Cretaceous Research 91 (2018) 287e298 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes The oldest representatives of Endomychidae (Coleoptera: Coccinelloidea) from the Upper Cretaceous Burmese amber * ** Wioletta Tomaszewska a, , Adam Slipinski b, Ming Bai c, Weiwei Zhang d, Dong Ren e, a Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warszawa, Poland b Australian National Insect Collection, CSIRO, Canberra, Australia c Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China d Three Gorges Entomological Museum, Chongqing 400015, China e College of Life Sciences, Capital Normal University, Xisanhuanbeilu 105, Haidian District, Beijing, 100048, China article info abstract Article history: Four new genera along with four new species belonging to three subfamilies of the family Endomychidae Received 24 April 2018 (Coleoptera: Coccinelloidea) are described, diagnosed and imaged from Upper Cretaceous Burmese Received in revised form amber: Burmalestes albertalleni Tomaszewska and Slipinski gen. et sp. nov., Cretolestes niger Tom- 22 June 2018 aszewska, Slipinski and Ren gen. et sp. nov. (Leiestinae), Cretaparamecus tarsalis Tomaszewska, Slipinski, Accepted in revised form 1 July 2018 Bai and Zhang gen. et sp. nov. (Merophysiinae), and Palaeomycetes foveolatus Tomaszewska, Slipinski and Available online 3 July 2018 Ren gen. et sp. nov. (Xenomycetinae). They represent the oldest discovered definitive fossil members of the Endomychidae and the first records of the superfamily Coccinelloidea from the Burmese amber. Keywords: Burmite Zemyna Tomaszewska nom. nov. is proposed as a replacement name for the recently described Laima Cenomanian Alekseev and Tomaszewska, 2018 (Endomychidae: Endomychinae) from the Eocene Baltic amber; Laima New taxa is the name already used in a fossil Brachiopoda (Gravitis, 1981). Replacement name © 2018 Elsevier Ltd. All rights reserved. 1. Introduction molecular studies (Hunt et al., 2007; Bocak et al., 2014; Zhang et al., 2018) and comprehensive Cucujoidea research by Robertson et al. The family Endomychidae, or handsome fungus beetles have (2015) have resulted in separation of the former Cerylonid Series always been considered as closely related to Coccinellidae sharing into an independent superfamily Coccinelloidea and restriction of pseudotrimerous tarsi (Tomaszewska, 2000a, 2010) and often Endomychidae by removing Anamorphidae, Mycetaeidae and aposematic colouration. Both families have been once combined in Eupsilobiidae as independent families. a higher-level taxon called Trimera, often placed at the end of the Current classification of Endomychidae is based on morpho- beetle system (Lawrence and Newton, 1995). However, since the logical cladistic analyses by Tomaszewska (2000a, 2005, 2010) and introduction of the phylogenetic classification of beetles (Crowson, a molecular approach by Robertson et al. (2015) recognising nine 1955) both Coccinellidae and Endomychidae were placed in the subfamilies: Cyclotominae, Danascelinae, Endomychinae, Epi- superfamily Cucujoidea (¼Clavicornia) and included in the derived pocinae, Leiestinae, Lycoperdininae, Merophysiinae, Pleganophor- group called Cerylonid Series. Original Cerylonid Series of Crowson inae and Xenomycetinae. (1955) was mostly defined by reduced tarsal formula (4-4-4 or 3-3- The family, as currently defined is a moderately large group of 3) and also included Alexiidae, Corylophidae, Cerylonidae, Dis- small or medium sized beetles comprising about 90 genera and colomatidae, Merophysiidae and Latridiidae. Pal and Lawrence 1600 species distributed throughout the world with a maximum (1986) added Bothrideridae to Cerylonid Series removed from diversity in the tropical and subtropical regions (Shockley et al., Colydiidae (Tenebrionoidea) and lately a new family Akalyptoi- 2009; Robertson et al., 2015). schiidae was added to the series by Lord et al. (2010). Ordinal level Endomychidae are mostly mycophagous. They are associated with a wide variety of fungi and their most frequent habitats are fungal growth, rotten wood and fungus-infested bark, where adults * Corresponding author. and larvae are often found feeding on fungal spores or hyphae. ** Corresponding author. Many endomychids live in leaf litter and feed on moulds and fungi E-mail addresses: [email protected] (W. Tomaszewska), Adam.Slipinski@ csiro.au (A. Slipinski), [email protected] (M. Bai), [email protected] developing on decaying plant products. Among the mould feeders, (W. Zhang), [email protected] (D. Ren). species of Holoparamecus and Mycetaea are pests of stored https://doi.org/10.1016/j.cretres.2018.07.001 0195-6671/© 2018 Elsevier Ltd. All rights reserved. 288 W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298 products, occurring in granaries and warehouses and causing mi- All beetle specimens studied during this project refer to the nor damages. Saula japonica Gorham is the only known predacious family Endomychidae having the following set of morphological endomychid, with both adults and larvae preying on scale insects characters: head with frontoclypeal suture; antennal grooves ab- on citrus trees (Sasaji, 1978). Many species of Pleganophorinae and sent; antenna 11-segmented with club composed of 2 or 3 anten- Merophysiinae occur in some association with termites and ants, nomeres; pronotum with transverse basal and longitudinal paired but little is known about their food habits. lateral sulci; visible portion of procoxae globular; procoxae usually Fossil taxa of handsome fungus beetles are poorly known and distinctly separated by prosternal process; mesocoxal cavities rarely studied (Shockley and Alekseev, 2014). Only four genera of externally open (closed in Merophysiinae); metaventral paired Endomychidae have been described from the Cenozoic ambers: postcoxal openings usually present; elytral epipleura well devel- Phymaphoroides Motschulsky, Palaeoestes Kirejtshuk and Nel, Gle- oped; tarsi 4-segmented; abdominal ventrite 1 at least as long as sirhanis Shockley and Alekseev and Laima Alekseev and Tom- ventrites 2 and 3 combined; abdominal postcoxal lines absent aszewska (¼Zemyna Tomaszewska nom. nov.). Alekseev and (Tomaszewska, 2000a; Robertson et al., 2015). Tomaszewska (2018) have also described two new species of the extant genus Trochoideus Westwood (Pleganophorinae) from the Subfamily LEIESTINAE Thomson, 1863 Eocene Baltic and Bitterfeld ambers. In the current paper the oldest fossil taxa of handsome fungus The following taxa are assigned to the subfamily Leiestinae, beetles from the Upper Cretaceous Burmese amber are described sharing a set of characters diagnostic for this subfamily: antennal and classified in the subfamilies Leiestinae, Merophysiinae and sockets not visible from above; antennal club loose, composed of Xenomycetinae. This discovery represents the second oldest record 3 antennomeres; triangular lateral sulci on the pronotum deep for the entire superfamily Coccinelloidea, superseded only by the and large; mesoventral process distinctly boat-shaped; meta- controversial latridiid beetle Tetrameropsis mesozoica Kirejtshuk ventrite with postcoxal pits, and tarsi 4-4-4 with simple tarso- and Azar (2008) from the Lebanese amber (see also Reike, 2012; meres. Moreover, the general appearance, the outline of antennae Kirejtshuk, 2013; Shockley and Alekseev, 2014). The present dis- and maxillary palpi of the species described here correspond well covery confirms results of McKenna et al. (2015) study indicating at with recent members of Leiestinae (Tomaszewska, 2000a, least Jurassic origin of Coccinelloidea. 2000b). Burmalestes Tomaszewska and Slipinski gen. nov. 2. Material and methods urn:lsid:zoobank.org:act:0F4442CC-6E6A-4EF7-A3D8- 3B0175317841 The study is based on four specimens of Endomychidae embedded in the Burmese amber originated from the Hukawng Type species: Burmalestes albertalleni Tomaszewska and Slipinski Valley of northern Myanmar (see details in Cruickshank and Ko, sp. nov. 2003: fig. 1). The age of the amber deposits generally considered Diagnosis. Burmalestes can be distinguished from all other extant to be the earliest Cenomanian (Grimaldi et al., 2002) or possibly and fossil members of the subfamily Leiestinae by the following latest Albian (Ross et al., 2010). The recently conducted U-Pb zircon combination of characters: 1) body comparatively small (about dating restricted its age at 98.79 ± 0.62, which is equivalent to the 1.3 mm long); 2) basal and lateral pronotal sulci clearly and deeply earliest Cenomanian, Late Cretaceous (see details in Shi et al., 2012). impressed; 3) antennomere 5 enlarged, much longer and distinctly The material was prepared using a razor table, polished with wider than antennomeres 4 or 6; 4) antennal club 3-segmented, emery papers with different grain sizes and finally lustrated with with antennomere 9 not grossly enlarged (e.g. Phymaphora); 5) polishing powder. Images were taken using a BK Lab Plus system prosternal process narrow and short, with procoxae subcon- http://www.duninc.com/bk-plus-lab-system.html and Leica tiguous; 5) abdomen with five freely articulated ventrites; and 6) M205C stereomicroscope with a drawing attachment; source im- hind wings well developed. ages were then aligned and stacked in Zerene Stacker and edited in Photoshop CS6. The type specimens are deposited in: Key Labora- Moreover, Burmalestes is