(Hemiptera, Sternorrhyncha) from the Middle Jurassic ...

16 downloads 0 Views 2MB Size Report
Apr 12, 2013 - American Museum Novitiates 3361, 1–72. GULLAN, P.J. & MARTIN, J.H., 2003. Sternorrhyncha (Jumping. Plant Lice, Whiteflies, Aphids, and ...
This article was downloaded by: [Capital Normal University] On: 30 May 2013, At: 17:17 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

Alcheringa: An Australasian Journal of Palaeontology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/talc20

Poljanka strigosa, a new species of Protopsyllidiidae (Hemiptera, Sternorrhyncha) from the Middle Jurassic of China a

a

Guang Yang , Yunzhi Yao & Dong Ren

a

a

Key Lab of Insect Evolution and Environmental Changes, Capital Normal University , Beijing , 100048 , PR China Published online: 12 Apr 2013.

To cite this article: Guang Yang , Yunzhi Yao & Dong Ren (2013): Poljanka strigosa, a new species of Protopsyllidiidae (Hemiptera, Sternorrhyncha) from the Middle Jurassic of China, Alcheringa: An Australasian Journal of Palaeontology, 37:1, 125-130 To link to this article: http://dx.doi.org/10.1080/03115518.2012.715325

PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

Poljanka strigosa, a new species of Protopsyllidiidae (Hemiptera, Sternorrhyncha) from the Middle Jurassic of China GUANG YANG, YUNZHI YAO and DONG REN

Downloaded by [Capital Normal University] at 17:17 30 May 2013

YANG, G., YAO, Y.Z. & REN, D., March 2013. Poljanka strigosa, a new species of Protopsyllidiidae (Hemiptera, Sternorrhyncha) from the Middle Jurassic of China. Alcheringa 37 , 125–130. ISSN 0311-5518. A new fossil species, Poljanka strigosa sp. nov., of the extinct family Protopsyllidiidae is described from the Middle Jurassic Jiulongshan Formation of Daohugou Village, Inner Mongolia, China. The new species is characterized by wings bearing long, stiff setae that are evident as stained impressions in the fine sedimentary rock. Comparison between Protopsyllidiidae and extant psylloids suggests that Protopsyllidiidae is probably closely related to extant psylloids. Guang Yang [[email protected]], Yunzhi Yao [[email protected]] (corresponding author) and Dong Ren [[email protected]], Key Lab of Insect Evolution and Environmental Changes, Capital Normal University, Beijing 100048, PR China. Received 14.5.2012; revised 2.7.2012; accepted 5.7.2012. Key words: Hemiptera, Sternorrhyncha, Protopsyllidiidae, psylloids, Middle Jurassic, Daohugou biota, Jiulongshan Formation.

EXTANT psylloids, or jumping plant-lice, are an important group of insects comprising about 4000 described species throughout the world and 1000 described species in China (Li 2011). Extant adult jumping plant lice are small insects, ranging in length from 0.2 to 10 mm (Gullan & Martin 2003, Santana & Burckhardt 2007, Ouvrard et al. 2010). Protopsyllidiidae is an extinct family of insects in the Hemiptera suborder Sternorrhyncha, currently comprising 31 described genera and 57 species (Tillyard 1917, Martynov 1937, Handlirsch 1938, Bekker-Migdisova 1959, 1960, Carpenter 1992, Ansorge 1996). This family is known from the Late Permian to Late Cretaceous (Davis 1942, Evans 1943a, b, 1956, Bekker-Migdisova 1985, Klimaszewski 1995, Grimaldi 2003, Grimaldi & Engel 2005). Klimaszewski (1964) argued that extant psylloids developed from fossil Protopsyllidiidae, as first proposed by Hennig (1969). Recently, we discovered two well-preserved fossil specimens attributed to the family, including one part and counterpart, from the Middle Jurassic Jiulongshan Formation at Daohugou Village, Ningcheng County, Inner Mongolia, China. Based on these specimens, we describe a new species assigned to Poljanka Klimaszewski, 1995. The Daohugou

Ó 2013 Association of Australasian Palaeontologists http://dx.doi.org/10.1080/03115518.2012.715325

fossiliferous beds are Middle Jurassic in age (ca 165 Ma; Ren et al. 2002, Wang et al. 2005, Gao & Ren 2006, Wang et al. 2009, Ren et al. 2010, Yang et al. 2011).

Material and methods This study is based on two fossil specimens (one is represented by part and counterpart), housed in the fossil insect collection of the Key Lab of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China. The specimens were examined both without alcohol and under alcohol using a Nikon SMZ800 dissecting microscope. Photos were taken using a Nikon Digital Camera DXM1200C. Line drawings were prepared with Adobe Photoshop CS2 graphic software. The wing venation nomenclature used in this paper is based on the scheme of Ossiannilsson (1992). Body length was measured along the midline from the anterior margin of the head to the tip of the abdomen. Body width was measured at the maximum width of the body. All measurements are provided in millimetres (mm).

Systematic palaeontology Order HEMIPTERA Linnaeus, 1758 Suborder STERNORRHYNCHA Amyot & Serville, 1843 Family PROTOPSYLLIDIIDAE Carpenter, 1931

GUANG YANG et al.

ALCHERINGA

Downloaded by [Capital Normal University] at 17:17 30 May 2013

126

Fig. 1. Poljanka strigosa sp. nov. A, B, Photographs of the holotype without alcohol; C, D, photographs of the holotype under alcohol; A, C, CNU-PSY-NN2011021p; B, D, CNU-PSY-NN2011021c. Scale bar ¼ 1 mm.

ALCHERINGA

HEMIPTERAN FROM THE MIDDLE JURASSIC OF CHINA

127

Downloaded by [Capital Normal University] at 17:17 30 May 2013

Fig. 2. Poljanka strigosa sp. nov. Photographs. Paratype, CNU-PSYNN2011003. Scale bar ¼ 1 mm.

Poljanka Klimaszewski, 1995 Type species. Cicadellopsis shurabensis Bekker-Migdisova, 1985. Poljanka strigosa Yang, Yao & Ren sp. nov. (Figs 1–4) Materials. Holotype, CNU-PSY-NN2011021PC (part and counterpart), a well-preserved specimen, with body mainly preserved and a rather clear forewing; paratype, CNU-PSY-NN2011003. Type locality, formation and age. Daohugou Village, Ningcheng County, Inner Mongolia, China; Jiulongshan Formation, Middle Jurassic. Etymology. The name of the new species refers to the stiff setae on the thorax and wings, which derives from the Latin strigosa (‘wire-haired’). Diagnosis. Scapus twice as thick as pedicelli, pedicelli 1.33 times thicker than flagellomeres, apical flagellomere swollen; femora about twice as thick as corresponding tibiae, hind tibiae 1.41 times longer than fore tibiae and 1.35 times longer than middle tibiae, first tarsomere of fore legs 1.07 times longer than second, basitarsi of middle legs 1.31 times length of distal segments, first tarsomere of hind legs 1.81 times longer than second; R1 of the fore wings without an apical fork, vein M þ Cu1 about 1.07 times longer than vein R, M1þ2 4.45 times longer than M, and M3þ4 4.35 times longer than M, cell m1 2.47 times longer than cell cu1, Cu1a obviously longer than Cu1b. Description. Head. The head is 1.27 times as wide as long; the vertex bears a few stiff setae; the eyes are very large and subround; the clypeus is approximately triangular; the anteclypeus and postclypeus are convex; and the rostrum is stout (Fig. 4C). The antennae are long and comprise 10 segments. The

scapus are globular and twice as thick as the pedicelli; the pedicelli are cylindrical and thicker than the flagellomeres. The flagellomeres are thin, with long and fine setae; the apical flagellomere is longest, swollen and divided into many subsegments (Fig. 4D). Thorax. The pronotum and mesothorax bear stiff setae. The junction of the pronotum and mesopraescutum bears a row of long and stiff setae. The mesothorax is large and the mesoscutum is especially well developed. Legs. The trochanter is large and approximately quadrate. All femora are stout, slightly enlarged distally, about twice as thick as the corresponding tibiae. Relatively sparse fine setae are present on the femora, except the middle femora, which bear five stiff setae. The tibiae are covered with relatively long and dense setae arranged in rows, and two stiff setae are visible at the distal apex of the middle and hind tibiae. The fore tibiae are about 1.58 times as long as the corresponding femora. The first tarsomere of the fore tarsi is slightly longer than the second. The middle tibiae are slightly longer than the fore tibiae and 1.64 times as long as the corresponding femora. The tarsi are 0.45 times the length of the corresponding tibiae. The hind legs are distinctly longer than the fore and middle legs. The tibiae are 1.75 times as long as the femora; the tarsi are distinctly longer than the fore and middle tarsi, and 0.48 times the length of the tibiae; two claws are present at the apex of the tarsi. Wings. The fore wing is elongate (Fig. 3B), narrow at the base, rounded at apex, with a marginal fringe of short, fine, dense setae. Moreover, the costal margin bears short or long stiff setae, and all veins support long, stiff, sharp setae on the upper surface; a small ring is evident around each setal base inserted on a vein (Fig. 4A). Vein R is very short; R1 is long, sinuous, without an apical fork. Vein Rs branches off R very proximally and is arched to the costal margin

128

GUANG YANG et. al

Downloaded by [Capital Normal University] at 17:17 30 May 2013

of the wing at the apex. Vein Rs is slightly longer than R1. Vein Cu1 is connected to M creating the common part M þ Cu1, and M þ Cu1 is connected to R forming the common stem R þ M þ Cu1. The M þ Cu1 vein stem is slightly longer than vein R. Vein M is short, straight, dividing into M1þ2 and M3þ4. Vein M1þ2 is arched to the wing apical angle and is slightly longer than M3þ4. The branches of vein M are 4.35–4.45 times longer than M. Vein Cu1 is very long, about 2.75 times longer than vein M, and forked into a very long, curved Cu1a and short, straight Cu1b. Vein Cu1a is 0.78 times the

ALCHERINGA length of Cu1 and 7.53 times the length of Cu1b. Cell m1 is obviously longer than cell cu1. Vein Cu2 is long and approximately parallel to Cu1. Vein A1þ2 is long and straight, except for its sinuous base. The anal break is visible at the apex of veins Cu2 and A1þ2 (Fig. 4B). Vein A3 is short. Veins A1þ2 and A3 are merged into a point. The hind wing is slightly shorter than the fore wing. Abdomen. Segments II–VIII are clearly visible. Short and fine setae are present on segments VII and VIII. Dimensions (in mm). Holotype, CNU-PSY-NN201 1021PC: Head length 0.30, width 0.37; antennal segment lengths I–X: 0.10, 0.06, 0.27, 0.30, 0.17, 0.20, 0.14, 0.10, 0.27, 0.47; pronotum length 0.33; mesopraescutum length 0.24; length of fore leg: femur 0.81, tibia 1.28, tarsomeres I–II: 0.29, 0.27; length of middle leg: femur 0.81, tibia 1.33, tarsomeres I–II: 0.34, 0.26; length of hind leg: femur 1.03, tibia 1.80, tarsomeres I–II: 0.56, 0.31; claw 0.06; forewing length 4.27, width 1.91; vein lengths: R 0.46, R1 2.28, Rs 2.98, M þ Cu1 0.49, M 0.60, M1þ2 2.67, M3þ4 2.61, Cu1 1.65, Cu1a 1.28, Cu1b 0.17; cell m1 2.64, cell cu1 1.07. Paratype, CNU-PSY-NN2011003: head length 0.32, width 0.38; antennal segment lengths I–X: 0.08, 0.06, 0.24, 0.28, 0.15, 0.20, 0.09, 0.10, 0.18, 0.26; length of middle leg: femur 0.69, tibia 1.12, tarsomeres I–II: 0.20, 0.19; length of hind leg: femur 0.80, tibia 1.88, tarsomeres I: 0.44; forewing length 3.88, width 1.41; vein lengths: R 0.28, R1 2.36, Rs 2.97, M þ Cu1 0.29, M 0.59, M1þ2 2.59, M3þ4 2.62, Cu1 1.50, Cu1a 1.65, Cu1b 0.24; cell m1 2.56, cell cu1 0.99.

Fig. 3. Poljanka strigosa sp. nov. Line drawings of holotype, CNUPSY-NN2011021. A, Body with wings; B, fore wing. Scale bars ¼ 1 mm.

Remarks. The new species is assigned to Poljanka based on the following features: apical flagellomere swollen; first tarsomere longer than second; R1 without an apical fork; vein M þ Cu1 slightly longer than vein R; Vein Cu1 forked into long Cu1a and short Cu1b.

Fig. 4. Poljanka strigosa sp. nov. Photographs, CNU-PSY-NN2011021. A, Setae on fore wing; B, anal break at apex of veins Cu2 and A1 þ 2; C, convex anteclypeus and postclypeus and stout rostrum; D, apical flagellomere of right antenna with many subsegments. Scale bar ¼ 1 mm.

1.07

3.99

2.94

?

?



?

þ

þ

þ –



– 1.31 þ

4.00

3.10 1.45 –

1985)

1985)

?

?

?

?

1.45 – 1985)

?

? 1985)

?

?

? ?

6.50

4.00–4.20





? –

– 2.80–3.20 1.47 þ 1985)

P. strigosa sp. nov. P. hirsuta Yang, Yao & Ren, 2012 P. kukalovae (Bekker-Migdisova, P. sharovi (Bekker-Migdisova, P. shurabensis (Bekker-Migdisova, P. ventriculosa (Bekker-Migdisova, P. sp. 1 (Bekker-Migdisova, P. sp. 2 (Bekker-Migdisova,

Species

1985)

?

þ

?



1.27 þ

3.00

þ

2.18 ? –



þ

4.45 2.39 þ þ 1.75 2.10 þ – þ –

þ –

4.27 4.94

þ þ

– –

Forewing M1 þ 2/M ratio Bifurcation of M before bifurcation of CuA Vein Cu1a and Cu1b straight Veins with setae Forewing length (mm) Ratio of hind tibiae to hind femora Vertex with setae Apical flagellomere divided into subsegments Apical flagellomere longer than flagellomere 7

Downloaded by [Capital Normal University] at 17:17 30 May 2013

Table 1. Distribution of characters differentating the new species and other species of Poljanka. ‘ þ ’ means in accordance with the corresponding character; ‘7’ means discordance with the corresponding character; ‘?’ means no fossil record.

HEMIPTERAN FROM THE MIDDLE JURASSIC OF CHINA

ALCHERINGA

129

In addition, Poljanka differs from Cicadellopsis Martynov, 1937 in the R1 lacking an apical fork (vs R1 forked into R1a and R1b) and from Carpenterella Bekker-Migdisova, 1968 and Karatavopsyllidium Bekker-Migdisova, 1968 in the Cu1 having two branches (vs Cu1 without fork). The new species can be distinguished from other species of Poljanka based on nine flagellomere, setal, leg ratio and wing characters (Table 1).

Discussion The relationships between Protopsyllidiidae and extant psylloids remain controversial. Klimaszewski (1964) and Bekker-Migdisova (1985) argued that extant psylloids developed from fossil Protopsyllidiidae. However, Grimaldi (2003) and Ouvrard et al. (2010) agreed that Protopsyllidiidae is basal to all extant Sternorrhyncha, and argued it is not closely related to extant psylloids. We find that important characters of extant psylloids are also found in some representatives of Protopsyllidiidae, e.g., antenna filiform, ten-segmented, bearing two terminal setae, in Postopsyllidium rebeccae Grimaldi et al. 2002, Grimaldi, 2003; Rs long, the commom stem of R þ M þ Cu1, vein M forked into M1þ2 and M3þ4, and vein Cu1 forked into Cu1a and Cu1b, in Protopsyllidium austral Tillyard, 1926 and Triassotha analis Evans, 1956; and the pterostigma present between the costal vein and R1, in Clavopsyllidium minutum Davis, 1942 and Protopsyllidium lynae Riek, 1976. Based on these shared characters, we hypothesize that Protopsyllidiidae were closely related to extant psylloids (Shcherbakov & Popov 2002). However, this hypothesis needs to be tested by future phylogenetic studies of Protopsyllidiidae and extant psylloids.

Acknowledgements We are grateful to Zhang Weiting, Gao Yan and Dong Qiuping in the Capital Normal University Key Lab, for their careful and critical comments on the manuscript. This research was funded by grants from the National Basic Research Program of China (973 Program; No. 2012CB821900); National Natural Science Foundation of China (No. 31172143, 31071964); the PHR Project of Beijing Municipal Commission of Education (No. 201107120); Fok Ying-Tong Education Foundation for Young Teachers in the Higher Education Institutions of China (No. 131021), and The General Program of Science and Technology Development Project of Beijing Municipal Education Commission of China (No. KM2012 10028016). References AMYOT, C.J.B. & SERVILLE, J.G., 1843. Deuxie`me partie. Homopte`res. Homoptera Latr. Histoire Naturelle des insectes. Hemipte`res 1843, 1–676.

Downloaded by [Capital Normal University] at 17:17 30 May 2013

130

GUANG YANG et al.

ANSORGE, J., 1996. Insekten aus dem Oberen Lias von Grimmen (Vorpommern, Norddeutschland). Neue Palaontologische Abhandlungen 2, 1–13. BEKKER-MIGDISOVA, E.E., 1959. Some new representatives of a group of Sternorrhyncha from the Permian and Mesozoic of the USSR. Materialy k osnovam paleontologii 3, 104–116. (in Russian) BEKKER-MIGDISOVA, E.E., 1960. New Permian Homoptera from European USSR. Trudy Paleontologicheskogo Instituta Akadamii Nauk SSSR 76, 1–112. (in Russian) BEKKER-MIGDISOVA, E.E., 1968. Protopsyllidiids and their morphology (Homoptera, Protopsyllidiidae). Jurassic Insects of Karatau, 87–98. (in Russian) BEKKER-MIGDISOVA, E.E., 1985. Fossil Psyllomorpha insects. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR 206, 1–93. (in Russian) CARPENTER, F.M., 1931. The Lower Permian insects of Kansas. Part 4. The order Hemiptera, and additions to the Paleodictyoptera and Protohymenoptera. American Journal of Science 22, 113– 130. CARPENTER, F.M., 1992. Treatise on Invertebrate Paleontology, Part R, Arthropoda 4, volume 3: Superclass Hexapoda. The Geological Society of America, Boulder, Colorado, and University of Kansas, Kansas, Lawrence, Kansas, xxii–655 pp. DAVIS, C., 1942. Hemiptera and Copeognatha from the Upper Permian of New South Wales. Proceedings of the Linnean Society of New South Wales 67, 111–122. EVANS, J.W., 1943a. Upper Permian Homoptera from New South Wales. Records of the Australian Museum 21, 180–198. EVANS, J.W., 1943b. Two interesting Upper Permian homoptera from New South Wales. Transactions of the Royal Society of South Australia 67, 7–9. EVANS, J.W., 1956. Palaeozoic and Mesozoic Hemiptera (insect). Australian Journal of Zoology 4, 165–258. GAO, K.Q. & REN, D., 2006. Radiometric dating of ignimbrite from Inner Mongolia provides no indication of a post-Middle Jurassic age for the Daohugou Beds. Acta Geologica Sinica 80, 42–45. GRIMALDI, D.A., 2003. First amber fossils of the extinct family Protopsyllidiidae, and their phylogenetic significance among Hemiptera. Insect Systematics and Evolution 34, 329–344. GRIMALDI, D.A. & ENGEL, M.S., 2005. Evolution of the Insects. Cambridge University Press, Cambridge, 722 pp. GRIMALDI, D.A., ENGEL, M.S. & NASCIMBENE, P.C., 2002. Fossiliferous Cretaceous amber from Myanmar (Burma): Its rediscovery, biotic diversity, and paleontological significance. American Museum Novitiates 3361, 1–72. GULLAN, P.J. & MARTIN, J.H., 2003. Sternorrhyncha (Jumping Plant Lice, Whiteflies, Aphids, and Scale Insects). In VINCENT, H.R. & CARDE, R.T., eds, Encyclopedia of Insects. Academic Press, Amsterdam, 1079–1089. HANDLIRSCH, A., 1938. Neue Untersuchungen u¨ber die fossilen Insekten mit Erga¨nzungen und Nachtra¨gen sowie Ausblicken auf phylogenetische, palaeogeographische und allgemein biologische Probleme. II Teil. Annalen des Naturhistorischen Museums in Wien 49, 1–240. HENNIG, W., 1969. Die Stammesgeschichte der Insecten. Senckenbergische Naturforschende Gesellschaft, Verlag von Waldemar Kramer, Frankfurt am Main, 436 pp. KLIMASZEWSKI, S.M., 1964. Studia nad układem systematycznym podrzedu Psyllodea. Annales Zoologici 22, 81–138.

ALCHERINGA KLIMASZEWSKI, S.M., 1995. Supplement to the knowledge of Protopsyllidiidae (Homoptera, Psyllomorpha). Prace naukowe Uniwersytetu S´laskiego w Katowicach 1477, Acta Biologica Silesiana 27 (44), 33–43. LI, F.S., 2011. Psyllidomorpha of China (Insecta: Hemiptera). Science Press, Beijing 1976 pp. (in Chinese, keys in English) LINNAEUS, C., 1758. Systema naturae per regna tria naturae, secundum classes, ordinus, genera, species, cum characteribus, differentiis, synonymis, locis. 10th ed., Vol. 1. Holmiae Salvii, 824 pp. MARTYNOV, A.B., 1937. Liassic insects of Shurab and Kizilkya Mongolia. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR 7, 1–232. OSSIANNILSSON, F., 1992. The Psylloidea (Homoptera) of Fennoscandia and Denmark. Fauna Entomologica Scandinavica 26, E.J. Brill, Leiden (Netherlands), 346 pp. OUVRARD, D., BURCKHARDT, D., AZAR, D. & GRIMALDI, D.A., 2010. Non-jumping plant-lice in Cretaceous amber (Hemiptera: Sternorrhyncha: Psylloidea). Systematic Entomology 35, 172– 180. REN, D., GAO, K.Q., GUO, Z.G., JI, S., TAN, J.J. & SONG, Z., 2002. Stratigraphic division of the Jurassic in the Daohugou area, Ningcheng, Inner Mongolia. Geological Bulletin of China 21, 584–591. (in Chinese with English Abstract) REN, D., SHIH, C.K., GAO, T.P., YAO, Y.Z. & ZHAO, Y.Y., 2010. Silent Stories—Insect Fossil Treasures from Dinosaur Era of The Northeastern China. Science Press, Beijing 322 pp. RIEK, E.F., 1976. New Upper Permian insects from Natal, South Africa. Annals of the Natal Museum 22, 755–790. SANTANA, D.L.Q. & BURCKHARDT, D., 2007. Introduced Eucalyptus psyllids in Brazil. Journal of Forest Research 12, 337–344. SHCHERBAKOV, D.E. & POPOV. YU.A., 2002. Superorder Cimicidea Laicharting, 1781 Order Hemiptera Linne´, 1758. The Bugs, Cicadas, Plantlice, Scale Insects, etc. (¼ Cimicida Laicharting, 1781, ¼ Homoptera Leach, 1815 þ Heteroptera Latreille, 1810). In History of Insects. RASNITSYN, A.P. & QUICKE, D.L.J., eds, Kluwer Academic Publishers, Dordrecht, pp. 143–157. TILLYARD, R.J., 1917. Permian and Triasic insects from New South Wales in the collection of Mr. John Mitchell. Proceedings of the Linnean Society of New South Wales 42, 720–756. TILLYARD, R.J., 1926. Upper Permian insects of New South Wales. Part 1. Introduction and the order Hemiptera. Proceedings of the Linnean Society of New South Wales 51, 1–30. WANG, X.L., ZHOU, Z.H., HE, H.Y., JIN, F., WANG, Y.Q., ZHANG, J.Y., WANG, Y., XU, X. & ZHANG, F.C., 2005. Stratigraphy and age of the Daohugou Bed in Ningcheng, Inner Mongolia. Chinese Science Bulletin 50, 2369–2376. WANG, Y.J., LIU, Z.Q. & REN, D., 2009. A new fossil lacewing genus and species from the Middle Jurassic of Inner Mongolia, China (Neuroptera: Osmylidae). Acta Palaeontologica Polonica 54, 557–560. YANG, Q., MAKARKIN, V. & REN, D., 2011. Two interesting new genera of Kalligrammatidae (Neuroptera) from the Middle Jurassic of Daohugou, China. Zootaxa 2873, 60–68. YANG, G., YAO, Y.Z. & REN, D., 2012. A new species of Protopsyllidiidae (Hemiptera, Sternorrhyncha) from the Middle Jurassic of China. Zootaxa 3274, 36–42.