A new Eocene Bacanius species (Histeridae: Dendrophilinae) from Baltic amber

A study of Baltic amber revealed a new fossil species of the genus Bacanius LeConte, 1853. Bacanius gorskii sp. nov. differs from the congener described previously from Eocene Rovno amber, B. kirejtshuki Sokolov and Perkovsky, in the shape of meso-metaventral suture, incomplete subhumeral stria, and the impunctate prosternal lobe (urn:lsid:zoobank.org:pub:EE9C6859-607A-41349037-8385601CF42F).


Introduction
The clown beetle family (Histeridae) is nearly cosmopolitan in distribution, and it is very diverse both ecologically and morphologically (Caterino and Vogler, 2002). Histerid beetles comprise over 4000 extant species. Ten subfamilies are presently recognized (Zhou et al., 2020): Antigracilinae, an extinct subfamily from the Lower Cretaceous, and the extant subfamilies Niponiinae, Abraeinae, Saprininae, Dendrophilinae, Onthophilinae, Tribalinae, Histerinae, Haeteriinae, and Chlamydopsinae. Histerid beetles, although generalist predators, have widely varied habitats and are associated with different substrates including dung, carrion, fungi, dead and dying trees, leaf litter or other decomposing vegetation, and symbioses with other animals, most spectacularly with social insects (Caterino and Vogler, 2002).
In this study, description of a new species, Bacanius gorskii sp. nov., an extinct Eocene representative of the tribe Bacaniini, is presented.

Material and methods
The studied Bacanius specimen no. 8619 is deposited in the private collection of Andrzej Górski (Bielsko-Biała, Poland) (CAG) and will subsequently be deposited in the collections of the Museum of Natural History, Institute of Systematics and Evolution of Animals, Polish Academy of Sciences (Kraków, Poland) (ISEA PAS) under collection number no. MP/4077/AG col./no. 8619.
The X-ray micro-CT (µCT) observations of specimen "8619" (CAG) were conducted at Daugavpils University, Daugavpils, Latvia (DU), using a Zeiss Xradia 510 Versa system. Scans were performed with a polychromatic X-ray beam at an energy of 40 kV and power of 3 W. Sample-todetector distance was set to 54.6 mm and source-to-sample distance of 18 mm was used. Tomographic slices were generated from 1601 rotation steps through a 360 • rotation, using a 4× objective, and exposure time during each projection was set to 12 s. Acquired images were binned (2×2×2), giving a voxel size of 1.67 µm. Images were imported into the Dragonfly PRO (ver. 2020.1) software platform for interactive segmentation and 3D visualization. Prior to the full scan, a 10 min warmup scan was conducted with the same scan parameters, except for rotation steps, which had been reduced to 201, and exposure time which was reduced to 3 s. The photographs were taken using a Nikon SMZ 745T stereomicroscope with a Nikon DS-Fi1 digital camera. Extended depth of field at high magnifications was achieved by combining multiple images from a range of focal planes using Helicon Focus v. 6.0.18 software, and the resulting images were edited to create figures using Adobe Photoshop CS5.

Type material
Holotype: no. 8619 [CAG], "HOLOTYPE/Bacanius gorskii sp. nov./des. Alekseev V.I. and Bukejs A. 2021"; adult, female. A complete beetle with partially exposed hind wings is included in a transparent, yellow amber piece with dimensions of 15 × 9 mm and maximum thickness of 1.5 mm and preserved without supplementary fixation. Syninclusions: few stellate Fagaceae trichomes.

Type stratum
A predominantly Bartonian age (41.3-37.9 Ma) is interpreted for the extinct central European resin-producing forests, which produced the amber that has eroded out of the Eocene blue earth layers (Bukejs et al., 2019). However, the vast majority of Baltic amber derives from the geological amber-bearing strata that have been assigned a Priabonian age (37.8-33.9 Ma) (Sadowski et al., 2017(Sadowski et al., , 2020.
Pronotum transverse, 1.85 times as wide as long, widest at base; punctation indistinct, very sparse and fine, with more distinct small punctures at posterior margin; lateral margins widely rounded, anterior and posterior margins convex. Antescutellar stria absent. Prohypomera impressed, sparsely covered with fine punctation. Prosternal lobe wide, convex, without distinct punctation. Suture between prosternal lobe and prosternal keel indistinct. Prosternal keel with concave base. Scutellum not visible.
Mesosventrite almost flat, narrow, about 2.8 times as wide as long, without distinct punctures. Meso-metaventral suture distinct, straight, without punctures. Metaventrite with flat, apparently impunctate disc; with coarse and dense punctation laterally, distance between punctures about 0.5-1.5 times the diameter of one puncture; postmesocoxal stria semicircular; posterior margin slightly convex.

Differential diagnosis
The new species is assigned to the subfamily Dendrophilinae based on (1) the prosternum with short lobe, laterally incised to receive the antennae, (2) non-costate elytra, and (3) antennal cavities on prosternum weakly defined. Placement in the tribe Bacaniini is based on a combination of the following characters: (1) the bent short propygidium (visible only ventrally) and large pygidium, (2) general habitus (small size, with round and convex body), (3) elytral disc without dorsal striae, and (4) scutellum not visible.
The specimen under study differs from representatives of the genus Cyclobacanius Müller, 1925 in that the elytra are without a distinct elliptical area on the elytral disc margined by punctation, and the metaventrite is without longitudinal lines. It differs from representatives of the genus Abraeomorphus Reitter, 1886 in lacking semicircular femoral lines on the metaventrite and from the genus Australanius Gomy, 2009 based on the absence of discal striation. The shape of the prosternal lobe in the new fossil species is similar to representatives of the anophthalmic and Sardinian endemic genus Sardulus Patrizi, 1955(Magrini, 2005Magrini and Fancello, 2005;Magrini et al., 2012;Magrini and Onnis, 2019); however, Bacanius gorskii sp. nov. differs from Sardulus based on the presence of eyes and elytral suhumeral stria. The new species from Baltic amber lacks an arcuate antescutellar stria on the pronotal base, in contrast to representatives of the subgenus Muellerister Cooman, 1936 (of the genus Bacanius); of the genus Neobacanius Müller, 1925; or of several representatives of Abraeomorphus. The subgenus Gomyister Mazur, 1984 (genus Bacanius) is characterized by the presence of a complete subhumeral stria (Mazur, 1984;Mazur and Sawoniewicz, 2008), but the new fossil species has an incomplete subhumeral stria. Based on the previously mentioned combination of characters, we tentatively place Bacanius gorskii sp. nov. in the nominative subgenus.
Bacanius (s.str.) gorskii sp. nov. differs from the Eocene Bacanius kirejtshuki Sokolov and Perkovsky, 2020 described from Rovno amber in that it possesses a straight, impunctate, finely impressed meso-metaventral suture (differing from the convex, coarsely punctate meso-metaventral suture of B. kirejtshuki); an apically reduced subhumeral stria (complete in B. kirejtshuki); and an impunctate prosternal lobe (sparsely punctate in B. kirejtshuki). The new species can be easily distinguished from the other two known histerid beetles in Baltic amber, Carcinops donelaitisi Alekseev, 2016 and Xestipyge ikanti Alekseev, 2016, based upon its convex and rounded body shape, absence of dorsal elytral striae, and distinctly smaller body size.

Derivatio nominis
Patronymic, this new species is named in honour of Andrzej Górski (Bielsko-Biała, Poland), collector of this amber piece.

Remarks
Sex of the examined specimen determined was based on micro-CT results. We were unable to identify any sclerotized structure resembling an aedeagus in the abdominal cavity, and therefore we presume the specimen in question is a female.

Discussion
The life history of extant Bacaniini is little known. Members of the tribe are mainly recorded living in rotting wood, sometimes in leaf litter or in nests of social insects like termites and ants (Kryzhanovskij and Reichardt, 1976), or even belonging to the hypogean and troglophilous faunas (Vomero, 1973;Magrini, 2005). Representatives of Bacaniini, like several other microhisterid taxa, have their mouthparts adapted to consume fungal spores, as well as the larvae of tiny arthropods (Kovarik and Caterino, 2000). A mixed sporophagous-predaceous diet in rotten wood of fallen and standing trees could be assumed as the ecological niche for the fossil species described in the present paper.
The Paleogene entomofauna of Baltic amber has been in the focus of palaeontological research for nearly a century. Dozens of new coleopteran taxa described yearly from this amber deposit show our fragmented knowledge of general coleopteran biodiversity in this lagerstätte, as well as the need for ongoing study of its inclusions. The newly described species is the third formally described from Baltic amber. Described histerid inclusions are also known from three fossil resins of different ages and locations (Chatzimanolis et al., 2006;Poinar and Brown, 2009;Caterino et al., 2015;Caterino and Maddison, 2018;Zhou et al., 2019;Jiang et al., 2020;Sokolov and Perkovsky, 2020) Data availability. All material included in the paper is deposited in the private collection of Andrzej Górski (Bielsko-Biała, Poland) (CAG) and will subsequently be deposited in the collection of the Museum of Natural History, Institute of Systematics and Evolution 98 V. I. Alekseev and A. Bukejs: A new Eocene Bacanius species of Animals, Polish Academy of Sciences (Kraków, Poland) (ISEA PAS), and all data are included in the description. X-ray microtomography volume renderings of the habitus and antenna of Bacanius gorskii sp. nov., holotype, 8619 (CAG) are available as Video Supplements.
Author contributions. VIA and AB designed the study, performed specimen identification and systematic placement, and prepared new species description. Both authors drafted the manuscript and suggestions that improved the manuscript.