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Extant terrestrial vertebrates, including birds, have a panoply of symbiotic relationships with many insects and arachnids, such as parasitism or mutualism. Yet, identifying arthropod¿vertebrate symbioses in the fossil record has been based largely on indirect evidence; findings of direct association between arthropod guests and dinosaur host remains are exceedingly scarce. Here, we present direct and indirect evidence demonstrating that beetle larvae fed on feathers from an undetermined theropod host (avian or nonavian) 105 million y ago. An exceptional amber assemblage is reported of larval molts (exuviae) intimately associated with plumulaceous feather and other remains, as well as three additional amber pieces preserving isolated conspecific exuviae. Samples were found in the roughly coeval Spanish amber deposits of El Soplao, San Just, and Peñacerrada I. Integration of the morphological, systematic, and taphonomic data shows that the beetle larval exuviae, belonging to three developmental stages, are most consistent with skin/hide beetles (family Dermestidae), an ecologically important group with extant keratophagous species that commonly inhabit bird and mammal nests. These findings show that a symbiotic relationship involving keratophagy comparable to that of beetles and birds in current ecosystems existed between their Early Cretaceous relatives., This work was supported by the project CRE, funded by the Spanish Agencia Estatal de Investigación (AEI)/European Regional Development Fund (FEDER)/European Union (EU) Grant CGL2017-84419, the project PGC2018-094034-B-C22 (Ministerio de Ciencia, Innovación y Universidades/AEI/FEDER, UE), the project CGL2014-52163, funded by the Spanish Ministry of Economy, Industry, and Competitiveness, and by the Consejería de Industria, Turismo, Innovación, Transporte y Comercio of the Gobierno de Cantabria through the public enterprise EL SOPLAO S.L. (research contract REF VAPC 20225428 to Centro Nacional Instituto Geológico y Minero de España-Consejo Superior de Investigaciones Científicas and research agreement #20963 with University of Barcelona, both for the period 2022 to 2025). S.Á.-P. is funded by a grant from the Secretary of Universities and Research of the Government of Catalonia and European Social Fund (2021FI_B2 00003), Results Amber Samples Studied Description of the Larval Exuviae Discussion Affinities of the Larvae Taphonomy Keratophagy Concluding Remarks Materials and Methods Amber Pieces Imaging Data, Materials, and Software Availability Acknowledgments Supporting Information References

Herhold, Hollister W, Davis, Steven R, Degrey, Samuel P, Grimaldi, David A (2023): COMPARATIVE ANATOMY OF THE INSECT TRACHEAL SYSTEM PART 1: INTRODUCTION, APTERYGOTES, PALEOPTERA, POLYNEOPTERA. Bulletin of the American Museum of Natural History 459 (1): 1-184, DOI: 10.5531/sd.sp.55, URL: http://dx.doi.org/10.5531/sd.sp.55

Diplurans are among the earliest hexapods in the geological record. These primitively wingless relatives of insects are infrequently encountered despite being pervasive in soil habitats. Two groups have disparate adaptations for hunting, one mechanical and the other chemical, in Japygoidea and Projapygoidea, respectively. Here, we report three genera (two new) and four species of fossil Projapygidae preserved in Mesozoic and Cenozoic ambers: Electroprojapyx alchemicus gen. et sp. nov. in mid-Cretaceous Myanmar amber, and Symphylurinopsis punctatus gen. et sp. nov. and two species (unnamed) of Symphylurinus in Miocene Dominican amber. The exceptionally preserved specimens possess cerci that are morphologically specialized for expelling a glandular substance on prey. The new Cretaceous E. alchemicus is a stem group to all living species of the family and provides conclusive Mesozoic evidence for a rare type of predation involving offensive (vs. defensive) secretions. The specimen was fossilized in a preying-and-spraying position with its presumptive meal, a Symphypleona springtail (Collembola), reflecting the behavioural predatory repertoire of modern relatives. Apart from one Cenozoic species, these rare specimens represent the only fossil record of the family Projapygidae. Our findings demonstrate that these basal hexapods were predators of small arthropods of ancient forest soils and enrich our understanding of palaeoecological associations and behavioural strategies paramount for the survival of species.

Jones, Lance E., Grimaldi, David A. (2022): Revision Of The Nearctic Species Of The Genus Amiota Loew (Diptera: Drosophilidae). Bulletin of the American Museum of Natural History 2022 (458): 1-181, DOI: 10.1206/0003-0090.458.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-458/issue-1/0003-0090.458.1.1/Revision-of-the-Nearctic-Species-of-the-Genus-Amiota-Loew/10.1206/0003-0090.458.1.1.full

Dead wood is a microcosm of biological diversity. Saproxylic insect diversity at the global level is immense, with estimates that up to one third of forest dwelling species rely on moribund or dead...

PREMISE Early Eocene ambers of the Cambay lignite in Gujarat, India, are well known for their diverse insect fauna and dispersed pollen, but the included flowers have received limited attention. The fossil record of Apocynaceae is relatively poor, and the distinctive floral characters of this family have not been recognized in the fossil record before. METHODS Remains of tiny flowers in amber were studied by micro-CT scanning, reflected light, and epifluorescence microscopy. RESULTS Flowers of Maryendressantha succinifera gen. et. sp. n. have actinomorphic, pentamerous, tubular corollas 2.2-2.3 mm wide, and 1.7-2.1 mm deep with sinistrorse aestivation and androecia consisting of a whorl of five stamens attached by short filaments to the lower half of the corolla tube. Anthers are ovate, rounded basally and apically tapered with their connectives convergent with one another in a conical configuration. The pollen is globose, psilate, tricolporate, and very small (10-11 µm). The combined characters indicate a position within the grade known as subfamily Rauvolfioideae. CONCLUSIONS These fossils, as the oldest remains of Rauvolfioids, complement the fossil records of Apocynoid and Asclepioid fossil seeds from other regions, demonstrating that the Apocynaceae were well established by the early Eocene, mostly consistent with prior divergence estimates for the phylogeny of this family. Potential pollinators, also preserved in the Cambay amber, include mosquitos, gnats, small moths, and stingless bees.

The first fossil Archaeidae in Cambay amber from India, of Eocene age, is documented. The inclusion is a spider exuvium and is placed as Myrmecarchaea based on the presence of elongated legs, a slightly elongated pedicel with lateral spurs, and a diastema between coxae III and IV that is similar to M. antecessor from Oise amber. The previous occurrences of the genus are from Baltic and Oise amber, both of Eocene age. Because most spiders, including Archaeidae, only molt as juveniles, the exuvium does not have adult features nor have distinct species-specific features, and a new taxon is not erected. This new record further extends the distribution of the family and genus to India 50–52 million years ago. Myrmecarchaea in Indian Cambay amber provides additional evidence that India in the Early Eocene had affinities with the Palearctic mainland rather than showing Gondwanan insularity.

The study of terrestrial arthopod fossils preserved with microscopic fidelity in amber and as permineralized replicas has been revolutionized by CT scanning. Fine preservation facilitates phylogenetic interpretation of fossils, but molecular divergence-time models still commonly use insufficient fossil calibrations, skewing estimates away from the direct (i.e. fossil, morphological) evidence. Interactions among terrestrial arthropods (predation, parasitoidism; phoresy, social symbionts) are briefly reviewed from the fossil record. Predation is the oldest and most widespread, originating with arachnids since probably the Silurian. The first phoretic arthropods were probably mites (Acari). Parasitoidism extends to the early Jurassic ~200 mya, with four main episodes proposed by [1•]. 100-myo Burmese amber, the most diverse Cretaceous paleobiota, is unique for our understanding of insect eusociality and interrelationships among terrestrial arthropods. Eusocial insect colonies are ecological sinks for thousands of symbiont species; ages of the major eusocial groups and some of their nest symbionts are discussed. Fossilized arthropod interrelationships in Miocene Dominican amber are presented as visual exemplars.

1. Dracula orchids mimic mushrooms in appearance and scent, and are pollinated by mushroom flies. This study examined the guilds of insect visitors to Dracula species and to co‐occurring mushrooms in an Ecuadorean cloud forest. Significant visitor overlap was documented (mostly in the genera Zygothrica and Hirtodrosophila; Drosophilidae). To test the brood‐site mimicry hypothesis, behavioural observations were also performed and rearing success was examined. 2. Many fly species that visit Dracula are also found on mushrooms. Furthermore, the presence of pollinia enabled us to document the fact that particular individuals visiting mushrooms also visit Dracula. Roughly two‐thirds of the visitors to these unusual flowers are fungal‐associated, and the rest appear to be Dracula specialists. 3. A variety of behaviours common on both host groups were observed, including courtship semaphoring, feeding, sheltering, defending territory, and mating. Given this suite of shared reproductive behaviour, it was hypothesised that flies may also oviposit in both substrates. Flies were reared from mushrooms, Dracula spp., and other flowers (controls) to determine which substrate led to the highest fitness for the flies. Very few flies emerged from the Dracula flowers and there was no species overlap between the emergent insects and the known pollinators. 4. Despite the fact that there is evidence for brood‐site mimicry, the Dracula situation is complex. All the flies could derive many of the same fitness benefits from the flowers as they do from mushrooms (shelter, food, mating), but they are clearly not gaining the same fitness benefit, in terms of progeny, on flowers compared with mushrooms.

Two African drosophilids have distinctive protarsal spines in both sexes: Drosophila spinipes Lamb, 1914 from the Seychelles and D. suma Burla, 1954 from the Ivory Coast. Both were later classified in the genus Hirtodrosophila Duda of the Zygothrica genus group, and suma synonymized under spinipes by Tsacas (2006). We resurrect suma from synonymy, define the spinipes species group, and add seven new species in the group from Africa and Madagascar: Drosophila cameroonensis n. sp., D. freidbergi n. sp., D. hypandrilata n. sp., D. jambiya n. sp., D. malagasy n. sp., D. nigrospinipes n. sp., and D. phalloserra n. sp. Another four species are reported but not described. Detailed morphology informs that these anthophilous flies do not belong to the Zygothrica genus group, and they are returned to the genus Drosophila (unplaced to subgenus), albeit primitive members with some very distinctive specialized features. All distribution records, published and new, are presented along with a key to all nine named species.

Diverse new basal (aschizan) Cyclorrhapha fossilized in amber are described from the Tertiary and Cretaceous, and their relationships are examined with character-based phylogenetic hypotheses for each family or family group. There are 18 new species in 15 genera (11 of them new) and four families plus the Syrphoidea. Fossils are from the Early Cretaceous of Lebanon, Late Cretaceous of New Jersey (United States) and Alberta (Canada), Eocene of the eastern Baltic coast, and Miocene of the Dominican Republic, but predominantly from the mid-Cretaceous of Myanmar. Stem-group Lonchopteroidea are Alonchoptera lebanica, n. gen., n.sp., and Lonchopterites burmensis, n. sp. Platypezidae include the stem groups Burmapeza radicis, n. gen., n. sp., Canadopeza biacrosticha, n. gen., n. sp., and Calvopeza divergens, n. gen., n. sp. An unnamed Microsania sp. is the first definitive Platypezidae in Baltic amber; Lebanopeza azari, n. gen., n. sp., is a stem group to the Microsaniinae and Melanderomyiinae. Chandleromyia anomala, n. gen., n. sp., is an anomalously derived Platypezinae from the Cretaceous, and two new species of the diverse Recent genus Lindneromyia are in Dominican amber (L. neomedialis and L. dominicana). Fossils of the relict family Ironomyiidae (with 3 living species from eastern Australia) include two stem-group genera with two new species each, all in Burmese amber: Palaeopetia dorsalis and P. terminus, Proironia (n. gen.) gibbera and P. burmitica. All other species of Palaeopetia are compression fossils from the Cretaceous of Asia and Eurasia. For Phoridae, a new defining feature is a stridulatum on the procoxa and profemur in both sexes, occuring in most fossil taxa where observable. New sciadocerines include Eosciadocera pauciseta, n. sp., a very large species in Baltic amber, and two stem groups in Burmese amber, Prophora dimorion, n. gen., n. sp., and a very small, undescribed taxon. Archiphora pria Grimaldi and Cumming in Turonian-aged New Jersey amber is transferred to Hennigophora Brown, based on evidence from a new specimen. Prioriphorinae (not taxonomically treated here) is a paraphyletic, Cretaceous grade to the very diverse, crown-group radiation of Euphorida that occurred in the Cenozoic. Two syrphoids occur in Burmese amber: Prosyrphus thompsoni, n. gen., n. sp. (an apparent stem group to the Syrphidae), and Aschizomyia burmensis, n. gen., n. sp. (with more ambiguous affinities). Several immatures of undetermined family are reported, one a probable phorid larva. No definitive Schizophora are yet known from the Cretaceous.

The genus Litoleptis has consisted of nine described species, seven of them Asian and only two in the New World: L. alaskensis Chillcott, known from two specimens from northwest Alaska, and L. chilensis Hennig, known from a male specimen from near Santiago, Chile. A third New World species is described here, Litoleptis tico, n. sp., based on a single female from Costa Rica. The species is unique for the genus in having a vestigial proboscis and lacking spermathecal accessory ducts and glands. Female terminalia are unknown for the other two New World species. A morphologically based, preliminary phylogeny of spaniines is provided, indicating Litoleptis is recently derived among spaniines and thus Rhagionidae; the Early Cretaceous Litoleptis fossilis is a stem group to the living species. A derived position of the genus, its apparently broad distribution, and an abundance in Japan where Litoleptis has been bred from liverworts (Imada and Kato, 2016a), all indicate that these flies are probably not at all relict, simply vastly undersampled because of a reliance on mass-collecting techniques.

The Chickaloon Formation in south-central Alaska contains rich coal deposits dated very close to the Paleocene-Eocene boundary, immediately beneath which occur dispersed nodules of amber along with abundant remains of Metasequoia, dicots, and monocots. The nodules are small (less than 10 mm in length), nearly 10,000 of which were screened, yielding several inclusions of fungi and plant fragments, but mostly terrestrial arthropods: 29 specimens in 10 orders and 13 families. The fungi include resinicolous hyphae and a dark, multiseptate hyphomycete. Plants include wood/bark fragments and fibers, and the microphylls of a bryophyte (probably a moss, Musci). Among the arthropods are arachnids: mites (Acari: Oribatida), Pseudoscorpionida, and the bodies and a silken cocoon of spiders (Araneae). Insecta include Blattodea, Thysanoptera, Hemiptera (Heteroptera and Aphidoidea), Coleoptera (Dermestidae: Megatominae), Trichoptera, Diptera (Chironomidae: Tanypodinae), and Hymenoptera (Formicidae: Formicinae). Nymphal aphids predominate (65% of the arthropod individuals), which were probably feeding on the source tree, likely Metasequoia. There is a bias in preservation toward small arthropods (mean body length 0.75 mm) that are surface-dwelling (nonwinged) stages and taxa. Chickaloon amber contains the most northerly fossil records of pseudoscorpions, thrips, Dermestidae, and Cenozoic ants and mites, so the deposit is contributing unique data on high-latitude paleodiversity of the Paleogene hothouse earth.

Species concepts are morphologically revised and updated for members of the mycophagous genus Hirtodrosophila Duda that occur in America north of Mexico. Photomicrographs of external features, illustrations of male and female terminalia, and detailed descriptions are provided for 12 species. One species, H. cinerea (Patterson and Wheeler) is known only from the original description; its status is uncertain. Species exclusively from the southwestern United States are H. grisea (Patterson and Wheeler), H. longala (Patterson and Wheeler), and H. orbospiracula (Patterson and Wheeler). Hirtodrosophila alabamensis (Sturtevant) and H. duncani (Sturtevant) are widespread throughout the eastern half of North America; the latter species is morphologically disparate for Hirtodrosophila but provisionally retained in the genus. Hirtodrosophila chagrinensis (Stalker and Spencer) is very rare, known only from two female specimens from the northern United States. Hirtodrosophila ordinaria (Coquillett) is the most widespread species of the genus in North America, occurring throughout the northern half of the continent up to Alaska; H. shaitanensis (Sidorenko) from far eastern Russia may be a junior synonym. A preliminary scheme of relationships in the H. melanderi species group (including H. ordinaria) is presented. Two species from Florida (H. pictiventris [Duda], H. prognatha [Sturtevant]) and one from Florida plus other Gulf states (H. thoracis [Williston]) are widespread throughout the Caribbean, Central America, and portions of South America. Hirtodrosophila mendeli (Mourao et al.), from Brazil, may be a junior synonym of H. prognatha. Two new species are described from southern Florida: H. florida, n. sp., and H. jaenikei n. sp., the latter in the nigrohalterata species complex. A key to the North American species is provided.

Three wasp (Hymenoptera: Vespidae) fossils in Cretaceous amber (Late Albian) of northern Myanmar are described. Two are new species of the Mesozoic genus Curiosivespa (Rasnitsyn): C. zigrasi sp.n. and C. striata sp.n. The third species, Protovespa haxairei gen.n. et sp.n., has a combination of features unique among Mesozoic Priorvespinae and the extant subfamilies. These well preserved fossils provide new morphological data for a cladistic analysis of the basal lineages of Vespidae. Results suggest that Euparagiinae is the sister group of all other Vespidae. The new genus Protovespa appears more closely related to extant Masarinae, Eumeninae and social wasps than to Priorvespinae. We assign it to a new subfamily: Protovespinae. Finally, fossil information combined with a phylogenetic tree shows that the main groups of Vespidae probably evolved during the Early Cretaceous. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1E7E4796-6E70-4D81-BB34-0FEEA765DC25.

Angiosperms and their insect pollinators form a foundational symbiosis, evidence for which from the Cretaceous is mostly indirect, based on fossils of insect taxa that today are anthophilous, and of fossil insects and flowers that have apparent anthophilous and entomophilous specializations, respectively. We present exceptional direct evidence preserved in mid-Cretaceous Burmese amber, 100 mya, for feeding on pollen in the eudicot genus Tricolporoidites by a basal new aculeate wasp, Prosphex anthophilos, gen. et sp. nov., in the lineage that contains the ants, bees, and other stinging wasps. Plume of hundreds of pollen grains wafts from its mouth and an apparent pollen mass was detected by micro-CT in the buccal cavity: clear evidence that the wasp was foraging on the pollen. Eudicots today comprise nearly three-quarters of all angiosperm species. Prosphex feeding on Tricolporoidites supports the hypothesis that relatively small, generalized insect anthophiles were important pollinators of early angiosperms., Grimaldi et al. report on a primitive stinging wasp preserved in mid-Cretaceous amber from Myanmar, encapsulated just after it fed on the pollen of a eudicot. Micro-CT shows a pollen mass in the wasp’s mouth cavity, supporting the idea that insect anthophiles were important pollinators of early angiosperms.

The great evolutionary success of angiosperms has traditionally been explained, in part, by the partnership of these plants with insect pollinators [1, 2, 3, 4, 5, 6]. The main approach to understanding the origins of this pervasive relationship has been study of the pollinators of living cycads, gnetaleans, and basal angiosperms [7]. Among the most morphologically specialized living pollinators are diverse, long-proboscid flies. Early such flies include the brachyceran family Zhangsolvidae, previously known only as compression fossils from the Early Cretaceous of China and Brazil. It belongs to the infraorder Stratiomyomorpha, a group that includes the flower-visiting families Xylomyidae and Stratiomyidae. New zhangsolvid specimens in amber from Spain (ca. 105 mega-annum [Ma]) and Myanmar (100 Ma) reveal a detailed proboscis structure adapted to nectivory. Pollen clumped on a specimen from Spain is Exesipollenites, attributed to a Mesozoic gymnosperm, most likely the Bennettitales. Late Mesozoic scorpionflies with a long proboscis have been proposed as specialized pollinators of various extinct gymnosperms, but pollen has never been observed on or in their bodies [8]. The new discovery is a very rare co-occurrence of pollen with its insect vector and provides substantiating evidence that other long-proboscid Mesozoic insects were gymnosperm pollinators. Evidence is thus now gathering that visitors and probable pollinators of early anthophytes, or seed plants, involved some insects with highly specialized morphological adaptations, which has consequences for interpreting the reproductive modes of Mesozoic gymnosperms and the significance of insect pollination in angiosperm success.

For slightly more than a century only the type specimen has been known for Pyrgometopa penicillata, a monotypic genus of Drosophilidae whose sole species has an ocellar triangle distended into a bizarre tubercle bearing a tuft of large, spine-like bristles in both sexes. Here we report additional specimens of both sexes of the species, recently rediscovered from Brazil, Peru and French Guyana, allowing us to state that Pyrgometopa is a junior synonym for Stegana, a large cosmopolitan genus. A detailed redescription of Stegana penicillata (Kertész) new combination is provided, including new morphological information, like the male and female terminalia, the egg and a short discussion is provided about its hallmark feature.

Two closely related and distinctive species of Schistostoma Becker, i.e. Schistostoma burmanicum sp. nov. and Schistostoma foliatum sp. nov., are described from two male specimens preserved in mid-Cretaceous Burmese amber and recorded as the first known fossil representatives of this extant genus of Microphorinae. Both species possess greatly enlarged lamellate fore tibiae and modified leg chaetotaxy that are presumably male secondary sexual features. The generic limits of Schistostoma are briefly discussed and a rationale for the inclusion of these new species is provided.

All 16 species of the family Eremochaetidae occur from the Late Jurassic to the mid-Cretaceous of eastern and Central Asia. The first species in amber, and the latest occurrence of the family, was recently described as Zhenia xiai, from the mid-Cretaceous of Myanmar, ca. 100 Ma. New observations of a finely preserved specimen allow refinement of the morphological interpretations in the original description. The female of Zhenia, for example, has the distinctive piercing oviscapt of the superfamily Archisargoidea, formed from modified cerci (not tergites 8 and 9 as originally reported). The pretarsus of Zhenia bears an enormous empodial pad and pair of pulvillae, but the claws are highly vestigial (contra Zhang et al., 2016). The fly was almost certainly a parasitoid. A cladistic analysis of 26 binary-state characters and six continuously variable characters, using 47 exemplar Archisargoidea species from most genera and all four families, and five outgroup Brachycera, has very poor support for mos...

A remarkable diversity of new nonempidoid orthorrhaphan flies from the mid-Cretaceous of Myanmar (Late Albian–Early Cenomanian, ca. 99 Ma) is presented, including 28 species (all but one new) in 22 genera (13 new), and at least 12 families. Two families are new; three genera are unplaced in Tabanomorpha and one unplaced within Brachycera. Comparisons are presented between the amber taxa and extensive lithified taxa from the Jurassic and Cretaceous of eastern Laurasia.In Stratiomyomorpha: A new species of Zhangsolvidae has color patterns and body shape that apparently mimic Vespidae or other stinging aculeate wasps. Diverse new Xylomyidae and Stratiomyidae are described, the latter with male terminalia preserved in detail. In Tabanomorpha: The genus Athericites Mostovski et al. is synonymized with Palaepangonius Ren, and a new species of Galloatherix Nel is described in which the female proboscis is much longer than that of the male. All three genera were attributed to Athericidae, but the amber species reveal they are more basal tabanomorphs. Described are a new genus of stem-group Tabanoidea and a new species of Cratotabanus Martins-Neto and Kucero-Santos (Tabanidae), previously known from the Cretaceous of Brazil and New Jersey. In Nemestrinoidea: Three species of Hirmoneura Meigen are the first Nemestrinidae known from amber; one species has long cerci typical of Recent species of the genus. A new species of the Mesozoic family Rhagionemestriidae reveals this family is closely related to Acroceridae, not Xylophagidae as previously proposed. In Archisargoidea: A new species of Tethepomyiidae is described, the family known only in Cretaceous amber from New Jersey, Spain, and Myanmar. In Asiloidea: Three new species and genera of Bombyliidae are described, two of them with abdominal setal “baskets” distinctive to females of higher bombyliids. A fourth new asiloid genus is a probable stem-group bombyliid. The recently described Pseudorhagio Zhang et al., is transferred from Tabanomorpha to Bombyliidae. The male of a new species of Burmapsilocephala Gaimari and Mostovski (Apsilocephalidae) has terminalia very similar to that of Evocoa (Evocoidae: Recent, Chile), corroborating close relationship of the two families. In Families incertae sedis: A new species of Hilarimorphites Grimaldi and Cumming (Apystomyiidae) is described, the family known only in New Jersey and Burmese amber and the Recent fauna of California, and a sister group to either Cyclorrhapha or Eremoneura. Mysteromyiidae and Eucaudomyiidae, new families, are described, based on highly modified species with vestigial mouthparts, reduced venation, and unique specializations.Discussion is presented on some morphological features in fossil and Recent Brachycera that have apparent phylogenetic significance but are rarely discussed. This study adds further data to the phenotypic diversity, geological history, and biogeography of a major radiation.

Species in the Drosophila bromeliae group visit flowers, where most or all of the species probably breed. They are not collected in significant numbers. The group has thus far included five Neotropical species: aguape Val and Marques, 1996 (from southern Brazil), bromeliae Sturtevant, 1921 (Cuba), bromelioides Pavan and Cunha, 1947 (Brazil), florae Sturtevant, 1921 (type locality Cuba, also reported from the Caribbean and Central Ameria), and speciosa Silva and Martins, 2004 (northern Brazil). Based on specimens from various museums, collections by the author, and detailed study of some types, the following revisions are made to the group: Drosophila florae is known only on the basis of the holotype female from Cuba, and bromeliae is widespread throughout Central America, the Caribbean, and northern South America; both species are redescribed in detail. Nine new species are described: Drosophila billheedi, n. sp. (from Trinidad); D. manni, n. sp. (Bolivia); D. mexiflora, n. sp. (Mexico, possibly ...

Across terrestrial ecosystems, modern ants are ubiquitous. As many as 94 out of every 100 individual arthropods in rainforests are ants, and they constitute up to 15% of animal biomass in the Amazon. Moreover, ants are pervasive agents of natural selection as over 10,000 arthropod species are specialized inquilines or myrmecomorphs living among ants or defending themselves through mimicry. Such impact is traditionally explained by sociality: ants are the first major group of ground-dwelling predatory insects to become eusocial, increasing efficiency of tasks and establishing competitive superiority over solitary species. A wealth of specimens from rich deposits of 99 million-year-old Burmese amber resolves ambiguity regarding sociality and diversity in the earliest ants. The stem-group genus Gerontoformica maintained distinct reproductive castes including morphotypes unknown in solitary aculeate (stinging) wasps, providing insight into early behavior. We present rare aggregations of workers, indicating group recruitment as well as an instance of interspecific combat; such aggression is a social feature of modern ants. Two species and an unusual new genus are described, further expanding the remarkable diversity of early ants. Stem-group ants are recovered as a paraphyletic assemblage at the base of modern lineages varying greatly in size, form, and mouthpart structure, interpreted here as an adaptive radiation. Though Cretaceous stem-group ants were eusocial and adaptively diverse, we hypothesize that their extinction resulted from the rise of competitively superior crown-group taxa that today form massive colonies, consistent with Wilson and Hölldobler's concept of "dynastic succession."

A hallmark of animals that are eusocial, or those with advanced sociality, is reproductive specialization into worker and queen castes. In the most derived societies, these divisions are essentially fixed and in some arthropods, include further specialization--a tripartite system with a soldier caste that defends the colony. Eusociality has originated numerous times among insects but is believed to have appeared first in the termites (Isoptera), in the Early Cretaceous. However, all termites known from the Cretaceous have, until now, only been winged reproductives (alates and dealates); the earliest soldiers and definitive workers were known from just the Miocene (ca. 17-20 million years ago [mya]). Here, we report six termite species preserved in Early Cretaceous (ca. 100 mya) amber from Myanmar, one described as Krishnatermes yoddha gen. et sp. nov., comprising the worker/pseudergate, winged reproductive, and soldier, and a second species, Gigantotermes rex gen. et sp. nov., based on one of the largest soldier termites yet known. Phylogenetic analysis indicates that Krishnatermes are in the basal "Meiatermes-grade" of Cretaceous termites. Workers/pseudergates of another four species are briefly described, but not named. One of these workers/pseudergates reveals that ants--the most serious enemies of modern termites--lived in close proximity to termites in the Burmese paleofauna. These discoveries demonstrate the Mesozoic antiquity of specialized termite caste systems and corroborate that among all social species, termites probably had the original societies.

Insects have long been thought to largely not require hemoglobins, with some notable exceptions like the red hemolymph of chironomid larvae. The tubular, branching network of tracheae in hexapods is traditionally considered sufficient for their respiration. Where hemoglobins do occur sporadically in plants and animals, they are believed to be either convergent, or because they are ancient in origin and their expression is lost in many clades. Our comprehensive analysis of 845 Hexapod transcriptomes, totaling over 38 Gbases, revealed the expression of hemoglobins in all 32 orders of hexapods, including the 29 recognized orders of insects. Discovery and identification of 1333 putative hemoglobins were achieved with target-gene BLAST searches of the NCBI TSA database, verifying functional residues, secondary- and tertiary-structure predictions, and localization predictions based on machine learning. While the majority of these hemoglobins are intracellular, extracellular ones were recovered in 38 species. Gene trees were constructed via multiple-sequence alignments and phylogenetic analyses. These indicate duplication events within insects and a monophyletic grouping of hemoglobins outside other globin clades, for which we propose the term insectahemoglobins. These hemoglobins are phylogenetically adjacent and appear structurally convergent with the clade of chordate myoglobins, cytoglobins, and hemoglobins. Their derivation and co-option from early neuroglobins may explain the widespread nature of hemoglobins in various kingdoms and phyla. These results will guide future work involving genome comparisons to transcriptome results, experimental investigations of gene expression, cell and tissue localization, and gas binding properties, all of which are needed to further illuminate the complex respiratory adaptations in insects.

The amber fossil record of land snails is poorly studied, with few described species relative to other invertebrate groups (e.g., insects and chelicerates). Recently discovered land snail amber fossils from Myanmar present an important opportunity to understand the tropical land snail fauna of the Cretaceous, which was a time when many terrestrial invertebrates were rapidly diversifying. We describe 12 new land snail species in 3 families from mid-Cretaceous Burmese amber: Eotrichophorus kachin gen. et sp. nov., Perissocyclos kyrtostoma gen. et sp. nov., Macropupina electricus gen. et sp. nov., Paleodiplommatina spelomphalos gen. et sp. nov., Xenostoma lophopleura gen. et sp. nov. and Assiminea striatura sp. nov. and six species of uncertain taxonomic affinity. We also discuss the fossil history of the Cyclophoroidea. We report the first mid-Cretaceous land snail species with periostracal hairs and discuss their adaptive significance, as well as the general paleoecology of Burmese mid-Cretaceous tropical land snails.

Study of all flies (Diptera) collected for one year from a four-hectare (150 x 266 meter) patch of cloud forest at 1,600 meters above sea level at Zurquí de Moravia, San José Province, Costa Rica (hereafter referred to as Zurquí), revealed an astounding 4,332 species. This amounts to more than half the number of named species of flies for all of Central America. Specimens were collected with two Malaise traps running continuously and with a wide array of supplementary collecting methods for three days of each month. All morphospecies from all 73 families recorded were fully curated by technicians before submission to an international team of 59 taxonomic experts for identification. Overall, a Malaise trap on the forest edge captured 1,988 species or 51% of all collected dipteran taxa (other than of Phoridae, subsampled only from this and one other Malaise trap). A Malaise trap in the forest sampled 906 species. Of other sampling methods, the combination of four other Malaise traps and an intercept trap, aerial/hand collecting, 10 emergence traps, and four CDC light traps added the greatest number of species to our inventory. This complement of sampling methods was an effective combination for retrieving substantial numbers of species of Diptera. Comparison of select sampling methods (considering 3,487 species of non-phorid Diptera) provided further details regarding how many species were sampled by various methods. Comparison of species numbers from each of two permanent Malaise traps from Zurquí with those of single Malaise traps at each of Tapantí and Las Alturas, 40 and 180 km distant from Zurquí respectively, suggested significant species turnover. Comparison of the greater number of species collected in all traps from Zurquí did not markedly change the degree of similarity between the three sites, although the actual number of species shared did increase. Comparisons of the total number of named and unnamed species of Diptera from four hectares at Zurquí is equivalent to 51% of all flies named from Central America, greater than all the named fly fauna of Colombia, equivalent to 14% of named Neotropical species and equal to about 2.7% of all named Diptera worldwide. Clearly the number of species of Diptera in tropical regions has been severely underestimated and the actual number may surpass the number of species of Coleoptera. Various published extrapolations from limited data to estimate total numbers of species of larger taxonomic categories (e.g., Hexapoda, Arthropoda, Eukaryota, etc.) are highly questionable, and certainly will remain uncertain until we have more exhaustive surveys of all and diverse taxa (like Diptera) from multiple tropical sites. Morphological characterization of species in inventories provides identifications placed in the context of taxonomy, phylogeny, form, and ecology. DNA barcoding species is a valuable tool to estimate species numbers but used alone fails to provide a broader context for the species identified.

Made available in DSpace on 2019-10-06T15:56:52Z (GMT). No. of bitstreams: 0 Previous issue date: 2018-12-01 Estimations of tropical insect diversity generally suffer from lack of known groups or faunas against which extrapolations can be made, and have seriously underestimated the diversity of some taxa. Here we report the intensive inventory of a four-hectare tropical cloud forest in Costa Rica for one year, which yielded 4332 species of Diptera, providing the first verifiable basis for diversity of a major group of insects at a single site in the tropics. In total 73 families were present, all of which were studied to the species level, providing potentially complete coverage of all families of the order likely to be present at the site. Even so, extrapolations based on our data indicate that with further sampling, the actual total for the site could be closer to 8000 species. Efforts to completely sample a site, although resource-intensive and time-consuming, are needed to better ground estimations of world biodiversity based on limited sampling. Entomology Section Natural History Museum of Los Angeles County, 900 Exposition Boulevard Royal British Columbia Museum and the American Museum of Natural History, 691-8th Ave. SE Department of Plant and Environmental Sciences Poole Agricultural Center Clemson University, 130 McGinty Court, E-143 Depto. de Biologia FFCLRP Universidade de São Paulo, Av. Bandeirantes 3900 Great Lakes Forestry Centre Canadian Forest Service Natural Resources Canada, 1219 Queen St. E. Australian Museum, 1 William Street Department of Natural Resource Sciences McGill University Macdonald Campus Canadian National Collection of Insects Invertebrate Biodiversity Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling Avenue Department of Biological Sciences Spaulding Hall University of New Hampshire Department of Biological Sciences Wright State University, 3640 Colonel Glenn Hwy Instituto Federal do Triângulo Mineiro – Campus Uberaba. Rua João Batista Ribeiro 4000 Distrito Industrial II Departamento de Zoologia Universidade Federal do Paraná Jardim das Américas Mississippi Entomological Museum Mississippi State University, 100 Old Highway 12, P.O. Drawer 9775 Department of Biology Western Kentucky University Integrative Research Center Field Museum of Natural History Independent Investigator California State Collection of Arthropods, 2683 Tam O’ Shanter Dr., El Dorado Hills California Department of Food and Agriculture California State Collection of Arthropods, 3294 Meadowview Rd. The Academy of Natural Sciences of Drexel University, 1900 Ben Franklin Parkway American Museum of Natural History, Central Park West at 79th St. Department of Entomology University of California Riverside, 900 University Ave. Zoological Museum Biodiversity Unit FI-20014 University of Turku Instituto de Ecología A.C. (INECOL) Red Ambiente y Sustentabilidad, Carretera Antigua a Coatepec 351, Col El Haya, Xalapa Station Linné, Ölands Skogsby 161 I. I. Schmalhausen Institute of Zoology of the National Academy of Sciences of Ukraine, Bogdan Chmielnicki St. 15 Universidad de Panama Department of Natural History University Museum of Bergen University of Bergen, P.O. Box 7800 Agriculture and Agri-Food Canada, 960 Carling Avenue School of Environmental Sciences University of Guelph Department of Entomology Smithsonian Institution, PO Box 37012, MRC 169 Natural History Museum of Denmark, Universitetsparken 15 Institute of Evolutionary Biology and Environmental Studies University of Zurich, Winterthurerstrasse 190 Systematic Entomology Laboratory USDA ARS c/o National Museum of Natural History, MRC-168, P.O. Box 37012 Departamento de Sistemática e Ecologia CCEN Universidade Federal da Paraíba, Castelo Branco, s/n Research Institute for Nature and Forest (INBO), Kliniekstraat 25 Research Group Terrestrial Ecology (TEREC) Ghent University, K.L.Ledeganckstraat 35 Entomology Unit Royal Belgian Institute for Natural Sciences (RBINS), Vautierstraat 29 USDA Forest Service Rocky Mountain Research Station Forestry Sciences Laboratory, 1648 S. 7th Avenue Department of Biological Sciences Bishop’s University, 2600 College Street UNESP - Univ Estadual Paulista Faculdade de Ciências Agrárias e Veterinárias Departamento de Morfologia e Fisiologia Animal; Via de Acesso Prof. Paulo Donato Castellane s/n Canadian National Collection of Insects & Canadian Food Inspection Agency OPL-Entomology, K.W. Neatby Bldg., C.E.F., 960 Carling Ave. Department of Biological Sciences University of Calgary, 2500 University Drive NW Finnish Museum of Natural History Zoology Unit University of Helsinki Washington State University, 2533 Inter Avenue Bohart Museum of Entomology University of California, One Shields Avenue Instituto Nacional de Biodiversidad (INBio) UNESP - Univ Estadual Paulista Faculdade de Ciências Agrárias e Veterinárias Departamento de Morfologia e Fisiologia Animal; Via de Acesso Prof. Paulo Donato Castellane s/n

The originally published version of this Article was updated shortly after publication to add the word ‘Ticks’ to the title, following its inadvertent removal during the production process. This has now been corrected in both the PDF and HTML versions of the Article.

A new genus and species of predaceous water bugs,Triassonepa solensisn. gen. n. sp., is described from the Triassic Cow Branch Formation of Virginia and North Carolina (USA) based on ~36 adult specimens and 51 nymphs. This species is the oldest known member of the extant family Belostomatidae. It is placed in a new genus based on the unique structure of the raptorial foreleg, in which the tarsus is elongate and opposed to the tibia + femur. The fossil record of this family is reviewed and the paleoenvironmental implications of the species assemblage preserved in the Cow Branch Formation are discussed.

David Grimaldi introduces amber and the fossils contained therein.

Background and Aims The first documented observation of pollination in Pleurothallidinae was that of Endre ́s, who noticed that the ‘viscid sepals’ of Specklinia endotrachys were visited by a ‘small fly’. Chase would later iden- tify the visiting flies as being members of the genus Drosophila. This study documents and describes how species of the S. endotrachys complex are pollinated by different Drosophila species. Methods Specimens of Specklinia and Drosophila were collected in the field in Costa Rica and preserved in the JBL and L herbaria. Flies were photographed, filmed and observed for several days during a 2-year period and were identified by a combination of non-invasive DNA barcoding and anatomical surveys. Tissue samples of the sepals, petals and labellum of Specklinia species were observed and documented by SEM, LM and TEM. Electroantennogram experiments were carried out on Drosophila hydei using the known aggregation pheromones ethyl tiglate, methyl tiglate and isopropyl tiglate. Floral compounds were analysed by gas chromatography–mass spectometry using those same pheromones as standards. Key Results Flowers of S. endotrachys, S. pfavii, S. remotiflora and S. spectabilis are visited and pollinated by several different but closely related Drosophila species. The flies are arrested by aggregation pheromones, including ethyl tiglate, methyl tiglate and isopropyl tiglate, released by the flowers, and to which at least D. hydei is very sen- sitive. Visible nectar drops on the adaxial surface of sepals are secreted by nectar-secreting stomata, encouraging male and female Drosophila to linger on the flowers for several hours at a time. The flies frequently show courtship behaviour, occasionally copulating. Several different Drosophila species can be found on a single Specklinia species. Conclusions Species of the S. endotrachys group share a similar pollination syndrome. There seem to be no spe- cies-specific relationships between the orchids and the flies. It is not expected that Specklinia species will hybridize naturally as their populations do not overlap geographically. The combination of pheromone attraction and nectar feeding is likely to be a generalized pollination syndrome in Pleurothallidinae. Ministerio del Ambiente y Energía/[]/MINAE/Costa Rica Sistema Nacional de Áreas de Conservación/[]/SINAC/Costa Rica Universidad de Costa Rica/[814-B1-239]/UCR/Costa Rica Universidad de Costa Rica/[814-B3-075]/UCR/Costa Rica UCR::Vicerrectoría de Investigación::Unidades de Investigación::Ciencias Agroalimentarias::Jardín Botánico Lankester (JBL)

New species of fossil Aphelinidae and Trichogrammatidae are described from middle Eocene (Lutetian) Baltic amber (41.3–47.8 Ma). A new subfamily, two new genera and three new species of Aphelinidae are described, with comments on their placement: Phtuaria fimbriae gen.n., sp.n. in Phtuariinae subf.n., Glaesaphytis interregni gen.n., sp.n. and Centrodora brevispinae sp.n. These represent the first described true fossil Aphelinidae. Four new species of Trichogrammatidae are described: Mirufens illusionis sp.n., Palaeogramma eos gen.n., sp.n., Pterandrophysalis plasmans sp.n. and Szelenyia terebrae sp.n., thus expanding our knowledge of fossil Trichogrammatidae beyond the single previously described species. The presence of recognizable extant genera of Aphelinidae and Trichogrammatidae in the Eocene suggests that the morphology of these genera has been relatively invariant despite highly variable conditions during and since the Eocene. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:E9AD60B6-3D56-4E74-AA54-F7B91F4FDC79.

The monophyletic family Zhangsolvidae comprises stout-bodied brachyceran flies with a long proboscis and occurring only in the Cretaceous, originally known in shale from the Early Cretaceous Laiyang Formation (Fm.) in China (Zhangsolva Nagatomi & Yang), subsequently from limestones of the Early Cretaceous Crato Fm. of Brazil. Cratomyoides Wilkommen is synonymized with Cratomyia Mazzarolo & Amorim, both from the Crato Fm.; Cratomyiidae is synonymized with Zhangsolvidae. Two genera and three species of Zhangsolvidae are described: Buccinatormyia magnifica Arillo, Penalver & Perez-de la Fuente, gen. et sp.n. and B. soplaensis Arillo, Penalver & Perez-de la Fuente, sp.n., in Albian amber from Las Penosas Fm. in Spain; and Linguatormyia teletacta Grimaldi, gen. et sp.n., in Upper Albian–Lower Cenomanian amber from Hukawng Valley in Myanmar. Buccinatormyia soplaensis and Linguatormyia teletacta are unique among all Brachycera, extant or extinct, by their remarkably long, flagellate antennae, about 1.6× the body length in the latter species. A phylogenetic analysis of 52 morphological characters for 35 taxa is presented, 11 taxa being Cretaceous species, which supports placement of the family within Stratiomyomorpha, although not to any particular family within the infraorder. This published work has been registered in Zoobank, http://zoobank.org/urn:lsid:zoobank.org:pub:F32CF887-7C37-45D5-BD6B-135FE9B729A7.

• Ceratomyrmex ellenbergeri Perrichot, et al. – an exceedingly unusual taxon described from four worker specimens of varying completeness. Retaining the same overall Cretaceous trap-jaw bauplan as other haidomyrmecines, C. ellenbergeri possesses a unique clypeal horn coated ventrally in elongate, tapered setae. The mandibles are correspondingly very long, apparently extended to pin prey between the mandibular points and lengthy clypeal projection. This configuration is a dramatic variant on the trap-jaw syndrome and would appear to confirm vertical mandibular articulation. Locality: Burmese amber.

Linguamyrmex vladi Barden & Grimaldi sp.n. http://zoobank.org/urn:lsid:zoobank.org:act:2C55A44F-974D- 4A33-BDC2-26269687577A Figs 1, 2 A, 3F���I, 5. Diagnosis. As for genus. Description. Head: measuring 0.90 mm postero-anteriorly along dorsal margin, 0.96 mm in length/depth from vertex of head to anterior margin of clypeus. Occipital foramen positioned highly dorsad, just under vertex of head. Postgena broadly depressed; postgenal suture visible, deeply furrowed. Vertex broadly rounded and glabrous with gena gradually tapered ventrally towards mandibular socket and oral opening with fine, sparse setae. Ocelli faintly visible on vertex, positioned dorsally. Eye situated high on head capsule and bulging in frontal view, ovoid, measuring 0.38 mm in length and 0.25 mm in width when viewed laterally. Three antennal segments fully preserved (scape 0.94 mm in length; pedicel 0.12 mm; flagellomere I 0.55 mm). Antennal socket approximately in line with ventral margin of eye; socket exposed and immediately flanking a medial frontal triangle (sensu Perrichot et al., 2016). Frontal triangle extends the vertex, contrasted with antennal sockets, which are present within cuticular depressions. Clypeal horn originating at both frontal triangle and clypeal stalk, both structures heavily sclerotized with cleared, membrane-like cuticle connecting from frontal triangle to stalk. Horn paddle-shaped, total length 0.64 mm, diameter 0.49 at greatest; narrow (0.04 mm) glabrous stalk 0.22 mm in length leading to setose pad; setose pad (0.42 in length) with long trigger hairs originating at pad base; dorsal margin of setose pad glabrous, underside coated in a large number of stout setae in centre and longer, more tapered setae along edges. Anterior margin of clypeus medially triangulate; distinct medial ridge extending to clypeal horn; lateral margins, beginning just above mandibular insertion, extending diagonally toward clypeal horn. Cheek-like lobes projecting anteroventrally above mandible insertion. Mandible scythe-like, comprise a linear basal margin (0.55 mm in length) and curved apical tooth (0.92 mm measured as a straight line from base to tip, ignoring curvature) meeting nearly at right angle; preserved with apical teeth in parallel, nearly touching. Basal portion of mandible with anterior flange-like expansion, concave inner margin coated with pointed setae; leading edge of anterior flange expansion smooth; apical tooth rounded broadly with slight point. Maxillary and labial palps not visible. Mesosoma. Pronotum broad, coated in fine setae; propleuron reduced, not visible in lateral view except where abutting head capsule anteriorly. Pronotal length 0.72 mm, measured along dorsal margin. Mesonotum 0.34 mm; metanotum 0.24 mm; propodeum 0.49 mm. Weber���s length 1.77 mm, mesosomal height 0.56 mm measured perpendicular to Weber���s length line at pronotum. Procoxal length 0.72 mm, max width 0.22; mesocoxal length 0.31, max width 0.21; metacoxal length 0.49, max width 0.26; protrochantal length 0.14, max width 0.11; mesotrochantal length 0.19, max width 0.11; metatrochantal length 0.19, max width 0.12; profemoral length 1.01, max width 0.13; mesofemoral length 1.09, max width 0.12; metafemoral length 1.48, max width 0.18; protibial length 1.09, max width 0.11; mesotibial length 1.30, max width 0.08; metatibial length 1.59, max width 0.08. Trochantellus present, length included in femur measurements. Three protibial spurs present, the largest approximately 2 �� length of other two; mesotibia with two spurs, the larger 2 �� the length of the smaller; two conspicuous setae of equal length positioned along anterior margin of mesotibial apex; metatibia with two spurs, the largest pectinate and ~ 3 �� the size of the smaller. Pretarsal claw with subapical tooth positioned closer to apex than to claw origin. Dorsal margin of propodeum gradually rounded, posterior margin with sheer face. Propodeal spiracle a dorsoventrally elongate slit situated medially. Metapleural gland opening gaping, present within horizontal invagination of cuticle, opening visible posteroventrally following well-developed bulla. Metasoma. Petiole 0.49 mm in length, 0.49 mm in height at greatest, pedunculate; not tergosternally fused, deep suture visible; anterior margin gradually sloping; with gradually rounded dorsal margin; broadly attached to gastral segment I (abd seg III) with striated helcium clearly visible. Lateral sulcus visible running anteroposteriorly along petiole, signifying incomplete fusion; ventral to sulcus lies thin, membraneous cuticular expansion with distinct anteriorly positioned keel. Sternite III with pointed projection on anterior margin exhibiting striations running laterally. Circumgastral constriction present between gastral segments I and II with dorsal, posteriorly extended notch. Gastral cuticle translucent, revealing heavily sclerotized and expanded apodemes of gastral sternites IV and V withdrawn into the gaster. Type material. Holotype, AMNH BuPH-01, in the Department of Invertebrate Zoology, American Museum of Natural History. Additional material examined. Although not named, three specimens, BuPH-02 (Figs 2 D, 3C, 7, Video S1), BuPH-03 (Figs 2 B, 3A, B, 4) and BuPH-04 (2C), were studied and are illustrated here. These three specimens probably represent additional species, particularly with respect to overall size and clypeal paddle composition. Moreover, these specimens possess mandibles that appear to interlock basally, joining the distal apical tooth portions of the mouthparts (Figs 3 D, E, 7, Video S1). As the inner surface of each mandible is concave, their joining forms a channel that is open dorsally near the pointed mandibular apex (Fig. 7). However, there remains some ambiguity regarding the reliability of measurements and identification of other discrete features due to incomplete preservation. We therefore delay description until more complete material is obtained. Measurements are provided here (in mm): (BuPH-02) [BuPH-03] {BuPH-04} Head length measured postero-anteriorly along dorsal margin (1.14) [0.87] {0.69}, head depth (1.48) [1.22] {1.2}, scape length (1.56) [1.36] {1.28}, paddle length (1.5) [1.52] {1.26}, paddle diameter (���) [~ 0.57] {���}, eye length (���) [���] {0.29}, eye width (���) [���] {0.19}, basal margin of mandible (0.71) [0.53] {0.39}, apical tooth (1.97) [1.57] {1.48}, pronotal length (1.08) [0.97] {0.8}, mesosomal height (~ 0.81) [~ 0.75] {0.66}, Weber���s length (2.64) [2.49] {1.99}, petiole length (0.82) [0.76] {0.60}. Unfortunately, no specimen retained all antennal segments, preventing a reliable comparison of scape length relative to all other segments. In addition, all specimens, including the type, are missing terminal abdominal segments. Etymology. In reference to Vlad III, or Vlad Dracula (c. 1429���1476), prince of a region of Romania then called Wallachia. His moniker, Vlad the Impaler, refers to his favoured and frequent method of execution, which inspired the vampirous character Count Dracula fictionalized by Bram Stoker in 1897. The patronym is in reference to the presumed impalement of prey by Linguamyrmex and its liquid diet (see later). Systematic comments. Haidomyrmecines have been recovered as monophyletic (Barden & Grimaldi, 2016) and have traditionally been placed as tribe within Sphecomyrminae (Bolton, 2003). The subfamily is defined largely by plesiomorphic features and is now suspected to be paraphyletic (Barden & Grimaldi, 2016). We tentatively follow this subfamilial assignment here but note that, following additional phylogenetic analyses, haidomyrmecines may be best served by the erection of a separate subfamily., Published as part of Phillip Barden, Hollister W. Herhold & David A. Grimaldi, 2017, A new genus of hell ants from the Cretaceous (Hymenoptera: Formicidae: Haidomyrmecini) with a novel head structure, pp. 837-846 in Systematic Entomology 42 on pages 839-842, DOI: 10.1111/syen.12253, http://zenodo.org/record/887095, {"references":["Perrichot, V., Wang, B. & Engel, M. S. (2016) Extreme morphogenesis and ecological specialization among Cretaceous basal ants. Current Biology, 26, 1468 - 1472.","Barden, P. & Grimaldi, D. A. (2016) Adaptive radiation in socially advanced stem-group ants from the Cretaceous. Current Biology, 26, 515 - 521.","Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute, 71, 1 - 370."]}

��� Haidomyrmex cerberus Dlussky ��� the first hell ant was named 20 years ago by the prolific paleomyrmecologist Gennady Dlussky (Dlussky, 1996). Haidomyrmex cerberus was described from a single partial worker specimen, providing the basis for what are now tribal synapomorphies for the Haidomyrmecini, established by Bolton (2003) and revised by Perrichot et al. (2016): unusual L-shaped mandibles and a bulging clypeus possessing a clypeal brush comprising dense, stout setae. Locality: Burmese amber, ~ 98.8 Ma, Kachin State, Myanmar (Shi et al., 2012), Published as part of Phillip Barden, Hollister W. Herhold & David A. Grimaldi, 2017, A new genus of hell ants from the Cretaceous (Hymenoptera: Formicidae: Haidomyrmecini) with a novel head structure, pp. 837-846 in Systematic Entomology 42 on page 838, DOI: 10.1111/syen.12253, http://zenodo.org/record/887095, {"references":["Dlussky, G. M. (1996) Ants (Hymenoptera: Formicidae) from Burmese amber. Paleontologicheskii Zhurnal, 30, 449 - 454.","Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute, 71, 1 - 370.","Perrichot, V., Wang, B. & Engel, M. S. (2016) Extreme morphogenesis and ecological specialization among Cretaceous basal ants. Current Biology, 26, 1468 - 1472.","Shi, G., Grimaldi, D. A., Harlow, G. E. et al. (2012) Age constraint on Burmese amber based on U - Pb dating of zircons. Cretaceous Research, 37, 155 - 163."]}

• Haidomyrmodes mammuthus Perrichot, et al. – described from both partial gyne and worker specimens, H. mammathus is the only alate specimen yet described (Perrichot et al., 2008). In addition, the type material is the only reported conspecific worker and putative queen caste for any Cretaceous taxon and provides important information regarding sociality in early ants. The description of multiple specimens from an additional locality confirmed beyond any doubt that the highly unusual features observed in H. cerberus were not preservational. Largely similar to H. cerberus, H. mammathus possesses ocelli, which have long been presumed to be an ancestral ant trait (Wilson et al., 1967b). Locality: Charentese amber, ~ 100 Ma, Charentes, France (Perrichot et al., 2010)

Genus Linguamyrmex Barden & Grimaldi, new genus http://zoobank.org/urn:lsid:zoobank.org:act:9F04FAEE-59DA- 4B3C-AB8B-EE0D234CC118 Figs 1–5, 7, Video S1. Diagnosis, worker. As in other haidomyrmecines (Haidomyrmex, Haidoterminus, Haidomyrmodes, Ceratomyrmex), head hypognathous-like with mandibles projecting primarily downward; mandible scythe-like, with flattened basal margin leading to a curved apical tooth that is expanded perpendicular to axial plane of head. Cephalic clypeal ‘horn’ present but abbreviated, differs from Ceratomyrmex by horn stalk of Linguamyrmex being glabrous, that of Ceratomyrmex with fine, stiff setae of various lengths; clypeal horn much shorter in Linguamyrmex, less than head length/depth, stalk short, with the expanded, flat, paddle-shaped setose pad comprising>50% total horn length; clypeal pad slightly trough-shaped ventrally, covered with very short, dense velcro-like vestiture; trigger hairs originate not at base of stalk as in Ceratomyrmex but near basal margin of setose pad; ocelli present. In addition, Linguamyrmex with first and second gastral segments with deep constriction between them (a gastral constriction is figured in description of Haidomyrmodes mammuthus but is less developed). Type species. L. vladi sp.n. Etymology. From Latin ‘lingua’, meaning tongue – in reference to the tongue-like clypeal projection – and the Greek ‘myrmex’, meaning ant.