Insects and crustaceans from the latest Early–early Middle Triassic of Poland

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Abstract

Two stratigraphical horizons in the Pałęgi clay-pit, a new Triassic paleontological site within Buntsandstein deposits (latest Olenekian–early Anisian in age) in the Holy Cross Mountains (Poland), have yielded arthropod faunas comprising ca. 400 fossil specimens assigned to two subphyla: Crustacea (class Branchiopoda and Maxillopoda) and Hexapoda (class Insecta). The Pałęgi arthropod assemblage is similar to that described from the Middle Triassic of France and Germany but is dominated by remains of conchostracans and cockroaches. This new fauna expands our knowledge of the latest Early–early Middle Triassic diversity of insects and freshwater arthropods in the Germanic Basin. The newly discovered fauna represents one of the oldest Mesozoic records of insects described from the Buntsandstein facies of Europe, and provides important information to better appreciate the process of ecosystem recovery after the Permian–Triassic extinction.

Highlights

► We describe a noteworthy Triassic arthropod assemblage from Germanic Basin. ► Arthropod fossils occur in deposits interpreted as freshwater ponds. ► This is the oldest record of insects from the Buntsandstein of Germanic Basin.

Introduction

Deposits with Early or Middle Triassic fossil insects and other terrestrial (freshwater) arthropods (e.g., limulids, triopsids, cycloids, conchostracans) are known from a selected number of exposures in Germany, France, Spain, European Russia (and Siberia, Urals), Kyrgyzstan, Mongolia and Australia (see Trümpy, 1957, Kozur and Seidel, 1983a, Kozur and Seidel, 1983b, Hauschke and Szurlies, 1998, Hauschke and Szurlies, 2006, Lozovsky et al., 2001, Durante and Luvsantseden, 2002, Jell, 2004, Gall and Grauvogel-Stamm, 2005, Shcherbakov, 2008a, Shcherbakov, 2008b, Béthoux et al., 2009, Bashkuev et al., 2011). Arthropod fossils generally are very rare in the Early–Middle Triassic continental deposits, except for numerous occurrences of conchostracans and ostracods (see Kozur and Weems, 2010). The succession of insect and small freshwater crustacean assemblages in the Early–Middle Triassic is typically well recognized based on material collected from sites in European Russia, the Urals, Siberia, China, Kyrgyzstan and Mongolia, but very rich and exceptionally preserved arthropod fossils were described from the early Middle Triassic of France and Germany (Gall and Grauvogel-Stamm, 2005, Bashkuev et al., 2011). The most interesting fossils are insects, which are good indicators of environmental changes and fluctuations in biodiversity. The most complete entomofaunal successions of the Late Permian and Early Triassic (across the Permian-Triassic boundary) have been found in European Russia (Shcherbakov, 2008a, Shcherbakov, 2008b).

In this study, we present a preliminary description of arthropod remains (conchostracans, cycloid crustacean and insects) collected from the Pałęgi clay-pit (Fig. 1), a new late Early–early Middle Triassic paleontological locality, which is situated in the easternmost part of the Germanic Basin. Triassic deposits from this site contain also fossils of other invertebrates (gastropods and bivalves), plants (both micro- and macrofossils), numerous invertebrate trace fossils and tetrapod bones and traces (Kuleta et al., 2006, Skawina and Niedźwiedzki, 2012).

Lower–Middle Triassic continental deposits are widely distributed in several basins of western and central Europe (Aigner and Bachmann, 1992), but they are often represented by the classic redbed sequences that are poorly fossiliferous. Arthropod remains (others than conchostracans) are very rare in the Late Permian–Middle Triassic succession of the Buntsandstein facies of Europe. They have been found only in a few sites (France, Germany, and Spain) and are usually not well preserved with the exception of material from the Middle Triassic ‘Grès à Voltzia’ Formation (Ansorge, 1994, Gall and Grauvogel-Stamm, 2005, Aristov and Zessin, 2009, Bashkuev et al., 2011).

The first conchostracans and other arthropod remains were discovered at Pałęgi in 2003 by one of the authors of this paper (GN) and were mentioned by Kuleta et al. (2006). The occurrence of fossil insects in Pałęgi was first mentioned by Krzemińska et al. (2007), at a meeting of the Section of Fossil Insects Polish Entomological Society in Gdańsk in 2007 (see Krzemińska et al., 2007). Their poor state of preservation did not allow the undisputed classification of the specimens to any insect group. The authors gathered thirty specimens and suggested that the specimens included representatives of Coleoptera, homopterous and heteropterous Hemiptera. Our exploration of insects in Pałęgi in 2010–2012 led to an assembly of many important new specimens, several of which are well preserved. In total, 20 specimens were found, including 19 preserved in the form of isolated wings and one as a body (without wings). Initially they were classified into three groups: Blattodea, Grylloblattida and Mecoptera. The most abundant specimens are Blattodea.

Section snippets

Geological, stratigraphical and palynological setting

The Pałęgi site is located within a pit of vitrified clay situated near Grzymałków about 7 km to the west from the town of Mniów (see Kuleta et al., 2006, Ptaszyński and Niedźwiedzki, 2006). The exposed sedimentary section in Pałęgi is comprised of the upper part of the Samsonów Formation, which is described as late Olenekian in age (Kuleta and Nawrocki, 2000, Kuleta et al., 2006). In the upper part of the Pałęgi exposure younger Middle Triassic deposits are also exposed, which was recently

Material and methods

The studied specimens were collected by all of the authors and collaborators. Specimens described in this contribution are housed at the Institute of Paleobiology, Polish Academy of Sciences, Warszawa (ZPAL PA.I). Reported measurements (in millimetres) were obtained using digital callipers and a micrometre eyepiece included in a binocular microscope. Photographs of specimens were taken using a digital camera Canon EOS 200 and AxioCam MR5 (Department of Zoology, University of Silesia, Katowice,

Taphonomy and fossil distribution

The assemblage was collected from two horizons from the middle part of the exposure and is dominated by conchostracan carapaces, which represent about 95% of collected arthropod fossils. The vast majority of collected insect remains from the Pałęgi clay-pit are represented by isolated wings, sometimes preserved partially, the size of which varies from 4 to 7 mm (11 rather complete and 8 partially preserved wings are found among a total of 20 insect fossils). Specimens of whole insects are very

Systematic palaeontology

Subphylum Crustacea Pennant, 1777

Class Branchiopoda Latreille, 1817

Order Conchostraca Sars, 1867

Suborder Spinicaudata Linder, 1945

The Pałęgi conchostracans are preserved mainly as highly flattened or as three-dimensional, silica-replaced specimens that have undergone relatively little crushing. Nearly 400 separate specimens (ZPAL PA.I/1–286) were collected from the seven stratigraphic levels from the middle part of the exposure. They have ovate to subovate valves with numerous, closely-spaced

Discussion

The Triassic period is heralded as one of the key stages in the evolution of insects (Grimaldi and Engel, 2005). Knowledge about Early and Middle Triassic insects and freshwater crustaceans is still insufficient, however. Only very few insects are known from the Early Triassic, but in the Middle Triassic onwards the number of fossil sites increases and in fact, the most Triassic insect faunas are from the Carnian or later (e.g., Fujiyama, 1991, Fraser et al., 1996, Wappler, 2001, Jell, 2004,

Conclusions

The Pałęgi arthropod assemblage has been recognized in the upper part of the Samsonów Formation and contains a unique record of a few species of conchostracans, probably a new species of cycloid crustacean and remains of three groups of insects (Blattodea, Grylloblattida and Mecoptera). The Pałęgi arthropod assemblage is similar to those described from the Middle Triassic of France and Germany. Evidence from sedimentology and palaeoecological data indicates that deposits from Pałęgi represent a

Acknowledgments

This study started in 2003, after an initial visit of one of the author of this paper (GN) in the Pałęgi clay-pit and first discoveries of arthropod fossils in this locality. Special thanks to Józef Szajn, Stanisław Piskorz, Maria Kuleta, Stanisława Zbroja (Kielce, Poland), Tadeusz Ptaszyński (Warszawa, Poland), Katarzyna Zaremba–Niedźwiedzka (Uppsala, Sweden), Tomasz Sulej (Warszawa, Poland), Marian Dziewiński (Warszawa, Poland), students from Scientific Group “Perisphinctes” (Department of

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