Phylogeny, molecular species delimitation, and biogeography of the cryptic subgenus Ulopsina (Hemiptera: Cicadellidae: Ulopinae)

Abstract Ulopinae are flightless or flight-limited, making them an ideal model insect group for investigating biogeographical history. Within Ulopinae, the subgenus Ulopsina Dai, Viraktamath and Zhang is the most speciose subgenus within the genus Ulopsina, which is itself the most speciose genus in the tribe Ulopini. This study presents the first molecular phylogenetic and biogeographical analyses of the subgenus Ulopsina based on four genes. Species delimitation analyses were performed by combining five molecular delimitation methods with morphological evidence. In addition, ecological niche modeling was used to simulate suitable habitat for the subgenus Ulopsina during the Last Glacial Maximum, the Mid-Holocene, the present day and into the future. Our results confirm the status of three described species of the subgenus Ulopsina and support the recognition of three new species: U. (U.) biprocessasp. n., U. (U.) furcatasp. n. and U. (U.) serratasp. n. and classified these species into two new species groups. Dating and biogeography analyses indicate that the subgenus Ulopsina likely originated during the early Miocene in southern Yunnan and southeast Xizang. Most speciation events in the subgenus Ulopsina occurred in the late Miocene, possibly triggered by several dispersal and vicariance events and influenced by the uplift process of the Qinghai-Tibet Plateau and by climate change. Ecological niche modeling results indicate that the precipitation of warmest quarter and isothermality are two key ecological factors influencing the current distribution pattern of the subgenus Ulopsina. As the climate continues warming, the subgenus Ulopsina may expand northward in subtropical China.