South African Lagerstätte reveals middle Permian Gondwanan lakeshore ecosystem in exquisite detail.

Continental ecosystems of the middle Permian Period (273-259 million years ago) are poorly understood. In South Africa, the vertebrate fossil record is well documented for this time interval, but the plants and insects are virtually unknown, and are rare globally. This scarcity of data has hampered studies of the evolution and diversification of life, and has precluded detailed reconstructions and analyses of ecosystems of this critical period in Earth's history. Here we introduce a new locality in the southern Karoo Basin that is producing exceptionally well-preserved and abundant fossils of novel freshwater and terrestrial insects, arachnids, and plants. Within a robust regional geochronological, geological and biostratigraphic context, this Konservat- and Konzentrat-Lagerstätte offers a unique opportunity for the study and reconstruction of a southern Gondwanan deltaic ecosystem that thrived 266-268 million years ago, and will serve as a high-resolution ecological baseline towards a better understanding of Permian extinction events.

A twig-like insect stuck in the Permian mud indicates early origin of an ecological strategy in Hexapoda evolution.

Full body impressions and resting traces of Hexapoda can be of extreme importance because they bring crucial information on behavior and locomotion of the trace makers, and help to better define trophic relationships with other organisms (predators or preys). However, these ichnofossils are much rarer than trackways, especially for winged insects. Here we describe a new full-body impression of a winged insect from the Middle Permian of Gonfaron (Var, France) whose preservation is exceptional. The elongate body with short prothorax and legs and long wings overlapping the body might suggests a plant mimicry as for some extant stick insects. These innovations are probably in relation with an increasing predation pressure by terrestrial vertebrates, whose trackways are abundant in the same layers. This discovery would possibly support the recent age estimates for the appearance of phasmatodean-like stick insects, nearly 30 million years older than the previous putative records. The new exquisite specimen is fossilized on a slab with weak ripple-marks, suggesting the action of microbial mats favoring the preservation of its delicate structures. Further prospections in sites with this type of preservation could enrich our understanding of early evolutionary history of insects.

New fossil discoveries illustrate the diversity of past terrestrial ecosystems in New Caledonia.

New Caledonia was, until recently, considered an old continental island harbouring a rich biota with outstanding Gondwanan relicts. However, deep marine sedimentation and tectonic evidence suggest complete submergence of the island during the latest Cretaceous to the Paleocene. Molecular phylogenies provide evidence for some deeply-diverging clades that may predate the Eocene and abundant post-Oligocene colonisation events. Extinction and colonization biases, as well as survival of some groups in refuges on neighbouring paleo-islands, may have obscured biogeographic trends over long time scales. Fossil data are therefore crucial for understanding the history of the New Caledonian biota, but occurrences are sparse and have received only limited attention. Here we describe five exceptional fossil assemblages that provide important new insights into New Caledonia's terrestrial paleobiota from three key time intervals: prior to the submersion of the island, following re-emergence, and prior to Pleistocene climatic shifts. These will be of major importance for elucidating changes in New Caledonia's floristic composition over time.

Sound vs. light: wing-based communication in Carboniferous insects.

Acoustic communication is well-known in insects since the Mesozoic, but earlier evidence of this behavior is rare. Titanoptera, an 'orthopteroid' Permian-Triassic order, is one of the few candidates for Paleozoic intersex calling interactions: some specimens had highly specialized broadened zones on the forewings, which are currently considered-despite inconclusive evidence-as 'resonators' of a stridulatory apparatus. Here we argue that the stridulatory apparatus hypothesis is unlikely because the Titanoptera lack a stridulatory file on their bodies, legs or wings. Instead, comparing these broadened zones with similar structures in extant locusts, flies, and fossil damselflies, we find evidence that the Titanoptera used their wings to produce flashes of light and/or crepitated sounds. Moreover, we describe the first Carboniferous (~310 Mya) Titanoptera, which exhibits such specialized zones, thus corresponding to the oldest record of wing communication in insects. Whether these communication systems were used to attract sexual partners and/or escape predators remain to be demonstrated.

Carbon speciation in organic fossils using 2D to 3D x-ray Raman multispectral imaging.

The in situ two-dimensional (2D) and 3D imaging of the chemical speciation of organic fossils is an unsolved problem in paleontology and cultural heritage. Here, we use x-ray Raman scattering (XRS)-based imaging at the carbon K-edge to form 2D and 3D images of the carbon chemistry in two exceptionally preserved specimens, a fossil plant dating back from the Carboniferous and an ancient insect entrapped in 53-million-year-old amber. The 2D XRS imaging of the plant fossil reveals a homogeneous chemical composition with micrometric "pockets" of preservation, likely inherited from its geological history. The 3D XRS imaging of the insect cuticle displays an exceptionally well preserved remaining chemical signature typical of polysaccharides such as chitin around a largely hollowed-out inclusion. Our results open up new perspectives for in situ chemical speciation imaging of fossilized organic materials, with the potential to enhance our understanding of organic specimens and their paleobiology.

Palaeozoic giant dragonflies were hawker predators.

The largest insects to have ever lived were the giant meganeurids of the Late Palaeozoic, ancient stem relatives of our modern dragonflies. With wingspans up to 71 cm, these iconic insects have been the subject of varied documentaries on Palaeozoic life, depicting them as patrolling for prey through coal swamp forests amid giant lycopsids, and cordaites. Such reconstructions are speculative as few definitive details of giant dragonfly biology are known. Most specimens of giant dragonflies are known from wings or isolated elements, but Meganeurites gracilipes preserves critical body structures, most notably those of the head. Here we show that it is unlikely it thrived in densely forested environments where its elongate wings would have become easily damaged. Instead, the species lived in more open habitats and possessed greatly enlarged compound eyes. These were dorsally hypertrophied, a specialization for long-distance vision above the animal in flight, a trait convergent with modern hawker dragonflies. Sturdy mandibles with acute teeth, strong spines on tibiae and tarsi, and a pronounced thoracic skewness are identical to those specializations used by dragonflies in capturing prey while in flight. The Palaeozoic Odonatoptera thus exhibited considerable morphological specializations associated with behaviours attributable to 'hawkers' or 'perchers' among extant Odonata.

Redefining the extinct orders Miomoptera and Hypoperlida as stem acercarian insects.

BACKGROUND: The systematic positions of the extinct insect orders Hypoperlida, Miomoptera and Permopsocida were enigmatic and unstable for nearly a century. The recent studies based on new material, especially from the Cenomanian Burmese amber, shed light on evolutionary history of Acercaria resolving Permopsocida as the stem group of Condylognatha. However, the knowledge of the remaining two orders differs significantly. RESULTS: In this study, we describe new specimens and evaluate morphology of various structures with emphasis on the mouthparts and wing venation. Our results are primary based on revisions of the type specimens with a proper delimitation of taxa Hypoperlida and Miomoptera followed by their significance for the evolutionary history of Acercaria. Three new genera as Belmomantis gen. nov., Elmomantis gen. nov., and Mazonopsocus gen. nov. are designated as members of Palaeomanteidae. The Pennsylvanian Mazonopsocus provides a minimum age for calibration, in accordance to the presence of crown acercarians during the late Carboniferous. CONCLUSIONS: This contribution demonstrates that Hypoperlida and Miomoptera are stem groups of Acercaria. The putative clade (Hypoperlida + Miomoptera) is appearing as potential sister group of (Psocodea + (Permopsocida + (Thripida + Hemiptera))).

3-D imaging reveals four extraordinary cases of convergent evolution of acoustic communication in crickets and allies (Insecta).

When the same complex trait is exhibited by closely related species, a single evolutionary origin is frequently invoked. The complex stridulatory apparatus present in the forewings of extant crickets, mole crickets, katydids, and prophalangopsids, is currently interpreted as sharing a single common origin due to their similarity and unique function. An alternative hypothesis of convergent evolution in these ensiferan groups has challenged this common view, but remained controversial because of competing interpretations of wing venation. Here we propose another hypothesis for the widely and long debated homology of ensiferan stridulatory apparatus, performing the first 3D reconstruction of hidden structures at the wing bases. This approach allowed defining the homology of each vein from its very origin rather than after its more distal characteristics, which may be subjected to environmental pressure of selection. The stridulatory apparatus involves different veins in these four singing clades. In light of the most recent phylogenetic evidence, this apparatus developed four times in Ensifera, illustrating extraordinary convergent evolutions between closely related clades, by far exceeding the number of evolutionary steps ever proposed for calling ability in this group.

The "wild shot": photography for more biology in natural history collections, not for replacing vouchers.

Recently a correspondence in Zootaxa (Ceríaco et al., 2016) with more than 450 signatories including taxonomists, curators and other taxonomy users from all continents has received wide attention and has stimulated extensive discussion (a true buzz) around the possible interpretations of the Code (ICZN) about photography in taxonomy (Researchgate website link). This short note was necessary to recall the necessity of preserved specimens as vouchers for taxonomy, in response to photography-based taxonomy (PBT) as defended by Pape et al. (2016), and in a broad sense, for all the life sciences. This had been widely discussed and argued by Dubois & Nemésio (2007) who concluded on the importance of vouchers in taxonomy. But if the subject of these papers and discussions are about photography as the only way to document a new species, none of them discussed really what photography could represent in enhancing knowledge in natural sciences based on collections of specimens including type series and in association with other media (video and sound).

Insect mimicry of plants dates back to the Permian.

In response to predation pressure, some insects have developed spectacular plant mimicry strategies (homomorphy), involving important changes in their morphology. The fossil record of plant mimicry provides clues to the importance of predation pressure in the deep past. Surprisingly, to date, the oldest confirmed records of insect leaf mimicry are Mesozoic. Here we document a crucial step in the story of adaptive responses to predation by describing a leaf-mimicking katydid from the Middle Permian. Our morphometric analysis demonstrates that leaf-mimicking wings of katydids can be morphologically characterized in a non-arbitrary manner and shows that the new genus and species Permotettigonia gallica developed a mimicking pattern of forewings very similar to those of the modern leaf-like katydids. Our finding suggests that predation pressure was already high enough during the Permian to favour investment in leaf mimicry.

A new permopsocidan genus and species from the Late Permian of Australia (Insecta: Acercaria: Psocidiidae).

The psocidiid Hypopsylla belmontensis gen. et sp. nov., new psocidiid genus and species, is described and figured from the Late Permian of New South Wales in Australia. This discovery extends the knowledge on the diversity of the small order Permopsocida whose members passed the Permian-Triassic boundary and became extinct in Cretaceous.

The oldest accurate record of Scenopinidae in the Lowermost Eocene amber of France (Diptera: Brachycera).

Eocenotrichia magnifica gen. et sp. nov. (Diptera: Scenopinidae: Metatrichini) is described and illustrated from the Lowermost Eocene amber of Oise (France) and represents the oldest definitive window fly fossil. The present discovery in the Earliest Eocene supports the Late Cretaceous-Paleocene age currently proposed for the emergence of Metatrichini.

A new genus and species of micro bee fly from the Earliest Eocene French amber (Diptera: Mythicomyiidae: Psiloderoidinae).

Mythicomyiidae, or micro bee flies, are tiny flies (0.5-5.0 mm) that are found throughout most parts of the world except the highest altitudes and latitudes (Greathead & Evenhuis 2001). Including all extinct and extant taxa, the Mythicomyiidae currently comprise more than 380 valid taxonomic species distributed among 30 genera. The subfamily Psiloderoidinae is especially well represented among the fossil Mythicomyiidae by seven Cretaceous or Cenozoic genera. We here describe a new genus and a new species of this subfamily based on fossils from the Earliest Eocene of Oise (France). A Psiloderoidinae, Proplatypygus matilei Nel & DePloëg, 2004, is already described in this amber. Another mythicomyiid, Eurodoliopteryx inexpectatus Nel, 2006, is the most frequent bombylioid in this amber (Nel & DePloëg, 2004; Nel, 2006).

New fossil insect order Permopsocida elucidates major radiation and evolution of suction feeding in hemimetabolous insects (Hexapoda: Acercaria).

With nearly 100,000 species, the Acercaria (lice, plant lices, thrips, bugs) including number of economically important species is one of the most successful insect lineages. However, its phylogeny and evolution of mouthparts among other issues remain debatable. Here new methods of preparation permitted the comprehensive anatomical description of insect inclusions from mid-Cretaceous Burmese amber in astonishing detail. These "missing links" fossils, attributed to a new order Permopsocida, provide crucial evidence for reconstructing the phylogenetic relationships in the Acercaria, supporting its monophyly, and questioning the position of Psocodea as sister group of holometabolans in the most recent phylogenomic study. Permopsocida resolves as sister group of Thripida + Hemiptera and represents an evolutionary link documenting the transition from chewing to piercing mouthparts in relation to suction feeding. Identification of gut contents as angiosperm pollen documents an ecological role of Permopsocida as early pollen feeders with relatively unspecialized mouthparts. This group existed for 185 million years, but has never been diverse and was superseded by new pollenivorous pollinators during the Cretaceous co-evolution of insects and flowers. The key innovation of suction feeding with piercing mouthparts is identified as main event that triggered the huge post-Carboniferous radiation of hemipterans, and facilitated the spreading of pathogenic vectors.

New Eocene damselflies and first Cenozoic damsel-dragonfly of the isophlebiopteran lineage (Insecta: Odonata).

The study of a new specimen of Petrolestes hendersoni from the Eocene Green Formation allows a more precise description of the enigmatic damselfly and the diagnosis of the Petrolestini. Petrolestes messelensis sp. nov. is described from the Eocene Messel Formation in Germany, extending the distribution of the Petrolestini to the European Eocene. The new damsel-dragonfly family Pseudostenolestidae is described for the new genus and species Pseudostenolestes bechlyi, from the Eocene Messel Formation. It is the first Cenozoic representative of the Mesozoic clade Isophlebioptera.

A new libelluloid family from the Eocene Green River Formation (Colorado, USA) (Odonata, Anisoptera).

The new family Urolibellulidae is proposed for the new genus and species Urolibellula eocenica, based on a fossil dragonfly from the Eocene Green River Formation (USA). This new taxon is considered as the sister group of the extant Libellulidae. As the oldest libellulid dragonfly is dated from the Turonian, the Urolibellulidae should also be at least Late Cretaceous.

The first African Anthracoptilidae (Insecta: Paoliida) near the Permian--Triassic boundary in Kenya.

The first findings of Antracoptilidae (Paoliida) from the Late Permian/ Early Triassic of Africa, i.e. the new genus Afrocladus comprising A. pumilio sp. nov. and A. kenyaensis sp. nov., are described from the Mombasa Basin of Maji ya Chumvi Formation in Kenya (Duruma sandstones). Both diagnoses are based on wing venation pattern. Their occurrences close to the P-T boundary possibly indicate the last appearence date of the group being known since early Pennsylvanian. Moreover, their significantly smaller wing sizes compared to known taxa suggest adaptations to different habitats and environmental conditions or different life strategy.

Exceptionally preserved insect fossils in the Late Jurassic lagoon of Orbagnoux (Rhône Valley, France).

The Late Kimmeridgian marine limestones of the area around Orbagnoux (Rhône, France) are well known for their fish fauna and terrestrial flora. Here we record the first insects and their activities (mines on leaves and trails in sediments) from these layers, including the oldest record of the gerromorphan bugs, as a new genus and species Gallomesovelia grioti, attributed to the most basal family Mesoveliidae and subfamily Madeoveliinae. These new fossils suggest the presence of a complex terrestrial palaeoecosystem on emerged lands near the lagoon where the limestones were deposited. The exquisite state of preservation of these fossils also suggests that these outcrops can potentially become an important Konservat-Lagerstätte for the Late Jurassic of Western Europe.

The first ant-termite syninclusion in amber with CT-scan analysis of taphonomy.

We describe here a co-occurrence (i.e. a syninclusion) of ants and termites in a piece of Mexican amber (Totolapa deposit, Chiapas), whose importance is two-fold. First, this finding suggests at least a middle Miocene antiquity for the modern, though poorly documented, relationship between Azteca ants and Nasutitermes termites. Second, the presence of a Neivamyrmex army ant documents an in situ raiding behaviour of the same age and within the same community, confirmed by the fact that the army ant is holding one of the termite worker between its mandibles and by the presence of a termite with bitten abdomen. In addition, we present how CT-scan imaging can be an efficient tool to describe the topology of resin flows within amber pieces, and to point out the different states of preservation of the embedded insects. This can help achieving a better understanding of taphonomical processes, and tests ethological and ecological hypotheses in such complex syninclusions.

The earliest known holometabolous insects.

The Eumetabola (Endopterygota (also known as Holometabola) plus Paraneoptera) have the highest number of species of any clade, and greatly contribute to animal species biodiversity. The palaeoecological circumstances that favoured their emergence and success remain an intriguing question. Recent molecular phylogenetic analyses have suggested a wide range of dates for the initial appearance of the Holometabola, from the Middle Devonian epoch (391 million years (Myr) ago) to the Late Pennsylvanian epoch (311 Myr ago), and Hemiptera (310 Myr ago). Palaeoenvironments greatly changed over these periods, with global cooling and increasing complexity of green forests. The Pennsylvanian-period crown-eumetabolan fossil record remains notably incomplete, particularly as several fossils have been erroneously considered to be stem Holometabola (Supplementary Information); the earliest definitive beetles are from the start of the Permian period. The emergence of the hymenopterids, sister group to other Holometabola, is dated between 350 and 309 Myr ago, incongruent with their current earliest record (Middle Triassic epoch). Here we describe five fossils--a Gzhelian-age stem coleopterid, a holometabolous larva of uncertain ordinal affinity, a stem hymenopterid, and early Hemiptera and Psocodea, all from the Moscovian age--and reveal a notable penecontemporaneous breadth of early eumetabolan insects. These discoveries are more congruent with current hypotheses of clade divergence. Eumetabola experienced episodes of diversification during the Bashkirian-Moscovian and the Kasimovian-Gzhelian ages. This cladogenetic activity is perhaps related to notable episodes of drying resulting from glaciations, leading to the eventual demise in Euramerica of coal-swamp ecosystems, evidenced by floral turnover during this interval. These ancient species were of very small size, living in the shadow of Palaeozoic-era 'giant' insects. Although these discoveries reveal unexpected Pennsylvanian eumetabolan diversity, the lineage radiated more successfully only after the mass extinctions at the end of the Permian period, giving rise to the familiar crown groups of their respective clades.