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Систематика, Филогения И Эволюция Гриллоблаттидовых Насекомых (Insecta: Grylloblattida)

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Систематика, Филогения И Эволюция Гриллоблаттидовых Насекомых (Insecta: Grylloblattida) РОССИЙСКАЯ АКАДЕМИЯ НАУК ДАЛЬНЕВОСТОЧНОЕ ОТДЕЛЕНИЕ БИОЛОГО–ПОЧВЕННЫЙ ИНСТИТУТ С. Ю. СТОРОЖЕНКО СИСТЕМАТИКА, ФИЛОГЕНИЯ И ЭВОЛЮЦИЯ ГРИЛЛОБЛАТТИДОВЫХ НАСЕКОМЫХ (INSECTA: GRYLLOBLATTIDA) ВЛАДИВОСТОК ДАЛЬНАУКА 1998 УДК 595.72 Стороженко С. Ю. Систематика, филогения и эволюция гриллоблаттидовых насе- комых (Insecta: Grylloblattida). – Владивосток: Дальнаука, 1998. – 207 с. Настоящая книга посвящена исследованию одного из наименее изученных отрядов насекомых в объеме мировой фауны. Основное содержание монографии условно разделяется на две части. В общей части рассматриваются история изучения гриллоблаттидовых насекомых, использованные материалы и методы, особенности морфологии, анатомии, биологии, экологии, геологического и географического распространения, предложена классификация отряда и показано его место в системе класса насекомых, на основе кладистического анализа разработана схема филогенеза и выявлены особенности эволюции гриллоблаттидовых. В систематической части книги даны определительные таблицы 45 семейств, 169 родов и 320 видов гриллоблаттидовых, описанных до 1997 г. включительно; для всех таксонов приведена полная синонимия. Книга предназначена для энтомологов, палеонтологов, систематиков–зоологов, преподавателей и студентов биологических и геологических факультетов высших учебных заведений. Библ. 337, илл. 487, табл. 1. Storozhenko S. Yu. Systematics, phylogeny and evolution of the grylloblattids (Insecta: Grylloblattida). – Vladivostok: Dal'nauka, 1998. – 207 p. The book treats one of the little-known insects order Grylloblattida. The book consists of two large parts. In general part the history of study of these insects, materials and methods of investigation, morphology, anatomy, biology, ecology, geological and geographical distribution of grylloblattids, or ice crawlers of the world are shortly considered. The system of the order Grylloblattida is proposed and the relationships between them and other insects orders are discussed. Based on cladistic analysis the phylogeny and evolution of grylloblattids are studied. In systematic part the keys to 45 families, 169 genera and 320 species (known up to 1997) as well as synonymy for all taxa are given. The book is introduced for entomologist, palaeontologist, systematic zoologist, teachers and students of the biological and geological departments of universities and colleges. Bibl. 337, ill. 487, tabl. 1. На обложке – Grylloblattina djakonovi kurentzovi (фото Ю.А. Чистякова) Ответственный редактор А. С. Лелей Рецензенты: В. В. Богатов, Е. А. Макарченко Издание книги осуществлено при поддержке Российского фонда фундаментальных исследований по проекту № 98–04–62021 ISBN 5–7442–0899–2. С.Ю.Стороженко, 1998 г Дальнаука, 1998 г. 3 ВВЕДЕНИЕ Гриллоблаттидовые (Grylloblattida) – небольшой отряд насекомых, известный с карбона до наших дней, до последнего времени оставался одним из наименее исследо- ванных отрядов полинеоптерного комплекса. Система и объем отряда существенно изменялись в последние десятилетия. Гриллоблаттидовые являются центральной группой надотряда веснянкообразных (Plecopteroidea), от которой произошли остальные формы. Поэтому выяснение филогенеза этого отряда и разработка его системы крайне важны для понимания филогенеза как веснянкообразных, так и всех полионеоптер. Изучение гриллоблаттид важно и в общебиологическом плане в связи с черезвычайно замедленным жизненным циклом и устойчивостью к низким температурам у рецентных видов, а также сохранением существенных элементов протометаболии у ряда ископае- мых форм. Сведения по морфологии и биологии гриллоблаттидовых разбросаны в обширной литературе, а вопросы филогении, классификации, геологического и географи- ческого распространения стали разрабатываться лишь в последнее время. Целью настоящей работы является разностороннее изучение отряда гриллоблат- тидовых с особым вниманием к филогении, классификации, биологии, экологии и распространению этих насекомых. Для этого было необходимо выяснить объем отряда и его положение в системе насекомых, разработать систему отряда (включая как ископаемые так и рецентные формы), выяснить филогенитические связи семейств, наметить основные тенденции в морфологической и экологической эволюции, собрать, дополнить и обобщить сведения о морфологии, образе жизни, биологии, экологии и распространении гриллоблаттидовых, ревизовать современных и ископаемых гриллоблаттид, составить определительные таблицы и аннотированный каталог семейств, родов и видов гриллоблаттидовых мировой фауны. Я искренне признателен В.В. Жерихину, А.Г. Пономаренко, А.П. Расницыну, Ю.А. Попову, Н.Д. Синиченковой, И.Д. Сукачевой (Палеонтологический институт РАН, Москва) и В.Г. Новокшонову (Пермский государственный университет) за любезно предоставленную возможность изучить коллекции ископаемых гриллоблаттидовых и ценные советы и замечания, А.В. Горохову (Зоологический институт РАН, Санкт– Петербург), А.С. Лелею (Биолого–почвенный институт ДВО РАН, Владивосток), H. Ando и T. Nagashima (Sugadaria Rasearch Center, Japan), с которыми обсуждались предварительные результаты, а также А.Ф. Емельянову (Зоологический институт РАН, Санкт–Петербург) и А.К. Бродскому (Сенкт–Петербургский государственный университет) за ценные критические замечания по рукописи. Я благодарен T. Yamasaki (Tokyo Metropolitan University), D. Otte (Academy of Natural Science of Philadelphia), R. Nickle (U.S. Natural Museum, Washington), за помощь во время изучения типовых материалов по современным гриллоблаттидам из США и Японии. Работа по обобщению литературных данных, ревизии мировой фауны и составлению каталога стала возможной благодаря помощи со стороны приславших мне оттиски или ксерокопии отсутствующих в России статей зарубежных коллег: S. Asahina, B. Baccetti, G. Fukusima, D.K.McE. Kevan, J. Kukalova–Peck, J. Namkung, D.C.F. Rentz, A. Szeptycky, V. R. Vickery и Chu– Yong Wang. Я благодарен С.К. Холину и П.Г. Немкову (Биолого–почвенный институт ДВО РАН, Владивосток) за помощь при математической обработке данных по распро- странению гриллоблаттидовых. Неоценимую помощь при проведении полевых исследований оказали сотрудники лаборатории энтомологии Биолого–почвенного инсти- тута ДВО РАН: В.Н. Макаркин, А.А. Мещеряков и другие. Я благодарен всем, кого не смог упомянуть здесь, за содействие в моей работе. 4 Глава 1. ИСТОРИЯ ИЗУЧЕНИЯ 1.1. История изучения современных гриллоблаттид Первый современный представитель отряда Grylloblattida был описан из провинции Альберта в Канаде и сразу привлек внимание энтомологов своеобразным сочетанием морфологических признаков, характерных для других отрядов насекомых, о чем свидетельствовало название типового вида - Grylloblatta campodeiformis (Walker, 1914). Вскоре после этого гриллоблаттиды были найдены в Японии (Caudell, King, 1924). Затем новые роды, виды и подвиды гриллоблаттид были описаны из Канады, США, Японии, Кореи, Китая и России (Caudell, 1924a; 1924b; Gurney, 1937; 1948; 1953; 1961; Kamp, 1963; 1979; Silvestri, 1927; 1931; Бей-Биенко, 1951; Asahina, 1959; 1961; Namkung, 1974а; 1974b; Правдин, Стороженко, 1977; Стороженко, Олигер, 1984; Szeptycki, 1987; Wang, 1987a; Стороженко, 1988а; 1996c). Общие обзоры распространения, морфологии, биологии современных гриллоблаттид и библиографии по ним содержаться в работах ряда авторов (Comstok, 1920; Imms, 1925; Crampton, 1927; Brues, Melander, 1932; Crampton, 1933; Walker, 1937; Gurney, 1937; 1948; 1953; Kamp, 1953; Brues et al., 1954; Imms, 1957; Kamp, 1963; Бей-Биенко, 1964; 1966; Fukushima, 1969a; 1969b; Key, 1970; Asahina, 1971; Yamasaki, 1971; Kamp, 1979; Ando et al., 1979; Kevan, 1979; Rentz, 1982; Namkung, 1982; Yamasaki, 1982a; Brown, 1982; Vickery, Kevan, 1983), а определительные таблицы родов и видов можно найти в ряде работ (Gurney, 1937; 1953; Правдин, Стороженко, 1977; Vickery, Kevan, 1983; Стороженко, 1986б; 1995). Автором был составлен аннотированный каталог, проведена ревизия семейства Grylloblattidae, на родовом уровне рассмотрена филогения и выявлены особенности формирования рецентной фауны (Стороженко, 1986а; 1988а; 1996d). Всего к настоящему времени известны 4 рода и 25 видов, однако таксономический период в изучении современных гриллоблаттид еще не закончен и следует ожидать описаний новых форм, особенно из России, Японии, Кореи и Канады. Морфология имаго изучалась главным образом на примере Grylloblatta campodei- formis и Galloisiana nipponensis. Лейтмотивом морфологических исследований являлось выяснение места гриллоблаттид в системе насекомых. Отмечено, что голова и ее придатки у Grylloblattidae - генерализованного типа, за исключением частично или полностью редуцированных глаз и отсутствия глазков (Crampton, 1917a; 1917c; 1923; 1926a; 1927; Walker, 1931; Crampton, 1933; Walker, 1933; Koyama, 1952; Matsuda, 1965; Fukushima, 1967; Nagashima, 1982a). Строение цервикальных склеритов позволяет сближать гриллоблаттид с эмбиями и уховертками (Crampton, 1926b). Морфология груди привлекала внимание ряда исследователей, отмечавших, что гомономность отделов груди связана с вторичной бескрылостью этих насекомых (Crampton, 1915; 1917a; 1926b; 1927; 1933; Walker, 1938; Fukushima, 1967; Rahle, 1970; Matsuda, 1970). Строение ног гриллоблаттид позволяло противопоставлять их собственно прямокрылым насекомым (Crampton, 1927, 1933; Walker, 1937; Fukushima, 1967). В строении брюшка гриллоблаттид отмечены ряд примитивных черт: отсутствие слияния 1-го стернита брюшка с заднегрудью, наличие отчлененных коксоподитов у самца и др. (Walker, 1919a; 1919b; Crampton, 1927; 1933; Walker, 1943; Matsuda, 1976; Fukushima, 1967). Асимметричные гениталии самца
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