Turczaninowia 2000 3(4) : 28–36
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ÓÄÊ 581.9
À.Á. Äîóýëüä
A. Doweld
ÂÎÑÑÒÀÍÎÂËÅÍÈÅ ÐÎÄÀ THOA Aublet (GNETACEAE)
REHABILITATION OF THE GENUS THOA Aublet (GNETACEAE)
Ñîâðåìåííûå èññëåäîâàíèÿ àíàòîìèè è ìîðôîëîãèè ðåïðîäóêòèâíûõ
îðãàíîâ îáîëî÷êîñåìåííûõ è, â ÷àñòíîñòè, ñåìåéñòâà ãíåòîâûõ, ïðåäñòàâëåííîãî
â ñîâðåìåííûõ ðóêîâîäñòâàõ îäíèì ðîäîì Gnetum L., âûÿâèëè íåîæèäàííóþ
ãåòåðîãåííîñòü èõ ñòðîåíèÿ ó ðàçíûõ âèäîâ ýòîãî ðîäà (Kapil, Rodin, 1969; Gillespie,
Nowicke, 1994). Ìýááåðëè (Mabberley, 1993) ïðèâåë äëÿ ñåìåéñòâà åùå îäèí ðîä
Vinkiella R. Johns nom. nud., êîòîðûé äî ñèõ ïîð íå îáíàðîäîâàí ñîãëàñíî òðåáîâàíèÿì ìåæäóíàðîäíîãî êîäåêñà áîòàíè÷åñêîé íîìåíêëàòóðû. Ïî-âèäèìîìó, ðå÷ü
øëà î äîâîëüíî ñïåöèàëèçèðîâàííûõ ìàëàçèéñêèõ ôîðìàõ ãíåòóìîâ, ó êîòîðûõ
ñåìåíà èìåþò äîâîëüíî íåîáû÷íûé âèä, ÷òî ïîçâîëèëî ïðåäïîëîæèòü î ñóùåñòâîâàíèè ðîäà, îòëè÷íîãî îò ðîäà Gnetum. Îäíàêî îòñóòñòâèå äåòàëüíûõ èññëåäîâàíèé âñåõ âèäîâ ýòîãî øèðîêî ðàñïðîñòðàíåííîãî ïàíòðîïè÷åñêîãî ðîäà íå ïîçâîëÿëî ïðîâåñòè ãðàíèöó ìåæäó ðàçíûìè ãðóïïàìè âèäîâ, ïðèóðî÷åííûõ ê ðàçíûì
ðåãèîíàì, – àôðèêàíñêîìó (G. africanum), þæíîàìåðèêàíñêîìó (G. nodiflora –
G. urens-ãðóïïà) è àçèàòñêîìó (G. gnemon-ãðóïïà, G. indicum–G. gnemonoidesãðóïïà).
Íà îñíîâàíèè ãåîãðàôè÷åñêîãî ðàñïðîñòðàíåíèÿ ðîäà Ìàðêãðàô (Markgraf,
1930) ïîñòðîèë îðèãèíàëüíóþ ñèñòåìó, âûäåëèâ äâå ñåêöèè (sect. Gnetum, sect.
Cylindrostachys) è ïîäðàçäåëèâ êàæäóþ èç íèõ åùå íà äâå äîïîëíèòåëüíûå
ïîäñåêöèè. Îäíàêî ýòà ñèñòåìà íå ïîëó÷èëà ñâîåãî ïîäòâåðæäåíèÿ íè â àíàòîìèè
ñåìÿí (Rodin, Kapil, 1969), íè â ïàëèíîëîãèè (Gillespie, Nowicke, 1994). Íàïðîòèâ,
áûëî âûÿâëåíî íåîæèäàííîå ñõîäñòâî ðàçíûõ äîâîëüíî óäàëåííûõ äðóã îò äðóãà
ãðóïï âèäîâ. Ñèñòåìà Ìàðêãðàôà òðåáóåò êîðåííîãî ïåðåñìîòðà, êîòîðûé ïðåäïîëàãàåòñÿ ïðåäïðèíÿòü â áëèæàéøåå âðåìÿ ñ ïîëó÷åíèåì äîïîëíèòåëüíûõ
ìàòåðèàëîâ èç ðåãèîíàëüíûõ ãåðáàðèåâ.
Ðîäèí è Êàïèë (Rodin, Kapil, 1969), èññëåäîâàâ àíàòîìèþ è ìîðôîëîãèþ
ñåìÿí ãíåòóìîâ, íåîæèäàííî âûÿâèëè ãåòåðîãåííîñòü ñòðîåíèÿ ñåìÿí.  ÷àñòíîñòè? ýòèì èññëåäîâàòåëÿì óäàëîñü óñòàíîâèòü, ÷òî òðîéíîé ïîêðîâ ñåìÿí
ñîõðàíÿåòñÿ òîëüêî ó åäèíñòâåííî äðåâåñíîãî âèäà G. gnemon, â òî âðåìÿ êàê ó
îñòàëüíûõ âèäîâ ðîäà, ïðåäñòàâëÿþùèõ ñîáîé ëèàíû, íàðóæíûå äâà ñåìåííûõ
ïîêðîâà ñðàñòàþòñÿ ìåæäó ñîáîé â îäíó ñëîæíóþ ñòðóêòóðó ñî ñêëåðèôèêàöèåé
âíóòðåííåé ÷àñòè ýòîãî íîâîîáðàçîâàíèÿ. Ìîðôîëîãè è ñèñòåìàòèêè â ïðîøëîì
âåêå ïîëàãàëè, ÷òî òðîéíîé ïîêðîâ ãíåòóìîâ âîçíèêàåò ïóò¸ì ðàñùåïëåíèÿ
åäèíñòâåííîãî èíòåãóìåíòà, ïðèñóùåãî äðóãèì ãîëîñåìÿííûì (Strasburger, 1872),
ëèáî ïðåäñòàâëÿåò ñîáîé ñëîæíóþ ñòðóêòóðó, ñîñòîÿùóþ èç äâóõ íàðóæíûõ
ýëåìåíòîâ ïðèöâåòíèêà (perianthium) è ñàìîãî âíóòðåííåãî, ñ÷èòàâøåãîñÿ
Äîóýëüä À.Á.: Âîññòàíîâëåíèå ðîäà Thoa Aublet (Gnetaceae)
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çàâÿçüþ (Beccari, 1877; Lignier, Tison, 1912) èëè âíóòðåííèì èíòåãóìåíòîì (Griffith,
1859; Benson, 1904a; Coulter, Chamberlain, 1917). Ïðèìå÷àòåëüíî, ÷òî íåêîòîðûå
àâòîðû (Blume, 1848; Eichler, 1875; Strasburger, 1872; Benson, 1904a) ïîëàãàëè,
÷òî íàðóæíûå ýëåìåíòû ïîêðîâíîé ñèñòåìû ãíåòóìîâ – ýòî âèäîèçìåíåííàÿ
çàâÿçü, ò.å. ãîìîëîã ïëîäîëèñòèêà. Ñîâðåìåííûå èññëåäîâàíèÿ ñòðîåíèÿ
ðåïðîäóêòèâíîãî ïîáåãà ãíåòóìîâ (Markgraf, 1971; Takaso, Bouman, 1986)
ïîäòâåðäèëè ñïðàâåäëèâîñòü îòíåñåíèÿ ñàìîãî âíóòðåííåãî ïîêðîâà ê
èíòåãóìåíòàëüíîé ñèñòåìå, ãîìîëîãè÷íîé èíòåãóìåíòó äðóãèõ ñåìåííûõ
ðàñòåíèé êàê ïðîèçâîäíîìó ñòåðèëèçîâàâøåéñÿ ÷àñòè ñïîðàíãèàòíîé ñèñòåìû
ðàñòåíèÿ (Benson, 1904b; Doweld, 1998). Äâà ïîêðîâà, ðàñïîëàãàþùèõñÿ êíàðóæè
îò ýòîãî èíòåãóìåíòà, ïðåäëîæåíî ñ÷èòàòü ôîëèàðíûìè ñòðóêòóðàìè, ò. ê. â
îòëè÷èå îò ïðèñóùåãî èíòåãóìåíòàì áàçèïåòàëüíîãî çàëîæåíèÿ, ýòè ñòðóêòóðû
ðàçâèâàþòñÿ íà ïîáåãå àêðîïåòàëüíî, êàê è âñå ôèëëîìû ïîáåãà âûñøåãî
ðàñòåíèÿ.  îòëè÷èå îò Ìàðòýíà (Martens, 1971), ïðîäîëæàþùåãî íàçûâàòü ýòè
ýëåìåíòû ïðèöâåòíèêàìè (perianthium), a ñàì ðåïðîäóêòèâíûé ïîáåã – öâåòêîì
è åãî ñîáðàíèå – ñîöâåòèåì, Òàêàñî è Áîóìàí (Takaso, Bouman, 1986) îñòîðîæíî
íàçûâàþò ýòè äâà ýëåìåíòà ñåìåííîé êîæóðû ïðîñòî «ïîêðîâàìè» (envelopes), à
ïîáåã – ñòðîáèëîì. Ê ñîæàëåíèþ, äëÿ áîëüøèíñòâà ñîâðåìåííûõ ãîëîñåìÿííûõ
ðàñòåíèé, èìåþùèõ ïðèíöèïèàëüíî îòëè÷íîå ñòðîåíèå ðåïðîäóêòèâíûõ îðãàíîâ
îò òàêîâûõ ñîâðåìåííûõ öâåòêîâûõ, äî ñèõ ïîð íå ñîçäàíî îðèãèíàëüíîé
òåðìèíîëîãèè, êîòîðàÿ áû ó÷èòûâàëà ãîìîëîãèè è àíàëîãèè ñòðîåíèÿ îðãàíîâ.
Èñïîëüçîâàíèå àíãèîñïåðìîèäíîé èëè ïòåðèäîôèòîèäíîé òåðìèíîëîãèé èëè,
ñêîðåå, èõ ñóððîãàòà (Meyen, 1984) äëÿ îïèñàíèÿ ñòðîåíèÿ ãîëîñåìÿííûõ òîðìîçèò
ðàçâèòèå íàøèõ çíàíèé îá ýâîëþöèè è ôèëîãåíåòè÷åñêèõ ñâÿçÿõ ýòèõ òàêñîíîâ,
÷àñòî ïðèâîäÿ ê îøèáî÷íûì îáîáùåíèÿì è èíòåðïðåòàöèÿì. Òàê, â ñîâðåìåííûõ
ðàáîòàõ ïî àíàòîìèè ñåìÿí ãíåòóìîâ ïðèõîäèòñÿ ñòàëêèâàòüñÿ ñ óïîòðåáëåíèåì
òåðìèíà «òåñòà» ïðèìåíèòåëüíî ê îáðàçîâàíèÿì ôîëèàðíîé ïðèðîäû (à íå
ñïîðàíãèàòíîé, êàê ó âñåõ äðóãèõ ñåìåííûõ è ïîêðûòîñåìÿííûõ â ÷àñòíîñòè).
Êàê ñëåäñòâèå, ñòðîåíèå ýòîé ñëîæíîé ñòðóêòóðû áëàãîäàðÿ ñõîäíîìó òåðìèíó
ñîâåðøåííî íåîáîñíîâàííî íàïðÿìóþ ñðàâíèâàåòñÿ ñ òàêîâûì ïîêðûòîñåìÿííûõ,
âûìåðøèõ òðèãîíîêàðïîâûõ (ìåäóëëîçîâûõ) è áåííåòòèòîâûõ. Ñòðîãî ãîâîðÿ, â
îòñóòñòâèå äîêóìåíòèðîâàííûõ è àðãóìåíòèðîâàííûõ äîêàçàòåëüñòâ ãîìîëîãèè
ýòîé «òåñòû» ãíåòóìîâ ñ òàêîâîé ïîêðûòîñåìÿííûõ (äëÿ êîòîðûõ ýòîò òåðìèí
áûë è ñîçäàí), à òàêæå äðóãèõ ãðóïï ñîâðåìåííûõ è èñêîïàåìûõ ñåìåííûõ, îò
óïîòðåáëåíèÿ ýòîãî ñòðîãî ìîðôîëîãè÷åñêîãî òåðìèíà â ïðèìåíåíèè ê äàííûì
ñòðóêòóðàì ñëåäóåò âîçäåðæàòüñÿ.
 îïèñàòåëüíóþ ìîðôîëîãèþ ðåïðîäóêòèâíîãî ïîáåãà ãíåòóìîâ ìû ïðåäëàãàåì ââåñòè ðÿä òåðìèíîâ, ñòðîãî îñíîâûâàÿñü íà ïîëó÷åííûõ ðàíåå äàííûõ î
ñòðîåíèè è ãîìîëîãèÿõ ñîñòàâëÿþùèõ åãî ñòðóêòóðíûõ ýëåìåíòîâ (Markgraf,
1971; Takaso, Bouman, 1986).  ÷àñòíîñòè, ñàìûé âíóòðåííèé ïîêðîâ ñåìåíè
ãíåòîâûõ ìû âñëåä çà áîëüøèíñòâîì ñîâðåìåííûõ èññëåäîâàòåëåé ðàññìàòðèâàåì êàê èíòåãóìåíò. Ìû ðàññìàòðèâàåì åãî êàê ãîìîëîã íàðóæíîãî èíòåãóìåíòà
ñåìåííûõ (â òîì ÷èñëå è ïîêðûòîñåìÿííûõ), ïðèíèìàÿ âî âíèìàíèå îáíàðóæåí-
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Turczaninowia 2000 3(4) : 28–36
íûå àíãëèéñêîé èññëåäîâàòåëüíèöåé Ìýðè Òîäåé (Thoday, 1921) ðóäèìåíòû
çàëîæåíèÿ êíóòðè îò äàííîãî èíòåãóìåíòà åùå äîïîëíèòåëüíîãî ïîêðîâà (÷èòàé:
âíóòðåííåãî èíòåãóìåíòà). Ýòî âïîëíå ñîãëàñóåòñÿ ñ ñîâðåìåííîé òî÷êîé çðåíèÿ
î ïðèìèòèâíîñòè áèòåãìè÷åñêîãî òèïà ñåìÿçà÷àòêà ñåìåííûõ ðàñòåíèé (Smith,
1964; Doweld, 1998) è ñïåöèàëèçèðîâàííîñòè îäíîïîêðîâíîãî, êîòîðûé è îáíàðóæåí
ó ãíåòóìîâ. Òàêèì îáðàçîì, ìû îïðåäåëÿåì ýòîò åäèíñòâåííûé èíòåãóìåíò êàê
ñîõðàíèâøèéñÿ íàðóæíûé èñõîäíîãî áèòåãìè÷åñêîãî ñåìÿçà÷àòêà ïðà-ãíåòîâûõ,
â êîòîðîì âíóòðåííèé èíòåãóìåíò, ïî-âèäèìîìó, ïîäâåðãñÿ àáîðòàöèè. Â ýòîé
ñâÿçè òåðìèí “òåñòà” íàìè îñòàâëåí òîëüêî çà ýòèì ñòðóêòóðíûì ýëåìåíòîì
ñòðîáèëà ãíåòóìîâ.
Ó÷èòûâàÿ ñ îïðåäåëåííîñòüþ óñòàíîâëåííóþ ãîìîëîãèþ äâóõ íàðóæíûõ
ïîêðîâîâ ñåìÿçà÷àòêà ãíåòîâûõ äåêóññàòíûì ïàðíûì ëèñòîâûì îáðàçîâàíèÿì
(Takaso, Bouman, 1986), ìû ïðåäëàãàåì êàæäóþ èç ýòèõ ôîëèàðíûõ ñèñòåì
íàçûâàòü ñïåöèàëüíûì òåðìèíîì – òåãóëîé (tegulà, lat., a tegule, Engl.), –
îïðåäåëÿÿ åå êàê äâóñîñòàâíîé (äèìåðíûé) èíòåãóìåíòîïîäîáíûé ôîëèàðíûé
ïîêðîâ, âîçíèêàþùèé â ïðîöåññå ýâîëþöèè îò äâóõ áðàêòåîçíûõ êðîþùèõ
îáðàçîâàíèé, ñðàñòàþùèõñÿ â åäèíóþ êîëüöåîáðàçíóþ ñòðóêòóðó âîêðóã ñåìåíè.
Äâå ñâîáîäíûå (íå ñðàñòàþùèåñÿ) òåãóëû îòìå÷àþòñÿ òîëüêî ó îäíîãî âèäà
ãíåòóìîâ – G. gnemon, – â òî âðåìÿ êàê ó âñåõ äðóãèõ âèäîâ (Thoday, 1921;
Rodin, Kapil, 1969) ýòè òåãóëû ñðàñòàþòñÿ, îáðàçóÿ åäèíûé ñïåöèàëèçèðîâàííûé
ïîêðîâ ñåìåíè ñ î÷åíü ñâîåîáðàçíûì àíàòîìè÷åñêèì ñòðîåíèåì (ñïåöèàëèçèðîâàííîé ñêëåðèôèêàöèåé âíóòðåííåé ÷àñòè, êîòîðàÿ, ïî íàøèì äàííûì, íå
îáíàðóæåíà áîëüøå íè ó êàêèõ äðóãèõ ãðóïï ñåìåííûõ). Òàêîé òèï ìû ïðåäëàãàåì
íàçâàòü ñèíòåãóëîé (syntegulà, lat., a syntegule, Engl.), ÷òîáû â îïèñàíèÿõ
àíàòîìè÷åñêîãî ñòðîåíèÿ ñåìÿí ãíåòóìîâ ñòðîãî ïðèäåðæèâàòüñÿ ðàçãðàíè÷åíèÿ
ãîìîëîãè÷íûõ è àíàëîãè÷íûõ ñòðóêòóð. Ñ òî÷êè çðåíèÿ óñòàíîâëåííûõ ãîìîëîãèé
ñîñòàâëÿþùèõ ðåïðîäóêòèâíîãî ïîáåãà ãíåòîâûõ àáñîëþòíî ëèøåíà îáîñíîâàííîñòè ïîïûòêà Òàêàñî è Áîóìàíà (ibid.) ãîìîëîãèçèðîâàòü ïàõèõàëàçó ïîêðûòîñåìÿííûõ ñî ñõîäíûì (ìîðôîãåíåòè÷åñêè) ñòàíîâëåíèåì ñèíòåãóëû ãíåòîâûõ è
òàêèì îáðàçîì ââåñòè íè÷åì íå îáîñíîâàííûé òåðìèí «ýíäîõàëàçà» ê ýòîìó
ñâîåîáðàçíîìó îáðàçîâàíèþ.
Ïðèìåíèòåëüíî ê îïèñàíèþ âñåãî ðåïðîäóêòèâíîãî ïîáåãà ãíåòîâûõ,
ñîñòîÿùåãî èç íóöåëëóñà, ïîêðûòîãî åäèíñòâåííûì (íàðóæíûì) èíòåãóìåíòîì,
è äâóõ ïàð ñâîáîäíûõ èëè ñðàñòàþùèõñÿ òåãóë, ìû íàñòîÿòåëüíî ðåêîìåíäóåì
îòêàçàòüñÿ îò óïîòðåáëåíèÿ òåðìèíà «öâåòîê», îãðàíè÷èâàÿ ïðèìåíåíèå ïîñëåäíåãî òîëüêî äëÿ ïîêðûòîñåìÿííûõ. Ïîëàãàÿ ñïðàâåäëèâûì ìíåíèå î âîçíèêøåé
àáñòðàêòíîñòè òåðìèíà «ñòðîáèë» â ñîâðåìåííîé ìîðôîëîãèè âûñøèõ ðàñòåíèé,
êîòîðûé ïðèìåíÿåòñÿ è â îïèñàíèÿõ ëþáûõ ñîáðàíèé ðåïðîäóêòèâíûõ ñòðóêòóð
îò õâîùåé è ïàïîðîòíèêîâ äî ñîâðåìåííûõ ãîëîñåìåííûõ è äàæå ïîêðûòîñåìÿííûõ, ìû ââîäèì ñîáñòâåííûé òåðìèí äëÿ «ñåìÿí» ãíåòîâûõ, êîòîðûå â
ýâîëþöèîííîì ïëàíå ïðåäñòàâëÿþò ñîáîé ìîðôîëîãè÷åñêîå íîâîîáðàçîâàíèå
òàêîãî æå ðàíãà, êàê ïëîä ïîêðûòîñåìåííûõ è øèøêà õâîéíûõ, íî êîòîðîå áûëî
íåçàñëóæåííî îáîéäåíî â îðãàíîãðàôèè ñåìåííûõ ðàñòåíèé èç-çà ìàëîãî
Äîóýëüä À.Á.: Âîññòàíîâëåíèå ðîäà Thoa Aublet (Gnetaceae)
31
êîëè÷åñòâà ïðåäñòàâèòåëåé ãíåòîâûõ (âñåãî îäèí ðîä) è èõ âíåøíåãî ñòðóêòóðíîãî
îäíîîáðàçèÿ. Íàìè ïðåäëàãàåòñÿ òåðìèí glandidium (îò ëàò. glandioideus –
æåëóäåîáðàçíûé) (ðóññê. – ãëàíäèäèé). Ñ÷èòàåì íåïðèåìëåìûì èñïîëüçîâàòü
òåðìèí «êîñòÿíêà» (Parlatore, 1868) ïðèìåíèòåëüíî ê ãëàíäèäèÿì ãíåòîâûõ,
ïîñêîëüêó ðàñïðîñòðàíåíèå êàðïîëîãè÷åñêîãî òåðìèíà íà ãîëîñåìÿííûå ðàñòåíèÿ
âíåñåò òîëüêî ïóòàíèöó è íåðàçáåðèõó. Ìû ñ÷èòàåì ñîâåðøåííî íåîïðàâäàííûì
ñ òî÷êè çðåíèÿ ìîðôîëîãèè âñåãî ðåïðîäóêòèâíîãî ïîáåãà ãíåòîâûõ èñïîëüçîâàòü
òåðìèí «øèøêîÿãîäà» (arcesthidium) (Spjut, 1994), êîòîðûé îáû÷íî ïðèìåíÿåòñÿ
äëÿ îïèñàíèÿ ñâîåîáðàçíîãî ðåïðîäóêòèâíîãî ïîáåãà ñåì. Cupressaceae, ïîñêîëüêó åãî èñïîëüçîâàíèå ìîæåò âûçâàòü íåîáîñíîâàííûå àññîöèàöèè ñ îáùèì
ïðîèñõîæäåíèåì è ñòðóêòóðíûì ñõîäñòâîì äàííûõ ðåïðîäóêòèâíûõ ñèñòåì.
Ó÷èòûâàÿ, ÷òî â ãëàíäèäèè ãíåòîâûõ ïðîèñõîäèò ñòàíîâëåíèå íîâîãî òèïà
ïîêðîâíûõ ñòðóêòóð (ñèíòåãóëû), ìû ïðåäëàãàåì êëàññèôèöèðîâàòü ãëàíäèäèè
ïî àíàëîãèè ñ ïëîäàìè ïîêðûòîñåìÿííûõ íà òàê íàçûâàåìûå âåðõíèå (superior),
èëè ãëàíäèäèè ñ äâóìÿ ïàðàìè òåãóë (òîëüêî G. gnemon), è íèæíèå ãëàíäèäèè
(inferior glandidia), â êîòîðûõ ïðîèñõîäèò ñðàñòàíèå òåãóë â ñèíòåãóëó, ÷òî â
ïðèíöèïå íàïîìèíàåò ñòàíîâëåíèå íèæíèõ ïëîäîâ àïïåíäèêóëÿðíîãî òèïà, ó
êîòîðûõ ïðîèñõîäèò ñðàñòàíèå ïëîäîëèñòèêîâ ñ íàðóæíûìè ôèëëîìíûìè
ýëåìåíòàìè (Douglas, 1944, 1957). Ïðåäïðèíÿòîå íàìè ðàçãðàíè÷åíèå òèïîâ
ãëàíäèäèåâ ïîìîæåò ïðàâèëüíî (ñ òî÷êè çðåíèÿ àíàëîãèé è ãîìîëîãèé) ñðàâíèâàòü
ìíîãî÷èñëåííûå èñêîïàåìûå îñòàòêè, ðÿä êîòîðûõ ñåãîäíÿ îòíîñÿò ê ãíåòîâûì
(Êðàñèëîâ, 1989; Doweld, 1996). Ñîáðàíèÿ ãëàíäèäèåâ âìåñòî íåïðèåìëåìîãî
òåðìèíà «ñîöâåòèå» ñëåäóåò íàçûâàòü inglandescentia (lat., ïî àíàëîãèè ñ
ñîöâåòèåì – inflorescentia, lat.; inglandescence, Engl. ñîãëàíäåòèå, ðóññê., ïî
àíàëîãèè ñ ñîöâåòèåì).
Ââåäåíèå íàìè íîâûõ òåðìèíîâ è ìîðôîëîãè÷åñêèõ ïîíÿòèé äëÿ îïèñàíèÿ
âûÿâëåííîãî ðàçíîîáðàçèÿ ñòðîåíèÿ ðåïðîäóêòèâíûõ îðãàíîâ ãíåòîâûõ íàõîäèò
ñâîå îòðàæåíèå è â ñèñòåìàòèêå ýòîé äðåâíåé ãðóïïû. Àíàòîìèÿ ñåìÿí è ïàëèíîëîãè÷åñêèå èññëåäîâàíèÿ Äæèëëåñïè è Íîâèêè (Gillespie, Nowicke, 1994) óêàçûâàþò íà àðõàè÷íîñòü åäèíñòâåííîãî äðåâåñíîãî ïðåäñòàâèòåëÿ ðîäà – âèäà
G. gnemon – è íà åãî òàêñîíîìè÷åñêóþ èçîëèðîâàííîñòü â ðîäå, òàê êàê âèä
îòëè÷àåòñÿ îò âñåõ äðóãèõ âèäîâ ãíåòóìîâ ïðèìèòèâíûìè âåðõíèìè ãëàíäèäèÿìè,
äðåâåñíûì (à íå ëèàíîâèäíûì) ãàáèòóñîì è ñòðîåíèåì ìèêðîñïîð. Ïî íàøåìó
ìíåíèþ, ñëåäóåò ïðèäàòü ðàíã îòäåëüíûõ ðîäîâ ýòèì äâóì ãðóïïàì âèäîâ – ñ
âåðõíèìè è íèæíèìè ãëàíäèäèÿìè, – ÷òîáû òàêèì îáðàçîì ïîä÷åðêíóòü
ñóùåñòâîâàíèå äâóõ ðàçíûõ íûíå ñóùåñòâóþùèõ òèïîâ ãíåòîèäíûõ ôðóêòèôèêàöèé. Íàèáîëåå ñòàðøèì ðîäîâûì íàçâàíèåì äëÿ ãðóïïû âèäîâ ñ íèæíèìè
ãëàíäèäèÿìè ÿâëÿåòñÿ ïîçàáûòîå òîëüêî â ñåðåäèíå XIX ñòîëåòèÿ ðîäîâîå
íàçâàíèå Thoa (Aublet, 1775), êîòîðîå øèðîêî èñïîëüçîâàëîñü â ëèòåðàòóðå äî
îáîáùàþùèõ ñèñòåìàòèêó ïðåäøåñòâóþùèõ ýïîõ ñèíîïñèñà õâîéíûõ Ýíäëèõåðà
(Endlicher, 1847) è Äåêàíäîëëåâñêîãî ïðîäðîìóñà (Parlatore, 1868). Íå ïåðåíîñÿ
ôîðìàëüíî âíóòðèðîäîâûå òàêñîíîìè÷åñêèå åäèíèöû, óñòàíîâëåííûå Ìàðêãðàôîì â 1930 ã., êîòîðûå íå ïîëó÷èëè ñâîåãî ïîäòâåðæäåíèÿ â ñîâðåìåííûõ
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Turczaninowia 2000 3(4) : 28–36
èññëåäîâàíèÿõ ðåïðîäóêòèâíûõ îðãàíîâ ãíåòîâûõ, íàìè ïðåäëîæåíû êîìáèíàöèè
äëÿ âñåõ âèäîâ, êîòîðûå ñåãîäíÿ ïðèçíàþòñÿ â ñîñòàâå ðîäà Gnetum, çà èñêëþ÷åíèåì G. gnemon.  ïîñëåäóþùåé ïëàíèðóåìîé ðåâèçèè ðîäà áóäóò ââåäåíû
âíóòðèðîäîâûå ïîäðàçäåëåíèÿ äëÿ îòðàæåíèÿ â ñèñòåìå ñîâðåìåííîé êàðòèíû
èððàäèàöèè ðîäà.
Thoa Aublet, 1775, Hist. Pl. Guiane, 2 : 874; Scop. 1777, Intr. Hist. Nat. : 265;
Juss. 1789, Gen. Pl. : 406; Gmel. 1791, Syst. : 826; Schreb. 1791, Gen. Pl. 2 : 650;
Batsch, 1802, Tab. : 178; St. Hil. 1805, Éxpos. Fam. 2 : 312; Willd. 1805, Sp. Pl. 4, 1 :
476; Pers. 1807, Ench. Bot. 2 : 573; Spreng. 1817, Anleit. Kenntn. Gew. 2, 1 : 361; id.
1825, Syst. 2 : 461; C. Agardh, 1825, Aphor. Bot. 14 : 204; Reichenb. 1828, Consp.
Regn. Veg. : 92; Blanco, 1837, Fl. Filipp. : 764. – Gnetum sect. Thoa Endl. 1847, Syn.
Conif. : 251. — Gnetum sect. Erecta Griffith, 1859, Trans. Linn. Soc. London, 22 :
308. – Gnetum sect. Scandentia Griffith, 1859, l.c. : 308. – Ula Rheede, 1688, Hort.
Malab. 7 : 41, tab. 22, nom. prae-Linn. – Abutua Loureir. 1790, Fl. Conchinch. 2 : 773–
774. – Gnemon Rumph. ex O. Kuntze, 1891, Rev. Gen. Pl. 2 : 796, p.p., excl. typo. –
Arthostema Necker, 1790, Elem. Bot. 2 : 280, nom. utique rejic., p.p. (excl. typo).
Ò è ï: Th. urens Aublet
34 âèäà:
Thoa acuta (Markgr.) Doweld comb. nov. – Gnetum acutum Markgr. 1972,
Fl. Males. Ser. 1, 6, 6 : 947. – Gnetum acutatum Markgr. 1966, Blumea, 13, 2 : 404,
nom. invalid., non Miq. 1861, Fl. Ind. Bat. Suppl. 588. Ãîëîòèï: P. S. Ashton 17804 (L).
Thoa africana (Welw.) Doweld comb. nov. – Gnetum africanum Welw.
1869, Trans. Linn. Soc. London, 27, 1 : 73.
Thoa arborea (Foxw.) Doweld comb. nov. – Gnetum arboreum Foxw. 1911,
Philipp. J. Sci. Bot. 6, 3 : 174.
Thoa buchholziana (Engl.) Doweld comb. nov. – Gnetum buchholzianum
Engl. 1907, Bot. Jahrb. Syst. 40, 1 : 519.
Thoa catasphaerica (H. Shao) Doweld comb. nov. – Gnetum
catasphaericum H. Shao, 1994, Guihaia, 14, 4 : 297.
Thoa contracta (Markgr.) Doweld comb. nov. — Gnetum contractum
Markgr. 1930, Bull. Jard. Bot. Buitenzorg, Sér. 3, 10, 4 : 470.
Thoa cuspidata (Blume) Doweld comb. nov. – Gnetum cuspidatum Blume,
1848, Rumphia, 4 : 5. – Gnetum longispicum Ridl. 1911, J. As. Soc. Straits, 60 : 63
(“longispica”). – Gnetum penangense Ridl. 1911, l.c. : 62.
Thoa diminuta (Markgr.) Doweld comb. nov. – Gnetum diminutum Markgr.
1930, Bull. Jard. Bot. Buitenzorg, Sér. 3, 10, 4 : 483.
Thoa edulis Willd. 1805, Sp. Pl. 4 : 477. – Gnetum ula Brongn. 1829, in Duperrey,
Voy. Coquille : 12.
Thoa globosa (Markgr.) Doweld comb. nov. – Gnetum globosum Markgr.
1971, Blumea, 19, 1 : 108. Ãîëîòèï: T. C. Whitmore F.R.I. 15262 (L).
Thoa gnemonoidea (Brongn.) Doweld comb. nov. – Gnetum gnemonoides
Brongn. 1829, in Duperrey, Voy. Coquille : 12. – Gnetum rumphianum Beccari, 1877,
Malesia, 1 : 182. — Gnetum macrocarpum Beccari, 1877, l.c. : 182. – Gnetum
Äîóýëüä À.Á.: Âîññòàíîâëåíèå ðîäà Thoa Aublet (Gnetaceae)
33
verrucosum Karst. 1893, Ann. Jard. Bot. Buitenzorg, 11 : 216. – Gnetum kerstingii
K. Schum. et Lauterb. 1901, Fl. Deutsch. Südsee : 157. – Gnetum wrayi Gamble,
1925, Kew Bull. 2 : 92.
Thoa hainanensis (C.Y. Cheng ex L.K. Fu, Y.F. Yu et M.G. Gilbert)
Doweld comb. nov. – Gnetum hainanense C.Y. Cheng ex L.K. Fu, Y.F. Yu et M.G.
Gilbert, 1999, Novon, 9, 2 : 187. – Gnetum hainanense C.Y. Cheng, 1975, Acta
Phytotax. Sin. 13, 4 : 88, nom. invalid. Ãîëîòèï: Trang et Fung 554 (PE).
Thoa indica (Loureir.) Doweld comb. nov. – Abutua indica Loureir. 1790,
Fl. Conchinch. 2 : 775. – Gnetum latifolium Blume, 1834, Tijdschr. Natuur. Gesch. 1,
3 : 160–161. – Gnetum minum Foxw. 1911, Philipp. J. Sci. Bot. 6, 3 : 176 (“minus”). –
Gnetum laxifrutescens Elmer, 1912, Leafl. Philipp. Bot. 4 : 1478. – Gnetum indicum
Merrill, 1917, Interpr. Rumph. Herb. Amboin. : 77.
Thoa klossii (Merrill ex Markgr.) Doweld comb. nov. – Gnetum klossii
Merrill ex Markgr. 1930, Bull. Jard. Bot. Buitenzorg, Sér. 3, 10, 4 : 478.
Thoa leptostachya (Blume) Doweld comb. nov. – Gnetum leptostachyum
Blume, 1848, Rumphia, 4 : 5.
Thoa leyboldii (Tulasne) Doweld comb. nov. – Gnetum leyboldii Tulasne,
1858, Ann. Sci. Nat. Bot. Sér. 4, 10 : 115. – Gnetum paraense Huber, 1902, Bol. Mus.
Paraense Hist. Nat. Ethnogr. 3 : 403.
Thoa loerzingii (Markgr.) Doweld comb. nov. – Gnetum loerzingii Markgr.
1930, Bull. Jard. Bot. Buitenzorg, Sér. 3, 10, 4 : 480.
Thoa macrostachya (J.D. Hook.) Doweld comb. nov. – Gnetum
macrostachyum J.D. Hook. 1890, Fl. Brit. Ind. 5 : 642.
Thoa markgrafii Doweld nom. nov. – Gnetum bosavicum Markgr. 1977,
Blumea, 23, 2 : 478, nom. invalid. (quoad descr. lat.). Ãîëîòèï: M. Jacobs 8783 (L).
Thoa microcarpa (Blume) Doweld comb. nov. – Gnetum microcarpum
Blume, 1848, Rumphia, 4 : 7. – Gnetum apiculatum Griff. 1854, Notul. Pl. Asiat. 4 :
30. – Gnetum sylvestre Gamble ex Ridl. 1925, Fl. Malay Penins. 5 : 275, nom. illeg.
non Brongn. 1829, in Duperrey, Voy. Coquille : 12.
Thoa montana (Markgr.) Doweld comb. nov. – Gnetum montanum Markgr.
1930, Bull. Jard. Bot. Buitenzorg, Sér. 3, 10, 4 : 466.
Thoa neglecta (Blume) Doweld comb. nov. – Gnetum neglectum Blume,
1848, Rumphia, 4 : 6.
Thoa nodiflora (Brongn.) Doweld comb. nov. – Gnetum nodiflorum Brongn.
1829, in Duperrey, Voy. Coquille : 12. – Gnetum amazonicum Tulasne, 1858, Ann. Sci.
Nat. Bot. Sér. 4, 10 : 116. – Gnetum dioicum Leyb. ex Tulasne, 1858, l.c. : 115, 116. –
Gnetum cruzianum Gleason, 1925, Bull. Torrey Bot. Club, 52 : 196. – Gnetum melinonii
Benoist, 1945, Bull. Mus. d’Hist. Nat. Paris, Sér. 2, 17, 1 : 66.
Thoa oblonga (Markgr.) Doweld comb. nov. – Gnetum oblongum Markgr.
1930, Bull. Jard. Bot. Buitenzorg, Sér. 3, 10, 4 : 471.
Thoa oxycarpa (Ridl.) Doweld comb. nov. – Gnetum oxycarpum Ridl. 1926,
Kew Bull. 2 : 94.
Thoa paniculata (Spruce ex Benth.) Doweld comb. nov. – Gnetum
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paniculatum Spruce ex Benth. 1856, Hook. J. Bot. 8 : 357. – Gnetum microstachyum
Spruce ex Benth. 1856, l.c. : 358.
Thoa parvifolia (Warb.) Doweld comb. et stat. nov. – Gnetum scandens
var. parvifolium Warb. 1900, Monsunia, 1 : 196. – Gnetum parvifolium (Warb.)
W.C. Cheng, 1964, Acta Phytotax. Sin. 9, 4 : 386.
Thoa raya (Markgr.) Doweld comb. nov. – Gnetum raya Markgr. 1967,
Blumea, 14, 1 : 284. Ãîëîòèï: Jugah Kudi S. 23801 (L).
Thoa ridleyi (Gamble ex Markgr.) Doweld comb. nov. – Gnetum ridleyi
Gamble ex Markgr. 1930, Bull. Jard. Bot. Buitenzorg, Ser. 3, 10, 4 : 479. – Gnetum
ridleyi Gamble ex Burkill et H.R. Henderson, 1925, Garden’s Bull. Straits Settl. 3 :
458, nom. nud.
Thoa scandens (Roxb.) Doweld comb. nov. – Gnetum scandens Roxb. 1814,
Hort. Bengal. : 66.
Thoa schwackeana (Taub. ex A. Schenk) Doweld comb. nov. – Gnetum
schwackeanum Taub. ex A. Schenk, 1893, in W. Schimper, Bot. Mitt. Tropen, 5, 2 :
249.
Thoa tenuifolia (Ridl.) Doweld comb. nov. – Gnetum tenuifolium Ridl. 1911,
J. As. Soc. Straits, 59 : 188.
Thoa urens Aublet 1775, Hist. Pl. Guiane, 2 : 874. – Gnetum thoa Brongn.
1829, in Duperrey, Voy. Coquille : 12. – Gnetum urens (Aubl.) Blume, 1834, Tijdschr.
Natuur. Gesch. 1, 3 : 162. – Gnetum pyrifolium Miq. ex Parlat. 1869, in DC. Prodr.
16, 2 : 350.
Thoa venosa (Spruce ex Benth.) Doweld comb. nov. – Gnetum venosum
Spruce ex Benth. 1856, Hook. J. Bot. 8 : 358.
Gnetum L. f. 1767, Mant. 1 : 18, 125; Syst. Nat. ed. 12 : 637. – Gnemon
Rumph. 1741, Herb. Amboin. 1 : 181, nom. prae-Linn. – Gnemon Rumph. ex O. Kuntze,
1891, Rev. Gen. Pl. 2 : 796. – Arthostema Necker, 1790, Elem. Bot. 2 : 280, nom.
utique rejic., p.p.
Ò è ï: G. gnemon L.
2 (?) âèäà: G. gnemon, (?) G. costatum K. Schum. 1889, in K. Schum. et Hollr.
Fl. Kais. Wilh. Land : 13.
Species incertae sedis:
G. cavaleriei Leveille, 1914/15, Fl. Kouy-Tcheou : 186. Ïðèìå÷àíèå: åäèíñòâåííûé ýêçåìïëÿð ýòîãî âèäà ïðåäñòàâëÿåò ñîáîé îáëèñòâåííûå ïîáåãè òîïîëÿ.
G. karstenianum Warb. 1900, Monsunia, 1 : 196–197. Ïðèìå÷àíèå:
îòñóòñòâóþò ðåïðîäóêòèâíûå îðãàíû, ïîçâîëÿþùèå èäåíòèôèêàöèþ âèäà.
G. nigrum Carrière, 1855, Traité Gen. Conif. : 545. Ïðèìå÷àíèå: îòñóòñòâóþò
ðåïðîäóêòèâíûå îðãàíû, ïîçâîëÿþùèå èäåíòèôèêàöèþ âèäà.
ËÈÒÅÐÀÒÓÐÀ
Êðàñèëîâ Â.À. Ïðîèñõîæäåíèå è ðàííÿÿ ýâîëþöèÿ öâåòêîâûõ ðàñòåíèé. – Ì., 1989. –
Äîóýëüä À.Á.: Âîññòàíîâëåíèå ðîäà Thoa Aublet (Gnetaceae)
35
264 ñ.
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Turczaninowia 2000 3(4) : 28–36
36
SUMMARY
A new system of classification of the genus Gnetum L. f. is proposed upon the base of
distinct reproductive morphology and pollen morphology in particular. All species of the genus,
having a dimerous system of seed coats, are transferred into a nearly forgotten genus Thoa
Aublet, which is an older available generic name for that group of species. The genus Gnetum
s.str. is re-circumscribed as consisting of two species only, G. gnemon L. and G. costatum K.
Schum., which are characterized by more primitive, trimerous system of seed coats. To put into
order the descriptive morphology of gnetoid reproductive shoots, a new system of terms is
introduced. The trimerous system is considered as most archaic; the inner envelope is treated
as a homologue of the outer integument of other seed plants (the inner one is probably aborted
in seed evolution and sometimes reported as rudiments within the outer one); the other coats
(“envelopes”) are considered as collar derivatives of the two pairs of originally decussate
bracteose foliar structures, which are fused within Thoa into a single compound structure. New
terms were proposed for a descriptive morphology of gnetoid fructifications: a tegule (tegula,
lat.), which is applied to outer and middle envelopes of the Gnetum seed, a syntegule (syntegula,
lat.) termed a compound single outer seed-coat consisting of two concrescent tegules (in a
Thoa seed), glandidium – a new term introduced instead of “flower”, “drupe”, “arcesthide” and
“seed” for gnetoid “seeds”, inglandescence (inglandescentia, lat. by analogy of inflorescentia,
lat., inflorescence) for the aggregations of male and female glandidia. A further re-classification
of glandidia is proposed: superior glandidia (having free tegules – in Gnetum s.str.) and inferior
ones (characterized by concrescent tegules – syntegules – in Thoa). A new system of
morphological terms should be used in the analysis of homological/analogical structures within
extant and extinct spermatophytes. 33 new nomenclatural combinations were validated.
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Ïîëó÷åíî 10.10.2000 ã.