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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
ABSTRACT
The nymphal stages of Palaeozoic insects differ significantly in morphology from those of their modern counterparts. Morphological details for some previously reported species have recently ...been called into question. Palaeozoic insect nymphs are important, however – their study could provide key insights into the evolution of wings, and complete metamorphosis. Here we review past work on these topics and juvenile insects in the fossil record, and then present both novel and previously described nymphs, documented using new imaging methods. Our results demonstrate that some Carboniferous nymphs – those of Palaeodictyopteroidea – possessed movable wing pads and appear to have been able to perform simple flapping flight. It remains unclear whether this feature is ancestral for Pterygota or an autapomorphy of Palaeodictyopteroidea. Further characters of nymphal development which were probably in the ground pattern of Pterygota can be reconstructed. Wing development was very gradual (archimetaboly). Wing pads did not protrude from the tergum postero‐laterally as in most modern nymphs, but laterally, and had well‐developed venation. The modern orientation of wing pads and the delay of wing development into later developmental stages (condensation) appears to have evolved several times independently within Pterygota: in Ephemeroptera, Odonatoptera, Eumetabola, and probably several times within Polyneoptera. Selective pressure appears to have favoured a more pronounced metamorphosis between the last nymphal and adult stage, ultimately reducing exploitation competition between the two. We caution, however, that the results presented herein remain preliminary, and the reconstructed evolutionary scenario contains gaps and uncertainties. Additional comparative data need to be collected. The present study is thus seen as a starting point for this enterprise.
Leanchoilia superlata is one of the best known arthropods from the middle Cambrian Burgess Shale of British Columbia. Here we re-describe the morphology of L. superlata and discuss its possible ...autecology. The re-description follows a standardized scheme, the descriptive matrix approach, designed to provide a template for descriptions of other megacheiran species.
Our findings differ in several respects from previous interpretations. Examples include a more slender body; a possible hypostome; a small specialised second appendage, bringing the number of pairs of head appendages to four; a further sub-division of the great appendage, making it more similar to that of other megacheirans; and a complex joint of the exopod reflecting the arthropod's swimming capabilities.
Different aspects of the morphology, for example, the morphology of the great appendage and the presence of a basipod with strong median armature on the biramous appendages indicate that L. superlata was an active and agile necto-benthic predator (not a scavenger or deposit feeder as previously interpreted).
Lobopodians, a nonmonophyletic assemblage of worm-shaped soft-bodied animals most closely related to arthropods, show two major morphotypes: long-legged and short-legged forms. The morphotype with ...stubby, conical legs has a long evolutionary history, from the early Cambrian 1 through the Carboniferous 2, 3, including the living onychophorans and tardigrades 4–6. Species with tubular lobopods exceeding the body diameter have been reported exclusively from the Cambrian 7–12; the three-dimensionally preserved Orstenotubulus evamuellerae from the uppermost middle Cambrian “Orsten” (Sweden) is the youngest long-legged lobopodian reported thus far 8. Here we describe a new long-legged lobopodian, Carbotubulus waloszeki gen. et sp. nov., from Mazon Creek, Illinois, USA (∼296 million years ago) 13. This first post-Cambrian long-legged lobopodian extends the range of this morphotype by about 200 million years. The three-dimensionally preserved specimen differs significantly from the associated short-legged form Ilyodes inopinata 2, of which we also present new head details. The discovery of a Carboniferous long-legged lobopodian provides a more striking example of the long-term survival of Cambrian morphotypes than, for example, the occurrence of a Burgess Shale-type biota in the Ordovician of Morocco 14 and dampens the effect of any major extinction of taxa at the end of the middle Cambrian 15, 16.
► The first post-Cambrian long-legged lobopodian was found in the late Carboniferous ► It is about 200 million years younger than other long-legged lobopodians ► This finding is an extreme example of a long-term survival of a Cambrian morphotype
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