Introduction

Brauckmann and Schlüter (1993) described the insect assemblage of Hammelburg, Unterfranken (Lower Franconia), which is hitherto the richest occurrence from the European Buntsandstein except for the Vosges (“Grès à Voltzia” Formation of the Upper Buntsandstein, NE France; see for example Gall and Grauvogel-Stamm 2005). They mentioned 17 specimens from the Myophoria beds (uppermost Röt, Early Anisian), belonging to Blattodea (Triassoblatta? sp.), Coleoptera: Permosynidae (cf. Ademosyne sp., aff. Meseumolpites sp.), Hemiptera: Auchenorrhyncha (Heseneuma hammelburgensis Brauckmann et Schlüter, 1993; Saaloscytina perreticulata Brauckmann et Schlüter, 1993; Ipsviciidae gen. et sp. indet.), Hemiptera: Sternorrhyncha (Dubiaphis curvata Brauckmann et Schlüter, 1993), Heteroptera fam., gen. et sp. indet. and also Planipennia fam., gen. et sp. indet. The latter three are referred here to Diptera, Hemiptera indet. (determined by D.E. Shcherbakov), and Grylloblattida: Chauliodites Heer, 1864, respectively.

The number of fossil insects collected from Hammelburg to other Buntsandstein outcrops in Lower Franconia and Southern Thuringia has greatly increased in recent years owing to the activity of the working group “Mainfränkische Trias” from Markt Euerdorf, Unterfranken, headed by Jürgen Sell. This collection includes more than 300 specimens by now, assigned to ten orders: to the orders listed above, Ephemeroptera, Glosselytrodea, Orthoptera, Mecoptera, Diptera and Archaeognatha were added.

In addition, Ansorge and Brauckmann (2008) recorded a grylloblattid insect from the “Strohgelbe Kalke” of Gambach am Main, one of the localities considered here, and identified it as Chauliodites picteti Heer, 1864. The total list of insects recorded from the Upper Bundsandstein of Lower Franconia and Southern Thuringia adjusted for the redefining of Brauckmann and Schlüter’s types is provided below.

 

Archaeognatha: Dasyleptidae: Dasyleptus sp.

Ephemeroptera: Misthodotidae:

 Triassodotes vogesiacus Sinitshenkova, Marchal-Papier, Grauvogel-Stamm et Gall, 2005

Ephemeroptera: Sharephemeridae:

 Hammephemera pulchra Sinitshenkova, gen. et sp. n.

Blattodea: Mesoblattinidae: ?Triassoblatta sp.

Grylloblattida: Chaulioditidae:

 Chauliodites picteti Heer, 1864

Orthoptera fam., gen. et sp. indet.

Hemiptera: Auchenorrhyncha: Saaloscytinidae:

 Saaloscytina perreticulata Brauckmann et Schlüter, 1993

Hemiptera: Auchenorrhyncha incertae sedis:

 Heseneuma hammelburgensis Brauckmann et Schlüter, 1993

Hemiptera: Auchenorrhyncha: Ipsviciidae gen. et sp. indet.

Coleoptera: ?Adephaga: Triaplidae: Triaplus sp.

Coleoptera: Archostemata:

    Cupedidae: Notocupes sp.

    Tricoleidae indet.

    Permosynidae (formal taxa):

      Ademosyne sp.

      Ulomites sp.

    Schizocoleidae (formal taxa):

        Polysitum sp.

        Pseudohydrophilites sp.

        Strongulites sp.

        Tillyardiopsis sp.

        Adikia sp.

        Metriorhynchus sp.

        Wollastonia sp.

        Shepherdia sp.

        Mesolobites sp.

Glosselytrodea: Jurinidae (s.l.) gen. et sp.

Mecoptera: Pseudopolycentropodidae:

    Pseudopolycentropus triasicus Papier, Nel et Grauvogel-Stamm, 1996

Diptera:

    Dubiaphis curvata Brauckmann et Schlüter, 1993

    Diptera gen. et sp.

All the new specimens referred to as SMTE are stored in the collection “Sammlung Mainfränkische Trias” in Markt Euerdorf, Unterfranken. Specimen MHI 1486 is deposited in the Dr. Hans Hagdorn’s collection in the Muschelkalkmuseum Ingelfingen, Baden-Württemberg.

Geological and stratigraphical setting

After being classified as Formation s7, the Röt-Formation (Upper Buntsandstein), was divided into four subformations. Röt 4 Subformation (Table 1) consists of “Rötquarzitschichten,” “Obere Röttonsteine” and “Myophorienschichten,” and corresponds to the subformation s7–4 (resolution no. 61/66 of the Permian-Triassic Subcommission, 2006/2007).

Table 1 Stratigraphic division of the Upper Buntsandstein (Röt 4) in Lower Franconia and South Thuringia
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“Rötquarzitschichten” are interpreted as having been developed in the tidal area (Geyer 2002) and correspond to the beginning of transgressive system tracts of Lower Muschelkalk (Aigner and Bachmann 1992).

The depositional environment of reddish silty clay marls of the “Obere Röttonsteine” is supposed to be similar to the recent playa/sabkha deposits, which were sometimes flooded by marine incursions. In the upper third section of the “Obere Röttonsteine” the vulgaris/costata bed (isochronous to the “Dolomitischen Grenzbank” in Thuringia) forms the most important key horizon in the Upper Buntsandstein in Lower Franconia (Mahler and Sell 1993).

The layers above the “vulgaris/costata” bed are characterized by some distinctive green to grey-yellow reduced horizons separated by red-brown, silty, crumbly clay marls. The “Münnerstädter bank” with Pseudocorbula subundata, Saurichthys sp., vertebrate and insect remains along with the vulgaris/costata bed is the horizon with the richest content of fossils within the “Obere Röttonsteine.”

The “Myophorienschichten” (Myophoria beds) form the uppermost part of the Buntsandstein section. In Lower Franconia they begin by the “Untere Dendritenschichten,” an interbedding of green-grey mudstones and marls, at the base with glauconitic material (Mahler et al. 1990), which are overlaid by sandstones separated by thin clay marl layers. The terminal bed of grey-green mudstones of 0.20–0.80 m thickness (“Myophorien-Hauptbank”: Schuster 1936) with bivalves, conchostracans, decapods and rare insect remains (beetle elytra) extends from southern Thuringia to northern Baden-Württemberg.

The overlaying, maximally 6.5 m thick, “Rotes Zwischenmittel” is an equivalent of Myophorientone of the Thuringian Basin. It is formed mainly by red clay marls, in which spreite burrows (Rhizocorallium sp.) are rarely to be found.

The “Rotes Zwischenmittel” are followed by the “Obere Dendritenschichten,” a sequence of interbedded gray-gray-green and yellow clay marls, mudstones and sandstones. At the top there are intraformational mudstone/marlstone breccias up to 0.50 m thickness, penetrated by calcite veins, often with halite crystals. In contrast to the “Untere Dendritenschichten,” bivalves here are rare and small sized. Most commonly lingulids, vertebrate remains (particularly Saurichthys sp.), crustaceans (Triops sp., conchostracans), and plant and insect remains are found.

The “Strohgelbe Kalke” form the top of the Myophoria beds. Yellow-grey to straw-yellow dolomitic marls to mudstones contain mainly conchostracans, plant and insect fossils.

After Kozur (1974) the Myophoria beds are considered to be the biostratigraphic equivalent of the “Grès à Voltzia” Formation in the southwest and of the lower Gogoliner Schichten Formation in the east. However, the biostratigraphic correlation of the “Grès à Voltzia” Formation is based only on the occurence of Myophoria vulgaris, which confirms at best the Bithynian age.

Localities and paleogeography

In the early Triassic the Germanic Basin resembled to a great extent that of the Zechstein sea. Due to tectonic processes at the beginning of the Anisian, a connection to the Tethys basin was established via the East Carpathian Gate (Fig. 1).

Fig. 1
figure 1

Paleogeography of the Germanic Basin during the Upper Buntsandstein (Röt 4). Modified after Ziegler (1990). 1–5, insect localities

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The marine sediments of the lower “Rötkalk” (equivalent of the “Dolomitischen Grenzbank” of Thuringia) with crinoids, Costatoria costata and M. vulgaris were deposited in areas of the East Carpathian Gate. In contrast, from the time of the formation of the vulgaris/costata bed, the region between Jena and Würzburg was situated within an area characterized by the frequent change of playa-, sabkha- and shallow-marine settings. This assumption, hitherto predominantly justified lithologically, is confirmed particularly by the frequent co-occurrence of terrestrial/freshwater, brackish and marine organisms.

Composition and taphonomy of insect assemblages

The ordinal composition of the insect collections from Upper Buntsandstein localities housed at the SMTE is summarized in the Table 2. The incomplete or poorly preserved specimens that cannot be definitely assigned to any particular order are listed as incertae sedis and excluded from percentage counts.

Table 2 Distribution of insect orders in Upper Buntsandstein localities
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All assemblages are strongly biased, with the Coleoptera comprising from 48 to 78% of collected material. They are represented mainly by isolated elytra (or rarely by isolated thoracic fragments); among other insects the strongly sclerotized elements such as tegmina or their parts (isolated clavi) of Auchenorrhyncha and Blattodea also prevail. Wings of other insects, as well as whole insects including beetles, are very rare.

Assemblages enriched with robust elytra of beetles, cockroaches, and cicadas, are typical for allochthonous taphocoenoses with a long transportation of remains by flowing water (Zherikhin 2002). Though some schizocoleid coleopterans are presumed to be surely aquatic, they are taphonomically similar to other beetles and are apparently allochthonous as well. The only possibly autochthonous elements in the assemblage are quite numerous Dasyleptus fossils (Fig. 2) found in the “Obere Dendritenschichten” of Hammelburg. They are mostly completely preserved (not as molting casts), including a number of specimens buried clustered on a small piece of rock. The data available imply at least a semiaquatic lifestyle of Dasyleptus (Rasnitsyn 1999; Rasnitsyn et al. 2004). There is no evidence of other aquatic insects, e.g., mayfly nymphs, which in contrast are very abundant in the Vosges.

Fig. 2
figure 2

Dasyleptus sp.; early Middle Triassic (Upper Buntsandstein, Röt Fm, “Strohgelbe Kalke”, Lower Anisian); Hammelburg, Lower Franconia. a–b Part and counterpart of specimen SMTE, 5825/2-436, lateral view. c Specimen SMTE, 5825/2-364, dorsal view. d Specimen SMTE, 5825/2-417, lateral view

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The insect assemblage of the Schwarza locality rather differs from other regions: Here three rather complete grylloblattid wings (in other localities grylloblattids are represented mostly by wing fragments) and a fragment of large orthopteran wing are found among a total of ten insect fossils. The fossil content of the Buntsandstein section of Schwarza has not yet been prospected in detail, but owing to these finds it looks to be the most promising locality for fossil insects.

Coleoptera

Beetles are mostly represented by diverse Schizocoleidae and Permosynidae. Among the beetles determined as cf. Ademosyne sp. by Brauckmann and Schlüter, only two specimens may actually belong to this genus (SMTE, 5825/2-43, 5825/2-31), while most others are rather similar to Ulomites Dunstan, 1924 (in Tillyard and Dunstan 1924), and one specimen is referable to Shepherdia Dunstan, 1924 (in Tillyard and Dunstan 1924) (Schizocoleidae). The specimen identified as aff. Meseumolpites sp. by Brauckmann and Schlüter belongs most probably to the genus Triaplus Ponomarenko, 1977 (in Arnoldi et al. 1977) (Triaplidae): no other known Triassic beetles had their opistognathous head so deeply bent under the prothorax.

Other coleopterans are represented by rare Cupedidae (single specimen of Notocupes sp.) and indeterminable Tricoleidae. There are not any Polyphaga as well as true Adephaga. Most determined genera are common in the Middle-Late Triassic and Early Jurassic.

The list of determined genera is provided in the Introduction section. But since many specimens are represented by poorly preserved isolated elytra, their further specification as well as the comprehensive systematic study of the beetle assemblage is needed.

Ephemeroptera

The finds of the fossil mayfly wings are quite rare. Therefore, the two complete wings with clear venation, recorded from the Upper Buntsandstein of Lower Franconia, are of great interest.

One wing (SMTE, 6123/1-28; Fig. 3), from “Strohgelbe Kalke” of Lengfurt, is referred to the family Misthodotidae and may be identified to the species level as Triassodotes vogesiacus Sinitshenkova, Marchal-Papier, Grauvogel-Stamm et Gall, 2005 from the Vosges in northeastern France (Sinitshenkova et al. 2005).

Fig. 3
figure 3

Triassodotes vogesiacus Sinitshenkova, Marchal-Papier, Grauvogel-Stamm et Gall, 2005; specimen SMTE, 6123/1-28; early Middle Triassic (Upper Buntsandstein, Röt Fm, “Strohgelbe Kalke,” Lower Anisian); Lengfurt, Lower Franconia. a Photograph, b drawing

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In the deposits of the “Grès à Voltzia” Formation in the Vosges, a rich mayfly fauna has been found. Two mayfly species, based on wings, belong to the families Misthodotidae and Toxodotidae (Sinitshenkova et al. 2005). Up to now this is the only case in which the typical Permian family Misthodotidae still occurs in Triassic deposits. The representatives of Misthodotidae and Toxodotidae are characterized by the presence of oval and almost homonomous wings.

The particular characters of the second wing make it possible to erect a new genus and species, named H. pulchra Sinitshenkova, gen. et sp. n. Due to the presence of a long CuA with intercalary veins forming a triad, it is included in the family Sharephemeridae Sinitshenkova, 2002. This family was originally established for the single species Sharephemera cubitalis Sinitshenkova, 2002, based on a unique find of a forewing from the Late Jurassic insect-bearing deposits in the Shar Teg locality in Mongolia (Sinitshenkova 2002). The new specimen renders a review of the family and the nominate genus Sharephemera, showing that the diagnosis of the family as given in the original paper was right. The new genus differs from Sharephemera in many features.

In the new species H. pulchra sp. n. the wing has an obviously expressed tornus, which is characteristic for a heteronomous flying apparatus. This means that hind wings of H. pulchra sp. n. were considerably shortened in comparison with the forewings. This feature is typical for an advanced state of the flying apparatus. However, the venation with CuA branching and forming a triad is ples iomorphic.

Order Ephemerida Latreille, 1810

(=Ephemeroptera Hyatt and Arms, 1890, s.l.)

Family Sharephemeridae Sinitshenkova, 2002

Emended diagnosis:

Small mayflies, forewing two times as long as wide. Base of forewing broad with developed anal area, cubital area narrow, CuA fork long with intercalary vein, CuP long and simple, anal veins long, cubital and anal veins straight and slightly curved.

Comparison:

The new family sharply differs from known families by the shape of the forewing with wide basal part, narrow cubital area, long CuA fork with intercalary vein, almost straight and slightly curved cubital and anal veins.

Genus Sharephemera Sinitshenkova, 2002

Type species:

Sh. cubitalis Sinitshenkova, 2002, from the Upper Jurassic lacustrine insect-bearing deposits from west of Shar Teg Mountain to southeast of Adzh-Bogdo in Gobi-Altai aymak of Mongolia.

Emended diagnosis:

Forewing. Anterior margin slightly concave, costal area wider at wing base than on its top. Anterior RS branch simple, posterior one branching. Forks of MA and CuA longer than their stems correspondingly. CuP straight, A1 branching, with a long intercalary longitudinal vein between its branches. Anal area obviously widened, tornus poorly expressed.

Genus Hammephemera Sinitshenkova, gen. n.

Etymology:

from Hammelburg.

Type species:

Hammephemera pulchra Sinitshenkova, n. sp.

Diagnosis:

Forewing. Anterior margin slightly convex, its top rounded. RS system with seven longitudinal veins, posterior branch of RS simple, anterior one branching. Forks of MA and CuA not longer than their stems correspondingly, CuP almost straight. Anal veins simple, A1 smoothly curved, A2 almost straight, moderately seen simple short A3. Anal area narrowed, tornus expressed and moved towards the middle of wing.

Comparison:

Hammephemera gen. n. differs well from Sharephemera by the convex anterior wing margin, branching of the anterior RS branch, shorter forks of MA and CuA, simple anal veins in more narrow anal area and by tornus moved towards the middle of the wing.

Hammephemera pulchra Sinitshenkova, sp. n.

Etymology:

from Latin pulchra, beautiful.

Holotype:

SMTE, 5825/2-365; negative impression of complete forewing with well-preserved longitudinal venation, crossveins not visible (Fig. 4).

Fig. 4
figure 4

Hammephemera pulchra Sinitshenkova, gen. et sp. n., holotype SMTE, 5825/2-365; early Middle Triassic (Upper Buntsandstein, Röt Fm, “Obere Dendritenschichten”, Lower Anisian); Hammelburg, Lower Franconia. a Photograph, b drawing

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Locality and Horizon:

Herolds-Berg N’Hammelburg, Lower Franconia; Myophoria beds (“Obere Dendritenschichten”), Röt Fm, Early Anisian, Bythinian, Middle Triassic.

Diagnosis:

as for the genus by monotypy.

Description:

Forewing length, 8.75 mm, maximum width, 4.06 mm. Costal area obviously wider than subcostal one at the wing base and in its top. Anterior RS branch forking twice and entering wing margin by seven branches. MA fork situated restrictedly proximal than third RS fork, MA fork shorter than its stem. MP branching near wing base slightly distad of first RS fork, its posterior branch running from MA with curve. CuA branching in the middle, CuP slightly curved near wing margin. Tornus situated almost at middle of wing.

Grylloblattida

All of the nine identifiable grilloblattid fossils found in the Upper Buntsandstein of Lower Franconia and Thuringia (including the specimen MHI 1486, deposited in the Muschelkalkmuseum Ingelfingen) belong to C. picteti Heer, 1864 (Chaulioditidae) (argumentation is given below in Remarks). In addition, there are four wing fragments belonging apparently to Chaulioditidae and possibly to Chauliodites as well. Thus, the grylloblattid assemblage is in contrast to that of the Vosges, where Blattogryllidae dominate (about 70% of all grylloblattids), while Chaulioditidae are rare (as well as Megakhosaridae and Mesorthopteridae) (Aristov et al. 2011).

Also, grylloblattids are relatively more abundant in Lower Franconia (Table 2) than in the Vosges: 5–11% in Hammelburg and Lengfurt; 4 specimens of 10 in Schwarza, versus 2 or <1% (Shcherbakov 2008; Aristov et al. 2011, respectively).

The family Chaulioditidae was erected by Handlirsch (1906) for C. picteti Heer, 1864 from the Middle Buntsandstein of Gödewitz near Salzmünde in Saxony-Anhalt. As it proved later, Chaulioditidae (especially the most widespread genus Chauliodites) are among the most important insect index fossils of the Late Permian-Early Triassic, which numerically dominated most of the Early Triassic insect assemblages. The most ancient findings of the family are known from the Middle and Late Permian of European Russia, and the youngest ones from the Middle Triassic of France (Vosges) and China (Aristov 2003, 2004, 2005, 2008, 2009; Aristov et al. 2011).

Order Grylloblattida Walker, 1914

Suborder Grylloblattina Walker, 1914

Family Chaulioditidae Handlirsch, 1906

Genus Chauliodites Heer, 1864

Chauliodites picteti Heer, 1864

Holotype:

(Heer 1864: Fig. 11; Handlirsch 1906: pl. 39, Fig. 16); Gödewitz near Salzmünde (S Wettin), Saxony-Anhalt; Middle Buntsandstein, Lower Triassic.

New material:

specimens SMTE, 5329/1-12 (Fig. 5) and 5329/1-39 (Fig. 6), each presented by part and counterpart of well-preserved forewing without anal area; Myophoria beds (“Obere Dendritenschichten”), Röt Fm, Lower Anisian, Bythinian, Middle Triassic; Schwarza, Thuringia; SMTE, 5329/1-1, forewing with apical and basal parts being destroyed; Myophoria beds (“Strohgelbe Kalke”), Röt Fm, Lower Anisian, Bythinian, Middle Triassic; Schwarza, Thuringia; SMTE, 5825/2-198, fragment of central part of forewing; Myophoria beds (“Strohgelbe Kalke”), Röt Fm, Lower Anisian, Bythinian, Middle Triassic; Herolds-Berg N’Hammelburg, Lower Franconia.

Fig. 5
figure 5

Chauliodites picteti Heer, 1864, specimen SMTE, 5329/1-12; early Middle Triassic (Upper Buntsandstein, Röt Fm, “Obere Dendritenschichten”, Lower Anisian); Schwarza, Thuringia. a Photograph, b drawing

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Fig. 6
figure 6

Chauliodites picteti Heer, 1864, specimen SMTE, 5329/1-39; early Middle Triassic (Upper Buntsandstein, Röt Fm, “Obere Dendritenschichten”, Lower Anisian); Schwarza, Thuringia. a Photograph, b drawing

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Specimens SMTE, 5825/2-21 (Brauckmann and Schlüter 1993: Fig. 10, described as Planipennia, fam., gen. et sp. indet.), from “Strohgelbe Kalke” of Hammelburg, and SMTE, 6123/1-26, 6123/1-31 and 6123/1-32 from “Strohgelbe Kalke” of Lengfurt, Lower Franconia, all represented by forewing fragments, also belong most likely to the same species.

Description:

Forewing length about 14; length/width ratio about 3:1. Anterior margin convex. Costal space at the beginning of RS is 2.5–3 times as wide as subcostal space. SC terminating before distal wing third. Fore branches of SC and R simple or dichotomous. R curving towards before beginning of RS. RS beginning in basal wing third, forked before the SC apex. M forked distally to base of RS. MA and MP two-branched. Space between the CuA branches narrowed in its distal half. CuA1 kinked at joints with crossveins. Interradial space at wing base narrow. Crossveins simple. Color pattern formed by dark spots in wing base, at base of RS and at pterostigma.

Remarks:

The described material is indistinguishable from the holotype of Ch. picteti as figured by Ansorge and Brauckmann (2008, Fig. 3b) and referred here to this species despite of the stratigraphic difference. Since the holotype of Ch. picteti is missing (Ansorge and Brauckmann 2008, p. 256), the final decision of the specific attribution of the described material should be deferred until new finds of conspecific wings in the type locality.

The specimen MHI 1486 from “Strohgelbe Kalke” of Gambach, described as Ch. cf. picteti by Ansorge and Brauckmann (2008), is only distinguished by the simple (or forking very late) MP and falls apparently within the observed intraspecific variability.

Mecoptera

The only specimen of Mecoptera from German Buntsandstein occurs from “Strohgelbe Kalke” of Hammelburg and is referred to the Vosges species P. triasicus Papier, Nel, et Grauvogel-Stamm, 1996. The family Pseudopolycentropodidae includes three genera, of which the most widespread, nominate genus Pseudopolycentropus is known from the Middle Triassic to Late Jurassic of Europe und Asia (Grimaldi et al. 2005), comprising only the minor component of the mecopteran fauna in almost all localities. The only exception is for the Vosges, where 19 of 22 mecopteran specimens belong to P. triasicus, while the rest fall within Permochoristidae (Prochoristella pilosa Papier, Nel et Grauvogel-Stamm, 1996), Liassophilidae [Laurentiptera gallica (Laurentiaux et Grauvogel, 1953) (in Laurentiaux 1953)] and Parachoristidae (Bashkuev, personal observation). In consideration of the shared species, it can be assumed that the Mecoptera assemblage of Lower Franconia was generally similar to the Vosges.

Order Mecoptera Packard, 1886

Family Pseudopolycentropodidae Handlirsch, 1925

Genus Pseudopolycentropus Handlirsch, 1906

Pseudopolycentropus triasicus Papier, Nel et Grauvogel-Stamm, 1996

Holotype:

Papier et al. (1996): Fig. 1; Northern Vosges Mountains, NE France; “Grès à Voltzia” Formation, Lower Anisian, Middle Triassic.

New material:

SMTE, 5825/2-199; complete forewing, part and counterpart (Fig. 7); Herolds-Berg N’Hammelburg, Lower Franconia; Myophoria beds (“Strohgelbe Kalke”), Röt Fm, Early Anisian, Bythinian, Middle Triassic.

Fig. 7
figure 7

Pseudopolycentropus triasicus Papier, Nel et Grauvogel-Stamm, 1996, specimen SMTE, 5825/2-199; early Middle Triassic (Upper Buntsandstein, Röt Fm, “Strohgelbe Kalke”, Lower Anisian); Hammelburg, Lower Franconia. a Photograph, b drawing

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Remarks:

Wing length, 5 mm, and maximum width 1.9. SC, pterostigma, free base of CuA and crossveins in anal area are not traceable because of poor preservation. The described specimen slightly differs from the holotype of P. triasicus in the narrow costal space and shorter CuA, reaching just the wing midlength. These deviations apparently fall within the range of intraspecific variability shown in the material from the Vosges.