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Description of the Male of Fossil Calomicrus Eocenicus Bukejs Et Bezdekˇ (Coleoptera: Chrysomelidae: Galerucinae) from Eocene Baltic Amber Using X-Ray Microtomography

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Description of the Male of Fossil Calomicrus Eocenicus Bukejs Et Bezdekˇ (Coleoptera: Chrysomelidae: Galerucinae) from Eocene Baltic Amber Using X-Ray Microtomography Foss. Rec., 23, 105–115, 2020 https://doi.org/10.5194/fr-23-105-2020 © Author(s) 2020. This work is distributed under the Creative Commons Attribution 4.0 License. Description of the male of fossil Calomicrus eocenicus Bukejs et Bezdekˇ (Coleoptera: Chrysomelidae: Galerucinae) from Eocene Baltic amber using X-ray microtomography Andris Bukejs1, Jan Bezdekˇ 2, Vitalii I. Alekseev3,4, Kristaps Kairišs1, and Ryan C. McKellar5,6,7 1Institute of Life Sciences and Technologies, Daugavpils University, Vien¯ıbas 13, Daugavpils, 5401, Latvia 2Department of Zoology, Mendel University, Zemedˇ elskᡠ1, 613 00 Brno, Czech Republic 3Shirshov Institute of Oceanology, Russian Academy of Sciences, Nahimovskiy prospekt 36, Moscow 117997, Russia 4Kaliningrad Regional Amber Museum, Marshal Vasilevskii square 1, Kaliningrad 236016, Russia 5Royal Saskatchewan Museum, 2445 Albert St., Regina, SK, S4P 4W7, Canada 6Biology Department, University of Regina, Regina, SK, S4S 0A2, Canada 7Department of Ecology & Evolutionary Biology, University of Kansas, Lawrence, Kansas 66045, USA Correspondence: Andris Bukejs ([email protected]) Received: 18 March 2020 – Revised: 23 April 2020 – Accepted: 1 May 2020 – Published: 11 June 2020 Abstract. A male representative of the extinct species Calomicrus eocenicus Bukejs et Bezdek,ˇ 2014 from mid- Calomicrus eocenicus Bukejs et Bezdek,ˇ 2014 (Chrysomel- dle Eocene Baltic amber (Bartonian, 41.2–37.8 Ma) (Bukejs idae: Galerucinae) is found and described for the first time and Bezdek,ˇ 2014); Calomicroides danicus Nadein, 2015 from Eocene Baltic amber using X-ray microtomography. from upper Eocene Danish (Scandinavian) amber (37.2– The aedeagus is well preserved within the body cavity of the 33.9 Ma) (Nadein et al., 2015); and Taimyraltica calcarata specimen, and it is illustrated in detail. This fossil species Nadein, 2017 from Upper Cretaceous Taimyr amber (85.8– exhibits distinct sexual dimorphism: the male has a smaller 84.9 Ma) (Nadein and Perkovsky, 2018). Additionally, mem- total body size, as well as a copula-adapted modification in bers of Galerucella Crotch, 1837; Luperus Geoffroy, 1762; abdominal ventrite 5 (apical margin deeply trilobed, with and Monolepta Chevrolat, 1836 have been reported from round medial fovea present); meanwhile the female is larger Baltic amber without detailed descriptions (Hope, 1836; in body size and has a simple abdominal ventrite 5 (with- Klebs, 1910; Bachofen-Echt, 1949; Hieke and Pietrzeniuk, out fovea, non-incised and widely rounded apically). Simi- 1984; Spahr, 1981). lar sexually dimorphic characters are typical for extant mem- Calomicrus eocenicus was originally described based on bers of the tribe Luperini, and this report is the first time that a single female specimen. Unfortunately, the holotype was they are described in an Eocene species. The known sexually lost during postal shipment. In the current paper, the male dimorphic characters present in Coleoptera within Eocene of C. eocenicus from Baltic amber is described. The male Baltic amber are briefly discussed. genitalia are examined and illustrated using X-ray microto- mography. The external sexually dimorphic characters are described, and the original species concept (based on the fe- male holotype only) is revised. Finally, the available data on 1 Introduction sexual dimorphism in fossil Coleoptera from Eocene Baltic amber are briefly discussed. Records of Galerucinae sensu stricto (excluding Alticini) from fossil resins are scarce. Until now only four species have been described: Leptonesiotes virkkii Santiago-Blay, Poinar et Craig, 1996 from middle Miocene Domini- can amber (20.43–13.65 Ma) (Santiago-Blay et al., 1996); Published by Copernicus Publications on behalf of the Museum für Naturkunde Berlin. 106 A. Bukejs et al.: Description of the male of fossil Calomicrus eocenicus Bukejs et Bezdekˇ 2 Geological setting were generated from 3001 rotational steps through a 360◦ rotation, using a 4× objective, and the exposure time during Baltic amber originally occurs within the “blaue Erde” each projection was set to 5 s. Acquired images were binned (“Blue Earth”) horizon, which can be found throughout the (2 × 2 × 2), giving a voxel size of 4.87 µm. Images were Baltic Sea coastal area in Europe (Kaliningrad region, Rus- imported into the Dragonfly PRO (version 4.1) software sia; Poland; Denmark; Sweden; Germany; and Lithuania) platform for interactive segmentation and 3D visualization. (Engel, 2001). Although most estimates of the age of Baltic amber have suggested that it is derived from the early part of the middle Eocene (Lutetian, 47.8–41.2 Ma) based largely 4 Systematic palaeontology on K–Ar dating (Ritzkowski, 1997), palynological biostratig- Family Chrysomelidae Latreille, 1802 raphy of the specific region where the sample originated suggests a younger, late Eocene (Priabonian, 37.8–33.9 Ma) Subfamily Galerucinae Latreille, 1802 age (Aleksandrova and Zaporozhets, 2008). An intermedi- ate, middle Eocene (mostly Bartonian, 41.2–37.8 Ma) age Tribe Luperini Gistel, 1848 was recently proposed for the extinct central European resin- Subtribe Luperina Gistel, 1848 producing forests according to the stratigraphy of the Sam- bian amber deposits (Bukejs et al., 2019). Following classi- Genus Calomicrus Dillwyn, 1829 cal views (e.g. Andrée, 1951; Poinar, 1992; Turkin, 1997), Baltic amber is thought to have been produced by Pinus suc- Calomicrus eocenicus Bukejs et Bezdek,ˇ 2014 cinifera (Göppert) Conwentz. Pinaceae along with different Figs. 1–4 coniferous (Sadowski et al., 2017) and deciduous (primarily fagacean) trees, dominated the mixed forest cover of northern and central Europe in the warm-temperate, humid conditions Material examined of the Eocene. More recent research on the chemical compo- sition of Baltic amber has also hypothesized that trees within Collection number RSKM_P3300.139, male. A complete the families Araucariaceae or Sciadopityaceae might be can- beetle is included in an elongate, transparent, yellow am- didates for the production of this amber deposit (Wolfe et ber piece with dimensions of 41 mm × 19 mm × 8 mm, and al., 2009; Lambert et al., 2014); however, the final chemical preserved without supplementary fixation. Syninclusions: composition of amber can be influenced by long-term diage- one specimen of Hymenoptera, a few stellate Fagaceae tri- netic processes, leaving the source tree for the deposit open chomes, numerous small detritus particles and gas vesicles. to debate. Strata Baltic amber from Eocene amber-bearing Blue Earth layers 3 Material and methods (a primarily Bartonian age is interpreted for the extinct cen- tral European resin-producing forests, according to Bukejs et The material examined is deposited in the Palaeontology al., 2019). Collection of the Royal Saskatchewan Museum (Regina, Saskatchewan, Canada) (RSKM specimen number prefix). Locality Observations of the studied beetle specimen were made using a Nikon SMZ745T stereomicroscope. Photographs Yantarny settlement (formerly Palmnicken), Sambian (Sam- were taken using a Visionary Digital imaging system, con- land) Peninsula, Kaliningrad region, Russia. sisting of a Canon EOS 5D camera with a Canon MP-E 65 mm macrophotography lens, attached to an automated Note camera lift with studio flash lighting. Extended depth of field at high magnifications was achieved by combining multiple Specimen RSKM_P3300.139 is morphologically similar to images from a range of focal planes using Helicon Focus the female holotype of Calomicrus eocenicus and is identi- 6.8.0 software, and the resulting images were edited to create fied as male, showing sexually dimorphic characters of ab- figures using Adobe Photoshop CS5. dominal ventrite 5 and a distinctly smaller body size (see The X-ray micro-CT observations of specimen Discussion). RSKM_P3300.139 were conducted at Daugavpils Uni- Description of male versity, Daugavpils, Latvia (DU), using a Zeiss Xradia 510 Versa system. Scans were performed with a polychro- Measurements: total body length 3.5 mm; pronotum length matic X-ray beam at an energy of 40 kV and power of 0.75 mm, pronotum maximum width 1.0 mm; elytra length 3 W. Sample-to-detector distance was set to 17.5 mm, and 2.7 mm, elytra maximum width 1.9 mm. source-to-sample distance was 47.7 mm. Tomographic slices Foss. Rec., 23, 105–115, 2020 https://doi.org/10.5194/fr-23-105-2020 A. Bukejs et al.: Description of the male of fossil Calomicrus eocenicus Bukejs et Bezdekˇ 107 Figure 1. Calomicrus eocenicus Bukejs et Bezdek,ˇ RSKM_P3300.139, photomicrographs, habitus: (a) dorsal view; (b) ventral view. Scale bars D 1.0 mm. https://doi.org/10.5194/fr-23-105-2020 Foss. Rec., 23, 105–115, 2020 108 A. Bukejs et al.: Description of the male of fossil Calomicrus eocenicus Bukejs et Bezdekˇ Figure 2. Calomicrus eocenicus Bukejs et Bezdek,ˇ RSKM_P3300.139, X-ray micro-CT renderings, habitus: (a) dorsal view; (b) ventral view; (c) frontal view; (d) left lateral view. Scale bars D 1.0 mm. Foss. Rec., 23, 105–115, 2020 https://doi.org/10.5194/fr-23-105-2020 A. Bukejs et al.: Description of the male of fossil Calomicrus eocenicus Bukejs et Bezdekˇ 109 Figure 3. Calomicrus eocenicus Bukejs et Bezdek,ˇ RSKM_P3300.139, X-ray micro-CT renderings: (a) elytra, dorsal view, with clipping planes exposing details beneath pronotum and right elytron; (b) pterothorax without legs, ventral view; (c) habitus without legs, caudal view. Scale bars D 0.5 mm. Abbreviations: f – fovea; p – pygidium; v1–v5 – abdominal ventrites 1–5 respectively. Body elongate, slender, slightly convex dorsally and ven- eter 1:3× transverse diameter. Distance between eyes equal trally; reddish-brown as preserved, shiny, with weak metal- to about 1:4× transverse diameter of one eye. Antennal calli lic lustre; glabrous dorsally, with very fine, short, recum- wide and elongate, slightly oblique, apparently contiguous; bent setation ventrally (distinctly visible on pygidium and supracallinal sulcus distinct. Frontal ridge narrow, and sharp. abdomen). Clypeus strongly transverse, with anterior margin concave. Head hypognathous, relatively small, without distinct Labrum transverse, about 1:4× length of clypeus; anterior punctures; head together with eyes slightly wider than an- margin almost straight.
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