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(Makarov Bokhovko).Pmd Russian Entomol. J. 14(4):263–274 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2005 Ïðååìñòâåííîñòü ñòðóêòóð õåòîìà ó ðàçâèâàþùèõñÿ ñ ãèïåð- ìåòàìîðôîçîì ëè÷èíîê Brachinus Weber (Coleoptera: Carabidae) Continuity of chaetom pattern in Brachinus-larvae developing with hypermetamophoSiS (Coleoptera: Carabidae) Ê.Â.Ìàêàðîâ1, Å.E.Áîõîâêî2 K.V.Makarov1, E.E.bokhovko2 1 Московский педагогический государственный университет, кафедра зоологии и экологии, ул. Кибальчича 6, корп. 5, Москва 129278, Россия. 2 Московский государственный университет им. М.В. Ломоносова, rафедра энтомологии, Москва 119992, Воробьёвы горы, Россия. 1 Moscow State Pedagogical University, Department of Zoology & Ecology, Kibaltschitcha str. 6, build. 5, Moscow 129278, Russia. 2 Moscow Lomonosov State University, Department of Entomology, Moscow 119992, Vorobiovy Gory, Russia. KEY WORDS: larvae, chaetotaxy, morphology, adaptations, Coleoptera, Carabidae, Brachinini. КЛЮЧЕВЫЕ СЛОВА: личинки, хетотаксия, морфология, адаптации, Coleoptera, Carabidae, Brachinini. ABSTRACT. Extrenal morphology and the chaeto- риты и плевриты гипертрофированы. Редукция и им- taxy of all larval stages of carabid beetle Brachinus мобилизация отделов конечностей не влияет на со- elegans Chaudoir, 1842 were studied. First larva of this став их хетома. Анализ хетотаксии позволил выделить beetle is triangulina-like, whereas larvae II and III are по меньшей мере две тенденции развития хетома в engorged, scarcely moving forms. In spite of hyperme- онтогенезе Brachinus: ослабление сенсорных и локо- tamorphosis, the great continuity of chaetome compo- моторных структур и развитие пассивно-опорного sition was revealed. The analysis of the topology of хетома абдоминальных вентритов и плевритов. Оцен- chaetome elements during larval development allowed ка трансформации адаптивных структур позволила to ascertain follows: 1) the strong modification of the провести отбор таксономически значимых призна- cephalic capsule is caused but the disproportional en- ков и подтвердить гипотезу о близости Brachinini к largement of basal regions of frontal and parietal scleri- группе Truncatipenne в составе Limbata conchifera. tes; 2) body tergits increase in size proportionally; 3) pleurites and ventrites become hypertrophied; 4) the Триба Brachinini — своеобразная группа жуже- reduction and immobilization of leg segments have not лиц с неясным систематическим положением. Нео- effect on the set of their chetome elements. There are бычные черты строения и образа жизни имаго по- two the most important tendencies in chaetome forma- служили причинной неоднократного изучения мор- tion in Brachinus larvae during the ontogenesis: the фологии и физиологии этой группы [Aneshanelsey weakening of sensorial and locomotory elements and et al., 1969; Eisner, 1959; Galián et al., 1990], а также the development of the supporting chaetome of abdo- обсуждения её родственных связей и таксономичес- minal ventrites and pleurites. The assessment of the кого статуса. Ранг этого таксона многократно ме- transformation of different adaptive and nonadaptive нялся — от уровня подтрибы до подсемейства и structures enabled to select the taxonomically valuable даже семейства, а в разных классификациях Brachinini features. Their application supports the hypothesis of сближались то с наиболее плезиоморфными груп- the proximity of Brachinini to the group Truncatipenne пами, то с наиболее продвинутыми [Крыжановский, within Limbata conchifera. 1983; Ball, 1979; Deuve, 1993; Ball et al., 1998]. Проблематичность этих трактовок усугубляется РЕЗЮМЕ. Описана морфология и хетотаксия ли- особенностями биологии Brachinini. Уже в конце чинок старших возрастов Brachinus elegans Chaudoir, XIX века были обнаружены личинки старших воз- 1842 показана высокая преемственность структур хе- растов Brachinus Weber, 1801, паразитировавшие тома в ходе гиперметаморфоза. Сравнение тополо- на куколках водных жуков [Wickham, 1893; цит. по: гии гомологичных элементов хетома позволило уста- Dimmok & Knab, 1904]. На основании аналогий в новить, что изменение формы головной капсулы оп- образе жизни личинок для Brachinus предполага- ределяется неравномерным разрастанием основания лось развитие с гиперметаморфозом и подвижной фронтального и париетальных склеритов, тергиты тела личинкой первого возраста. Позднее были описа- пропорционально увеличиваются в размерах, а вент- ны личинки первого возраста родов Pheropsophus 264 К. В. Макаров, Е. Е. Боховко Solier, 1833 [Boldori, 1939; van Emden, 1919, 1920], и ку рода Brachinus и онтогенетические изменения Brachinus [van Emden, 1942; Habu & Sadanaga, 1965; хетома на примере нескольких видов. Wautier, 1963, 1964], которые не питались и вели Основой для работы послужили личинки перво- себя подобно триунгулинами Meloidae. Полностью го возраста, выведенные в лаборатории в разные цикл развития и строение личинок старших возрас- годы от имаго Brachinus crepitans и Brachinus тов североамериканского Brachinus pallidus Erwin, plagiatus Reiche, 1868. Личинки старших возрастов 1965 были изучены Ирвином [Erwin, 1966, 1967]. В Brachinus elegans Chaudoir, 1842 были собраны в дальнейшем биология этого и нескольких близких 2004 г. в Краснодарском крае Е.Е. Боховко. Материал видов была изучена очень подробно [Juliano, 1983, хранится в коллекции кафедры зоологии и экологии 1984, 1985a, b, 1986a, b]. Оказалось, что их метамор- Московского педагогического государственного фоз удлинён и насчитывает пять личиночных ста- университет (МПГУ) в 70% этиловом спирте, часть дий, при этом старшие возраста малоподвижны, личинок для исследования хетотаксии заключена в обладают редуцированными конечностями и орга- препараты с жидкостью Фора-Берлезе или эупара- нами чувств и питаются куколками водных жуков лом.* Номенклатура хетома приведена по общепри- (Hydrophilidae, Gyrinidae), разыскиваемых триунгу- нятой схеме [Bousquet & Goulet, 1984]. линами. Эти данные, вместе с краткими наблюде- ниями о развитии Pheropsophus jesoensis A.Mora- Морфология личинок witz, 1862 [Habu & Sadanaga, 1965], долгое время оставались единственными сведения о биологии Личинка первого возраста личинок Brachinini и вызывали немало сомнений, поскольку местообитания многих Brachinus никак ÌÀÒÅÐÈÀË. Brachinus crepitans: Þ Àçåðáàéäæàí, Òàëûø, áåðåã ð. Èñòèñó÷àé, 1.II.1988 leG. Â.Äóøåíêîâ, âûâåäåíû не связаны с водоёмами. Развитие европейских 23.III.2988, Í.Êîçëîâ — 11L1 (¹43.2–1.1 F), 28L1 (¹43.2– Brachinus crepitans (Linné, 1758) и B. explodens 1.2 Al); Þ Àçåðáàéäæàí, Ëåíêîðàíñêèé ð-í, îêð. ï.Ìèêîëàí, (Duftschmid, 1812) было изучено лишь в последние áåðåã ð.Ëåíêîðàí÷àé 23.IV.2004 leG. Ê.Ìàêàðîâ, âûâåäåíû годы [Saska & Honek, 2004]. При этом выяснилось, 13.V.2004, À.Ìàòàëèí — 3L1 (¹43.2–1.3 Al), 1L1 (¹43.2– 1.4 Eu); breeding by F.I. van Emden from adults: Croydon, Surrey, что пищей для личинок старших возрастов служат leG. R.L. Coi[.] — 1L1 (¹43.2–1.5 F); Brachinus plagiatus: куколки других видов жужелиц. Ìîëäàâèÿ, Êàãóëüñêèé ð-í, îêð. ñ.Ðîøó; âûâåäåíû â ëàáîðàòîðèè Такие особенности образа жизни сильно ослож- 19.VI.1984 leG. Â.Êàðïîâà — 1L1 (¹43.2–2.1 F). няют изучение преимагинальных стадий Brachinini. ОПИСАНИЕ. Мелкие, камподеовидные, депигменти- К настоящему времени есть данные по морфоло- рованные, безглазые личинки с относительно короткими гии личинок первого возраста шести европейских ротовыми придатками и урогомфами. [Wautier, 1963, 1964] и одного японского вида Окраска светлая, лишь голова и мандибулы коричне- вато-желтые, остальные части тела едва пигментированы, Brachinus [Habu & Sadanaga, 1965], двух видов часто склериты брюшка едва отличимы от окружающей Aptinus Bonelli, 1810 [Wautier & Viala, 1967; Hovorka, их кутикулы. 1996] и нескольких палеотропических Pheropsophus Микроскульптура в виде чешуеобразных кутику- [van Emden, 1919, 1920; Boldori, 1939; Habu, 1986; лярных шипиков развита на боковых поверхностях парие- Habu & Sadanaga, 1965; Rajagopal & Kumar, 1993; тальных склеритов, в латеральных частях тергитов груди Qu, 1996]. Именно признаки личинок первого возра- и, особенно, брюшка. Вероятно, они выполняют функ- ста используются и в определительных таблицах цию яйцевых зубчиков, отсутствующих у личинок [Bøving & Craighead, 1931; van Emden, 1942; Шарова, Brachinus. Кроме того, отдельные шипики развиты на 1958, 1964; Hůrka, 1978; Arndt, 1991; Luff, 1993; поверхности стипеса, прементума, тазиков ног и X сег- Makarov, 1994]. Известны краткие описания личи- мента брюшка. Голова (Рис. 1, 2) слегка удлинённая, параллельнос- нок старших возрастов B. pallidus [Erwin, 1967] и Ph. торонняя, без выраженных борозд, глаза отсутствуют, jesoensis [Habu & Sadanaga, 1965; Habu, 1986], при- антеннальное кольцо редуцировано. По литературными чём в последнем случае авторы использовали и данным, личинки I возраста ряда видов Brachinus облада- некоторые данные по хетотаксии. ют более или менее развитыми глазными пятнами или Однако детали морфологии личинок Brachinini глазами [Wautier & Viala, 1964; Erwin, 1967]. Фронталь- почти не изучены, неполные сведения о хетотаксии ные швы едва изогнуты, кпереди сильно утончаются, имеются только для личинки Aptinus bombarda эпикраниальный шов очень короткий. Назале без зубцов, (Illiger, 1880) [Hovorka, 1996]. Хетотаксия и детали лишь намечено в виде слабо выступающей широкой ок- строения личинок старших возрастов остаются прак- руглой лопасти (Рис. 3). тически не изученными. В связи с этим значитель- Хетотаксия головы характеризуется отсутствием до- полнительных щетинок и редукцией ряда сенсилл генера- ный интерес представляет как подробное описание лизованного набора
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