Research 59 (2016) 1e9

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Cretaceous Research

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Short communication Two new glaphyrids (Coleoptera, ) from the Jehol Biota, China

** * Hongyun Zhao a, Ming Bai b, , Chungkun Shih a, Dong Ren a, a Key Lab of Evolution and Environmental Change, College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing 100048, China b Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, China article info abstract

Article history: Two new species, Cretohypna puncta sp. nov. and Cretohypna robusta sp. nov., of are Received 11 May 2015 described and illustrated. These fossils were collected from the Jehol Biota, Lower Cretaceous Yixian Received in revised form Formation of Liutiaogou Village in Ningcheng, Inner Mongolia, China. Up to date, 18 fossil species in 6 24 October 2015 genera of Glaphyridae have been reported, among which ten species are from China. Respective keys to Accepted in revised form 29 October 2015 the Mesozoic genera of Glaphyridae and to species of Cretohypna Yan, Nikolajev & Ren, 2012 are pro- Available online 22 November 2015 vided. This is the first report of spiracles in fossil Glaphyridae leading to a hypothesis that the character of the 7th abdominal spiracles present in pleural membrane is a plesiomorphy for Glaphyridae. Keywords: © New species 2015 Elsevier Ltd. All rights reserved. Mesozoic Lower Cretaceous Glaphyridae

1. Introduction Cretaceous of Russia; while Glaphyrus and Lichnanthe are discov- ered in Cenozoic of Germany and America. Fossil species of Gla- The Glaphyridae MacLeay, 1819 is a small family comprising ten phyridae have been described from four countries: Germany, genera with over 200 species and subspecies. Extinct species can be Russia, China and America, while about 90% of species from Russia found in six genera, among which four genera (Cretoglaphyrus and China. Nikolajev, 2005, Lithohypna Nikolajev, Wang & Zhang, 2011, Creto- Herein we describe two new species Cretohypna puncta sp. nov. hypna Yan, Nikolajev & Ren, 2012 and Mesohypna Nikolajev & Ren, and Cretohypna robusta sp. nov., based on four well-preserved 2013) are extinct genera, while two genera (Glaphyrus Latreille, fossils, from the Lower Cretaceous Yixian Formation near Liutiao- 1807 and Lichnanthe Burmeister, 1844) have both extinct and extant gou Village, Ningcheng County, Chifeng City, Inner Mongolia, China. species. The other four genera (Anthypna Eschscholtz, 1818, The new findings offer new evidence to broaden our understanding Amphicoma Latreille, 1807, Eulasia Truqui, 1848 and Pygopleurus of morphological characters, diversity and speciose of Mesozoic Motschlsky, 1860) only contain extant species (Nikolajev et al., glaphyrids and to elucidate the evolution of spiracles of glaphyrids. 2011). A list of described fossil species of Glaphyridae is summarized in Table 1 (Heer, 1862; Wickham, 1910; Carlson, 1980; Carlson, 2002; 2. Material and methods Nikolajev, 2005; Krell, 2007; Nikolajev and Ren, 2011; Nikolajev et al., 2011; Nikolajev and Ren, 2012; Yan et al., 2012b; Nikolajev The study is based on four specimens (including two with parts and Ren, 2013). As shown in Table 1, three genera of Cretohypna, and counterparts) collected near Liutiaogou Village, Ningcheng Lithohypna and Mesohypna are only distributed in the Lower County, Chifeng City, from Yixian Formation of Inner Mongolia, Cretaceous of China; Cretoglaphyrus is only distributed in the Lower China (Fig 1). Yixian Formation, belonging to the famous Jehol Biota (Ren et al., 1997, 2010, 2012), has yielded a number of fossil , such as Odonata (Zhang et al., 2006), Chresmodidae (Zhang et al.,

* Corresponding author. 2008), Coleoptera (Bai et al., 2012, 2013; Yan et al., 2012a,b; Yu ** Corresponding author. et al., 2013, 2015a,b), Hemiptera (Chen et al., 2014; Yao et al., E-mail addresses: [email protected] (M. Bai), [email protected] (D. Ren). 2014), Neuroptera (Shi et al., 2015), Hymenoptera (Li et al., 2014; http://dx.doi.org/10.1016/j.cretres.2015.10.026 0195-6671/© 2015 Elsevier Ltd. All rights reserved. 2 H. Zhao et al. / Cretaceous Research 59 (2016) 1e9

Table 1 Described fossil species of Glaphyridae (Coleoptera: Scarabaeoidea) from around the world.

Species Chronostratigraphy Locality Reference

Glaphyrus Heer, 1862 Glaphyrus antiquus Heer, 1862 Miocene Germany Heer (1862) Glaphyrus ancestralis Nikolajev & Ren, 2011 Lower Cretaceous China Nikolajev and Ren (2011) Cretoglaphyrus Nikolajev, 2005 Cretoglaphyrus calvescens Nikolajev, 2005 Lower Cretaceous Russia Nikolajev (2005) Cretoglaphyrus leptopterus Nikolajev, 2005 Lower Cretaceous Russia Nikolajev (2005) Cretoglaphyrus olenguicus Nikolajev, 2005 Lower Cretaceous Russia Nikolajev (2005) Cretoglaphyrus rohdendorfi Nikolajev, 2005 Lower Cretaceous Russia Nikolajev (2005) Cretoglaphyrus transbaikalicus Nikolajev, 2005 Lower Cretaceous Russia Nikolajev (2005) Cretoglaphyrus zherikhini Nikolajev, 2005 Lower Cretaceous Russia Nikolajev (2005) Cretohypna Yan, Nikolajev & Ren 2012 Cretohypna cristata Yan, Nikolajev & Ren 2012 Lower Cretaceous China Yan et al. (2012b) Cretohypna robusta Zhao, Bai, Shih & Ren, sp. nov. Lower Cretaceous China This study Cretohypna puncta Zhao, Bai, Shih & Ren, sp. nov. Lower Cretaceous China This study Lithohypna Nikolajev & Ren, 2012 Lithohypna chifengensis Nikolajev, Wang & Zhang, 2011 Lower Cretaceous China Nikolajev et al. (2011) Lithohypna lepticephala Nikolajev & Ren, 2012 Lower Cretaceous China Nikolajev and Ren (2012) Lithohypna longula Nikolajev & Ren, 2012 Lower Cretaceous China Nikolajev and Ren (2012) Lithohypna tuberculata Nikolajev & Ren, 2012 Lower Cretaceous China Nikolajev and Ren (2012) Lithohypna yuxiana Nikolajev & Ren, 2012 Lower Cretaceous China Nikolajev and Ren (2012) Lichnanthe (Wickham, 1910) Lichnanthe defuncta (Wickham, 1910) upper Eocene America Wickham (1910); Carlson (1980) Mesohypna Nikolajev & Ren, 2013 Mesohypna lopatini Nikolajev & Ren, 2013 Lower Cretaceous China Nikolajev and Ren (2013)

Updated from Yan et al., 2012b.

Wang et al., 2015) and Pseudopulicidae (Gao et al., 2012), which are housed in the Key Lab of Insect Evolution & Environmental provided new information to enhance our understanding of the Changes, College of Life Sciences, Capital Normal University, Beijing, taxonomy and evolution of the respective groups. The age of Yixian China (Dong Ren, Curator). Formation is regarded as (latest Barremian to earliest Aptian) and linked to a radioisotope-dating of ca. 125 Ma. 3. Systematic Paleontology (Yang et al., 2007; Zhu et al., 2007; Chang et al., 2009; Zhou and Wang, 2010; Pan et al., 2013). Order Coleoptera Linnaeus, 1758 The specimens were examined using a Leica MZ 16.5 dissecting Superfamily Scarabaeoidea Latreille, 1802 microscope connected to a Leica DFC500 digital camera and illus- Family Glaphyridae Macleay, 1819 trated with the aid of a drawing tube attachment. The figures were A key to the five genera of fossil Glaphyridae from the Mesozoic drawn by CorelDraw 12.0 and Photoshop CS 5.1. All type specimens is given here.

Fig. 1. Map showing the fossil locality. H. Zhao et al. / Cretaceous Research 59 (2016) 1e9 3

Key to the Mesozoic genera of Glaphyridae with apical 3-antennomere club, club loose. Clypeus front margin distinctly straight; labrum and mandibles prominent, extending 1. Body elongate; labrum relatively large, approximately twice as beyond apex of clypeus, anterior edge of labrum straight and wide as long…………………………………………………………2 distinctly wider than long, nearly four times as wide as long; Body compact and flat; labrum short, over twice as wide as maxillary palpus with 3 segments. long……………………………………………………………………3 Pronotum: Nearly subquadrate, 1.9 times wider than long, the 2. Labrum somewhat bilobed, sinuate in the middle; scutellum U- widest part near its base. Anterior margin concave, with lateral shaped; pygidium visible beyond elytra; protarsus longer than margins convex and posterior margin slightly protruding. Covered protibia………………Lithohypna Nikolajev, Wang & Zhang, 2011 with irregular punctures (Fig. 2A). Labrum front margin straight; scutellum triangular; pygidium Scutellum: Triangular, nearly as wide as long, covered with not beyond elytra; protarsus subequal to protibia……… irregular punctures. …………………………………Mesohypna Nikolajev & Ren, 2013 Wing: Elytra widest near the base, convexly constricted to the 3. Mesepimera well visible in dorsal view between the pronotum apex in the apical quarter, dehiscent. Full of irregular punctures and elytra……………………………………………………………..4 (Fig. 2A). Hind wing with the RP, RA3þ4,RA4þRP1 veins preserved Mesepimera not visible in dorsal view between the pronotum on the fossil. and elytra…………………………Cretoglaphyrus Nikolajev, 2005 Leg: Protibia dilated apically and three teeth on outer margin of 4. Clypeus with two or three teeth on anterior margin; elytra the protibia (Fig. 2E, F), relative lengths (base to apex) of five pro- smooth or with four distinct longitudinal carinae; apex of male tarsomeres 32: 21: 21: 27: 70 (Fig. 2G); mesotibia without ridges mesotibia without lamellate……………Glaphyrus Latreille, 1807 and outer apical angle of mesotibia simple, without lobe teeth or Clypeus with straight anterior margin; elytra without longitu- spines, mesotibia slender than metatibia; metatibia with two dinal carinae; apex of male mesotibia lamellate………… ridges, the apical spurs differing in length. ………………………...... Cretohypna Yan, Nikolajev & Ren, 2012 Abdomen: Pygidium exposed beyond apices of elytra. The 6th and 7th abdominal spiracles present and located in pleural mem- Genus Cretohypna Yan, Nikolajev & Ren, 2012 brane (Figs. 2D and 3C, F). Type species: Cretohypna cristata Yan, Nikolajev & Ren, 2012 Remarks. Compared with Cretohypna cristata Yan, Nikolajev & Ren, Key to known species of Cretohypna Yan, Nikolajev & Ren, 2012, C. puncta. sp. nov. has irregular punctures in pronotum and 2012 elytra while the spurs at the end of metatibia not of equal length. 1. Pronotum and elytra smooth, without obvious punctures; apex of Dimensions of holotype (in mm). Body 16.3 long and 6.3 wide (at the mesotibia lamellate…………………………………………………2 base of elytra). Head 2.8 in length and 3.4 in width as preserved; Pronotum and elytra with irregular punctures; apex of male pronotum length 3.4, width 6.3; elytra 8.4 in length, 4.0 in width. mesotibia not lamellate………………………………Cretohypna Dimensions of paratypes (in mm). CNU-COL-NN2014003, body 17.2 puncta sp. nov. long and 6.5 wide; CNU-COL-NN2014004p/c, body 15.6 long and 2. Clypeus wider than long; spurs of metatibia subequal in 7.2 wide. length………………Cretohypna cristata Yan, Nikolajev & Ren, 2012 Clypeus nearly as wide as long; spurs of metatibia differing Cretohypna robusta Zhao, Bai, Shih & Ren, sp. nov. (Figs. 4 e 5) distinctly in length…………………...Cretohypna robusta sp. nov. Etymology. The specific name is derived from the Latin word Cretohypna puncta Zhao, Bai, Shih & Ren, sp. nov. (Figs. 2 e 3) robustus, referring to the strong body. Etymology. The specific name is derived from the Latin word Type material. Holotype, No. CNU-COL-NN2014001p/c, a well- “punct-” referring to the irregular punctures in pronotum and preserved specimen collected from the Yixian Formation, near elytra. Liutiaogou Village, Ningcheng County, Chifeng City, Inner Mongolia, Type material. Holotype, CNU-COL-NN2014002, sex unknown, China, sex unknown. Housed in the Key Lab of Insect Evolution & deposited in CNUB, a well-preserved specimen in dorsal view. Environmental Changes, College of Life Sciences, Capital Normal Paratypes, CNU-COL-NN2014003, sex unknown, dorsal view, a University, Beijing, China. well-preserved specimen with antennae and part of hind wing Locality and horizon. Liutiaogou Village, Ningcheng County, Chifeng (Fig. 3E, F, G). CNU-COL-NN2014004p/c, sex unknown, part and City, from Yixian Formation of Inner Mongolia, China, Lower counterpart, a well-preserved specimen with part of hind wing Cretaceous (uppermost Barremian to lowermost Aptian), ca. (Fig. 3A, B, C, D). 125 Ma. Locality and horizon. Liutiaogou Village, Ningcheng County, Chifeng Diagnosis. Body large, elongate oval and compact. The antennae City, from Yixian Formation of Inner Mongolia, China, Lower with 10 antennomeres and a 3-antennomere lamellare club. Mes- Cretaceous (uppermost Barremian to lowermost Aptian), ca. ofemur slender than metafemur. Metatibia with transverse ridges 125 Ma. on outer margins. The spurs at the end of metatibia differing Diagnosis. Body oval, compact and flat. Antennae capitate, with distinctly in length. Mesopretarsus and metapretarsus with two apical club. Anterior margin of clypeus straight. Frons with dense long, thin and symmetrical claws. punctures. Sides of prothorax moderately to strongly curved. Pro- Description. Body large, elongate oval and compact, 23.2 mm long, notum and elytra full of irregular and big punctures. Mesotibia 11.1 mm wide (at the base of elytra) (Fig. 4A, B). without ridges and slender than metatibia. Metatibia with two Head: Slightly wider than long, narrower than pronotum. Eyes ridges, the apical spurs differing in length. not protuberant. The antennae with 10 antennomeres, with a 3- Description. Body oval, compact and flat, 16.3 mm long and 6.3 mm antennomere lamellare club, club loose, scape with long setae wide (at the base of elytra) (Fig. 2A, B). (Fig. 4C, D). Clypeus front margin round, with a transverse carina. Head: Nearly as wide as long, obviously narrower than prono- Labrum and mandibles prominent, produced beyond apex of tum, with dense punctuate on the frons (Fig. 2C). Antennae capitate clypeus, labrum broad and about four times as wide as long. Maxilla 4 H. Zhao et al. / Cretaceous Research 59 (2016) 1e9

Fig. 2. Holotype of Cretohypna puncta Zhao, Bai, Shih & Ren, sp. nov. CNU-COL-NN2014002. A, Photo. B, Line drawing. C, Head. D, Abdomen. E, F, Protibia and Protarsus (E: yellow rectangle in A; F: red rectangle in A). G, Metatarsus (blue rectangle in A). A, B, Scale bars ¼ 2 mm. CeG, Scale bars ¼ 1 mm. (Online figure in color). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article). H. Zhao et al. / Cretaceous Research 59 (2016) 1e9 5

Fig. 3. Photos of paratypes of Cretohypna puncta sp. nov. AeD, CNU-COL-NN2014004p/c. C, Spiracles on the abdomen (red rectangle in A). D, Protibia and Protarsus (purple rectangle in A). EeG, CNU-COL-NN2014003. F, Spiracles on the abdomen (blue rectangle in E). G, Antennae (green rectangle in E). A, B, E, Scale bars ¼ 2 mm. C, D, F, G, Scale bars ¼ 1 mm. (Online figure in color). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article).

with setae, maxillary palpus with 3 segments. mesotibia lamellate (Fig. 4F), metatibia strong and dilated at the Pronotum: Subtrapezoidal, obviously wider than head, widest at apex, transverse ridges on the mesotibia and metatibia, two spurs base, anterior margin concave and lateral sides slightly convex, at the end of metatibia differing distinctly in length (Fig. 4I); pro- posterior angles of pronotum moderately round. coxa with setae (Fig. 4H), mesocoxal cavities oval, slightly sepa- Scutellum: Triangular, 1.8 times as wide as long. rated; protarsus preserved with only three tarsomeres in this Wing: Elytron long and narrow, 2.4 times as long as wide, specimen, mesotarsus and metatarsus with five segments and their without longitudinal carina convexly constricted at the apical pretarsi with two symmetrical claws (Fig. 4E). Mesotarsus length quarter and dehiscent at the apex. Hind wing with RP, RA3þ4, ratio of each segment (base to apex) 4.73: 1: 1.43: 1.37: 2.60; RA4þRP1 veins preserved on the fossil. metatarsus length ratio of each segment (base to apex) 3.61: 1.42: Leg: Robust legs. Protibia with 3 teeth on outer margin, apex of 1.20: 1: 1.37 (Fig. 4I). 6 H. Zhao et al. / Cretaceous Research 59 (2016) 1e9

Fig. 4. Photos of holotype of Cretohypna robusta Zhao, Bai, Shih & Ren, sp. nov. CNU-COL-NN2014001p/c. A, Dorsal view. B, Ventral view. C, Setae on the scape (red rectangle in A). D, Antennae (yellow rectangle in A). E, Mesotarsus (green rectangle in A). F, Mesotibia (dark blue rectangle in B). G, Spiracles on the abdomen (purple rectangle in A). H, Setae on the coxa (under alcohol. pink rectangle in B). I, Metatarsus (light blue rectangle in B). Scale bars ¼ 1 mm. (Online figure in color). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article). H. Zhao et al. / Cretaceous Research 59 (2016) 1e9 7

Fig. 5. Line drawings of holotype of Cretohypna robusta sp. nov. A, Dorsal view. B, Ventral view. Scale bars ¼ 1 mm.

Abdomen: Six visible sternites and the first ventrite not lamellate, six visible sternites and the first sternites not obscured by obscured by hind coxae, pygidium exposed beyond apices of elytra. hind coxae. These two new species are not placed in other genera The 7th abdominal spiracles preserved and located in pleural based on the following reasons: anterior margin of clypeus with membrane (Fig. 4G). two or three teeth in Claphyrus Latreille, 1807, but clypeus has straight anterior margin for the two new species. Labrum front Remarks. Compared with Cretohypna cristata Yan, Nikolajev & Ren, margin straight, scutellum triangular and protarsus shorter than 2012, C. robusta sp. nov. has a longer clypeus, and the spurs at the protibia in the two new species don't allow them to be placed in end of metatibia differing distinctly in length. Compared with Lithohypna Nikolajev, Wang & Zhang, 2011. Mesoepimeron visible C. puncta. sp. nov., C. robusta sp. nov. doesn't have irregular punc- between the pronotum and elytra in dorsal view of the two new tures in pronotum and elytra, but has two transverse ridges on the species is different from those of Cretoglaphyrus Nikolajev, 2005. mesotibia. Mesohypna Nikolajev & Ren, 2013 has metatarsi longer than met- atibiae, while metatarsi are shorter than metatibiae for the two Dimensions of holotype (in mm). Body 23.2 long, 11.1 wide (at the new species. base of elytra). Head 3.8 in length and 4.3 in width as preserved; A total of 18 species has been described so far, among which 16 pronotum length 4.1, width 9.2; elytra 13.5 in length, 5.7 in width. species reported from the Lower Cretaceous, one from the Miocene another from the upper Eocene (Table 1). The age distribution suggests a high diversity of Glaphyridae during the Early Creta- 4. Discussion ceous. In addition, 13 species out of the 16 previously described have preserved bodies and elytra with the exception of Cretogla- Cretohypna puncta sp. nov. is classified in Cretohypna Yan, phyrus leptopterus, C. transbaikalicus, C. calvescens having only one Nikolajev & Ren, 2012 based on the following characters: body elytron preserved. Our findings not only broaden the species di- compact, pronotum subquadrate shaped with concave anterior versity of Glaphyridae, but also provide new morphological char- margin, scutellum triangular, metatarsus shorter than corre- acters for future phylogenetic study and new information to sponding tibia, and elytra not covering pygidium. Cretohypna enhance our understanding of the evolution of Glaphyridae. robusta sp. nov. belongs to Cretohypna based on pronotum with This is the first report on the spiracles in fossil Glaphyridae concave anterior margin and convex lateral and posterior margins, (Figs. 2D, 3C, F and 4G), which are also rarely preserved in other mesocoxae moderately separated, apex of male mesotibia insect fossils. The positions of spiracles have been used in the 8 H. Zhao et al. / Cretaceous Research 59 (2016) 1e9 higher systematics, which could be classified into two groups: References Laparostic and Pleurostic (Zheng and Gui, 1999; Jameson and Ratcliffe, 2002). For extant Glaphyridae, the 7th abdominal spira- Bai, M., Ren, D., Yang, X.K., 2012. Prosinodendron krelli from the Yixian Formation, China: a missing link among Lucanidae, Diphyllostomatidae and Passalidae cles are located in either pleural membrane or tergite, while the 6th (Coleoptera: Scarabaeoidea). Cretaceous Research 34 (1), 334e339. http:// or 5th abdominal spiracles located in pleural membrane (Carlson, dx.doi.org/10.1016/j.cretres.2011.11.017. 2002). Only two of the abdominal spiracles are preserved in our Bai, M., Beutel, R.G., Shih, C.K., Ren, D., Yang, X.K., 2013. Septiventeridae, a new and ancestral fossil family of Scarabaeoidea (Insecta: Coleoptera) from the Late C. puncta sp. nov. specimens: the 6th and 7th abdominal spiracles to Early Cretaceous Yixian Formation, China. Journal of Systematic are present in pleural membrane, but the 5th abdominal spiracles Palaeontology 11 (3), 359e374. are not preserved. For the Cretohypna robusta sp. nov., the 7th Carlson, D.C., 1980. Taxonomic revision of Lichnanthe Burmeister (Coleoptera: e abdominal spiracles are also present in pleural membrane. Based Scarabaeidae). The Coleopterists Bulletin 34 (2), 177 208. Carlson, D.C., 2002. Glaphyridae MacLeay, 1819. In: Arnett, R.H., Thomas, M.C., on these limited data, it is hypothesized that the character of the Skelley, P.E., Frank, J.H. (Eds.), American , : Scarabaeoidea 7th abdominal spiracles present in pleural membrane is a plesio- through Curculionoidea, vol. 2. CRC Press, Boca Raton, pp. 37e38. morphy for Glaphyridae while the character of those present in Chang, S.C., Zhang, H.C., Renne, P.R., Fang, Y., 2009. High-precision 40Ar/39Ar age for the Jehol Biota. Palaeogeography, Palaeoclimatology, Palaeoecology 280, tergite is an apomorphy. 94e104. The spiracles are covered by elytra in modern glaphyrids. Chen, D., Yao, Y.Z., Ren, D., 2014. A new species of fossil Procercopidae(Hemiptera, Although the spiracles can be observed in our fossil specimens due Cicadomorpha) from the Lower Cretaceous of Northeastern China. Cretaceous Research 52, 402e406. to a slight opening of elytra, it is highly possible the spiracles are not Gao, T.P., Shih, C.K., Xu, X., Wang, S., Ren, D., 2012. Mid-Mesozoic Flea-like ecto- exposed and would be covered by elytra when the elytra are closed. parasites of feathered or Haired Vertebrates. Current Biology 22, 732e735. € Many species of the extant adult Glaphyridae are brightly Heer, O., 1862. Beitrage zur Insektenfauna Oeningens Coleoptera Geodaphagen, Hydrocanthariden, Gyriniden, Brachelytren, Clavicornen, Lamellicornen und colored, covered with distinct setae, resembling bees and bumble Buprestiden. Natuurkundige Verhandelingen van de Hollandsche Maatschappij bees, and often observed near flowers or foliage, the pronotum and der Wetenschappen te Haarlem 16 (2), 90. elytra of some extant glaphyrids have punctures under the setae Jameson, M.L., Ratcliffe, B.C., 2002. Superfamily Scarabaeoidea Latreille, 1802. In: Arnett, R.H., Thomas, M.C., Skelley, P.E., Frank, J.H. (Eds.), American Beetles, (Carlson, 2002). However, among the 18 fossil species reported, Polyphaga: Scarabaeoidea through Curculionoidea, vol. 2. CRC Press, Boca including our new species, the setae are only preserved on the Raton, pp. 1e5. elytra surface of Lichnanthe defuncta (Wickham, 1910; Carlson, Krell, F.T., 2007. Catalogue of Fossil Scarabaeoidea (Coleoptera: Polyphaga) of the 1980) and on the scape, maxilla and coxae of Cretohypna robusta Mesozoic and Tertiary e Version 2007. Denver Museum of Nature and Science, p. 81. Technical Report 8. sp. nov. All others do not have distinct and dense setae preserved. In Li, L.F., Shih, C.K., Ren, D., 2014. New fossil evaniids (Hymenoptera, Evanioidea) from addition, the obviously irregular punctures are found in Cretohypna the Yixian Formation of western Liaoning, China. Cretaceous Research 47, e puncta sp. nov., which is very similar to modern glaphyrids. 48 55. Nikolajev, G.V., 2005. Scarab beetles of the subfamily Glaphyrinae (Coleoptera, Scarabaeidae) in the Lover Cretaceous of Transbaikalia (Plastinchatousye zhuki 5. Conclusions podsemeystva Glaphyrinae (Coleoptera, Scarabaeidae) v nizhnem mele Zabay- kal'ya). Zhivotnyy mir Dal'nego Vostoka 5, 57e66 (In Russian with English abstract). We describe two new species, Cretohypna puncta sp. nov. and Nikolajev, G.V., Ren, D., 2011. The oldest species of the genus Glaphyrus Latr. Cretohypna robusta sp. nov., from the Lower Cretaceous Yixian (Coleoptera: Scarabaeoidea: Glaphyridae) from the Mesozoic of China. Pale- ontological Journal 2, 179e182. Formation. So far, 18 fossil species in 6 genera of Glaphyridae have Nikolajev, G.V., Ren, D., 2012. New species of the genus Lithohypna Nikolajev, been reported and 5 genera from the Mesozoic. Respective keys to Wang et Zhang, 2011 (Coleoptera, Scarabaeidae, Glaphyridae) from the Yixian species of Cretohypna and the Mesozoic genera of Glaphyridae are Formation, China. Evraziatskiy entomologicheskiy zhurnal 11 (3), 209e211þ212þI. provided. Our new fossils have clearly preserved spiracles, which Nikolajev, G.V., Ren, D., 2013. A new Glaphyridae genus (Coleoptera: Scar- are described for the first time in fossil Glaphyridae. It is hypoth- abaeidae) from the Yixian Formation. Caucasian Entomological Bulletin 9 (1), esized that the character of the 7th abdominal spiracles present in 62e64. pleural membrane is a plesiomorphy for Glaphyridae while the Nikolajev, G.V., Wang, B., Zhang, H.C., 2011. A new fossil genus of the family Gla- phyridae (Coleoptera: Scarabaeoidea) from the Lower Cretaceous Yixian For- character of those present in tergite is an apomorphy. New infor- mation. Zootaxa 2811, 47e52. mation and data provide new morphological characters to enhance Pan, Y.H., Sha, J.G., Zhou, Z.H., Fürsich, F.T., 2013. The Jehol Biota: definition and our knowledge of diversity and speciose of Mesozoic glaphyrids distribution of exceptionally preserved relicts of a continental Early Cretaceous ecosystem. Cretaceous Research 44 (0), 30e38. http://dx.doi.org/10.1016/ and lay the foundation for future phylogenetic study of j.cretres.2013.03.007. Glaphyridae. Ren, D., Guo, Z.G., Lu, L.W., Ji, S.A., Tang, W., Jing, Y.G., Fang, X.S., Ji, Q., 1997. A further contribution to the knowledge of the Upper Jurassic Yixian Formation in western Liaoning. Geological Review 43, 449e459. Acknowledgments Ren, D., Shih, C.K., Gao, T.P., Yao, Y.Z., Zhao, Y.Y., 2010. Silent Story e Insect Fossil Treasures from Dinosaur Era of the Northeastern China. Science Press, Beijing, p. 332. We thank the editor and two anonymous reviewers for their Ren, D., Shih, C.K., Gao, T.P., Yao, Y.Z., Zhao, Y.Y., 2012. Insect Fossil Treasures from advice, suggestion and comments in improving our manuscript. We the Mesozoic of the Northeastern China. Science Press, Beijing, p. 409 (in sincerely thank Dr. Adam Slipinski from CSIRO Ecosystem Sciences Chinese). Shi, C.F., Winterton, S.L., Ren, D., 2015. 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