The First Cretaceous Epyrine Wasp (Hymenoptera: Bethylidae): A New Genus and Species from Early Cenomanian Kachin Amber.

The Epyrinae are the second largest subfamily of Bethylidae and the most diverse in the fossil record. However, although six of the nine bethylid subfamilies are known during the Cretaceous (either as compression or amber fossils), the Epyrinae were hitherto unknown before the lower Eocene. In this contribution, we report the discovery of the oldest member of this group, based on a female specimen from the early Cenomanian amber of Kachin, Myanmar. We describe and illustrate a new genus and species, Hukawngepyris setosus gen. et sp. nov. The new genus is compared with the other epyrine genera and characterized by a unique combination of characters not known in the subfamily. Hukawngepyris setosus gen. et sp. nov. is especially unique in the configuration of the forewing venation, with a complete 2r-rs&Rs vein, curved towards the anterior wing margin, and the presence of three proximal and three distal hamuli. The key to the genera of Epyrinae is accommodated to include the newly erected genus.

The Angiosperm Terrestrial Revolution buffered ants against extinction.

With ~14,000 extant species, ants are ubiquitous and of tremendous ecological importance. They have undergone remarkable diversification throughout their evolutionary history. However, the drivers of their diversity dynamics are not well quantified or understood. Previous phylogenetic analyses have suggested patterns of diversity dynamics associated with the Angiosperm Terrestrial Revolution (ATR), but these studies have overlooked valuable information from the fossil record. To address this gap, we conducted a comprehensive analysis using a large dataset that includes both the ant fossil record (~24,000 individual occurrences) and neontological data (~14,000 occurrences), and tested four hypotheses proposed for ant diversification: co-diversification, competitive extinction, hyper-specialization, and buffered extinction. Taking into account biases in the fossil record, we found three distinct diversification periods (the latest Cretaceous, Eocene, and Oligo-Miocene) and one extinction period (Late Cretaceous). The competitive extinction hypothesis between stem and crown ants is not supported. Instead, we found support for the co-diversification, buffered extinction, and hyper-specialization hypotheses. The environmental changes of the ATR, mediated by the angiosperm radiation, likely played a critical role in buffering ants against extinction and favoring their diversification by providing new ecological niches, such as forest litter and arboreal nesting sites, and additional resources. We also hypothesize that the decline and extinction of stem ants during the Late Cretaceous was due to their hyper-specialized morphology, which limited their ability to expand their dietary niche in changing environments. This study highlights the importance of a holistic approach when studying the interplay between past environments and the evolutionary trajectories of organisms.

Trait-Based Paleontological Niche Prediction Recovers Extinct Ecological Breadth of the Earliest Specialized Ant Predators.

AbstractPaleoecological estimation is fundamental to the reconstruction of evolutionary and environmental histories. The ant fossil record preserves a range of species in three-dimensional fidelity and chronicles faunal turnover across the Cretaceous and Cenozoic; taxonomically rich and ecologically diverse, ants are an exemplar system to test new methods of paleoecological estimation in evaluating hypotheses. We apply a broad extant ecomorphological dataset to evaluate random forest machine learning classification in predicting the total ecological breadth of extinct and enigmatic hell ants. In contrast to previous hypotheses of extinction-prone arboreality, we find that hell ants were primarily leaf litter or ground-nesting and foraging predators, and by comparing ecospace occupations of hell ants and their extant analogs, we recover a signature of ecomorphological turnover across temporally and phylogenetically distinct lineages on opposing sides of the Cretaceous-Paleogene boundary. This paleoecological predictive framework is applicable across lineages and may provide new avenues for testing hypotheses over deep time.

Multiple drivers and lineage-specific insect extinctions during the Permo-Triassic.

The Permo-Triassic interval encompasses three extinction events including the most dramatic biological crisis of the Phanerozoic, the latest Permian mass extinction. However, their drivers and outcomes are poorly quantified and understood for terrestrial invertebrates, which we assess here for insects. We find a pattern with three extinctions: the Roadian/Wordian (≈266.9 Ma; extinction of 64.5% insect genera), the Permian/Triassic (≈252 Ma; extinction of 82.6% insect genera), and the Ladinian/Carnian boundaries (≈237 Ma; extinction of 74.8% insect genera). We also unveil a heterogeneous effect of these extinction events across the major insect clades. Because extinction events have impacted Permo-Triassic ecosystems, we investigate the influence of abiotic and biotic factors on insect diversification dynamics and find that changes in floral assemblages are likely the strongest drivers of insects' responses throughout the Permo-Triassic. We also assess the effect of diversity dependence between three insect guilds; an effect ubiquitously found in current ecosystems. We find that herbivores held a central position in the Permo-Triassic interaction network. Our study reveals high levels of insect extinction that profoundly shaped the evolutionary history of the most diverse non-microbial lineage.

Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae).

Fossils are critical for understanding the evolutionary diversification, turnover, and morphological disparification of extant lineages. While fossils cannot be sequenced, phenome-scale data may be generated using micro-computed tomography (µ-CT), thus revealing hidden structures and internal anatomy, when preserved. Here, we adduce the male caste of a new fossil ant species from Miocene Ethiopian amber that resembles members of the Aneuretinae, matching the operational definition of the subfamily. Through the use of synchrotron radiation for µ-CT, we critically test the aneuretine-identity hypothesis. Our results indicate that the new fossils do not belong to the Aneuretinae, but rather the Ponerini (Ponerinae). Informed by recent phylogenomic studies, we were able to place the fossils close to the extant genus Cryptopone based on logical character analysis, with the two uniquely sharing absence of the subpetiolar process among all ponerine genera. Consequently, we: (1) revise the male-based key to the global ant subfamilies; (2) revise the definitions of Aneuretinae, Ponerinae, Platythyreini, and Ponerini; (3) discuss the evolution of ant mandibles; and (4) describe the fossils as †Desyopone hereon gen. et sp. nov. Our study highlights the value of males for ant systematics and the tremendous potential of phenomic imaging technologies for the study of ant evolution.

Cretaceous Antodicranomyia (Diptera: Limoniidae) and their paleohabitat.

New representatives of the Cretaceous cranefly genus Antodicranomyia (Diptera: Limoniidae) are reported from Albian-Cenomanian Charentese (French) amber. The newly reported specimens allow for an emended diagnosis of the type species A. azari, as well as the description of a new species, Antodicranomyia rubra sp. nov., which is mostly distinguished from the type species by features of its wing venation, antennae, and genitalia. As a rare, extinct genus known only from French amber, Antodicranomyia is compared with its closest relative genera Antocha, Dicranomyia and Antohelia. The evolutionary implications and paleohabitat of Antodicranomyia are discussed. The new discovery adds to the knowledge of the crane flies' diversity and evolution in the mid-Cretaceous.

The mid-Miocene Zhangpu biota reveals an outstandingly rich rainforest biome in East Asia.

During the Mid-Miocene Climatic Optimum [MMCO, ~14 to 17 million years (Ma) ago], global temperatures were similar to predicted temperatures for the coming century. Limited megathermal paleoclimatic and fossil data are known from this period, despite its potential as an analog for future climate conditions. Here, we report a rich middle Miocene rainforest biome, the Zhangpu biota (~14.7 Ma ago), based on material preserved in amber and associated sedimentary rocks from southeastern China. The record shows that the mid-Miocene rainforest reached at least 24.2°N and was more widespread than previously estimated. Our results not only highlight the role of tropical rainforests acting as evolutionary museums for biodiversity at the generic level but also suggest that the MMCO probably strongly shaped the East Asian biota via the northern expansion of the megathermal rainforest biome. The Zhangpu biota provides an ideal snapshot for biodiversity redistribution during global warming.

Specialized Predation Drives Aberrant Morphological Integration and Diversity in the Earliest Ants.

Extinct haidomyrmecine "hell ants" are among the earliest ants known [1, 2]. These eusocial Cretaceous taxa diverged from extant lineages prior to the most recent common ancestor of all living ants [3] and possessed bizarre scythe-like mouthparts along with a striking array of horn-like cephalic projections [4-6]. Despite the morphological breadth of the fifteen thousand known extant ant species, phenotypic syndromes found in the Cretaceous are without parallel and the evolutionary drivers of extinct diversity are unknown. Here, we provide a mechanistic explanation for aberrant hell ant morphology through phylogenetic reconstruction and comparative methods, as well as a newly reported specimen. We report a remarkable instance of fossilized predation that provides direct evidence for the function of dorsoventrally expanded mandibles and elaborate horns. Our findings confirm the hypothesis that hell ants captured other arthropods between mandible and horn in a manner that could only be achieved by articulating their mouthparts in an axial plane perpendicular to that of modern ants. We demonstrate that the head capsule and mandibles of haidomyrmecines are uniquely integrated as a consequence of this predatory mode and covary across species while finding no evidence of such modular integration in extant ant groups. We suggest that hell ant cephalic integration-analogous to the vertebrate skull-triggered a pathway for an ancient adaptive radiation and expansion into morphospace unoccupied by any living taxon.

† Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicoidea).

Fossils provide primary material evidence for the pattern and timing of evolution. The newly discovered "beast ants" from mid-Cretaceous Burmite, †Camelosphecia gen. nov., display an exceptional combination of plesiomorphies, including absence of the metapleural gland, and a series of unique apomorphies. Females and males, represented by †C. fossor sp. nov. and †C. venator sp. nov., differ in a number of features which suggest distinct sexual biologies. Combined-evidence phylogenetic analysis recovers †Camelosphecia and †Camelomecia as a clade which forms the extinct sister group of the Formicidae. Notably, these genera are only known from alate males and females; workers, if present, have yet to be recovered. Based on ongoing study of the total Aculeata informed by the beast ant genera, we provide a brief diagnosis of the Formicoidea. We also provide the first comprehensive key to the major groupings of Mesozoic Formicoidea, alongside a synoptic classification in which †Zigrasimeciinaestat. nov. and †Myanmyrma maraudera comb. nov. are recognized. Finally, a brief diagnosis of the Formicoidea is outlined.

A Late Cretaceous amber biota from central Myanmar.

Insect faunas are extremely rare near the latest Cretaceous with a 24-million-year gap spanning from the early Campanian to the early Eocene. Here, we report a unique amber biota from the Upper Cretaceous (uppermost Campanian ~72.1 Ma) of Tilin, central Myanmar. The chemical composition of Tilin amber suggests a tree source among conifers, indicating that gymnosperms were still abundant in the latest Campanian equatorial forests. Eight orders and 12 families of insects have been found in Tilin amber so far, making it the latest known diverse insect assemblage in the Mesozoic. The presence of ants of the extant subfamilies Dolichoderinae and Ponerinae supports that tropical forests were the cradle for the diversification of crown-group ants, and suggests that the turnover from stem groups to crown groups had already begun at ~72.1 Ma. Tilin amber biota fills a critical insect faunal gap and provides a rare insight into the latest Campanian forest ecosystem.

Production and preservation of resins - past and present.

Amber is fossilised plant resin. It can be used to provide insights into the terrestrial conditions at the time the original resin was exuded. Amber research thus can inform many aspects of palaeontology, from the recovery and description of enclosed fossil organisms (biological inclusions) to attempts at reconstruction of past climates and environments. Here we focus on the resin itself, the conditions under which it may have been exuded, and its potential path to fossilisation, rather than on enclosed fossils. It is noteworthy that not all plants produce resin, and that not all resins can (nor do) become amber. Given the recent upsurge in the number of amber deposits described, it is time to re-examine ambers from a botanical perspective. Here we summarise the state of knowledge about resin production in modern ecosystems, and review the biological and ecological aspects of resin production in plants. We also present new observations on conifer-derived resin exudation, with a particular focus on araucarian conifer trees. We suggest that besides disease, insect attacks and traumatic wounding from fires and storms, other factors such as tree architecture and local soil conditions are significant in creating and preserving resin outpourings. We also examine the transformation of resin into amber (maturation), focusing on geological aspects of amber deposit formation and preservation. We present new evidence that expands previous understanding of amber deposit formation. Specific geological conditions such as anoxic burial are essential in the creation of amber from resin deposits. We show that in the past, the production of large amounts of resin could have been linked to global climate changes and environmental disruption. We then highlight where the gaps in our knowledge still remain and potential future research directions.

Marsupial brood care in Cretaceous tanaidaceans.

Parental care in animal evolution has long fascinated biologists, but tracing this complex of behavioural repertoires is challenging, as these transitory states often leave no corporeal traces as fossils. Among modern invertebrates, the tanaidaceans (Malacostraca: Peracarida), a lineage of marsupial crustaceans, show an interesting variety of brooding strategies. Here we report on fossil tanaidaceans from the Cretaceous of Spain and France that provide conclusive evidence for marsupial care of brood-offspring. Two exceptionally preserved female specimens of Alavatanais carabe and A. margulisae from Late Albian Peñacerrada I amber (Spain) possess four pairs of rudimentary oostegites, indicating formation of a marsupium. From Recent data, given the taxonomic distribution of a marsupium of four pairs of oostegites, we hypothesize that this may be plesiomorphic for the Tanaidomorpha. We also report on a peculiar tanaidacean specimen referable to the fossil family Alavatanaidae, Daenerytanais maieuticus gen. et sp. nov., from Early Cenomanian La Buzinie amber (France), preserved with its marsupial pouch and content. Our discoveries provide early evidence of the peracarid reproductive strategy, as seen in modern Tanaidacea, and argue that this form of parental care may have played a role in the diversification of the lineage during this period.

100-million-year-old conifer tissues from the mid-Cretaceous amber of Charente (western France) revealed by synchrotron microtomography.

BACKGROUND AND AIMS: Terrestrial plant remains in fossilized tree resin are relatively common. However, histology and preservation of plants entombed in Cretaceous ambers remain poorly known. We report an exquisitely preserved conifer leafy axis from 100-million-year-old opaque amber of western France that is assignable to Glenrosa carentonensis Moreau, Néraudeau, Tafforeau & Dépré. The purpose of this paper is to discuss the taphonomy and the use of microtomography for studies of palaeobotanical remains in amber. METHODS: A leafy axis was examined using propagation phase-contrast X-ray synchrotron microtomography with voxel sizes of 14·9, 1·4, and 0·7 µm. KEY RESULTS: The conifer leafy axis described is preserved in three dimensions. Despite desiccation of the specimen within the surrounding amber, the cuticle, as well as most of inner tissues, is preserved in three dimensions down to the cellular level. Epidermis, palisade parenchyma, spongy parenchyma, transfusion tracheids and vascular bundles are clearly distinguished. CONCLUSIONS: Gross morphology and histology of the specimen were revealed using synchrotron microtomography, allowing an unprecedented resolution for the study of soft-bodied plants entombed in amber. The study reveals a peculiar combination of authigenic and duripartic preservation as well as permineralization, and highlights the complexity of taphonomic processes that can occur in amber inclusions. This fossil demonstrates the difficulty of studying amber-preserved plant remains under certain conditions.

Debris-carrying camouflage among diverse lineages of Cretaceous insects.

Insects have evolved diverse methods of camouflage that have played an important role in their evolutionary success. Debris-carrying, a behavior of actively harvesting and carrying exogenous materials, is among the most fascinating and complex behaviors because it requires not only an ability to recognize, collect, and carry materials but also evolutionary adaptations in related morphological characteristics. However, the fossil record of such behavior is extremely scarce, and only a single Mesozoic example from Spanish amber has been recorded; therefore, little is known about the early evolution of this complicated behavior and its underlying anatomy. We report a diverse insect assemblage of exceptionally preserved debris carriers from Cretaceous Burmese, French, and Lebanese ambers, including the earliest known chrysopoid larvae (green lacewings), myrmeleontoid larvae (split-footed lacewings and owlflies), and reduviids (assassin bugs). These ancient insects used a variety of debris material, including insect exoskeletons, sand grains, soil dust, leaf trichomes of gleicheniacean ferns, wood fibers, and other vegetal debris. They convergently evolved their debris-carrying behavior through multiple pathways, which expressed a high degree of evolutionary plasticity. We demonstrate that the behavioral repertoire, which is associated with considerable morphological adaptations, was already widespread among insects by at least the Mid-Cretaceous. Together with the previously known Spanish specimen, these fossils are the oldest direct evidence of camouflaging behavior in the fossil record. Our findings provide a novel insight into early evolution of camouflage in insects and ancient ecological associations among plants and insects.

Extreme Morphogenesis and Ecological Specialization among Cretaceous Basal Ants.

Ants comprise one lineage of the triumvirate of eusocial insects and experienced their early diversification within the Cretaceous [1-9]. Their ecological success is generally attributed to their remarkable social behavior. Not all ants cooperate in social hunting, however, and some of the most effective predatory ants are solitary hunters with powerful trap jaws [10]. Recent evolutionary studies predict that the early branching lineages of extant ants formed small colonies of ground-dwelling, solitary specialist predators [2, 5, 7, 11, 12], while some Cretaceous fossils suggest group recruitment and socially advanced behavior among stem-group ants [9]. We describe a trap-jaw ant from 99 million-year-old Burmese amber with head structures that presumably functioned as a highly specialized trap for large-bodied prey. These are a cephalic horn resulting from an extreme modification of the clypeus hitherto unseen among living and extinct ants and scythe-like mandibles that extend high above the head, both demonstrating the presence of exaggerated morphogenesis early among stem-group ants. The new ant belongs to the Haidomyrmecini, possibly the earliest ant lineage [9], and together these trap-jaw ants suggest that at least some of the earliest Formicidae were solitary specialist predators. With their peculiar adaptations, haidomyrmecines had a refined ecology shortly following the advent of ants.

Chemical Characterization and Botanical Origin of French Ambers.

The molecular composition of 10 Cretaceous and one Eocene ambers from France was analyzed by infrared spectroscopy, solid-state (13)C nuclear magnetic resonance spectroscopy, and thermochemolysis gas chromatography-mass spectrometry. The terpenoids identified in the samples were used as biomarkers for the botanical origin of the resins. The Cretaceous samples, comprising the so-called Alpine, Anjou, Charentese, Provence, Pyrenean, and Vendean ambers, ranged from the Albian-Cenomanian transition to the early Santonian (100 to 85 Ma) and correspond to class Ib resins typical of conifers. The extinct conifer family Cheirolepidiaceae was proposed as the plant source of Pyrenean and brown Charentese ambers. Araucariaceae or Cheirolepidiaceae were the plant sources of the Cenomanian Alpine, Anjou, and yellow Charentese ambers. The Santonian ambers of Provence and Vendée were found to derive from the Cupressaceae. The Eocene Oise amber (ca. 53 Ma) is a class Ic resin typical of angiosperms and was produced by a Fabaceae. The evolution of resin sources from the early Cretaceous to the Eocene periods is discussed. Finally, a possible fingerprint hitherto unveiled is proposed for cheirolepidiaceous resins, defined by the simultaneous presence of phenolic diterpenoids, labdanoic acids, callitrisate structures, and their respective derivatives.

An ant-associated mesostigmatid mite in Baltic amber.

Fossil mesostigmatid mites (Acari: Parasitiformes: Mesostigmata) are extremely rare, and specimens from only nine families, including four named species, have been described so far. A new record of Myrmozercon sp. described here from Eocene (ca 44-49 Myr) Baltic amber represents the first-and so far only-fossil example of the derived, extant family Laelapidae. Significantly, modern species of this genus are habitually myrmecophilous and the fossil mite described here is preserved attached to the head of the dolichoderine ant Ctenobethylus goepperti (Mayr, 1868). It thus offers the oldest unequivocal evidence for an ecological association between mesostigmatid mites and social insects in the order Hymenoptera.

The genus Macroteleia Westwood in Middle Miocene amber from Peru (Hymenoptera, Platygastridae s.l., Scelioninae).

A new species of the scelionine genus Macroteleia Westwood (Platygastridae s.l., Scelioninae) is described and figured from a female beautifully preserved in Middle Miocene amber from Peru. Macroteleia yaguarum Perrichot & Engel, sp. n., shows a unique combination of characters otherwise seen independently within its congeners. It is most similar to the modern M. surfacei Brues, but differs from it by the non-foveolate notauli, the contiguous punctures of the vertex, and the continuous propodeum. The new species is the first New World fossil of the genus, suggesting a Cretaceous origin for the group and a relatively old age of the South American, tropical African, and Australian faunas, and a younger age of the modern Holarctic faunas.

The first ibis fly in mid-Cretaceous amber of France (Diptera: Athericidae).

A new genus and species of ibis fly is described from an isolated wing in amber from the Late Albian-Early Cenomanian of Charentes, southwestern France. Galloatherix incompletus gen. et sp. n., is the first Athericidae fossilized in Cretaceous amber, and only the eighth Mesozoic species. It adds to the diverse aquatic and semiaquatic paleobiota already identified from Charentese amber.