Taxonomic review and phylogenetic inference elucidate the evolutionary history of Mesozoic Procercopidae, with new data from the Cretaceous Jehol Biota of NE China (Hemiptera, Cicadomorpha)

Abstract

The Mesozoic family Procercopidae is widely treated as the ancient group of Cercopoidea and a transitional unit to recent lineages, but its evolution and diversity are vague due to fragmentary fossil record and confusing taxonomic history. Herein, an extensive taxonomic review of Procercopidae is presented and some new fossils are reported from the Lower Cretaceous Yixian Formation of NE China. As a result, Chengdecercopis Hong, 1983 is transferred from Procercopidae to Sinoalidae; Procercopis longipennis Becker-Migdisova, 1962 and P shawanensis Zhang, Wang and Zhang, 2003 are transferred to Procercopina Martynov, 1937, resulting in Procercopina longipennis (Becker-Migdisova, 1962), comb. n. and P shawanensis (Zhang, Wang and Zhang, 2003), comb. n.; Luanpingia senjituensis Hong, 1984 is transferred to Stellularis Chen, Yao and Ren, 2015, leading to Stellulari senjituensis (Hong, 1984), comb. n.; Anthoscytina macula Hu, Yao and Ren, 2014 is transferred to Sinocercopis Hong, 1982, and Sunoscytinopteris (Scytinopteridae) and Cathaycixius (Cixiidae) are treated as junior homonym names of Sinocercopis, leading to Sinocercopis macula two new species are erected: Stellularis bineuris Chen and Wang, sp. n. and S minutus Chen and Wang, sp. n. Our cladistic analysis based on wing (tegmen and hind wing) characteristics recovers the high-level relationships within Cercopoidea: Sinoalidae + (Procercopidae + (Cercopionidae + modern cercopoids)). Within the family Procercopidae, the cladistic analysis reveals that the Middle to Late Jurassic Titanocercopis and Jurocercopis and the Cretaceous Cretocercopis occupy the basal position, and a gradual change in wing venation can be recognized from the Early Jurassic Procercopis and Procercopina to the Jurassic Anthoscytina, and then to the Cretaceous Stellularis and Sinocercopis. The two Cretaceous genera, sharing wing traits with extant | 175 CHEN Et al.

New data on Jurassic Procercopidae (Hemiptera: Cercopoidea) from northeastern China

“…Cretaceous procercopids such as Sinocercopis and Stellularis became further reduced the MP to single‐branched (Chen, Wang, et al, 2015). Anthoscytina is related to Cretaceous Sinocercopis and Stellularis by similar shape of tegmen and reduced venation, but distinctly differs from two Cretaceous genera in bearing a hind wing retained two‐branched MP (hind wing MP unbranched in Sinocercopis and Stellularis ) (Chen et al, 2020). Most of the Jurassic procercopids share the hind wing with MP bearing two free branches, but Cretaceous procercopids possess the hind wing with MP 3+4 fused with MP 1+2 apically in Cretocercopis from the Lower Cretaceous of China, or MP unbranched such as Burmocercopis from the mid‐Cretaceous Burmese amber (Ren, 1995; Chen et al, 2020; Fu et al, 2020), which confirm vein MP reduced in number is one of the trends of evolution of froghoppers.…”

“…Most of the Jurassic procercopids share the hind wing with MP bearing two free branches, but Cretaceous procercopids possess the hind wing with MP 3+4 fused with MP 1+2 apically in Cretocercopis from the Lower Cretaceous of China, or MP unbranched such as Burmocercopis from the mid‐Cretaceous Burmese amber (Ren, 1995; Chen et al, 2020; Fu et al, 2020), which confirm vein MP reduced in number is one of the trends of evolution of froghoppers. Meanwhile, the phylogenetic relationship within Procercopidae was proposed by Chen et al (2020), using a Bayesian analysis based on 32 morphological characters. The result shows that Procercopina and Procercopis occupied the basal position, and Anthoscytina with sister relationship to three Cretaceous genera (i.e., Sinocercopis , Stellularis , Cercopion ) and modern Cercopoidea (Chen et al, 2020).…”

“…Meanwhile, the phylogenetic relationship within Procercopidae was proposed by Chen et al (2020), using a Bayesian analysis based on 32 morphological characters. The result shows that Procercopina and Procercopis occupied the basal position, and Anthoscytina with sister relationship to three Cretaceous genera (i.e., Sinocercopis , Stellularis , Cercopion ) and modern Cercopoidea (Chen et al, 2020).…”

Insect‐plant‐fungus interactions in mycorrhizal associations, with a focus on spittlebugs and ectomycorrhizal host plants

“…Spittlebugs appear in the fossil record in the earliest Jurassic, about 200 mya, and probably date to the late Triassic (Chen et al, 2019). This is consistent with molecular phylogeny estimates that spittlebugs diverged from cicadas, their closest living relatives, about 225 mya (K. P. Johnson et al, 2018, fig.…”

“…S1). Several spittlebug lines existed in the Jurassic and Cretaceous periods (Chen et al, 2019), one eventually giving rise to extant lineages 75 to 110 mya in the mid‐Cretaceous (Shcherbakov, 2002), followed by diversification of modern forms in the Paleogene (Chen et al, 2019; K. P. Johnson et al, 2018, fig. S1; Paladini et al, 2018, figs.…”