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Abstract Much of what we know about terrestrial life during the Carboniferous Period comes from Middle Pennsylvanian (~315–307 Mya) Coal Measures deposited in low-lying wetland environments1–5. We know relatively little about terrestrial ecosystems from the Early Pennsylvanian, which was a critical interval for the diversification of insects, arachnids, tetrapods, and seed plants6–10. Here we report a diverse Early Pennsylvanian trace and body fossil Lagerstätte (~320–318 Mya) from the Wamsutta Formation of eastern North America, distinct from coal-bearing deposits, preserved in clastic substrates within basin margin conglomerates. The exceptionally preserved trace fossils and body fossils document a range of vertebrates, invertebrates and plant taxa (n = 131), with 83 distinct foliage morphotypes. Plant-insect interactions include what may be the earliest evidence of insect oviposition. This site expands our knowledge of early terrestrial ecosystems and organismal interactions and provides ground truth for future phylogenetic reconstructions of key plant, arthropod, and vertebrate groups.

Abstract The higher classification of termites requires substantial revision as the Neoisoptera, the most diverse termite lineage, comprise many paraphyletic and polyphyletic higher taxa. Here, we produce an updated termite classification using genomic-scale analyses. We reconstruct phylogenies under diverse substitution models with ultraconserved elements analyzed as concatenated matrices or within the multi-species coalescence framework. Our classification is further supported by analyses controlling for rogue loci and taxa, and topological tests. We show that the Neoisoptera are composed of seven family-level monophyletic lineages, including the Heterotermitidae Froggatt, Psammotermitidae Holmgren, and Termitogetonidae Holmgren, raised from subfamilial rank. The species-rich Termitidae are composed of 18 subfamily-level monophyletic lineages, including the new subfamilies Crepititermitinae, Cylindrotermitinae, Forficulitermitinae, Neocapritermitinae, Protohamitermitinae, and Promirotermitinae; and the revived Amitermitinae Kemner, Microcerotermitinae Holmgren, and Mirocapritermitinae Kemner. Building an updated taxonomic classification on the foundation of unambiguously supported monophyletic lineages makes it highly resilient to potential destabilization caused by the future availability of novel phylogenetic markers and methods. The taxonomic stability is further guaranteed by the modularity of the new termite classification, designed to accommodate as-yet undescribed species with uncertain affinities to the herein delimited monophyletic lineages in the form of new families or subfamilies.

The rare bee genus Habrophorula Lieftinck, 1974 is recorded for the first time from Vietnam. The genus is represented by a new species, Habrophorula belladeceptrix Tran, Engel & Nguyen, sp. nov., from Cao Bang Province and can be most easily confused with H. nigripes Wu from China. The species is most easily differentiated by the unique form of the male terminalia but can also be distinguished by differences in integumental and setal coloration. A revised key is provided to the species of the genus. Females of the new species were collected at flowers of Saurauia roxburghii Wall. and Saurauia napaulensis DC. (Actinidiaceae); males were collected at flowers of Lantana camara L. (Verbenaceae).

Clubtail dragonfly Lindenia heeri sp. nov., is described and figured as the first fossil representative of the extant lindeniine genus Lindenia, based on a finely preserved forewing from the mid-Miocene lacustrine maar of Öhningen, Germany. The new species differs from the type and only species of Lindenia tetraphylla in the dark brown pterostigma covering ten cells vs. only five. Otherwise its forewing venation is identical to that of the modern species. The paleoclimatic data fit well with the climatic preferences of extant Lindenia tetraphylla, consistent with the generally warmer climate of that region during that period of time, allowing for more Mediterranean or even subtropical elements of the flora and fauna to persist.

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 Parallelothrips separatus gen. et sp. nov. and Didymothrips abdominalis gen. 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.

Studying insect fossils, particularly those preserved as compressions in sedimentary matrices, can be difficult due to the taphonomic processes that often result to poor preservation and contrast of structures compared to the embedding matrix. To address this, we propose a user-friendly and simple methodology based on UV-light to study insect fossils and select specimens of interest for more advanced imagery exploration. While UV-light imaging has been previously applied to compressions of arthropod fossils, it typically involved laser light sources. Our approach allows the investigation of fossils using an affordable, compact, and portable UV-light source, along with a simple and replicable low-cost protocol. • The methodology is based on UV-light induced natural fluorescence of sediment and fossil remains. • UV-light is effective on compression fossils to gain natural contrast and enhance observation of body structures like veins or setae on wings. • UV-light is effective to reveal palaeoecological information such as pollen grains preserved on specimens, especially near or on putative pollinator or pollen-eating taxa.

Abstract 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.

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.

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: Allorhynchium latum 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.

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.

The vibroacoustic and chemical alarm communication in the wood roach Cryptocercus and in 20 termite species are studied to reveal that the Neoisoptera species have developed the most sophisticated communication system, which could contribute towards their ecological success.

A new species of the orchid bee genus Eufriesea Cockerell (Apidae: Apinae: Euglossini) is described and figured from the Islas Marías of Nayarit State, México in the Pacific. Eufriesea insularis sp. nov., is a member of the coerulescens species group and is restricted to Islas Marias. The species is readily recognized by its dark blue integument with purple iridescence, black pubescence, dark wings, and clypeus green with purple hues and a prominent elevated ridge along the midline. The new species is known only from the female.

As climatic and other impactful environmental changes continue to gain momentum pollination, services are poised to be harmed, and wild bee species are not an exception. In the present study, maximum entropy (MaxEnt) modeling was used to predict the potential climatic niches of five wild bee species, namely, Chalicodoma flavipes, Chalicodoma sicula, Coelioxys coturnix, Megachile minutissima, and Osmia submicans (all of Megachilidae: Megachilinae). The Maxent model performed better than random for the five species, and all model predictions were significantly robust, giving ratios above null expectations. Under future climate change scenarios, the Maxent model predicted habitat loss for C. flavipes, C. sicula, and M. minutissima in North Africa and habitat loss for O. submicans in Europe and North Africa in all scenarios. Conversely, the study showed that the cleptoparasitic bee Co. coturnix would expand their suitable habitat in most scenarios in Europe, Asia, and the United States, although this species would also suffer habitat loss in North Africa in two scenarios. Between the present situation and future scenarios, the potential distribution for all species decreased in their suitable habitat, with the exception of Co. coturnix. The present results are of considerable value for informed conservation programs and policy decisions regarding wild pollinators.

The little-known megachiline genus Noteriades Cockerell, 1931 is recorded from Vietnam for the first time. A new species, Noteriades hangkia Tran, Engel & Nguyen sp. nov. is described and figured based on a series of females collected from the provinces of the northern and central highlands of Vietnam. The genus is briefly discussed and a new subtribe is established, Noteriadina Engel, Tran & Nguyen subtrib. nov. of Megachilini. Lastly, an identification key and distribution map are provided for those species occurring in Southeast Asia.

Braconid parasitoid wasps are a widely diversified group today, while their fossil record from the Mesozoic is currently poorly known. Here, we describe Utrillabracon electropteron Álvarez-Parra & Engel, gen. et sp. nov., from the upper Albian (Lower Cretaceous) amber of San Just in the eastern Iberian Peninsula. The holotype specimen is incomplete, although the forewing and hind wing venation are well preserved. The new taxon is assigned to the subfamily †Protorhyssalinae (Braconidae) and, based on characteristics of the wing venation, seems to be closely related to Protorhyssalus goldmani Basibuyuk & Quicke, 1999 and Diorhyssalus allani (Brues, 1937), both from Upper Cretaceous ambers of North America. We discuss the taxonomy of the Cretaceous braconids, considering †Seneciobraconinae as a valid subfamily. We also comment on possible relationships within †Protorhyssalinae, although a phylogenetic analysis is necessary. Additionally, a checklist is included of braconids known from Cretaceous ambers.

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.

A new genus of minute stingless bees (Meliponini: Hypotrigonina) is described from Southeast Asia. Ebaiotrigona Engel & Nguyen, gen. nov., is based on the type species Lisotrigona carpenteri Engel, recorded from Vietnam, Thailand, Laos, Cambodia, and southern China. The species was previously considered an enigmatic member of Lisotrigona Moure, but is removed to a new genus based on its unique male terminalia that differs considerably from that of Lisotrigona and instead shares resemblances with Austroplebeia Moure, and the presence of yellow maculation (also similar to that of Austroplebeia). It is possible that Ebaiotrigona is the extant sister group of Austroplebeia, but this requires confirmation by future phylogenetic analyses. Based on available biological observations, Ebaiotrigona carpenteri could not be confirmed as lachryphagous as is well documented from the tear-drinking species of Lisotrigona and Pariotrigona Moure.

Mantis lacewings (Neuroptera: Mantispidae) are prominent and charismatic predatory representatives of Insecta. Nevertheless, representatives of the group are surprisingly scarce in Paleogene deposits after a relative abundance of specimens known from Cretaceous. Here we present Mantispa? damzenogedanica sp. nov., representing the first adult of Mantispidae described from Baltic amber and the only Eocene adult mantispid hitherto preserved in amber. The new fossil species is also among the earliest representatives of Mantispinae, certainly the oldest adult of this group described from amber. Additionally, we discuss the changes through time in the ecological morphospace within Mantispidae based on the morphological diversity (≈disparity) of the raptorial legs. Possible explanations for the post-Cretaceous decline in the morphological diversity of mantis lacewings are posited.

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.

Two new species of wood wasps (Anaxyelidae), Orthosyntexis elegans gen. et sp. nov. and Orthosyntexis thanti sp. nov., are described from mid-Cretaceous Kachin amber. Orthosyntexis gen. nov. exhibits characters and character combinations unique to the family, such as the combination of two mesotibial spurs, a length ratio of forewing 1Rs to 1M

Two new species of Archidermaptera are described and figured from the Middle Jurassic Jiulonghsan Formation of Daohugou, Inner Mongolia, China. Aneuroderma oiodes gen. & sp. nov. is described in the family Protodiplatyidae and Sinopalaeodermata concavum sp. nov. is established in the family Dermapteridae. Both new species share the typical characters of the extinct suborder Archidermaptera (e.g., pentamerous metatarsi, filiform and multimerous cerci, externalized ovipositor). Aneuroderma gen. nov. is compared with other genera of the Protodiplatyidae, while S. concavum sp. nov. allows us to emend the diagnosis of the genus Sinopalaeodermata. We briefly discuss the diversity of Archidermaptera and challenges to understanding relationships among this mid-Mesozoic diversity.

This novel review of analytical methods for pot-honey research was intended to provide concise references to a 35-day post-harvest experiments at 30 °C, in an integrated study. Diverse methods were selected from specialized literature, from the AOAC (Association of Official Analytical Chemists), and the International Honey Commission. Besides the geographical and seasonal origin, the pot-honey I.D. consists of entomological and botanical identifications, the latter performed by acetolyzed or natural melissopalynology. The methods of this integrative study included: 1. Physicochemical analysis (Aw, color, moisture, pH, free acidity, lactone acidity, total acidity, hydroxymethylfurfural (HMF), and sugars by highperformance liquid chromatography HPLC), 2. Targeted proton nuclear magnetic resonance 1H-NMR metabolomics (sugars, ethanol, HMF, aliphatic organic acids, amino acids, and botanical markers), 3. Biochemical composition (flavonoids, polyphenols), 4. Antioxidant activity (ABTS 2,2'-azino-bis(3-ethylbenzothiazoline-6-sulfonic acid-free radical scavenging assay, DPPH 2,2-diphenyl-1-picrylhydrazyl radical scavenging assay, ferric reduction assay FRAP), 5. Microbial counts (aerobic plate, yeast and mold, Bacillus, and lactic acid bacteria count), 6. Honey microbiome profiling via independent-culture method: high-throughput bacteria and fungi based on amplicon sequencing approaches, 7. Sensory evaluation (odor, aroma, taste, persistence), and 8. Honey authenticity and biosurfactant tests by an interphase emulsion. A further section was included to provide basic information on the results obtained using each method. This was needed to explain the interacting components derived from pot-honey processing within the stingless bee nest and post-harvest transformations.

Burrowing bugs are distinctive, beetle-like insects of the pentatomoid family Cydnidae, noteworthy for their morphological specializations for digging and a hemiedaphic life history. However, less is known about their biological significance and the early origin of soil dwelling. Direct fossil evidence illuminating the evolutionary history of soil dwelling in cydnids is extremely rare. In this study, we report four new species of the burrowing bug subfamily Amnestinae from mid-Cretaceous Burmese amber, including two exhibiting specialized bulldozing and digging morphological traits on the anterior of the head and forelegs. Associated morphological features and phylogenetic placement indicate that Acanthamnestus represents the earliest unequivocal soil-dwelling cydnids and pushes back the geological record of hemiedaphic true bugs to 99 Ma. Environmental evidence, the distribution of host plants, and the fossils provide a new window for understanding the early origin of soil habits in Amnestinae and may be linked to the appearance of Moraceae.

In several insect eusocial lineages, e.g., some aphids, thrips, ants, some stingless bees, and termites, task specialization is brought to its climax with a sterile soldier caste solely devoted to colony defense. In Isoptera, while the reproductives are defenseless, the soldiers have unique morpho-physiological specializations whose origin and evolution remain unresolved. Here we report on two instances of Cretaceous fossil termite reproductives belonging to different families († Valkyritermes inopinatus gen. et sp. nov. and an unpublished specimen from the Crato Formation), with intriguing phragmotic soldier-like heads and functional wings. These individuals, herein called Valkyries, are the first termite reproductives known with defensive features and suggest that phragmosis arose at least in the Early Cretaceous. Valkyries resemble modern neotenic soldiers except for their complete wings. Their discovery supports the hypothesis that the division between reproductive (indicated by the winged condition of Valkyries) and defensive tasks (indicated by the phragmotic head) has not always been complete in termite history. We explore two alternative scenarios regarding the origin of Valkyries (i.e., relatively recent and convergent origins vs. plesiomorphic condition) and discuss how they might relate to the development of soldiers. We argue that, in both cases, Valkyries likely evolved to face external threats, a selective pressure that could also have favored the origin of soldiers from helpers. Valkyries highlight the developmental flexibility of termites and illustrate the tortuous paths that evolution may follow.

Beetles constitute the most biodiverse animal order with over 380 000 described species and possibly several million more yet unnamed. Recent phylogenomic studies have arrived at considerably incongruent topologies and widely varying estimates of divergence dates for major beetle clades. Here, we use a dataset of 68 single-copy nuclear protein-coding (NPC) genes sampling 129 out of the 193 recognized extant families as well as the first comprehensive set of fully justified fossil calibrations to recover a refined timescale of beetle evolution. Using phylogenetic methods that counter the effects of compositional and rate heterogeneity, we recover a topology congruent with morphological studies, which we use, combined with other recent phylogenomic studies, to propose several formal changes in the classification of Coleoptera: Scirtiformia and Scirtoidea sensu nov., Clambiformia ser. nov. and Clamboidea sensu nov., Rhinorhipiformia ser. nov., Byrrhoidea sensu nov., Dryopoidea stat. res., Nosodendriformia ser. nov. and Staphyliniformia sensu nov., and Erotyloidea stat. nov., Nitiduloidea stat. nov. and Cucujoidea sensu nov., alongside changes below the superfamily level. Our divergence time analyses recovered a late Carboniferous origin of Coleoptera, a late Palaeozoic origin of all modern beetle suborders and a Triassic–Jurassic origin of most extant families, while fundamental divergences within beetle phylogeny did not coincide with the hypothesis of a Cretaceous Terrestrial Revolution.

A new species of one of the basal families among extant Dermaptera, Pygidicranidae, is described from mid-Cretaceous amber of Myanmar based on two females and a male. Astreptolabis laevis sp. nov., belongs to the extinct subfamily Astreptolabidinae, sharing the diagnostic combination of features typical of this group, such as the well-developed compound eyes, large pronotum, and straight and tubular cerci. The discovery of a male with its genitalia partly exerted permits characterization of traits for the subfamily and provides further information on the uniqueness and affinities of the subfamily. In addition, the extended hind wing allows for a comparison between the folding mechanism between these fossils and their modern counterparts, demonstrating considerable conservatism in hind wing evolution among Dermaptera.

Summary: This protocol explains how to extract pollen from fossil insects with subsequent descriptions of pollen treatment. We also describe how to document morphological and ultrastructural features with light-microscopy and electron microscopy. It enables a taxonomic assignment of pollen that can be used to interpret flower-insect interactions, foraging and feeding behavior of insects, and the paleoenvironment. The protocol is limited by the state of the fossil, the presence/absence of pollen on fossil specimens, and the availability of extant pollen for comparison.For complete details on the use and execution of this protocol, please refer to Wappler et al. (2015), Ulrich and Grímsson (2020), and Wedmann et al. (2021).

Litsea is a group of evergreen trees or shrubs in the laurel family, Lauraceae. Species of the genus are widely used for a wide range of medicinal and industrial aspects. At present, most studies related to the gene resources of Litsea are restricted to morphological analyses or features of individual genomes, and currently available studies of select molecular markers are insufficient. In this study, we assembled and annotated the complete chloroplast genomes of nine species in Litsea, carried out a series of comparative analyses, and reconstructed phylogenetic relationships within the genus. The genome length ranged from 152,051 to 152,747 bp and a total of 128 genes were identified. High consistency patterns of codon bias, repeats, divergent analysis, single nucleotide polymorphisms (SNP) and insertions and deletions (InDels) were discovered across the genus. Variations in gene length and the presence of the pseudogene ycf1Ψ, resulting from IR contraction and expansion, are reported. The hyper-variable gene rpl16 was identified for its exceptionally high Ka/Ks and Pi values, implying that those frequent mutations occurred as a result of positive selection. Phylogenetic relationships were recovered for the genus based on analyses of full chloroplast genomes and protein-coding genes. Overall, both genome sequences and potential molecular markers provided in this study enrich the available genomic resources for species of Litsea. Valuable genomic resources and divergent analysis are also provided for further research of the evolutionary patterns, molecular markers, and deeper phylogenetic relationships of Litsea.

Taxonomic notes are provided on species of the uncommonly encountered ripiphorid subfamily Pelecotominae. Zapotecotoma sumichrasti gen. et sp. nov., is described from southern Mexico based on a unique male likely collected in the later part of the mid-19th Century. The discovery of additional species of the South African genus Clinops Gerstaecker permit a revised diagnosis and distinction of the group from the eastern Mediterranean genus Scotoscopus Brenske and Reitter, resurrected status. Two new species of Clinops are established: Clinops inexpectatus sp. nov. (northeast of Durban near Swaziland) and C. perpessus sp. nov. (region of Durban), and Scotoscopus spectabilis (Schaufuss) is newly recorded for the Peloponnese in Greece.

A new genus of melitturgine bees (Panurginae: Melitturgini) is described and figured from central Saudi Arabia. Belliturgula najdica Engel, gen. et sp. n., is similar in several respects to the genus Flavomeliturgula Patiny in that both have a greatly elongate glossa that is longer than the face and is truncate apically. The former differs in the greatly elongate and flattened first labial palpomere, the remaining palpomeres unmodified, the greatly protuberant clypeus, and long labrum lacking dense setal patches, among other traits. In addition, remarks are made on the composition of Flavomeliturgula, with Meliturgula deserta Warncke (Iran, Pakistan), removed to Khuzimelissa Engel, gen. n., resulting in the new combination Khuzimelissa deserta (Warncke), comb. n. Khuzimelissa differs from Flavomeliturgula by the unmodified labial palpomeres (lacking the elongate first three labial palpomeres of the latter genus), the acute glossa (broadly truncate in the latter), glossa shorter than head length (greater than head length in the latter), the abundant, white, squamiform setae of the mesoscutum (sparse, erect setae in the latter), cleft pretarsal claws (simple in the latter), presence of metasomal setal bands (lacking in the latter), and apically emarginate pygidial plate (acutely rounded in the latter). The species of Flavomeliturgula are organized into three subgenera, including Koreshomelissa Engel, subgen. n., and Freyamelissa Engel, subgen. n., and these may eventually warrant generic status once phylogenetic work on the tribe has been completed. A key to the Arabian genera of Panurginae is provided, along with an augmented key to the genera of subtribe Meliturgulina.

Apis mellifera jemenitica is the indigenous race of honey bees in the Arabian Peninsula and is tolerant to local drought conditions. Experiments were undertaken to determine the differences in associative learning and memory of honey bee workers living in the arid zone of Saudi Arabia, utilizing the proboscis extension response (PER). These experiments were conducted on the indigenous race (A. m. jemenitica) along with two introduced European races (A. m. carnica and A. m. ligustica). The data revealed that A. m. jemenitica is amenable to PER conditioning and may be used in conditioning experiments within the olfactory behavioral paradigm. The results also demonstrated that the three races learn and retain information with different capacities relative to each other during the experimental time periods. Native Arabian bees (A. m. jemenitica) exhibited significantly lower PER percentage during second and third conditioning trials when compared to exotic races. Apis mellifera jemenitica also exhibited reduced memory retention at 2 h and 24 h when compared to A. m. carnica and A. m. ligustica. Therefore, the native Arabian bees were relatively slow learners with reduced memory retention compared to the other two races that showed similar learning and memory retention. Three or five conditioning trials and monthly weather conditions (October and December) had no significant effects on learning and memory in A. m. jemenitica. These results emphasized a novel line of research to explore the mechanism and differences in associative learning as well as other forms of learning throughout the year among bee races in the harsh arid conditions of Saudi Arabia. This is the first study in Saudi Arabia to demonstrate inter-race differences regarding olfactory associative learning between native Arabian bees and two introduced European honey bee races. Keywords: Associative learning, Memory formation, Olfaction, Proboscis extension response, Honey bee races, Saudi Arabia

As one of the largest families of beetles (Coleoptera), the Staphylinidae (rove beetles and their relatives) are rich not only in extant species but also in a comparatively robust fossil record. Despite this preponderance of available fossil material, fossils of the diverse subfamily Osoriinae remain rare. Here, we describe a new ososriine species, Priochirus trisclerite sp. nov., from the mid-Cretaceous amber of Myanmar. The new specimen is similar to the only other definitive fossil of the genus, Priochirus thayerae Yamamoto 2019, and both are placed in the extinct subgenus Eopriochirus subgen. nov. The new species differs noticeably in a number of morphological details in relation to the submentum, gular sutures and protibial crenulae. The new fossil provides further evidence for understanding the radiation of staphylinoid beetles.

Summary: Mimicry is ubiquitous in nature, yet understanding its origin and evolution is complicated by the scarcity of exceptional fossils that enable behavioral inferences about extinct animals. Here we report bizarre true bugs (Hemiptera) that closely resemble beetles (Coleoptera) from mid-Cretaceous amber. The unusual fossil bugs are described as Bersta vampirica gen. et sp. nov. and Bersta coleopteromorpha gen. et sp. nov. and are placed into a new family, Berstidae fam. nov. The specialized mouthparts of berstids indicate that they were predaceous on small arthropods. Their striking beetle-like appearance implies that they were either involved in defensive mimicry or mimicked beetles to attack unsuspecting prey. The latter would represent the first case of aggressive mimicry in the invertebrate fossil record. These findings enrich our understanding of the paleoecological associations and extinct behavioral strategies of Mesozoic insects.

Larvae of the Myrmeleontiformia, which include antlions, are not well preserved in much of the fossil record. Here, Badano et al. describe a collection of predatory myrmeleontiform larvae from Cretaceous amber, resolving their evolutionary relationships and inferring their ecology.

An unusual and well-preserved fossil staphylinid is described and figured from a single specimen in Upper Cretaceous Burmese amber. Gollandia planata gen. et sp. n. is tentatively placed in the extant oxyteline tribe Coprophilini, although it lacks a few characteristic features of present-day members of the group, likely indicating it to be either a stem group of the tribe or prove to be distinct pending future discoveries. The discovery of this genus suggests that early oxytelines were more morphologically diverse during the Cretaceous and their evolutionary history was more complicated than previously documented. Tribal placement as regards fossil oxyteline taxa is discussed.

While fossils of honey bees (Apini: Apis Linnaeus) are comparatively abundant in European Oligocene and Miocene deposits, the available material from Asia is scant and represented by only a handful of localities. It is therefore significant to report a new deposit with a fossil honey bee from southern China. Apis (Synapis) dalica Engel & Wappler, sp. n., is described and figured from Middle Miocene sediments of Maguan County, southeastern Yunnan Province, China. This is the first fossil bee from the Cenozoic of southern China, and is distinguished from its close congeners present at the slightly older locality of Shanwang, Shandong in northeastern China. The species can be distinguished on the basis of wing venation differences from other Miocene Apis.

A new braconid wasp from the Upper Cretaceous (Cenomanian) amber of the Hukawng Valley in Kachin State, Myanmar is described and figured from a unique female. Seneciobracon novalatus Engel & Huang, gen. et sp. n., is placed in a distinct subfamily, Seneciobraconinae Engel & Huang, subfam. n., owing to the presence of a unique combination of primitive protorhyssaline-like traits, with an otherwise more derived wing venation. The fossil is discussed in the context of other Cretaceous Braconidae.

A new species of the carpenter bee genus Xylocopa Latreille (Xylocopinae: Xylocopini) is described and figured from two localities in southern Saudi Arabia. Xylocopa (Koptortosoma) sarawatica Engel, sp. n. is a relatively small species similar to the widespread X. pubescens Spinola, but differs in the extent of maculation in males, setal coloration of both sexes, and male terminalia. A revised key to the species of Xylocopa in Saudi Arabia is provided.

The first gynandromorph of the insect order Zoraptera is reported. A gynandromorph of Zorotypus brasiliensis Silvestri is described from a likely parthenogenetic population in the Atlantic Forest around the border of the Brazilian states of Minas Gerais, Rio de Janeiro, and São Paulo. As has been previously surmised for this species, the sample consisted of only females, tending to support the hypothesis of thelytokous parthenogenesis for this population. The gynandromorph is largely female in outward appearance, but exhibits male secondary sexual traits on the left side of the apical abdominal segments. The left (male) antenna consists of antennomeres of different proportions to those of the right (female) antenna. The internal genitalia and associated sclerites, however, are female although some are augmented in their form relative to normal-type females. Comments are made on possible factors resulting in the development of the gynandromorph. This is the 16th insect order with gynandromorphism reported. Keywords: Angel insects, Brazil, Gynandromorphism, Soil-lice, Telythoky

A second species of protorhyssaline wasps (Braconidae) is described and figured from inclusions in Upper Cretaceous (Turonian) amber of the Raritan Formation in New Jersey, USA. Rhetinorhyssalites emersoni, gen. n., sp. n., is distinguished from other protorhyssalines, particularly the contemporaneous Protorhyssalus goldmani.

A new species of fossil bumble bee (Apinae: Bombini) is described and figured from Early Miocene (Burdigalian) deposits of the Most Basin at the Bílina Mine, Czech Republic. Bombus trophonius sp. n., is placed within the subgenus Cullumanobombus Vogt and distinguished from the several species groups therein. The species is apparently most similar to the Nearctic B. (Cullumanobombus) rufocinctus Cresson, the earliest-diverging species within the clade and the two may be related only by symplesiomorphies. The age of the fossil is in rough accordance with divergence estimations for Cullumanobombus.

Abstract 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.

A remarkable teratological female of Polistes (Fuscopolistes) dorsalis neotropicus Bequaert, 1940 (Vespidae: Polistinae) is described and illustrated. The specimen lacks all three external dorsal ocelli but is normally developed in almost every other aspect. Additionally, similar findings in other Hymenoptera are briefly discussed, as are the consequences and the reasons that might cause the random loss of ocelli.

The Late Palaeozoic insect superorder Palaeodictyopterida exhibits a remarkable disparity of larval ecomorphotypes, enabling these animals to occupy diverse ecological niches. The widely accepted hypothesis presumed that their immature stages only occupied terrestrial habitats, although authors more than a century ago hypothesized they had specializations for amphibious or even aquatic life histories. Here, we show that different species had a disparity of semiaquatic or aquatic specializations in larvae and even the supposed retention of abdominal tracheal gills by some adults. While a majority of mature larvae in Palaeodictyoptera lack unambiguous lateral tracheal gills, some recently discovered early instars had terminal appendages with prominent lateral lamellae like in living damselflies, allowing support in locomotion along with respiratory function. These results demonstrate that some species of Palaeodictyopterida had aquatic or semiaquatic larvae during at least a brief period of their post-embryonic development. The retention of functional gills or gill sockets by adults indicates their amphibious lifestyle and habitats tightly connected with a water environment as is analogously known for some modern Ephemeroptera or Plecoptera. Our study refutes an entirely terrestrial lifestyle for all representatives of the early diverging pterygote group of Palaeodictyopterida, a greatly varied and diverse lineage which probably encompassed many different biologies and life histories.

A remarkable teratological female of Megascolia (Regiscolia) maculata flavifrons (Fabricius, 1775) (Scoliidae: Scoliinae: Scoliini) with a supernumerary median ocellus is described and illustrated. While supernumerary ocelli have been reported before from Diptera, Orthoptera, and Hymenoptera, this is the first record of such a malformation from a scoliid wasp. Four other teratological scoliid wasps have been reported in the literature but all were gynandromorphs. A brief summary of known records of supernumerary ocelli among insects is provided.

A brief overview of those bee genera and subgenera occurring in the Kingdom of Saudi Arabia is provided. In total 22 genera are currently documented in the literature for the country, although a further 17 genera undoubtedly occur here based on extralimital records. The lists provided here will serve as a baseline for biodiversity studies on the bee fauna of the country that are presently underway. The anthidiine genus Icteranthidium Michener (Megachilidae) and the dasypodaine genus Promelitta Warncke (Melittidae) are newly recorded for the country.

Ten amber inclusions of male Ripidiinae (Coleoptera: Ripiphoridae) are reported from Early Miocene deposits of the Dominican Republic and compared with extant species of Neorrhipidius Viana, 1958 from Argentina and Paraguay and Quasipirhidius Zaragoza Caballero, 1991 from Mexico. Neorrhipidius seicherti sp. n. and Quasipirhidius luzziae sp. n. are described and illustrated. Both species are characterised by 11-segmented antennae with eight distal antennomeres uniflabellate, mouthparts reduced to maxillary palpi represented by long styli with fused basal palpomeres, by metathoracic wings without crossveins and tarsal formula of 5-5-4. Neorrhipidius seicherti sp. n. differs from Quasipirhidius luzziae sp. n. by the presence of postocular ommatidia and by the shape of metascutellum. The distribution of fossil and extant Ripidiinae in Central America is briefly discussed. Furthermore, the frequent occurrence of Quasipirhidius luzziae sp. n. in amber as syninclusions suggests possibly unique synchronisation of emergence unknown in extant taxa or more likely intraspecific aggregative behaviour by males prior to mating with receptive females.

A new species of inocelliid snake-fly, Fibla carpenteri, is described and figured frorn Middle to Upper Eocene Baltic amber. The species belongs to the nominate subgenus and is the largest amber snake-fly currently known. The aberrant fossil species Inocellia peculiaris is transferred to a new genus, EIectrinocellia, and placed in a new subfamily, the Electrinocelliinae, sister to the remainder of the Inocelliidae.