Vol. 77 No. 2 ACTA GEOLOGICA SINICA June 2003 141

Kalligrammatid Lacewings from the Upper Jurassic Daohugou Formation in Inner Mongolia, China

ZHANG Junfeng Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, Jiangsu 21 0008, E-mail: [email protected]

Abstract A new species, referable to a new genus, is erected, and named the Sinokalligramma jurassicum gen. et sp. nov. It is the second finding of kalligrammatids in the Daohugou Formation. The origin and migration of the family Kalligrammatidae are discussed. The geological age and stratigraphic correlation of the Daohugou and Karabastau Formations are briefly reviewed and reassessed.

Key words: kalligrammatid lacewings, new taxa, age, origin and migration, Jurassic, Inner Mongolia, China

1 Introduction animal and botanic fossils to be of the known Barremian Yixian Formation (Wang et al., 2000; Wang, 2000). Later, Kalligrammatids belonging to the family they classified the rocks as the Early Cretaceous basal Kalligrammatidae within Neuroptera, Insecta are extinct, parts of the Yixian Formation or Dabeigou Formation, an specialized and large insects that lived during the underlying stratigraphic unit of the Yixian Formation Mesozoic. Up to date, twenty-one species, referable to (Wang et al., 2002). Others argued that they are of the Late eleven genera, are recognized as kalligrammatids Aalenian or Early Bajocian Jiulongshan Formation @en throughout eastern Asia, central Asia and western Europe and Oswald, 2002), or Upper Jurassic (Ji and Yuan, 2002). (Scudder, 1886; Walther, 1904; Handlirsch, 1906, 1919; On the basis of an analysis of new biostratigraphic data, Cockerell, 1928; Martynova, 1947; Panfilov, 1968, 1980; Zhang (2002) believed that fossil entomofaunas in the Ponomarenko, 1984, 1992; Janembowski, 1984, 2001; Daohugou and Karabastau Formations in Karatau, Whalley, 1988; Carpenter, 1992; Lambkin, 1994; Ren and Kazakhstan can be correlated, and they are Guo, 1996; Ren and Oswald, 2002). Some of them are contemporaneous or nearly so, belonging to the Oxfordian reassessed, and do not belong to the family or Kimmeridgian in age. Hence, a new stratigraphic unit, Kalligrammatidae. Some taxa are based on badly the Daohugou Formation, was set up. The insect preserved specimens, and their systematic position is assemblage was named the Daohugou Biota or pre-Jehol uncertain at familial, or even ordinal level (Zhang and Biota, which is earlier in age than the Jehol Biota beyond Zhang, 2003). The undoubted representatives of doubt. kalligrammatids appeared only in the Late Jurassic and It should be pointed out that the age of the Yixian Early Cretaceous, and it is uncertain that they can be Formation has been the focus of much discussion among extended into the undoubted Middle Jurassic. Heretofore, geologists and paleontologists. Some researchers regarded five species referable to three genera of kalligrammatids it to be Late Jurassic, or Late Jurassic-Early Cretaceous, or have been recorded respectively from the Yixian Early Cretaceous (Zhang, 1999; Ji. 2002; Ji et al.. 2003; Formation at Huangbanjiegou in Chaomidianzi of Beipiao. Rougier et al.. 2003). This author thought that it is to be Jiaoning Province, China (Ren and Guo, 1996) and the latest Late Jurassic-earliest Early Cretaceous based on new Daohugou Formation near Daohugou village in Shantou materials of insect fossils from Yixian Formation (Zhang, Town of the Ningcheng Area, Chifeng, Inner Mongolia, 1999; Zhang and Zhang, 2001). Despite various China (Ren and Oswald, 2002). viewpoints mentioned above, the insect assemblages in the New fossils belonging to kalligrammatids were recently Yixian and Dabeigou Formations should belong recovered from the Daohugou Formation. They are of a doubtlessly to the Jehol Biota, and younger in age than new species, referable to a new genus, within that of the Daohugou Formation, because those Kalligrammatidae. The geological age and stratigraphic researchers who regarded the Yixian Formation to be Late correlation of the Daohugou Formation are, at the present Jurassic also held that they must not be older than the time, controversial. Some researchers considered these Tithonion (Sun et al., 1998; Lo et al., 1999; Jiang et al., freshwater lacustrine sedimentary rocks that bear rich 2000, Ji, 2002; etc.). 142 Late Jurassic Kalligramrnatid from Inner Mongolia, China In the earliest articles the age of the Karabastau are most likely coeval and equivalent to the Oxfordian or Formation was given as the lower Dogger of the Middle Kimmeridgian. Jurassic, but, for several dozens years, was regarded as It is a complex and comprehensive problem to make a Upper Jurassic (Rohdendorf, 1962, 1968) or stratigraphic correlation in non-marine sedimentary rocks Kimmeridgian (Hedg, 1981) or and identify their geological ages. To carry out a study Oxfordian-Kimmeridgianof the Upper Jurassic (Zherikhin based on biostratigraphic data of various animal and and Gratshev. 1993; Mostovski 1996). Recently, however, botanic groups, an all-around investigation, analysis and some Russian paleoentomologists considered the age of deduction would be required, and there is a precondition these non-marine sedimentary rocks to be Middle or in identifying exactly the systematic positions of taxa. The uppermost Middle Jurassic to pre-Tithonian of the Upper more abundant the short-lived groups used, the more Jurassic (Rasnitsyn et al. 1998; Mostovski 1998). But, amrate the results obtained. The conclusion would be not they have not interpreted how the rocks could be Middle reliable if it is based only on very few groups, or one and or uppermost Middle Jurassic. However, many Russian two known genera and species because there are some paleoentomologists thought it to be Upper Jurassic taxa that can be attributed to longish living insects. For (Belayeva et al., 2002). It seems to be in the right that the example, a mayfly, Ephemempsis trisetalis Eichwald, Karabastau Formation is older in age than the pertains to a permanently dominant and geographically Archaeopteryx fauna from the Early T~thonian of widespread species within the Jehol Biota, and existed Solnhofen, Germany, and thus, the suggestion that the age respectively in the Dabeigou, Yixian and Jiufutang of the upper limit of the Karabastau Formation is formations in Liaoning Province, China. It is evident, pre-Tithonian is well founded. But, it is questionable however, that the latter is undoubtedly younger in age than whether it can be extended into the early or middle Middle the first two that include deposits of different ages. or uppermost Middle Jurassic because, contrary to the Likewise, Mesolygaeius laiyangensis Ping remains with a known early or middle Middle Jurassic entomofaunas in semi-aquatic bug that was discovered respectively in the China and Russia, the insect assemblage in the Karabastau Laiyang Formation of Shandong, Lushangshufen Formation possesses distinct differences in the collective (=Lushangfen) Formation of western Beijing, Yixian characteristics. Many special groups have been merely Formation of Liaoning and Shahai Formation of Inner discovered in the latter, such as Prodiplatidae referable to Mongolia, China. It is clearly incorrect that the Yixian and Archidermaptera within Dermaptera, Protoscelinae Shahai Formations are coeval because there is another referable to Chrysomelidae within Coleoptera. intercalary stratigraphic unit, the Jiufutang Formation, Kalligrammatidae within Neuroptera, Mesoraphidiidae between them, and thus the latter must be younger in age within Raphidioptera, Nemestrinidae within Diptera, than the former. The aquatic beetle, Coptoclava longipoda Praeaulacinae referable to Praeaulacidae, and Sphecidae Ping, co-existed in the Dabeigou and Dalazi Formations in within Hymenoptera. These familial or subfamilial groups Hebei and Jilin Provinces, China. But the former is much are hardly existent in the undoubted early and middle older in age than the latter, and cannot be correlated. This Middle Jurassic entomofaunas; and accidentally, very few author deems that there were many aquatic and of the forerunners that belong to one or two of the semi-aquatic insects in the Mesozoic belonging to the above-mentioned groups occurred. Nevertheless, they longish-lived species, and they cannot serve as the proofs usually bloomed in the Late Jurassic-Early Cretaceous that the strata bearing these insects are contemporary. The Jehol Biota, such as Kalligrammatidae, Mesoraphidiidae, evolution velocity of insects is faster than other Nemestrinidae and Sphicidae. Therefore, a invertebrate and botanic groups, therefore, it seems to be biostratigraphic correlation and insect phylogenesis and also unreliable to determine the age and make evolution suggest that the Karabastau Formation was stratigraphic correlation based merely on very few groups deposited later than the early and middle Middle Jurassic or one and two known genera and species. But, there are but earlier than latest Late Jurassicearliest Early exceptions to most vertebrate groups because they usually Cretaceous @re-'Iithonian to Beniasian), viz., during the belong to the short-lived animal. This is the reason why early or middle Late Jurassic (Oxfordian or some researchers (e.g. Jiang et al., 2000) disagree with the Kimmeridgian). There is still an open question whether or deduction proposed by Swisher et al. (1999) that the fauna not the rocks were deposited in the Callovian. This of the lower Yixian Formation is at least 20 Ma younger problem needs to be further investigated. than the Archaepteryx fauna of Solnh6fen. Germany. It is The discovery of almost all these special groups hardly possible that the primitive avifauna and the extinct mentioned above also in the Daohugou Formation family Compsopthidae of Coelumsauria within indicates that the Karabastau and Daohugou Formations Thempoda, Dinosauria shared a very low evolution speed Vol. 77 No. 2 ACIAGEOLOGICA SINICA June 2003 143 or even lay in a logiam during at least 20 Ma. The reason is most likely that the absolute age of the 2 'hxonomy and Description Jurassic-Cretaceous boundary is inaccurate, i.e., the age of the Archuepteryx fauna of Solnhofen, Gennany is much Class Insecta Linnaeus, 1758 younger than 150 Ma, or the dating result of 124.6 Ma Order Neuroptera Linnaeus, 1758 (Swisher et al., 1999) for the age of the Lower Yixian Family Kalligrammatidae Handlirsch, 1906 Formation is doubtful. In the same way, before the Sinokulligramma Zhang gen. nov. discovery of a new representative of anurognathids (Rhamphorhynchchoiodea, Pterosauria) from the Holotype: Sinokalligramma jurassicum Zhang gen. et Daohugou Formation (Wang et al., 2002), the family sp. nov. Anurignathidae appears to be a short-lived group in the Difgnosis: Hindwing short and broad, obtusely Karabastau Formation in Karatau, Kazakhstan, the triangular. Eye-spot well developed. Humeral recurrent Amhaepteryx fauna of Solnhofen, Germany and the Yixian vein (Vr) absent. C clearly arched, with costal area Formation in Liaoning, China. They lived only in the Late moderately broadened, costal veinlets sinuate and forked. Jurassic or, at most, Late Jurassic to earliest Early Sc and R1almost joining near wing apex. R1pectinately Cretaceous. The anurignathid group will become a very forked anteriorly near wing apex. Rs with 7 primary long-lived chronogroup with an interval of about 40 Ma if branches, basal two or three of them forked deeply. MA the age of the Daohugou Formation is regarded as Late dichotomously forked near wing middle. Basal cell (bc) Aalenian to Early Bajocian. On the other hand, it seems to between Mpl and MP2 at base of wing absent. MP well be impossible that the age of the Daohugou Formation is developed, with 4 pectinately elongated branches so young according to the basic characteristics and anteriorly, each of them dichotomously forked for several relationship to other known members of the new times. CuA pectinately forked shallowly, and terminated at anurognathid representative described and discussed in hind margin of wing. CUPwell developed, with a serious Wang et al. (2002). of pectinate branches posteriorly. 1A with a few branches, This paper describes a new species referable to a new its stem running parallel to last branch of CuA for a long genus, which is the second finding of Kalligrammatidae distance. 2A well developed, with its stem running parallel from the Daohugou Biota. It is demonstrated that to 1A for a long distance. Crossveins very numerous over kalligrammatids are not a rare group in the Daohugou entire wing. Formation and, represent an older representative that lived, Distribution and geological age: Inner Mongolia, at least, in the early or middle Late Jurassic. It is not Late Jurassic. certain that it existed in the Middle Jurassic. Consequently, kalligrammatids originated in Asia, probably in Sinokalligrammajurassicum Zhang sp. nov. northeastern China, as an undoubted harbinger of (Pl. I-1,2 and 3) kalligrammatids is recovered from the Haifanggou Formation in Haifeng Town of Beipiao, Liaoning Province. Holotype: Almost complete left hindwing (part and It has been recognized as the earliest member of the genus counterpart), registration No. H99072-I and H99072-XI, Kalligramma (Zhang and Zhang, 2003). Most specialists deposited in the Nanjing Institute of Geology and hold that the Haifanggou Formation is of early or Palaeontology, Chinese Academy of Sciences. early-middle Middle Jurassic age (Lin, 1976; CGRSTLPC, Description: Hindwing 59.7 mm long, width 49.6 1978; Mi et al., 1980; Hong, 1983) although Pan (1981) mm, yellowish brown but basal half of wing darkish regarded it as probably Middle to Late Jurassic. Lately, brown. A strip-shaped marking located near the hind more researchers claimed that the age of this formation is margin of the wing, and nearly parallel to it. Eye-spot questionable, and needs to be restudied (Ji and Yuan, 2002; located beyond the middle of the wing, longitudinally Zhang, 2002a). Kalligrammatids appeared in northeastern suboval (height Lq:er than width), fuscous, with two China during the latest Middle Jurassic-earliest Late rather small, white markings arranging longitudinally in Jurassic times, and radiated rapidly from eastern Asia to the middle of the eye-spot, and two ring-like markings western Europe. During the early or middle Late Jurassic encircling the eye-spot. Sc, R1 and Rs straight, not joining they arrived in central Asia, and by the beginning of the each other near the wing apex, but linking up with much late Late Jurassic (early Tithonian), in western Europe. short crossveins between them. Costal area with broadest They existed up to the Early Cretaceous. located before the midwing, and at least three times as broad as the subcostal area. Rs dichotomously forked, with basal two branches forked deeply, remains shallowly 144 Late Jurassic Kalligrammatid from Inner Mongolia, China zhw forked near the hind margin of the wing. Stem of MA and costal area moderately broadened, Sc not joining RI straight, dichotomously forked for several times. First near wing apex, eye-spot well developed, and main branch of MP (MPI)straight, forked near the base of the longitudinal veins, such as MA, MP, CuA, CUP and A, wing, other branches (MP2) distinctly curved downwards showing specialized characteristics in a manner of beyond midway, branches nearly parallel to each other, furcation and in wing venation. The detailed discussion is each dichotomously forked for several times, and with as follows. So far as the feature that Sc does not join R1 is numerous veinlets near the margin of the wing, its breadth concerned, this new genus is similar to that of (MP area) about three times as broad as MA and CuA Kalligrarnmina from the Karabastau Formation in Karatau areas, respectively. CuA forked shallowly, with stem of Kazakhstan, but may be easily distinguished from the curved downwards beyond midway, and running parallel latter by C distinctly arched and costal area moderately to the stem of MP. Stem of CUPrunning parallel to CuA, broadened where all costal veinlets run sinuately and and forked near the base of the wing, with 6 primary furcately, R1 with several pectinate branches anteriorly branches, running somewhat convergent towards the hind near the wing apex, primary basal branches of Rs forked margin of the wing, each branch dichotomously forked for deeply, MA forked near the middle of wing, and several times near the hind margin of the wing. 1A straight, cmssveins denser over the entire wing. In the wing forked shallowly, with three or four branches, each venation and shape, this new genus shares close dichotomously forked for several times near the hind resemblance to Kalligrammula from Karatau, Kazakhstan margin of the wing. 2A pectinately forked posteriorly, With and Solnhofen in Germany, especially the Karatau species, numerous veinlets, occupied most parts of the hind margin K. karatavica Martynova, 1947 (Pdilov, 1968). It differs of the wing, and obviously more than veinlets of 1A. 3A from the latter in having eye-spot, and special character short, only forked twice. Crossveins extremely dense, that Sc does not meet R1 near the wing apex. sometimes formed erose network, but wanting near the On the other hand, the well known Kalligramma, a margin of the wing. The characters of the hindwing geographically widespread genus in eastern Asia, central venation as shown in P1.1-3 in detail. Asia and western Europe, possesses similarity in wing Locality and horizon: Daohugou Village in Shantou venation to Sinokalligramma gen. nov., but by C straight, Town of the Ningcheng Area, Chifeng, Inner Mongolia, with costal area narrow and subequaling to each other in China; Upper Jurassic (Oxfordian-Kimmeridgian) anterior and posterior in breadth, Sc joining R1 near wing Daohugou Formation. apex, and primary branches of CUP not convergent towards the wing margin. the former can be differentiated 3 Comparison and Discussion from the latter. Lithogramma was based on an incomplete forewing This new species and new genus, Sinokalligramma from the Karabastau Formation. It is different from this jurassicum, may be placed in the family Kalligrammatidae new genus by the identifying features: MA with many by having the following familial diagnoses: hindwing branches and forked near the base of the wing as well remarkably large (nearly 60 mm long), with its costal area Sc linking up with R, near the wing apex. moderately narrower (not distinctly broadened) than that In having narrow costal area that subequals subcostal of representatives in other related families, although a area in breadth, a few primary branches of main little broadened than that of known elements of longitudinal veins, Rs forked apically and crossvein sparse kalligrammatids; MA forked shallowly (at about the over the entire wing, Meioneurites may be distinguishable middle of the wing); MP extensively forked, sending from the Chinese new genus. several main branches anteriorly, and crossveins very Sophogrammu within the Jehol Biota includes three numerous over the entire wing (except areas near the species that are established based on forewings (Ren and margin of the wing). Guo, 1996). It is unlike this new genus by the following Up to date, eight genera are usually regarded as characters: eye-spot wanting, humeral recurrent veins (Vr) kalligrammatids: Angumgramma Ponomarenko, 1984, present, Sc joining RI near the wing apex, a basal cell @c) Kalligramma Walther, 1904, Kalligranunina Panflov, present between MP, and MP2 at the base of the wing. 1980, Kalligramla Handlirsch. 1919, Lithogramma near the wing margin MA, CuA and 1A areas broader than Panfilov, 1968, Meioneurites Handlirsch, 1906, MP, CUPand 2A areas, respectively. Sophogramma Ren et Guo, 1996 and Kallibemembius Ren It may be separated from Kallibemembius, a new genus et Oswald, 2002. This new genus may be separated from recently recovered from the same locality and horizon in all of the known genera within Kalligrammatidae by the the Daohugou Formation, by the triangular, shortened and combination of diagnostic characters: C clearly arched, broadened hindwing, Sc not joining R1 near wing apex, Vol. 77 No. 2 ACTA GEOLOGICA SINICA June 2003 145 primary basal branches of Rs forked deeply, and both CUP der rezenten Formen. Leipzig: Wilhelm Engelmann, 473478, and 2A strongly developed, the latter with a series of 610-612. Handlirsch. A., 1919. Em neue Kalligrammatidae (Newoptera) pectinate branches posteriorly. aus dem Solnhiifen Plattenkalke. Senkenbergiana,1: 61-63. Angarogramma is known from the Middle-Upper Hennig W., 1981. Insect Phylogeny. New York: John Wdey & Jurassic Uda Formation in Transbaikal of Siberia, Russia Sons, 514. (Ponomarenko, 1984). It is based on badly preserved Hong Youchong, 1983. Middle Jurassic Fossil Insects in North specimens of a pair of fore- and hindwings. Its generic China. Beijiug: Geological Publishing House, 223 (in diagnosis seems to be too poorly known to make Chinese with English summary). Jarzembowski, E. A., 1984. Early Cretaceous insects from comparison with this new genus. Until new information southern England. Modern Geology, 9: 71-93. becomes available, the diagnostic characteristics of this Jamembowski, E. A., 2001. Geology and Palaeontology: A new genus are uncertain, for example, whether or not Sc Wealden fossil lacewing. In: Rowlands, M.L. (4.).Tunbridge joining R1 near wing apex, eye-spot absent or present, and Wells and Rusthall Commons, a History and Natural History. the state of furcation of main longitudinal veins and the Tunbridge Wells: Tunbridge Wells Museum and Art Gallary, 48-58. distribution of crossveins. To sum up, this new genus Ji Qiang. 2002. On the Mesozoic Jehol Biota of China. shares a closer similarity in wing venation to that of Geological Review, 48(3): 290-296 (in Chinese with English Kalligramma. meanwhile, they have certain differences in summary). taxonomic features. Hence, it is reasonable to found a new Ji Qiang and Yuan Chongxi, 2002. Discovery of two kinds of genus, and the comparison between species is now protofeathered pterosaurs in the Mesozoic Daohugou Biota in impossible because there is only a single species referable the Ningcheng Region and its stratigraphic and biologic significances. Geological Review, 48(2): 221-224 (in Chinese this genus. to with English summary). Ji Qiang. Ji Shu’an, You Hailu, Zhang Jianping, Zhang Hongbin, Acknowledgements Zhang Nanjun. Yuan Chongxi and Ji Xinxin, 2003. An Early Cretaceous Avialian Bird, Shenzhouraptor sinensis, from The research was supported by the Pilot Project of Western Liaoning, China. Acta Geologicu Sinica (English Edition), 77(1): 21-27. Knowledge Innovation Program of the Chinese Academy Jiang Xingfu, Chen Jinhua, Cao Meizhen and Komatsu, T.. 2000. of Sciences (grant no. KzCX2-114) and the Major Basic A discussion on the age of the feathered dinosaurs-bearing Research Project of the Ministry of Science and beds of Liaoning. China. Acta Palaeontologica Sinica, 39(2): Technology, China (grant no. G2000077700). 307-3 11 (in Chinese with English summary). Lambkin, K. J., 1994. Palparites deichmuelleri Handlirsch from Tithonian Solnhofen Plattenkalk belongs to the Manucript received June 2 1,2002 the Kalligrammatidae (Insects: Newoptera). Pabntologische accepted 14,2002 Nov. Zeitschn?, 68(ln) : 163-166. edited by Wang Wen and Xie Guanglian Lin Qibin, 1976. The Jurassic fossil insects from western Liaoning. Acta Palaeonotologica Sinica, 15( 1): 97-116 (in Chinese with English summary). References Lo Chinghua, Chen Peiji, TSOUTungyu, Sun Shensu and Lee Belayeva, N. V., Blagoderov, V. A., Dmitriev, V. Yu., Eskov. K. Chiyu, 1999. Age of Sinosaumpteryx and Yu., Gorokhov, A. V., Ivanov, V. D., Klug, N. Yu., Kozlov, M. Confuci~sornie~Ar?~Arlaser single-grain and K-Ardating V., Lukashevich, E. D., Mostovski, M. B., Novokshonov, V. G, of the YiXian Formation, NE China. Geochimica, 28(4): Ponomarenko, A. G. Popov, Yu. A.. Pritykina, L. N., 405409. Rasnitsyn, A. P., Shcherbakov, D. E.. Sinitshenkova, N. D., Martynova, 0. M., 1947. Kalligrammidae (Newoptera) from the Jurassic shales of Karatau. Doklady Akademii Nauk Storozhenko, S. Yu., Sukatsheva, I. D., Vishniakova, V. N., SSSR. Vdansv, P. and Zherikhin, V. V., 2002. History of insects. 58(9): 2055-2058 (in Russian). Dordrecht, The Netherlank Kluwer Academic Publishers. Mi Jiarong, Xu Kaizhi, Zhang Chuanbou, Chang Jianping and 517. Yao Peiyi, 1980. The Mesozoic strata of Beipiao Dishict, Liaoning. Journal the Changchun Geological Institute, 4 Carpenter, E M., 1992. Treatise on Invertebrate Paleonrology of Part R Volume 4: Superrlass Hemp&. The 18-37 (in Chinese). Kansas: Mostovski, M. B., 1996. New species of the genus Mesosolva Geological Society of America and the University of Kansas, Hong, 1983 (Diptera, Archisargidae) from the Jurassic of 351-352. Cockerell, T. D. A., 1928. The Jurassic insects of Turkestan. Kazakhstan and Mongolia. Museum of Northern Arizona Psyche, 35(2): 126-130. Bullletin, 60,329-332. CGRSTLPC (Compiling Group for Regional Stratigraphy Table Mostovski, M. B., 1998. A brief review of brachcemus flies of Liaoning Province, China), 1978. Regional Stratigraphy (Diptera, Brachycera) in the Mesozoic, with descriptions of Table of Northeastern China, Volume Liaoning Province. some curious taxa. In: Zherikhin, V. V. (4.).Proceedings of Beijing: Geological Publishing House, 101-145 (in Chinese). the First Palaeoentomological Confemnce Moscow 1998. Handlirsch, A., 1906. Die Fossilen Insekten und die Phylogenie Bratislava: AMBA Projects Publication, 103-1 10. Pan Kuang, 1981. A note on the Mesozoic of Yanliao Region. I46 Late Jurassic Kalligrammatid from Inner Mongolia, China =g Bulletin of the Geological Society Limning Pmvince, China, Wang Yuan, 1999. Cretaceous age for the feathered dinosaurs 1: 26-52 (in Chinese with English summary). of Liaoning, China. Nature, 400: 5841. Panfiiov, D. V.. 1968. Kalligrammatids (Neuroptera: Walther, J., 1904. Die Fauna der Solnhtjfener Plattenbalke Kalligrammatidae) from the Jurassic deposits of Karatau. In: bionomisch betrachter. Denkschnfien &r Medizinisch- Rohndendorf, B. B.(ed.), Jurassic insects of Karatau. Moscow: natunvissenschaftlichen Gesellschafrzu ha.11: 135-214. Nauka, 166-174 (in Russian). Wang Xiaolin, Wang Yuanqing. Zhang Fucheng, Zhang Panflov, D. V.. 1980. New representatives of lacewings Jiangyong, Zhou Zhonghe. Jim Fan, Hu Yaoming. Gu Gang, (Neuroptera) from the Jurassic of Karatau. In: Dolint V. G, and Zhang Haichun, 2000. Vertebrate biostratigraphy of the Panfilov, D. V.. Ponomarenko, A. G, and F’ritykina, L. N. Lower Cretaceous Yian Formation in Lingyuan western (4s.). Mesozoicfossil insects. Kiev: Naukova Dumka, 82-1 11 Liaoning and its neighboring southern Nei Mongol (Inner (in Russian). Mongolia), China. Vertebrata PalAsiatica, 38: 81-99 (in Ponomarenko, A. G. 1984. Neuroptera from the Jurassic in Chinese). eastern Asia. Paleontologicheskij uluml. 3: 64-73 (in Wang Xiaolin. Zhou Zhonghe and Zhang Fucheng. 2002. Russian). Discovery of the feathered pterosaur from Jehol biota. Ponomarenko. A. G, 1992. New lacewings (hsecta. Newoptera) Chinese Science Bulletin, 47(1) : 54-58 (in Chinese). from the Mesozoic of Mongolia. Transactions of the Joint Wang Yuan, 2000. A new salamander (Amphibia: Caudata) hm Russian-Mongolian Palaeontological Expedition, 41: 101-1 11 the Early Cretaceous Jehol Biota. Vertebrata PalAsiatica, 38: (in Russian). 100-103 (in Chinese). Rasnitsn, A. P., Janembowski, A. E., and Ross, A. J., 1998. Whalley, P. E. S.. 1988. Mesozoic Neuroptera and Raphidioptera Wasps (Insecta: Vespida = Hymenoptera) from the Purbeck (Insecta) in Britain. Bulletin of the British Museum (Natural and Wealden (Lower Cretaceous) of southern England and History) Geology, 44( 1): 4543. their biostratigraphical and palaeoenvironmental significance. Zhang Junfeng, 1999. Aeschnidiid nymphs from the Jehol biota Cretaceous Research, 19: 329-390. (latest Jurassic-Early Cretaceous), China, with a discussion of Ren Dong, and Guo Ziguang, 1996. On the new fossil genera and the family Aeschnidiidae (Insecta, Odonata). Cretaceous species of Newoptera (Insects) from the Late Jurassic of Research, 20(6): 8 13-827. northeast China. Acta Zootaxonomica Sinica. 21(4): 461-479. Zhang Junfeng, 2002a. Discovery of Daohugou Biota (pre-Jehol Ren Dong, and Oswald, J. D.. 2002. A new genus of Biota) with a discussion on its geological age. Journal of kalligrammatid lacewings from the Middle Jurassic of China Stratigraphy. 26(3): 173-177. 215 (in Chinese with English (Neuroptera: Kalligrammatidae). Stuttgarter Beitrdge zur summary). Naturkunde Sene B (Geologic und Paldontologie),3 17: 1-8. Zhang Junfeng, 2002b. The most primitive earwigs (lnsecta: Rohdendorf, B. B.. 1962. Fundamentals of paleontology. Dermaptera: Archidermaptera) from Upper Jurassic of Nei Arthropoda: Tracheata and Chelicerata Moscow: Nayka. 560 Monggol Autonomous Region, northern China. Acta (in Russian). MicropalaeontologicaSinica 19(4): 348-362. Rohdendorf, B. B., 1968. Jurassic insects of &ratau. Moscow: Zhang Junfeng, and Bang Haichun, 2001.New findings of Nayka, 252 (in Rassian). larval and adult aeschnidiids (Insecta: Odonata) in the Yixian Rougier, G W., Ji Qiang and Novacek, M. J., 2003. A New Formation, Liaoning Province. China. Cretaceous Research, Symmetrodont Mammal with Fur Impressions from the 22(4): 443450. Mesozoic of China. Acta Geologica Sinica (English Edition), Zhang Junfeng and Zhang Haichun, 2003. Kalligramma 77( 1): 7-14. jurarchegonium sp. nov. (Neuroptera: Kalligrammatidae) from Scudder, S. H., 1886. A review of Mesozoic cockroaches. the Middle Jurassic of northeastern China. Oriental Insects, 37: Memoirs of the Boston Society of Nature History. 3: 439485. 301-308. Sun Ge, Dilcher, D. L., Zheng Shaolin and Zhou Zhekun, 1998. Zherikhin, V. V., and Gratshev, V. G., 1993. Obrieniidae fam. In search of the First Flower: A Jurassic Angiosperm, nov., the Oldest Mesozoic weevils (Coleoplerii. Atrhaefnrcnrs, from Northeast China. Science, 282 Curculionoidea). Paleontology Jorrrnal. 27: 50-69. 16921695. Swisher III, C. C., Wang Yuanqing, Wang Xiaolin. Xu Xing and 147 Late Jurassic Kallierammatid from Inner Mongolia. China Zhane

Plate I

R 3A

3 'CuA Explanation of Plate

Plate I 1. Sinokalligramma jurassicum Zhang gen. et sp. nov. holotype, left hindwing, part, x 1.4, No. H99072-I1 2. holotype, left hindwing, counterpart, x 1.4, No. H99072-I1 3. Sinokalligramma jurassicum Zhang gen. et sp. nov., holotype, left hindwing, drawing after specimen No. H99072-1