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Юрские Сетчатокрылые (Insecta: Neuroptera) Центральной Азии

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Юрские Сетчатокрылые (Insecta: Neuroptera) Центральной Азии РОССИЙСКАЯ АКАДЕМИЯ НАУК ПАЛЕОНТОЛОГИЧЕСКИЙ ИНСТИТУТ им. А.А. Борисяка на правах рукописи Храмов Александр Валерьевич ЮРСКИЕ СЕТЧАТОКРЫЛЫЕ (INSECTA: NEUROPTERA) ЦЕНТРАЛЬНОЙ АЗИИ 25.00.02 Палеонтология и стратиграфия Диссертация на соискание ученой степени кандидата биологических наук Научный руководитель: доктор биологических наук Пономаренко Александр Георгиевич Москва - 2014 Оглавление ВВЕДЕНИЕ............................................................................................................................. стр. 4 Глава 1. История изучения юрских Neuroptera................................................................стр.7 Глава 2. Отряд Neuroptera..................................................................................................стр. 11 2.1. Система и биология современных Neuroptera....................................................... стр. 11 2.2. Строение крыльев и номенклатура жилкования Neuroptera............................. стр. 14 2.3. Палеонтологическая летопись Neuroptera.............................................................. стр. 17 Глава 3. Материалы и методы.......................................................................................... стр. 31 3.1. Коллекции юрских Neuroptera и их обработка...................................................... стр. 31 3.2. Описание местонахождений юрских Neuroptera Центральной Азии................ стр. 32 Глава 4. Обзор фаун юрских Neuroptera Центральной Азии..................................... стр. 42 4.1. Согюты (Киргизия)...................................................................................................... стр. 42 4.2. Сай-Сагул (Киргизия)................................................................................................. стр. 43 4.3. Саук-Таньга (Киргизия)............................................................................................. стр. 48 4.4. Ошин-Боро-Удзюр-Ула (Монголия)......................................................................... стр. 50 4.5. Баян-Тэг (Монголия)................................................................................................... стр. 50 4.6. Бахар (Монголия) .........................................................................................................стр. 51 4.7. Хоутийн-Хотгор (Монголия)...................................................................................... стр. 52 4.8. Шар-Тэг (Монголия)................................................................................................... стр. 53 4.9. Каратау (Казахстан)..................................................................................................... стр. 54 Глава 5. Систематическая часть.......................................................................................стр. 79 Глава 6. Сравнительный анализ фаун юрских Neuroptera Центральной Азии....стр. 137 6.1. Neuroptera средней-верхней юры Центральной Азии и Китая......................... стр. 137 2 6.1.1. Количественный состав фаун Даохугоу и Каратау.......................................... стр. 137 6.1.2. Фауны Каратау, Даохугоу и монгольских местонахождений (Бахар, Хоутийн- Хотгор, Шар-Тэг).............................................................................................................. стр. 140 6.2. Юрская и среднеюрская-нижнемеловая когорты Neuroptera...........................стр. 141 6.3. Фауны юрских-нижнемеловых Neuroptera и климатический фактор............стр. 144 6.4. Список видов юрских Neuroptera Центральной Азии........................................ стр. 147 6.5. Распространение родов Neuroptera, найденных в юре Центральной Азии.... стр. 152 ЗАКЛЮЧЕНИЕ................................................................................................................. стр. 154 СПИСОК ЛИТЕРАТУРЫ............................................................................................... стр. 155 3 ВВЕДЕНИЕ Актуальность темы исследования. Сетчатокрылые (Neuroptera) - это небольшой отряд насекомых с полным превращением, насчитывающий в настоящеее время около 5 750 видов (Winterton et al., 2010) в составе 16 рецентных семейств, причем три из них, Chrysopidae, Hemerobiidae и Coniopterygidae, имеют хозяйственное значение как естественные враги сельскохозяйственных вредителей, таких как тли и червецы. До сих пор специалистам не удалось выработать консенсуса касательно филогении и системы данного отряда, первые находки которого приходятся на раннюю пермь. Для решения этой проблемы важно привлекать не только молекулярные данные и сведения по морфологии современных сетчатокрылых, но и ископаемый материал. Изучение юрских сетчатокрылых для этой цели представляется необходимым, поскольку в юре впервые начинают появляться представители рецентных семейств Neuroptera, а также возникают многие вымершие семейства, имевшие широкое распространение, например, Kalligrammatidae. Центральная Азия (в настоящей работе мы понимаем под Центральной Азией среднеазиатские республики бывшего СССР и Монголию) особенно богата местонахождениями юрских сетчатокрылых - по совокупному числу находок Neuroptera юрского возраста (около 900 экз.) Центральная Азия уступает лишь Китаю (более 2000 экз.) и значительно превосходит Европу (около 260 экз.) и Сибирь (около 20 экз.), из других регионов юрские сетчатокрылые неизвестны. До сих пор юрские сетчатокрылые Центральной Азии изучались лишь выборочно, а из целого ряда местонахождений сборы сетчатокрылых не были изучены вовсе. Тем не менее, максимально полный список найденных таксонов и данные о количественном составе фаун юрских сетчатокрылых Центральной Азии необходимы для сопоставления их с юрскими и меловыми фаунами сетчатокрылых из других регионов. Особенно важно такое сопоставление в свете изучения двух крупнейших мезозойских лагерштеттов, открытых в Китае в 1990-е годы - Даохугоу (бат-келовей) и Исяня (апт). В данных лагеррштеттах обнаружена богатейшая фауна насекомых, птерозавров, динозавров и млекопитающих. Сделанные там находки имеют ключевое значение для понимания эволюции целого ряда макротаксонов: например, из Даохугоу известны древнейшие плацентарные млекопитающие, из Исяня - древнейшие птицы и покрытосеменные растения. Поскольку в Даохугоу и Исяне найдены и активно изучаются богатые фауны сетчатокрылых, то их сравнение с юрскими сетчатокрылыми из местонахождений 4 Центральной Азии становится весьма актуальной задачей. Учитывая географическую близость Центральной Азии и Китая, такое сравнение имеет особый смысл для уточнения характеристик Даохугоу и Исяня, таких как возраст, тафономия и палеоклимат. Географически центральноазиатские местонахождения занимают промежуточное положение между китайскими и европейскими, причем для последних, поскольку они образованы морскими отложениями, известен точный возраст. Изучение сетчатокрылых наряду с другими насекомыми юры Центральной Азии помогло бы сопоставить верхнеюрское местонахождение Зольнгофен (Германия) с Даохугоу и Исянем и тем самым уточнить возраст и другие особенности китайских лагерштеттов. Анализ фауны юрских Neuroptera Центральной Азии может многое сказать и о самих центральноазиатских местонахождениях. Для этого необходимо принять во внимание все известные комплексы сетчатокрылых из юры и мела (их обзор приводится в разделе 1.3). Особый интерес представляет сравнение близковозрастных местонахождений Каратау и Даохугоу, откуда собраны самые обильные из имеющихся юрские фауны сетчатокрылых. Степень разработанности темы исследования. До сих пор из юрских местонахождений Центральной Азии было известно 44 рода и 54 вида сетчатокрылых, включая 7 видов, описанных по крайне фрагментарному материалу. Данных о количественном составе фаун юрских Neuroptera не существовало. Цель и задачи работы. Целью работы было изучение юрских сетчатокрылых, найденных на территории Центральной Азии. Для ее достижения были поставлены следующие задачи: - Охарактеризовать фауны сетчатокрылых из всех известных на сегодня юрских местонахождений Центральной Азии, их таксономическое разнообразие и количественный состав - Сопоставить юрские фауны сетчатокрылых Центральной Азии с юрскими и нижнемеловыми фаунами из других регионов Евразии - Выявить основные этапы изменений таксономического состава сетчатокрылых Центральной Азии в юрское время Научная новизна. Определен таксономический и количественный состав фаун сетчатокрылых из девяти центральноазиатских местонахождений, причем сетчатокрылые из пяти местонахождений (Сай-Сагул, Саук-Таньга, Шар-Тэг, Бахар и Хоутийн-Хотгор) изучены впервые. Также впервые предварительно охарактеризован количественный 5 состав крупнейшей фауны юрских Neuroptera из местонахождения Даохугоу (Китай). В ходе работы описано 16 родов и 33 вида юрских сетчатокрылых Центральной Азии, тем самым число видов юрских сетчатокрылых, известных из данного региона, увеличено в 1,5 раза. Ископаемое семейство Mesithonidae сининомизировано с рецентным Mantispidaе. Представители семейств Grammolingiidae и Saucrosmylidae впервые найдены за пределами типового местонахождения Даохугоу, что в совокупности с другими находками помогло связать его с монгольскими местонахождениями Бахар, Шар-Тэг и Хоутийн-Хотгор. Ряд таксонов юрских сетчатокрылых Центральной Азии исходя из возраста их первых и последних находок были сгруппированы в две когорты - юрскую и среднеюрскую- нижнемеловую. Теоретическая и практическая значимость. Полученные результаты могут быть использованы при реконструкции филогении отряда сетчатокрылых в целом и эволюции отдельных семейств. Данные о сетчатокрылых, найденных в центральноазиатских юрских
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