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Two new species of Kalligrammula Handlirsch, 1919 (Insecta, Neuroptera, Kalligrammatidae) from the Jurassic of China and Kazakhstan

Published online by Cambridge University Press:  22 September 2015

Qing Liu,
Aleksandr V. Khramov,
Haichun Zhang and
Edmund A. Jarzembowski
Qing Liu
Affiliation:
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, 210008, PR China 〈qingliu1029@yeah.net〉, 〈hczhang@nigpas.ac.cn〉
Aleksandr V. Khramov
Affiliation:
Borissiak Paleontological Institute of the Russian Academy of Sciences, Moscow 117997, Russia 〈a-hramov@yandex.ru〉
Haichun Zhang
Affiliation:
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, 210008, PR China 〈qingliu1029@yeah.net〉, 〈hczhang@nigpas.ac.cn〉
Edmund A. Jarzembowski
Affiliation:
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, 210008, PR China 〈qingliu1029@yeah.net〉, 〈hczhang@nigpas.ac.cn〉Department of Earth Sciences, The Natural History Museum, London SW7 5BD, UK 〈jarzembowski2@live.co.uk〉
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Abstract

Two new species of Kalligrammula Handlirsch, 1919 (Insecta, Neuroptera, Kalligrammatidae) are described as K. lata n. sp. and K. karatensis n. sp. based on two nearly complete hindwings respectively from the Middle–Upper Jurassic of Daohugou, China and the Upper Jurassic of Karatau, Kazakhstan. Limnogramma Ren, 2003 is considered to be a junior synonym of Kalligrammula based on the hindwing venation. The distribution of kalligrammatid genera is discussed.

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Type
Articles
Information
Journal of Paleontology , Volume 89 , Issue 3 , May 2015, pp. 405 - 410
Copyright
Copyright © 2015, The Paleontological Society 

Introduction

The butterfly-like Kalligrammatidae (Insecta, Neuroptera) is a family of most impressive Mesozoic lacewings. The earliest described kalligrammatids are known from the Middle Jurassic Haifanggou Formation of Yujiagou and the Middle–Upper Jurassic Daohugou beds (Callovian–Oxfordian) of Daohugou in China (Ren and Oswald, Reference Ren and Oswald2002; Zhang and Zhang, Reference Zhang and Zhang2003; Makarkin et al., Reference Makarkin, Ren and Yang2009; Yang et al., Reference Yang, Makarkin and Ren2011; Liu et al., Reference Liu, Zheng, Zhang, Wang, Fang and Zhang2014), and the latest is probably Makarkinia adamsi Martins-Neto, Reference Martins-Neto1992 from the Lower Cretaceous Crato Formation of Brazil (late Aptian–early Albian) (Makarkin et al., Reference Makarkin, Ren and Yang2009).

Seventeen genera can be assigned to Kalligrammatidae: Kalligramma Walther, Reference Walther1904; Meioneurites Handlirsch, Reference Handlirsch1906; Palparites Handlirsch, Reference Handlirsch1906; Kalligrammula Handlirsch, Reference Handlirsch1919; Lithogramma Panfilov, Reference Panfilov1968; Makarkinia Martins-Neto, 1992, Sophogramma Ren et Guo, Reference Ren and Guo1996; Kallihemerobius Ren et Oswald, Reference Ren and Oswald2002; Oregramma Ren, Reference Ren2003; Sinokalligramma Zhang, Reference Zhang2003; Protokalligramma Yang et al., Reference Yang, Makarkin and Ren2011; Apochrysogramma Yang et al., Reference Yang, Makarkin and Ren2011; Huiyingogramma Liu et al., Reference Liu, Zheng, Zhang, Wang, Fang and Zhang2014; Affinigramma Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014; Stelligramma Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014; Abrigramma Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014; and Ithigramma Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014. Two additional monotypic genera have also been attributed to the Kalligrammatidae including Kalligrammina Panfilov, Reference Panfilov1980 from the Upper Jurassic of Karatau, Kazakhstan and Angarogramma Ponomarenko, Reference Ponomarenko1984 from the Middle–Upper Jurassic of the Uda River, Transbaikalia, Russia. The first genus, a miniature kalligrammatid-like lacewing (forewing length 30 mm), however, is close to Ectopogramma Engel et al., Reference Engel, Huang and Lin2011 (Aetheogrammatidae) from Daohugou and they probably belong to the same family (Engel et al., Reference Engel, Huang and Lin2011). The second genus was established based on a poorly preserved forewing with crossveins indiscernible and the size (only 36 mm in length) is much smaller than that of typical kalligrammatids, so the attribution of Angarogramma to Kalligrammatidae is doubtful.

Among the known kalligrammatid genera, 11 genera have a restricted range, while six genera are widespread (at least found to co-occur in two different fossil localities): the Jurassic Meioneurites and Huiyingogramma, the Jurassic–Cretaceous Kalligramma and Kalligrammula, and the Cretaceous Sophogramma and Oregramma (Fig. 1). Meioneurites is present in Karatau (South Kazakhstan, Panfilov, Reference Panfilov1968; Engel, Reference Engel2005) and Solnhofen (Bavaria, Germany; Handlirsch, Reference Handlirsch1906). Huiyingogramma is represented by one species from Daohugou (Ningcheng, Inner Mongolia, China; Liu et al., Reference Liu, Zheng, Zhang, Wang, Fang and Zhang2014) and by an undescribed species from Karatau (a photograph of this specimen is published by Ponomarenko in Rasnitsyn and Quicke, Reference Rasnitsyn and Quicke2002, fig. 254, p. 192, labeled as “Kalligramma turutanovae O. Martynova”). Kalligramma is the most common kalligrammatid lacewing known from at least five localities: Daohugou (Liu et al., Reference Liu, Zheng, Zhang, Wang, Fang and Zhang2014), Yujiagou (Beipiao, Liaoning, China; Zhang and Zhang, Reference Zhang and Zhang2003), Huangbanjigou (Beipiao, Liaoning, China; Ren and Guo, Reference Ren and Guo1996), Karatau (Martynova, Reference Martynova1947; Panfilov, Reference Panfilov1968), and Solnhofen (Walther, Reference Walther1904). K. roycrowsoni Jarzembowski, Reference Jarzembowski2001 from the Lower Cretaceous of the Wealden Supergroup (England, Valanginian, Wadhurst Clay Formation) is represented by a fragmentary forewing with eye-spot and its generic affiliation is questionable. The genus Sophogramma occurs in the English Purbecks (Jepson et al., Reference Jepson, Makarkin and Coram2012), Huangbanjigou (Ren and Guo, Reference Ren and Guo1996; Yang et al., Reference Yang, Zhao and Ren2009), Pingquan (Hebei, China; Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014), and Baissa (Buryatia, Russia; Makarkin, Reference Makarkin2010) and Oregramma is only known from the Yixian Formation of Liutiaogou (Ningcheng, Inner Mongolia, China; Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014) and Huangbanjigou (Ren, Reference Ren2003; Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014).

Figure 1

Distribution of the family Kalligrammatidae.

In this study, we describe two new species assigned to Kalligrammula, K. lata n. sp. and K. karatensis n. sp., based on two nearly complete hindwings respectively from the Middle–Upper Jurassic of Daohugou, China and the Upper Jurassic of Karatau, Kazakhstan. Consequently, the genus Kalligrammula becomes one of the most widespread kalligrammatid genera. It was initially established by Handlirsch (Reference Handlirsch1919) for the single species K. senckenbergiana Handlirsch, Reference Handlirsch1919 from Solnhofen, Germany. The holotype of K. senckenbergiana is an impression of a pair of nearly complete forewings (only one drawn by Handlirsch) and a hindwing. Later, K. karatavica Martynova, Reference Martynova1947 (re-described by Panfilov, Reference Panfilov1968) is from the Upper Jurassic of Karatau and K. atra Ponomarenko, Reference Ponomarenko1992 from the Lower Cretaceous of Anda Khuduk in Mongolia. K. karatavica is represented by a complete but poorly preserved hindwing and greatly differs from K. senckenbergiana in having a hindwing with CuA deeply forked and CuP simple. Unfortunately, the holotype of this species is covered by epoxy resin that has darkened over the decades, so it is difficult to re-study the type specimen. The description of K. atra is based on fragmented material. We exclude both of these species from Kalligrammula and treat them as genera incertae sedis.

Material and methods

The specimen NIGP 156192 described in this paper was collected from the Middle–Upper Jurassic Daohugou Beds at Daohugou Village, Wuhua Township, Ningcheng County, Chifeng City, Inner Mongolia, China. The Daohugou beds, consisting of grey tuff, tuffaceous siltstone and mudstone, are now considered to be one of the most important Mesozoic insect Lagerstätten (Rasnitsyn and Zhang, Reference Rasnitsyn and Zhang2004; Ponomarenko and Ren, Reference Ponomarenko and Ren2010). The fossil insects at this locality are commonly preserved as compressions in grey tuffaceous siltstones, found together with small freshwater spinicaudatan (Wang et al., Reference Wang, Li, Zhang and Fang2009). The fossil-bearing beds at Daohugou were previously considered to be Bathonian to Callovian in age (e.g., Yang et al., Reference Yang, Makarkin, Winterton, Khramov and Ren2012) and to belong to the Jiulongshan Formation (e.g., Ren et al., Reference Ren, Labandeira, Santiago-Blay, Rasnitsyn, Shih, Bashkuev, Logan, Hotton and Dilcher2009). However, recent radiometric dating suggests that the Daohugou beds are of Callovian–Oxfordian age (Liu et al., Reference Liu, Liu and Zhang2006, Reference Liu, Kuang, Jiang, Peng, Xu and Sun2012; Cohen et al., Reference Cohen, Finney and Gibbard2013; Wang et al., Reference Wang, Hu, Zhang, Zheng, He, Deng, Wang, Zhou and Zhu2013) and should be correlated with the Lanqi Formation (or Tiaojishan Formation) (Liu et al., Reference Liu, Liu and Zhang2006, Reference Liu, Kuang, Jiang, Peng, Xu and Sun2012; Chang et al., Reference Chang, Zhang, Renne and Fang2009; Wang et al., Reference Wang, Hu, Zhang, Zheng, He, Deng, Wang, Zhou and Zhu2013). In addition, the aquatic insects Mesobaetis sibirica and Mesoneta antiqua, which were previously considered to be present in the Daohugou beds, the Haifanggou Formation, and the Jiulongshan Formation indicating that these strata are of the same age, proved to be absent from Jurassic deposits in China. There are, in fact, completely different aquatic insect assemblages in these strata: Zhang and Kluge, Reference Zhang and Kluge2007; Zhang, Reference Zhang2010. The specimen PIN No. 2784/1032 was collected from the Upper Jurassic Karabastau Formation, consisting of lacustrine deposits. Several nearby outcrops of the Karabastau Formation, known as the locality Karatau, are situated near the Big Karatau Range in South Kazakhstan. The most characteristic lithology at Karatau is the so-called lithographic shales: highly laminated, fissile claystone that provides an exceptional preservation of fossil compressions, including more than 18,000 insects. The age of the Karabastau Formation is not precisely established. Spore and pollen analysis indicates a late Callovian–Kimmeridgian (Doludenko and Orlovskaya, Reference Doludenko and Orlovskaya1976) or Oxfordian–Kimmeridgian (Doludenko et al., Reference Doludenko, Sakulina and Ponomarenko1990) age for these deposits.

The photographs of specimen NIGP 156192 were taken using a Canon EOS 5D digital camera and the picture of specimen PIN no. 2784/1032 was taken with a Nikon D200 digital camera. The line drawings were prepared on photographs using the image-editing software CorelDRAW 12. The traditional venational terminology of Comstock (Reference Comstock1918) (sensu Wootton, Reference Wootton2003) with the recent interpretation of Oswald (Reference Oswald1993) and Archibald and Makarkin (Reference Archibald and Makarkin2006) is used in this study. Wing vein abbreviations are: C, costa; Sc, subcosta; R, radius; R1, first branch of R; Rs, radial sector; Rs1, most basal branch of Rs; M, media; MA, media anterior; MP, media posterior; Cu, cubitus; CuA, cubitus anterior; CuP, cubitus posterior; 1A–3A, first to third anal veins. Specimen NIGP 156192 is housed at the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, and specimen PIN no. 2784/1032 is at the Borissiak Paleontological Institute of the Russian Academy of Science in Moscow, Russia.

Systematic paleontology

Class Insecta Linnaeus, Reference Linnaeus1758

Order Neuroptera Linnaeus, Reference Linnaeus1758

Family Kalligrammatidae Handlirsch, Reference Handlirsch1906

Genus Kalligrammula Handlirsch, Reference Handlirsch1919

Type species

Kalligrammula senckenbergiana Handlirsch, Reference Handlirsch1919 from the Upper Jurassic (lower Tithonian) of Solnhofen, Germany.

Other species

Kalligrammula mira (Ren, Reference Ren2003) from the Lower Cretaceous Yixian Formation of Huangbanjigou, China; K. hani (Makarkin et al., Reference Makarkin, Ren and Yang2009), K. mongolica (Makarkin et al., Reference Makarkin, Ren and Yang2009), and K. lata n. sp. from the Middle–Upper Jurassic Daohugou beds of Daohugou, China; K. karatensis n. sp. from the Upper Jurassic Karabastau Formation of Karatau, Kazakhstan.

Diagnosis

Deeply bifurcated CuP in hindwings (CuP shallowly forked in other genera, except for Abrigramma Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014), stem of Rs originating from R near the wing base.

Remarks

Among the known kalligrammatid genera, three genera possess a deeply forked CuP vein in the hindwing including Kalligrammula Handlirsch, Reference Handlirsch1919, Limnogramma Ren, Reference Ren2003, and Abrigramma Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014. The stem of Rs originates very distally from R in the hindwing of A. calophleba Yang et al., Reference Yang, Wang, Labandeira, Shih and Ren2014, the type species of Abrigramma, but originates from R near the base of the wing in both Kalligrammula and Limnogramma. The genus Limnogramma is composed of three species, L. mirum Ren, Reference Ren2003, the type species, from the Lower Cretaceous Yixian Formation; and L. hani Makarkin et al., Reference Makarkin, Ren and Yang2009 and L. mongolicum Makarkin et al., Reference Makarkin, Ren and Yang2009 from the Middle–Upper Jurassic of Daohugou. All three species were described only based on hindwings. The distinguishing characteristic of Limnogramma is a “double” CuP vein that forks immediately after its connection with CuA and, additionally, CuP2 is close to 1A. Although the forewing venation of K. senckenbergiana needs re-examination, a deeply forked CuP vein is clearly visible in the hindwing (Fig. 2). Therefore Limnogramma is considered herein as a subjective synonym of Kalligrammula. Furthermore, Oregramma aureolusa Yang et al. Reference Yang, Wang, Labandeira, Shih and Ren2014 also possesses a relatively deeply forked CuP vein in the hindwing. However, the vein CuP bifurcates mid-way and, besides, the vein R1 is strongly branched distally, which makes the species easily distinguished from Kalligrammula spp.

Figure 2

Holotype of Kalligrammula senckenbergiana Handlirsch, Reference Handlirsch1919. (1, 3) part; (2, 4) counterpart. (1, 2) basal part of hindwing respectively from (3) and (4) (indicated in black box). Scale bar is 5 mm for (1, 2) and 30 mm for (3, 4).

Kalligrammula lata new species

Figure 3

Figure 3

Kalligrammula lata n. sp., (1, 2) holotype, a well-preserved left hindwing (NIGP156192a, b); (3) line drawing of holotype NIGP156192a. Scale bar is 20 mm.

Holotype

A well-preserved and nearly complete left hindwing (NIGP 156192a, b), part and counterpart.

Diagnosis

Hindwing triangular. Costal space narrow; costal veinlets slightly sinuous; Rs with 12 primary branches; MA simple; 3A long.

Occurrence

Daohugou Village, Wuhua Township, Ningcheng County, Chifeng City, Inner Mongolia, China; Daohugou beds, Middle–Upper Jurassic.

Description

Left hindwing triangular and broad, 71 mm long and 57 mm wide as preserved. Costal margin slightly curved anteriorly apically. Outer margin slight undulate. Inner and outer margins partially damaged, and costal margin with apex missing. Eye-spot fuscous and subrounded, located slightly distal of wing mid-length, with several small unpigmented spots apically. Costal space relatively narrow. Costal veinlets slightly sinuous, with several distal branches forked and remainder simple; crossveins present between costal veinlets. Subcostal space and R1 space of about equal width, with widely spaced crossveins. Sc and R1 fused apically, extending four-fifths of wing length. Rs with 12 primary branches preserved, running parallel and forked distally several times. MA simple and forked distally. MP with eight branches preserved and dichotomously forked. CuA forked distally. CuP forked very basally, CuP1 with four branches. 1A stem long, very close to CuP and forked distally. 2A long and pectinately branched. 1A stem parallel with 2A stem for long distance. 3A present and long.

Etymology

From the Latin ‘latus’ for broad.

Kalligrammula karatensis new species

Figure 4

Figure 4

Kalligrammula karatensis n. sp., (1) holotype, a nearly complete left hindwing (PIN No. 2784/1032); (2) line drawing of holotype PIN No. 2784/1032. Scale bar is 10 mm.

Holotype

A well-preserved and nearly complete left hindwing (PIN no. 2784/1032).

Diagnosis

Hindwing triangular. Costal space narrow, costal veinlets nearly straight, Rs with six primary branches, MA forked basally, CuA forked distally, CuP forked very basally, 1A forked distally.

Occurrence

Karatau, Kazakhstan; Karabastau Formation, Upper Jurassic.

Description

Left hindwing triangular and broad, 48 mm long and 30 mm wide as preserved. Costal, outer, and inner margins mostly damaged. Eye-spot fuscous and subrounded, located slightly distal of wing mid-length. Dozens of small unpigmented spots located around the eye-spot and along the wing outer margin. Costal space narrow. Costal veinlets relatively straight and simple. Subcostal space wider than R1 space with widely spaced crossveins. Sc and R1 fused apically. Rs with six primary branches preserved, branches running parallel and forked distally several times. MA forked basally. MP with five branches preserved and dichotomously forked. CuA with six branches. CuP forked very basally, CuP1 with five branches. 1A stem long, very close to CuP and forked distally. 2A long and pectinately branched.

Etymology

The specific name refers to the occurrence of the type specimen at Karatau, Kazakhstan.

Discussion

The two new species are undoubtedly assigned to Kalligrammula Handlirsch, Reference Handlirsch1919 due to a combination of the hindwing characters: costal space narrow with relatively straight costal veinlets which are connected by few crossveins, stem of Rs originating from R near the base of the wing, MA simple, MP multibranched, CuP forked very basally with CuP2 close to 1A, 1A forked distally and an eye-spot well-developed.

Kalligrammula lata n. sp. can be distinguished from the four known congeneric species in having slightly sinuous costal veinlets, an Rs vein with 12 primary branches, and a slightly undulate outer margin. It is also notable that the vein 3A is present and long in K. lata n. sp. but is not detected in all other known species. Kalligrammula karatensis n. sp. is different from all other congeneric species in having Rs with six primary branches, basally forked MA, and dozens of small unpigmented spots located along the wing outer margin.

Labandeira (Reference Labandeira2010) pointed out that kalligrammatids had long proboscises ranging from 11 to 20–25 mm long. So it could be supposed that some of these lacewings were specialized pollinators like other Mesozoic insects with siphonate mouthparts, for example scorpionflies in the families Mesopsychidae, Aneuretopsychidae, and Pseudopolycentropodidae. Representatives of these three families with long proboscises, as well as kalligrammatids, first appeared in the Middle Jurassic (viz. Daohugou), and their extinction during the late Early Cretaceous coincided with the decline and extinction of a number of gymnosperm taxa at the time of the angiosperm radiation (Ren et al., Reference Ren, Labandeira, Santiago-Blay, Rasnitsyn, Shih, Bashkuev, Logan, Hotton and Dilcher2009). If kalligrammatids also formed associations with gymnosperms, the widespread presence of Kalligramma and Kalligrammula in the Middle Jurassic–Lower Cretaceous of Europe and Asia could be due to the similarity of floras in the low and mid latitudes of Laurasia. All representatives of both genera are known from the territory of the Euro-Sinian paleofloristic province, which is characterized by the abundance of Bennettitales (significantly with “flowers”) and Cheirolepidiaceae, and scarcity of Czekanowskiales and Ginkgoaceae (Vakhrameev, Reference Vakhrameev1991).

Acknowledgments

We are grateful to Q.-P. Lei (Changzhou Museum, Changzhou, Jiangsu, China) for her kind help and to S. Tränkner and C. Franz (Senckenberg Gesellschaft für Naturforschung, Frankfurt, Germany), who have taken photos of the K. senckenbergiana holotype. This research was supported by the National Basic Research Program of China (grant no. 2012CB821900), National Natural Science Foundation of China (grant nos. 41272013, 41002006, J1210006), the Chinese Academy of Sciences Visiting Professorship for Senior International Scientists (grant no. 2011T2Z04), and the RFFI grant no. 13-04-01839A.

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Figure 0

Figure 1Distribution of the family Kalligrammatidae.

Figure 1

Figure 2Holotype of Kalligrammula senckenbergiana Handlirsch, 1919. (1, 3) part; (2, 4) counterpart. (1, 2) basal part of hindwing respectively from (3) and (4) (indicated in black box). Scale bar is 5 mm for (1, 2) and 30 mm for (3, 4).

Figure 2

Figure 3Kalligrammula lata n. sp., (1, 2) holotype, a well-preserved left hindwing (NIGP156192a, b); (3) line drawing of holotype NIGP156192a. Scale bar is 20 mm.

Figure 3

Figure 4Kalligrammula karatensis n. sp., (1) holotype, a nearly complete left hindwing (PIN No. 2784/1032); (2) line drawing of holotype PIN No. 2784/1032. Scale bar is 10 mm.