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(Prokin Et Al).Pmd Russian Entomol. J. 22(4): 259–274 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2013 New beetle larvae (Coleoptera: Coptoclavidae, Caraboidea, Polyphaga) from the Upper Triassic of Germany Íîâûå ëè÷èíêè æóêîâ (Coleoptera: Coptoclavidae, Caraboidea, Polyphaga) èç âåðõíåãî òðèàñà Ãåðìàíèè A.A. Prokin1,2, K.V. Makarov3 , A.G. Ponomarenko4 & A.S. Bashkuev4 A.A. Ïðîêèí1,2, Ê.Â. Ìàêàðîâ3, À.Ã. Ïîíîìàðåíêî4, À.Ñ. Áàøêóåâ4 1 Papanin Institute for Biology of Inland Waters, Russian Academy of Sciences, Yaroslavl Oblast, Nekouzsky District, Borok 152742, Russia. ´ 2 Venevitinovo Research and Educational Centre, Voronezh State University, Universitetskaya pl., 1, Voronezh 394006, Russia. E-mail: [email protected]. 3 Zoology & Ecology Department, Moscow Pedagogical State University, Kibalchicha Str., 6, Bld. 5, Moscow 129164, Russia. E-mail: [email protected]. 4 Borissiak Palaeontological Institute, Russian Academy of Sciences, Profsoyuznaya ul., 123, Moscow 117997, Russia. E-mail: [email protected] (AGP), [email protected] (ASB). 1 Институт Биологии внутренних вод им. И.Д. Папанина РАН, пос. Борок 152742, Ярославская обл., Некоуский р-н. 2 Биологический учебно-научный центр «Веневитиново» Воронежского государственного университета, Воронеж 394006, Университетская пл., д. 1. 3 Московский педагогический государственный университет, кафедра зоологии и экологии, Москва, 129164 ул. Кибальчича д.6 корп. 5. 4 Палеонтологический институт им. А.А. Борисяка РАН, Москва 117997, ул. Профсоюзная, д. 123. KEY WORDS: Coleoptera, larvae, Triassic, Germany, Coptoclavidae. Caraboidea, Polyphaga, evolution. КЛЮЧЕВЫЕ СЛОВА: Coleoptera, личинки, триас, Германия, Coptoclavidae. Caraboidea, Polyphaga, эволюция. ABSTRACT. Three new larvae, †Protonectes ger- which is near-contemporaneous with a number of other manicus gen. et sp.n., †Carabilarva triassica sp.n., assemblages known in European Triassic: the richest and †Larvula triassica sp.n., are described from the one occurs in the “Grès à Voltzia” Formation of the Triassic of Germany (Coburger Sandstein, Hassberge Vosges Mountains [Gall & Grauvogel-Stamm, 2005], Formation, Middle Keuper). These taxa are the earliest the others are known from the Buntsandstein of Mallor- known larvae of the family Coptoclavidae, subfamily ca [Zessin, 2008] and from the Holy Cross Mountains in Caraboidea, and suborder Polyphaga, respectively. The Poland [Żyła et al., 2013]. From the younger deposits of evolutionary changes of Coptoclavidae and all Dytis- the Germanic Basin, only sparse and non-diverse in- coidea and life forms of the earliest known larvae of sects have been described, mainly from the Lower and ground beetles (in the broad sense) are discussed. Upper Keuper (Rhaetian) [Geyer & Kelber, 1987; Brauckmann & Schlüter, 1993, and references cited РЕЗЮМЕ. Три новые личинки: †Protonectes there]. Quite recently, Barth et al. [2011] reported a germanicus gen. et sp.n., †Carabilarva triassica sp.n., diverse insect fauna from the new late Middle Keuper †Larvula triassica sp.n. описаны из триаса Германии (Late Norian) fossil site at the nearby Langenberg, (Кобургский песчаник, свита Хасберге, средний кей- Lower Saxony, which is currently the richest Late Trias- пер). Данные таксоны являются наиболее древними sic insect fauna known in Germany. Among others, из известных личинок семейства Coptoclavidae, над- aquatic larvae of beetles were mentioned, but remain семейства Caraboidea и подотряда Polyphaga, соот- unstudied. ветственно. В статье рассматриваются эволюцион- In addition, insect remains are known from the ные преобразования личинок коптоклавид и дитис- Middle Keuper Coburger Sandstein (Hassberge For- коидов в целом; жизненные формы древнейших mation, Upper Carnian) in Lower Franconia. The личинок жужелиц (в широком в смысле). Coburger Sandstein crops out at numerous quarries in the Hassberge area and shows a remarkable interbed- Introduction ded succession of fluvial channel sandstones interca- lated with fine-laminated lacustrine and floodplain The Triassic insects of Germany are rare and rather sediments. The beds are rich in fossils [Kelber & poorly studied. The most abundant insect assemblage Nitsch, 2005], including, besides insects, particularly was discovered in the Upper Buntsandstein (Lower abundant remains of Triops cancriformis [Kelber, Anisian) of Lower Franconia [Bashkuev et al., 2012], 1999], as well as conchostracans, fish remains, thero- 260 A.A. Prokin, K.V. Makarov , A.G. Ponomarenko & A.S. Bashkuev pod tracks, invertebrate ichnofossils, and diverse mac- Results roflora. Insect remains collected in Coburger Sandstein Descriptions of new taxa (mainly from a quarry south of Eltmann) number several hundred specimens, mostly beetles [J. Sell, pers. comm. Order Coleoptera to A.S. Bashkuev, 2013], but all the material is housed Suborder Adephaga in private collections and has never been studied by Superfamily Dytiscoidea specialists. Only a few specimens were mentioned in Family Coptoclavidae Ponomarenko, 1961 publications: e.g., an odonate referred to as Triadotypus cf. guillaumei Grauvogel et Laurentiaux, 1952 [Bechly, †Protonectes Prokin et Ponomarenko, gen.n. 2013, in press] and a cupedoid elytron [Kelber in Re- Figs 1–7 imann & Schmidt-Kaler, 2002: 16], which can be as- TYPE SPECIES: †Protonectes germanicus sp.n. signed to the genus Zygadenia Handlirsch, 1906. DIAGNOSIS: Larva. Head somewhat wider than long, A number of insect remains (about 40 specimens) narrowing anteriorly of eyes, nasale not protruding anteri- were collected by Jürgen Sell (SMTE) and one of the ad. Mandibles without retinaculum, with two cutting sur- authors (ASB) at the quarry Schönbachsmühle near faces and blunt tab medially on outer margin. Epicranial Ebelsbach in 2012 and 2013. This collection includes suture narrow; frontal sutures weakly sinuate. Length of antenna equal to maximum head width. Head subequal in fragments of orthopterans and cockroaches, single length to prothorax. Pronotum almost twice as long as specimens of Thysanoptera, Trichoptera, and possi- meso- or metanotum. Tracheae without pronounced taeni- ble Raphidioptera, elytra of Cupedidae (not only dia; commissures and connectives not visible; spiracles present Zygadenia, but also other morphotypes), and diverse in posterior part of mesothorax. Sparse swimming hairs larvae, including several well-preserved beetle lar- present on femora of fore- and midlegs and tarsi of hindlegs. vae described herein. This collection includes also Raptorial forelegs almost as long as maximum body width; elytra similar in shape and dark coloration pattern to natatorial hindlegs twice as long as maximum body with. those of Holcoptera Handlirsh, 1906, known from COMPARISON: The new genus is especially similar in the Triassic of Europe and North America. There are the body shape, level of “constructive perfection” of legs reasons to suggest that these elytra belong to copto- structure of †Daohugounectes primitivus Wang, Ponoma- renko et Zhang, 2009 from the Middle Jurassic of China, and clavid beetles. In addition, a beetle hindwing of mem- differs from it in the mandibles without retinaculum and with bers of Coptoclavidae have been found at this site. a blunt tab on outer margin, shorter epicranial suture, and Characters, which support assignment this hind wihgs sparser swimming hairs, which are absent on mesotibia, to Coptoclavidae are: straight and long vein Cu2, the mesotarsus and metatibia. The new genus is also similar in basal position of the common base vertex branches the structure of thoracic apodemes to †Stygeonectes jurassi- CuP and AA1+2, as well as large, long and convex cus Ponomarenko, 1977, and differs from it in the structure of forward 2nd anal cell [Fedorenko, in press]. mandibles and sparser swimming hairs of the legs. Compar- ison with the other known species of Mesozoic dytiscoid larvae and members of recent families is given below (table). Material and methods ETYMOLOGY: From traditional latinization of the Greek πρώτοs = protos (first) and the nectes, traditional latinization The newly described fossils are deposited in the of the Greek νήκτης = nektes (swimmer). collection “Sammlung Mainfränkische Trias” in Markt †Protonectes germanicus Prokin et Ponomarenko, Euerdorf, Lower Franconia (SMTE). The fossils are sp.n. preserved as impressions in laminated clayey shale of Figs 1–7 lacustrine genesis. Two larvae (SMTE 5930/1–65 A MATERIAL: Holotype SMTE 5930/1–65A, part and counterpart. and B) are preserved on the same slab and somewhat LOCALITY AND HORIZON. Quarry Schönbachsmühle near overlapping: the head of the latter was uncovered after Ebelsbach, Lower Franconia; Coburger Sandstein, Hassberge For- fine preparation of the pronotum of the coptoclavid mation, Middle Keuper, Upper Carnian. larva. The same slab contains also a mayfly nymph and DESCRIPTION (Figs 1–2, 5–9). Medium-sized larva, several head shields of Triops. with incompletely preserved abdomen. Body terete, widest at Fossils used for comparison are deposited in the level of abdominal segments 3 and 4. Antennomere 2 longer than antennomeres 3 and 4 together; antennomere 3 three collections of the Borissiak Paleontological Institute, times as long as antennomere 4; antennomere 1 wider than Russian Academy of Sciences, Moscow, Russia (PIN) long, twice as wide as any other antennomere. Maxillary and Capital Normal University, Beijing, China (CNU). palpus with homonomous palpomeres and equal in length to Collections of recent beetles larvae are deposited in mandible. Ocelli at boundary between anterior and medial
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