Palaeoentomology 003 (2): 180–187 ISSN 2624-2826 (print edition) https://www.mapress.com/j/pe/ Article PALAEOENTOMOLOGY Copyright © 2020 Magnolia Press ISSN 2624-2834 (online edition) PE https://doi.org/10.11646/palaeoentomology.3.2.7 http://zoobank.org/urn:lsid:zoobank.org:pub:1AB433B7-1FE2-46DC-9870-CACF1499B76F A new genus and species representing the first leafhopper (Hemiptera: Cicadellidae) from Eocene Rovno amber CHRISTOPHER H. DIETRICH1, * & EVGENY E. PERKOVSKY2, 3 1Illinois Natural History Survey, Prairie Research Institute, University of Illinois, 1816 S. Oak St., Champaign, IL 61820, USA �[email protected]; https://orcid.org/0000-0003-4005-4305 2Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, B. Khmelnitskogo 15, Kiev, 01601, Ukraine �[email protected]; https://orcid.org/0000-0002-7959-4379 3Borissiak Paleontological Institute of the Russian Academy of Sciences, Profsoyuznaya Str. 123, Moscow, 117997, Russia *Corresponding author. Abstract Sakhalinian amber was also described recently (Dietrich & Perkovsky, 2019). Unlike the more recent (Miocene) Rovnotettix brachypterus gen. et sp. nov. is described Dominican amber, in which most of the leafhoppers and illustrated from Eocene Rovno (Ukraine) amber and tentatively placed in subfamily Bathysmatophorinae, tribe can be accommodated in modern genera (Dietrich & Malmaemichungiini. The new fossil is the first cicadellid Vega, 1995), Eocene-age amber has yielded genera that described from Rovno amber from the Volyn region and mostly represent extinct lineages with combinations of the second brachypterous leafhopper adult described from morphological characters not found in the modern fauna. the fossil record. It does not appear to be closely related to Thus, these faunas document an important transitional Brevaphrodella Dietrich & Gonçalves from Eocene Baltic stage in the evolution of leafhoppers. amber, indicating that multiple flightless cicadellid lineages, now mostly associated with dry habitats with patchy Rovno amber, dated at 35–37 Ma, is a southern vegetation, had evolved in Europe by the Eocene. analogue of the better known and more intensively studied Baltic amber (Perkovsky et al., 2007; Nadein et al., Keywords: Homoptera, Auchenorrhyncha, morphology, 2016). Over the past 20 years, 292 new species have been taxonomy, flightless described from Rovno amber (Perkovsky & Rasnitsyn, 2013; Jałoszyński & Perkovsky, 2016; Perkovsky, 2015, 2018; Lyubarsky & Perkovsky, 2020, in press; Colombo Introduction et al., 2020, in press). Nearly all Rovno amber inclusions studied so far have originated from Klesov and the Horyn Although leafhoppers (Cicadellidae) are currently among River basin (Perkovsky et al., 2010; Perkovsky, 2017), the most diverse and abundant families of herbivorous but new collections have recently been made from the insects, their fossil record remains poorly studied with more western basins of the Styr and Stokhod rivers and major gaps. Improved knowledge of leafhopper fossils especially the Veselukha river floodplain in between. is needed not only to elucidate morphological character These new collections (mostly from Voronki and Velyki evolution through time but also to improve calibrations for Telkovichi) have yielded a number of new species of ongoing molecular divergence time analyses of this group beetles, neuropterans and snakeflies (Jałoszyński & (Dietrich et al., 2017, Skinner et al., 2019). The oldest Perkovsky, 2019; Legalov et al., 2019; Makarkin & true leafhoppers are recorded from the Lower Cretaceous Perkovsky, 2020; Perkovsky & Makarkin, 2019, 2020), of Brazil and Australia (Hamilton, 1990, 1992) and as well as some species previously recorded from Baltic the first leafhoppers from Cretaceous amber have only amber (Perkovsky & Olmi, 2018; Martynova et al., recently been described (Poinar & Brown, 2017; Wang 2019) or from Baltic and Bitterfeld amber (Radchenko et al., 2018; Chen et al., 2019). In contrast, many species & Perkovsky, 2018, 2020, in press). Inclusions from the and genera of Cicadellidae have been described from Volyn region were not previously reported, although the late Eocene Baltic amber (reviewed by Szwedo, 2002; first lacertid from Rovno amber (deposited in a private Dietrich & Gonçalves, 2014; Dietrich & Thomas, 2018). collection) was found at an even more western locality. The first genus and species of Cicadellidae from Eocene Some new taxa have also been described from nearby 180 Submitted: 8 Feb. 2020; accepted by D.-Y. Huang: 9 Mar. 2020; published: 30 Apr. 2020 Belarus (Perkovsky, 2017 and references therein) and the Systematic palaeontology Zhitomir region (Fedotova & Perkovsky, 2015, 2017). The new genus and species of leafhopper described below Order Hemiptera Linnaeus, 1758 was collected in the river basin next to the Stokhod large Suborder Auchenorrhyncha Duméril, 1805 river basin from a site 1.5 km west of Kovel situated on Infraorder Cicadomorpha Evans, 1946 the river Turia (Volyn region). Some other inclusions Superfamily Membracoidea Rafinesque, 1815 known from the same recently-discovered locality include Family Cicadellidae Latreille, 1825 numerous Phoridae, including Aenigmatias Meinert, Subfamily Bathysmatophorinae Anufriev, 1978 1890, the first Rovno amber dragonfly (Vasilenko, pers. Tribe Malmaemichungiini Kwon, 1983 com.), and a non-biting midge belonging to a boreal genus, Apsectrotanypus cf. pecularis (V. Baranov, pers. Rovnotettix gen. nov. com.). (Figs 1, 2) Hemipterans (except Germaraphis aphids: Perkovsky & Wegierek, 2018 and references therein) Type species. Rovnotettix brachypterus sp. nov., by from Rovno amber remain understudied in comparison monotypy. with hymenopterans. Only two heteropteran species Etymology. The genus name, a masculine noun, (Putschkov & Popov, 2003; Herczek et al., 2013), one was formed by combining the name of the fossil amber, aleyrodid genus and species (Drohojowska et al., 2015), Rovno, with -tettix, Greek for “cicada”; a common suffix and one dictyopharid species also recorded from Baltic used for cicadellid genus names. amber (Emeljanov & Shcherbakov, 2011) have been Diagnosis. The new genus differs from other fossil and described so far, compared to 54 named hymenopteran extant Cicadellidae in having the following combination species (nearly half of the known Rovno fauna). Seventy- of traits: head wider than pronotum with anterior margin four hymenopteran genera are recorded. One tribe and angulate in dorsal and lateral views, vertex extended one subfamily of Hymenoptera are recorded in the Eocene behind compound eye; ocelli absent; lorum large, flat and only from Rovno amber and 21 of the 74 hymenopteran extended to lateral margin of face; gena emarginate below genera recorded from Rovno amber are the only known eye exposing proepisternum; rostrum extended past middle Eocene representatives of these genera (Perkovsky, 2018; coxae; forewing short, heavily sclerotized and elytralike Radchenko & Dlussky, 2019; Colombo et al., 2020 in with venation not delimited, apex obliquely truncate; hind press; Radchenko & Perkovsky, 2020, in press). If the femur with apical and preapical pair of macrosetae, tibia tentative placement of the new cicadellid described below with alternating short and long setae in row PV; male in Malmaemichungiini can be verified, it would represent subgenital plates separate from each other and articulated the third tribe of insects recorded in the fossil state known to valve, tapered in ventral view, compressed and curved only from Rovno amber (Legalov et al., 2018). dorsad distally, with scattered macrosetae. The new genus and species described below is only Rovnotettix differs from Brevaphrodella in its more the second hemipteran genus and the first leafhopper elongate, less depressed overall body form and narrow known only from Rovno amber. The fossil is also gena, which does not conceal the proepisternum and important because it is only the second brachypterous somewhat longer forewings (extended to sternite VI rather adult leafhopper described from the fossil record and than V). The specimen photographed by Szwedo (2002) is quite different from the previously described one, appears to be more similar to Rovnotettix in overall body Brevaphrodella nigra Dietrich & Gonçalves, 2014 from form but its wings are also shorter, exposing abdominal Baltic amber, in several respects. segment III and the (male?) genital capsule is much more elongated. Rovnotettix brachypterus sp. nov. Material and methods Material. Holotype male SIZK Kov-1. Eocene Rovno Morphological terminology follows Dietrich (2005). amber. Ukraine: Volyn Region, 1.5 km west of Kovel. Photographs were taken using a Leica Z16 APO The holotype inclusion is very well preserved with microscope equipped with a Leica DFC 450 camera and little apparent distortion and most parts of the exoskeleton processed by LAS Core software. clearly visible. The anterior view of the head and front The holotype is housed in the Schmalhausen Institute legs and the lateral and dorsal views of the abdominal of Zoology, National Academy of Sciences of Ukraine terminalia are obscured by fractures. (SIZK). FIRST LEAFHOPPER FROM ROVNO AMBER Palaeoentomology 003 (2) © 2020 Magnolia Press • 181 FIGURE 1. Rovnotettix brachypterus gen. et sp. nov., male holotype. A, Lateral habitus. B, Dorsal habitus. C, Ventral habitus. D, Enlarged lateral view of head and thorax. E, Male terminalia, ventral view. F, Right front tibia and tarsus, posterolateral view. G, Right hind leg,