Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ
Аллозимный анализ неритид рода Theodoxus Montfort, 1810 подтвердил эволюционно-генетическую дискретность трех видов: T. astrachanicus, T. fluviatilis, T. danubialis, тогда как T. danasteri, T. sarmaticus, T. velox и T. euxinus по данным генного маркирования оказались конспецифичными T. fluviatilis....
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| Veröffentlicht in: | Вестник зоологии |
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| Datum: | 2011 |
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| Format: | Artikel |
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Інститут зоології ім. І.І. Шмальгаузена НАН України
2011
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| Назва журналу: | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| Zitieren: | Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ / С.В. Межжерин, Ю.В. Тарасова, Е.И. Жалай // Вестник зоологии. — 2011. — Т. 45, № 6. — С. 531–540. — Бібліогр.: 15 назв. — рос. |
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Digital Library of Periodicals of National Academy of Sciences of Ukraine| _version_ | 1859861173408628736 |
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| author | Межжерин, С.В. Тарасова, Ю.В. Жалай Е.И. |
| author_facet | Межжерин, С.В. Тарасова, Ю.В. Жалай Е.И. |
| citation_txt | Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ / С.В. Межжерин, Ю.В. Тарасова, Е.И. Жалай // Вестник зоологии. — 2011. — Т. 45, № 6. — С. 531–540. — Бібліогр.: 15 назв. — рос. |
| collection | DSpace DC |
| container_title | Вестник зоологии |
| description | Аллозимный анализ неритид рода Theodoxus Montfort, 1810 подтвердил эволюционно-генетическую дискретность трех видов: T. astrachanicus, T. fluviatilis, T. danubialis, тогда как T. danasteri, T. sarmaticus, T. velox и T. euxinus по данным генного маркирования оказались конспецифичными T. fluviatilis. К такому же выводу приводят и результаты морфометрического анализа раковины и половой системы.
Allozyme analysis of neritid mollusks of the genus Theodoxus Montfort, 1810 shows evolutionary and genetical discreteness of the three species T. astrachanicus, T. fluviatilis, T. danubialis, whereas T. danasteri, T. sarmaticus, T. velox and T. euxinus species were found to be conspecific with T. fluviatilis according to the results of gene labeling. The results of morphometric analysis of the shell and genital system lead support this conclusion.
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ÓÄÊ 594:575.8(477)
ÌÎËËÞÑÊÈ ÐÎÄÀ THEODOXUS (GASTRAPODA,
NERITIDAE) ÓÊÐÀÈÍÛ: ÌÎÐÔÎËÎÃÈ×ÅÑÊÈÉ
È ÀËËÎÇÈÌÍÛÉ ÀÍÀËÈÇ
Ñ. Â. Ìåææåðèí1, Þ. Â. Òàðàñîâà2, Å. È. Æàëàé1
1 Èíñòèòóò çîîëîãèè èì. È. È. Øìàëüãàóçåíà ÍÀÍ Óêðàèíû,
óë. Á. Õìåëüíèöêîãî, 15, Êèåâ, 01601 Óêðàèíà
Å-mail: mezh@izan.kiev.ua
2 Æèòîìèðñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò èì. Èâàíà Ôðàíêî,
óë. Á. Áåðäè÷åâñêàÿ, 40, Æèòîìèð, 10002 Óêðàèíà
Ïîëó÷åíî 11 èþíÿ 2010
Ïðèíÿòî 10 íîÿáðÿ 2011
Ìîëëþñêè ðîäà Theodoxus (Gastrîpoda, Neritidae) Óêðàèíû: ìîðôîëîãè÷åñêèé è àëëîçèìíûé àíà-
ëèç. Ìåææåðèí Ñ. Â., Òàðàñîâà Þ. Â, Æàëàé Å. È. – Àëëîçèìíûé àíàëèç íåðèòèä ðîäà
Theodoxus Montfort, 1810 ïîäòâåðäèë ýâîëþöèîííî-ãåíåòè÷åñêóþ äèñêðåòíîñòü òðåõ âèäîâ:
T. astrachanicus, T. fluviatilis, T. danubialis, òîãäà êàê T. danasteri, T. sarmaticus, T. velox è T. euxi-
nus ïî äàííûì ãåííîãî ìàðêèðîâàíèÿ îêàçàëèñü êîíñïåöèôè÷íûìè T. fluviatilis. Ê òàêîìó æå
âûâîäó ïðèâîäÿò è ðåçóëüòàòû ìîðôîìåòðè÷åñêîãî àíàëèçà ðàêîâèíû è ïîëîâîé ñèñòåìû.
Êëþ÷ å âûå ñ ë î â à: ñèñòåìàòèêà, Theodoxus, àëëîçèìíàÿ èçìåí÷èâîñòü, ìîðôîìåòðèÿ,
Óêðàèíà.
Mollusks of the Genus Theodoxus (Gastrîpoda, Neritidae) of Ukraine: the Morphological Analysis and
Analysis of Allosymes. Mezhzherin S. V., Tarasova J. V., Zhalay E. I. – Allozyme analysis of neritid
mollusks of the genus Theodoxus Montfort, 1810 shows evolutionary and genetical discreteness of the
three species T. astrachanicus, T. fluviatilis, T. danubialis, whereas T. danasteri, T. sarmaticus, T. velox
and T. euxinus species were found to be conspecific with T. fluviatilis according to the results of gene
labeling. The results of morphometric analysis of the shell and genital system lead support this conclu-
sion.
Ke y wo r d s: systematics, Theodoxus, allozyme variation, morphometry, Ukraine.
Ââåäåíèå
Ñèñòåìàòèêà ìîëëþñêîâ-íåðèòèä ðîäà Theodoxus Montfort, 1810 òðàäèöèîííî ÿâëÿåòñÿ ïîëåì
òàêñîíîìè÷åñêîé íåîïðåäåëåííîñòè è îáëàñòüþ îáøèðíûõ íàó÷íûõ äèñêóññèé ìàëàêîëîãîâ.
Ïðè÷èíîé ýòîãî ÿâëÿåòñÿ íåîáû÷àéíàÿ äëÿ ïðåñíîâîäíûõ áðþõîíîãèõ èçìåí÷èâîñòü îêðàñêè ðàêî-
âèíû, èìåþùàÿ âûðàæåííûé äèñêðåòíûé õàðàêòåð. Â íàñòîÿùåå âðåìÿ â àêâàòîðèÿõ Óêðàèíû âûäå-
ëÿþò íå ìåíåå 6 ôîðì, êîòîðûì ñèñòåìàòèêè ñòðîãî òèïîëîãè÷åñêîé øêîëû ïðèäàþò ðàíã âèäîâ
(Àíèñòðàòåíêî, Àíèñòðàòåíêî, 2001; Îïðåäåëèòåëü…, 2004). Âìåñòå ñ òåì ìîðôîãåíåòè÷åñêèå èññëå-
äîâàíèÿ, ïðîâåäåííûå åùå â 50-õ ãîäàõ ÕÕ ñò. (Neumann, 1959 a, b), ïîêàçàëè, ÷òî èçìåí÷èâîñòü
îêðàñêè ñàìîãî ðàñïðîñòðàíåííîãî âèäà T. fluviatilis – íè ÷òî èíîå, êàê ýïèãåíåòè÷åñêèé ïîëèìîð-
ôèçì, âûçâàííûé ìîäèôèöèðîâàííûìè óñëîâèÿìè ñðåäû (ñîëåíîñòüþ âîäû, òåìïåðàòóðîé è ò. ä.),
äåéñòâèåì ãåíîâ, îòâå÷àþùèõ çà îêðàñêó ðàêîâèíû. Ýêñïåðèìåíòû ïîêàçàëè, ÷òî ïåðåõîä ñ ñåêòî-
ðàëüíîé íà ïîïåðå÷íîïîëîñàòóþ îêðàñêó èëè ïÿòíèñòóþ âûçâàí àëüòåðíàòèâíûìè ñîñòîÿíèÿìè
íåñêîëüêèõ ãåíîâ, õàðàêòåð ýêñïðåññèè êîòîðûõ çàâèñèò îò ïîêàçàòåëåé ñðåäû. Ïîýòîìó îñîáè îäíî-
ãî è òîãî æå âèäà, îáèòàþùèå â ðå÷íîé èëè ñîëîíîâàòîé âîäå, ñ íèçêèì èëè âûñîêèì çíà÷åíèåì ðÍ
èëè ïðîñòî â ðàçíûõ òåìïåðàòóðíûõ óñëîâèÿõ, ìîãóò îòëè÷àòüñÿ ïî îêðàñêå â ñòåïåíè íå ìåíüøåé,
÷åì êëàññè÷åñêèå âèäû.
Îáøèðíûé âíóòðèâèäîâîé ïîëèìîðôèçì, ìàñêèðóþùèé òàêñîíîìè÷åñêè âàæíûå îòëè÷èÿ,
ïðèâåë ê òîìó, ÷òî ïðîâåäåíèå ÷åòêîé ãðàíèöû ìåæäó âèäàìè ëóíîê ïî ìîðôîëîãè÷åñêèì ïðèçíà-
êàì îêàçàëîñü çàäà÷åé êðàéíå ñëîæíîé.  ðåçóëüòàòå âèäîâîé ñîñòàâ, ïðèâîäèìûé òîëüêî äëÿ ôàóíû
Vestnik zoologii, 45(6): 531—540, 2011 Ìîðôîëîãèÿ
Óêðàèíû, ïîñòîÿííî ìåíÿëñÿ. Â. È. Æàäèí (1933, 1952) óêàçûâàë ÷åòûðå âèäà T. danubialis, T. fluvia-
tilis, T. pallasi è T. transversalis. À. Ë. Ïóòü (1972) â ýòîò ñïèñîê åùå äîáàâèë
T. dniestroviensis. ß. È. Ñòàðîáàãîòîâ (1970) ïðèâîäèò äëÿ Óêðàèíû òðè âèäà: T. danubialis, T. fluviatilis
è T. prevostianus. Â êîíöå 90-õ ãîäîâ ÕÕ ñò. ñòîðîííèêàìè óçêîé êîíöåïöèè âèäà (Àíèñòðàòåíêî è
äð., 1999; Àíèñòðàòåíêî, Àíèñòðàòåíêî, 2000) ïðîâåäåíà ðåâèçèÿ, â ðåçóëüòàòå êîòîðîé âñå ðàçíîîá-
ðàçèå ôîðì ñâåäåíî ê øåñòè âèäàì, ñðåäè êîòîðûõ îòñóòñòâóåò T. danubialis, óïîìèíàâøèéñÿ ðàíåå â
ôàóíå Óêðàèíû âñåìè èññëåäîâàòåëÿìè áåç èñêëþ÷åíèÿ.  ïîñëåäíåì îïðåäåëèòåëå áåñïîçâîíî÷íûõ
ôàóíû Ðîññèè (Îïðåäåëèòåëü…, 2004) êîëëåãè ïîääåðæàëè èìåííî ýòó êîíöåïöèþ. Òåì íå ìåíåå
íåäàâíèå èññëåäîâàíèÿ ïðåäïðèíÿòûå â íèçîâüÿõ Äóíàÿ (Æàëàé è äð., 2008), äîêàçàëè íàëè÷èå åùå
è T. danubialis, à ýòî îçíà÷àåò, ÷òî â ôàóíå Óêðàèíû ïî ïîñëåäíèì äàííûì äîëæíî íàñ÷èòûâàòüñÿ
ñåìü âèäîâ ýòîãî ðîäà. Âìåñòå ñ òåì çàïàäíîåâðîïåéñêèå èññëåäîâàòåëè (Glöer, 2002; Glöer, Meier-
Brook, 2003), òðàäèöèîííî ïðèäåðæèâàþùèåñÿ øèðîêîé êîíöåïöèè, äëÿ áàññåéíîâ ðåê Óêðàèíû
óêàçûâàþò âîçìîæíîñòü ïðèñóòñòâèÿ òîëüêî òðåõ âèäîâ T. danubialis, T. fluviatilis è T. transversalis.
Òàêèì îáðàçîì, íà ñåãîäíÿ ìîæíî òîëüêî ñêàçàòü: ñêîëüêî áûëî èññëåäîâàòåëåé, ñòîëüêî æå
áûëî è ìíåíèé î âèäîâîì ñîñòàâå ýòîãî ðîäà. Èìåííî ýòè ïðîòèâîðå÷èÿ çàñòàâëÿþò ñ îñîáûì âíè-
ìàíèåì îòíåñòèñü ê ýòèì íåîáû÷àéíî èçìåí÷èâûì ìîëëþñêàì, ïðîâåäÿ èõ öåëåíàïðàâëåííîå èññëå-
äîâàíèå, áàçèðóþùååñÿ íà áèîõèìè÷åñêîì ãåííîì ìàðêèðîâàíèè è ìíîãîìåðíîé ñòàòèñòè÷åñêîé
îáðàáîòêå äàííûõ ìîðôîìåòðèè ðàçíûõ ñèñòåì ïðèçíàêîâ.
Ìàòåðèàë è ìåòîäû
Îñíîâîé èññëåäîâàíèÿ ïîñëóæèëè âûáîðêè ñåìè ïðåäïîëàãàåìûõ âèäîâ ìîëëþñêîâ, ñîáðàííûå
ëåòîì è îñåíüþ 2007—2009 ãã. â îñíîâíûõ ðå÷íûõ ñèñòåìàõ Óêðàèíû (òàáë. 1). Ïðåäâàðèòåëüíîå óñòà-
íîâëåíèå âèäîâîé ïðèíàäëåæíîñòè ìîëëþñêîâ áûëî ïðîâåäåíî ïî îñíîâíûì äèàãíîñòè÷åñêèì ïðè-
çíàêàì: õàðàêòåðó ðèñóíêà, öâåòó è ôîðìå ðàêîâèíû è åå îòäåëüíûõ ÷àñòåé (ðèñ. 1).
532 Ñ. Â. Ìåææåðèí, Þ. Â. Òàðàñîâà, Å. È. Æàëàé
Òàáëèö à 1. Ìåñòà ñáîðà âûáîðîê ñåìè ïðåäïîëàãàåìûõ âèäîâ íåðèòèä ðîäà Theodoxus
Ta b l e 1. Sample localities of species of genus Theodoxus
Âèä ¹ Ìåñòî âûáîðêè
Êîîðäèíàòû, î
ñåâ. øèðîòà /
âîñò. äîëãîòà
n
T. fluviati-
lis
1 ð. Òåòåðåâ, Æèòîìèð 50.2500 / 28.6667 20
2 ð. Çáðó÷, ñ. Ñêàëà Ïîäîëüñêàÿ 48.8500 / 26.2000 12
3 ð. Ãóéâà, ñ. Ïðÿæåâî 50.1852 / 28.6761 20
4 ð. Ðîñòàâèöà, ñ. Øàìðàåâêà 49.7719 / 29.8327 20
5 ð. Ðîñü, ñ. Ïîëîâåöêîå 49.5288 / 31.0102 20
6 ð. Ïðîòîêà, ã. Áåëàÿ Öåðêîâü 49.8019 / 30.1151 10
7 ð. Ðîñü, ñ. Ãàðáóçèí 49.4460 / 31.3359 21
8 ð. Äíåïð, ã. Ðæèùåâ 49.9656 / 31.0360 11
9 ð. Äíåïð, ñ. Ñîêèðíîå 49.1869 / 32.1147 22
10 ð. Äíåïð, ñ. Ñòðàõîëåñüå 51.0772 / 30.3938 7
11 ð. Äóíàé, ã. Âèëêîâî 45.4000 / 29.5833 50
12 ð. Ñåâåðñêèé Äîíåö, ã. Èçþì 49.2128 / 37.2569 11
T. danaste-
ri
13 ð. Ãîðûíü, ñ. Êîçëèí 50.7222 / 26.4292 20
14 ð. Äíåïð, ñ. Ñîêèðíîå 49.1870 / 32.1147 10
15 ð. Ðîñü, ñ. Ãàðáóçèí 49.4461 / 31.3360 10
16 ð. Äíåïð, ã. Ðæèùåâ 49.9656 / 31.0361 12
17 ð. Ãíèëîé Òèêè÷, ñ. Ãóëÿéïîëå 47.6631 / 36.2612 10
T. sarmati-
cus
18 Äíåñòðîâñêèé ëèìàí, ã. Áåëãîðîä-Äíåñòðîâñêèé 46.1868 / 30.3284 11
19 ð. Äíåñòð, ñ. Ìàÿêè 46.4122 / 30.2731 22
20 ð. Þæíûé Áóã, ñ. Âàðâàðîâêà 49.5362 / 35.2083 10
21 Áóãñêèé ëèìàí, ñ. Ñòàðàÿ Áîãäàíîâêà 46.8413 / 31.9086 12
T. velox 22 ð. Äíåïð, ã. Õåðñîí 46.6333 / 32.5833 27
23 ð. Èíãóëåö, ñ. Äàðüåâêà 46.7507 / 32.7887 25
24 Çáóðüåâñêèé ëèìàí, ñ. Ðûáàëüñêîå 46.4747/ 32.2091 30
25 ð. Êîíêà, ã. Öþðóïèíñê 46.6200 / 32.7200 20
26 ð. Êîíêà, ã. Ãîëàÿ Ïðèñòàíü 46.5166 / 32.5167 10
T. euxinus 27 Äíåñòðîâñêèé ëèìàí, ã. Áåëãîðîä Äíåñòðîâñêèé 46.1867 / 30.3284 20
T. astrac-
hanicus
28 Óòëþêñêèé ëèìàí c. Ñòåïîê 46.3630 / 35.3616 32
T. danu-
bialis
29 ð. Äóíàé, ã. Âèëêîâî 45.4000/ 29.5833 47
533Ìîëëþñêè ðîäà Theodoxus (Gastrîpoda, Neritidae) Óêðàèíû…
Ðèñ. 1. Âíåøíèé âèä è îêðàñêà ðàêîâèíû ñåìè ïðåäïîëàãàåìûõ âèäîâ ìîëëþñêîâ ðîäà Theodoxus:
1 – T. fluviatilis; 2 – T. danasteri; 3 – T. sarmaticus; 4 – T. velox; 5 – T. euxinus; 6 – T. astrachanicus;
7 – T. danubialis.
Fig. 1. Habitus and coloration of the shell in seven supposed species of the genus Theodoxus: 1 – T. fluvi-
atilis; 2 – T. danasteri; 3 – T. sarmaticus; 4 – T. velox; 5 – T. euxinus; 6 – T. astrachanicus; 7 – T. danu-
bialis.
Ìåòîäîì ýëåêòðîôîðåçà â 7,5%-íîì ïîëèàêðèëàìèäíîì ãåëå âîäíûõ ýêñòðàêòîâ ìÿãêèõ òêàíåé
òåëà ìîëëþñêîâ èññëåäîâàíà àëëîçèìíàÿ èçìåí÷èâîñòü ñëåäóþùèõ ôåðìåíòîâ: àñïàðòàòàìèíîòðàí-
ñôåðàçû (Aat), ìàëàòäåãèäðîãåíàçû (Mdh) è íåñïåöèôè÷åñêîé ýñòåðàçû (Es), êîäèðóåìûõ ñîîòâåò-
ñòâóþùèìè ëîêóñàìè Aat, Mdh, Es-1, Es-2, Es-3.
Ìîðôîìåòðè÷åñêèé àíàëèç ïðîâîäèëè ïî äâóì ãðóïïàì ïðèçíàêîâ. Êîíõèîëîãè÷åñêèå – èçìå-
ðÿëè ñ ïîìîùüþ øòàíãåíöèðêóëÿ: âûñîòà (ShH – shell heigth) è øèðèíà (ShW – shell width) ðàêî-
âèíû, âûñîòà (AH – aperture heigth) è øèðèíà (AW – aperture width) óñòüÿ, äëèíà ðàêîâèíû (ShL –
shell length), øèðèíà êîëóìåëÿðíîãî îòâîðîòà (ClW – columellar lip width), øèðèíà 1-ãî îáîðîòà
ðàêîâèíû (Wh1W – whorl-1 width), øèðèíà 2-ãî îáîðîòà ðàêîâèíû (Wh2W – whorl-2 width). Òàêèì
æå ñïîñîáîì èçìåðÿëè øèðèíó ïîÿñêà êðûøå÷êè (HBW – horny «belt» width) è øèðèíó åå èçâåñòêî-
âîé ÷àñòè (ÑPW – calcareous plate width). Ïî ýòèì ïàðàìåòðàì ðàññ÷èòûâàëè èíäåêñû ðàêîâèíû è
êðûøå÷êè.
Êðîìå òîãî, àíàëèçèðîâàëàñü è ïîëîâàÿ ñèñòåìà, â ÷àñòíîñòè áûëè çàäåéñòâîâàíû ñëåäóþùèå
ïðîìåðû ìóæñêîé è æåíñêîé ïîëîâûõ ñèñòåì: äëèíà (PrL – prostate length) è øèðèíà (PrW – pro-
state width) ïðîñòàòû, äëèíà (COL – copulatory organ length) è øèðèíà (COW – copulatory organ
width) êîïóëÿòèâíîãî îðãàíà, äëèíà (CBL – copulatory bursa receptacle length) è øèðèíà (CBW –
copulatory bursa receptacle width) ðåçåðâóàðà êîïóëÿòèâíîé ñóìêè, äëèíà (SRL – spermatotheca recep-
tacle length) è øèðèíà (SRW – spermatotheca receptacle width) ðåçåðâóàðà ñåìÿïðèåìíèêà.
Còàòèñòè÷åñêàÿ îáðàáîòêà äàííûõ ïðîâåäåíà ïî ïàêåòó Statistica v. 6.0.
Ðåçóëüòàòû è îáñóæäåíèå
Àëëî çèìíûé àíàëè ç
 ðåçóëüòàòå ïðîâåäåííûõ èññëåäîâàíèé ýâîëþöèîííî-ãåíåòè÷åñêàÿ äèñ-
êðåòíîñòü áûëà ïîäòâåðæäåíà äëÿ òðåõ èç ñåìè âèäîâ íåðèòèä è çàêëþ÷àëàñü îíà
â ôèêñàöèÿõ àëüòåðíàòèâíûõ àëëåëåé ïî ïÿòè èç øåñòè èçó÷åííûõ ëîêóñîâ
(òàáë. 2).
Íåðèòèäà äóíàéñêàÿ (T. danubialis) õàðàêòåðèçîâàëàñü ôèêñàöèÿìè àëüòåðíà-
òèâíûõ ïî îòíîøåíèþ ê íåðèòèäå ïðåñíîâîäíîé (T. fluviatilis) àëëåëåé òðåõ ëîêó-
ñîâ (Aat, Es-2, Es-3), à ïî îòíîøåíèþ ê íåðèòèäå àñòðàõàíñêîé (T. astrachani-
cus) – äâóõ (Mdh, Es-3). Ïðè÷åì ó ïîñëåäíåãî âèäà ëîêóñ Mdh ïîëèìîðôåí, è
ðàñïðåäåëåíèå åãî ãåíîòèïîâ â âûáîðêå ÷åòêî îòâå÷àëî çàêîíó Õàðäè-Âàéíáåðãà
[Mdh125/125 = 17 (12,5); Mdh125/150 = 16 (15); Mdh150/150 = 4 (4,5); (χ2 = 1,74; d.f. = 2;
p > 0,05]. Ãåíåòè÷åñêàÿ äèâåðãåíöèÿ T. fluviatilis è T. astrachanicus íàøëà îòðàæå-
íèå â ôèêñàöèÿõ àëëåëåé òðåõ ëîêóñîâ (Aat, Mdh, Es-3). Ñëåäóåò ñêàçàòü, ÷òî
534 Ñ. Â. Ìåææåðèí, Þ. Â. Òàðàñîâà, Å. È. Æàëàé
Òàáëèö à 2. Aëëîçèìíàÿ èçìåí÷èâîñòü ñåìè ïðåäïîëàãàåìûõ âèäîâ ìîëëþñêîâ ðîäà Theodoxus
Ta b l e 2. Allozyme variation of seven supposed species of genus Theodoxus
* Ðàñøèôðîâêà íîìåðîâ âûáîðîê ïðèâåäåíà â òàáëèöå 1.
Ïðèìå÷ àíèå. Ëîêóñ Es-1 – èíâàðèàíòíûé.
Ë
îê
óñ
À
ëë
åë
ü
Âèä
T. fluviatilis T. danasteri T. sarmaticus T. velox
T
.
eu
xi
nu
s
T
.
as
tr
ac
ha
ni
cu
s
T
.
da
nu
bi
al
is
1* 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Aat 90 1 1
100 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
Es-3 100 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
110 1 1
Es-2 100 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
110 1
Mdh 100 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1
125 0,4
150 0,6
ïîëó÷åííûå ðàçëè÷èÿ íà óðîâíå àëëîçèìîâ â ïðåäåëàõ 33—50% îòíîñÿòñÿ ê ðàç-
ðÿäó ñóùåñòâåííûõ, à ó ïîçâîíî÷íûõ îáû÷íî îòâå÷àþò óðîâíþ ðîäîâûõ
(Ìåææåðèí, Ìîðîçîâ-Ëåîíîâ, 1995, 1996), íî â ëþáîì ñëó÷àå ñâèäåòåëüñòâóåò î
äîñòàòî÷íî äðåâíåé äèâåðãåíöèè ýòèõ âèäîâ. Òîãäà êàê 4 âèäà: T. danasteri, T. sar-
maticus, T. velox è T. euxinus, âûäåëåííûå â ñàìîå ïîñëåäíåå âðåìÿ èç T. fluviati-
lis, íå èìåëè ïî èññëåäîâàííûì àëëîçèìàì íèêàêèõ îòëè÷èé íè äðóã îò äðóãà,
íè îò íîìèíàòèâíîãî âèäà, ÷òî äàåò îñíîâàíèå ñ÷èòàòü ýòè ïÿòü ôîðì êîíñïå-
öèôè÷íûìè.
Ìîðôîëî ãè÷ å ñêèé àí àëè ç
Êà÷ å ñ ò â åííûå ïðè çíàêè. Õîòÿ äëÿ êàæäîãî èç ïðåäïîëàãàåìûõ ñåìè
âèäîâ õàðàêòåðåí ñâîé ðèñóíîê ðàêîâèíû, òåì íå ìåíåå ìîæíî âûäåëèòü äâà
ïðåâàëèðóþùèõ òèïà îêðàñêè ðàêîâèíû (ðèñ. 1). Ïåðâûé – êðàï÷àòûé è ñåêòî-
ðàëüíûé ñâîéñòâåíåí ãðóïïå âèäîâ, òðàäèöèîííî îòíîñèìûõ ê T. fluviatilis s. l.
Ïðè÷åì ó îäíèõ âèäîâ íà òåìíîì ôîíå ðàñïîëàãàþòñÿ ñâåòëûå êðóãëûå èëè
îâàëüíûå ïÿòíà (T. fluviatilis s. str., T. danasteri, T. sarmaticus, T. velox), òîãäà êàê
ó äðóãèõ áûëà ñåòêà èç òåìíûõ, ñëèâàþùèõñÿ ìåæäó ñîáîé ëèíèé íà ñâåòëîì
ôîíå (T. euxinus). Âòîðîé òèï – ýòî ïîïåðå÷íàÿ èñ÷åð÷åííîñòü ðàêîâèíû. Ïðè
ýòîì äëÿ T. astrachanicus õàðàêòåðíû òîíêèå òåìíûå ïîïåðå÷íûå ëèíèè íà ñâåò-
ëîì ôîíå, à äëÿ T. danubialis – íà îòíîñèòåëüíî ñâåòëîì ôîíå øèðîêèå çèãçàãî-
îáðàçíûå òåìíûå ïîïåðå÷íûå ïîëîñû, êîòîðûå òàêæå íå ñëèâàþòñÿ ìåæäó ñîáîé.
Äåëàåò ïîõîæèìè äðóã íà äðóãà äâà ïîñëåäíèõ âèäà è ñâåòëî-ñåðûé öâåò
ïîÿñêà íà êðûøå÷êå, òîãäà êàê ó äðóãèõ ïÿòè (êîòîðûõ ìîæíî ðàññìàòðèâàòü êàê
T. fluviatilis s. l.) îí – îðàíæåâûé.
Êîëè÷å ñ ò â åííûå ïðè çíàêè. Ïðîâåäåííûé äèñêðèìèíàíòûé àíàëèç
ïî ñîâîêóïíîñòè ïðèçíàêîâ ðàêîâèíû êðûøå÷êè è ïîëîâîé ñèñòåìû ñåìè ïðåä-
ïîëàãàåìûõ âèäîâ íåðèòèä ïîêàçàë, ÷òî èç 18 èññëåäîâàííûõ ïðèçíàêîâ äîñòî-
âåðíûé âêëàä â ìåæãðóïïîâóþ èçìåí÷èâîñòü âíîñÿò òîëüêî 10 èç íèõ: âñå ïðî-
ìåðû ðàêîâèíû (ShW), ïðîìåðû êðûøå÷êè, à èç ïðîìåðîâ ïîëîâîé ñèñòåìû
òîëüêî îäèí – PrW (òàáë. 3). Ïðè ýòîì ñàìûì íàäåæíûì ðàçäåëèòåëüíûì ïðè-
çíàêîì ñðåäè ìîëëþñêîâ ýòîãî ðîäà ÿâëÿåòñÿ øèðèíà ïîÿñêà êðûøå÷êè (HBW).
Íàäåæíîé ñòåïåíüþ äèñêðèìèíàöèè õàðàêòåðèçóþòñÿ òîëüêî äâà (T. astrac-
hanicus è T. danubialis) èç ñåìè âèäîâ, óðîâåíü èõ îïðåäåëåíèÿ äîñòèãàåò 100%.
Îñòàëüíûå ïÿòü âèäîâ, òðàäèöèîííî îòíîñèìûå ê T. fluviatilis s. l., èìåëè ñðåä-
íèé óðîâåíü äèñêðèìèíàöèè îêîëî 65%. Ýòî îçíà÷àåò, ÷òî ïî ñîâîêóïíîñòè ïðè-
çíàêîâ ýòè ïÿòü âèäîâ íå äèàãíîñòèðóþòñÿ, ÷òî ñòàíîâèòñÿ îñîáåííî î÷åâèäíûì
ïðè ãðàôè÷åñêîì èçîáðàæåíèè ðåçóëüòàòîâ äèñêðèìèíàíòíîãî àíàëèçà – îáëà-
êà ðàññåèâàíèÿ îñîáåé ýòèõ ïÿòè âèäîâ, ïåðåêðûâàþùèõñÿ ïðàêòè÷åñêè ïîë-
íîñòüþ (ðèñ. 2). Ïðèâåäåíèå ïðåäïîëàãàåìûõ âèäîâ (T. fluviatilis s. str., T. dana-
535Ìîëëþñêè ðîäà Theodoxus (Gastrîpoda, Neritidae) Óêðàèíû…
Òàáëèö à 3. Âêëàä êàæäîãî èç ïðèçíàêîâ (÷àñòíàÿ Ëÿìáäà) è åãî äîñòîâåðíîñòü (ð-óðîâåíü) â
ìåæãðóïïîâóþ èçìåí÷èâîñòü ñåìè ïðåäïîëàãàåìûõ âèäîâ ðîäà Theodoxus
Ta b l e 3. Contributions of the each character (partial Lambda) and its significance (p-level) in intergroup vari-
ation of seven supposed species of Theodoxus
Ïðèìå÷àíèå. Âêëàä ïðèçíàêîâ: ShW, PrL, PrW, COL, COW, CBL, CBW, SRL, SRW áûë íå
äîñòîâåðíûì.
Ïðèçíàê ×àñòíàÿ Ëÿìáäà ð-óðîâåíü
Wh1W 0,93 0,012
Wh2W 0,78 0,000
HBW 0,58 0,000
ÑPW 0,83 0,000
PrW 0,90 0,002
Ïðèçíàê ×àñòíàÿ Ëÿìáäà ð-óðîâåíü
ShH 0,91 0,003
ShL 0,95 0,035
AH 0,87 0,000
AW 0,81 0,000
ClW 0,82 0,000
536 Ñ. Â. Ìåææåðèí, Þ. Â. Òàðàñîâà, Å. È. Æàëàé
Ðèñ. 2. Èçìåí÷èâîñòü ñåìè ïðåäïîëàãàåìûõ âèäîâ ðîäà Theodoxus ïî êîìïëåêñó ïðèçíàêîâ ðàêîâè-
íû, êðûøå÷êè è îðãàíîâ ïîëîâîé ñèñòåìû â ïðîñòðàíñòâå ïåðâûõ äâóõ êàíîíè÷åñêèõ ïåðåìåííûõ
(ROOT_1, ROOT_2).
Fig. 2. Variation of seven supposed species of the genus Theodoxus by complex of shell, plate and reproduc-
tive organs in the space of the two first canonical variables (ROOT_1, ROOT_2).
Òàáëèö à 4. Êëàññèôèêàöèîííàÿ ìàòðèöà, ïîëó÷åííàÿ ïóòåì äèñêðèìèíàíòíîãî àíàëèçà äëÿ ñåìè
ïðåäïîëàãàåìûõ âèäîâ ðîäà Theodoxus ïî ïðîìåðàì ðàêîâèíû, êðûøå÷êè è îðãàíîâ ïîëîâîé ñèñòåìû
Ta b l e 4. Classification matrix obtained by means of discriminant analysis of seven supposed species of genus
Theodoxus by measurements of shell, plate and organs of the reproductive system
Âèä % ¹ 1 2 3 4 5 6 7
T. danubialis 100 1 20 0 0 0 0 0 0
T. fluviatilis 65 2 0 13 3 1 3 0 0
T. danasteri 55 3 0 5 11 2 1 1 0
T. euxinus 75 4 0 0 2 15 2 1 0
T. velox 70 5 0 0 1 4 14 1 0
T. sarmaticus 60 6 0 4 1 0 3 12 0
T. astrachanicus 100 7 0 0 0 0 0 0 20
Âñ å ã î 75 20 22 18 22 23 15 20
Òàáëèö à 5. Íàäåæíîñòü äèñêðèìèíàöèè òðåõ ðåàëüíûõ âèäîâ ðîäà Theodoxus, îöåíåííàÿ ïî äîëå (%)
ïðàâèëüíî îïðåäåëåííûõ îñîáåé, ïî ðàçíûì ãðóïïàì ïðèçíàêîâ
Ta b l e 5. Reliability of discrimination of three real species of genus Theodoxus estimated by the part of cor-
rectly identified specimens
Âèä Âñå ïðèçíàêè Ðàêîâèíà Êðûøå÷êà Ïîëîâàÿ ñèñòåìà
T. danubialis 100 75 75 0
T. fluviatilis s. l. 100 100 89 96
T. astrachanicus 100 100 75 85
Âñ å ã î 100 96,4 85 80,8
537Ìîëëþñêè ðîäà Theodoxus (Gastrîpoda, Neritidae) Óêðàèíû…
Ðèñ. 3. Ñðåäíèå çíà÷åíèÿ è äîâåðèòåëüíûé èíòåðâàë îòíîøåíèÿ øèðèíû èçâåñòêîâîé ÷àñòè êðûøå÷-
êè ê øèðèíå ïîÿñêà êðûøå÷êè (CPW/HBW) ñåìè ïðåäïîëàãàåìûõ âèäîâ ðîäà Theodoxus: 1 –
T. danubialis; 2 – T. fluviatilis; 3 – T. danasteri; 4 – T. sarmaticus; 5 – T. velox; 6 – T. euxinus; 7 –
T. astrachanicus.
Fig. 3. Average values and sample deviations of CPW/HBW index of seven supposed species of genus
Theodoxus: 1 – T. danubialis; 2 – T. fluviatilis; 3 – T. danasteri; 4 – T. sarmaticus; 5 – T. velox; 6 –
T. euxinus; 7 – T. astrachanicus.
1 2 3 4 5 6 7
Ïðåäïîëàãàåìûé âèä
1,5
2,0
2,5
3,0
3,5
4,0
4,5
5,0
5,5
6,0
6,5
C
P
W
/H
B
W
Mean
Ðèñ. 4. Ñðåäíèå çíà÷åíèÿ è äîâåðèòåëüíûé èíòåðâàë îòíîøåíèÿ äëèíû ðàêîâèíû ê åå øèðèíå
(ShL/ShW) ñåìè ïðåäïîëàãàåìûõ âèäîâ ðîäà Theodoxus: 1 – T. fluviatilis; 2 – T. danasteri; 3 – T. sar-
maticus; 4 – T. velox; 5 – T. euxinus; 6 – T. astrachanicus; 7 – T. danubialis.
Fig. 4. Average values and sample deviations of ShL/ShW index of seven supposed species of genus
Theodoxus: 1 – T. fluviatilis; 2 – T. danasteri; 3 – T. sarmaticus; 4 – T. velox; 5 – T. euxinus; 6 – T. astra-
chanicus; 7 – T. danubialis.
Mean
1 2 3 4 5 6 7
Ïðåäïîëàãàåìûé âèä
0,40
0,42
0,44
0,46
0,48
0,50
0,52
0,54
0,56
0,58
0,60
0,62
0,64
0,66
0,68
Sh
L/
Sh
W
538 Ñ. Â. Ìåææåðèí, Þ. Â. Òàðàñîâà, Å. È. Æàëàé
Ðèñ. 5. Ñðåäíèå çíà÷åíèÿ è äîâåðèòåëüíûé èíòåðâàë îòíîøåíèÿ øèðèíû óñòüÿ ê øèðèíå ðàêîâèíû
(AW/ShW) ñåìè ïðåäïîëàãàåìûõ âèäîâ ðîäà Theodoxus: 1 – T. danubialis; 2 – T. fluviatilis; 3 –
T. danasteri; 4 – T. sarmaticus; 5 – T. velox; 6 – T. euxinus; 7 – T. astrachanicus.
Fig. 5. Average values and sample deviations of AW/ShW index of seven supposed species of genus
Theodoxus: 1 – T. danubialis; 2 – T. fluviatilis; 3 – T. danasteri; 4 – T. sarmaticus; 5 – T. velox; 6 – T.
euxinus; 7 – T. astrachanicus.
1 2 3 4 5 6 7
Ïðåäïîëàãàåìûå âèäû
0,22
0,24
0,26
0,28
0,30
0,32
0,34
0,36
0,38
0,40
0,42
0,44
0,46
0,48
A
W
/S
hW
Mean
Ðèñ. 6. Ðàñïðåäåëåíèå çíà÷åíèé èíäåêñà êðûøå÷êè CPW/HBW ñåìè ïðåäïîëàãàåìûõ âèäîâ ðîäà
Theodoxus.
Fig. 6. Distribution of CPW/HBW index values in seven supposed species of the genus Theodoxus.
0
2
4
6
8
10
12
1-1,5 1,5-2 2-2,5 2,5-3 3-3,5 3,5-4 4-4,5 4,5-5 5-5,5 5,5-6 6-6,5 6,5-7 7-7,5 7,5-8 8-8,5
Ê
îë
è
֌
ñò
âî
î
ñî
áå
é
fluviatilis danastrei euxinus velox
sarmaticus danubialis astrachanicus
steri, T. sarmaticus, T. velox, T. euxinus) â îäèí – T. fluviatilis s. l. â ñëó÷àå èñïîëü-
çîâàíèÿ âñåé ñîâîêóïíîñòè ïðèçíàêîâ äàåò 100%-íóþ äèàãíîñòèêó òðåõ âèäîâ
(òàáë. 4). Ïðè ýòîì èñïîëüçîâàíèå êîíêðåòíûõ ãðóïï ïðèçíàêîâ ïðèâîäèò ê
áîëåå îãðàíè÷åííîìó óñïåõó èõ îïðåäåëåíèÿ, êîòîðîå ïî ïðîìåðàì ðàêîâèíû
ñîñòàâëÿåò â ñðåäíåì 96,4%, êðûøå÷êè – 85% è îðãàíîâ ïîëîâîé ñèñòåìû –
81% (òàáë. 5). Ïðè÷åì â ñëó÷àå ïîñëåäíåé ãðóïïû ïðèçíàêîâ îñîáè T. danubialis
íèñêîëüêî íå îòëè÷àþòñÿ îò T. fluviatilis.
Òàêèì îáðàçîì, ïîëó÷åííûå ðåçóëüòàòû îäíîçíà÷íî äîêàçûâàþò, ÷òî íà
óðîâíå ñîâîêóïíîñòè ïðîìåðîâ ðàêîâèíû è ïîëîâîé ñèñòåìû òàê æå ÷åòêî, êàê
è ïî àëëîçèìàì, äèàãíîñòèðóþòñÿ òîëüêî òðè âèäà: T. astrachanicus, T. danubialis
è T. fluviatilis s. l., à ñîîòâåòñòâåííî îñòàëüíûå ÷åòûðå: T. danasteri, T. sarmaticus,
T. velox, T. euxinus äîëæíû ðàññìàòðèâàòüñÿ ïðîñòî êàê ìîðôû ïîñëåäíåãî.
Ïðîâåäåííûé äèñïåðñèîííûé àíàëèç ïîêàçàë, ÷òî âèäîâàÿ ïðèíàäëåæíîñòü
îêàçûâàåò â âûñøåé ñòåïåíè äîñòîâåðíîå âëèÿíèå íà èçìåí÷èâîñòü öåëîãî ðÿäà
èíäåêñîâ ðàêîâèíû, êðûøå÷êè è ïîëîâîé ñèñòåìû (òàáë. 6). Ïðè ýòîì ñàìûì
âûñîêèì óðîâíåì ãåíåðàëèçîâàííîé äèñïåðñèè õàðàêòåðèçóþòñÿ ñëåäóþùèå òðè
èíäåêñà: ïðîïîðöèè êðûøå÷êè ÑPW/HBW è ðàêîâèíû AW/ShW, ShL/ShW.
Çíà÷åíèÿ ïåðâûõ äâóõ ïðèçíàêîâ â âûñøåé ñòåïåíè âûñîêî äîñòîâåðíî îòëè-
÷àþòñÿ â ñðàâíåíèÿõ T. astrachanicus – T. danubialis – T. fluviatilis s. l. ìåæäó
ñîáîé (ðèñ. 3, 5), à ïî òðåòüåìó – ÷åòêî âû÷ëåíÿåòñÿ T. astrachanicus (ðèñ. 4),
òîãäà êàê âèäû â ïðåäåëàõ T. fluviatilis s. l. íå îòëè÷àþòñÿ. Ïðè ýòîì êàæäûé èç
ýòèõ ïðèçíàêîâ ñàì ïî ñåáå íå ìîæåò áûòü èñïîëüçîâàí êàê äèàãíîñòè÷åñêèé èç-
çà çíà÷èòåëüíûõ òðàíñãðåññèé (ðèñ. 6).
539Ìîëëþñêè ðîäà Theodoxus (Gastrîpoda, Neritidae) Óêðàèíû…
Òàáëèö à 6. Ïàðàìåòðû äèñïåðñèîííîãî àíàëèçà, îöåíèâàþùåãî ìåæâèäîâóþ èçìåí÷èâîñòü ïðèçíàêîâ
ïðåäñòàâèòåëåé ðîäà Theodoxus
Ta b l e 6. Parameters of variance analysis which estimate interspecies variation of characters of genus
Theodoxus representatives
Âëèÿíèå äîñòîâåðíî íà óðîâíå: ** – ð < 0,01; *** – ð < 0,001.
1 Èçìåí÷èâîñòü îöåíèâàëàñü â ïðåäåëàõ ãðóïïû èç ñåìè (T. danasteri, T. sarmaticus, T. velox è
T. euxinus, T. fluviatilis, T. danubialis, T. astrachanicus) ïðåäïîëàãàåìûõ âèäîâ.
2 Èçìåí÷èâîñòü îöåíèâàëàñü â ïðåäåëàõ òðåõ (T. fluviatilis, T. danubialis, T. astrachanicus) âèäîâ.
3 Èçìåí÷èâîñòü îöåíèâàëàñü â ïðåäåëàõ T. fluviatilis s. l. (T. danasteri, T. sarmaticus, T. velox è
T. euxinus, T. fluviatilis s. str.).
Ïðèçíàê
Theodoxus (7 âèäîâ)1 Theodoxus (3 âèäà)2 T. fluviatilis s. l. (5 âèäîâ)3
F ν1 ν2 р F ν1 ν2 р F ν1 ν2 ð
ShW, ìì 14,2 6 133 31,9 2 137 4,9 4 95 ***
ShH / ShW 1,4 6 133 *** 2,9 2 137 *** 0,8 4 95
ShL / ShW 29,0 6 133 *** 89,2 2 137 *** 0,1 4 95
AH / ShW 5,4 6 133 *** 14,6 2 137 *** 1,0 4 95
AW / ShW 43,4 6 133 *** 126,3 2 137 *** 1,7 4 95
ClW / ShW 17,9 6 133 *** 55,2 2 137 *** 0,1 4 95
Wh1W / ShW 1,0 6 133 0,5 2 137 2,1 4 95
Wh2W / ShW 12,9 6 133 *** 29,2 2 137 *** 6,0 4 95 ***
PrL / ShW 6,0 6 133 *** 10,4 2 137 *** 4,6 4 95 **
PrW / PrL 12,0 6 133 *** 26,0 2 137 *** 3,5 4 95 **
COL / PrL 5,5 6 133 *** 2,6 2 137 7,4 4 95 ***
COW / PrL 3,4 6 133 ** 0,5 2 137 4,9 4 95 **
CBL / PrL 0,3 6 133 0,6 2 137 0,1 4 95
CBW / PrL 0,6 6 133 0,9 2 137 0,4 4 95
SRL / PrL 5,8 6 133 *** 17,7 2 137 *** 0,1 4 95
SRW / PrL 6,8 6 133 *** 20,4 2 137 *** 0,3 4 95
ÑPW / HBW 64,8 6 133 *** 196,2 2 137 *** 1,1 4 95
Çàêëþ÷åíèå
Òàêèì îáðàçîì, ïðîâåäåííîå èññëåäîâàíèå íåçàâèñèìûõ ñèñòåì ïðèçíàêîâ,
àëëîçèìîâ è ìîðôîëîãèè ðàêîâèíû, äàëè îäèíàêîâûå ðåçóëüòàòû, êîòîðûå ìîãóò
áûòü ñôîðìóëèðîâàíû â âèäå äâóõ îñíîâíûõ âûâîäîâ.
1. Èç ñåìè ïðåäïîëàãàåìûõ äëÿ àêâàòîðèé Óêðàèíû âèäîâ ðåàëüíîé ãåíåòè-
÷åñêîé è ìîðôîëîãè÷åñêîé äèñêðåòíîñòüþ îáëàäàþò òîëüêî òðè: ëóíêà ðå÷íàÿ
(T. fluviatilis), ëóíêà äóíàéñêàÿ (T. danubialis) è ëóíêà àñòðàõàíñêàÿ (T. astrachani-
cus).
2. ×åòûðå ïðåäïîëàãàåìûå îòäåëüíûìè èññëåäîâàòåëÿìè «âèäà» (T. danaste-
ri, T. sarmaticus, T. velox è T. euxinus) íå èìåþò íè àëëîçèìíûõ ìàðêåðîâ, íè ìîð-
ôîëîãè÷åñêèõ ïðèçíàêîâ çà èñêëþ÷åíèåì îêðàñêè ðàêîâèíû, êîòîðûå áû
îòëè÷àëè èõ äðóã îò äðóãà è âñåõ èõ îò T. fluviatilis, à ïîòîìó ïðîñòî ÿâëÿþòñÿ
ìîðôàìè è äîëæíû ðàññìàòðèâàòüñÿ òîëüêî êàê ñèíîíèìû ïîñëåäíåãî âèäà.
Àíèñòðàòåíêî Î. Þ., Ñòàðîáàãàòîâ ß. È., Àíèñòðàòåíêî Â. Â. Ìîëëþñêè ðîäà Theodoxus
(Gastropoda, Pectinibranchia, Neritidae) Àçîâî-×åðíîìîðñêîãî áàññåéíà // Âåñòí. çîîëîãèè. –
1999. – 33, ¹ 3. – Ñ. 11—19.
Àíèñòðàòåíêî Â. Â. Àíèñòðàòåíêî Î. Þ. Êëàññ Ïàíöèðíûå èëè Õèòîíû, êëàññ Áðþõîíîãèå –
Cyclobranchia, Scutibranchia è Pectinibranchia (÷àñòü). – 2001. – 240 c. – (Ôàóíà Óêðàèíû.
Ìîëëþñêè ; Ò. 29, âûï. 1, êí. 1).
Æàäèí Â. È. Ïðåñíîâîäíûå ìîëëþñêè ÑÑÑÐ. – Ë., 1933. – 232 ñ.
Æàäèí Â. È. Ìîëëþñêè ïðåñíûõ è ñîëîíîâàòûõ âîä ÑÑÑÐ. – Ì. ; Ë. : Èçä-âî ÀÍ ÑÑÑÐ, 1952. –
376 ñ.
Æàëàé Å. È., Ìåææåðèí Ñ. Â., Øóáðàò Þ. Â., Ãàðáàð À. Â. Î âèäîâîì ñîñòàâå ìîëëþñêîâ ðîäà
Theodoxus (Gastropoda, Neritidae) áàññåéíà Íèæíåãî Äóíàÿ: ðåøåíèå ïðîáëåìû ïóòåì àíàëèçà
àëëîçèìîâ // Íàóê. â³ñíèê Óæãîðîä. óí-òó. Ñåð. Á³îë. – 2008. – Âèï. 23. – Ñ. 205—208.
Ìåææåðèí Ñ. Â., Ìîðîçîâ-Ëåîíîâ Ñ. Þ. Ãåíåòè÷åñêàÿ äèôôåðåíöèàöèÿ òàêñîíîâ ìëåêîïèòàþùèõ:
îöåíêà ïî áèîõèìè÷åñêèì ãåííûì ìàðêåðàì // Æóðí. îáù. áèîë. – 1995. – 56, ¹ 1. –
Ñ.71—96.
Ìåææåðèí Ñ.Â., Ìîðîçîâ-Ëåîíîâ Ñ.Þ. Ðàçíîìàñøòàáíîñòü ãåííîé äèôôåðåíöèàöèè òàêñîíîâ âèäî-
âîãî è ðîäîâîãî óðîâíåé è áèîëîãè÷åñêàÿ ðàçíîêà÷åñòâåííîñòü îòðÿäîâ ìëåêîïèòàþùèõ //
Æóðí. îáù. áèîë. – 1996. – 57, ¹ 2. – Ñ. 79—94.
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| id | nasplib_isofts_kiev_ua-123456789-65840 |
| institution | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| issn | 0084-5604 |
| language | Russian |
| last_indexed | 2025-12-07T15:46:00Z |
| publishDate | 2011 |
| publisher | Інститут зоології ім. І.І. Шмальгаузена НАН України |
| record_format | dspace |
| spelling | Межжерин, С.В. Тарасова, Ю.В. Жалай Е.И. 2014-07-03T20:07:56Z 2014-07-03T20:07:56Z 2011 Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ / С.В. Межжерин, Ю.В. Тарасова, Е.И. Жалай // Вестник зоологии. — 2011. — Т. 45, № 6. — С. 531–540. — Бібліогр.: 15 назв. — рос. 0084-5604 https://nasplib.isofts.kiev.ua/handle/123456789/65840 594:575.8(477) Аллозимный анализ неритид рода Theodoxus Montfort, 1810 подтвердил эволюционно-генетическую дискретность трех видов: T. astrachanicus, T. fluviatilis, T. danubialis, тогда как T. danasteri, T. sarmaticus, T. velox и T. euxinus по данным генного маркирования оказались конспецифичными T. fluviatilis. К такому же выводу приводят и результаты морфометрического анализа раковины и половой системы. Allozyme analysis of neritid mollusks of the genus Theodoxus Montfort, 1810 shows evolutionary and genetical discreteness of the three species T. astrachanicus, T. fluviatilis, T. danubialis, whereas T. danasteri, T. sarmaticus, T. velox and T. euxinus species were found to be conspecific with T. fluviatilis according to the results of gene labeling. The results of morphometric analysis of the shell and genital system lead support this conclusion. ru Інститут зоології ім. І.І. Шмальгаузена НАН України Вестник зоологии Морфология Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ Mollusks of the Genus Theodoxus (Gastrоpoda, Neritidae) of Ukraine: the Morphological Analysis and Analysis of Allosymes Article published earlier |
| spellingShingle | Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ Межжерин, С.В. Тарасова, Ю.В. Жалай Е.И. Морфология |
| title | Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ |
| title_alt | Mollusks of the Genus Theodoxus (Gastrоpoda, Neritidae) of Ukraine: the Morphological Analysis and Analysis of Allosymes |
| title_full | Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ |
| title_fullStr | Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ |
| title_full_unstemmed | Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ |
| title_short | Моллюски рода Theodoxus (Gastrоpoda, Neritidae) Украины: морфологический и аллозимный анализ |
| title_sort | моллюски рода theodoxus (gastrоpoda, neritidae) украины: морфологический и аллозимный анализ |
| topic | Морфология |
| topic_facet | Морфология |
| url | https://nasplib.isofts.kiev.ua/handle/123456789/65840 |
| work_keys_str_mv | AT mežžerinsv mollûskirodatheodoxusgastropodaneritidaeukrainymorfologičeskiiiallozimnyianaliz AT tarasovaûv mollûskirodatheodoxusgastropodaneritidaeukrainymorfologičeskiiiallozimnyianaliz AT žalaiei mollûskirodatheodoxusgastropodaneritidaeukrainymorfologičeskiiiallozimnyianaliz AT mežžerinsv mollusksofthegenustheodoxusgastropodaneritidaeofukrainethemorphologicalanalysisandanalysisofallosymes AT tarasovaûv mollusksofthegenustheodoxusgastropodaneritidaeofukrainethemorphologicalanalysisandanalysisofallosymes AT žalaiei mollusksofthegenustheodoxusgastropodaneritidaeofukrainethemorphologicalanalysisandanalysisofallosymes |