A new cockroach (Insecta: Blattaria: Blattulidae) from the Lower Cretaceous Laiyang Formation of China

Cretaceous Research 101 (2019) 17e22

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A new cockroach (Insecta: Blattaria: Blattulidae) from the Lower Cretaceous Laiyang Formation of China Tao Chen a, b, Shaohua Liu a, b, Xiaojie Lei b, c, Lei Chen a, b, * a

Shandong Provincial Key Laboratory of Depositional Mineralization & Sedimentary Minerals, Shandong University of Science and Technology, Qingdao, Shandong 266590, China b State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology and Center for Excellence in Life and Paleoenvironment, Chinese Academy of Sciences, 39 East Beijing Road, Nanjing 210008, China c University of Science and Technology of China, No.96, JinZhai Road Baohe District, Hefei, Anhui 230026, China

a r t i c l e i n f o

a b s t r a c t

Article history: Received 6 December 2018 Received in revised form 31 January 2019 Accepted in revised form 3 March 2019 Available online 8 March 2019

The Blattulidae is an extinct family of cosmopolitan Mesozoic cockroaches with a low generic diversity. Habroblattula laiyangensis sp. nov. from the Lower Cretaceous Laiyang Formation of Shandong Province of China with well-preserved both hind wings is only closely related to the H. drepanoides Wang, Liang & Ren, 2007 from the Early Cretaceous Yixian Formation, sharing the venation pattern, while being significantly smaller, and provides new characteristics of ancient Blattulidae. In addition, H. laiyangensis sp. nov. resembles distribution patterns of a closely related complex of “Piniblattella” yixianensis Gao, Shih & Ren, 2018 with “Mesoblattina” sinica Ping, 1928, suggesting a spatiotemporal paleobioprovincial relation among Laiyang and Yixian formations, former probably representing more warmer southern localities. Laiyangia Grabau, 1923 and Sinoblatta Grabau, 1923 are indigenous to Laiyang and lack analogues also in numerous Cretaceous Mongolian and Russian sites evidencing a degree of isolation. © 2019 Published by Elsevier Ltd.

Keywords: Fossil insect Mesozoic Blattodea New species

1. Introduction Blattaria (commonly known as cockroaches) are one of the most important urban health pests (Tan, 1980; Vr
sanský et al., 2002; Liang et al., 2006). They firstly appeared in the Middle Pennsylvanian in the Carboniferous fossil recorded about 320 Ma and flourished rapidly throughout the earth (Tan, 1980; Vr
sanský et al., 2002; Liang et al., 2006). Cockroaches with about 4300 known living (Roth, 2003; Vr
sanský, 2008) and 1500 fossil species (about 500 more species than the estimated by Vr
sanský et al. (2002)) (Vr
sanský, 2008), and the variety of species with about 110,000 collected adult fossil specimens are second only to the largest number of Coleoptera fossils (Carpenter, 1953, 1992; Liang et al., 2006). Linnaeus marked the beginning when he established Blatta Linnaeus, 1758, the genus of cockroaches (Linnaeus, 1758; Wang and Che, 2010). Blattulidae was attributed to Mesoblattinidae for over a century. Vishniakova (1982) promoted Blattulidae into a family, and the families Blattulidae and Mesoblattinidae Handlirsch, 1906 very similar only superficially (Liang et al., 2006).

* Corresponding author. E-mail address: [email protected] (L. Chen). https://doi.org/10.1016/j.cretres.2019.03.003 0195-6671/© 2019 Published by Elsevier Ltd.

The Blattulidae is a dominant extinct family of cosmopolitan Mesozoic cockroaches. Nevertheless, their diversity at the generic level is poordonly 13 genera are known in their greater than 100 myr history (Cifuentes et al., 2006; Vr
sanský, 2005a; Vr
sanský et al., 2017; Wang et al., 2007). Mesozoic cockroaches are best studied in Myanmar amber due to enormous discoveries during last years (Bai et al., 2016, 2018; Grimaldi and Ross, 2004; Hinkelman, 2019; Ko
c arek, 2018; Li and Huang, 2018; Mlynský et al., 2018; Poinar, 1999, 2009; Poinar and  and Sendi, 2018; Qiu et al., 2019; Sendi Brown, 2017; Podstrelena
 and Lei, 2017; Vr
sanský and Bechly, and Azar, 2017; Smídov a 2015; Vr
sanský and Wang, 2017; Vr
sanský et al., 2017, 2018a, b, 2019a, b). To these vast Late Cretaceous data, it is necessary to establish an Early Cretaceous reference point like done in Lebanese amber (Vr
sanský, 2003b, 2004; Anisyutkin and Gorochov, 2008; Sendi and Azar, 2017). One of Early Cretaceous data is related Yixian Formation with numerous cockroaches, few of which were formally described (see Table 4). Habroblattula drepanoides Wang et a., 2007 was included, a representative of the genus Habroblattula Wang et al., 2007 now found in Laiyang Formation. Therefore it is essential to collect more data on the Early Cretaceous cockroaches of other localities in China (summarized by Liang et al., 2006), in addition to those already summarized from Mongolia.

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T. Chen et al. / Cretaceous Research 101 (2019) 17e22

Table 1 Comparison with Habroblattula drepanoides Wang et al., 2007, Sinoblattaria laiyangensis Grabau, 1923 and Laiyangia paradoxiformis Grabau, 1923; body type. Head length/width(mm)

Body length/width(mm)

Forewing length/width(mm)

Hind wing length/width(mm)

Habroblattula harpeodes Sinoblattaria laiyangeasis

2.0e2.7/2.0e2.2 2.0/3.0

11.0e12.0/3.2e4 –/–

10.0e11.0/– –/–

Laiyangia paradoxiformis Habroblattula laiyangensis sp. nov.

2.0/1.3 –/–

–/– 17.0(end of last preserved segment of abdomen)/– 17.0/– –/–

12.0/3.8 –/–

–/– 7.3/3.6

Table 2 Comparison with Habroblattula drepanoides Wang et al., 2007; venation.

Habroblattula drepanoides Habroblattula laiyangensis sp. nov.

R1

Rs

M

CuA

CuP

A

5e6 6

9e11 6 or 8

2e5 4 or 6

6e9 6

1 1

1 1

Table 3 Cockroaches from the Lower Cretaceous Laiyang Formation. Genus

Species

Sinoblatta Laiyangia

Sinoblatta laiyangensis Grabau, 1923 Laiyangia paradoxifonnis Grabau, 1923 Laiyangia delicatula Zhang, 1985 Habroblattula laiyangensis sp. nov.

Habroblattula

Table 4 Blattulids from the Early Cretaceous Yixian Formation. Genus

Species

Habroblattula Elisama

Habroblattula drepanoides Wang et al., 2007 Elisama extenuata Ren, 1995 Elisama hyalina Wang, 2007 Elisama cuboides Wang, 2007 Blattula delicatula Ren, 1995 Blattula platypa Ren, 1995 Macaroblattula ellipeoides Wang, 2007

Blattula Macaroblattula

The third Member of the Laiyang Formation consists of 250e705 m of calcareous sandstone, siltstone and fine-granted sandstone intercalated with thin, paper-like, insect-rich shales representing the facies of a large (several hundred km2) lake in the Laiyang basin. The diversity and preservation of fossil insects in Laiyang are excellent. It provides a reliable basis for deepening the research on the late Mesozoic insect community in the East Asia on restoring the overall features of palaeogeography, palaeoclimatology and palaeoecology (Zhang, 1991). In this paper, Habroblattula laiyangensis sp. nov. is described based on a wellpreserved fossil with two hind wings from the Laiyang Formation and provides new characteristics of ancient Blattulidae. 2. Material and methods The specimen was collected from the Laiyang Formation, Nanligezhuang Village, Tuanwang Town, Laiyang City, Shandong Province, China (Fig. 1). The Lower Cretaceous Laiyang Formation (Zhang et al., 2015; Jin et al., 2018; Zhao et al., 2018) with a depositional age of ca. 121e120 Ma (early Aptian) consists primarily of fluvial and lacustrine siltstones and sandstones facies interbedded with shale and volcaniclastic rocks (Wang et al., 2015; Zhao et al., 2018). The specimen is deposited in Shandong University of Science and Technology, Qingdao, Shandong, China. Specimen was

examined dry and cleaned up with alcohol, photograph was taken using a LEICA M125; and optimized using allowed all-surface improvements with Photoshop CS6. Line drawing was performed with image editing software CorelDraw X8. Venational terms follow Comstock and Needham (1899), currently used in cockroach wing nomenclature (Vr
sanský et al., 2018a, b, 2019b). See also Li et al. (2018) for modest elaboration of cockroach wing system.

3. Systematic paleontology Order Blattaria Latreille, 1810. Family Blattulidae Vishniakova, 1982. Genus Habroblattula Wang et al., 2007. Type species. Habroblattula drepanoides Wang et al., 2007 by original designation. Other species. Habroblattula laiyangensis sp. nov. by present designation. Diagnosis. (on the basis of hind wing after Wang 2007). Sc simple, nearly straight and unbranched, reaching costal margin gently. Radius (R) branching into R1 and Rs; R1 branches short; pterostigma present; Rs branches numerous. M rich. CuA with many simple branches, CuP simple. Intercalary veins and cross-veins present. Habroblattula laiyangensis sp. nov (Fig. 2). Derivation of name. The nomen triviale laiyangensis refers to Laiyang Formation where the specimen was collected. Gender, feminine. Holotype. Fossil SKD-LYQ-201808089a (Fig. 2). Well-preserved two hind wings and faint metathorax rudiments. Type locality and horizon. Nanligezhuang Village, Tuanwang Town, Laiyang City, Shandong Province, China; Lower Cretaceous (upper Barremianelower Aptian) Laiyang Formation. Description. Hind wings medium-size, 7.3 mm long as preserved and 3.6 mm wide in mid length, with distinct and numerous intercalary veins and cross-veins. Anterior margin arched. Apex sharp. Sc simple, long and nearly straight. R1 with pterostigma, reaching anterior margin slightly beyond 1/2 of costal margin, giving off 6 veins ending at margin (one on the wing on the left side is secondarily branched). Rs differentiated, curved, ending in costal margin, extensively branched, with 6 branches on the left and 8 branches on the right. M with 4 branches on the left and 6 branches on the right; anteriormost branch of M ending at apex. Cubitus (Cu) branched before middle of wing; CuA with 6 simple descending branches, ending in outer margin; CuP sigmoidal, simple. A1 present in remigium, sigmoidal and with blind branch descending basally. Vannus short, and half the wing length, with 1 forked and numerous unforked anal veins. Character of preservation. Two hind wings outspread, with rudiments of metathorax.

T. Chen et al. / Cretaceous Research 101 (2019) 17e22

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Fig. 1. A, location map of type locality; B, outcrop at Nanligezhuang Village.

4. Discussion The species can be categorized within Corydioidea on the basis of the venations scheme (Sc1, R1þRs, branched M, CuA branched, CuP1, A1 branched) and more specifically within Blattulidae on the basis of lack of CuA secondarily branches. The distinct pterostigma and apex coloration, branching R1, M with a robust base, cubital area with cross veins and forked A1 prove that the new species belong to Habroblattula Wang et al., 2007, but H. laiyangensis sp. nov. differs from H. drepanoides Wang et al., 2007 by small size, less Rs and M of the tertiary branches (see Tables 1 and 2). Closely related Laiyangia Grabau, 1923 differs in lack of coloration, sigmoidally curved anteriormost R1 and Kridla Vr
sanský, 2005 has coloration more extensive. The specimen (Fig. 3) is unique in the length of hind wing, and the M of the hind wing with tertiary branches. Habroblattula laiyangensis sp. nov. is closely related to H. drepanoides Wang et al., 2007 from the Lower Cretaceous of Yixian Formation. It can be further distinguished from all other blattulids in the hind wing with the Sc reaching 1/2 of the wing length, less Rs, more M, and possibly by the Cubital area without cross veins (see Table 2). The predominantly fine-grained sediments of the Laiyang Formation suggest a subdued topography with hydrophilous equisetalean marshland around the lake margin. There existed a large number of phytophagous, parasitic, predatory and saprophagous insects throughout the forest and marsh zones (Zhang, 1985; Zhang et al., 1986; Zhang, 1992). The discovery of a large number of specific insect combinations confirmed this conclusion. Cockroaches from the Lower Cretaceous Laiyang Formation include four species of three genera: Sinoblatta laiyangensis Grabau, 1923, Laiyangia paradoxifonnis Grabau, 1923, Laiyangia delicatula Zhang, 1985 and now also Habroblattula laiyangensis sp. nov. These record (see Table 3) suggest that the diversity was not extremely reduced. Taphonomically, rather small species were preserved and more or less complete specimens suggest a short predepositional

transport. Preservation of both hind wings is rare, which more frequently occurs only in Kholbotu Gol in Mongolia (Vr
sanský, 2008). In most well studied locality Bon Tsagaan Nuur in Mongolia (comparable age and region), only 11 both hind wings were preserved among 671 specimens (Vr
sanský, 2003a), but notably all of them belonging to Blattulidae. Also neither the new species, nor previously described specimens bear a single wing deformity (Vr
sanský, 2005b; Vr
sanský et al., 2017) confirming the lack of diversification point during the Laiyang times. Compared with Yixian Formation, the diversity of Blattulidae in the Laiyang Formation has declined. Four genera and seven species of the family Blattulidae have been reported in Yixian Formation (see Table 4), and there also had found other families like Ectobiidae (Gao et al., 2018), Mesoblattinidae (Wei & Ren, 2013) and Caloblattinidae (Wang & Ren, 2013). Laiyang Formation is not very diverse, the number of genera and species is much lower than in Yixian Formation. On the other hand, Laiangia with two species is indigenous to Laiyang place. This shows that there is a large gap between the diversity of Yixian Formation and Laiyang Formation. It is preliminary to judge the overall connection between Laiyang and Yixian faunas, because Habroblattula Wang et al., 2007 and several ectobiid genera represented by “Piniblattella” yixianensis Gao et al., 2018 and “Mesoblattina” sinica Ping, 1928 (synapomorphic in terminal forewing branchelets, long clavus and symplesiomorphic in branched Sc) are found only in these two formations, while they were apparently absent in numerous Mongolian and Russian sites. Due to the wide stratigraphy in Mongolian sites (Upper JurassiceUpper Cretaceous), this doubtfully relate to temporal differences. As a result, it relates either to communication between these two faunas, or to the warmer climate in more southern localities. Because of the rarity of Blattulidae fossils in Laiyang Biota, it is necessary to erect the new species according to hind wings to reveal the diversity of Laiyang Biota. We expect to find forewings or more complete fossils in the near future.

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T. Chen et al. / Cretaceous Research 101 (2019) 17e22

Fig. 2. Photograph of holotype SKD-LYQ-201808089a, Habroblattula laiyangensis sp. nov. Scale bar ¼ 2 mm.

Fig. 3. Line drawing of hind wing of holotype SKD-LYQ-201808089a, Habroblattula laiyangensis sp. nov. Abbreviations: Sc, Subcosta; R1, the first Radial; Rs, Radial Sector; M, Media; CuA, the first Cubitus; CuP, the second Cubitus; A1, the first Anal vein. Scale bar ¼ 2 mm.

T. Chen et al. / Cretaceous Research 101 (2019) 17e22

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sanský for their useful comments on the manuscript. We are also thankful to the editor and anonymous reviewers for their invaluable suggestions. This research was supported by the National Key Research and Development Program of China (2016YFC0601001), National Natural Science Foundation of China (41502019, 41572010, 41622201), Shandong University of Science and Technology Research Fund (2015RCJJ009, 2015YQJH102, 2015TDJH101). References Anisyutkin, L.N., Gorochov, A.V., 2008. A new genus and species of the cockroach family Blattulidae from Lebanese amber (Dictyoptera, Blattina). Paleontological Journal 42 (1), 43e46. Bai, M., Beutel, R.G., Klass, K.D., Zhang, W.W., Yang, W.K., Wipfler, B., 2016. yAlienopterada new insect order in the roachemantodean twilight zone. Gondwana Research 39, 317e326. €rnig, M., et al., 2018. A new CretaBai, M., Beutel, R.G., Zhang, W.W., Wang, S., Ho ceous insect with a unique cephalo-thoracic scissor device. Current Biology 28, 438e443. Carpenter, F.M., 1953. The geological history and evolution of insects. American Scientist 41, 256e270. Carpenter, F.M., 1992. Order Blattaria Latreille, 1810. In: Kaesler, Roger L. (Ed.), Treatise on invertebrate Palaeontology. Part R.Arthropoda 4 (3). Superclass Hexapoda. The Geological Society of America, Inc.and the University of Kansas, Boulder, Colorado and Lawrence, pp. 134e137. les-Soriano, E., Cifuentes-Ruiz, P., Vr
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Appendix A. Supplementary data Supplementary data to this article can be found online at https://doi.org/10. 1016/j.cretres.2019.03.003.