Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых
В сравнительном и функциональном аспектах проанализированы некоторые морфологические особенности таза и бедра представителя бескилевых — нанду (Rhea americana). Определена высокая степень корреляции между шириной таза и шири¬ной грудины у представителей 35 видов 10 отрядов килегрудых птиц. В приложе...
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| Veröffentlicht in: | Вестник зоологии |
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| Datum: | 2008 |
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| Format: | Artikel |
| Sprache: | Russisch |
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Інститут зоології ім. І.І. Шмальгаузена НАН України
2008
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| Zitieren: | Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых / И.А. Богданович // Вестник зоологии. — 2008. — Т. 42, № 4. — С. 347–354. — Бібліогр.: 45 назв. — рос. |
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Digital Library of Periodicals of National Academy of Sciences of Ukraine| _version_ | 1860011428404002816 |
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| author | Богданович, И.А. |
| author_facet | Богданович, И.А. |
| citation_txt | Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых / И.А. Богданович // Вестник зоологии. — 2008. — Т. 42, № 4. — С. 347–354. — Бібліогр.: 45 назв. — рос. |
| collection | DSpace DC |
| container_title | Вестник зоологии |
| description | В сравнительном и функциональном аспектах проанализированы некоторые морфологические особенности таза и бедра представителя бескилевых — нанду (Rhea americana). Определена высокая степень корреляции между шириной таза и шири¬ной грудины у представителей 35 видов 10 отрядов килегрудых птиц. В приложении к морфо¬логии таза нанду и наиболее близкородственным представителям группы указанная зависимость позволяет, в частности, высказаться в пользу первичности их нелетания.
Some peculiarities of Rhea americana pelvis and femur are analyzed in comparative and functional aspects. High level of correlation between the width of the pelvis and that of the sternum is shown to occur in representatives of 35 species and 10 orders of the carinate birds. In respect to pelvis morphology of Rhea and close related ratitae, it allows to suppose in particular the primary character of its flightlessness.
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ÓÄÊ 598.221.2:591.47
ÍÅÊÎÒÎÐÛÅ ÎÑÎÁÅÍÍÎÑÒÈ ÑÊÅËÅÒÀ ÍÀÍÄÓ
RHEA AMERICANA (RHEIFORMES, RHEIDAE)
 ÑÂÅÒÅ ÝÂÎËÞÖÈÈ ÁÅÑÊÈËÅÂÛÕ
È. À. Áîãäàíîâè÷
Èíñòèòóò çîîëîãèè èì. È. È. Øìàëüãàóçåíà ÍÀÍ Óêðàèíû,
óë. Á. Õìåëüíèöêîãî, 15, Êèåâ, 01601 Óêðàèíà
E-mail: ibogdanovich@rambler.ru
Ïðèíÿòî 29 ìàÿ 2008
Íåêîòîðûå îñîáåííîñòè ñêåëåòà íàíäó Rhea americana (Rheiformes, Rheidae) â ñâåòå ýâîëþöèè
áåñêèëåâûõ. Áîãäàíîâè÷ È. À. — Â ñðàâíèòåëüíîì è ôóíêöèîíàëüíîì àñïåêòàõ ïðîàíàëèçè-
ðîâàíû íåêîòîðûå ìîðôîëîãè÷åñêèå îñîáåííîñòè òàçà è áåäðà ïðåäñòàâèòåëÿ áåñêèëåâûõ —
íàíäó (Rhea americana). Îïðåäåëåíà âûñîêàÿ ñòåïåíü êîððåëÿöèè ìåæäó øèðèíîé òàçà è øèðè-
íîé ãðóäèíû ó ïðåäñòàâèòåëåé 35 âèäîâ 10 îòðÿäîâ êèëåãðóäûõ ïòèö. Â ïðèëîæåíèè ê ìîðôî-
ëîãèè òàçà íàíäó è íàèáîëåå áëèçêîðîäñòâåííûì ïðåäñòàâèòåëÿì ãðóïïû óêàçàííàÿ çàâèñè-
ìîñòü ïîçâîëÿåò, â ÷àñòíîñòè, âûñêàçàòüñÿ â ïîëüçó ïåðâè÷íîñòè èõ íåëåòàíèÿ.
Êëþ÷åâûå ñ ëîâ à: ïîëåò, ýâîëþöèÿ, áåñêèëåâûå.
Some Skeletal Peculiarities of Rhea americana (Rheiformes, Rheidae) with Respect of Ratitae
Evolution. I. A. Bogdanovich. — Some peculiarities of Rhea americana pelvis and femur are analyzed
in comparative and functional aspects. High level of correlation between the width of the pelvis and
that of the sternum is shown to occur in representatives of 35 species and 10 orders of the carinate
birds. In respect to pelvis morphology of Rhea and close related ratitae, it allows to suppose in partic-
ular the primary character of its flightlessness.
Ke y wo r d s: flight, evolution, ratitae.
Ââåäåíèå
Ýâîëþöèÿ è ôèëîãåíèÿ ïàëåîãíàò è îñîáåííî íåëåòàþùèõ áåñêèëåâûõ (Ratitae) äîñòàòî÷íî
äàâíî ÿâëÿåòñÿ îáüåêòîì èññëåäîâàíèé. Ãëàâíûå âîïðîñû — ñòåïåíü ðîäñòâà ñ ëåòàþùèìè
êèëåãðóäûìè è ïåðâè÷íîñòü èëè âòîðè÷íîñòü íåëåòàíèÿ. Íà îïðåäåëåííîì ýòàïå èòîãè èçûñêàíèé
áûëè ïîäâåäåíû â âèäå òðåõ ãèïîòåç (Thulborn, 1984): 1 — áåñêèëåâûå — íàñëåäíèêè êèëåãðóäûõ
îòíîñèòåëüíî äàâíåãî âîçðàñòà (Beer de, 1956; Cracraft, 1974); 2 — áåñêèëåâûå ÿâëÿþòñÿ ãðóïïîé
ñðåäíåãî ýâîëþöèîííîãî âîçðàñòà è ïðîèñõîäèëè îò ëåòàþùèõ ïòèö, áîëåå ïðèìèòèâíûõ, ÷åì
Ñarinates (Tucker, 1938; McGovan, 1982); 3 — áåñêèëåâûå — ýòî äðåâíÿÿ ãðóïïà, êîòîðàÿ ïðîèçîøëà
îò ïðèìèòèâíûõ íåëåòàþùèõ ïòèö (Lowe, 1928; Friant, 1968). Ðåçóëüòàòû áîëåå ïîçäíèõ
èññëåäîâàíèé ïîääåðæèâàëè êàê ïåðâóþ (McGowan, 1986), òàê è âòîðóþ (Houde, 1986) èëè òðåòüþ
(Feduccia, 1986) èç ïðèâåäåííûõ ãèïîòåç. Êðîìå òîãî, èììóíîëîãè÷åñêîå èññëåäîâàíèå ïðèâåëî
èññëåäîâàòåëåé (Stapel et al., 1984) ê çàêëþ÷åíèþ î òîì, ÷òî Ratitae íå ïðåäêîâàÿ èëè ïðîèçâîäíàÿ,
à ñåñòðèíñêàÿ ãðóïïà ïî îòíîøåíèþ ê êèëåãðóäûì, â ÷àñòíîñòè êóðî- è ãóñåîáðàçíûì. Ïîäîáíûé
âûâîä ïîääåðæèâàåòñÿ ðÿäîì äðóãèõ ðàáîò (Äçåðæèíñêèé, 2000; Sibley, Ahlquist, 1990; Cracraft, 2000).
Âîçìîæíîñòü âûâåäåíèÿ ìîðôîëîãè÷åñêèõ ïðèçíàêîâ íåîãíàò èç èõ ñîñòîÿíèÿ ó ïàëåîãíàò
(Kurochkin, 1995) ñâèäåòåëüñòâóåò, â ÷àñòíîñòè, î ïðîèñõîæäåíèè ñèëîâîãî ïîëåòà ïîñëå äèâåðãåíöèè
Ratitae è Galliformes îò áàçîâîãî ñòâîëà ïòèö.
Íå ñòàâÿ ñïåöèàëüíûõ çàäà÷ ïî ðåøåíèþ ñïîðíûõ âîïðîñîâ ôèëîãåíèè áåñêèëåâûõ, ìû ëèøü
ïîïûòàëèñü ïðîàíàëèçèðîâàòü íåêîòîðûå ïðèçíàêè ñêåëåòà îäíîãî èç íàèáîëåå ãåíåðàëèçîâàííûõ
(èñõîäÿ èç ñòðîåíèÿ êðûëà è âòîðè÷íîñòè íåëåòàíèÿ, Cracraft, 1974) ïðåäñòàâèòåëåé ãðóïïû â àñïåêòå
îáñóæäàåìîé ïðîáëåìû.
Vestnik zoologii, 42(4): 347–354, 2008
È. À. Áîãäàíîâè÷
Ìàòåðèàë è ìåòîäû
Ìàòåðèàëîì èññëåäîâàíèÿ ïîñëóæèë ïîñòêðàíèàëüíûé ñêåëåò íàíäó (Rhea americana). Â
êà÷åñòâå ñðàâíèòåëüíîãî ìàòåðèàëà ïðèâëå÷åíû äàííûå ïî ðàíåå èçó÷åííûì ëåòàþùèì ïòèöàì, â
÷àñòíîñòè êóðîîáðàçíûì (Áîãäàíîâè÷, 1997), êàê íàèáîëåå ãåíåðàëèçîâàííûì êèëåãðóäûì.
Ìîðôîìåòðè÷åñêèå äàííûå äëÿ îïðåäåëåíèÿ êîððåëÿòèâíîé ñâÿçè ìåæäó ìîðôîëîãè÷åñêèìè
ïðèçíàêàìè ïîëó÷åíû íà ñêåëåòàõ ïòèö, îòëè÷àþùèõñÿ ëîêîìîòîðíûìè àäàïòàöèÿìè è
ñèñòåìàòè÷åñêè, èç îðíèòîëîãè÷åñêîé êîëëåêöèè ÇÈÍ ÐÀÍ. Äëÿ ïîëó÷åíèÿ ïàðàìåòðîâ ðåãðåññèè è
âåëè÷èíû êîýôôèöèåíòà êîððåëÿöèè èñïîëüçîâàíà ïðîãðàììà Sigma Plot 8.0
Ïðèìåíåíà ìåòîäèêà òðàäèöèîííîé ñðàâíèòåëüíîé ìîðôîëîãèè, äîëæíîå ìåñòî êîòîðîé ñðåäè
äðóãèõ áèîëîãè÷åñêèõ äèñöèïëèí óáåäèòåëüíî àðãóìåíòèðîâàíî (Gans, 1986; Duncker, 1989) íà ôîíå
îáùåé óâëå÷åííîñòè êëàäèñòèêîé — íåñîìíåííî óäîáíûì, íî è äîñòàòî÷íî îãðàíè÷åííûì
(Òàòàðèíîâ, 1984; Gans, 1986 è äð.) ìåòîäîì àíàëèçà ìîðôîëîãè÷åñêèõ äàííûõ.
Àíàòîìè÷åñêèå íàçâàíèÿ ïðèâåäåíû ñîãëàñíî “Nomina Anatomica Avium” (1979).
Ñïèñîê âèäîâ èç ìóçåéíûõ ôîíäîâ ÇÈÍ ÐÀÍ, èñïîëüçîâàííûõ â ðàáîòå: Anser albifrons (¹ 3),
Anser erythropus (¹1980), Tetraogallus caucasicus (¹ 249), Tetraogallus himalayensis (¹ 243), Phasianus
colchicus (¹ 963), Francolinus francolinus (¹ 1057), Coturnix japonicus (¹ 1580), Alectoris kakelik
(¹ 932), Perdix perdix (¹ 805), Numida meleagris (¹ 1421), Lagopus lagopus (¹ 23), Lyrurus tetrix
(¹ 1768), Bonasa umbellus (¹ 2580), Tetrao parvirostris (¹ 240), Centrocercus urophasianus (¹ 135),
Canachites canadensis (¹ 1448), Pedioecetes phasianellus (¹ 2577/195), Lagopus leucurus (¹ 2581/135),
Crax nigra (¹ 425), Ortalis vetula (¹ 451), Crax globulosa (¹ 386), Macrocephalon maleo (¹ 413),
Megapod. gen. ? sp. ? (¹ 1447), Megapod. gen. ? sp. ? (¹ 2231), Tinamus sp. ? (¹ 427), Crex crex
(¹ 1767), Fulica atra (¹ 1464), Columba rupestrris (¹ 260/64), Goura victoria (¹ 2708), Columba
palumbus (¹ 1940), Pterocles orientalis (¹ 2557), Accipiter gentilis (¹?), Charadrius leschenaultii (¹ 1766),
Cuculus canorus (¹ 1956), Turdus philomachus (¹ 1656).
Ðåçóëüòàòû è îáñóæäåíèå
Ñðåäè ñêåëåòíûõ ïðèçíàêîâ àïïàðàòà íàçåìíîé ëîêîìîöèè íàíäó íàèáîëåå
èíòåðåñíûì ïîêàçàëîñü ñòðîåíèå òàçà. Ïðåæäå âñåãî, îáðàùàåò íà ñåáÿ âíèìàíèå
åãî êðàéíå âûðàæåííàÿ îáùàÿ ñóæåííîñòü (ðèñ. 1). Ñðåäè èçó÷åííûõ íàìè ðàíåå
ïòèö 34 âèäîâ 5 îòðÿäîâ äîðñàëüíàÿ øèðèíà òàçà (ìåæäó âåðòëóæíûìè âïàäèíà-
ìè) âàðüèðóåò îò 34,4% åãî äëèíû ó ãóñÿ äî 98,3% ó âÿõèðÿ (Áîãäàíîâè÷, 1997),
òîãäà êàê ó íàíäó îíà ñîñòàâëÿåò 19%. Ïðè ýòîì ñëåäóåò îòìåòèòü ñîâåðøåííî
ñïåöèôè÷åñêóþ åãî ÷åðòó: ïîäâçäîøíûå êîñòè äîðñàëüíî ïî÷òè ñìûêàþòñÿ ïî
ñðåäèííîé ëèíèè ïîçâîíî÷íèêà (ðèñ. 1).
Ðàíåå íàìè áûëà îòìå÷åíà çíà÷èòåëüíàÿ êîððåëÿöèÿ ìåæäó îòíîñèòåëüíîé
øèðèíîé òàçà è îòíîñèòåëüíîé øèðèíîé ãðóäíîé êëåòêè ïòèö (Áîãäàíîâè÷,
1993), çàâèñÿùåé, â ïåðâóþ î÷åðåäü, îò øèðèíû ãðóäèíû. Õîðîøî èçâåñòíî, ÷òî
ñîáñòâåííî îñíîâàíèå ãðóäèíû (corpus sterni) ÿâëÿåòñÿ ó ïòèö ïåðâè÷íûì ïî
îòíîøåíèþ ê êèëþ (Fell, 1925). Íàðÿäó ñ îòðîñòêàìè ãðóäèíû è ìåìáðàíîé èõ
ñîåäèíÿþùåé, òåëî ãðóäèíû ñëóæèò ìåñòîì ïðèêðåïëåíèÿ ãðóäíîé è íàäêîðà-
êîèäíîé ìûøö — îñíîâíûõ ìûøö, îïóñêàþùèõ è ïîäíèìàþùèõ êðûëî (Ñû÷,
1985). Îáðàçîâàíèå îòðîñòêîâ è âûðåçîê, çàòÿíóòûõ ñîåäèíèòåëüíîòêàííûìè
ìåìáðàíàìè — ýêîíîìèÿ êîñòíîé òêàíè (Øòåãìàí, 1958), èëè òî÷íåå — ðåçóëü-
òàò îáùåé òåíäåíöèè ðåäóêöèè êîñòíîé òêàíè äëÿ îáëåã÷åíèÿ ñêåëåòà (Ñû÷,
1999). Âî âñÿêîì ñëó÷àå, íàì êàæåòñÿ âïîëíå óáåäèòåëüíûì íàïðàâëåíèå ýâîëþ-
öèè ãðóäèíû îò ïëàñòèíîîáðàçíîé ê «âûðåçàíîé» â ñâÿçè ñ ñîâåðøåíñòâîâàíèåì
ëåòíûõ êà÷åñòâ. Õîòÿ îòñóòñòâèå ïîñòåðîëàòåðàëüíûõ îòðîñòêîâ ãðóäèíû (ðàâíî,
êàê è êèëÿ) ó áåñêèëåâûõ èíîãäà ðàññìàòðèâàåòñÿ êàê ðåçóëüòàò ïîòåðè ýòèõ
ñòðóêòóð è îïðåäåëÿåòñÿ ïîýòîìó êàê ïðîäâèíóòîå ñîñòîÿíèå, äîñòèãíóòîå ïîñëå
ïîòåðè ñïîñîáíîñòè ê ïîëåòó (Cracraft, 1974).  êà÷åñòâå êîíòðàðãóìåíòà ìîæíî
ïðèâåñòè ðåçóëüòàòû èññëåäîâàíèé ìîðôîëîãèè âòîðè÷íî íåëåòàþùèõ
ïðåäñòàâèòåëåé ëåòàþùèõ ïòèö. Ó íåëåòàþùåãî ïàñòóøêà Óýêà (Gallirallus aus-
tralis), íàïðèìåð, ñîõðàíÿåòñÿ ïðàêòè÷åñêè ïîëíûé íàáîð ìûøö ïëå÷åâîãî ïîÿñà
è êðûëà (McGowan, 1986). Ñðàâíåíèå ñ áåñêèëåâûìè ñî çíà÷èòåëüíî ìåíüøèì
êîëè÷åñòâîì ìûøö ïîçâîëèëî àâòîðó âûñêàçàòüñÿ â ïîëüçó ïðîèñõîæäåíèÿ
348 È. À. Áîãäàíîâè÷
ïîñëåäíèõ îò ïðåäêîâ ñ ñèëüíî îãðàíè÷åííîé ñïîñîáíîñòüþ ê ïîëåòó. Î÷åâèäíî,
÷òî îòìå÷åííûé ôàêò ñîõðàíåíèÿ «ëåòàòåëüíûõ» ÷åðò ìîðôîëîãèè ìîæíî îòíåñ-
òè è ê ñêåëåòíûì ñòðóêòóðàì.
Íà íàø âçãëÿä, ëîãè÷íî ïðåäïîëîæèòü, ÷òî íà÷àëüíûé ýòàï óñèëåíèÿ ïåêòî-
ðàëüíûõ ìûøö ó áèïåäàëüíûõ ïðåäêîâ ïòèö áûë ñîïðÿæåí ñ ðàñøèðåíèåì ãðó-
äèíû. Ê ñîæàëåíèþ, îñòàåòñÿ íå âïîëíå ïîíÿòíûì ôóíêöèîíàëüíî-ýêîëîãè-
÷åñêèå ðåçîíû âîçìîæíîãî óñèëåíèÿ óêàçàííûõ ìûøö è ãðóäíûõ êîíå÷íîñòåé â
öåëîì ïîñëå èõ îñâîáîæäåíÿ îò îïîðû. Ýòîò âîïðîñ òåñíî ñâÿçàí ñ ïðîáëåìîé
ïðîèñõîæäåíèÿ áèïåäàëèçìà, çàñëóæèâàþùåé âïîëíå ñàìîñòîÿòåëüíîãî ðàññìîò-
ðåíèÿ.  äàííîì àñïåêòå î÷åíü èíòåðåñíî èññëåäîâàíèå òåðîïîäíîãî àâèìèìà
(Êóðçàíîâ, 1987). Êðàéíå âûðàæåííîå ñõîäñòâî èõ ãðóäíûõ (âîçìîæíî îïåðåí-
íûõ) êîíå÷íîñòåé ñ ïòè÷üèì êðûëîì ïðåäïîëàãàåò ñîâåðøåííî îïðåäåëåííûå
äâèæåíèÿ, ïîìîãàþùèå ïðè áåãå, à òàêæå âñïàðõèâàíèå è ïîäëåòûâàíèå, ïîäîá-
íûå òàêîâûì ïëîõî ëåòàþùèõ êóðèíûõ (Êóðçàíîâ, 1987).  êà÷åñòâå àíàëîãèè
äîáàâèì, íàïðèìåð, ñïîñîáíîñòü ïðèìèòèâíûõ êóðèíûõ — êðàêñîâ — çàïðûãè-
âàòü ñ çåìëè (ñ ïîìîùüþ êðûëüåâ) íà íèæíèå âåòâè äåðåâüåâ (Bent, 1932).
Ïðåäïîëàãàÿ îáÿçàòåëüíîå íàëè÷èå «äðåâåñíîãî» ýòàïà â ðàííåé ýâîëþöèè ïòèö
(Áîãäàíîâè÷, 2000), îñìåëèìñÿ ïðåäïîëîæèòü, ÷òî èìåííî åãî îòñóòñòâèå â
èñòîðèè àâèìèìèä ìîãëî áûòü îäíèì èç ôàêòîðîâ, îïðåäåëèâøèõ äèâåðãåíöèþ
èõ ïðåäêîâ ñ òàêîâûìè ïòèö.
Ôóíêöèîíèðîâàíèå ãðóäíûõ êîíå÷íîñòåé àâèìèìà â óïîìÿíóòûõ âûøå
ðåæèìàõ ñ ðàçâèòèåì ñîîòâåòñòâóþùèõ ìûøö, î÷åâèäíî, íå ìîãëî íå îòðàçèòüñÿ
â ÷àñòíîñòè íà ðàñøèðåíèè ãðóäèíû. Åå îïèñàíèå, ê ñîæàëåíèþ, îòñóòñòâóåò, íî
íå ìåíåå èíòåðåñíûì â ýòîé ñâÿçè êàæóòñÿ îñîáåííîñòè ñòðîåíèÿ êðåñòöà è òàçà
àâèìèìà. Óñèëåíèå êðåñòöà (â äàííîì ñëó÷àå 7 ñðîñøèõñÿ ìåæäó ñîáîé ïîçâîí-
êîâ) ìîæíî ñ÷èòàòü äîñòàòî÷íî îáùåé òåíäåíöèåé, ïîñêîëüêó îíî â ðàçíîé
ñòåïåíè õàðàêòåðíî äëÿ áèïåäàëüíûõ äèíîçàâðîâ (Ðîæäåñòâåíñêèé, 1964;
Andreas, 1997 è äð.), îáåñïå÷èâàÿ óâåëè÷åíèå æåñòêîñòè ïðîòèâ èçãèáàíèÿ. Ó
àâèìèìà, êðîìå òîãî, îòìå÷åíû õîðîøî âûðàæåííûå ïîïåðå÷íûå îòðîñòêè, èç
349Íåêîòîðûå îñîáåííîñòè ñêåëåòà íàíäó Rhea americana…
Ðèñ. 1. Òàç ôàçàíà Phasianus colchicus (1) è íàíäó Rhea americana (2) ñ äîðñàëüíîé ñòîðîíû.
Fig. 1. Pelves of Phasianus colchicus (1) and Rhea americana (2) in dorsal view.
êîòîðûõ 4-é è 5-é îñîáåííî øèðîêèå è ñðîñøèåñÿ ìåæäó ñîáîé ïî÷òè íà âñåì
ïðîòÿæåíèè (Êóðçàíîâ, 1987). Î÷åâèäíî, ÷òî èìåííî ýòà îñîáåííîñòü îáóñëî-
âëèâàåò óâåëè÷åííóþ ñðåäè òåðîïîä øèðèíó òàçà, ìàêñèìàëüíóþ â îáëàñòè
âåðòëóæíûõ âïàäèí (Êóðçàíîâ, 1987).
Äëÿ óòî÷íåíèÿ ïîëó÷åííûõ ðàíåå äàííûõ (Áîãäàíîâè÷, 1993) ìû ïðîâåëè
áîëåå êîððåêòíîå èññëåäîâàíèå êîððåëÿòèâíîé ñâÿçè — ìåæäó øèðèíîé òàçà â
îáëàñòè âåðòëóæíûõ âïàäèí è øèðèíîé ãðóäèíû â îáëàñòè ãðóäíûõ îòðîñòêîâ
(ðèñ. 2).
Âûñîêóþ ñòåïåíü êîððåëÿöèîííîé ñâÿçè ìåæäó îáñóæäàåìûìè ìîðôîëîãè-
÷åñêèìè ïðèçíàêàìè ìû ïîïûòàëèñü îáüÿñíèòü ñëåäóþùèì îáðàçîì. Èçâåñòíî,
÷òî ïî ëèíèè ôîðìèðîâàíèÿ «ïòè÷üåé» ìîäåëè ëîêîìîòîðíîãî àïïàðàòà èìåëî
ìåñòî èçìåíåíèå îðèåíòàöèè áåäðà îò áëèçêîé ê âåðòèêàëüíîé (ñîõðàíåííîé ó
áèïåäàëüíûõ äèíîçàâðîâ) ê áîëåå ãîðèçîíòàëüíîé ñî ñìåùåíèåì ïðèîðèòåòà
äâèæåíèÿ êîíå÷íîñòè â ïîëüçó ñãèáàíèÿ—ðàçãèáàíèÿ â êîëåííîì ñóñòàâå
(Alexander, 1985; Chiappee, 1995 è äð.). Ïðè ýòîì ïðîòðàêöèÿ áåäðà ó ïòèö
ñî÷åòàåòñÿ ñ åãî îòâåäåíèåì äëÿ «îáõîæäåíèÿ» ãðóäíîé êëåòêè ñ áîêîâ (ðèñ. 3).
Íà íàø âçãëÿä, ðàñøèðåíèå ãðóäèíû ÿâëÿëîñü âåäóùèì ôàêòîðîì ðàñøèðåíèÿ
òàçà (çà ñ÷åò ýëåìåíòîâ ñèíñàêðóìà), ïîñêîëüêó ñòåïåíü îòâåäåíèÿ áåäðà âïîëíå
îïðåäåëåííî îãðàíè÷åíà ñòðîåíèåì òàçîáåäðåííîãî ñóñòàâà.
Ïîìèìî èçëîæåííîé âûøå, íåîáõîäèìî ñêàçàòü î âîçìîæíî èíîé ïðè÷èíå
ðàñøèðåíèÿ ãðóäíîé êëåòêè ïòèö, ñâÿçàííîé ñ åå «äûõàòåëüíîé» ôóíêöèåé, à
èìåííî: ñîçäàíèå êàðêàñà äëÿ êðóïíûõ ëåãî÷íûõ ìåøêîâ ñ èçìåíÿåìûì îáüåìîì
è ñèñòåìîé âåíòèëÿöèè, îáåñïå÷èâàþùåé âûñîêóþ ýôôåêòèâíîñòü ãàçîîáìåíà â
ëåãêèõ (Äçåðæèíñêèé, 2005). Îäíàêî ôîðìèðîâàíèå òàêîé äûõàòåëüíîé ñèñòåìû
íàì êàæåòñÿ áîëåå ïîçäíèì ïðèîáðåòåíèåì, ñïîñîáñòâóþùèì ïîâûøåíèþ
óðîâíÿ âîçìîæíîñòåé óæå èìåþùåãîñÿ àêòèâíîãî ïîëåòà.
Êðàéíå ìåäëåííàÿ îññèôèêàöèÿ ãðóäèíû Ratitae, èíîãäà íåïîëíàÿ äàæå âî
âçðîñëîì ñîñòîÿíèè, ñáëèæàåò èõ ñ ðåïòèëèÿìè è ðàññìàòðèâàåòñÿ â êà÷åñòâå
350 È. À. Áîãäàíîâè÷
Ðèñ. 2. Ðåãðåññèÿ äîðñàëüíîé øèðèíû òàçà íà øèðèíó ãðóäèíû (îáúÿñíåíèÿ â òåêñòå).
Fig. 2. Regression of dorsal width of pelvis on width of sternum (explanations in the text).
ïðèìèòèâíîãî äëÿ ãðóïïû ïðèçíàêà (Elzanowski, 1989). Ó÷èòûâàÿ ïðåäïîëà-
ãàåìûå íàìè ìîðôîëîãè÷åñêèå âçàèìîñâÿçè, ôîðìó òàçà íàíäó (åãî êðàéíå
ìàëóþ äîðñàëüíóþ øèðèíó) òàêæå ìîæíî ïðèíÿòü êàê ïðèìèòèâíóþ.
Ñîõðàíåíèå óçêîãî òàçà îñîáåííî âàæíî ïî ëèíèè èñêëþ÷èòåëüíî íàçåìíûõ
áåãàþùèõ ïòèö, ïîñêîëüêó îáåñïå÷èâàåò ñðàâíèòåëüíî ìàëûå ïëå÷î äåéñòâèÿ è
ñîîòâåòñòâåííî ìîìåíò ñèëû, âîçíèêàþùåé ïðè òîë÷êå áåãóùåé ïòèöû. Âåðîÿò-
íîå, ñêîððåëèðîâàííîå ñ ðàñøèðåíèåì (ïî òîé èëè èíîé ïðè÷èíå) ãðóäíîé
êëåòêè, ðàñøèðåíèå òàçà ó ëåòàþùèõ ïòèö ñ óìåíüøåíèåì åãî ïðî÷íîñòè
«êîìïåíñèðîâàëîñü» ðàçâèòèåì àïïàðàòà ïîëåòà. Îäíî èç âîçìîæíûõ ñëåäñòâèé
òàêîãî ðàñøèðåíèÿ — âîçìîæíîñòü óâåëè÷åíèÿ ðàçìåðà ÿèö — âïîëíå ïðî-
ãðåññèâíîãî ïðèçíàêà.
Ïðè îòñóòñòâèè êàêîãî-ëèáî ôóíêöèîíàëüíî-ýêîëîãè÷åñêîãî ñõîäñòâà
ìåæäó ñïåöèàëèçèðîâàííûìè «íàçåìíèêàìè» — Ratitae — è íå ìåíåå ñïåöèà-
ëèçèðîâàííûìè â îòíîøåíèè âîäíîé ëîêîìîöèè Hesperornithidae ïðèíöè-
ïèàëüíîå ñõîäñòâî â ñòðîåíèè èõ òàçà ïî îáñóæäàåìîìó ïðèçàêó (Marsh, 1880;
Cracraft, 1982; ðèñ. 1) âïîëíå ëîãè÷íî, íà íàø âçãëÿä, îáüÿñíèòü èõ ôèëîãåíå-
òè÷åñêîé áëèçîñòüþ. Ïîäîáíîå ïðåäïîëîæåíèå áûëî âûñêàçàíî Ì. Ôþðáðèí-
ãåðîì (Furbringer, 1888) â îòíîøåíèè Hesperornithidae è Struthionidae, Rheidae.
Èìåííî ýòèõ ïðåäñòàâèòåëåé áåñêèëåâûõ è Dromaeus (êîòîðûõ îáüåäèíÿåò, â
÷àñòíîñòè, è êðàéíå âûðàæåííàÿ ñóæåííîñòü òàçà). À. Ñóäèëîâñêàÿ (1931)
ïðåäïîëîæèòåëüíî ñ÷èòàåò ïðîèçîøåäøèìè îò îáùåãî ïðåäêà. Íàëè÷èå ñðàâíè-
òåëüíî øèðîêîãî òàçà ó Aepyornis (Cracraft, 1974) ìîãëî áû ñäåëàòü íàøè
ïîñòðîåíèÿ íåñêîëüêî ñîìíèòåëüíûìè ïðè óñëîâèè äîêàçàííîé ìîíîôèëå-
òè÷íîñòè Ratitae. Îäíàêî âîïðîñ ýòîò áûë è îñòàåòñÿ ñïîðíûì êàê â îòíîøåíèè
ïàëåîãíàò â öåëîì, òàê è áåñêèëåâûõ â ÷àñòíîñòè (Furbringer, 1888; Kurochkin,
1995 è äð.). Èììóíîëîãè÷åñêèå äîêàçàòåëüñòâà ìîíîôèëåòè÷íîñòè Ratitae ïðèâå-
äåíû, íàïðèìåð, â ðàáîòå Ñ. Ñòåéïåëÿ ñ ñîàâòîðàìè (Stapel et al., 1984), õîòÿ
ìàòåðèàëîì çäåñü òàêæå ïîñëóæèëè ëèøü óïîìÿíóòûå âûøå ïðåäñòàâèòåëè
(Struthio, Rhea, Dromaeus).
Íàëè÷èå âïîëíå êðûëîâèäíûõ ãðóäíûõ êîíå÷íîñòåé ó ÿâíî íåëåòàâøèõ
òåðîïîäíûõ äèíîçàâðîâ (Êóðçàíîâ, 1987; Xu et al., 1999 è äð. ), ïî êðàéíåé ìåðå,
351Íåêîòîðûå îñîáåííîñòè ñêåëåòà íàíäó Rhea americana…
Ðèñ. 3. Ïîëîæåíèå ïðîòðàãèðîâàííîãî áåäðà ïî îòíîøåíèþ ê ãðóäíîé êëåòêå (âèä ñçàäè): f — femur;
tt — tibiotarsus; pr. th. — processus thoracicus; pr. cl. — processus caudolateralis; c. st — corpus sterni.
Fig. 3. Position of protracted femur concerning to thorax (posterior view): f — femur; tt — tibiotarsus;
pr. th. — processus thoracicus; pr. cl. — processus caudolateralis; c. st — corpus sterni.
çàìåòíî ñíèæàåò öåííîñòü ýòîãî ïðèçíàêà êàê äîêàçàòåëüñòâà âòîðè÷íîñòè
íåëåòàíèÿ ó áåñêèëåâûõ. Òî æå ìîæíî ñêàçàòü â îòíîøåíèè ïèãîñòèëÿ, îïèñàí-
íîãî äëÿ Struthio camelus (Beer de, 1956). Îòñóòñòâóåò îí êàê ó ïðåäïîëîæèòåëüíî
âòîðè÷íî íåëåòàþùèõ ãåñïåðîðíèñîâ è áîëüøèíñòâà áåñêèëåâûõ, òàê è ó
íåêîòîðûõ ëåòàþùèõ òèíàìóîáðàçíûõ (Bellairs, Jenkin, 1960), îäíàêî îáíàðóæåí
ó òåðîïîäíîãî îâèðàïòîçàâðà (Barsbold et al., 2000).
Íå ìåíåå èíòåðåñíîé îñîáåííîñòüþ òàçà íàíäó ÿâëÿåòñÿ óäëèíåííîñòü åãî
ïîñòàöåòàáóëÿðíîãî îòäåëà, êîòîðûé ñóùåñòâåííî äëèííåå ïðåàöåòàáóëÿðíîãî
(ïîäîáíîå ñîñòîÿíèå ó àôðèêàíñêîãî ñòðàóñà, Cracraft, 1974) çà ñ÷åò ñåäàëèùíîé
êîñòè (ðèñ 1). Ñðåäè ëåòàþùèõ ïòèö ýòîò ïðèçíàê õàðàêòåðåí äëÿ âîäîïëàâàþ-
ùèõ è ôîðìèðîâàëñÿ â ñîîòâåòñòâèè ñ òðåáîâàíèÿìè ê òåëó, êîòîðîå íàõîäèòñÿ
â âîäå (Êóðî÷êèí, 1971 è äð.). Òîãäà êàê äëÿ áîëåå àäàïòèðîâàíûõ ê íàçåìíîìó
ïåðåäâèæåíèþ âèäîâ õàðàêòåðíà ïðèìåðíî ðàâíàÿ äëèíà ïðå- è ïîñòàöåòàáó-
ëÿðíîãî îòäåëîâ èëè ïðåèìóùåñòâåííàÿ äëèíà ïåðâîãî (Soni, Saxena, 1980;
Áîãäàíîâè÷, 1997). Âîçìîæíî, «íåàäàïòèâíàÿ», íà ïåðâûé, âçãëÿä óäëèíåííîñòü
ïîñòàöåòàáóëÿðíîãî îòäåëà òàçà ó íàíäó âçàèìîñâÿçàíà, â ÷àñòíîñòè, ñ êàóäà-
ëèçàöèåé ìåñòà ïðîêñèìàëüíîé ôèêñàöèè çàäíåáåäðåííûõ ìûøö — ðåòðàêòîðîâ
áåäðà. Íå îñòàíàâëèâàÿñü çäåñü íà ñðàâíèòåëüíîé ìèîëîãèè íàíäó, êîòîðàÿ
çàñëóæèâàåò îäåëüíîé ïóáëèêàöèè, îòìåòèì òîëüêî, ÷òî îáå ïîðöèè m. caudo-
ilio-femoralis è m. cruris flexor lateralis íà÷èíàþòñÿ îò íàèáîëåå êàóäàëüíîé ÷àñòè
ñåäàëèùíîé êîñòè. Óäëèíåíèå èõ âîëîêîí ïðåäïîëàãàåò ñðàâíèòåëüíî áîëüøóþ
àìïëèòóäó è ñêîðîñòü ñîêðàùåíèÿ è ñîîòâåòñòâåííî ñãèáàòåëüíî-ðàçãèáàòåëüíûõ
äâèæåíèé â òàçîáåäðåííîì ñóñòàâå. Îíè èìåëè áîëüøîå çíà÷åíèå â ëîêîìîöèè
äèíîçàâðîâ (Chiappee, 1995; Jones et al., 2000) è ñðàâíèòåëüíî îãðàíè÷åíû ó
352 È. À. Áîãäàíîâè÷
Ðèñ. 4. Ïðîêñèìàëüíûé êîíåö áåäðåííîé êîñòè íàíäó Rhea americana (1) è ôàçàíà Phasianus colchicus
(2) c äîðñàëüíîé (A) è êðàíèàëüíîé (Á) ñòîðîíû: cr. tr. — crista trochanteris; fac. art. ant. — facies
articularis antitrochanterica; cap. fem. — caput femoris; col. fem. — collum femoris.
Fig. 4. Proximal end of femur of Rhea americana (1) and Phasianus colchicus (2) from dorsal (A) and cranial
(Á) view: cr. tr. — crista trochanteris; fac. art. ant. — facies articularis antitrochanterica; cap. fem. — caput
femoris; col. fem. — collum femoris.
21
A
B
ñîâðåìåííûõ ëåòàþùèõ ïòèö (Cracraft, 1971; Ñû÷ è äð., 1985), ÷òî ñâÿçàíî ñ
èçìåíåíèåì îðèåíòàöèè áåäðà. Ê ñîæàëåíèþ, ó íàñ íåò ïðÿìûõ äàííûõ îá
îðèåíòàöèè áåäðà ó íàíäó. Îäíàêî î çíà÷èòåëüíîì ðàçìàõå óêàçàííûõ äâèæåíèé
ìîæåò ñâèäåòåëüñòâîâàòü âåëè÷èíà äîðñàëüíîé ñóñòàâíîé ïîâåðõíîñòè áåäðà,
çàìåòíî óäëèíåííîé êðàíèîêàóäàëüíî â ñðàâíåíèè ñ èçó÷åííûìè íàìè ðàíåå
ëåòàþùèìè ïòèöàìè (ðèñ. 4).
Çàêëþ÷åíèå
Èçëîæåííûå íàìè ñîîáðàæåíèÿ íå ïðåäïîëàãàþò êàêèõ-ëèáî îäíîçíà÷íûõ
âûâîäîâ îòíîñèòåëüíî ýâîëþöèè è ôèëîãåíèè èçó÷åííîãî ïðåäñòàâèòåëÿ áåñêè-
ëåâûõ. Îäíàêî ïðîâåäåííûé àíàëèç, êîòîðûé íåñêîëüêî îòëè÷àåòñÿ îò áîëü-
øèíñòâà èçâåñòíûõ íàì ðåøåíèé ïî ýòèì âîïðîñàì, ïîçâîëÿåò âûñêàçàòüñÿ â
ïîëüçó ãèïîòåçû î ãëóáîêîé ïðèìèòèâíîñòè íàíäó è ïåðâè÷íîñòè åãî íåëåòàíèÿ.
Áîãäàíîâè÷ È. À. Êîððåëÿòèâíûå âçàèìîñâÿçè íåêîòîðûõ ìîðôîëîãè÷åñêèõ ïðèçíàêîâ àïïàðàòà
íàçåìíîé ëîêîìîöèè ïòèö // Âåñòí. çîîëîãèè. — 1993. — ¹ 4. — Ñ. 75–77.
Áîãäàíîâè÷ È. À. Àïïàðàò íàçåìíîé ëîêîìîöèè òåòåðåâèíûõ (Tetraonidae, Galliformes) è äðóãèõ
êóðîîáðàçíûõ. Ìîðôî-ýêîëîãè÷åñêàÿ õàðàêòåðèñòèêà // Âåñòí. çîîëîãèè. — 1997. — Îòä.
âûï. ¹ 3. — 152 ñ.
Áîãäàíîâè÷ È. À. Òðàíôîðìàöèè ñòîïû â ðàííåé ýâîëþöèè ïòèö // Âåñòí. çîîëîãèè. — 2000. — 34,
¹ 4–5. — Ñ. 123–127.
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| id | nasplib_isofts_kiev_ua-123456789-65484 |
| institution | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| issn | 0084-5604 |
| language | Russian |
| last_indexed | 2025-12-07T16:41:53Z |
| publishDate | 2008 |
| publisher | Інститут зоології ім. І.І. Шмальгаузена НАН України |
| record_format | dspace |
| spelling | Богданович, И.А. 2014-06-26T05:26:34Z 2014-06-26T05:26:34Z 2008 Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых / И.А. Богданович // Вестник зоологии. — 2008. — Т. 42, № 4. — С. 347–354. — Бібліогр.: 45 назв. — рос. 0084-5604 https://nasplib.isofts.kiev.ua/handle/123456789/65484 В сравнительном и функциональном аспектах проанализированы некоторые морфологические особенности таза и бедра представителя бескилевых — нанду (Rhea americana). Определена высокая степень корреляции между шириной таза и шири¬ной грудины у представителей 35 видов 10 отрядов килегрудых птиц. В приложении к морфо¬логии таза нанду и наиболее близкородственным представителям группы указанная зависимость позволяет, в частности, высказаться в пользу первичности их нелетания. Some peculiarities of Rhea americana pelvis and femur are analyzed in comparative and functional aspects. High level of correlation between the width of the pelvis and that of the sternum is shown to occur in representatives of 35 species and 10 orders of the carinate birds. In respect to pelvis morphology of Rhea and close related ratitae, it allows to suppose in particular the primary character of its flightlessness. ru Інститут зоології ім. І.І. Шмальгаузена НАН України Вестник зоологии Морфология Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых Some Skeletal Peculiarities of Rhea americana (Rheiformes, Rheidae) with Respect of Ratitae Evolution Article published earlier |
| spellingShingle | Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых Богданович, И.А. Морфология |
| title | Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых |
| title_alt | Some Skeletal Peculiarities of Rhea americana (Rheiformes, Rheidae) with Respect of Ratitae Evolution |
| title_full | Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых |
| title_fullStr | Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых |
| title_full_unstemmed | Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых |
| title_short | Некоторые особенности скелета нанду Rhea americana (Rheiformes, Rheidae) в свете эволюции бескилевых |
| title_sort | некоторые особенности скелета нанду rhea americana (rheiformes, rheidae) в свете эволюции бескилевых |
| topic | Морфология |
| topic_facet | Морфология |
| url | https://nasplib.isofts.kiev.ua/handle/123456789/65484 |
| work_keys_str_mv | AT bogdanovičia nekotoryeosobennostiskeletanandurheaamericanarheiformesrheidaevsveteévolûciibeskilevyh AT bogdanovičia someskeletalpeculiaritiesofrheaamericanarheiformesrheidaewithrespectofratitaeevolution |