Large cicada fossil (Cicadidae, Cicadinae, Tacuini) from the Middle Pleistocene Shiobara Group, Nasushiobara, Tochigi, Japan

Abstract

In this study, we report a cicada fossil from the Middle Pleistocene (Chibanian) of the Shiobara Group in Nasushiobara, Tochigi, Japan. The specimen, preserving both wings and the body, represents a large individual with an estimated total length of 62 mm (head to forewing apex). Its body size, operculum morphology, and wing venation most closely resemble the extant Auritibicen flammatus (Distant, 1892). However, it differs from that species in its elongated wings and absence of forewing infuscation. Therefore, we tentatively identified the fossil as Auritibicen cf. flammatus (Distant, 1892). Morphological differences between a fossil and extant species may indicate that the fossil represents either an extinct species or an ancestral form of the extant species. Cicada fossils are extremely rare, with only 45 species described to date, all from the Cenozoic. Most are fragments of wings, with few specimens preserving the entire body. The specimen reported here is relatively well-preserved among cicada fossils and is the largest known specimen with the entire body intact. A detailed description of this fossil provides important insights into cicada evolution and speciation. Being from the Middle Pleistocene, it is also the youngest occurrence of a fossil cicada.

Introduction

Singing cicadas (Cicadidae, Hemiptera) are a diverse group of hemipteran insects distributed worldwide, comprising approximately 3,400 species across more than 520 known genera (Dmitriev et al., 2022). Cicadas are relatively large insects known for their extensive sound production systems, and most spend their long larval stage underground, with a short adult lifespan (Williams and Simon, 1995). Consequently, cicada fossils are extremely rare. Most fossil discoveries consist of wing fragments, with complete body fossils being particularly uncommon. Notable examples include Dominicicada youngi Poinar and Kritsky, 2011 and Minyscapheus dominieanus Poinar et al., 2012 from Dominican amber; Meimuna protopalifera Fujiyama, 1969 from Japan; Cryptotympana incasa Zhang et al., 1994 and Cryptotympana miocenica Zhang and Zhang, 1990 from China; Eoplatypleura messelensis Jiang et al., 2025 from Messel, Germany; and Cicada aff. lodosi Boulard, 1979 from Willershausen, Germany (Moulds et al., 2023). Most of these specimens are small, with none exceeding 60 mm in total length.

Moulds (2018) reviewed the fossil record of 33 true cicada species (Cicadidae) from the Late Cretaceous to Late Pleistocene. Since then, significant progress has been made in cicada fossil research. The oldest reported cicadid fossil was initially thought to be a first instar nymph preserved in Kachin amber from the Middle Cretaceous in Myanmar, representing the only Mesozoic record (Poinar and Kritsky, 2011). However, this was re-evaluated by Jiang et al. (2024), who assigned it to the family Tettigarctidae. The oldest confirmed cicadid fossil is Davispia bearcreekensis (Cooper, 1941) from the Late Paleocene of Montana, USA (Cooper, 1941).

Cicadid fossils are currently known only from the Cenozoic, with 45 confirmed species (Jiang et al., 2025). These include: one from the Paleocene, four from the Eocene, four from the Oligocene, and 22 from the Miocene, none of which have been identified as extant species; eleven from the Pliocene, including two recently described extinct species from the Late Pliocene of Willershausen, Germany (Moulds et al., 2023; Jiang et al., 2025) and seven from the Pleistocene, all of which are from Japan. These include Meimuna sp., Graptopsaltria sp., Euterpnosia sp. and Tanna? sp. (Fujiyama, 1979, 1982; Inoue, 1986) from the Early Pleistocene and Graptopsaltria aff. nigrofuscata Motschulsky, 1866, Auritibicen bihamatus Motschulsky, 1861 and Yezoterpnosia nigricosta Motschulsky, 1866 (Fujiyama, 1969, 1979) from the Middle Pleistocene. All of these have been identified or compared with extant species, and no extinct species have been discovered from the Pleistocene to date.

Here, we report a third cicadid fossil that was recently discovered from the Middle Pleistocene Shiobara Group in Nasushiobara, Tochigi, Japan. Group. The fossil is relatively well-preserved, with both wings and the body intact, and its estimated total length of 62 mm makes it the largest cicadid fossil found to date.

Material and methods

The fossil was discovered on June 30, 2025 by two elementary school sisters, Maho Shinomiya and Yuno Shinomiya. The rock containing the fossil was a small fragment of siltstone from the Miyajima Formation (Shiobara Group). It was excavated from the fossil quarry of the Konoha Fossils Museum in Nasushiobara, Tochigi, Japan, and was sold as educational material.

The Shiobara Group is a Middle Pleistocene (Chibanian: Suganuma et al., 2021) lacustrine deposit (ca. 0.3 million years ago) that is widely distributed across Nasushiobara, Tochigi, Japan. Numerous plant, vertebrate, insect, and spider fossils have been identified from these strata. For geological and paleontological background, refer to Onoe (1989), Tuzino et al. (2009), and Aiba (2015). Over 100 insect fossils have been reported from the Shiobara Group (Aiba and Hayashi, 2024), including two cicadid taxa: Auritibicen bihamatus Motschulsky, 1861 (Fujiyama, 1979) and Yezoterpnosia nigricosta (Fujiyama, 1969).

The specimen (KYFSI253) is reposited at the Konoha Fossils Museum in Nasushiobara, Tochigi, Japan. It was examined under a Leica M205 C stereo microscope (Leica Corporation, Wetzlar, Germany). Photographs and measurements were obtained using a Leica MC170HD microscope camera with Leica Application Suite Version 4.1.3. Images were sharpened and adjusted for contrast and tonality using Adobe Photoshop version CS6 (Adobe Systems Incorporated, San Jose, CA, USA). We follow the morphological terminology of Moulds (2005, 2012), with abbreviations for wing venation as follows: a1–a8, apical cells 1–8; C = costal vein; CuA = cubitus anterior vein; CuA₁ = first branch of the cubitus anterior vein; M = median vein; M₁–M₄ = first to fourth branch of the median vein; m = medial crossvein; m-cu = mediocubital crossvein; n = node; R = radius; R + Sc = radius + subcostal vein; r = radial crossvein; rc = radial cell; r-m = radiomedial crossvein; RP = radius posterior.

Fossil body and total length were estimated with reference to the morphology of Auritibicen intermedius Mori, 1931 (male: Wang et al., 2018, fig. 12), the type species of the genus Auritibicen.

Systematic paleontology

Order Hymenoptera Linnaeus, 1758

Family Cicadidae Latreille, 1802

Subfamily Cicadinae Latreille, 1802

Tribe Tacuini Distant, 1904

Genus Auritibicen Lee, 2015

Auritibicen cf. flammatus (Distant, 1892)

Figures 1, 2

Figure 1. Photographs Auritibicen cf. flammatus (Distant, 1892), KYFSI253a and KYFSI253b. A, habitus, ventral side (KYFSI253a); B, head, pronotum and mesonotum, dorsal side (KYFSI253b), red arrow indicates continuous W-shaped marking; C, enlarged view of crossvein m; D, enlarged part of crossvein m-cu; E, enlarged view of posterior forewing; F, enlarged abdominal sternite, red arrow indicates spiracles. Abbreviations: a2–a5, apical cells 2–5; b3, base of apical cell 3; b4, base of apical cell 4; m, medial crossvein; m-cu, mediocubital crossvein; r, radial crossvein; r-m, radiomedial crossvein; III–VII, abdominal sternites III–VII.

Figure 2. Line drawing Auritibicen cf. flammatus (Distant, 1892), KYFSI253. A, habitus, ventral side; B, head, pronotum and mesonotum, dorsal side; C, forewing; D, hindwing. Abbreviations: a1–a8, apical cells 1–8; CuA, cubitus anterior; CuA₁, first branch of the cubitus anterior vein; M, median vein; m, medial crossvein; M₁–M₄ = first to fourth branch of the median vein; mc, medial cell; m-cu, mediocubital crossvein; n, node; r, radial crossvein; rc, radial cell; r-m, radiomedial crossvein; RP, radius posterior; R+Sc, radius+subcostal vein; u1–u3, ulnar cells 1–3; I–VII, abdominal sternites II–VII. All scale bars indicate 5 mm.

Specimen.—KYFSI253, consisting of part and counterpart slabs preserving ventral (KYFSI253a: Figure 1A) and dorsal (KYFSI253b: Figure 1B) portions of the body, respectively. The visible surface of KYFSI253a reveals the head, prosternum, mesosternum, opercula, right forewing, partial right hindwing, partial right foreleg, and abdominal sternites II–VII. The head, pronotum, and mesonotum are visible on KYFSI253b.

Locality and horizon.—The specimens were collected from a laminated siltstone horizon of the Middle Pleistocene (Chibanian) Miyajima Formation (Shiobara Group) exposed in a fossil quarry located at the Konoha Fossils Museum (36°58′52″N, 139°48′23″E) in Nasushiobara, Tochigi, Japan.

Description.—A large-bodied male preserved from head to abdominal sternite VI. Preserved body length 35.80 mm. Estimated body length 43 mm. Estimated total length (head to forewing apex) 62 mm.

Head (Figures 1A, B; 2A, B): Left eye missing, slightly sclerotized, fuscous. Length 2.3 mm and width 16.3 mm (estimated including eyes). Antennae not preserved. Ocelli not visible. Clypeus slightly visible.

Thorax (Figures 1A, B; 2A, B): Pronotum twice length of head (length 4.7 mm and width 15.5 mm). Pronotal collar and cruciform elevation not preserved. Mesonotum sclerotized with continuous W-shaped marking (length 12.5 mm and width 13.5 mm) (Figures 1B, 2B).

Wings (Figures 1A; 2A, C, D): Hyaline without any infuscation. Forewing long and narrow with eight apical cells; apical and basal portion not preserved; estimated length 50 mm and maximum width 12.8 mm; vein C and vein R+Sc completely preserved and overlap thickly; part of vein M preserved near base; vein M₁₊₂ completely preserved and parallel to vein M₃₊₄;. r and r-m crossveins faintly preserved, base of apical cell 3 and 4 preserved (Figure 1E); m and m-cu crossveins preserved (Figure 1C, D). Only apex of hindwing preserved overlapping under forewing with six apical cells (Figure 2D).

Abdomen (Figures 1A, F; 2A): Cylindrical, abdominal sternites II to base of VII preserved, apices of sternites VII and VIII not preserved; spiracles present on right side of sternite III and from sternites IV to VII; sternites II and III widest (maximum width 18.7 mm) but boundary between them not clearly visible; sternites IV to VI almost same length. Opercula relatively short, slightly sclerotized with overlapping inner margins, rounded apically.

Discussion

KYFSI253 exhibits several diagnostic characteristics (Lee, 2015; Wang et al., 2018) that support its referral to the genus Auritibicen. These include a large, robust body; pronotum distinctly longer than the head; mesonotum with a central W-shaped marking; abdominal segment II wider than the head (including eyes); and overlapping inner margins of male opercula. The wings are hyaline, with the forewings showing eight apical cells and the hindwings six apical cells.

The genus Auritibicen was reviewed by Wang et al. (2018) and currently comprises 24 known species distributed throughout East Asia (primarily temperate regions), including Japan, South Korea, China, and Russia (Table 1).

Table 1. Comparison of male of the modern species of Auritibicen (referring to Wang et al., 2018) with the fossil Auritibicen cf. flammatus (This study). Body length (mm), forewing length (L: mm), forewing width (W: mm) and the ratio of L to W (L/W) in males. Opercula: S, not extending beyond posterior margins of abdominal sternite II; L, extending beyond posterior margins of abdominal sternite II. Infuscation: “―”, no infuscation; “*”, weak infuscation; “**”, distinct infuscation. + = estimated value. ++ = referring to Hayashi and Saisho, 2015.

Taxon	Body length (mm)	forewing length (L: mm)	forewing width (W: mm)	ratio of L to W (L/W)	opercula	infuscation				Distribution
						r, r‑m	m, m‑cu	base of a3	base of a4
Auritibicen aethus Wang et al., 2018	44.6	51.7	16.6	3.1	L	**	**	**	**	China (Zhejiang, Fujian)
Auritibicen atrofasciatus (Kirkaldy, 1909)	28.6–30.3	34.5–36.9	12.0–12.5	3.0	S	**	*	―	―	China (Sichuan, Yunnan, Fujian, Tibet)
Auritibicen bihamatus (Motschulsky, 1861)	35.4–36.2	43.0–43.4	14.2–14.5	3.0	L	**	**	**	**	Japan, Russia
Auritibicen chujoi (Esaki, 1935)	34.3–34.4	41.0–43.0	12.8–14.6	2.9	S	*	*	*	*	Taiwan
Auritibicen curvatus Wang et al., 2018	38.1–42.5	46.5–52.5	14.7–17.0	3.1	L	**	*	*	*	China (Shaanxi)
Auritibicen daoxianensis Wang et al., 2018	38.2	47.7	14.8	3.2	L	**	―	―	―	China (Hunan)
Auritibicen esakii (Kato, 1958)	34.9	42.1	14.2	3.0	L	**	**	**	**	Japan (endemic to Yakushima Is. south of Kyushu)
Auritibicen flammatus (Distant, 1892)	39–44++	49.7–52.0	16.6–17.4	3.0	S	**	*	*	*	China (Hubei, Sichuan, Shaanxi), Japan, Korea
Auritibicen cf. flammatus (Distant, 1892)	43+	50+	12.8	3.9	S	―	―	―	―	Japan (This study)
Auritibicen flavomarginatus (Hayashi, 1977)	35.3	40.9	13.3	3.1	L	**	*	―	―	Taiwan
Auritibicen gracilis Wang et al., 2018	32.6–33.5	37.2–37.5	12.5–13.1	2.9	S	**	―	―	―	China (Sichuan)
Auritibicen intermedius (Mori, 1931)	36.8	41.3	13.8	3.0	S	**	**	*	―	Russia (Primorsky Region), Korea
Auritibicen jai (Ouchi, 1938)	44.4	51.6	18.2	2.8	L	**	―	―	―	China (Hebei, Shaanxi, Zhejiang, Jiangxi)
Auritibicen japonicus (Kato, 1925)	40–46++	51.9–52.1	16.0–17.7	3.0	L	**	**	**	**	China (Shaanxi, Jiangxi), Japan, Korea
Auritibicen kyushyuensis (Kato, 1926)	34–40++	42.3–44.1	14.0–15.2	2.9	S	**	**	*	―	Japan
Auritibicen leechi (Distant, 1890)	38.0–42.6	45.3–47.7	15.5–16.0	3.0	L	**	**	―	―	China (Sichuan, Yunnan)
Auritibicen lijiangensis Wang et al., 2018	32.5	37.2	12.8	2.9	S	*	―	―	―	China (Yunnan)
Auritibicen pallidus Wang et al., 2018	35.7	45.7	15.2	3.0	L	*	―	―	―	China (Sichuan)
Auritibicen parvus Wang et al., 2018	28.4	35.1	11.3	3.1	L	*	―	―	―	China (Sichuan)
Auritibicen pekinensis (Haupt, 1924)	34.2	40.2	13.4	3.0	S	*	*	―	―	China (East Tibet)
Auritibicen purus Wang et al., 2018	31.0–31.8	36.2–39.4	11.5–12.4	3.2	L	―	―	―	―	China (Yunnan)
Auritibicen rotundus Wang et al., 2018	38.4	48.6	16	3.0	S	**	―	―	―	China (Hubei)
Auritibicen septatus Wang et al., 2018	38.6	47.7	15.7	3.0	L	*	―	―	―	China (Yunnan)
Auritibicen slocumi (Chen, 1943)	35.2	40.7	14.1	2.9	L	**	*	*	―	China (Sichuan, Guangxi)
Auritibicen tsaopaonensis (Chen, 1943)	32.1–33.0	38.0–39.4	13.1	3.0	L	**	**	**	**	China (Sichuan, Yunnan)

The opercula of KYFSI253 are incompletely preserved, and the boundary between abdominal sternites II and III is not visible. However, as the opercula are relatively short and rounded apically, they likely do not extend beyond the posterior margins of abdominal sternite II. According to Wang et al.’s (2018) key, six Auritibicen species have overlapping, apically rounded male opercula that do not extend beyond the posterior margins of abdominal sternite II: A. chujoi (Esaki, 1935), A. flammatus (Distant, 1892), A. intermedius (Mori, 1931), A. kyushyuensis (Kato, 1926), A. lijiangensis Wang et al., 2018 and A. rotundus Wang et al., 2018. Of these, A. intermedius and A. lijiangensis are clearly distinguishable from KYFSI253 due to their small body size. Furthermore, A. rotundus can be distinguished from KYFSI253 by its discontinuous W-shaped markings. KYFSI253 is therefore most comparable to both A. flammatus and A. kyushyuensis. However, A. kyushyuensis is smaller and exhibits distinct infuscation on the m and m-cu crossveins. Therefore, morphologically, the fossil most closely resembles A. flammatus, a species currently found across Japan, the Korean Peninsula, and China (Hubei, Sichuan and Shaanxi) (Wang et al., 2018).

However, the elongated forewings of KYFSI253 distinguish it from other species. Whereas the length-to-maximum width ratio ranges from 2.8 to 3.2 in all other species, this ratio is 3.9 in KYFSI253 (Table 1). Nevertheless, this difference may be due to deformation during fossilization and cannot serve as the sole distinguishing feature.

Another difference between KYFSI253 and A. flammatus lies in the infuscation morphology. A. flammatus exhibits infuscation on the r and r-m crossveins, with weaker infuscation on the m and m-cu crossveins and the bases of apical cells 3 and 4 (Wang et al., 2018). In contrast, no infuscation was observed on any of the veins of KYFSI253. When cicada fossils are well preserved, their infuscation and other markings are often clearly visible. For example, Graptopsaltria inaba from the upper Miocene of Tottori, Japan has clearly preserved markings (Fujiyama, 1982). Similarly, Meimuna sp. from the upper Pliocene of Hyogo, Japan has clearly preserved infuscation, and both Auritibicen bihamatus (Motschulsky, 1861) and Yezoterpnosia nigricosta (Motschulsky, 1866) from the same locality of the Shiobara Group as KYFSI253 also show clearly preserved infuscation (Fujiyama, 1979; 1982). Based on this evidence, it is likely that if infuscation had been present on this fossil, it would be preserved. Therefore, it is inferred that the taxon represented by KYFSI253 lacks infuscation, or if present, it was faint in life. However, designating KYFSI253 as a new species based on the absence of infuscation alone is not sufficient, as individual or geographical variations may be present. In fact, A. flammatus is known to exhibit many individual variations (Hayashi and Saisho, 2015). Therefore, we tentatively identify the fossil as A. cf. flammatus.

Wang et al. (2018) conducted phylogenetic analyses of Auritibicen using both morphological and molecular data. In the morphological analysis, they selected Chremistica ochracea (Walker, 1850) as the outgroup, whereas in the molecular analysis, they selected Lyristes plebejus (Scopoli, 1763) and Lyristes gemellus (Boulard, 1988) as outgroups. In both analyses, A. flammatus was recovered as the earliest species to diverge from the outgroup. Based on molecular data, this divergence was estimated to have occurred approximately 12.16 million years ago. All three outgroup species lack infuscation on their forewings (Wang et al., 2018). This suggests that the fossil may represent an ancestral form of A. flammatus predating the development of infuscation after diverging from the outgroup.

The genus Auritibicen includes 18 species from East Asia, most of which occur in the mountainous regions of southeastern China: eight species in Sichuan and six species in Yunnan (Wang et al., 2018). These species have restricted distributions and are considered Ice Age relicts (Wang et al., 2018). Among 14 species from this region, 12 lack or exhibit weak infuscation on the m and m-cu crossveins. This suggests that the absence or reduction of infuscation may represent a primitive trait within this genus.

In summary, KYFSI253 most closely resembles A. flammatus, but differences in infuscation suggest that it may represent an ancestral form of this species. According to Wang et al. (2018), many Auritibicen species diversified during the Pleistocene, and discovery of this fossil from the Middle Pleistocene bolsters that hypothesis. KYFSI253 provides crucial evidence for the early diversification of Auritibicen and broader cicadid evolution.

Acknowledgements

We would like to express our gratitude to Chika Shinomiya for providing this fossil and to Maho Shinomiya and Yuno Shinomiya for discovering it. Additionally, we are grateful to the editor and the two anonymous reviewers for their critical comments on this manuscript.

Author contributions

H. A. initiated the study, drafted the manuscript, and compiled all the figures. M. H. revised the manuscript and provided taxonomic input. All authors contributed to the writing of the manuscript.

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