Early Pennsylvanian Lagerstätte reveals a diverse ecosystem on a subhumid, alluvial fan.

Much of what we know about terrestrial life during the Carboniferous Period comes from Middle Pennsylvanian (~315-307 Mya) Coal Measures deposited in low-lying wetland environments1-5. We know relatively little about terrestrial ecosystems from the Early Pennsylvanian, which was a critical interval for the diversification of insects, arachnids, tetrapods, and seed plants6-10. Here we report a diverse Early Pennsylvanian trace and body fossil Lagerstätte (~320-318 Mya) from the Wamsutta Formation of eastern North America, distinct from coal-bearing deposits, preserved in clastic substrates within basin margin conglomerates. The exceptionally preserved trace fossils and body fossils document a range of vertebrates, invertebrates and plant taxa (n = 131), with 83 distinct foliage morphotypes. Plant-insect interactions include what may be the earliest evidence of insect oviposition. This site expands our knowledge of early terrestrial ecosystems and organismal interactions and provides ground truth for future phylogenetic reconstructions of key plant, arthropod, and vertebrate groups.

Endophytic ancestors of modern leaf miners may have evolved in the Late Carboniferous.

Endophytic feeding behaviors, including stem borings and galling, have been observed in the fossil record from as early as the Devonian and involve the consumption of a variety of plant (and fungal) tissues. Historically, the exploitation of internal stem tissues through galling has been well documented as emerging during the Pennsylvanian (c. 323-299 million years ago (Ma)), replaced during the Permian by galling of foliar tissues. However, leaf mining, a foliar endophytic behavior that today is exhibited exclusively by members of the four hyperdiverse holometabolous insect orders, has been more sparsely documented, with confirmed examples dating back only to the Early Triassic (c. 252-250 Ma). Here, we describe a trace fossil on seed-fern foliage from the Rhode Island Formation of Massachusetts, USA, representing the earliest indication of a general, endophytic type of feeding damage and dating from the Middle Pennsylvanian (c. 312 Ma). Although lacking the full features of Mesozoic leaf mines, this specimen provides evidence of how endophytic mining behavior may have originated. It sheds light on the evolutionary transition to true foliar endophagy, contributes to our understanding of the behaviors of early holometabolous insects, and enhances our knowledge of macroevolutionary patterns of plant-insect interactions.

Comment on Marden (2013): "reanalysis and experimental evidence indicate that the earliest trace fossil of a winged insect was a surface skimming neopteran".

Marden's (2013) reanalysis of Knecht et al. (2011) suggesting that specimen SEMC-F97 is the result of the skimming behavior of a neopteran insect and, more importantly, fossil evidence of "… surface skimming as a precursor to the evolution of flight in insects" (Marden 2013) is found to be deficient on three fronts: (1) the principal specimen was never viewed firsthand which led to significant morphological misinterpretations; (2) poorly designed and executed neoichnological experiments led to incredulous results; and (3) the assumption that this specimen is fossil evidence supporting the surface skimming hypothesis of the origin of insect flight despite the fact that since its induction into the literature that hypothesis has been refuted based on significant paleontological, phylogenetic, genetic, and developmental evidence.

Late Carboniferous paleoichnology reveals the oldest full-body impression of a flying insect.

Insects were the first animals to evolve powered flight and did so perhaps 90 million years before the first flight among vertebrates. However, the earliest fossil record of flying insect lineages (Pterygota) is poor, with scant indirect evidence from the Devonian and a nearly complete dearth of material from the Early Carboniferous. By the Late Carboniferous a diversity of flying lineages is known, mostly from isolated wings but without true insights into the paleoethology of these taxa. Here, we report evidence of a full-body impression of a flying insect from the Late Carboniferous Wamsutta Formation of Massachusetts, representing the oldest trace fossil of Pterygota. Through ethological and morphological analysis, the trace fossil provides evidence that its maker was a flying insect and probably was representative of a stem-group lineage of mayflies. The nature of this current full-body impression somewhat blurs distinctions between the systematics of traces and trace makers, thus adding to the debate surrounding ichnotaxonomy for traces with well-associated trace makers.