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Морфология Спор Некоторых Представителей Подсемейства Pteridoideae Семейства Pteridaceae А.В

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Морфология Спор Некоторых Представителей Подсемейства Pteridoideae Семейства Pteridaceae А.В НАУЧНЫЙ ЖУРНАЛ РАСТИТЕЛЬНЫЙ МИР АЗИАТСКОЙ РОССИИ Растительный мир Азиатской России, 2014, № 2(14), с. 29–36 h p://www.izdatgeo.ru УДК 582.394 МОРФОЛОГИЯ СПОР НЕКОТОРЫХ ПРЕДСТАВИТЕЛЕЙ ПОДСЕМЕЙСТВА PTERIDOIDEAE СЕМЕЙСТВА PTERIDACEAE А.В. Ваганов1, А.П. Шалимов1, Д.Н. Шауло2 1Алтайский государственный университет, 656049, Барнаул, просп. Ленина, 61, e-mail: [email protected] 2 Центральный сибирский ботанический сад СО РАН, 630090, Новосибирск, ул. Золотодолинская, 101, e-mail: [email protected] Методом сканирующей электронной микроскопии (СЭМ) проведено сравнительное исследование шес- ти представителей подсемейства Pteridoideae C. Chr. ex Crabbe, Jermy et Mickel семейства Pteridaceae E.D.M. Kirchn.: Jamesonia auriculata A.F. Tryon, Pteris aspericaulis Wallich ex J. Agardh, P. biaurita L., P. te ne ra Kaulf., P. umbro s a R. Br., Taenitis blechnoides (Willd.) Sw. В результате морфологических описаний выявлены признаки, позволяющие судить о принадлежности исследуемых видов к одному подсемейству – Pteridoideae. Ключевые слова: подсемейство Pteridoideae, семейство Pteridaceae, Pteris, Jamesonia, Taenitis, морфология спор, сканирующая электронная микроскопия (СЭМ). SPORE MORPHOLOGY OF SOME REPRESENTATIVES OF PTERIDACEAE SUBFAM. PTERIDOIDEAE A.V. Vaganov1, A.P. Shalimov1, D.N. Shaulo2 1Altai State University, 656049, Barnaul, Lenina str., 61, e-mail: [email protected] 2Central Siberian Botanical Garden, SB RAS, 630090, Novosibirsk, Zolotodolinskaya str., 101, e-mail: [email protected] Th e method of scanning electronic microscopy (SEM) carries out relative research of ten representatives of subfamily Pteridoideae C. Chr. ex Crabbe, Jermy et Mickel family Pteridaceae E.D.M. Kirchn.: Jamesonia auriculata A.F. Tryon, Pteris aspericaulis Wallich ex J. Agardh, P. biaurita L., P. te ne ra Kaulf., P. umbrosa R. Br., Taenitis blechnoides (Willd.) Sw. As a result, morphological descriptions revealed signs that would reveal the ownership of the studied species to one subfamily – Pteridoideae. Key words: subfamily Pteridoideae, family Pteridaceae, Pteris, Jamesonia, Taenitis, morphology of the spores, scanning electronic microscopy (SEM). ВВЕДЕНИЕ Cемейство Pteridaceae E.D.M. Kirchn. считается Smith et al., 2006; Christenhusz et al., 2011) позволило одним из самых сложных по числу неразрешенных доказать с учетом глубокого анализа ранее предла- вопросов в систематике его таксономических рангов. гаемых систем принадлежность к семейству Pterida- Ввиду слабой изученности представителей этого се- ceae пяти подсемейств: Cryptogrammoideae S. Linds.; мейства, а также отсутствия доказательных методик Pteridoideae C. Chr. ex Crabbe, Jermy et Mickel; его окончательный объем не установлен. Представи- Ceratopteridoideae (J. Sm.) R.M. Tryon; Vittarioideae тели семейства Pteridaceae распространенны на всех (C. Presl) Crabbe, Jermy et Mickel; Cheilanthoideae континентах, за исключением Антарктиды. Макси- W.C. Shieh. При построении системы семейства све- мальное видовое разнообразие наблюдается в тропи- дения по отдельным генам хлоропластной и ядерной ческих и субтропических широтах Старого и Нового ДНК использовались для того, чтобы воссоздать меж- Света. родственные связи в системе семейства Pteridaceae с Система семейства Pteridaceae на фоне разраба- помощью филогенетических деревьев. тываемых систем папоротников неоднократно дора- Подсемейства Ceratopteridoideae и Cheilanthoideae, батывалась. В настоящее время использование мето- возводимые рядом авторов в ранг семейств, считают- дов молекулярной генетики для систематических за- ся наиболее устойчивыми в мировой системе папо- дач в ряде капитальных трудов (Schuettpelz et al., 2007; ротников, в том числе ввиду наличия стабильных © А.В. Ваганов, А.П. Шалимов, Д.Н. Шауло, 2014 29 признаков внешней морфологии. В подсемействах статочно внимания в крупных сводках, посвященных Vittarioideae, Cryptogrammoideae и Pteridoideae сложи- этой группе папоротников (Belling, Heusserl, 1957; лась обратная ситуация – до настоящего времени не Nayar, Devi, 1966, 1967; Chen, Huang, 1974; Wagner, существует однозначных доказательств обоснован- 1974). Однако в этих работах приводятся описания ности включения в состав указанных подсемейств лишь по некоторым представителям подсемейства ряда папоротников. В процессе изучения папоротни- Pteridoideae. Снимки спор в статьях A.J. Belling и ков (так сложилось исторически) в подсемейство C.J. Heusserl, а также B.K. Nayar с S. Devi пригодны Pteridoideae включали роды, которые не могли по ряду лишь для описания формы и снятия очень “грубых” причин обоснованно отнести к той или иной таксоно- замеров при биометрических исследованиях. По мической категории. снимкам, представленным в этих работах, невозмож- Так, в работе (Chen, Huang, 1974) по спорам папо- но выявить и описать подробно морфологические ротников семейства Pteridaceae с территории Формо- структуры, строение лезур, экваториальные складки, зы (Тайвань) из-за использования “сырой” и не дора- бугорки на сторонах спор и поверхность экзоспория. ботанной системы папоротников этого семейства, а Точное установление размеров всех структур спор и также неверного понимания его объема была оши- их описание необходимо для выявления общих при- бочно предложена классификация морфологии спор. знаков на межродовом, видовом и межсекционных В итоге для данного семейства разработана классифи- уровнях. Таким образом, в работах полувековой дав- кация с двумя типами спор – билатеральные и трех- ности описания спор, как и сами снимки, содержат лучевые тетраэдрические. Ввиду этого необходимо недостаточно информации о морфологии спор этой сравнить результаты наших анатомо-морфологичес- сложной группы папоротников и требуют современ- ких и молекулярно-генетических исследований с но- ного высокотехнологичного оборудования и дополне- выми сведениями по систематике той или иной таксо- ния новыми данными. номической группы. Однако обзорные работы по морфологии спор Кроме сведений, получаемых методами молеку- папоротников семейства Pteridaceae позволяют все же лярной генетики для выяснения филогенетических получить общие сведения по ряду стабильных при- границ родов, и установления более четких различий знаков, характерных подсемейству Pteridoideae: тетра- между ними, в последние годы часто применяют ре- эдрические споры с трехлучевой лезурой, округло- зультаты микрофотографирования сканирующим треугольные или треугольно-округлые, они имеют микроскопом. Микроснимки спор более информа- четкий экзоспорий с ярко выраженной скульптурой тивны, они несут больше сведений для решения спор- поверхности и полное отсутствие периспория, по- ных вопросов в систематике папоротников сложных верхность экзоспория от шероховатой до крупных групп. бугорков и валиковидных утолщений (Belling, Heus- Морфологии спор отдельных представителей serl, 1957; Nayar, Devi, 1966, 1967; Chen, Huang, 1974; подсемейства Pteridoideae в свое время уделялось до- Wagner, 1974). МАТЕРИАЛ И МЕТОДЫ ИССЛЕДОВАНИЯ Материалом для исследования послужили образ- Все образцы спор исследовали в режиме высоко- цы спор, отобранные с гербарного материала в Герба- го вакуума. Сканирование образцов осуществляли рии Ботанического института имени В.Л. Комарова при увеличении от 400 (общий вид спор) до 14 000 РАН (г. Санкт-Петербург). Для исследования было (поверхность экзоспория). отобрано 6 образцов по 6 видам подсемейства Pteri- При анализе спор использовали следующие doideae с территории Центральной и Восточной Азии, морфологические признаки: 1 – экваториальный а также Южной Америки. Изучения спор проводили диаметр, мкм; 2 – полярная ось; 3 – длина лезуры; в лаборатории Института водных и экологических 4 – ширина лезуры; 5 – ширина экваториальной проблем (ИВЭП СО РАН, г. Барнаул) на сканирующем складки; 6 – ширина валиковидного утолщения; 7 – электронном микроскопе Hitachi-S 3400 N фирма Hi- толщина валиковидной складки на дистальной сто- tachi High-Technologies Corp. и на микроскопе JEOL роне споры; 8 – толщина валиковидной складки на JSM-6390LA Analytical Scanning Electron Microscope в проксимальной стороне споры; 9 – ширина вали- Центре коллективного пользования БИН РАН. Спо- ковидного утолщения около лезуры; 10 – диаметр ры наносили на двусторонний скотч, крепили на ме- бугорков на проксимальной стороне споры; 11 – таллические предметные столики диаметром 10 мм. Поверхность спор обрабатывали золотопалладие- диаметр выростов на проксимальной стороне споры; вой смесью в вакуумной напылительной установке 12 – диаметр выростов на дистальной стороне споры; Emitech SC 7620/QT S около 6 мин, а в напылительной 13 – ширина валиковидного утолщения по контуру установке JEOL SPUTTER JFC-1100 – 7–10 мин. дистальной стороны споры; 14 – ширина валико- 30 видного утолщения по контуру проксимальной споры. Измерения проводили в 25-кратной повтор- стороны споры; 15 – поперечная складка по углам ности. РЕЗУЛЬТАТЫ И ОБСУЖДЕНИЕ Ниже приводятся описания и оригинальные мик- стороне споры находятся бугорки, слегка приподни- рофотографии спор видов из подсемейства Pteridoi- мающиеся над поверхностью спор диаметром 2.02 (от deae. 1.2 до 2.7) мкм. Ширина экваториальной складки 6.03 1. Jamesonia auriculata A.F. Tryon (рис. 1, а–г). Спо- (от 4.6 до 7.6) мкм. По обе стороны от экваториальной ры в проксимально- и дистально-полярном положе- складки с проксимальной стороны валиковидная ниях округло-треугольные, лопастные. Экваториаль- складка по всему контуру составляет 7.16 (от 5.2 до ный диаметр 42.38 (от 40.9 до 44.6) мкм. Полярная ось 8.4) мкм шир., а с дистальной – 7.52 (от 6.5 до 8.3) мкм 25.7 (от 23.4 до 28.1) мкм. В экваториальном положе- шир. Бугорки
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