Defensive glands in Stylotermitidae (Blattodea, Isoptera).

The large abundance of termites is partially achieved by their defensive abilities. Stylotermitidae represented by a single extant genus, Stylotermes, is a member of a termite group Neoisoptera that encompasses 83% of termite species and 94% of termite genera and is characterized by the presence of the frontal gland. Within Neoisoptera, Stylotermitidae represents a species-poor sister lineage of all other groups. We studied the structure of the frontal, labral and labial glands in soldiers and workers of Stylotermes faveolus, and the composition of the frontal gland secretion in S. faveolus and Stylotermes halumicus. We show that the frontal gland is a small active secretory organ in soldiers and workers. It produces a cocktail of monoterpenes in soldiers, and some of these monoterpenes and unidentified proteins in workers. The labral and labial glands are developed similarly to other termite species and contribute to defensive activities (labral in both castes, labial in soldiers) or to the production of digestive enzymes (labial in workers). Our results support the importance of the frontal gland in the evolution of Neoisoptera. Toxic, irritating and detectable monoterpenes play defensive and pheromonal functions and are likely critical novelties contributing to the ecological success of these termites.

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.

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.

New stenurothripid thrips from mid-Cretaceous Kachin amber (Thysanoptera, Stenurothripidae).

Hitherto, only two species of the thysanopteran suborder Terebrantia have been reported from mid-Cretaceous Kachin amber (Myanmar). This is here expanded through the discovery of two new genera and species, described and figured as Parallelothripsseparatusgen. et sp. nov. and Didymothripsabdominalisgen. et sp. nov., both of the family Stenurothripidae. Both taxa have key apomorphies of the Stenurothripidae, allowing for a confident assignment as to family. Both species have characteristic comb-like anteromarginal setae, which are discussed along with structural differences between the two sexes. Cycad pollen was found on the thrips' bodies, providing further evidence that Thysanoptera were pollinators of gymnosperms during the mid-Cretaceous.

The taxonomic identity of Trigona lacteipennis Friese, 1900 and description of a new bee species of the Trigona cilipes (Fabricius, 1804) group (Hymenoptera: Apidae: Meliponini).

A critical study of type material for Trigona lacteipennis Friese, 1900 and allied species is presented along with a discussion of literature data. This work reveals that T. lacteipennis is a junior subjective synonym of Melipona paupera Provancher, 1889 (new synonymy), a species currently classified placed in the genus Frieseomelitta. Similarly, T. lehmanni Friese, 1901 and T. parastigma Cockerell, 1918 are also junior subjective synonyms of Melipona paupera (new synonymies). Morphological characters, photographic illustrations of type specimens, and data on distribution are provided for Frieseomelitta paupera. Also, we describe a new species of Trigona cilipes species group, Trigona (Aphaneuropsis) silveirai sp. nov., which is very similar to the former and had been misidentified with it.

The earliest fossil mosquito.

Female mosquitoes are among the most notorious blood-feeding insects, sometimes causing severe allergic responses or vectoring a variety of microbial pathogens.1,2 Hematophagy in insects is likely a feeding shift from plant fluids, with the piercing-sucking mouthparts serving as suitable exaptation for piercing vertebrates' skin. The origins of these habits are mired in an often-poor fossil record for many hematophagous lineages,3,4 particularly those of sufficient age, as to give insights into the paleoecological context in which blood feeding first appeared or even to arrive at gross estimates as to when such shifts have occurred. This is certainly the case for mosquitoes, a clade estimated molecularly to date back to the Jurassic.5 The known Mesozoic Culicidae are Late Cretaceous, assigned to the modern Anophelinae or to the extinct Burmaculicinae, sister to other Culicidae, all with mouthparts of a modern type. Here, we report the discovery, in Lower Cretaceous amber from Lebanon, of two conspecific male mosquitoes unexpectedly with piercing mouthparts, armed with denticulate sharp mandibles and laciniae. These male fossils were likely hematophagous. They represent a lineage that diverged earlier than Burmaculicinae, extending the definitive occurrence of the family into the Early Cretaceous and serving to narrow the ghost-lineage gap for mosquitoes.

Independent wing reductions and losses among stick and leaf insects (Phasmatodea), supported by new Cretaceous fossils in amber.

BACKGROUND: Phasmatodea (stick and leaf insects) play a central role on the debate regarding wing reduction and loss, and its wings are putative reacquisition from secondarily wingless ancestors based solely on extant species. A pivotal taxon in this respect is the species-poor Timematodea, consisting of approximately 21 wingless extant species, which form the sister group of all remaining winged or wingless stick and leaf insects, the Euphasmatodea. RESULTS: Herein, the new fossils of Timematodea from mid-Cretaceous Kachin amber are reported, with winged and wingless species co-occurring. The palaeogeographic distributions of all fossils of Holophasmatodea are summarized, showing their wide paleo-distributions. The phylogenetic analysis based on morphological characters confirms the earliest-diverging lineage of winged Breviala cretacea gen. et sp. nov. in Timematodea, and the possible relationships among all families of Holophasmatodea. These are critical for the reconstruction of patterns of wing evolution in early Phasmatodea. CONCLUSIONS: The new fossils suggest that Timematodea once had wings, at least during the mid-Cretaceous. The palaeogeographic occurrences imply that Timematodea probably have been widely distributed since at least the Jurassic. The phylogenetic analysis with the ancestral-state reconstruction of wings indicates that the common ancestors of Holophasmatodea were winged, the reductions and losses of wings among Timematodea and Euphasmatodea have occurred independently since at least the Cretaceous, and the reduction or loss of the forewing earlier than the hind wings.

Phylogenomics of weevils revisited: data curation and modelling compositional heterogeneity.

Weevils represent one of the most prolific radiations of beetles and the most diverse group of herbivores on land. The phylogeny of weevils (Curculionoidea) has received extensive attention, and a largely satisfactory framework for their interfamilial relationships has been established. However, a recent phylogenomic study of Curculionoidea based on anchored hybrid enrichment (AHE) data yielded an abnormal placement for the family Belidae (strongly supported as sister to Nemonychidae + Anthribidae). Here we reanalyse the genome-scale AHE data for Curculionoidea using various models of molecular evolution and data filtering methods to mitigate anticipated systematic errors and reduce compositional heterogeneity. When analysed with the infinite mixture model CAT-GTR or using appropriately filtered datasets, Belidae are always recovered as sister to the clade (Attelabidae, (Caridae, (Brentidae, Curculionidae))), which is congruent with studies based on morphology and other sources of molecular data. Although the relationships of the 'higher Curculionidae' remain challenging to resolve, we provide a consistent and robust backbone phylogeny of weevils. Our extensive analyses emphasize the significance of data curation and modelling across-site compositional heterogeneity in phylogenomic studies.

Stingless bee classification and biology (Hymenoptera, Apidae): a review, with an updated key to genera and subgenera.

Stingless bees (Meliponini) are a ubiquitous and diverse element of the pantropical melittofauna, and have significant cultural and economic importance. This review outlines their diversity, and provides identification keys based on external morphology, brief accounts for each of the recognized genera, and an updated checklist of all living and fossil species. In total there are currently 605 described extant species in 45 extant genera, and a further 18 extinct species in nine genera, seven of which are extinct. A new fossil genus, Adactylurina Engel, gen. nov., is also described for a species in Miocene amber from Ethiopia. In addition to the systematic review, the biology of stingless bees is summarized with an emphasis on aspects related to their nesting biology and architecture.

New Earwigs from the Middle Jurassic Jiulongshan Formation of Northeastern China (Dermaptera).

Two new genera and species of Dermaptera are described from the Middle Jurassic Jiulongshan Formation of Daohugou, Inner Mongolia, China: Applanatiforceps angustus gen. et sp. nov. in the archidermapteran family Protodiplatyidae, and Ekpagloderma gracilentum gen et sp. nov. in the eodermapteran family Semenoviolidae. Applanatiforceps shares the typical characters of the extinct suborder Archidermaptera (e.g., pentamerous meta tarsi, filiform and multimerous cerci) and externalized ovipositor. The family identity of the Protodiplatyidae can be further distinguished by comparing this new genus with other genera of the Protodiplatyidae. As a result of its large compound eyes, tegmina without venation, body sparsely setose, legs rather short and slender, and shape of the veinless tegmina, Ekpagloderma is classified in the subfamily Aglyptodermatinae. Ekpagloderma not only has the typical features of the Aglyptodermatinae, but also exhibits a more primitive slender segmented cerci, which is different from all other genera of Eodermaptera. In fact, the diversity of Eodermaptera as known today indicates some of the challenges in understanding the suborder and whether or not it is monophyletic as historically construed, or if the separation of Turanodermaptera is justified.

The potter wasp genus Allorhynchium from Vietnam, with descriptions of three new species and a new country record (Hymenoptera, Vespidae, Eumeninae).

Species of the potter wasp genus Allorhynchium van der Vecht (Eumeninae: Odynerini) occurring in Vietnam are presented. Seven species have been recorded from Vietnam. Of them, three species are described as new to science: Allorhynchiumlatum Nguyen, Tran & MT Nguyen, sp. nov., A.moerum Nguyen & AD Nguyen, sp. nov., and A.setosum Nguyen & Engel, sp. nov., and one species, A.argentatum (Fabricius, 1804), is recorded from Vietnam for the first time. An updated key to the Oriental species of the genus is presented.

300 Million years of coral treaders (Insecta: Heteroptera: Hermatobatidae) back to the ocean in the phylogenetic context of Arthropoda.

Among hundreds of insect families, Hermatobatidae (commonly known as coral treaders) is one of the most unique. They are small, wingless predaceous bugs in the suborder Heteroptera. Adults are almost black in colour, measuring about 5 mm in body length and 3 mm in width. Thirteen species are known from tropical coral reefs or rocky shores, but their origin and evolutionary adaptation to their unusual marine habitat were unexplored. We report here the genome and metagenome of Hermatobates lingyangjiaoensis, hitherto known only from its type locality in the South China Sea. We further reconstructed the evolutionary history and origin of these marine bugs in the broader context of Arthropoda. The dated phylogeny indicates that Hexapoda diverged from their marine sister groups approximately 498 Ma and that Hermatobatidae originated 192 Ma, indicating that they returned to an oceanic life some 300 Myr after their ancestors became terrestrial. Their origin is consistent with the recovery of tropical reef ecosystems after the end-Triassic mass extinction, which might have provided new and open niches for them to occupy and thrive. Our analyses also revealed that both the genome changes and the symbiotic bacteria might have contributed to adaptations necessary for life in the sea.

The bee genus Anthidiellum in Vietnam: descriptions of five new species and the first male of Anthidiellumcoronum (Hymenoptera, Megachilidae).

The Vietnamese fauna of bees in the Anthidiellum Cockerell (Megachilinae, Anthidiini) is reviewed. Seven species are recognized, representing two subgenera. Five new species are described and figured as: Anthidiellum (Clypanthidium) nahang Tran, Engel & Nguyen, sp. nov., A. (Pycnanthidium) ayun Tran, Engel & Nguyen, sp. nov., A. (P.) chumomray Tran, Engel & Nguyen, sp. nov., A. (P.) flavaxilla Tran, Engel & Nguyen, sp. nov., and A. (P.) cornu Tran, Engel & Nguyen, sp. nov. from the northern and central highlands of Vietnam. Two previously described species are newly recorded for the fauna: A. (P.) carinatum (Wu) and A. (P.) coronum (Wu), with the male of the latter species described and illustrated for the first time. An identification key is provided for all species of Anthidiellum occurring in Vietnam.

Insect pollination in deep time.

Inferring insect pollination from compression fossils and amber inclusions is difficult because of a lack of consensus on defining an insect pollinator and the challenge of recognizing this ecological relationship in deep time. We propose a conceptual definition for such insects and an operational classification into pollinator or presumed pollinator. Using this approach, we identified 15 insect families that include fossil pollinators and show that pollination relationships have existed since at least the Upper Jurassic (~163 Ma). Insects prior to this can only be classified as presumed pollinators. This gives a more nuanced insight into the origin and evolution of an ecological relationship that is vital to the establishment, composition and conservation of modern terrestrial ecosystems.

Impact of Wood Age on Termite Microbial Assemblages.

The decomposition of wood and detritus is challenging to most macroscopic organisms due to the recalcitrant nature of lignocellulose. Moreover, woody plants often protect themselves by synthesizing toxic or nocent compounds which infuse their tissues. Termites are essential wood decomposers in warmer terrestrial ecosystems and, as such, they have to cope with high concentrations of plant toxins in wood. In this paper, we evaluated the influence of wood age on the gut microbial (bacterial and fungal) communities associated with the termites Reticulitermes flavipes (Rhinotermitidae) (Kollar, 1837) and Microcerotermes biroi (Termitidae) (Desneux, 1905). We confirmed that the secondary metabolite concentration decreased with wood age. We identified a core microbial consortium maintained in the gut of R. flavipes and M. biroi and found that its diversity and composition were not altered by the wood age. Therefore, the concentration of secondary metabolites had no effect on the termite gut microbiome. We also found that both termite feeding activities and wood age affect the wood microbiome. Whether the increasing relative abundance of microbes with termite activities is beneficial to the termites is unknown and remains to be investigated. IMPORTANCE Termites can feed on wood thanks to their association with their gut microbes. However, the current understanding of termites as holobiont is limited. To our knowledge, no studies comprehensively reveal the influence of wood age on the termite-associated microbial assemblage. The wood of many tree species contains high concentrations of plant toxins that can vary with their age and may influence microbes. Here, we studied the impact of Norway spruce wood of varying ages and terpene concentrations on the microbial communities associated with the termites Reticulitermes flavipes (Rhinotermitidae) and Microcerotermes biroi (Termitidae). We performed a bacterial 16S rRNA and fungal ITS2 metabarcoding study to reveal the microbial communities associated with R. flavipes and M. biroi and their impact on shaping the wood microbiome. We noted that a stable core microbiome in the termites was unaltered by the feeding substrate, while termite activities influenced the wood microbiome, suggesting that plant secondary metabolites have negligible effects on the termite gut microbiome. Hence, our study shed new insights into the termite-associated microbial assemblage under the influence of varying amounts of terpene content in wood and provides a groundwork for future investigations for developing symbiont-mediated termite control measures.

Impact of genus (Geotrigona, Melipona, Scaptotrigona) in the targeted 1H-NMR organic profile, and authenticity test by interphase emulsion of honey processed in cerumen pots by stingless bees in Ecuador.

The biodiversity of Ecuadorian stingless bees is almost 200 species. Traditional pot-honey harvest in Ecuador is mostly done from nests of the three genera selected here Geotrigona Moure, 1943, Melipona Illiger, 1806, and Scaptotrigona Moure, 1942. The 20 pot-honey samples collected from cerumen pots and three ethnic honeys "abeja de tierra", "bermejo", and "cushillomishki" were analyzed for qualitative and quantitative targeted 1H-NMR honey profiling, and for the Honey Authenticity Test by Interphase Emulsion (HATIE). Extensive data of targeted organic compounds (41 parameters) were identified, quantified, and described. The three honey types were compared by ANOVA. Amino acids, ethanol, hydroxymethylfurfural, aliphatic organic acids, sugars, and markers of botanical origin. The number of phases observed with the HATIE were one in Scaptotrigona and three in Geotrigona and Melipona honeys. Acetic acid (19.60 ± 1.45 g/kg) and lactic acid (24.30 ± 1.65 g/kg) were particularly high in Geotrigona honey (in contrast to 1.3 g/kg acetic acid and 1.6 g/kg lactic acid in Melipona and Scaptotrigona), and with the lowest fructose + glucose (18.39 ± 1.68) g/100g honey compared to Melipona (52.87 ± 1.75) and Scaptotrigona (52.17 ± 0.60). Three local honeys were tested using PCA (Principal Component Analysis), two were assigned with a correct declared bee origin, but "bermejo" was not a Melipona and grouped with the Scaptotrigona cluster. However after HCA (Hierarchical Cluster Analysis) the three honeys were positioned in the Melipona-Scaptotrigona cluster. This research supports targeted 1H-NMR-based profiling of pot-honey metabolomics approach for multi-parameter visualization of organic compounds, as well as descriptive and pertained multivariate statistics (HCA and PCA) to discriminate the stingless bee genus in a set of Geotrigona, Melipona and Scaptotrigona honey types. The NMR characterization of Ecuadorian honey produced by stingless bees emphasizes the need for regulatory norms. A final note on stingless bee markers in pot-honey metabolites which should be screened for those that may extract phylogenetic signals from nutritional traits of honey. Scaptotrigona vitorum honey revealed biosurfactant activity in the HATIE, originating a fingerprint Honey Biosurfactant Test (HBT) for the genus in this set of pot-honeys.

Diversity, Form, and Postembryonic Development of Paleozoic Insects.

While Mesozoic, Paleogene, and Neogene insect faunas greatly resemble the modern one, the Paleozoic fauna provides unique insights into key innovations in insect evolution, such as the origin of wings and modifications of postembryonic development including holometaboly. Deep-divergence estimates suggest that the majority of contemporary insect orders originated in the Late Paleozoic, but these estimates reflect divergences between stem groups of each lineage rather than the later appearance of the crown groups. The fossil record shows the initial radiations of the extant hyperdiverse clades during the Early Permian, as well as the specialized fauna present before the End Permian mass extinction. This review summarizes the recent discoveries related to the documented diversity of Paleozoic hexapods, as well as current knowledge about what has actually been verified from fossil evidence as it relates to postembryonic development and the morphology of different body parts.

Alarm communication predates eusociality in termites.

Termites (Blattodea: Isoptera) have evolved specialized defensive strategies for colony protection. Alarm communication enables workers to escape threats while soldiers are recruited to the source of disturbance. Here, we study the vibroacoustic and chemical alarm communication in the wood roach Cryptocercus and in 20 termite species including seven of the nine termite families, all life-types, and all feeding and nesting habits. Our multidisciplinary approach shows that vibratory alarm signals represent an ethological synapomorphy of termites and Cryptocercus. In contrast, chemical alarms have evolved independently in several cockroach groups and at least twice in termites. Vibroacoustic alarm signaling patterns are the most complex in Neoisoptera, in which they are often combined with chemical signals. The alarm characters correlate to phylogenetic position, food type and hardness, foraging area size, and nesting habits. Overall, species of Neoisoptera have developed the most sophisticated communication system amongst termites, potentially contributing to their ecological success.

High acoustic diversity and behavioral complexity of katydids in the Mesozoic soundscape.

Acoustic communication has played a key role in the evolution of a wide variety of vertebrates and insects. However, the reconstruction of ancient acoustic signals is challenging due to the extreme rarity of fossilized organs. Here, we report the earliest tympanal ears and sound-producing system (stridulatory apparatus) found in exceptionally preserved Mesozoic katydids. We present a database of the stridulatory apparatus and wing morphology of Mesozoic katydids and further calculate their probable singing frequencies and analyze the evolution of their acoustic communication. Our suite of analyses demonstrates that katydids evolved complex acoustic communication including mating signals, intermale communication, and directional hearing, at least by the Middle Jurassic. Additionally, katydids evolved a high diversity of singing frequencies including high-frequency musical calls, accompanied by acoustic niche partitioning at least by the Late Triassic, suggesting that acoustic communication might have been an important driver in the early radiation of these insects. The Early-Middle Jurassic katydid transition from Haglidae- to Prophalangopsidae-dominated faunas coincided with the diversification of derived mammalian clades and improvement of hearing in early mammals, supporting the hypothesis of the acoustic coevolution of mammals and katydids. Our findings not only highlight the ecological significance of insects in the Mesozoic soundscape but also contribute to our understanding of how acoustic communication has influenced animal evolution.