The earliest known holometabolous insects.

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

Predictability of Craniofacial Skeletal Age with Geometric Morphometries.

AIM: This study aims to estimate skeletal age of craniofacial shape obtained from cone beam computed tomography (CBCT)-defined facial and basicranial landmarks using geometric morphometrics method in a random sample of growing patients, and explore the correlation between craniofacial shape and skeletal age as determined from hand and wrist radiograph. MATERIALS AND METHODS: Generalized Procrustes analysis (GPA) of craniofacial shape with estimation of centroid size was performed on CBCTs of 48 growing patients (mean age 11.7 ± 1.5 years). Greulich and Pyle method for skeletal age assessment were used for correlation with centroid size. Correlation among the variables relied on Pearson's coefficient and its 95% confidence interval was estimated. The model's R2 was calculated, (Cook's distances, Mahalanobis distances, leverage values, and studentized residuals) and multiple regression analysis performed using the Statistical Package for the Social Sciences (SPSS) version 22. RESULTS: Mean skeletal age was 11.9 ± 2.4 years. Centroid size (151.5 ± 7.2) was significantly correlated with chronological age (R = 0.616, 95% CI 0.355-0.789, p < 0.01) and skeletal age (R = 0.605, 95 % CI 0.331-0.794, p < 0.01). CONCLUSION: A new equation for determining craniofacial skeletal age was developed, using the centroid size of the craniofacial frame, gender, and the known chronological age. CLINICAL SIGNIFICANCE: A CBCT may be used for skeletal age assessment without additional hand wrist radiograph.

The first Mesozoic microwhip scorpion (Palpigradi): a new genus and species in mid-Cretaceous amber from Myanmar.

A fossil palpigrade is described and figured from mid-Cretaceous (Cenomanian) amber from northern Myanmar. Electrokoenenia yaksha Engel and Huang, gen. n. et sp. n., is the first Mesozoic fossil of its order and the only one known as an inclusion in amber, the only other fossil being a series of individuals encased in Pliocene onyx marble and 94-97 million years younger than E. yaksha. The genus is distinguished from other members of the order but is remarkably consistent in observable morphological details when compared to extant relatives, likely reflecting a consistent microhabitat and biological preferences over the last 100 million years.

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.

Preliminary inventory of terrestrial Heteroptera from Lebanon (Hemiptera: Leptopodomorpha, Cimicomorpha and Pentatomomorpha).

This paper gives, for the first time, a faunistic list of terrestrial Heteroptera (Leptopodomorpha, Cimicomorpha and Pentatomomorpha) known to occur in Lebanon. The list is based on examination and identification by the first author of specimens preserved in various collections and on reliable published records. A total of 231 species and subspecies belonging to 161 genera and 26 families are recorded for the country. Among these species, 71 belonging to 60 genera and 11 families are not cited in the Catalogue of the Heteroptera of the Palaearctic Region (Aukema & Rieger, 1995-2006; Aukema et al., 2013; Aukema 2018-2022), as being present in Lebanon, and thus may be considered as new for the country. A list of 18 species of terrestrial Heteroptera housed at the Lebanese University Collection is also given but these identifications need to be verified.

The first Early Cretaceous representative of the fly family Tipulidae from the lower Barremian dysodiles of Lebanon (Diptera).

Leptotarsus reyi sp. nov., first representative of the family Tipulidae from the Lower Cretaceous of Lebanon, is characterised, described and illustrated from the Early Barremian dysodile of Jdeidet Bkassine, South Lebanon.

Palaeoparasycorax globosus Stebner et al., 2015 from the mid-Cretaceous amber of Myanmar: Description of a mating pair.

A mating pair of Palaeoparasycorax globosus Stebner et al., 2015 from the mid-Cretaceous Burmese amber is illustrated and described. The male is redescribed, and the female genitalia of this species is described for the first time. This discovery allows for a better understanding of this fossil sycoracines characteristics and further demonstrates the bioinclusion richness of the mid-Cretaceous Burmese amber. A list of all Cretaceous Psychodidae is given.

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.

First record of fossil psocodeans in copula from mid-Cretaceous Burmese amber.

We report the first discovery of barklice preserved in copula from amber dating back to the mid-Cretaceous, thus documenting the oldest preserved reproductive behaviour of Psocodea. The new finding provides new insight into the evolution of copulatory behaviour in Trogiomorpha. Moreover, we describe and illustrate the new fossil material of Burmempheria densuschaetae (= Latempheria kachinensis) in the extinct family Empheriidae and Longiantennum fashengi in the extinct family Archaeatropidae, two diverse and widely distributed groups during the Cretaceous. No significant sexual dimorphism is recorded, apart from variation in size where males are larger. The taxonomy of Cretaceous empheriids from Burmese amber is revised and discussed.

A transitional fossil mite (Astigmata: Levantoglyphidae fam. n.) from the early Cretaceous suggests gradual evolution of phoresy-related metamorphosis.

Metamorphosis is a key innovation allowing the same species to inhabit different environments and accomplish different functions, leading to evolutionary success in many animal groups. Astigmata is a megadiverse lineage of mites that expanded into a great number of habitats via associations with invertebrate and vertebrate hosts (human associates include stored food mites, house dust mites, and scabies). The evolutionary success of Astigmata is linked to phoresy-related metamorphosis, namely the origin of the heteromorphic deutonymph, which is highly specialized for phoresy (dispersal on hosts). The origin of this instar is enigmatic since it is morphologically divergent and no intermediate forms are known. Here we describe the heteromorphic deutonymph of Levantoglyphus sidorchukae n. gen. and sp. (Levantoglyphidae fam. n.) from early Cretaceous amber of Lebanon (129 Ma), which displays a transitional morphology. It is similar to extant phoretic deutonymphs in its modifications for phoresy but has the masticatory system and other parts of the gnathosoma well-developed. These aspects point to a gradual evolution of the astigmatid heteromorphic morphology and metamorphosis. The presence of well-developed presumably host-seeking sensory elements on the gnathosoma suggests that the deutonymph was not feeding either during phoretic or pre- or postphoretic periods.

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.

Mammalian hairs in Early Cretaceous amber.

Two mammalian hairs have been found in association with an empty puparium in a approximately 100-million-year-old amber (Early Cretaceous) from France. Although hair is known to be an ancestral, ubiquitous feature in the crown Mammalia, the structure of Mesozoic hair has never been described. In contrast to fur and hair of some Jurassic and Cretaceous mammals preserved as carbonized filaments, the exceptional preservation of the fossils described here allows for the study of the cuticular structure. Results show the oldest direct evidence of hair with a modern scale pattern. This discovery implies that the morphology of hair cuticula may have remained unchanged throughout most of mammalian evolution. The association of these hairs with a possible fly puparium provides paleoecological information and indicates peculiar taphonomic conditions.

First Report of the Ticks Haemaphysalis punctata Canestrini et Fanzago, 1878, Haemaphysalis parva (Neumann, 1897) and Dermacentor marginatus (Sulzer, 1776) (Acari, Amblyommidae) from Humans in Lebanon.

PURPOSE: Knowledge on ticks infesting humans is scarce for the middle East. In this work, tick specimens (Acari: Amblyommidae) infesting humans in Lebanon were identified. METHODS: Ticks that were found on humans were received in the Lebanese University, Faculty of Sciences. The specimens were preserved in alcohol for their further morphological identification. RESULTS: Three tick species were identified: a red sheep tick Haemaphysalis punctata Canestrini et Fanzago, 1878, a Mediterranean ear tick H. parva (Neumann, 1897), and an ornate sheep tick Dermacentor marginatus (Sulzer, 1776); all isolated from human hosts. CONCLUSION: This is the first report of Haemaphysalis punctata, H. parva and Dermacentor marginatus infesting humans from Central and North Lebanon.

A new subfamily of Trichoceridae (Diptera: Tipulomorpha) from Early Cretaceous Lebanese amber.

The first find of Trichoceridae in the Mesozoic of Gondwana and in Cretaceous ambers worldwide, Ewaurista pusilla gen. et sp. nov. is separated into Ewauristinae subfam. nov. demonstrating some venational traits in common with Limoniidae and a peculiar sexual dimorphism. The diagnosis of Trichoceridae is emended, and relationships of the family are discussed.

First levantine fossil murines shed new light on the earliest intercontinental dispersal of mice.

Recent extensive field prospecting conducted in the Upper Miocene of Lebanon resulted in the discovery of several new fossiliferous localities. One of these, situated in the Zahleh area (Bekaa Valley, central Lebanon) has yielded a particularly diverse vertebrate fauna. Micromammals constitute an important part of this assemblage because not only do they represent the first Neogene rodents and insectivores from Lebanon, but they are also the only ones from the early Late Miocene of the Arabian Peninsula and circumambient areas. Analyses of the murines from Zahleh reveal that they belong to a small-sized early Progonomys, which cannot be assigned to any of the species of the genus hitherto described. They are, thereby, shown to represent a new species: Progonomys manolo. Morphometric analyses of the outline of the first upper molars of this species suggest a generalist and omnivorous diet. This record sheds new light onto a major phenomenon in the evolutionary history of rodents, which is the earliest dispersal of mice. It suggests that the arrival of murines in Africa got under way through the Levant rather than via southern Europe and was monitored by the ecological requirements of Progonomys.

Forensically Relevant Blow Flies in Lebanon Survey and Identification Using Molecular Markers (Diptera: Calliphoridae).

Calliphoridae are among the first insects associated to decomposing animal remains. We have collected 1,841 specimens of three calliphorid genera: Calliphora, Lucilia, and Chrysomya, from different Lebanese localities as a first step in implementing a database of insects of forensic relevance for the country. Blow-flies are crucial for the estimation of the postmortem interval. DNA-based identification is a rapid and accurate method, often used for morphologically similar species, especially for immatures or incomplete specimens. In this study, we test the suitability of three genetic markers to identify adults and immature stages of calliphorids, viz., mitochondrial cytochrome c oxidase subunit I (COI) barcode, a region including partial sequences of mitochondrial Cyt-b-tRNAser-ND1, and second internal transcribed spacer (ITS2) region of nuclear ribosomal DNA. Forty Lebanese specimens of various developmental stages (egg, larva, wandering third instar, pupa, newly emerged adult, and mature adult) were identified among the three calliphorid genera: Calliphora, Lucilia, and Chrysomya, and compared with published sequences to confirm their specific assignation. Phylogenetic analyses showed the robustness of ITS2 and COI to identify calliphorids at species level. Nevertheless, ITS2 failed to discriminate Lucilia caesar (Linnaeus) (Diptera, Calliphoridae) from Lucilia illustris (Meigen) (Diptera, Calliphoridae), and COI had a similar issue with Lucilia sericata (Meigen) (Diptera, Calliphoridae) and Lucilia cuprina (Wiedemann) (Diptera, Calliphoridae). Thus, these two markers are complementary. This work contributes new nucleotide sequences for Lebanon. It is a first step in implementing a molecular database of forensic relevant insects for the country.

A soil-carrying lacewing larva in Early Cretaceous Lebanese amber.

Diverse organisms protect and camouflage themselves using varied materials from their environment. This adaptation and associated behaviours (debris-carrying) are well known in modern green lacewing larvae (Neuroptera: Chrysopidae), mostly due to the widespread use of these immature insects in pest control. However, the evolutionary history of this successful strategy and related morphological adaptations in the lineage are still far from being understood. Here we describe a novel green lacewing larva, Tyruschrysa melqart gen. et sp. nov., from Early Cretaceous Lebanese amber, carrying a preserved debris packet composed by soil particles entangled among specialised setae of extremely elongate tubular tubercles. The new morphotype has features related to the debris-carrying habit that are unknown from extant or extinct green lacewings, namely a high number of tubular tubercle pairs on the abdomen and tubular tubercle setae with mushroom-shaped endings that acted as anchoring points for debris. The current finding expands the diversity of exogenous materials used by green lacewing larvae in deep time, and represents the earliest direct evidence of debris-carrying in the lineage described to date. The debris-carrying larval habit likely played a significant role during the initial phases of diversification of green lacewings.

Contribution to the knowledge of Saprinus Erichson, 1834 of forensic relevance from Lebanon (Coleoptera, Histeridae).

Many histerid beetles are necrophilous on carrion during both active and advanced stages of decomposition. In this study, 13 species of Saprinus were recorded on carrion from Lebanon, containing eight that are new for the Lebanese fauna. The following Saprinus species are newly recorded from Lebanon: 1) Saprinus (S.) caerulescens caerulescens (Hoffmann, 1803); 2) S. (S.) calatravensis Fuente, 1899; 3) S. (S.) chalcites (Illiger, 1807); 4) S. (S.) godet (Brullé, 1832); 5) S. (S.) maculatus (P. Rossi, 1792); 6) S. (S.) strigil Marseul, 1855; 7) S. (S.) submarginatus J. Sahlberg, 1913; and 8) S. (S.) tenuistrius sparsutus Solsky, 1876. The peak activity was recorded, key for the species is provided, and habitus images and male genitalia are illustrated in order to facilitate their taxonomic identifications. Saprinus species are diverse and common on animal carcass; they were likewise collected from a human cadaver in Lebanon. Preliminary comments on biology and distribution of the studied species are given. Our paper represents the first faunistic study on Histeridae from Lebanon. A rigorous research program regarding the biology of Saprinus in Lebanon and the neighbouring countries would greatly improve the knowledge of the diversity, activity, and possible forensic value of Saprinus.

Dental and Skeletal Age Estimations in Lebanese Children: A Retrospective Cross-sectional Study.

AIMS AND OBJECTIVES: Assessing dental and bone ages is frequently required in a wide range of fields such as odontology, forensic science, as well as orthopedics. The aim of this study was to evaluate applicability of two methods of bone age assessment and two methods of dental age (DA) assessment for Lebanese children. MATERIALS AND METHODS: Skeletal age (SA) of 260 orthodontic patients (124 males, 136 females divided into four groups each) was consecutively assessed using Greulich and Pyle and Fishman's SMI methods. DA was evaluated using both Demirjian's and Willem's methods. Mean age was 11.89 ± 1.38 years for males and 11.75 ± 1.58 years for females. Data were collected and statistically analyzed using the SPSS software (IBM SPSS Statistics, version 21, USA). The differences between estimated DA, estimated SA, and chronological age (CA) were compared by gender and age group. RESULTS: Greulich and Pyle method showed nonsignificant difference with CA in male sample, while in both assessment methods, the difference between skeletal and CAs is significant in female sample. Results of Willem's method in the whole sample suggested a statistically nonsignificant difference, when compared to CA. Demirjian's method delivered higher mean value than Willem's assessment in both genders. CONCLUSIONS: Greulich and Pyle method is accurate for SA assessment in males and only in one group of females, while it significantly overestimates age in all other female groups. Willem's method is more suitable to assess DA in both genders. A strong correlation exists between both dental and skeletal assessment methods and CA.