Expanding the Mesozoic Record of Early Brachyceran Fly Larvae, including New Larval Forms with Chimera-Type Morphologies.

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.

Morphological diversity in true and false crabs reveals the plesiomorphy of the megalopa phase.

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.

Fossils in Myanmar amber demonstrate the diversity of anti-predator strategies of Cretaceous holometabolan insect larvae.

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.

The Morphological Diversity of Dragon Lacewing Larvae (Nevrorthidae, Neuroptera) Changed More over Geological Time Scales Than Anticipated.

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.

Quantitative analysis of lacewing larvae over more than 100 million years reveals a complex pattern of loss of morphological diversity.

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.

An Expanded View on the Morphological Diversity of Long-Nosed Antlion Larvae Further Supports a Decline of Silky Lacewings in the Past 100 Million Years.

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.

A hatching aphidlion-like lacewing larva in 100 million years old Kachin amber.

We report a fossil aphidlion-like larva preserved with its egg case in 100 million year old Kachin amber, Myanmar. It appears to have been enclosed very shortly after hatching, especially when comparing it with extant aphidlions during hatching. Although hatching aphidlion-like larvae are known from amber from other localities, this is the first case from Myanmar amber, despite the comparably high number of lacewing larvae known from the latter.

The Morphological Diversity of Antlion Larvae and Their Closest Relatives over 100 Million Years.

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.

100 Million-year-old straight-jawed lacewing larvae with enormously inflated trunks represent the oldest cases of extreme physogastry in insects.

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.

The Diversity of Aphidlion-like Larvae over the Last 130 Million Years.

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.

The first fossil immature of Elmidae: an unusual riffle beetle larva preserved in Baltic amber.

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.

Changes in the Morphological Diversity of Larvae of Lance Lacewings, Mantis Lacewings and Their Closer Relatives over 100 Million Years.

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.

Fossil dragonfly-type larva with lateral abdominal protrusions and implications on the early evolution of Pterygota.

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.

New extreme morphologies as exemplified by 100 million-year-old lacewing larvae.

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.

Intraspecific variation in the Cambrian: new observations on the morphology of the Chengjiang euarthropod Sinoburius lunaris.

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.

A new fossil mantis shrimp and the convergent evolution of a lobster-like morphotype.

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.

Morphological changes during the post-embryonic ontogeny of mesothelan spiders and aspects of character evolution in early spiders.

Most morphological studies focus on adult specimens, or if developmental studies are pursued, especially in Euarthropoda, they focus on embryonic development. Araneae (spiders) is one of these groups, in consequence with understudied post-embryonic development. Here we present aspects of the post-embryonic stages of different species of Mesothelae, sister group to the remaining spiders (when fossil species are not taken into account). We used different imaging methods and measured different external morphological structures to detect possible ontogenetic changes. One structure exhibiting post-embryonic changes is the chelicera. Here the significant change occurs between the last immature stage and the adult, yet only in males. For the spinnerets, we could not detect ontogenetic changes, but instead a high variability in length and width, probably due to their lack of pivot joints between the elements. The strongest morphological change during ontogeny occurred on the sternum, which begins with a rather roundish shape in the first stage and changes to being fairly elongate in shape in the last immature stages and the adult. This specific sternum shape only occurs in adults of mesothelan spiders, while opisthothelan spiders have a broader sternum also in the adult. We discuss our results in an evolutionary context, also taking into account recent finds of fossil spiders.

An unusual 100-million-year old holometabolan larva with a piercing mouth cone.

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.

First African thylacocephalans from the Famennian of Morocco and their role in Late Devonian food webs.

Thylacocephalans are enigmatic arthropods with an erratic Palaeozoic and Mesozoic fossil record. In many of the few localities where they occur, they are quite abundant. This also holds true for the Famennian Thylacocephalan Layer in the Maider (eastern Anti-Atlas of Morocco), a small epicontinental basin hosting some strata with taphonomic properties of a conservation deposit yielding exceptionally preserved gnathostomes and non-vertebrates. In a thin argillaceous interval in the earliest middle Famennian, thylacocephalans occur in such great numbers that they became eponyms of this unit. Therein, we discovered a new taxon of thylacocephalans, Concavicaris submarinus sp. nov., which represent the oldest records of thylacocephalans from Africa. In the CT-imagery, the holotype of Concavicaris submarinus sp. nov. revealed anatomical details including its eyes, appendages and other soft parts. Sedimentary facies and faunal composition of the Thylacocephalan Layer suggest that these animals populated the water column above the low-oxygen sea floor. Thus, thylacocephalans likely represented an important component of the diet of chondrichthyans and placoderms, which are quite common as well. The abundance of thylacocephalans in other conservation deposits like the Cleveland Shale (USA) and the Gogo Formation (Australia) underline their pivotal role in Late Devonian pelagic food webs.

Untangling the Gordian knot-further resolving the super-species complex of 300-million-year-old xiphosurids by reconstructing their ontogeny.

The group Xiphosurida (horseshoe "crabs") is today only represented by four species. However, in the fossil record, several dozen species have been described, especially from the Carboniferous (about 300 million years ago). Several species have been interpreted as representatives of Euproops or Belinurus, but there is ongoing discussion which of these species are valid and how they can be differentiated. Recent studies suggested that differences in the timing of individual development could provide information for species distinction, exemplified by studies on Euproops danae (Mazon Creek, USA) and Euproops sp. ("Piesproops"; Piesberg, Germany). For this study, we reinvestigated all Carboniferous xiphosurids from the British Coal Measures stored in the collections of the Natural History Museum London. Size comparisons of the specimens revealed nine size groups; the smaller specimens were originally labelled as Belinurus, the larger ones as Euproops. The nine size groups exhibit five different morphotypes differing in structures surrounding the posterior shield (= thoracetron): spines of different lengths and, in larger specimens, a more or less developed flange. Two of these morphotypes show significantly longer spines than the remaining specimens and could be conspecific as E. anthrax. The remaining specimens are interpreted as growth series of another species, presumably of E. rotundatus. An ontogenetic flange formation is also known from E. danae and the "Piesproops", but the timing differs between all three species. In E. rotundatus, the flange develops rather late, but then comparably abruptly, which makes this development more metamorphic in relation to development in the other species.