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Морфология Пыльцевых Зерен Некоторых Родов Из Триб Trigonotideae И Myosotideae (Boraginaceae)

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Морфология Пыльцевых Зерен Некоторых Родов Из Триб Trigonotideae И Myosotideae (Boraginaceae) «Растительный мир Азиатской России», 2008, № 1, 37–51 УДК 582.948 © 2008 МОРФОЛОГИЯ ПЫЛЬЦЕВЫХ ЗЕРЕН НЕКОТОРЫХ РОДОВ ИЗ ТРИБ TRIGONOTIDEAE И MYOSOTIDEAE (BORAGINACEAE) О. Д. НикифОрОва MORPHOLOGY OF POLLEN GRAINS OF SOME GENERA OF THE TRIBES TRIGONOTIDEAE AND MYOSOTIDEAE (BORAGINACEAE) O. D. NikifOrOva Центральный сибирский ботанический сад СО РАН, 630090 Новосибирск, ул. Золотодолинская, 101 Central Siberian Botanical Garden, SB RAS, 630090 Novosibirsk, Zolotodolinskaya st., 101 Fax: +7(383) 330-19-86, e-mail: [email protected] Подведены итоги исследования морфологии пыльцевых зерен с помощью сканирующего электронного микроскопа видов в родах Mertensia, Pseudomertensia, Trigоnotis, Myosotis, Strophiostoma, Trigonocаryum из триб Trigonotideae и Myosotideae. Впервые изучена морфология пыльцевых зерен 27 азиатских и северо- американских видов рода Mertensia и 5 видов гималайского рода Pseudomertensia. Все изученные роды относятся к группе единного родства, так как для них характерен один тип пыльцевых зерен: коконооб- разные, с большим или меньшим сужением в области экватора, 6-бороздно-3-оровые, с чередующими- ся оровыми и безоровыми бороздами. Наиболее близкое родство проявляется между родами Myosotis, Strophiostoma, Trigonocаryum, так как у них не только одинаковый тип пыльцевых зерен, но и сходная по- верхность экзины: она ровная и гладкая, с перфорацией на полюсах. Пыльцевые зерна родов Mertensia, Pseudomertensia и Trigоnotis существенно отличаются скульптурированной поверхностью экзины, на ко- торой расположены бугорки. Морфология пыльцевых зерен рода Brachybotrys показывает родство с ро- дом Trigоnotis. Ключевые слова: Boraginaceae, триба Trigonotideae, Mertensia, Pseudomertensia, Trigоnotis, Myosotis, морфо- логия, пыльцевые зерна. The results of study of pollen morphology of the genera Mertensia Roth, Pseudomertensia Riedl, Trigоnotis Stev., Myosotis L., Strophiostoma Turcz., Trigonocаryum Stev. of the tribes Trigonotideae and Myosotideae by scanning electron microscopy have been summed up. For the first time morphology of pollen grains of 27 North-American and Asian species of the genus Mertensia Roth and 5 species of the genus Pseudomertensia from the Himalayas has been studied. All genera studied have 6-heterocolpate pollen grains (3-colpi alternate with 3-pseudocolpi) and belong to a unified evolutionary line as they have an identical type of pollen grains. Key words: Boraginaceae, tribe, Trigonotideae, Mertensia, Pseudomertensia, Trigоnotis, Myosotis, morphology, pol- len grains. А.П. Декандоль (A.P. de Candolle, 1846), а позднее Данная подтриба описана на русском языке и ока- М. Гюрке (Guerke, 1893) роды Mertensia Roth, Trigоno- залась невалидной. В.Р. Григорьев (1964) на осно- tis Stev., Myosotis L. включали в трибу Lithospermeae вании изучения семейства Boraginaceae методом DC. семейства Boraginaceae Juss. Во «Флоре СССР» анализа морфологического строя (Виноградов, М.Г. Попов (1953а) вслед за Х. Рейхенбахом 1947), предложил указанные выше роды отнести (Reichenbach, 1857) род Myosotis включил в само- к трибе Myosotideae, а ее разделить на подтрибы: стоятельную трибу Myosotideae Reichenb., а роды Myosotidinae, Trigonotidinae M. Pop., Bothriosperminae Mertensia, Trigоnotis и Brachybotrys Maxim. выде- M. Pop., но и эти предложения оказались невалид- лил в подтрибу Trigonotidinae трибы Lithospermeae. ными. Х. Ридл (H. Riedl, 1968) подтрибу Trigonotideae © Российская академия наук, Сибирское отделение; Центральный сибирский ботанический сад, 2008 узаконил в ранге трибы Trigonotidinae, значительно Hargrove, Simpson, 2003; Овчинникова, 2006; и др.). расширил ее состав, а роды Mertensia, Trigоnotis и Наиболее полные сведения о палиноморфологии Brachybotrys включил в состав типовой подтрибы. большинства родов семейства Boraginaceae содер- Для выявления и уточнения родственных связей жатся в работе Е.М. Аветисян (1956). Под световым родов в трибе Trigonotideae мною изучена морфо- микроскопом (СМ) ею изучены основные морфо- логия пыльцевых зерен (п. з.) видов разных родов логические характеристики п. з. разных родов данной трибы. Значение палиноморфологических этого семейства, но их видовое разнообразие не- признаков в систематике семейства Boraginaceae велико. Поэтому на современном этапе для целей оценивается исследователями высоко (Аветисян, таксономии необходимо пополнять базу данных 1956; Nowicke, Skvarla, 1974; Попова, Земскова, по морфологии п. з. за счет изучения большинс- 1990, 1995; Земскова, Попова, 1991; Bigazzi, Selvi, тва видов из разных родов с использованием ска- 1997, 2000; Khatamsaz, 2001; Никифорова, 2002; нирующего электронного микроскопа (СЭМ). МАТЕРИАЛ И МЕТОДЫ ИССЛЕДОВАНИЯ Материалом для работы послужили образ- тывали следующие признаки: форма и размеры цы пыльцы, отобранные с гербарных листов, хра- п. з., размеры полярной оси, экваториального диа- нящихся в Гербариях Ботанического института метра в узкой и широкой частях, форма и разме- им. В.Л. Комарова (БИН РАН, С.-Петербург, LE), ры оровых и безоровых борозд, размеры оры. Все Центрального сибирского ботанического сада СО измерения проводились масштабной линейкой РАН (Новосибирск, NSK, NS), Южносибирского на микрофотографиях п. з., при этом измеряли ботанического сада (г. Барнаул, SSBG) и Научно- 3–5 п. з., а в таблицу вносили крайние показатели производственного объединения «Ботаника» размеров. Для описания использовали терминоло- (г. Ташкент, TASH). гию, принятую в работах Л.А. Куприяновой (1996), Пыльца исследовалась на СЭМ марки Multiscan Куприяновой, Л.А. Алешиной (1967), Т.Н. Попо- 200 GS, LEO 420 при увеличении ×10 000 и ×15 000 вой, Е.А. Земсковой (1995). Описания иллюстриро- после напыления медью. При описании п. з. учи- ваны микрофотографиями. РЕЗУЛЬТАТЫ И ИХ ОБСУЖДЕНИЕ Genus Mertensia Roth морфологические характеристики п. з. 7 видов в Род Mertensia Roth включает около 45 видов, роде Mertensia, в том числе 5 сибирских и сибир- которые почти поровну распределены между ско-монгольских видов (M. serrulata, M. stylosa, Северной Америкой и Азией с небольшим пре- M. davurica, M. pallasii, M. sibirica), циркумполярного обладанием в Северной Америке. Единственная M. maritima и памиро-алайского M. dschagastanica. система рода, охватывающая основное разнообра- Она показала, что для изученных видов Mertensia зие видов, принадлежит М.Г. Попову (1953б). Он характерны п. з. коконообразной формы, 12.6– разделил род Mertensia на 6 секций: Typomertensia 16.8 мкм дл. полярной оси, 6-бороздно-3 оровые, M. Pop., Steenhammera A. Gray, Neuranthia Williams, из которых оровые борозды чередуются с тремя бе- Mertensianthe (M. Pop.) M. Pop., Eumertensia и зоровыми. Оры элипсоидальные, расположенные Oreocharis (Decne.) M. Pop., а многовидовую секцию на экваторе. Eumertensia (Mertensia) — на группы близкородс- С помощью СЭМ исследована морфология твенных видов (greges), для которых дал латинский п. з. 27 видов из 5 секций рода Mertensia из разных диагноз. Во «Флоре СССР» (Попов, 1953а) состав районов Евразии и Северной Америки. Для всех азиатских видов, объединенных в greges, идентичен изученных видов характерны коконообразные п. з. сериям (series), но они оказались невалидными. В с большим или меньшим сужением в экватори- настоящее время по правилам Международной альной части или сфероидальные с едва заметным номеклатуры greges являются незаконными таксо- сужением в области экватора, 8.1–12.5(17.0) мкм нами (Международный кодекс…, 2001). дл. полярной оси. П. з. с полюсов округлые или Первые сведения о палиноморфологии видов в 6-угольно-округлые. Три более широкие оровые роде Mertensia содержатся в работе Аветисян (1956). борозды чередуются с безоровыми бороздами. Под световым микроскопом ею изучены основные Безоровые борозды узко- или широколанцетные, 38 края борозд зернистые (гранулированные), с выра- По данным Аветисян (1956) п. з. M. maritima женной неровностью. Оровые борозды к экватору сильно сужено в области экватора, что совпадает с расширяются, приобретая ромбовидную форму. нашими данными. Оры расположены по экватору, округлые, округ- 3. M. asiatica Macbr. [Дальний Восток], Порт Аян ло-эллипсоидальные или узко-эллипсоидальные, у Охотского моря, берег моря, на гальке 28 VII 1912, их поверхность гранулированная. У большинства Ф.В. Соколов (LE). — Литоральный притихоокеан- видов поверхность экзины бугорчатая, форма бу- ский вид. горков одинаковая, но их плотность разная. Чаще П. з. коконообразные, с сильным сужением в бугорки рассеяны по всей поверхности, реже — области экватора, оровая борозда чуть короче бе- только на полюсах. зоровой, широкоромбическая, по краям грану- В таблице приведены основные палиноморфо- лированная; безоровая борозда узколанцетная с логические характеристики п. з. исследованных ви- сомкнутыми краями. Ора выпуклая. Поверхность дов, которые расположены в порядке принимаемой экзины плотнобугорчатая, в области экватора глад- нами системы рода Mertensia (Попов, 1953б), за ис- кая. Морфологически сходны с п. з. M. maritima, но ключением секции Oreocharis, которую мы призна- отличаются большими размерами полярной оси и ем в качестве самостоятельного рода Pseudomertensia. экваториального диаметра (рис. 1, 3). П. з. изученных видов различаются формой, разме- рами, соотношением длины оровых и безоровых Sect. Neuranthia Williams борозд (оровая борозда равна, короче или длин- 4. M. bella Piper. [США], Oregon, Cascade нее безоровой), формой и размерами оры, а также Range, № 2701, 14 VI 1966, N.H. Holmgren (LE). — скульптурированностью поверхности экзины. Североамериканский сонорский вид запада и юго- Ниже приводим местонахождения исследован-
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