A new genus of moths (Lepidoptera: Gracillarioidea: Douglasiidae) in Myanmar amber

A new genus and species of moths, Tanyglossus orectometopus n. gen. et sp. (Lepidoptera: Gracillarioidea), is described from mid-Cretaceous Myanmar amber. Autapomorphies for the new genus are small size (body length of only 0.9 mm), long proboscis greater than body length, presence of ocelli, 3-segmented porrect labial palps, 5-segmented curved maxillary palps, no eye cap, lanceolate wings with reduced venation and a body and wing covering of several types of scales. Based on the available characters, the fossil is placed in the small, aberrant family Douglasiidae and represents the first fossil member of this family. This study adds to our scant knowledge of the diversity of Mesozoic Glossata.     

On the phylogenetic position of the palaeopteran Syntonopteroidea (Insecta: Ephemeroptera), with a new species from the Upper Carboniferous of England

A new syntonopterid, Anglolithoneura magnifica gen. et sp. n., is described from a siderite concretion (nodule) from the Late Carboniferous (Langsettian) of Lancashire County (UK). The new genus is diagnosed on hind wing venation and compared with other syntonopterid genera. The new species is the first syntonopterid formally described from the Late Carboniferous of Europe. The systematic positions of other potential Syntonopteroidea (Miracopteron mirabile, Bojophlebia prokopi, and specimens described in 1985 by J. Kukalová-Peck from Obora in the Czech Republic) are reconsidered. Wing venation synapomorphies are proposed for the Syntonopteroidea (sensu novo), and for a potential clade ((Ephemeroptera+Syntonopteroidea)+Odonatoptera) separated from the Palaeodictyopterida. The close relations of the new species with Lithoneura lameerei Carpenter, 1938 from Mazon Creek (Illinois, USA) provide additional support for a Euramerican connection during the Late Carboniferous.     

On the edge of parasitoidism: a new Lower Cretaceous woodwasp forming the putative sister group of Xiphydriidae + Euhymenoptera

The fossil woodwasp Cratoenigma articulata gen. et sp.n. (Insecta: Hymenoptera) is described from the Lower Cretaceous Crato formation of Brazil. This fossil cannot be placed in any existing superfamily, but its putative phylogenetic position within Hymenoptera is discussed in detail on the basis of relevant thoracic, abdominal and wing venation characters. These characters are critically evaluated and compared with those of extant and fossil Hymenoptera. The phylogenetic position of C. articulata sp.n. is investigated relative to extant Xyelidae, Tenthredinoidea s.l., Pamphilioidea, Cephidae, Siricoidea, Xiphydriidae, Orussidae and Apocrita, and also to Mesozoic Gigasiricidae, Myrmiciidae, Daohugoidae, Sepulcidae, Anaxyelidae, Paroryssidae and Ephialtitidae. Based on the presence of a synapomorphic transscutal articulation, a plesiomorphic unconcealed mesopostnotum and autapomorphic hindwing venation (cu‐a distinctly basal to fork between M and Cu), C. articulata sp.n. most likely forms the sister group of Xiphydriidae + Euhymenoptera. This would place it well within Unicalcarida, i.e. the clade in which the transition from endophytic to parasitoid lifestyle evolved.     

The oldest bee fly in the French Paleocene (Diptera: Bombyliidae)

Paleolomatia menatensis gen. and sp. n., oldest Bombyliidae sensu stricto, is described from the Paleocene of Menat (France). The new genus, based on the wing venation, is attributed to the rather ‘derived’ subfamily Lomatiinae, strongly supporting a Late Cretaceous age for the diversification of the pollinator bee flies, in relation with the floristic changes and the angiosperm radiation that occurred at the same time.     

Paramesopsocidae,a new Mesozoic psocid family (Insecta: Psocodea “Psocoptera”: Psocomorpha)

Paramesopsocus lu n. gen., n. sp. and Paramesopsocus adibi n. sp. are respectively described from the Early Cretaceous amber of Lebanon and from the Late Jurassic limestone of Karatau (Kazakhstan). They are placed within the suborder Psocomorpha, and in the Mesozoic extinct family Paramesopsocidae n. fam. A cladistic phylogeny for Psocomorpha is given including our fossil taxa. The discovery of these new taxa demonstrates the necessity of a deep cladistic redefinition of the currently admitted major subdivisions of this suborder.     

Two new species of Archaeohelorus (Hymenoptera,Proctotrupoidea, Heloridae) from the Middle Jurassic of China

Two new fossil species, Archaeohelorus polyneurus sp. n. and A. tensus sp. n., assigned to the genus Archaeohelorus Shih, Feng & Ren, 2011 of Heloridae (Hymenoptera), are reported from the late Middle Jurassic, Jiulongshan Formation of Inner Mongolia, China. Based on the well-preserved forewings and hind wings of these specimens, the diagnosis of the Archaeohelorus is emended: forewing 2cu-a intersecting Cu and Rs+M at the same point or postfurcal, and hind wing may have tubular veins C, Sc+R, R, Rs, M+Cu, M and Cu distinct, or simplified venation. The new findings also elucidate the evolutionary trend of forewing and hind wing venation and body size for the Heloridae from the late Middle Jurassic to now.     

Traits and evolution of wing venation pattern in paraneopteran insects

Two different patterns of wing venation are currently supposed to be present in each of the three orders of Paraneoptera. This is unlikely compared with the situation in other insects where only one pattern exists per order. We propose for all Paraneoptera a new and unique interpretation of wing venation pattern, assuming that the convex cubitus anterior gets fused with the common stem of median and radial veins at or very near to wing base, after separation from concave cubitus posterior, and re‐emerges more distally from R + M stem. Thereafter, the vein between concave cubitus posterior and CuA is a specialized crossvein called “cua‐cup,” proximally concave and distally convex. We show that despite some variations, that is, cua‐cup can vary from absent to hypertrophic; CuA can re‐emerge together with M or not, or even completely disappear, this new interpretation explains all situations among all fossil and recent paraneopteran lineages. We propose that the characters “CuA fused in a common stem with R and M”and “presence of specialized crossvein cua‐cup” are venation apomorphies that support the monophyly of the Paraneoptera. In the light of these characters, we reinterpret several Palaeozoic and early Mesozoic fossils that were ascribed to Paraneoptera, and confirm the attribution of several to this superorder as well as possible attribution of Zygopsocidae (Zygopsocus permianus Tillyard, 1935) as oldest Psocodea. We discuss the situation in extinct Hypoperlida and Miomoptera, suggesting that both orders could well be polyphyletic, with taxa related to Archaeorthoptera, Paraneoptera, or even Holometabola. The Carboniferous Protoprosbolidae is resurrected and retransferred into the Paraneoptera. The genus Lithoscytina is restored. The miomopteran Eodelopterum priscum Schmidt, 1962 is newly revised and considered as a fern pinnule. In addition, the new paraneopteran Bruayaphis oudardi gen. nov. et sp. nov. is described fromthe Upper Carboniferous of France (see Supporting Information). J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

Mesozoic diversity of relict subfamily Kempyninae (Neuroptera: Osmylidae)

A new genus and two new species of Kempyninae (Osmylidae), Ponomarenkius excellens gen. et sp. nov. and Arbusella magna sp. nov., are described from the Middle-Upper Jurassic of Daohugou (China). The finding demonstrates that extant kempynins, a typical Gondwanan group, are overshadowed by their Mesozoic relatives in taxonomic diversity and variety of wing venation. Striped patterns of both newly described species are an example of cryptic and disruptive coloration, possibly pinnate leaf mimesis. Eyespot patterns of Arbusella magna sp. nov. can be explained as a result of sexual selection. The abundance and distribution of the Mesozoic kempynins are discussed.

http://zoobank.org/urn:lsid:zoobank.org:pub:CF33DEB4-0A69-407D-B1B2-1EC6537E4095     

Pronemouridae fam. nov. (Insecta: Plecoptera), the stem group of nemouridae and notonemouridae,from the Middle Jurassic of Inner Mongolia,China

Abstract: A new family, Pronemouridae fam. nov., with a new genus, Pronemoura gen. nov., and five new species of fossil stoneflies are described from Daohugou village (Middle Jurassic), Inner Mongolia, China: P. shii sp. nov., P. angustithorax sp. nov., P. longialata sp. nov., P. minuta sp. nov. and P. peculiaris sp. nov. Wing venation of pronemourinids presents some or considerable variety in the fore and hind wings of the same or different individuals. Pronemourinidae, retaining plesiomorphic characters (short, multisegmented cerci and CuA with forks), should be treated as the stem group of Nemouridae and Notonemouridae. Based on fossil data, we propose a model in which the extant plecopterid distribution began in the Early Cretaceous. Nemouridae and Notonemouridae must have occurred in Asia and begun to migrate from here by at least the Early Cretaceous.     

New braconid wasps from French Cretaceous amber (Hymenoptera, Braconidae): synonymization with Eoichneumonidae and implications for the phylogeny of Ichneumonoidea

Two new fossils of Braconidae are described from Albian-Cenomanian amber of south-western France, Protorhyssalodes arnaudi gen. n., sp. n., and Aenigmabracon capdoliensis gen. n., sp. n. The former appears superficially similar to the type genus and species of the extinct sub-family Protorhyssalinae, from Turonian New Jersey amber specimens, and the latter both to Protorhyssalus and to members of the extinct family Eoichneumonidae. However, both new taxa display unique combinations of wing venation characters making confident assignment to sub-family impossible. Indeed, they are the first braconids ever known to possess both vein 2-CU and a distinct trace of vein 2-1A on hindwing. The new fossil taxa are incorporated into a morphological analysis of extinct and extant ichneumonoids. As a result of the analyses we synonymize the Eoichneumonidae with the Braconidae.     

Zoraptera wing structures: evidence for new genera and relationship with the blattoid orders (Insecta: Blattoneoptera)

Abstract. The order Zoraptera has traditionally been thought to contain only one family (Zorotypidae) and one genus ( Zorotypus Silvestri). An analysis of known zorapteran wings shows that the wing venation contains character sets indicative of the existence of seven genera: Zorotypus, Brazilozoros gen.n., Centrozoros gen.n., Floridazoros gen.n., Latinozoros gen.n., Meridozoros gen.n. and Usazoros gen.n. The wing venation of Meridozoros leleupi (Weidner) from the Galapagos Islands, Ecuador and Venezuela is described here for the first time.
The major wing structures show that Zoraptera belong to the blattoid lineage. Head and abdomen characters indicate that Zoraptera probably diverged from the Blattoneoptera stock early, almost certainly before the (Protelytroptera + Dermaptera) line, and much before the (Isoptera + (Blattodea + Mantodea)) line. A homologized wing vein system is proposed for the Isoptera.
The homologized wing vein system is based on the hypothesis that the Pterygota originated with the development of protowings, which then diverged through separate but characteristic adaptations for flapping flight. Therefore the basic wing venation pattern is monophyletic, but the changes in wing musculature, articulation and basic braces between main veins are different in the major (super-ordinal) pterygote lineages (Pleconeoptera, Orthoneoptera, Blattoneoptera, Hemineoptera and Endoneoptera). Thus, these characters provide an extremely useful, almost untapped, source of data for higher-level systematics. Both higher-level and lower-level wing characters have been applied here to the phylogeny of Zoraptera and are discussed.     

Moltenia rieki n. gen., n. sp. (Hymenoptera: Xyelidae?, a tentative sawfly from the Molteno Formation (Upper Triassic), South Africa

The fore wing of the new genus and speciesMoltenia rieki of the order Hymenoptera, tentatively belonging to the family Xyelidae, is described from the Molteno Formation (Upper Triassic) of South Africa. This fossil is a representative from the early evolutionary history of this order and the oldest Hymenoptera from central Gondwanaland. However, the plesiomorphic characters in its wing venation do not allow a clear assignment to a recent subgroup of the Hymenoptera, although generally it can be assumed that the Xyelidae had a much higher diversity during the Triassic than today.     

The Gomphaeschninae (Odonata: Aeshnidae): new fossil genus, reconstructed phylogeny, and geographical history

ABSTRACT. Three new species of fossil Anisoptera (dragonflies) are reported from Tiffanian (Late Palaeocene) sediments of the Paskapoo Formation, near Red Deer, Alberta, Canada. The three species are all assigned to the new genus Alloaeschna , which is classified in the subfamily Gomphaeschninae.
Phylogenetic analysis of gomphaeschnine wing venation suggests that the subfamily is paraphyletic, with derived gomphaeschnines such as Boyeria and Linaeschna more closely related to the Brachytroninae and Aeshninae than are other gomphaeschnines. One of the most primitive gomphaeschnines, and the oldest fossil form, is the Jurassic Morbaeschna. Parallelism, convergence and reversal in the evolution of features of the wing venation is common.
The new genus Alloaeschna is at about the same evolutionary grade as the old-world genus Oligoaeschna and the new-world genus Gomphaeschna , and is one of the more primitive known members of the Gomphaeschninae. The new species are the first recorded Palaeocene gomphaeschnines and the oldest known gomphaeschnines from the Americas.
Several extant genera have fossil representatives on continents different from those on which they now survive, suggesting repeated crossing of land bridges and/or widespread ancestral species prior to the separation of the continents, and subsequent extinction leading to present relict distributions.     

An early winged crown group stick insect from the Early Eocene amber of France (Insecta,Phasmatodea)

The new stick insect family Gallophasmatidae, based on Gallophasma longipalpis gen. et sp.n. , from the Earliest Eocene French amber has a pattern of tegmina venation typical of Archaeorthoptera, also present in at least some Mesozoic ‘Phasmatodea’. On the other hand, Gallophasma displays in its body anatomy some apomorphies of the extant Euphasmatodea, e.g. fusion of metatergum and abdominal tergum 1, correlated with the reduction of abdominal sternum 1 to lateral triangular sclerites. A unique autapomorphy of Gallophasma is the presence of annulated and apparently multi‐segmented or pseudo‐segmented cerci; all other Phasmatodea have one‐segmented cerci. The venation of the tegmina of Gallophasma differs from that of extant winged Phasmatodea in the plesiomorphic absence of a knob‐like dorsal eversion. This and other differences in the wing venation between extant and extinct Phasmatodea might have been caused by the loss of wings at some point in the evolutionary history of the order and their secondary gain in a subclade of the extant phasmids.     

Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology

Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species ?Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re‐interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem‐Mantodea, as are some other ‘protorthopterous’ taxa. This hypothesis implies that the total‐group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous ‘Protorthoptera’ are stem‐representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock‐crawlers), suggesting a survivorship of several main Pterygota lineages at the end‐Permian extinction event higher than previously expected. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 79–113.     

Phylogeny,landmark analysis and the use of wing venation to study the evolution of social wasps (Hymenoptera: Vespidae: Vespinae)

Wing venation provides useful characters with which to classify extant and fossil insects. Recently, quantification of its shape using landmarks has increased the potential of wing venation to distinguish taxa. However, the use of wing landmarks in phylogenetic analyses remains largely unexplored. Here, we tested landmark analysis under parsimony (LAUP) to include wing shape data in a phylogenetic analysis of hornets and yellow jackets. Using 68 morphological characters, nine genes and wing landmarks, we produced the first total‐evidence phylogeny of Vespinae. We also tested the influence of LAUP parameters using simulated landmarks. Our data confirmed that optimization parameters, alignment method, landmark number and, under low optimization parameters, the initial orientation of aligned shapes can influence LAUP results. Furthermore, single landmark configurations never accurately reflected the topology used for data simulation, but results were significantly close when compared to random topologies. Thus, wing landmark configurations were unreliable phylogenetic characters when treated independently, but provided some useful insights when combined with other data. Our phylogeny corroborated the monophyly of most groups proposed on the basis of morphology and showed the fossil Palaeovespa is distantly related to extant genera. Unstable relationships among genera suggest that rapid radiations occurred in the early history of the Vespinae.     

Taxonomic review and phylogenetic inference elucidate the evolutionary history of Mesozoic Procercopidae,with new data from the Cretaceous Jehol Biota of NE China (Hemiptera,Cicadomorpha)

The Mesozoic family Procercopidae is widely treated as the ancient group of Cercopoidea and a transitional unit to recent lineages, but its evolution and diversity are vague due to fragmentary fossil record and confusing taxonomic history. Herein, an extensive taxonomic review of Procercopidae is presented and some new fossils are reported from the Lower Cretaceous Yixian Formation of NE China. As a result, Chengdecercopis Hong, 1983 is transferred from Procercopidae to Sinoalidae; Procercopis longipennis Becker-Migdisova, 1962 and P shawanensis Zhang, Wang and Zhang, 2003 are transferred to Procercopina Martynov, 1937, resulting in Procercopina longipennis (Becker-Migdisova, 1962), comb. n. and P shawanensis (Zhang, Wang and Zhang, 2003), comb. n.; Luanpingia senjituensis Hong, 1984 is transferred to Stellularis Chen, Yao and Ren, 2015, leading to Stellulari senjituensis (Hong, 1984), comb. n.; Anthoscytina macula Hu, Yao and Ren, 2014 is transferred to Sinocercopis Hong, 1982, and Sunoscytinopteris (Scytinopteridae) and Cathaycixius (Cixiidae) are treated as junior homonym names of Sinocercopis, leading to Sinocercopis macula (Hu, Yao and Ren, 2014), comb. n., S lushangfenensis (Hong, 1984), comb. n., S pustulosis (Ren, 1995), comb. n., and S trinervis (Ren, 1995), comb. n. Additionally, two new species are erected: Stellularis bineuris Chen and Wang, sp. n. and S minutus Chen and Wang, sp. n. Our cladistic analysis based on wing (tegmen and hind wing) characteristics recovers the high-level relationships within Cercopoidea: Sinoalidae + (Procercopidae + (Cercopionidae + modern cercopoids)). Within the family Procercopidae, the cladistic analysis reveals that the Middle to Late Jurassic Titanocercopis and Jurocercopis and the Cretaceous Cretocercopis occupy the basal position, and a gradual change in wing venation can be recognized from the Early Jurassic Procercopis and Procercopina to the Jurassic Anthoscytina, and then to the Cretaceous Stellularis and Sinocercopis. The two Cretaceous genera, sharing wing traits with extant cercopoids, likely represent transitional forms between Procercopidae and recent Cercopoidea; however, they are very similar to their Jurassic relatives in body structures, suggesting it is applicable to attribute them to Procercopidae. Furthermore, our analysis suggests that the extinction of Procercopidae and the origin and early diversification of modern Cercopoidea approximately coincided with the rise and explosive radiation of angiosperms in the late Early Cretaceous and onwards.     

Primary feather lengths may not be important for inferring the flight styles of Mesozoic birds

Chan, N.R., Dyke, G.J. & Benton, M.J. 2013: Primary feather lengths may not be important for inferring the flight styles of Mesozoic birds. Lethaia, Vol. 46, pp. 146–152. Although many Mesozoic fossil birds have been found with primary feathers preserved, these structures have rarely been included in morphometric analyses. This is surprising because the flight feathers of modern birds can contribute approximately 50% of the total wing length, and so it would be assumed that their inclusion or exclusion would modify functional interpretations. Here we show, contrary to earlier work, that this may not be the case. Using forelimb measurements and primary feather lengths from Mesozoic birds, we constructed morphospaces for different clades, which we then compared with morphospaces constructed for extant taxa classified according to flight mode. Consistent with older work, our results indicate that among extant birds some functional flight groups can be distinguished on the basis of their body sizes and that variation in the relative proportions of the wing elements is conservative. Mesozoic birds, on the other hand, show variable proportions of wing bones, with primary feather length contribution to the wing reduced in the earlier diverging groups. We show that the diverse Mesozoic avian clade Enantiornithes overlaps substantially with extant taxa in both size and limb element proportions, confirming previous morphometric results based on skeletal elements alone. However, these measurements cannot be used to distinguish flight modes in extant birds, and so cannot be used to infer flight mode in fossil forms. Our analyses suggest that more data from fossil birds, combined with accurate functional determination of the flight styles of living forms is required if we are to be able to predict the flight modes of extinct birds. □Birds, flight, morphospace, Mesozoic, wing.