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Coleoptera: Archostemata: Ommatidae) ARTICLE A miniaturized ommatid beetle in mid‑Cretaceous Burmese amber (Coleoptera: Archostemata: Ommatidae) Yan‑Da Li¹²⁵; Shûhei Yamamoto³⁶; Di‑Ying Huang¹⁷ & Chen‑Yang Cai¹⁴⁸ ¹ State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, and Center for Excellence in Life and Palaeoenvironment, Chinese Academy of Sciences, Nanjing, China. ² School of Life Sciences, Peking University, Beijing, China. ³ Hokkaido University Museum, Sapporo, Japan. ⁴ School of Earth Sciences, University of Bristol, Bristol, United Kingdom. ⁵ ORCID: http://orcid.org/0000-0002-9439-202X. E-mail: [email protected] ⁶ ORCID: http://orcid.org/0000-0002-4162-8457. E-mail: [email protected] ⁷ ORCID: http://orcid.org/0000-0002-5637-4867. E-mail: [email protected] ⁸ ORCID: http://orcid.org/0000-0002-9283-8323. E-mail: [email protected] (corresponding author) Abstract. Recent findings of ommatids from mid-Cretaceous Burmese amber have greatly increased our knowledge on the Mesozoic diversity of Ommatidae. Here, we report the first distinctly miniaturized ommatid species, Miniomma chenkuni gen. et sp. nov., entombed in mid-Cretaceous amber from northern Myanmar. This new fossil species is characterized by its small body size (less than 2 mm long) and subglobular metacoxae. Our discovery of miniaturization in extinct Ommatidae suggests a high ecological diversity of this family in the Mesozoic. Key‑Words. Archostemata; Ommatidae; Miniomma; Miniaturization; Burmese amber. INTRODUCTION our understanding of the diversity and disparity of Mesozoic ommatids. Ommatidae is a small family in the beetle Miniaturization, or the evolution towards ex- suborder Archostemata, with only three extant tremely small body size, is a commonly found phe- genera, Omma Newman and Beutelius Escalona nomenon in animals (Hanken & Wake, 1993), and et al. in Australia, and Tetraphalerus Waterhouse insects in particular (Polilov, 2016; Minelli & Fusco, in South America (Hörnschemeyer & Beutel, 2016; 2019). The upper size limit for microinsects have Lawrence & Escalona, 2019; Escalona et al., 2020). been somewhat arbitrarily determined to 2 mm, Fossil records have revealed that ommatids had partly based on the threshold of pumilistic degen- a much higher diversity during the Mesozoic, as eration (Polilov, 2016). Most of these microinsects well as wider distribution (Cai & Huang, 2017; belong to Coleoptera (beetles) or Hymenoptera Kirejtshuk, 2020). Morphologically, Ommatidae (ants, bees and wasps) (Polilov, 2016). Even though is most similar to the family Cupedidae. Recent miniaturization is quite common in beetles, such molecular phylogenetic analyses, however, have a phenomenon was previously unknown in any revealed a closer affinity between Ommatidae extant or fossil representatives in the beetle family and Micromalthidae (McKenna et al., 2015, 2019). Ommatidae. Here, we report the first miniaturized Historically, Mesozoic ommatids are only repre- member of Ommatidae from Burmese amber, which sented by compression or impression fossils, in not only increases our knowledge on the morpho- which detailed structures are not well preserved. logical diversity of this family, but also points to a Recently, various exquisitely preserved omma- high ecological diversity of Mesozoic ommatids. tids have been described from mid-Cretaceous Burmese amber, including at least 21 species in 11 genera (generic assignment according to original MATERIAL AND METHODS authors) (Ross, 2019, 2020; Jarzembowski et al., 2020; Kirejtshuk, 2020). These amber inclusions The Burmese amber specimens studied here with astonishingly fine details greatly enhanced were derived from amber mines near Noije Bum Pap. Avulsos Zool., 2020; v.60: e20206063 ISSN On-Line: 1807-0205 http://doi.org/10.11606/1807-0205/2020.60.63 ISSN Printed: 0031-1049 http://www.revistas.usp.br/paz ISNI: 0000-0004-0384-1825 http://www.scielo.br/paz Edited by: Simone Policena Rosa http://zoobank.org/11A06909-C340-472C-830A-3E00246FCE93 Received: 08/09/2020 Accepted: 20/10/2020 Published: 17/11/2020 Pap. Avulsos Zool., 2020; v.60: e20206063 Li, Y-D. et al.: A tiny ommatid from Burmese amber 2/7 Village (26°20 N, 96°36 E), Hukawng Valley, Kachin State, “Omma”, the type genus of Ommatidae. The name is neu- ′ ′ northern Myanmar. The holotype is deposited in the ter in gender. Nanjing Institute of Geology and Palaeontology (NIGP), Chinese Academy of Sciences, Nanjing, China. The para- types are deposited in Oregon State Arthropod Collection Miniomma chenkuni Li, Yamamoto & Cai sp. nov. (OSAC), Oregon State University, Corvallis, OR, United (Figs. 1‑4) States of America. The amber pieces were trimmed with a small table saw, ground with emery papers of different Material: Holotype, NIGP173375, sex unknown. Paratypes, grain sizes, and finally polished with polishing powder. OSAC_0002900274, OSAC_0002900275, OSAC_0002900276 Photographs under incident light were taken with and OSAC_0002900277, sex unknown. The specimens a Zeiss Discovery V20 stereo microscope. Widefield OSAC_0002900274-6 are in one piece of amber. fluorescence images were captured with a Zeiss Axio Imager 2 light microscope combined with a fluores- Locality and horizon: Amber mine located near Noije cence imaging system. Confocal images were obtained Bum Village, Tanai Township, Myitkyina District, Kachin with a Zeiss LSM710 confocal laser scanning microscope. State, Myanmar; unnamed horizon, mid-Cretaceous, Images under incident light and widefield fluorescence Upper Albian to Lower Cenomanian (Mao et al., 2018). were stacked in Helicon Focus 7.0.2 or Zerene Stacker 1.04. Confocal images were manually stacked in Adobe Diagnosis: As for the genus. Photoshop CC. The holotype was also imaged using high-resolution X-ray microtomography (Zeiss Xradia Description: Body minute, 1.2-2.0 mm long, 0.4-0.7 mm 520 Versa) at the micro-CT laboratory of NIGP. Based wide, tuberculate, with thin setae. on the comparatively small size of the fossil specimen, a CCD-based 4× objective was used, providing isotropic Head (Figs. 3D, 4A-B) prognathous, moderately voxel sizes of 2.1437 μm with the help of geometric mag- elongate, constricted posteriorly to form a neck; dorsal nification. During the scanning, the acceleration voltage surface with Y-shaped depression (see Kirejtshuk et al., for the X-ray source was 40 kV. To improve signal-to-noise 2016), but without distinct protuberances. Temples ratio, 3,001 projections over 360° were collected, and moderately long, not prominent. Compound eyes entire the exposure time for each projection was 4 s. The to- and not protruding, finely facetted, without interfacetal mographic data were analyzed using VGStudio MAX 3.0. setae. Antennal insertions lateral. Antennal grooves ab- Images were further processed in Adobe Photoshop CC to sent. Antenna extending beyond anterior prothoracic enhance contrast. margin, but not reaching posterior prothoracic margin, 11-segmented and subfiliform, with thin hairs; anten- nomere 3 slightly longer than 4; antennomere 11 near- RESULTS ly twice as long as 10. Labrum transverse, with apex slightly concave. Mandible with vertically aligned teeth. Systematic Paleontology Maxillary and labial palps relatively short, not reaching mandibular apex when anteriorly directed. Prementum Order Coleoptera Linnaeus, 1758 subtrapezoidal. Gular sutures invisible. Suborder Archostemata Kolbe, 1908 Pronotal disc (Fig. 4A) longer than wide, with a Family Ommatidae Sharp & Muir, 1912 shallow medial groove; anterior corners rounded. Genus Miniomma Li, Yamamoto & Cai gen. nov. Prosternum (Fig. 4B) in front of coxae moderately long, probably fused with propleura. Prosternal process in- Type species: Miniomma chenkuni Li, Yamamoto & Cai complete, apically acute. Procoxal cavities contiguous. sp. nov. Elytra (Figs. 3F, 4C) 1.7 times as long as wide combined, with rows of window punctures, probably without mac- Diagnosis: Body minute. Head moderately elongate, ulae; longitudinal ridges (primary veins) weakly raised. without distinct protuberances; compound eyes not Procoxae (Fig. 4B) subglobular. Metacoxae protruding; antennae subfiliform; antennomere 3 slight- (Figs. 3E, 4D) subglobular, not extending laterally. Femora ly longer than 4; antennomere 11 nearly twice as long stout. Tibiae thinner, without well-developed tibial spurs. as 10; antennal grooves absent. Pronotal disc with a Tarsi five-segmented; tarsomeres simple, not bilobed; shallow medial groove; propleura fused with proster- tarsomeres 1-4 subequal in length; tarsomere 5 more num; procoxae contiguous; metacoxae subglobular, not than twice as long as tarsomere 4; pretarsal claws simple. extending laterally. Elytra non-serrate, with very narrow Abdomen with five coplanar ventrites, separated by epipleura, and with indistinct longitudinal ridges (prima- wide grooves; ventrite 1 longer than others; ventrite 5 ry veins). Abdominal ventrites abutting; ventrite 5 slight- slightly longer than penultimate one, with broadly gen- ly longer than penultimate one, with broadly gently ar- tly arcuate posterior margin. cuate posterior margin. Etymology: The species is named after Dr. Chen-Kun Etymology: The generic name is derived from the Jiang, a young evolutionary botanist, who helped the English “mini‑”, meaning small, and the generic name first author enormously on his previous research project. Li, Y-D. et al.: A tiny ommatid from Burmese amber Pap. Avulsos Zool., 2020; v.60: e20206063 3/7 Figure 1. General habitus of Miniomma chenkuni gen. et sp. nov.,
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