Integrative Taxonomy of the Spinous Assassin Bug Genus Sclomina (Heteroptera: Reduviidae: Harpactorinae) Reveals Three Cryptic Species Based on DNA Barcoding and Morphological Evidence

SIMPLE SUMMARY: The assassin bugs (Heteroptera: Reduviidae) are one of the largest and morphologically most diverse families of true bug, having essential impacts on forest ecosystems as predators. The spinous reduviid genus Sclomina exhibits shape mimicry and protective coloration adapted to the spinous Rubus plant that they inhabit. The genus Sclomina shows gradual morphological variability, so its morphological classification is still unresolved, and its biology is almost unknown. In this study, DNA barcodes and morphological evidence were combined to accurately divide the species of a comprehensive collection sampled in South China and North Vietnam. We found three cryptic species. The biological information and mimicry behavior uncover their successive evolutionary survival strategies. ABSTRACT: Sclomina Stål, 1861 (Heteroptera: Reduviidae: Harpactorinae) is endemic to China and Vietnam, with only two species, Sclomina erinacea Stål, 1861 and Sclomina guangxiensis Ren, 2001, characterized by spinous body and dentate abdominal connexivum. However, due to variable morphological characteristics, Sclomina erinacea, which is widely distributed in South China, is possibly a complex of cryptic species, and Sclomina guangxiensis was suspected to be an extreme group of the S. erinacea cline. In the present study, we conducted species delimitation and phylogenetic analyses based on the mitochondrial cytochrome c oxidase subunit I (COI) gene sequences of 307 Sclomina specimens collected from 30 sampling localities combined with morphological evidence. The result showed that all samples used in this study were identified as five species: Sclomina guangxiensis is a valid species, and Sclomina erinacea actually includes three cryptic species: Sclomina xingrensis P. Zhao and Cai sp. nov., Sclomina pallens P. Zhao and Cai sp. nov., and Sclomina parva P. Zhao and Cai sp. nov. In this paper, the genus Sclomina is systematically revised, and the morphological characteristics of the five species are compared, described, and photographed in detail. We elucidate the evolutionary history of Sclomina based on results of estimated divergence time. The body shape and coloration (green in nymph and brown in adult) of Sclomina match their environment and mimic the Rubus plants on which they live. The symbiotic relationship between Sclomina and spinous Rubus plants is presented and discussed.