Research Article Journal of Orthoptera Research 2024, 33(1): 21-26 A new long-winged pygmy grasshopper in Eocene Baltic amber raises questions about the evolution of reduced tegmenulda in Tetrigidae (Orthoptera) Josip SkEso! 3", Niko KASALO23", M. JARED THomas*>*, Sam W. HEAbs** 1 IUCN/SSC Grasshopper Specialist Group, Zagreb, Croatia. 2 SIGTET—Special Interest Group Tetrigidae, Karlsruhe, Germany. 3 University of Zagreb, Faculty of Science, Zagreb, Croatia. 4 Center for Paleontology, Illinois Natural History & State Geological Surveys, Prairie Research Institute, University of Illinois at Urbana-Champaign, 1816 South Oak Street, Champaign, Illinois 61820, USA. 5 Illinois State Geological Survey, Prairie Research Institute, University of Illinois at Urbana-Champaign, 615 East Peabody Drive, Champaign, Illinois 61820, USA. Corresponding authors: Niko Kasalo (niko.kasalo5 @gmail.com), Josip Skejo (skejo.josip@gmail.com, jskejo@biol.pmf.hr) Academic editor: Daniel Petit | Received 18 April 2023 | Accepted 30 May 2023 | Published 9 January 2024 https://zoobank.org/1A982C52-DF7C-42FE-A D3B-376A3380829E Citation: Skejo J, Kasalo N, Thomas MJ, Heads SW (2024) A new long-winged pygmy grasshopper in Eocene Baltic amber raises questions about the evolution of reduced tegmenula in Tetrigidae (Orthoptera). Journal of Orthoptera Research 33(1): 21-26. https://doi.org/10.3897/jor.33.105144 Abstract Extant pygmy grasshoppers (Tetrigidae) that possess wings have the forewings reduced into scale-like tegmenula, while hind wings remain fully developed. Rusmithia gorochovi gen. et sp. nov. (Tetrigidae, Batrachideinae, Rusmithini trib. nov.) is described based on a single adult female holotype from Lithuanian Baltic amber, from the Bartonian- Priabonian age, some 40 million years ago, and this is the only known tetrigid in which tegmenula or tegmina (the forewings) extend as far as half the length of the hind femur. Besides this very unique trait, other characters of Rusmithia gen. nov. indicate similarity with extant and especially fossil Batrachideinae (genus Danatettix Thomas, Skejo & Heads, 2019). Because of the strong differences this genus and Danatettix have with American Batrachideinae, they are assigned to a new tribe, European Batrachideinae or Rusmithini trib. nov. Acrydium bachofeni (Zeuner, 1937) might belong to this or a sibling genus based on its very long tegmenula or Succinotettix chopardi Piton, 1938, based on its 19-segmented antennae; neither species is tranferred as their types could not be examined. Keywords Amber, autapomorphy, Baltic, European Batrachideinae, fossil, plesiomor- phy, tegmina, Tetrigidae Introduction Although many have dedicated their lives to studying them, our knowledge of extant pygmy grasshoppers (Orthoptera, * All the authors are of equal contribution. Tetrigidae) is still scarce (Cigliano et al. 2023). Given this, it is unsurprising that the available information on extinct species of the family can easily be summarized in a single table (Table 1). Such a small dataset paints a tragically incomplete picture of their evolution—a picture that is slowly being painted one fossil at a time (Sharov 1968, Thomas et al. 2019). Most of the extant tetrigid species have fore- and hind wings. The forewings of these species are reduced to tiny scales and are called tegmenula in pygmy grasshoppers, as opposed to tegmina in winged crickets and grasshoppers. The hind wings of pygmy grasshoppers are functional and are referred to as alae, as in other orthopteran families (Harz 1969, Devriese 1991, 1999). The long- est tegmenula have been found in tPrototetrix Sharov, 1968 and tArchaeotetrix Sharov, 1968 (Sharov 1968), although it is not clear whether they represent stem Tetrigidae or specialized Locustop- sidae (Heads et al. 2014, Thomas et al. 2019). It is also not clear how long the wings were relative to body size, as the wings are the only well-preserved part of species in these genera (Sharov 1968). Because of +Prototetrix and tArchaeotetrix, it is believed that the evolution of pygmy grasshoppers included a reduction in the size of forewings (Sharov 1968). Herein, we describe a new genus and species, tRusmithia gorochovi gen. et sp. nov. (Batrachideinae: tRusmithini trib. nov.), an extinct tetrigid from Lithuania found in Baltic amber. With tegmenula reaching the mid-portion of the hind femora, this species is the new record holder for the longest tegmenula, which makes it an invaluable snapshot of the evolutionary his- tory of Tetrigidae. Copyright Josip Skejo et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 22 J. SKEJO, N. KASALO, M.J. THOMAS AND S.W. HEADS Table 1. A checklist of fossil Tetrigoidea, sorted by taxonomy and approximate age, with location of the type specimens. An asterix indicates that it is unclear where the type is deposited. Updated from Thomas et al. (2019). CLASSIFICATION LOCALITY AGE Tetrigidae: Batrachideinae: Batrachideini 1 Eotetrix unicornis Gorochov, 2012 Green River Fm, Wyoming, USA Middle Eocene Tetrigidae: Batrachideinae: Rusmithini trib. nov. 2 Danatettix hoffeinsorum Thomas, Heads & Skejo, 2019 Baltic amber Middle Eocene 3 Rusmithia gorochovi gen. et sp. nov., this study Baltic amber Middle Eocene Tetrigidae: Tetriginae North Jutland, Denmark North Jutland, Denmark Early Eocene Early Eocene 4 Eozaentetrix wittecki Zessin, 2017 5 Eozaentetrix furi Zessin, 2017 Tetrigidae: Metrodorinae 6 Antilotettix electrum Heads, 2009 7 Baeotettix lottiae Heads, 2009 8 Electrotettix attenboroughi Heads & Thomas, 2014 Tetrigidae incertae sedis ) Archaeotetrix locustopseiformis Sharov, 1968 10 Prototetrix reductus Sharov, 1968 11* = Acrydium (?) bachofeni (Zeuner, 1937)* 12* — Succinotettix chopardi Piton, 1938* 13* — Tettigidea (?) gracilis (Heer, 1865)* Material and methods The holotype of Rusmithia gorochovi gen. et sp. nov. was pur- chased on eBay by Ru Smith who then contacted the first author. The female holotype is deposited in Ru Smith's private collection, York, UK. The widest possible age range of Baltic amber is 25 to 43 million years (Sadowski et al. 2017), but it is currently unclear whether the younger deposits are simply redeposits of the older material, so the usual age estimate is Lutetian (47.8-41.2 Ma) to Priabonian (37.8- 33.9 Ma) (Seyfullah et al. 2018), with most of the material dated to the Priabonian (Sadowski et al. 2020). We refer to our fossil as belonging to the Middle Eocene following the Lutetian—Priabonian estimates. The holotype was photographed and drawn by Ru Smith. Characters relevant for subfamily classification are number of antennal segments (>19 in Batrachideinae, <18 in other subfamilies), shape of paranota (rectangular in Batrachideinae, triangular in other subfamilies), and morphology of the legs (with sulcate dorsal margin in Batrachideinae, carinated in other subfamilies). As a result, the new genus is assigned to Batrachideinae. Taxonomy follows that of the Orthoptera species file (Cigliano et al. 2023). Morphological terminology follows Tumbrinck (2014), and nomenclature is in accordance with the International Code of the Zoological Nomenclature (ICZN 1999). Results Taxonomic part Family Tetrigidae Rambur, 1838 Subfamily Batrachideinae Bolivar, 1887 Type genus.—Batrachidea Serville, 1838. Diagnosis.—Antennae with more than 19 segments, paranota rectangular, mid-femora with sulcate dorsal margin, female sper- matheca with two diverticula. Dominican amber Early Miocene Dominican amber Early Miocene Dominican amber Early Miocene Turga Fm, Transbaikalia, Russia Early Cretaceous Early Cretaceous Middle Eocene Middle Eocene Late Miocene Turga Fm, Transbaikalia, Russia Baltic amber Baltic amber Oeningen, Switzerland Composition.—Batrachideini (Southern and Northern America), Cassitettigini (Africa and SE Asia), Bufonidini (New Guinea, Australia, New Caledonia, and Solomon islands) (Storozhenko 2019), and Rusmithini trib. nov. (Eocene Europe, extinct). Tribe Rusmithini Skejo, Kasalo, Thomas & Heads, trib. nov. https://zoobank.org/38C79F95-AA70-452E-832D-2D2825D950D6 Diagnosis.—Antenna composed of 19 or more antennomeres. Pronotum smooth and flat. Anterior margin of the pronotum ob- tusely triangular, not spine-like and not projected above the head. Tegmenula and alae present. Legs elongated. Mid-femora sulcate. Genicular and antegenicular teeth of the hind femora minute, genicular notch almost indistinct. Type genus.—Rusmithia Skejo, Kasalo, Thomas & Heads, gen. nov. Composition.—Two genera (Danatettix and Rusmithia), two spe- cies. Danatettix hoffeinsorum Thomas, Skejo & Heads, 2019 and the herewith described Rusmithia gorochovi Skejo, Kasalo, Thomas & Heads, gen. et sp. nov. Enigmatic Acrydium bachofeni (Zeuner, 1937), characterized by long wings (Zeuner 1937) might also be- long to this tribe as well as the genus Succinotettix Piton, 1938 (with one species, S. chopardi Piton, 1938) whose antennae are reported to have at least 19 segments (Piton 1938). However, we refrain from transferring these genera to Rusmithini as we were unable to examine the type specimens. Genus Rusmithia Skejo, Kasalo, Thomas & Heads, gen. nov. https://zoobank.org/2112E38B-6726-40EB-8AA3-6129D1CA04BD Type species. —Rusmithia gorochovi sp. nov. Diagnosis. —Antenna with 20 segments (or more, some articulations between antennomeres are not clear). Tegmenula extremely long (for a member of Tetrigidae), reaching the mid-length of the hind femur. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) J. SKEJO, N. KASALO, M.J. THOMAS AND S.W. HEADS Composition.—A single species, Rusmithia gorochovi Skejo, Kasalo, Thomas & Heads, sp. nov., described herein. Acrydium bachofeni (Zeuner, 1937) may also belong to this genus if the long tegmen- ula (5 mm in length), cited in the original description, are taken into account (Zeuner 1937). However, the type of A. bachofeni has not been examined by us, and we are unsure of its location. Etymology.—The new genus-group name is patronymic and honors Dr. Ru Smith, who kindly made the important specimen available to science. The name is feminine. Rusmithia gorochovi Skejo, Kasalo, Thomas & Heads, sp. nov. https://zoobank.org/92DD1977-586C-41E2-9024-E585BCE90B24 Figs 1 and 2 Type specimen.—Holotype: Russian Federation ¢ adult female (Figs 1 and 2); Kaliningradskaya oblast’, Yantarny, Anna mine; Ru Smith’s collection. Syninclusions: a fly (Diptera) belonging to the family Sciaridae (det. A.J. Ross). 23 Etymology.—The new species name is patronymic and honours Dr. Andrei V. Gorochov, world-renowned expert on fossil Orthoptera. Diagnosis.—As for the genus. Description.—Head (Fig. 2B, C, D). In frontal view: Eyes globose. Top margin of compound eye a little above vertex. Lateral and transverse carina forming an acute angle below which surface of exoskeleton more granular; some air bubble encapsulation appar- ent in this part. Frontal costa prominent. Frontal costa bifurcates at approximately middle of the compound eye height. Scutellum vaguely bottle-shaped; the section between the eyes approximately as narrow as an antennal groove. Below eyes, scutellum progres- sively widened up to the bottom margin of the antennal groove and then progressively but slightly narrowed; section below eyes double the length of the one between the eyes. Paired ocelli placed a little above the bottom margin of the compound eye; median ocellus occluded by debris. Top margin of the antennal groove at the level of the bottom margin of the compound eye. In dorsal Fig. 1. Rusmithia gorochovi gen. et sp. nov. Female holotype, lateral habitus. Photo and drawing credit: Ru Smith, used with permission. Scale bar: 10 mm. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) J. SKEJO, N. KASALO, M.J. THOMAS AND S.W. HEADS Maia Ode es eS a DH ae Fs Fig. 2. Rusmithia gorochovi gen. et sp. nov., female holotype, details. Photo credit: Ru Smith, used with permission. A. Dorsal view of the pronotum; B. Dorsal view of the head; C. Anterior view of the head; D. Antenna; E. Lateral view of the anterior half of the body; E. Ovipositor; G. Hind femur; H. Forewing. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) J. SKEJO, N. KASALO, M.J. THOMAS AND S.W. HEADS view: Vertex highly granulated and wider than the compound eye. Anterior margin of the vertex slightly dorsal to the anterior margin of the compound eyes; frontal costa protrudes slightly anterior to the level of the compound eyes. Lateral and transverse carinae form acute triangular shapes that enclose shallow triangular fossulae. Medial carina barely visible throughout the length of the vertex. The compound eyes not touching anterior margin of pronotum. In lateral view: Frontal costa prominent. Scutellum very prominent. Most of the view obscured by encapsulated air bubbles and debris. Pronotum (Fig. 2A, B, E). Macropronotal form. In dorsal view: Entire surface tuberculated. Anterior margin of the prono- tum projected forwards in the form of an obtuse triangle. Prozonal carinae parallel. Interhumeral carinae long, converging dorsally. Median carina present throughout the length of the pronotum. Lateral area large dorsal to the tegmina.Pronotal apex occluded by debris. Wings surpass the pronotal apex but it is unclear by how much as the apex is broken. In lateral view: Paranotum rec- tangular. Ventral sinus in the form of an obtuse angle; tegminal sinus in the form of a nearly right angle. Infrascapular area narrow and of unclear length. Median carina forms a hump between the shoulders; anteriorly, there is a lower hump between the prozona and metazona, and the anterior process forms yet another hump, smaller than the preceding one. Dorsal to the interhumeral hump, the median carina is straight. Wings (Fig. 2E, H). Alae well-developed, reaching past the pronotal apex. Tegmina unusually wide and long, a little less than half of the length of alae. Tegmina width more than half of that of the hind femur. Radius straight, very disctinct. Subcosta parallel to the radius. Other veins not discernable. Legs (Fig. 2E, G). Fore- and mid-femora thin, rectangular in cross-section. Foretibiae with small teeth distally. Hind femora long and narrow with small genicular and tiny antegenicular teeth. Ovipositor (Fig. 2F). Short, serrated. Both ovipositor valves bulging in the middle. Dorsal valve wider than ventral valve, but of same length. Measurements.—Pronotum length 17 mm; body length 18 mm; hind femur length 9 mm; hind femur width 2 mm; tegmen length 6 mm. Discussion The long tegmenula of Rusmithia gorochovi are peculiar in that the radius and costa are clearly visible, unlike in any other known tetrigid. A similar pattern of venation can be observed in Tridacty- loidea (Sharov 1968), which modern analyses have reconstructed as basal Caelifera (Song et al. 2020). This seems to imply that this morphology may be ancestral in Tetrigoidea, but such a conclu- sion cannot be drawn at the moment for several reasons. First, it has been shown in different taxa that reduction in wing size is a relatively common occurrence and is followed by a reduction in wing venation, with only the veins crucial for wing armature persisting (Perfilieva 2010, Zikié et al. 2017). This means that it is impossible to exclude convergence based only on the present data. Second, Danatettix hoffeinsorum, a European relative of Rusmithia gorochovi, and Eotetrix unicornis, an American fossil Batrachideinae, both have typical tetrigid tegmenula (Gorochov and Labandeira 2012, Thomas et al. 2019), which casts doubt on the continu- ous persistence of this morphology through time. Acrydium (?) bachofeni is stated as having long tegmenula by Zeuner (1937), meaning that it could belong to Rusmithia or a new closely related genus. Examining the type of this species would help elucidate 25 the taxonomy of European Batrachideinae and the evolution of tegmina in this group. Third, Archaeotetrix and Prototetrix, which are much older than Rusmithia, also exhibit long tegmenula but with different venation (Sharov 1968), although their affinities remain obscure. The most parsimonious explanation is that Ru- smithia belongs to a line of Batrachideinae in which the longer teg- menula were retained, which may represent an intermediary state between the fully winged ancestral form and the more modern, reduced forms. This implies the possibility that tegminal reduc- tion occurred at least twice in Tetrigoidea. More research is needed to clarify the evolution of Tetrigoidea and their wings. Baltic amber is a gold mine for fossil arthropods (Weitschat and Wichard 2010), comparable to the number of plant taxa de- scribed from the same deposits (Sadowski et al. 2020). A recent botanical study by Sadowski et al. (2020) revealed the presence of various habitats in the Baltic region, ranging from coastal swamps to mixed mesophytic conifer-angiosperm forests with a wide range of transitionary habitats between them. Similar habi- tats are found today in the warm-temperate regions of East Asia and North America, and they house numerous species of modern Tetrigidae, including Batrachideinae (Cigliano et al. 2023). Thus, with relative certainty, we can conclude that Rusmithia gorochovi likely spent its time in warm and wet habitats feeding on algae, mosses, and detritrus in unknown proportions (Hochkirch et al. 2000, Kuyavova et al. 2020). The fly trapped together with the hol- otype of R. gorochovi (Fig. 3) belongs to the family Sciaridae (det. Andrew Ross), which feed on detritus and fungi (Gerhardt and Hribar 2019), indicating their shared habitat. The fossil record of Tetrigidae is scant, and much more work needs to be done. Most notably, the placement of Archaeotetrix locustopseiformis and Prototetrix reductus within the taxonomy of Tetrigoidea is unclear (Sharov 1968, Thomas et al. 2019), if they indeed belong to this superfamily. Because the venation of the tegmina of these species differs from both the extant and other known extinct tetrigids, it should be closely examined as it could indicate that these two species are only distantly related to the other known tetrigids. Furthermore, the types of Acrydium (?) ba- chofeni, Succinotettix chopardi, and Heer's (1856) Tettigidea (?) gra- cilis could not be examined, and their affinities remain unclear. If and when they are found, they will undoubtedly shed much- needed light on the evolutionary history of Tetrigidae. Fig. 3. A fly belonging to the family Sciaridae, encapsulated to- gether with Rusmithia gorochovi gen. et sp. nov. 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