Research Article |
Corresponding author: Dong Ren ( rendong@mail.cnu.edu.cn ) Academic editor: Fabian Haas
© 2021 Shurong Xiong, Michael S. Engel, Lifang Xiao, Dong Ren.
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.
Citation:
Xiong S, Engel MS, Xiao L, Ren D (2021) New archidermapteran earwigs (Dermaptera) from the Middle Jurassic of Inner Mongolia, China. ZooKeys 1065: 125-139. https://doi.org/10.3897/zookeys.1065.72720
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Two new species of Archidermaptera are described and figured from the Middle Jurassic Jiulonghsan Formation of Daohugou, Inner Mongolia, China. Aneuroderma oiodes gen. & sp. nov. is described in the family Protodiplatyidae and Sinopalaeodermata concavum sp. nov. is established in the family Dermapteridae. Both new species share the typical characters of the extinct suborder Archidermaptera (e.g., pentamerous metatarsi, filiform and multimerous cerci, externalized ovipositor). Aneuroderma gen. nov. is compared with other genera of the Protodiplatyidae, while S. concavum sp. nov. allows us to emend the diagnosis of the genus Sinopalaeodermata. We briefly discuss the diversity of Archidermaptera and challenges to understanding relationships among this mid-Mesozoic diversity.
Dermapteridae, earwigs, new genus, new species, Protodiplatyidae, systematic palaeontology
The Dermaptera (earwigs) are, like all organisms, an interesting mosaic of primitive and derived traits – on the one hand they have a typical ‘orthopteroid’ habitus with chewing mouthparts, while on the other extant species have specialized cerci modified as forceps; a vestigial ovipositor; reduced tarsal count; a shortened and tegminized forewing; a unique hind wing composed of a greatly enlarged anal fan, reduced remigium, and distinctive folding pattern; and have lost ocelli (
Excluding the Protelytroptera, the earliest definitive earwigs are classified in the suborder Archidermaptera, distinguished from other suborders by the pentamerous metatarsi, frequent presence of venation in the forewing tegmina, which are often longer, and a prominent externalized ovipositor (
Hierarchical classification of Archidermaptera and summary of tarsal formulae, where known. Interrogative marks (?) indicate missing data.
Genus | Species | Data | |||
---|---|---|---|---|---|
Suborder Archidermaptera Bey-Bienko, 1936 | Superfamily Protodiplatyoidea Martynov, 1925 | Family Dermapteridae Vishniakova, 1980 | Genus Brevicula Whalley, 1985 | B. gradus Whalley, 1985 | 4/5-4/5-5* |
B. maculata |
?-?-? | ||||
Genus Dacryoderma Engel, 2021 | D. teres (Tihelka, 2019) | ?-?-? | |||
Genus Dermapteron Martynov, 1925 | D. incerta Martynov, 1925 | ?-?-? | |||
Genus Dimapteron |
|||||
D. corami |
?-?-? | ||||
Genus Jurassimedeola Zhang, 2002 | J. orientalis Zhang, 2002 | ?-?-? | |||
Genus Palaeodermapteron |
P. dicranum |
5-5-5 | |||
Genus Phanerogramma Cockerell, 1915 |
P. australis |
?-?-? | |||
P. dunstani |
?-?-? | ||||
P. gouldsbroughi |
?-?-? | ||||
P. heeri (Giebel, 1856) | ?-?-? | ||||
P. kellyi Tihelka, 2019 | ?-?-? | ||||
Genus Sinopalaeodermata Zhang, 2002 | S. concavum sp. nov. | 5-5-5 | |||
S. neimonggolense Zhang, 2002, nom. emend. | 5-5-5 | ||||
Genus Trivenapteron |
T. moorei |
?-?-? | |||
Genus Valdopteron |
V. woodi |
?-?-? | |||
Family Protodiplatyidae Martynov, 1925 | Genus Abrderma |
A. gracilentum |
?-?-5 | ||
Genus Aneuroderma gen. nov. | A. oiodes sp. nov. | 5-5-5 | |||
Genus Archidermapteron Vishniakova, 1980 | A. martynovi Vishniakova, 1980 | 4-4-5 | |||
Genus Asiodiplatys Vishniakova, 1980 | A. speciosus Vishniakova, 1980 | 4-4-5 | |||
Genus Barbderma |
B. oblonguatum |
?-?-? | |||
Genus Longicerciata Zhang, 1994 | L. mesozoica Zhang, 1994 | 5-5-5 | |||
L. rumpens Zhang, 1994 | 5-5-5 | ||||
Genus Microdiplatys Vishniakova, 1980 | M. campodeiformis Vishniakova, 1980 | 4-4-5 | |||
M. oculatus Vishniakova, 1980 | 4-4-5 | ||||
M. perfectus Vishniakova, 1985 | 4-4-5 | ||||
Genus Perissoderma |
P. triangulum |
5-5-5 | |||
Genus Protodiplatys Martynov, 1925 | P. fortis Martynov, 1925 | 4-4-5 | |||
P. gracilis Vishniakova, 1980 | 4-4-5 | ||||
P. mongoliensis Vishniakova, 1986 | 4-4-5 | ||||
Genus Sinoprotodiplatys |
S. ellipsoideuata |
5-5-5 | |||
S. zhangi |
5-5-5 | ||||
Family Turanoviidae Engel, 2003 (Turanodermaptera Engel, 2019) | Genus Turanovia Vishniakova, 1980 | T. incompleta Vishniakova, 1980 | ?-?-? |
Herein we describe a new genus and species of Protodiplatyidae and a new species of Dermapteridae, both preserved in the Middle Jurassic Jiulongshan Formation of Inner Mongolia Province, China. This discovery increases the diversity of Archidermaptera and complements our limited understanding of this suborder.
Three specimens were collected from the Middle Jurassic Jiulongshan Formation at Daohugou Village, Ningcheng County, Inner Mongolia, northeastern China. The age of the fossil deposit is approximately 164–165 Ma (
Suborder Archidermaptera Bey-Bienko, 1936
Moderate-sized earwigs, with numerous setose and distinctively sculptured (densely punctate-granulose throughout, particularly on head and thorax). Head broad, nearly as wide as anterior border of pronotum, posterior margin nearly straight. Antenna with 20 antennomeres; scape robust and slightly broader than remaining antennomeres; pedicel slightly longer than wide; all flagellomeres longer than wide. Compound eyes large and situated at posterior temples; ocelli absent. Dorsal surface without Y-shaped ecdysial cleavage scar. Pronotum approximately oval, anterior and posterior margins subequal in width, lateral margin convex and rounded. Tegmina without longitudinal veins; tegmina and squamata covering abdominal segment II. Legs with abundant short setae; femora carinulate; all tarsi pentamerous (i.e., tarsal formula 5-5-5 rather than the 4-4-5 of some genera); pretarsal claws simple. Female with exposed ovipositor. Pygidium small. Cerci filiform and long, with about 30 cercomeres.
The generic name is a combination of the Greek prefix a– (ᾰ–, alpha privativum designating negation), neûron (νεῦρον, meaning, “nerve”), and dérma (δέρμᾰ, genitive dérmatos, meaning, “skin” – an allusion to the leathery tegmina and from which the ordinal name is derived, Dermaptera literally meaning, “skin wings”), referencing the absence of tegminal venation, a rare feature among Archidermaptera. The gender of the name is neuter.
As for the genus (vide supra).
Holotype, a completely preserved female,
Jiulongshan Formation (Middle Jurassic); Daohugou Village, Wuhua Township, Ningcheng County, Inner Mongolia, China.
Adult female, preserved in both dorsal and ventral aspects. Body with numerous setae and distinctively sculptured (densely punctate-granulose throughout, particularly on head and thorax). Total length as preserved (excluding antennae, ovipositor, and cerci) about 10.75 mm. Head medial length from clypeal apex to posterior border 1.57 mm, maximum width (across level of compound eyes) 1.56 mm, prognathous; maxillary palpus pentamerous, ca 1.33 mm long (Fig.
Holotype of Aneuroderma oiodes gen. et sp. nov.
The specific epithet is the Greek neuter adjective ōiôdes (ᾠῶδες, meaning, “oval” or “egg-like”), as a reference to the ovoid pronotum.
The new genus is placed within Protodiplatyidae on the basis of the characteristic filiform antenna with 17–23 antennomeres; pedicel and flagellomere I subequal in size; pentamerous metatarsus; cerci elongate, slender, and multimerous; and externalized ovipositor in females.
Holotype of Aneuroderma oiodes gen. et sp. nov.
Key diagnostic characters of Aneuroderma gen. nov. are summarized in Table
Summary and comparisons of key characters for several described genera of Protodiplatyidae. Interrogative marks (?) indicate data unavailable.
Genus | Head | Number of antennomeres | Pronotum | Tegmina | Pygidium | Cerci (ratio lengths cercus/body, cercomeres) |
---|---|---|---|---|---|---|
Abrderma | Broader than pronotum | 17–19 | Elliptical, anterior and posterior margins subequal in width | With venation, reaching second abdominal segment | Small | 0.2, ? |
Archidermapteron | Narrower than pronotum | 17–19 | Reniform, broad notch anteriorly | With venation, reaching fourth abdominal segment | Transverse, trapeziform | 1.0, at least 40 |
Asiodiplatys | Narrower than pronotum | 22 | With shallow, broad notch anteriorly | Without venation, not reaching second abdominal segment | ? | 0.5, 40 |
Barbderma | ? | 19 | Oblong or trapezoidal, anterior and posterior margins subequal in width | With venation, reaching first abdominal segment | Small | 0.42, ? |
Longicerciata | Broader than pronotum | 26 | Transverse, anterior margin wider than posterior margin | Without venation, reaching fourth abdominal segment | ? | 1.0, at least 36 |
Microdiplatys | Broader than pronotum | 19 | Transverse | With venation, reaching fourth abdominal segment | Not protruding | 1.0, at least 36 |
Perissoderma | Narrower than pronotum | 17 | Elliptical, anterior margin wider than posterior margin | With venation, reaching third abdominal segment | Small | 0.5, ? |
Protodiplatys | Narrower than pronotum | 17–18 | Transverse, notch in front, broadly rounded posteriorly | With venation, reaching fourth abdominal segment | Transverse, trapeziform | 0.5, no more than 40 |
Sinoprotodiplatys | Narrower than pronotum | 18 | Anterior and posterior margins subequal in width | Without venation, reaching fourth abdominal segment | ? | 0.8, 20 |
Aneuroderma gen. nov. | As wide as pronotum | 20 | Oval, anterior and posterior margins subequal in width | Without venation, reaching second abdominal segment | Small | 0.5, 30 |
Sinopalaeodermata Zhang, 2002: 351. Type species: Sinopalaeodermata neimonggolensis Zhang, 2002, nomen imperfectum [recte neimonggolense], by original designation.
[modified from
Included species. Aside from the type species, the genus currently includes S. concavum Xiong, Engel & Ren, sp. nov. (infra).
The new species can be distinguished from the type species, Sinopalaeodermata neimonggolense (note that the name Sinopalaeodermata is neuter, not feminine, as dérmata is the neuter nominative plural of dérma; and given that the specific epithet is adjectival it must still agree in gender with the generic name) by the relatively straight apical margin of the penultimate sternum (in S. neimonggolense the penultimate sternum has a concave margin); the roughly reniform pronotum, with the anterior margin concave medially the posterior margin weakly convex, and lateral margins rounded (in S. neimonggolense the pronotum is approximately rectangular, with the anterior margin almost as wide as the posterior margin, and the lateral margins relatively straight and parallel to each other); the tegmina with a more pronounced concave arc marginally at the apex of Rs (in S. neimonggolense the margin is more sloping rather than deeply concave); and M does not extend to near the apex of CuA , with CuP terminating more proximal to CuA (even before the tangent with M) (in S. neimonggolense M terminates more proximally and CuP extends to the apex of CuA).
Holotype. A completely preserved female,
Jiulongshan Formation (Middle Jurassic); Daohugou Village, Wuhua Township, Ningcheng County, Inner Mongolia, China.
Adult female, preserved in both dorsal and ventral aspects. Total length as preserved (excluding antennae, cerci, and valvulae) about 18.02 mm. Body with sparse pubescence and punctate. Head medial length from clypeal apex to posterior border 2.08 mm, maximum width (across level of compound eyes) 2.27 mm, triangular. Compound eye large, ovate, located near posterior margin of head; compound eye length 0.91 mm; width between compound eyes 2.58 mm. Ocelli comparatively small. Cervix with large anterior and posterior cervical sclerites, anterior sclerite slightly larger than posterior sclerite. Pronotum approximately reniform, medial length 1.46 mm, maximum width 2.58 mm, anterior width 1.64 mm, posterior width 2.07 mm, anterior margin concave and posterior margin arched, lateral margins convexly rounded. Mesoscutellum large, elliptical, entirely exposed. Tegmina present, not truncated, length 6.73 mm, maximum width 2.55 mm, with medially sinuate anterior (lateral) margin and straight posterior (mesal) margin. Veins simple, Rs curved anterior margin, fading out just before margin; M simple, basally and apically straight, gently curved medially; Cu with two branches (CuA and CuP), CuP terminates proximal to CuA; A1 and A2 simple and straight, running parallel to each other and posterior margin, terminating apically (Fig.
The specific epithet is taken from the Latin adjective concavus (meaning, “concave”), in reference to the more pronounced concave margin to the tegmina relative to the type species.
Both of the new species described herein are easily recognized as archidermapterans owing to the pentamerous metatarsi (and more than three pro- and mesotarsomeres); elongate, flexible, and multimerous cerci; and the externalized ovipositor. Jurassic earwigs are known mostly from four assemblages: the Jiulongshan flora of the Middle Jurassic, the England flora of the Early and Late Jurassic, and the Karatau flora of the Late Jurassic (
Tarsal formulae have been used to distinguished significant groups of fossil Dermaptera. All Neodmerpatera and Eodermaptera have three tarsomeres (3-3-3), while the number of tarsomeres is greater, where known, among Archidermaptera (Table
Based on three well-preserved fossil specimens from the Middle Jurassic, we describe a new genus and two new species, Aneuroderma oiodes gen. et sp. nov. (Protodiplatyidae) and Sinopalaeodermata concavum sp. nov. (Dermapteridae). We make extensive comparisons with other Archidermaptera and discuss challenges in understanding relationships among these Triassic and Jurassic lineages.
We express our gratitude to the Editorial Board of Zookeys, and in particular Dr Fabian Haas. We thank Dr Petr Kocarek and another anonymous reviewer for their valuable comments on this manuscript. We are grateful to Yue Mao (Capital Normal University) for helpful discussions and advice with the project. D.R. was supported by grants from the National Natural Science Foundation of China (grant nos. 31730087, 32020103006 and 41688103). The authors declare no competing interests.