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Историческое Развитие Парнопалых (Artiodactyla) Северной Евразии И Этапы Эв

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Историческое Развитие Парнопалых (Artiodactyla) Северной Евразии И Этапы Эв ÝÂÎËÞÖÈß ÁÈÎÑÔÅÐÛ È ÁÈÎÐÀÇÍÎÎÁÐÀÇÈß 4. ÏÀËÅÎÝÊÎËÎÃÈß È ÝÂÎËÞÖÈß ÑÎÎÁÙÅÑÒ ÓÄÊ 569.73: 415 (551.77) Èñòîðè÷åñêîå ðàçâèòèå ïàðíîïàëûõ (Àrtiodactyla) Ñåâåðíîé Eâðàçèè è ýòàïû ýâîëþöèè èõ ñîîáùåñòâ â êàéíîçîå È.À. Âèñëîáîêîâà Ïàëåîíòîëîãè÷åñêèé èíñòèòóò ÐÀÍ, Ìîñêâà E-mail: [email protected] Ïðîàíàëèçèðîâàíà ýâîëþöèÿ ñîîáùåñòâ ïàðíîïàëûõ â êàéíîçîå è âûäåëåíû îñíîâíûå ýòàïû â èíòåðâàëå îò 56 äî 0,4 ìëí. ëåò íàçàä. Ðîñò ðàçíîîáðàçèÿ ïàðíîïàëûõ òåñíî ñâÿçàí ñ ýâîëþöèåé è äèâåðñèôèêàöèåé ðàñòèòåëüíîãî ïîêðîâà è èçìåíåíèÿìè êëèìàòà è ëàíä- øàôòîâ. Îñíîâíûå ðóáåæè ïðåîáðàçîâàíèÿ ñîîáùåñòâ â ïàëåîãåíå ñèíõðîííû êðóïíåé- øèì áèîñôåðíûì ñîáûòèÿì, à ïëèîöåí-ïëåéñòîöåíîâûå èçìåíåíèÿ îòðàæàþò êðóïíûå êîëåáàíèÿ êëèìàòà â ñåâåðíîì ïîëóøàðèè. Êëþ÷åâûå ñëîâà: ïàðíîïàëûå, ýâîëþöèÿ, îñíîâíûå ýòàïû, áèîðàçíîîáðàçèå, êàéíîçîé, Ñåâåðíàÿ Åâðàçèÿ. Îòðÿä ïàðíîïàëûõ (Artiodactyla) îäíî èç ìàãèñòðàëüíûõ íàïðàâëåíèé ýâîëþöèè ïëàöåíòàðíûõ ìëåêîïèòàþùèõ (Eutheria). Âîçíèêíóâ â ìåçîçîå, îí äîìèíèðóåò ñðåäè êðóï- íûõ ìëåêîïèòàþùèõ ïëàíåòû íàðÿäó ñ ïðèìàòàìè è õèùíûìè. Ýòî äèíàìè÷íî ðàçâèâàâ- øàÿñÿ ãðóïïà ìëåêîïèòàþùèõ õîðîøî ïðåäñòàâëåíà â èñêîïàåìîì ñîñòîÿíèè è óíèêàëüíà äëÿ èçó÷åíèÿ çàêîíîìåðíîñòåé ìàêðîýâîëþöèè è èçìåíåíèÿ áèîðàçíîîáðàçèÿ. Èñòîðèÿ ïàðíîïàëûõ òåñíî ñâÿçàíà ñ ýâîëþöèåé áèîñôåðû è ýâîëþöèåé ðàñòèòåëüíîãî ïîêðîâà Çåìëè è ðàçâîðà÷èâàëàñü íà ôîíå èçìåíåíèé î÷åðòàíèé ìàòåðèêîâ, ñîñòàâà àòìîñôåðû è åå öèðêóëÿöèè, êëèìàòà è ëàíäøàôòîâ. Õîðîøèå èíäèêàòîðû ïîñëåäíèõ, ýòè â îñíîâíîì ðàñòèòåëüíîÿäíûå ìëåêîïèòàþùèå ñ óñïåõîì èñïîëüçóþòñÿ ïðè ðåêîíñòðóêöèè ïðèðîä- íîé ñðåäû ïðîøëûõ ýïîõ è â áèîñòðàòèãðàôèè. Áîëüøîå ðàçíîîáðàçèå è õîðîøàÿ èçó÷åí- íîñòü ïàðíîïàëûõ ïîçâîëÿþò èñïîëüçîâàòü èõ äëÿ ïîçíàíèÿ èñòîðèè íàçåìíûõ áèîöåíî- çîâ êàéíîçîÿ è îñîáåííîñòåé ôîðìèðîâàíèÿ ñîâðåìåííûõ áèîìîâ.  ñòàòüå àíàëèçèðóþòñÿ ñóêöåññèè êîìïëåêñîâ ïàðíîïàëûõ, ñóùåñòâîâàâøèõ â Ñå- âåðíîé Åâðàçèè (íà òåððèòîðèè Ðîññèè, Ìîëäàâèè, Óêðàèíû, Ãðóçèè, Êàçàõñòàíà, Êèð- ãèçèè, Òàäæèêèñòàíà, Ìîíãîëèè è Ñåâåðíîãî Êèòàÿ) îò ýîöåíà äî ñåðåäèíû ñðåäíåãî ïëåéñòîöåíà (560,4 ìëí. ëåò íàçàä). Äàííûå ïî ïàðíîïàëûì ýòîé òåððèòîðèè èíòåðåñíû â ñâÿçè ñ òåì, ÷òî çäåñü ôèêñè- ðóþòñÿ âàæíûå ýòàïû è ñîáûòèÿ èõ èñòîðè÷åñêîãî ðàçâèòèÿ è ñòàíîâëåíèÿ ìíîãèõ ãðóïï, â ÷àñòíîñòè, ôîðìèðîâàíèå áîðåàëüíûõ è ñóáàðêòè÷åñêèõ ôîðì, íåêîòîðûõ ñîâðåìåí- íûõ ðîäîâ è âèäîâ. Êðîìå òîãî, ôàóíû ýòîé òåððèòîðèè áûëè ñâÿçóþùèì çâåíîì ìåæäó Âèñëîáîêîâà È.À. Èñòîðè÷åñêîå ðàçâèòèå ïàðíîïàëûõ 417 ôàóíàìè Åâðîïû, Ñåâåðíîé Àìåðèêè, Þæíîé Àçèè è Àôðèêè. Çäåñü ïðîõîäèëè âàæíûå ôàóíèñòè÷åñêèå îáìåíû è ïðîëåãàëè ìíîãèå ìèãðàöèîííûå ïóòè. Çíà÷èòåëüíûé âêëàä â èçó÷åíèå ïàðíîïàëûõ âíåñëè À.À. Áîðèñÿê, Ì.Â. Ïàâëîâà Ë.È. Àëåêñååâà, Â.Ñ. Áàéãóøåâà, Ì.È. Áóð÷àê-Àáðàìîâè÷, À.Ê. Âåêóà, Í.Ê. Âåðåùàãèí, À.ß. Ãîäèíà, Å.Ë. Äìèòðèåâà, Å.Ë. Êîðîòêåâè÷, À.Í. Ëóíãó, Ï.À. Òëåóáåðäèíà, Á.À. Òðî- ôèìîâ, Ê.Ê. Ôëåðîâ è ß.È. Õàâåñîí. Ïîëó÷åííûå àâòîðîì äàííûå ïî ìîðôîëîãèè, ýâîëþöèè è áèîðàçíîîáðàçèþ ïàëåî- ãåíîâûõ, íåîãåíîâûõ è ðàííåïëåéñòîöåíîâûõ ïàðíîïàëûõ Ñåâåðíîé Åâðàçèè ñóùåñòâåí- íî óòî÷íèëè è äîïîëíèëè èìåâøèåñÿ ñâåäåíèÿ è ïîçâîëèëè ïåðåñìîòðåòü ìíîãèå êëþ- ÷åâûå âîïðîñû, êàñàþùèåñÿ èñòîðè÷åñêîãî ðàçâèòèÿ ïàðíîïàëûõ è èõ ñîîáùåñòâ.  ñòàòüå ïðîñëåæèâàþòñÿ îñíîâíûå ìîìåíòû èñòîðè÷åñêîãî ðàçâèòèÿ è âîçðàñòàíèÿ ñðåäîîáðàçóþùåé ðîëè Artiodañtyla â êàéíîçîå Ñåâåðíîé Åâðàçèè, âûäåëÿþòñÿ îñíîâ- íûå ðóáåæè è àíàëèçèðóþòñÿ îñíîâíûå ýòàïû ýâîëþöèè èõ ñîîáùåñòâ, èçìåíåíèÿ èõ áèîðàçíîîáðàçèÿ îò àáèîòè÷åñêèõ è áèîòè÷åñêèõ ôàêòîðîâ. Ïîñëåäîâàòåëüíûå ýòàïû îõàðàêòåðèçîâàíû îïðåäåëåíûìè àññîöèàöèÿìè òàêñîíîâ è äàííûìè ïî ïîÿâëåíèþ è ðàñïðîñòðàíåíèþ ðÿäà òàêñîíîâ. Èññëåäîâàíèÿ ïðîâîäèëèñü â ðàìêàõ îòå÷åñòâåííûõ ïðîãðàìì è ìåæäóíàðîäíûõ ïðîåêòîâ. Èñòîðè÷åñêîå ðàçâèòèå ïàðíîïàëûõ â îëèãîöåíå Àçèè ñóùåñòâåííî óòî÷íè- ëèñü â õîäå ðàáîò ïî ïðîåêòó ÞÍÅÑÊÎ ¹ 326 «Ïåðåõîä îò îëèãîöåíà ê ìèîöåíó â Ñå- âåðíîì ïîëóøàðèè» è â ðàìêàõ ñîâåòñêî-ìîíãîëüñêîãî ñîòðóäíè÷åñòâà è àâñòðèéñêî- ìîíãîëüñêîãî ïðîåêòà (_W_: P-10505-GEO). Èçó÷åíèå íåîãåíîâûõ è ïëåéñòîöåíîâûõ ôàóí âåëîñü ïî ïðîãðàììå Åâðîïåéñêîãî íàó÷íîãî ôîíäà «Îáñòàíîâêà è äèíàìèêà ýêî- ñèñòåì íåîãåíà Åâðàçèè» (EEDEN), â ðàìêàõ ìåæäóíàðîäíîãî ïðîåêòà «Íåîãåí Ñòàðîãî Ñâåòà» (NOW), àâñòðèéñêîãî ïðîåêòà «Èçìåíåíèÿ ìèîöåíîâûõ ýêîñèñòåì Âîñòî÷íûõ Àëüï ïî ïîçâîíî÷íûì» (_W_ P 15724), ìåæäóíàðîäíîãî ïðîåêòà ïî ñðàâíèòåëüíîìó èçó÷åíèþ ôàóí Åâðîïû. Áëàãîäàðÿ ýëåêòðîííûì áàçàì äàííûõ, ñîçäàííûì ñîâìåñòíî ñ çàðóáåæíûìè êîëëåãàìè, ïðîàíàëèçèðîâàíû îñíîâíûå ðóáåæè è ñîáûòèÿ â èñòîðèè ôàóí íåîãåíà è ïëåéñòîöåíà. Ïðåäñòàâëåííàÿ àâòîðîì ìîäåëü ýâîëþöèè ñîîáùåñòâ ïàðíîïàëûõ îñíîâàíà íà èçó- ÷åíèè áîãàòåéøåé êîëëåêöèè Ïàëåîíòîëîãè÷åñêîãî èíñòèòóòó ÐÀÍ, à òàêæå ìàòåðèàëîâ êðóïíåéøèõ ìóçååâ ìèðà: Àìåðèêàíñêîãî ìóçåÿ åñòåñòâåííîé èñòîðèè, Íàöèîíàëüíîãî ìóçåÿ åñòåñòâåííîé èñòîðèè â Ïàðèæå, ìóçååâ åñòåñòâåííîé èñòîðèè â Ëîíäîíå, Ïåêè- íå, óíèâåðñèòåòîâ Áåðêëè, Êàíçàñà, Ëèîíà, Ìîíïåëüå è äð. Íà ðàçíûõ ýòàïàõ ìîåé íàó÷íîé æèçíè ìåíÿ ïîääåðæèâàëè òàêèå êîðèôåè ïàëåîòå- ðèîëîãèè, êàê Â.È. Ãðîìîâ, Ê.Ê. Ôëåðîâ, Ý. Ýéíö, Ã.-Â. Êàëüêå è Ð. Òåäôîðä, ê êîòîðûì ÿ âñåãäà èñïûòûâàþ ãëóáîêóþ ïðèçíàòåëüíîñòü. Îñîáóþ áëàãîäàðíîñòü çà ïîñòîÿííóþ ïîääåðæêó ÿ õî÷ó âûðàçèòü À.Þ. Ðîçàíîâó, çàìå÷àòåëüíîìó ÷åëîâåêó è âûäàþùåìóñÿ ó÷åíîìó, êîòîðîìó ÿ ïîñâÿùàþ ýòó ðàáîòó. Ñòàòüÿ âûïîëíåíà ïî ïðîãðàììå Ïðåçèäèóìà ÐÀÍ 11 «Íàó÷íûå îñíîâû ñîõðàíåíè áèî- ðàçíîîðàçèÿ Ðîññèè» (ïðîåêò «Ýâîëþöèÿ è ñðåäîîáðàçóþùàÿ ôóíêöèÿ áèîðàçíîîáðàçèÿ»), ïðè ïîääåðæêå ÐÔÔÈ (ïðîåêò 05-04-48493á) è Ïðåçèäåíòà Ðîññèè (ÍØ-6228.2006.4). Îñíîâíûå íàïðàâëåíèÿ ýâîëþöèè Àrtiodactyla Ñòðóêòóðà îòðÿäà. Îòðÿä Artiodactyla îáúåäèíÿåò òðè ïîäîòðÿäà: âåðáëþäîâ (Tylopoda), íåæâà÷íûõ, èëè ñâèíîîáðàçíûõ (Suiformes) è æâà÷íûõ (Ruminantia).  êàé- íîçîå Ñåâåðíîé Åâðàçèè ê Suiformes ïðèíàäëåæàëè øåñòü ñåìåéñòâ ðÿäà àðõàè÷íûõ íàä- ñåìåéñòâ (Diacodexeidae, Helohyidae, Dichobunidae, Raoellidae, Antracotheriidae, 418 4. Ïàëåîýêîëîãèÿ è ýâîëþöèÿ ñîîáùåñòâ Entelodontidae) è äâà ñåìåéñòâà íàäñåìåéñòâà Suoidea: áåãåìîòû (Hippopotamidae) è ñâè- íüè (Suidae). Ruminantia ïðåäñòàâëåíû äåñÿòüþ ñåìåéñòâàìè äâóõ èíôðàîòðÿäîâ: íèç- øèå æâà÷íûå, èëè òðàãóëèíû (Tragulina) è âûñøèå æâà÷íûå (Pecora).  ñîñòàâå òðàãóëèí âûäåëÿþòñÿ äâà èíôðàîòðÿäà: Traguloidea è Hypertraguloidea. Tragulina ïðåäñòàâëåíû ÷åòûðüìÿ ñåìåéñòâàìè: Archaeomerycidae, Praetragulidae, Gelocidae è Lophiomerycidae, à âûñøèå æâà÷íûå ïÿòüþ: êàáàðãè (Moschidae), ïàëåîìåðèöèäû (Palaeomerycidae), æè- ðàôû (Giraffidae), îëåíè (Cervidae) è ïîëîðîãèå (Bovidae).  ñîâðåìåííîé ôàóíå Åâðà- çèè äîìèíèðóþò äâà ïîñëåäíèõ ñåìåéñòâà è ïðèñóòñòâóþò Suidae è Tragulidae.  ñîâðå- ìåííîé ôàóíå ìèðà ê ïàðíîïàëûì ïðèíàäëåæèò îêîëî 90 ðîäîâ è 150 âèäîâ, èç íèõ æâà÷íûì îêîëî 76 ðîäîâ è 130 âèäîâ. Ñðåäè êðóïíûõ ìëåêîïèòàþùèõ ïî ÷èñëó âè- äîâ ïàðíîïàëûå óñòóïàþò òîëüêî ïðèìàòàì è õèùíèêàì. Ïîäîòðÿä ìîçîëåíîãèõ (Tylopoda), êîòîðîìó èíîãäà ïðèäàþò ðàíã ñàìîñòîÿòåëüíîãî îòðÿäà, ïðåäñòàâëåí ñåìåéñòâîì âåðáëþäîâ (Camelidae). Âåðáëþäû (ðîä Paracamelus) âñåëÿþòñÿ â Ñåâåðíóþ Åâðàçèþ èç Àìåðèêè òîëüêî â êîíöå ïîçäíåãî ìèîöåíà. Àðõàè÷- íûå ãðóïïû ìîçîëåíîãèõ (àìôèìåðèöèäû è êàéíîòåðèè), õàðàêòåðíûå äëÿ îëèãîöåíà Çàïàäíîé Åâðîïû, íà ðàññìàòðèâàåìîé òåððèòîðèè íå âñòðå÷åíû. Ýêîëîãè÷åñêîå çíà÷åíèå. Ýâîëþöèÿ Artiodactyla ñâÿçàíà ñ îñâîåíèåì ðàñòèòåëüíûõ ðåñóðñîâ ïëàíåòû. Îñíîâíàÿ ìàññà èñêîïàåìûõ è ñîâðåìåííûõ Artiodactyla ðàñòè- òåëüíîÿäíûå, ëèøü Suoidea îáúåäèíÿþò ãðóïïû âñåÿäíûõ. Íàðÿäó ñ äðóãèìè æèâîòíû- ìè, Artiodactyla àêòèâíî ó÷àñòâóþò â ïîääåðæàíèè ãîìåîñòàçà áèîñôåðû, â âûñâîáîæäå- íèè CO èç ðàñòåíèé è ïîääåðæàíèè îïòèìàëüíîãî äëÿ æèçíè ñîîòíîøåíèÿ Î /ÑÎ â 2 2 2 àòìîñôåðå, à òàêæå êðóãîâîðîòå áèîãåííûõ âåùåñòâ. ×ðåçâû÷àéíî âûñîêà èõ ðîëü â ïîääåðæàíèè ãîìåîñòàçà íà óðîâíå áèîöåíîçîâ, áèî- ìîâ è ìåñòîîáèòàíèé. Áóäó÷è ïåðâè÷íûìè êîíñóìåíòàìè, îíè èãðàþò àêòèâíóþ ðîëü â ïèùåâûõ öåïÿõ, îáåñïå÷èâàÿ ïåðåíîñ ýíåðãèè îò ðàñòåíèé ê õèùíèêàì è ÷åëîâåêó. Ïàð- íîïàëûå äåìîíñòðèðóþò øèðîêèé ñïåêòð ýêîëîãè÷åñêèõ àäàïòàöèé. Ýòà ãðóïïà îòíî- ñèòñÿ ê ïåðìåàíòàì, ïèùåâûå ðåñóðñû êîòîðûõ îõâàòûâàþò ðàçíûå òèïû è ÿðóñû ðàñòè- òåëüíîñòè. Ìèãðàöèè è âèäîâîå ðàçíîîáðàçèå îáåñïå÷èâàþò ðàâíîìåðíîå èñïîëüçîâà- íèå âñåãî ñïåêòðà ðàñòèòåëüíîñòè. Íàèáîëåå âûñîêî ñïåöèàëèçèðîâàííûå ðàñòèòåëüíî- ÿäíûå âûñøèå æâà÷íûå (Pecora) îáëàäàþò ñëîæíûì ÷åòûðåõêàìåðíûì æåëóäêîì, õî- ðîøî ïðèñïîñîáëåííûì äëÿ ïåðåâàðèâàíèÿ ðàñòèòåëüíîãî ìàòåðèàëà. Óâåëè÷åíèå ðàçíîîáðàçèÿ Artiodactyla âî âðåìåíè íàïðàâëåíî íà ïîääåðæàíèå ãîìå- îñòàçà áèîñôåðû è óñòîé÷èâîñòè áèîöåíîçîâ. Ïðîèñõîæäåíèå. Ïî âñåé âèäèìîñòè, ïîÿâëåíèå ïàðíîïàëûõ ñâÿçàíî ñ êðóïíåéøåé áèîñôåðíîé ïåðåñòðîéêîé â êîíöå ìåëà (ðèñ. 1). Ñîâðåìåííûå ïàëåîíòîëîãè÷åñêèå, öèòîãåíåòè÷åñêèå è ìîëåêóëÿðíûå äàííûå ñâèäåòåëüñòâóþò îá î÷åíü äðåâíåì ïðîèñ- õîæäåíèè îòðÿäà Artiodactyla è åãî ðàííåì ðàçâåòâëåíèè íà îñíîâíûå ñòâîëû (Todd, 1975; Prothero et al., 1988; Shimamura et al., 1997; Vislobokova, 1998; Vislobokova, Trofimov, 2002), î êîòîðûõ ïðåäïîëàãàë åùå Â.Î. Êîâàëåâñêèé (Kowalevsky, 1873-1874, 1875). Ñî- ãëàñíî êàëèáðîâêå ìîëåêóëÿðíûõ ÷àñîâ àäàïòèâíàÿ ðàäèàöèÿ äðåâíåéøèõ ýóòåðèåâ, ïðè- âåäøàÿ ê ïîÿâëåíèþ îòðÿäîâ, ïðîèçîøëà áîëåå 100 ìëí. ëåò íàçàä (Shimamura et al., 1997). Åñëè ïðèíÿòü âî âíèìàíèå ïðîäîæèòåëüíîñòü ñóùåñòâîâàíèÿ ïîñëåäíåé òðèàäû Ruminantia (Cervidae, Giraffidae è Bovidae), êîòîðàÿ ñîñòàâëÿåò 34 ìëí. ëåò, è ïðåä- ïîëîæèòü, ÷òî ïðîäîëæèòåëüíîñòü äâóõ ïðåäûäóùèõ áûëà ïðèìåðíî òàêîé æå, òî äàæå áåç ó÷åòà âîçìîæíîãî óñêîðåíèÿ òåìïîâ ýâîëþöèè, ïîëó÷èòñÿ äàòà, áëèçêàÿ ê öèôðå, âû÷èñëåííîé ïî ìîëåêóëÿðíûì äàííûì. Ïðîäîæèòåëüíîñòü ïåðâîé òðèàäû Âèñëîáîêîâà È.À. Èñòîðè÷åñêîå ðàçâèòèå ïàðíîïàëûõ 419 Ý Ðèñ. 1. Ñõåìà ýâîëþöèè Artiodactyla â êàéíîçîå Âíóòðåííåé Àçèè. Ñòðåëêè ïîêàçûâàþò
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