Palaeontographica Abteilung B-palaophytologie, Dec 13, 2023
Endophytic oviposition, the reproductive behavior of insects to insert eggs into dead or living p... more Endophytic oviposition, the reproductive behavior of insects to insert eggs into dead or living plant tissue, and exophytic oviposition, the deposition of eggs on plants, are only rarely documented from the Pennsylvanian. In the present paper, ten oviposition types from the Piesberg quarry near Osnabrück, Germany (Osnabrück Formation, Middle Pennsylvanian, latest Moscovian, Asturian/Westphalian D) are described. The most abundant are large oviposition scars on sphenophyte stems, which probably housed single eggs. For these, we introduce Megaovoidus igen. nov., including the ichnospecies Megaovoidus piriformis igen. nov. et isp. nov. and M. compactus igen. nov. et isp. nov. A third ichnotaxon is Megaovoidus foveolatus igen. nov. et comb. nov., initially described as “Equisetites” foveolatus G.Roselt from the Late Triassic of Germany. A second new ichnogenus, Paleoincisurus igen. nov., is established for long and narrow slit-like ovipositional damages, which likely housed long rows of insect eggs. The very long scars of P. hellmundi igen. nov. et isp. nov. are associated with sphenophyte stems. For slit-like damages on Sphenopteris sp. cf. S. crepini fronds, the ichnotaxon Paleoincisurus muelleri igen. nov. et isp. nov. is introduced. We also provide evidence for three ichnospecies on leaves of the pteridosperm Macroneuropteris scheuchzeri: Paleoovoidus isp. cf. P. marginatus and two new types of oviposition damage (Paleoovoidus irregularis isp. nov. and P. parvus isp. nov.). Another oviposition damage, provisionally named “Ichnotaxon indet. A”, shows affinities to Paleoovoidus contactus, but is only fragmentarily documented. Finally, exophytic insect eggs, arranged in rows on an unidentified leaf, are described as Palaexovoidus lineatus isp. nov. Our study revealed a high abundance and diversity of endophytic oviposition concerning egg case morphology, oviposition patterns, and host plants of different habitats already in the Middle Pennsylvanian. As a result, a novel, different view on the early evolutionary history of reproductive strategies of late Paleozoic insects is indicated. Eventually, the establishment of an improved ichnotaxobase for oviposition traces according to the latest recommendations for modern ichnotaxonomy is another aim of our study.
Endophytic oviposition, the reproductive behavior of insects to insert eggs into dead or living p... more Endophytic oviposition, the reproductive behavior of insects to insert eggs into dead or living plant tissue, and exophytic oviposition, the deposition of eggs on plants, are only rarely documented from the Pennsylvanian. In the present paper, ten oviposition types from the Piesberg quarry near Osnabrück, Germany (Osnabrück Formation, Middle Pennsylvanian, latest Moscovian, Asturian/Westphalian D) are described. The most abundant are large oviposition scars on sphenophyte stems, which probably housed single eggs. For these, we introduce Megaovoidus igen. nov., including the ichnospecies Megaovoidus piriformis igen. nov. et isp. nov. and M. compactus igen. nov. et isp. nov. A third ichnotaxon is Megaovoidus foveolatus igen. nov. et comb. nov., initially described as “Equisetites” foveolatus G.Roselt from the Late Triassic of Germany. A second new ichnogenus, Paleoincisurus igen. nov., is established for long and narrow slit-like ovipositional damages, which likely housed long rows of insect eggs. The very long scars of P. hellmundi igen. nov. et isp. nov. are associated with sphenophyte stems. For slit-like damages on Sphenopteris sp. cf. S. crepini fronds, the ichnotaxon Paleoincisurus muelleri igen. nov. et isp. nov. is introduced. We also provide evidence for three ichnospecies on leaves of the pteridosperm Macroneuropteris scheuchzeri: Paleoovoidus isp. cf. P. marginatus and two new types of oviposition damage (Paleoovoidus irregularis isp. nov. and P. parvus
isp. nov.). Another oviposition damage, provisionally named “Ichnotaxon indet. A”, shows affinities to Paleoovoidus contactus, but is only fragmentarily documented. Finally, exophytic insect eggs, arranged in rows on an unidentified leaf, are described as Palaexovoidus lineatus isp. nov. Our study revealed a high abundance and diversity of endophytic oviposition concerning egg case morphology, oviposition patterns, and host plants of different habitats already in the Middle Pennsylvanian. As a result, a novel, different view on the early evolutionary history of reproductive strategies of late Paleozoic insects is indicated. Eventually, the establishment of an improved ichnotaxobase for oviposition traces according to the latest recommendations for modern ichnotaxonomy is another aim of our study.
Arthropod borings are commonly described from pteridophyte and gymnosperm wood in the late Palaeo... more Arthropod borings are commonly described from pteridophyte and gymnosperm wood in the late Palaeozoic, but they are almost unknown from calamitalean stems. In this paper, a new type of boring in calamitalean stems from two German localities is reported. These are the mine dumps of Plötz near Halle (Saale), Saale Basin, (Wettin Subformation, Latest Pennsylvanian, Gzhelian, Stephanian C) and the Piesberg quarry near Osnabrück, Lower Saxony Basin, (Osnabrück Formation, Middle Pennsylvanian, latest Moscovian, Asturian/Westphalian D). Most borings were found in marginal parenchyma of the internodes. They run longitudinally through the pith. The borings are preserved as three-dimensional casts that protrude on the surface of considerably compressed pith casts. This unique preservation of the boring casts required special taphonomic conditions, such as rapid burial coupled with different sediment infilling of the borings and the pith cavity, as well as anoxic conditions to prevent decomposition of the non-resistant parenchyma. Most borings are between 3 to 5 mm wide and contain two classes of invertebrate coprolites: the smaller coprolites are sub-spheroidal in shape and measure 37 to 74 x 37 to 63 μm in diameter. The larger coprolites are also sub-spheroidal and range between 88 to 158 x 68 to 123 μm in diameter. The coprolites, the morphology of the borings, as well as the feeding strategy of the arthropods, suggest that the tunnel system was most likely produced by small millipedes (Myriapoda) and probably also visited by oribatid mites.
Sandiella herbigi sp. nov. is the first record of the family Eugereonidae Handlirsch, 1906 (Insec... more Sandiella herbigi sp. nov. is the first record of the family Eugereonidae Handlirsch, 1906 (Insecta: Palaeodictyoptera) in the Pennsylvanian (Late Carboniferous: Westphalian D) sequence of the Piesberg quarry near Osnabrück, Lower Saxony, Germany. It is represented by a single fragment of a mesothoracic wing with the typical coarse reticulation in this family. The species is mainly characterized by the following features: (i) the shape of the mesothoracic wing is extremely narrow, (ii) the posterior subcostal vein is ending before the wing apex, (iii) all the main veins show a typical bend close to the posterior margin. The new wing is the second most ancient record of the Eugereonidae.
Piesberg quarry is famous for its Upper Carboniferous plant and arthropod fossils, including seve... more Piesberg quarry is famous for its Upper Carboniferous plant and arthropod fossils, including several holotypes of flying insects. The high degree of maturity of the Piesberg strata, such as the presence of anthracitic coal, quartzite, and large quartz crystals, led to controversies over a possible underlying thermal anomaly. The Piesberg is of further importance for correlation between the deep underground of northwestern Germany and the Ruhr basin coal field, as well as the Pennsylvanian coal areas in North America, and for investigations of Upper Carboniferous tight gas fields. The importance of the Piesberg for international geosciences has been enabled through its very rich mining history beginning in the Middle Ages and its long scientific history beginning at the end of the 18th century. Today, the Piesberg is not only one of the largest active quarry sites in Europe, but also a local recreation and hiking area. While the Museum am Scholerberg in Osnabruck protects its paleont...
Arthropod borings are commonly described from pteridophyte and gymnosperm wood in the late Palaeo... more Arthropod borings are commonly described from pteridophyte and gymnosperm wood in the late Palaeozoic, but they are almost unknown from calamitalean stems. In this paper, a new type of boring in calamitalean stems from two German localities is reported. These are the mine dumps of Plötz near Halle (Saale), Saale Basin, (Wettin Subformation, Latest Pennsylvanian, Gzhelian, Stephanian C) and the Piesberg quarry near Osnabrück, Lower Saxony Basin, (Osnabrück Formation, Middle Pennsylvanian, latest Moscovian, Asturian/Westphalian D). Most borings were found in marginal parenchyma of the internodes. They run longitudinally through the pith. The borings are preserved as three-dimensional casts that protrude on the surface of considerably
compressed pith casts. This unique preservation of the boring casts required special taphonomic conditions, such as rapid burial coupled with different sediment infilling of the borings and the pith cavity, as well as anoxic conditions to prevent decomposition of the non-resistant parenchyma. Most borings are between 3 to 5 mm wide and contain two classes of invertebrate coprolites: the smaller coprolites are sub-spheroidal in shape and measure 37 to 74 x 37 to 63 μm in diameter. The larger coprolites are also sub-spheroidal and range between 88 to 158 x 68 to 123 μm in diameter. The coprolites, the morphology of the borings, as well as the feeding strategy of the arthropods, suggest that the tunnel system was most likely produced by small millipedes (Myriapoda) and probably also visited by oribatid mites.
Specimens of Euproops sp. (Xiphosura, Chelicerata) from the Carboniferous Piesberg quarry near Os... more Specimens of Euproops sp. (Xiphosura, Chelicerata) from the Carboniferous Piesberg quarry near Osnabrück, Germany, represent a relatively complete growth series of 10 stages. Based on this growth sequence, morphological changes throughout the ontogeny can be identified. The major change affects the shape of the epimera of the opisthosoma. In earlier stages, they appear very spine-like, whereas in later stages the bases of these spine-like structures become broader; the broadened bases are then successively drawn out distally. In the most mature stage known, the epimera are of trapezoidal shape and approach each other closely to form a complete flange around the thoracetron (=fused tergites of the opisthosoma). These ontogenetic changes question the taxonomic status of different species of Euproops, as the latter appear to correspond to different stages of the ontogenetic series reconstructed from the Piesberg specimens. This means that supposed separate species could, in fact, represent different growth stages of a single species. It could alternatively indicate that heterochrony (=evolutionary change of developmental timing) plays an important role in the evolution of Xiphosura. We propose a holomorph approach, i.e., reconstructing ontogenetic sequences for fossil and extant species as a sound basis for a taxonomic, phylogenetic, and evolutionary discussion of Xiphosura.
Palaeontographica Abteilung B-palaophytologie, Dec 13, 2023
Endophytic oviposition, the reproductive behavior of insects to insert eggs into dead or living p... more Endophytic oviposition, the reproductive behavior of insects to insert eggs into dead or living plant tissue, and exophytic oviposition, the deposition of eggs on plants, are only rarely documented from the Pennsylvanian. In the present paper, ten oviposition types from the Piesberg quarry near Osnabrück, Germany (Osnabrück Formation, Middle Pennsylvanian, latest Moscovian, Asturian/Westphalian D) are described. The most abundant are large oviposition scars on sphenophyte stems, which probably housed single eggs. For these, we introduce Megaovoidus igen. nov., including the ichnospecies Megaovoidus piriformis igen. nov. et isp. nov. and M. compactus igen. nov. et isp. nov. A third ichnotaxon is Megaovoidus foveolatus igen. nov. et comb. nov., initially described as “Equisetites” foveolatus G.Roselt from the Late Triassic of Germany. A second new ichnogenus, Paleoincisurus igen. nov., is established for long and narrow slit-like ovipositional damages, which likely housed long rows of insect eggs. The very long scars of P. hellmundi igen. nov. et isp. nov. are associated with sphenophyte stems. For slit-like damages on Sphenopteris sp. cf. S. crepini fronds, the ichnotaxon Paleoincisurus muelleri igen. nov. et isp. nov. is introduced. We also provide evidence for three ichnospecies on leaves of the pteridosperm Macroneuropteris scheuchzeri: Paleoovoidus isp. cf. P. marginatus and two new types of oviposition damage (Paleoovoidus irregularis isp. nov. and P. parvus isp. nov.). Another oviposition damage, provisionally named “Ichnotaxon indet. A”, shows affinities to Paleoovoidus contactus, but is only fragmentarily documented. Finally, exophytic insect eggs, arranged in rows on an unidentified leaf, are described as Palaexovoidus lineatus isp. nov. Our study revealed a high abundance and diversity of endophytic oviposition concerning egg case morphology, oviposition patterns, and host plants of different habitats already in the Middle Pennsylvanian. As a result, a novel, different view on the early evolutionary history of reproductive strategies of late Paleozoic insects is indicated. Eventually, the establishment of an improved ichnotaxobase for oviposition traces according to the latest recommendations for modern ichnotaxonomy is another aim of our study.
Endophytic oviposition, the reproductive behavior of insects to insert eggs into dead or living p... more Endophytic oviposition, the reproductive behavior of insects to insert eggs into dead or living plant tissue, and exophytic oviposition, the deposition of eggs on plants, are only rarely documented from the Pennsylvanian. In the present paper, ten oviposition types from the Piesberg quarry near Osnabrück, Germany (Osnabrück Formation, Middle Pennsylvanian, latest Moscovian, Asturian/Westphalian D) are described. The most abundant are large oviposition scars on sphenophyte stems, which probably housed single eggs. For these, we introduce Megaovoidus igen. nov., including the ichnospecies Megaovoidus piriformis igen. nov. et isp. nov. and M. compactus igen. nov. et isp. nov. A third ichnotaxon is Megaovoidus foveolatus igen. nov. et comb. nov., initially described as “Equisetites” foveolatus G.Roselt from the Late Triassic of Germany. A second new ichnogenus, Paleoincisurus igen. nov., is established for long and narrow slit-like ovipositional damages, which likely housed long rows of insect eggs. The very long scars of P. hellmundi igen. nov. et isp. nov. are associated with sphenophyte stems. For slit-like damages on Sphenopteris sp. cf. S. crepini fronds, the ichnotaxon Paleoincisurus muelleri igen. nov. et isp. nov. is introduced. We also provide evidence for three ichnospecies on leaves of the pteridosperm Macroneuropteris scheuchzeri: Paleoovoidus isp. cf. P. marginatus and two new types of oviposition damage (Paleoovoidus irregularis isp. nov. and P. parvus
isp. nov.). Another oviposition damage, provisionally named “Ichnotaxon indet. A”, shows affinities to Paleoovoidus contactus, but is only fragmentarily documented. Finally, exophytic insect eggs, arranged in rows on an unidentified leaf, are described as Palaexovoidus lineatus isp. nov. Our study revealed a high abundance and diversity of endophytic oviposition concerning egg case morphology, oviposition patterns, and host plants of different habitats already in the Middle Pennsylvanian. As a result, a novel, different view on the early evolutionary history of reproductive strategies of late Paleozoic insects is indicated. Eventually, the establishment of an improved ichnotaxobase for oviposition traces according to the latest recommendations for modern ichnotaxonomy is another aim of our study.
Arthropod borings are commonly described from pteridophyte and gymnosperm wood in the late Palaeo... more Arthropod borings are commonly described from pteridophyte and gymnosperm wood in the late Palaeozoic, but they are almost unknown from calamitalean stems. In this paper, a new type of boring in calamitalean stems from two German localities is reported. These are the mine dumps of Plötz near Halle (Saale), Saale Basin, (Wettin Subformation, Latest Pennsylvanian, Gzhelian, Stephanian C) and the Piesberg quarry near Osnabrück, Lower Saxony Basin, (Osnabrück Formation, Middle Pennsylvanian, latest Moscovian, Asturian/Westphalian D). Most borings were found in marginal parenchyma of the internodes. They run longitudinally through the pith. The borings are preserved as three-dimensional casts that protrude on the surface of considerably compressed pith casts. This unique preservation of the boring casts required special taphonomic conditions, such as rapid burial coupled with different sediment infilling of the borings and the pith cavity, as well as anoxic conditions to prevent decomposition of the non-resistant parenchyma. Most borings are between 3 to 5 mm wide and contain two classes of invertebrate coprolites: the smaller coprolites are sub-spheroidal in shape and measure 37 to 74 x 37 to 63 μm in diameter. The larger coprolites are also sub-spheroidal and range between 88 to 158 x 68 to 123 μm in diameter. The coprolites, the morphology of the borings, as well as the feeding strategy of the arthropods, suggest that the tunnel system was most likely produced by small millipedes (Myriapoda) and probably also visited by oribatid mites.
Sandiella herbigi sp. nov. is the first record of the family Eugereonidae Handlirsch, 1906 (Insec... more Sandiella herbigi sp. nov. is the first record of the family Eugereonidae Handlirsch, 1906 (Insecta: Palaeodictyoptera) in the Pennsylvanian (Late Carboniferous: Westphalian D) sequence of the Piesberg quarry near Osnabrück, Lower Saxony, Germany. It is represented by a single fragment of a mesothoracic wing with the typical coarse reticulation in this family. The species is mainly characterized by the following features: (i) the shape of the mesothoracic wing is extremely narrow, (ii) the posterior subcostal vein is ending before the wing apex, (iii) all the main veins show a typical bend close to the posterior margin. The new wing is the second most ancient record of the Eugereonidae.
Piesberg quarry is famous for its Upper Carboniferous plant and arthropod fossils, including seve... more Piesberg quarry is famous for its Upper Carboniferous plant and arthropod fossils, including several holotypes of flying insects. The high degree of maturity of the Piesberg strata, such as the presence of anthracitic coal, quartzite, and large quartz crystals, led to controversies over a possible underlying thermal anomaly. The Piesberg is of further importance for correlation between the deep underground of northwestern Germany and the Ruhr basin coal field, as well as the Pennsylvanian coal areas in North America, and for investigations of Upper Carboniferous tight gas fields. The importance of the Piesberg for international geosciences has been enabled through its very rich mining history beginning in the Middle Ages and its long scientific history beginning at the end of the 18th century. Today, the Piesberg is not only one of the largest active quarry sites in Europe, but also a local recreation and hiking area. While the Museum am Scholerberg in Osnabruck protects its paleont...
Arthropod borings are commonly described from pteridophyte and gymnosperm wood in the late Palaeo... more Arthropod borings are commonly described from pteridophyte and gymnosperm wood in the late Palaeozoic, but they are almost unknown from calamitalean stems. In this paper, a new type of boring in calamitalean stems from two German localities is reported. These are the mine dumps of Plötz near Halle (Saale), Saale Basin, (Wettin Subformation, Latest Pennsylvanian, Gzhelian, Stephanian C) and the Piesberg quarry near Osnabrück, Lower Saxony Basin, (Osnabrück Formation, Middle Pennsylvanian, latest Moscovian, Asturian/Westphalian D). Most borings were found in marginal parenchyma of the internodes. They run longitudinally through the pith. The borings are preserved as three-dimensional casts that protrude on the surface of considerably
compressed pith casts. This unique preservation of the boring casts required special taphonomic conditions, such as rapid burial coupled with different sediment infilling of the borings and the pith cavity, as well as anoxic conditions to prevent decomposition of the non-resistant parenchyma. Most borings are between 3 to 5 mm wide and contain two classes of invertebrate coprolites: the smaller coprolites are sub-spheroidal in shape and measure 37 to 74 x 37 to 63 μm in diameter. The larger coprolites are also sub-spheroidal and range between 88 to 158 x 68 to 123 μm in diameter. The coprolites, the morphology of the borings, as well as the feeding strategy of the arthropods, suggest that the tunnel system was most likely produced by small millipedes (Myriapoda) and probably also visited by oribatid mites.
Specimens of Euproops sp. (Xiphosura, Chelicerata) from the Carboniferous Piesberg quarry near Os... more Specimens of Euproops sp. (Xiphosura, Chelicerata) from the Carboniferous Piesberg quarry near Osnabrück, Germany, represent a relatively complete growth series of 10 stages. Based on this growth sequence, morphological changes throughout the ontogeny can be identified. The major change affects the shape of the epimera of the opisthosoma. In earlier stages, they appear very spine-like, whereas in later stages the bases of these spine-like structures become broader; the broadened bases are then successively drawn out distally. In the most mature stage known, the epimera are of trapezoidal shape and approach each other closely to form a complete flange around the thoracetron (=fused tergites of the opisthosoma). These ontogenetic changes question the taxonomic status of different species of Euproops, as the latter appear to correspond to different stages of the ontogenetic series reconstructed from the Piesberg specimens. This means that supposed separate species could, in fact, represent different growth stages of a single species. It could alternatively indicate that heterochrony (=evolutionary change of developmental timing) plays an important role in the evolution of Xiphosura. We propose a holomorph approach, i.e., reconstructing ontogenetic sequences for fossil and extant species as a sound basis for a taxonomic, phylogenetic, and evolutionary discussion of Xiphosura.
Uploads
isp. nov.). Another oviposition damage, provisionally named “Ichnotaxon indet. A”, shows affinities to Paleoovoidus contactus, but is only fragmentarily documented. Finally, exophytic insect eggs, arranged in rows on an unidentified leaf, are described as Palaexovoidus lineatus isp. nov. Our study revealed a high abundance and diversity of endophytic oviposition concerning egg case morphology, oviposition patterns, and host plants of different habitats already in the Middle Pennsylvanian. As a result, a novel, different view on the early evolutionary history of reproductive strategies of late Paleozoic insects is indicated. Eventually, the establishment of an improved ichnotaxobase for oviposition traces according to the latest recommendations for modern ichnotaxonomy is another aim of our study.
compressed pith casts. This unique preservation of the boring casts required special taphonomic conditions, such as rapid burial coupled with different sediment infilling of the borings and the pith cavity, as well as anoxic conditions to prevent decomposition of the non-resistant parenchyma. Most borings are between 3 to 5 mm wide and contain two classes of invertebrate coprolites: the smaller coprolites are sub-spheroidal in shape and measure 37 to 74 x 37 to 63 μm in diameter. The larger coprolites are also sub-spheroidal and range between 88 to 158 x 68 to 123 μm in diameter. The coprolites, the morphology of the borings, as well as the feeding strategy of the arthropods, suggest that the tunnel system was most likely produced by small millipedes (Myriapoda) and probably also visited by oribatid mites.
isp. nov.). Another oviposition damage, provisionally named “Ichnotaxon indet. A”, shows affinities to Paleoovoidus contactus, but is only fragmentarily documented. Finally, exophytic insect eggs, arranged in rows on an unidentified leaf, are described as Palaexovoidus lineatus isp. nov. Our study revealed a high abundance and diversity of endophytic oviposition concerning egg case morphology, oviposition patterns, and host plants of different habitats already in the Middle Pennsylvanian. As a result, a novel, different view on the early evolutionary history of reproductive strategies of late Paleozoic insects is indicated. Eventually, the establishment of an improved ichnotaxobase for oviposition traces according to the latest recommendations for modern ichnotaxonomy is another aim of our study.
compressed pith casts. This unique preservation of the boring casts required special taphonomic conditions, such as rapid burial coupled with different sediment infilling of the borings and the pith cavity, as well as anoxic conditions to prevent decomposition of the non-resistant parenchyma. Most borings are between 3 to 5 mm wide and contain two classes of invertebrate coprolites: the smaller coprolites are sub-spheroidal in shape and measure 37 to 74 x 37 to 63 μm in diameter. The larger coprolites are also sub-spheroidal and range between 88 to 158 x 68 to 123 μm in diameter. The coprolites, the morphology of the borings, as well as the feeding strategy of the arthropods, suggest that the tunnel system was most likely produced by small millipedes (Myriapoda) and probably also visited by oribatid mites.