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Em[Ginkgo Gomolitzkyana N. Nosova, Sp. Nov. Из Среднею УДК 561.46:551.762 Палеоботаника. 2012. Т. 3. С. 62—91 ЖЕНСКИЕ РЕПРОДУКТИВНЫЕ СТРУКТУРЫ GINKGO GOMOLITZKYANA N. NOSOVA, SP. NOV. ИЗ СРЕДНЕЮРСКИХ ОТЛОЖЕНИЙ АНГРЕНА (УЗБЕКИСТАН) Н. В. Носова Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург Впервые из юрских отложений Средней Азии (Узбекистан, Ангрен) описаны женские репродук- тивные структуры Ginkgo. Выделен новый вид — Ginkgo gomolitzkyana N. Nosova, sp. nov. Изучено эпи- дермальное строение воротничка, расположенного в основании семязачатка, а также строение на- ружной и внутренней эпидермы интегумента, эпидермы нуцеллуса и строение мегаспоровой мем- браны семязачатков (семян). Ключевые слова: Ginkgo, семязачатки, семена, интегумент, нуцеллус, мегаспоровая мембрана, кутикула, средняя юра, Узбекистан. ВВЕДЕНИЕ В течение последних трех десятилетий описано несколько хорошо сохранившихся ископаемых женских репродуктивных структур Ginkgo L. В мезозое выделено три вида из средней юры и один — из раннего мела. Среднеюрские виды описаны из Китая — Ginkgo yimaensis Zhou et Zhang (Zhou, Zhang, 1989) и Ginkgo sp. (Deng et al., 2004), и Швеции — G. ginkgoidea (Tralau) Yang, Friis et Zhou (Yang et al., 2008). Раннемеловой вид — G. apodes Zheng et Zhou (Zheng, Zhou, 2004) — происходит из Китая. Все мезозойские виды найдены в ассоциации с сильно рассеченными листьями Ginkgo и Ginkgoites. В отличие от юрских и меловых видов единственный палеоценовый гинкго с изученными женскими репродуктивными структурами — G. cranei Zhou, Quan et Liu из США — найден совместно с цельнокрайными листьями Ginkgo (Crane et al., 1990; Zhou et al., 2012). Исследование ископаемых находок женских репродуктивных структур Ginkgo значительно расширило представление о разно- образии Ginkgoales и внесло заметный вклад в решение вопросов эволюции гинкговых (Zhou, 1991; Zhou, 1997; Zhou, 2009; Zhou et al., 2012). Как и у всех голосеменных, семязачаток гинкго состоит из мегаспорангия (нуцеллуса), окружен- ного интегументом. Однако в отличие от остальных голосеменных, за исключением саговниковых, для него характерна ранняя дифференциация интегумента, которая происход ит задолго до оплодот- ворения. Интегумент гинкго состоит из трех слоев. Наружный слой интегумента — саркотеста — мя- систый, достигает в толщину 5—6 мм и покрыт эпидермой с устьицами. Осенью саркотеста окраши- вается в янтарно-желтый цвет. Средний слой — склеротеста — твердый, образует «косточку» созрев- шего семязачатка, а впоследствии — семени. Примыкающий к нуцеллусу внутренний, самый тонкий слой — эндотеста. Как известно, семязачаток превращается в семя после оплодотворения. По степени развития интегумента у гинкго невозможно отличить стерильный семязачаток от оплодотворенного, не сде- лав поперечного среза. Известно, что развитие зародыша, а нередко и оплодотворение происходит у G. biloba уже в опавших с дерева семязачатках. При описании ископаемого материала Ginkgo не- которые авторы используют термин «семязачаток», если он найден прикрепленным к семяножке, и «семя» — для дисперсных семян. В данной статье все остатки (прикрепленные к семяножке и дис- персные) описываются как «семязачаток (семя)». Описываемый материал происходит из отложений ангренской свиты угольного месторождения Ангрен (Узбекистан). Систематический состав флоры свидетельствует о том, что возраст отложений ангренской свиты не древнее средней юры (Сикстель, 1953; Гомолицкий, Лобанова, 1969; Гомолиц- кий и др., 1981), предположительно аален-байос (Troitsky, Gomolitzky, 1996; Носова, 1998). Описан- ные в данной статье растительные остатки представлены семязачатками (семенами), два из которых сидят на семяножках с воротничками. Из-за фрагментарности материала полная длина семяножки 62 неизвестна, как не установлено количество и расположение семязачатков на одной генеративной оси. Саркотеста сохранилась в виде углефицированного слоя толщиной 0,25—0,65 мм, под которым рас- положена «косточка», образованная склеротестой. На некоторых образцах сохранились отдельные «косточки». В процессе мацерации углистый слой саркотесты растворяется, остаются только смоля- ные тельца и наружная кутикула, несущая на себе отпечаток строения эпидермы, покрывающей сар- котесту. Склеротеста растворяется полностью. Кроме наружной кутикулы интегумента после маце- рации остаются также внутренняя кутикулы интегумента (эндотесты), кутикула нуцеллуса и мега- споровая мембрана. На основе полученных данных по строению наружной и внутренней эпидермы интегумента, эпи- дермы нуцеллуса и мегаспоровой мембраны семязачатков (семян) выделен новый вид Ginkgo go- molitzkyana N. Nosova, sp. nov., который заметно отличается от ранее описанных ископаемых видов Ginkgo и от современного G. biloba. Представленный в данной статье материал является первой находкой женских репродуктивных структур Ginkgo в мезозойских отложениях Средней Азии. МАТЕРИАЛ И МЕТОДЫ Сборы материала на угольном месторождении Ангрен (41°01' с. ш., 70°08' в. д., рис. 1), г. Ангрен, Ташкентская область, Узбекистан, проводились с 1976 по 1981 год сотрудниками лаборатории па- леоботаники Ботанического института АН СССР: Л. Ю. Буданцевым, Д. В. Громыко, Г. В. Делле и В. А. Самылиной. Образцы хранятся в Ботаническом институте им. В. Л. Комарова (БИН) РАН (Санкт-Петербург): колл. БИН №№ 813, 820—823. Описываемый в данной статье материал представлен многочисленными объемно сохранивши- мися семязачатками (семенами) и их фрагментами, из них четырнадцать подверглись мацерации. Растительные остатки мацерировались по стандартной методике, в дальнейшем их эпидермаль- ное строение изучалось при помощи светового микроскопа (СМ) — Carl Zeiss Axio Scope A1, и ска- нирующего электронного микроскопа (СЭМ) — JSM-6390 LA. При описании строения мегаспоровой мембраны использовалась терминология, принятая Ч. Чжоу (Zhou, 1993). Рис. 1. Географическое положение г. Ангрен. 63 СИСТЕМАТИКА Класс GINKGOOPSIDA Порядок GINKGOALES GINKGO L., 1771 Ginkgo gomolitzkyana N. Nosova, sp. nov. Табл. I—VIII Название вида — в честь палеоботаника Н. П. Гомолицкого. Голотип. Колл. БИН № 821, экз. 45-15(1); семязачаток с воротничком на семяножке, Узбекистан, Ташкентская область, Ангрен, ангренская свита, средняя юра. — Табл. I, фиг. 1а. Holotype. Сoll. BIN № 821, spec. 45-15(1); ovule with collar on pedicel, Uzbekistan, Tashkent region, Angren, the Angrenian Formation, the Middle Jurassic. — Pl. I, fi g. 1a. Диагноз. Семязачаток расположен на семяножке с воротничком. Диаметр воротничка 2,5—4 мм. Устьица на наружной части воротничка многочисленные — 65—75 на 1 мм2. П обочные клетки без папилл. Апертурные стенки побочных клеток часто утолщены, образуя кутикулярный валик во- круг апертуры. Семязачатки (семена) в очертании округлой или овальной формы c закругленным основанием и острой верхушкой. Длина семязачатков (семян) 5,3—9,2 мм, ширина 4—7,5 мм. Толщи- на саркотесты 0,25—0,65 мм. Толщина наружной кутикулы интегумента 1,0—5,5 мкм. Устьица ред- кие — 2—12 на 1 мм2. Большинство побочных клеток без папилл, редко встречаются устьица с прок- симальными папиллами. Апертурные стенки побочных клеток часто утолщены, образуя кутикуляр- ную складку вокруг апертуры. Антиклинальные стенки клеток прямые, периклинальные — ровные. Эпидермальные клетки нуцеллуса удлиненные с закругленными углами. Мегаспоровая мембрана толщиной 2-5 мкм. Альвеолярный слой сложен расположенными косо или параллельно к подстила- ющему слою ветвящимися и соединяющимися друг с другом столбиками, образующими сеть. Diagnosis. Ovule attached to pedicel with a collar. Collar diameter 2,5—4,5 mm. Stomata on the outer collar surface numerous — 65—75 per 1 mm2. Subsidiary cells lacking papillae. Aperture walls of the sub- sidiary cells often thickened forming a cuticular rim around the aperture. Seeds rounded to ovate in outline with rounded base and pointed apex. Ovule (seed) length 5,3—9,2 mm, width 4—7,5 mm. Thickness of the sarcotesta 0,25—0,65 mm. Thickness of the outer cuticle of the integument 1,0—5,5 μm. Stomata sparse — 2—12 per 1 mm2. Most of subsidiary cells lacking papillae, occasionally stomata with proximal papillae oc- cur. Aperture walls of the subsidiary cells often thickened forming a cuticular fold around the aperture. An- ticlinal cell walls straight, periclinal smooth. Epidermal cells of nucellus elongated with rounded corners. Megaspore membrane 2—5 μm thick. Patterned layer is composed of bacula arranged obliquely or parallel to the foot layer, branching and connected to one another to form a network. Описание. В коллекции имеются многочисленные семязачатки (семена) и их фрагменты (табл. I, II). На двух образцах сохранились небольшие — около 1 мм длиной и 0,7—0,8 мм шириной — фрагменты семяножек, расширяющиеся в чашевидную структуру — воротничок, на котором распо- ложен семязачаток (табл. I, фиг. 1, 3). Высота воротничков 1—1,8 мм, диаметр 2,5—4 мм. На одном из образцов рядом с семязачатком на семяножке находятся еще два отдельных семязачатка (семени) (табл. I, фиг. 1), на втором — три семязачатка (семени) (табл. I, фиг. 3). По расположению семязачат- ков (семян) можно предположить, что они (три на одном образце и четыре — на другом) сидели на одной генеративной оси. Кутикула на наружной поверхности воротничка тонкая. Большинство клеток эпидермы корот- кие, четырех-, пяти- и шестиугольные. Антиклинальные стенки клеток прямые или неравномерно утолщенные. Периклинальные стенки клеток часто выпуклые (табл. III, фиг. 9). Углы клеток острые или закругленные. Устьица многочисленные, разбросаны по всей поверхности, ориентированы бес- порядочно. На 1 мм2 приходится 65—75 устьиц. Устьичные комплексы неполно амфициклические или
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