Cretaceous Research 91 (2018) 287e298

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Cretaceous Research

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The oldest representatives of (Coleoptera: ) 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 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. (), 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 research by Robertson et al. The family Endomychidae, or handsome fungus have (2015) have resulted in separation of the former Cerylonid Series always been considered as closely related to sharing into an independent superfamily Coccinelloidea and restriction of pseudotrimerous tarsi (Tomaszewska, 2000a, 2010) and often Endomychidae by removing Anamorphidae, 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- 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 , , , Dis- small or medium sized beetles comprising about 90 genera and colomatidae, Merophysiidae and . Pal and Lawrence 1600 species distributed throughout the world with a maximum (1986) added to Cerylonid Series removed from diversity in the tropical and subtropical regions (Shockley et al., Colydiidae () 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 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 distinguished from the Cenozoic fossil tory of Insect Evolution & Environmental Changes, Capital Normal taxa as follows: (1) from Phymaphoroides by much smaller body, University in Beijing, Three Gorges Entomological Museum, antennal club simple with antennomere 9 not enlarged and infla- Chongqing, China, and Museum and Institute of Zoology, Warsaw, ted, and by only weakly lobed tarsi; (2) from Glesirhanis by much Poland. smaller body, longer antennae reaching beyond base of elytra, The following measurements were made and are used in de- antennomeres 3 and 4 subequal in size, and shorter and bearing scriptions: total length e from apical margin of clypeus to apex of setigerous punctures elytra; (3) from Palaeoestes by shorter body, elytra; pronotal length e from the middle of anterior margin to distinctly marked lateral and basal pronotal sulci, tarsomere 1 not margin of basal foramen; pronotal width e at widest part; elytral shorter than 2 or 3, and the antenna with pedicel and scape length e across sutural line including scutellum; elytral width e differently shaped and sized; (4) from Cretolestes gen. nov. by across both elytra at the widest part. antennomere 5 swollen, much longer and wider than anten- The morphological terminology and the generic attribution and nomeres 4 or 6, pronotum basally narrower than base of elytra, and comparison with extant and extinct taxa follows: Strohecker the prosternal process not separating procoxae. (1953), Tomaszewska (2000a, 2000b, 2010). Burmalestes differs from the extant genera of Leiestinae by much All taxonomic acts established in the present work have been smaller body and by the following characters: (1) from Leiestes registered in ZooBank (see below), together with the electronic Chevrolat by antennomere 3 and 4 subequal in size, pronotum with publication LSID: urn:lsid:zoobank.org:pub:49EA8C4D-2D96- deep basal sulcus and abdominal ventrite 1 as long as remaining 420C-A8FD-50BCF9AC0D22. four ventrites combined; (2) from Panaleies Tomaszewska by pu- bescent dorsum, antennomere 9 slightly but distinctly smaller than 3. Systematic palaeontology antennomere 10, pronotum with lateral margins smooth and abdomen with five visible ventrites; (3) from Rhanidea Strohecker Family ENDOMYCHIDAE Leach, 1815 by antennomere 3 and 4 subequal in size, pronotal disc without pair W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298 289 of longitudinal, subparallel grooves and abdomen with five visible Mesoventrite bicarinate with intercoxal process somewhat ventrites; (4) from Panamomus Gorham by very narrow prosternal boat-shaped; mesoventral pits not apparent; mesocoxal cavities process, well developed hind wings, elytra without rows of punc- circular, narrowly separated with mesoventral process about 0.5 tures, and abdomen with five ventrites; (5) from Phymaphora times as wide as coxal diameter; lateral closure of mesocoxal cav- Newman by male antennal club not modified, prosternal process ities not visible. Scutellum moderately large, transverse, length very narrow and the abdominal ventrite 1 as long as remaining four about 0.5 times width. Elytra 1.25 times as long as wide, convex; ventrites combined; (6) from Stethoranis Blaisdell by prosternal irregularly punctate, punctures denser than those on head and process very narrow, hind wings well developed, and much longer pronotum, about 1.5e2.0 diameters apart; sutural stria fine, entire; abdominal ventrite 1. epipleuron narrow, incomplete apically. Metaventrite transverse, nearly twice as broad as long; more than 1.5 times longer than Etymology. The name of the new genus is derived from a combi- mesoventrite; intercoxal process straight between mesocoxae; nation of Burma, referring to the traditional name of Myanmar with postcoxal pits not apparent; punctation moderately dense and -lestes, derived from Leiestes. The gender is masculine. comparatively coarse. Metacoxae broadly separated. Hind wings Composition. The new genus is monotypic. well developed. Legs comparatively long. Femur subclavate, pubescent. Tibiae Burmalestes albertalleni Tomaszewska and Slipinski sp. nov. widening towards apices without apparent modifications, covered urn:lsid:zoobank.org:act:0B238EA2-974B-406C-B577- with short pubescence; tibial spurs absent. Tarsal formula 4-4-4. AABF935E657B Tarsi long, tarsomeres 1e3 gradually slightly shorter and weakly (Figs. 1AeD) lobed; tarsomere 4 about as long as tarsomeres 1e3 combined. Material. Holotype: No. MIZ 7/2018/1 (Museum and Institute of Abdomen with five ventrites; ventrite 1 as long as remaining Zoology, PAS). Presumably male. The holotype is embedded in a four ventrites combined. Ventrites 2, 3 and 4 subequal in length; small, flattened, subquadrate piece (9.0 8.7 2.6 mm) of clear- terminal ventrite weakly rounded apically. Postcoxal lines absent. yellow coloured Burmese amber (burmite). Complete specimen. Abdominal punctation (especially on ventrites 2e5) sparser and Syninclusions are present in form of two small organic pieces. distinctly smaller than those on metathorax. Locality and horizon. Myanmar, Kachin State, Hukawng Valley, Burmese amber; Upper Cretaceous, lowermost Cenomanian. Remarks. The holotype is recognized as a male, based on modifi- Etymology. The species name is dedicated to Mr. Albert Allen, who cation on the antennae and hardly rounded/subtruncate apically brought an attention of the first author to this inclusion. terminal ventrite of abdomen. Diagnosis. As stated for the new genus. Cretolestes Tomaszewska, Slipinski and Ren gen. nov. Description. Body length 1.3 mm; width (at widest point in about urn:lsid:zoobank.org:act:93508C80-C55C-4DBE-90B8- middle part of elytra) 0.58 mm. Body elongate-oval (about 2.3 times 9E1BA9985F89 as long as wide) and convex; dorsal surfaces shiny and shortly pubescent; brownish-black with appendages (antennae, mouth- Type species: Cretolestes niger Tomaszewska, Slipinski and Ren sp. parts and legs) brown. nov. Diagnosis. Cretolestes can be distinguished from all other extant and Head transverse, retracted into prothorax to the hind margin of fossil members of Leiestinae by the following combination of eyes, finely punctate with punctures 2.0e3.0 diameters apart, characters: 1) body comparatively small (about 1.2 mm long); 2) provided with pale pubescence. Clypeus transverse and flat. Eyes basal and lateral pronotal sulci clearly and deeply impressed; 3) large, rounded, prominent and coarsely faceted. Antennal in- antennomeres 3e8 bead-like and loosely joined; 4) antennal club sertions not visible. Antenna 11-segmented, relatively long, 3-segmented, simple with antennomere 9 not grossly enlarged (e.g. extending at least to bases of elytra; scape stout, slightly larger than Phymaphora); 5) prosternal process broadly separating procoxae pedicel; antennomeres 3 and 4 subequal, bead-like, about as long (about 0.6 times as wide as coxal diameter); 5) abdomen with five as wide, much shorter than pedicel; antennomere 5 swollen, freely articulated ventrites; and 6) metaventrite much longer than 2.5e3.0 times longer and about 1.6 times wider than antennomeres mesoventrite (but hind wings not apparent in the studied 4 or 6; antennomeres 6e8 similar to 3 and 4, although anten- specimen). nomere 7 slightly longer than neighbouring ones; antennomeres 9e11 form elongate, narrow, loose and scarcely flattened club; Specifically Cretolestes can be distinguished from the other fossil antennomere 9 slightly but distinctly shorter and narrower than Leiestinae as follows: (1) from Glesirhanis by distinctly less elongate antennomere 10. Maxillary palpomeres 1 and 3 very short; palpo- habitus, antennomere 3 and 4 subequal in size, antennomere 9 mere 2 at least twice as long as 1 or 3; terminal palpomere nearly as smaller than 10 and elytra with setigerous punctures; (2) from long as palpomeres 2 and 3 combined, widest near basal third then Phymaphoroides by much less elongate body, antennal club simple tapering, weakly rounded apically, about 1.4 times as long as wide with antennomere 9 not enlarged and by tarsomeres 1 and 2 in widest part. Labial terminal palpomere subcylindrical, about as apparently not lobed; (3) from Palaeoestes by less elongate body, long as wide, distinctly longer than palpomere 2, subtruncate at antenna with pedicel differently shaped and sized than scape, apex. antennomere 9 distinctly shorter and narrower than 10, pronotal Prothorax 0.7 times as long as wide; widest near middle, slightly lateral and basal sulci distinctly marked and tarsi with tarsomere 1 constricted at basal fourth; disc rather finely punctate with in- not being shorter than 2 or 3; (4) from Burmalestes gen. nov. by terspaces about 3.0 diameters apart. Lateral margins finely antennomeres 4e6 subequal in size, the pronotum as wide basally as bordered, evenly rounded from apex towards base; anterior angles base of elytra and prosternal process distinctly separating procoxae. blunt, posterior angles nearly right-angled; disc evenly convex; Cretolestes can be distinguished from the extant genera of the lateral sulci in form of triangular, concave impressions, basal sulcus subfamily Leiestinae by the following other characters: (1) from deep. Posterior margin very weakly bisinuate, anterior margin Leiestes Chevrolat by antennomeres 3 and 4 subequal in length, straight. Prosternal process very narrow with procoxae contiguous, pronotum with deep basal sulcus and abdominal ventrite 1 as long circular in outline. Prosternum in front of coxae about 1.2 times as as following 3.5 ventrites combined; (2) from Panaleies long as coxal diameter. 290 W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298

Fig. 1. Burmalestes albertalleni gen. et sp. nov., holotype, specimen numer: MIZ 7/2018/1. A, photograph of dorsal habitus; B, photograph of ventral habitus; C, drawing of head, antenna and pronotum, dorsal view; D, drawing of habitus, ventral view. Scale bars ¼ 0.5 mm (A, C), 0.1 mm (B, D). W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298 291

Tomaszewska by dorsally pubescent body, antennomere 9 Meso- and metathorax. Mesoventrite carinate with intercoxal distinctly smaller than antennomere 10, pronotum with lateral process somewhat boat-shaped; mesoventral pits not apparent; margins smooth and abdomen with five ventrites; (3) from Rha- mesocoxal cavities circular, separated by about one coxal diameter; nidea Strohecker by antennomeres 3 and 4 subequal, pronotal disc open laterally. Scutellum comparatively small, strongly transverse, without pair of longitudinal, subparallel grooves and abdomen with 1.6 times wider than long, rounded apically. Elytra elongate, 1.3 five ventrites; (4) from Panamomus Gorham by prosternal process times as long as wide, convex; irregularly, sparsely punctate, distinctly separating procoxae, metaventrite much longer than punctures 2.5e4.0 diameters apart; sutural stria fine but well mesoventrite, elytra without rows of punctures, and abdomen with visible, entire; epipleuron narrow, incomplete apically. Metaven- five ventrites; (5) from Phymaphora Newman by antennal club in trite transverse, 1.5 times as wide as long; 2.4 times longer than male not modified, prosternal and mesoventral processes relatively mesoventrite and 1.4 times longer than abdominal ventrite 1; broad, and abdominal ventrite 1 as long as following three ventrites intercoxal process straight between mesocoxae; one oval pit latero- combined; (6) from Stethoranis Blaisdell by prosternal and meso- posteriorly to each mesocoxa visible; punctation moderately dense ventral processes relatively broad, and abdominal ventrite 1 as long and comparatively coarse. Metacoxae broadly separated, about 1.7 as three following ventrites combined. diameter apart. Hind wings not apparent. Legs comparatively long and slender. Femora subclavate, pu- Etymology. The name of the new genus is a combination of the bescent. Tibiae widening towards their apices without apparent Creto-, referring to the Cretaceous age of the deposit with -lestes, modifications, covered with short, dense pubescence; tibial spurs derived from Leiestes. The gender is masculine. absent. Tarsal formula 4-4-4. Tarsi long, simple; tarsomeres 1e3 Composition. The new genus is monotypic. gradually shorter; tarsomere 4 as long as tarsomeres 1e3 combined. Abdomen with five ventrites; ventrite 1 as long as following 3.5 Cretolestes niger Tomaszewska, Slipinski and Ren sp. nov. ventrites combined. Apex of terminal ventrite arcuate. Postcoxal urn:lsid:zoobank.org:act:1B965320-E1E8-4361-BBB0- lines absent. Abdominal punctation (especially on ventrites 2e5) 78A223F8A49E sparser and finer than those on metaventrite. (Figs. 2AeD) Material. Holotype: No. CNU008071 (Capital Normal University). Remarks. Based on a simple, not modified antennae and rounded Sex unknown. The beetle specimen is embedded in a small, flat- terminal ventrite of abdomen we presume that the holotype can be tened, long oval piece (14.1 8.8 2.5 mm) of clear-yellow col- a female. The mouthparts are hardly visible in this specimen. oured Burmese amber (burmite). The right hind leg is missing. Subfamily MEROPHYSIINAE Seidlitz, 1872 Syninclusions are represented by numerous air bubbles, organic pieces and well preserved sp. The characters of Cretaparamecus do not correspond perfectly Locality and horizon. Myanmar, Kachin State, Hukawng Valley, with any subfamily of Endomychidae. However, the overall body Burmese amber; Upper Cretaceous, lowermost Cenomanian. size and shape, and subsequently listed characters are shared with Etymology. The species name is derived from the Latin adjective the Holoparamecus-line (Tomaszewska, 2000a; Robertson et al., niger referring to its black body colour. 2015), and we assign it to the subfamily Merophysiinae pending Diagnosis. As stated for the new genus. further research. The following characters of Cretaparamecus are Description. Body length ¼ 1.2 mm; width (at widest point in about shared with members of Merophysiinae: antenna 11-segmented middle part of elytra) ¼ 0.53 mm. Habitus elongate-oval (about with two segmented club; frontoclypeal suture straight; sub- 2.20 times as long as wide) and convex; dorsal surface shiny and antennal grooves absent; antennal sockets not visible from above; shortly pubescent; body uniformly blackish. pronotum narrowest at base with lateral margins smooth; base of the pronotum without sulci and with characteristic transverse tu- Head transverse, partially retracted into prothorax with eyes bercles (although weakly marked); mesocoxal cavities narrowly well exposed, finely punctate, and punctures provided with pale closed laterally; hind coxae oval; trochanters elongate. pubescence. Frontoclypeal suture present. Clypeus transverse and The main difference between Cretaparamecus and extant Mer- flat. Eyes large, rounded, prominent and coarsely faceted. Antennal ophysiine are 4-segmented tarsi with tarsomeres 1 and 2 weakly insertions not visible. Antenna 11-segmented, moderately long, lobed as compared to tarsi 3-segmented with simple tarsomeres in extending almost to base of elytra; scape stout, slightly larger than extant genera of the subfamily (however Colovocerida Belon ac- pedicel; antennomeres 3e8 bead-like, 3e6 about as long as wide, cording to original description (Belon, 1884) has 4-segmented tarsi 7e8 scarcely transverse, all much shorter than pedicel; anten- with shortened tarsomere 3). nomeres 9e11 form elongate, narrow, loose and scarcely flattened club; antennomere 9 distinctly shorter and narrower than anten- Cretaparamecus Tomaszewska, Slipinski, Bai and Zhang gen. nov. nomere 10; terminal antennomere obliquely truncate at apex. urn:lsid:zoobank.org:act:FED84588-8F37-496C-8807- Maxillary terminal palpomere widest near basal third then 82FED0CCA2EF tapering, weakly rounded apically. Prothorax 0.7 times as long as wide; widest at base; nearly as Type species: Cretaparamecus tarsalis Tomaszewska, Slipinski, Bai wide basally as base of elytra; lateral edges of pronotum narrowly and Zhang sp. nov. but distinctly bordered, evenly rounded from apex towards basal Diagnosis. Cretaparamecus can be easily distinguished from all fourth where pronotum is slightly constricted; anterior angles known extant Merophysiinae genera by the following combination blunt, posterior angles nearly right-angled; disc evenly convex; of characters: antennomere 5 in male greatly enlarged, bulbous; lateral sulci in form of triangular, concave impressions, basal sulcus tarsi 4-segmented with tarsomeres 1 and 2 weakly lobed; fore tibia well visible. Posterior margin of pronotum weakly bisinuate, in male with strongly marked large tooth; eyes comparatively anterior margin straight. Pronotum rather finely punctate. Pros- large; and hind wings well developed. ternal process relatively broad, about 0.5 times as wide as coxal diameter, reaching posterior margin of coxae; procoxa circular in Additionally, apart from all above mentioned characters, Cre- outline. Prosternum in front of coxae 1.3 times as long as coxal taparamecus can be specifically distinguished from: (1) diameter. 292 W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298

Fig. 2. Cretolestes niger gen. et sp. nov., holotype, specimen numer: CNU008071. A, photograph of dorsal habitus; B, photograph of ventral habitus; C, drawing of head, antennae and pronotum, dorsal view; D, drawing of habitus, ventral view. Scale bars ¼ 0.1 mm. W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298 293

Holoparamecus Curtis by the base of pronotum with only weakly (pubescence not apparent). Colour dark brown with appendages marked tubercles or impressions; (2) Pseudevolocera Champion by slightly paler. 11-segmented antenna with 2-segmened club, the prosternum smooth without fossae and metaventrite without postcoxal lines; Head sparsely and moderately densely punctate. Eyes compar- (3) Colovocerida Belon by abdominal ventrite 1 without postcoxal atively large, oval, coarsely faceted. Antennal sockets concealed by lines; (4) Pseudoparamecus Brethes by 11-segmented antenna sides of frons, not visible from above; antennal grooves absent. with 2-segmened club and at least mid and hind coxae well Antenna comparatively long, reaching beyond base of prono- separated; (5) Lycoperdinella Champion by 11-segmented antenna tum, composed of 11 antennomeres with club formed by two ter- with 2-segmened club and the pronotum with lateral margins minal antennomeres; scape stout, slightly larger than pedicel; hardly bordered and smooth; (6) Rueckeria Arriaga-Varela et al. by antennomeres 3e4 and 6e9 bead-like, transverse, subequal in size 11-segmented antenna with 2-segmened club, pronotum hardly to each other, much shorter than antennomere 2; antennomere 5 bordered and smooth laterally, abdominal ventrite 1 without large, swollen, more than 4.0 times longer and nearly 3.0 times postcoxal lines and hind wings well developed; (7) Merophysia wider than antennomeres 4 or 6; antennomeres 10e11 form Lucas by 11-segmented antenna with 2-segmened club, head and elongate, narrow, loose and scarcely flattened club with anten- prosternum smooth without foveae, frontoclypeal suture straight nomere 10 about as long as wide, and antennomere 11 longer than and hind wings well developed; (8) Reitteria Leder by much wide and conical at apex. smaller body (below 1 mm), 11-segmented antenna with 2- Frontoclypeal suture straight. Clypeus transverse, flat. Maxilla segmened club, frontoclypeal suture straight, lateral margins of with palpomere 3 short; palpomere 2 at least twice as long as pronotum distinctly constricted basally and hind wings well palpomere 3; terminal palpomere about 1.35 times as long as pal- developed; (9) Cholovocera Motschulsky by 11-segmented an- pomere 2, subcylindrical, somewhat bulbous at basal third then tenna with 2-segmened club, frontoclypeal suture straight and weakly tapering apically, apex weakly rounded. Labium with pal- hind wings well developed and lateral margins of the pronotum pomere 1 very small; palpomere 2 transverse, somewhat oval, distinctly constricted basally; (10) Hexasternum Rücker by 11- weakly inflated; terminal palpomere largest, weakly oval, sub- segmented antenna with 2-segmened club, different shape of truncate at apex. Mentum trapezoidal, nearly twice as wide as long; the pronotum lacking lateral carinae, and abdomen with five finely punctate. ventrites; (11) Lathrapion Rücker, by 11-segmented antenna with Prothorax weakly transverse, about 1.3 times as wide as long, 2-segmened club and elytral sutural stria complete; (12) Lixella widest at anterior fourth, constricted at basal fourth; punctation Peyerimhoff by 11-segmented antenna with 2-segmened club, and pubescence of pronotal surface hardly visible due to numerous frontoclypeal suture straight, lateral margins of the pronotum small air bubbles; lateral edges hardly bordered, smooth; anterior distinctly constricted basally, prosternum smooth without foveae margin straight, with anterior angles rounded; posterior angles and mesoventral process narrower than coxal diameter; (13) right-angled to weakly acute. Pronotal disc weakly convex. Pronotal Pythlarhinus Dajoz by 11-segmented antenna with 2-segmened base with a pair of weakly transverse, oval impressions/tubercles, club, frontoclypeal suture present, mid and hind coxae well with area around tubercles depressed. Prosternal process narrow separated, and elytra with well visible complete sutural striae but separates front coxae. Procoxa circular in outline. Prosternum in (according to description (Dajoz, 1970) Pythlarhinus has antenna front of coxa about 2.3 times longer than coxal longitudinal composed of 12-antennomeres with 3 antennomeres of club, diameter. frontoclypeal suture invisible, pro- and mid coxae contiguous, and Meso- and metathorax. Scutellum small, transverse, widely sutural striae are absent). rounded apically. Mesoventrite with intercoxal process elongate, narrow, subparallel and widest in its mid length, bordered/carinate Etymology. This new genus-group name is composed of “Creta-“ laterally, distinctly separates mesocoxae (about 0.5 times as wide as (part of the name Cretaceous indicating the origin of this taxon) and coxal diameter) reaching half of their length; mesoventral pits not a part of the name Holoparamecus (referring to similarity between apparent (postcoxal area of mesoventrite poorly visible). Mesocoxa both genera). The gender is masculine. weakly oval in outline, its cavity narrowly closed outwardly by Composition. The new genus is monotypic. sterna. Meso-metaventral junction of straight-line type. Elytron elongate, about 1.6 times as long as wide, convex, irregularly and Cretaparamecus tarsalis Tomaszewska, Slipinski, Bai and Zhang moderately coarsely punctate with punctures 2e3 diameters apart; sp. nov. epipleuron narrow, incomplete at apex. urn:lsid:zoobank.org:act:12624136-8423-4AC4-9D98- Sutural stria sharply defined, complete. Metaventrite transverse, D6FB9C5EECC7 about 1.6 times as wide as long, and 1.9 times as long as meso- (Figs. 3AeF) ventrite, weakly convex; anterior margin rather thick; discrimen Material. Holotype: No. BU-001534, currently housed in the Insti- extending to at least half length of metaventrite; metaventral tute of Zoology, Chinese Academy of Sciences but will eventually be postcoxal pits absent. Metacoxae transversely oval, separated by deposited in the Three Gorges Entomological Museum, Chongqing. about 1.3 of coxal longitudinal diameter. Hind wings present and The holotype is embedded in a small, flattened, rectangular piece well developed, much longer than elytra, with inner margins sur- (8.3 4.2 1.7 mm) of clear-yellow coloured Burmese amber rounded with long hairs. (burmite). Complete specimen. Syninclusions are represented by Legs long and comparatively slender. Femur subclavate, pubes- one specimen of Collembola. cent, widest near apical third, about twice as wide as tibia; tibiae Locality and horizon. Myanmar, Kachin State, Hukawng Valley, pubescent, gradually widening towards tarsus, fore tibia in male Burmese amber; Upper Cretaceous, lowermost Cenomanian. with large tooth in about apical third; tibial spurs absent. Tarsal Etymology. The name of this new species refers to 4-segmented formula 4-4-4. Tarsi long, with tarsomeres 1e3 gradually shorter; tarsi, an unusual character for the subfamily Merophysiinae. tarsomeres 1 and 2 weakly lobed. Tarsomere 4 slightly shorter than Diagnosis. As stated for the new genus. 1e3 combined. Claws simple. Description. Length 0.95 mm; greatest width (at widest point in Abdomen with five freely articulated ventrites; ventrite 1 about middle part of elytra) ¼ 0.38 mm. Body elongate, about 2.5 slightly shorter than three following ventrites combined and about times longer than wide, weakly convex; shiny, smooth, glabrous 0.75 times as long as metaventrite, without postcoxal lines; 294 W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298

Fig. 3. Cretaparamecus tarsalis gen. et sp. nov., holotype, specimen numer: BU-001534. A, photograph of dorsal habitus; B, drawing of antenna; C, photograph of ventral habitus; D, photograph of hind half of body with part of hind wings and aedeagus visible, dorsal view; E, drawing of head with antennae and mouthparts, and prothorax with parts of fore and mid legs, ventral view; F, drawing of pro-, meso- and metathorax with fore and mid legs, ventral view. Scale bars ¼ 0.5 mm (A, B, D), 0.1 mm (C, E, F). W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298 295 ventrites 2e4 almost subequal in length. Terminal ventrite with Head transverse, retracted into prothorax to the hind margin of subtruncate apex. Abdominal punctation (especially on ventrites eyes, densely and finely punctate. Eyes large, oval, prominent, 2e5) distinctly sparser and finer than those on metathorax. coarsely faceted. Antennal sockets visible from above. Antenna Aedeagus seems to be pushed out during compression and resin longer than head and thorax together, reaching beyond elytral congealment. It is visible as weakly curved and possessing internal shoulder, composed of 11 antennomeres with loose, narrow, sclerites. moderately flattened club composed of 3 antennomeres; anten- nomeres 1e3 elongate, antennomere 4 and 6 subquadrate, anten- Subfamily XENOMYCETINAE Strohecker, 1962 nomere 5 enlarged, swollen, somewhat trapezoidal in shape and nearly as long as wide (measured in widest/anterior margin), The following genus is assigned to the subfamily Xenomycetinae 2.5e3.0 times longer and about 3.0 times wider than antennomeres (Tomaszewska, 2000a, 2005) based on its general appearance 4 or 6; antennomeres 7e10 transverse; terminal antennomere including body colour and the following set of diagnostic charac- nearly as long as wide, obliquely truncate apically. Frontoclypeal ters, such as: body long-oval and moderately convex, antenna suture hardly visible on the specimen. Clypeus flat. Maxilla with longer than head and prothorax together with antennal club terminal palpomere at least 2 times as long palpomere 3, sub- composed of three rather loosely articulated antennomeres, cylindrical and subtruncate at apex. maxilla with terminal palpomere long, subcylindrical, pronotum Prothorax about as long as wide, widest at base and at anterior distinctly bordered laterally, sutural striae on elytra well visible, third, somewhat constricted in between. Pronotum distinctly pro- and mesocoxae narrowly separated while hind coxae widely bordered laterally; basal sulcus well marked, lateral sulci moder- separated, metaventrite with discrimen almost complete and with ately large, triangularly elongate and rather deep; anterior angles postcoxal pits, abdominal ventrite 1 nearly as long or longer than weakly produced, blunt; posterior angles almost right-angled. 2e5 combined, and tarsi 4-4-4 simple or weakly pseudotrimerous. Posterior margin of pronotum bisinuate, anterior margin straight. Pronotal disc moderately convex, rather finely punctate with in- Palaeomycetes Tomaszewska, Slipinski and Ren gen. nov. terspaces 1.0e3.0 diameters apart. Prosternal process invisible on urn:lsid:zoobank.org:act:CB5B1B78-3D89-4C82-9AF9- the specimen but seems to be very narrow as procoxae look 090BBA54C85A contiguous, circular in outline. Prosternum in front of coxae about 1.4 times as long as procoxal longitudinal diameter. Type species: Palaeomycetes foveolatus Tomaszewska, Slipinski and Meso- and metathorax. Scutellum small, transverse (about Ren sp. nov. 1.35 times wider than long), somewhat heart-shaped. Mesoven- Diagnosis. The new genus may be separated from Xenomycetes Horn, trite hardly visible on the specimen but with intercoxal process 1880 (the only known so far extant genus of the subfamily Xen- elongate, carinate and narrow (about 0.5 times as wide as omycetinae) by the following combination of characters: 1) body mesocoxal diameter); narrowly separates mesocoxae, not extends much smaller (about 2.0 mm long); 2) antennomere 5 enlarged, beyond them. Mesocoxa circular in outline, its cavity outwardly swollen; 3) basal and lateral sulci on pronotum clearly and compar- open. atively deeply impressed; 4) anterior margin of pronotum not Elytron elongate, about 1.9 times as long as wide, convex, bordered; 5) paired postcoxal pits on metaventrite verylarge, notably irregularly punctate; shoulders prominent; sutural striae well larger than mesocoxal diameter; 6) abdomen with ventrite 1 longer visible, entire; epipleuron narrow, incomplete apically. Metaven- than ventrites 2e5 combined; and 7) hind wings well developed. trite weakly transverse and about 2.7 times longer than mesoven- Etymology. The new genus-group name Palaeomycetes is composed trite; weakly convex, comparatively coarsely punctate (punctures of “palaeo” (meaning old, ancient) and “-mycetes”, a part of the about 2.0e3.0 diameters apart); with two pairs of oval, very large name Xenomycetes, the only extant, habitually similar xen- postcoxal pits (larger than mesocoxal diameter); discrimen almost omycetine genus from North America. The gender is masculine. complete; intercoxal process straight between mesocoxae. Meta- Composition. The new genus is monotypic, represented by the type coxae transverse, separated by about coxal longitudinal diameter. species only. Hind wing visible (somewhat folded) from one side of specimen, presumably well developed. Palaeomycetes foveolatus Tomaszewska, Slipinski and Ren sp. nov. Legs comparatively long. Femur subclavate, pubescent, widest urn:lsid:zoobank.org:act:38677EF6-C658-472E-AA72- near apical third, about twice as wide as tibia; tibia gradually, D244D1CE732D weakly widening towards tarsus, without apparent modifications, (Figs. 4AeF) covered with short pubescence; tibial spurs absent. Tarsal formula Material. Holotype: No. CNU008072 (Capital Normal University). 4-4-4. Tarsi long, with weakly lobed tarsomeres 1e3; tarsomeres Sex unknown. The beetle specimen is embedded in a small, flat- 1e3 subequal in length. Tarsomere 4 longer than 1e3 combined. tened, semirectangular piece (20.1 6.2 2.4 mm) of clear-yellow Claws simple. coloured Burmese amber (burmite). Complete specimen. Synin- Abdomen with five ventrites; ventrite 1 longer than remaining clusions are represented by piece of wood and remnants of insect four ventrites together and about as long as metaventrite; post- leg. coxal pits not apparent. Ventrites 2, 3 and 4 subequal in length. Locality and horizon. Myanmar, Kachin State, Hukawng Valley, Apex of terminal ventrite arcuate. Postcoxal lines on ventrite 1 Burmese amber; Upper Cretaceous, lowermost Cenomanian. absent. Abdominal punctation (especially on ventrites 2e5) sparser Etymology. The name of this new species refers to the large post- and distinctly smaller than those on metathorax. coxal pits on the metaventrite. Remarks. The antenna illustrated on Fig. 4D does not match well Diagnosis. As stated for the new genus. the digital image (Fig. 4A) in which the antennomeres look slimmer Description. Body length ¼ 2.00 mm; width (at widest point in about and more elongate. This discrepancy is a result of different angles middle part of elytra) ¼ 0.65 mm. Habitus long-oval (about 3.07 used to execute the drawing and images. The exact position of an times as long as wide) and moderately convex; dorsal surface shiny amber piece used for executing the drawing (Fig. 4D) was not and shortly pubescent. Body uniformly dark brown; confusedly suitable for a good quality photograph. punctate with punctures provided with pale pubescence. 296 W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298

Fig. 4. Palaeomycetes foveolatus gen. et sp. nov., holotype, specimen numer: CNU008072. A, photograph of dorsal habitus; B, drawing of head and pronotum, dorsal view; C, drawing of metatarsus; D, drawing of antenna; E, photograph of ventral habitus; F, drawing of ventral habitus. Scale bars ¼ 0.5 mm. W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298 297

4. Taxonomic changes author is grateful to Al Allen (Boise, Idaho, USA) for donating an amber piece with leiestine inclusion (described here as Burmalestes Zemyna Tomaszewska, nomen novum albertalleni) to the collection of Museum and Institute of Zoology, urn:lsid:zoobank.org:act:9893B393-D29B-40A0-A2FF- PAS in Warsaw. This project is partly supported by grants from the 2B29DA5DA581 National Natural Science Foundation of China (grant nos. 31730087 and 31672323), Program for Changjiang Scholars and Innovative Laima Alekseev and Tomaszewska, 2018: 12 (nec Laima Gravitis, Research Team in University (IRT-17R75) and Project of High-level 1981). Type species, by original designation: Laima andreei Teachers in Beijing Municipal Universities (IDHT20180518). James Alekseev and Tomaszewska, 2018. Robertson and one anonymous reviewer are greatly appreciated for Etymology. This new genus-group name is a theonym and derived their careful and thorough review of our manuscript. from Zemyna, the name of an Earth goddess in the Baltic pagan religion. The gender is feminine. References

5. Discussion Alekseev, V.I., Tomaszewska, W., 2018. New handsome fungus beetles (Coleoptera: Coccinelloidea: Anamorphidae, Endomychidae) from European amber of the e Representatives of Endomychidae in fossil record have appeared Upper Eocene. Palaeontologia Electronica 21.1.6A, 1 23. https://doi.org/10. 26879/832. in several publications over time (Klebs, 1910; Strohecker, 1953; Belon, M.J., 1884. Description d'un Coleopt ere nouveau du Chili. Annales de la Spahr, 1981; Carpenter, 1992; Shockley et al., 2009) but very few Societ e Entomologique de Belgique 28, 163e165. Cenozoic taxa have been formally described so far (Shockley and Bocak, L., Barton, C., Crampton-Platt, A., Chesters, D., Ahrens, D., Vogler, A., 2014. Building the Coleoptera tree-of-life for >8000 species: composition of public Alekseev, 2014; Alekseev and Tomaszewska, 2018). Of the prop- DNA data and fit with Linnaean classification. Systematic Entomology 39, erly described taxa, Palaeoestes Kirejtshuk and Nel, 2009 (French 97e110. Oise amber) and Phymaphoroides Motschulsky, 1856 (Baltic amber) Burakowski, B., Slipinski, S.A., 2000. The larvae of Leiestinae with notes on the phylogeny of Endomychidae (Coleoptera: Cucujoidea). Annales Zoologici 50 (4), represent extinct genera of Leiestinae and Laima Alekseev and 559e573. Tomaszewska, 2018 (¼Zemyna Tomaszewska nom. nov.), the only Carpenter, F.M., 1992. Treatise on Invertebrate Paleontology. Part R, Arthropoda 4, known extinct genus of the subfamily Endomychinae. Two species Vol. 4. Superclass Hexapoda. Geological Society of America, Boulder, Colorado, pp. 1e655. of Trochoideus described by Alekseev and Tomaszewska (2018) are Crowson, R.A., 1955. The Natural Classification of the Families of Coleoptera. the only Eocene species classified in the recent genus of the sub- Nathaniel Lloyd, London, p. 187. family Pleganophorinae. Cruickshank, R.D., Ko, K., 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. Journal of Asian Earth Sciences 21, 441e455. Interestingly, the species known (described and undescribed) Dajoz, R., 1970. Etude des Coleopt eres Lathridiidae de l'Afrique Intertropicale. from the Cretaceous Burmese amber represent much greater Annales Musee Royal de l'Afrique Centrale. Serie ine8. Sciences Zoologiques taxonomic diversity than the taxa from the Eocene Baltic and Bit- 184, 1e49. terfield ambers described so far, with the subfamilies Mer- Gravitis, V.A., 1981. Bezzamkovye brakhiopody [Brachiopods]. In: Sorokin, V.S. (Ed.), Devon i karbon Pribaltiki [Devonian and Carboniferous of the Baltic Region]. ophysiinae and Xenomycetinae not recorded from the Eocene Zinatne, Riga, pp. 448e463 [in Russian]. deposits. Grimaldi, D.A., Engel, M.S., Nascimbene, P.C., 2002. Fossiliferous Cretaceous amber There appears to be continuous diversity of the Leiestinae in the from Myanmar (Burma): its rediscovery, biotic diversity, and paleontological significance. American Museum Novitates 3361, 1e72. Burmese, French and Baltic ambers, which is currently small and Hunt, T., Bergsten, J., Levkanicova, Z., et al., 2007. A comprehensive phylogeny of exclusively Holarctic group (Tomaszewska, 2000a, 2000b). Biology beetles reveals the evolutionary origins of a superradiation. Science 21, e of Leiestinae is not well known but their larvae are subcylindrical 1913 1916. Kirejtshuk, A.G., 2013. Taxonomic names, in Current knowledge of Coleoptera and feed internally in the basidiomycete fungi or in fungi infested (Insecta) from the Lower Cretaceous Lebanese amber and taxonomical notes for sapwood (Burakowski and Slipinski, 2000). The occurrence of some Mesozoic groups. Terrestrial Reviews 6, 103e134. Merophysiinae in the Burmese amber may be connected to the Kirejtshuk, A.G., Azar, D., 2008. New taxa of beetles (Insecta, Coleoptera) from Lebanese amber with evolutionary and systematic comments. Alavesia 2, increased availability of termite and ant colonies inhabited by many 15e46. species in this subfamily. However, the typical symphile, Trochoi- Kirejtshuk, A.G., Nel, A., 2009. New genera and species of (Coleoptera, deus is so far known only from the Eocene deposits. ) from lowermost Eocene French amber. Denisia 26, 103e118. fl Klebs, R., 1910. Über Bernsteineinschlüsse im allgemeinen und die Coleopteren An observed bulbous and in ated antennomere 5 in members of meiner Bernsteinssammlung. Schriften der Physikalischeokonomischen€ all three endomychid subfamilies discovered in the Burmese amber Gesellschaft zu Konigsberg€ 51, 217e242. is functionally not fully understood fenomenon. Lawrence, J.F., Newton, A.F., 1995. Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). In: Pakaluk, J., Slipinski, S.A. (Eds.), Biology, Phylogeny and Classification of Cole- 6. Conclusions optera. Papers celebrating the 80th Birthday of Roy A. Crowson, Vol. 2. Muzeum i Instytut Zoologii PAN, Warszawa, pp. 779e1006. Our discovery of four new genera of Endomychidae from the Leach, W.E., 1815. Entomology. In: Brewster, D. (Ed.), Edinburgh Encyclopedia, Vol. 9. Edinburgh, pp. 57e172. Upper Cretaceous Burmese amber represents the first Mesozoic Lord, N., Hartley, C.S., Lawrence, J.F., McHugh, J.V., Whiting, M.F., Miller, K.B., 2010. fossil record for the family. The newly described genera represent Phylogenetic analysis of the minute brown scavenger beetles (Coleoptera: three extant subfamilies, Leiestinae, Merophysiinae and Xen- Latridiidae), and recognition of a new beetle family, Akalyptoischiidae fam. N. (Coleoptera: Cucujoidea). Systematic Entomology 35, 753e763. omycetinae exposing rich palaeodiversity of handsome fungus McKenna, D.D., Wild, A.L., Kanda, K., Bellamy, C.L., Beutel, R.G., Caterino, M.S., et al., beetles in the Cretaceous period. This discovery is of potential 2015. The beetle tree of life reveals that Coleoptera survived end of Permian significance for understanding the phylogenetic relationships be- mass extinction to diversify during the cretaceous terrestrial revolution. Sys- tematic Entomology 40 (4), 835e880. tween Endomychidae and other related groups of Coccinelloidea Motschulsky, V.I., 1856. Voyages. Lettres de M. de Motschulsky a M. Menetries. No. very poorly represented among the Mesozoic fossils. 3. New York le 15 Juillet 1654. Etudes Entomologiques 5, 21e38. Pal, T.K., Lawrence, J.F., 1986. A new genus and subfamily of mycophagus Bothri- deridae (Coleoptera: Cucujoidea) from the Indo-Australian Region, with notes Acknowledgements on related families. Journal of the Australian Entomological Society 25, 185e210. We are grateful to Hong Pang (Sun Yat-sen University), Zhenhua Reike, H.-P., 2012. Neue Arten und Anmerkungen zu Latridiidae (Coleoptera) aus Baltischem Bernstein. Latridiidae 9, 7e23. Liu and Cate Lemann (CSIRO) for their help with cutting and pol- Robertson, J.A., Slipinski, S.A., Moulton, M., Shockley, F.W., Giorgi, A., Lord, N.P., ishing amber pieces and photographs of the specimens. The first McKenna, D.D., Tomaszewska, W., Forrester, J., Miller, K.B., Whiting, M.F., 298 W. Tomaszewska et al. / Cretaceous Research 91 (2018) 287e298

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