Holometabolan larvae are a major part of the animal biomass and an important food source for many animals. Many larvae evolved anti-predator strategies and some of these can even be recognized in ...fossils. A Lagerstätte known for well-preserved holometabolan larvae is the approximately 100-million-year-old Kachin amber from Myanmar. Fossils can not only allow to identify structural defensive specializations, but also lifestyle and even behavioral aspects. We review here the different defensive strategies employed by various holometabolan larvae found in Kachin amber, also reporting new cases of a leaf-mining hymenopteran caterpillar and a hangingfly caterpillar with extensive spines. This overview demonstrates that already 100 million years ago many modern strategies had already evolved in multiple lineages, but also reveals some cases of now extinct strategies. The repetitive independent evolution of similar strategies in distantly related lineages indicates that several strategies evolved convergently as a result of similar selective pressures.
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•Review of different defensive strategies of holometabolan larvae in Kachin amber•New cases including hymenopteran and hangingfly caterpillars were found•Modern and now extinct strategies were present in Cretaceous in multiple lineages•Strategies may have evolved convergently as result of similar selective pressures
Entomology; Evolutionary biology; Paleobiology
Diptera are one of the four megadiverse groups of holometabolan insects. Flies perform numerous ecological functions, especially in their larval stages. We can assume that this was already the case ...in the past; however, fly larvae remain rare in most deposits. Here we report new dipteran larvae preserved in Cretaceous (about 99 Ma) Kachin amber from Myanmar and, even older, Jurassic (about 165 Ma) compression fossils from China. Through light microscopy and micro-CT scanning we explore their peculiar morphology and discuss their possible phylogenetic affinities. Several larvae seem to represent the lineage of Stratiomyomorpha. A few others present characters unique to Xylophagidae (awl-flies), as well as to Athericidae (water sniper-flies), resulting in a chimeric morphology. Understanding the exact relationships of most of these specimens with a particular lineage remains challenging, since they differ considerably from any other known dipteran larvae and present some unique traits. Additionally, we report new specimens of
Chen et al., 2014, supposedly parasitic larvae, most likely representatives of Athericidae. These new findings offer valuable insights into the evolution of the early diversification of the brachyceran flies and underscore the importance of immature stages in understanding the evolutionary history and ecology of flies.
Mantodeans or praying mantises are flying insects and well known for their raptorial behaviour, mainly performed by their first pair of thoracic appendages. We describe here a new, exceptionally ...preserved specimen of the early mantodean
Grimaldi, 2003 from the famous 110 million years old Crato Formation, Brazil. The incomplete specimen preserves important morphological details, which were not known in this specific form before for this species or any other representative of Mantodea. Unlike in modern representatives or other fossil forms of Mantodea not only the first pair of thoracic appendages shows adaptations for predation. The femora of the second pair of thoracic appendages bear numerous strong, erect spines which appear to have a sharp tip, with this strongly resembling the spines of the first pair of thoracic appendages. This indicates that individuals of
likely used at least two pairs of thoracic appendages to catch prey. This demonstrates that the prey-catching behaviour was more diverse in early forms of praying mantises than anticipated.
Representatives of Schizomida, known as short-tailed whip scorpions, are an understudied group within Megoperculata. They are found subterraneanly in tropical and sub-tropical regions. They lack ...eyes, have superior mobility, and possess a flagellum which is relevant to their mating, but in general little is known about their general biology. Fossil representatives of Schizomida from a variety of time periods are available for study. Using 23 fossil specimens (14 of which are described here for the first time) and 86 extant individuals from the literature, changes in both body size and flagella shape in Schizomida were compared over time. Measurements of prosoma length and leg length were used as a proxy for body size, and a comparative size analysis was carried out. Individuals from the Cretaceous period were found to be significantly smaller than extant individuals, contrary to our expectations. For flagellum shape, images were used to create reconstructions, which were then analysed using an elliptic Fourier analysis followed by a principal component analysis. The morphological diversity of the male flagellum shape was found to have decreased between the Cretaceous and modern fauna. We discuss potential explanations for our findings, although a greater understanding of the general biology of Schizomida is required to properly interpret our results.
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.
Elmidae, riffle beetles, have both adult and immature stages that show specializations for water environments. Fossils of adults of Elmidae are already known from amber, however a record of immatures ...was so far lacking. We report here the first fossil larva of Elmidae, preserved in Baltic amber. To be able to access details of the body hidden by inclusions and "Verlumung" we conducted, in addition to optical documentation methods, micro-CT and synchrotron documentation methods. The larva is characterised by prominent dorso-lateral and lateral processes and a plate-like ventral operculum at the end of the abdomen. The new fossil has similarities in the general body shape and the prominent characters with some modern larvae of Elmidae. The posterior protrusions on the trunk end possibly represent gills, which would imply that fossil larvae of Elmidae also led a water-related life style similar to modern representatives.
Patterned transcranial magnetic stimulation (TMS) such as theta burst stimulation (TBS) or quadri-pulse stimulation (QPS) can induce changes in cortico-spinal excitability, commonly referred to as ...long-term potentiation (LTP)-like and long-term depression (LTD)-like effects in human motor cortex (M1). Here, we aimed to test the plasticity-inducing capabilities of a novel protocol that merged TBS and QPS. 360 bursts of quadri-pulse TBS (qTBS) were continuously given to M1 at 90% of active motor threshold (1440 full-sine pulses). In a first experiment, stimulation frequency of each burst was set to 666 Hz to mimic the rhythmicity of the descending cortico-spinal volleys that are elicited by TMS (i.e., I-wave periodicity). In a second experiment, burst frequency was set to 200 Hz to maximize postsynaptic Ca2+ influx using a temporal pattern unrelated to I-wave periodicity. The second phase of sinusoidal TMS pulses elicited either a posterior-anterior (PA) or anterior-posterior (AP) directed current in M1. Motor evoked potentials (MEPs) were recorded before and after qTBS to probe changes in cortico-spinal excitability. PA-qTBS at 666 Hz caused a decrease in PA-MEP amplitudes, whereas AP-qTBS at 666 Hz induced an increase in mean AP-MEP amplitudes. At a burst frequency of 200 Hz, PA-qTBS and AP-qTBS produced an increase in cortico-spinal excitability outlasting for at least 60 minutes in PA- and AP-MEP amplitudes, respectively. Continuous qTBS at 666 Hz or 200 Hz can induce lasting changes in cortico-spinal excitability. Induced current direction in the brain appears to be relevant when qTBS targets I-wave periodicity, corroborating that high-fidelity spike timing mechanisms are critical for inducing bi-directional plasticity in human M1.
Larvae of malacostracan crustaceans represent a large fraction of modern day zooplankton. Plankton is not only a major part of the modern marine ecosystem, but must have played an important role in ...the ecosystems of the past as well. Unfortunately, our knowledge about plankton composition of the past is still quite limited. As an important part of today's zooplankton, malacostracan larvae are still a rarity in the fossil record; many types of malacostracan larvae dominating the modern plankton have so far not been found as fossils. Here we report a new type of fossil malacostracan larva, found in the 150 million years old lithographic limestones of southern Germany (Solnhofen Lithographic Limestones). The three rather incomplete specimens mainly preserve the telson. A pronounced middle spine on the posterior edge of these specimens indicates that they are either larval forms of a clawed lobster or of an axiidean lobster, or of a closer relative to one of the two groups. The tergo-pleura are drawn out into distinct spines in one specimen, further supporting the interpretation as a larva of a clawed lobster or an early relative. The telson morphology also shows adaptations to a prolonged planktic life style, the latero-posterior edges are drawn out into distinct spines. Similar adaptations are known in larvae of the modern homarid lobster
, not necessarily indicating a closer relationship, but convergent life styles. The new finds provide an important new insight into the composition of Mesozoic zooplankton and demonstrate the preservation potential of lithographic limestones.
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