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Abstract 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. more...

Abstract Brachyura and Anomala (or Anomura), also referred to as true and false crabs, form the species-rich and globally abundant group of Meiura, an ingroup of Decapoda. The evolutionary success of both groups is sometimes attributed to the process of carcinization (evolving a crab-like body), but might also be connected to the megalopa, a specific transitional larval phase. We investigate these questions, using outline analysis of the shields (carapaces) of more than 1500 meiuran crabs. We compare the morphological diversity of different developmental phases of major ingroups of true and false crabs. We find that morphological diversity of adults is larger in false crabs than in true crabs, indicating that taxonomic diversity and morphological diversity are not necessarily linked. The increasing morphological disparity of adults of true and false crabs with increasing phylogenetic distance furthermore indicates diverging evolution of the shield morphology of adult representatives of Meiura. Larvae of true crabs also show larger diversity than their adult counterparts, highlighting the importance of larvae for biodiversity studies. The megalopa phase of Meiura appears to be plesiomorphic, as it overlaps between true and false crabs and shows little diversity. Causes may be common evolutionary constraints on a developmental phase specialized for transitioning. more...

Modern insect decline sparks interest in similar declines of the past. Neuroptera, the group of lacewings, has been suggested to have undergone decline after the Cretaceous, and quantitative studies of larval morphology have supported this view. One of the groups within Neuroptera seeing a major decline is that of split-footed lacewings, Nymphidae. We here report 13 new larval specimens of split-footed lacewings from about 100-million-year-old Kachin amber, Myanmar. We explore how enlarging the data set to 60 specimens and expanding the investigated morphological structures affect the recognition of decline. The expanded data set has an increased morphospace, indicating a lack of saturation. The decline is, therefore, likely even larger than so far anticipated for Nymphidae. The study of head and mouthparts provides the strongest signal for a loss of diversity, which is feasible due to the direct coupling to feeding ecology. Analysis of the trunk region cannot recover a signal for loss of diversity. Sampling issues play a role here: the trunk of the larvae is less often well preserved, and the fossil larvae are more likely in early developmental stages due to the constraints of preservation. more...

Abstract Convergent evolution is a common phenomenon, independently leading to similar morphologies in different evolutionary lineages. Often similar functional demands drive convergent evolution. One example is the independent evolution of grasping structures in different lineages of Euarthropoda, though the exact morphology of these grasping structures varies significantly. In this study, we investigated grasping apparatuses with two movable counteracting structures as well as some related structures, exemplified by the stylets (compound structures of mouthparts) of aphidlion-like larvae (part of Neuroptera or lacewings) and the cerci of immature earwigs (Dermaptera). For the stylets of aphidlion-like larvae, studies have pointed to a significant loss in morphological diversity in the last 100 million years. We used quantitative morphology to evaluate if a similar process has also occurred in the cerci of earwigs. The cerci of extant immature earwigs exhibit two distinct types of morphologies in the modern fauna: elongated cerci divided into several ringlets with a feeler-type function, and pincer-like stout cerci. In some fossil immature earwigs, however, the cerci are generally elongated but undivided and roughly occupy the morphospace between those of the two modern cerci types; hence this fossil cerci morphology appears to have been lost. To some extent, a comparable loss is also found in certain lacewing larvae. Outgroup comparisons suggest that the morphologies no longer present today are in fact not ancestral, but instead specialised, hence their loss is possibly resulting from disruptive evolution in earwigs as well as lacewings. We discuss the possible functions of these specialised grasping structures. more...

Beetle larvae represent important components of the modern-day fauna. This should have been the case in the past as well. Yet, fossil beetle larvae are rare, or at least are rare in the literature, as identifying a beetle larva to a narrower taxonomic group is very challenging. This is even more complicated if prominent features have evolved convergently in several lineages. Yet, even in such cases, an ecological interpretation of the fossils is possible if the convergent character is coupled to a specific life habit. For example, different, not closely related, beetle larvae that possess setiferous processes. We here report on three beetle larvae, one from Miocene Mexican and two from Cretaceous Kachin amber, Myanmar. These larvae possess setiferous processes, most similar to the processes of modern representatives of Cucujiformia, especially of the groups Endomychidae, Erotylidae, Cerylonidae and Coccinellidae. Considering the shape of the entire habitus, we see the most similarities between the new larvae and the modern larvae of Endomychidae. However, the new larvae and the larvae of modern representatives differ in certain aspects, most prominently in the body size. The fossils are smaller than their extant counterparts with setiferous processes. Hence the fossils could represent larvae of Endomychidae, but the case remains unclear. Despite this uncertainty, we suggest a lifestyle of the fossil larvae as fungus-eaters on rotting wood. This lifestyle is not only known from extant larvae of Endomychidae, but also from other larvae with similar processes. more...

The larval life phase often represents the major share of the life span of holometabolans, such as beetles or flies. Therefore, knowledge of these larvae is crucial concerning ecological functions, but also concerning the evolutionary history of the different groups. In the holometabolan group Neuroptera (lacewings), larvae have numerous specialisations, among them the venom-injecting mouthparts, the stylets. Another such very characteristic feature of many lacewing larvae is the empodium, an attachment structure at the tip of the leg, which varies in shape in the larvae of the different lacewing groups. One type of empodium is the elongate trumpet-shaped type, which is found in several lacewing groups today as well as in many fossil representatives, especially well visible due to preservation in amber. Based on the pattern of phylogenetic distribution of the elongate trumpet-shaped empodium, different reconstructions of the evolution of this structure have been put forward, some suggesting possibly convergent evolution, others repetitive losses. In this study, new lacewing larvae preserved in 100-million-year-old amber from Myanmar are presented, which expand the number of lineages with an elongate trumpet-shaped empodium. Based on these new data, the implications on the evolutionary history of the different lineages within Neuroptera are discussed. The fossils demand for an earlier evolution in some reconstructions, but especially for more and later losses of this structure. more...

Summary: 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. more...

Abstract Loss of biodiversity and especially insect decline are widely recognised in modern ecosystems. This decline has an enormous impact due to the crucial ecological roles of insects as well as their economic relevance. For comparison, the fossil record can provide important insights on past biodiversity losses. One group of insects, for which a significant decline over the last 100 million years has often been postulated, but not demonstrated quantitatively, is Neuroptera (lacewings). Many adult lacewings are pollinators, while the larvae are mostly predators, which becomes very obvious from their prominent stylet-like mouthparts. We investigated the fossil record of larvae of all neuropteran lineages as well as a large share of extant neuropteran larvae. Based on these, we performed an outline analysis of the head with stylets. This analysis provides a quantitative frame for recognising the decline of lacewings since the Cretaceous, indicating also a severe loss of ecological roles. more...

Lacewing larvae in the Cretaceous were more diverse in appearance than they are today, best documented by numerous fossils preserved in amber. One morphotype of an unusual larva from about 100 Ma old Kachin amber (Myanmar) was formally recognised as a distinct group called Ankyloleon. The original description erected a single formal species, Ankyloleon caudatus. Yet, it was indicated that among the five original specimens, more species were represented. We here report five new specimens. Among these is the so far largest as well as the so far smallest specimen. Based on this expanded material we can estimate certain aspects of the ontogenetic sequence and are able to recognise a second discrete species, Ankyloleon caroluspetrus sp. nov. We discuss aspects of the biology of Ankyloleon based on newly observed details such as serrations on the mandibles. Long and slender mouthparts, legs and body together with a weakly expressed outer trunk segmentation provide indications for a lifestyle hunting for prey in more confined spaces. Still many aspects of the biology of these larvae must remain unclear due to a lack of a well comparable modern counterpart, emphasising how different the fauna of the Cretaceous was. more...

IntroductionEuchelicerata is a diverse group encompassing Xiphosura, Chasmataspidida, Eurypterida, and Arachnida. Xiphosura represents an extant group with a rich fossil record dating back to the Ordovician period. Xiphosurans are often referred to as “living fossils” due to their seemingly unchanged morphology over millions of years. Numerous studies have contributed to the understanding of xiphosuran development, revealing changes in the timing and rate of their growth. These changes have been mainly associated with the freshwater invasion of early xiphosuran forms. However, limited research has been conducted to compare the developmental patterns of xiphosurans with other euchelicerates inhabiting aquatic environments.MethodsThis study compares the developmental patterns of xiphosurans with that of the fossil clades of eurypterids and chasmataspidids. By incorporating environmental and phylogenetic information within ancestral state reconstruction analyses, and then testing different evolutionary scenarios, the influence of the environment on the evolution of developmental patterns of euchelicerates is examined.ResultsThe results confirm that the developmental changes in Xiphosura throughout their evolutionary history are correlated with the exploitation of different environments. However, the inclusion of eurypterids and chasmataspidids indicates that the entirety of changes seen for Xiphosura represent only a small portion of the total variability recovered for euchelicerates.DiscussionOur results emphasize the importance of considering phylogenetic relationships and outgroup comparisons to understand the evolutionary dynamics of Xiphosura. more...

Resin is a plastic-like product of trees. Older occurrences of such resin are referred to as amber and are considered fossil resin. Younger resins are termed copals. Even younger ones have been dubbed defaunation resins. Non-fossil resins remain in a terminological limbo, often referred to as “sub-fossils”. We report two lepidopteran caterpillars preserved in non-fossil resin: one from Madagascar, one from Brazil. Prominent hairs (=setae) and spines (=spine-like setae) of the specimens make it likely that they represent larvae of Erebidae (e.g., tussock moths and others). So far, most known caterpillars preserved in resins are either “naked” or bear protective cases; only few are armoured with spines or hairs. In particular, long-haired caterpillars such as the ones reported here are so far almost absent. Only one specimen with comparable setae has been reported from 15-million-year-old Dominican amber, but no significant details of this specimen are accessible. We briefly also review the record of caterpillars known from the Holocene, recognising that it is very sparse. The new specimens demonstrate that very hairy caterpillars can readily be preserved in resins in fine detail. Furthermore, the specimens increase the known size range of caterpillars preserved in resins, with one measuring more than 12 mm. more...

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 Qiyia jurassica 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. more...

Abstract Physogastry is a phenomenon occurring in Euarthropoda and describes an extreme inflation of (parts of) the trunk. It is best known from ticks, termite queens, or honey-pot ants, but can also be found in several other representatives of Euarthropoda. Physogastry has so far rarely been seen in the fossil record. We describe here an example of physogastry in two lacewing larvae (Neuroptera) enclosed in a single piece of Kachin amber (ca. 100 Ma old). We measured head and trunk ratios of different physogastric and non-physogastric representatives of Euarthropoda. Plotting these ratios shows that the new larvae, which display quite extremely inflated trunks, are very similar to ticks or honey-pot ants, but also to certain lacewing larvae of the group Berothidae (beaded lacewings). Outline analysis of head capsule and mouthparts (stylets) further suggests a position within Berothidae. Physogastry is presumed to be linked with living in confined spaces such as wood galleries or soil. Indeed, at least some larvae of Berothidae are known to live inside termite nests for part of their larval life phase, a habit the new larvae may also have had. The new record represents the oldest case of extreme physogastry in insects known to date. more...

Mantis lacewings (Neuroptera: Mantispidae) are prominent and charismatic predatory representatives of Insecta. Nevertheless, representatives of the group are surprisingly scarce in Paleogene deposits after a relative abundance of specimens known from Cretaceous. Here we present Mantispa? damzenogedanica sp. nov., representing the first adult of Mantispidae described from Baltic amber and the only Eocene adult mantispid hitherto preserved in amber. The new fossil species is also among the earliest representatives of Mantispinae, certainly the oldest adult of this group described from amber. Additionally, we discuss the changes through time in the ecological morphospace within Mantispidae based on the morphological diversity (≈disparity) of the raptorial legs. Possible explanations for the post-Cretaceous decline in the morphological diversity of mantis lacewings are posited. more...

During ontogenetic development, insects can undergo quite drastic changes (metamorphosis) until the adult stage is reached. A substantial part of this development in one group of Insecta, Holometabola, takes place during the pupa stage. Despite the pupa being recognised as an important phase, rather few depictions of pupae exist in the literature. We report here the first find of a fossil pupa of the lacewing group Mantispidae. The specimen represents an exuvia and is enclosed in Ukrainian Rovno amber, Eocene in age (c. 35–40 million years). We review the entire record of extant pupae of Mantispidae depicted in the literature or in online image repositories. With the aid of elliptic Fourier analysis, we compare the outline of the femur of the foreleg (raptorial appendage in the adults) of pupae and adults of Mantispidae. The pupae are all very similar concerning the femur, while the adults show a larger morphological diversity, particularly the extinct forms. Furthermore, our results indicate that the forelegs do not become increasingly complex throughout ontogenetic stages, but instead undergo an indirect development. According to the low variation in morphology seen in the pupa stage in Mantispidae, it is plausible that it represents a phylotypic stage for the group, i.e. a phase characterised by a significantly lower variability than other stages. more...

Abstract Morphological identification of planktic crustacean larvae is required in many scientific contexts, such as ecology or taxonomy. Due to a still low availability of genetic sequences for many ingroups of Eumalacostraca, this task is still more feasible by morphological methods. Our understanding of eumalacostracan larval morphology is challenged by phenotypic variability. We investigated four eumalacostracan ingroups: Galatheidae, Hippoidea, Raninidae and Stomatopoda. Representatives of all four groups develop through spine-bearing planktic larval stages. Incorporating dorsal and lateral shield outlines into three-dimensional shape analysis of the shields, we compare specimens from the wild with laboratory-reared specimens. Using graphical and statistical analysis methods, we find that at least the lateral morphology of the shields of Hippoidea and Raninidae seems to be too strongly dependent on phylogeny to show phenotypic variability with our current sample size, but Hippoidea do show phenotypic variability in their dorsal shield morphology. In Galatheidae and Stomatopoda, a clear difference in shield morphology can be found between wild-caught and laboratory-reared specimens. This difference likely represents phenotypic variability. The exact environmental signals causing this phenotypic variability are still unknown, but some candidates are discussed. more...

Neuroptera, the group of lacewings, is well known to have been more diverse in the past, offering to study patterns of biodiversity loss over time. This loss of diversity has been quantitatively established by the morphological diversity of lacewing larvae. Here, we explore in more detail the diversity of lacewing larvae with tooth-bearing mouthparts. All these larvae are representatives of Myrmeleontiformia, the group of antlion-like lacewings. Today, larvae of several major ingroups bear teeth on their mouthparts: (1) owllions (formerly Ascalaphidae and Myrmeleontidae; taxonomic status is currently unclear); (2) Nymphidae; (3) Crocinae (mostly in younger larvae); and (4) Nemopterinae (only micro teeth). In addition, there are several now extinct larval types with teeth known from Cretaceous ambers (about 100 million years old). These larvae also possess several plesiomorphic characters, indicating that they were part of the early diversification of Myrmeleontiformia. We report numerous new specimens of these now extinct forms and provide a quantitative morphological comparison of head and mouthpart shapes, demonstrating that some of these Cretaceous larvae possessed morphologies not represented in the extant fauna. The resulting pattern is complex, indicating that at least some extinct morphologies have been later replaced by modern-day antlions due to convergent evolution. more...

Abstract Larvae of the group Holometabola (beetles, wasps, flies, moths and others) differ significantly in their morphology from their corresponding adults. In most larvae, appendages and other structures protruding from the body (antennae, palps, legs, trunk processes) appear less elongate than in their corresponding adults, providing the impression that these larvae are restricted to a certain degree in developing more elongate structures. We provide here numerous counterexamples of larvae of lacewings (Neuroptera). These include different forms of elongated antennae, mandibles, maxillae, labial palps, legs, trunk processes and neck regions. Most of these examples are larvae preserved in different types of 100 million-year-old amber. The longest neck region was found in an extant specimen. All these examples demonstrate that certain branches of Neuroptera indeed had larval forms that possessed strongly elongated structures. Hence there is no principal constraint that hinders holometabolan larvae to develop such structures. more...

Nevrorthidae, the group of dragon lacewings, has often been considered a relic group. Today, dragon lacewings show a scattered distribution, with some species occurring in southern Europe, Japan, Australia, and one in China. The idea that this distribution is only a remnant of an originally larger distribution is further supported by fossils of the group preserved in ambers from the Baltic region (Eocene, ca. 35–40 MaBP) and Myanmar (Kachin amber, Cretaceous, ca. 100 MaBP). Larvae of the group are slender and elongated and live mostly in water. Yet, larvae are in fact very rare. So far, only slightly more than 30 larval specimens, counting all extant and fossil larvae, have been depicted in the literature. Here, we report numerous additional specimens, including extant larvae, but also fossil ones from Baltic and Kachin amber. Together with the already known ones, this sums up to over 100 specimens. We analysed quantitative aspects of the morphology of these larvae and compared them over time to identify changes in the diversity. Despite the enriched sample size, the data set is still unbalanced, with, for example, newly hatched larvae (several dozen specimens) only known from the Eocene. We expected little change in larval morphology over geological time, as indicated by earlier studies. However, on the contrary, we recognised morphologies present in fossils that are now extinct. This result is similar to those for other groups of lacewings which have a relic distribution today, as these have also suffered a loss in diversity in larval forms. more...

Abstract Background The Chengjiang biota from southwest China (518-million-years old, early Cambrian) has yielded nearly 300 species, of which more than 80 species represent early chelicerates, crustaceans and relatives. The application of µCT-techniques combined with 3D software (e.g., Drishti), has been shown to be a powerful tool in revealing and analyzing 3D features of the Chengjiang euarthropods. In order to address several open questions that remained from previous studies on the morphology of the xandarellid euarthropod Sinoburius lunaris, we reinvestigated the µCT data with Amira to obtain a different approach of visualization and to generate new volume-rendered models. Furthermore, we used Blender to design 3D models showing aspects of intraspecific variation. Results New findings are: (1) antennulae consist of additional proximal articles that have not been detected before; (2) compared to other appendages, the second post-antennular appendage has a unique shape, and its endopod is comprised of only five articles (instead of seven); (3) the pygidium bears four pairs of appendages which are observed in all specimens. On the other hand, differences between specimens also have been detected. These include the presence/absence of diplotergites resulting in different numbers of post-antennular appendages and tergites and different distances between the tip of the hypostome and the anterior margin of the head shield. Conclusions Those new observations reveal intraspecific variation among Chengjiang euarthropods not observed before and encourage considerations about possible sexual dimorphic pairs or ontogenetic stages. Sinoburius lunaris is a variable species with respect to its morphological characters, cautioning that taxon-specific variabilities need to be considered when exploring new species. more...

Larvae of lacewings (Neuroptera) are known to be fierce predators. According to the morphology of fossil forms thisseems to have been the case already in the Early Cretaceous. While being predators, lacewing larvae are also food itemsfor other organisms. Here we report two pieces of amber from Myanmar providing instances of such cases. In one amberpiece several isolated stylets of lacewing larvae are preserved closely associated together. The most likely interpretationis that a predator preying on lacewing larvae has regurgitated or defecated these non-digestible pieces, yet the identityof the predator remains unclear. The other amber piece preserves a larva resembling modern day larvae of split-footedlacewings (Nymphidae). The larva has projections on its trunk, allowing it to wear a camouflaging cloak. In the headregion, a mite (Acari) is attached to the larva; more precisely, the entire anterior body region of the mite is apparentlyinserted into the lacewing larva. The mite is smaller than the larva. It is known from the modern fauna that stage 1 larvaeof Ascalaphidae can be attacked also by rather small predators, such as ants. The mite can therefore well be interpretedas a true predator instead of a parasite, especially considering the unusual mode of attachment. We briefly review interactionsof lacewing larvae with other organisms represented in amber from Myanmar and add two new pieces to thepuzzle of reconstructing the trophic interactions in the 100-million-year old amber forest. more...

Neuroptera is one of the smaller ingroups of Holometabola, the ingroup of Insecta characterised by “complete” metamorphosis. Neuroptera comprises about 6000 species in the modern fauna, but appears to have been more diverse in the past. While adults distantly resemble certain moths or damselflies, the larval forms of Neuroptera are mostly fierce predators with prominent venom-injecting stylets. The most well-known of these larvae are probably those of antlions. Antlions and their closer relatives (silky lacewings, split-footed lacewings, ribbon-winged lacewings, spoon-winged lacewings, and owlflies) form a distinct monophyletic ingroup of Neuroptera, Myrmeleontiformia, hence the antlion-like forms. The fossil record of antlion-like larvae dates back far into the Cretaceous; many forms are known by exceptionally wellpreserved specimens entrapped in amber. The oldest fossil record of a neuropteran larva (not an antlion-like form) comes from Lebanese amber. Interestingly, the supposedly oldest record of an antlion-like larva is preserved in rock and comes from the famous Lower Cretaceous Crato Formation. We re-evaluate this fossil based on high-resolution composite photography. Due to the non-availability of many key characters, standard procedures for identifying the specimen to a more narrow ingroup remains challenging. Therefore, we used a morphometric approach. A combination of non-metric multidimensional scaling (NMDS), parallel coordinate plots and discriminant function analysis indicates that the fossil is a representative of the group Ascalaphidae (owlflies) + Myrmeleontidae (antlions). We discuss implications of this result for the fossil record of neuropteran larvae. These include the rather derived morphology of the oldest fossil larva of Myrmeleontiformia in contrast to previous expectations. Furthermore, fossils from soil dwellers can not only be expected to be found in amber, but also as compression fossils. more...

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. more...

Beetle larvae contribute to the overall biomass with a great share, yet they often stay unnoticed and underexplored. Larvae of the group Scraptiidae, also called false flower beetles, lead a life hidden in the wood, not easily accessible for observers. There, they contribute to wood decomposition and carbon cycling. Even though their ecological role is of great importance, these larvae have been comparably rarely studied. This is true for extant as well as fossil representatives of this group. It seems that this knowledge gap is not based on the limited availability of material but results from insufficiently studied material. Here we report new specimens, of which seven are extant and twelve are fossil. Fossil specimens are either from 40-million-year-old Baltic amber (Eocene) or 100-million-year-old Myanmar amber (Cretaceous), the latter representing the oldest record of these larvae. All specimens considered here possess a large, elongated terminal end. We performed an outline analysis of the shape of this terminal end for all so far known larval specimens sufficiently well preserved (in total 33 specimens: 17 extant, 14 Eocene, 2 Cretaceous). There is a recognisable difference between Eocene and extant specimens, yet it remains unclear whether this is due to different represented larval stages or an effect of evolution. more...

Larvae of soldier flies and their closest relatives (Diptera: Stratiomyomorpha) are important decomposers of organic material, including wood, that take part in carbon cycling. They also play a certain role in the modern-day animal and human food industry, representing economic value. Larvae of Stratiomyomorpha are considered to be rather rare in the fossil record. Indeed, only very few larvae have been reported so far. Here, we demonstrate that larvae of Stratiomyomorpha are in fact abundant in the Cretaceous, at least in Myanmar amber (about 100 million years old), based on more than 100 amber pieces containing larvae. The specimens could be differentiated into three morphotypes, two already described in a previous work, and a new one. For one morphotype, three larval stages could be distinguished by analysing the dimensions of the head capsules. A quantitative analysis of body shapes indicates a lower morphological diversity of the fossil sample in comparison to the extant fauna, but suggests that they might have had a different ecology in the past. It appears that the data set is not yet saturated, i.e., that more fossil larvae of this group, including different morphologies, are expected to be found. more...

Lacewings have been suggested to be a relict group. This means that the group of lacewings, Neuroptera, should have been more diverse in the past, which also applies to many ingroups of Neuroptera. Psychopsidae, the group of silky lacewings, is one of the ingroups of Neuroptera which is relatively species-poor in the modern fauna. Larvae of the group Psychopsidae, long-nosed antlions, can be easily identified as such in being larvae of antlion-like lacewings without teeth in their stylets (=compound structure of mandible and maxilla), with empodia (=attachment structures on legs) and with a prominent forward-protruding labrum. Therefore, such larvae can also be recognised in the fossil record. An earlier study demonstrated a decline in the morphological diversity of long-nosed antlion larvae over the past 100 million years. Here, we report several dozen new long-nosed antlion larvae and expand the earlier quantitative study. Our results further corroborate the decline of silky lacewings. Yet, a lack of an indication of saturation indicates that we have still not approached the original diversity of long-nosed antlions in the Cretaceous. more...

Summary: Aquatic larvae are known in three early branches of Pterygota: Ephemeroptera (mayflies), Plecoptera (stoneflies), and Odonata (dragonflies, damselflies). A common origin of these larvae has been suggested, yet also counterarguments have been put forward, for example, the different position of larval gills: laterally on the abdomen in Ephemeroptera, terminally in Odonata, variably in Plecoptera. We discuss recent fossil findings and report a new dragonfly-type larva from Kachin amber (Myanmar), which possesses ancestral characters such as a terminal filum, maintained in ephemeropterans, but lost in modern odonatan larvae. The new larva possesses lateral protrusions on the abdominal segments where in other lineages gills occur. Together with other fossils, such as a plecopteran retaining lateral gills on the abdomen, this indicates that lateral protrusions on the abdomen might have well been an ancestral feature, removing one important argument against the idea of an aquatic larva in the ground pattern of Pterygota. more...

Achelatan lobsters, also known as spiny and slipper lobsters, develop via a highly specialised larval form. This special larva, phyllosoma, is flat, translucent, possesses elongate legs and can grow to enormous sizes. Although these larvae may appear very fragile, they are well-known as fossils. Thousands of specimens have been found in the lithographic limestone of Southern Germany (Tithonian, Upper Jurassic, about 150 mya). At least three types of fossil, but modern-appearing phyllosoma larvae are known. Additionally, fossil larvae that possess only some of the characters of modern-day phyllosoma larvae are known from the same Lagerstätte, but also from the younger limestone beds of Lebanon. Here we report a new achelatan fossil from the older Posidonia Shale (Toarcian, Lower Jurassic, 175–183 mya). The specimen shows certain characters of a phyllosoma larva, but other characters appear like those of post-phyllosoma stages of achelatan lobsters. This specimen is therefore the oldest occurrence of an achelatan lobster larva. We compare the new specimen with other fossil larvae with such mixed or “intermetamorphic” morphologies. more...

Abstract Background Biological diversity is a hot topic in current research, especially its observed decrease in modern times. Investigations of past ecosystems offer additional insights to help better understand the processes underlying biodiversity. The Cretaceous period is of special interest in this context, especially with respect to arthropods. During that period, representatives of many modern lineages appeared for the first time, while representatives of more ancient groups also co-occurred. At the same time, side branches of radiating groups with ‘experimental morphologies’ emerged that seemed to go extinct shortly afterwards. However, larval forms, with their morphological diversity, are largely neglected in such studies, but may provide important insights into morphological and ecological diversity and its changes in the past. Results We present here a new fossil insectan larva, a larval lacewing, in Cretaceous amber, exhibiting a rather unusual, ‘experimental’ morphology. The specimen possesses extremely large (in relation to body size) mandibulo-maxillary piercing stylets. Additionally, the labial palps are very long and are subdivided into numerous elements, overall appearing antenniform. In other aspects, the larva resembles many other neuropteran-type larvae. Conclusions We provide a comparison that includes quantitative aspects of different types of neuropteran larvae to emphasise the exceptionality of the new larva, and discuss its possible relationships to known lineages of Neuroptera; possible interpretations are closer relationships to Dilaridae or Osmylidae. In any case, several of the observed characters must have evolved convergently. With this new find, we expand the known morphological diversity of neuropterans in the Cretaceous fauna. more...

Abstract Background Calmanostraca is a group of branchiopod eucrustaceans, with Triops cancriformis and Lepidurus apus as most prominent representatives. Both are regularly addressed with the inaccurate tag “living fossil”, suggesting that the morphology has remained stable for several millions of years. Yet, T. cancriformis and L. apus represent only a fraction of the morphological diversity occurring in Calmanostraca, comprising the two groups Notostraca and Kazacharthra. Notostracans, commonly called tadpole shrimps, comprise the two groups Lepidurus and Triops with their elongated and rather narrow (in dorsal view) head shields. Kazacharthrans are exclusively fossil calmanostracans with broad and rather short shields, known from the Jurassic and Triassic period. One formation where fossil calmanostracans have been found is the Yixian Formation of northeastern China (Lower Cretaceous, 125–121 million years). It is part of the Jehol Group, an ecosystem known for its exceptionally well-preserved fossils, including vertebrates and plants, but also diverse arthropods. Two calmanostracan species have to date been described from the Yixian Formation, Jeholops hongi and Chenops yixianensis. Results We describe here a new calmanostracan crustacean from the Yixian Formation, Notostraca oleseni, and additionally a simple tool using a morphospace analysis to delineate different species. Measurements characterising the shield and trunk proportions of different calmanostracan species were performed, data were size-corrected, and used for this morphospace analysis to compare the different morphologies. As sclerotised body parts are more likely to be preserved in fossils than soft tissue, shields and parts of the trunk are in many cases the only morphological structures available for study. Therefore, the present analysis represents a simple tool for distinguishing between different species, as well as allowing the inclusion of specimens that are only preserved fragmentarily. Additionally, it provides a tool to demarcate the kazacharthran-like specimen described, but not formally named, by Wagner et al. (Paleontol Res. 22:57–63, 2018). Hence, we amended the description and name the species Calmanostraca hassbergella. Conclusion Our results indicate a large diversity in shield and trunk morphology in calmanostracans, in contrast to their often claimed highly conserved and uniform morphology. Especially extinct forms such as Notostraca oleseni add up to this result and point to the species richness and morphological diversity within Calmanostraca. more...

Abstract Representatives of Hippoidea, often called sand crabs or mole crabs, are an ingroup of Anomala. These marine crustaceans inhabit the tropical and subtropical coasts of the world, yet some also appear in temperate climates. Their adults are specialized for digging and living in sandy substrates. Hippoidean zoea-type larvae are planktic and reach large sizes up to a few centimetres. These larvae transform into megalopa larvae, strongly resembling the adult, mediating the transition to the benthic lifestyle of the adult. We reconstructed outlines in dorsal view of over 80 shields of hippoideans, including representatives of Blepharipodidae (sister group to all others), Albuneidae, and Hippidae and including adults, megalopa-type, and zoea-type larvae from all three ingroups. We conducted a morphological analysis on this data using an elliptic Fourier transformation and principal component analysis. We used the results of the analysis to discuss the life history of hippoideans and the special function of megalopae, which often lack emphasis in current research. Early stage zoea larvae, megalopae, and adults show a linear gradient in their morphological development according to our analysis. However, late stage zoea larvae deviate from this pattern, possibly due to their specialization to a long-lasting planktic life. Lastly, we discuss the influence of phenotypic plasticity in hippoidean zoea larvae. more...

The process of interpreting and evaluating a fossil is a difficult task. Isopoda is a species rich group of peracarid eumalacostracans which represent quite a challenge when found as fossils, independent of whether we are working with fragmentary or more complete specimens. Here we describe a new fossil species of crustacean, Platuropodus odysseus n. gen. n. sp., from the Irati Formation, Permian of Paraná Basin, Brazil. After misinterpretations, the fossil taxon is recognised here as a representative of Isopoda. The new species presents characters found in Phreatoicidea, Asellota and Oniscidea in a unique combination for the fossil and extant record, such as two pairs of sub-chelate anterior trunk appendages, a short region after the anus and flat uropods. This chimera-like morphology and a morphometric analysis of the sub-chelae indicate convergent evolution in the early diversification of Isopoda. The morphological diversification present in the Palaeozoic and Mesozoic fossil record of Eumalacostraca indicate a “push of the past” effect in different ingroups of Peracarida. more...

Eumalacostracan crustaceans all have a more or less stereotypic body organisation in the sense of tagmosis. Originally, this included a head with six segments (ocular segment plus five appendage-bearing segments), a thorax region with eight segments, and a pleon with six segments. Interestingly, despite these restrictions in variability in terms of tagmosis, the morphological diversity within Eumalacostraca is rather high. A group providing representative examples that are commonly known is Decapoda. Decapodan crustaceans include shrimp-like forms, lobster-like forms and crab-like forms. The stem species of Eucarida, the group including Decapoda and Euphausiacea, presumably possessed a rather shrimp-like morphology, quite similar to the stem species of Eumalacostraca. Also two other lineages within Eumalacostraca, namely Hoplocarida (with the mantis shrimps as modern representatives) and Neocarida (with the sister groups Thermosbaenacea and Peracarida) evolved from the shrimp-like body organisation to include a lobster-like one. In this study, we demonstrate that the stepwise evolution towards a lobster morphotype occurred to a certain extent in similar order in these three lineages, Hoplocarida, Eucarida and Peracarida, leading to similar types of derived body organisation. This evolutionary reconstruction is based not only on observations of modern fauna, but especially on exceptionally preserved Mesozoic fossils, including the description of a new species of mantis shrimps bridging the morphological gap between the more ancestral-appearing Carboniferous forms and the more modern-appearing Jurassic forms. With this, Mesozoic eumalacostracans represent an important (if not unique) ‘experimental set-up’ for research on factors leading to convergent evolution, the understanding of which is still one of the puzzling challenges of modern evolutionary theory. more...

Within Neuroptera, the group of lacewings, the ingroup Nevrorthidae is special in several aspects: 1) the group may represent the sister group to all other neuropterans; 2) the group is quite species-poor with only 19 extant species known so far; 3) its representatives show a disjunct relict distribution; 4) it has very unusual appearing larvae. These larvae are very elongate, almost worm-shaped. Our overall knowledge of these larval forms is still very limited. We here review all known occurrences of extant larvae, amended by fossil larvae, all preserved in amber. So far there have been only 17 extant larval specimens of Nevrorthidae depicted in the literature. We also present new fossil larvae, with this reaching a total number of 16 known fossil larvae of Nevrorthidae. When plotting measured lengths, larvae cluster into more than three clusters. Also reconstructed size gains between these clusters indicate that Nevrorthidae might in fact develop through more than three larval stages, unlike other lacewings. A special subdivision of the trunk segments observed in many larvae indicates a specialisation for moving in confined spaces. Comparison of syn-inclusions and literature data make it likely that not all larvae lived in running waters, as previously assumed. Overall our knowledge remains rather limited, yet the data allow to draw some new conclusions about the life and evolution of these larval forms. more...

Lacewing larvae are mostly predatory, highly mobile larval forms of Insecta. The modern fauna yields several morphotypes of lacewing larvae, each closely associated with a distinct evolutionary lineage within the group of lacewings, Neuroptera. Back in the Cretaceous, about 100 million years ago, many of these larval forms had already evolved. Additionally, many larval forms seem to have been present that are now extinct. We report here a new form, which appears to be extinct now. This new larva has a prominent forward projecting labrum like larval forms of Nevrorthidae and Psychopsidae. It furthermore has (again similar to the latter two) curved venom-injecting stylets formed by mandibles and maxillae. We used quantitative outline analysis to compare the new larva to those of Nevrorthidae and Psychopsidae. The results of this analysis demonstrate that the new larva differs in all aspects of head capsule shape from all known larvae of Nevrorthidae. Its head shape is more similar to that of many larvae of Psychopsidae, yet also here the new larva differs recognisably in one principal component. Also qualitative differences clearly differentiate the new larva from already known ones. Hence, the new larva represents a new, so far unknown morphotype of lacewing larva. This finding adds to the growing 'zoo' of unusual lacewing larvae back in the Cretaceous, indicating that form diversity and ecological diversity of lacewings were much higher 100 million years ago. more...

When the evolution of Euarthropoda is discussed, often the lineage of Chelicerata s. str. is assumed to be the more ‘primitive’ or ‘basal’ part of the tree, especially when compared to the other major lineage, Mandibulata. This claimed primitiveness is (at least partly) based on the assumption that different morphological structures are still in an ancestral state and did not evolve any further. One of these sets of structures is the feeding apparatus, which has been stated to be highly advanced in Mandibulata, but not ‘properly’ developed, or at least not to such a high degree, within Chelicerata s. str. In this study, I reinvestigate the feeding apparatus of different ingroups of Euchelicerata, with a focus on assumed ‘primitive’ groups such as Eurypterida and Trigonotarbida. The basis of this study is a large amount of material from different museum collections, with fossils with the entire feeding apparatuses being exceptionally well preserved. Based on high-resolution micro-photography and three-dimensional imaging, it is possible to resolve fine details of the feeding apparatuses. The results make clear that the feeding apparatuses of different ingroups of Euchelicerata are highly specialised and often possess morphological structures comparable to those of the feeding apparatuses of representatives of Mandibulata, apparently convergently evolved. Though the reconstruction of the evolution of the feeding apparatus within Euchelicerata is to a certain degree hampered by unclear phylogenetic relationships, there was clearly a shortening of the feeding apparatus from posterior (i.e. only the anterior appendages being involved in the feeding apparatus), probably linked to the colonisation of land in Arachnida. more...

Ant mimicry, i.e., the mimicking of ant workers by another organism, is a widespread phenomenon among different groups of Euarthropoda, including spiders and different insect species. One example of ant mimicry occurs among praying mantises (Mantodea); here the first stage nymphs have been recorded to perform ant mimicry. In this study, we investigated different nymphal instars of Miomantis binotata for possible morphological similarities to ants. The different instars were compared as stages supposed to perform ant mimicry would differ morphologically from those stages not supposed to resemble ants. The specimens were investigated under different microscopic settings and measurements were performed. Our results do not show significant differences concerning morphological measurements or shape of structures between the different nymphal instars of M. binotata. One prominent difference between stage one nymphs and later stages occurs in the colouration of the body, which is very dark in the earliest nymph. This difference might explain why young nymphs of Miomantis binotata are interpreted as ant-mimicking, despite the apparent lack of other morphological resemblances. more...

Holometabola is a hyperdiverse group characterised by a strong morphological differentiation between early post-embryonic stages (= larvae) and adults. Adult forms of Holometabola, such as wasps, bees, beetles, butterflies, mosquitoes or flies, are strongly differentiated concerning their mouth parts. The larvae most often seem to retain rather plesiomorphic-appearing cutting-grinding mouth parts. Here we report a new unusual larva preserved in Burmese amber. Its mouth parts appear beak-like, forming a distinct piercing mouth cone. Such a morphology is extremely rare among larval forms, restricted to those of some beetles and lacewings. The mouth parts of the new fossil are forward oriented (prognathous). Additionally, the larva has distinct subdivisions of tergites and sternites into several sclerites. Also, the abdomen segments bear prominent protrusions. We discuss this unusual combination of characters in comparison to the many different types of holometabolan larvae. The here reported larva is a new addition to the ‘unusual zoo’ of the Cretaceous fauna including numerous, very unusual appearing forms that have gone extinct at the Cretaceous–Palaeogene boundary. more...

Among lacewings (Neuroptera), representatives of the groups Ascalaphidae (owlflies) and Myrmeleontidae (antlions) are likely the most widely known ones. The exact taxonomic status of the two groups remains currently unclear, each may in fact be nested in the other group. Herein, we refer to the group including representatives of both with the neutral term “owllion”. Owllion larvae are voracious ambush hunters. They are not only known in the extant fauna, but also from the fossil record. We report here new findings of a fossil owlfly larva from Eocene Baltic amber, as well as several owlfly-like larvae from Cretaceous Kachin amber, Myanmar. Based on these fossils, combined with numerous fossil and extant specimens from the literature, collections, and databases, we compared the morphological diversity of the head and mouthpart shapes of the larvae of owllions in the extant fauna with that of owllion-like larvae from three time slices: about 100 million years ago (Cretaceous), about 40 million years ago (Eocene), and about 20 million years ago (Miocene). The comparison reveals that the samples from the Eocene and Miocene are too small for a reliable evaluation. Yet, the Cretaceous larvae allow for some conclusions: (1) the larval morphological diversity of owllion larvae increased over time, indicating a post-Cretaceous diversification; (2) certain morphologies disappeared after the Cretaceous, most likely representing ecological roles that are no longer present nowadays. In comparison, other closely related lineages, e.g., silky lacewings or split-footed lacewings, underwent more drastic losses after the Cretaceous and no subsequent diversifications. more...

Abstract Background Adult mantis lacewings, neuropteran holometabolan insects of the group Mantispidae, possess anterior walking legs transformed into prey-catching grasping appendages reminiscent of those of praying mantises. While adult mantis lacewings are hence active “wait-and-catch” predators, the larvae of many mantis lacewings have a quite different biology: first-stage larvae seek out female spiders, mount them, and either wait until the spider has produced an egg sac or, in some cases, choose a female already bearing an egg sac. The larva then enters the egg sac and feeds on the eggs. While first stage larvae are highly mobile with comparably long legs and a certain degree of dorso-ventral flattening (“campodeiform”), larval stages two and three are almost immobile, grub-like, and simply remain within the egg sac. Fossils of mantis lacewings are relatively rare, fossils of larval mantis lacewings are even rarer; only a single larva sitting on a juvenile spider has been described from ca. 50 million year old Baltic amber. Results Here we describe a second occurrence of a larval mantis lacewing from significantly older Burmese amber, about 100 million years old. The specimen is preserved in a position right at the leg of a spider, similar to modern-day larvae that are about to mount their prospective host. The claws of the larva can be seen to grab around the leg of the spider. Conclusions We discuss how reliable these fossils are as indicators of palaeo-parasitism, and in which aspects the behaviour of mantis lacewing larvae in general indeed represents parasitism. While the specimen appears to be about to board the spider, it may not necessarily represent a parasite in the strict sense. Evaluating the actual ecological role of a fossil heavily depends on comparison to modern forms, and not all modern-day larvae of Mantispidae are parasites. We therefore provide a closer look into the known feeding habits of modern mantis lacewing larvae. more...

Aphidlions are larvae of certain lacewings (Neuroptera), and more precisely larvae of the groups Chrysopidae, green lacewings, and Hemerobiidae, brown lacewings. The name ‘aphidlion’ originates from their ecological function as specialised predators of aphids. Accordingly, they also play an economic role as biological pest control. Aphidlions have, mostly, elongated spindle-shaped bodies, and similarly to most lacewing larvae they are equipped with a pair of venom-injecting stylets. Fossils interpreted as aphidlions are known to be preserved in amber from the Cretaceous (130 and 100 million years ago), the Eocene (about 35 million years ago) and the Miocene (about 15 million years ago) ages. In this study, new aphidlion-like larvae are reported from Cretaceous amber from Myanmar (about 100 million years old) and Eocene Baltic amber. The shapes of head and stylets were compared between the different time slices. With the newly described fossils and specimens from the literature, a total of 361 specimens could be included in the analysis: 70 specimens from the Cretaceous, 5 from the Eocene, 3 from the Miocene, 188 extant larvae of Chrysopidae, and 95 extant larvae of Hemerobiidae. The results indicate that the diversity of head shapes remains largely unchanged over time, yet there is a certain increase in the diversity of head shapes in the larvae of Hemerobiidae. In certain other groups of Neuroptera, a distinct decrease in the diversity of head shapes in larval stages was observed. more...

Dictyoptera, which comprises cockroaches, termites and mantids, is a quite successful group of insects in evolutionary terms with a long fossil record—roachoid insects were already abundant 315 million years ago in the Carboniferous forests. One of the most remarkable autapomorphies of extant dictyopterans, and possibly a major factor for their persisting success, is the ability to produce oothecae. Despite the robustness of this sort of egg package, fossils of oothecae are very rare, the oldest direct evidences being from the Cretaceous Crato Formation in Brazil (115 mya). The ability to produce oothecae is presumably linked to a specific ovipositor morphology, including a significant length reduction. Hence, ovipositor morphology can indirectly inform about the reproductive strategy of a species. Herein we describe the ovipositor morphology of various fossil forms of dictyopteran insects. Early fossil roachoids, in contrast to the modern forms, possessed a very long and prominent ovipositor, reminiscent of the ovipositor in orthopterans (Ensifera), indicating that these forms laid individual, rather small eggs into a substrate. We present examples from different fossil deposits, which show the entire range of ovipositor morphologies, from very long forms over forms with ovipositors partly reduced in length to modern-appearing morphologies. Most remarkably, different shapes of ovipositors seem to be present in roachoids in the fauna of the 115 million years old Crato Formation—species with long prominent ovipositors co-existed with species with a reduced short and broad ovipositor. Additionally, females that carry oothecae attached to their abdomen indicate a third type of ovipositor: a further reduced ovipositor as seen in modern forms, which already allowed the internal production of oothecae. more...

Abstract We present a documentation of the morphological details of two larval stages of mantis shrimps. Documentation was done using the autofluorescence capacities of the cuticle. This is the first time that morphological details of late mantis shrimp larvae are documented in great detail via photography, including all parts of the body up to the proximal elements of the appendages, and not presented as line drawings; it is the second time for mantis shrimp larvae in general. The description is presented as a standardized descriptive matrix. Documentation and description style are adjusted to facilitate comparison with fossil representatives of mantis shrimps, but also their extant counterparts, as well as specimens in the wider framework of Malacostraca and Eucrustacea. Through an exemplary comparison with fossil mantis shrimps, we provide indications about the early evolutionary history of the group. Through an out-group comparison, we identify several possible evolutionary changes of developmental timing, i.e., heterochrony, which could explain some morphological specialisations of mantis shrimps. more...

Larvae, and especially fossil larvae, are challenging to deal with from a purely taxonomic view. Often one cannot determine which species the larvae belong to. Yet, larvae can still contribute to various scientific questions. Especially morphological traits of a fossil larva can be highly informative for reconstructing character evolution. Also the occurrence of specific larval types and larval characters in time and the disappearance of such forms can well be reconstructed also without being able to narrow down the phylogenetic relationship of a larva very far. Here, we report two new beetle larvae preserved in Baltic amber which are identified as representatives of Scraptiidae, based on an enlarged terminal end (‘9th abdomen segment’); this is only the third record of such larvae. In comparison to modern forms, the terminal ends of the two new fossil larvae is even larger in relation to the remaining body than in any known larva. Unfortunately, our knowledge of such larvae in the modern fauna is very limited. Still, one of the two already known fossil larvae of Scraptiidae also has a very long terminal end, but not as long as those of the two new fossils. These three fossil larvae therefore seem to possess a specific morphology not known from the modern fauna. This might either mean that they (1) represent a now extinct larval morphology, a phenomenon well known in other euarthropodan lineages, or that (2) these forms represent a part of the larval phase not known from modern day species as they have not been described yet; such cases occur in closely related lineages. In any case, the fossils expand the known diversity of larval morphologies. more...

Neuroptera, the group of lacewings, comprises only about 6000 species in the modern fauna, but is generally assumed to have been more diverse and important in the past. A major factor of the modern-day ecological diversity of the group, and supposedly in the past as well, is represented by the highly specialised larval forms of lacewings. Quantitative analyses of the morphology of larvae revealed a loss of morphological diversity in several lineages. Here we explored the diversity of the larvae of mantis lacewings (Mantispidae), lance lacewings (Osmylidae), beaded lacewings (Berothidae and Rhachiberothidae, the latter potentially an ingroup of Berothidae), and pleasing lacewings (Dilaridae), as well as fossil larvae, preserved in amber, resembling these. We used shape analysis of the head capsule and stylets (pair of conjoined jaws) as a basis due to the high availability of this body region in extant and fossil specimens and the ecological importance of this region. The analysis revealed a rather constant morphological diversity in Berothidae. Mantispidae appears to have lost certain forms of larvae, but has seen a drastic increase of larval diversity after the Cretaceous; this is in contrast to a significant decrease in diversity in adult forms. more...

Abstract Mantis shrimps are prominent predatory crustaceans. Their larvae, although morphologically very differently-appearing from their adult counterparts, are already predators; yet, unlike the adults they are not benthic. Instead they are part of the plankton preying on other planktic organisms. Similar to some types of lobsters and crab-like crustaceans the planktic larvae of mantis shrimps can grow quite large, reaching into the centimeter range. Nonetheless, our knowledge on mantis shrimp larvae is still rather limited. Recently new types of giant mantis shrimp larvae with “extreme morphologies” have been reported. Here we describe another type that qualifies to be called “extreme”. Comparative measurements of certain morphological structures on selected known larvae support the exceptionality of the new specimen. It differs in several aspects from the original four types of extreme mantis shrimp larvae described by C. Haug et al. (2016). With this fifth type we expand the known morphological diversity of mantis shrimp larvae and also contribute to our still very incomplete, although growing, knowledge of this life phase. more...

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 Santanmantis axelrodi 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 S. axelrodi 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. more...

The early fossil record of insects is scarce, with only few finds in the Devonian. All these finds appear problematic and controversial, partly due to incomplete preservation and challenging interpretation of many structures. We provide details of one of these important forms, Rhyniognatha hirsti from the famous Rhynie Chert Lagerstätte with up-to-date 3D imaging techniques. The fossil has been interpreted as the remains of one of the earliest flying insects. The specimen mainly preserves the remains of the head. The structures of the mandibles have been used as a main argument for an interpretation as an insect, but these are in fact less easy to interpret. New observed structures include the remains of a head capsule and an additional pair of mouth parts. Structures formerly suggested to represent remains of the head capsule or apodemes are more likely to be representing glands of ectodermal origin. The newly observed structures do not support an interpretation as an insect. Instead they make the interpretation as a myriapod more likely, possibly as a centipede. Centipede remains from the Rhynie Chert are known from scutigeromorphs. We therefore point out that R. hirsti could be interpreted as an early centipede. more...

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 Nephrops norvegicus, 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. more...

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. more...