Jun-Feng Zhang a & Hai-Chun Zhang a a Nanjing Institute of Geology and Palaeontology, Chinese. Available online: 05 Jan 2012

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1 This article was downloaded by: [Nanjing Institute of Geology and Palaeontology] On: 20 March 2012, At: 00:13 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: Mortimer House, Mortimer Street, London W1T 3JH, UK Oriental Insects Publication details, including instructions for authors and subscription information: Kalligramma Jurarchegonium, sp. nov. (Neuroptera: Kalligrammatidae) from the middle Jurassic of northeastern China Jun-Feng Zhang a & Hai-Chun Zhang a a Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, , P. R. China Available online: 05 Jan 2012 To cite this article: Jun-Feng Zhang & Hai-Chun Zhang (2003): Kalligramma Jurarchegonium, sp. nov. (Neuroptera: Kalligrammatidae) from the middle Jurassic of northeastern China, Oriental Insects, 37:1, To link to this article: PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: 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.

2 Oriental Insects, Vol. 37: , May KALLIGRAMMA JURARCHEGONIUM, SP. NO (NEUROPTERA: KALLIGRAMMATIDAE) FROM THE MIDDLE JURASSIC OF NORTHEASTERN CHINA JUN-FENG ZHANG & HAI-CHUN ZHANG Nanjing Institute ofgeology and Palaeontology, Chinese Academy of Sciences, Nanjing , P. R. China ABSTRACT. A fossil wing of a species from the Haifanggou Formation of Beipiao City in Liaoning Province, northeastern China is described as Kalligramma jurarchegonium, sp. nov. It may be the oldest and the second largest species so far known within the family Kalligrammatidae, an extinct member of the order Neuroptera. Although the age of net-winged insect-bearing beds is controversial, this specimenbelongs probably to the latest Middle Jurassic or earliest Late Jurassic. Key words: Gigantism, fossil insect, Neuroptera, Kalligrammatidae, New Species, Geological Age, China. Introduction The fossil insect deposit within the Haifanggou Formation, near the village of Yujiagou in Haifeng Town, near Beipiao City in Liaoning Province, China has yielded thousands of specimens recovered by Chinese paleontologists (Lin, 1976; Hong, 1983). The kalligrammatid-bearing bed has yielded a large number offossil insects, including the following 12 orders: Ephemeroptera, Odonata, Blattaria, Plecoptera, Orthoptera, Hemiptera, Coleoptera, Neuroptera, Mecoptera, Trichoptera, Diptera and Hymenoptera (Lin, 1976; Hong, 1983). Almost all of these fossil insects are of small or medium size; large insects are exceptionally rare. Recently, however, a large fore wing was found, attributable to the genus Kalligramma Walther, 1904, ofthe Kalligrammatidae (Neuroptera). Most Chinese specialists have thought that the uppermost parts ofthe insect-bearing beds ofthe Haifanggou Formation were ofearly or early-middle Middle Jurassic age (Lin, 1976; CGRSTLPC, 1978; Mi et al., 1980; Hong, 1983; Wang et ai., 1987; Sun & Zheng, 2000) although Pan (1980) regarded it as being Middle to Late Jurassic. The latter interpretation is probably correct because the new species described in this paper shares a remarkable resemblance in wing venation to that ofkalligramma liaoningense Ren & Guo, 1996 (originally Kalligramma liaoningensis) from the uppermost Upper Jurassic or earliest Lower Cretaceous ofthe Yixian formation from Beipiao, China (Zhang, 1999; Zhang & Zhang, 2001) and of Kalligramma haeckeli Walther, 1904 from the Lower Tithonian of Solnhofen in Germany. Based on known biogeographical information, we tend to believe that the stratigraphic range ofthe genus Kalligramma is narrow; the uppermost beds ofthe Haifanggou Formation that have yielded these new net-winged insects thus is most likely

3 302 Oriental Insects Vol. 37 to be uppermost Middle Jurassic or lowermost Upper Jurassic in age, equivalent to the Callovian or Oxfordian interval. Materials and Methods Specimen H99071, a nearly complete right fore wing in shale, is well preserved (Figs. 1, 2), although parts of the opposite side were damaged during collection. Only one preserved impression is present. Description, photograph, and drawings were obtained without application of any liquid on the surface of the holotype. The drawing was made with a camera lucida and the photographwas taken using a stereomicroscope, a Seagull Z-8 equipped with Zeiss camera lens. Systematics Order NEUROPTERA Linnaeus, 1758 Family KALLIGRAMMATIDAE Handlirsch, 1906 Genus KALLIGRAMMA Walther, 1904 Kalligrammajurarchegonium, sp. nov. (Figs. 1,2) Holotype: CHINA: Liaoning Province: Beipiao City: Haifeng Town: Yujiagou, Coli. I. Zhang (Zhang Coli.). Specimen H99071, A right fore wing housed in Nanjing Institute ofgeology and Palaeontology, Chinese Academy ofsciences (Nanjing, Jiangsu Province, China). Type locality: Haifanggou Formation outcrop, Village of Yujiagou in Haifeng Town, Beipiao City, Liaoning Province, China. Stratum typicum: Specimen discovered in lacustrine shales from the uppermost Haifanggou Formation of uppermost Middle Jurassic or lowest Upper Jurassic age, corresponding to an age of 156 to 169 million years. Etymology: The "jur" refers to "Jurassic", and "archegonium" to "initial". Description: A right fore wing, yellowish brown, nearly triangular, parts of wing covered with irregular markings. Wing 105 mm long, 69.2 mm wide, with proximal and two distal margins abraded, nearly rounded. Vein Sc with 78 branches, among them several basic branches simple and others dichotomously forked. Vein Rs+RI with numerous branches that run towards wing margin. Present are 86 short crossveins between veins Rs and R I. Vein Rs with 12 primary branches, each running rectilinearly and forking late with an exception of the 6th which forks somewhat more basally. Vein MA forking prior to midsection of wing, provided with few but straight branches. Vein MP extensively branched near wing margin, divided into six dichotomous pectinate branches anteriorly, with the 2nd vein bifurcated several times. Vein Cu with short stem, and dichotomous branches. Vein CuA with many branches and area subtended by CuA widened terminally. Vein CuP branched minimally compared with vein CuA, with few branches, and area subtended by CuP in wing margin narrower than that ofcua. Fork ofvein 1A close to fork

4 2003 Zhang & Zhang: On fossil Neuroptera 303 of vein CuP. Vein 1A with five branches. Vein 2A with many pectinated dichotomous branches posteriorly, each re-forked several times, and the area encompassed by vein 2A adjacent the wing margin distinctly widened. Vein 3A poorly preserved. Crossveins very numerous, covering the entire wing with exception ofwing margins. Wing with a subcircular eye-spot, situated beyond midpoint ofwing. Wing venation detailed in figure 1. Discussion This new species, Kalligramma jurarchegonium, may be placed in the family Kalligrammatidae by having the following familial diagnoses: fore wing remarkably large, with its costal area ofmoderate width (not distinctly widened); vein Sc joining R near wing apex; MA branched only near wing margins; MP extensively branched, sending several main branches anteriorly and crossveins very numerous over entire wing. The Kalligrammatidae is a specialized group that lived during the late Mesozoic. Nineteen species, referable to ten genera, are recognized as kalligrammatids: Angarogramma incertum Ponomarenko, 1984 (originally A. incerta), Besobrasovia latissima Cockerell, 1928, Kalligramma f1exuosum Panfilov, 1968, K. haeckeli Walther, 1904 (Handlirsch, 1906), K. liaoningense Ren & Guo, 1996, K mu/tinerve Panfilov, 1968, K. sharovi Panfilov, 1968, K. turutanovae Martynova, 1947, Kalligrammina areolata Panfilov, 1980, Kalligrammu/a karatavica Martynova, 1947, K. senckenbergiana Handlirsch, 1919, Lithogramma ocu/atum Panfilov, 1968, Meioneurites schlosseri Handlirsch, 1906, M vil/osus Panfilov, 1968, Palparites deichmuelleri Handlirsch, 1906 (Lambkin, 1994), Paractinophlebia curtisii (Scudder, 1886) (Whalley, 1988), Sophogramma eucallum Ren & Guo, 1996 (originally S. eucalla), S. papilionaceum Ren & Guo, 1996 (originally S. papilionacea) and S. plecophlebium Ren & Guo, 1996 (originally S. p/ecoph/ebia). Of these Kalligramma haeckeli, Kalligrammu/a senckenbergiana, and Meioneurites schlosseri are known from the Lower Tithonian (corresponding to approximately 150 million years) of Solnhofen in Germany. Ka/ligramma turutanovae, K. multinerve, K. f1exuosum, K. sharovi, Ka/ligrammu/a karatavica, Meioneurites villosus, Lithogramma oculatum and Kalligrammina areo/ata were recovered from the Karabastau Formation at Karatau in Kazakhstan. The age of the bed that yielded the Karatau kalligrammatids is debatable. It could be Kimmeridgian (corresponding to approximately 150 to 156 million years age) (Hennig, 1981) or Oxfordian to Kimmeridgian (corresponding to approximately 150 to 163 million years age) (Zherikhin & Gratshev, 1993; Mostovski, 1996). The suggestion that the age ofthese fossiliferous rocks is Middle or uppermost Middle Jurassic to pre-tithonian (corresponding to approximately 150 to 169 or 150 to 188 million years age respectively) (Rasnitsyn, Jarzembowski & Ross, 1998; Mostovski, 1998) is not accepted here. The time span represented by the Karabastau Formation is too long to define this unit, and it is difficult to make a biostratigraphical correlation to other fossil sites based on this information. In addition, there are two known insect assemblages from the Haifanggou and Uda Formations in China and Russia, respectively, which are assigned from Middle or uppermost Middle Jurassic to earliest Late Jurassic age. Both deposits share close similarities in overall entomofaunal biofacies, and taxa differ sharply from those ofthe Kara-

5 304 Oriental Insects Vol. 37 bastau Formation, apparently exhibiting more primitive characters within their respective lineages. Meanwhile, the Chinese species of Kalligramma liaoningense, Sophogramma papilionaceum, S.plecophlebium and S. eucallum were found from the Yixian Formation in Beipiao of Liaoning, China. The age of the fossiliferous strata has been the focus of much controversy among Chinese and other geologists and paleontologists. Some researchers regarded this formation to be Late Jurassic age (corresponding to approximately 145 million years age) (Ji & Ji, 1996; Ji et al. 1998; Sun et ai., 1998; Lo et al., 1999a, 1999b; Jiang et ai., 2000) but others have argued that it is ofearly Cretaceous age (corresponding to approximately 124 million years age) (Swisher et al., 1998, 1999; Xu et al., 1999; Wang et ai., 2000; Wang, 2000). Recently, however, the authors maintain that the Yixian Formation is of latest Late Jurassic or earliest Early Cretaceous age based on new biostratigraphical data (Zhang, 1999; Zhang & Zhang, 2001). Alternatively, the poor preservation of Besobrasovia latissima and Palparites deichmuelleri from the lower Tithonian of Solnhofen, Germany, prevents an accurate assessment of systematic position, though these taxa have been regarded as kalligrammatids (Cockerell, 1928; Lambkin, 1994). Paractinophlebia curtisii from Upper Liassic of Alderton, England, was described originally as an isolated wing of a cockroach (Scudder, 1886), but has been transferred provisionally to the Kalligrammatidae by Whalley (1988). This taxon is known only from an isolated, fragmentary wing. It is difficult to see from the author's illustration how it might be related to Kalligrammatidae or even Neuroptera. Its status remains uncertain. Angarogramma incertum from the Uda Formation ofuppermost Middle Jurassic to lowest Upper Jurassic age, from Transbaikalia, Russia, also has an uncertain geological age (Ponomarenko, 1984; Kalugina & Kovalev, 1985). This specimen was based on badly preserved fore- and hind wings and the lack of crossveins disallows identification of kalligrammatid relationships. Material from the Early Cretaceous of England and Mongolia (Jarzembowski, 1984; Ponomarenko, 1992) are referable to the Kalligrammatidae, but, these specimens are too incomplete and badly preserved for more definitive assignment. A kalligrammatid fossil from lower Cretaceous of Spain (Martinez-Delclos & Ruiz de Loizaga, 1993) has not been officially described. Consequently, on the basis ofthe specimens mentioned above, known representatives ofkalligrammatidae occurred only during the Late Jurassic (Oxfordian to Tithonian), and most likely extended to the Early Cretaceous; it is questionable whether kalligrammatids existed during the latest Middle Jurassic. This new species may be assigned to the genus Kalligramma Walther, 1904 because of Rs of fore wing with 12 primary branches and a conspicuous eye-spot. Until now, six species have been placed in the genus Kalligramma, all ofwhich apparently existed during the Late Jurassic (Oxfordian? to Tithonian or Kimmeridgian to Tithonian). Thus the new species, Kalligramma jurarchegonium, from the Haifanggou Formation in Haifeng of Beipiao, Laioning, China, represents one of the oldest definitive members of the Kalligrammatidae. The new species is similar in wing venation to that of K. liaoningense, but can be distinguishable from the latter by greater wing size, veins Rs+R 1 and Rs with more branches, CuA area wider than that of CuP in wing margins, and vein 1A with fewer branches. Alternatively, K. jurarchegonium sp. nov. shares remarkable resemblance in

6 2003 Zhang & Zhang: On fossil Neuroptera 305 wing venation to K. haeckeli. It is separated from the latter by a shorter and wider fore wing, more branches ofvein Sc, including several simple veins at the base, more branches of'rs-r], vein MA forked more basally, and vein Cu with short stem. The Solnhofen species K haeckeli possesses a fore wing 122 mm long, and with an estimated wingspan ofnearly 270 to 280 mm. The fore wing length ofk jurarchegonium sp. nov. is 105 mm and with an estimated wingspan of nearly 230 to 240 mm. It thus represents the second largest known net-winged insect. Based on fossil evidence, Kalligrammatidae is thought to be an extinct group that possessed specialized wing venation. By the Upper Jurassic and possibly into the Lower Cretaceous, there was a marked increase in the diversity ofthis group. The evolutionary lineage of Kalligrammatidae seems to be obscure, but it is likely that they occupied northeastern China during Middle or latest Middle Jurassic to earliest Late Jurassic times. Acknowledgments The research was supported by the Major Basic Research Projects of Ministry of Science and Technology, China (grant no. G ), the Pilot Project ofknowledge Innovation Program, Chinese Academy of Sciences (grant no. KZCX2-114) and the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (grant no ). CARPENTER, F. M., References Treatise on invertebrate paleontology, Part R Arthropoda 4, Volume 3: Superclass Hexapoda, pp The Geological Society ofamerica, Denver, Colorado & The University of Kansas, Lawrence, Kansas. CGRSTLPC (COMPILING GROUP FOR REGIONAL STRATIGRAPHY TABLE OF LIAONING PROVINCE, CHINA), Regional stratigraphy table of northeastern China, Volume Liaoning Province, pp Geological Publishing House, Beijing (in Chinese). COCKERELL, T. D. A., The Jurassic insects of Turkestan. Psyche, 35(2): HANDLlRSCH, A., Die Fossilen Insekten und die Phylogenie der rezenten Formen, pp , pp Wilhelm Engelmann, Leipzig. HANDLlRSCH, A., Eine neue Kalligrammatidae (Neuroptera) aus dem Solnhofen Plattenkalke. Senkenbergiana, 1: HENNIG, w., Insect phylogeny, 514 pp. John Wiley and Sons, Chichester. HONG, Y, Middle Jurassic fossil insects in North China, 223 pp. Geological Publishing House, Beijing (in Chinese, with English summary). JARZEMBOWSKI, E. A., Early Cretaceous insects from southern England. Modem Geology, 9: n, Q., CORRIE, P. C., NORELL, M.A.& n, S., Two feathered dinosaurs from northeastern China. Nature, 393: Jl, Q. & Jl, S., Discovery ofthe earliest bird fossils in China and the origin ofbirds. Chinese Geology, 10: (in Chinese). nang, F., CHEN, J., CAO, M. & KOMATSU, T., A discussion on the age of the feathered dinosaurs-bearing beds of Liaoning, China. Acta Palaeontologica Sinica, 39(2): (in Chinese, with English summary). LAMBKIN, K. J., Palparites deichmuelleri Handlirsch from the Tithonian Solnhofen Plattenkalk belongs to the Kalligrammatidae (Insecta: Neuroptera). Palaontologische Zeitschrift, 68(112) : LIN, Q., The Jurassic fossil insects from western Liaoning. Acta Palaeonotologica Sinica, 15(1): (in Chinese, with English summary). LO, C., CHEN, P., TSOU, T., SUN, S. & LEE, c., 1999a. Age of Sinosauropteryx and Confuciusornis-40Arf 9 Ar laser single-grain and K-Ar dating of the Yixian Formation, NE China. Geochimica, 28(4): (in Chinese, with English summary). LO, C., CHEN, P., TSOU, T., SUN, S. & LEE, c, 1999b. 4OArp 9 Ar laser single-grain and K-Ar dating of the Yixian Formation, NE China. Palaeoworld, 11:

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8 CuA MP, lomm, Fig. 1.Kalligrammajurarchegonium, sp. nov., right fore wing. Camera lucida drawing of specimen H too.:l o Col:l N ::r (J'q N ::r $l:> CfCl o en en Z CD "1 s <'+ CD "1 $l:> C.:l o -:J

9 308 Zhang & Zhang: On fossil Neuroptera Vol. 37 Fig. 2. Kalligrammajurarchegonium, sp. nov., photograph (original size), H99071.