÒÛ˘ÂÒÚ‚Û˛Ú Ë ‡ÁÎË˜Ëfl ‚Ó ‚Â-ÏÂÌË (Ì‡ÔËÏÂ, Ó‰ËÌ Ë ÚÓÚ ÊÂ Ó‰ ÏÓÊÂÚ ·˚Ú¸Ï‡ÒÒÓ‚˚Ï ‚ Ó‰ÌÓÏ ‚ÂÍÂ Ë ·ÓÎÂÂ Â‰ÍËÏ – ‚ ‰Û„ÓÏ,˜ÚÓ ÓÚ‡Ê‡ÂÚÒfl Ì‡ ‚ÂÓflÚÌÓÒÚË Â„Ó ÔÓÔ‡‰‡ÌËfl ‚ ÎÂ-ÚÓÔËÒ¸). èÓ˝ÚÓÏÛ Í‡Ê‰ÓÏÛ Ó‰Û ‚ Í‡Ê‰ÓÏ ‚ÂÍÂ, ‚ÚÂ˜ÂÌËÂ ÍÓÚÓÓ„Ó Ó‰ ÒÛ˘ÂÒÚ‚Ó‚‡Î, ÔËÒ‚‡Ë‚‡ÂÚÒfl‚ÂÎË˜ËÌ‡ pij = r1 × pi, „‰Â r1 – ÒÎÛ˜‡ÈÌÓÂ ˜ËÒÎÓ.

ÖÒÎË ‚ÂÎË˜ËÌ‡ r2 × pij ÓÍ‡Á˚‚‡ÂÚÒfl ÏÂÌ¸¯Â ËÁÌ‡-˜‡Î¸ÌÓ Á‡‰‡ÌÌÓÈ ÔÓÎ¸ÁÓ‚‡ÚÂÎÂÏ ‚ÂÎË˜ËÌ˚, ı‡‡Í-ÚÂËÁÛ˛˘ÂÈ ËÌÚÂÌÒË‚ÌÓÒÚ¸ ‚˚ÏË‡ÌËfl (ÔÓÒÚÓflÌ-ÌÛ˛ ‰Îfl ‚ÒÂı Ó‰Ó‚ Ë ‚ÂÍÓ‚), ÚÓ Ó‰ ‚˚ÏË‡ÂÚ, ÌÂÔÂÂÈ‰fl ËÁ ‚ÂÍ‡ j ‚ ‚ÂÍ j + 1. ÅÎ‡„Ó‰‡fl ÌÂÊÂÒÚÍÓÈÔÓÎÓÊËÚÂÎ¸ÌÓÈ Ò‚flÁË ÏÂÊ‰Û pij Ë ‚ÂÓflÚÌÓÒÚ¸˛ ÔÂ-ÂıÓ‰‡ Ó‰‡ ‚ ÒÎÂ‰Û˛˘ËÈ ‚ÂÍ ÏÓ‰ÂÎ¸ Û˜ËÚ˚‚‡ÂÚÚÓÚ Ó˜Â‚Ë‰Ì˚È Ù‡ÍÚ, ˜ÚÓ Û Ú‡ÍÒÓÌ‡ ÂÒÚ¸ Ò‚ÓÈÒÚ‚‡(Ú‡ÍËÂ, Í‡Í Ï„ÌÓ‚ÂÌÌ‡fl ˜ËÒÎÂÌÌÓÒÚ¸ Ë ÔÎÓ˘‡‰¸‡Â‡Î‡), ÍÓÚÓ˚Â ÍÓÂÎËÛ˛Ú Ó‰ÌÓ‚ÂÏÂÌÌÓ Ò ‚Â-ÓflÚÌÓÒÚ¸˛ Ë Â„Ó ÔÂÂıÓ‰‡ ‚ ÒÎÂ‰Û˛˘ËÈ ‚ÂÍ, Ë ÚÓ-„Ó, ̃ ÚÓ Â„Ó ÓÒÚ‡ÚÍË ·Û‰ÛÚ Ó·Ì‡ÛÊÂÌ˚ Ô‡ÎÂÓÌÚÓÎÓ-„‡ÏË ‚ ÓÚÎÓÊÂÌËflı ‰‡ÌÌÓ„Ó ‚ÂÍ‡.

èÓfl‚ÎÂÌËÂ ÌÓ‚˚ı Ó‰Ó‚ ÏÓ‰ÂÎËÓ‚‡ÎÓÒ¸ ‰‚ÛÏfl‡ÁÌ˚ÏË ÒÔÓÒÓ·‡ÏË, ÒÓÓÚ‚ÂÚÒÚ‚Û˛˘ËÏË ‰‚ÛÏ ·‡-ÁÓ‚˚Ï ÏÓ‰ÂÎflÏ ‰Ë‚ÂÒËÙËÍ‡ˆËË: ˝ÍÒÔÓÌÂÌˆË‡Î¸-ÌÓÈ Ë ÎÓ„ËÒÚË˜ÂÒÍÓÈ. Ç ÔÂ‚ÓÏ ÒÎÛ˜‡Â ˜ËÒÎÓ Ó-‰Ó‚, ÔÓfl‚Ë‚¯ËıÒfl ‚ ‰‡ÌÌÓÏ ‚ÂÍÂ, ÓÔÂ‰ÂÎflÎÓÒ¸ Í‡Ík1N, „‰Â N – ̃ ËÒÎÓ Ó‰Ó‚, ÔÂÂ¯Â‰¯Ëı ‚ ‰‡ÌÌ˚È ‚ÂÍË ÔÂ‰˚‰Û˘Â„Ó, k1 – Á‡‰‡ÌÌ‡fl ÔÓÎ¸ÁÓ‚‡ÚÂÎÂÏ ‚ÂÎË-˜ËÌ‡, ı‡‡ÍÚÂËÁÛ˛˘‡fl ÚÂÏÔ ˝ÍÒÔÓÌÂÌˆË‡Î¸ÌÓÈ‰Ë‚ÂÒËÙËÍ‡ˆËË. ÇÓ ‚ÚÓÓÏ ÒÎÛ˜‡Â ˜ËÒÎÓ ÌÓ‚˚ıÓ‰Ó‚ ÓÔÂ‰ÂÎflÎÓÒ¸ Í‡Í k2N*(Nmax – N).

í‡ÍËÏ ÔÛÚÂÏ „ÂÌÂËÓ‚‡Î‡Ò¸ “Â‡Î¸Ì‡fl” ‰ËÌ‡-ÏËÍ‡ ‡Á‚ËÚËfl ÏÓ‰ÂÎ¸ÌÓÈ ·ËÓÚ˚. á‡ÚÂÏ ÓÒÛ˘ÂÒÚ‚-ÎflÎÓÒ¸ ÏÓ‰ÂÎËÓ‚‡ÌËÂ “Ò·Ó‡ Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍÓ-„Ó Ï‡ÚÂË‡Î‡”. èË ̋ ÚÓÏ Á‡‰‡‚‡ÎËÒ¸ ‡ÁÎË˜Ì˚Â ‚Â-ÎË˜ËÌ˚ “ÔÓÎÌÓÚ˚ ÎÂÚÓÔËÒË” c. ùÚÓÚ ÔÓÍ‡Á‡ÚÂÎ¸,‚‡¸ËÛ˛˘ËÈ ÓÚ ÌÛÎfl (ÌÛÎÂ‚‡fl ÒÓı‡ÌÌÓÒÚ¸ ËÒÍÓ-Ô‡ÂÏ˚ı) ‰Ó Â‰ËÌËˆ˚ (ÔÓÎÌ‡fl ÒÓı‡ÌÌÓÒÚ¸), ÓÚ‡-Ê‡ÂÚ ÒÓ‚ÓÍÛÔÌÓÒÚ¸ ‚ÒÂı Ù‡ÍÚÓÓ‚, ‚ÎËfl˛˘Ëı Ì‡ÔÓÎÌÓÚÛ ÔÂ‰ÒÚ‡‚ÎÂÌÌÓÒÚË ‚ Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍËıÍÓÎÎÂÍˆËflı Â‡Î¸ÌÓ„Ó ‡ÁÌÓÓ·‡ÁËfl, ÒÛ˘ÂÒÚ‚Ó‚‡‚-¯Â„Ó ‚ ‰‡ÌÌÓÏ ‚ÂÍÂ: ‡ÒÔÓÒÚ‡ÌÂÌÌÓÒÚ¸ ÓÚÎÓÊÂ-ÌËÈ ‰‡ÌÌÓ„Ó ‚ÓÁ‡ÒÚ‡, ÒÚÂÔÂÌ¸ ÎËÚËÙËˆËÓ‚‡ÌÌÓ-ÒÚË ÔÓÓ‰, ËÒÒÎÂ‰Ó‚‡ÚÂÎ¸ÒÍÓÂ ÛÒËÎËÂ Ë Ú.‰. ÄÌ‡ÎË-ÁËÓ‚‡ÎË ‡ÁÎË˜Ì˚Â ‚‡Ë‡ÌÚ˚ ËÁÏÂÌÂÌËfl ÔÓÎÌÓÚ˚Ò ‚ÓÁ‡ÒÚÓÏ ÓÚÎÓÊÂÌËÈ: 1) ÔÓÒÚÓflÌÌ‡ ‚ ÚÂ˜ÂÌËÂ ‚ÒÂ-„Ó ËÌÚÂ‚‡Î‡ ‚ÂÏÂÌË, 2) ÎËÌÂÈÌÓ ÒÌËÊ‡ÂÚÒfl Ò ‚ÓÁ-‡ÒÚÓÏ ÓÚÎÓÊÂÌËÈ, 3) ÒÌËÊ‡ÂÚÒfl ÔÓ ̋ ÍÒÔÓÌÂÌˆË‡Î¸-ÌÓÏÛ Á‡ÍÓÌÛ (‚Ì‡˜‡ÎÂ ·˚ÒÚÓ, Á‡ÚÂÏ ‚ÒÂ ÏÂ‰ÎÂÌ-ÌÂÂ), 4) ÒÌËÊ‡ÂÚÒfl ÔÓ „ËÔÂ·ÓÎË˜ÂÒÍÓÏÛ Á‡ÍÓÌÛ:‚Ì‡˜‡ÎÂ ÓÌ‡ Ó˜ÂÌ¸ ‚˚ÒÓÍ‡, Á‡ÚÂÏ Ò‡ÁÛ Ì‡˜ËÌ‡ÂÚÒÚÂÏËÚÂÎ¸ÌÓ Û·˚‚‡Ú¸ Ë ÓÒÚ‡ÂÚÒfl Í‡ÈÌÂ ÌËÁÍÓÈÌ‡ ÔÓÚflÊÂÌËË ·ÓÎ¸¯ÂÈ ˜‡ÒÚË ‚ÂÏÂÌÌÓ„Ó ËÌÚÂ-‚‡Î‡. ÑÎfl ÔÓÒÎÂ‰ÌÂ„Ó (ÒÓÚÓ„Ó) ‚ÂÍ‡, ÍÓÚÓ˚È Ò˜Ë-

Ú‡ÎÒfl ‡Ì‡ÎÓ„Ë˜Ì˚Ï ÒÓ‚ÂÏÂÌÌÓÒÚË („ÓÎÓˆÂÌÛ),ÔÓÎÌÓÚ‡ ‚ÒÂ„‰‡ Á‡‰‡‚‡Î‡Ò¸ ‡‚ÌÓÈ Â‰ËÌËˆÂ (‰ÎflËÏËÚ‡ˆËË ‚ ÏÓ‰ÂÎË ˝ÙÙÂÍÚ‡ “ÔËÚflÊÂÌËfl ÒÓ‚Â-ÏÂÌÌÓÒÚË”). èÓÔ‡‚¯ËÏË ‚ ÍÓÎÎÂÍˆËË, ÒÓ·‡ÌÌ˚ÂËÁ ÓÚÎÓÊÂÌËÈ ‚ÂÍ‡ j, Ò˜ËÚ‡ÎËÒ¸ ÚÂ Ó‰˚, Û ÍÓÚÓ˚ı‚ÂÎË˜ËÌ‡ pij ‚ ‰‡ÌÌÓÏ ‚ÂÍÂ ÔÂ‚˚¯‡Î‡ ‚ÂÎË˜ËÌÛ1 – c. Ç ÂÁÛÎ¸Ú‡ÚÂ Â‡Î¸Ì˚È ËÌÚÂ‚‡Î ÒÚ‡ÚË„‡-ÙË˜ÂÒÍÓ„Ó ‡ÒÔÓÒÚ‡ÌÂÌËfl Û ÌÂÍÓÚÓ˚ı Ó‰Ó‚ÓÍ‡Á˚‚‡ÎÒfl ÛÒÂ˜ÂÌÌ˚Ï, ÔÂ˚‚ËÒÚ˚Ï, ËÎË Ó‰ ‚Ó-‚ÒÂ ÌÂ ÔÓÔ‡‰‡Î ‚ “ÍÓÎÎÂÍˆËË”. ç‡ ÓÒÌÓ‚Â ˝ÚËı ‰‡Ì-Ì˚ı ÒÚÓËÎÒfl „‡ÙËÍ “Ì‡·Î˛‰‡ÂÏÓÈ” ‰ËÌ‡ÏËÍË‡ÁÌÓÓ·‡ÁËfl, ÔË˜ÂÏ Ó‰ Ò˜ËÚ‡ÎÒfl ÒÛ˘ÂÒÚ‚Ó‚‡‚-¯ËÏ Ì‡ ÔÓÚflÊÂÌËË ‚ÒÂ„Ó ËÌÚÂ‚‡Î‡ ÓÚ Â„Ó ÔÂ‚Ó-„Ó ‰Ó ÔÓÒÎÂ‰ÌÂ„Ó ÔÓfl‚ÎÂÌËfl ‚ Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍËıÍÓÎÎÂÍˆËflı. èË ˝ÚÓÏ Ó‰˚, ‰ÓÊË‚¯ËÂ ‰Ó ÒÓ‚Â-ÏÂÌÌÓÒÚË (100-„Ó ËÌÚÂ‚‡Î‡) Ë ÔÓÔ‡‚¯ËÂ ‚ Ô‡ÎÂÓÌ-ÚÓÎÓ„Ë˜ÂÒÍËÂ ÍÓÎÎÂÍˆËË ıÓÚfl ·˚ ‚ Ó‰ÌÓÏ ËÁ ‚ÂÍÓ‚(Ò 1-„Ó ÔÓ 99-È), Ò˜ËÚ‡ÎËÒ¸ ÒÛ˘ÂÒÚ‚Ó‚‡‚¯ËÏË ‚ ÚÂ-˜ÂÌËÂ ‚ÒÂ„Ó ‚ÂÏÂÌË ÓÚ ˝ÚÓ„Ó ‚ÂÍ‡ ‰Ó ÒÓ‚ÂÏÂÌÌÓ-ÒÚË (˝ÙÙÂÍÚ “ÔËÚflÊÂÌËfl ÒÓ‚ÂÏÂÌÌÓÒÚË”).

éˆÂÌÍ‡ Ô‡‡ÏÂÚÓ‚ ÏÓ‰ÂÎË. é Â‡ÎËÒÚË˜ÌÓÒÚË‚ıÓ‰Ì˚ı Ô‡‡ÏÂÚÓ‚ ÏÓ‰ÂÎË ÏÓÊÌÓ ÒÛ‰ËÚ¸ ÔÓ ÒÓ-ÓÚ‚ÂÚÒÚ‚Ë˛ ÂÁÛÎ¸Ú‡ÚÓ‚ ÏÓ‰ÂÎËÓ‚‡ÌËfl Â‡Î¸Ì˚Ï‰‡ÌÌ˚Ï. Ç ÎËÚÂ‡ÚÛÂ ÔÂ‰ÎÓÊÂÌÓ ÌÂÒÍÓÎ¸ÍÓ ÒÔÓ-ÒÓ·Ó‚ ÍÓÎË˜ÂÒÚ‚ÂÌÌÓÈ ÓˆÂÌÍË ÒÚÂÔÂÌË ÔÓÎÌÓÚ˚ÎÂÚÓÔËÒË, ‰‡˛˘Ëı ‚ ˆÂÎÓÏ ÒıÓ‰Ì˚Â ÂÁÛÎ¸Ú‡Ú˚.êËÒÛÌÓÍ 8,‡ ‰‡ÂÚ ÔÂ‰ÒÚ‡‚ÎÂÌËÂ Ó Â‡Î¸ÌÓÈ ‚ÂÎË-˜ËÌÂ ÌÂÔÓÎÌÓÚ˚ ÎÂÚÓÔËÒË Ì‡ Ó‰Ó‚ÓÏ Ë ÒÂÏÂÈ-ÒÚ‚ÂÌÌÓÏ ÛÓ‚Ìflı. ÑÎfl ÏÓÒÍËı ÊË‚ÓÚÌ˚ı, ËÏÂ˛-˘Ëı ÒÍÂÎÂÚ, ÒÓı‡ÌflÂÏÓÒÚ¸ ÒÂÏÂÈÒÚ‚ ÒÓÒÚ‡‚ÎflÂÚ ‚ÒÂ‰ÌÂÏ ÔËÏÂÌÓ 50–80%, Ó‰Ó‚ – 20–60% (·ÂÒ-ÒÍÂÎÂÚÌ˚Â ÊË‚ÓÚÌ˚Â ÒÓÒÚ‡‚Îfl˛Ú ÌË˜ÚÓÊÌÛ˛ ‰Ó-Î˛ ËÒÍÓÔ‡ÂÏÓ„Ó ÏÓÒÍÓ„Ó ‡ÁÌÓÓ·‡ÁËfl Ë ÌÂ ‚ÎËfl-˛Ú Ì‡ Ó·˘ËÈ ‚Ë‰ ÍË‚˚ı). ÑÎfl ÚÂı ‚‡ÊÌÂÈ¯Ëı„ÛÔÔ ÍÓÌÚËÌÂÌÚ‡Î¸Ì˚ı Ó„‡ÌËÁÏÓ‚ (‚˚Ò¯ËÂ ‡Ò-ÚÂÌËfl, ÚÂÚ‡ÔÓ‰˚, Ì‡ÒÂÍÓÏ˚Â), ÍÓÚÓ˚Â Ô‡ÍÚË˜Â-ÒÍË ÔÓÎÌÓÒÚ¸˛ ÓÔÂ‰ÂÎfl˛Ú ÙÓÏÛ ÍË‚˚ı ‡ÁÌÓ-Ó·‡ÁËfl îäÅ, ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË, ÔÓ Í‡ÈÌÂÈ ÏÂÂÌ‡ ÛÓ‚ÌÂ ÒÂÏÂÈÒÚ‚, Ì‡ıÓ‰ËÚÒfl ‚ ÚÂı ÊÂ ÔÂ‰ÂÎ‡ı,ıÓÚfl ˝ÚÓ Ë ÔÓÚË‚ÓÂ˜ËÚ ËÌÚÛËÚË‚Ì˚Ï ÓÊË‰‡ÌËflÏ(ÔËÌflÚÓ Ò˜ËÚ‡Ú¸, ˜ÚÓ ÏÓÒÍ‡fl Ô‡ÎÂÓÌÚÓÎÓ„Ë˜Â-ÒÍ‡fl ÎÂÚÓÔËÒ¸ “ÎÛ˜¯Â” ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ) (Benton,Simms, 1995). ç‡ÔËÏÂ, ËÁ ÒÓ‚ÂÏÂÌÌ˚ı ÒÂÏÂÈÒÚ‚Ì‡ÒÂÍÓÏ˚ı ÌÂ ÏÂÌÂÂ 60% ÔÂ‰ÒÚ‡‚ÎÂÌ˚ ‚ Ô‡ÎÂÓÌ-ÚÓÎÓ„Ë˜ÂÒÍÓÈ ÎÂÚÓÔËÒË (Labandeira, Sepkoski,1993). éÚÌÓÒËÚÂÎ¸Ì‡fl ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË Ì‡ÒÂÍÓ-Ï˚ı Ë ‚˚Ò¯Ëı ‡ÒÚÂÌËÈ, ‚ÂÓflÚÌÓ, Ó·ÂÒÔÂ˜Ë‚‡ÂÚÒflËı ‚˚ÒÓÍÓÈ ˜ËÒÎÂÌÌÓÒÚ¸˛ Ë ·ËÓÏ‡ÒÒÓÈ, ÚÂÚ‡ÔÓ‰ –Ï‡ÍÒËÏ‡Î¸Ì˚Ï “ËÒÒÎÂ‰Ó‚‡ÚÂÎ¸ÒÍËÏ ÛÒËÎËÂÏ”.

ùÚË ‰‡ÌÌ˚Â ÔÓÏÓ„‡˛Ú ÓˆÂÌËÚ¸ ÒÚÂÔÂÌ¸ Â‡ÎË-ÒÚË˜ÌÓÒÚË ËÏËÚ‡ˆËÓÌÌÓ„Ó ÏÓ‰ÂÎËÓ‚‡ÌËfl. í‡Í, Ì‡-ÔËÏÂ, ÂÒÎË ËÁ 2000 “ÒÓ‚ÂÏÂÌÌ˚ı” Ó‰Ó‚ (ÒÛ˘Â-ÒÚ‚Û˛˘Ëı ‚ ÒÓÚÓÏ ‚ÂÍÂ), ÚÓÎ¸ÍÓ 20 ËÎË 100 (1–5%)ÔÓÔ‡ÎË ‚ ËÒÍÓÔ‡ÂÏÛ˛ ÎÂÚÓÔËÒ¸, ÏÓÊÌÓ Û‚ÂÂÌÌÓ„Ó‚ÓËÚ¸ Ó ÚÓÏ, ˜ÚÓ ÌÂÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË ·˚Î‡ ‚ÏÓ‰ÂÎË ÒËÎ¸ÌÓ Á‡‚˚¯ÂÌ‡.

êÂÁÛÎ¸Ú‡Ú˚ ÏÓ‰ÂÎËÓ‚‡ÌËfl (ËÒ. 8,·-Ë) ÔÓÍ‡-Á˚‚‡˛Ú, ˜ÚÓ ÔË ˝ÍÔÓÌÂÌˆË‡Î¸ÌÓÈ ‰Ë‚ÂÒËÙËÍ‡-

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êËÒ. 8. ÇÎËflÌËÂ ‰ËÙÙÂÂÌˆË‡Î¸ÌÓÈ ÌÂÔÓÎÌÓÚ˚ Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍÓÈ ÎÂÚÓÔËÒË Ì‡ ı‡‡ÍÚÂ Ì‡·Î˛‰‡ÂÏÓÈ ‰ËÌ‡ÏËÍË ‡Á-ÌÓÓ·‡ÁËfl. ‡ – ÒÓı‡ÌflÂÏÓÒÚ¸ Ó‰Ó‚ Ë ÒÂÏÂÈÒÚ‚ ÏÓÒÍËı ÊË‚ÓÚÌ˚ı ‚ Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍÓÈ ÎÂÚÓÔËÒË. ãËÌËfl – ÒÓÓÚÌÓ¯Â-ÌËÂ ÒÓı‡ÌflÂÏÓÒÚË Ó‰Ó‚ Ë ÒÂÏÂÈÒÚ‚, ‚˚˜ËÒÎÂÌÌÓÂ Ì‡ ÓÒÌÓ‚Â ‡Ì‡ÎËÁ‡ ÍÓÎË˜ÂÒÚ‚ÂÌÌ˚ı ÒÓÓÚÌÓ¯ÂÌËÈ ̃ ËÒÎ‡ Ú‡ÍÒÓÌÓ‚ ‡Á-ÌÓ„Ó ‡Ì„‡ ‚ îåÅ (å‡ÍÓ‚, 2003). íÓ˜ÍË – ˝ÏÔËË˜ÂÒÍËÂ ÓˆÂÌÍË ÒÓı‡ÌflÂÏÓÒÚË ÒÂÏÂÈÒÚ‚ Ë Ó‰Ó‚, ÔÓÎÛ˜ÂÌÌ˚Â ÔÛÚÂÏÒÓÔÓÒÚ‡‚ÎÂÌËfl ‰ÓÎË ÒÓ‚ÂÏÂÌÌ˚ı ÒÂÏÂÈÒÚ‚, ÔÂ‰ÒÚ‡‚ÎÂÌÌ˚ı ‚ ËÒÍÓÔ‡ÂÏÓÈ ÎÂÚÓÔËÒË, Ò ‚ÂÓflÚÌÓÒÚ¸˛ ÒÓı‡ÌÂÌËfl Ó‰Ó‚,‚˚˜ËÒÎÂÌÌÓÈ ÔÓ ÏÂÚÓ‰ËÍÂ îÛÚ‡ – ê‡ÛÔ‡ (Foote, Raup, 1996; Foote, 1997) ‚ ‚‡ÊÌÂÈ¯Ëı „ÛÔÔ‡ı ÏÓÒÍËı ÊË‚ÓÚÌ˚ı (Foote,Sepkoski, 1999). ÇË‰ÌÓ, ̃ ÚÓ ̋ ÏÔËË˜ÂÒÍËÂ ÚÓ˜ÍË Ì‡ „‡ÙËÍÂ ÎÂÊ‡Ú Ò‡‚ÌËÚÂÎ¸ÌÓ ÌÂ‰‡ÎÂÍÓ ÓÚ ÍË‚ÓÈ, ̃ ÚÓ Ò‚Ë‰ÂÚÂÎ¸ÒÚ‚Û-ÂÚ Ó ıÓÓ¯ÂÈ ÒÚÂÔÂÌË ÒÓ‚Ô‡‰ÂÌËfl ÂÁÛÎ¸Ú‡ÚÓ‚, ÔÓÎÛ˜ÂÌÌ˚ı ‡ÁÌ˚ÏË ÏÂÚÓ‰‡ÏË Ë Ì‡ ÓÒÌÓ‚Â ‡ÁÌ˚ı Ï‡ÒÒË‚Ó‚ ‰‡ÌÌ˚ı. · –Ë – ‚ÎËflÌËÂ ÌÂÔÓÎÌÓÚ˚ ÎÂÚÓÔËÒË Ë ˝ÙÙÂÍÚ‡ “ÔËÚflÊÂÌËfl ÒÓ‚ÂÏÂÌÌÓÒÚË” Ì‡ ÙÓÏÛ ÍË‚˚ı ‡ÁÌÓÓ·‡ÁËfl (ÂÁÛÎ¸Ú‡Ú˚ËÏËÚ‡ˆËÓÌÌÓ„Ó ÏÓ‰ÂÎËÓ‚‡ÌËfl). åÓ‰ÂÎËÓ‚‡Î‡Ò¸ ˝ÍÒÔÓÌÂÌˆË‡Î¸Ì‡fl (·,„,Â,Á) Ë ÎÓ„ËÒÚË˜ÂÒÍ‡fl (‚,‰,Ê,Ë) ‰Ë‚ÂÒËÙËÍ‡ˆËfl



ˆËË ·ËÓÚ˚ (ËÒ. 8,·) ‰‡ÊÂ Ó˜ÂÌ¸ ÌËÁÍ‡fl ÔÓÎÌÓÚ‡ ÎÂ-ÚÓÔËÒË ÌÂ ÏÓÊÂÚ ËÁÏÂÌËÚ¸ ‚Ë‰ ÍË‚˚ı ‡ÁÌÓÓ·‡-ÁËfl: ÓÌË ÓÒÚ‡˛ÚÒfl ˝ÍÒÔÓÌÂÌˆË‡Î¸Ì˚ÏË Í‡Í ÔËÂ‡ÎËÒÚË˜Ì˚ı ÁÌ‡˜ÂÌËflı ÔÓÎÌÓÚ˚ ÎÂÚÓÔËÒË(ËÒ. 8,„, ÚÓÎÒÚ‡fl ÎËÌËfl), Ú‡Í Ë ÔË Á‡ÌËÊÂÌÌ˚ı Ì‡ÔÓfl‰ÓÍ (ËÒ. 8,„, ÚÓÌÍ‡fl ÎËÌËfl). ì·˚‚‡ÌËÂ ÔÓÎÌÓ-Ú˚ ÎÂÚÓÔËÒË Ò ‚ÓÁ‡ÒÚÓÏ ÚÓÊÂ ÌÂ ÏÓÊÂÚ Á‡ÒÚ‡‚ËÚ¸ÍË‚˚Â Á‡ÏÂÚÌÓ ÓÚÍÎÓÌËÚ¸Òfl ÓÚ ˝ÍÔÓÌÂÌˆË‡Î¸ÌÓ-„Ó ÚÂÌ‰‡. éÌË ÓÒÚ‡˛ÚÒfl ÒÚÓ„Ó ̋ ÍÒÔÓÌÂÌˆË‡Î¸Ì˚-ÏË, ıÓÚfl ‚ÓÁ‡ÒÚ‡˛Ú ÌÂÒÍÓÎ¸ÍÓ ÍÛ˜Â, ˜ÂÏ “Â‡Î¸-Ì‡fl” ÍË‚‡fl (ËÒ. 8,Â). Ñ‡ÊÂ ÂÒÎË ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒËÒ ‚ÓÁ‡ÒÚÓÏ ‚Ì‡˜‡ÎÂ Û·˚‚‡ÂÚ ·˚ÒÚÓ, ‡ Á‡ÚÂÏ ÂÂÒÌËÊÂÌËÂ Á‡ÏÂ‰ÎflÂÚÒfl ÔÓ ˝ÍÒÔÓÌÂÌˆË‡Î¸ÌÓÏÛ Á‡-ÍÓÌÛ, ÍË‚‡fl ‡ÁÌÓÓ·‡ÁËfl ‚ÒÂ ‡‚ÌÓ ÓÒÚ‡ÂÚÒfl ˝ÍÒ-ÔÓÌÂÌˆË‡Î¸ÌÓÈ, ‚ ÚÓÏ ̃ ËÒÎÂ Ë ÚÓ„‰‡, ÍÓ„‰‡ ÔÓÎÌÓÚ‡ÎÂÚÓÔËÒË Ì‡ ·ÓÎ¸¯ÂÈ ˜‡ÒÚË “Ù‡ÌÂÓÁÓfl” ÓÍ‡Á˚‚‡-ÂÚÒfl Á‡ÌËÊÂÌÌÓÈ ÔÓ˜ÚË Ì‡ ÔÓfl‰ÓÍ (ËÒ. 8,Á, ÚÓÎ-ÒÚ‡fl ÎËÌËfl).

èË ÎÓ„ËÒÚË˜ÂÒÍÓÈ ‰Ë‚ÂÒËÙËÍ‡ˆËË ·ËÓÚ˚(ËÒ. 8,‚) ÍË‚˚Â ‡ÁÌÓÓ·‡ÁËfl ÓÍ‡Á˚‚‡˛ÚÒfl ·ÓÎÂÂ˜Û‚ÒÚ‚ËÚÂÎ¸Ì˚ÏË Í ÌÂÔÓÎÌÓÚÂ ÎÂÚÓÔËÒË. ãÓ„ËÒÚË-˜ÂÒÍ‡fl ÍË‚‡fl ÏÓÊÂÚ “‡ÒÔflÏËÚ¸Òfl” ÔË ÌÂ ÏÂÌfl-˛˘ÂÈÒfl Ò ‚ÓÁ‡ÒÚÓÏ, ÌÓ Ó˜ÂÌ¸ ÌËÁÍÓÈ ÔÓÎÌÓÚÂ ÎÂ-ÚÓÔËÒË (˝ÚÓ Ë ÂÒÚ¸ ˝ÙÙÂÍÚ “ÔËÚflÊÂÌËfl ÒÓ‚ÂÏÂÌ-ÌÓÒÚË”) (ËÒ. 8,‰, ÚÓÌÍ‡fl ÎËÌËfl); ÚÓ ÊÂ Ì‡·Î˛‰‡ÂÚÒflË ÔË ÎËÌÂÈÌÓÏ ÒÌËÊÂÌËË ÔÓÎÌÓÚ˚ Ò ‚ÓÁ‡ÒÚÓÏ(ËÒ. 8,Ê, ÚÓÎÒÚ‡fl ÎËÌËfl). ÖÒÎË Á‡ÌËÁËÚ¸ ÁÌ‡˜ÂÌËflÔÓÎÌÓÚ˚ ÎÂÚÓÔËÒË ‰Îfl ÌÂ‰‡‚ÌËı ˝ÔÓı Ì‡ ÔÓfl‰ÓÍ ËÁ‡‰‡Ú¸ ÎËÌÂÈÌÓÂ ÒÌËÊÂÌËÂ ÔÓÎÌÓÚ˚ ‰Ó ÌÛÎfl ‚ Ì‡˜‡-ÎÂ “Ù‡ÌÂÓÁÓfl”, Ì‡·Î˛‰‡ÂÏ‡fl ÍË‚‡fl ‡ÁÌÓÓ·‡-ÁËfl ÏÓÊÂÚ ÒÚ‡Ú¸ ‰‡ÊÂ ÌÂÏÌÓ„Ó ‚Ó„ÌÛÚÓÈ, ÌÓ Ë ‚˝ÚÓÏ ÒÎÛ˜‡Â ÓÌ‡ ÌÂ ÒÚ‡ÌÓ‚ËÚÒfl ÔÓıÓÊ‡ ÌË Ì‡ ̋ ÍÒÔÓ-ÌÂÌÚÛ, ÌË ÚÂÏ ·ÓÎÂÂ Ì‡ „ËÔÂ·ÓÎÛ (ËÒ. 8,Ê, ÚÓÌÍ‡flÎËÌËfl). íÓ ÊÂ Ò‡ÏÓÂ Ì‡·Î˛‰‡ÂÚÒfl Ë ÔË ˝ÍÒÔÓÌÂÌ-ˆË‡Î¸ÌÓÏ ÒÌËÊÂÌËË ÔÓÎÌÓÚ˚ ÎÂÚÓÔËÒË Ò ‚ÓÁ‡Ò-ÚÓÏ (ËÒ. 8,Ë, ÚÓÎÒÚ‡fl ÎËÌËfl).

Ö‰ËÌÒÚ‚ÂÌÌ‡fl ÒËÚÛ‡ˆËfl, ÔË ÍÓÚÓÓÈ ÍË‚‡fl‡ÁÌÓÓ·‡ÁËfl ‚ÒÂ-Ú‡ÍË ÔËÓ·ÂÚ‡ÂÚ ÒıÓ‰ÒÚ‚Ó Ò „Ë-ÔÂ·ÓÎÓÈ – ˝ÚÓ ÒÎÛ˜‡È, ÍÓ„‰‡ ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË ‚ÌÂ‰‡‚ÌËÂ ˝ÔÓıË ‚˚ÒÓÍ‡ (ÒÓÓÚ‚ÂÚÒÚ‚ÛÂÚ Â‡ÎËÒÚË˜-Ì˚Ï ÓˆÂÌÍ‡Ï ËÎË ÌÂÏÌÓ„Ó Á‡ÌËÊÂÌ‡), Á‡ÚÂÏ ÒÚÂ-ÏËÚÂÎ¸ÌÓ Û·˚‚‡ÂÚ ÔÓ „ËÔÂ·ÓÎË˜ÂÒÍÓÏÛ Á‡ÍÓÌÛ, ËÓÒÚ‡ÂÚÒfl Í‡ÈÌÂ ÌËÁÍÓÈ ‚ ÚÂ˜ÂÌËÂ ·ÓÎ¸¯ÂÈ ˜‡ÒÚËÙ‡ÌÂÓÁÓfl (ËÒ. 8,Á,Ë, ÚÓÌÍËÂ ÎËÌËË).

ç‡ ÒÂ„Ó‰Ìfl¯ÌËÈ ‰ÂÌ¸ ÌÂÎ¸Áfl ÚÓ˜ÌÓ ÒÍ‡Á‡Ú¸, Í‡Í‚ ‰ÂÈÒÚ‚ËÚÂÎ¸ÌÓÒÚË ÏÂÌflÂÚÒfl ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË Ò‚ÓÁ‡ÒÚÓÏ. é‰ÌË ÒÔÂˆË‡ÎËÒÚ˚ ÔÓÎ‡„‡˛Ú, ˜ÚÓ ÓÌ‡ÔËÏÂÌÓ Ó‰ËÌ‡ÍÓ‚‡ ‰Îfl ·ÓÎ¸¯ÂÈ ˜‡ÒÚË Ù‡ÌÂÓ-ÁÓfl ËÎË ÔÂÚÂÔÂ‚‡ÂÚ ÌÂÌ‡Ô‡‚ÎÂÌÌ˚Â ı‡ÓÚË˜Â-ÒÍËÂ ÍÓÎÂ·‡ÌËfl, ‰Û„ËÂ ‰ÓÔÛÒÍ‡˛Ú, ˜ÚÓ ÓÌ‡ ÔÓÒÚÂ-

ÔÂÌÌÓ ÒÌËÊ‡ÂÚÒfl Ò ‚ÓÁ‡ÒÚÓÏ ÓÚÎÓÊÂÌËÈ. Ç Î˛·ÓÏÒÎÛ˜‡Â ˝ÚÓ ÒÌËÊÂÌËÂ, ÒÛ‰fl ÔÓ ‚ÒÂÏÛ, ÌÂ Ì‡ÒÚÓÎ¸ÍÓÁÌ‡˜ËÚÂÎ¸ÌÓ, ˜ÚÓ·˚ Á‡ÏÂÚÌÓ ÔÓ‚ÎËflÚ¸ Ì‡ ÍË‚˚Â„ÎÓ·‡Î¸ÌÓ„Ó ‡ÁÌÓÓ·‡ÁËfl, ÔÓ Í‡ÈÌÂÈ ÏÂÂ Ì‡ÛÓ‚ÌÂ ÒÂÏÂÈÒÚ‚. í‡ÍÓÈ ‚˚‚Ó‰ ·˚Î Ò‰ÂÎ‡Ì ÅÂÌÚÓ-ÌÓÏ Ë Â„Ó ÍÓÎÎÂ„‡ÏË Ì‡ ÓÒÌÓ‚Â ÒÓÔÓÒÚ‡‚ÎÂÌËfl ‰‡Ì-Ì˚ı Ô‡ÎÂÓÌÚÓÎÓ„Ë˜ÂÒÍÓÈ ÎÂÚÓÔËÒË Ò ÙËÎÓ„ÂÌÂÚË-˜ÂÒÍËÏË ÂÍÓÌÒÚÛÍˆËflÏË (Benton et al., 2000). çÂÚÌËÍ‡ÍËı ÓÒÌÓ‚‡ÌËÈ ÔÂ‰ÔÓÎ‡„‡Ú¸, ˜ÚÓ ÔÓÎÌÓÚ‡ ÎÂ-ÚÓÔËÒË Û·˚‚‡ÂÚ Ò ‚ÓÁ‡ÒÚÓÏ ÔÓ „ËÔÂ·ÓÎË˜ÂÒÍÓÏÛÁ‡ÍÓÌÛ, Ú.Â. (ÔËÌËÏ‡fl ‚Ó ‚ÌËÏ‡ÌËÂ ÙÓÏÛ Â‡Î¸-Ì˚ı ÍË‚˚ı ·ËÓ‡ÁÌÓÓ·‡ÁËfl) Ó˜ÂÌ¸ ‚˚ÒÓÍ‡ ‰ÎflÌÂÓ„ÂÌ‡ Ë ˜ÂÁ‚˚˜‡ÈÌÓ ÒÚÂÏËÚÂÎ¸ÌÓ, ‚Ó ÏÌÓ„Ó‡Á ÒÌËÊ‡ÂÚÒfl ‚ ÚÂ˜ÂÌËÂ Ô‡ÎÂÓ„ÂÌ‡ Ë ÏÂÎ‡, ‡ Á‡ÚÂÏ‚ ÚÂ˜ÂÌËÂ ÔÂ‚ÓÈ ÔÓÎÓ‚ËÌ˚ ÏÂÁÓÁÓfl Ë ‚ÒÂ„Ó Ô‡ÎÂÓ-ÁÓfl ÓÒÚ‡ÂÚÒfl ÌË˜ÚÓÊÌÓ ÌËÁÍÓÈ ÔÓ Ò‡‚ÌÂÌË˛ Ò ÌÂÓ-„ÂÌÓÏ. ÑÎfl ÚÓ„Ó ˜ÚÓ·˚ Ó·˙flÒÌËÚ¸ „ËÔÂ·ÓÎË˜Â-ÒÍËÈ ÓÒÚ ‡ÁÌÓÓ·‡ÁËfl Ú‡ÍËÏË Í‡ÈÌÂ Ï‡ÎÓ‚ÂÓ-flÚÌ˚ÏË Ì‡Ô‡‚ÎÂÌÌ˚ÏË ËÁÏÂÌÂÌËflÏË ÔÓÎÌÓÚ˚ÎÂÚÓÔËÒË Ò ‚ÓÁ‡ÒÚÓÏ, ÌÂÓ·ıÓ‰ËÏÓ Ú‡ÍÊÂ ‰ÓÔÛ-ÒÚËÚ¸, ˜ÚÓ ˝ÚË ËÁÏÂÌÂÌËfl ÔÓËÒıÓ‰flÚ ÒËÌıÓÌÌÓ Ë‚ ÏÓÒÍÓÈ, Ë ‚ ÍÓÌÚËÌÂÌÚ‡Î¸ÌÓÈ „ÂÓÎÓ„Ë˜ÂÒÍÓÈ ÎÂ-ÚÓÔËÒË.

í‡ÍËÏ Ó·‡ÁÓÏ, ÂÁÛÎ¸Ú‡Ú˚ ÏÓ‰ÂÎ¸Ì˚ı ˝ÍÒÔÂ-ËÏÂÌÚÓ‚ ÔÓÍ‡Á˚‚‡˛Ú, ˜ÚÓ ÌË ˝ÙÙÂÍÚ “ÔËÚflÊÂ-ÌËfl ÒÓ‚ÂÏÂÌÌÓÒÚË”, ÌË Û·˚‚‡˛˘‡fl Ò ‚ÓÁ‡ÒÚÓÏÓÚÎÓÊÂÌËÈ ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË ÌÂ ÏÓ„ÛÚ ÔÂ‚‡ÚËÚ¸ÌÂ„ËÔÂ·ÓÎË˜ÂÒÍÛ˛ ÍË‚Û˛ ‡ÁÌÓÓ·‡ÁËfl ‚ „Ë-ÔÂ·ÓÎË˜ÂÒÍÛ˛ ÔË ÒÍÓÎ¸ÍÓ-ÌË·Û‰¸ Â‡ÎËÒÚË˜Ì˚ıÍÓÏ·ËÌ‡ˆËflı ‚ıÓ‰Ì˚ı Ô‡‡ÏÂÚÓ‚ Ë ‰ÓÔÛ˘ÂÌËÈ.

ëèàëéä ãàíÖêÄíìêõ

ä‡ÔËˆ‡ ë.è., 1992. å‡ÚÂÏ‡ÚË˜ÂÒÍ‡fl ÏÓ‰ÂÎ¸ ÓÒÚ‡ Ì‡-ÒÂÎÂÌËfl ÏË‡ // å‡Ú. ÏÓ‰ÂÎËÓ‚‡ÌËÂ. í. 4. ‹ 6.C. 65–79.

ä‡ÔËˆ‡ ë.è., 1999. é·˘‡fl ÚÂÓËfl ÓÒÚ‡ ˜ÂÎÓ‚Â˜ÂÒÚ‚‡.å.: ç‡ÛÍ‡. 190 Ò.

äÓÓÚ‡Â‚ Ä.Ç., å‡ÎÍÓ‚ Ä.ë., ï‡ÎÚÛËÌ‡ Ñ.Ä., 2005‡.á‡ÍÓÌ˚ ËÒÚÓËË: å‡ÚÂÏ‡ÚË˜ÂÒÍÓÂ ÏÓ‰ÂÎËÓ‚‡ÌËÂËÒÚÓË˜ÂÒÍËı Ï‡ÍÓÔÓˆÂÒÒÓ‚ (ÑÂÏÓ„‡ÙËfl. ùÍÓ-ÌÓÏËÍ‡. ÇÓÈÌ˚). å.: ìêëë / äÓÏäÌË„‡. 344 Ò.

äÓÓÚ‡Â‚ Ä.Ç., å‡ÎÍÓ‚ Ä.ë., ï‡ÎÚÛËÌ‡ Ñ.Ä., 2005·.äÓÏÔ‡ÍÚÌ‡fl Ï‡ÚÂÏ‡ÚË˜ÂÒÍ‡fl Ï‡ÍÓÏÓ‰ÂÎ¸ ÚÂıÌË-ÍÓ-˝ÍÓÌÓÏË˜ÂÒÍÓ„Ó Ë ‰ÂÏÓ„‡ÙË˜ÂÒÍÓ„Ó ‡Á‚ËÚËflåË-ëËÒÚÂÏ˚ (1–1973 „„.) / èÓ‰ Â‰. å‡ÎÍÓ‚‡ ë.û.,äÓÓÚ‡Â‚‡ Ä.Ç. àÒÚÓËfl Ë ÒËÌÂ„ÂÚËÍ‡: å‡ÚÂÏ‡ÚË-˜ÂÒÍÓÂ ÏÓ‰ÂÎËÓ‚‡ÌËÂ ÒÓˆË‡Î¸ÌÓÈ ‰ËÌ‡ÏËÍË. å.:ìêëë / äÓÏäÌË„‡. ë. 6–48.

äÓÓÚ‡Â‚ Ä.Ç., å‡ÎÍÓ‚ Ä.ë., ï‡ÎÚÛËÌ‡ Ñ.Ä., 2007‡.á‡ÍÓÌ˚ ËÒÚÓËË. å‡ÚÂÏ‡ÚË˜ÂÒÍÓÂ ÏÓ‰ÂÎËÓ‚‡ÌËÂ

·ËÓÚ˚. ç‡ ‚ÒÂı „‡ÙËÍ‡ı ‡ÁÌÓÓ·‡ÁËÂ ÔÓÍ‡Á‡ÌÓ Ì‡ ËÌÚÂ‚‡ÎÂ ÓÚ 1-„Ó ‰Ó 99-„Ó ‚ÂÍ‡ (ÔÓ „ÓËÁÓÌÚ‡Î¸ÌÓÈ ÓÒË). èÓ ‚ÂÚË-Í‡Î¸ÌÓÈ ÓÒË – ˜ËÒÎÓ Ó‰Ó‚. ·,‚ – “Â‡Î¸ÌÓÂ” ‡ÁÌÓÓ·‡ÁËÂ. „,‰,Â,Ê: ÚÓÎÒÚ˚Â ÎËÌËË – “Ì‡·Î˛‰‡ÂÏÓÂ” ‚ ËÒÍÓÔ‡ÂÏÓÈ ÎÂÚÓ-ÔËÒË ‡ÁÌÓÓ·‡ÁËÂ ÔË Â‡ÎËÒÚË˜ÌÓÈ ‚ÂÎË˜ËÌÂ ÔÓÎÌÓÚ˚ ÎÂÚÓÔËÒË; ÚÓÌÍËÂ ÎËÌËË – ÚÓ ÊÂ, ÔË ‚ÂÎË˜ËÌÂ ÔÓÎÌÓÚ˚ ÎÂÚÓ-ÔËÒË, Á‡ÌËÊÂÌÌÓÈ Ì‡ ÔÓfl‰ÓÍ ÔÓ Ò‡‚ÌÂÌË˛ Ò Â‡ÎËÒÚË˜Ì˚ÏË ˝ÏÔËË˜ÂÒÍËÏË ÓˆÂÌÍ‡ÏË ‰Îfl Ó„‡ÌËÁÏÓ‚ ÒÓ ÒÍÂÎÂÚÓÏ. „,‰ –ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË ÌÂ ÏÂÌflÂÚÒfl Ò ‚ÓÁ‡ÒÚÓÏ ÓÚÎÓÊÂÌËÈ. Â,Ê – ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË ÎËÌÂÈÌÓ ÒÌËÊ‡ÂÚÒfl Ò ‚ÓÁ‡ÒÚÓÏ ÓÚÎÓÊÂ-ÌËÈ (‰Ó ÌÛÎÂ‚ÓÈ ÔÓÎÌÓÚ˚ ‚ 1-Ï ‚ÂÍÂ). Á,Ë – ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË Ò ‚ÓÁ‡ÒÚÓÏ ÒÌËÊ‡ÂÚÒfl ‚Ì‡˜‡ÎÂ ·˚ÒÚÓ, Á‡ÚÂÏ ‚Òfi ÏÂ‰ÎÂÌÌÂÂ(ÓÚ Â‡ÎËÒÚË˜Ì˚ı ÁÌ‡˜ÂÌËÈ ‚ 99-Ï ‚ÂÍÂ ‰Ó Á‡ÌËÊÂÌÌ˚ı Ì‡ ÔÓfl‰ÓÍ ‚ 1-Ï ‚ÂÍÂ); ÚÓÎÒÚ‡fl ÎËÌËfl – ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË ÒÌËÊ‡-ÂÚÒfl ÔÓ ˝ÍÒÔÓÌÂÌˆË‡Î¸ÌÓÏÛ Á‡ÍÓÌÛ, ÚÓÌÍ‡fl ÎËÌËfl – ÔÓÎÌÓÚ‡ ÎÂÚÓÔËÒË ÒÌËÊ‡ÂÚÒfl ÔÓ „ËÔÂ·ÓÎË˜ÂÒÍÓÏÛ Á‡ÍÓÌÛ.

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Hyperbolic growth of marine and continental biodiversity through

the Phanerozoic and community evolution

A. V. Markov1, A. V. Korotayev2

1 Paleontological Institute, RAS 117997 Moscow, ul. Profsoyuznaya, 123

e-mail: [email protected] 2 Russian State Humanitarian University

125993 Moscow, Miusskaya plosh., 6e-mail: [email protected]

Among diverse models that are used to describe and interpret the changes in global biodiversity through thePhanerozoic, the exponential and logistic models (traditionally used in population biology) are the most popu-lar. As we have recently demonstrated (Markov, Korotayev, 2007), the growth of the Phanerozoic marine biodi-

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versity at genus level correlates better with the hyperbolic model (widely used in demography and macrosoci-ology). Here we show that the hyperbolic model is also applicable to the Phanerozoic continental biota at genusand family levels, and to the marine biota at species, genus, and family levels. There are many common featuresin the evolutionary dynamics of the marine and continental biotas that imply similarity and common nature ofthe factors and mechanisms underlying the hyperbolic growth. Both marine and continental biotas are charac-terized by continuous growth of the mean longevity of taxa, by decreasing extinction and origination rates, bysimilar pattern of replacement of dominant groups, by stepwise accumulation of evolutionary stable, adaptableand “physiologically buffered” taxa with effective mechanisms of parental care, protection of early develop-mental stages, etc. At the beginning of the development of continental biota, the observed taxonomic diversity was substantial-ly lower than that predicted by the hyperbolic model. We suggest that this is due, firstly, to the fact that, duringthe earliest stages of the continental biota evolution, the groups that are not preserved in the fossil record (suchas soil bacteria, unicellular algae, lichens, etc.) played a fundamental role, and secondly, to the fact that the con-tinental biota initially formed as a marginal portion of the marine biota, rather than a separate system. The hy-perbolic dynamics is most prominent when both marine and continental biotas are considered together. Thisfact can be interpreted as a proof of the integrated nature of the biosphere. In the macrosociological models, thehyperbolic pattern of the world population growth arises from a non-linear second-order positive feedback be-tween the demographic growth and technological development (more people – more potential inventors – fastertechnological growth – the carrying capacity of the Earth grows faster – faster population growth – more people –more potential inventors, and so on). Based on the analogy with macrosociological models and diverse paleontological data, we suggest that thehyperbolic character of biodiversity growth can be similarly accounted for by a non-linear second-order posi-tive feedback between the diversity growth and community structure complexity. The feedback can work viatwo parallel mechanisms: 1) decreasing extinction rate (more taxa – higher alpha diversity, or mean number oftaxa in a community – communities become more complex and stable – extinction rate decreases – more taxa,and so on) and 2) increasing origination rate (new taxa facilitate niche construction; newly formed niches canbe occupied by the next “generation” of taxa). The latter possibility makes the mechanisms underlying the hy-perbolic growth of biodiversity and human population even more similar, because the total ecospace of the bio-ta is analogous to the “carrying capacity of the Earth” in demography. As far as new species can increaseecospace and facilitate opportunities for additional species entering the community, they are analogous to the“inventors” of the demographic models whose inventions increase the carrying capacity of the Earth. The hy-perbolic growth of the Phanerozoic biodiverstiy suggests that “cooperative” interactions between taxa can playan important role in evolution, along with generally accepted competitive interactions. Due to this “coopera-tion”, the evolution of biodiversity acquires some features of a self-accelerating process. Macroevolutionary“cooperation” reveals itself in: 1) increasing stability of communities that arises from alpha diversity growth;2) ability of species to facilitate opportunities for additional species entering the community.

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