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

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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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    RECOMMENDED INSECT CLASSIFICATION FOR UGA ENTOMOLOGY CLASSES (2020) In an effort to standardize the hexapod classification systems being taught to our students by our faculty in multiple courses across three UGA campuses, I recommend that the Entomology Department adopts the basic system presented in the following textbook: Triplehorn, C.A. and N.F. Johnson. 2005. Borror and DeLong’s Introduction to the Study of Insects. 7th ed. Thomson Brooks/Cole, Belmont CA, 864 pp. This book was chosen for a variety of reasons. It is widely used in the U.S. as the textbook for Insect Taxonomy classes, including our class at UGA. It focuses on North American taxa. The authors were cautious, presenting changes only after they have been widely accepted by the taxonomic community. Below is an annotated summary of the T&J (2005) classification. Some of the more familiar taxa above the ordinal level are given in caps. Some of the more important and familiar suborders and families are indented and listed beneath each order. Note that this is neither an exhaustive nor representative list of suborders and families. It was provided simply to clarify which taxa are impacted by some of more important classification changes. Please consult T&J (2005) for information about taxa that are not listed below. Unfortunately, T&J (2005) is now badly outdated with respect to some significant classification changes. Therefore, in the classification standard provided below, some well corroborated and broadly accepted updates have been made to their classification scheme. Feel free to contact me if you have any questions about this classification.
  • Auditory and Vibratory Sense of Crickets Application Note

    Auditory and Vibratory Sense of Crickets Application Note

    Technical Paper Auditory and Vibratory Sense of Crickets Polytec Technical Papers A Aerospace B Audio & Acoustics C Automotive Development D Data Storage G General Vibrometry M Microstructure Testing P Production Testing S Scientific & Medical T Structural Testing U Ultrasonics The Origin of Sound-Processing Elements in Ensifera Using Laser Vibrometry In the life of insects, vibrational signals mediate important information that is used in various contexts, from pair formation to detection of predators or finding prey. Therefore, insects are equipped with both extremely sensitive receptor organs in the legs for detection of substrate vibrations and the underlying neural network enabling recog- nition and localization of the signallers in a complex environment. Without the use of special equipment to detect those signals, the intriguing world of insect vibrations would remain hidden to humans, which mostly communicate by sight and sound. Introduction Insects in the group called Ensifera produce sounds for com- munication. These insects, which include crickets and bush crickets, have evolved ears in their legs from the pre-existing vibratory organs. In our study, we investigated whether their auditory and vibratory sense may share a common origin at the level of the central nervous system. Therefore, we studied the network of vibration-sensitive interneurons in a primitive, deaf cave cricket (Fig. 1) and compared our results with well known results for auditory neurons of hearing Ensifera. Fig. 1: The cave-dwelling cricket Troglophilus neglectus (Ensifera, Rhaphidophoridae) has organs specialized for detection of substrate vibrations located below the “knee” joint of all legs. Advancing Measurements by Light www.polytec.com Experimental Set-up In our experiments, the first thoracic segment of the nerve chord in the cave cricket was penetrated by intracellular record- ing electrodes, while the insect’s front legs were vibrated by sinusoidal pulses delivered by two mini shakers.
  • EPPO Reporting Service

    EPPO Reporting Service

    ORGANISATION EUROPEENNE EUROPEAN AND MEDITERRANEAN ET MEDITERRANEENNE PLANT PROTECTION POUR LA PROTECTION DES PLANTES ORGANIZATION EPPO Reporting Service NO. 10 PARIS, 2012-10-01 CONTENTS _______________________________________________________________________ Pests & Diseases 2012/203 - First report of Anthonomus eugenii in the Netherlands 2012/204 - First report of Aromia bungii in Italy 2012/205 - Drosophila suzukii continues to spread in Europe 2012/206 - First report of Drosophila suzukii in Germany 2012/207 - First report of Drosophila suzukii in Croatia 2012/208 - First report of Drosophila suzukii in the United Kingdom 2012/209 - First report of Drosophila suzukii in Portugal 2012/210 - First report of Drosophila suzukii in the Netherlands 2012/211 - Situation of Drosophila suzukii in Belgium 2012/212 - First report of Aculops fuchsiae in Belgium 2012/213 - First report of Carpomya incompleta in France 2012/214 - Outbreaks of Bemisia tabaci in Finland 2012/215 - Update on the situation of Diabrotica virgifera virgifera in the Czech Republic 2012/216 - Synchytrium endobioticum no longer found in Northern Ireland, United Kingdom 2012/217 - Outbreak of Anacridium melanorhodon arabafrum in Qatar 2012/218 - Ralstonia solanacearum detected in the Czech Republic 2012/219 - First report of „Candidatus Liberibacter solanacearum‟ on carrots in France, in association with Trioza apicalis 2012/220 - Occurrence of „Candidatus Phytoplasma pyri‟ is confirmed in Belgium 2012/221 - „Syndrome des basses richesses‟ detected in Germany: addition to
  • The Insects: Structure and Function: Fifth Edition R

    The Insects: Structure and Function: Fifth Edition R

    Cambridge University Press 978-0-521-11389-2 - The Insects: Structure and Function: Fifth Edition R. F. Chapman Edited by Stephen J. Simpson and Angela E. Douglas Frontmatter More information The Insects Structure and Function FIFTH EDITION The Insects has been the standard textbook in the field since the first edition was published over 40 years ago. Building on the strengths of Chapman’s original text, this long-awaited new edition has been revised and expanded by a team of eminent insect physiologists, bringing it fully up to date for the molecular era. The chapters retain the successful structure of the earlier editions, focusing on particular functional systems rather than on taxonomic groups and making it easy for students to delve into topics without extensive knowledge of taxonomy. The focus is on form and function, bringing together basic anatomy and physiology and examining how these relate to behavior. This, combined with nearly 600 clear illustrations, provides a comprehensive understanding of how insects work. Now also featuring a richly illustrated prologue by George McGavin, this is an essential text for students, researchers and applied entomologists alike. R. F. Chapman (1930–2003) was an eminent insect physiologist and Professor in the Division of Neurobiology at the University of Arizona. His first four editions of The Insects have formed the standard text in the field for more than 40 years. Stephen J. Simpson is ARC Laureate Fellow in the School of Biological Sciences and Academic Director of the Perkins Centre for the study of obesity, diabetes and cardiovascular disease at the University of Sydney.
  • Bryophyte Ecology Table of Contents

    Bryophyte Ecology Table of Contents

    Glime, J. M. 2020. Table of Contents. Bryophyte Ecology. Ebook sponsored by Michigan Technological University 1 and the International Association of Bryologists. Last updated 15 July 2020 and available at <https://digitalcommons.mtu.edu/bryophyte-ecology/>. This file will contain all the volumes, chapters, and headings within chapters to help you find what you want in the book. Once you enter a chapter, there will be a table of contents with clickable page numbers. To search the list, check the upper screen of your pdf reader for a search window or magnifying glass. If there is none, try Ctrl G to open one. TABLE OF CONTENTS BRYOPHYTE ECOLOGY VOLUME 1: PHYSIOLOGICAL ECOLOGY Chapter in Volume 1 1 INTRODUCTION Thinking on a New Scale Adaptations to Land Minimum Size Do Bryophytes Lack Diversity? The "Moss" What's in a Name? Phyla/Divisions Role of Bryology 2 LIFE CYCLES AND MORPHOLOGY 2-1: Meet the Bryophytes Definition of Bryophyte Nomenclature What Makes Bryophytes Unique Who are the Relatives? Two Branches Limitations of Scale Limited by Scale – and No Lignin Limited by Scale – Forced to Be Simple Limited by Scale – Needing to Swim Limited by Scale – and Housing an Embryo Higher Classifications and New Meanings New Meanings for the Term Bryophyte Differences within Bryobiotina 2-2: Life Cycles: Surviving Change The General Bryobiotina Life Cycle Dominant Generation The Life Cycle Life Cycle Controls Generation Time Importance Longevity and Totipotency 2-3: Marchantiophyta Distinguishing Marchantiophyta Elaters Leafy or Thallose? Class
  • Karyotype of Karoophasma Biedouwense (Austrophasmatidae)

    Karyotype of Karoophasma Biedouwense (Austrophasmatidae)

    Eur. J. Entomol. 112(4): 599–605, 2015 doi: 10.14411/eje.2015.093 ISSN 1210-5759 (print), 1802-8829 (online) First chromosomal study of Mantophasmatodea: Karyotype of Karoophasma biedouwense (Austrophasmatidae) DOROTA LACHOWSKA-CIERLIK1, ANNA MARyAńSKA-Nadachowska2, VALENTINA KuZNETSOvA3 and MIKE PICKER4 1 Institute of Zoology, Jagiellonian university, Cracow, Poland; e-mail: [email protected] 2 Institute of Systematic and Evolution of Animals, PAS, Cracow, Poland; e-mail: [email protected] 3 Zoological Institute Russian Academy of Sciences, St. Petersburg, Russia; e-mail: [email protected] 4 Department of Biological Sciences, university of Cape Town, Rondebosch, Cape Town, South Africa; e-mail: [email protected] Key words. Mantophasmatodea, Karoophasma biedouwense, FISH, heterochromatin, heelwalkers, meiosis, ribosomal genes, sex determination system, telomeres Abstract. We have investigated for the first time the chromosomes of Karoophasma biedouwense, a species belonging to the Manto- phasmatodea, a recently discovered order of carnivorous insects. Our study has revealed that males of this species display testes with numerous seminal tubes (follicles), as in other Polyneoptera, and short tubular seminal vesicles embedded in a utricular gland. The karyotype consists of 2n = 12A + X monocentric and biarmed, meta/submetacentric chromosomes (fundamental number of arms: FN = 26) with blocks of heterochromatin around centromeres. The autosomes are classified into two size groups, one represented by a single, very large pair of autosomes, the other by five smaller pairs which constitute a continuous series gradually decreasing in size. Among “monocentric” orders of Polyneoptera, K. biedouwense shares its low chromosome number, 2n = 13, as also found with some Orthoptera (Acridoidea, Grylloidea, Gryllacridoidea).