Another Amber First: a Tiny Tetraphalerin Beetle

Cretaceous Research 78 (2017) 84e88

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Cretaceous Research

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Short communication Another amber ﬁrst: A tiny tetraphalerin beetle (Coleoptera: Archostemata) in Myanmar birmite

* 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, Nanjing 210008, China b Department of Earth Sciences, Natural History Museum, 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, University of Hong Kong, Hong Kong Special Administrative Region, China article info abstract

Article history: A new tetraphalerin beetle, Tetraphalerus lindae sp. nov. (Insecta: Coleoptera: Archostemata) is described Received 26 April 2017 from mid-Cretaceous Burmese amber from northern Myanmar. This is the ﬁrst species of this Jurassic- Received in revised form recent genus of archaic beetles to be described from amber inclusions, and is the ﬁrst tetraphalerin 17 May 2017 cupedid from Burmese amber. This small, unusual Cretaceous Tetraphalerus is considered to belong to the Accepted in revised form 22 May 2017 T. bruchi species group of this now relict South American genus. Available online 31 May 2017 © 2017 Elsevier Ltd. All rights reserved.

Keywords: Burmese amber Myanmar Coleoptera Cupedid Tetraphalerus New species

1. Introduction Archostematans are, nevertheless, scarce in Burmese amber which is surprising considering that most recent archostematans are With over 300,000 described species, beetles (Insecta: Coleop- associated with wood in both active stages (larval and adult) and tera) are by far the largest order in the animal kingdom; the sometimes the adults visit flowers. Here we describe a unique Cupedina, however, is the smallest and most archaic beetle ‘sub- archaic beetle from this deposit which shows affinity with the still- order’, totalling just over 100 living species, and now commonly living tetraphalerins, the sister of the ommatins or typical omma- split into the smaller suborders Archostemata and Myxophaga tines (Yan, Beutel and Ponomarenko, 2017). Opinion is divided as to (Hornschemeyer,€ 2005). Several hundred fossil species have been whether ommatines are a subfamily of cupedids sensu lato, the described from the Permian onwards and archostematans are reticulated beetles (Kirejtshuk and Ponomarenko, 2015), or a notable constituents of Mesozoic insect faunas, even occurring in separate family (Beutel et al., 2008). For consistency with work on regions from where they have now vanished (such as Europe; the Burmese ommatins (Jarzembowski et al., 2017), we consider Kirejtshuk and Ponomarenko, 2015). Such finds are usually pre- cupedids and ommatines in the broad sense, cupedids including served as adpressions, some exceptionally well preserved as in ommatines, and ommatines including tetraphalerins and omma- northeastern China, but they have also been discovered recently as tins (as well as extinct brochocoleins) as tribes (cf. Tan et al., 2012). amber inclusions in northern Myanmar (Xia et al., 2015). The beetle described below belongs to an uncommon species, only a single example being known so far from over 100,000 inclusions examined. Extant tetraphalerins are also rare insects, two species now only occurring in South American brushland, unlike in * Corresponding author. State Key Laboratory of Palaeobiology and Stratigraphy, þ Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, the Mesozoic when they were widespread across Eurasia (20 Nanjing 210008, China. species; Soriano and Delclos, 2006). Like the ommatins, they are a E-mail address: [email protected] (E.A. Jarzembowski). http://dx.doi.org/10.1016/j.cretres.2017.05.023 0195-6671/© 2017 Elsevier Ltd. All rights reserved. E.A. Jarzembowski et al. / Cretaceous Research 78 (2017) 84e88 85 relict group of “living fossils”,no‘Tertiary’ tetraphalerins having yet metafemora short, not extending past edge of body; elytral disc been described, Tetraphalerites Crowson, 1962 now being consid- with little relief; rows of window punctures present on elytra but ered a brochocolein, the extinct sister of the ommatins and tetra- generally inconspicuous. phalerins (Kirejtshuk et al., 2016). Remarks. At the time of Ponomarenko's 1969 classical monograph of fossil archostematans, only a single species of Cretaceous Tetra- 2. Geological setting phalerus was known (discussed below), the others being earlier (Jurassic) or recent. The diagnosis is therefore augmented by Burmese amber (burmite or birmite) from northern Myanmar Ponomarenko and Yan (2012) and Tan et al. (2012) combining contains the most diverse biota in amber (fossil resin) known from adpression and living features, the inclusion described herein the Cretaceous (Wang et al., 2015). 34 orders are represented, adding small size (medium size is about 12 mm body length, cf. beetles being one of the most diverse, but most species are unde- Ponomarenko and Yan, 2012), the eye protuberance (‘P2’), and head € scribed. The amber is currently dated stratigraphically and radio- length: width ratio (cf. Hornschemeyer, 2009). The diagnosis is metrically from late Albian to early Cenomanian (Cruickshank and abridged, e.g. excluding family group characters, viz. adjacent Ko, 2003; Ross et al., 2010; Shi et al., 2012). The amber shows signs procoxae present and abdomen with flat (coplanar) ventrites, as of reworking and we therefore consider the age as mid Cretaceous well as fourth tarsomere not being bilobed, and long third anten- (circa 100 Ma). nomere present, plus relatively short antennae. Amber has been found in several districts of Myanmar over the years, but the current supply is from Myitkyina District, Kachin Earlier, Ponomarenko (1969) cautioned that, notwithstanding State, in the Hukawng Valley of northern Myanmar: a working the development of ridges and furrows on the forebody, the only mine is located near Noije Bum Village in Tanaing (Tanai) Township readily observable difference between fossil tetraphalerins and (Kania et al., 2015: fig. 1; Jarzembowski et al., 2017: fig. S1). The ommatins is the short temples on the latter (not exceeding half the amber is prepared for the foreign jewellery trade and inclusions are eye diameter, as recently seen in Burmese amber Omma lii affected by polishing as well as natural deformation so that addi- Jarzembowski, Wang and Zhang, 2017). In recent cladistics ana- € tional specimens for taxonomic study are often useful. lyses, Hornschemeyer (2009) recognized seven possible autapomorphies for crown-group tetraphalerins, but Tan et al. 3. Material and method (2012), including fossils (adpressions), recognised only two (pres- ence of antennal grooves ventrolaterally and lack of gular sutures The specimens were examined under an Olympus SZX7 binoc- on head). Unfortunately, as noted earlier, these head characters can fi ular microscope with fibreoptics and top and bottom illumination; be dif cult to observe, even in amber inclusions. The wide labrum they were photographed with a Zeiss Axiocam 506 digital camera and short metafemur were suggested proxies to which may now be with Combine ZP software mounted on a Zeiss AX10 Zoom.v16 added the head ratio (in combination) and shape of the eye pro- binocular microscope. Drawings were prepared from both photo- tuberance (see above). graphs and specimen by hand (EAJ). Only standard degreasing and Tetraphalerus lindae wetting were undertaken during examination; glycerol under a sp. nov. cover slip was applied temporarily to reduce surface interference. (urn:lsid:zoobank.org:act:BDE6FC25-64B0-4CA5-96AC- For morphology, we follow the terminology in Jarzembowski et al., D3345164B437) 2017: figs 2, S2. Drawing conventions are: solid line, distinct Figs. 1, 2 margin; dashed, indistinct or damaged; dotted, extrapolated. The Derivation of name. After Linda Wang (Nanjing), daughter of B. W. abbreviations used are NIGP and NIGPAS, Nanjing Institute of Ge- Holotype. NIGP 166152, beetle body in tumbled and polished small, ology and Palaeontology, Chinese Academy of Sciences. yellow, amber cabochon. Locality and horizon. Probably mined near Noije Bum Village, 4. Systematic palaeontology Tanaing Township, Myitkyina District, Kachin State, Myanmar, 26 150 N., 96 330 E.; unnamed horizon, mid Cretaceous, upper Albian Class INSECTA Linnaeus, 1758 or lower Cenomanian. Order COLEOPTERA Linnaeus, 1758 Diagnosis. Very small Cretaceous species of Tetraphalerus with Suborder ARCHOSTEMATA Kolbe, 1908 elongate prothorax as well as head, ridge-like paired longitudinal Family CUPEDIDAE Laporte, 1836 s.l. protuberances on the latter developing spines as well as tubercles, Subfamily OMMATINAE Sharp & Muir, 1912 clavate antennae reaching prothorax anteriorly, large anterior Tribe TETRAPHALERINI Crowson, 1962 ventrite on abdomen with posterior ventrite only a little larger than penultimate one, and moderately well-developed elytral venation Remark. With the exclusion of Tetraphalerites (Section 1) this tribe and epileuron on the broad hindbody. (sometimes subfamily) is now monobasic. Description. Small, black beetle, 4.7 mm long (from anterior end of Genus Tetraphalerus Waterhouse, 1901 mandibles to posterior end of folded elytra), 1.3 mm wide (across Type species: Tetraphalerus wagneri Waterhouse, 1901 by mono- folded elytra). Body flattened (dorsoventrally); hirsute, covered typy; recent, Brazil. with small setae (especially on fronto-clypeal area and fronting labrum); spurred and spinose locally (notably above and behind the Diagnosis. Small to medium-size beetles with dorso-ventrally eyes). Cuticle generally coarsely tuberculate. compressed body; head long (1.3[þ] longer than wide), antennal grooves sometimes visible; antennae with apical anten- Head some 1.4 longer than wide with well-developed neck; nomere (flagellomere) rounded distally, appearing inflated; eyes temples subrectangular, short; eyes elongate and rounded laterally, forward with distance between posterior end of head and eyes prominent ventrally, appearing small dorsally as each overlapped equal to or more than half the eye diameter; ridge-like protuber- by longitudinal, ridge-like protuberance (P2) above eye which ance above eye present; mandibles very long and distinctly pro- continues posteriorly (P3) adorned by 3/4 backward-directed truding (prognathous); labrum broad, as wide as clypeus and spines. Antennae longer than head but not reaching middle of truncate anteriorly, gula sutures present or absent; meso- and 86 E.A. Jarzembowski et al. / Cretaceous Research 78 (2017) 84e88

Fig. 1. Tetraphalerus lindae sp. nov., photographs of holotype (under glycerol) in A, dorsolateral view; B, close up of forebody; C, ventral view. E.A. Jarzembowski et al. / Cretaceous Research 78 (2017) 84e88 87

small Cretaceous species start at 5.7 mm (Tetraphalerus mongolicus Ponomarenko, 1986) from the Lower Cretaceous (Berriasian-Val- anginian) of western Mongolia (Gurban-Erenny-Nuru). Unfortu- nately, judging from the photograph (Ponomarenko, 1986: pl. VI, fig. 1), the unique specimen of this adpressed species is poorly preserved although, in addition to the antennae, it differs in the proportions of the ventrites and in having a squat (short and broad) prothorax. The original report states, however, that it is comparable to a somewhat larger species at 6e6.5 mm long, Tetraphalerus verrucosus Ponomarenko, 1966 from the Valanginian-Hauterivian of Baissa (Siberia). T. verrucosus is better preserved judging by the excellent photograph (by D. E. Shcherbakov) in Ponomarenko (2002: fig. 237). A similar elytral venation is clearly visible in the Siberian species, but the prothorax is ‘almost square’ unlike the elongated shape of T. lindae sp. nov., plus the head is broader. Whilst recognizing that Mesozoic tetraphalerin fossils resem- bled each other more than any recent species, Ponomarenko (1969) nevertheless recognized two species groups (‘series’) dating back to the Upper Jurassic based on extant forms: the type species of the genus (T. wagneri) and Tetraphalerus bruchi Heller, 1913 from Argentina. They differed in head morphology, the former group Fig. 2. Tetraphalerus lindae sp. nov., holotype, drawings of dorsal (left) and ventral with paired transverse protuberances and latter with longitudinal (right) views. Representative window punctures shown in oblique strip on left elytron. ones. T. verrucosus falls in the T. bruchi group, as does T. lindae sp. nov.; in the former species, however, the protuberances appear prothorax, 11-segmented, clavate, inserted on side of head; 3rd linear rather than ridged as in the latter, and the eyes of the latter antennomere slightly elongated (longer than 2nd and 4th), 11th are located more posteriorly on the head, as in the Jurassic pro- rounded. Lateral insertion of 1st antennomere (scape) overlapped genitor (Ponomarenko, 1969: fig. 54). T. lindae sp. nov. also differs by ridge-like protuberance (P1). Lateral notch between P1 and P2 from T. bruchi itself (in addition to smaller size) in that the latter has broad, suggestive of antennal groove. Labrum broad, straight shorter (filiform) antennae only just reaching the pronotum, the anteriorly. Mandibles pointing forward with three vertical teeth eyes are located more anteriorly on the head, the prothorax is less (although only two visible dorsoventrally), i.e. tridentate. Maxillary rounded, and the hindbody narrower (see Beutel et al., 2008; palps narrow and extending beyond labrum. Hünefeld et al., 2011). The elongate prothorax of T. lindae sp. nov. Thorax. Pronotum longer than broad with rounded corners/ actually resembles that of T. wagneri (Hornschemeyer,€ 2005). edges anteriorly, slightly obtuse posteriorly, only slightly wider than head, much narrower than abdomen. Procoxae posteromesal 6. Conclusion on prothorax. Profemur thick and extending beyond edge of prothorax; protibia thinner, shorter and spurred; protarsus shorter The solitary find of Tetraphalerus lindae sp. nov. in Burmese again, not lobed, 5th tarsomere elongated and clawed. Meso- amber, the first amber tetraphalerin inclusion and youngest metathoracic junction obscured by amber fracture but mesocoxae described fossil tetraphalerin to date, shows a unique combination large, adjacent and mesofemur not extending beyond edge of of features distinguishing it from both recent and fossil species hindbody. Metaventrite broad, apparently short like small meso- (Sections 4, 5). It can, nevertheless, be tentatively assigned to the ventrite. Metatrochantin broad, spindle-shaped. Metacoxa very Tetraphalerus bruchi species group which ranged from Spain to large, subtriangular. Metafemur broad and longer than spurred China and Siberia in the Early Cretaceous as shown by the metatibia but not extending to epipleuron; metatarsus long and adpression fossil record (Soriano and Delclos, 2006; Tan and Ren, thin also with elongated first tarsomere. 2009). Today, Tetraphalerus is confined to a few countries in Elytron overlapping slightly waisted hindbody; base and humerus South America with a much reduced species diversity. With a fossil (shoulder) slightly curved and rounded; apex broadly rounded; gap in the ‘Tertiary’, it resembles a marginalised Gondwanan relic lateral (anterior) margin slightly curved; disc flattened with three which belies its Laurasian past. The pattern is not unique. For weak longitudinal veins (1A-M with intercalaries) and short 2A; example, living moss bugs (Hemiptera: Coleorrhyncha) are now discal fold fairly abrupt below M; epipleural rim moderately narrow, confined to the southern continents, but have a Mesozoic fossil tapering posteriorly; six discernible rows of rounded window record in Eurasia as adpression and inclusion fossils, the pattern punctures on disc and about four above epipleural rim, lower one being attributed to the mid-Cretaceous biotic crisis (Szwedo et al., with larger, somewhat elongated and irregular maculated cells. 2011). Small fossils can thus contribute to our understanding of Abdomen rounded apically with coplanar ventrites, 1st and 5th insect biogeography. Never numerous, more well-preserved ex- longer than others, 1st markedly so, 5th only slightly longer than amples are needed of Tetraphalerus to observe fine details including 4th. antennal grooves, gula sutures and terminalia. Tiny inclusions in Burmese amber are already augmenting the sedimentary fossil 5. Comparison record.

T. lindae sp. nov. differs from all other congeners in having Acknowledgements clavate as opposed to moniliform or ﬁliform antennae (cf. Tan and Ren, 2009: 295). At less than 5 mm in body length, it is a very small We thank Messrs Xiangdong Zhao (NIGPAS) and Fred Clouter species for the genus, recent ones being 11e15 mm long (UK) for help with images, and Lorraine Portch (NHMUK) for library (Ponomarenko, 1969) and large ones from the Cretaceous (all Early) assistance. We are indebted to two anonymous reviewers and the occasionally exceeding 20 mm (Soriano and Delclos, 2006). The editor for their time and comments. 88 E.A. Jarzembowski et al. / Cretaceous Research 78 (2017) 84e88

This research (grant nos in parentheses) was supported by the Linnaeus, C., 1758. Systema naturae per regni tria naturae, secundum classes, Chinese Academy of Sciences (XDPBO5), the National Natural Sci- ordines genera, species, cum characteribus, differentiis, synonymis, locis (10th revised ed.), vol. 1. Laurentius Salvius, Stockholm, Sweden iv þ 824 pp. ence Foundation of China (41572010, 41622201, 41688103) and the Ponomarenko, A.G., 1966. New beetles of the family Cupedidae (Coleoptera) from Youth Innovation Promotion Association of CAS (2011224). This is a Mesozoic deposits of Transbaikalia. Revue d'Entomologie de l'URSS 45, e Leverhulme Emeritus Fellowship contribution for EAJ. 138 143. Ponomarenko, A.G., 1969. Historical development of the archostematan co- leopterans. Trudy Paleontologicheskogo Instituta 125, 1e241 (in Russian). Ponomarenko, A.G., 1986. Scarabaeida (¼Coleoptera). In: Rasnitsyn, A.P. (Ed.), In- References sects in the Early Cretaceous Ecosystems of West Mongolia. Nauka, Moscow, Russia, pp. 84e107 (in Russian). Beutel, R.G., Ge, S., Hornschemeyer,€ T., 2008. On the head morphology of Tetra- Ponomarenko, A.G., 2002. Superorder Scarabaeidea Laicharting, 1781. In: phalerus, the phylogeny of Archostemata and the basal branching events in Rasnitsyn, A.P., Quicke, D.L.J. (Eds.), History of Insects. Kluwer, Dordrecht, The e Coleoptera. Cladistics 24, 270e298. Netherlands, pp. 164 176. Crowson, R.A., 1962. Observations on the beetle Family Cupedidae, with de- Ponomarenko, A.G., Yan, E.V., 2012. Revision of some early Mesozoic beetles from e scriptions of two new fossil forms and a key to the recent genera. Annals and China. Acta Palaeontologica Sinica 51 (4), 475 490. Magazine of Natural History (13) 5, 147e157, pls III, IV. Ross, A., Mellish, C., York, P., Crighton, B., 2010. Burmese amber. In: Penney, D. (Ed.), fi Cruickshank, R.D., Ko, K., 2003. Geology of an amber locality in the Hukawng Valley, Biodiversity of Fossils in Amber from the Major World Deposits. Siri Scienti c e northern Myanmar. Journal of Asian Earth Sciences 21, 441e455. Press, Manchester, pp. 208 235. Heller, K.M., 1913. Ein neue cupedide. Wiener entomologische Zeitung 32, 235e237. Sharp, D., Muir, F.A.G., 1912. The comparative anatomy of the male genital tube in e Hornschemeyer,€ T., 2005. Archostemata Kolbe, 1908. Handbuch der Zoologie IV, Coleoptera. Transactions of the Entomological Society of London 60, 477 642. pp. 29e42. Shi, G., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Yang, M., Lei, W., Li, Q., Li, X., Hornschemeyer,€ T., 2009. The species-level phylogeny of archostematan beetles e 2012. Age constraint on Burmese amber based on U-Pb dating of zircons. e where do Micromalthus debilis and Crowsoniella relicta belong? Systematic Cretaceous Research 37, 155 163. Entomology 34, 533e558. Soriano, C., Delclos, X., 2006. New cupedid beetles from the Lower Cretaceous of Hünefeld, F., Marvaldi, A.E., Müller, B., Lawrence, J.F., Beutel, R.G., 2011. The male Spain and the palaeogeography of the family. Acta Palaeontologica Polonica 51, e postabdomen of the “ancestral” archostematan beetle Tetraphalerus bruchi 185 200. fi Heller, 1913 (Ommatidae) and its phylogenetic significance. Arthropod Struc- Szwedo, J., Azar, D., Ziade, K., 2011. The rst Progonocimididae (Insecta: Hemiptera: ture and Development 40 (2), 146e158. Coleorrhyncha) from Lower Cretaceous Lebanese amber. Insect Systematics and e Jarzembowski, E.A., Wang, B., Zheng, D., 2017. A new ommatin beetle (Insecta: Evolution 42, 161 177. Coleoptera) with unusual genitalia from mid-Cretaceous Burmese amber. Tan, J., Ren, D., 2009. Mesozoic Archostematan Fauna from China. Science Press, þ Cretaceous Research 71, 113e117. Beijing, China v 347 pp. Kania, I., Wang, B., Szwedo, J., 2015. Dicranoptycha Osten Sacken, 1860 Diptera, Tan, J., Wang, Y., Ren, D., Yang, X., 2012. New fossil species of ommatids (Coleoptera: Limoniidae) from the earliest Upper Cretaceous Burmese amber. Cretaceous Archostemata) from the Middle Mesozoic of China illuminating the phylogeny Research 52, 522e530. of Ommatidae. BMC Evolutionary Biology 12 (113), 19 pp. Kirejtshuk, A.G., Ponomarenko, A.G., 2015. Taxonomical list of fossil beetles of the Wang, B., Xia, F., Wappler, T., Simon, E., Zhang, H., Jarzembowski, E.A., Szwedo, J., suborders Cupedina, Carabina and Scarabaeina (Part 1) (Accessed 3 October 2015. Brood care in a 100-million-year-old scale insect. eLife 4, e05447. 2015). www.zin.ru/Animalia/Coleoptera/eng/paleosy0.htm. Waterhouse, C.O., 1901. Two new genera of Coleoptera belonging to the Cupesidae e Kirejtshuk, A.G., Nel, A., Kirejtshuk, P.A., 2016. Taxonomy of the reticulate beetles of and Prionidae. Annals and Magazine of Natural History (7) 42, 520 523. the Subfamily Cupedinae (Coleoptera: Archostemata) with a review of their Xia, F., Yang, G., Zhang, Q., Shi, G., Wang, B., 2015. Amber Lives Through Time and þ historical development. Invertebrate Zoology 13 (2), 61e190. Space. Science Press, Beijing viii 198 pp. ϯ Kolbe, H., 1908. Mein System der Coleopteren. Zeitschrift für wissenschaftliche Yan, E.V., Beutel, R.G., Ponomarenko, A.G., 2017. Peltosynidae, a new beetle family fi Insekten-Biologie 4, 116e400. from the Middle-Late Triassic of Kyrgyzstan: its af nities with Polyphaga Laporte, F. L. de, 1836. Histoire naturelle des animaux articules, annelides, crustaces, (Insecta, Coleoptera) and the groundplan of this megadiverse suborder. Journal arachnids, myriapodes et insects. R. Dumenil, Paris, France, 324 pp. of Systematic Palaeontology (eprint).