Mesozoic larva in amber reveals the venom delivery system and the palaeobiology of an ancient lineage of venomous insects (Neuroptera).

The larvae of Neuroptera are predators that feed by injecting bioactive compounds into their prey and then suctioning the fluids through modified mouthparts. We explore the evolutionary history of this feeding structure through the examination of a new fossil larva preserved in Late Cretaceous Kachin amber, which we describe as new genus and species, Electroxipheus veneficus gen et sp. nov. X-ray phase-contrast microtomography enabled us to study the anatomy of the larva in 3D, including the structure of the mouthparts and that of the venom delivery system. The specimen exhibited a unique combination of morphological traits not found in any known fossil or extant lacewing, including an unusual structure of the antenna. Phylogenetic analyses, incorporating a selection of living and fossil larval Neuroptera and enforcing maximum parsimony and Bayesian inference, identified the larva as belonging to the stem group Mantispoidea. The larva shows that the anatomy of the feeding and venom-delivery apparatus has remained unchanged in Neuroptera from the Cretaceous to the present. The morphology of the specimen suggests that it was an active predator, in contrast with the scarcely mobile, specialized relatives, like mantispids and berothids.

Cretaceous amber inclusions illuminate the evolutionary origin of tardigrades.

Tardigrades are a diverse phylum of microscopic invertebrates widely known for their extreme survival capabilities. Molecular clocks suggest that tardigrades diverged from other panarthropods before the Cambrian, but their fossil record is extremely sparse. Only the fossil tardigrades Milnesium swolenskyi (Late Cretaceous) and Paradoryphoribius chronocaribbeus (Miocene) have resolved taxonomic positions, restricting the availability of calibration points for estimating for the origin of this phylum. Here, we revise two crown-group tardigrades from Canadian Cretaceous-aged amber using confocal fluorescence microscopy, revealing critical morphological characters that resolve their taxonomic positions. Formal morphological redescription of Beorn leggi reveals that it features Hypsibius-type claws. We also describe Aerobius dactylus gen. et sp. nov. based on its unique combination of claw characters. Phylogenetic analyses indicate that Beo. leggi and Aer. dactylus belong to the eutardigrade superfamily Hypsibioidea, adding a critical fossil calibration point to investigate tardigrade origins. Our molecular clock estimates suggest an early Paleozoic diversification of crown-group Tardigrada and highlight the importance of Beo. leggi as a calibration point that directly impacts estimates of shallow nodes. Our results suggest that independent terrestrialization of eutardigrades and heterotardigrades occurred around the end-Carboniferous and Lower Jurassic, respectively. These estimates also provide minimum ages for convergent acquisition of cryptobiosis.

First fossil species of family Hyidae (Arachnida: Pseudoscorpiones) confirms 99 million years of ecological stasis in a Gondwanan lineage.

Burmese amber preserves a diverse assemblage of Cretaceous arachnids, and among pseudoscorpions (Arachnida: Pseudoscorpiones), ten species in five families have already been named. Here, we describe a new fossil species from Burmese amber in the pseudoscorpion family Hyidae, providing detailed measurements, photographs and 3D-models from synchrotron scanning. Based on morphology, the new fossil, Hya fynni sp. nov. is placed in the genus Hya, and is nearly identical to extant species in the genus, except for the position of trichobothrium est on the pedipalpal chela, thereby indicating extreme morphological stasis in this invertebrate lineage over the last 99 million years. Hya fynni represents the first described fossil species in Hyidae, and the third described Burmese fossil in the superfamily Neobisioidea. It also joins the garypinid, Amblyolpium burmiticum, in representing the oldest fossil records for extant pseudoscorpion genera. Considering proposed divergence dates, the newly described fossil species bolsters a Gondwanan origin for Hyidae, and provides evidence for the "Late Jurassic Rifting" hypothesis for the Burma Terrane, in which this landmass rifted from Gondwana in the Late Jurassic and collided with Eurasia by the Cretaceous/Eocene. Like Hya species today, H. fynni likely inhabited humicolous microhabitats in tropical forests on the Burma Terrane, supporting ecological niche stasis for this family since the Mesozoic.

Cretaceous crown male ant reveals the rise of modern lineages.

Most described Mesozoic ants belong to stem groups that existed only during the Cretaceous period. Previously, the earliest known crown ants were dated to the Turonian (Late Cretaceous, ca. 94-90 million years ago (Ma)) deposits found in the USA, Kazakhstan, and Botswana. However, the recent discovery of an alate male ant in Kachin amber from the earliest Cenomanian (ca. 99 Ma), representing a new genus and species, Antiquiformica alata, revises the narrative on ant diversification. Antiquiformica can be distinctly differentiated from all known male stem ants by its geniculate antennae with elongated scape, extending far beyond the occipital margin of the head and half the length of the funiculus, as well as its partly reduced forewing venation. Furthermore, the combination of a one-segmented waist with a well-developed node, elongated scape extending beyond the occipital margin, and reduced forewing venation, particularly the completely reduced m-cu and rs-m crossveins and absence of rm and mcu closed cells, firmly places the fossil within the extant subfamily Formicinae. Fourier transform infrared spectroscopy (FTIR) confirmed that the amber containing Antiquiformica alata originated from the Kachin mines in Myanmar. This discovery significantly revises our understanding of the early evolution of Formicinae. The presence of Antiquiformica in Cenomanian amber indicates that the subfamily Formicinae emerged at least by the start of the Late Cretaceous, with crown ants likely originating earlier during the earliest Cretaceous or possibly the Late Jurassic, although paleontological evidence is lacking to support the latter hypothesis.

Three taphonomic stories of three new fossil species of Darwin wasps (Hymenoptera, Ichneumonidae).

Amber captures a snapshot of life and death from millions of years in the past. Here, the fate of three fossil Darwin wasps in Baltic amber is virtually dissected with the help of micro-CT scanning, to better understand the taphonomic processes that affected their preservation. The states of the fossils range from nearly perfect preservation, including remains of internal organs, to empty casts that were strongly affected by decomposition. We describe the three specimens as new taxa, Osparvis aurorae gen. et sp. nov., Grana harveydenti gen. et sp. nov. and Xorides? romeo sp. nov. Based on the taphonomic and morphological interpretations, we conclude that two specimens were trapped alive, and the third ended up in resin post-mortem. The morphology and classification of the specimens provide clues regarding their ecology, and we discuss their likely hosts and parasitation modes. Taken together, our three wasp fossils showcase how an integrative analysis of amber taphonomy, taxonomic association and morphology can shed light onto past biodiversity and offer valuable insights for interpreting their evolutionary history.

Compound osteoderms preserved in amber reveal the oldest known skink.

Scincidae is one of the most species-rich and cosmopolitan clades of squamate reptiles. Abundant disarticulated fossil material has also been attributed to this group, however, no complete pre-Cenozoic crown-scincid specimens have been found. A specimen in Burmite (99 MYA) is the first fossil that can be unambiguously referred to this clade. Our analyses place it as nested within extant skinks, supported by the presence of compound osteoderms formed by articulated small ostedermites. The specimen has a combination of dorsal and ventral compound osteoderms and overlapping cycloid scales that is limited to skinks. We propose that this type of osteoderm evolved as a response to an increased overlap of scales, and to reduced stiffness of the dermal armour. Compound osteoderms could be a key innovation that facilitated diversification in this megadiverse family.

Chemical and palaeoentomological evidence of a relationship between early Eocene Belgian and Oise (France) ambers.

Of the early Eocene amber deposits known across the world, Belgian amber has been mostly absent from the relevant literature. We reinvestigated amber held in the palaeobotanical collection of the Royal Belgian Institute of Natural Sciences, Brussels, which derived from three localities in Belgium that originated from two geographical areas (Leval-Trahegnies and Orp-le-Grand). Using Fourier transform infrared (FTIR) spectroscopy we show the close chemical relationship of Belgian amber to the early Eocene Oise amber from the Paris Basin, and highlight the potential effect of weathering on the amber chemistry. The amber derives from a very similar botanical source as the Oise amber (Combretaceae or Leguminosae-Caesalpinioideae), but from different coeval basins. The two Leval-Trahegnies localities provided amber that exhibit different stages of weathering (heavily fissured and crazed, darkened) and lacking any inclusions. The Orp-le-Grand locality provided the least weathered amber, with one amber piece containing two inclusions: a mite and a new genus and species of hemipteran (Cativolcus uebruum gen. et sp. nov.), and a second one that preserved the impression of insect wings pressed into the surface.

Eocene amber provides the first fossil record and bridges distributional gap in the rare genus Robsonomyia (Diptera: Keroplatidae).

Until now, the genus Robsonomyia was represented by two extant species: R. reducta Matile & Vockeroth, 1980 from North America and R. sciaraeformis (Okada, 1939) from Asia. This paper presents the first fossil members of the genus Robsonomyia, which is also the first record from Europe. Two new fossil species from Baltic amber are described: R. baltica Pelczynska, Krzeminski & Blagoderov, sp. nov. and R. henningseni Pelczynska, Krzeminski & Blagoderov, sp. nov.. The presence of fossil Robsonomyia spp. on the European continent suggests Holarctic distribution of the genus in the past. We also discuss possible pathways of its intercontinental dispersion.

Ant evolution: Amber revelations of extinction, survival and recovery.

Ant fossils from the Cretaceous are rare but critical for understanding the early evolution of this incredibly successful group of animals. New amber fossils fill important gaps, revealing patterns of death, survival, and radiation around the end Cretaceous extinction.

Early evolution of Anamorphidae (Coleoptera: Coccinelloidea): the oldest known anamorphid beetles from Upper Cretaceous amber of northern Myanmar and the first report of potential glandular pores in the family.

In order to place newly discovered fossil taxa (Palaeosymbius gen. nov. with P. groehni and P. mesozoicus spp. nov.) from the mid-Cretaceous amber from northern Myanmar, we investigated the relations of extant and extinct lineages of the coccinellid group of Coccinelloidea with emphasis on the family Anamorphidae. We assembled a taxonomic sampling of 34 taxa, including 15 genera and 19 species of Anamorphidae, the most comprehensive sampling of Anamorphidae at the generic level in a phylogenetic analysis. A morphological dataset of 47 characters was built as well as a molecular alignment of 7140 bp including fragments of eight genes (12S, 16S, 18S, 28S, COI, COII, H3 and CAD). Five anamorphid and one endomychid species were sequenced for the first time and added to the dataset. We performed parsimony-based analysis of the morphological dataset and Bayesian inference analysis of the combined matrix (morphological plus molecular data). Our results confirm that Palaeosymbius belongs to Anamorphidae and represents the oldest known member of this family so far. Among Anamorphidae, Symbiotes (with extant and known Eocene species) was recovered as the most probable closest relative of Palaeosymbius. Our morphological studies additionally revealed the presence of probable glandular openings in the anterolateral corners of the pronotal margins in Asymbius sp. and Anamorphus sp., representing the first report of secretory openings in the family Anamorphidae. Similar openings are found in other cucujiform beetles such as Cryptophagidae and Boganiidae with possible defensive purposes.

Group mating in Cretaceous water striders.

Fossilized mating insects are irreplaceable material for comprehending the evolution of the mating behaviours and life-history traits in the deep-time record of insects as well as the potential sexual conflict. However, cases of mating pairs are particularly rare in fossil insects, especially aquatic or semi-aquatic species. Here, we report the first fossil record of a group of water striders in copulation (including three pairs and a single adult male) based on fossils from the mid-Cretaceous of northern Myanmar. The new taxon, Burmogerris gen. nov., likely represents one of the oldest cases of insects related to the marine environment, such as billabongs formed by the tides. It exhibits conspicuous dimorphism associated with sexual conflict: the male is equipped with a specialized protibial comb as a grasping apparatus, likely representing an adaptation to overcome female resistance during struggles. The paired Burmogerris show smaller males riding on the backs of the females, seemingly recording a scene of copulatory struggles between the sexes. Our discovery reveals a mating system dominated by males and sheds light on the potential sexual conflicts of Burmogerris in the Cretaceous. It indicates the mating behaviour remained stable over long-term geological time in these water-walking insects.

Prolonged faunal turnover in earliest ants revealed by North American Cretaceous amber.

All ∼14,000 extant ant species descended from the same common ancestor, which lived ∼140-120 million years ago (Ma).1,2 While modern ants began to diversify in the Cretaceous, recent fossil evidence has demonstrated that older lineages concomitantly occupied the same ancient ecosystems.3 These early-diverging ant lineages, or stem ants, left no modern descendants; however, they dominated the fossil record throughout the Cretaceous until their ultimate extinction sometime around the K-Pg boundary. Even as stem ant lineages appear to be diverse and abundant throughout the Cretaceous, the extent of their longevity in the fossil record and circumstances contributing to their extinction remain unknown.3 Here we report the youngest stem ants, preserved in ∼77 Ma Cretaceous amber from North Carolina, which illustrate unexpected morphological stability and lineage persistence in this enigmatic group, rivaling the longevity of contemporary ants. Through phylogenetic reconstruction and morphometric analyses, we find evidence that total taxic turnover in ants was not accompanied by a fundamental morphological shift, in contrast to other analogous stem extinctions such as theropod dinosaurs. While stem taxa showed broad morphological variation, high-density ant morphospace remained relatively constant through the last 100 million years, detailing a parallel, but temporally staggered, evolutionary history of modern and stem ants.

New Canadian amber deposit fills gap in fossil record near end-Cretaceous mass extinction.

Amber preserves an exceptional record of tiny, soft-bodied organisms and chemical environmental signatures, elucidating the evolution of arthropod lineages and the diversity, ecology, and biogeochemistry of ancient ecosystems. However, globally, fossiliferous amber deposits are rare in the latest Cretaceous and surrounding the Cretaceous-Paleogene (K-Pg) mass extinction.1,2,3,4,5 This faunal gap limits our understanding of arthropod diversity and survival across the extinction boundary.2,6 Contrasting hypotheses propose that arthropods were either relatively unaffected by the K-Pg extinction or experienced a steady decline in diversity before the extinction event followed by rapid diversification in the Cenozoic.2,6 These hypotheses are primarily based on arthropod feeding traces on fossil leaves and time-calibrated molecular phylogenies, not direct observation of the fossil record.2,7 Here, we report a diverse amber assemblage from the Late Cretaceous (67.04 ± 0.16 Ma) of the Big Muddy Badlands, Canada. The new deposit fills a critical 16-million-year gap in the arthropod fossil record spanning the K-Pg mass extinction. Seven arthropod orders and at least 11 insect families have been recovered, making the Big Muddy amber deposit the most diverse arthropod assemblage near the K-Pg extinction. Amber chemistry and stable isotopes suggest the amber was produced by coniferous (Cupressaceae) trees in a subtropical swamp near remnants of the Western Interior Seaway. The unexpected abundance of ants from extant families and the virtual absence of arthropods from common, exclusively Cretaceous families suggests that Big Muddy amber may represent a yet unsampled Late Cretaceous environment and provides evidence of a faunal transition before the end of the Cretaceous.

Properhyssalus, a new fossil genus of braconid wasps close to the genus Rhyssalus Haliday (Hymenoptera: Braconidae: Rhyssalinae) from Baltic amber.

The new braconid genus and species from the subfamily Rhyssalinae, Properhyssalus szechowskii Belokobylskij, gen. et sp. nov., from late Eocene Baltic amber are described and illustrated. The differences between the new genus and the type species of Rhyssalus Haliday, 1833, Rh. clavator Haliday, 1833, are provided. The position of the previously described from Baltic amber species Rhyssalus brevicornis Brues, 1933 and Rh. rugosus Brues, 1933, as well as Palaeorhyssalus dubitosus Brues, 1933, are discussed.

A new species of Eocene fossil Malthodes (Coleoptera: Cantharidae) with a rarely observed aedeagus.

A new Malthodes (Coleoptera, Cantharidae, Malthininae) in Eocene amber from Poland (Baltic amber) is illustrated and described here: Malthodes (s. str.) maximiliani sp. nov. The new species of soldier beetle is characterized by its small body size and last tergites and last sternite distinct from similar taxa. Notably, the specimen has a partially visible aedeagus, a characteristic rarely seen in fossil Malthodes of the nominotypical subgenus; it has been observed in only three fossil species of this subgenus.

First record of the parasitoid family Ismaridae (Hymenoptera, Diaprioidea) from Eocene Baltic and Rovno ambers with the description of a new genus and two new species.

A new genus, Lubomirus gen. nov., of the family Ismaridae (Hymenoptera, Diaprioidea) from the late Eocene Baltic and Rovno ambers is described and illustrated. The new genus is the second known genus of the family Ismaridae. The two new species, Lubomirus masneri sp. nov. from Rovno amber and Lubomirus victori sp. nov. from Baltic amber, are described. The diagnosis of the Ismaridae family and the single extant genus of the familyIsmarus Halidayare clarified. The systematic position of Cretapria Fujiyama, 1994 is discussed.

Extinct and extant termites reveal the fidelity of behavior fossilization in amber.

Fossils encompassing multiple individuals provide rare direct evidence of behavioral interactions among extinct organisms. However, the fossilization process can alter the spatial relationship between individuals and hinder behavioral reconstruction. Here, we report a Baltic amber inclusion preserving a female-male pair of the extinct termite species Electrotermes affinis. The head-to-abdomen contact in the fossilized pair resembles the tandem courtship behavior of extant termites, although their parallel body alignment differs from the linear alignment typical of tandem runs. To solve this inconsistency, we simulated the first stage of amber formation, the immobilization of captured organisms, by exposing living termite tandems to sticky surfaces. We found that the posture of the fossilized pair matches trapped tandems and differs from untrapped tandems. Thus, the fossilized pair likely is a tandem running pair, representing the direct evidence of the mating behavior of extinct termites. Furthermore, by comparing the postures of partners on a sticky surface and in the amber inclusion, we estimated that the male likely performed the leader role in the fossilized tandem. Our results demonstrate that past behavioral interactions can be reconstructed despite the spatial distortion of body poses during fossilization. Our taphonomic approach demonstrates how certain behaviors can be inferred from fossil occurrences.

Changes in coastal ecosystems affected by overburden dumping from amber open-cut mining on the Sambia Peninsula (Baltic Sea).

The Sambia Peninsula (Kaliningrad region) is historically well known for its amber mining. The 2019 year was the last year of direct overburden disposal into the Baltic Sea as a part of technological amber mining process. The extremely high-suspended particulate matter concentrations during that disposal were recorded immediately after the discharge of significant volumes of pulp and reached 200 mg/L. The impact of pulp discharge had sequentially suppressed plankton communities development due to the high content of suspended solids and afterwards stimulated plankton development due to the glauconite infusion. Cladocera were the most sensitive group to the effects of suspended matter. According to the preliminary forecast, when the pulp discharge stops, the restoration of plankton communities may take from 1 to 2 seasons to 1 year for different groups. This is due to the timing of the removal of fine suspended particulate matter from sediments and the possibility of secondary entry during resuspension.

Stem chewing lice on Cretaceous feathers preserved in amber.

Phthirapteran lice (true lice or parasitic lice) are a major group of ectoparasitic insects living on their bird or mammal hosts during their entire life cycle.1 Due to their highly specialized lifestyles, they are extremely poorly represented in fossil records.2 Molecular clock estimations have speculated extensively about the origin time of parasitic lice,3,4 yet none have been confirmed unequivocally. Herein, we report a new family of stem chewing lice, based on two adult insects associated with several semiplume feathers preserved within a piece of Kachin amber from the mid-Cretaceous. They display some defining characteristics of the Amblycera, an early-diverging lineage of the crown lice group. These features include a wingless body, chewing mouthparts, narrow and small thorax, and short tarsus with elongated euplantulae. Our phylogenetic analysis places the new taxa in the Amblycera, and the discovery thus pushes back the lice fossil records by at least 55 million years. Furthermore, the new specimens show primitive characters such as compressed and club-shaped terminal segments of antennae, maxillary and labial palps, and unmodified femora of hind legs, providing key information for the evolutionary relationship between free-living booklice and parasitic lice. This suggests that some ectoparasitic characters defining the crown lice group might have evolved among amblyceran and non-amblyceran lice in parallel. These newly described fossil specimens imply at least a Cretaceous age of Phthiraptera.

A new mayfly subfamily sheds light on the early evolution and Pangean origin of Baetiscidae (Insecta: Ephemeroptera).

The family Baetiscidae Edmunds & Traver, 1954 is recognisable among mayflies due to its bizarre larvae, equipped with a robust and spiked thoracic notal shield covering part of the abdomen up to sixth segment. Originally being described as extant species from the USA and Canada, Baetiscidae were later found in the fossil record as well, specifically in Cretaceous of Brazil and Eocene Baltic amber. An enigmatic piece of fossil evidence are two larvae from the Early Cretaceous Koonwarra Fossil Bed in Australia, which have been presumed as attributable to Baetiscidae and briefly discussed in previous studies. In the present contribution, we reinvestigate these fossils and confirm their attribution to the family Baetiscidae. These larvae are depicted and described as Koonwarrabaetisca jelli gen. et sp. nov. and Koonwarrabaetisca duncani sp. nov. For both Cretaceous genera Protobaetisca Staniczek, 2007 and Koonwarrabaetisca gen. nov. we establish a new subfamily Protobaetiscinae subfam. nov. within the family Baetiscidae, based on the presence of markedly shortened thoracic sterna. The phylogenetic position of newly described subfamily is clarified using a cladistic analysis; Protobaetiscinae subfam. nov. forms a monophyletic clade, sister to Baetiscinae. The confirmation of the distribution of Baetiscidae in the Cretaceous of Australia suggests almost worldwide distribution of this family in the deep time. Given their limited dispersal abilities, this distributional pattern can be best explained by the Pangean origin for this family, moving the time of their origin at least to the Early Jurassic. The larvae of Koonwarrabaetisca gen. nov. exhibit the same ecomorphological specialization as the rest of Baetiscidae, that supporting with a high probability their lifestyle similar to extant Baetisca Walsh, 1862. The larvae probably lived in the flowing water with stony substrate densely covered by filamentous algae, and in the places of accumulation of dead plant and algae matter during the last instars. Thus, Koonwarrabaetisca gen. nov. could be the allochthonous component in mayfly fauna of the Koonwarra paleolake.