Research 80 (2017) 86e90

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Cretaceous Research

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Short communication A new reticulated (Coleoptera: Cupedidae) with aedeagus preserved from mid-Cretaceous amber of Myanmar

* Edmund A. Jarzembowski a, b, , Bo Wang a, c, Daran Zheng a, d a State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Rd, Nanjing 210008, People's Republic of China b Department of Earth Sciences, Natural History Museum, Cromwell Rd, London SW7 5BD, UK c Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, 1, Beichen West Road, Beijing 100101, China d Department of Earth Sciences, The University of Hong Kong, Hong Kong Special Administrative Region, China article info abstract

Article history: A rare archaic beetle, Barbaticupes combertiae n. gen. n. sp. (Insecta: Coleoptera: : Cupe- Received 23 May 2017 didae) is described from mid-Cretaceous burmite from northern Myanmar. This beetle's distinctive form Received in revised form includes dense setae (small hairs) and protuberances on the head and merging rows of window cells on 14 August 2017 the elytra (wing cases). It is the first true or typical reticulated beetle (cupedine) to be found in Burmese Accepted in revised form 30 August 2017 amber displaying male genitalia. Available online 1 September 2017 © 2017 Elsevier Ltd. All rights reserved.

Keywords: Fossil beetle Myanmar New genus New species

1. Introduction English Weald and northeastern China: this is a little surprising considering the modern association of these with wood, but With well over 300,000 described species, beetles (Coleoptera there may be a size filter involved, or even competition with new Linnaeus, 1758) are easily the largest order in the kingdom. terrestrial fauna, both of which need to be investigated. The Cupedina Ponomarenko, 1973, however, is the smallest beetle The beetle described herein belongs to an uncommon species ‘suborder’, totaling only about 135 living species, and now with only two known examples from an estimated 100,000þ commonly split into the even smaller suborders Archostemata amber inclusions. It is nevertheless a typical reticulated beetle, Kolbe, 1908 and Crowson, 1955 (Hornschemeyer,€ 2016). i.e.cupedine, a group sometimes considered to represent a separate Several hundred species of fossil cupedinans have been described family, e.g. Lawrence and Slipinski (2013). The reticulated beetles from the onwards and the archaic archostematans are a (cupedids) are so called because of their clathrate elytra (wing cases notable element of Mesozoic faunas, even occurring in places often showing a lattice pattern) and are the dominant arch- from where they have now vanished, such as the south of England ostematans both at the present day and in the late Mesozoic, (Kirejtshuk and Ponomarenko, 2015). Such finds are often pre- although in the latter they frequently belong to extinct taxa. The served as adpressions (‘rock’) fossils, but they are now turning up as cupedines are traditionally split three ways into the still-living amber inclusions in Myanmar, also known as Burma (Jarzembowski priacmins and cupedins and fossil mesocupedins (or their sub- et al., 2017b). Archostematans are nevertheless scarce in Burmese family equivalents): on this system, the new find shows affinity amber compared with earlier Cretaceous deposits, such as in the with the first of the three. These formal tribes (sometimes elevated to subfamilies) are, however, considered unworkable in the recent study by Kirejtshuk et al. (2016), so are referred to only informally here. Recent workers also prefer a narrower definition of the type * genus Fabricius, 1801 recognizing segregates such as Teno- Corresponding author. State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 merga Neboiss, 1984 (Lawrence and Slipinski, 2013); these are East Beijing Rd, Nanjing 210008, People's Republic of China. indicated in parentheses below. E-mail address: [email protected] (E.A. Jarzembowski). http://dx.doi.org/10.1016/j.cretres.2017.08.015 0195-6671/© 2017 Elsevier Ltd. All rights reserved. E.A. Jarzembowski et al. / Cretaceous Research 80 (2017) 86e90 87

2. Geological setting Derivation of name. Combination of barbati, Latin (genitive) for bearded, and generic name Cupes; neuter. (amber from northern Myanmar, birmite or Diagnosis. Small, hairy fossil cupedid, about half a centimetre long, burmite) contains the most diverse biota in amber known from the with broad head and long antennae and tarsi; four prominent head Cretaceous and has undergone a recent revival of interest (Wang protuberances (P1 2, P2 2); eyes rounded, moderately large et al., 2015). Thirty-four orders of are represented, beetles with transverse diameter not more than half of distance between being one of the most diverse, but the majority of species are them; temples pointed, curved anteriorly around eyes with brush- undescribed. The fossil resin is dated stratigraphically and radio- like setal fringes; pronotum subquadrate, wider than long, with metrically as late Albian or early (Cruickshank and Ko, peaked median longitudinal ridge, anterior angles pointed and 2003; Ross et al., 2010; Shi et al., 2012). A high degree of roundness produced, posterior ones obtuse, and sides coarsely serrated; elytra of the amber and Gastrochaenolites borings on the surface indicate wider than pronotum, elongated and parallel-sided anteriorly, each that it was reworked before deposition and the age is therefore with ten longitudinal rows of conspicuous, rounded window cells considered as mid-Cretaceous, circa 100 Ma. Amber has been found decreasing to eight or nine rows in posterior half of elytron below in several districts of Myanmar, but the current supply is from discal fold. Myitkyina District, Kachin State, in the Hukawng Valley of northern Comparison. The new genus is readily differentiated on the form of Myanmar; an active mine is located near Noije Bum Village, Tanaing the forebody like other archostematans in Burmese amber. The (Tanai) Township (Kania et al., 2015, fig. 1; Jarzembowski et al., setal fringes, however, may not be evident if found as articulated 2017a; fig. S1). The amber is extracted and worked by local peo- adpression fossils, e.g. amongst the numerous beetles from late ple and is prepared for the foreign jewelry trade so that inclusions Mesozoic Asian lake deposits. The elytral features will separate it such as the holotype described below may be over prepared. With from other archostematans if discovered in disarticulated assem- the current trade interest in this deposit, however, useful additional blages, e.g. elytral concentrates in UK Lower Cretaceous paralic finds may be expected. deposits. Merging of the outer cell rows is also seen in some recent cupedine genera (Cupes [] and Rhipsideigma Neboiss, fi fi 3. Material and method 1984) but is con ned to the anterior one fth of the elytron.

The specimens were examined under an Olympus SZX7 binoc- The tuberculate cuticle, exposed metatrochantins, long ular microscope with fibreoptics and top and bottom illumination; antennae inserted on top of head, expanded pronotum, bilobed they were photographed with a Zeiss Axiocam 506 digital camera fourth tarsomeres and tegular abdominal ventrites readily identify with Combine ZP software mounted on a Zeiss AX10 Zoom.v16 the new genus as an archostematan and cupedine sensu lato ‘ ’ binocular microscope. Only standard degreasing and wetting were excluding mesocupedins ; several Cretaceous genera have two undertaken during examination and glycerol under a cover slip was pairs of head protuberances but these are not as prominent, even applied temporarily to reduce surface interference. Drawings were allowing for differential preservation (Tan and Ren, 2009). No prepared from both photographs and specimens by hand (EAJ). closely similar form is evident in the monographical treatment of Drawing conventions are: solid line, distinct margin; dashed, fossil cupedines (including Cenozoic as well as Mesozoic species) indistinct or damaged; dotted, extrapolated; dashed and dotted, by Kirejtshuk et al. (2016). Nevertheless, parameres with apical folded. The abbreviations used are NIGP, Nanjing Institute of Ge- hooks and no ventro-marginal spines, as in the uniquely preserved ‘ ’ ology and Palaeontology, Chinese Academy of Sciences; P1, P2 head aedeagus, are autapomorphies of crown priacmins and bifurcate protuberances behind the antennal insertions and over the eyes processes are present in the recent genus Paracupes Kolbe, 1898 € respectively. (Vulcano and Pereira, 1975; Hornschemeyer, 2009). Our discovery For morphology, we follow terminology used in Jarzembowski is perhaps not surprising considering -like and Cupes-like et al. (2017a) for the dorsum (Fig. S1) and Jarzembowski et al. aedeagi have been reported in some late Mesozoic adpressions (2016) for the venter (Figs. S2 and S3). The prefixes pro-, meso- (Kirejtshuk et al., 2016). South American Paracupes, however, and and meta- on segments refer to the fore, middle and hind legs Paracupes-like Mallecupes Jarzembowski, Wang and Zheng, 2017b respectively and on sclerites (plates) to their corresponding in Burmese amber have only one pair of head protuberances thoracic segments. (P1s); also, the temples lack hairy extensions in the large-eyed All taxonomic acts established in the present work have been Mallecupes, and the antennal bases (scapes) are directed laterally registered in ZooBank LSID (see below), together with the elec- and not forward as in the smaller Barbaticupes, although the latter tronic publication under urn:lsid:zoobank.org:pub:EBADFEC4- may be partly taphonomic (Jarzembowski et al., 2017b). Paracupes, FB6A-4D5E-82F1-F9326939E8AF unlike Barbaticupes, has ten unforshortened rows of window cells. Finally, the Paracupes-like Cupopsis Kirejtshuk, Nel and Kirejtshuk, 2016 from the Upper Cretaceous of the USA only has nine rows of 4. Systematic palaeontology window cells, unlike the basic ten in Barbaticupes, in addition to differences in head morphology, notably the absence of P2s, and Class: INSECTA Linnaeus, 1758 rounded head with elongate, large eyes. The setose processes Order: COLEOPTERA Linnaeus, 1758 bordering the eyes in Barbaticupes are distinct from the corre- Suborder: ARCHOSTEMATA Kolbe, 1908 sponding pointed spines in Mallecupoides Jarzembowski, Wang & Family: CUPEDIDAE Laporte, 1836 Zheng, in press. We therefore propose a new genus. Subfamily: CUPEDINAE Laporte, 1836 Remark. The family and subfamily are used in the broad paleonto- Barbaticupes combertiae sp. nov. logical sense, i.e. including ommatines and priacmins respectively. (urn:lsid:zoobank.org:act:F5A9EEBB-379C-4E9E-8CE1- 1AED99F26231) Barbaticupes gen. nov. (Figs. 1, 2) (urn:lsid:zoobank.org:act:133B5D28-7EF3-422D-A33D-5DC7BD6C3DD0) Derivation of name. After the late Claudia Comberti of Oxford Uni- Type species. Barbaticupes combertiae gen. and sp. nov., by versity, researcher in climate change. monotypy. 88 E.A. Jarzembowski et al. / Cretaceous Research 80 (2017) 86e90

Fig. 1. Barbaticupes combertiae gen. nov. and sp. nov., holotype; Noije Bum, mid Cretaceous. A, B photographs and C line drawing of dorsal (left) and ventral view (right); D, E photograph and drawing of aedeagus (dorsal view). Setal fringe indicated by outline on forebody; window cells omitted in C (see A, B); obscured area (over prepared) indicated by double lines (C, top right).

Holotype. NIGP 166314, beetle body with aedeagus preserved in Locality and horizon. Reportedly mined near Noije Bum Village, yellow amber, partially obscured and polished away (front, left). Tanaing Township, Myitkyina District, Kachin State, northern Paratype. NIGP 166315, beetle body in tumbled and polished, yel- Myanmar, (26 150 N., 96 330 E.); unnamed horizon, mid Creta- low and brown amber cabochon with other insect (beetle and bug) ceous, upper Albian or lower Cenomanian. inclusions and fern hairs. Other material. None known. E.A. Jarzembowski et al. / Cretaceous Research 80 (2017) 86e90 89

Fig. 2. Barbaticupes combertiae gen. and sp. nov., paratype; Noije Bum, mid Cretaceous. A, oblique photograph and B, explanatory line drawing of dorsolateral view with window cells in rows 7e10 to show merging; setation omitted on forebody for clarity (i.e. in front of elytra).

Diagnosis. As for genus (key taxonomic features are head architec- spines; posterior angles obtuse, bluntly pointed; sides with several ture (setose processess in the temple-ocular area; prominent P1s coarse serrations. Prothoracic sutures (notopleural and sterno- and P2s), foreshortened elytral cell rows (distal merging in sub- pleural) just discernible. Pterothorax ventrally with subtrapezoidal costal (outer) area), and apparently forward-directed antennae. metaventrite (indicated by metakatepisternal (posterior) suture Description. Small cupedine, body elongate (length: width ratio >3:1), and incomplete metanapleural (lateral) clefts) and short, broad flattened, hirsute (covered with small setae, especially anteriorly), metatrochantins. Elytra (forewings) elongate, overlapping hind- spines and spurs developed locally (especially on tibio-tarsi of legs). body, shoulders (humeri) rectangular and rounded like elytral Cuticle black or brown and slightly tuberculate, latter passing into apices; ten longitudinal rows of comparatively large, slightly serration locally (especially on edge of elytron). maculated (laterally tuberculated), window cells present, the outermost two rows (nine and ten) merging below discal fold at six Head short and broad with well developed, laterally rounded tenths of length of elytron from base and then merging with row eyes; also well developed, rounded or pointed protuberances pre- eight at some nine tenths. Elytral disc (upper surface) flattened/ sent behind antennal bases and mesad of eyes; temples bluntly slightly curved with three longitudinal veins dorsally (first anal pointed, wider than eyes and curved anteriorly with brush-like (A1), anterior cubital (CuA) and media (M)) as well as intercalaries, fringe of dense setae. Neck pronounced and narrower than head. inner two veins joining first then outer one before elytral apex. Antennae long, inserted on top of head, reaching mid length of Epipleural rim narrow, without cells, epipleuron widening slightly body, 11-segmented, filiform; scape (first attached antennal anteriorly. Hindwings folded, small portion protruding on paratype. segment) long and broad; third antennomere (segment) not longer Legs. Mesocoxa long, metacoxa broad and triangular with small than second and fourth combined; last (11th) spindle shaped. trochanter; procoxae apparently separated. Femora stout, profemur Mandibles (jaws) well developed, curved anteriorly, inner dentition extending beyond lateral edge of prothorax, mesofemur extending unclear. Maxilla long, palp extending beyond mandibles. slightly beyond lateral edge of hindbody but metafemur not beyond edge of abdomen. Tibiae thin, meso- and metatibiae clearly spurred Thorax. Prothorax broader than long, wider than head, narrower and slightly curved; metatibia, unlike mesotibia, longer than cor- than hindbody. Pronotum laterally depressed and ridged medially responding femur. Tarsi with distinctly elongated first (articulated) with single peak; anterior angles extended as pointed, sharp tarsomeres; second and third tarsomeres short, especially third; 90 E.A. Jarzembowski et al. / Cretaceous Research 80 (2017) 86e90 fourth bilobed (underneath) and 5th slightly elongated, clawed; Cruickshank, R.D., Ko, K., 2003. Geology of an amber locality in the Hukawng Valley, e posterior tarsi longer than corresponding tibiae. northern Myanmar. Journal of Asian Earth Sciences 21, 441 455. Fabricius, J.C., 1801. Systema Eleutheratorum secundum ordines, genera, species: Abdomen. Elongate, apex acute, bluntly rounded; ventrite 5 (pos- adiectis synonymis, observationibus, descriptionibus. Tomus II. Impensis bib- terior-most) twice as long as ventrite 4, equal in length to 3 and 4 liopoli academici novi, Kiliae, Germany, 607 pp. € combined; ventrite 1 complete, shorter than 5, longer than other Hornschemeyer, T., 2009. The species-level phylogeny of archostematan beetlese fl where do Micromalthus debilis and Crowsoniella relicta belong? Systematic ventrites; all ventrites slightly tegular (i.e. not at). Entomology 34, 533e558. Male genitalia (paratype only). Partially extruded behind elytral Hornschemeyer,€ T., 2016. Archostemata Kolbe, 1908. Handbuch der Zoologie IV, apices; aedeagus trilobed, penis (median lobe) not extending 41e43. fl Hünefeld, F., Marvaldi, A.E., Müller, B., Lawrence, J.F., Beutel, R.G., 2011. The male beyond anking parameres (lateral lobes); small, inward pointing, postabdomen of the “ancestral” archostematan beetle Tetraphalerus bruchi subapical hook present on each paramere; dorsally, two elongate Heller, 1913 (Ommatidae) and its phylogenetic significance. Struc- (bifurcate) processes extend slightly beyond parameres. ture and Development 40 (2), 146e158. Measurements. Body length (from tip of mandibles to apex of elytra) Jarzembowski, E.A., Yan, E.V., Wang, B., Zhang, H., 2013. Brochocolein beetles (Insecta: Coleoptera) from the Lower Cretaceous of northeast China and 4.7e5.7 mm, width (across (folded) elytra) 1.2e1.3 mm; aedeagus southern England. Cretaceous Research 44, 1e11. 0.2 mm long. Jarzembowski, E.A., Wang, B., Zheng, D., 2016. An amber double first: a new bro- Remarks. The genital details should help to differentiate this species chocolein beetle (Coleoptera: Archostemata) from northern Myanmar. Pro- ceedings of the Geologists' Association 127, 676e680. from other amber congeners if found; the bifurcate process in the Jarzembowski, E.A., Wang, B., Zheng, D., 2017a. A new ommatin beetle (Insecta: paratype is long, as in recent Paracupes sensu stricto (Vulcano and Coleoptera) with unusual genitalia from mid-Cretaceous Burmese amber. fi Cretaceous Research 71, 113e117. Pereira, 1975: g. 28) versus short in Cupes [Tenomerga](Sharp and fi fi Jarzembowski, E.A., Wang, B., Zheng, D., 2017b. The rst cupedine beetle from Muir, 1912: pl. LIX, g. 104, d). Burmese amber (Coleoptera: Cupedidae). Comptes Rendus Palevol 16, Occurrence: Currently only known from the type locality. 241e247. Jarzembowski, E. A., Wang, B., Zheng, D., In Press. A new spiny reticulated beetle (Coleoptera: Cupedidae) from Cretaceous Burmese amber. Proceedings of the Geologists' Association.. 2017 5. Conclusion Kania, I., Wang, B., Szwedo, J., 2015. Dicranoptycha Osten Sacken, 1860 (Diptera, Limoniidae) from the earliest Upper Cretaceous Burmese amber. Cretaceous Research 52, 522e530. Ever since Sharp and Muir's (1912) monograph, no description Kirejtshuk, A.G., Nel, A., Kirejtshuk, P.A., 2016. of the reticulate beetles of of a beetle species has been deemed complete without reference to the Subfamily Cupedinae (Coleoptera: Archostemata) with a review of their the male genitalia, and the phylogenetic value of this in archaic historical development. Invertebrate Zoology 13, 61e190. archostematans is beginning to be appreciated (Hünefeld et al., Kirejtshuk, A.G., Ponomarenko, A.G., 2015. Taxonomical list of fossil beetles of the suborders Cupedina, Carabina and Scarabaeina (Part 1). http://www.zin.ru/ 2011). Obtaining such information in fossil beetles, however, is animalia/coleoptera/eng/paleosy0.htm (accessed 3.10.2015). generally much harder, even if the genital capsule happens to be Kolbe, H., 1908. Mein System der Coleopteren. Zeitschrift für wissenschaftliche e extruded by fluid expulsion during diagenesis, as in this case. The Insekten-Biologie 4, 116 400. Kolbe, H.J., 1898. Eine neue Gattung der Cupesiden. Entomologische Nachrichten 24 first discovery of a cupedine aedeagus in Burmese amber is there- (12), 179e180. fore significant and Barbaticupes n. gen. not only adds to the Laporte, F. L. De, 1836. Histoire naturelle des animaux articules, annelides, crustaces, arachnids, myriapodes et insects. R Dumenil, Paris, France, 324 pp. Cretaceous diversity of the reticulated beetles sensu stricto, but also Lawrence, J.F., Slipinski, A., 2013. Australian Beetles. Volume 1: morphology, clas- demonstrates the value of small fossil inclusions. In this case, the sification and keys. Commonwealth Scientific and Industrial Research Organi- genitalia suggest affinity with extant Paracupes sensu stricto (type: zation Publishing, Collingwood, Victoria, Australia, 576 pp. Paracupes brasiliensis Kolbe, 1898) despite the relatively specialized Linnaeus, C., 1758. Systema naturae per regni tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. gross morphology of Barbaticupes. Recent work, however, suggests Tomus I. (10th revised ed.). Laurentius Salvius, Stockholm, Sweden iv þ that the living species and Cretaceous fossils previously referred to 824 pp. Paracupes are not closely related (Kirejtshuk et al., 2016). The exact Neboiss, A., 1984. Reclassification of Cupes Fabricius (s. lat.), with descriptions of fi new genera and species (Coleoptera: Cupedidae). Systematic Entomology 9, af nities therefore need further investigation but living arch- 443e477. ostematans, like Burmese amber inclusions, are not often found Ponomarenko, A.G., 1973. On subdividing the order Coleoptera into suborders. In: and the DNA of Paracupes has yet to be described to help resolve Problems of Insect Palaeontology. Lectures from the 24th Annual Readings in e generic relationships. There is growing evidence of early involve- Memory of N. A. Kholodkovsky, 1 2 April, 1971. Nauka, Leningrad, Russia, pp. 78e89 (in Russian). ment by beetles in pollination (Wang et al., 2013) therefore the Ross, A., Mellish, C., York, P., Crighton, B., 2010. Burmese amber. In: Penney, D. (Ed.), brush-like hairy fringes of any new finds of Barbaticupes should be Biodiversity of Fossils in Amber From the Major World Deposits. Siri Scientific e checked accordingly. Press, Manchester, pp. 208 235. Sharp, D., Muir, F.A.G., 1912. The comparative anatomy of the male genital tube in Coleoptera. Transactions of the Entomological Society of London 60, Acknowledgements 477e642. Shi, G., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Yang, M., Lei, W., Li, Q., Li, X., 2012. Age constraint on Burmese amber based on U-Pb dating of zircons. We thank Xiangdong Zhao, Jingjing Tan (NIGPAS) and Fred Cretaceous Research 37, 155e163. Clouter (UK) for help with images, and Bobbie Winter-Burke Tan, J.J., Ren, D., 2009. Mesozoic archostematan fauna from China. Science Press, (NHMUK) for library assistance. We are indebted to two anony- Beijing, China v þ 347 pp. Vulcano, M.A., Pereira, F.S., 1975. Cupesidae (Coleoptera). Arquivos do Instituto mous reviewers and the editor for their time and comments. Biologico de Sao~ Paulo 42, 31e68. This research (grant nos in parentheses) was supported by the Wang, B., Xia, F., Wappler, T., Simon, E., Zhang, H., Jarzembowski, E.A., Szwedo, J., National Natural Science Foundation of China (41572010, 41622201, 2015. Brood care in a 100-million-year-old scale insect. eLife 4, e05447. Wang, B., Zhang, H., Jarzembowski, E.A., 2013. Early Cretaceous angiosperms and 41688103), the Youth Innovation Promotion Association of CAS beetle evolution. Frontiers in Plant Science 4, 1e6. (2011224) and CAS (XDPB05). This is a Leverhulme Emeritus Fellowship contribution for EAJ.

References Appendix A. Supplementary data

Crowson, R.A., 1955. The natural classification of the families of the Coleoptera. N. Supplementary data related to this article can be found at http://dx.doi.org/10. Lloyd, London, UK, 187 pp. 1016/j.cretres.2017.08.015.