Wing base structure supports Coleorrhyncha + Auchenorrhyncha (Insecta: Hemiptera)

The phylogenetic placement of the moss bugs (Insecta: Hemiptera: Coleorrhyncha) has been highly controversial. Many apparent morphological apomorphies support the close relationship between Coleorrhyncha and Heteroptera (=true bugs). However, a recent phylogenomic study strongly supported a sister group relationship between Coleorrhyncha and Auchenorrhyncha (planthoppers, leafhoppers, treehoppers, spittlebugs, and cicadas). To test these two alternative hypotheses, we examined the fore‐ and hindwing base structure of the only known extant macropterous species of Coleorrhyncha using binocular and confocal laser scanning microscopes and analyzed the data selected from the wing base phylogenetically. When full morphological data including the wing base characters were analyzed, the sister group relationship between Coleorrhyncha + Heteroptera was supported, agreeing with previous consensus based on morphology. In contrast, when only wing base characters were analyzed separately, the clade Coleorrhyncha + Auchenorrhyncha was recovered, in agreement with the result from the phylogenomic study. The membranous condition of the proximal median plate in the forewing was identified as a potential synapomorphy of the latter grouping, and the absence of the tegula was excluded as a potential synapomorphy of Coleorrhyncha and Heteroptera.     

The first Ironomyiidae from mid‐Cretaceous Burmese amber provides insights into the phylogeny of Phoroidea (Diptera: Cyclorrhapha)

Macalpinomyia jiewenae gen. et sp.n. is described from the mid‐Cretaceous (～99 Ma) amber of Myanmar. Macalpinomyia jiewenae is the first Oriental representative of the enigmatic family Ironomyiidae (Diptera: Phoroidea), currently known from a single extant genus restricted to southeastern Australia, plus a monotypic genus from Canadian amber and three controversial genera based on impression fossils from China, Mongolia and Russia. A phylogenetic analysis of all Phoroidea families, including all ironomyiid extant and extinct genera, corroborates the monophyly of Ironomyiidae, and Macalpinomyia gen.n. is assigned to the subfamily Sinolestinae. Cretonomyiinae subfam.n. , is erected to accommodate the basal lineage of Ironomyiidae. Lebambromyia acrai Grimaldi & Cumming, previously placed in Ironomyiidae, is supported as an early branching lineage of Platypezidae. Our topology proposes that Ironomyiidae is sister to the remaining Phoroidea. The phylogenetic results, in combination with the fossil ages and relevant molecular divergence time analysis, suggests that Ironomyiidae probably originated at least in the Berriasian of the Early Cretaceous (～140 Ma). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:5DFFC944‐1350‐418E‐BCDC‐BB87FC013D5D .     

Phylogeny and historical biogeography of Gondwanan moss-bugs (Insecta: Hemiptera: Coleorrhyncha: Peloridiidae)

The moss bugs of the Peloridiidae, a small group of cryptic and mostly flightless insects, is the only living family in Coleorrhyncha (Insecta: Hemiptera). Today 37 species in 17 genera are known from eastern Australia, New Zealand, New Caledonia and Patagonia, and the peloridiids are thereby a group with a classical southern Gondwanan distribution. To explicitly test whether the present-day distribution of the Peloridiidae actually results from the sequential breakup of southern Gondwana, we provide the first total-evidence phylogenetic study based on morphological and molecular characters sampled from about 75% of recognized species representing 13 genera. The results largely confirm the established morphological phylogenetic context except that South American Peloridium hammoniorum constitutes the sister group to the remaining peloridiids. A timescale analysis indicates that the Peloridiidae began to diversify in the land mass that is today's Patagonia in the late Jurassic (153 Ma, 95% highest posterior density: 78–231 Ma), and that splitting into the three extant well-supported biogeographical clades (i.e. Australia, Patagonia and New Zealand/New Caledonia) is consistent with the sequential breakup of southern Gondwana in the late Cretaceous, indicating that the current transoceanic disjunct distributions of the Peloridiidae are best explained by a Gondwanan vicariance hypothesis.     

A new genus of mantidflies discovered in the Oriental region,with a higher‐level phylogeny of Mantispidae (Neuroptera) using DNA sequences and morphology

A remarkable new genus and two new species of Mantispidae (Neuroptera) are described from the Oriental region. Allomantispa Liu, Wu, Winterton & Ohl gen.n. , currently including A. tibetana Liu, Wu & Winterton sp.n. and A. mirimaculata Liu & Ohl sp.n. The new genus is placed in the subfamily Drepanicinae based on a series of morphological characteristics and on the results of total evidence phylogenetic analyses. Bayesian and Parsimony analyses were undertaken using three gene loci (CAD, 16S rDNA and COI) combined with 74 morphological characters from living and fossil exemplars of Mantispidae (17 genera), Rhachiberothidae (two genera) and Berothidae (five genera), with outgroup taxa from Dilaridae and Osmylidae. The resultant phylogeny presented here recovered a monophyletic Mantispidae with ?Mesomantispinae sister to the rest of the family. Relationships among Mantispidae, Rhachiberothidae and Berothidae support Rhachiberothidae as a separate family sister to Mantispidae. Within Mantispidae, Drepanicinae are a monophyletic clade sister to Calomantispinae and Mantispinae. In a combined analysis, Allomantispa gen.n. was recovered in a clade comprising Ditaxis McLachlan from Australia, and two fossil genera from the Palaearctic, ?Promantispa Panfilov (Kazakhstan; late Jurassic) and ?Liassochrysa Ansorge & Schlüter (Germany; Jurassic), suggesting a highly disjunct and relictual distribution for the family. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:464B06E8‐47E6‐482E‐8136‐83FE3B2E9D6B .     

The oldest known clerid fossils from the Middle Jurassic of China,with a review of Cleridae systematics (Coleoptera)

Three fossil beetles recorded from Inner Mongolia, China (Callovian, Middle Jurassic, approximately 165 Ma) are assigned to Cleridae and constitute the earliest known representatives of this family. Two of the fossils are described as Protoclerus korynetoides gen. et sp.n. and the third as Wangweiella calloviana gen. et sp.n. Ninety‐six morphological characters of the larvae and adults of 17 extant clerid representatives and seven cleroid families as outgroups were analysed under maximum parsimony with, and without, fossil species included in the matrix. The results indicate that Protoclerus gen.n. is a separate lineage within the family, whereas Wangweiella gen.n. is sister to the rest of Epiclininae. The resulting trees are compared with the most recent morphological and molecular phylogenies of Cleridae. The trees presented appear congruent with the latest molecular phylogenies. The evolution of Cleridae, as well as their biogeography and systematics, are briefly reviewed. A complete list of clerid fossil species described to date is provided. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:32240D3D-2A7A-41F6-B485-A66F2265CB8A .     

New fossil Ochteridae (Hemiptera: Heteroptera: Ochteroidea) from the Upper Mesozoic of north‐eastern China,with phylogeny of the family

Specimens of the velvety shore bugs (Hemiptera: Ochtheridae) occur in the Early Cretaceous Yixian Formation in north‐eastern China. We describe two new genera and three new species –Pristinochterus ovatus Yao, Zhang & Ren sp. nov., Floricaudus multilocellus Yao, Ren & Shih gen.n. et sp.n. and Angulochterus quatrimaculatus Yao, Zhang & Ren gen.n. et sp.n. – from this deposit. A cladistic analysis based on a combination of fossil and extant taxa, and their morphological characters, clarifies the phylogenetic status of the new fossils and allows the reconstruction of relationships within the family Ochteridae. Two main clades within Ochteridae are recognized from the cladistic analysis: Pristinochterus Yao Cai & Ren and Floricaudus Yao, Ren & Shih gen.n. form a monophyletic lineage; and Angulochterus Yao, Zhang & Ren gen.n. emerges as a sister group to all extant velvety shore bugs. A key to all fossil and extant genera of Ochteridae is provided.     

Early evolution of Cupedidae revealed by a mid‐Cretaceous reticulated beetle from Myanmar (Coleoptera: Archostemata)

Cupedidae, the most species‐rich family of the archaic suborder Archostemata, were abundant, diverse and widespread in the Mesozoic, yet little is known about the early evolution and biogeography. This stems, in part, from a lack of exceptionally preserved fossils from the Mesozoic and of formal phylogenetic study of both extant and extinct taxa. Here we describe and illustrate a new fossil from mid‐Cretaceous Burmese amber, and provide a phylogeny combining both fossils and all known extant genera of Archostemata. A dataset of 43 ingroup taxa and four outgroup taxa based on 110 morphological characters was analysed under parsimony. The results indicate that Priacma LeConte and Paracupes Kolbe, as well as the Cretaceous genera Barbaticupes Jarzembowski et al. and Mallecupes Jarzembowski et al., together form a sister clade to the rest of Cupedidae. Priacma megapuncta sp.n. is attributed to the relict North American Priacma by the presence of distinct subtruncate elytral apices, lateral elytral margins with two rows of sharp teeth, and peculiar fixing epipleural folds near the elytral apices. Our discovery of the first fossil species of Priacma in Burmese amber reveals the antiquity and wider distribution of the genus in the late Mesozoic. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:313565C2‐4F42‐48BD‐8720‐F379DE202868 .     

Early lineages of Vespidae (Hymenoptera) in Cretaceous amber

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 .     

Phylogeny of the rove beetle tribe Gymnusini sensu n. (Coleoptera: Staphylinidae: Aleocharinae): implications for the early branching events of the subfamily

The rove beetle subfamily Aleocharinae is the largest subfamily of animals known in terms of species richness. Two small aleocharine tribes, Gymnusini and Deinopsini, are believed to be a monophyletic clade, sister to the rest of the Aleocharinae. Although the phylogenetic relationships of the extant lineages have been well investigated, the monophyly of Gymnusini has been questioned due to a series of previous studies and the recent discovery of the aleocharine †Cretodeinopsis Cai & Huang (Deinopsini) from mid‐Cretaceous Burmese amber. Using an additional specimen of †Cretodeinopsis and well‐preserved specimens of †Electrogymnusa Wolf‐Schwenninger from Eocene Baltic amber, we present here two types of morphology‐based phylogenetic analyses, employing all extant/extinct genera of Gymnusini and Deinopsini for the first time. The maximum parsimony and Bayesian analyses recovered a monophyletic clade of the two tribes combined, but each analysis suggested nonmonophyly of Gymnusini. In agreement with the results of the present study, we synonymize Deinopsini syn.n. under Gymnusini sensu n. , by priority. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:F09EB444‐C6CA‐4525‐A986‐3CFC826F5877 .     

Vibrational signalling in a Gondwanan relict insect (Hemiptera: Coleorrhyncha: Peloridiidae)

Ancient, long-extinct floras and faunas can be reassembled through fossils and phylogenetics, and even palaeo-environments can be reconstructed with the aid of palaeoclimatology. However, very little is known about the sound-scape of the past. Of what kind were the first biologically meaningful sounds and vibrations ever emitted and perceived? The earliest signals in the history of life were probably produced by arthropods making use of the mechanical properties of their exoskeleton. Here, we report an observation of vibrational signalling in the coleorrhynchan Hackeriella veitchi, a representative of a Gondwanan relict insect lineage which is still extant in the Queensland rainforest. Our finding suggests that vibrational signalling by tymbal organs is ancestral for the Hemiptera (exclusive of Sternorrhyncha)--the song of the Coleorrhyncha was a likely element of the acoustic environment in the Permian moss forests and had possibly changed little since.     

New species of Primozygodactylus from Messel and the ecomorphology and evolutionary significance of early Eocene zygodactylid birds (Aves,Zygodactylidae)

Zygodactylids (Zygodactylidae) are the extinct sister taxon of passerines (Passeriformes) and among the more abundant small arboreal birds in the early Eocene German fossil site Messel. Four species of the taxon Primozygodactylus have previously been identified and here two new species are described. In addition, new fossils of the poorly known P. eunjooae are reported. The fossils corroborate the presence of two very long central tail feathers for Primozygodactylus, and the feathering of the taxon corresponds with that of extant birds foraging in scrub and undergrowth. Species diversity of zygodactylids falls short of that of passerines, the most species-rich extant avian clade, and in bill shapes they also do not reach the diversity seen in extant Passeriformes. Because zygodactylids closely resemble passerines in skeletal morphology, the evolutionary success of passerines is unlikely to be due to particular skeletal features. Passerines and zygodactylids coexisted in Europe from the early Oligocene to the middle Miocene, and both groups probably differed in ecological characteristics. The zygodactyl foot of zygodactylids may have represented an adaptation for clinging to tree trunks, and if nesting in tree cavities, zygodactylids may have succumbed to competition for safe nesting places with avian or mammalian competitors.

http://zoobank.org/urn:lsid:zoobank.org:pub:7F142141-7E1F-4568-89BF-E2363D128C36     

Diverse oxyporine rove beetles from the Early Cretaceous of China (Coleoptera: Staphylinidae)

Two new genera and species, Protoxyporus grandis gen. et sp.n. and Cretoxyporus extraneus gen. et sp.n. , are described and figured on the basis of two well‐preserved impression fossils from the Lower Cretaceous Yixian Formation of China. The two genera display a combination of traits that are diagnostic of the extant staphylinid subfamily Oxyporinae: large and prominent mandibles, distinctly separated mesocoxae, abdominal tergites III–VII each with a pair of basolateral ridges, and abdominal intersegmental membranes with ‘brick‐wall’ pattern. Protoxyporus differs from extant Oxyporus in retaining the following plesiomorphic features: an elongate intercoxal process (or longitudinal carina) between metacoxae, relatively narrowly separated mesocoxae, less developed metaventral anterior process, and long infraorbital ridges. Cretoxyporus is morphologically very similar to Oxyporus, but retains distinct elongate infraorbital ridges. The discovery of two new oxyporine genera suggests that the early oxyporines were more diverse and their evolutionary history more complicated than previously documented. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:11503976‐D577‐45D2‐ACCB‐8226782EF817 .     

A molecular phylogeny of Bopyroidea and Cryptoniscoidea (Crustacea: Isopoda)

Epicaridean isopods are parasitic on other crustaceans. They represent a diverse group of highly derived taxa in two superfamilies and 10 families. Little work has been done on the phylogeny of these parasites because of the difficulty in defining homologous characters for adults above the genus level. The females exhibit morphological reduction of characters and the males have few distinguishing characters. Moreover, epicarideans have only rarely been included in past studies of isopod phylogeny. Our objective was to derive a phylogeny of epicaridean taxa based on 18S rDNA, then use that phylogeny to examine the relationships of the bopyrid subfamilies, bopyroid families and epicarideans to cymothoid isopods. We tested the monophyly of the Epicaridea, evaluated hypotheses on relationships among epicaridean families and subfamilies, examined the evolution of the abdominal mode of infestation on caridean, gebiidean, axiidean and anomuran hosts and examined coevolution between epicarideans and their crustacean hosts. The molecular phylogeny indicated that Epicaridea were monophyletic with respect to Cymothooidea. Bopyroidea formed a monophyletic group without Dajidae and Entophilinae (now as Entophilidae). Both latter taxa grouped with Cryptoniscoidea, and this group was the sister taxon to the redefined Bopyroidea in all trees. The bopyrid subfamily Ioninae is the sister taxon to the other bopyrid subfamilies (except Entophilidae). Ioninae was elevated to family status but found not to be monophyletic; a new subfamily, Keponinae, was erected for all genera formerly placed in Ioninae except the type genus. The abdominal mode of parasitism appears to have evolved independently among the subfamilies. Coevolution between host and parasite phylogenies showed extensive incongruence, indicating frequent host-switching as a general pattern in Epicaridea.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:30ECFB13-2795-494E-AABE-6B5F84A57A67     

New fossils from China elucidating the phylogeny of Praesiricidae (Insecta: Hymenoptera)

A new subfamily of Praesiricidae (Pamphilioidea), Decorisiricinae subfam.n. , is erected based on three new genera: Decorisiricius gen.n. , Limbisiricius gen.n. and Brevisiricius gen.n. Two new species – Decorisiricius patulus gen. et sp.n. and D. longus sp.n. – from the Lower Cretaceous Yixian Formation and three species –Limbisiricius aequalis gen. et sp.n. , Limbisiricius complanatus sp.n. and Brevisiricius partialis gen. et sp.n. – from the Middle Jurassic Jiulongshan Formation, are described. Based on these well‐preserved new fossil specimens and previously published data, the nonmonophyly of Praesiricidae is confirmed and the phylogenetic relationships of species of Praesiricidae are analysed for the first time. Two main clades within Praesiricidae are recognized from the cladistic analysis: Decorisiricinae subfam.n. forms a monophyletic lineage, with the remaining members of Praesiricidae plus Megalodontes (Megalodontesidae) forming its sister group. The two subfamilies Archoxyelydinae and Praesiricinae are discarded with no strong supported synapomorphic characters based on phylogenetic research. A key to all genera of Praesiricidae is provided. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:38D703ED‐127A‐4DB0‐8153‐8D78AF4AC212 .     

New clues on anuran evolution: the oldest record of an extant hyloid clade in the Oligocene of Patagonia

Despite the enormous diversity of the extant South American batrachofauna, its fossil record is sparse. Few pre-Neogene anuran remains have been attributed to extant taxa and, although some have been considered neobatrachians their relationships with extant groups are still mostly uncertain. In this context, a series of fossils recently collected in Oligocene sediments of central Patagonia are assigned to Odontophrynidae. This assignation is supported by an exclusive combination of characters, including extensive frontoparietals, in medial contact and deflecting posteriorly, with crescent-shaped posterolateral projections; robust pterygoid anterior ramus, reaching the planum antorbitale, projecting dorsally; notched premaxillary pars palatine; dentate vomers, narrowly separated medially; robust neopalatines, narrowly separated medially, bearing a conspicuous ridge, and with expanded lateral edges. The well-preserved fossil specimens were considered representative of a new genus and species, Chachaiphrynus lynchi. The presence of an Oligocene Odontophrynidae in central Patagonia, about 500 km south from the southern-most extant distribution of the group, would be associated with the more benign climate inferred in this region during the Paleogene. At the same time, this record represents the single pre-Neogene certain record of an extant clade deeply nested in Hyloidea, providing an exceptional high-quality calibration point to temporally frame the anuran phylogeny.

http://www.zoobank.org/urn:lsid:zoobank.org:act:AA043DC2-BCA5-4301-8FB0-FEB56B0F4404     

Pleurambus strongylus n. gen., n. sp. (Coleoptera: Belidae) in Dominican amber

A new genus and species of belid weevils, Pleurambus strongylus Poinar and Legalov, n. gen., n. sp. (Coleoptera: Belidae) in the tribe Allocorynini is described from Dominican amber. Belid fossils are quite rare and the new species is the first fossil member of the family from the Neotropics. This discovery suggests that belid diversity in Hispaniola during the Tertiary was higher than at present since no extant belids have been reported from the West Indies.http://www.zoobank.org/urn:lsid:zoobank.org:pub:DF79FF39-634D-4BD7-AD0B-F5FB3BBE1081     

A new extant family of primitive moths from Kangaroo Island,Australia, and its significance for understanding early Lepidoptera evolution

We report the first discovery since the 1970s of a new extant family (Aenigmatineidae fam.n. ) of homoneurous moths, based on the small Aenigmatinea glatzella sp.n . from Kangaroo Island off southern Australia. It exhibits a combination of extraordinary anatomical characters, and, unlike most homoneurous moths, its larva is a conifer‐feeder (stem mining in Callitris, Cupressaceae). While the adult's mouthparts are strongly regressed, evidence from other morphological characters and from a Bayesian analysis of 25 genetic loci convincingly places the taxon among Glossata (‘tongue moths’). An unexpected tongue moth clade including Acanthopteroctetidae and Neopseustidae, suggested with low support in recent molecular analyses, remarkably becomes strongly supported when Aenigmatinea is included in the molecular analysis; the new taxon becomes subordinated in that clade (as sister group to Neopseustidae) and the clade itself appears as the sister group of all Heteroneura, representing the vast majority of all Lepidoptera. Including Aenigmatinea into the analysis thereby strengthens the surprising indication of non‐monophyly of Myoglossata, and the new phylogeny requires an additional number of ad hoc assumptions of convergence/character reversals in early Lepidoptera evolution. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:44393B52‐1889‐431A‐AB08‐6BBCF8F946B8 .     

The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites

Five finely preserved termites from the mid‐Cretaceous (Cenomanian) amber of Myanmar provide new information allowing a reanalysis of the phylogeny of basal termites. The Mastotermitidae family is recovered as monophyletic, and a redefined Hodotermitidae sensu lato is also monophyletic to include Archotermopsidae, Hodotermitidae and Stolotermitidae. Such a phylogenetic relationship agrees with the results from previous molecular phylogeny. Alongside these findings, there are many taxa that can only be shown to be termites with no other phylogenetically informative data. These form a comb of ‘grade groups’ emerging in the Late Jurassic. The new amber specimens are described as two new species. Anisotermes xiai gen. et sp.n. is described from multiple castes and has symplesiomorphic characteristics: large body size, a broad pronotum, well‐developed reticulated veins, and a large anal lobe of the hindwings. It shares wing features with the other new species, Mastotermes monostichus sp.n. Both new taxa are assigned to the Mastotermitidae, as they are shown to have synapomorphies that unite the family. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1AD5CECA‐27B7‐48D5‐88DC‐CEC5150962D7 .     

Molecular phylogeny and generic‐level taxonomy of the widespread palaeotropical ‘Heteropsis clade’ (Nymphalidae: Satyrinae: Mycalesina)

The mycalesine butterfly genus Heteropsis Westwood, 1850 (Satyrinae: Mycalesina) has recently been conceived to be represented in three major palaeotropical regions (Madagascar, Africa and Asia), but there has been no formal taxonomic treatment covering this entire group. Studies aimed at understanding the evolutionary success of Mycalesina in the Old World tropics have been hampered by the lack of both a robust phylogeny and a stable nomenclature for this satyrine subtribe. Here, we present a well‐supported molecular phylogeny based on 10 genes and 133 exemplar taxa, representing almost all known species groups of Heteropsis (s.l.), and including all but four known species in Madagascar. We also combine sequences of the exemplars with a morphological matrix of 428 characters. The widespread ‘Heteropsis clade’ is confirmed as monophyletic, but lineages in different geographic regions also form endemic and well‐supported clades with deep divergences among them. Here we establish this group as comprising three genera, Heteropsis (Malagasy region only), Telinga Moore, 1880 (Asia), and Brakefieldia gen.n. (Africa). We recover the genera Telinga and Brakefieldia as sisters with high support. Each genus is taxonomically characterized and a revised synonymic checklist is appended with new combinations and some changes in rank. With a well‐resolved topology and updates to the taxonomy of the group, researchers are now in a position to explore the drivers of the spectacular radiation of the group, notably in Madagascar, where the highest phenotypic and species diversity occurs. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:AAF9F440‐A2D6‐4483‐BF35‐9BC074D9D29B .     

Long‐proboscid brachyceran flies in Cretaceous amber (Diptera: Stratiomyomorpha: Zhangsolvidae)

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, Peñalver & Pérez‐de la Fuente, gen. et sp.n. and B. soplaensis Arillo, Peñalver & Pérez‐de la Fuente, sp.n. , in Albian amber from Las Peñosas 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 .