New species of Palaeobolbomyia Kovalev and Ussatchovia Kovalev (Diptera, Brachycera, Rhagionidae) from the Callovian-Oxfordian (Jurassic) Daohugou biota of China: Biostratigraphic and paleoecologic implications

Three new species of fossil snipe flies (Insecta, Diptera, Rhagionidae) from the Daohugou Formation of Chifeng City in Nei Monggol Autonomous Region (= Inner Mongolia), northeastern China are described as Palaeobolbomyia sinica nov. sp., Ussatchovia robusta nov. sp. and Ussatchovia gracilenta nov. sp. They represent the first records of the genera Palaeobolbomyia Kovalev and Ussatchovia Kovalev in China and have implications for stratigraphic correlation of the formations in which they are found. Although the precise age of the Daohugou Formation is currently unresolved, it is most likely Callovian-Oxfordian (latest Middle Jurassic-earliest Late Jurassic) rather than early Middle Jurassic (late Aalenian-early Bajocian) or Early Cretaceous.     

Yanliaoa, an extinct genus of Cupressaceae s. l. from the Middle Jurassic,northeastern China

Yanliaoa is a common fossil in the Middle Jurassic of western Liaoning, eastern Inner Mongolia and northern Hebei Province, China. It is an important element of the Yanliao biota. The genus was established by Pan in 1977 for fossil plants from the Middle Jurassic Haifanggou Formation in Xiasanjiaochengzi, western Liaoning Province, and in present paper, the genus Yanliaoa is studied based on new material. Pan never designated a type specimen and his fossil material cannot be located. We designate a type specimen here for Yanliaoa, so that the genus name Yanliaoa remains valid. Yanliaoa sinensis Pan emend. Tan et al., is found in the same locality and formation as the lost specimens, Y. sinensis of Pan, 1977. Yanliaoa daohugouensis n. sp., a new species with epidermal anatomy, is from the Middle Jurassic Daohugou, Inner Mongolia. A holotype is also selected from the new material for this new species. Characters of the leafy shoots and ovulate cones of Yanliaoa are emended. The epidermal anatomy of this genus is described for the first time. Compared with other extant and extinct species of Cupressaceae s. l., the current species can be distinguished from any known species both by the leafy shoot characters and its epidermal anatomy. It further indicates that Yanliaoa is an extinct and endemic conifer found in the Middle Jurassic of northeastern China.     

The first find of chrysomelids (Insecta: Coleopetra: Chrysomeloidea) from Callovian-Oxfordian Daohugou biota of China

Tarsomegamerus mesozoicus nov. gen., nov. sp., an almost complete specimen of leaf beetle from the Callovian-Oxfordian of Daohugou Formation in Inner Mongolia, China is described. Tarsomegamerus mesozoicus is assigned to the Protoscelinae (Chrysomelidae, Coleoptera). This new leaf beetle represents the first record of chrysomeloids from Mesozoic of China and it extends the geographical distribution of Protoscelinae from central Asia into eastern Asia. This find provides new fossil material for studying the origin and evolution of Chrysomeloidea and by making a stratigraphic correlation of the correlative non-marine sedimentary rocks. Although the age of the Daohugou Formation is debatable it is most likely Callovian-Oxfordian (latest Middle Jurassic-earliest Late Jurassic) rather than early Middle Jurassic (late Aalenian-early Bajocian) or Early Cretaceous.     

A New Fossil of Necrotauliidae (Insecta: Trichoptera) from the Jiulongshan Formation of China and Its Taxonomic Significance

Background

Acisarcuatus variradius gen. et sp. nov., an extinct new species representing a new genus, is described from the Middle Jurassic Jiulongshan Formation in Daohugou Village, Inner Mongolia, China.

Methodology/Principal Findings

In this paper, we revised the diagnosis of Necrotauliidae Handlirsch, 1906. One new genus and species of Necrotauliidae is described. An analysis based on the fossil morphological characters clarified the taxonomic status of the new taxa.

Conclusions/Significance

New fossil evidence supports the viewpoint that the family Necrotauliidae belongs to the Integripalpia.     

An exquisitely preserved harvestman (Arthropoda,Arachnida, Opiliones) from the Middle Jurassic of China

Sclerosomatids constitute the largest family of the arachnid order Opiliones, and one of the two families commonly found in the temperate regions of the northern Hemisphere. Harvestmen have a sparse fossil record in the Mesozoic, with only two species known from the Jurassic, one of them poorly preserved and none with precise phylogenetic placement. Here we report a new fossil, Mesobunus dunlopi sp. nov., from the Middle Jurassic (approx. 165 Mya) of Daohugou, Inner Mongolia, China. The new species is related to another genus of the same formation, but the preservation quality and details of the penis and pedipalps allow us to place them in the extant sclerosomatid subfamilies Gagrellinae or Leiobuninae. The first recognisable fossil in this subfamily highlights morphological stasis over ca. 165 Mya and the finding of this species along with lacustrine insects suggests a life mode similar to that of some modern sclerosomatids, and a possible connection between morphological and ecological stasis.     

Two new species of Trichoceridae from the Middle Jurassic Jiulongshan Formation of Inner Mongolia,China

Two new species, Eotrichocera (Archaeotrichocera) longensis sp. n. and Eotrichocera (Archaeotrichocera) amabilis sp. n. of Trichoceridae are described based on a combination of the following characters: Sc ending proximad of the forking of R₂, shape of d cell and A₂ rather short and bending sharply toward posterior margin. These fossil specimens were collected from the Middle Jurassic Jiulongshan Formation of Daohugou in Inner Mongolia, China.     

The earliest fossil record of Panorpidae (Mecoptera) from the Middle Jurassic of China

The early history of Panorpidae (Mecoptera) is poorly known due to sparse fossil records. Up to date, only nine fossil species have been described, all from the Paleogene, except the Early Cretaceous Solusipanorpa gibbidorsa Lin, 1980. However, we suggest S. gibbidorsa is too incompletely preserved to permit even family classification. A new genus with two new species, Jurassipanorpa impunctata gen. et sp. n. and Jurassipanorpa sticta sp. n., are described based on four well-preserved specimens from the late Middle Jurassic Jiulongshan Formation of Daohugou, Inner Mongolia, China. These two new species are the earliest fossil records of Panorpidae. The new genus is erected based on a combination of forewing characters: both R₁ and Rs₁ with two branches, 1A reaching posterior margin of wing distad of the forking of Rs from R_1, and no crossveins or only one crossvein between veins of 1A and 2A. In all four specimens, long and robust setae ranging from 0.09 to 0.38 mm in length and pointing anteriorly, are present on anal veins of forewings. The function of these setae is enigmatic.     

First description of axymyiid fossils (Insecta: Diptera: Axymyiidae)

Two species of fossil insects from the Daohugou Formation of Chifeng City in Nei Monggol Autonomous Region, northeastern China are described, and recognized as Psocites pectinatus (Hong, 1983) nov. emend., nov. transl. and P. fossilis nov. sp. They are the oldest representatives of the family Callovian or Axymyiidae, and first described of this family in the Mesozoic. Although early the age of the fly-bearing beds is controversial, it is probably Oxfordian or Kimmeridgian (Late Jurassic) rather than Early Cretaceous or Middle Jurassic.     

Anomopterellidae Restored,with Two New Genera and Its Phylogeny in Evanioidea (Hymenoptera)

Background

Anomopterellidae was originally classified as a family within the Evanioidea, and later lowered to a subfamily, Anomopterellinae, of Praeaulacidae. Up to date, only Rasnitsyn 1975, with four species, was assigned to Anomopterellinae. Due to their special wing venation and their metasomal attachment similar to those known in Evanioidea, the systematic position of Anomopterellinae in Evanioidea has been in contention.

Principal Findings

Here we report a new fossil genus Synaphopterella gen. nov. and six species from the Middle Jurassic of China and transfer Anomopterella stenocera Rasnitsyn, 1975, from Upper Jurassic of Kazakhstan, to Choristopterella gen. nov. We place these three genera in the restored family Anomopterellidae and provide a key to known genera and species.

Conclusions/Significance

Based on new fossil specimens and phylogenetic analyses, Praeaulacidae has the most basal position in Evanioidea and it is justifiable to restore Anomopterellidae Rasnitsyn, 1975 as a full family. Comparing the size of all described anomopterellids from China, Mongolia and Kazakhstan, we conclude that the species from China have larger bodies and forewings. Diversity of the Praeaulacidae and Anomopterellidae in the late Middle Jurassic of Daohugou suggests that Evanioidea appeared at least before the late Middle Jurassic.     

Insect herbivory and plant defense on ginkgoalean and bennettitalean leaves of the Middle Jurassic Daohugou Flora from Northeast China and their paleoclimatic implications

Interactions between terrestrial arthropods and plants play a significant role in terrestrial ecosystems. Research on plant–insect interactions through geologic time provides valuable information for studying insect behavior and plant structure, understanding their coevolution, as well as analyzing climate change. In this paper, we choose fossil ginkgoalean and bennettitalean leaves as the plant hosts to study insect herbivory in the Middle Jurassic Daohugou area. Seven damage types of four functional feeding groups have been identified. Of the four functional feeding groups, margin feeding is the most common, indicating an abundance of insects with chewing mouthparts. Ginkgoalean leaves, probably because of their chemical defense, suffered less severe insect damage than bennettitalean leaves. Physical defense has also been observed in various genera of the bennettitalean leaves. Significantly, leaves of Anomozamites had a shaggy indumentum on the abaxial leaf surface and long stiff hairs along the rachis protecting them from insect herbivory. Our results indicate that the climate in the Middle Jurassic of the Daohugou area was relatively warm and humid. This work contributes to the study of plant–insect coevolution in the Daohugou Biota and provides more proxy data for understanding the Middle Jurassic paleoclimate and paleoenvironment in Daohugou area.     

Jurassic lacewings (Insecta: Neuroptera) of Western Siberia

A lacewing assemblage from the Middle Jurassic of the West Siberian Kubekovo locality, the only know Jurassic lacewing fauna of Russia, is discussed. One new genus and two new species of Osmylidae from Kubekovo, Kubekius multiramosus gen. et sp. nov. and Epiosmylus longus sp. nov. are described. Thaumatomerobius mirabilis Ponomarenko, 1985 is transferred to the family Saucrosmylidae, which has only been recorded in the Middle Jurassic of Daohugou.     

A new genus of Chifengiinae (Orthoptera: Ensifera: Prophalangopsidae) from the Middle Jurassic (Jiulongshan Formation) of Inner Mongolia,China

The Middle Jurassic Ashangopsis daohugouensis, gen. et sp. n., of the prophalangopsid subfamily Chifengiinae is described based on two well-preserved specimens collected from the fossil locality near Daohugou at Ningcheng County, Inner Mongolia, Northeast China. The new genus is characterized by the particular shape of its pronotum, its hind leg, and stridulatory veins that only concern the cubital area but not the anal area, unlike in other Chifengiinae.     

A new species of Plesioblattogryllus Huang, Nel et Petrulevičius (Grylloblattida: Plesioblattogryllidae) from the Middle Jurassic of China

A new species of grylloblattid insects, Plesioblattogryllus minor, sp. nov. (Gryllobattida: Plesioblattogryllidae), is described from the Middle Jurassic Daohugou locality (Inner Mongolia, China). The value of the diagnostic features of the families Plesioblattogryllidae and Blattogryllidae is briefly discussed.     

Tettigarctidae (Insecta: Hemiptera: Cicadoidea) from the Middle Jurassic of Inner Mongolia, China

Three new species of Tettigarctidae from the Middle Jurassic of Daohugou, China are described: Shuraboprosbole daohugouensis nov. sp., S. minuta nov. sp., and S. media nov. sp. The revised diagnostic characters of Shuraboprosbole Becker-Migdisova are provided based on well-preserved whole-bodied fossil tettigarctids. So far, only four definitive species of Tettigarctidae are recorded from the Mesozoic of China. The pronotum concealing major of mesonotum seems to be an autapomorphy of Tettigarctinae. The “cicadellid-like tarsi” found in Cicadellidae and Brazilian Tettigarctidae are probably a parallel evolution.     

Revision and description of water boatmen from the Middle-Upper Jurassic of Northern and Northeastern China (Insecta: Hemiptera: Heteroptera: Corixidae)

Four adult and immature water boatmen from the Middle Jurassic and uppermost Middle Jurassic or lowermost Upper Jurassic Jiulongshan, Haifanggou and Daohugou formations in the northern and northeastern China are described as Yanliaocorixa chinensis (Lin, 1976), Jiulongshanocorixa genuina gen. et sp. nov., and Daohugocorixa vulcanica gen. et sp. nov., referred to the extinct subfamily Velocorixinae, and Karataviella popovi sp. nov., referred to the extant subfamily Diaprepocorinae. All the forms have previously been identified as one species, Yanliaocorixa chinensis. This species, found only in the Haifanggou Formation, is absent from the Jiulongshan Formation and other sites in China. The systematic position of corixids recorded earlier from the Mesozoic of China is reviewed and reassessed. Assemblages of dominant aquatic insects from various Jurassic nonmarine sedimentary basins in northern and northeastern China are described for the first time. They are entirely different between the Jiulongshan, Haifanggou, and Daohugou formations. The first formation is probably older than the other two.     

A remarkable new genus of Procercopidae (Hemiptera: Cercopoidea) from the Middle Jurassic of China

Jurocercopis grandis, a new genus and species belonging to Procercopidae (Hemiptera: Cercopoidea), is described, based on seven specimens from the Middle Jurassic of Daohugou, China. It differs distinctly from other genera as follows: body and wings obviously large; pronotum with posterior margin straight; tegmen with costal margin convex at about basal 0.4 wing length; hind basitarsomere with a row of apical small teeth.     

Snipe flies (Diptera: Rhagionidae) from the Daohugou Formation (Jurassic), Inner Mongolia,and the systematic position of related records in China

Abstract: Four new species referred to three new genera and one known genus of fossil snipe flies (Family Rhagionidae) from the Daohugou Formation of Chifeng City in Inner Mongolia, north‐eastern China, are described as Daohugorhagio elongatus gen. et sp. nov., Parachrysopilus jurassicus gen. et sp. nov., Sinorhagio sinuatus sp. nov. and Trichorhagio gregarius gen. et sp. nov. These taxa represent only a fraction of the rhagionid assemblage in the Daohugou biota. They contribute towards the high diversity and abundance of snipe flies during the latest Middle Jurassic – earliest Late Jurassic. The diagnosis of Sinorhagio K. Zhang, Yang and Ren, 2006 is revised based on the new results. Many genera and species from the Lower Cretaceous of China have been assigned to Rhagionidae. Most of these do not actually belong to this family: specifically Mesorhagiophryne incerta Hong and Wang, 1990, M. robusta Hong and Wang, 1990, Mesostratiomyia laiyangensis Hong and Wang, 1990, Stratiomyopsis robusta Hong and Wang, 1990, Longhuaia orientalis Hong, Wang and Sun, 1992, Basilorhagio venusius Ren, 1995. Pauromyia oresbia Ren, 1998 is transferred to an archisargid genus as Sharasargus oresbius (Ren, 1998) comb. nov.     

A specialized new species of Ashicaulis (Osmundaceae,Filicales) from the Jurassic of Liaoning,NE China

A new species of structurally preserved fern rhizome, Ashicaulis plumites (Osmundaceae, Filicales), is described from the Middle Jurassic Tiaojishan Formation in western Liaoning Province, NE China. The new species is characterized by a peculiar sclerenchyma mass in the petiolar vascular bundle concavity. This sclerenchyma mass varies from a linear-shape to a mushroom-like shape with a remarkable outward protuberance, which distinguishes the present new species from other Ashicaulis species. Such a protuberance is very rare among osmundaceous ferns, and should represent a unique type for sclerenchymatous tissue in the osmundaceous vascular bundle concavity. Recognition of the peculiar structure of this new fossil species enriches anatomical diversity of permineralized osmundaceous ferns, indicating that the family Osmundaceae might have experienced a remarkable diversification during the Middle Jurassic in NE China. The new species show anatomical similarities to Osmunda pluma Miller from the Palaeocene of North America. The occurrence of A. plumites in the Middle Jurassic of China provides a new clue for understanding the evolution of some members of the living subgenus Osmunda.     

Morphology and microstructure of Pellites hamiensis nov. sp., a Middle Jurassic liverwort from northwestern China and its evolutionary significance

Pelliales is a basal lineage of the simple thalloid liverworts. Here, we describe a simple thalloid liverwort from the Middle Jurassic Xishanyao Formation of the Turpan–Hami Basin, Xinjiang Uygur Autonomous Region in northwestern China. The thalloid fossil liverwort is described as a new species: Pellites hamiensis (Pelliaceae, Pelliales). Pellites hamiensis nov. sp. resembles the extant species of Pelliaceae in having ribbon-like segments, conspicuous costae that appear as ridges on the ventral surface, rectangular cells with thickened walls adjacent to the margin, and elongated cells in the middle part of the thallus. The occurrence of P. hamiensis nov. sp. extends the fossil records of Pellites back to the Aalenian-Bajocian or Bajocian (ca. 170 Ma) and suggests that the divergence time of the Pelliaceae is no later than the Middle Jurassic.