Female-biased froghoppers (Hemiptera, Cercopoidea) from the Mesozoic of China and phylogenetic reconstruction of early Cercopoidea

Abstract

Cercopoidea is a diverse insect group, but its early evolution, disparity and ecology remain unclear. Juroala daohugouensis Chen & Wang gen. et sp. nov., from the Middle–Upper Jurassic of north-eastern China, is established herein and described on the basis of 42 whole-bodied fossils, representing a new subfamily, Juroalinae subfam. nov. of the primitive family Sinoalidae. Chengdecercopis Hong, 1983 is transferred from Procercopidae to this new subfamily and Stictocercopis Fu & Huang, 2018 is also attributed to the new subfamily. Fangyuanini Chen & Wang trib. nov. is erected for Fangyuania Chen, Szwedo & Wang, 2018 from mid-Cretaceous Burmese amber. Our cladistic analyses recover the relationships within the Cercopoidea as follows: (Juroalinae + (Sinoalini + Fangyuanini)) + (Procercopidae + (Cercopionidae + five modern families)). The evolutionary history and morphological diversification in the Mesozoic is discussed based on our phylogenetic reconstruction. In addition, the abundant material in the present study not only indicates high intra-specific or even intra-individual variation caused by biological and/or taphonomic factors, but also reveals an extremely female-biased propensity, suggesting that the new taxon probably had adaptations to special palaeoenvironments in physiology, ecology and ethology (e.g. parthenogenesis and sociability).

http://zoobank.org/urn:lsid:zoobank.org:pub:5EDD7D33-D509-4330-A01F-F41DF664E1C2

Keywords:

• Cercopoidea
• Mesozoic
• taxonomy
• phylogeny
• female bias

Acknowledgements

The authors sincerely thank Limei Lin, Junqiang Zhang and Yameng Li for their constructive comments on an earlier version of the manuscript. This research was supported by the National Natural Science Foundation of China (41502007; 41472023; 41702012; 41572010; 41622201; 41688103), the Natural Scientific Foundation of Shandong Province (ZR2013DQ017; ZR2016DB24), and China Postdoctoral Science Foundation (2015M580480). This is a Leverhulme Emeritus Fellowship contribution for EAJ.

Supplemental material

Supplemental material for this article can be accessed at: https://doi.org/10.1080/14772019.2019.1587526

Figure 5 Details of Juroala daohugouensis Chen & Wang gen. et sp. nov. A, head of paratype NIGP169641. B, right antenna of paratype NIGP169648. C, right antenna of paratype NIGP169663. D, mouthpart of paratype NIGP169663. E, right foreleg of paratype NIGP169648. F, right middle leg of paratype NIGP169648. G, right hind tibia of paratype NIGP169660. H, left hind tibia of paratype NIGP169660. I, male pygofer of paratype NIGP169650. J, female pygofer of paratype NIGP169663. B–J under alcohol. Probable joints between segments of antennae are indicated by arrows. Lateral spines on tibia are indicated by triangles. Abbreviations: tyl., tylus; cro., crown; oce., ocellus; sca., scape; ped., pedicel; fla., flagellum; pos., postclypeus; ros., rostrum; fem., femur; tib., tibia; tar., tarsus; VII, sternite VII; VIII, sternite VIII; pyg., pygofer; gv., genital valve; gs., genital style; ovi., ovipositor. Scale bars = 1 mm.

Figure 6 Scatter plots showing morphometric statistics of tegmina of Juroala daohugouensis Chen & Wang gen. et sp. nov. A, bivariate plots of wing length and width. B, ratios of branching position of longitudinal veins and mid-point position of crossveins to tegmen length. BC, ScP, R, MP, CuA, MP_1 + 2, MP_3 + 4: branching position of vein MP + CuA (basal cell length), RA + ScP, R + ScP, MP, CuA, MP_1 + 2, MP_3 + 4/tegmen length; ir, r-mp, imp, mp-cua: mid-point positon of crossvein ir, r-mp, imp, mp-cua/tegmen length. ‘N’ represents the sample size.

Figure 7 The most parsimonious tree inferred from maximum parsimony analysis with character mapping. White circles indicate homoplasious characters and black circles indicate non-homoplasious characters. Numbers above branches are character numbers, and below branches are character state changes. Labelled nodes (hollow circles) refer to specific clades discussed in the text.

Associate Editor: Vladimir Blagoderov